MEMOIRES

DU MUSEUM

NATIONAL

D’HISTOIRE

NATURELLE

TOME 172

ZOOLOGIE

1997

Resultats des Campagnes MUSORSTOM

Volume 16

CAMPAGNE

FRANC O-IND ONE SIENNE

KARUBAR

Coordonne par

Alain CROSNIER & Philippe BOUCHET

Publie avec le concours du Ministere des Affaires Etrangeres

Source : MNHN, Paris

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Ce volume des Resultats des Campagnes MUSORSTOM est dedie a M. Jean-Michel CHASSER1AUX qui,

alors Delegue aux Affaires Internationales au Ministere de la Recherche et de la Technologie, a assure de 1989

a 1993 la presidence frangaise du Comite mixte franco-indonesien en oceanologie.

Pendant cette periode, il s'est attache au developpement de cette cooperation et a soutenu l' organisation de

nombreuses campagnes d la mer, dont la campagne KARUBAR, sur les navires "Banina Java" qui venaient d'etre

livres par la France a I'Indonesie.

Professeur d'Universite en mathematiques, Jean- Michel CHASSER1AUX est actuellement Directeur des

Relations Internationales de I'INRIA (Institut National de Recherche en Infonnatique et en Automatique).

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, nett., (A), 133 : 1-525, 126 fig., 37 pi. (1986). ISBN : 2-85653-136-9.

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

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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.

Source : MNHN, Paris

resultats dcs campagnes

Volume 16

Source : MNHN, Paris

ISBN : 2-85653-506-2

ISSN : 1243-4442

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MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE

TOME 172

ZOOLOGIE

Resultats des Campagnes MUSORSTOM

Volume 16

CAMPAGNE FRANCO-INDONESIENNE

KARUBAR

Coordonne par

Alain CROSNIER & Philippe BOUCHET

Musdum national d'Histoire naturelle

Laboratoire de Biologie des Invertebres marins et Malacologie

55 rue Buffon

75005 Paris

Publie avec le concours du Ministere des Affaires Etrangeres

EDITIONS

DU MUSEUM

PARIS

1997

Source :

SOMMAIRE

Pages

1. La campagne Karubar en Indonesie, au large des lies Kai et Tanimbar . 9

Alain CROSNIER, Bertrand RICHER DE FORGES & Philippe BOUCHET

2. Cnidaria Anthozoa : Scleractiniaires sans zooxanthelles des Philippines

et d'Indonesie (en anglais) . 27

Stephen D. Cairns & Helmut Zibrowius

3. Mollusca Bivalvia : Pectinoidea (Propeamussiidae et Pectinidae) de I'lndonesie

orientale (en anglais) . 245

Henk H. DlJKSTRA & Woro W. KASTORO

4. Mollusca Gastropoda : Les Muricidae recoltes lors de la campagne Karubar

en Indonesie orientale (en anglais) . 287

Roland HOUART

5. Mollusca Gastropoda : Cancellariidae de la mer d'Arafura recoltes durant

la campagne KARUBAR (en anglais) . 295

Andre Verhecken

6. Mollusca Gastropoda : Nouveaux Turridae (Conoidea) de Test de I'lndonesie

(en anglais) . 325

Alexander SYSOEV

7. Mollusca Cephalopoda : Pieuvres bathyales (200-1000 m) des mers de Banda

et d'Arafura (Octopodidae et Alloposidae) (en anglais) . 357

Mark D. Norman, F.G. Hochberg & C.C. Lu

8. Crustacea Decapoda : Stylodactylidae recoltes en Indonesie, aux lies Wallis

et Futuna et au Vanuatu (campagnes Karubar. MUSORSTOM 7 et 8). Donnces

com p le men ta ires sur les Stylodactylidae recoltes en Nouvelle-Caledonie . 385

Regis Cleva

9. Crustacea Decapoda : Palinuridae, Scyllaridae et Nephropsidae recoltes

en Indonesie lors de la campagne KARUBAR. Cle d'identification des especes

du genre Metanephrops (en anglais) . 409

Tin-Yam CHAN

10. Crustacea Decapoda : Pagures de la famille des Paguridae recoltes lors de

la campagne Karubar en Indonesie (en anglais) . 433

Patsy A. McLaughlin

11. Crustacea Decapoda : Parapaguridae recoltes lors de la campagne KARUBAR

en Indonesie. Description de deux especes nouvelles (en anglais) . 573

Rafael Lemaitre

12. Crustacea Decapoda : Especes des genres Agononida Baba & de Saint Laurent, 1995

et Munida Leach, 1820 (Galatheidae) recoltees lors de la campagne Karubar

(en anglais) . 597

Enrique Macpherson

13. Crustacea Decapoda : Ethusinae (Dorippidae) recoltes principalement lors

de la campagne Karubar (en anglais) . 613

Chen Huilian (H. L. Chen)

14. Echinodermata Crinoidea : Les Pentacrines recoltees lors de la campagne Karubar

en Indonesie . 627

Nadia AMEZIANE

Source : MNHN Paris

CONTENTS

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

The Karubar Cruise in Indonesia, off the Kai and Tanimbar Islands (in French)

Alain Crosnier, Bertrand Richer DE FORGES & Philippe Bouchet

Cnidaria Anthozoa: Azooxanthellate Scleraetinia from the Philippine

and Indonesian Regions .

Stephen D. CAIRNS & Helmut ZIBROWIUS

Mollusca Bivalvia: Pectinoidea (Propeamussiidae and Pectinidae) from eastern

Indonesia .

Henk H. DlJKSTRA & Woro W. KASTORO

Mollusca Gastropoda: The Muricidae collected during the Karubar Cruise

in eastern Indonesia .

Roland HOUART

Mollusca Gastropoda: Arafura Sea Cancellariidae collected during the KARUBAR

Cruise .

Andre VERHECKEN

Pages

. 9

.... 27

... 245

287

295

Mollusca Gastropoda : New deep-water turrid gastropods (Conoidea) from eastern

Indonesia . . .

Alexander SYSOEV

Mollusca Cephalopoda : Mid-depth octopuses (200-1000 m) of the Banda

and Arafura Sea (Octopodidae and Alloposidae) .

Mark D. NORMAN, F.G. HOCHBERG & C.C. LU

Crustacea Decapoda: Stylodactylidae collected in Indonesia, the Wallis

and Futuna Islands and the Vanuatu (Karubar, MUSORSTOM 7 and 8 cruises).

Additional information on the Stylodactylidae from New Caledonia (in French)

Regis CLEVA

Crustacea Decapoda: Palinuridae, Scyllaridae and Nephropidae collected in

Indonesia by the KARUBAR Cruise, with an identification key for the species of

Metanephrops .

Tin-Yam CHAN

Crustacea Decapoda: Hermit crabs of the family Paguridae from the KARUBAR

Cruise in Indonesia .

Patsy A. McLaughlin

Crustacea Decapoda : Parapaguridae from the Karubar Cruise in Indonesia,

with description of two new species .

Rafael Lemaitre

Crustacea Decapoda: Species of the genera Agononida Baba & de Saint Laurent,

1995 and Munida Leach, 1820 (Galatheidae) from the Karubar Cruise

Enrique Macpherson

Crustacea Decapoda: Ethusinae (Dorippidae), mainly from the Karubar Cruise . 613

Chen Huilian (H. L. CHEN)

Echinodermata Crinoidea: Pentacrinidae collected during the KARUBAR Cruise

in Indonesia (in French) .

Nadia Ameziane

Source :

*ESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 16 - RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 - RESULT AT.'

La campagne KARUBAR en Indonesie,

au large des iles Kai et Tanimbar

Alain CROSNIER* Bertrand RICHER DE FORGES **

cfc

Philippe BO U CHET *

* Laboratoire de Biologie des Invertebres marins et Malacologie

Museum national d'Histoire naturelle

55 rue Buffon. 75005 Paris

** ORSTOM

B.P. A5, Noumea Cedex

Nouvelle-Caledonie

RESUME

La eampagne tranco-indonesienne Karubar, faite h bord du navire de recherche indondsien "Baruna Java I", s’est

deroulde dans 1 est de l indones.e. en mer de Banda et d'Arafura. au large des lies Kai et Tanimbar. Les prospections ont

ont dt^effectuiJs6 ba‘hya C' Quatre-ving>-onze dragages et chalutages. & des profondeurs comprises entre 200 et 1200 m,

ABRIDGED ENGLISH VERSION

The Karubar cruise in Indonesia, off the Kai and Tanimbar Islands.

Despite a long-established scientific reputation as a hotspot of marine species richness, the seas of Indonesia remain

P°° y .k.n°Wn ln lerms of their deep-sea fauna. Since the landmark “ Siboga " expedition a century ago, comparatively

FvnJo I13 ‘keP;searexPloratlon has been carried out in the archipelago. A remarkable exception was the Danish

Th Mill, u tn°wusPelled, Ka'J Islands, conducted in 1922 by an equally remarkable man. Professor

MfipTrwvrv’ ^N" 00 h°ard the Ambolna ■ an ''old' deep-draught, primitively equipped vessel” (Wolff, 1967).

hr fnirnH .h„Sc?1mn PurPose,was 10 flnd a suitable location for the establishment of a Scandinavian tropical station. When

,,1°. " . Slalked cnn0lds. elasipods and other deep-sea creatures occurred as shallow as 200-400 m around the Kai

nmii'ri suggested that this was an ideal place to build a marine laboratory to study the abyssal fauna. Morthnsen’s

projeci was never realised, but his dream has remained alive in the heart of many marine zoologists to this day.

Kai^TS,nin?K^A "/RlC aE^DE Forges' B- & Bouchet, P„ 1997. — La campagne Karubar en Indondsie, au large des Iles

nntn Hi_. bi J"' A0 ^°oN1ER * P- BouCHET (eds)- Resultats des campagnes MUSORSTOM, Volume 16. Mem. Mus.

natn. Hist, nat., 172 : 9-26. Pans ISBN 2-85653-506-2.

Source : MNHN, Paris

10

A. CROSNIER, B. RICHER DE FORGES & P. BOUCHET

In the 1980s, new opportunities for collaborative oceanographic research in partnership with Indonesian scientists

were formalised through a joint agreement between the Indonesian and French governments. A series of three research

vessels, "Banina Jaya" 1, 2 and 3 were built in France and became operational in Indonesia in 1990-91. As we submitted

to the ad hoc committee a proposal for deep-sea biological exploration, our suggestion to target the Kai Islands met the

approval of the Indonesian government, which insisted that the seas of Eastern Indonesia had to receive special

attention. The Karubar (a contraction of the names of the Kai, Aru and Tanimbar Islands) project was thus bom.

The purpose of the expedition was to:

(a) document the composition of the deep-sea fauna and

(b) investigate potential economic resources in the untapped deep-sea benthos, with special emphasis on shrimps.

This dual goal was reflected in the composition of the scientific staff, which included zoologists as well as fisheries

biologists from several Indonesian and French institutions: Pusat Penelitian dan Pengembangan Oseanologi LIP1 (P30

LIPI) [Institute of Oceanology of the Indonesian Academy of Sciences], Balai Penelitian Perikanan Laut

(BALITKANLAUT) [Indonesian Institute of Fisheries], Badan Pengkajian dan Penerapan Teknologi (BPPT) [Indonesian

Ministry of Science and Technology], the Institut Fran^ais de Recherche Scientifique pour le Ddveloppement en

Cooperation (ORSTOM) [French Institute of scientific Research for Development through Cooperation], and the Musdum

National d'Histoire Naturelle (MNHN) [French National Museum of Natural History]. A total of 15 Indonesian and 7

French staff took part in the expedition, with Dr K. Moosa (P30 LIPI) as senior scientist and Lt. Col. Handoko the

commanding captain (see Appendix 1 for composition of staff).

The expedition was initially planned to depart from Ambon on October 7, 1991 and last 27 days. However, a series of

incidents, partly technical, delayed departure until October 21 and, as a result, only 13 days of work were available. Hence

our initial programme could not be carried out in full and we had to cancel the transects near the Aru Islands. A total of

91 stations, of which 85 were successful, were carried out near the Kai and Tanimbar Islands, at depths between 200 and

1200 m: 18 dredge hauls and 19 trawls near Kai. 10 dredges and 44 trawls near Tanimbar (List of stations: Appendix 2).

Incidentally, this is exactly the same number of hauls carried out by the "Siboga" expedition at depths below 200 m,

which points to the importance of the Karubar expedition, despite its short duration. It should be noted, however, that

the "Siboga" sampled many more stations in very deep water: 30 "Siboga" stations are deeper than 1000 m. versus only

5 Karubar stations at such depths.

The 13 papers included in the present volume provide much new information about Cnidaria (Scleractinia), Mollusca

(Bivalvia, Gastropoda and Cephalopoda), Crustacea (Decapoda) and Echinodermata (Crinoidea). In addition, over

20 papers published elsewhere are based wholly or in part on the zoological collections made during Karubar (List of

publications: Appendix 3). Undoubtedly, they are many more to come, which will document and describe the still little-

known fauna of the Indonesian archipelago. From a zoological point of view. Karubar can therefore be considered to

have been very successful, and we may only wish for further such cruises and volumes of expedition reports.

From the point of view of fisheries, however, the results of the Karubar expedition have been somewhat

disappointing. Considering the general geomorphology of the region, we did not expect vast trawlable bottoms near the

Kai islands, but the bathyal slope East of the Tanimbars appeared much more promising, especially in the context of

commercial fisheries off the Australian northwest shelf. Indeed, we encountered extensive flat areas suited for commercial

trawling. However, populations of echinoderms (holothurians, echinoids and ophiuroids) are so abundant that they

constitute the main catch. Furthermore, the very soft muddy bottoms and strong currents do not facilitate trawling

operations. We did encounter a number of potentially valuable commercial species of crustaceans (peneids: Aristeus,

Penaeopsis, Haliporoides, Metapenaeopsis, Hymenopenaeus, Hadropenaeus', and pandalids: Heterocarpus, Plesionika),

but they are apparently never abundant. One exception might be species of Metanephrops at depths between 250 and

300 m, but our initial results need to be confirmed by commercial deep-sea trawlers.

As noted above, Mortensen (1923) had suggested that the Kai Islands would be "an ideal place” for a tropical marine

laboratory, because of the “rich and varied fauna of genuine abyssal forms [occuring] over the whole of the large plateau

of 2-400 m depth". Are the results of the Karubar expedition in line with Mortensen's enthusiasm? At this stage, only

part of the zoological material has been studied and the results are rather contrasting. For instance, Cairns & Zibrowius

(this volume) report the highest regional diversity of azooxanthellate Scleractinia: 125 species were collected near the

Kai Islands versus 69 species, for example, near Lubang Island, Philippines, which has been intensively sampled during

the Musorstom 1, 2 and 3 expeditions. Also, considering the difference in sampling effort, the diversity in species of

Pectinoidea (Mollusca, Bivalvia) appears greater in the Arafura Sea than in New Caledonia (DlJKSTRA, this volume).

Conversely, several nominal species of pentacrinid crinoids previously recorded from the area are now shown to be

ecophenotypes and the pentacrinid crinoid fauna consequently appears less diverse than that recorded elsewhere in

Indonesia (Am£ziane, this volume). Probably such contrasting patterns of species richness in different zoological groups

only reflect preferences for different bottom types.

The occurrence of stalked crinoids in comparatively shallow water, which had so much impressed Mortensen, is

confirmed by the Karubar expedition, with the shallowest record of Saracrinus at 290 m (245 m by Mortensen). This is

admittedly very shallow by temperate North Atlantic standards, but is not exceptional in tropical waters. For instance,

pentacrinids occur from 290 m and deeper in New Caledonia (Bourseau et al., 1991) and from 185 m and deeper in the

Source : MNHN. Paris

CAMPAGNE KARUBAR

1 1

Philippines (Bourseau & Roux, 1989). In fact, we suspect that the Kai Islands appeared so exceptional to Mortensen

because, despite being a much-travelled marine biologist, he had only limited experience of tropical deep-sea faunal

assemblages. In the eyes of a European zoologist, stalked crinoids were abyssal animals and their occurence in shallow

waters called for a special explanation. Instead, we suggest that the occurrence of stalked crinoids, as well as other

markers of the deep-sea fauna, such as elasipods and echinothurids, in the 200-500 m depth interval is the norm at

tropical latitudes in the Indo-Pacific. The pectinoid bivalves provide limited evidence that the shallowest occurrence of

certain species is shallower in the Arafura Sea than in New Caledonia, through the opposite is true for other species. In

conclusion, based on the evidence available and our own field experience in other tropical Indo-Pacific regions we regard

the Kai Islands as a rich, but not exceptional, place.

La campagne Karubar, appellation provenant d'une contraction des noms des lies Kai, Aru et Tanimbar,

toutes situees en mers de Banda et d'Arafura dans lesquelles s'est effectuee la campagne, a ete progranmiee dans le

cadre de la cooperation franco-indonesienne en oceanographie et cofinancee par les deux parties.

Les objectifs principaux de cette campagne etaient l'etude de la faune bathyale et une premiere estimation des

ressources de la pente continentale en crevettes et poissons commercialisables, dans Test de l'lndonesie.

La campagne, prevue pour 27 jours, s'est faite sur le navire indonesien de recherches "Banina Jaya /",

magmltque unite construite en France par la CMN, de 60 m de longueur, jaugeant 700 tx. et dont c'etait la

premiere campagne de recherche en biologie.

Le navire pouvant embarquer, sans difficult^, une vingtaine de scientifiques, l’equipe indonesienne etait

nombreuse et regroupait quatre chercheurs du BBPT (Badan Pengkajian dan Penerapan Teknologi) que Ton peut

assimiler a noire Ministere de la Recherche et de la Technologie, six du BALITKANLAUT (Balai Penelitian

Perikanan Laut) qui correspond a notre ancien Office des Peches et cinq du P30 LIPI (Pusat Penelitian dan

Pengembangan Oseanologi LIPI) qui est 1'Institut d'Oceanologie dependant de 1'Academie des Sciences

indonesienne, plus un de la Direction des Peches.

12

A. CROSNIER, B. RICHER DE FORGES & P. BOUCHET

Du cotd fran^ais, on trouvait trois chercheurs de l'ORSTOM et trois chercheurs du Museum national d'Histoire

naturelle, a Paris, plus un maitre d'equipage de l'ORSTOM venu mettre son savoir-faire a la disposition de

l'equipage indondsien pour le greement et la manoeuvre des engins de peches (dragues, chaluts h perche et chalut a

panneaux). La presence de ce dernier devait se reveler d'autant plus utile que l'equipage du navire etait exclusivement

compose de militaires, le navire etant arme par la Marine nationale indonesienne.

Fig. 2. Quelques-uns des membres de l'6quipage et de I'equipe scientifique. De gauche it droite : Kapten Daryento

commandant en second; Kasim Moos a, chef de mission; Mayor Goenadi, chef mecanicien; k moitie cach<§ Kapten

Sarwono; Dwi Listyo Rayahu, carcmologiste; Yunus Soselisa, ichtyologiste; Lt. Col. Handoko. commandant;

Zaenal Arifin; Burhanudin. ichtyologiste; W.W. Kastoro. malacologiste; de dos Michel Potier. ichtyologiste-

Alain CROSNIER, responsable de l'dquipe franfaise; Mashiwara; Ali Kusnin; Albert Le Crom, maitre d’equipage

DEROULEMENT DE LA CAMPAGNE

A I origine le Baruna Jaya 1 " devait appareiller de Jakarta le 2 octobre et rejoindre Ambon en cinq jours de mer.

La mission scientifique devait embarquer alors et la campagne debuter le 7 octobre.

Mais ll n en a pas ete ainsi a la suite de toute une serie de contre-temps : defaillance d'un transitaire livrant le

materiel envoye de France avec 19 jours de retard, lenteurs administratives pour I'obtention du visa de travail du

maitre d equipage venu de Nouvelle-Caledonie, petits problemes materiels divers avec les engins du bord (poulie

compteuse, treuil, sondeurs) ont retardd l'appareillage de Jakarta jusqu'au 12 octobre. II etait alors decide que deux

des chercheurs fran?ais et le maitre d'equipage feraient la traversee Jakarta- Ambon a bord du " Baruna Jaya I" afin

d assurer la mise en place de tout le materiel durant la traversee.

Le 17 octobre, le "Baruna Jaya 1 " arrivait a Ambon et I'equipe embarquee etait complete le 19 octobre au

matin, avec I amvee du chef de I'equipe indonesienne. L'appareillage etait alors prevu pour le soir meme a 17h00.

CAMPAGNE KARUBAR

13

Cependant des problemes d’avitaillement cn eau et en vivres, ainsi qu'une aimable invitation du Commandant de

la Marine a Ambon, le 19 au soir, obligeaient de remettre a nouveau l'appareillage qui, ne pouvant avoir lieu un

dimanche, etait alors fixe au 21 et a 7h00 du matin, les appareillages de nuit etant interdits.

Finalement cet appareillage, un peu laborieux, avait bien lieu h cette date et a cette heure.

Lors de 1 etablissement du programme de campagne 27 jours de mer avaient ete prdvus, mais une fois en mer, le

Commandant nous prevenait que le navire ne pouvait rester plus de 14 jours en mer, car au dete l'eau douce

viendrait a manquer. Si Ton enlevait les 3 jours de mer necessaires aux transits, la duree de travail utile se trouvait

ramende h 1 1 jours, ce qui etait loin du programme etabli. Finalement un rationnement de l'eau etant accepte, la

duree de la sortie etait portee a 19 jours, ce qui permettait d'avoir 16 jours de travail en mer.

En fait cette duree a, par la suite, etd ramenee h 13 jours, un message de I'Amiraute indonesienne ayant demande

au Commandant de se derouter, lors du retour, sur Timor pour y recuperer un engin suspect trouvd en mer (qui s'est

revele etre un courantometre enregistreur).

Durant ces 13 jours, nous avons bendficie d'une mer pratiquement toujours calme, ce qui a considerablement

facilite les operations de dragages et de chalutages, qui etaient nouvelles pour l'equipage.

Plusieurs types d’engins ont dte utilises :

— drague Waren,

— drague epibenthique,

— chalut a perche,

— chalut h crevettes ^ panneaux.

Le travail ddbutait ^ 5h00 le matin pour s'arreter, au moins en ce qui concemait l'equipage, a 22h00.

Ftci. 3. Un chalut vient d'etre remonte : premiers tris sur la plage arridre. Au premier plan, de gauche h droite : Ali

Kusnin, Albert Le Crom, Michel Potier, Alain Crosnier, Aznam Azis, Philippe Bouchet, Lt. Col. Handoko.

14

A. CROSNER. B. RICHER DE FORGES & P. BOUCHET

Le 22 octobre a 8h00, ayant bendficie de courants favorables et la machine ayant ete un peu poussee, lc "Baruna

Java I" parvenail aux lies Kai, au sud de I'ile Taam, et nous effectuions le premier dragage, cntre 156 et 305 m, au

voisinage des stations 192 du "Challenger" et 46 de l'expedition danoise aux ties Kai, faite en 1922 et dirigee par le

Professeur MORTENSEN. Ces stations ont ete signalees comme ayant permis des recoltes zoologiques

particulierement riches. En fait, le fond compose de sable detritique grassier avec des debris de coraux et de

nombreux articles d 'Halimeda, nous fournissait une faune intdressante mais pas aussi riche que celle recoltee par le

"Challenger" et MORTENSEN.

Fig. 4. — Bernard M£tivier gree une drague Waren.

On peut d'ailleurs mentionner ici que nos rdcoltes n'ont que tres rarement correspondu h cclles indiquees par

MORTENSEN. En particulier dans des zones signalees comme riches par MORTENSEN, il nous a souvent ete

impossible de mettte un engin a l'eau, compte tcnu de la nature tourmentee des fonds. Ceci est-il du a des

positionnements peu exacts ?

La journee du 22 octobre etait consacree a une serie de sept dragages et chalutages a perche dans cette zone. Les

fonds s'y revelaient assez rugueux mais possibles a travailler, encore que les traits de chalut a perche n'aient guere

pu exceder 15 minutes.

Durant la nuit, nous empruntions le chenal separant les lies Tayandu des Petites Kai qui se montrait tres

accidente et reconnaissions, au sondeur, la zone situee au nord de ce chenal. Une cuvette chalutable de 10 milles sur

10 milles, a des profondeurs variant de 300 a 400 metres, y etait reperee; elle est toutefois parcourue, vers

132°35'E, par une faille nord-sud dont la profondeur atteint 80 m et qui doit done etre evitee.

La journee du 23 octobre etait consacree a l'exploration de cette cuvette. Le chalutagc s'y revelait facile (nous

avons pu y effectuer un trait de chalut k panneaux d'une heure sans probleme). Les recoltes y ont ete diversifies

mais si, en particulier, de nombreux crustaces commercialisables : langoustines ( Metanephrops ), crevettes peneides

( Penaeopsis , Haliporoides, Metapenaeopsis, Hymenopenaeus , Hadropenaeus), Pandalidae ( Heterocarpus ,

Plesionika) y etaient bien representees, aucune espece n'a ete trouvee en abondance.

Les 24 et 25 octobre, nous allions vers l'est et explorions la zone se trouvant au nord du chenal separant les

Petites Kai de la Grande Kai (6 dragages dont 1 epibenthique, 4 chalutages a perche, 1 chalutage a panneaux). Cette

zone se montre aisement dragable et chalutable par endroits et plus difficilement & d'autres. Parfois la presence

Source : MNHN. Pans

CAMPAGNE KARUBAR

15

d'enormes buttes de vase, notamment vers 600 m, a provoque l'interruption du trait. Cette zone ne fournissait pas

d'indices plus encourageants que la precedente en ce qui conceme les especes commercialisables; bien entendu.

comme elle s’etend a des profondeurs superieures a la precedente, les grosses crevettes peneides ( Aristeus ) y sont

presentes, de meme que l’enorme langoustine Metanephrops neptunus, mais pas, d'apres ce que nous avons pu voir,

en quantites commerciales.

Le 26 octobre, nous explorions le grand

chenal scparant les Petites Kai de la Grande Kai

(6 dragages, 3 chalutages a perche), dont la

profondeur, dans sa partie la plus resserree,

n'excede pas 400 m. Les fonds, couverts d’une

vase gluante avec souvent des affleurements

rocheux, se montrerent peu interessants. Par

ailleurs de forts courants, variables, rendirent les

manoeuvres difficiles ne nous incitant pas a

perseverer dans cette zone. Lors de cette journee,

la drague, crochee, ne put etre recuperee qu'apres

plus d'une heure d'efforts.

La journee du 27 octobre etait consacree a

une serie de 5 chalutages a perche dans le sud des

ties Kai, sur des fonds souvent assez accidentes

et peu faciles a travailler, mais aussi, parfois,

reguliers. C'est ainsi que les chalutages CP 35

et CP 36 ont pu durer une heure chacun, a des

profondeurs variant entre 390 et 500 m pour le

premier et 210 a 270 m pour le second,

permettant d'cxcellentes et abondantes captures.

De meme que dans le chenal, de forts courants

ont, dans cette zone, perturbe les operations.

Les journees passant, il etait alors decide de

rallier les Ties Tanimbar, seconde zone dont

l'expioration nous avait ete assignee. Ce transit

etait effectue dans la nuit du 27 au 28 octobre.

Trois radiales Etaient prevues dans la zone

s'etendant au sud-est de ces Ties.

Les fonds y different totalement de ceux

Fig. 5. Arrivee sur le pont d'une drague Warfen pleine de vase. rencontres aux Ties Kai. D'une maniere generate,

aux profondeurs explores (200 a 1550 m,

maximum de la profondeur dans cette zone) nous nous trouvions devant une pente douce et reguliere, avec toutefois

quelques variations notamment entre 850 et 1200 m ou la pente devient frequemmeent plus raide. Si les fonds

etaient durs aux profondeurs les plus faibles, ils etaient recouverts de vase a partir de 250-310 m. Cette vase, grise,

est souvent tres molle et rend les chalutages difficiles, d'autant que de forts courants (jusqu'a 2,7 nceuds) dont la

force se modifie rapidement, rendent difficiles le maintien du cap et surtout de la vitesse du navirc. Ces variations

de vitesse ont d'ailleurs provoque la perte de tout un train de chalutage, les panneaux s'etant enfouis dans la vase a

la suite d'une diminution trop importante de la vitesse.

Dix dragages, dont 3 epibenthiques, et 44 chalutages, dont 6 a panneaux, ont ete effectues dans cette zone.

Parmi les fonds les plus spectaculaires, on peut citer ceux a Virgularia (Pennatulacea) vers 290 m, ceux a

rachiopodes et scleractiniaires trouves entre 350 et 450 m essentiellement et ceux a echinodermes, tres nombreux

et diversifies. Vers 400-450 m, les holothuries, dont les molpadides et les elasipodes, etaient particulierement

abondantes, tandis que vers 800-1000 m on trouvait de tres nombreux oursins (Echinidae, Echinothuridae, Cidaridae

Spatangidae) et vers 1000 m des fonds a ophiures (Ophiuridae) et Hyalonoecia (Polychacta).

16

A. CROSNIER, B. RICHER DE FORGES & P. BOUCHET

Les crinoides (. Saracrinus ), celebres dans la litterature par les recoltes du "Challenger aux lies Kai, retrouvtfs

dans ces memes ties par l'expedition KARUBAR jusqua 430 m de profondeur, n'ont ete trouves au large des .les

Tanimbar que jusqu'h 300 m de profondeur, ceci s'expliquant sans doute par la nature des fonds, une vase tres

molle, au dela de cette profondeur. • .

Dans toute la vaste plaine de vase ainsi exploree, de nombreuses especes de crustaces qui pourraient etre

commercialisms ont 6l6 recoltes, notamment plusieurs especes de langoustmes ( Metanephrops arafurensis ,

M. neptunus, M. sibogae, M. velutinus), mais aucune n'a malheuseusement 6t6 prise en quantile significative.

Le 5 novembre au soir le travail en mer se terminait, comme nous l'avons mentionne plus haut, plus tot que

prevu. Le 6 novembre le "Banina Jaja 1 " etait de retour a Ambon et les equ.pes scientifiques debarquaient.

RESULTATS

Quatre-vingt-onze chalutages et dragages, dont quatre-vingt-cinq ont dte reussis, ont ete faits entre 200 et

1200 m.

Du point de vue des peches commercials, les nSsultats sont ddcevants. S'il etait peu Evident de trouver des

zones chalutables suffisamment etendues aux Ties Kai, compte tenu de la geomorpholog.e de cette zone, on pouyait

esperer, par contre, que la pente bathyale prolongeant le plateau continental de l'lnan Jaya, & Test des lies

Tanimbar, serait prometteuse. Certes les zones chalutables y sont vastes, mais la presence d'une vase grise souvent

molle et gluante, l'existence de forts courants et la presence de tres importantes concentrations d echinodermes

(holothuries, oursins et ophiures) encombrant les fonds et colmatant les chaluts, ne sont pas faits pour rendre

rentable une peche industrielle. Des especes interessantes se trouvent sur ces fonds, mais ll semblerait que ce soit

toujours en quantite assez faible. Ce n'est que dans la partie sud de la zone prospectee que des apparences de

langoustines, peut-etre un peu plus encourageantes, ont ete observees sur les fonds de 250 it 300 m. Ceci dit, il ne

faut pas oublier les conditions dans lesquelles nous avons travaille et il est bien certain qu'une campagne faite par

un chalutier mene par des professionnels de la peche profonde, comme il en existe en Australie, serait maintenant

souhai table.

Source : MNHN, Paris

CAMPAGNE KARUBAR

17

Si les resultats ont dtd decevants au plan du ddveloppement des peches, il n'en est pas de meme au plan de la

connaissance de la faune bathyale.

Avant nous plusieurs grandes expeditions ont travailld sur la faune d'eau profonde de l'lndonesie • celles du

"CMlenser (1872-1876), de la "Siboga" (1899-1900), de la "Galathea" (1950-1952), de Mortensen aux lies Ka“

(1922), sans oublier la campagne Corindon (1980) et l'expddition Snellius 2 en 1984 Beaucoup de ces

expddmons n'ont fan que traverser l'lndonesie, en y effectuant un nombre rdduit ^operations de peche. Quant i la

Siboga qui i est la plus connue en ce qui concerne 1'Indondsie, elle n'a effectud qu'un nombre de peches profondes

comparable k celui atteint par la campagne Karubar. Cette simple comparison situe bien rapport de la campagne

Karubar k notre connaissance de la faune bathyale indondsienne, encore qu'il faille la nuancer : si la "Siboga" n'a

pas fait, durant son people de pres de deux ans, plus de stations au del* de 200 m que nous en 13 jours elle a

effectud environ 30 stations h plus de 1000 m contre 5 seulement en ce qui concerne Karubar.

Le matdriel recoltd a dtd reparti de la manure suivante :

— la totahte des poissons ainsi, dans les dchinodermes, que la totality des astdries ont dtd conservdes par les

chercheurs mdondsiens.

— le reste du materiel a ete expedid pour tri et dtude au Musdum national d'Histoire naturelle, etant entendu

qu aprds etude une partie significative des rdcoltes serait renvoyde, identifiee, en Indonesie pour servir de collection

de rdference.

Fig. 7. — Deux trieuses achamdes : W.W. Kastoro h gauche, Dwi Listyo Rahayu il droite.

Les venues en France de chercheurs indondsiens pour participer aux tris et aux dtudes dtaient programmees. Au

moment ou nous dcnvons ces lignes, deux chercheurs sont ainsi venus, 1'un a deux reprises.

es avant la publication du prdsent volume, de nombreux travaux, essentiellement sur les crustaces, ont deiil

consacrds, en tout ou partie, k l'etude des recoltes de la campagne Karubar. Les 25 articles parus, dont la lisle

flen fnneXe,’ confl™ent I'onginalitd et l'interet de la faune recoltee. S'y ajoutent maintenant les 13 articles

T UtreS travaux SOnt en C0urs de r^daclion e‘ des collections triees attendent des preneurs

Lh/r r?5' ans quelques anndes’ de la campagne Karubar quant a la connaissance de la faune

oatnyale indondsienne apparaitra pleinement.

f('923),avait su^re que les lies Kai serait un endroit ideal pour un laboratoire consacre a la

Of .h 81 T tr0pica e’ k cause de "the nch and varied fauna of genuine abyssal forms [occuring] over the whole

of the large plateau of 2-400 m depth”. Les resultats de la campagne Karubar appuient-.ls ce, enthousiasme de

Source :

18

A. CROSNIER. B. RICHER DE FORGES & P. BOUCHET

MORTENSEN ? Actuellement seule une partie des recoltes zoologiques ont 6l6 etudiees et les rdsultats sont quelque

peu contradictoires. Par exemple Cairns et Zibrowius (dans ce volume) mentionnent le grand nombre d’espdces

de scleractiniaires sans zooxanthelles trouvees dans cette region : 125 espdces ont ete recoltdes pres des ties Kai

contre environ 69 au voisinage de l'tle de Lubang, aux Philippines, ou de nombreuses recoltes ont ete faites lors

Fig. 8. — Quelques-uns des participants & la campagne. De gauche & droite : 1. Le Let. Kol. Handoko, Alain Crosnier et

Mohammad Kasim Moosa. — 2. Le Professeur Jacques Forest et Dwi Listyo Rayayu examinant le rare pagure Tisea

grandis. — 3. Philippe BOUCHET et W.W. KaSTORO. — 4. Aznam AZIS, Michel Potier & Mohammad Kasim MOOSA. —

5. Bertrand Richer de Forges, au fond a gauche le Pr Jacques Forest. — Zaenal Arifin et Albert Le Crom.

Source : MNHN Paris

CAMPAGNE KARUBAR

19

des campagnes MUSORSTOM 1, 2 et 3. De meme, si Ton considere les densites differen.es de recol.es, le nombre des

especes de Pect.no.dea (mollusques bivalves) parait plus grand en mer d'Arafura qu'en Nouvelle-Caledonie

(DlJKSTRA, dans ce volume). Par centre, .1 a ete montre (Ameziane dans ce volume) que plus.eurs especes de

crinoides pedoncules (pentaennes) signalees de la region de lies Kai n'etaient en fait que des ecophenotypes si bien

que la faune : de : ce groupe, autour des lies Km, apparait moins diversif.ee que dans d'autres regions de l'Indonesie II

est vraiscmblable que ces resultats son., en fait, largement en liaison avec la presence de fonds de nature plus ou

moms diversifiee. K

MoRT^r^ crinoides pedoncules dans des eaux relativemen. peu profondes, qui avait tellement impressionne

MORTENSEN, est confirmee par la campagne Karubar avec la recolte de Saracrinus a 290 m (245 m par

MORTENSEN). De telles profondeurs son. tres faibles si Ton se refere a ce que I'on observe dans l'A.Ian.ique Nord

mais n ont rien d exceptionnel en mers tropicales. Par exemple des pentacrines on. ete trouves a partir de 290 m en

Nouvelle-Caledonie (BOURSEAU « aL, 1991) et de 185 m aux Philippines (Bourseau & Roux, 1989) II est

vraisemblable que MORTENSEN s'etait si fortement en.housiasme pour les lies Kai parce que, malgre ses nombreux

voyages en tant que biologiste mar.n, il n'ava.t qu'une experience limitee de la faune d'eau profonde tropicale Aux

yeux d un zoolog, ste europeen, les crinoides pedoncules etaient des animaux abyssaux e. leur presence en'eaux

relativement peu profondes necessity une explication particuliere, alors que leur presence de meme que celle

d autres marqueurs de la faune d'eau profonde, tels que les elasipodes et les echinothurides, est la norme entre 200 e.

500 m dans I Indo-Pacilique tropical. Les mollusque bivalves Pectinoidea ne fournissent pas d'evidence nette •

certaines especes se trouvent & des profondeurs moindres en mer d'Arafura qu'en Nouvelle-Caledonie, mais e'est le

contraire qui est observe pour d'autres. En definitive, en se basant sur les resultats disponibles, et egalement sur

notre experience d autres regions tropicales de I'Indo-Pacifique, nous sommes amenes a considerer les ties Kai

comme une region riche mais non exceptionnelle.

n -f" f0rme dC conclusi°n’ nous aimerions exprimer le souhait que la campagne Karubar ne demeure pas une

operation, somme toute bien reussie, mais unique. A la demande des autorites indonesiennes, une demande de

" r, TAR e" mCr dC T'm°r 3 ^ dt3blie en 0Ct0bre 1993- Evaluee favorablement du cote frames il

r PrnaireS indon*icns- Si le Pr6sem vo,ume' en a«iran, a nouveau

attention sur 1 interet de 1 etude de la faune bathyale indonesienne, pouvai. donner une nouvelle dynamique a ce

projet, il aurait alors joue un role au deli de toutes nos esperances.

REFERENCES

BOIJS;jeP/„Af R°HUX'r-M-' ~ Ech,noderma,a: Les Crinoides pedoncules de Nouvelle-

J'His,oire teS S sdr t R5S13il2a29d3e33 amPagneS MUS°RST°M' V°'Ume 8- Mim0lres du National

B° Resultats des Cairam 1989- ~ Echmodermes: Crinoides Pentacrinidae (Musorstom 2 & Corindon 2). In:

143:1 13-201. P 8 M S0RST0M’ Volume 4- Memoires du Museum National d'Hisioire Naturelle, Paris, ser. A,

DaniSh EXPedili°n ‘° ,he Kd ISl3ndS 1921 Videnskabelige fra Dansk Naturhistorisk

WOCopenhaRe'n96325^nDanSke Ekspedi,ioner p3 verdenshavene [Danish expeditions on the seven seas], Rhodos,

Source :

20

A. CROSNIER. B. RICHER DE FORGES & P. BOUCHET

ANNEXES

LISTE DES PARTICIPANTS A LA CAMPAGNE KARUBAR

Partie indonesienne

Chef de mission: M. Kasim Moosa (P30-LIPI).

Autres participants :

Appartenant au P30-LIPI :

Chercheurs : Aznam AZIS, BURHANUDIN, W.W. KASTORO, Dwi Listyo RAYAYU.

Appartenant au Balitkanlut :

Chercheurs : Bambang SUMIONO, MAHISWARA, Yunus SOSELISA.

Techniciens : Zaenal ARIFIN, Nasir MADJID.

Patron de Peche : Ali KUSNIN.

Appartenant au BPPT :

Chercheurs : Djunaedi Muljawan.

Techniciens : Abdul Haris, SURATMAN, Tri SETIADI.

Appartenant a la direction des Peches :

AZMI

Partie fran^aise

Appartenant a l'ORSTOM : Alain CROSNIER, Michel POTIER, Bertrand RICHER DE FORGES,

Albert LE CROM (maitre d'equipage).

Appartenant au Museum national d'Histoire naturelle, Paris : Philippe BOUCHET, Jacques FOREST,

Bernard Metiver.

Officiers indonesiens de P equipage du "Baruna Jaya 1"

Let. Kol. HANDOKO, commandant; Mayor GOENADI, chef mecanicien; Kapten DARYANTO, commandant en

second; Kapten SARWONO; Lettu Ishak ISKANDAR; Letda Budi SlSWANTO; Letda NURYADI; Letda Agus

Maryono; Letda Supendi; Serka Wachid Mullah.

LISTE DES STATIONS DE LA CAMPAGNE KARUBAR

Les majuscules se trouvant avant le numero de la station indiquent 1'engin utilise : DW : Drague Waren;

ED : drague epibenthique; CP : chalut a perche; CC : chalut a panneaux (crevettes)

Source : MNHN Paris

CAMPAGNE KARUBAR

Fig. 9. — Zones (hachurees) prospectees lors de la campagne Karubar.

Fig. 10. — Positions des stations effectives autour des lies Kai.

22

A. CROSNIER. B. RICHER DE FORGES & P. BOUCHET

Fig. 11. — Positions des stations effectuees autour des ties Tanimbar.

Source : MNHN, Paris

CAMPAGNE KARUBAR

24

A. CROSNER, B. RICHER DE FORGES & P. BOUCHET

LISTE DES PUBLICATIONS FAITES TOUT OU EN PARTIE

D'APRES LES RECOLTES DE LA CAMPAGNE KARUBAR

Mollusques

Sysobv, A. & BOUCHET, P., 1996. — Taxonomic reevaluation of Gemmuloborsonia Shuto, 1989 (Gastropoda: Conoidea),

with a description of new Recent deep-water species. Journal of Molluscan Studies, 62: 75-87.

Bouchet, P. & Sysoev, A., (sous presse). — Revision of the Recent species of Buccinaria (Gastropoda: Conoidea), a

genus of deep-water turrids of Tethyan origin. Venus, Japanese Journal of Malacology.

Source : MNHN . Paris

CAMPAGNE KARUBAR

25

Crustacea Cirripedia

Grygier, M. J. & CAIRNS, S. D., 1996. — Suspected neoplasm in the deep-sea corals (Scleractinia: Oculinidae:

Madrepora spp.) reinterpreted as galls caused by Petrarca madreporae n. sp. (Crustacea: Ascothoracida: Patrarcidae).

Diseases of aquatic Organisms, 24 : 61-69.

BUCKERIDGE J. S„ 1994. — Cirripedia Thoracica : Verrucomorpha of New Caledonia, Indonesia, Wallis and Futuna

Islands. In : A. Crosnier (ed.), Rdsultats des Campagnes MUSORSTOM, Volume 12. Memoires du Museum national

d'Histoire naturelle, 161 : 87-125.

Crustacea Amphipoda

Lowry, J. K. & Stoddart, H. E., 1993. — Crustacea Amphipoda : Lysianassoids from Philippine and Indonesian waters.

In : A. CROSNIER (ed.), Rdsultats des Campagnes MUSORSTOM, Volume 10. Memoires du Museum national d'Histoire

naturelle, 156 : 55-109.

Crustacea Pearacarida

Casanova, J.-P., 1996. — Crustacea Mysidacea : Les Lophogastridds d'Indondsie, de Nouvelle-Caledonie et des lies

Wallis et Futuna. In : A. Crosnier (ed.), Rdsultats des Campagnes Musorstom, Volume 15. Memoires du Museum

national d'Histoire naturelle, 168 : 125-146.

Crustacea Euphausiacea

Casanova, B., 1996. — Crustacea Euphausiacea : Euphausiacds du Pacifique sud-ouest tropical (Nouvelle-Caledonie, Ties

Wallis et Futuna, Indondsie). Morphologie fonctionnelle et Biogdographie. In : A. Crosnier (ed.), Rdsultats des

Campagnes MUSORSTOM, Volume 15. Memoires du Museum national d'Histoire naturelle, 168 : 167-195.

Crustacea Decapoda Dendrobranchiata

Crosnier, A., 1994. — Crustacea Decapoda : Penaeoidea rdcoltes lors de la campagne Karubar en Indondsie. In :

A. Crosnier (ed.), Rdsultats des Campagnes MUSORSTOM, Volume 12. Memoires du Museum national d'Histoire

naturelle, 161 : 351-365.

Crustacea Decapoda Caridea

Bruce, A. J., 1996. — Crustacea Decapoda : Palaemonoid shrimps from the Indo-West Pacific region mainly from New

Caledonia. In : A. Crosnier (ed.), Rdsultats des Campagnes Musorstom, Volume 15. Memoires du Museum national

d'Histoire naturelle, 168 : 197-267.

Chan, T.-Y., 1996. — Crustacea Decapoda Crangonidae : Revision of the three closely related crangonid genera, Aegeon

Agassiz, 1846, Pontocaris Bate, 1888, and Parapontocaris Alcock, 1901. In : A. Crosnier (ed.), Rdsultats des

Campagnes MUSORSTOM, Volume 15. Memoires du Museum national d'Histoire naturelle, 168 : 269-336.

Crustacea Decapoda Anomura

Forest, J., 1995. — Revision du genre Trizopagurus Forest, 1952 (Diogenidae), avec l’dtablissement de deux genres

nouveaux. In : A. Crosnier (ed.), Rdsultats des Campagnes Musorstom, Volume 13. Memoires du Museum national

d'Histoire naturelle, 163 : 9-149.

De Saint Laurent, M. & Poupin, J., 1996. — Crustacea Anomura : Les espdces indo-ouest pacifiques du genre Eumunida

Smith, 1880 (Chirostylidae). Description de six espdces nouvelles. In : A. Crosnier (ed.), Rdsultats des Campagnes

MUSORSTOM, Volume 15. Memoires du Museum national d'Histoire naturelle, 168 : 337-385.

Macpherson, E., 1993. — Crustacea Decapoda : Species of the genus Paramunida Baba, 1988 (Galatheidae) from the

Philippines, Indonesia and New Caledonia. In : A. Crosnier (ed.), Rdsultats des Campagnes Musorstom, Volume 10.

Memoires du Museum national d'Histoire naturelle, 156 : 443-473.

Baba, K. & de Saint Laurent, M., 1996. — Crustacea Decapoda : Revision of the genus Bathymunida Balss, 1914,

and description of six related genera (Galatheidae). In : A. Crosnier (ed.), Rdsultats des Campagnes Musorstom,

Volume 15. Memoires du Museum national d'Histoire naturelle, 168 : 433-502.

Crustacea Decapoda Brachyura

Guinot, D., 1993. — Donndes nouvelles sur les crabes primitifs (Crustacea Decapoda Brachyura Podotremata). Comptes

Rendus de I'Academie des Sciences, Sciences de la Vie, 316 : 1225-1232.

McLay, C. L., 1993. — Crustacea Decapoda : The sponge crabs (Dromiidae) of New Caledonia and the Philippines with a

review of the genera. In : A. Crosnier (ed.), Rdsultats des Campagnes Musorstom, Volume 10. Memoires du Museum

national d'Histoire naturelle, 156 : 111-251.

Source :

26

A. CROSNIER. B. RICHER DE FORGES & P. BOUCHET

Tavares, M., 1993. — Crustacea Decapoda : Les Cyclodorippidae et Cymonomidae de l'lndo-Ouest-Pacifique it

l'exclusion du genre Cymonomus. In : A. Crosnier (ed.), Resultats des Campagnes Musorstom, Volume 10,

Memoires du Museum nalional d'Histoire nalurelle , 156 : 253-313.

Guinot, D., 1995. — Crustacea Decapoda Brachyura : Revision de la famille des Homolodromiidae Alcock, 1899. In :

A. Crosnier (ed.), R6sultats des Campagnes Musorstom, Volume 13. Memoires du Museum nalional d'Histoire

nalurelle, 163 : 155-282.

Guinot, D. & Richer deForges, B., 1995. — Crustacea Decapoda Brachyura : Revision de la famille des Homolidae de

Haan, 1839. In : A. Crosnier (ed.), Resultats des Campagnes Musorstom, Volume 13. Memoires du Museum nalional

d'Histoire nalurelle. 163 : 283-517.

Manning, R. B., 1993. — A new deep-sea crab, genus Chaceon, from Indonesia (Crustacea: Decapoda: Geryonidae).

Raffles Bulletin of Zoology, 41 (2) : 169-172.

Moosa, M. K., 1996. — Crustacea Decapoda : Deep-water swimming crabs from the South-West Pacific, particularly New

Caledonia (Brachyura, Portunidae). In : A. Crosnier (ed.), Resultats des Campagnes Musorstom, Volume 15.

Memoires du Museum national d'Histoire nalurelle, 168 : 503-530.

Richer de Forges, B., 1995. — Nouvelles recoltes et nouvelles especes de Majidae de profondeur du genre Oxypleurodon

Miers, 1886. Crustaceana, 68 (1) : 43-60.

Richer de Forges, B., 1996. — The genus Platypilumnus Alcock and description of P. jamiesoni n. sp. from New

Caledonia (Crustacea, Decapoda. Brachyura). Records of the Australian Museum, 48 : 1-6.

Pycnogonides

Stock, J. H., 1994. — Indo-West Pacific Pycnogonida collected by some major oceanographic expeditions. Beaufortia,

44 (3) : 17-77.

Tuniciers

Monniot, C., 1993. — Tunicata : Sur trois especes d'ascidies bathyales recoltees au cours de la campagne franco-

indon£sienne Karubar. In : A. Crosnier (ed.), Resultats des Campagnes Musorstom, Volume 10. Memoires du

Museum national d'Histoire naturelle. 158 : 355-359.

Source : MNHN. Paris

JLTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS Dt

Cnidaria Anthozoa: Azooxanthellate Scleractinia

from the Philippine and Indonesian Regions

Stephen D. CAIRNS

National Museum of Natural History

NHB-163, W-329

Washington, D. C. 20560, U. S. A

&

Helmut ZIBROWIUS

Station Marine d'Endoume

Rue de la Batterie des Lions

13007 Marseille. France

ABSTRACT

A total ot 206 species of azooxanthellate Scleractinia are listed from the Philippine-Indonesian region, 176 of which

are reported as new records. The newly reported specimens originate primarily from the MUSORSTOM 1-3 and Karubar

expeditions, but also include specimens collected by the "Albatross", Danish Expedition to the Kei Islands, Snellius 2

Expedition, "Galathea", Mortensen's Java-South Africa Expedition, "Hakuho Maru", "Siboga", Corindon 2, and Estase 2

expeditions as well as some others. In all, approximately 15,600 specimens from some 640 stations are reported, the

new records including the description of 26 new species and 3 new genera. Also, 4 new combinations and 1 new name are

proposed ( Caryophyllia crosnieri for C. elongata Cairns, 1993, non Duncan, 1873).

The distribution and bathymetric ranges of the 206 species known from the combined Philippine-Indonesian region

are tabulated. 157 azooxanthellate species are now known from the Philippine Islands, with highest local diversities off

Lubang Island (69 species) and the Sulu Archipelago (66 species). 174 species are known from the Indonesian region, the

region with the highest diversity being the Banda Sea (138 species), specifically the Kai Islands (125 species). To a large

degree, these high levels of diversity reflect the intensity of sampling effort. 65 (or 31.5%) of the Philippine-Indonesian

species also occur in the Indian Ocean; 77 species (or 37%) off Japan; 67 species (or 32.5%) in the Australia and

New Zealand region; and 47 species (or 23%) among the other western Pacific Islands. Only 1 1 of these species (or 5.3%)

occur as tar east as continental eastern Pacific, and 1 1 also occur in the Atlantic Ocean. These corals occur from

0 to 2570 m, the deepest being Flabellum conuis. The highest diversity of species (123 species) has been found in the

200-400 m depth range.

Cairns, S. D. & Zibrowius, H., 1997. — Cnidaria Anthozoa: Azooxanthellate Scleractinia from the Philippine and

Indonesian Regions. In: A. Crosnier & P. Bouchet (cds), Resultats des Campagnes MUSORSTOM, Volume 16. Mem. Mus.

natn. Hist. not.. 172: 27-243. Paris ISBN 2-85653-506-2.

Source : MNHN, Paris

28

S. D. CAIRNS & H. ZIBROWIUS

The first example of sweeper tentacles in a deep-water coral is reported for Madrepora arbuscula. Examples of

commensal/symbiotic relationships are reported to occur with petrarcid ascothoracidan crustaceans (6 coral hosts),

acrothoracican cirripede crustaceans (4 coral hosts), eunicid polychaetes (4 coral hosts), and lumbrinerid polychaetes

(11 coral hosts). Several cases of epifauna living on live corals are the brachiopod Discradisca Stella and the gastropod

Malluvium sp. attached to Truncatoflabellum mortenseni, and a stalked suberitid sponge growing on Truncatoflabellum

paripavoninum.

RESUME

Cnidaria Anthozoa : Scleractiniaires sans zooxanthelles des Philippines et d'lndonesie.

206 espdces de scldractiniaires sans zooxanthelles sont recensdes dans la rdgion des Philippines et de I'lndondsie.

Pour 176 d'entre elles, des donndes nouvelles sont apportdes. Le matdriel etudie provient principalement des campagnes

Musorstom 1 d 3 (1976, 1980, 1985) aux Philippines et Karubar (1991) dans l'est de I'lndondsie. Diverses autres

collections sont dgalement prises en compte, dont notamment: "Challenger " (1874), "Siboga" (1899-1900),

"Albatross" (1908-1909), expedition danoise aux ties Kei (1922), expedition de Th. Mortensen it Java (1929),

"Galathea " (1951), "Hakuho-Maru " (1972, 1973, 1985), SNELLIUS 2 (1984), campagnes CORINDON 2 (1980) et Estase 2

(1984). Les dchantillons dtudies sont au nombre de 15.600 et proviennent d'environ 640 stations.

Trois genres nouveaux sont decrits {Confluphyllia, Ericiocyathus, Sympodangia) et 26 espdces nouvelles

( Balanophyllia crassiseptum, B. generatrix, B. serrata, Caryophyllia cornulum, C. karubartca, C. octonaria, C. secta,

C. unicristata, Confluphyllia juncta, Deltocyathus philippinensis, D. Stella, Endopachys bulbosa, Ericiocyathus

echinatus, Fungiacyathus fissidiscus, Madrepora minutiseptum, Rhizosmilia elata, Stephanocyathus regius,

Sympodangia albatrossi, Trochocyathus apertus, T. brevispina, T. discus, T. longispina, T. semperi, "Tropidocyathus"

labidus, Truncatoflabellum angustum, T. mortenseni). Un nom nouveau est introduit ( Caryophyllia crosnieri, nomen

novum) et 4 combinaisons nouvelles rdsultent de transferts dans un autre genre ( Colangia moseleyi, Premocyathus

dentiformis, Trochocyathus burchae, Deltocyathoides orientalis). Une espdce etrange, aux affinites incertaines ( incertae

sedis), est presentde, mais non decrite formellement.

Un tableau resume la repartition geographique et bathymetrique des 206 espdces recensdes dans la rdgion des

Philippines et de I'lndondsie. Aux Philippines (157 espdces), la diversitd est la plus dldvde autour de 1'ile Lubang

(69 espdces) et dans l'archipel de Sulu (66 espdces). En Indondsie (174 espdces), elle est la plus dldvde dans la mer de

Banda (138 espdces) et plus spdcialement aux ties Kai (125 espdces). Ces remarquables richesses en espdces refldtent aussi

une prospection particulidrement intense dans les secteurs en question. 65 des 206 espdces inventoriees dans la rdgion des

Philippines et de I'lndondsie vivent aussi dans l'ocdan Indien tropical, 77 au Japon, 67 autour de l'Australie ou au nord de

la Nouvelle-Zelande, et 47 autour d'autres iles du Pacifique occidental. Mais seulement 1 1 de ces espdces sont connues dans

le Pacifique oriental et 1 1 espdces dgalement dans l'ocdan Atlantique. La repartition en profondeur des espdces etudides

s'dtale entre 0 et 2700 m, Flabellum conuis etant l'espdce qui atteint la plus grande profondeur. L'intervalle de 200-400 m

est habitd par environ les 2/3 des espdces. Le genre Leptopenus, au squelette extremement fragile, a did reconnu pour la

premidre fois dans des fonds de 300 m seulement,

De nombreuses espdces prdsentent un mode de multiplication asexude: division transversale, fractionnement suivi de

rdgdndration, ddtachement de bourgeons. Pour la premidre fois des tentacules spdciaux, particulidrement allongds

("sweeper tentacles"), ont ete reconnus chez un scleractiniaire bathyal ( Madrepora arbuscula)', auparavant ce type de

tentacule dtait connu seulement chez divers scldractiniaires rdcifaux. Divers types dissociations symbiotiques plus ou

moins specialisdes ont ete reconnus. Les partenaires sont les suivants: crabes Cryptochiridae (avec Phyllangia

papuensis), crustacds ascothoracides causant des galles (dans Madrepora oculata, Balanophyllia carinata, Balanophyllia

sp., Deltocyathoides orientalis, Dendrophyllia sp. cf. D. ijimai, Flabellum lamellulosum)', cirripddes acrothoraciques

perforants (dans Balanophyllia crassiseptum, Balanophyllia sp., Javania lamprotichum, Tetliocyathus virgatus)',

polychdtes Eunicidae lids aux ddformations de colonies de Madrepora et de Neohelia, leurs tubes y dtant incorpores;

polychdte Lumbrineris flabellicola corrodant la surface du squelette (de Balanophyllia sp., Caryophyllia grayi,

C. spinicarens, C. spinigera, C. transversalis, Conotrochus brunneus, Flabellum lamellulosum, F. patens, Flabellum

sp., Rhizotrochus typus, Dendrophylliidae).

Des cas plus remarquables d'epifaune installee sur des coraux vivants, mais n’en ddpendant probablement pas, sont le

brachiopode Discradisca Stella et le gastropode Malluvium sp. sur Truncatoflabellum mortenseni, et un spongiaire

Suberitidae pddonculd sur Truncatoflabellum paripavoninum.

CONTENTS

Introduction . 29

List of Abbreviations . 29

Combined List of Stations and of Species obtained per Station . 30

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

29

Historical Review (Indonesian Region) . 55

Material . 57

Methods . 57

Commensal Relationships . 58

Distribution . 59

Regional Diversity . 59

Regional Affinities . 65

Depth Distribution . 65

Systematic Account . 66

Family Pocilloporidae . 66

Family Fungiacyathidae . 68

Family Micrabaciidae . 73

Family Rhizangiidae . 78

Family Oculinidae . 79

Family Anthemiphylliidae . 86

Family Caryophylliidae . 87

Family Turbinoliidae . 140

Family Guyniidae . 150

Family Flabellidae . 150

Family Dendrophylliidae . 175

INCERTAE SEDIS . 198

Acknowledgements . 198

References . 198

INTRODUCTION

This work may be considered as the concluding part 2 of the partial revision by Cairns (1989a) of the

azooxanthellate Scleractinia of the Philippines and adjacent waters. Whereas Cairns included only 58 Philippine

species from 5 families, this work lists 206 species in 1 1 families from the Philippine-Indonesian region,

including additional records of most species reported in Part 1. This work differs conceptually from Part 1

(CAIRNS, 1989a) in that it covers a larger geographic region, including most of Indonesia, and was based on a

much larger and varied collection.

LIST OF ABBREVIATIONS

Museums:

AMS Australian Museum, Sydney.

MNHN Museum national d'Histoire naturelle, Paris.

MoNZ Museum of New Zealand Te Papa Tonga-rewa, Wellington (formerly the National Museum of

New Zealand: NMNZ).

NHM The Natural History Museum, London [formerly the British Museum (Natural History): BMNH],

NMW Naturhistorisches Museum, Wien.

NNM Nationaal Natuurhistorisch Museum, Leiden (formerly the Rijksmuseum van Natuurlijke

Historic: RMNH).

NMNH National Museum of Natural History, Smithsonian, Washington. D. C. (formerly the United

States National Museum: USNM).

NZOI New Zealand Oceanographic Institute, Wellington.

Source :

30

S. D. CAIRNS & H. ZIBROWIUS

ORI Ocean Research Institute, Tokyo.

POLIPI Puslitbang Oseanologi (National Institute of Oceanology), Jakarta.

ZMA Zoologisch Museum, Amsterdam.

ZMB Zoologisches Museum, Berlin.

ZMUC Zoologisk Museum, Kdbenhavn.

Expeditions and Vessels:

CORINDON 2 French-Indonesian expedition (1980) that collected mainly in the Makassar Strait.

DEKI Danish Expedition to the Kei Islands (1922).

ESTASE 2 French expedition (1984) that collected in the Philippines.

KARUBAR French-Indonesian expedition (1991) that collected in the southeastern Banda Sea.

Named for the Kai, Aru, and Tanimbar Islands.

MUSORSTOM Cruises organized jointly by the Museum National d'Histoire Naturelle and Institut

Fran?ais de Recherche Scientifique pour le Developpement en Cooperation

(formerly: Office de la Recherche Scientifique et Technique d'Outre-Mer,

= ORSTOM).

SIPHILEXP Smithsonian Institution Philippine Expedition (1978).

TM R/V "Tansei Maru."

Morphological Terms:

D:H

GCD

GCD:H

GCD:LCD

H:D

LCD

LEL:H

PD:GCD

SCI

SEM

SSI

Sx, Cx, Px

Sx>Sy

Ratio of diameter to height of a solitary corallum.

Greater calicular diameter.

Ratio of greater calicular diameter to height of a solitary corallum.

Ratio of greater calicular diameter to lesser calicular diameter.

Ratio of height to diameter of a solitary corallum.

Lesser calicular diameter.

Ratio of lateral edge length to height of a solitary corallum, i.e., of a Flabellum.

Ratio of pedicel diameter to greater calicular diameter of a solitary corallum.

Septal concavity index: ratio of distance from thecal edge to point of greatest septal inflection

to length of thecal face along that septum.

Scanning Electron Microscopy.

Septal sinuosity index: ratio of amplitude of sinuosity of lower inner edge of a major septum

to the thickness of same septum (see Cairns, 1989b).

Septa, costae, or pali (respectively) of cycle designated by numerical subscript.

In the context of a septal formula, septa of cycle x wider than septa of cycle y.

COMBINED LIST OF STATIONS AND OF SPECIES OBTAINED PER STATION

This list provides the data of all stations mentioned in this report (ships, respectively cruises, in alphabetical

order) and of those stations of the "ALBATROSS" Philippines Expedition and of cruises MUSORSTOM 1, 2, 3

(Philippines) and CORINDON 2 (Indonesia) that were previously mentioned by CAIRNS (1989a). For some of the

stations previously mentioned by Cairns (1989a) the data are herein corrected.

The following abbreviations are used and placed after the species names:

CSD: (= confused station data) species found labeled as from this station, but certainly from another station

because of the unlikely depth;

C89: already reported by Cairns (1989a);

C89*: reported by Cairns (1989a), but under another name (see synonymy, remarks).

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

31

"Acheron" New Zealand National Museum (= NZNM),

Stn BS441. — 28.10.1975, Kermadec Isl., Raoul Isl., 3.7 km off Nugent Isl., 366-402 m: Truncatoflabellum

angustum.

"Akademik Oparin"

Stn 18. — 3.12.1990, 1 3°56. 1 'S, 140°57.4'E, 36 m, Gulf of Carpentaria: Truncatoflabellum spheniscus.

"Albatross" Philippines Expedition

Sin 5106. — 9.1.1908, 14°23'55"N, 120°32'33"E, 68 m, SW Luzon: Placotrochus laevis C89.

Stn 5107. — 9.1.1908, 14°24'30"N, 120°33'40"E, 51 m, SW Luzon: Placotrochus laevis C89.

Stn 5110. — 15.1.1908, 13°59'20"N, 120°15'45"E, 247 m, 15.0°C, SW Luzon: Balanophyllia cornu,

Deltocyathus vaughani, Flabellum (U.) deludens C89, Fungiacyathus (F.) stephanus C89.

Stn 5113. — 17.1.1908, 13°5r30"N, 120°50'30"E, 291 m, SW Luzon: Fungiacyathus (B.) variegatus C89.

Stn 5116. — 20.1.1908, 13°4r00"N, 120°47'05"E, 366 m, 10.1°C, SW Luzon: Caryophyllia (C.) secta,

Flabellum (F.) lamellulosum C89, Flabellum (F.) magnificum C89.

Stn 5117. — 21.1.1908, 13°52'22"N, 120°46'22"E, 216 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum C89, Flabellum (U.) deludens C89.

Stn 51 18. — 21.1.1908, 13°48'45"N, 120°41'51"E, 291 m, SW Luzon: Caryophyllia (A.) spinicarens, Flabellum

(F.) magnificum C89, Flabellum (U.) deludens C89.

Stn 5123. — 2.2.1908, 13°12'45"N, 121°38'45"E, 518 m, E Mindoro: Madrepora oculata.

Stn 5124. — 2.2.1908, 12°52'00"N, 121°48'30"E, 514 m, E Mindoro: Flabellum (U.) japonicum C89, Madrepora

oculata.

Stn 5130. — 5.2.1908, 7°35'00,,N, 122°04’45"E, 187 m, 15.1°C, W Mindanao: Leptopsammia crassa.

Stn 5133. — 6.2.1908, 7°41'00"N, 122°01'00"E, 70 m, W Mindanao: Asterosmilia marchadi, Balanophyllia

imperialis, Balanophyllia stimpsonii, Deltocyathoides orientalis C89* (ex P. australiensis) CSD?,

Endocyathopora laticostata C89, Endopachys grayi, Flabellum (F.) politum C89 CSD?, Notocyathus venustus

C89 CSD?, Placotrochus laevis C89, Trochocyathus (T.) burchae, Trochocyathus (T.) cooperi, Trochocyathus

(T.) semperi, Tropidocyathus lessonii C89.

Stn 5134. — 7.2.1908, 6°44'45"N, 121°48'00"E, 46 m, Jolo: Balanophyllia carinata, Cyathelia axillaris,

Endocyathopora laticostata C89, Placotrochus laevis C89.

Stn 5135. — 7.2.1908, 6°1 1'50"N, 121o08'20"E, 294 m, Jolo: Balanophyllia gemma, Deltocyathus magnificus.

Stn 5136. — 14.2.1908, 6°04'20"N, 120°59'20"E, 40 m, Jolo: Flabellum (F.) politum C89 CSD, Flabellum (U.)

deludens C89 CSD, Trochocyathus (T.) cooperi.

Stn 5137. — 14.2.1908, 6°04'25"N. 120°58'30"E, 37 m, Jolo: Balanophyllia stimpsonii, Deltocyathoides

orientalis CSD, Trochocyathus (T.) cooperi, Truncatoflabellum incrustation C89* (ex T. formosum).

Stn 5139. — 14.2.1908, 6°06'00"N, 121°02'30"E, 37 m, Jolo: Balanophyllia carinata.

Stn 5141. — 15.2.1908, 6°09'00"N, 120°58'00"E, 53 m, Jolo: Truncatoflabellum aculeatum.

Stn 5142. — 15.2.1908, 6°06'10"N, 121°02'40"E, 38 m, Jolo: Trochocyathus (T.) cooperi, Trochocyathus (T.)

semperi.

Stn 5143. — 15.2.1908, 6°05'50"N, 121°02'15"E, 35 m, Jolo: Balanophyllia stimpsonii, Trochocyathus (T.)

cooperi.

Stn 5144. — 15.2.1908, 6°05’50"N, 121°02'15"E, 35 m, Jolo: Placotrochus laevis C89. Trochocyathus (T.)

cooperi.

Stn 5145. — 15.2.1908, 6°04’30"N, 120°59'30"E, 42 m, Jolo: Sphenotrochus (S.) hancocki C89,

Truncatoflabellum irregulare C89.

Stn 5146. — 16.2.1908, 5°46'40"N, 120°48'50"E, 44 m, Siasi: Balanophyllia imperialis, Deltocyathoides

orientalis C89* (ex P. australiensis ) CSD, Truncatoflabellum phoenix, Trochocyathus (T.) cooperi.

Stn 5147, — 16.2.1908, 5°41'40"N, 120°47'10"E, 38 m, Siasi: Trochocyathus (T.) cooperi, Truncatoflabellum

phoenix.

Stn 5151. 18.2.1908, 5°24'40"N, 120°27'15"E, 44 m, Tawitawi group: Balanophyllia carinata, Premocyathus

dentiformis, Trochocyathus (T. ) cooperi.

Source :

32

S. D. CAIRNS & H. ZIBROWIUS

Stn 5152. — 18.2.1908, 5°22'55"N, 120°15'45"E, 62 m, Tawitawi group: Asterosmilia marchadi, Balanophyllia

carinata, Deltocyathoides orientalis C89* (ex P. australiensis), Notocyathus conicus C89.

Stn 5153. — 19.2.1908, 5°18'10MN, 120°02'55"E, 90 m, Tawitawi group: Fungiacyathus (B.) paliferus C89.

Stn 5155. — 19.2.1908, 5°13'40"N, 119°57'20"E, 22 m, Tawitawi group: Premocyathus dentiformis.

Stn 5156. — 21.2.1908, 5°12’50"N, 1 19°55’55"E, 33 m, Tawitawi group: Asterosmilia marchadi , Balanophyllia

carinata, Balanophyllia stimpsonii, Trochocyathus (T.) apertus, Truncatoflabellum aculeatum C89.

Stn 5159. — 21.2.1908, 5°1 1'50"N, 1 19°54'00"E, 18 m, Tawitawi group: Truncatoflabellum phoenix.

Stn 5161. — 22.2.1908, 5°10'15"N, 1 19°53’00"E, 29 m, Tawitawi group: Truncatoflabellum aculeatum C89.

Stn 5162. — 22.2.1908, 5°10'00"N, 119°47’30"E, 421 m, 11.6°C, Tawitawi group: Anthemiphyllia dentata,

Deltocyathoides orientalis C89* (ex P. australiensis), Fungiacyathus (B.) paliferus C89, Idiotrochus kikutu

C89, Madracis cf pharensis, Notocyathus conicus C89, Tropidocyathus lessonii C89, Truncatoflabellum

formosum C89, Truncatoflabellum' phoenix, Truncatoflabellum pusillum, Truncatoflabellum sp. C89*

(ex T. formosum).

Stn 5164. — 24.2.1908, 5°01’40"N, 1 19°52'20"E, 33 m, Tawitawi group: Asterosmilia marchadi, Balanophyllia

carinata, Balanophyllia stimpsonii, Paracyathus rotundatus, Trochocyathus (T.) apertus, Truncatoflabellum

aculeatum C89.

Stn 5171. — 28.2.1908, 5°05'00"N, 1 19°28'00"E, 458 m, 1 1.9°C, Sulu Sea: Conotrochus brunneus.

Stn 5172. — 5.3.1908, 6°03T5"N, 120°35'30"E, 582 m, Jolo: Caryophyllia (C.) rugosa, Deltocyathoides

orientalis C89* (ex P. australiensis), Idiotrochus kikutii C89, Peponocyathus minimus C89*

(ex P. folliculus).

Stn 5173. — 5.3.1908, 6°02’55"N, 120°53’00"E, 340 m, Jolo: Flabellum (F.) patens C89.

Stn 5174. — 5.3.1908, 6°03'45"N, 120°57'00"E, 37 m, Jolo: Balanophyllia imperials.

Stn 5178. — 25.3.1908, 12°43’00"N, 122°06'15"E, 143 m, Romblon: Anthemiphyllia dentata, Asterosmilia

marchadi, Deltocyathoides orientalis C89* (ex P. australiensis), Flabellum (F.) politum C89, Fungiacyathus

(B.) paliferus C89, Idiotrochus kikutii C89, Letepsammia formosissima C89, Notocyathus conicus C89,

Stephanophyllia neglecta C89, Tropidocyathus lessonii C89, "Tropidocyathus" pileus C89, Trochocyathus (T.)

philippinensis, Truncatoflabellum pusillum C89.

Stn 5179. — 25.3.1908, 12°38'15"N, 122°12'30"E, 68 m, 24.3°C, Romblon: Incertae sedis, Tropidocyathus

lessonii C89, Truncatoflabellum phoenix.

Stn 5197. — 9.4.1908, 9°52'30"N, 123°40'45"E, 318 m, 12.4°C, W Bohol: Flabellum (U.) deludens C89.

Stn 5198. — 9.4.1908, 9°40'50"N, 123°39'45"E, 403 m, 12.2°C, W Bohol: Caryophyllia (A.) spinicarens,

Deltocyathus magnificus, Deltocyathus vaughani, Flabellum (U.) japonicum C89, Fungiacyathus (F.)

stephanus C89.

Stn 5201. — 10.4.1908, 10°10’00"N, 125°04T5"E, 1014 m, 11.6°C, S Leyte: Madrepora oculata.

Stn 5202. — 10.4.1908, 10°12’00"N, 125°04T0"E, 919 m, S Leyte: Flabellum (U.) deludens, Madrepora oculata,

Trochocyathus (T.) cooperi CSD.

Stn 5212. — 24.4.1908, 12°04T5"N, 124°04'36"E, 198 m, 15.5°C, E Masbate: Flabellum (F.) lamellulosum

C89, Flabellum (F.) politum C89.

Stn 5213. — 20.4.1908, 12°15'00"N, 123°57'30"E, 146 m, E Masbate: Deltocyathoides orientalis C89*

(ex P. australiensis), Stephanophyllia neglecta C89, Trochocyathus (T.) philippinensis, Truncatoflabellum

candeanum C89.

Stn 5217. — 22.4.1908, 13°20'00"N, 123°14T5"E, 192 m, 17.3°C, between Luzon and Burias: Caryophyllia (C.)

rugosa, Conotrochus brunneus, Deltocyathoides orientalis C89* (ex P. australiensis), Flabellum (F.) politum

C89, Gardineria philippinensis C89, Letepsammia formosissima C89, Madracis sp. A, Peponocyathus

minimus C89* (ex P. folliculus), Premocyathus dentiformis, Thrypticotrochus multilobatus C89,

Trochocyathus (T.) philippinensis.

Stn 5221. — 24.4.1908, 13°38T5"N, 12r48'15"E, 353 m, 16.9°C, between Luzon and Marinduque: Flabellum

(U.) japonicum C89.

Stn 5222. — 24.4.1908, 13°38’30"N, 12r42'45"E, 357 m, 17.1°C, between Luzon and Marinduque: Flabellum

(U.) japonicum C89.

Stn 5236. — 11.5.1908, 8°50'45"N, 126°26'52"E, 903 m, 5.1°C, NE Mindanao: Peponocyathus minimus.

Stn 5244. — 14.5.1908, 6°52'05"N, 126°14I15,,E, 313 m, SE Mindanao: Rhizosmilia elata.

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTIN1A

33

Stn 5248. — 18.5.1908, 7°07'05"N, 125°40'24"E, 33 m, SE Mindanao, Gulf of Davao: Eguchipsammia wellsi.

Stn 5249. — 18.5.1908, 7°06'06"N, 125°40'08"E, 42 m, SE Minanao, Gulf of Davao: Eguchipsammia gaditana,

Eguchipsammia wellsi, Truncatoflabellum formosum C89, Truncatoflabellum incrustatum C89.

Stn 5250. — 18.5.1908, 7°05'07"N, 125°39'45"E, 42 m, SE Mindanao, Gulf of Davao: Truncatoflabellum

incrustatum C89.

Stn 5251. — 18.5.1908, 7°05'12"N, 125°39'35"E, 37 m, SE Mindanao, Gulf of Davao: Truncatoflabellum

incrustatum C89.

Stn 5253. — 18.5.1908, 7°04'48"N, 125°39'38"E, 51 m, SE Mindanao, Gulf of Davao: Truncatoflabellum

aculeatum, Truncatoflabellum incrustatum C89.

Stn 5255. — 18.5.1908, 7°03,00"N, 125°39'00"E, 183 m, SE Mindanao, Gulf of Davao: Cyathelia axilllaris,

Javania insignis C89, Rhizosmilia sagamiensis, Thalamophyllia tenuescens.

Stn 5256. —22.5.1908, 7°21'45"N, 124°07'15"E, 289 m, S Mindanao, Illana Bay: Caryophyllia (A.) spinicarens,

Flabellum (U.) deludens C89.

Stn 5260. — 3.6.1908, 12025'35"N, 121°31'35"E, 428 m, 10.8°C, SE Mindoro: Caryophyllia (A.) spinicarens,

Fungiacyathus (B.) granulosus C89.

Stn 5265. — 6.6.1908, 13°41’15"N, 120°00'50"E, 247 m, SW Luzon: Caryophyllia (C.) secta, Flabellum (U.)

marenzelleri C89, Truncatoflabellum formosum C89, Truncatoflabellum incrustatum C89.

Stn 5268. — 8.6.1908, 13°42'00''N, 120°57'15"E, 311 m, SW Luzon: Balanophyllia cornu, Cyathelia axillaris,

Deltocyathoides orientalis C89* (ex P. australiensis), Endopachys grayi, Flabellum (U.) marenzelleri C89,

Madracis cf pharensis.

Stn 5272. — 14.7.1908, 14°00'00"N, 120°22'30"E, 216 m, 14.1°C, SW Luzon: Notocyathus conicus C89.

Stn 5273. — 14.7.1908, 13°58'45"N, 120°21'35"E, 209 m, SW Luzon: Caryophyllia (A.) spinigera, Deltocyathus

andamanicus, Flabellum (F.) lamellulosum C89, Flabellum (U.) deludens C89, Stephanocyathus (A.) spiniger.

Stn 5277. — 17.7.1908, 13°56’55"N, 120°13'45"E, 146 m, 14.8°C, SW Luzon: Asterosmilia marchadi,

Conotrochus brunneus, Deltocyathoides orientalis C89* (ex P. australiensis), Madracis asanoi, Notocyathus

conicus C89, Peponocyathus folliculus C89, "Tropidocyathus" pileus C89, Truncatoflabellum pusillum C89.

Stn 5278. — 17.7.1908, 14°00'10"N, 120°17'15"E, 187 m, 15.3°C, SW Luzon: Caryophyllia (A.) spinigera,

Fungiacyathus (B.) variegatus C89, Flabellum (U.) deludens C89, Letepsarnmia formosissima C89.

Stn 5279. — 17.7.1908, 13°57,30"N, 120o22'15"E, 214 m, SW Luzon: Dendrophyllia arbuscula.

Stn 5280. — 17.7.1908, 13°55'20"N, 120°25’55"E, 353 m, 9.8°C, SW Luzon: Balanophyllia cornu,

Dendrophyllia arbuscula, Flabellum (F.) magnificum C89.

Stn 5281. — 18.7.1908, 13°52'45"N, 120°25'00"E, 368 m, 10.2°C, SW Luzon: Balanophyllia cornu, Flabellum

(F.) magnificum C89, Flabellum (U.) messum C89.

Stn 5282. — 18.7.1908, 13°53'00"N, 120°26'45"E, 454 m, 8.6°C, SW Luzon: Deltocyathoides orientalis C89*

(ex P. australiensis), Fungiacyathus (B.) granulosus C89.

Stn 5283. — 18.7.1908, 13°48’30"N, 120°28'40"E, 512 m, 8.2°C, SW Luzon: Fungiacyathus (B.) granulosus

C89, Rhombopsammia niphada C89, Truncatoflabellum paripavoninum C89.

Stn 5284. — 18.7.1908, 13°42'05"N, 120°30’45"E, 772 m, 5.7°C, SW Luzon: Flabellum (U.) sexcostatum C89,

Truncatoflabellum paripavoninum C89.

Stn 5289. — 22.7.1908, 13°41'50"N, 120°58'30"E, 315 m, SW Luzon: Flabellum (F.) patens C89, Flabellum

(U.) marenzelleri C89, Truncatoflabellum formosum C89, Truncatoflabellum incrustatum C89.

Stn 5297. — 24.7.1908, 13°41’20"N, 120°58'00"E, 362 m, SW Luzon: Flabellum (U.) deludens C89.

Stn 5298. — 24.7.1908, 13°43'25"N, 120°57'40"E, 256 m, SW Luzon: Flabellum (U.) deludens C89, Flabellum

(U.) marenzelleri C89.

Stn 5301. — 8.8.1908, 20°37'00"N, 1 15o43'00"E, 381 m, 10.3°C, South China Sea, SE Hong Kong:

Caryophyllia (A.) spinicarens, Conotrochus brunneus, Flabellum (F.) patens C89, Thrypticotrochus

multilobatus C89.

Stn 5310. — 4.11.1908, 21°33'00"N, 116°13’00"E, 183 m, 18.6°C, South China Sea, SE Hong Kong:

Balanophyllia parvula, Caryophyllia (C.) hawaiiensis.

Stn 5311. — 4.11.1908, 21°33'00"N, 116°15'00"E, 161 m, South China Sea, SE Hong Kong: Asterosmilia

marchadi, Deltocyathoides orientalis C89* (ex P. australiensis). Dendrophyllia alcocki. Flabellum (F.) politum

C89, Idiotrochus kikutii C89, Madracis sp. A, Notocyathus venustus C89. Peponocyathus minimus C89*

Source :

34

S. D. CAIRNS & H. ZIBROWIUS

(ex P folliculus ), Sphenotrochus (S.) hancocki C89, Stephanophyllia fungulus C89, Trochocyathus (T.)

philippinensis, Tropidocyathus lessonii C89, Truncatoflabellum candeanum C89, Truncatoguyma irregularis

C89.

Stn 5312 — 4.11.1908, 21°30'00"N, 116°32'00"E, 256 m. 14.2°C, South China Sea, SE Hong Kong:

Conotrochus funicolumna, Deltocyathoides orientalis C89, Flabellum (F.) pavoninum C89, Flabellum (F.)

politum C89 (not 5212), Notocyathus venustus C89, Peponocyathus minimus , Sphenotrochus (S.) hancocki

C89, Stephanophyllia fungulus C89, "Tropidocyathus" pileus C89, Truncatoflabellum carinatum C89.

Stn 5313. — 4.11.1908, 21°30'00"N, 116°43'00"E, 274 m, 12.0°C, South China Sea, SE Hong Kong:

Anthemiphyllia frustum, Caryophyllia (C.) quadragenaria, Deltocyathoides orientalis C89* (ex

P. australiensis), Flabellum (F.) politum, Peponocyathus minimus, Sphenotrochus (S.) hancocki C89,

Truncatoflabellum carinatum C89.

Stn 5314. — 5.11.1908, 2r4r00"N, 116°46'00"E, 223 m, 15.3°C, South China Sea, SE Hong Kong:

Caryophyllia (A.) spinicarens, Deltocyathoides orientalis C89, Deltocyathus magnificus, Fungiacyathus (R.)

variegatus C89, Madrepora oculata, Trochocyathus (T.) philippinensis, Truncatoflabellum candeanum C89,

Truncatoflabellum carinatum C89.

Stn 5315. — 5.11.1908, 21°40’00"N, 116o58’00"E, 271 m, 12.4°C, South China Sea, SE Hong Kong:

Deltocyathoides orientalis C89* (ex P. australiensis ), Peponocyathus minimus, Sphenotrochus (S.) hancocki

C89

Stn 5317. — 5.11.1908, 21°36'00"N, 117°27'00"E, 421 m, 10.3°C, South China Sea, SE Hong Kong:

Caryophyllia (C.) diomedeae, Caryophyllia (C.) quadragenaria, Deltocyathoides orientalis C89*

(ex P. australiensis), Enallopsammia marenzelleri, Madrepora oculata, "Tropidocyathus" pileus C89.

Stn 5318. — 5.11.1908, 21°32'00"N, 117o46'00"E, 622 m. South China Sea, SE Hong Kong: Conotrochus

funicolumna, Deltocyathoides orientalis C89* (ex P. australiensis ), Fungiacyathus (B.) turbinolioides C89.

Stn 5327. — 12.11.1908, 18°3r30"N, 122°03'00"E, 362 m, N Luzon: Madrepora oculata.

Stn 5331. — 22.1 1.1908, 15°36'45"N, 1 19°47'45"E, 326 m, 12.6°C, W Luzon: Trochocyathus (A.) longispina.

Stn 5336. — 18.12.1908, 1 1°37'45"N, 1 19°46'E, 84 m, Linapacan Strait: Dactylotrochus cervicornis (fide Wells,

1954).

Stn 5348. — 27.12.1908, 10°57'45"N, U8°38’15"E, 686 m, 13.6°C, NW Palawan: Caryophyllia (C.) scobinosa,

Fungiacyathus (F.) stephanus C89, Madrepora oculata, Rhombopsammia niphada C89, Stephanocyathus (O.)

weberianus.

Stn 5349. — 27.12.1908, I0°54'00"N, 1 18°26'20"E, 1336 m, 4.8°C, NW Palawan: Fungiacyathus (F.) stephanus

C89, Madrepora oculata, Stephanocyathus (O.) weberianus.

Stn 5353. — 1.1.1909, 7°50'45"N, 116°43’15"E, 271 m, S Balabac: Caryophyllia (A.) spinigera.

Stn 5355. — 5.1.1909, 8°08’10"N, 1 17°19'15"E, 81 m, N Balabac: Trochocyathus (T.) cooperi.

Stn 5357. — 5.1.1909, 8°06'00"N, 117°17'10"E, 124 m, N Balabac: Asterosmilia marchadi, Eguchipsammia

wellsi, Javania insignis C89, Endopachys grayi, Rhizotrochus typus C89.

Stn 5367. — 22.2.1909, 13°34'37"N, 121°07’30"E, 329 m, SW Luzon: Balanophyllia cornu, Cyathelia axillaris.

Stn 5369. — 24.2.1909, 13°48'00"N, 121°43'00"E, 194 m, Marinduque: Caryophyllia (A.) spinigera, Flabellum

(U.) deludens C89, Letepsammia formosissima C89, Stephanocyathus (A.) spiniger, Truncatoflabellum

candeanum C89.

Stn 5371. — 24.2.1909, 13°49'40"N, 12r40'15"E, 152 m, Marinduque: Caryophyllia (A.) spinigera,

Letepsammia formosissima C89, Stephanocyathus (A.) spiniger, Truncatoflabellum candeanum C89.

Stn 5372. — 24.2.1909, 13°49'12"N, 121°36'09"E, 274 m, Marinduque: Caryophyllia (A.) spinigera, Flabellum

(U.) deludens C89 (not 5392), Stephanocyathus (A.) spiniger.

Stn 5373. — 2.3.1909, 13°40'00"N, 121o31'10"E, 619 m, 1 1.0°C, Marinduque: Flabellum (U.) japonicum C89,

Madrepora oculata.

Stn 5374. — 2.3.1909, 13°46'45"N, 121°35'08"E, 348 m, Marinduque: Caryophyllia (A.) spinicarens,

Caryophyllia (A.) spinigera, Deltocyathus magnificus, Flabellum (U.) deludens C89.

Stn 5376. — 2.3.1909, 13°42'50"N, 121°5r30''E, 165 m, Marinduque: Caryophyllia (A.) spinigera, Letepsammia

formosissima C89, Truncatoflabellum candeanum C89.

Stn 5378. — 4.3.1909, 13°17’45"N, 122°22'00"E, 723 m, 10.2°C, Marinduque: Madrepora oculata.

Stn 5380. — 4.3.1909, 13°02'45"N, 122°29'00"E, depth?, Marinduque: Notocyathus venustus C89, CSD

(net lost).

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

35

Stn 5381. — 6.3.1909, 13°14'15"N, 122°44'45"E, 161 m, S Luzon, Gulf of Ragai: Cary’ophyllia (A.) grayi,

Deltocyathoides orientalis C89* (ex P. australiensis), Madracis asanoi, Madrepora oculata, Trochocyathus (T. )

philippinensis.

Stn 5387. — 1 1.3.1909, 12°54’40"N, 123°20'30"E, 382 m, 11.3°C, between Luzon and Burias: Flabellum (U.)

deludens C89.

Stn 5391. — 13.3.1909, 12°13'15"N, 124°05'03"E, 216 m, between Masbate and Samar: Batanophyllia cornu,

Flabellum (F.) lamellulosum C89, Flabellum (F.) politum C89, Truncatoflabellum candeanum C89.

Stn 5392. — 13.3.1909, 12°13'35"N, 124°02'48"E, 247 m, between Masbate and Samar: Balanophyllia cornu,

Deltocyathoides orientalis C89* (ex P. australiensis), Flabellum (F.) lamellulosum C89, Flabellum (F.)

politum C89.

Stn 5393. — 13.3.1909, 12°03'30"N, 124°03'36"E, 249 m, between Masbate and Samar: Balanophyllia cornu,

Deltocyathoides orientalis C89* (ex P. australiensis), Flabellum (F.) lamellulosum C89, Flabellum (F)

politum C89, Truncatoflabellum candeanum C89.

Stn 5398. — 15.3.1909, 1 1°35'12''N, 124°13'48"E, 209 m, between Masbate and Leyte: Deltocyathoides orientalis

C89* (ex P. australiensis), Letepsammia formosissima C89, Madracis sp. A, Sympodangia albatrossi.

Stn 5403. — 16.3.1909, 11°10'00"N, 124°17’15"E, 333 m, 13.2°C, between Cebu and Leyte: Caryophyllia (A.)

spinicarens, Deltocyathoides orientalis C89* (ex P. australiensis), Deltocyathus andamanicus, Flabellum (U.)

deludens C89, Fungiacyathus (B.) variegatus C89, Madrepora oculata, Notocyathus conicus C89.

Stn 5405. — 17.3.1909, 10°49’20"N, 124°24'23"E, 479 m, W Leyte: Madrepora oculata.

Stn 5406. — 17.3.1909, 10°49'03"N, 124°22'30"E, 545 m, W Leyte: Madrepora oculata.

Stn 5407. — 17.3.1909, 10°51'38"N, 124°20’54"E, 640 m, W Leyte: Madrepora oculata.

Stn 5408. — 18.3.1909, 10°40,15',N, 124°15'00"E, 291 m, 13.0°C, between Cebu and Leyte: Cary’ophyllia (A.)

spinicarens, Flabellum (U.) deludens C89, Madrepora oculata.

Stn 5411. — 23.3.1909, 10°10'30"N, 123°51'15"E, 265 m, 12.9°C, between Cebu and Bohol: Caryophyllia ( A .)

spinicarens, Flabellum (U.) deludens C89, Madrepora oculata.

Stn 5412. — 23.3.1909, 10o09'15''N, 123°52’00"E, 296 m, 12.7°C, between Cebu and Bohol: Cary’ophyllia (A.)

spinicarens, Deltocyathus andamanicus, Flabellum (F.) magnificum C89, Flabellum (U.) deludens C89.

Stn 5417. — 25.3.1909, 10°10'00"N, 123°53'15"E, 302 m, 12.4°C, between Cebu and Bohol: Caryophyllia (A.)

spinicarens, Deltocyathus andamanicus, Flabellum (U.) deludens C89, Madrepora oculata.

Stn 5418. — 25.3.1909, 10°08'50"N, 123°52'30"E, 291 m, 12.4°C, between Cebu and Bohol: Cary’ophyllia (A.)

spinicarens, Caryophyllia (A.) spinigera, Flabellum (U.) deludens C89, Madrepora oculata.

Stn 5419. — 25.3.1909, 9°58'30"N, 123°46'00"E, 320 m, 12.5°C, between Cebu and Bohol: Flabellum (U.)

deludens C89.

Stn 5423. — 31.3.1909, 9°38'30"N, 121°11'00"E, 930 m, 9.9°C, Sulu Sea, Cagayan Isl.: Deltocyathus rotulus,

Madrepora oculata, Ericiocyathus echinatus, Rhombopsammia squiresi C89.

Stn 5424. — 31.3.1909, 9°37'50"N, 121°12'37"E, 622 m, 10.2°C, Sulu Sea, Cagayan Isl.: Fungiacyathus (B.)

turbinolioides, Madrepora oculata, Peponocyathus minimus, Rhombopsammia squiresi C89.

Stn 5425. — 31.3.1909, 9°37'45"N, 121°irOO"E, 906 m, 9.7°C, Sulu Sea, Cagayan Isl.: Deltocyathus rotulus,

Fungiacyathus (F.) stephanus C89, Madrepora oculata, Ericiocyathus echinatus.

Stn 5426. — 3.4.1909, 9°12'00"N, 1 18°28'00"E, 49 m, SE Palawan: Stephanophyllia neglecta C89.

Stn 5428. — 3.4.1909, 9°13'00"N, 118°51’15"E, 2022 m, 9.8°C, SE Palawan: Enallopsammia rostrata,

Flabellum (U.) ?conuis C89, Madrepora oculata.

Stn 5429. — 5.4.1909, 9°41’30"N, 118°50'22"E, 1402 m, E Palawan: Deltocyathus rotulus, Ericiocyathus

echinatus, Rhombopsammia squiresi C89 (not 5425).

Stn 5438. — 8.5.1909, 15°54'42"N, 1 19°44'42''E, 544 m, 7.9°C, W Luzon: Deltocyathus rotulus.

Stn 5439. — 9.5.1909, 15°58’15"N, 1 19°40'20"E, 1720 m, 2.6°C, W Luzon: Deltocyathus rotulus.

Stn 5440. — 10.5.1909, 16°33'52"N, 1 19°52'54"E, 315 m, 1 1.8°C, W Luzon: Stephanocyathus (A.) spiniger.

Stn 5444. — 3.6.1909, 12043'51"N, 124°50'50"E, 564 m, 7.4°C, N Samar: Caryophyllia (C.) scobinosa,

Deltocyathus magnificus, Stephanocyathus (O.) weberianus, Stephanocyathus (S.) regius.

Stn 5445. — 3.6.1909, 12°44'42"N, 124°59'50"E, 701 m, 6.8°C, N Samar: Caryophyllia (C.) scobinosa,

Deltocyathus rotulus, Fungiacyathus (F.) stephanus C89, Stephanocyathus (O.) weberianus, Stephanocyathus

(S.) regius.

36

S. D. CAIRNS & H. ZIBROWIUS

Sin 5447. — 4.6.1909, 13°28'00"N, 123°46'18ME, 567 m, 7.4°C, SE Luzon: Caryophyllia ( C .) scobinosa,

Deltocyathus rotulus, Stephanocyathus (S.) regius.

Stn 5453. — 7.6.1909, 13°12'00"N, 123°49,18”E, 267 m, SE Luzon: Flabellum (U.) deludens C89.

Stn 5454. — 7.6.1909, 13°12’00"N, 123°50'30"E, 280 m, SE Luzon: Flabellum (U.) deludens C89.

Stn 5483. — 30.7.1909, 10°27'30"N, 125°19’15"E, 135 m, E Leyte: Truncatoflabellum sp. C89* (ex T.

formosum).

Stn 5484. — 30.7.1909, 10°28’00"N, 125°20'00"E, 139 m, E Leyte: Truncatoflabellum incrustation C89* (ex T.

formosum).

Stn 5487. — 31.7.1909, 10°02'45"N, 125°05'33"E, 1340 m, 1 1.3°C, SE Leyte: Ericiocyathus echinatus.

Stn 5499. — 4.8.1909, 8°41'30"N, 124°35'40"E, 1014 m, 1 1.8°C, N Mindanao: Enallopsammia rostrata.

Stn 5505. — 5.8.1909, 8°37'15"N, 124°36'00"E, 403 m, N Mindanao: Caryophyllia (A.) spinicarens,

Deltocyathus philippinensis, Flabellum (F.) lamellulosum C89, Flabellum (U.) japonicum C89.

Stn 5506. — 5.8.1909, 8°40'00"N, 124°3r45''E, 479 m, 11.8°C, N Mindanao: Deltocyathus magnificus,

Deltocyathus philippinensis. Flabellum (U.) deludens C 89, Trochocyathus (A.) longispina, "Tropidocyathus"

pileus C89.

Stn 5508. — 5.8.1909, 8°17'24"N, 124°H'42"E, 494 m, 11.8°C, N Mindanao: Caryophyllia (A.) spinicarens.

Deltocyathus magnificus. Flabellum (II.) deludens C89, Gardineria philippinensis C89, "Tropidocyathus"

pileus C89.

Stn 5510. — 7.8.1909, 8°16'00"N, 124°03'50"E, 774 m, 11.7°C, N Mindanao: Fungiacyathus (F.) stephanus

C89.

Stn 5512. — 7.8.1909, 8°16'02"N, 123°58'26"E, 814 m, 1 1.6°C. N Mindanao: Ericiocyathus echinatus.

Stn 5513. — 7.8.1909, 8°16'45"N, 124°02'48"E, 924 m, 1 1.6°C, N Mindanao: Madrepora oculata, Notocyathus

conicus C89, Ericiocyathus echinatus, Rhombopsammia squiresi C89.

Stn 5516. — 9.8.1909, 8°46'00"N, 123°32'30"E, 320 m, 12.4°C, NW Mindanao: Madrepora oculata.

Stn 5519. — 9.8.1909, 8°47’00"N, 123°31'15"E, 333 m, 12.4°C, NW Mindanao: Crispatotrochus rubescens.

Stn 5523. — 10.8.1909, 8°48'44"N, 123°27'35"E, depth?, NW Mindanao: Flabellum (F.) magnificum C89,

Flabellum (U.) marenzelleri.

Stn 5527. — 1 1.8.1909, 9°22’30"N, 123°42'40"E, 717 m, 11.8°C, between Bohol and Siquijor: Caryophyllia (A.)

spinicarens, Caryophyllia (C.) octonaria, Deltocyathus magnificus. Deltocyathus rotulus, Flabellum (F.)

politum C89, Flabellum (U.) japonicum C89, Madrepora oculata, Truncatoflabellum paripavoninum C89.

Stn 5528. — 1 1.8.1909, 9°24'45"N, 123°39'15"E, 803 m, 11.7°C, between Bohol and Siquijor: Truncatoflabellum

paripavoninum C89.

Stn 5529. — 11.8.1909, 9°23'45"N, 123°39'30"E, 807 m, 11.7°C, between Bohol and Siquijor: Madrepora

oculata, Deltocyathus vaughani, Truncatoflabellum paripavoninum C89.

Stn 5535. — 19.8.1909, 9°20'30"N, 123°23'45"E, 567 m, 1L8°C, between Negros and Siquijor: Caryophyllia

(A.) spinicarens, Flabellum (U.) japonicum C89.

Stn 5536. — 19.8.1909, 9015'45MN, 123°22'00"E, 511 m, 11.9°C, between Negros and Siquijor: Caryophyllia

(A.) spinicarens, Deltocyathus magnificus, Deltocyathus vaughani, Flabellum (U.) deludens C89, Flabellum

(U.) japonicum C89.

Stn 5537. — 19.8.1909, 9°H'00"N, 123°23'00"E, 465 m, 11.9°C, between Negros and Siquijor: Caryophyllia

(A.) spinicarens, Flabellum (U.) japonicum C89, Trochocyathus (A.) longispina.

Stn 5538. — 19.8.1909, 9°08'15"N, 123°23'20"E, 468 m, ll.8°C, between Negros and Siquijor: Caryophyllia

(A.) spinicarens, Deltocyathus philippinensis, Flabellum (U.) japonicum C89.

Stn 5541. — 20.8.1909, 8°49'38"N, 123°34'30"E, 401 m, 1 1.8°C, NW Mindanao: Caryophyllia (A.) spinicarens,

Deltocyathoides orientalis C89* (ex P. australiensis), Madrepora oculata, Stephanocyathus (A.) spiniger.

Stn 5543. — 20.8.1909, 8°47'15"N, 123°35'00"E, 296 m, 12.5°C, NW Mindanao: Balanophyllia generatrix,

Coenosmilia arbuscula, Madrepora oculata.

Stn 5545. — 15.9.1909, 6o04'45"N, 121o20'20"E, 209 m, 14,6°C, Jolo: Stephanocyathus (A.) spiniger.

Stn 5551. — 17.9.1909, 5°54’48"N, 120°44'24"E, 353 m. 11.8°C, Jolo: Caryophyllia (C.) scobinosa,

Conotrochus funicolumna.

Stn 5554. — 18.9.1909, 5°52'27"N, 120°52'18"E, 46 m, Jolo: Tubastraea micranthus.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

37

Stn 5564. — 20.9.1909, 5°50'00"N, 120°31’00"E, 432 m, 11.3°C, between Jolo and Tawitawi: Caryophyllia (C.)

scobinosa, F label turn (U.) japonicum C89.

Stn 5565. — 21.9.1909, 5°51'42"N, 120°30'30"E, 445 m, 1 1.3°C, between Jolo and Tawitawi: Caryophyllia (A.)

spinicarens, Caryophyllia (C.) scobinosa, Deltocyathoides orientalis.

Stn 5567. — 21.9.1909, 5°48'00"N, 120°33'45"E, 490 m, 1 1.1°C, between Jolo and Tawitawi: Caryophyllia (C.)

secta, Conotrochus funicolumna, Deltocyathoides orientalis C89* (ex P. australiensis ), Deltocyathus

magnificus, Flabellum (U.) deludens C89, Notocyathus venustus C89, Trochocyathus (T.) gardineri,

"Tropidocyathus" pileus C89, Truncatoflabellum angustum.

Stn 5569. — 22.9.1909, 5°33T5"N, 120°15'30"E, 554 m, 11.3°C, N Tawitawi: Deltocyathoides orientalis C89*

(ex P. australiensis ), Notocyathus venustus C89, Stephanophyllia neglecta C89.

Stn 5574. — 23.9.1909, 5o30'45"N, 120°07'57"E, 622 m, N Tawitawi: Madrepora oculata.

Stn 5576. — 23.9.1909, 5°25'56"N, 120°03'39"E, 507 m, 11.8°C, N Tawitawi: Deltocyathoides orientalis C89*

(ex P. australiensis ), Notocyathus venustus C89, Tlirypticotrochus multilobatus C89.

Stn 5577. — 23.9.1909, 5°20'36"N, 1 19°58'51"E, 439 m, 12.4°C, N Tawitawi: Deltocyathoides orientalis C89*

(ex P. australiensis), Flabellum (F.) patens C89, Fungiacyathus (B.) paliferus C89, Peponocyathus folliculus

C89.

Stn 5579. — 25.9.1909, 4°54T5"N, 119°09'52"E, 320 m, 12.9°C, NE Borneo, Sabah: Fungiacyathus (B.)

paliferus C89, Notocyathus venustus, Peponocyathus minimus.

Stn 5580. — 25.9.1909, 4°52'45"N, 1 19°06'45ME, 296 m, 13.2°C, NE Borneo, Sabah: Javania insignis C89,

Rhizotrochus typus C89.

Stn 5582. — 26.9.1909, 4°19'54"N, 118°58’38"E, 1629 m, 3.5°C, NE Borneo, Sabah: Deltocyathus rotulus,

Fungiacyathus (F.) stephanus C89, Placotrochides scaphula C89.

Stn 5584. — 27.9.1909, 4o17'40"N, 118°57'42"E, 534 m, 6.8°C, NE Borneo, Sabah: Anthemiphyllia dentata,

Deltocyathoides orientalis C89* (ex P. australiensis), Idiotrochus kikutii C89, Javania pachytheca, Madrepora

arbuscula, Peponocyathus folliculus C89.

Stn 5585. — 28.9.1909, 4o07'00"N, 118°49'54"E, 871 m, 5.1°C, NE Borneo, Sabah: Aulocyathus recidivus,

Deltocyathus rotulus, Flabellum (U.) messutn C89, Leplopenus sp. A C89, Stephanocyathus (S.) regius.

Stn 5586.’— 28.9.1909. 4°06'50"N, 1 18o47'20"E, 635 m, 6.7°C, NE Borneo, Sabah: Aulocyathus recidivus,

Conotrochus funicolumna, Crispatotrochus rugosus, Deltocyathus rotulus, Deltocyathoides orientalis C89* (ex

P. australiensis), Dendrophyllia alcocki, Flabellum (F.) magnificum C89, Fungiacyathus (B.) turbinolioides

C89, Goniocorella dumosa, Idiotrochus kikutii C89, Lochmaeotrochus oculeus, Madrepora oculata,

Notocyathus sp. C89 (ex N. conicus), Peponocyathus minimus, Placotrochides scaphula C89, Stephanocyathus

( O .) weberianus, Stephanophyllia fungulus C89, Sympodangia albatrossi, Truncatoflabellum paripavoninum

C89.

Stn 5587. — 28.9.1909, 4°10'35"N, 118°37T2"E, 759 m, 6.7°C, NE Borneo, Sabah: Fungiacyathus (F.)

stephanus C89, Stephanocyathus (O.) weberianus.

Stn 5589. — 29.9.1909, 4°12'10MN, 1 18°38’08"E, 366 m, 7.6°C, NE Borneo. Sabah: Caryophyllia (C.) grandis,

Flabellum (U.) messum C89, Fungiacyathus (B.) granulosus C89, Paraconotrochus zeidleri, Placotrochides

scaphula C89, Stephanocyathus (A.) explanans, Truncatoflabellum paripavoninum C89.

Stn 5590. — 29.9.1909, 4°fo'50"N, 1 18°39'35"E, 567 m, 6.8°C, NE Borneo, Sabah: Caryophyllia (C.) grandis,

Flabellum (F.) magnificum C89, Fungiacyathus (B.) granulosus C89, Paraconotrochus zeidleri,

Truncatoflabellum paripavoninum C89.

Stn 5591. — 29.9.1909, 4°11'48"N, 118°38'20"E, 476 m, NE Borneo, Sabah: Deltocyathus rotulus, Flabellum

(U.) messum C89, Placotrochides scaphula C89.

Stn 5592. — 29.9.1909, 4°12'44"N, 118027'44"E, 558 m, 6.3°C, NE Borneo, Sabah: Caryophyllia (C.) grandis,

Deltocyathoides orientalis C89* (ex P. australiensis), Fungiacyathus (B.) granulosus C89, Lochmaeotrochus

oculeus, Paraconotrochus zeidleri, Trochocyathus (A.) longispina, Truncatoflabellum aculeatum.

Stn 5593. — 29.9.1909, 4°02’40"N, 1 1 8° 1 1'20"E, 70 m, NE Borneo, Sabah: Caryophyllia (A.) grayi, Endopachys

grayi, Rhizotrochus typus C89, Truncatoflabellum candeanum C89.

Stn 5601. — 13.11.1909, 1°13'10"N, 125°17'05"E, 1400 m, NE Celebes. Moluccan Sea: Deltocyathus rotulus,

Stephanocyathus (O.) weberianus.

Stn 5605. — 16.11.1909, 0°21'33"N, 121o34T0"E, 1184 m, Celebes, Gulf of Tomini: Fungiacyathus (F.)

stephanus C89.

38 S. D. CAIRNS & H. ZIBROWIUS

Stn 5606. — 17.11.1909, 0°16'28"N, 121°33'30"E, 1526 m. Celebes, Gulf of Tomini: Caryophyllia (C.)

comulum.

Stn 5618. — 27.11.1909. 0°37’00"N, 127°15'00"E, 763 m, Molucca Passage: Fungiacyathus (B.) sibogae C89,

Fungiacyathus (F.) stephanus C89.

Stn 5619. — 27.1 1.1909, 0°35'00"N, 127°14'40"E, 796 m, Molucca Passage. Fungiacyathus (B.) sibogae C89,

Fungiacyathus (F.) stephanus C89.

Stn 5622. — 29.1 1.1909, 0°19’20"N, 127°28'30"E, 503 m, Makian: Caryophyllia (A.) spinicarens.

Stn 5625. — 29.11.1909, 0°07'00"N, 127°28’00"E, 421 m, Kayoa: Caryophyllia (A.) spinicarens, Madrepora

oculata.

Stn 5626. — 29.1 1.1909, 0°07'30"N, 127°29'00"E, 485 m, Kayoa: Caryophyllia (A.) spinicarens.

Stn 5630. — 2.12.1909, 0°56'30"S, 128°05'00"E, 1041 m, S Patiente Strait, Doworra: Fungiacyathus (F.)

stephanus C89.

Stn 5634. — 3.12.1909, 1°54’00"S, 127°36'00"E, 602 m, Pitt Passage, Gomomo: Caryophyllia (C.) diomedeae,

Javania pachytheca.

Stn 5636. — 3.12.1909, 1°55'00"S, 127°42’30"E, 2309 m, Pitt Passage, Gomomo: Caryophyllia (C.) diomedeae.

Stn 5645. — 16.12.1909, 5°29’06"S, 122°36’06"E, 377 m, Butung Strait: Madrepora oculata.

Stn 5647. — 16.12.1909, 5°34'00"S, 122°18'15"E, 950 m, Butung Strait: Flabellum (U.) messum C89.

Stn 5648. — 16.12.1909, 5°35'00"S, 122°20’00"E, 1023 m, 4.0°C, Butung Strait: Fungiacyathus (B.) sibogae

C89, Fungiacyathus (F.) stephanus C89, Deltocyathus rotulus, Stephanocyathus (O.) weberianus,

Truncatoflabellum paripavoninum C89.

Stn 5650. — 17.12.1909, 4°53'45"S, 121°29’00"E, 988 m, 4.5°C, S Celebes, Gulf of Boni: Caryophyllia (C.)

scobinosa, Stephanocyathus (O.) weberianus, Stephanocyathus <S.) regius.

Stn 5656. — 19.12.1909, 3°17'40"S, 120°36'45"E, 886 m, 5.1°C, S Celebes, Gulf of Boni: Caryophyllia (C.)

diomedeae, Truncatoflabellum paripavoninum C89.

Stn 5658. — 19.12.1909, 3°32'40"S, 120°31’30"E, 933 m, 5.1°C, S Celebes. Gulf of Boni: Truncatoflabellum

formosum C89, Truncatoflabellum paripavoninum C89.

Stn 5668. — 29.12.1909, 2°28'15"S, 118°49'00"E, 1649 m, 3.4°C, Macassar Strait: Deltocyathus rotulus,

Fungiacyathus (F.) stephanus C89.

Stn 5670. — 30.12.1909, 1°19'00"S, 1 18°43’00"E, 2161 m, 3.4°C, Macassar Strait: Caryophyllia (C.) comulum,

Fungiacyathus (F.) stephanus C89, Stephanocyathus (S.) regius.

Stn 5671. — 30.12.1909, 1°05'00"S, 118°56'00"E, 1647 m, 3.4°C, Macassar Strait: Stephanocyathus (O.)

weberianus.

"Alpha Helix"

Stn 1769. — date?, 4°32’S, 129°52'E, 25 m, Banda: Tubastraea micranthus.

Stn 79-M21 . — 7.6.1979, 8°45.0'S, 144°05.8'E, 55 m. Gulf of Papua: Paracyathus rotundatus.

Stn 79-M26. — 10.6.1979, 9°29.7'S, 147°06.6'E, 1 m, Papua-New Guinea, Port Moresby: Endopsammia

philippensis.

Stn 79-M48. — 18.6.1979, 6°41.7'S 147°53.1'E, 0-8 m, Papua-New Guinea, Cape Cretin: Endopsammia

philippensis.

Stn 79-M59. — 21.6.1979, 3°23.8'S, 143°40.7'E, 1-6 m, Papua-New Guinea, Schouten Isl.: Endopsammia

philippensis.

Stn 79-M122. — 15.7.1979, 1°40.9'N, 127°32.2'E, 0-3 m, NW Halmahera: Tubastraea micranthus.

Stn 79-M140. — 23.7.1979, 9°36.5'N, 123°53’E, 14-20 m, SW Bohol: Culicia stellata.

"Challenger"

Stn 174. — 3.8.1874, Fiji, Kandavu, 384 m: Caryophyllia (A.) dentata.

Stn 190. — 12.9.1874, 8°56'S, 136°05'E, 90 m, Arafura Sea: Paracyathus sp.

Stn 192. — 26.9.1874, 5°42'S, 132°25'E, 236 m, Kai Isl.: Balanophyllia cornu, Balanophyllia rediviva,

Caryophyllia ( C.) transversalis.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

39

Stn 194. — 29.9.1874, 4°33'S, 129°58'E, 366 m or 659 m, Banda: Madrepora arbuscula.

Stn 201. — 26.10.1874, Philippines, Basilan Strait, 187 m: Balanophyllia gemma, Balanophyllia parvula.

Stn 219. — 10.3.1874, 1°50'S, 146°42'E, 274 m, Admiralty Isl.: Paraconotrochus zeidleri.

Corindon 2, "Coriolis"

Stn 208. — 31.10.1980, 0°14.6'S, 1 17°52.0'E, 150 m, W Makassar Strait: Caryophyllia (A.) grayi.

Stn 210. — 31.10.1980, 0°12.6'S, 1 17°53.5'E, 338 m, W Makassar Stait: Fungiacyathus (F.) paliferus,

Peponocyathus minimus.

Stn 216. — 1.1 1.1980, 0°40.1'N(!), 117°51.4'E, 96 m, W Makassar Strait: Asterosmilia marchadi, Caryophyllia

(A.) grayi, Flabellum (F.) pavoninum, Truncatoflabellum formosum.

Stn 220. — 2.1 1.1980, 0°13.6'S, 1 18°12.3'E, 2350 m, W Makassar Strait: Caryophyllia (C.) cornulum.

Stn 235. — 4.11.1980, 0°04.7'S, 119°48.4'E, 1110 m, E Makassar Strait: Madrepora minutiseptum CSD.

Notocyathus conicus C89, Peponocyathus minimus.

Stn 240. — 5.1 1.1980, 0°37.6'S, 1 19°33.5'E, 675 m, E Makassar Strait: Caryophyllia (C.) ambrosia ambrosia,

Caryophyllia (C.) scobinosa, Rhombopsammia squiresi.

Stn 241. — 6.11.1980, 0°57.7'S, 1 19° 15.3'E, 1525-1550 m, E Makassar Strait: Caryophyllia (C.) cornulum,

Deltocyathus rotulus, Stephanocyathus (S.) regius.

Stn 248. — 6.1 1.1980, 0°54.2'S, 1 19°28.7'E, 170 m, E Makassar Strait: Balanophyllia imperialis, Balanophyllia

rediviva, Caryophyllia (C.) hawaiiensis, Cyathohelia axillaris, Deltocyathoides orientalis, Javania insignis,

Madracis sp. A, Madrepora minutiseptum, Neohelia cf. porcellana, Rhizosmilia sagamiensis, Thalamophyllia

tenuescens, Trochocyathus (T. ) philippinensis, Truncatoflabellum pusillum.

Stn 251. — 6.1 1.1980, 0°53.7'S, 1 19°29.6'E, 65 m, E Makassar Strait: Asterosmilia marchadi, Caryophyllia (A.)

grayi, Trochocyathus (T.) semperi.

Stn 260. — 6. 1 1 .1980, 1°56.9'S, 1 19°17.6'E, 15-50 m, E Makassar Strait: Truncatoflabellum aculeatum.

Stn 261. — 6.1 1.1980, 1°56.8'S, 1 19°16.8'E, 60 m, E Makassar Strait: Asterosmilia marchadi, Caryophyllia (A.)

grayi.

Stn 263. — 6.1 1.1980, 1°56.8'S, 1 19°16.7'E, 80 m, E Makassar Strait: Asterosmilia marchadi.

Stn 266. — 7.11.1980, 1°56.6’S, 119°15.8'E, 95 m, E Makassar Strait: Asterosmilia marchadi, Balanophyllia

desmophyllioides.

Stn 275. — 7.1 1.1980, 1°53.9’S, 1 19°13.7'E, 530 m, E Makassar Strait: Fungiacyathus (F.) paliferus.

Stn 286. — 9.11.1980, 2°04.4'S, 118°46.9’E, 1710-1730 m, E Makassar Strait: Caryophyllia (C.) cornulum,

Deltocyathus rotulus, Fungiacyathus (F.) stephanus.

Stn 292. — 10.11.1980, 2°37.2'S, 117°53.0'E, 46 m, Central Makassar Strait: Balanophyllia carinata,

Truncatoflabellum aculeatum.

Danish Expedition to the Kei Islands (- Deki)

Stn unnumbered. — 22.2.1922, Bay of Amboina, 91 m: Neohelia cf. porcellana.

Stn unnumbered. — 3.3.1922, Bay of Amboina, 91 m: Endocyathopora laticostata.

Stn unnumbered. — 3.6.1922, Banda, Komkir, 75-90 m: Balanophyllia carinata.

Stn unnumbered. — 3.6. 1922, Banda, Waling Bezar, 30 m: Incertae sedis.

Stn unnumbered. — Between Neira and Lontor, Banda Isl., 70-90 m: Caryophyllia (A.) dentata.

Stn 1. — 30.3.1922, 5°34'S, 132°50'E, 370 m, Kai Isl.: Enallopsammia pusilla.

Stn 2. — 31.3.1922, 5°32'S, 132°27'E, 180-222 m, Kai Isl.: Caryophyllia (A.) dentata, Fungiacyathus (F.)

paliferus, Letepsammia formosissima, Stephanocyathus (A.) spiniger.

Stn 3. — 31.3.1922, 5°32'S, 132°36'E, 245 m, Kai Isl.: Caryophyllia (A.) dentata, Caryophyllia (C.) crosnieri,

Caryophyllia (C.) rugosa, Caryophyllia (C.) transversalis, Conotrochus brunneus, Crispatotrochus rubescens,

Deltocyathus magnificus, Dendrophyllia arbuscula, Flabellum (F.) lamellulosum, Flabellum (F.) magnificum,

Flabellum (F.) politum, Fungiacyathus (F.) paliferus, Letepsammia formosissima, Rhombopsammia niphada,

Stephanocyathus (A.) spiniger, Truncatoflabellum pusillum.

Stn 4. — 3.4.1922, 5°31'40"S, 132°26'E, 250 m, Kai Isl.: Caryophyllia (A.) grayi, Conotrochus brunneus,

Letepsammia formosissima, Stephanocyathus (A.) spiniger, Tubastraea micranthus CSD.

Source :

40

S. D. CAIRNS & H. ZIBROWIUS

Stn 5. — 4.4.1922, 5°31'30"S, 132°38'E, 90-250 m, Kai Isl.: Conotrochus brunneus, Fungiacyathus (F.)

paliferus, Rhizotrochus typus, Stephanocyathus (A.) spiniger.

Stn 6_ _ 4.4.1922, 5°32'S, 132°36'30"E, 210 m, Kai Isl.: Asterosmilia marchadi, Caryophyllia (A.) dentata,

Caryophyllia (A.) grayi, Caryophyllia (C.) transversalis, Conotrochus brunneus, Deltocyathoides orientalis,

Deltocyathus magnificus, Deltocyathus rotulus, Fungiacyathus (F.) paliferus, Notocyathus comcus,

Stephanophyllia complicata, Stephanophyllia fungulus, Trochocyathus (T.) philippinensis, Trochocyathus (T.)

semperi, Stephanocyathus (A.) spiniger.

Sm 7. — 5.4.1922, 5°38'30"S, 132°26'E, 196 m, Kai Isl.: Balanophyllia cornu, Balanophyllia crassiseptum,

Deltocyathoides orientalis, Dendrophyllia arbuscula, Trochocyathus (T.) caryophylloides.

Stn 8. — 5.4.1922, 5°39'S, 132°26'E, 300 m, Kai Isl.: Trochocyathus (T.) rhombocolumna.

Stn JO. —6.4.1922, off Doelah, 50 m, Kai Isl.: Asterosmilia marchadi, Balanophyllia carinata, Balanophyllia

stimpsonii, Peponocyathus folliculus, Placotrochus laevis, Trochocyathus (T.) burchae, Trochocyathus (T.)

apertus, Tubastraea micranthus.

Sm 12. — 9.4.1922, 5°30'S, 132°35'E, 325 m, Kai Isl.: Balanophyllia generatrix, Caryophyllia (C.) crosnieri,

Deltocyathus magnificus, Enallopsammia pusilla, Flabellum magnificum, Trochocyathus (A.) brevispina.

Sm 13. —9.4.1922, 5°31'S, 132°36'30"E, 275 m, Kai Isl.: Trochocyathus (A.) brevispina.

Stn 14. — 10.4.1922, S of Doe Roa, 40 m, Kai Isl.: Balanophyllia carinata, Balanophyllia stimpsonii,

Truncatoflabellum aculeatum.

Stn 15. — 10.4.1922, S of Doe Roa, ca. 5-20 m, Kai Isl.: Trochocyathus (T.) cooperi.

Sm 17. — 12.4.1922, 5°34'40"S, 132°35'E, 100 m, Kai Isl.: Trochocyathus (T.) cooperi.

Stn 18. — Doe Roa Strait, 40 m, Kai Isl.: Balanophyllia carinata, Balanophyllia stimpsonii, Placotrochus laevis.

Stn 20. — 14.4.1922, Doe Roa Bassin, 50 m, Kai Isl.: Balanophyllia carinata.

Stn 22. — 15.4.1922, 5°30'40"S, 132°51'E, 340 m, Kai Isl.: Madrepora oculata, Stephanocyathus (A.) spiniger.

Stn 24. — 15.4.1922, 5°37’S, 132°56'E, 100 m, Kai Isl.: Balanophyllia carinata, Caryophyllia (C.) lamellifera,

Cyathelia axillaris, Deltocyathoides orientalis, Dendrophyllia arbuscula, Fungiacyathus (B.) variegatus,

Fungiacyathus (F.) paliferus, Javania insignis, Madracis cf. pharensis, Notocyathus venustus, Phyllangia

papuensis, Rhizotrochus typus, Trochocyathus (T.) cooperi, Trochocyathus (T.) philippinensis.

Stn 25. — 16.4.1922, 5°34'20"S, 132°55'E, 85 m, Kai Isl.: Caryophyllia (A.) grayi, Caryophyllia (C.)

hawaiiensis, Rhizotrochus typus.

Stn 26. — 16.4.1922, 5°38'S, 132°55'20"E, 90 m, Kai Isl.: Deltocyathoides orientalis, Rhizotrochus typus.

Stn 27. — 17.4.1922, 2 miles N of Elat, 60-70 m, Kai Isl.: Rliizosmilia elata, Rhizotrochus typus.

Stn 30. — 18.4.1922, between Doe Roa and Kai Doelah, 40 m, Kai Isl.: Balanophyllia stimpsonii.

Stn 31. — 18.4.1922, Doe Roa Bassin, 50 m, Kai Isl.: Balanophyllia carinata.

Stn 32. — 22.4.1922, 5°32'20"S, 132°34'E, 260 m, Kai Isl.: Caryophyllia (C.) transversalis.

Stn 33. —22.4.1922, 5°31'S, 132°34’E, 285 m, Kai Isl.: Madrepora oculata.

Stn 35. — 23.4.1922. Bay N of Noehoe Roa, 32 m, Kai Isl.: Asterosmilia marchadi.

Stn 38. — 24.4.1922, NE of Doe Roa, 35 m, Kai Isl.: Balanophyllia carinata.

Stn 41. — 25.4.1922, 5°28’40"S, 132°28'E, 245 m, Kai Isl.: Caryophyllia (C.) transversalis, Deltocyathus

magnificus, Flabellum (F.) lamellulosum, Flabellum (F.) magnificum, Letepsammia formosissima,

Stephanocyathus (A.) spiniger, Trochocyathus (A.) brevispina.

Stn 42. — 26.4.1922, 5°35'S, 132°29'E, 225 m, Kai Isl.: Caryophyllia (A.) spinigera, Caryophyllia (C.)

transversalis, Conotrochus brunneus, Flabellum (F.) magnificum, Fungiacyathus ( F.) paliferus, Letepsammia

formosissima, Stephanocyathus (A.) spiniger.

Stn 44. — 30.4.1922, 5°39'S, 132°23'E, 268 m, Kai Isl.: Alatotrochus rubescens, Caryophyllia (A.) grayi,

Caryophyllia (C.) transversalis, Conotrochus funicolumna, Deltocyathoides orientalis, Deltocyathus

magnificus, Deltocyathus suluensis, Letepsammia formosissima, Notocyathus venustus, Placotrochides laevis,

Trochocyathus (T.) philippinensis.

Stn 45. — 1.5.1922, 5°48'30"S, 132°14'E, 270 m, Kai Isl.: Madrepora oculata.

Stn 46. — 2.5.1922, 5°47'20"S, 132°13'E, 300 m, Kai Isl.: Balanophyllia gemma, Balanophyllia parvula,

Caryophyllia (C.) quadragenaria, Caryophyllia (C.) transversalis, Conotrochus brunneus, Deltocyathoides

orientalis, Deltocyathus magnificus, Flabellum (U.) deludens, Idiotrochus kikutii, Letepsammia formosissima,

Thrypticotrochus multilobatus, Truncatoflabellum pusillum.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

41

Stn 48. — 3.5.1922, 5°40'10"S, 130°21'E, 263 m, Kai Isl.: Balanophyllia cornu, Bourneotrochus stellulatus,

Caryophyllia (A.) dentata, Caryophyllia (C.) transversalis, Conotrochus brunneus, Crispatotrochus rubescens,

Fungiacyathus (B.) variegatus, Rhizotrochus flabelliformis.

Stn 49. — 3.5.1922, 5°37'10"S, 132°23'E, 245 m, Kai Isl.: Alatotrochus rubescens, Balanophyllia cornu,

Caryophyllia (A.) grayi, Deltocyathoides orientalis, Endopachys grayi, Flabellum ( U.) deludens, Fungiacyathus

(F.) paliferus, Fungiacyathus (F.) stephanus, Letepsammia formosissima, Stephanocyathus (A.) spiniger,

Stephanophyllia fungulus, Trochocyathus (T.) philippinensis, Tropidocyathus lessonii, "Tropidocyathus"

pileus.

Stn 50. — 4.5.1922, 5°34'S, 132°25'40"E, 233 m, Kai Isl.: Balanophyllia gigas, Caryophyllia (C.) transversalis,

Flabellum (F.) lamellulosum, Fungiacyathus (B.) sibogae, Fungiacyathus (B.) variegatus, Madrepora oculata,

Notocyathus venustus, Stephanocyathus (A.) spiniger, Stephanophyllia complicata, Stephanophyllia neglecta.

Stn 51. — 7.5.1922, 5°46'30"S, 132°51'E, 348 m, Kai Isl.: Deltocyathus magnificus, Deltocyathus suluensis.

Stn 52. — 7.5.1922, 5°46'S, 132°49'35"E, 352 m, Kai Isl.: Conotrochus funicolumna, Deltocyathus suluensis,

Flabellum (F.) lamellulosum, Flabellum (U.) marenzelleri, Lochmaeotrochus oculeus, Madrepora oculata,

Paraconotrochus zeidleri, Stephanocyathus (A.) spiniger, Stephanophyllia complicata, Stephanophyllia

fungulus, "Tropidocyathus" labidus.

Stn 53. — 9.5.1922, 5°36'S, 132°55'E, 85 m, Kai Isl.: Caryophyllia (A.) grayi, Caryophyllia (C.) hawaiiiensis,

Deltocyathoides orientalis, Flabellum (F.) politum, Rhizotrochus typus, Trochocyathus (T.) cooperi,

Trochocyathus (T.) semperi, Tropidocyathus lessonii. Truncatoflabellum pusillum.

Stn 54. — 9.5.1922, 5°34'S, 132°55'E, 85 m, Kai Isl.: Flabellum (F.) politum, Rhizotrochus typus,

Thalamophyllia tenuescens, Truncatoflabellum candeanum.

Stn 56. — 10.5.1922, 5°30'20"S, 132°51'E, 345 m, Kai Isl.: Flabellum (U.) hoffmeisteri, Lochmaeotrochus

oculeus, Rhizotrochus flabelliformis.

Stn 57. — 10.5.1922, 5°32’S, 132°49'25"E, ca. 200 m, Kai Isl.: "Cryptotrochus" venustus, Deltocyathus

suluensis, Fungiacyathus (B.) sibogae.

Stn 58. — 12.5.1922, 5°29'S, 132°37'E, 290 m, Kai Isl.: Balanophyllia generatrix, "Cryptotrochus" venustus,

Conotrochus brunneus, Deltocyathus rotulus, Deltocyathus suluensis, Fungiacyathus (F.) paliferus,

Letepsammia formosissima, Madrepora oculata, Truncatoflabellum candeanum.

Stn 59. — 12.5.1922, 5°28'S, 132°36'E, 385 m, Kai Isl.: Balanophyllia gemma, Caryophyllia (C.) quadragenaria,

Crispatotrochus rubescens, Confluphyllia juncta, Deltocyathus andamanicus, Enallopsammia pusilla,

Madrepora oculata, Polymyces wellsi, Rhizotrochus flabelliformis, Trochocyathus (T.) rhombocolumna.

Stn 60. — 14.5.1922, S of Doe Roa, 25 m, Kai Isl.: Truncatoflabellum aculeatum.

Stn 61. — 14.5.1922, between Doe Roa and Kai Doelah, 50 m, Kai Isl.: Enallopsammia pusilla.

Stn 62. — 15.5.1922, 5°29'25"S, I32°50'E, 290 m, Kai Isl.: Cary-ophyllia (A.) spinicarens, Caryophyllia (C.)

transversalis, Deltocyathus suluensis, Deltocyathus vaughani, Notocyathus venustus.

Stn 63. — 16.5.1922, 5°32'S, 132°36'25"E, ca. 250 m, Kai Isl.: Balanophyllia crassiseptum, Caryophyllia (A.)

dentata, Caryophyllia (C.) transversalis, Conotrochus brunneus, Flabellum (F.) lamellulosum, Flabellum (F.)

politum, Fungiacyathus (B.) variegatus, Fungiacyathus (F.) paliferus, Letepsammia formosissima,

Stephanocyathus (A.) spiniger, Truncatoflabellum pusillum.

Stn 64. — 26.7.1922, 5°51'S, 106°22'E, 35 m, Java Sea: Balanophyllia carinata, Balanophyllia stimpsonii,

Placotrochus laevis.

Stn 65. — 27.7.1922, 5°52’05"S, 106°17'E, 25 m, Java Sea: Placotrochus laevis.

Stn 66. — 27.7.1922, 5°54'S, 106°12'E, 24 m, Java Sea: Placotrochus laevis.

Stn 67. — 27.7.1922, 5°48'S, 106°12'E, 38 m, Java Sea: Placotrochus laevis, Truncatoflabellum spheniscus.

Stn 68. — 27.7.1922, 5°47’S, 106°14'E, 55 m, Java Sea: Balanophyllia stimpsonii, Paracyathus rotundatus.

Stn 69. — 27.7.1922, 5°47'S, 106°17'E, 50 m, Java Sea: Placotrochus laevis.

Stn 70. — 28.7.1922, 5°40'S, 106°21'E, 35m, Java Sea: Balanophyllia carinata.

Stn 71. — 28.7.1922, 5°40'S, 106°08'E, 54 m, Java Sea: Paracyathus rotundatus, Truncatoflabellum spheniscus.

Stn 73. — 28.7.1922, 5°57'S, 105°57'E, 30 m, Sunda Strait: Eguchipsammia gaditana.

Stn 74. — 29.7.1922, 6°03’S, 105°54'E, 30 m, Sunda Strait: Rhizopsammia nuda, Truncatoflabellum spheniscus.

Stn 82. — 30.7.1922, 6°38'S, 105°21'E, 35 m, Sunda Strait: Balanophyllia carinata, Balanophyllia stimpsonii,

Paracyathus rotundatus, Trochocyathus (T.) burchae, Truncatoflabellum aculeatum.

42 S. D. CAIRNS & H. ZIBROWIUS

Stn 84. — 31.7.1922, 5°55'S, 105°31'E, 38 m, Sunda Strait: Balanophyllia carinata, Balanophyllia stimpsonii,

Truncatoflabellum aculeatum.

Stn 89. — 31.7.1922, 5°55'S, 105°31'E, 38 m, Sunda Strait: Placotrochus laevis, Truncatoflabellum aculeatum.

Sm 90. — 1.8.1922, 5°55'S, 105°30’E, 36 m, Sunda Strait: Trochocyathus (T.) cooperi, Truncatoflabellum

aculeatum. Truncatoflabellum spheniscus.

Stn 91. — 1.8.1922, 5°53’S, 105°27'E, 42 m, Sunda Strait: Placotrochus laevis.

Stn 92. — 1.8.1922, 5°49'S, 105°29'E, 32 m, Sunda Strait: Truncatoflabellum aculeatum.

Sm 95. — 1.8.1922, 5°44'S, 105°20'E, 25 m, Sunda Strait: Trochocyathus (T.) cooperi.

Sm 100. — 3.8.1922, 5°49’S, 105°25'E, 54 m, Sunda Strait: Balanophyllia stimpsonii.

Stn 103. — 4.8.1922, 5°52'S, 106°05'E, 52 m, Sunda Strait: Trochocyathus (T.) cooperi. Truncatoflabellum

spheniscus.

Stn 104. — 4.8.1922, 5°52’S, 105°04'E, 38 m, Java Sea: Cyathelia axillaris. Rhizopsammia nuda.

Stn 105. — 5.8.1922, 5°56'S, 106°07'E, 13 m, Java Sea: Cyathelia axillaris.

Stn 106. — 5.8.1922, 5°50'S, 106°16'E, 32 m, Java Sea: Truncatoflabellum spheniscus.

Stn 1 10. — 5.8.1922, 5°25'S, 105°53'E, 12 m, Java Sea: Placotrochus laevis.

Stn 1 16. — 7.8.1922, 5°57’S, 106°34'E, 22 m, Java Sea: Placotrochus laevis.

Estase 2, "Jean Charcot"

Stn 42/CP6. — 5.12.1984, 4°38.00'N, 1 19°49.00’E, 2570 m, S Sibutu: Flabellum (U.) conuis.

"Galathea"

Stn 330. — 15.5.1951, 4 miles SE Singapore, 40 m: Truncatoflabellum spheniscus.

Stn 423. — 25.7.1951, 10°27'N, 124°18'E, 750 m, E Cebu: Caryophyllia (A.) spinicarens.

Stn 436. — 9.8.1951. — 10°12'N, 124°14’E, 710 m, E Cebu: Caryophyllia (A.) spinicarens. Deltocyatlius

magnificus. Flabellum (U.) japonicum, Madrepora oculata.

Stn 443. — 16.8.1951, 8C48'N, 124°09'E, 1500 m, N Mindanao: Madrepora oculata.

Stn 476. — 1 1.9.1951, 9°04'S, 114°43'E, 1555 m, S Bali: Deltocyathus rotulus.

Stn 477. — 9°01'N, 114°48'E, 780 m, S Bali: Peponocyathus minimus.

Stn 480. — 19.9.1951, 8°49'S, 1 15°00'E, 440 m, S Bali: Fungiacyathus (F.) variegatus.

Stn 485. — 12.9.1951, 8°48'S, 1 15°16'E, 62 m, S Bali: Blastotrochus nutrix.

Stn 488. — 12.9.1951, 8°49'S, 1 15°19’E, 202 m, S Bali: Balanophyllia desmophyllioides.

Stn 489. — 13.9.1951, 7°38'S, 1 16°08'E, 1 160 m, N Lombok: Flabellum (U.) conuis.

Stn 490. — 14.9.1951, 5°25'S, 1 17°03'E, 570-545 m, SW Sulawesi: Cryptotrochus javanus (see Cairns, 1988),

Fungiacyathus (B.) turbinolioides, Madrepora oculata, Premocyathus dentiformis.

Stn 500. — 25.9.1951, 7°34'N, 132°44'E, 390 m, Arafura Sea, E Tanimbar Isl.: Balanophyllia cornu.

Conotrochus funicolumna, Deltocyathoides orientalis, Paraconotrochus zeidleri, Deltocyathus suluensis,

Rhizotrochus flabelliformis, "Tropidocyathus" labidus.

Stn 501. — 27.9.1851, 10°43'S, 139°17'E, 54 m, Arafura Sea: Truncatoflabellum spheniscus.

"Gazelle"

Stn 37. — 25.8.1875, 6°29.5'S, 154°45'E, 88 m, Bougainville Isl.: Phyllangia papuensis.

"Hakuho Maru"

Stn KH72-1-8. — 25.5.1972, 8°44.6'N, 1 19°05.4'E, 2030 m, Sulu Sea: Flabellum (U.) conuis.

Stn KH72-1-20. — 10.6.1972, 5°40.9'N, 119°46.3'E, 460-514 m, Sulu Sea, Sibutu Passage: Alatotrochus

rubescens, Anthemiphyllia dentata, Conotrochus brunneus, Conotrochus funicolumna, Deltocyathoides

orientalis, Deltocyathus magnificus, Enallopsammia pusilla, Fungiacyathus (F.) turbinolioides, Madrepora

oculata, Notocyathus conicus, Stephanophyllia fungulus, "Tropidocyathus" pileus.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

43

Stn KH72-1-26. — 18-19.6.1972, 9°27.0'S, 127°58.6'E, 610-690 m, SE Timor: Caryophyllia (C.) ambrosia

ambrosia, Fungiacyathus (F.) stephanus, Stephanocyathus ( 0 .) weberianus, Stephanocyathus (S.) regius.

Stn KH72-1-28. — 19.6.1972, 9°34.4'S, 128°06.0'E, 295-296 m, SE Timor: Balanophyllia cornu, Conotrochus

funicolumna, Deltocyathus magnificus, Flabellum (F.) lamellulosum, Flabellum (F.) patens, Flabellum (U.)

marenzelleri, Madrepora oculata, Letepsammiaformosissima.

Stn KH72-1-29. — 7.6.1972, 12°17.3'S, 129°40.9'E, 49-52 m, NW Australia, Beagle Gulf: Truncatoflabellum

aculeatum, Truncatoflabellum spheniscus.

Stn KH72-1-30. — 25.6.1972, 12°24.8'S, 128°00.1'E, 115 m, NW Australia, Beagle Gulf: Balanophyllia

imperialis, Truncatoflabellum aculeatum, Truncatoflabellum spheniscus.

Stn KH72-1-50. — 10-11.7.1972, 6°51.6'N, 108°47.2'S, 132-137 m, South China Sea: Caryophyllia (A.) grayi,

Letepsammia formosissima, Stephanocyathus (A.) spiniger.

Sin KH72-1-52. — 11.7.1972, 7°26.3'N, 109°14.9'E, 265-286 m, South China Sea: Caryophyllia (A.)

spinicarens, Conotrochus brunneus, Flabellum (F.) lamellulosum, Flabellum (U.) deludens, Madrepora oculata.

Stn KH73-2-44-2. — 18.3.1973, 21°42.1'N, 1 17°36.8'E, 412-430 m, South China Sea: Crispatotrochus

rubescens, Dendrophyllia alcocki, Desmophyllum dianthus, Enallopsammia pusilla, Goniocorella dumosa,

Lochmaeotrochus oculeus, Madrepora oculata.

Stn KH85-1-A1. — 12.2.1985, 5°47.3’S, 1 19°35.4'E, 250-285 m, S Celebes: Flabellum (U.) marenzelleri.

Stn KH85-1-A2. — 12.2.1985, 5°47.1'S, 119°35.5'E, 250-285 m, S Celebes: Balanophyllia cornu, Balanophyllia

desmophyllioides, Conotrochus brunneus, Flabellum (U.) marenzelleri.

Karubar, "BarunaJaya I"

Stn 1. — 22.10.1991, 5°46'45"S, 132°11T0"E, 156-305 m, Kai Isl.: Anthemiphyllia dentata, Caryophyllia (C.)

hawaiiensis, Conotrochus brunneus, Deltocyathus Stella, Flabellum (F.) pavoninum, Letepsammia superstes,

Stephanophyllia neglecta, Tropidocyathus lessonii, Truncatoflabellum formosum, Truncatoflabellum

mortenseni.

Stn 2. — 22.10.1991, 5°47'00"S, 132°H'35"E, 209-300 m, Kai Isl.: Alatocyathus rubescens, Anthemiphyllia

frustum, Caryophyllia (C.) quadragenaria, Caryophyllia (C.) transversalis, Conotrochus brunneus,

"Cryptotrochus" venustus, Deltocyathoides orientalis, Deltocyathus rotulus, Deltocyathus Stella, Deltocyathus

suluensis, Endopachys grayi, Flabellum (F.) politum, Flabellum (U.) marenzelleri, Guynia annulata,

Idiotrochus kikutii, Letepsammia formosissima, Notocyathus conicus, Notocyathus venustus, Peponocyathus

minimus, Stephanophyllia complicata, Thrypticotrochus multilobatus, Trochocyathus (A.) brevispina,

Trochocyathus (T.) discus, Trochocyathus (T.) philippinensis, "Tropidocyathus" labidus, Tropidocyathus

lessonii, "Tropidocyathus" pileus, Truncatoflabellum angustum, Truncatoflabellum dens.

Stn 3. — 22.10.1991, 5°47'40"S, 132°12'H"E, 278-300 m, Kai Isl.: Alatocyathus rubescens, Bourneotrochus

stellulatus, Caryophyllia (C.) diomedae, Caryophyllia (C.) quadragenaria, Caryophyllia (C.) secta, Conotrochus

brunneus, "Cryptotrochus" venustus, Deltocyathoides orientalis, Deltocyathus magnificus, Deltocyathus

suluensis, Fungiacyathus (B.) fissidiscus, Fungiacyathus (B.) granulosus, Fungiacyathus (F.) ?paliferus

(irregular), Guynia annulata, Idiotrochus kikutii, Leptopenus sp., Letepsammia formosissima, Notocyathus

venustus, Peponocyathus minimus, Stephanophyllia complicata, Thrypticotrochus multilobatus,

Trochocyathus (A.) brevispina, Trochocyathus (T.) discus, Trochocyathus philippinensis, "Tropidocyathus"

labidus, "Tropidocyathus" pileus, Truncatoflabellum angustum, Truncatoflabellum formosum.

Stn 5. — 22.10.1991, 5°46'39"S, 132°20'04"E, 285-323 m, Kai Isl.: Caryophyllia crosnieri, Flabellum (F.)

patens, Gardineria paradoxa, Stephanophyllia complicata.

Stn 7. — 22.10.1991, 5°47'35"S, 132°20’39"E, 282-287 m, Kai Isl.: Balanophyllia cornu, Bourneotrochus

stellulatus, Caryophyllia (C.) transversalis, Conotrochus brunneus, "Cryptotrochus" venustus, Deltocyathoides

orientalis, Deltocyathus magnificus, Deltocyathus suluensis, Fungiacyathus (B.) granulosus, Fungiacyathus

(B.) fissidiscus, Fungiacyathus (F.) ?paliferus (irregular), Guynia annulata, Idiotrochus kikutii, Leptopenus

sp. A, Letepsammia formosissima, Letepsammia superstes, Notocyathus venustus, Peponocyathus minimus,

Stephanophyllia complicata, Thrypticotrochus multilobatus, Trochocyathus (A.) brevispina, "Tropidocyathus"

labidus, "Tropidocyathus" pileus, Truncatoflabellum phoenix.

Stn 9. — 23.10.1991, 5°19'21"S, 132°30'35"E, 361-389 m, Kai Isl.: Flabellum (U.) hoffmeisteri, Madrepora

oculata.

44

S. D. CAIRNS & H. ZIBROWIUS

Stn 10. —23.10.1991, 5°26'11"S, 132°27'37"E, 329-389 m, Kai Isl.: Caryophyllia (A.) karubarica, Flabellum

(U.) hoffmeisteri.

Stn 12. — 23.10.1991, 5°25'23"S, 132°36'59"E, 412-434 m, Kai Isl.: Caryophyllia (A.) karubarica, Flabellum

(F.j lamellulosum, Flabellum (F.) magnificum, Flabellum (U.) hoffmeisteri, Fungiacyathus (F.) stephanus.

Sm 13. — 24.10.1991, 5°26'27"S, 132°37'37"E, 393-417 m, Kai Isl.: Balanophyllia desmophyllioides,

Balanophyllia gigas, Caryophyllia (A.) karubarica, Deltocyathoides orientalis, Enallopsammia rostrata,

Flabellum (U.) hoffmeisteri, Lochmaeotrochus oculeus, Madrepora oculata, Polymyces wellsi.

Stn 15. — 24.10.1991, 5°17'38"S, 132°40'51"E, 214-221 m, Kai Isl.: Anthemiphyllia frustum, Caryophyllia (P.)

dentiformis, Conotrochus brunneus, Deltocyathoides orientalis, Flabellum (F.) politum, Fungiacyathus (F.)

?paliferus (irregular), Guynia annulata, Idiotrochus kikutii, Leptopenus sp„ Notocyathus conicus, Notocyathus

venustus, Peponocyathus folliculus, Peponocyathus minimus, Stephanophyllia complicata, Trochocyathus

philippinensis, "Tropidocyathus" labidus, Truncatoflabellum pusillum.

Stn 16. — 24.10.1991, 5°17’06"S, 132051'19"E, 315-348 m, Kai Isl.: Balanophyllia cornu, Balanophyllia

generatrix, Crispatotrochus rubescens, Madrepora oculata, Tethocyathus virgatus.

Stn 18. — 24.10.1991, 5°17'49"S, 133°00'51"E, 205-212 m, Kai Isl.: Balanophyllia cornu, Balanophyllia

desmophyllioides, Balanophyllia parvula, Bourneotrochus stellulatus, Caryophyllia (C.) lamellifera,

Caryophyllia (C.) rugosa, Conotrochus brunneus, Dactylotrochus cervicornis, Deltocyathoides orientalis,

Deltocyathus Stella, Deltocyathus suluensis, Dendrophytlia alcocki, Eguchipsammia gaditana, Fungiacyathus

(F.) paliferus, Idiotrochus kikutii, Letepsammia superstes, Madrepora arbuscula, Madrepora minutiseptum,

Notocyathus conicus, Peponocyathus minimus, Premocyathus dentiformis, Sympodangia albatrossi,

Thrypticotrochus multilobatus, Trochocyathus (T.) philippinensis, Truncatoflabellum angustum,

Truncatoflabellum pusillum.

Stn 19. — 25. 10. 1991, 5°15'52"S, 1 33°00'0 1"E, 576-604 m, Kai Isl.: Madrepora oculata.

Stn 20. — 25.10.1991, 5°16'30"S, 132°58'20"E, 768-810 m, Kai Isl.: Fungiacyathus (F.) stephanus,

Rhombopsammia niphada, Stephanocyathus ( O. ) weberianus.

Stn 21. — 25.10.1991, 5°16'25"S, 132°59'03"E, 688-694 m, Kai Isl.: Caryophyllia (C.) hawaiiensis,

Deltocyathus rotulus, Fungiacyathus (F.) stephanus, Rhombopsammia niphada.

Stn 22. — 25.10.1991, 5°16'23"S, 133°00'23"E, 85-124 m, Kai Isl.: Balanophyllia rediviva, Caryophyllia (C.)

hawaiiensis, Cyathelia axillaris, Deltocyathoides orientalis, Guynia annulata, Idiotrochus kikutii, Javania

insignis, Madracis sp. A, Notocyathus venustus, Peponocyathus minimus, Thalamophyllia tenuescens,

Truncatoflabellum phoenix.

Stn 25. — 26.10.1991, 5°31'30"S, 132°50'40"E, 318-352 m, Kai Isl.: Enallopsammia pusilla, Confluphyllia

juncta.

Stn 27. — 26.10.1991, 5°34'22"S, 132°51’29"E, 304-314 m, Kai Isl.: Tethocyathus virgatus, Trochocyathus (T.)

rhombocolumna.

Stn 28. — 26.10.1991, 5°31'27"S, 132°54'07"E, 448-468 m, Kai Isl.: Deltocyathoides orientalis, Deltocyathus

vaughani, Fungiacyathus (F.) ? paliferus (irregular), Guynia annulata.

Stn 29. — 26.10.1991, 5°35'49"S, 132°55’44"E, 181-184 m, Kai Isl.: Deltocyathoides orientalis, Guynia

annulata, Madracis sp. A.

Stn 30. — 26.10.1991, 5°37'34"S, 132°55'34"E, 111-116 m, Kai Isl.: Caryophyllia lamellifera, Cyathelia

axillaris, Dendrophytlia cf. ijimai, Flabellum (F.) pavoninum, Thalamophyllia tenuescens, Truncatoflabellum

mortenseni.

Stn 31. — 26.10.1991, 5°39'39"S, 132°50'4r’E, 288-289 in, Kai Isl.: Caryophyllia (C.) crosnieri, Conotrochus

brunneus, Deltocyathus magnificus, Deltocyathus suluensis, Flabellum (F.) patens, Flabellum (F.) politum,

Fungiacyathus (B.) variegatus, Notocyathus venustus, Truncatoflabellum angustum.

Stn 32. — 26.10.1991, 5°46'31"S, 132°50'42"E, 170-206 m. Kai Isl.: Balanophyllia desmophyllioides,

Caryophyllia (C.) crosnieri, Conotrochus brunneus, Deltocyathus Stella, Javania insignis, Madrepora arbuscula,

Notocyathus conicus, Trochocyathus (T.) caryophylloides, Trochocyathus (T.) philippinensis, "Tropidocyathus"

labidus.

Stn 35. — 27.10.1991, 6°07'22"S, 132°43'45"E, 390-502 m, Kai Isl.: Deltocyathus suluensis, Flabellum (U.)

marenzelleri, Flabellum (U.) hoffmeisteri, Fungiacyathus (F.) stephanus, Letepsammia formosissima,

Madrepora oculata, Stephanocyathus (A.) explanans, "Tropidocyathus" pileus.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

45

Stn 36. _ 27.10.1991, 6o05'50"S, 132°44'29''E, 210-268 m, Kai Isl.: Conotrochus brunneus, Deltocyathus

suluensis, Flabellum (F.) lamellulosum, Flabellum (F.) magnificum, Flabellum (U.) marenzelleri,

Fungiacyathus (F.) stephanus, Labyrinthocyathus sp. A, Letepsammia formosissima, "Tropidocyathus" pileus.

§tn 39. _ 28.10.1991, 7°45'43"S, 132°28'22"E, 466-477 m, Tanimbar Isl.: Caryophyllia (A.) karubarica,

Caryophyllia (A.) unicristata, Caryophyllia (C.) grandis, Deltocyathus vaughani, Flabellum (U.) hoffmeisteri,

Fungiacyathus (F.) stephanus, Madrepora oculata, Rhombopsammia niphada, Stephanocyathus (A.) explanans,

Truncatoflabellum paripavoninum.

Stn 40. _ 28.10.1991, 7°47'53"S, 132°28'19"E, 442-468 m, Tanimbar Isl.: Caryophyllia (A.) unicristata,

Caryophyllia (C.) ambrosia ambrosia, Caryophyllia (C.) grandis, Deltocyathus vaughani, Flabellum (U.)

hoffmeisteri, Fungiacyathus (B.) granulosus, Rhombopsammia niphada, Stephanocyathus (A.) explanans.

Stn 42. — 28.10.1991, 7°49’48"S, 132°43'29"E, 350-353 m, Tanimbar Isl.: Madrepora oculata.

Stn 44. — 29.10.1991, 7°52’27"S, 132°48'24"E, 291-295 m, Tanimbar Isl.: Conocyathus zelandiae,

Deltocyathoides orientalis, Javania sp., Peponocyathus minimus, Tethocyathus virgatus, Truncatoflabellum

angustum, Truncatoflabellum phoenix, Truncatoflabellum pusillum.

Stn 49. — 29.10.1991, 7°59'51"S, 132°58'50"E, 206-209 m, Tanimbar Isl.: Balanophyllia gemma, Balanophyllia

desmophyllioides, Balanophyllia crassiseptum, Balanophyllia parvula, Caryophyllia (A.) dentata,

Deltocyathoides orientalis, Fungiacyathus (F.) ?paliferus (irregular), Idiotrochus kikutii, Javania sp.,

Notocyathus venustus, Peponocyathus minimus, Trochocyathus (T.) caryophylloides, Trochocyathus (T.)

rhombocolumna.

Stn 50. — 29.10.1991, 7°59'09"S, 133°01'56"E, 184-185 m, Tanimbar Isl.: Balanophyllia gemma, Balanophyllia

crassiseptum, Letepsammia superstes, Notocyathus venustus, Trochocyathus (T.) caryophylloides.

Stn 56. — 31.10.1991, 8°12'39"S, 132°01T5"E, 549-552 m, Tanimbar Isl.: Caryophyllia (C.) ambrosia

ambrosia, Deltocyathus rotulus, Deltocyathus vaughani, Flabellum (U.) messum, Fungiacyathus (B.)

granulosus, Fungiacyathus (F.) stephanus, Madrepora oculata, Truncatoflabellum paripavoninum.

Stn 57. — 31.10.1991, 8°15'39"S, 131°56'38"E, 603-622 m, Tanimbar Isl.: Caryophyllia (C.) ambrosia

ambrosia, Flabellum (U.) messum, Truncatoflabellum paripavoninum.

Stn 58. — 31.10.1991, 8°2r47"S, 132°00'55"E, 457-461 m, Tanimbar Isl.: Caryophyllia (A.) karubarica,

Caryophyllia (A.) unicristata, Fungiacyathus (B.) granulosus, Stephanocyathus (A.) explanans.

Stn 59. — 31.10.1991, 8°20'01"S, 132°09'32"E, 399-405 m, Tanimbar Isl.: Caryophyllia (A.) unicristata,

Caryophyllia (C.) grandis, Conotrochus funicolumna, Deltocyathus magnificus, Deltocyathus vaughani,

Flabellum (F.) lamellulosum, Flabellum (F.) magnificum, Flabellum (U.) deludens, Flabellum (U.)

hoffmeisteri, Fungiacyathus (B.) granulosus, Fungiacyathus (F.) stephanus, Madrepora oculata,

Paraconotrochus zeidleri, Rhombopsammia niphada, Stephanocyathus (A.) explanans.

Stn 61. — 1.11.1991, 9°05'09"S, 132°44'35"E, 235 m, Tanimbar Isl.: Balanophyllia cornu, Deltocyathoides

orientalis, Peponocyathus minimus, Stephanocyathus (A.) spiniger, Trochocyathus (T.) caryophylloides.

Stn 62. — 1.11.1991, 9o02T0"S, 132°43'05"E, 245-251 m, Tanimbar Isl.: Caryophyllia (A.) unicristata,

Caryophyllia (A.) spinigera, Cary’ophyllia (C.) grandis, Endopachys bulbosa, Flabellum (U.) deludens,

Fungiacyathus (F.) stephanus, "Tropidocyathus" pileus.

Stn 63. — 1.1 1.1991, 8°59’59"S, 132°56'40"E, 213-214 m, Tanimbar Isl.: Flabellum (U.) deludens.

Stn 65. — 1.1 1.1991, 9°14'0r'S, 132°28’28"E, 174-176 m, Tanimbar Isl.: Caryophyllia (A.) dentata, Flabellum

(F.) politum, Flabellum (U.) deludens, Placotrochus laevis, Truncatoflabellum spheniscus.

Stn 67. — 1.11.1991, 8°58'59"S, 132°07'20"E, 233-246 m, Tanimbar Isl.: Caryophyllia (C.) transversalis,

Crispatotrochus rubescens, Deltocyathus suluensis, Endopachys bulbosa, Flabellum (F.) lamellulosum,

Flabellum (F.) magnificum, Letepsammia formosissima, "Tropidocyathus" pileus.

Stn 68. — 1.11.1991, 8°55’09"S, 132°03T3"E, 280-296 m, Tanimbar Isl.: Caryophylita (C.) transversalis.

Stn 69. — 2.11.1991, 8°45T7"S, 13I°5T35"E, 356-367 m, Tanimbar Isl.: Caryophyllia (A.) unicristata,

Conotrochus brunneus, Deltocyathus magnificus, Flabellum (F.) lamellulosum, Flabellum (F.) magnificum,

Flabellum (U.) hoffmeisteri, Madrepora oculata, Paraconotrochus zeidleri.

Stn 70. — 2.11.1991, 8°39T4"S, 131°49T6"E, 410-411 m, Tanimbar Isl.: Cary’ophyllia (A.) unicristata,

Caryophyllia (C.) grandis, Deltocyathus magnificus, Flabellum (F.) lamellulosum, Flabellum (F.)

magnificum, Flabellum (U.) hoffmeisteri, Rhombopsammia niphada, Stephanocyathus (A.) explanans,

Truncatoflabellum paripavoninum.

46

S. D. CAIRNS & H. Z1BR0WIUS

Stn 71. — 2.11.1991, 8°39’39"S, 131°42’29"E, 477-480 m, Tanimbar Isl.: Caryophyllia (A.) karubarica,

Caryophyllia (A.) unicristata, Caryophyllia (C.) grandis, Conotrochus brunneus, Deltocyathus magnificus,

Deltocyathus vaughani, Flabellum (U.) hoffmeisteri, Rhombopsammia niphada, Stephanocyathus explanans,

Truncatoflabellum paripavoninum.

Sm 72. — 2.1 1.1991, 8°33'19"S, 131°35'10"E, 676-699 m, Tanimbar Isl.: Caryophyllia (C.) ambrosia ambrosia,

Flabellum (U.) messum, Stephanocyathus (A.) explanans.

Sm 75. — 3.1 1.1991, 8°46'52"S, 131°33'37"E, 451 m, Tanimbar Isl.: Caryophyllia (C.) grandis, Flabellum (U.)

hoffmeisteri, Rhombopsammia niphada, Stephanocyathus (A.) explanans.

Sm 76. — 3.1 1.1991, 8°49’08"S, 131°35'36"E, 400 m, Tanimbar Isl.: Caryophyllia (A.) unicristata, Deltocyathus

magnificus, Deltocyathus vaughani, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus.

Stn 77. — 3.11.1991, 8°55'38"S, 131°29'12"E, 347-351 m, Tanimbar Isl.: Conotrochus brunneus, Deltocyathus

suluensis, Flabellum (F.) lamellulosum, Flabellum (F.) magnificum. Flabellum ( U .) hoffmeisteri, Madrepora

oculata, Paraconotrochus zeidleri.

Stn 79. — 3.11.1991, 9°13'34"S, 131°22'35"E, 239-250 m, Tanimbar Isl.: Caryophyllia (A.) spinigera,

Caryophyllia (C.) transversalis, Endopachys bulbosa, Flabellum (F.) lamellulosum, Flabellum (U.) deludens.

Stn 82. — 4.1 1.1991, 9°30’00"S, 131°02'41"E, 215-218 m, Tanimbar Isl.: Balanophyllia generatrix.

Stn 84. — 4.1 1.1991, 9°22'41"S, 131°07'17"E, 246-275 m, Tanimbar Isl.: Caryophyllia (C.) transversalis.

Sm 85. — 4.11.1991, 9°22'51"S, 131°12'04"E, 239-244 m, Tanimbar Isl.: Flabellum (F.) magnificum,

Letepsammia formosissima, "Tropidocyathus" pileus.

Stn 86. — 4.1 1.1991, 9°23'59"S, 131°14'29"E, 222-226 m, Tanimbar Isl.: Cladopsammia echinata, Conocyathus

zelandiae, Crispatotrochus rubescens, Dendrophyllia arbuscula, Gardineria philippinensis, Javania sp.,

Letepsammia formosissima, Rhizosmilia elata, Stephanocyathus (A.) spiniger, Trochocyathus (T.)

cary’ophylloides, Trochocyathus (T.) rhombocolumna.

Stn 87. — 5.11.1991, 8°48’13"S, 130o46'37"E, 1016-1024 m, Tanimbar Isl.: Caryophyllia (C.) ambrosia

ambrosia, Deltocyathus rotulus, Rhombopsammia squiresi, Stephanocyathus (A.) explanans,

Truncatoflabellum paripavoninum.

Stn 89. — 5.11.1991, 8039'41"S, 1 3 1 °05'25,,E, 1048-1084 m, Tanimbar Isl.: Caryophyllia (C.) ambrosia

ambrosia, Deltocyathus rotulus, Rhombopsammia squiresi, Stephanocyathus (O.) weberianus.

Stn 91. — 5.1 1.1991, 8°44'54"S, 131°03'10"E, 884-890 m, Tanimbar Isl.: Caryophyllia ( C.) ambrosia ambrosia,

Caryophyllia (C.) scobinosa, Deltocyathus rotulus, Flabellum (U.) messum, Stephanocyathus (O.) weberianus,

Truncatoflabellum paripavoninum.

MORTENSEN'S JAVA (-SOUTH AFRICA) EXPEDITION

Stn 1. — 3.4.1929, 7°34'S, 1 14°18'E, 100 m, N Bali: Caryophyllia (A.) grayi.

Stn 2. — 3.4.1929, 7°33'S, 1 14°36'E, 200 m, N Bali: Caryophyllia (A.) spinigera.

Stn 5. — 5.4.1929, 8°23'S, 114°29’E, 70 m, Bali Strait: Asterosmilia marchadi, Balanophyllia carinata,

Balanophyllia stimpsonii, Caryophyllia (A.) grayi, Endopachys grayi, Stephanophyllia neglecta,

Tropidocyathus lessonii, Trochocyathus (T.) burchae, Truncatoflabellum mortenseni.

Stn 6. — 5.4.1929, 8°26'S, 114°29'E, 70 m, Bali Strait: Asterosmilia marchadi, Balanophyllia carinata,

Caryophyllia (A.) grayi, Endopachys grayi, Stephanophyllia neglecta, Truncatoflabellum mortenseni.

Stn 8. — 6.4.1929, 8°23'S, 114°24'E, 50 m, Bali Strait: Asterosmilia marchadi, Balanophyllia carinata,

Balanophyllia stimpsonii, Caryophyllia (A.) grayi, Endopachys grayi, Stephanophyllia neglecta,

Tropidocyathus lessonii, Truncatoflabellum mortenseni.

Stn 9. — 6.4.1929, 8°30-35’S, 114°28'E, 70-150 m, Bali Strait: Asterosmilia marchadi, Caryophyllia (A.) grayi,

Endocyathopora laticostata, Trochocyathus (T.) burchae, Trochocyathus (T.) apertus, Truncatoflabellum

mortenseni.

Stn 15. — 10.4.1929, 7°29'S, 114°49'E, 240 m, N Bali: Balanophyllia cornu, Caryophyllia (C.) secta,

Deltocyathus suluensis, Lochmaeotrochus oculeus, Tethocyathus virgatus.

Stn 18. — 1 1.4.1929, 7°15'S, 1 14°45'E, 100 m, N Bali: Notocyathus venustus, Truncatoflabellum mortenseni.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACT1N1A

47

MORTENSEN'S PACIFIC EXPEDITION

Stn unnumbered. — 14.3.1914, W Mindanao, Zamboanga Peninsula, 301-373 m: Caryophyllia (C.) secta.

Stn unnumbered. — 19.3.1914, Sulu Archipelago, Jolo, 46 m: Leptopsammia stokesiana.

MUSORSTOM 1, "VAUBAN"

Stn 2. — 19.3.1976, 14°02.8’N, 120°18.8'E, 182-187 m, SW Luzon: Letepsammia formosissima,

Stephanocyathus (A.) spiniger.

Stn 3. — 19.3.1976, 14°01.7'N, 120°16.0'E, 183-185 m, SW Luzon: Balanophyllia cornu, Bcilcmophyllia

desmophyllioides, Balanophyllia rediviva.

Stn 4. — 19.3.1976, 14°01.8’N, 120°17.2'E, 182-194 m, SW Luzon: Letepsammia formosissima.

Stn 5. — 19.3.1976, 14°01.5'N, 120°23.5'E, 200-215 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum (U.)

deludens.

Sm 9. — 19.3.1976, 14°01.8'N, 120°17.6'E, 180-194 m, SW Luzon: Flabellum (F.) lamellulosum, Flabellum

(U.) deludens, Letepsammia formosissima.

Stn 10. — 19.3.1976, 13°59.8'N, 120°18.2'E, 187-205 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(U.) deludens, Letepsammia formosissima.

Stn 11. — 20.3.1976, 13°59.8'N, 120°23.7'E, 217-230 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (U.) deludens.

Stn 12. — 20.3.1976, 14°00.8'N, 120°20.5'E, 187-210 m, SW Luzon: Flabellum (F.) lamellulosum, Flabellum

(U.) deludens, Letepsammia formosissima.

Stn 13. — 20.3.1976, 14°00.5'N, 120°17.0'E, 190 m, SW Luzon: Flabellum (F.) lamellulosum, Flabellum (U.)

deludens, Fungiacyathus (B.) variegatus, Trochocyathus (T.) philippinensis, "Tropidocyathus" pileus.

Stn 14. — 20.3.1976, 14°00.2'N, 120°17.2'E, !90'm, SW Luzon: Balanophyllia cornu, Fungiacyathus (B.)

variegatus, Letepsammia formosissima, Trochocyathus (T.) philippinensis.

Stn 15. — 20.3.1976, 14°00.3'N, 120°18.0'E, 188-192 m, SW Luzon: Flabellum (U.) deludens, Fungiacyathus

(B.) variegatus.

Stn 20. — 21.3.1976, 13°59.2'N, 120°20.3'E, 208-222 m, SW Luzon: Cary’ophyllia (A.) spinicarens,

Cary’ophyllia (A.) spinigera, Flabellum (U.) deludens, Letepsammia formosissima.

Stn 24. — 22.3.1976, 14°00.0'N, 120°18.0'E, 189-209 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (U.) deludens, Letepsammia formosissima.

Stn 25. — 22.3.1976, 14°02.7'N, 120°20.3'E, 191-200 m, SW Luzon: Cary’ophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Letepsammia formosissima,

Stephanocyathus (A.) spiniger.

Stn 27. — 22.3.1976, 13°59.8'N, 120°18.6'E, 188-192 m, SW Luzon: Balanophyllia serrata, Flabellum (F.)

lamellulosum.

Stn 31. — 22.2.1976, 14°00.0'W, 120°16.0'E, 187-195 m, SW Luzon: Flabellum (F.) lamellulosum.

Stn 32. — 23.2.1976, 14°02.2'W, 120°17.7'E, 184-193m, SW Luzon: Asterosmilia marchadi, Balanophyllia

crassisepta, Flabellum (F.) lamellulosum, Tethocyathus virgatus.

Stn 35. — 23.3.1976, 13°59.0'N, 120°18.5'E, 186-187 m, SW Luzon: Letepsammia formosissima,

Trochocyathus (T.) philippinensis.

Stn 40. — 24.3.1976, 13°57.4'N, 120°27.8'E, 265-287 m, SW Luzon: Cary’ophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (U.) deludens, Flabellum (U.) marenzelleri, "Tropidocyathus" pileus.

Stn 42. —24.3.1976, I3°55.1'N, 120°28.6’E, 379-407 m, SW Luzon: "Tropidocyathus" pileus.

Stn 43. — 24.3.1976, 13°50.5’N, 120°28.0'E, 448-484 m, SW Luzon: Fungiacyathus (B.) granulosus.

Stn 44. — 24.3.1976, 13°46.9'N, 120°29.5'E, 592-610 m, SW Luzon: Caryophyllia (C.) scobinosa, Fungia¬

cyathus (B.) granulosus, Truncatoflabellum paripavoninum.

Stn 45. — 24.3.1976, 13°46.0'N, 120°23.8'E, 100-180 m, SW Luzon: Caryophyllia (A.) grayi.

Stn 47. — 25.3.1976, 13°40.7'N, 120°30.0’E, 685-757 m, SW Luzon: Caryophyllia (C.) scobinosa, Flabellum

(U.) sexcostatum, Rhombopsammia niphada, Truncatoflabellum paripavoninum.

48

S. D. CAIRNS & H. Z1BR0WIUS

Stn 49. — 25.3.1976, 13°49.1'N, 119°59.8'E, 750-925 m, SW Luzon: Caryophyllia (C.) ambrosia ambrosia,

Caryophyllia (C.) diomedeae, Fungiacyathus (F.) stephanus, Madrepora oculata, Stephanocyathus (O.)

weberianus, Stephanocyathus (S.) regius.

Stn 50. — 25.3.1976, 13°49.2'N, 120°01.8'E, 415-510 m, SW Luzon: Fungiacyathus (B.) granulosus,

Trochocyathus (A.) longispina.

Stn 54. — 26.3.1976, 13°54.2'N, 119°57.9'E, 975-1125 m, SW Luzon: Caryophyllia (C.) ambrosia ambrosia,

Deltocyathus rotulus, Fungiacyathus (F.) stephanus, Stephanocyathus (O.) weberianus, Stephanocyathus (S.)

regius.

Stn 55. — 26.3.1976, 13°55.0'N, 120°12.5’E, 194-200 m, SW Luzon: Flabellum (F.) lamellulosum,

"Tropidocyathus" pileus.

Stn 56. — 26.3.1976, 13°53.1'N, 120°08.9'E, 129-134 m, SW Luzon: Caryophyllia (A.) grayi, Truncatoflabellum

candeanum.

Stn 57. — 26.3.1976, 13°53.1'N, 120°13.2'E, 96-107m, SW Luzon: Asterosmilia marchadi, Balanophyllia

rediviva, Endopachys grayi, Trochocyathus (T.) cooperi.

Stn 58. — 26.3.1976, 13°58.0'N, 120°13.7'E, 143-178 m, SW Luzon: "Tropidocyathus" pileus.

Stn 61. — 27.3.1976, 14°02.2'N, 120°18.1'E, 184-202 m, SW Luzon: Balanophyllia cornu, Balanophyllia sp.,

Caryophyllia (A.) spinigera, Flabellum (F.) lamellulosum, Flabellum (U.) deludens, Letepsammia

formosissima, Stephanocyathus (A.) spiniger.

Stn 62. — 27.3.1976, 13°59.5'N, 120°15.6'E, 179-194 m, SW Luzon: Javania lamprotichum, "Tropidocyathus"

pileus, Truncatoflabellum candeanum.

Stn 63. — 27.3.1976, 14°00.8'N, 120°15.8'E, 191-195 m, SW Luzon: Balanophyllia cornu, Balanophyllia

desmophyllioides, Balanophyllia serrata, Gardineria philippinensis, Javania lamprotichum, Stephanocyathus

(A.) spiniger, Tethocyathus virgatus, Trochocyathus (T.) caryophylloides.

Stn 64. — 27.3.1976, 14°00.5'N, 120°16.3’E, 194-195 m, SW Luzon: Balanophyllia desmophyllioides,

Caryophyllia (C.) octonaria, Letepsammia formosissima, Trochocyathus (T.) philippinensis, Tropidocyathus

lessonii, Truncatoflabellum angustum, Truncatoflabellum candeanum.

Stn 65. — 27°3. 1976, 14°00.0'N, 120°19.2'E, 194-202 m, SW Luzon: Balanophyllia desmophyllioides,

Balanophyllia serrata, Javania lamprotichum, Rhizosmilia robusta, Stephanocyathus (A.) spiniger.

Stn 68. — 27.3.1976, 14°00.8'N, 120°17.4'E, 183-199 m, SW Luzon: Flabellum (F.) lamellulosum, Flabellum

(U.) deludens.

Stn 69. — 27.3.1976, 13°58.8'N, 120°17.3'E, 187-199 m, SW Luzon: Balanophyllia serrata.

Stn 71. — 28.3.1976, 14°09.3'N, 120°26.2'E, 174-204m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum (F.)

lamellulosum.

Stn 72. — 28.3.1976, 14°1 1.8'N, 120°28.7'E, 122-127 m, SW Luzon: Caryophyllia (A.) grayi, Caryophyllia (A.)

spinigera, Trochocyathus (T.) philippinensis, Truncatoflabellum candeanum.

Stn 73. — 28.3.1976, 14°15.0’N, 120°31.2'E, 70-76 m, SW Luzon: Asterosmilia marchadi.

Musorstom 2, "Coriolis"

Stn 1. — 20.11.1980, 14°00.3'N, 120°19.3'E, 188-198 m, SW Luzon: Balanophyllia gigas, Caryophyllia (A.)

spinigera, Flabellum (F.) lamellulosum, Flabellum ( U.) deludens, Letepsammia formosissima.

Stn 2. — 20.11.1980, 14°01.0'N, 120°17.1'E, 184-186 m, SW Luzon: Alatotrochus rubescens,

Balanophyllia gigas, Caryophyllia (C.) octonaria, Flabellum (F.) politum, Letepsammia formosissima,

Tethocyathus virgatus, Trochocyathus (T.) caryophylloides, Truncatoflabellum candeanum, Truncatoflabellum

formosum.

Stn 4. — 20.11.1980, 14°01.2'N, 120°18.4'E, 183-190 m, SW Luzon: Caryophyllia (A.) spinigera,

Fungiacyathus (B.) variegatus, Letepsammia formosissima.

Stn 6. — 20.11.1980, 13°56.5'N, 120°20.7’E, 136-152 m, SW Luzon: Flabellum (F.) politum, Letepsammia

formosissima, Trochocyathus (T.) philippinensis, "Tropidocyathus" pileus, Truncatoflabellum candeanum.

Stn 8. — 21.1 1.1980, 13°55.0'N, 120°20.0'E, 85-90 m, SW Luzon: Asterosmilia marchadi.

Stn 9. — 21.11.1980, 13°53.4'N, 120°20.7'E, 66 m, SW Luzon: Balanophyllia stimpsonii, Paracyathus

rotundatus.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

49

Stn 10. — 21.11.1980, 14°00.1'N, 120°18.5'E, 188-195 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (F.) politum, Flabellum (U.) deludens, Trochocyathus (T.) philippinensis,

"Tropidocyathus" pileus, Truncatoflabellum candeanum.

Stn 11. — 21.11.1980, 14°00.4'N, 120°19.7'E, 194-196 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(U.) deludens, Letepsammia formosissima, Notocyathus conicus C89.

Stn 12. — 21.1 1.1980, 14°01.0'N, 120°19.7'E, 197-210 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Letepsammia formosissima,

Stephanocyathus (A.) spiniger.

Stn 13. — 21.11.1980, 14°00.5'N, 120°20.7'E, 193-200 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(U.) deludens, Letepsammia formosissima.

Stn 15. — 21.11.1980, 13°55.1'N, 120°28.4'E, 326-330 m, SW Luzon: Balanophyllia cornu, Caryophyllia (C.)

crosnieri, Caryophyllia (C.) diomedeae, Crispatotrochus rubescens, Flabellum (F.) magnificum, Flabellum (U.)

deludens, Letepsammia formosissima.

Stn 17. — 22.11.1980, 14°00.0'N, 120°17.1'E, 174-193 m, SW Luzon: Balanophyllia desmophyllioides,

Trochocyathus (T.) philippinensis.

Stn 18. — 22.11.1980, 14°00.0'N, 120°18.6'E, 188-195 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Letepsammia formosissima, Stephanocyathus (A.) spiniger.

Stn 19. — 22.11.1980, 14°00.5'N, 120°16.5'E, 189-192 m, SW Luzon: Flabellum (F.) lamellulosum.

Stn 20. — 22.11.1980, 14°00.9'N, 120°18.1'E, 185- 192m, SW Luzon: Caryophyllia ( A .) spinigera, Flabellum

(U.) deludens.

Stn 21. — 22.11.1980, 14°00.2'N, 120°17.8'E, 191-192 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum ( U.) deludens, Stephanocyathus (A.) spiniger.

Stn 24. — 23.1 1.1980, 13°37.2'N, 120°42.3'E, 640-647 m, SW Luzon: Fungiacyathus (F.) stephanus.

Stn 25. — 23.11.1980, 13°39.0'N, 120°42.6'E, 520-550 m, SW Luzon: Caryophyllia (C.) scobinosa,

Fungiacyathus (B.) granulosus, Fungiacyathus (F.) stephanus, Placotrochides scaphula C89, Rhombopsammia

niphada, Truncatoflabellum paripavoninum.

Stn 26. — 23.11.1980, 13°49.6'N, 120°51.0'E, 299-300 m, SW Luzon: Flabellum (F.) lamellulosum,

Stephanocyathus (A.) spiniger.

Stn 27. — 23.1 1.1980, 13°41.5’N, 120°50.1'E, 95-100 m, SW Luzon: Balanophyllia gigas.

Stn 29. — 23.11.1980, 13°42. l’N, 120°50. l’E, 119-204 m, SW Luzon: Caryophyllia (A.) grayi, Endopachys

grayi, Tropidocyathus lessonii.

Stn 32. — 24.11.1980, 13°40.5'N, 120°53.9'E, 192-220 m, SW Luzon: Balanophyllia cornu, Balanophyllia

parvula, Balanophyllia sp., Caryophyllia (C.) secta, Conotrochus brunneus, Crispatotrochus rugosus,

Gardineria musorstomica C89, Idiotrochus kikutii C89, Javania insignis, Letepsammia superstes,

Trochocyathus (T.) cary’ophylloides, Truncatoflabellum formosum.

Stn 33. — 24.11.1980, 13°32.3'N, 121°07.5'E, 130-137 m, S Luzon: Anthemiphyllia dentata, Balanophyllia

desmophyllioides, Balanophyllia gemma, Balanophyllia rediviva, Balanophyllia sp., Caryophyllia (C.)

hawaiiensis, Caryophyllia (C.) rugosa, Conotrochus brunneus, Deltocyathoides orientalis C89* (ex

P. australiensis), Deltocyathus Stella, Dendrophyllia cf. ijimai, Dendrophylliidae (colonial), Guynia annulata

C89, Idiotrochus kikutii C89, Javania insignis, Letepsammia formosissima, Notocyathus conicus C89,

Stephanophyllia neglecta, Tethocyathus virgatus, Trochocyathus (T. ) philippinensis, Tropidocyathus lessonii

C89, Thrypticotrochus multilobatus C89, Truncatoflabellum mortenseni, Truncatoflabellum pusillum.

Stn 34. — 24.1 1.1980, 13°27.9’N, 121°12.0'E, 155-167 m, N Mindoro: Balanophyllia sp.

Stn 36. — 24.11.1980, 13°31.4'N, 121°23.9'E, 569-595 m, S Luzon: Flabellum (U.) japonicum, Madrepora

oculata.

Stn 39. — 25.1 1.1980, 13°02.8'N, 122°37.1'E, 1030-1 190 m, S Luzon: Madrepora oculata.

Stn 40. — 25.11.1980, 13°07.7'N, 122°39.1'E, 280-440 m, S Luzon: Flabellum (U.) deludens, Flabellum (U.)

japonicum.

Stn 44. — 26.1 1.1980, 13°23.2'N, 122°20.7'E, 760-820 m, S Luzon: Flabellum (U.) japonicum, Trochocyathus

(A.) longispina.

Stn 45. 26.1 1.1980, 13°26.8'N, 122°18.5'E, 447-500 m, S Luzon: Caryophyllia (A.) spinicarens.

Stn 46. — 26.11.1980, 13°25.7'N, 122°17.0'E, 445-520 m, S Luzon: Flabellum (U.) deludens, Caryophyllia (A.)

spinicarens.

50

S. D. CAIRNS & H. ZIBROWIUS

Stn 47. — 26.11.1980. 13°33.0’N, 122°10.1'E, 81-84 m, S Luzon: Asterosmilia marchadi, Caryophyllia (A.)

grayi, Phyllangia papuensis, Trochocyathus (T.) cooperi.

Stn 49. — 26.11.1980, 13°38.4'N, 121°44. l'E, 416-425 m, S Luzon: Flabellum (U.) japonicum , Mcidrepora

oculala.

Stn 53. — 27.1 1.1980, 13°59.2'N, 120°18.3'E, 215-216m, SW Luzon: Javania lamprotichum.

Stn 62. — 29.11.1980, 14°00.4'N, 120°17.0'E, 186-189 m, SW Luzon: Caryophyllia (A.) spinigera,

Letepsammia formosissima, Stephanocyalhus (A.) spiniger.

Stn 63. — 29.11.1980, 14°07.3'N, 120°15.0'E, 215-230 m, SW Luzon: Caryophyllia (A.) spinicarens,

Caryophyllia (A.) spinigera, Conotrochus brunneus, Deltocyalhus andamanicus, Flabellum (F.) lamellulosum,

Flabellum (U.) deludens, Letepsammia formosissima, "Tropidocyathus" pileus, Truncatoflabellum angustum.

Stn 64. — 29.1 1.1980, 14°01.5'N, 120°18.9'E, 191-195 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Fungiacyathus (B.) variegatus, Flabellum (U.) deludens, Letepsammia formosissima.

Sm 66. — 29.11.1980, 14°00.6'N, 120°20.3'E, 192-209 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (U.) deludens C89, Fungiacyathus (B.) variegatus, Letepsammia formosissima.

Stn 68. — 29.1 1.1980, 14°01.9'N, 120°18.8'E, 195-199 m, SW Luzon: Balanophyllia cornu, Caryophyllia (A.)

spinigera, Flabellum (F.) lamellulosum, Flabellum (F.) politum, Flabellum (U.) deludens, Letepsammia

formosissima, Truncatoflabellum candeanum.

Stn 75. — 1.12.1980, 13°50.5'N, 120°30.3'E, 300-330 m, SW Luzon: Flabellum (F.) lamellulosum, Flabellum

(F.) magnificum.

Stn 77. — 1.12.1980, 13°48.8'N, 120°30.3'E, 529-552 m, SW Luzon: Truncatoflabellum paripavoninum.

Stn 78. — 1.12.1980, 13°49.1'N, 120°28.0'E, 441-550 m, SW Luzon: Flabellum (F.) magnificum, Flabellum

(U.) sp.

Stn 82. — 2.12.1980, 13°46.1’N, 120°28.4'E, 550 m, SW Luzon: Fungiacyathus (B.) granulosus.

Stn 83. — 2.12.1980, 13°55.2'N. 120°30.5’E, 318-320 m, SW Luzon: Caryophyllia (A.) spinicarens,

Deltocyathus magnificus, Flabellum (F.) lamellulosum, Flabellum (U.) deludens, "Tropidocyathus" pileus.

Musorstom 3, "Coriolis"

Stn 86. — 31.5.1985, 14°00.4'N, 120°17.8'E, 187-192 m, SW Luzon: Alatotrochus rubescens, Caryophyllia (A.)

spinigera, Flabellum (F.) lamellulosum, Flabellum (F.) politum, Letepsammia formosissima.

Stn 87. — 31.5.1985, 14°00.6'N, 120°19.6'E, 191-197 m, SW Luzon: Aulocyathus ?juvenescens, Caryophyllia

(A.) spinigera, Flabellum (F.) lamellulosum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Guynia

annulata, Letepsammia formosissima, Truncatoflabellum pusillum.

Stn 88. — 31.5.1985, 14°00.5'N, 120°17.4'E, 183-187 m, SW Luzon: Alatotrochus rubescens, Balanophyllia

cornu, Caryophyllia (C.) octonaria, Caryophyllia (A.) spinigera, Deltocyathus andamanicus, Flabellum (F.)

lamellulosum, Flabellum (F.) politum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Javania

lamprotichum, Letepsammia formosissima, Rhizosmilia sagamiensis, Trochocyathus (T.) caryophylloides,

Trochocyathus (T.) philippinensis, Tropidocyathus lessonii C89, "Tropidocyathus" pileus, Truncatoflabellum

candeanum, Stephanocyathus (A.) spiniger.

Stn 89. — 31.5.1985, 14°01.0'N, 120°17.1'E, 187-191 m, SW Luzon: Madrepora oculata.

Stn 90. — 31.5.1985, 14°00.1'N, 120°18.6'E, 195 m, SW Luzon. Caryophyllia (C.) octonaria, Caryophyllia (A.)

spinigera, Letepsammia formosissima, Truncatoflabellum candeanum.

Stn 91. — 31.5.1985, 14°00.1’N, 120°17.8'E, 190-203 m, SW Luzon: Alatotrochus rubescens, Caryophyllia (A.)

spinigera, Flabellum (F.) lamellulosum, Flabellum (U.) deludens, Letepsammia formosissima,

Stephanophyllia neglecta C89, Truncatoflabellum candeanum.

Stn 92. — 31.5.1985, 14°03.0'N, 120°11.5'E, 224 m, SW Luzon: Caryophyllia (A.) spinicarens, Caryophyllia

(A.) spinigera, Conotrochus brunneus, Deltocyathus andamanicus, Flabellum (F.) lamellulosum, Flabellum

(U.) deludens, Letepsammia formosissima, "Tropidocyathus" pileus, Truncatoflabellum angustum,

Truncatoflabellum candeanum.

Stn 93. — 1.6.1985, 13°48.6’N, 120°02.4’E, 540 m, SW Luzon: Deltocyathus suluensis.

Stn 94. — 1.6.1985, 13°47.4'N, 120°03.4'E, 842 m, SW Luzon: Caryophyllia (C.) ambrosia ambrosia,

Deltocyathus rotulus, Javania lamprotichum, Polymyces wellsi.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

51

Stn 95. — 1.6.1985, 13°55.8'N, 119°59.3'E, 865 m, SW Luzon: Caryophyllia (C.) diomedeae, Flabellum (U.)

japonicum, Fungiacyathus (B.) variegatus.

Stn 96. — 1.6.1985, 14°00.3'N, 120°17.3'E, 190-194 m, SW Luzon: Alatotrochus rubescens, Balanophyllia

cornu, Caryophyllia (A.) spinigera, Flabellum (F.) lamellulosum, Flabellum (F.) politum, Flabellum (U.)

deludens, Fungiacyathus (B.) variegatus, Letepsammia formosissima C89, Stephanocyathus (A.) spiniger,

Trochocyathus (T.) philippinensis, "Tropidocyathus" pileus, Truncatoflabellum candeanum, Truncatoflabellum

pusillum.

Stn 97. — 1.6.1985, 14°00.7'N, 120°18.8'E, 189-194 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum (F.)

lamellulosum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Letepsammia formosissima,

Stephanocyathus (A.) spiniger.

Stn 98. — 1.6.1985, 14°00.2'N, 120°17.9’E, 194-205 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum (F.)

lamellulosum, Flabellum (F.) politum, Flabellum (U.j deludens, Letepsammia formosissima, Stephanocyathus

(A.) spiniger, Trochocyathus (T.) philippinensis.

Stn 99. — 1.6.1985, 14°01.0'N, 120°19.5'E, 196-204 m, SW Luzon: Caryophyllia (A.) grayi, Caryophyllia (A.)

spinigera, Flabellum (F.) lamellulosum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Letepsammia

formosissima, Stephanocyathus (A.) spiniger, "Tropidocyathus" pileus, Truncatoflabellum candeanum.

Stn 100. — 1.6.1985, 14°00.0'N, lt)°17.6’ E, 189-199 m, SW Luzon: Balanophyllia cornu, Caryophyllia (C.)

octonaria, Caryophyllia (A.) spinigera, Flabellum (F.) lamellulosum, Flabellum (F.) politum, Flabellum (U.)

deludens, Fungiacyathus (B.) variegatus, Letepsammia formosissima, Thrypticotrochus multilobatus.

Stn 101. — 1.6.1985, 14°00.1'N, 120°19.2'E, 194-196 m, SW Luzon: Caryophyllia (C.) octonaria, Caryophyllia

(A.) spinigera, Flabellum (F.) lamellulosum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Guynia

annulata, Letepsammia formosissima.

Stn 102. — 1.6.1985, 14°00.8'N, 120°17.8'E, 192 m, SW Luzon: Alatotrochus rubescens, Asterosmilia marchadi,

Balanophyllia cornu, Caryophyllia (C.) octonaria, Caryophyllia (A.) spinigera, Conotrochus brunneus,

Deltocyathoides orientalis, Endopachys grayi, Flabellum (F.) lamellulosum, Flabellum (F.) politum,

Flabellum (U.) deludens, Fungiacyathus (F.) paliferus, Fungiacyathus (B.) variegatus, Guynia annulata,

Idiot rochus kikutii, Letepsammia formosissima, Notocyathus conicus C89, Stephanocyathus (A.) spiniger,

Stephanophyllia neglecta C89, Trochocyathus (T.) philippinensis, Truncatoflabellum candeanum,

Truncatoflabellum pusillum.

Stn 103. — 1.6.1985, 14°00.4'N, 120°18.1'E, 193-200 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(U.) deludens, Letepsammia formosissima, Stephanocyathus (A.) spiniger.

Stn 105. — 1.6.1985, 13°52.6'N, 120°29.6'E, 398-417 m, SW Luzon: Balanophyllia cornu, Madrepora oculata.

Stn 106. — 2.6.1985, 13°47.0'N, 120°30.3'E, 640-668 m, SW Luzon: Fungiacyathus (B.) granulosus, Madrepora

oculata, Notocyathus conicus, Truncatoflabellum paripavoninum.

Stn 107. — 2.6.1985, 14°01.9’N, 120°27.9’E, 111-115 m, SW Luzon: Asterosmilia marchadi, Caryophyllia (A.)

grayi, Flabellum (F.) politum, Letepsammia formosissima, Truncatoflabellum candeanum.

Stn 108. — 2.6.1985, 14°01.1'N, 120°17.9'E, 188-195 m, SW Luzon: Anthemiphyllia dentata, Balanophyllia

cornu, Balanophyllia serrata, Balanophyllia pan'ula, Caryophyllia (A.) grayi, Caryophyllia (A.) spinigera,

Flabellum (F.j lamellulosum, Flabellum (F.) politum, Flabellum (U.) deludens, Fungiacyathus (B .) variegatus,

Letepsammia formosissima C89, Tethocyathus virgatus, Trochocyathus (T.) philippinensis, Tropidocyathus

lessonii, "Tropidocyathus" pileus, Truncatoflabellum candeanum.

Stn 109. 2.6.1985, 14°00.2'N, 120°17.6'E, 188-190 m, SW Luzon: Caryophyllia (A.) spinigera, Conotrochus

brunneus, Flabellum (F.) lamellulosum, Flabellum (F.) politum, Flabellum (U.) deludens, Letepsammia

formosissima C89, Stephanocyathus (A.) spiniger, "Tropidocyathus" pileus, Truncatoflabellum candeanum,

Truncatoflabellum pusillum.

Stn 110. — 2.6.1985, 13°59.5'S, 120°18.2'E, 187-193 m, SW Luzon: Flabellum (F.) politum, Letepsammia

formosissima, Truncatoflabellum candeanum.

Stn 111. 2.6.1985, 14°00.1'N, 120°17.5'E, 193-205 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Letepsammia formosissima.

Stn 112. — 2.6.1985, 14°00.2'N, 120°19.2'E, 187-199 m, SW Luzon: Caryophyllia (A.) spinigera, Flabellum

(F.) lamellulosum, Flabellum (U.) deludens, Fungiacyathus (B.) variegatus, Letepsammia formosissima.

52 S. D. CAIRNS & H. ZIBROWIUS

Sln 116. — 3.6.1985, 12°32.2'N, 120°46.4'E, 804-812 m, SW Mindoro: Fungiacyathus (F.) stephanus,

Rhombopsammia niphada.

Stn 1 17. — 3.6.1985, 12°31.2’N, 120°39.3'E, 92-97 m, SW Mindoro: Balanophyllia rediviva, Caryophyllia (C.)

hawaiiensis, Conotrochus brunneus, Deltocyathoides orientalis, Dendrophyllia cf. ijimai, Guynia annulata,

Idiotrochus kikutii, Madracis cf. pharensis, Rhizosmilia sagamiensis.

Stn 119. — 3.6.1985, 11°59.7’N, 121°12.7'E, 320-327 m, S Mindoro: Caryophyllia (A.) spinicarens.

Sin 120. — 3.6.1985, 12°05.6'N, 1 2 1 ° 15.6’E, 219-220 m, S Mindoro: Balanophyllia cornu, Caryophyllia (A.)

spinigera, Deltocyathus magnificus, Flabellum (U.j deludens, Stephanocyathus (A.) spiniger.

Stn 121. — 3.6.1985, 12°08.3’N, 121°17.3’E, 73-84 m, S Mindoro: Fungiacyathus (B.) variegatus.

Stn 122. — 4.6.1985, 12°20.0'N, 121°4I.6’E, 673-675m, SE Mindoro: Flabellum (U.) japonicum, Madrepora

oculata.

Stn 123. — 4.6.1985, 12°10.6'N, 121°45.5'E, 700-702 in, NW Panay: Flabellum (U.) japonicum, Madrepora

oculata.

Stn 124. — 4.6.1985, 12°02.6'N, 121°35.3'E, 120-123 m, between Panay and Mindoro: Caryophyllia (A.) grayi,

Letepsammia formosissima, Trochocyathus (T.) philippinensis, "Tropidocyathus" pileus, Truncatoflabellum

mortenseni.

Stn 125. — 4.6.1985, 11°57.7’N, 121°28.5'E, 388-404 m, between Panay and Mindoro: Caryophyllia ( A .)

spinicarens.

Stn 126. — 4.6.1985, 11°49.2'N, 121°22.1'E, 266 m, between Panay and Mindoro: Anthemiphyllia dentata,

Balanophyllia cornu, Confluphyllia juncta, Conotrochus brunneus, Caryophyllia (C.) secta, Deltocyathus

andamanicus, Flabellum (F.) patens, Letepsammia formosissima, Madrepora arbuscula, Truncatoflabellum

angustum.

Stn 128. — 5.6.1985, 1 1°49.7'N, 121°41.2'E, 815-821 m, NW Panay: Flabellum ( U.) japonicum, Madrepora

oculata.

Stn 130. — 5.6.1985, 11°36.7’N, 121°43.5'E, 178-195 m, NW Panay: Anthemiphyllia dentata, Deltocyathus

andamanicus, Flabellum (F.) lamellulosum, Fungiacyathus (F.) paliferus, Truncatoflabellum angustum.

Stn 131. — 5.6.1985, 11°36.6'N, 121°43.0'E, 120-122 m, NW Panay: Anthemiphyllia dentata, Asterosmilia

marchadi, Balanophyllia sp., Caryophyllia (A.) grayi, Caryophyllia (C.) lamellifera, Cyathelia axillaris,

Dendrophyllia arbuscula, Endopachys grayi, Flabellum (F.) pavoninum, Fungiacyathus (F.) paliferus, Guynia

annulata, Javania insignis, Letepsammia formosissima, Madracis cf. pharensis, Rhizosmilia sagamiensis,

Rhizotrochus typus, Stephanophyllia neglecta C89, Thalamophyllia tenuescens, Trochocyathus (T.)

philippinensis, Tropidocyathus lessonii C89, Truncatoflabellum mortenseni, Truncatoflabellum formosum.

Stn 133. — 5.6.1985, 1 1°57.8’N, 121°52.25'E, 334-390 m, NW Panay: Balanophyllia cornu, Flabellum (F.) sp.

Stn 134. — 5.6.1985, 12°01.1'N, 121°57.3'E, 92-95 m, N Panay: Caryophyllia (C.) lamellifera, Madracis cf.

pharensis, Rhizosmilia sagamiensis, Trochocyathus (T.) maculatus.

Stn 135. — 5.6.1985, 11°58.6'N, 122°01.8'E, 486-551 m, N Panay: Flabellum (F.) lamellulosum, Flabellum (F.)

magnificum.

Stn 137. — 6.6.1985, 12°03.5'N, 122°05.8'E, 56 m, N Panay: Eguchipsammia wellsi, Trochocyathus (T.)

cooperi, Truncatoflabellum phoenix.

Stn 138. — 6.6.1985, 1 1°53.8'N, 122°15.9'E, 252-370 m, N Panay: Deltocyathus andamanicus, Caryophyllia (A.)

spinicarens.

Stn 139. — 6.6.1985, 11°52.9'N, 122°14.7’E, 240-267 m, N Panay: Caryophyllia (A.) spinicarens, Caryophyllia

(A.) spinigera, Conotrochus brunneus, Deltocyathoides orientalis, Deltocyathus andamanicus, Flabellum (U.)

deludens, Letepsammia formosissima, Madrepora oculata.

Stn 140. — 6.6.1985, 1 1°42.6’N, 122°34.5'E, 93-99 m, N Panay: Asterosmilia marchadi, Caryophyllia (A.) grayi,

Fungiacyathus (B.) variegatus, Guynia annulata, Trochocyathus (T.) semperi.

Stn 142. — 6.6.1985, 11°47.3'N, 123°01.5'E, 26-27 m, N Panay: Balanophyllia imperialis, Eguchipsammia

wellsi, Truncatoflabellum aculeatum.

Stn 143. — 7.6.1985, 11°28.3’N, 124°11.6'E, 205-214 m, NW Leyte: Flabellum (F.) lamellulosum, Flabellum

(F.) politum, Letepsammia formosissima, Stephanocyathus (A.) spiniger, Truncatoflabellum angustum,

Truncatoflabellum candeanum.

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

53

Stn 145. — 7.6.1985, 11°01.6'N, 124°04.2'E, 214-246 m, NE Cebu: Caryophyllia (A.) spinicarens, Flabellum

(U.) deludens.

"SlBOGA"

Stn 12. — 14.3.1899, 7°15'S, 115°15.6'E, 289 m, Bali Sea, S Kangeang Isl., Bali Sea: Caryophyllia (C.)

transversalis, Madrepora arbuscula.

Stn 41. — 3.4.1899, 7°25'S, 1 17°50.5'E, 96 m, Pulau Tenga (= Paternoster) Isl.: Balanophyllia generatrix.

Stn 49a. — 14.4.1899, 8°23.5'S, 119°04.6'E, 69 m, E Sumbawa, Sapeh Strait: Fungiacyathus (F.) paliferus,

Truncatoflabellum sp.

Stn 52. — 20.4.1899, 9°03.4'S, 1 19°56.7'E, 959 m, SW Flores: Sabinotrochus bipatella (= Stephanocyathus ).

Stn 59. — 26.4.1899, 10°22.7'S, 123°16.5'E, 390 m, W entrance Samau Strait: Caryophyllia (C.) diomedeae,

Premocyathus dentiformis.

Stn 91. — 22.6.1899, E Borneo, Moeras reef, max. 54 m: Placotrochus laevis.

Stn 95. — 26.6.1899, 5°43.5'N, 1 19°40'E, 522 m, Sulu Isl.: Balanophyllia gemma, Caryophyllia (C.) diomedeae,

Deltocyathus philippinensis, Flabellum (U.) japonicum, Fungiacyathus (F.) stephanus, Madrepora oculata,

Truncatoflabellum dens.

Stn 100. — 29.6.1899, 6°H'N, 120°37.5'E, 450 m, Sulu Isl.: Deltocyathus philippinensis.

Stn 102. — 1.7.1899, 6°04.1'N, 120°44'E, 535 m, Sulu Isl., "dredge full of fine yellow sand, nearly no animals :

Balanophyllia generatrix CSD?, Thrypticotrochus multilobatus CSD?

Stn 105. — 4,7.1899, 6°08'N, 121°19'E, 275 m, Sulu Isl.: Crispatotrochus rubescens, Javania ?lamprotichum.

Stn 116. — 12.7.1899, 0°58.5'N, 122°42.5'E, 72 m, N Sulawesi, W Kwandang Bay entrance: Placotrochus laevis.

Stn 133. — 25-27.7.1899, Talaut Isl., Salibabu Isl., Lirung, anchorage, max. 36 m: Placotrochus laevis.

Stn 150. — 1 1.7.1899, 0°06'N, 129°07.2'E, 1089 m, E Halmahera: Conotrochus brunneus CSD?

Stn 156. — 15.8.1899, 0°29.2'S, 130°05.3'E, 469 m, SW Waigeu: Madrepora oculata.

Stn 159. — 16.8.1899, 0°59. l'S, 129°48.8'E, 411 m, SE Halmahera: Lochmaeotrochus oculeus.

Stn 204. — 20.9.1899, 4°20'S, 122°58'E, 75-94 m, N entrance Buton Strait: Caryophyllia (A.) grayi, Flabellum

(F.) politum.

Stn 211. — 25.9.1899, 5°40.7'S, 120°45.5'E, 1158 m, NE Saleyer: Sabinotrochus flatiliseptis

(= Stephanocyathus).

Stn 212. — 26.9.1899, 5°54.5'S, 120°19.2'E, 462 m, Saleyer: Flabellum (U.) hoffmeisteri, Placotrochides

scaphula.

Stn 231. — 14-18. 1 1 .1899, Ambon, anchorage, 40 m: Tubastraea coccinea.

Stn 240. — 22.1 1.-1.12.1899, Banda, anchorage, 9-45 m: Blastotrochus nutrix, Tubastraea micranthus.

Stn 251. — 8.12.1899, 5°28.4'S, 132°00.2'E, 204 m, Kai Isl.: Balanophyllia panmla, Deltocyathus suluensis,

Flabellum (F.) patens.

Stn 253. — 10.12.1899, 5°48.2'S, 132°13'E, 304 m, Kai Isl.: Truncatoflabellum sp.

Stn 256. — 1 1.12.1899, 5°26.6'S, 132°32.5'E, 397 m, Kai Isl.: Caryophyllia (C.) transversalis, Paracyathus sp.

Stn 258. — 12-16.12.1899, Kai Isl.., Tual anchorage, 22 m: Leptopsammia crassa.

Stn 259. — 16.12.1899, 5°29.2'S, 132°52.5'E, 487 m, Kai Isl.: Desmophyllum dianthus, Lochmaeotrochus

oculeus, Madrepora oculata.

Stn 260. 16.12.1899, 5°36.5'S, 132°55.2'E, 90 m, Kai Isl.: Cyathelia axillaris, Flabellum (F.) politum,

Rhizotrochus ty’pus.

Stn 262. 18.12.1899, 5°53.8'S, 132°48.8'E, 560 m, Kai Isl.: Trochocyathus (T.) brevispina.

Stn 266. 19.12.1899, 5°56.5'S, 132°47.7'E, 595 m, Kai Isl.: Enallopsammia pusilla.

Stn 273. 23-26.12.1899, NE Aru Isl., Pulu Jedan, anchorage, 13 m: Placotrochus laevis.

Stn 274. — 26.12.1899, 5°28.2'S, 134°53.9'E, 57 m, NE Aru Isl.: Truncatoflabellum formosum.

Stn 277. 9-1 1.1.1900, Banda Sea, Dammer Isl., Kullewatti (Sollot) Bay, 45 m: Dendrophyllia arbuscula.

Stn 279. 1 1-13.1.1900, Banda Sea, Roma Isl., Rumah-Kuda Bay, 36 m: Placotrochus laevis.

Stn 289. 20.1.1900, 9°00.3'S, 126°24.5’E, 1 12 m, S Timor: Caryophyllia (A.) grayi, Neohelia cf. porcellana.

Stn 297. 27.1.1900, 10°39'S, 123°40'E, 520 m, E Rotti Isl.: Balanophyllia cornu, Placotrochides scaphula.

Source :

54

S. D. CAIRNS & H. ZIBROWIUS

Stn 299. — 27-29.1.1900, 10°52.4'S, 123°01.I'E, max. 34 m (dive), S Rotti Isl., Boeka or Cyrus Bay:

Truncatoflabellum spheniscus.

Stn 303. — 2-5.1.1900, Samau Isl., Haingsisi, max. 36 m: Truncatoflabellum incrustatum, Truncatoflabellum

irregulare.

Stn 305. — 8.2.1900, mid-channel in Solor Strait off Kampong Menanga, 1 13 m: Neohelia cf. porcellana.

Stn 310. — 12.2.1900, 8°30'S, 1 19°07.5'E, 73 m, E Sumbawa: Cyathelia axillaris, Eguchipsammia gaditana.

Stn 315. — 17-18.2.1900, Pulau Tenga (Paternoster) Isl., anchorage east of Sailus Besar, max. 36 m:

Blastotrochus nutrix.

Smithsonian Institution Marquesas Expedition, "Pele"

Stn TH1. — 28.9.1967, 10°S, 139°10'W, 75-79 m, Marquesas: Trochocyathus (T.) cooperi.

Smithsonian Institution Philippine Expedition (= Siphilexp), "Stinc, Ray" (mainly)

Stn 78-CAC189. — 7.5.1978, Cebu, Tanon Strait, Pescador Isl., 12-18 m: Tubastraea micranthus.

Stn 78-CAC194. — 1 1.5.1978, 9°04'15"N, 123°16'10"E, 1-6 m, SE Negros: Tubastraea coccinea.

Stn 78-SP1-1. — 8.5.1978, 9°25'15"N, 123°18'10"E, 1-5 m, SE Negros: Tubastraea diaphana.

Stn 78-SP40. — 11.6.1978, 9°31'14"N, 123°40'00"W, depth? (bought), Bohol Strait: Stephanocyathus (A.)

spiniger.

Stn 78-T10. — 6.6.1978, 1 1°35'46"N, 123°55'32"E, 75 m, Visayan Sea: Asterosmilia marchadi.

Stn 78-T14 . — 6.6.1978, 1 1°34'45"N, 123°52'08"E, 84 m, Visayan Sea: Balanophyllia carinata.

Snellius 2 Expedition, "Tyro" (mainly)

Stn 81.2. — 28.8.1984, 6°59'S, 131°30'E, 340 m, Tanimbar Isl.: Anthemiphyllia frustum, Bourneotrochus

stellulatus, Caryophyllia (C.) quadragenaria, Deltocyathoides oriental is, Javania pachy theca, Notocyathus

venustus, Peponocyathus minimus.

Stn D2. — 3.3.1985, 6°57.9'S, 131°39.7'E, 91 m, Tanimbar Isl.: Deltocyathoides orientalis, Idiotrochus kikutii,

Notocyathus venustus, Truncatoflabellum phoenix.

Stn 4.019. — 9.9.1984, 5°57.5'S, 123°46.5'E, 285-305 m, Tukang Besi Isl., S Karang Kaledupa: Fungiacyathus

(F.) paliferus, Stephanophyllia fungulus.

Stn 4.032. — 10.9.1984, 5°52.5'S, 123°58.5'E, ca. 385 m, Tukang Besi Isl., NW Binongko: Truncatoflabellum

dens.

Stn 4.033. — 10.9.1984, 5°52.5'S, 123°58.5'E, 250-290 m, Tukang Besi Isl., NW Binongko: Anthemiphyllia

dentata, Fungiacyathus (F.) paliferus.

Stn 4.034. — 10.9.1984, 5°52.5'S, 123°58.5'E, 280 m, Tukang Besi Isl., NW Binongko: Anthemiphyllia dentata,

Deltocyathus Stella, Fungiacyathus (F.) paliferus.

Stn 4.039. — 10.9.1984, 5°54'S, 123°57.7'E, ca. 525 m, Tukang Besi Isl., NW Binongko: Stephanophyllia

neglecta.

Stn 4.056. — 14.9.1984, 9°54'S, 120°44.8'E, 125 m, NE Sumba, E Melolo: Conocyathus zelandiae.

Stn 4.057. — 14.9.1984, 9°52.8'S, 120°44.7'E, 154 m, NE Sumba, E Melolo: Letepsammia formosissima.

Stn 4.066. — 16.9.1984, 9°53'S, 120°53’E, 295 m, NE Sumba: Crispatotrochus rubescens.

Stn 4.070. — 17.9.1984, 8°36'S, 1 19°31.2'E, rocky shore, E Komodo, Teluk Slawi: Endopsammia philippensis.

Stn 4.099. — 19.9.1984, 8°29'S, 1 19°38.2'E, 81 m, E Komodo: Truncatoflabellum aculeatum.

Stn 4.100. — 19.9.1984, 8°28.6'S, 1 19°37.3'E, 91 m, E Komodo: Javania insignis, Rhizopsammia nuda.

Stn 4.104. — 20.9.1984, 8°25.3'S, 1 19°36.2’E, 140-150 m, NE Komodo: Neohelia cf. porcellana.

Sm 4.105. — 20.9.1984, 8°25.7'S, 119°37.8'E, 105-120 m, NE Komodo: Neohelia cf. porcellana.

Stn 4.106. — 20.9.1984, 8°26.9'S, 1 19°37.9'E, 80 m, NE Komodo: Cyathelia axillaris, Rhizotrochus typus.

Stn 4.115. — 21.9.1984, 8°19.4’S, 1 18°15.3'E, 60-75 m, N Sumbawa, Bay of Sanggar: Rhizotrochus typus.

Stn 4.130. — 23.9.1984, 8°17.9'S, 1 18°17.8'E, 700-730 m, N Sumbawa, Bay of Sanggar: Deltocyathus rotulus.

Stn 4.134. — 25.9.1984, 6°31'S, 121°08.2'E, 53-59 m, NE Taka Bone Rate (Tiger Isl.), SE Tarupa Kecil:

Truncatoflabellum aculeatum.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

55

Stn 4.144. — 26.9.1984, 6°26.7'S, 121°10'E, 730-850 m, NE Taka Bone Rate (Tiger Isl.), NE Tarupa Besar:

Enallopsammia pusilla, Madrepora oculata.

Stn 4.173. — 1.10.1984, 6°28.5S, 120°24.3'E, 300-340 m, SW Salayer, NW Pulau Bahuluang: Deltocyathus

andamanicus.

Stn 4.174. — 1.10.1984, 6°28.1 S, 120°24.2'E, 330 m, SW Salayer, NW of Pulau Bahuluang: Fungiacyathus (F.)

variegatus.

Stn 4.196. — 9.10.1984, 6°23.0'S, 120°26.5'E, 150-200 m, SW Salayer, off Tanjung Batu Kerapo: Madrepora

minutiseptum.

Stn 4.226. 17.10.1984, 6°32 S, 121°10.5'E, NE Taka Bone Rate (Tiger Isl.), N Pulau Tinanja, scuba diving

along reef edge: Thalamophyllia tenuescens CSD?

Stn 4.228. — 15.10.1984, 6°32.1'S, 121°07.5'E, 60 m, NE Taka Bone Rate (Tiger Isl.), N Pulau Tarupa Kecil:

Balanophyllia carinata, Balanophyllia stimpsonii, Truncatoflabellum aculeatum.

Stn 4.232. — 16.10.1984, 6°32. 1 'S, 121°09.0'E, 59 m, NE Taka Bone Rate (Tiger Isl.), S Pulau Tarupa Kecil:

Balanophyllia stimpsonii, Placotrochides laevis, Truncatoflabellum aculeatum.

Stn 4.234. — 17.10.1984, 6°31.6'S, 121°07.5'E, 58 m, NE Taka Bone Rate (Tiger Isl.), S Pulau Tarupa Kecil:

Balanophyllia carinata, Balanophyllia stimpsonii, Placotrochides laevis, Trochocyathus (T.) cooperi,

Truncatoflabellum aculeatum.

Stn 4.235. — 18.10.1984, 6°32.7’S, 121°08.7'E, 53-57 m, NE Taka Bone Rate (Tiger Isl.), S Pulau Tarupa

Kecil : Asterosmilia marchadi, Balanophyllia stimpsonii, Placotrochides laevis.

"Tangaroa" New Zealand Oceanographic Institute (= NZOI),

Stn G3. — 27.9.1966, 26°25.0'S, 167°15.0’E, 710 m, Norfolk Ridge: Flabellum (U.) sp.

Stn K858. — 30.7.1974, 30°34.2'S, 178°29.8'W, 465 m, Kermadec Isl.: Truncatoflabellum angustum.

Stn Q47. — 24.5.1978, 33°06'S, 156°1 l’E, 135 m, Tasman Sea, Taupo Seamount: Dendrophyllia cf. ijimai.

Stn T243. — 24.3.1982, 30°05.0'S, 178°15.0'W, 1035 m, Kermadec Isl.: Stephanocvathus (S.) regius.

Stn U582. — 5.2.1988, 31°52.0’S, 172°26.5'E, 988-1058 m. Three Kings Ridge: Flabellum (U.) sp.

"Tansei Maru"

Stn KT86-16-F. — 3.11.1986, 31°55.3'N, 133°23.9'E, 2576-2603 m, S Shikoku: Caryophyllia (C.) cornulum,

Flabellum (U.) conuis.

Stn KT90-13-T6. — 3.9.1990, 32°14.23’N, 134°01.3'E, 2547-2565 m, S Shikoku: Carvophyllia (C.) cornulum.

Stn KT93-09-AM6. — 22.6.1993, 28°20.48'N, 129°40.18'E, 107-108 m, Ryukyu, Amami: Trochocyathus (T.)

philippinensis.

Stn KT93-09-AM7. — 22.6.1993, 28°I7.07'N, 129°40.15'E, 191-196 m, Ryukyu, Amami: Guynia annulata,

Trochocyathus (T.) philippinensis.

Stn KT93-09-AM8. 25.6.1993, 28°1 l'N, 129°43'E, 422-425 m, Ryukyu, Amami: "Tropidocyathus" labidus.

"Te Vega "

Stn 1-54. 25.9.1963, 1°08.6'N, 128°01'E, 46-55 m, Halmahera: Balanophyllia imperialis, Balanophyllia

stimpsonii.

HISTORICAL REVIEW (INDONESIAN REGION)

Perhaps the earliest azooxanthellate corals reported from the Indonesian region were those collected by the

H. M. S. "Challenger" at stations 191-198 and 214, as reported by MOSELEY (1876, 1881). Sixteen species were

described, including 1 1 from one station (" Challenger " stn 192) at the Kai Islands at 256 m, 9 of those new

species. The Challenger specimens are deposited at the BMNH and were examined by both authors.

56

S. D. CAIRNS & H. ZIBROWIUS

The historically most significant collection of Indonesian deep-water corals, reported in at least 8 publications,

was that of the "Siboga" expedition of 1899-1900, which made 323 stations throughout the Indonesian region and

Sulu Archipelago (Weber, 1902; Tydeman, 1902). ALCOCK (1902a, b, July) published 2 preliminary accounts

in which he described 45 new species of deep-water corals from this expedition. Although the descriptions are

adequate, he did not include illustrations of these species or station numbers for the specimens. Just one month

later (ALCOCK, 1902c, August) his report on the deep-water ( i.e ., over 100 fathoms, 183 m) corals of the "Siboga

Expedition ("Siboga" Report 16a) appeared, in which he published verbatim the description of 37 of the 45 species

he described the previous month, 3 additional new species (Paracyathus pruinosus, Flabellum laciniatum var.

messum, and Pourtalosmilia dumosa ), 26 previously described species, and 7 lots identified only to the generic

level (a total of 73 azooxanthellate species). Illustrations and station data were included for all taxa in the "Siboga"

report, and, although ALCOCK (1902c: 2) noted that most of the new species had been published in a "preliminary

communication ... of the Journal of the Netherland Zoological Society", the 2 earlier publications were largely

forgotten until 1991 (HOEKSEMA & BEST, 1991). It is unknown why ALCOCK omitted 8 species from his Siboga

report that were included in his preliminary accounts. These species were shallow water (0-183 m)

azooxanthellates, on which he perhaps intended to write a separate "Siboga" report (ALCOCK, 1902c: 2);

nonetheless, 2 of these 8 species were later redescribed by van der HORST (1922) inadvertently as new species

using ALCOCK's original names: Dendrophyllia florulenta and Leptopsammia poculum (the latter described as

Endopsammia poculum by ALCOCK, 1902a). Even VAN DER HORST (1922) was apparently unaware of ALCOCK’s

preliminary papers. Only one of the other 6 species was ever cited; Flabellum weberi, a junior synonym of Javania

insignis was cited by CAIRNS (1989a, 1994), based on a collection label, not the ALCOCK publication. The

remaining 5 species: Trochocyathus weberi (= ? T. cooperi), T. cavatus, Endopachys weberi (= E. grayi),

Heteropsammia pisum , and Rhodopsammia (= Balanophyllia) corniculans , all described in Alcock (1902a), were

not subsequently reported. Aside from the 2 species redescribed by VAN DER HORST, the types of only 2 of the

other 6 species are now known to exist at the ZMA: Flabellum weberi and Endopachys weberi.

In a paper describing the shallow- water fungiids collected by the "Siboga", van der Horst (1921, "Siboga"

Report 16b) also reported 2 specimens of Bathyactis (= Fungiacyathus) palifera; and in a paper dedicated to the

study of Fungia patella, BOSCHMA (1923, "Siboga" Report 16d) also described a new Indonesian species,

Stephanophyllia neglecta, from a "Siboga " station off the Kai Islands. Another paper on "Siboga" azooxanthellates

(VAN DER Horst, 1922, "Siboga" Report 16c) was an account of the dendrophylliids of the expedition, which

included 36 species from Indonesian waters, 1 1 of these new; however, as mentioned above, 2 of these 1 1 new

species were previously described by ALCOCK (1902a). Most "Siboga" specimens are deposited at the ZMA (see

VAN SOEST, 1979 for type deposition) and have been examined by the authors.

Two more papers include reidentifications of "Siboga" specimens: ZIBROWIUS (1973) reidentified the "Siboga"

Dendrophyllia reported by ALCOCK (1902c) in the context of a revision of the genus Enallopsammia; and

ZIBROWIUS & Grygier (1985) reidentified two lots of Balanophyllia reported by VAN DER HORST (1922).

BEDOT (1907) reported Cyathelia axillaris from the Ceram Sea (depth unknown), a species previously reported

by Moseley (1881) from the adjacent Molucca Sea.

Azooxanthellate Scleractinia collected on the Danish Expedition to the Kai Islands (DEKI) in 1922 have been

reported in 3 publications. BOSCHMA (1923) listed 1 species, Stephanophyllia formosissima-, VAN DER HORST

(1926) reported 5 dendrophylliid species, including the new species Rhizopsammia nuda\ and BOSCHMA (1953)

reported Tubastraea aurea ( = T. coccinea and T. faulkneri) from the Kai Islands and Ambon. However, the bulk of

the specimens from that expedition are reported herein and are deposited at the NNM and ZMK (see Material).

More recent reports of Indonesian azooxanthellates from miscellaneous sources include: Polycyathus

furanaensis from Sulawesi (VERHEIJ & BEST, 1987); Anthemiphyllia dentata from the Banda Sea (BEST &

HOEKSEMA, 1987); Cryptotrochus javanus from the Java Sea (CAIRNS, 1988); and a revision of Heterocyathus and

Heteropsammia from the Indonesian region (HOEKSEMA & BEST, 1991).

A common theme in previous expedition reports has been a high diversity of deep-water species from the Kai

Islands, and the Indonesian region in general, as evidenced by the collections of the "Challenger", "Siboga", and the

Danish Expedition to the Kai Islands (Deki). This is confirmed herein on the basis of collections from the

SNELLIUS 2 and KARUBAR expeditions. ALCOCK (1902c: 3) alluded to the Kai Islands as second only to the Sulu

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

57

Archipelago in coral biodiversity; however, as a result of this study, 122 species are known from the Kai Islands,

and only 65 from the Sulu Archipelago.

Works containing references to fossil azooxanthellates from the Indonesian region include: Gerth (1921)

Miocene of Java; UMBGROVE (1938), Pleistocene of Talaud; Umbgrove (1950), Pleistocene of Java; and

Boekschoten et al. (1989), Pliocene-Pleistocene of Nias.

A review of previous records of azooxanthellate Scleractinia from the Philippines can be found in Cairns

(1989a).

MATERIAL

This study is based on the examination of approximately 15,600 coral specimens collected throughout the

Philippine-Indonesian region from some 640 stations. A specimen equates to an individual solitary corallum or a

colony or branch of a colonial species. Many of the new specimens were collected by the French and French-

Indonesian expeditions: MUSORSTOM cruises 1-3 (1976, 1980, 1986, respectively), which concentrated on

southwestern Luzon (vicinity of Lubang Island and Verde Island Passage), and also did some collecting near

southern Luzon (Mindoro and Panay) (FOREST, 1981, 1985, 1989); CORINDON 2 (1980), on the R.V. Coriolis,

which collected in Makassar Strait (Moosa, 1984); and Karubar (1991) on the R.V. Banina Jaya 1 , which made

extensive deep-water collections in the Banda Sea (Kai Islands) and Arafura Sea (southeast of Tanimbar Islands)

(CROSNIER, Richer de Forges & BOUCHET, 1997). Many of the MUSORSTOM 1 stations were made in a small

area north of Lubang Island near "Albatross" stn 5278 (the type locality of Neoglyphea inopinata), a "living fossil"

decapod crustacean. The French collections are divided among the MNHN, NMNH, and Puslitbang Oseanologi,

Jakarta. Many additional specimens from the Philippines and Banda Sea also came from the "Albatross" expedition

(1906-1909; see ANONYMOUS, 1910 for a history and station list of the expedition), and are deposited at the

NMNH. A remarkably large and diverse collection of azooxanthellates was also made available from the Danish

Expedition to the Kei Islands made in 1922 by Th. MORTENSEN and H. BOSCHMA. This expedition collected not

only near the Kai Islands but also in the Java Sea (MORTENSEN, 1923). Specimens are deposited primarily in the

NNM and to a lesser extent at the ZMK. Few specimens of shallow water azooxanthellates were found at the NNM

from the Snellius" expedition to Indonesia in 1930, but many upper slope specimens were available for study at

the NNM from the Snellius 2 (R/V Tyro) expedition of 1984, which made deep-water stations throughout the

Indonesian region (van DER Land & SUKARNO, 1986). Specimens from the following expeditions were also

examined. Galathea expedition (1950-1952), throughout the Philippine and Indonesian region (see Bruun, 1957;

WOLFF, 1964), deposited at the ZMK; Mortensen's Java-South African Expedition (1929-1930), primarily from the

Bali Strait, deposited at the ZMK; stations made by the "Hakuho Maru" (1972-1985) in the South China Sea,

Philippine, and Indonesian waters (see Nishiwaki, 1974; HORIKOSHI et al., 1983; HORIKOSHI & Ohta, 1987),’

deposited at the ORI and NMNH; and specimens from "Siboga" stations (1899-1900) in Indonesian waters that

were not previously reported (see Tydeman, 1902).

In addition to the newly reported specimens listed above, all previously reported specimens from the "Siboga"

xpedition (ALCOCK, 1902a-c; Horst, 1921, 1922; BOSCHMA, 1923) and specimens from the Indonesian and

Philippine stations of the "Challenger" Expedition (stations 191-213) were re-examined by the first author in 1994

an > the second author on various occasions. Semper's (1872) Philippine specimens were examined by the

second author.

Although not politically part of Indonesia or the Philippines, specimens from the eastern coast of Sabah,

alaysia (e.g., Darvel Bay, Celebes Sea) were included in this study, and their depth ranges included with those of

the Indonesian region.

METHODS

Species descriptions and illustrations are provided only for those species described as new or for those for which

equate cscription previously existed. Shorter diagnoses are provided for the remaining species for which new

ma ena is reported, with an indication as to where to Find a more complete description. New material was collected

Source :

58

S. D. CAIRNS & H. ZIBROWIUS

of 176 (85%) of the 206 species known from the Philippine-Indonesian region, the remaining 30 species being

indicated in Table 1 by an asterisk but not discussed in the text. Most of these 30 are rarely collected species,

indeterminable species (e.g., those described but not illustrated by ALCOCK, 1902a, b), or probable junior

synonyms of species better known under other names (e.g., Balanophyllia dubia, B. ovalis, B. parallela). Although

some species of Heterocyathus and Heteropsammia occur in this region and are acknowledged to be

azooxanthellate, they are not included in this report.

Species synonymies are complete unless otherwise indicated with a reference to a more complete account;

however, it was attempted to include in the synonymies all references to specimens reported from the Philippine-

Indonesian region. When possible, all historical records were verified, but when material was unavailable and the

published account unclear, the synonymy entry and corresponding distribution record are queried.

In the "Material examined" sections, the number of specimens examined follows the station number, followed

by the museum of deposition, and its catalog number, if any. Holotypes and paratypes are deposited primarily at

the MNHN and NMNH, as well as Puslitbang Oseanologi - LIPI, Indonesia, in the case of Karubar material.

Essentially, only new material is reported in the Material Examined sections, not types or previously reported

specimens.

In order to avoid erroneous depth ranges for species as a result of bathymetrically wide-ranging trawls, a

confirmed depth range is employed in this paper, which is defined as the deepest shallow to the shallowest deep

component of all trawls considered. For example, if a species was trawled at a station indicating 20-300 m and

again at a station indicating 250-500 m, the confirmed depth range is 250-300 m, a depth range within which it

most likely was collected.

The SEM and most conventional photography was done by the first author, the former on a Cambridge

Stereoscan 100 in the SEM Laboratory of the NMNH.

COMMENSAL RELATIONSHIPS

Several types of specialized coral symbionts (other than simple epibionts) have been found associated with

various members of the species-rich fauna of the Philippines and Indonesia studied here.

Lumbrinerid polychaete eroding the coral skeleton. — This association has been described in detail

by ZIBROWIUS et al. (1975) on the basis of material from the northeastern Atlantic and the southwestern Indian

Ocean (South Africa), with additional records from Madagascar, the China Sea, and Japan. The coral-skeleton-

eroding polychaete Lumbrineris flabellicola (Fage, 1936) inhabits a soft tube exteriorly attached to the host, and

causes a superficial to deep erosion of the coral skeleton. The worm itself is easily lost by the mechanical con¬

straints of dredging and the subsequent manipulations, but frequently empty tube fragments remain attached to the

coral, or a corrosion trace can be detected on the coral even after the worm and tube have disappeared. This associa¬

tion occurs in the Philippines and Indonesia. Worms obtained during cruise MUSORSTOM 2 in the Philippines have

been compared by T. MlURA with Lumbrineris flabellicola from the northeastern Atlantic and Japan and have been

found to be the same species (L. flabellicola) in these widely distant areas (T. MlURA, in litt., 1989).

The following species have been found to be the coral partner of this association, specimens still bearing the

worm (WO), or still having empty tubes fragments attached (ET), or showing only a characteristic erosion trace

left over (TR):

Ca ryophyl lia (C.) transversalis: DEKI stn 32 (WO).

Caryophyllia (A.) grayi : MUSORSTOM 2 stn 29 (ET); MUSORSTOM 3 stn 131 (ET, TR).

Caryophyllia (A.) spinigera : MUSORSTOM 2 stn 63 (WO).

Caryophyllia (A.) spinicarens: "Albatross" stn 5256 (ET), stn 5418 (TR), stn 5535 (ET), stn 5536 (TR),

stn 5538 (TR); MUSORSTOM 1 stn 20 (TR?); MUSORSTOM 2 stn 63 (ET).

Conotrochus brunneus : MUSORSTOM 3 stn 92 (ET, TR).

Flabellum (F.) patens: Karubar stn 31 (TR).

Flabellum (F.) lamellulosum: MUSORSTOM 1 stn 27 (TR?), stn 31 (TR); MUSORSTOM 2 stn 63 (ET);

MUSORSTOM 3 stn 86 (WO), stn 92 (WO, TR).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

59

Flabellum (F.) sp.: Musorstom 3 stn 133 (ET).

Rhizotrochus typus: MUSORSTOM 3 stn 131 (ET).

Balanophyllia sp.: MUSORSTOM 2 stn 33 (WO).

Dendrophylliidae, colonial: MUSORSTOM 2 stn 33 (WO).

Eunicid polychaete causing deformation of the coral colony. — Some colonies of Madrepora

oculata from Indonesia (" Albatross " stn 5645; Karubar stn 56) show deformations similar to those found in

colonies of Madrepora oculata , Lophelia pertusa , and Solenosmilia variabilis from the northeastern Atlantic, in

which the parchment-like tube of Eunice norvegica (Linnaeus, 1767) is overgrown by the coral coenosteum and

incorporated into the colony (Zibrowius, 1980). We have no information whether the deformations of Indonesian

M. oculata is caused by the same Eunice species. Similar deformations characterize all colonies of Madrepora

arbuscula and of Madrepora minutiseptum studied here. Even though no overgrown soft tube has been formally

identified in this material and no worm been extracted, it is presumed that the commensal organism is an eunicid

polychaete. Overgrown parchment-like Eunice tubes have also been found in some colonies of Neohelia sp.

cf. N. porcellana.

Acrothoracican cirripede crustacean boring the coral skeleton. — Acrothoracican cirripeds may

bore the skeleton of live corals and when penetrating through the wall cause the polyp to deposit additional wall

material that is intended to seal off the borer. The orifice of the burrow may migrate upward along the growing

coral (Grygier & Newman, 1985). Orifice motility is particularly marked in a specimen of Javania

lamprotichum (MUSORSTOM 2 stn 53) bored by 4 large acrothoracids. Other species bored alive are Tethocyathus

virgatus (MUSORSTOM 3 stn 108), Balanophyllia crassiseptum (Karubar stn 50) and Balanophyllia sp.

(Musorstom l stn. 61; Musorstom 2 stn 32; Musorstom 3 stn 131).

Ascothoracidan crustacean inducing a skeleton gall. — The most common aspect of this

association has been described in detail by Zibrowius & Grygier (1985), who already reported some examples

from the Philippines and Indonesia: "internal galls" are recognizable as a spongy proliferation of the columella that

covers the underlying cavity occupied by the parasite. The list from the Philippines and Indonesia now includes :

Deltocyathoides orientalis (" Albatross " stn 5178, 5313, 5314, 5315, 5317, 5403, 5569); Flabellum lamellulosum

(MUSORSTOM 2 stn 83); Balanophyllia carinata ("Siboga" stn 240); Balanophyllia sp.

(Musorstom 2 stn 34); Balanophyllia sp. (MUSORSTOM 3 stn 131); and Dendrophyllia sp. cf. D. ijimai

(Musorstom 2 stn 33). A newly recognized expression (Grygier & Cairns, 1996) of ascothoracidan gall

induction are abnormally hypertrophied corallites in Madrepora oculata (". Albatross " stn 5529; Deki stn 50;

"Hakuho Maru" stn KH-73-2-44-2; KARUBAR stn 9, 13, 19, 77).

Cryptochirid crab inhabiting a crypt in the coral skeleton. — Cryptochirid (formerly

hapalocarcinid) crabs are obligate symbionts of scleractinians. The crypts (or in some cases cage-like galls) they

inhabit are due to dissolution of the coral skeleton and to induced modified coral growth (ZIBROWIUS, 1982;

Zibrowius & Gili, 1990). Previous to these authors, cryptochirids had always been considered as typical of the

reef fauna, but new deep-water species continue to be discovered. Zibrovia galea Kropp & Manning, 1995, has thus

been found on Phyllangia papuensis from the Philippines (MUSORSTOM 2 stn 47) and from Madagascar.

DISTRIBUTION

Regional Diversity. — Vaughan & Wells (1943), ALCOCK (1902c), and Cairns (1989a) have all

suggested that the highest diversity of deep-water corals (azooxanthellates) occurs in the Philippine-Indonesian

region. ALCOCK mentioned the Sulu Archipelago and the Kai Islands as areas with a particularly diversified fauna.

With the possible exception of the New Caledonia-Chesterfield Islands region, which even though a much smaller

region, may have more species, the Philippine-Indonesian region does have the highest recorded number of

azooxanthellate species, i.e., 206 species (Table 1).

Source :

60

S. D. CAIRNS & H. ZIBROWIUS

Table 1. — Geographic distribution and bathymetric ranges of all azooxanthellate species known from the

Philippine-Indonesian region.

Key to areas: 1 = Japan and/or Ryukyu Islands. 2 = South China Sea, 3 = Philippine region. 4 = Celebes Sea and/or

Makassar Strait. 5 = Molucca. Halmahera, and/or Ceram Seas, 6 = Banda Sea (including Teluk Bone), 7 = Arafura,

Timor, and/or Savu Seas, 8 = Flores and/or Bali Seas, 9 = Java Sea, 10 = Indian Ocean, 1 1 = western and central Pacific

islands (including Hawaiian Islands), 12 = Australia and/or New Zealand, 13 = eastern Pacific, 14 = Atlantic Ocean.

Bathymetric ranges (given in meters) only for records from Philippine-Indonesian region.

Symbols: * no new records of these species and not included in text; ** no new records of these species but included in

text; # distribution unknown; + fossil occurrence only.

Source : MNHN Paris

AZOOX ANTH ELL ATE SCLERACTINIA

61

Source

62

S. D. CAIRNS & H. ZIBROWIUS

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

63

GUYNIIDAE

Guynia annulala Duncan, 1872

FLABELL1DAE

Flabellum (F.) pavoninum Lesson, 1831

F. (F.) magnificum Marenzeller, 1904

F. (F.) patens Moseley, 1881

F. IF.) lamellulosum Alcock, 1902

F. (F.) politum Cairns, 1989

F. (U.) deludens Marenzeller. 1904

F. IU.) marenzelleri Cairns, 1989

F. IU.) japonicum Moseley, 1881

F. (U.) hoffmeisteri Cairns & Parker, 1992

F. IU.) messum Alcock, 1902

F. IU.) sp.

F. IU.) sexcostatum Cairns, 1989

F. IU.) conuis Moseley, 1881

Polymyces wellsi Cairns, 1991

Rhizotrochus typus ME & H, 1848

"R." flabelliformis Cairns, 1989

Gardineria philippinensis Cairns, 1989

*G. hawaiiensis Vaughan, 1907

G. paradoxa (Pourtal6s, 1868)

Javania insignis Duncan, 1876

J. lamprotichum (Moseley, 1880)

J. pachytheca Cairns, 1995

J. sp.

Truncatoflabellum spheniscus (Dana, 1846)

T. aculeatum (ME & H, 1848)

*T. crassum (ME & H, 1848)

*71 stokesi (ME & H, 1848)

*71 cumingi (ME & H, 1848)

T. candeanum (ME & H, 1848)

71 incrustatum Cairns, 1989

71 irregulare (Semper, 1 872)

71 paripavoninuni (Alcock, 1894)

71 formosurn Cairns, 1989

71 pusillum Cairns, 1989

71 dens (Alcock, 1902)

71 phoenix Cairns, 1995

T. mortenseni sp. nov.

71 angustum sp. nov.

Blastotrochus nutrix ME & H, 1848

Placotrochides scaphula Alcock, 1902

Placotrochus leavis ME & H, 1848

DENDROPHYLLUDAE

Bulanophyllia carinata (Semper, 1 872)

1 2

x x

Philippine-lndonesian

Region

3 4 5 6 7 8 9

x

X

X X

X

10 11 12 13 14

XXX

X

Depth (m)

97-194

96-156

225-567

204-439

187-486

40-288

176-480

240-390

425-1060

345-477

368-949

441-550

685-772

1160-2570

385-842

70-296

263-390

192-494

192-220

285-323

73-296

191-842

534-601

209-291

30-174

11-81

unknown

256

46-55

70-290

30-415

18-42

411-1022

42-933

85-300

300-522

18-421

50-156

195-490

11-62

462-1628

12-289

33-100

Source

64

S. D. CAIRNS & H. ZIBROWIUS

TOTALS: Area 1: 77. — Area 2: 60. — Area 3: 157. — Area 4: 66. — Area 5: 30. — Area 6: 138. — Area 7: 83. —

Area 8: 51. — Area 9: 20. — Area 10: 64. — Area 11: 47. — Area 12: 67. — Area 13: 11. — Area 14: 11.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

65

Currently, 157 species of azooxanthellate Scleractinia are known from the Philippine Islands (Table 1), which,

when combined with the 410 species of zooxanthellate species known from this country (Veron & HODGSON,

1989), results in 567 species of Scleractinia from this archipelago. 69 of the 157 azooxanthellate species (44%) are

known from a small region north and northeast of Lubang Island, Verde Island Passage, which was intensively

sampled by MUSORSTOM 1, 2 and 3. 66 azooxanthellate species (42%) of the Philippine fauna are recorded from

the Sulu Archipelago, mainly as a result of the "Siboga" and "Albatross" expeditions early in the century.

Cairns (1989a) reported no azooxanthellate species from the eastern coasts of the Philippines (i.e., Pujada Bay,

Mindanao to the Batan Islands); however, the present study indicates 10 such species, primarily found in Leyete

and Lagonoy Gulfs: Fungiacyathus stephanus, Caryophyllia scobinosa, Stephanocyathus regius, S. weberianus,

Deltocyathus magnificus, Rhizosmilia elongata, Peponocyathus minimus, Flabellum deludetis, Truncatoflabellum

incrustatum , and T. formosum.

A total of 174 azooxanthellate species are known from the Indonesian region (Table 1), the most species-rich

subregion being the Banda Sea (138 species) and the most species-rich archipelago the Kai Islands, from which

125 species are known. As can be seen from Table 1, 66 species are known from the Celebes Sea (Sabah) and

Makassar Strait; only 30 from the combined region of the Molucca, Halmahera, and Ceram Seas; 83 from the

Arafura, Timor, and Savu Seas; 5 1 from the Flores and Bali Seas; and 20 from the Java Sea. It is highly probable

that the wide variation in numbers of species for various seas reflects the unequal collecting effort. For example,

the impressive total from the Kai Islands reflects collections made by the "Challenger", "Siboga", Danish

Expedition to the Kei Islands, and the Karubar expeditions, making it one of the most intensively sampled upper

slope areas in the world. Conversely, little collecting has been done in the Java Sea and even less off the southern

coast of Java, which results in low numbers for these regions.

Regional Affinities. — The Philippine-Indonesian region lies at the heart of the largest and most diverse

tropical marine province — the Indo-Polynesian — extending from the Persian Gulf to the Tuamotu Archipelago

(Briggs, 1974). 65 of the 206 species known from this region (31.5%) are also known from the tropical regions

of the Indian Ocean. ALCOCK (1902c) was surprised to discover that only 10% of his Indonesian "Siboga" species

also occurred in the Indian Ocean; later, Cairns (1989a) was able to report an overlap of 25%, and the current ratio

of 31.5% will probably continue to increase as Indian Ocean corals become better known. An even higher number

of Philippine-Indonesian species are found to the north in the warm temperate region of southern Japan (77/206, or

37%). This may be due to the better-known nature of the Japanese fauna (Cairns, 1994) and/or suggests that the

boundaries that distinguish shallow-water tropical and temperate regions do not coincide with boundaries of deep¬

water organisms. The number of shared species with southern tropical regions (i.e., Australia and ridges/islands

north of New Zealand) is similar to that shared with the Indian Ocean (i.e., 67/206, or 32.5%). Only 47 (or 23%)

of the Philippine-Indonesian species are known from other western and central Pacific islands (e.g., Pelau, New

Guinea, Fiji, Hawaiian Islands), which, except for the Hawaiian Islands (Vaughan, 1907; Cairns, 1984), have

been poorly sampled. Only 1 1 Philippine-Indonesian species occur as far east as the west coast of the Americas

(Table 1): of these, 4 are restricted to the Pacific Ocean, 4 have an Indo-Pacific distribution, and 3 are cosmopolitan

in distribution. Of the 1 1 species that also occur in the Atlantic, most (9) are cosmopolitan species, 1 occurs in

the eastern Atlantic and Pacific (i.e., Asterosmilia marchadi), and 1 (i.e., Gardineria paradoxa) is known only from

the western Atlantic and western Pacific. The disjunct nature of the last 2 categories may be the result of inadequate

collection.

Depth Distribution. — 198 azooxanthellate species were scored for their occurrence in 8 bathymetric

zones. The remaining 8 of the 206 species known from this region are not considered because their depth

distribution is unknown or uncertain (Table 1). The results were: 0-100 m (82 species), 100-200 m (89 species),

200-400 m (123 species), 400-600 m (79 species), 600-800 m (40 species), 800-1000 m (25 species), 1000-

2000 m (19 species), and over 2000 m (4 species). When these data are graphed (depth range on the x-axis,

number of species on the y-axis), a bell-shaped curve results, in this case peaking at the 200-400 m category (62%)

and gradually attenuating with greater depth to only 2.5% at over 2000 m. This is consistent with a peak diversity

ol 53% at 200-300 fathoms (= 366-549 m) for Philippine azooxanthellates given by VAUGHAN & WELLS (1943,

table 2), although their data were limited to only 51 species. A similar unpublished analysis of the southwest

66

S. D. CAIRNS & H. ZIBROWIUS

Indian Ocean (Cairns & Keller, 1993) and New Zealand (Cairns, 1995) azooxanthellates results in similar bell¬

shaped curves, but in the case of the southwest Indian Ocean (N = 97), peaking at a shallower depth of 100-200 m

(55%), and for the New Zealand fauna (N=104) at a greater depth of 400-600 m (54%). The bell-shaped maximum

at various depths in different parts of the world is probably correlated with regional water characteristics, especially

temperature.

Although the depth record for a scleractinian coral is 6328 m (KELLER, 1976), only 4 species are known from

the Philippine-Indonesian region at depths slightly in excess of 2000 m: Flabellum conuis (1160-2570 m),

Caryophyllia cornulum (1525-2350 m), Stephanocyathus regius (563-2160 m), and Enallopsammia rostrata (417-

2021 m).

SYSTEMATIC ACCOUNT

Order SCLERACTINIA

Suborder ASTROCOENIINA

Family POCILLOPORIDAE Gray, 1842

Genus MADRAC1S H. Milne Edwards & Haime, 1849

Madracis asanoi Yabe & Sugiyama, 1936

Figs 1 a-d

Madrasis (sic) asanoi Yabe & Sugiyama, 1936: 349, figs 4-4a.

Madracis palaoensis Yabe & Sugiyama, 1936: 349, figs 5-5a; 1941: 71, pi. 62, figs 1-1 a (new synonym).

Madracis asanoi - Yabe & Sugiyama, 1941: 71, pi. 61. fig. 4, pi, 62, fig. 2. — ? Eguchi, 1968: Cl 1.

Material EXAMINED. — Philippines. "Albatross"', stn 5277, 1 branch (USNM 96671). — Sin 5381, 1 branch

(USNM 96672).

Type Locality. — West channel of Pelau barrier reef, Pelau, 160 m.

Description. — Corallum ramose, sparsely branched in 3 dimensions; distal branches blunt, 3. 5-4.0 mm in

diameter. Calices circular to slightly elliptical; GCD from 1.07-/. 62-2. 04, with large calices directly adjacent to

some of the smallest. Calices at branch tips relatively close to one another (e.g., 0.3 mm apart), this intercalicular

distance increasing to 1.0- 1.5 mm on large-diameter branches. Coenosteum covered with short (0.15-0.17 mm in

height) spines arranged in rows, 1 row occurring between closely packed distal calices and 3 or 4 rows between

spaced out calices on larger-diameter branches.

Ten primary septa 0.20-0.25 mm exsert and 0.25-0.30 mm in width, their inner edges fused to the massive

columella. In some larger calices traces of secondary septa occur in some interseptal chambers formed by the

primary septa, these secondaries irregular in development, quite small, and having dentate inner edges. Columella

a large dome-shaped structure occupying most of calice and rising slightly above coenosteal level. In centre of

dome is a wedge-shaped columellar lamella that rises as high as primary septa.

Remarks. — Madracis asanoi is similar to M. interjecta Marenzeller, 1907, which is known only from the

Red Sea, but differs in having calices of very unequal size — the calices of M. interjecta being of a uniform size.

M. asanoi is distinguished from M. kauaiensis Vaughan, 1907 (Hawaiian Islands) by having thicker and blunt

terminal branches; a shallower fossa; and nonuniform calicular sizes. The record of M. asanoi from Japan (EGUCHI,

1968) is queried because of its delicate, attenuate branching; deep calicular fossa; and large calices (up to 3 mm).

Madracis palaoensis is considered to be synonymous with M. asanoi given the similarity evident in their re¬

spective illustrations; the differences cited by Yabe & Sugiyama (1936) are considered as intraspecific variation.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

67

DISTRIBUTION. — Philippines: Lubang Island; Ragay Gulf, Luzon; 146-161 m. Elsewhere: Palau; ? Sagaini

Bay, Honshu; 1 10-183 m.

Madracis sp. cf. M. pharensis (Heller, 1868)

Figs 1 g-h

Madracis sp. - Wells, 1954: 414, pi. 99, fig. 5.

Madracis pharensis - WELLS, 1983: 224, pi, 16, figs 1, 5-6.

Madracis sp. cf. M. pharensis - Cairns, 1991: 6, pi. 1, figs b-e, pi. 11, fig. g (synonymy). — Gardiner & Waugh,

1939 : 229.

Madracis sp. A - Cairns, 1994: 36-37 (in part: USNM 88378, not pi. 13, figs c-f).

MATERIAL EXAMINED. — Philippines. "Albatross": sin 5162, 1 (USNM 96674). — Stn 5268, 1 (USNM 96675).

Musorstom 3: stn 1 17, 1 (USNM 96676). — Stn 131, 1 (MNHN). — Sin 134, 5 (MNHN).

Indonesia. Deki: stn 24, 1 (ZMUC).

Fiji. Korolevu, 27 m, 10 colonies (USNM 96673).

Gulf of California. Las Animas, La Paz, Gulf of California, 12 m, 1 colony (USNM 93920).

Type Locality. — Hvarski Kanal, Adriatic Sea, 36 m.

DIAGNOSIS. — Coralla exclusively encrusting, forming relatively small colonies (usually less than

20 corallites) shaped as: thin, irregular masses; stoloniferous ribbons; or small nodules. Perimeter of colonies

usually enclosed by a smooth, low epithecal lip. Corallites polygonal in shape and 1.5-2. 5 mm in diameter, each

calice sharing a thin (0.07-0.09 mm) common wall with its adjacent calices. Intercalicular walls bear tall (up to

0.2 mm) triangular spines, one corresponding to each of the 20 septa in corallite.

Septa decamerally arranged in 2 cycles (20 septa), the 10 primary septa rising as high as the costal spines, but

their outer edges separated from the costal spines by a deep notch. Inner edges of primary septa smooth and

vertical, each primary bearing a discrete paliform lobe; however, paliform lobes of 5 primary septa are wider and

taller than the lobes of the other 5 alternate primaries, the larger lobes being 0.12-0.15 mm in width, the smaller,

0.06-0.07 mm (Fig. lh). Septal faces covered with tall (40-50 (im), blunt spines. Secondary septa 1/2 to 2/3

width of primaries and much less thick, having dentate to laciniate inner edges. Columella a massive pointed style

rising well above paliform lobes as high as the exsert septa. Columellar style varies from cylindrical to slightly

elliptical or flattened in cross section and is surrounded by a crown of 10 Pi.

Remarks. — Three species of Madracis are characterised as having exclusively encrusting coralla: M. decactis

(Lyman, 1859); M. kirbyi Veron & Pichon, 1976; and M. pharensis. M. pharensis is distinguished from the other

two by having well-developed secondary septa and a distinct ring of Pi. M. kirbyi is also reported from the

Philippines (VERON & HODGSON, 1989), but is not discussed herein because it is considered to be a

zooxanthellate.

Although no skeletal differences could be found between Atlantic and Pacific populations of Madracis reported

herein, the second author believes that characteristics of the soft parts of these populations may ultimately place

the Atlantic and Indo-Pacific populations in different species or subspecies.

Distribution. — Philippines: Verde Island Passage; Mindoro Strait; Sibuyan Sea; Sulu Sea (west of Panay

and Sulu Archipelago); 95-421 m. Indonesia: Banda Sea (Kai Islands); 85-124 m. Elsewhere: Marshall Islands;

Galapagos; Fiji; Gulf of California; 5-343 m.

Madracis sp. A

Figs 1 e-f

Madracis asperula ? - Gardiner & Waugh. 1939: 229. — Durham & Barnard, 1952: 14-15. pi. 1, figs 2 a-b. — Wells,

1983: 224. [Not Madracis asperula H. Milne Edwards & Haime, 1849].

Madracis sp. cf. M. asperula - Cairns, 1991: 5-6, pi. 1, fig. a.

68

S. D. CAIRNS & H. ZIBROWIUS

Madracis sp. A - Cairns & Keller, 1993: 228-229, pi. 3, figs A-B (in part: not USNM 91499). — Cairns, 1994: 36-37,

pi. 13, figs c-f (in part: not USNM 88378).

MATERIAL EXAMINED. — Philippines. " Albatross stn 5217, 3 branches (USNM 96679). — Stn 5311,

1 branch (USNM 96680). — Stn 5398, 2 branches (USNM 96681).

Indonesia. Corindon 2: stn 248, 1 branch (USNM 96684).

KARUBAR: stn 22, 1 fragment (POLIPI). — Stn 29, 1 fragment (MNHN).

Diagnosis. — Corallum ramose, sparsely branched in all directions; distal branches slender and attenuate,

about diameter of a calice. Calices circular to slightly elliptical, 1.75-2.5 mm in GCD, and usually separated by

0.4-0. 5 mm of coenosteum, although calices more crowded near branch tips. Coenosteum spinose. Septa

decamerally arranged in 2 cycles (20 septa). The 10 primary septa are exsert, having spinose faces, each bearing a

small axial paliform lobe. The 10 secondary septa are rudimentary, each expressed as a row of small spines. Fossa

shallow. Columella consists of a central, circular platform from which a pointed, laterally compressed style

projects. 10 Pi encircle the columellar style at the edge of the circular platform.

Remarks. — Madracis sp. A was compared to M. asperula H. Milne Edwards & Haime, 1849 (type locality:

Madeira), primarily because both species have slender branches and decameral symmetry; however, Madracis sp.

A consistently differs in having rudimentary secondary septa and a ring of small Pi. Madracis sp. A is more

similar to M. profunda Zibrowius, 1980 (northeastern Atlantic, 112-327 m), both species having slender branches,

10 primary and 10 small secondary septa. M. sp. A appears to differ in having a distinct crown of Pi. In calicular

features, Madracis sp. A is similar to M. pharensis , but differs significantly in having a ramose corallum.

DISTRIBUTION OF similar Forms. — Philippines: Ragay Gulf, Luzon; Samar Sea; 161-208 m. Indonesia:

Makassar Strait; Banda Sea (Kai Islands); 124-181 m. Elsewhere: southwestern Indian Ocean; Arabian Sea;

Galapagos; 46-274 m.

Suborder FUNGIINA

Superfamily FUNGIOIDEA Dana, 1846

Family FUNGIACYATHIDAE Chevalier, 1987

Genus FUNGIACYATHUS Sars, 1872

Subgenus FUNGIACYATHUS (FUNGIACYATHUS) Sars, 1872

Discussion. — The nominate subgenus has recently been defined (Cairns, 1989a) as including those species

that have 5 cycles of septa (96 septa), the subgenus Bathyactis reserved for those species having 4 cycles of septa

(48). However, the second author suggests that a more meaningful division of the genus would be based on the

characteristic of their septal edges, the nominate subgenus, defined by type species F. fragilis, having corrugated

septa with sinuous edges, and the subgenus Bathyactis, defined by the type species F. symmetricus, having planar

septa with straight edges. Under this suggestion, 3 Recent species would be placed in the nominate subgenus:

F. fragilis, F. stephanus, and F. pliciseptus Keller, 1981, only the last having 4 cycles of septa. The

15 remaining Recent species would be placed in the subgenus Bathyactis, all but 3 having 4 cycles of septa.

Fungiacyathus (F.) stephanus (Alcock, 1893)

Bathyactis symmetrica - Moseley, 1881: 189 (in part: "Challenger" stn 194). — ALCOCK, 1902c: 37 (in part: "Siboga"

stn 12 and 18). [Not Fungia symmetrica Pourtales, 1871].

Bathyactis stephanus Alcock, 1893: 149, pi. 5, figs 12, 12a; 1902c: 38.

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

69

Bathyactis sibogae Alcock, 1902a: 108 (in part); 1902c: 38 (in part: "Siboga" stn 95 and large specimen of 57 mm

GCD).

Fungiacyathus (F.) stephanus - Zou et al„ 1988: 195. — Cairns. 1989a: 7-9, pi. 1, figs a-k, pi. 2, figs a-b (synonymy);

1994: 37. pi. 13, figs g-i; 1995: 31-32, pi. 1, figs a-c. — Cairns & Keller. 1993: 230.

MATERIAL EXAMINED. — Philippines. "Siboga": stn 95, 1 with flat base (ZMA).

Musorstom 1: stn 49, 1 with flat base (MNHN). — Stn 54, 1 with flat base (USNM 96694).

MUSORSTOM 2: stn 24, 17 with concave bases (MNHN). — Stn 25, 26 with concave bases (MNHN).

Musorstom 3: stn 116, 20 with concave bases: 5 (MNHN), 15 (USNM 96696).

Indonesia. "Hakuho Maru": stn KH72-1-26, 4 with flat bases (USNM 96698).

DEKI: stn 49, 1 (NNM 22472).

CORINDON 2: stn 286, 1 with flat base (MNHN).

Karubar: stn 12, 2 with flat bases (USNM 96686). — Stn 20, 3 with concave bases (POL1P1). — Stn 21, 4 with flat

bases: 3 (MNHN), 1 (POLIPI). — Stn 35, I with flat base (USNM 96687). — Stn 36, 2 with flat bases (MNHN). —

Stn 39, 2 with concave bases (MNHN). — Stn 56, 1 with concave base (USNM 96693). — Stn 59, 9 with fiat bases

(MNHN). — Stn 62, 2 with fiat bases (USNM 96691).

Type Locality. — "Investigator" stn 133: 15°43'30"N, 81°19,30',E (off Kristna Delta, Bay of Bengal),

1240 m.

DIAGNOSIS. — Two forms of the species occur, one having a flat base and another with a highly concave base.

Corallum large and fragile, the flat-based form up to 62 mm in diameter (Karubar stn 36); concave-based coralla

usually smaller and may have a marginal shelf. Costae consist of thin, finely serrate ridges. Septa hexamerally

arranged in 5 full cycles, the inner septal edges highly sinuous. Each Si bears 20-23 trabecular ridges on each face

and is linked to adjacent septa by 12-15 T-shaped synapticulae. Septal canopies porous and rudimentary. Small P2

usually present adjacent to columella. Columella crispate and flat.

Remarks. — Fungiacyathus stephanus is distinguished from F. paliferus (Alcock, 1902), the only other

Indonesian species having 5 cycles of septa, by having a larger, more fragile corallum; sinuous septa; and smaller

P2. It is more fully described and illustrated by Cairns (1989a, 1994).

Distribution. — Philippines: Lubang Island; Verde Island Passage; Bohol Sea; Sulu Sea (Palawan and Sulu

Archipelago); South China Sea (Palawan); 245-1335 m. Indonesia: Makassar Strait; Halmahera Sea; Banda Sea

(Kai Islands, Banda Islands, and Tukangbesi Islands); Arafura Sea (southeast of Tanimbar Islands); Timor Sea

(south of Leti Islands); Bali Sea; 245-1977 m. Elsew here: Malaysia (Celebes Sea off Sabah); southwestern Indian

Ocean; Bay of Bengal; Lord Howe Rise; Norfolk and Kermadec Ridges; Japan (Honshu, Kyushu, and Ryukyu

Islands); 446-2000 m.

Fungiacyathus (F.) paliferus (Alcock, 1902)

Bathyactis palifera Alcock, 1902a: 108; 1902c: 38, pi. 5, figs 34, 34a. — Van DER Horst, 1921: 38. — FaUSTINO, 1927:

214, pi. 71, figs 1-2.

Bathyactis symmetrica - ALCOCK, 1902c: 37 (in part: "Siboga" stn 95). — FAUSTINO, 1927: 213 (in part: "Siboga"

stn 95). [Not Fungia symmetrica PourtaRs, 1871],

Bathyactis kikaiensis Yabe & Eguchi, 1942b: 138, 155-156, pi. 12. figs 6-7.

Fungiacyathus (F.) paliferus - Cairns, 1989a: 9-10, pi. 2, figs c-i, pi. 3. figs a-c (synonymy); 1994: 37-38, pi. 14,

figs a-e. — Cairns & Parker, 1992: 6-7, pi. 1, figs a-b. — Cairns & Keller, 1993: 230.

Not Bathyactis palifera - Latypov, 1986: 266, 268 [probably a juvenile fungiidj.

Not Fungiacyathus palifera - KELLER, 1976: 33-34, pi. 1, figs 1-2 [= Fungiacyathus sp., see CAIRNS, 1994].

Material examined. — Philippines. Musorstom 3: stn 102, 2 (USNM 96700). — Stn 130, 2 (MNHN). —

Stn 131, 2 (MNHN).

Indonesia. "Siboga": stn 49a, 1 (ZMA Coel. 706).

Deki: stn 2. 1 (NNM 22475). — Stn 3, 2 (NNM 22476). — Stn 5, 1 (NNM 22477). — Stn 6, 1 (NNM 22478). —

Stn 24, 1 (NNM 22479). — Stn 42, 1 (NNM 22482). — Stn 49, 4 (NNM 22481). — Stn 58, 1 (NNM 22483). — Stn 63,

1 (NNM 22482).

70

S. D. CAIRNS & H. ZIBROWIUS

CORINDON 2: stn 210, 1 (MNHN). — Sin 275, 1 irregular fragment (MNHN).

SNELLIUS 2: stn 4.019, 2 (NNM 22529). — Stn 4.033, 3 (NNM 22528). — Stn 4.034, 2 (NNM 22484).

Karubar: stn 2, 1 (MNHN). — Stn 3, 10 irregular fragments (MNHN). — Stn 7, 15 irregular fragments (MNHN). —

Stn 15, 6 irregular fragments (MNHN). — Stn 18, 1 (USNM 96699). — Stn 28, 2 irregular fragments (MNHN). — Stn 49,

2 irregular fragments (MNHN).

Type Locality. — Sulu Sea and Moluccas, 141-350 m.

Diagnosis. — Corallum robust, up to 21 mm in calicular diameter, having a slight to only slightly concave

base covered with rounded, granular costae. 5 cycles of hexamerally arranged, straight, planar septa. Each Si bears

15-20 trabecular ridges on each face and is linked to its 2 adjacent septa by 9-14 solid synapticulae. Septal canopies

porous and rudimentary. Well-developed P2 and sometimes even P3 present. Columella crispate and rudimentary.

Remarks. — Comparisons to the other species in this subgenus known from the Indonesian region,

F. stephanus (Alcock, 1893), are made in that account and by CAIRNS (1989a, table 1). F. paliferus is more fully

described and illustrated by Cairns (1989a, 1994) and Cairns & Parker (1992).

Small (up to 5 mm CD) irregularly regenerated specimens thought to be F. paliferus were obtained from

Karubar stations 3, 7, 15, 28, and 49.

Distribution. — Philippines : Lubang Island; Sulu Sea (west of Panay and Sulu Archipelago); 90-522 m.

Indonesia : Makassar Strait; Halmahera Sea; Banda Sea (Kai and Tukangbesi Islands; Teluk Bone, Sulawesi); Flores

Sea (Sumbawa); 69-530 m. Elsewhere : Malaysia (Celebes Sea off Sabah); southwestern Indian Ocean; Great

Australian Bight; Japan (Honshu, Korea Strait, and Ryukyu Islands); 70-823 m.

Subgenus FUNGI ACYATHUS (BATHYACTIS) Moseley, 1881

Fungiacyathus (B.) sibogae (Alcock, 1902)

Bathyactis sibogae Alcock, 1902a: 108; 1902c: 38 (in part: "Siboga" stn 175). — FAUSTINO, 1927: 214 (in part:

"Siboga" stn 175). — Van Soest, 1979: 109 (in part: "Siboga" stn 175), pi. 2, figs 1-2.

Bathyactis symmetrica - ALCOCK, 1902c: 37 (in part: "Siboga" stn 208). (Not Fungia symmetrica PourtalLs, 1871],

Bathyactis stabilis Gardiner & Waugh, 1939: 231-232, text-figs 1-2.

Fungiacyathus sibogae - Wells, 1977: 7. — Cairns, 1989a: 10-11, pi. 3, figs d-k, pi. 4, figs a-c. — Cairns & Keller,

1993: 229-230 (synonymy).

Material EXAMINED. — Indonesia. Deki: stn 50, 1 (NNM 22474). — Stn 57, 5 (NNM 22473).

Type Locality. — "Siboga" stn 175: 2°37.7'S, 130°33.4'E (Ceram Sea), 1914 m.

Diagnosis. — Corallum fragile, up to 14.5 mm in GCD; base flat, covered with narrow ridged, serrate costae.

Septa hexamerally arranged in 4 cycles of planar septa, each Si composed of 14-19 trabeculae, the innermost 5 or 6

forming tall exsert spines. Synapticular plates T-shaped, approximately 6 per Si. Columella papillose and rounded.

Remarks. — It was previously thought that 2 of the paralectotypes of Bathyactis sibogae , those from

"Siboga" stn 95 and 159, were missing (Cairns, 1989a: 11); however, in 1994 the specimen from "Siboga"

stn 95 was found in the collections of the ZMA (Coel. 5098). It was mounted on a board along with a specimen

of Flabellum japonicum in a bottle of preservative, filed under the name of the latter species. It is the flat-based

form of Fungiacyathus stephanus and measures about 48 mm in diameter. F. sibogae is more fully described and

illustrated by Cairns (1989a), which also includes a key to the species from this region.

Distribution. — Indonesia : Molucca Sea; Ceram Sea; Banda Sea (Kai Islands and southeast of Sulawesi);

200-1914 m. Elsewhere: southwest Indian Ocean; 463-1948 m (CAIRNS & KELLER, 1993).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

71

Fungiacyathus (B.) granulosus Cairns, 1989

Bathyactis symmetrica - Alcock, 1902c: 37 (in part: "Siboga" stn 59). [Not Fungia symmetrica Pourtales, 1871].

Fungiacyathus (B.) granulosus Cairns, 1989a: 11, pi. 4, figs d-i; 1994: 39, pi. 15, figs d-e.

MATERIAL EXAMINED. — Philippines. Musorstom 1: stn 43, 5 (USNM 96705). — Stn 44. 13 (MNHN). —

Sin 50, 1 (MNHN).

MUSORSTOM 2: stn 25, 20 (USNM 96707). — Stn 82, 14 (MNHN).

Musorstom 3: stn 106, 6 (MNHN).

Indonesia. Karubar: stn 3, 1 (USNM 96708). — Stn 7, 7 + 8 juveniles (USNM 96702). — Sin 40, 3 (USNM

96704). — Stn 56. 1 (POLIPI). — Stn 58, 1 (MNHN). — Stn 59, 2 (MNHN).

Type Locality. — "Albatross" stn 5590: 4°10'50"N, 1 18°39'35"E (off Sabah, Celebes Sea), 567 m.

Diagnosis. — Corallum robust, up to 24.5 mm in calicular diameter (Karubar stn 40), with a flat to

slightly concave base covered with rounded, granular costae. Septa hexamerally arranged in 4 cycles, each Si

having 21-24 coarsely dentate trabecular ridges on each face and linked to its adjacent septa by 6-9 Y-shaped

synapticulae. Septa planar. Septal canopies porous, but well developed. Paliform lobes absent, but well-developed

trabecular spines present on all septa. Columella large and tuberculate.

Remarks. — As noted by Cairns (1989a), F. granulosus differs from F. sibogae in having a granular base

and beaded costae that are rounded to triangular in cross section, not covered with thin, serrate ridges as in

F. sibogae. Also, F. granulosus attains a larger diameter (24 mm vs 15 mm) and has more robust inner trabecular

spines, especially those of the S2- F. granulosus is more fully described and figured by Cairns (1989a).

Distribution. — Philippines-. Lubang Island; Verde Island Passage; Tablas Strait, Mindoro; 428-640 m.

Indonesia: Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar Islands); Savu Sea (Timor); 287-567 m.

Elsewhere: Malaysia (Celebes Sea off Sabah); northern Ryukyu Islands; 402-410 m.

Fungiacyathus (B.) variegatus Cairns, 1989

Fungiacyathus fragilis - Wells, 1984: 205-206 (in part: USGS 24918, pi. 1. figs 1-2). [Not Fungiacyathus fragilis

G.O. Sars, 1872],

Fungiacyathus (B.) variegatus Cairns, 1989a: 11-12, pi. 5, figs a-h; 1994: 38-39, pi. 15, figs a-b.

Material EXAMINED. — Philippines. Musorstom 1: stn 13, 2 (MNHN). — Stn 14. 3 (MNHN). — Stn 15,

4 (MNHN). — Stn 25, 1 (MNHN).

Musorstom 2: stn 4, 1 (USNM 96710). — Stn 12, 1 ((USNM 96711). — Stn 64, 2 (MNHN). — Stn 66, 1 (MNHN).

Musorstom 3: stn 87, 3 (MNHN). — Stn 88, 1 (USNM 96713). — Stn 95, I (USNM 96714). — Stn 96. 1 (USNM

96715). — Stn 97, 2 (MNHN). — Stn 99. 7 (MNHN). — Stn 100, 7 (USNM 96718). — Stn 101. 21: 5 (MNHN),

16 (USNM). — Stn 108, 1 (USNM 96720). — Stn 111, 3: 2 (MNHN), 1 (USNM 96721). — Stn 112, 2 (MNHN). —

Stn 121, 1 (MNHN). — Stn 140, 3 (USNM 96722).

Indonesia. Deki: stn 24, 1 (NNM 22485). — Stn 48, 1 (NNM 22486). — Stn 50, 1 (ZMUC), 6 (NNM 22487). —

Stn 63, 8 (NNM 22488).

"Galathea": stn 480, 9 (ZMUC).

Snellius 2: stn 4.174, 1 (NNM 22490).

Karubar: stn 31, 1 (POLIPI). — Stn 76, 1 (MNHN).

Type Locality. — "Albatross" stn 5113: 13°52'N, 120°51'E (Verde Island Passage, Luzon), 291 m.

Diagnosis. — Corallum delicate, up to 10.3 mm in calicular diameter (Karubar stn 76), with a flat to

slightly concave base. Centre of base granular, but edge of base covered with thin, serrate costal ridges. Septa

hexamerally arranged in 4 cycles, each Si bearing 15-17 low dentate trabecular ridges per face and linked to their

72

S. D. CAIRNS & H. ZIBROWIUS

adjacent septa by 3 or 4 solid synapticulae. Septa planar. Septal canopies well developed, solid, and inclined.

Paliform lobes absent, but Si trabecular spines well developed. Columella rudimentary.

Remarks. — Fungiacyathus variegatus is distinguished by its relatively small size and its broad, solid

canopies that unite the inner edges of the S3 to their common S2, and the S4 to their common S3. In well-

preserved coralla Si -2 are pigmented a dark brown, but this pigmentation was present on only about 1/3 of the

specimens listed above, the colour apparently fading after the death of the coral. Small coralla (< 4 mm GCD) are

stellate in shape, their Si -2 projecting beyond the otherwise circular calicular perimeter. This species is more fully

described and illustrated by Cairns (1989a).

DISTRIBUTION. — Philippines: Lubang Island; Verde Island Passage; Sibuyan Sea; Visayan Sea; Sulu Sea

(Semirara Islands); 84-333 m. Indonesia: Banda Sea (Kai Islands); Arafura Sea (south of Tanimbar Islands); Flores

Sea (Selayar Island, Sulawesi); Bali Strait; 100-440 m. Elsewhere: South China Sea (Hong Kong); Japan (Kyushu

and Ryukyu Islands); 422-715 m. Pleistocene of Vanuatu (Wells, 1984).

Fungiacyathus (B.) turbinolioides Cairns, 1989

Fungiacyathus (B.) turbinolioides Cairns, 1989a: 12-13, pi. 6, figs a-g.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5424. 1 (USNM 96724).

"Galathea": stn 490, 1 (ZMUC).

"Hakuho Mart,": stn KH72-1-20, 1 (USNM 96723).

TYPE Locality. — "Albatross" stn 5586: 4°06’50"N, 1 18°47'20"E (off Sabah, Celebes Sea), 635 m.

DIAGNOSIS. — Corallum robust, relatively small (up to 10.3 mm calicular diameter), with a flat to slightly

concave base covered with rounded granular costae. Costae separated by deep intercostal furrows. Septa hexamerally

arranged in 4 cycles, each Si bearing 20-25 low, serrate trabecular ridges and joined to adjacent septa by 3 or

4 T-shaped synapticulae. Septal edges straight. Septal canopies absent. No paliform lobes, but trabecular spines

robust. Columella well developed, tuberculate.

Remarks. — Fungiacyathus turbinolioides is distinguished from all other species in the genus by having deep

intercostal furrows, which suggests a resemblance to a turbinoliid. It is more fully described and illustrated by

Cairns (1989a).

DISTRIBUTION. — Philippines: Sulu Sea (Cagayan Islands and Sulu Archipelago); 514-622 m. Indonesia: Java

Sea; 570-635 m. Elsewhere: Malaysia (Celebes Sea off Sabah); South China Sea (northeast of Pratas Island);

622 m.

Fungiacyathus (B.) fissidiscus sp. nov.

Figs 2 a-d

MATERIAL EXAMINED/TYPES. — Indonesia. Karubar: stn 3, 8 paratypes (MNHN). — Stn 7, holotype and

140paratypes (MNHN); 31 paratypes (POLIPI); 99 paratypes (USNM 96725).

Type Locality. — Karubar stn 7: 5°47'35"S, 132°20'39"E (Kai Islands, Banda Sea), 282-287 m.

ETYMOLOGY. — The species name fissidiscus (Latin findere, to split + discus, a circular plate) refers to the

tendency of this species to reproduce by fragmentation. The name is considered as a noun in apposition.

Description. — Corallum reproduces primarily by fragmentation (see Remarks), resulting in wedge-shaped

pieces of 6-12 septa that ultimately regenerate into a roughly circular corallum containing 48 septa. Largest

completely regenerated specimen (holotype) 4.9 mm in diameter and 1.7 mm in height. Base flat, often featureless

at centre, but toward peripheral edge formed into well-defined, rounded, granular costae. Costae up to 0.27 mm

wide, separated by deep intercostal furrows about 0.05 mm wide.

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACT1NIA

73

Septa hexamerally arranged in 4 cycles. Si extend to centre of fossa, each septum bearing 8-10 trabecular

spines, the innermost spines short, flattened, and curved inward toward fossa; the outer spines are much more

robust and cylindrical in cross section (0.20 mm in diameter). S2 similar to Si in shape, but smaller, extending

only 3/4 distance to centre of fossa, bearing only 7-9 trabecular spines that are less robust than those on the Si.

S3 smallest septa, extending only 1/5 distance to centre of fossa and quite low, bearing only 2 or 3 small

trabecular spines. S4 dimorphic in size, those adjacent to Si being almost as large as Si, bearing 7-9 trabecular

spines, the inner edges of each pair within a system fusing near centre of fossa. S4 adjacent to S2 much smaller

than those adjacent to Si and only marginally larger than the S3, i.e., extending 1/4 distance to centre of fossa,

bearing 4 or 5 trabecular spines. All septa have straight upper edges and highly granular faces. Face granules up to

80pm in height and occasionally bifid, the granules aligned in low ridges only at upper septal edges along

trabecular spines. Approximately 5 solid synapticular plates connect each Si to its adjacent S4, the 4th plate from

corallum centre being the tallest, rising almost to the upper edge of S4. Only 2 synapticular plates connect S2 to

their adjacent S4, and no synapticulae were noted on the S3. Fossa shallow; no columella.

Remarks. — The lack of synapticulae between S3 and their adjacent S4 creates radial lines of weakness

throughout a corallum that are presumed to facilitate the fragmentation process.

Three other species of Fungiacyathus are characterised by having highly regenerative coralla: F. crispus

(Pourtales, 1871): Atlantic, 183-1010 m; F. fissilis Cairns, 1984: Hawaiian Islands, 212-503 m; and F. demand

Cairns & Parker, 1992: South Australia, 190-770 m. F. fissidiscus is distinguished from all 3 in having

intercostal furrows, granular costae, and dimorphic S4. F. fissidiscus is perhaps more similar to F. turbinolioides

Cairns, 1989 (see above), both species having a similar costal structure and septal ornamentation; however, it

differs in having dimorphic S4, a smaller corallum, a regenerative corallum, and in lacking a columella.

Distribution. — Indonesia: Banda Sea (Kai Islands); 282-287 m.

Family MICRABACIIDAE Vaughan, 1905

Genus LEPTOPENUS Moseley, 1881

Leptopenus sp. A

Figs 2 e-f

Leptopenus sp. A - Cairns, 1989a: 14-15, pi. 7a-f.

Material EXAMINED. — Indonesia. Karubar: stn 7, 1 fragment (MNHN).

Remarks. — One small specimen, a wedge-shaped fragment measuring 3.3 mm in calicular radius, is reported

herein. It appears to be the same species described by Cairns (1989a) from the Celebes Sea. Based on this small

fragment, nothing can be added to the previous description. Four other fragments, too small to identify, are also re¬

ported from Karubar stations 3 (USNM 96728) and 15 (USNM 96729). It is noteworthy that these specimens

represent the shallowest record for Leptopenus (i.e., 214-300 m), a genus customarily found at depths in excess of

2000 m.

Distribution. — Indonesia : Banda Sea (Kai Islands). Malaysia: Celebes Sea (off Sabah); 287-871 m.

Genus LETEPSAMMIA Yabe & Eguchi, 1932

Letepsammia formosissima (Moseley, 1876)

Stephanophyllia formosissima Moseley, 1876: 561-562; 1881: 201-204, pi. 4, fig. 11. pi. 13, figs 6-7, pi. 16.

}&*->• Alcock, 1902c: 39 (in part: "Siboga" stn 95). — BOSCHMA, 1923: 144-145, pi. 10. fig. 31. — Faustino,

1927: 244-245, pi. 77, figs 7-8.

74

S. D. CAIRNS & H. ZIBROWIUS

Letepsammia formosissima - Owens, 1986b: 486-487. — Cairns, 1989a: 15-18, pi. 6, fig. j, pi. 7, figs g-i, pi. 8,

figs a-d (synonymy); 1995: 36-37, pi. 3, figs f-g. — Cairns & Parker, 1992: 8-9, pi. 1, figs f, h.

Not Stephanophyllia formosissima var. - ALCOCK, 1902c: 39-40 [= Rhombopsammia squiresi Owens, 1986].

Not Letepsammia formosissima - CAIRNS & Keller, 1993: 230-231. pi. 3. fig. D [= L franki Owens, 1994], — Cairns,

1994: 40-41. pi. 15, figs c, f [= L. superstes (Ortmann, 1888)].

MATERIAL EXAMINED. — Philippines. Musorstom 1: stn 2, 1 (USNM 96568). — Stn 3, 1 (USNM 96579).

Sm 4. 1 (USNM 96557). — Stn 9, 6 (MNHN). — Stn 10, 9 (MNHN). — Sin 12, 9 (MNHN). — Stn 14, 1 (USNM 96555).

— Stn 20, 3 (USNM 96567). — Stn 24, 18 (MNHN). — Stn 25, 17 (MNHN). — Stn 35, 2 (MNHN). — Stn 61, 57:

1 (MNHN), 56 (USNM 96556). — Stn 62, 8 (USNM 96562). — Stn 64, 1 (MNHN).

Musorstom 2: stn 1, 16 (MNHN). — Stn 2, 27 (MNHN). — Stn 4. 3 (USNM 96551). — Stn 6. 1 (USNM 96566). —

Stn 10 40- 29 (MNHN), 11 (USNM 96575). — Stn 11, 6 (USNM 96553). — Stn 12, 9 (MNHN). — Stn 13, 4 (USNM

96552) — Stn 15. 1 (MNHN). — Stn 18, 11: 6 (USNM 96576), 5 (BMNH 1992.8.11.12). — Stn 33, 6 (MNHN). -

Stn 62. 4 (?). — Stn 63, 3: 1 (MNHN), 2 (USNM 96550). — Stn 64, 22 (USNM 96559). — Stn 66, 7 (USNM 96573). —

SU1 Mu SORSTOM^ 3 / stn 86, 13 (MNHN). — Stn 87, 15 (USNM 96570). — Stn 88, 42 (MNHN). — Stn 90, 8 (MNHN). —

Stn 91, 31 (MNHN). — Stn 92, 5 (MNHN). — Stn 96, 116: 63 (MNHN), 46 (USNM 81879), 7 (BMNH 1992.8.1 1.1 1). —

Stn 97 39' 23 (MNHN), 16 (USNM 96565). — Stn 98, 36: 6 (MNHN), 30 (USNM 96563). — Stn 99, 28: 17 (MNHN),

1 1 (USNM 96549). — Stn 100. 34: 19 (MNHN), 15 (USNM 96548). — Stn 101, 25 (MNHN). — Stn 102, 27 (MNHN).

— Stn 103 15- 8 (MNHN), 7 (USNM 96571). — Stn 107, 1 (MNHN). — Stn 108, 62: 17 (MNHN), 45 (USNM 81877). —

Stn 109, 78: 54 (MNHN), 17 (USNM 81878). 7 (BMNH 1992.8.1 1.13). — Stn 1 10, 2 (MNHN). — Stn 111. 17 (MNHN).

— Stn 112, 20 (USNM 96561). — Stn 124, 1 (MNHN). — Stn 126, 3 (MNHN). — Stn 131, 5 (MNHN). — Stn 139,

1 (MNHN). — Stn 143, 81 (USNM 81878).

Indonesia. Deki: stn 2. 1 (NNM 22502). — Stn 3, 7 (NNM 22501). — Stn 4. 1 (NNM 22503). — Stn 41, 25 (NNM

22504). — Stn 42, 1 (NNM 22505). — Stn 44, 1 (NNM 22506). — Stn 46. 2 (NNM 22507). — Stn 49, 2 (NNM 22509).

— Stn 58, 2 (NNM 22511). — Stn 63, 4 (NNM 22512).

"Hakuho Mam": stn KH72-1-28, 1 (USNM 96580).

Snellius 2: stn 4.057, 1 (NNM 22513).

Karubar: stn 2, 9 (MNHN). — Stn 3, 8 (USNM 96577). — Stn 7, 5: 1 (MNHN), 4 (USNM 96578). — Stn 35,

1 (MNHN). — Stn 36, 1 1 (MNHN). — Stn 67, 2 (POLIPI). — Stn 85, 2 (POL1PI). — Stn 86, 8: 2 (MNHN), 6 (USNM

96569).

South China Sea. "Hakuho Maru": stn KH72-1-50, 2: 1 (USNM 96581), 1 (ORI).

Type Locality. — Philippines and Indonesia, 174-236 m.

Diagnosis. — Discoidal corallum up to 50.8 mm in GCD (Karubar stn 35); base flat to slightly convex;

D:H up to 4.9 in large specimens. Thin (0.06-0.07 mm), ridged costae bear very small teeth or short spines,

producing a finely serrate edge; intercostal region quite wide (3-6 times costal width) and porous, the synapticular

bars connecting each costa to its 2 alternating, adjacent septa clearly visible in basal view through intercostal

region. A low, marginal shelf, up to 4 mm wide, present on large, well-preserved specimens. Septa arranged in

typical micrabaciid fashion (Cairns, 1989a, text-fig. 2), attaining the 120-septa stage at a GCD of 17-20 mm and

often maintaining this number; however, a large syntype of GCD 38 mm has 144 septa, and the largest known

specimen of GCD 51 mm has 228 septa. Si independent and unbranched, having a smooth upper, inner edge, but a

spinose peripheral edge. Si of small specimens highly porous, but as corallum increases in size they develop a

more solid, lamellar upper, inner edge — retaining their porosity only on their lower half near the base. S2 also

unbranched but not independent, a pair of S3 fusing to each S2 near the columella. Each S3 bifurcates repeatedly,

producing the majority of the septa. Columella elongate, spongy, and often densely fused.

Remarks. — Letepsammia formosissima was the coral most commonly collected on the MUSORSTOM cruises,

taken at 55 stations ranging from 1 15-390 m. A more complete description and illustrations of this species are

given by Cairns (1989a) and Cairns & Parker (1992), and a comparison to Rhombopsammia niphada is given

in the account of that species.

Another species, L. franki Owens, 1994, occurs in the southwestern Indian Ocean where it had occasionally

been ascribed to L. formosissima (see Cairns, 1989a; Cairns & Keller, 1993; Owens, 1994). L. franki differs

from the latter by its papillose columella and coarse septal dentation that give the corallum a distinctly beaded

appearance.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

75

DISTRIBUTION. — Philippines: common from Lubang Island to the Bohol Sea; 1 15-390 m. Indonesia: Banda

Sea (Kai Islands); Arafura Sea (south of Tanimbar Islands); Timor Sea (south of Leti Islands); Savu Sea (west of

Timor); 154-390 m. Elsewhere: western Pacific to Hawaiian Islands, including South China Sea (Charlotte Bank);

97-457 m.

Letepsammia superstes (Ortmann, 1888)

Stephanophyllia superstes Ortmann, 1888; 160-161, pi. 6, fig. 5. — Owens, 1986a: 487.

Stephanophyllia (Letepsammia) japonica Yabe & Eguchi, 1934a; 281, figs 1-3; 1942b: 156-157, pi. 12, figs 8a-c.

Letepsammia formosissima forma superstes - CAIRNS, 1994: 40, pi. 15, figs c. f.

Letepsammia superstes - Cairns, 1995: 34-35, pi. 2, figs f-i (synonymy).

Not Stephanophyllia (Letepsammia) japonica - Zou, 1988: 75, pi. 5, fig. 7 [= Stephanophyllia neglecta Boschma,

1923].

Material EXAMINED. — Philippines. Musorstom 2: stn 32, 3 (USNM 96733).

Indonesia. Karubar: stn 1, 12 (MNHN). — Stn 7, 1 (USNM 96730). — Stn 18, 1 (USNM 96731). — Stn 50,

1 (USNM 96732).

Type Locality. — Sagami Bay, Japan, 183-366 m.

Remarks. — Although collected on the same type of bottom and often together mixed with the more common

L. formosissima , L. superstes differs in having a smaller corallum with fewer septa. The largest known specimen

(MUSORSTOM 2 stn 32) is only 22 mm in calicular diameter and has 96 septa, a size at which L. formosissima

would have 120 septa. Although similar to L. formosissima, L. superstes differs in having a papillose columella,

more robust trabecular spines, and a tendency to have a patellate (vs a flat based) corallum. The species is more

fully described by Cairns (1994) as L. formosissima forma superstes and by Cairns (1995).

DISTRIBUTION. — Philippines: Verde Island Passage; 192-220 m. Indonesia: Banda Sea (Kai Islands); Arafura

Sea (east of Tanimbar Islands); 185-282 m. Elsewhere: Japan (Honshu and northern Ryukyu Islands); Korea Strait;

South China Sea (Hong Kong); Kermadec Ridge; 77-710 m. Pleistocene of Ryukyu Islands.

Genus RHOMBOPSAMMIA Owens, 1986

Rhombopsammia niphada Owens, 1986

Rhombopsammia niphada Owens, 1986a: 252-255, figs 2b, 3a-d. — Cairns 1989a: 19-20, pi. 9, figs d-i, pi. 10.

figs a-b, text-fig. 2 (synonymy); 1994: 41, pi. 15, figs i-k, pi. 16, fig. e.

Material examined. — Philippines. Musorstom l: stn 47, 1 (USNM 96738).

Musorstom 2: stn 25, 1 (MNHN).

Musorstom 3: stn 1 16, 1 (MNHN).

Indonesia. Deki: stn 3, 1 (NNM 22493).

Karubar: stn 20, 3 (POLIPI). — Stn 21, 15 (MNHN). — Stn 39, 14 (MNHN). — Stn 40, 15 (USNM 96735). —

Stn 59, 17 (MNHN). — Stn 70, 7: 6 (MNHN). 1 (USNM 96737). — Stn 71, 3 (MNHN). — Stn 75, 4 (MNHN).

Type Locality. — "Albatross" stn 4911: 31°38'30"N, 129°19'E (East China Sea, off Kyushu), 715 m.

Diagnosis. — Discoidal corallum up to 46 mm in diameter (Karubar stn 59); base usually Hat. Costae thin

(0.06-0.07 mm) ridges. Intercostal regions much wider (about 0.45 mm) than costae, traversed by thin

synapticulae, which produce a series of pores in each elongate space. Marginal shelf present but not wide and often

damaged. Septa arranged in typical micrabaciid fashion (Cairns, 1989a, text-fig. 2), both septa and costae up to

M 1 in number, which alternate in position. Columella elongate and spongy.

76

S. D. CAIRNS & H. ZIBROWIUS

Remarks. — The similarities between R. niphada and Letepsammia formosissima have been noted previously

(Owens, 1986a; Cairns, 1989a). R. niphada differs in having solid Si that bear faint vertical ridges (vepreculae)

on their faces, whereas the SI of L. formosissima are highly porous at a small calicular diameter and become more

solid with age, but do not bear vepreculae, and always maintain some porosity adjacent to the base. R. niphada

usually has 144 septa, whereas L. formosissima usually has 120 septa, but as many as 228. R. niphada has

well-developed septal canopies and reduced trabecular spines, whereas L. formosissima has well-developed septal

spines on the S3 and S3' and no canopies. Finally, R. niphada is characteristic of deeper water than

L. formosissima, all records of the former between 405 and 804 m, of the latter species, 97-457 m. A more

detailed description and illustrations of this species are found in OWENS (1986a) and Cairns (1989a).

DISTRIBUTION. — Philippines: Verde Island Passage; Mindoro Strait; Palawan Passage; 512-804 m.

Indonesia : Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar Islands); 405-768 m. Elsewhere: Japan

(Honshu, Kyushu, and Ryukyu Islands); 660-783 m.

Rhombopsammia squiresi Owens, 1986

Slephanophyllia formosissima var. - ALCOCK, 1902c: 39-40. [See Cairns, 1989a].

Rhombopsammia squiresi Owens, 1986a: 250-252, figs la-d, 2a. — Cairns, 1989a: 18-19, pi. 8, figs e-j, pi. 9, figs a-c

(synonymy).

MATERIAL EXAMINED. — Indonesia. Corindon 2: stn 240, 3 (MNHN).

Karubar: stn 87. 1 (MNHN), 1 (POLIPI). — Stn 89, 2 (USNM 96741).

Type Locality. — "Albatross" stn 5423: 9°38'30"N, 121°H'E (Cagayan Island, Philippines), 929 m.

REMARKS. — Previously known from only 5 specimens from 5 stations, 7 additional specimens of this rarely

collected species are reported herein from the Indonesian region, the largest (Karubar stn 89) 41.2 mm in

calicular diameter. R. squiresi is distinguished from R. niphada by its distinctive marginal shelf, wherein each

septum projects as a broad, flat, spongy mass; and by having only 96 septa.

Distribution. — Philippines: Sulu Sea (Cagayan Islands and Palawan); Iligan Bay, Bohol Sea. The

Philippine depth range is 905-1401 m, not 622-1401 as previously reported (Cairns, 1989a). Also, the paratype

mentioned by CAIRNS (1989a) from "Albatross" stn 5424, should read "Albatross" stn 5429. Indonesia: Makassar

Strait; Timor Sea (southwest of Tanimbar Islands and southeast of Timor); 675-1048 m.

Genus STEPHANOPHYLLIA Michelin, 1841

Slephanophyllia fungulus Alcock, 1902

Slephanophyllia fungulus Alcock, 1902b: 122; 1902c: 40, pi. 5, figs 35a-b. — Faustino, 1927: 245-246,

pi. 77, figs 9-11. — Zou et al., 1988: 195. — Cairns, 1989a: 21-23, pi. 10, figs c-k, pi. 11, figs a-b,

text-fig. 3 (synonymy). — Cairns & Keller, 1993: 231. — Cairns, 1994: 41-42, pi. 16, figs a-d, f-g

(synonymy).

Stephanophyllia complicata - Alcock, 1902c: 40 (in part: 1 of 3 specimens from "Siboga" stn 59). [Not

Slephanophyllia complicata Moseley, 1876].

Material EXAMINED. — Philippines. "Hakuho Maru": stn KH72-1-20, 7 (USNM 96743).

Indonesia. Deki: stn 6, 66, (NNM 22545). — Stn 49, 115 (NNM 22547). — Stn 52, 1 (NNM 22548).

SNELLIUS 2: stn 4019, 5 (NNM 22550).

Type Locality. — "Siboga" stn 100: 6°1 l'N, 120°37.5'E (Sulu Archipelago), 450 m.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

77

Diagnosis. — Corallum discoidal, with a thick, flat base and slightly upturned edges; D:H = 1. 9-2.6. Largest

known specimen (" Albatross " stn 5586, Celebes Sea) 15.6 mm in diameter. Costae flat, ornamented with a medial

row of coarse granules close to epicentre, but outward from basal centre each costa bears 2 rows of smaller

granules, one row on each edge of the costa. No marginal shelf. Adjacent septal faces fused together by massive,

elongate mural synapticulae (fulturae sensu Gill, 1979), producing a sturdy, robust corallum. 96 septa arranged in

typical micrabaciid fashion (Cairns, 1989a, text-fig. 3). Septal edges straight; septal face granules wide-based,

equilateral triangular-shaped spines. Columella massive, lenticular to rectangular in cross section, often surrounded

by additional papillae.

Remarks. — Stephanophyllia fungulus is distinguished from the other 2 Recent species in the genus by its

massive columella and its thick, upturned base. A more complete description and comparisons to the other Recent

species can be found in Cairns (1989a, table 2; 1994).

DISTRIBUTION. — Philippines: Sulu Sea (Sulu Archipelago); 450-514 m. Indonesia: Banda Sea (Kai and

Tukangbesi Islands); Savu Sea (Timor); 210-635 m. Elsewhere: Malaysia (Celebes Sea off Sabah); widespread

from southwestern Indian Ocean to Japan, including South China Sea (north of Pratas Island); 73-256 m.

Stephanophyllia neglecta Boschma, 1923

Fungia patella - VAN der Horst, 1921: 57 (in part: "Siboga" stn 260). [Not Madrepora patella Ellis & Solander, 1786],

Stephanophyllia neglecta Boschma, 1923: 144-145, pi. 10, figs 28-30, — Cairns, 1989a: 23-24, pi. 11, figs c-j

(synonymy).

Material EXAMINED. — Philippines. MUSORSTOM 2: stn 33, 1 (USNM 81866).

Musorstom 3: stn 91. 1 (USNM 81865). — Stn 102, 46 (USNM 81859). — Stn 131, 1 (USNM 81864).

Indonesia. Deki: stn 50, 1 (NNM).

Mortensen s Java-S.A. Expedition: stn 5, 54: 48 (ZMUC), 6 (USNM 96746). — Stn 6, 1 (ZMUC). — Stn 8,

2 (ZMUC).

Snellius 2: stn 4.039, 2 (NNM 22758).

Karubar: stn 1, 1 (MNHN).

Type Locality. — "Siboga" stn 260: 5°36.5’S, 132°55.2'E (Kai Islands, Banda Sea), 90 m.

Diagnosis. — Corallum discoidal, with a thin, flat to highly convex base; D:H = 2. 8-3.2. Largest known

specimen (MORTENSEN'S Java Exp. stn 5) 12.2 mm in calicular diameter and 5.1 mm in height. Costal

ornamentation as in S. fungulus. No marginal shelf. Synapticulae variable in shape: circular, elliptical, or quite

elongate. 96 septa arranged in typical micrabaciid fashion. Septal edges straight; septal face granules blunt to

clavate cylindrical spines. Columella variable, usually papillose but in some coralla lamellar.

Remarks. — Stephanophyllia neglecta is more fully described and illustrated by Cairns (1989a), who also

compared it to the other Recent species.

Distribution. — Philippines: Lubang Island; Verde Island Passage; Sibuyan Sea; Samar Sea; Tablas Strait;

Sulu Sea (Sulu Archipelago and Palawan); 49-555 m. Indonesia: Banda Sea (Kai and Tukangbesi Islands); Bali

Strait; 50-525 m.

Stephanophyllia complicata Moseley, 1876

Stephanophyllia complicata Moseley, 1876: 558-561, text-fig.; 1881: 198-201, pi. 4, fig. 12. pi. 13, figs 3-5. —

ALCOCK, 1902c: 40 (in part: "Siboga" stn 256). — Cairns, 1989a: 21, pi. 12, figs a-b; 1995: 37-38, pi. 3, fig. h.

pi. 4, figs a-e (synonymy). — Cairns & Keller, 1993: 231-232.

Stephanophyllia japonica - Wells, 1984: 207, pi. 1, figs 5-6. Not Stephanophxllia japonica Yabe & Eguchi. 1934.

78

S. D. CAIRNS & H. ZIBROWIUS

MATERIAL EXAMINED. — Indonesia. Deki: stn 6. i (NNM 22542). — Stn 50, 7 (NNM). — Stn 52, 4 (NNM

22543).

Karubar: stn 2, 51 (MNHN). — Stn 3, 46 (POLIPI). — Stn 5, 1 (USNM 96748). — Stn 7, 66 (USNM 96749). —

Stn 15, 1 (MNHN).

TYPE Locality. — "Challenger" stn 192: 5°42'S, 132°25'E (Kai Islands, Banda Sea), 236 m.

DIAGNOSIS. — Corallum discoidal, with a thin, flat base; D:H = 2. 5-2. 7. Corallum up to 18 mm in calicular

diameter. Costal ornamentation as in S. fungulus Alcock, At calicular edge of well-preserved specimens, costae

are slightly upturned, and bifid, extending about 0.5 mm beyond calicular perimeter and producing a small

marginal shelf. Synapticulae circular to elliptical in cross section. 96 septa arranged in typical micrabaciid fashion,

the S 1-2 having straight edges, but the S3 meandering toward the calicular edge. Septal faces covered with narrow-

based isosceles triangular-shaped granules. Columella a prominent, but thin lamella, sometimes divided into

papillae or sub-lamellar elements at its summit.

Remarks. — All specimens reported above are essentially topotypic, being collected very close to the type

locality in the Kai Islands. S. complicata is distinguished from the other species in the genus (see Cairns, 1989a,

table 2) by having a thin, lamellar columella, and a narrow marginal shelf. It is described in greater detail by

Cairns (1995) based on specimens from the New Zealand region.

Distribution. — Indonesia: Banda Sea (Kai Islands); 210-397 m. Elsewhere: western Indian Ocean; Norfolk

and Three Kings Ridges; 229-1 137 m.

Suborder FAV13NA

Superfamily FAVIOIDEA Gregory, 1900

Family RHIZANGIIDAE d'Orbigny, 1851

Genus CULICIA Dana, 1846

Culicia stellata Dana, 1846

Figs 3 a-b

Culicia stellata Dana, 1846: 377, pi. 28, figs 5a-d. — Nemenzo, 1976: 252, pi. 9, figs 2-3.

Culicia truncata Dana, 1846: 378, pi. 28, figs 7, 7a (new synonym).

Culicia japonica Yabe & Eguchi, 1936: 167-168, figs 1-3 (new synonym). — Cairns, 1994: 42, pi. 17, figs a-e

(synonymy).

MATERIAL EXAMINED. — Philippines. "Alpha Helix": stn 79-M140, 1 large colony (USNM 80029).

Malaysia. Kota Kinabalu, Sabah, depth unknown, 2 colonies (USNM 78565).

Type Locality. — Singapore, South China Sea (depth not given).

DESCRIPTION. — Philippine corallum consists of 50-60 corallites produced by extratentacular, reptoid budding.

Stolons linking corallites thin and flat, 1. 5-2.0 mm wide; corallites spaced 1-5 mm apart. Stolons often covered by

encrusting epifauna. Corallites circular to slightly elliptical in cross section, up to 3.8 mm in GCD, and 4.3 mm

in height. Corallites epithecate when well-preserved, with a very thin (50 pm), almost translucent, smooth upper

rim that rises above upper outer septal edges (fig. 3b).

Septa hexamerally arranged in 4 cycles, the last cycle never complete, 34-42 being the most common septal

complement. Si independent, each composed of a tall but narrow upper lobe, a vertical inner edge, and 1 or

2 laciniate teeth on its lower, inner margin. S2-3 equal in size and shape, the inner edges of the 2 S3 and 1 S2 in

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

79

each system fusing near the columella. S2-3 not lobate as the Si, but bearing 3 or 4 laciniate teeth that grade into

the columellar elements. S4 rudimentary, consisting of a discontinuous row of spines, each spine 1/3 to 1/4 height

of the adjacent lower cycle septum. Fossa of moderate depth. Columella papillose, consisting of 5-9 elements

similar in size and shape to innermost septal teeth.

REMARKS. — Dana (1846: 377) described C. stellata as having corallites "3 lines high, and 1.5 broad" and

24 septa per corallite. Since a "line" is 1/12 inch (2.12 mm), the corallites are inferred to be about 6.4 mm in

height and 3.2 mm in GCD. Examination of Dana's holotype (USNM 185) shows it to have shorter (4.3 mm

max.) and broader (to 3.8 mm) corallites, most of which have septa of the 4th cycle, up to 42 septa/corallite.

Likewise, his description of C. truncata implies corallites of 2.6 mm in height and GCD, and 24 septa; whereas

the type (USNM 183) has corallites up to 3.5 mm in GCD and none over 2 mm in height, and most having some

pairs of S4. C. truncata would appear to be a specimen of C. stellata in which the corallites remained slightly

lower than normal, possibly for an environmental reason. The description and figures of C. japonica , as well as the

Japanese specimens reported by Cairns (1994), are also consistent with C. stellata.

Distribution. — Philippines: Cebu and Bohol; 14-20 m. Elsewhere: South China Sea (Singapore; Kota

Kinabalu, Sabah); Japan (Honshu, Kyushu, Ryukyu Islands, and Korea Strait); Fiji; 5-100 m.

Family OCULINIDAE Gray, 1847

Genus MADREPORA Linnaeus, 1758

Madrepora oculata Linnaeus, 1758

Madrepora oculata Linnaeus, 1758: 798. — Zibrowius, 1974b: 762-766, pi. 2, figs 3-5 (synonymy). — Cairns, 1982:

15, pi. 3, figs 4-6 (synonymy); 1991: 9-10, pi. 2, fig. j, pi. 3, figs a-d (synonymy); 1994: 18-19, pi. 3, figs f-h

(synonymy); 1995: 41, pi. 5, figs e-f. pi. 6, figs a-b. — Cairns & Keller, 1993: 233.

Lophohelia tenuis Moseley, 1881: 180-181, pi. 8, figs 11-14.

Amphihelia oculata - Alcock, 1902c: 35. — Marenzeller, 1904a: 308-310, pi. 14, fig. 1.

Amphihelia ramea - Alcock, 1902c: 35. [See Zibrowius, 1980: 39 for a discussion of Madrepora ramea],

Amphihelia arbuscula - Alcock, 1902c: 35 (in part: "Siboga" 95, 156). [Not Lophohelia arbuscula Moseley, 1881).

Amphihelia tenuis - ALCOCK, 1902c: 36.

Sclerolielia formosa - Alcock, 1902c: 36. [See Zibrowius, 1974a: 570 for a discussion of Madrepora formosa].

Desmophyllum sp. - ALCOCK, 1902c: 28.

Madrepora alcocki Faustino, 1927: 106 ( nom . nov. for M. ramea Duncan, 1873; see Zibrowius, 1980: 39).

Madrepora tenuis - Faustino, 1927: 107-108, pi. 14, figs 2, 5. — ZIBROWIUS, 1974b: 765 (discussion).

Lophelia tenui (sic) - Hu, 1987: 40-41, pi. 2, figs 8-10, 12.

MATERIAL EXAMINED. — Philippines. "Siboga": stn 95 (ZMA Coel. 6455).

"Albatross": stn 5123, 50 branches (USNM M235386). — Stn 5124, 2 branches (USNM 96608). — Stn 5201,

3 branches (USNM 96602). — Stn 5202, 1 (USNM 96612). — Stn 5327, 7 branches (USNM 96631). — Stn 5348,

1 (USNM 96625). — Stn 5349, 1 (USNM 96627). — Sin 5373, 5 branches (USNM 96607). — Stn 5378, 5 branches

(USNM 96606). — Stn 5381, 6 branches (USNM 96630). — Stn 5403, 1 (USNM 96621). — Stn 5405. 10 fragments

(USNM 96618). — Stn 5406, 4 branches (USNM 96609). — Stn 5407, 13 branches (USNM 96622). — Stn 5408,

2 branches (USNM 96616). — Stn 5411, 1 (USNM 96626). — Stn 5417, 1 (USNM 96619). — Stn 5418, 2 branches

(USNM 96617). — Stn 5423, 2 (USNM 96620). — Stn 5424, 2 branches (USNM 96613). — Stn 5425, 1 branch (USNM

96602). — Stn 5428, 1 branch (USNM 96598). — Stn 5513, 5 branches (USNM 96604). — Stn 5516, 1 (USNM 96615).

— Stn 5527, 4 branches (USNM 96594). — Stn 5529, 7 branches (USNM 96593). — Stn 5541, 2 branches (USNM

96614). — Stn 5543, 5 branches (USNM 96632). — Stn 5574, 3 branches (USNM 96633).

Galathea": stn 436 (ZMUC). — Stn 443 (ZMUC).

"Hakuho Man,": stn KH72-1-20, 20 branches: 10 (OR1), 1 (USNM 96628).

Musorstom I: stn 49. 1 branch (USNM 96601).

Musorstom 2: stn 36 (MNHN). — Stn 39 (MNHN). — Stn 49 (MNHN).

80

S. D. CAIRNS & H. ZIBROWIUS

Musorstom 3: stn 89 (MNHN). — Stn 105, 2 branches (USNM 96597). — Sin 106 (MNHN). — Stn 122, 3 branches

(USNM 96599). — Stn 123 (MNHN). — Stn 128, 10 branches (USNM 96596). — Stn 139, 1 branch (USNM 96600).

Indonesia. "Siboga": stn 156 (ZMA). — Stn 259 (ZMA Coel. 5523).

"Albatross": stn 5586, 1 (USNM 96629). — Stn 5625, 8 branches (USNM 96623). — Stn 5645, 3 colonies (USNM

96592).

DEKi: stn 22, 2 branches (NNM 22723). — Stn 33 (NNM 22724). — Stn 45 (NNM 22726). — Stn 50 (NNM 22728)

— Stn 52 (NNM 22729). — Stn 58 (NNM 22730 and ZMUC). — Stn 59 (NNM 22731).

"Galathea": stn 490 (ZMUC).

"Hakuho Maru": stn KH72-1-28, 10 branches (USNM 96603).

Snellius 2: stn 4.144, 1 colony (NNM).

Karubar: stn 7, 2 (MNHN). — Stn 9, many branches (USNM 96582, 96635). — Stn 13, many branches (MNHN). —

Stn 16, 3 branches: 2 (MNHN), 1 (POLIPI). — Stn 19, 7 branches (USNM 96584). — Stn 35, 3 branches (POLIPI). -

Stn 39, 1 (USNM 96585). — Stn 42, 8 branches (POLIPI). — Stn 56, 3 branches (POLIPI). — Stn 59, many branches

(POLIPI). — Stn 69, many branches (MNHN and USNM 96582). — Stn 77, 3 branches (MNHN and USNM 96634).

South China Sea. " Albatross ": stn 5314, 2 branches (USNM 96610). — Stn 5317, 4 branches (USNM 96605).

"Hakuho Maru": stn KH72-1-52, 6 branches (USNM 96628). — Stn KH73-2-44-2, 20 branches (USNM 96586

96587).

Type Locality. — Tyrrhenian Sea and off Sicily (depth not given).

Diagnosis and Remarks. — Madrepora oculata is a widespread, variable species. Many names have beei

used for it (Zibrowius, 1974b) and more recently various forms have also been distinguished (Cairns, 1991). Thi

most commonly collected form occurring in the Philippine/Indonesian region is characterized by having delicate

uniplanar colonies formed of regular, nonanastomosing, sympodially budded corallites (M. tenuis of MOSELEY

1881 and form beta of Cairns, 1991). Its coenosteum is often light beige, finely granular, and longitudinally

striate. It is often attached to long hexactinellid spicules. Corallites 2. 3-3.0 mm in diameter, containing 3 ful

cycles of hexamerally arranged, narrow septa (Si>S2>S3), the CSi-2 usually slightly exsert but extending onl)

about 1 mm down from calice as thin, ridged costae. Inner edges of septa finely dentate to highly laciniate. Septa

faces may also bear tall, slender spines. Fossa of distal corallites usually deep, containing a papillose columella.

Specimens from two lots (Karubar stn 56 and "Albatross" stn 5645) differ from the form described above it

that they live in association with a symbiotic eunicid polychaete, which causes the coral to form a bushy corallun

of highly anastomotic branches.

Specimens from seven lots (Karubar stns 9, 13, 19, 77; "Albatross" stn 5529; "Hakuho Maru" stn 73-2-44

2; and DEKI stn 50) bear some bell-shaped, hypertrophied corallites, as much as 23 mm in GCD and having up tc

82 porous septa. Once reported as the first neoplasms in Scleractinia (SQUIRES, 1965), these abnormally large

corallites are now known to be the manifestation of a parasitic petrarcid ascothoracidan crustacean (Grygier &

Cairns, 1996).

A more complete description and illustrations of this species are found in Zibrowius (1974b) and Cairns

(1982, 1991).

Distribution. — Philippines: common throughout Philippines from Lubang Island to Bohol Sea; Sulu

Archipelago; 161-2021 m. Indonesia: Molucca Sea; Halmahera Sea; Banda Sea (Kai Islands and Sulawesi); Arafura

Sea (southeast of Tanimbar Islands); Timor Sea (south of Leti and Timor Islands); Savu Sea; Flores Sea (Selayar

Island, Sulawesi); Java Sea; 1 12-984 m. Elsewhere: Malaysia (Celebes Sea off Sabah); cosmopolitan, except for

continental Antarctica; 15-1500 m.

Madrepora arbuscula (Moseley, 1881)

Figs 3 c-g

Not Madrepora arbuscula Dana, 1846: 474 [= Acropora sp.].

Lophohelia arbuscula Moseley, 1881: 180, pi. 8, figs 9-10.

Amphihelia arbuscula - Alcock, 1902c: 35 (in part: "Siboga" stn 12).

Madrepora arbuscula - FAUST1NO, 1927: 107 (in part). — ZIBROWIUS, 1974b: 765.

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

81

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn 126, 2 large colonies (USNM 96750) and 13 branch

fragments (MNHN).

Indonesia. " Challenger ": stn 194, holotype (BMNH 1880.11.25.96).

" Siboga stn 12, 2 branches (ZMA).

"Albatross": stn 5584, 1 branch (USNM 96757).

Karubar: stn 18, 17 branch fragments (USNM 96752). — Stn 32, 4 branches: 1 (POLIPI), 3 (MNHN).

Type Locality. — "Challenger" stn 194: 4°3 1 'S, 129°57'E (Banda Island, Banda Sea), 366 or 658 m.

Description. — Corallum massive, the largest colony (MUSORSTOM 3 stn 126) 26 cm in height, having a

stout, dense basal stem 5.0 cm in diameter. Lower 1/3 to 1/2 of colony consists only of basal stem and individual

corallites that bud directly from the coenosteum, but upper corallum ramifies into anastomosing branches, each of

which contains a central tubular cavity about 5x7 mm in diameter that is presumed to be inhabited by a eunicid

polychaete. Calices 2.5-3. 1 mm in diameter, circular to slightly elliptical, tending to concentrate on one face of the

corallum (here defined as anterior). Whereas corallites do occur on the posterior face, they are fewer in number,

often nonfunctional, filled in with stereome, and usually completely absent from the lower 10 cm of basal stem

due to incorporation within basal coenosteal layering. Coenosteum on distal branches remarkably smooth and

porcellaneous (not granular) even at a magnification of x 2500; however, larger-diameter branches and worn

coenosteum sometimes reveal very shallow longitudinal striae that delimit perfectly flat costae. Septocostal

ridges absent from calicular edges. Corallum white; however, coenosteum immediately adjacent to calice a dark

brown.

Septa hexamerally arranged in 3 complete cycles: S i >S2-3- Si only about 0.4 mm exsert, extremely narrow

(0.10-0.15 mm), having entire (nondentate) inner edges and irregularly granular septal faces. S2-3 nonexsert, only

about 0.06 mm (60 )tm) in width, and otherwise similar to the Si. Deep in fossa the lower, inner edges of the

6 Si and sometimes some of the S2 widen and join one another in centre of fossa, forming a short styliform

columella (Fig. 3f). Deep in fossa S3 are very narrow, each pair within a system bending toward and fusing to its

common S2. Fossa deep and spacious, due to the very narrow septa.

Each polyp possesses 1 long sweeper tentacle, up to 1 1 mm long in the preserved state, the other more typical

tentacles being only 1-2 mm in length. This is the first case of sweeper tentacles being observed in a deep-water

coral.

Remarks. — Madrepora arbuscula differs from other species of Madrepora in several significant characters.

It has: 1) a very massive basal stem, 2) no coenosteal granules, which produces a porcellaneous texture, 3) white

coenosteum but brown-edged calices, 4) extremely narrow septa with no inner edge dentition, 5) a rudimentary

styliform columella (that of M. oculata is papillose), and 6) sweeper tentacles. Although tentacle length of

Madrepora generally had no special attention, the second author has occasionally observed that live M. oculata had

short, uniform-length tentacles. It was therefore surprising to see that M. arbuscula from MUSORSTOM 3 stn 126

had one long sweeper tentacle per polyp in addition to normal-sized, shorter tentacles. This was seen on the freshly

dredged live colonies kept in seawater, and can still be seen after preservation in formalin.

In addition to the ubiquitous spirocysts, both the normal and sweeper tentacles of M. arbuscula (MUSORSTOM

stn 3-126) contain b-rhabdoids (type 1), p-rhabdoids D (type 1), and holotrichs (type I). According to DEN Hartog

(1977), the cnidae of sweeper tentacles are always different from that of normal tentacles, so M. arbuscula may

present a unique condition among the approximately 14 scleractinian species now known to have sweeper tentacles

(Williams, 1991). Nematocyst identifications and terminology follows the review by Pirez and Pitombo (1992)

and is considered only preliminary — the quantification of sizes remains to be tabulated.

Madrepora arbuscula (Moseley, 1881) is a junior secondary homonym of M. arbuscula Dana, 1846, but because

Dana s species is now considered to be in the genus Acropora, according to the ICZN (article 59c) Moseley’s

name does not have to be replaced.

Distribution. Philippines: Sulu Sea (Semirara Islands); 266 m. Indonesia: Banda Sea (Kai Islands and

Banda Islands); Bali Sea; 212-658 m; Malaysia (Celebes Sea off Sabah).

82

S. D. CAIRNS & H. ZIBROWIUS

Madrepora minutiseptum sp. nov.

Figs 4 a-d, 5 a-b

Amphihelia infundibulifera - Saville Kent, 1871: 276-277, pi. 24, figs 4, 4a, 4b. — Quelch, 1886: 26, 53. [Not Oculint

infundibulifera Lamarck, 1816: 286; 1836: 457 (= stylasterid), see Remarks],

MATERIAL EXAMINED/TYPES. — Indonesia. Ternate, from local governor S.C.J. Musschenbroek tc

"Challenger" naturalists, October 1874, 1 paratype colony (BMNH 1886.12.9.57).

CORINDON 2: stn 235 (confused station data!), 1 branch fragment with 6 calices, paratype (USNM 96756). —

Stn 248, 1 small branch with ca. 20 calices, paratype (POL1PI) and 1 small colony with remarkably different calices, bu

possibly the same species, see description (MNHN).

Snellius 2: stn 4.196, 1 colony (holotype) (NNM 22734).

Karubar: stn 18, 3 tiny fragments altogether 4 calices (USNM 96755).

Taiwan (= Formosa). From R. SwiNHOE, 3 paratype colonies (BMNH 1865.12.15.1), 6 paratype colonies (BMN1

1870.5.9.21-22-24-25-26-27). — 1 colony (AMS G 7014; probably transfer from BMNH). — coll. FRIES, I paratyp<

colony (NMW 8227; old number 3180, registered in 1884).

Japan. From F. van Heukelom, 1 colony (ZMA Coel. 7400). — No details, 2 colonies (ZMA Coel. 138, 139). — N<

details, 1 colony (USNM 96754, ex ZMA Coel. 137, transfer Sept. 1994).

No origin indicated. Formerly Gerresheim collection, seen by Ehrenberg, 1 colony (ZMB 582).

Type Locality. — SNELLIUS 2 stn 4.196: 6°23'S, 120°26.5'E (southwest of Salayer, Flores Sea)

150-200 m.

ETYMOLOGY. — The species name minutiseptum (Latin minutus, small + septum , a fence, bar), alludes to th<

very small septa of this species. The name is treated as a noun in apposition.

DESCRIPTION. — Corallum arborescent, irregularly branching, often tending to form subflabellate colonies

Larger colonies (holotype; Ternate specimen) 20-25 cm in height. A strongly developed trunk and main branche

are hollow, containing a central canal with lateral openings (by analogy with Madrepora oculata interpreted a:

tubular growth induced by an eunicid polychaete). Inner diameter of these tubes generally ca 3.5 mm. Anastomose

of branches frequent. Thin distal or peripheral branches distinctly zigzag-shaped, with corallites sympodially buddei

(1 or 2 on the parent corallite), on trunk and main branches corallites immersed into sclerenchyme. Corallum solid

with smooth surface; no costae. White, except for pigmented calicular edge or upper 1/2 of sympodial corallites

Pigmentation reddish brown to dark brown, extending throughout the skeleton thickness at the pigmented level

not only superficial; also extending inside calice on septa. Pigmentation of septa decreasing in depth of calice

Corallites on thin branches about equal in height and width, 1. 5-2.5 mm in calicular diameter, with a narrow base

and up to 2.5 mm in height. Corallites distinctly infundibuliform, with wide "empty" fossa because of the low

septa. Septa are essentially reduced to a row of flattened septal edge teeth coalescent laterally with granules of the

reduced septal faces. Septa hexamerally arranged in 3 complete cycles, as ridges of low papillae rather than distinct

lamellae. Si to S3 slightly decreasing in width, with a correspondingly slight decrease in exsertness above the

calicular edge. No columella, bottom of fossa covered by small papillae similar to those of the higher and more

peripheral parts of the septa.

CORINDON 2 stn 248 provided 1 small branch (21 mm in length) with the eunicid-induced deformation and

comprising ca. 20 calices, of the typical aspect. But from the same station there is another small colony about

15 mm in height with the same type of deformation, the calices of which are exceptionally small (1 mm or less in

diameter), with only 12 septa. These septa are similar to those in typical M. minutiseptum with 24 septa (i.e., as

in the branch from CORINDON 2 stn 248) by being very low and denticulate. This unique specimen may be an

initial colony of the same species, dwarfed for some reason. For this reason it is not given paratype status.

Remarks. — Although differing considerably from the type of the genus, Madrepora oculata , the species in

question is attributed herein, provisionally, to the genus Madrepora on the basis of a series of characters: the small

sized corallites arranged in sympodial branches, septa in 3 cycles, the general shape of colonies, and the regular

presence of a not yet observed symbiont which causes the tubular growth deformation and which is presumed to be

an eunicid polychaete. M. minutiseptum differs from all congeners by its very low septa which are hardly more

than a denticulate ridge. Like M. arbuscula , it is partly pigmented (near the calicular edge).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

83

The species described here under Madrepora minutiseptum had recognizably been characterized and illustrated by

Saville Kent (1871) as a scleractinian under the name Amphihelia infundibulifera, on the basis of "specimens in

the British Museum, collected at Formosa by Consul Swinhoe" (material which we have seen). Saville Kent

was mistaken in being convinced that his material was "evidently identical with the species described .. by

Lamarck", i.e., Oculina infundibulifera Lamarck, 1816, presumed to come from the Indian Ocean. Lamarck's

(1816: 286) text in Latin and French (the same in the 2nd edition, 1836: 457, without additions) is brief and im¬

precise, as usual for most coral descriptions in the early 19th century. A critical analysis of this text strongly

suggests that Lamarck's O. infundibulifera is not a scleractinian but a stylasterid coral (at that time stylasterids

were not yet recognized as a distinct group). O. infundibulifera was said to be close to the "following" species,

O. flabelliformis Lamarck, 1816 (now Stylaster flabelliformis; for a detailed description, including of

Lamarck's type, see Boschma, 1957b); to be almost flabellate; and to have very small zigzag-shaped branches

aside the thicker main ones, both of which are coalescent. All this characterizes a stylasterid rather than a

scleractinian.

Lamarck's O. infundibulifera was mentioned by H. Milne Edwards & Haime (1857: 131) as a

problematical stylasterid under the combination Allopora (Stylaster) infundibulifera. Having not seen

Lamarck's type, they referred to Dana (1848) for this interpretation. Lamarck's type of O. infundibulifera

appears lost. It is not with the LAMARCK collection at the Museum national d'Histoire naturelle, Paris. It probably

has never been there since Lamarck's text does not indicate "Mus. no.", his usual way of referring to specimens

present in the MNHN collection.

Many (not all) secondary quotations of Lamarck's species are listed by Boschma (1957a: 62-63, under

Amphelia infundibulifera) who, following Saville Kent (1871), considered it as a scleractinian. But, given the

strong arguments for Lamarck's O. infundibulifera being a stylasterid, this name cannot be used for Saville

Kent's species which is a Madrepora ( = Amphihelia).

Should there be another old name available for Saville Kent's scleractinian? Ehrenberg (1834: 302-303)

distinguished 4 new varieties of Oculina virginea (this name dates back to Madrepora virginea Linnaeus, 1758, an

unidentifiable taxon): pachyclados, leptoclados, tubulifera, immersa. As typical for Ehrenberg's taxa, the

characterization of these 4 varieties is too brief to allow adequate identification based on his text alone, O. virginea

being itself a confused taxon since the beginning.

The Berlin Museum possesses 5 colonies referable to 2 species of Madrepora (= Amphihelia) and considered as

being part of the museum's initial collection studied by Ehrenberg (1834). All are without indication of origin,

as common at that time of "natural history cabinets", and 4 of them are ascribed to GERRESHEIM, an early

19th century collector. The 2 species in question are: Madrepora oculata Linnaeus, 1758 (see Cairns, 1979;

Zibrowius, 1980) represented by 4 colonies, and M. "infiindibulifera" sensu Saville Kent, 1871, represented by

one colony. Only one colony (of M. oculata) is accompanied by a label in EHRENBERG's handwriting, reading

Oculina virginea Lam. (but no variety is indicated). The 4 other colonies (3 M. oculata; 1 M. "infundibulifera"

sensu Saville Kent) all have later 19th century, not original, labels reading Oculina virginea var. leptoclados.

It cannot be elucidated whether the identification of colonies of 2 distinct species to var. leptoclados was by

Ehrenberg himself or was due to some later confusion. Ehrenberg's characterization ( ramis tenuioribus ; Latin:

with thinner branches) of var. leptoclados (Greek: thin branch), opposed to the characterization ( ramis crassioribus ;

Latin: with thicker branches) of var. pachyclados (Greek: thick branch) is of no help, especially since no

authentically labeled sample of var. pachyclados is present that could be the reference. Accordingly the only colony

(ZMB 582, Gerresheim) of M. "infundibulifera" sensu Saville Kent in the old Berlin collection should not be

considered as the type of Ehrenberg’s var. leptoclados and this name is rejected as unavailable for SAVILLE

Kents species. The latter is here named Madrepora minutiseptum because of its very reduced low septa.

Distribution. — Remarkably, this species is represented by quite a number of colonies in several old

museum collections (Berlin, Amsterdam, London, Sydney, Wien), received in the 19th century. A few more

specimens were collected in the 1980's. Although there is no modern record from the area, Taiwan (= Formosa) as

the origin of a series of colonies at the BMNH received from "consul Swinhoe" is surely correct since Robert

S\ tnhoe (1836-1877) had indeed been on duty in China for many years, including in Taiwan (in 1858, 1861 and

1864-1866, especially at Tansui, in the north of the island; see Mearns & Mearns, 1988).

84

S. D. CAIRNS & H. ZIBROWIUS

In Indonesia the species has been recorded from 4 distinct areas: from Salayer, type locality (Snellius 2: 1 50-

200 m); from Temate (" Challenger " expedition: undetailed); from Makassar Strait (CORINDON 2 stn 235, although

the depth of 1 1 10 m is certainly confused); and from the Kai Islands (KARUBAR stn 18: 205-212 m).

Confirmed distribution. — Taiwan, Japan and Indonesia; 150-212 m.

Genus CYATHELIA H. Milne Edwards & Haime, 1849

Cyathelia axillaris (Ellis & Solander, 1786)

Madrepora axillaris Ellis & Solander, 1786: 153, pi. 13, fig. 5.

Cyathohelia axillaris - MOSELEY, 1876: 547; 1881: 175-176. — Bassett-Smith, 1890: 367. — BEDOT, 1907: 145

pi. 15, figs 1-3.

Cyathelia axillaris - NEMENZO, 1979: 11-12, pi. 3, fig. 3. — CAIRNS, 1994: 43-44, pi. 18, figs a-c (synonymy).

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5134, 1 (USNM 96758). — Stn 5255, 1 (USNM 96759)

— Stn 5268, 1 (USNM 96760). — Stn 5367, 5 (USNM 96761).

MUSORSTOM 3: stn 131, 2 (USNM 96763).

Indonesia. "Siboga": stn 260, 1 (ZMA Coel. 6438). — Stn 310, 1 colony (ZMA Coel. 6619); unknown station

1 colony (ZMA Coel. 6439).

DEKi: stn 24, 2, (NNM 22449). — Stn 104, 1 (NNM 22448). — Stn 105, 1 (NNM 22443). — Unnumbered station

Ambon, 25-130 m, 1 (NNM 22445-47).

CORINDON 2: stn 248, 2 (MNHN).

Snellius 2: stn 4.106, I (NNM 22444).

Karubar: stn 22, 1 (USNM 96762). — Stn 30, 1 (USNM 96763).

Type Locality. — Eastern Indian Ocean (depth not given).

Diagnosis. — Corallum sparsely branched, resulting in small, robust, bushy colonies, the largest knowr

7.5 cm in height, consisting of about 100 corallites (EGUCHI, 1968). Extratentacular branching essentially

sympodial, 2 buds usually originating on opposite sides of a terminal corallite, the parent corallite eventually

becoming immersed in thick coenosteum in resultant branch axil. Corallites relatively large, up to 11 mm GCD

Corallum light brown, the corallites usually a darker shade. Septa hexamerally arranged in 4 cycles: Si-2>S3>S4

A crown of 12 thick pali (P1-2) encircle the columella and a second, slightly more recessed crown of 12 P3 stand

higher in the fossa, resembling the calice of a Trochocyathus. Columella papillose.

REMARKS. — Among the colonial azooxanthellate corals known from this region, Cyathelia is distinguished

by its distinctive branching: two large corallites budding from opposite sides of a parent corallite, the parent

corallite often becoming constricted. Also distinctive, the coenosteum of this species is white to beige, whereas

the calice and near-calice coenosteum are a darker brown. A complete description is given by NEMENZO (1979) and

Cairns (1994).

DISTRIBUTION. — Philippines: Verde Island Passage; Bohol Sea (Negros); Davao Gulf; Sulu Sea (Zamboanga

Peninsula); 46-329 m. Indonesia : Makassar Strait; Molucca Sea; Ceram Sea (south of Obi Islands); Banda Sea (Kai

and Ambon Islands); Flores Sea (Sumbawa); Java Sea (Sunda Strait); 13-170 m. Elsewhere: South China Sea

(Tizard Bank); northern Indian Ocean; Japan (Honshu and northern Ryukyu Islands); Korea Strait; 15-366 m.

Genus NEOHELIA Moseley, 1881

Neohelia sp. cf. N. porcellana Moseley, 1881

Figs 5 c-e, g-h

Material EXAMINED. — Indonesia. "Siboga": stn 289, 1 colony (USNM 96766). — Stn 305, 1 colony (ZMA).

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

85

Deki: unnumbered station in Bay of Ambon, 91 m, 1 broken colony (ZMUC). — Unnumbered station in Bay of

Ambon, 46-55 m, 1 colony (NNM 22680).

Corindon 2: stn 248, 1 colony (MNHN).

Snellius 2: stn 4.104, several colonies (NNM 22681), 1 colony (USNM 96765). — Stn 4.105, 2 colonies (NNM

22733).

DESCRIPTION. — Corallum up to 5 cm in height, forming a very thin, easily fractured encrustation around the

parchment-like tube of a eunicid polychaete or gorgonian axis. Only very short, slender branches consisting

of 3-5 corallites project from the basal encrustation. If encrusting a polychaete tube, the terminal aperture

created by the polychaete is approximately 2.8 mm in diameter; however, additional circular to elliptical

pores of variable diameter (0.4- 1.6 mm) penetrate the coenosteum providing feeding apertures for the worm.

Calices circular but variable in diameter, ranging from 0.70-1.50 mm. Theca white, with very faint, longitudinal

costal striations. Coenosteum of some coralla also densely covered with small, blunt papillae up to 70 pm

in diameter and 80 pm in height, similar to the "granular echinulations" illustrated by Pratt (1900: pi. 62,

fig. 2).

Septa hexamerally arranged in 3 generally full cycles: Si>S2>S3; in some corallites from Ambon (ZMUC),

1 or 2 pairs of S3 are sometimes missing, resulting in 20-22 septa. Si highly exsert (up to 0.25 mm) and quite

narrow (about 0.11 mm). Vertical inner edges of Si slightly thickened and sinuous deep in fossa, where they

extend to the centre of fossa. S2 less exsert and narrower (about 0.6 mm) than the Si, not extending quite as far

toward centre of fossa. S3 least exsert and least wide (about 0.4 mm) septa, attenuating before they reach base of

fossa. Fossa relatively shallow and quite wide, resulting from the small size of the septa; fossa with a horizontal

floor. Columella rudimentary, consisting of several irregularly shaped papillae, the papillae often an extension of

the inner Si septal edges.

Remarks. — Both Moseley (1881) and Pratt (1900) reported their Vanuatu specimens of N. porcellana

to have exclusively pentameral symmetry in 3 cycles (5:5:10), resulting in 20 septa, although PRATT (1900: 592)

acknowledged that "there is a tendency for them to lose their symmetry of arrangement". Likewise, most of

Wells' (1984) Pleistocene Vanuatu specimens (USNM) have 20 septa, but several corallites were found

to have 24 septa arranged in three cycles. Whereas most of the Ambon corallites have hexameral symmetry

in 3 cycles (24 septa), the 3rd cycle is occasionally incomplete, resulting in 20, 22, or 24 septa. Because

of the difference in septal symmetry (pentameral resulting in 20 septa for previously reported

Neohelia vs. hexameral resulting in 24 septa) the Indonesian specimens are not definitively identified as

N. porcellana.

Some early authors (PRATT, 1900; HICKSON, 1903) believed that N. porcellana was able to produce a horny

membrane underlying its calcareous coenosteum. In fact, similar to Madrepora , Neohelia may overgrow the

productions of other organisms and especially the membrane-like tubes of symbiotic eunicid polychaetes.

However, more recently, polychaetes have been found within Madrepora axes (CAIRNS, 1991, 1995) and, indeed, a

dried eunicid was found in the tube of the Neohelia from Ambon, suggesting that the polychaete, not the coral,

secretes the homy membrane.

WELLS (1984) synonymised Neohelia with Madrepora without explanation. The rudimentary columella in

Neohelia and its distinctive coenosteal papillae would seem to justify the retention of this genus. Neohelia

porcellana is described in great detail by Pratt (1900), including characteristics of the soft parts.

Distribution. — Indonesia : Makassar Strait; Banda Sea (Bay of Ambon: Solar Strait); Timor Sea (southeast

of Timor); Lintah Strait, west of Flores; 55-170 m.

Typical pentameral Neohelia porcellana is known from Vanuatu (Api Island, type locality) (Fig. 50 and

Pleistocene of Kere and Navaka River (WELLS, 1984); Loyalty Islands (PRATT, 1900; HICKSON, 1903); and New

Caledonia. Wells (1984) listed N. porcellana from northwestern Australia (140-141 m), but we can find no

documentation of this record. According to MOSELEY (1881) the types were collected at "Challenger" stn 177 at

63 fathoms (=115 m), but according to TIZZARD et al. (1885, cruise narrative) it is 130 fathoms (= 238 m).

Vaughan & Wells (1943) indicate a range of 91-1 15 m for the species, but again without documentation. The

only reliable depth range for the species is that of the types: 1 15-238 m.

86

S. D. CAIRNS & H. ZIBR0W1US

Family ANTHEM I PH YLL1 1 DAE Vaughan, 1907

Genus ANTHEMIPHYLLIA Pourtales, 1878

Anthemiphyllia dentata (Alcock, 1902)

Discotrochus dentaius Alcock, 1902a: 104; 1902c: 27, pi. 4, figs 26, 26a. — FAUSTINO, 1927: 63, pi. 7, figs 1-2.

Anthemiphyllia dentata - BEST & HOEKSEMA. 1987: 398-399, figs 9a-c. — Zou et ai, 1988: 195. — Cairns & Parker

1992: 16-17, pi. 4, figs e-f (synonymy). — Cairns & Keller, 1993: 233, pi. 3, fig. E. — Cairns, 1994: 44, pi. 18

figs d-f (synonymy); 1995: 41-42. pi. 6, figs c-g (synonymy).

Anthemiphyllia dentatus - Yabe & EGUCHI, 1941b: 213, figs la-b.

Deltocyathus andamanicus - Keller, 1982: 52 (in part: pi. 1 [= 4], figs 3-4, "Dimitri Mendeleev" sin 1411). [No

Deltocyathus andamanicus Alcock, 1898].

Not Discotrochus sp. - ALCOCK, 1902c: 27-28 [= undescribed Anthemiphyllia having costal spines].

Not Anthemiphyllia dentata - Cairns. 1984: 1 1, pi. 1, figs F-G [= undescribed Anthemiphyllia, see CAIRNS, 1994],

MATERIAL EXAMINED. — Philippines. " Albatross stn 5162, 2 (USNM 96767). — Stn 5178, 1 (USNM 96768).

"Hakuho Maru ": stn KH72-1-20, 1 (USNM 96774).

Musorstom 2: stn 33, 7 (MNHN).

Musorstom 3: stn 108, 1 (USNM 96771). — Stn 126, 2 (USNM 96772). — Stn 130, 1 (MNHN). — Stn 131

11 (MNHN).

Indonesia. "Albatross": stn 5584, 1 (USNM 96770).

Snellius 2: stn 4.033, 1 (NNM 22494). — Stn 4.034, 3 (NNM 18013, mentioned by Best & Hoeksema, 1987).

Karubar: stn 1, 10 (MNHN).

Type Locality. — Sulu Sea, 350-522 m.

DIAGNOSIS. — Corallum discoidal, with a flat to slightly bowl-shaped base; largest Philippine specimer

(MUSORSTOM 3 stn 126) 21.3 mm in calicular diameter. Corallum usually free, but with a circular scar oi

irregularity 2-6 mm in diameter at centre of base. Costae rounded and granular, separated by shallow intercostal

furrows, that, in large coralla, are bisected by very thin ridges. Septa hexamerally arranged in 5 cycles, the 5tl

cycle complete only in large specimens. Septal formula: Si-2>S3>S4>Ss- Si-2 quite thick, highly exsert, bearing

7-11 coarse septal lobes. Like the Si-2, the S3 also reach the columella but are less thick, having more numerous,

finer teeth. S4 much smaller than S3, having laciniate inner edges. S5 rudimentary and highly laciniate. Fossa

shallow; columella papillose.

Remarks. — Anthemiphyllia dentata is distinguished from other solitary, discoidal, azooxanthellate species

from this region by having very thick Si -2 with coarse septal dentition.

Distribution. — Philippines: Lubang Island; Verde Island Passage; Sibuyan Sea; Sulu Sea (Semirara Islands.

Sulu Archipelago); 122-522 m. Indonesia: Banda Sea (Kai and Tukangbesi Islands); 280-534 m. Elsewhere:

Malaysia (Celebes Sea off Sabah); South China Sea (west of Palawan; north of Pratas Islands); widespread from

southwestern Indian Ocean to southern Japan, the Kermadecs, and Tasmania; 50-570 m.

Anthemiphyllia frustum Cairns, 1994

Figs 6 a-b

Anthemiphyllia frustum Cairns, 1994: 44-45, pi. 18, figs g-i, pi. 19, figs a-b.

Material EXAMINED. — Indonesia. Snellius 2: stn 81.2, 3 (NNM 23073).

Karubar: stn 2, 1 (MNHN). — Stn 15, 24: 6 (MNHN), 18 (USNM 96776).

South China Sea. "Albatross": stn 5313, 1 (USNM 96769).

Type Locality. — 30°59'N, 130°32'E (Osumi Strait, southern Kyushu, Japan), 237-241 m.

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

87

DIAGNOSIS. — Anthocyathus: base flat, its calicular diameter being less than its basal diameter. Largest known

specimen (SNELLIUS 2 stn 81.2) 10.2 mm in calicular diameter and 7.4 mm in height, consisting of 2 anthocyathi

that have not completely separated (Fig. 6a). A faint, circular detachment scar 5-6 mm in diameter present in centre

of base. Septa hexamerally arranged in 4 cycles (Si>S2>S3>S4), being thick and closely spaced. Adjacent septa

strongly fused by 3 or 4 vertical synapticular plates, similar to those illustrated for Fungiacyathus turbinolioides

(CAIRNS, 1989a, pi. 6, fig. f), but only visible in a longitudinal fracture of a damaged corallum. All septa bear

massive triangular teeth for their entire length. Columella papillose, composed of 10-15 massive, granular

papillae. Anthocaulus unknown.

DISTRIBUTION. — Indonesia : Banda Sea (Kai and Tanimbar Islands); 209-340 m. Elsewhere : South China Sea

(Pratas Islands); Osumi Strait (south of Kyushu, Japan); 237-274 m.

Suborder CARYOPHYLLIINA

Superfamily CARY OPH YLLIOIDEA Dana, 1846

Family CARYOPHYLLIIDAE Dana, 1846

Genus CARYOPHYLLIA Lamarck, 1816

Subgenus CARYOPHYLLIA (CARYOPHYLLIA) Lamarck, 1816

Key to the 13 species of Caryophyllia (Cary ophy Ilia)

known from the Philippine/Indonesian region

Note: Additional undescribed and/or unreported species of Caryophyllia probably also occur in this region. Thus

this key should be considered as only a guide to the more common species included in this review.

1 . Corallum attached (ceratoid, trochoid, or subcylindrical) . 2

— Corallum free (unattached), usually ceratoid . 1 2

2. Septa arranged hexamerally (6 or 12 primary septa) . 3

— Septa arranged pentamerally, septamerally, octamerally, decamerally, or tetradecamerally

(5, 7, 8, 10, or 14 primary septa) . 7

3. Penultimate cycle of septa (S3) equal to or wider than last cycle (S3^S4) . 4

— Penultimate cycle of septa (S3) less wide than last (S4) cycle (S3<S4) . 6

4. Corallum firmly attached to substratum . 5

— Corallum rises from a slender pedicel that bears a small detachment scar . C. secta

5. Theca transversely ridged; septa often brown . C. lamellifera

— Theca longitudinally costate; corallum white . C. diomedeae

6. Fossa quite deep; columella rudimentary . C. crosnieri

— Fossa not deep; columella well developed . C. panda

7. Septa arranged octomerally . 8

— Septa arranged pentamerally, decamerally, or tetradecamerally . 9

8. Theca transversely ridged; Si>S2>S3; S1-2 and pali quite sinuous . C. rugosa

— Theca costate or porcellaneous; Si>S3>S2; septa and pali only moderately sinuous .

. C. octonaria

88

S. D. CAIRNS & H. ZIBROWIUS

9. Corallum with 5 large primaries (pentameral symmetry); theca and septa usually bear a

mottled pigmentation . C. hawaiiensis

— Corallum with 10-14 large primary septa . 10

10. Corallum with 10 primary septa (decameral symmetry) . C. quadragenaria

— Corallum with 14 (rarely 16) primary septa . 11

1 1 . Theca transversely ridged; septa usually darkly pigmented . C. lamellifera

— Theca longitudinally costate; septa not pigmented . C. transversalis

12. Corallum with 96 or more septa . 13

— Corallum with less than 96 septa (usually 48-72) . 1 4

13. S4 wider than S5; pedicel may be present; known depth range 251-567 m ... C. grandis

— S5 equal to or wider than S4; pedicel absent; known depth range deeper, 468-1048 m .

. C. ambrosia

14. Calicular margin lanceted; 48 septa usually present; depth range 353-1276 m .

. C. scobinosa

— Calicular margin serrate; usually 48-72 septa; known depth range deeper, 1525-2623 m ...

. C. cornulum

Caryophyllia (C.) diomedeae Marenzeller, 1904

Caryophyllia ephyala - Alcock, 1902c: 9. [= ? C. ephyala Alcock in Wood-Mason & Alcock, 1891].

Caryophyllia diomedeae Marenzeller, 1904b: 79-80, pi. 1. fig. 2. — Cairns, 1991: 11-12, pi. 4, figs c-e (synonymy

1995: 49-50, pi. 9, figs a-d (synonymy).

MATERIAL EXAMINED. — Philippines. "Siboga": stn 95, 1 (ZMA).

"Albatross": stn 5317, 2 (USNM 96778).

Musorstom 1: stn 49, 2 (USNM 96781).

Musorstom 2: stn 15, 1 (MNHN).

Musorstom 3: stn 95. 2 (MNHN).

Indonesia. "Siboga": stn 59, 1 (ZMA).

"Albatross": stn 5634, 1 (USNM 96789). — Stn 5656, 3 (USNM 96780).

Karubar: stn 3, 1 (MNHN).

Type Locality. — "Albatross" stn 3358: 6°30'N, 81°44’W (off Coiba Island, Panama), 1043 m.

DIAGNOSIS. — Corallum conical, distally flared; largest known corallum 29.9 mm in GCD (Cairns, 1995),

but largest from Indonesian region only 12 mm in GCD and 21 mm in height. PD:GCD = 0.26-0.40. Costae fla

and poorly distinguished, covered with inconspicuous granules, sometimes porcellaneous. Septa hexamerally ar¬

ranged in 4 full cycles: Si-2>S3>S4- S3 have highly sinuous inner edges, those of S 1-2,4 less sinuous. Septa only

moderately exsert. A tight crown of 12 P3 encircle a fascicular columella of 3-12 slender lamellae. Fossa shallow.

Remarks. — Among the 56 Recent species of Caryophyllia listed by Cairns (1991), the largest subset

(19 species) are those species having attached coralla and hexamerally arranged septa in 4 cycles. C. diomedeae

belongs to this morphological subset, as do the first 3 species in the account of this genus, the 4th (C. lamellifera )

having both hexameral and heptameral symmetry. C. diomedeae can be distinguished from other species in this

region using the key.

DISTRIBUTION. — Philippines: Lubang Island; Verde Island Passage; Sulu Sea (Sulu Archipelago); 330-

865 m. Indonesia: Ceram Sea (south of Obi Islands); Banda Sea (Kai Islands); Teluk Bone (Sulawesi); Savu Sea

(Timor); 300-885 m. Elsewhere: widespread, including throughout Pacific from Panama to Tasmania; 225-

2200 m.

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

89

Caryophyllia (C.) crosnieri nom. nov.

Caryophyllia elongata Cairns in Cairns & Keller, 1993: 236-237, pi, 4, figs A-B. — Cairns, 1995: 52, pi. 10,

figs d-f. [Not Caryophyllia clavus var. elongata Duncan, 1873],

MATERIAL EXAMINED. — Philippines. Musorstom 2: stn 15, 1 (MNHN).

Indonesia. Deki: stn 3, 2 (NNM 22885). — Stn 12, 3 (NNM 22884).

Karubar: stn 5, 1 (MNHN). — Stn 31, 7 (USNM 96785). — Stn 32, 1 (POLIP1).

TYPE Locality. — "Vityaz" stn 2716: 33°17'S, 44°55'E (off Walter's Shoal, Madagascar Plateau), 630-

680 m.

DIAGNOSIS. — Corallum ceratoid to subcylindrical; largest known specimen (Karubar stn 31) 8.8 x

10.8 mm in calicular diameter and 14.1 mm in height. PD:GCD = 0.50-0.81. Theca porcellaneous, covered with

low, rounded granules. Corallum white, but often having a light brown calicular edge. Septa hexamerally arranged

in 4 complete cycles: Si>S2>S4^S3- Si highly exsert, forming triangular calicular lancets. Inner septal edges

only slightly sinuous. A tight crown of 12 slender P3 encircle a fascicular columella composed of 3-9 very slender

twisted lamellae. P3 appear to be paired within each system. Fossa extremely deep and narrow.

REMARKS. — Caryophyllia crosnieri is more fully described and illustrated by CAIRNS & KELLER (1993) and

CAIRNS (1995) as C. elongata. The name elongata Cairns in CAIRNS & KELLER (1993) is a junior primary

homonym of Caryophyllia clavus var. elongata Duncan, 1873, and thus is replaced herein. This species is renamed

for Alain CROSNIER (MNHN), one of the driving forces behind the MUSORSTOM expeditions and the resultant

publications. The species also occurs in Madagascar, Indonesia, Philippines, and New Caledonia, areas where he

contributed to the collection of deep-water benthos.

Caryophyllia crosnieri is distinguished from its congeners by having an extremely deep fossa and small "paired"

pali (see key).

Distribution. — Philippines : Mindoro Strait; 326-330 m. Indonesia: Banda Sea (Kai Islands); 206-296 m.

Elsewhere: Madagascar Plateau; Kermadec and Three Kings Ridges; New Caledonia region; 165-680 m.

Caryophyllia (C.) secta sp. nov.

Figs 6 c-e

Material EXAMINED/TYPES. — Philippines. "Albatross": stn 5116, I paratype (USNM 96786). — Stn 5265,

holotype (USNM 96787) and 1 paratype (USNM 96788). — Stn 5567, I paratype (USNM 96789).

Mortensen'S Pacific Expedition: Zamboanga Peninsula, 160-200 fv (= 301-373 m), 14 March 1914, 2 paratypes

(NNM 22771).

Musorstom 2: stn 32, 8 paratypes (MNHN).

Musorstom 3: stn 126, 2 paratypes (USNM 96790).

Indonesia. Mortensen'S Java-S. A. Expedition: stn 15, 1 paratype (ZMUC).

Karubar: stn 3, 1 paratype (MNHN).

Type Locality. — "Albatross" stn 5265: 13°41'N, 120°00'E (Lubang Island, Philippines), 247 m.

Etymology. — The species name secta (Latin sectus, cut) refers to the base of the anthocyathus stage, which

appears to be cut from the anthocaulus.

Description. — Anthocaulus stage unknown, probably not collected because of its small size. Corallum

(anthocyathus) elongate-conical (edge angle 24°-36°), straight, and always unattached, the pedicel narrowing to a

small circular to elliptical detachment scar 2.0-2.5 x 2. 2-3.4 mm in diameter. Holotype 15.7 x 12.9 mm in

calicular diameter and 21 mm in height; largest calice (KARUBAR stn 3) 17.6 mm in GCD, and largest corallum

(Musorstom 3 stn 126) 30.0 mm in height. Calice elliptical: GCD:LCD = 1.19-7.24-1.39 (N=6). Costae flat to

onij slightly convex, each about 0.85 mm wide near calice and separated by very thin, shallow striae. Costae

90

S. D. CAIRNS & H. ZIBROWIUS

covered with low, rounded granules — 4 or 5 occurring across width of a costa near calice. Thin porcellaneou

epithecal bands present on well-preserved coralla. Corallum white to light reddish-brown.

Septa usually hexamerally arranged in 4 complete cycles of 48 septa according to formula: Si-2>S3>S4; onb

one corallum (Karubar stn 3) having 14 sectors, or 56 septa and 14 pali. Septa evenly and widely spaced

0.6-0.7 mm from one another. Si -2 only slightly exsert (1.1 -1.2 mm), extending about 3/4 distance to columella

having moderately sinuous inner edges. S3 and S4 equally exsert (about 0.8 mm), the S3 about 3/4 width of S1-2

having very sinuous inner edges. S4 3/4 to 4/5 width of an S3, inner edges moderately sinuous. Septal face

covered with small, pointed granules arranged in closely-spaced, parallel rows in a fan system radiating from th

thecal wall. An elliptical crown on 12 lamellar P3 encircle the elongate columella, each palus having a slightl

sinuous inner and outer edge and measuring 1.0- 1.3 mm in width. Fossa shallow to moderate in depth. Columell

consists of 4-9 slender, twisted lamellae.

Remarks. — The nature of the base of C. secta is unique within Caryophyllia. If only one specimen wer

found to be truncate, with a small, basal scar, it might be interpreted as an accidental fracture from the substraturr

but its presence in all specimens reported above suggests that it is a normal feature of the species. Basal scars ar

assumed to have resulted from transverse division from a small, attached anthocaulus stage; however, no such stag

has yet been identified for this species.

Distribution. — Philippines: Lubang Island; Verde Island Passage; Sulu Sea (Semirara Islands am

Zamboanga Peninsula); 220-366 m. Indonesia-. Banda Sea (Kai Islands); Bali Sea; 240-278 m.

Caryophyllia (C.) lamellifera Moseley, 1881

Caryophyllia lamellifera Moseley, 1881: 140-141, pi. 1, figs 7a-b. — Cairns, 1995: 51-52 , pi. 9, fig. i, pi. 1(,

figs a-c (synonymy).

Material EXAMINED. — Philippines. MUSORSTOM 3: stn 131, 2 (USNM 96791). — Stn 134, 5 (MNHN).

Indonesia. Deki: stn 24, 1 (NNM 22747).

Karubar: stn 18, 2 (MNHN). — Stn 30, 1 (MNHN).

Type Locality. — "Challenger" stn 170: 29°55'S, 178°14'W (north of Macauley Island, Kermadec Ridge),

1152 m.

DIAGNOSIS. — Corallum elongate-conical to trochoid; largest known specimen 15.5 mm in GCD (CAIRNS

1995); largest reported herein (MUSORSTOM 3 stn 134) 11.5 mm in GCD and 17.2 mm in height. PD:GCD =

0.38-0.53. Theca glisteny, covered with closely spaced, transverse ridges. Most coralla have brown thecal striping

and brown Si -2. Septal symmetry variable, most specimens reported above having 4 cycles of heptamerally

arranged septa (56 septa, 14 pali), but one small specimen (Karubar stn 18) having hexamerally arranged septa

(48 septa, 12 pali). Septal formula: Si>S2>S3>S4, the S1-2 highly exsert, forming triangular calicular lancets

with their flanking S4. Inner edges of all septa slightly sinuous. 12 to 14 robust P3 encircle a fascicular columella

composed of 1-19 very slender twisted lamellae. Fossa of moderate depth.

REMARKS. — This species is more fully described and illustrated by CAIRNS (1995). It is distinguished by its

relatively large, pigmented corallum and its transverse thecal ridging (see key).

DISTRIBUTION. — Philippines: northwestern Panay (Sulu Sea and Tablas Strait); 95-120 m. Indonesia: Banda

Sea (Kai Islands); 100-212 m. Elsewhere: Kermadec Ridge; southern Norfolk Ridge; Lord Howe Island; Taupo

Tablemount, Tasman Sea; 89-1 152 m.

Caryophyllia (C.) transversalis Moseley, 1881

Figs 6 f-h

Caryophyllia clavus var. transversalis Moseley. 1881: 134-135, pi. 1, figs 2, 2a. — Alcock. 1902c: 9-10.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

91

MATERIAL EXAMINED. — Indonesia. "Challenger": stn 192, syntype of C. clavus transversalis (BMNH

1880.1 1.25.23).

"Siboga": stn 12, 1 (ZMA). — Stn 256, 1 (ZMA).

Deki: stn 3, 24 (NNM 22704). — Stn 6, 2 (NNM 22703). — Stn 32, 1 (NNM 22705). — Stn 41, 1 (NNM 22706). —

Stn 42, 7 (NNM 22707). — Stn 44, 12 (NNM 22708). — Stn 46, 2 (NNM 22709). — Stn 48, 2 (NNM 22710). — Stn 50,

18 (NNM 22712). — Stn 62 , 1 (NNM). — Stn 63, 1 (NNM 22713).

Karubar: stn 2, 8 (USNM 96793). — Stn 7, 6 (POLIPI). — Stn 67, 2 (MNHN). — Stn 68, 4: 2 (MNHN), 2 (USNM

96794). _ stn 79, 4 (POLIPI). — Stn 84, 4: 3 (MNHN), 1 (USNM 96796).

TYPE Locality. — "Challenger" stn 192: 5°42'S, 132°25'E (Kai Islands, Banda Sea), 235 m.

Description. — Corallum trochoid (edge angle 53°-65°), straight, and always attached through a narrow

pedicel 1.6-2. 9 mm in diameter (PD:GCD = 0.11-0.19). Substratum often a small scaphopod, gastropod, or

bivalve shell. Calice elliptical: GCD:LCD = 1.21-7.27-1.36 (N=6). Largest known specimen (Karubar stn 79)

21.6 x 15.9 mm in calicular diameter and 23.8 mm in height, with a pedicel diameter of 2.7 mm. Costae flat to

slightly convex, each about 1 .0 mm wide near the calice of a large specimen, and separated by thin, shallow striae.

Costae covered with low, rounded, glisteny granules — 4 or 5 occurring across a costa. Corallum near calice of

large specimens beige, becoming white or discoloured 4-5 mm below calicular edge.

Septa, even of coralla as small as 8.8 mm GCD, arranged accordingly: 14:14:28 (56 septa and 14 pali);

however, 1 large corallum from Karubar stn 67 has 16 primary septa and 16 pali, and altogether in 60 septa.

Primary septa highly exsert (up to 3.5 mm), with straight, vertical inner edges that attain the columella low

in fossa. Secondary septa least exsert (about 1.3 mm), about 3/4 width of a primary, with highly sinuous inner

edges. Tertiary septa about 2 mm exsert, each pair adjacent to a primary fusing to that primary in a low, triangular

calicular lancet, producing a serrate calicular margin. Tertiary septa about 4/5 width of a secondary, with straight

inner edges. An elliptical crown of 14-16 P3 encircles an elongate columella, each palus about 1.7 mm wide,

having highly sinuous edges. Fossa of moderate depth, containing a columella of 9-15 slender, closely-spaced

twisted lamellae.

Remarks. — Caryophyllia transversalis is similar to C. secta sp. nov., but can be distinguished by its

attached, trochoid (vs unattached, ceratoid) corallum; lanceted calicular edge; and tendency to have 14 (vs 12)

primary septa. Among the Recent Caryophyllia , attached species having decatetrameral symmetry (x 14) are not

common, only three being listed by Cairns (1991, table 3), all of those species having a range of septal

symmetry that includes decatetrameral.

Distribution. — Indonesia : Banda Sea (Kai Islands); Arafura Sea (south of Tanimbar Islands); Bali Sea; 210-

397 m.

Caryophyllia (C.) rugosa Moseley, 1881

Caryophyllia rugosa Moseley, 1881: 141-143, pi. 1, figs 8a-b. — FAUSTINO, 1927: 70-71, pi. 8, figs 12-14. — Cairns

& Keller, 1993: 236, pi. 3, fig. 1. — Cairns, 1994: 47, pi. 20, fig. i, pi. 21, fig. a (synonymy); 1995: 43-44, pi. 6,

fig. h, pi. 7, figs a-c.

Material EXAMINED. — Philippines. "Albatross": stn 5172, 1 (USNM 96797). — Stn 5217, 1 (USNM 96798).

Musorstom 2: stn 33, 1 (MNHN).

Indonesia. Deki: stn 3, 1 (NNM 22746).

Karubar: stn 18, 9: 5 (MNHN), 4 (USNM 96799).

Type Locality. — "Challenger" stns 192 and 201: Banda and Sulu Seas, 187-230 m.

Diagnosis. Corallum elongate-conical to cylindrical, firmly attached through a thick pedicel (PD:GCD =

0.3-0.6), largest known corallum only 8.5 mm in GCD (Cairns, 1994). Theca covered with fine transverse ridges.

ci 'a octamerally arranged in 3 cycles (Si>S2>S3), resulting in 32 septa. Inner edges of Si and S2 extremely

sinuous. Septal faces bear blunt granules, some fused into short carinae. A crown on 8 very sinuous pali occurs

92

S. D. CAIRNS & H. ZIBROWIUS

before the secondary septa, encircling a fascicular columella composed of 1-4-15 slender, twisted lamellae. Foss i

shallow.

Remarks. — Caryophyllia rugosa is a very common azooxanthellate coral found throughout the Indo-We t

Pacific, characterised by having a relatively small corallum, 3 cycles of very sinuous, octamerally arranged sept ,

and a transversely ridged theca. The 5 species of attached Caryophyllia having octamerally symmetrical septa ai

discussed in the following species account.

DISTRIBUTION. — Philippines-. Verde Island Passage; Ragay Gulf; Sulu Sea (Sulu Archipelago); 137-581 n

Indonesia: Banda Sea (Kai Islands); 205-256 m. Elsewhere: widespread in Indo-West Pacific from South Africa i i

the Hawaiian Islands, including Japan and the Kermadec Islands; 71-508 m.

Caryophyllia (C.) octonaria sp. nov.

Figs 7 a-b

MATERIAL EXAMINED/TYPES. — Philippines. "Albatross": stn 5527, 1 paratype (USNM 97060).

Musorstom 1: stn 64, holotype (MNHN).

Musorstom 2: stn 2, 2 paratypes (MNHN).

Musorstom 3: stn 88, 3 paratypes (USNM 96802). — Stn 90, 1 paratype (USNM 96803). — Stn 100, 3 paratyp< s

(MNHN). — Stn 101, 1 paratype (MNHN). — Stn 102, 8 paratypes (USNM 96804).

Type Locality. — Musorstom 1 stn 64: 14°01'N, 120°16’E (Lubang Island, Philippines), 194-195 m.

ETYMOLOGY. — The species name octonaria (Latin octonarius, consisting of 8 units) refers to the octamer 1

symmetry of this species.

DESCRIPTION. — Corallum conical, straight to slightly bent, and always attached by a slender pedic I

(PD:GCD = 0.26-0.5/-0.40). Coralla always attached to gastropod or bivalve shells; one dead corallun

(MUSORSTOM 2 stn 2) secondarily incorporated into a Xenophora shell. Calice slightly elliptical: GCD:LCD =

1.06-/. 08- 1.1 3. Largest known specimen (the holotype) 7.8 x 8.4 mm in calicular diameter, 13.9 mm in heigh ,

and 2.2 mm in pedicel diameter. Costae not well defined, the theca often smooth and porcellaneous near calic .

Costae on lower corallum equal in width (0. 5-0.6 mm) and flat, separated by very shallow, narrow intercostal strire

and covered with low, rounded granules. Theca light grey-brown or reddish-brown; septa, pali, and columella

usually white.

Septa octamerally arranged in 3 complete cycles (32 septa) in all specimens examined. Eight primary sepia

relatively highly exsert (1.7-2. 2 mm), extend about 3/4 distance to columella, having moderately sinuous inner

edges. Remaining septa equally exsert (1.0- 1.2 mm), the 8 secondaries being about 5/6 width of a primary, also

having moderately sinuous inner edges. Tertiary septa of small coralla equal to or slightly less wide than

secondaries, but at a later stage its tertiaries become slightly wider than its secondaries (Si>S3>S2). Inner edges of

tertiaries only slightly sinuous. Septal faces covered with prominent, pointed granules. Fossa quite shallow,

containing an elliptical crown of 8 sinuous pali before the secondaries, each palus 1.0- 1.1 mm wide. The fascicular

columella consists of a field of 6-9 twisted elements strongly fused together basally.

Remarks. — There are 4 previously described octamerally symmetrical, attached Caryophyllia: C. rugosa

Moseley, 1881; C. barbadensis Cairns, 1979; C. octopali Vaughan, 1907; and C. marmorea Cairns, 1984, all of

which are characterised by having their tertiaries equal to or wider than their secondaries. C. octonaria is similar lo

C. octopali, but differs in having a narrower pedicel (PD:GCD = 0.26-0.40 vs >0.5 for C. octopali ), a more flared

corallum (that of C. octopali is elongate and subcylindrical), and in having relatively exsert septa. In the last

character C. octonaria resembles C. octopali var. incerta Vaughan, 1907, but that form differs in having hexameral

or heptameral symmetry, a very reduced columella, and a quasicolonial habit.

Distribution. — Philippines: Lubang Island; Bohol Sea; 186-194 m.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

93

Caryophyllia (C.) hawaiiensis Vaughan, 1907

Caryophyllia hawaiiensis Vaughan, 1907: 76, pi. 5, figs 4a-b. — CAIRNS, 1984: 11; 1995: 44-45 , pi. 7, figs d-f.

MATERIAL EXAMINED. — Philippines. " Albatross stn 5310, 1 (USNM 96805).

Musorstom 2: stn 33, 3 (MNHN).

Musorstom 3: stn 117, 1 (MNHN).

Indonesia. Deki: stn 25, 1 (NNM 22744). — Stn 53, 1 (NNM 22745).

Corindon 2: stn 248, 2 (MNHN).

Karubar: stn 1, 1 (POLIP1). — Stn 22, 1 (USNM 96807).

Type Locality. — "Albatross" stn 3838: 21°04'05"N, 157°10'35"W (off Molokai, Hawaiian Islands), 168-

388 m.

Diagnosis. — Corallum elongate-conical and attached; largest known specimen 11.6 mm in GCD (Cairns,

1995); largest specimen reported herein (" Albatross " stn 5310) 8.4 mm in GCD. PD:GCD = 0.34-0.49. Costae

absent or poorly defined, the theca porcellaneous and covered with low, rounded granules. Theca and most septa

speckled with dark brown pigmentation. Septa pentamerally arranged in 4 cycles (5:5:10:20, Si>S2>S4>S3)

resulting in 40 septa and 10 pali. The asymmetrical arrangement of the 5 Si results in a pentagonal disposition of

major septa, with only one of the Si being aligned with the greater calicular axis. S] highly exsert, forming tall

calicular lancets. A crown of 10 P3 encircles a fascicular columella consisting of 6-18 slender, twisted lamellae.

Remarks. — Caryophyllia hawaiiensis is unique among the Caryophyllia in having pentamerally arranged

septa. It is more fully described and illustrated by Cairns (1995).

DISTRIBUTION. — Philippines: Verde Island Passage; Midoro Strait; 97-130 m. Indonesia: Makassar Strait;

Banda Sea (Kai Islands); 85-170 m. Elsewhere: South China Sea (Vereker Banks near Pratas Island); Hawaiian

Islands; Kermadec Ridge; 126-279 m.

Caryophyllia (C.) quadragenaria Alcock, 1902

Caryophyllia quadragenaria Alcock, 1902a: 91-92; 1902c: 10, pi. 1, figs 4, 4a. — Cairns, 1994: 46-47, pi. 20,

figs c-h, pi. 41, figs c-d (synonymy); 1995: 45-46, pi. 7, figs g-h (synonymy).

Material EXAMINED. — Indonesia. Deki: stn 46, 2 (NNM 22741). — Stn 59, 1 (NNM 23080).

Snellius 2: stn 81.2, 1 (NNM 23079).

Karubar: stn 2, 16 (MNHN). — Stn 3, 5 (USNM 96811).

South China Sea. "Albatross": stn 5313, 2 (USNM 96809). — Stn 5317, 6 (USNM 96810).

Type Locality. — "Siboga" stns 90, 251, and 289: Makassar Strait, Banda, and Timor Seas, 54-281

m.

Diagnosis. Corallum elongate-conical and attached, straight, and relatively small; largest Indonesian

specimen (Karubar stn 3) 8.9 x 12.0 mm in calicular diameter and 15.4 mm in height. PD:GCD = 0.1 1-0.39.

Costae well defined only near calice. Septa decamerally arranged in 3 cycles (10: 10:20, 40 septa; Si>S3>S2). Ten

broad, highly sinuous P2 encircle a fascicular columella of 3-1 1 twisted lamellae.

Remarks. Comparisons of C. quadragenaria to the other 6 species of Caryophyllia that have decameral

1995)1Ctry ^ ma^C ^ CAIRNS (1995), and the species is more fully described and illustrated by CAIRNS (1994,

LUSTribution. — Indonesia: Makassar Strait; Banda Sea (Kai and Tanimbar Islands); Timor Sea (Timor); 1 12-

i i T \ Jewliere '■ South China Sea (north of Pratas Island); Japan (Honshu, Shikoku, and northern Ryukyu

Islands); New Zealand; 54-296 m.

94

S. D. CAIRNS & H. ZIBROWIUS

Caryophyllia (C.) scobinosa Alcock, 1902

Caryophyllia cultrifera Alcock, 1902a: 89-90; 1902c: 7-8, pi. 1, figs 1, la. — FAUSTINO, 1927: 67-68, pi. 8, figs 8-9.

Caryophyllia scobinosa Alcock, 1902a: 90; 1902c: 8, pi. 1, figs 2, 2a. — FAUSTINO, 1927: 68-69, pi. 8, figs 10-11. --

Cairns, 1994: 45-46 (in part: pi. 20, figs a-b); 1995: 52-53, pi. 10, figs g-i, pi. 11, figs a-c (synonymy).

MATERIAL EXAMINED. — Philippines. " Albatross stn 5348, 1 (USNM 96812). — Stn 5444, 1 (USNM 96813

— Stn 5445, 12 (USNM 96814). — Stn 5447, 14 (USNM 96815). — Stn 5551, 4 (USNM 96816). — Stn 556‘

1 (USNM 96817). — Stn 5565, 2 (USNM 96818).

MUSORSTOM 1 : stn 44, 1 (MNHN). — Stn 47, 10 (MNHN).

Musorstom 2: stn 25, 1 (USNM 96821).

Indonesia. "Albatross": stn 5650, 2 (USNM).

CORINDON 2: stn 240, 1 (USNM 96822).

Karubar: stn 91, 2 (POL1PI).

TYPE Locality. — "Siboga" stns 45 and 102: Flores and Sulu Seas, 535-794 m.

Diagnosis. — Corallum ceratoid (usually curved between 45°-90°) and free; rarely more than 20 mm in GCE

Ci -2 ridged, C3.4 flat and granular. Septa hexamerally arranged in 4 complete cycles: Si-2>S4^S3- Si -2 highl

exsert, forming calicular lancets with adjacent pairs of S4. A crown of wide, lamellar P3 encircles a fasciculi -

columella of 4-14 relatively broad, twisted elements.

Remarks. — Caryophyllia scobinosa is more fully described, illustrated, and discussed by Cairns (199‘ ,

1995), and compared to Caryophyllia cornulum sp. nov. in the following account.

The second author believes that this taxon is a species complex, composed of several morphologically distint 1

species that are also bathymetrically distinct.

DISTRIBUTION. — Philippines: Lubang Island; South China Sea (Palawan); Lagonoy Gulf; Sulu Sea (Sul 1

Archipelago); 353-1270 m. Indonesia: Makassar Strait; Banda Sea (Gulf of Bone, Sulawesi); Timor Sea (south cf

Babar Islands); 675-988 m. Elsewhere: southwestern Indian Ocean; Queensland; Lord Howe Rise; Tonga an I

Samoa; 535-1276 m.

Caryophyllia (C.) cornulum sp. nov.

Figs 7 d-e

MATERIAL EXAMINED/TYPES. — Indonesia. "Albatross": stn 5606, 2 paratypes (USNM 96823). — Stn 5670,

16paratypes (USNM 96824).

Corindon 2: stn 220, holotype (MNHN), 12 paratypes (POLIPI). — Stn 241, 1 paratype (USNM 96826). — Stn 286,

1 paratype (MNHN).

Japan. "Tansei-Maru": stn KT86-16-F, 2 paratypes (USNM 96827). — Stn KT90-13-T6, 6 paratypes (USNM

96828).

Type Locality. — Corindon 2 stn 220: 0°13.8'S, 1 18°12.7'E (Makassar Strait), 2350 m.

ETYMOLOGY. — The species name (Latin cornulum , small horn) refers to the shape of the corallum. The name

is treated as a noun in apposition.

Description. — Corallum ceratoid, unattached, and rarely curved more than 45°, the base of all but

1 specimens being strongly eroded; the base of one specimen (CORINDON 2 stn 220) is 1.5 mm in diameter and

shows 6 protosepta in the basal disc. Corallum of most specimens medium to small in size, the holotype only

15.3 x 17.9 mm in calicular diameter and 16.5 mm in height; however, the 8 Japanese specimens are larger, one

up to 27 mm in GCD. Calice slightly elliptical: GCD:LCD = 1.06-1.18. Costae well preserved only near calice,

where they are about 0.6 mm wide, separated by thin, shallow striae and covered with small granules. Below 2 mm

Source : MNHN Paris

AZOOX ANTHELL ATE SCLERACTINIA

95

from the calicular edge the corallum is usually eroded, becoming grayish in color. Primary costae sometimes

slightly ridged.

Septal arrangement variable, the most common complement being: 14:14:28, and 14 pali, shared by 1 1 of the

30 specimens analyzed. Nine specimens have 16 primary septa (up to 64 septa); the 8 Japanese specimens have

only 12 primary septa (48 septa); 1 has 13 primary septa (52 septa); and 1 has 18 primary septa (72 septa). It is

common, however, for 1 or more sectors to lack its tertiary pair of septa and corresponding palus, resulting in

septal and palar complements that fall short of the expected for that number of primary septa (e.g., a corallum

having 16 primary and 16 secondary septa, may have only 28 or 30 (instead of 32) tertiary septa and only 14 pali).

Primary septa little exsert (1.0- 1.1 mm), extending only about 1/2 distance to columella, having straight to

slightly sinuous inner edges. Secondary and tertiary septa equally exsert (0.7-0.8 mm), producing a low, serrate

calicular margin. Secondaries about 1/2 width of a primary septum, having slightly sinuous inner edges; tertiaries

equal to or slightly wider than a secondary, having slightly sinuous inner edges. A crown of 12-18 (usually 14-16)

wide pali occurs before the secondary septa, the pali usually wider (2.2-2. 8 mm) than septa they border. Palar edges

slightly sinuous, their faces bearing a granulation coarser than that of septa. Columella fascicular, composed of 1-7

(usually 2 or 3) broad, twisted elements, often arranged in a line.

Remarks. — Caryophyllia cornulum is similar to C. scobinosa but differs in having: much less exsert septa

that do not form calicular lancets; a tendency to have 14-16 primary septa (vs 12 for the typical form); a less

curved corallum with a strongly eroded base; and a deeper bathymetric range (1525-2576 m vs 353-1276 m for

C. scobinosa). Although some large specimens of C. scobinosa occasionally have 14 or 16 primary septa

(CAIRNS, 1995), even small specimens of C. cornulum have 14-16 primary septa, only the Japanese specimens

having 12 primary septa.

DISTRIBUTION. — Indonesia: Makassar Strait; Gulf of Tomini, Sulawesi; 1525-2350 m. Elsewhere : Shikoku,

Japan; 2576-2603 m.

Caryophyllia (C.) ambrosia Alcock, 1898

Caryophyllia ambrosia Alcock, 1898: 12, pi. 1, figs 1, la. — ZlBROWtus, 1980: 63-65, pi. 25, figs A-K (synonymy). —

Cairns & Keller, 1993: 234, pi. 3, fig. 14 (synonymy). — Cairns, 1994: 48, pi. 21, figs d-h; 1995: 53-54, pi. 1 1,

figs d-e.

Material EXAMINED. — Philippines. Musorstom 1: stn 49, 4 (MNHN). — Stn 54, 1 (MNHN).

Musorstom 3: stn 94, 1 (MNHN).

Indonesia. "Hakuho Maru": stn KH72-1-26, 3: 1 (USNM 96835), 2 (ORI).

Corindon 2: stn 240, 4 (USNM 96834).

Karubar: stn 40, 1 (USNM 96829). — Stn 56, 3 (MNHN). — Stn 57. 1 (POLIP1). — Stn 72, 1 (POLIPI). — Stn 87.

10 (USNM 96832). — Stn 89, 3 (MNHN). — Stn 91, 2 (MNHN).

Type Locality. — Laccadive Sea, Arabian Sea, 1829-1957 m.

Diagnosis. — Corallum massive (GCD up to 60 mm) and curved, having a pointed and/or eroded base. Ci-3

usually slightly ridged. Coralla contain 18-30 (but usually 24) primary septa, an equal number of secondaries, and

twice that number of tertiaries, resulting in calices with 72-120 (but usually 96) septa. Tertiary septa equal to or

wider than secondaries, but always more exsert than secondaries, fusing with their flanking primary septum to

form highly exsert lancets. A broad palus occurs before each secondary septum, each of the 18-24-30 pali usually

wider than the septa they border. Fascicular columella composed of numerous, broad, twisted elements.

Remarks. — Cairns (1994, 1995) redescribed what he believed to be the nominate subspecies of

C. ambrosia. It appears to be a widespread Pacific and Atlantic species with regional variation regarding corallum

size and number of septa (Cairns, 1995). The Philippine/Indonesian specimens are the largest thus far recorded, up

to 60 mm in GCD and containing 130 septa, which makes them most similar to those reported from New Zealand

96

S. D. CAIRNS & H. ZIBROWIUS

(CAIRNS, 1995). However, the second author believes that "C. ambrosia" may represent a species complex, th

coralla of which are similar to virtually indistinguishable, and the species of this complex having more restricte 1

geographic and bathymetric ranges. Characteristics of the soft parts or molecules may be necessary to resolve th'

taxonomic issue.

C. ambrosia is compared to C. grandis in the account of the latter species.

DISTRIBUTION. — Philippines: Lubang Island; 842-975 m. Indonesia: Makassar Strait; Arafura Sea (south <

Tanimbar Islands); Timor Sea (south of Leti Islands); 468-1048 m. Elsewhere : amphi-Atlantic; Indo-West Pacifn

including Japan and New Zealand; 311-2670 m.

Caryophyllia (C.) grandis Gardiner & Waugh, 1938

Figs 7 g-h

Caryophyllia grandis Gardiner & Waugh, 1938: 177, pi. 1, fig. 2. — Cairns & KELLER, 1993: 234 (synonymy).

MATERIAL EXAMINED. — Indonesia. "Albatross": stn 5589, 3 (USNM 96837). — Stn 5590, 15 (USNM 96836

— Stn 5592, 1 (USNM 86838).

Karubar: stn 39, 2 (USNM 96839). — Stn 40, 4 (POL1PI). — Stn 59, 8 (MNHN). — Stn 62, I (MNHN). — Stn 7< ,

3 (MNHN). — Stn 71, 4 (MNHN). — Stn 75, 2 (MNHN).

Type Locality. — John Murray Expedition stn 145: 4°58'42"S, 73°16'24"E (Fadiffolu Atoll), 494 m.

DIAGNOSIS. — Corallum large (GCD up to 50 mm), sometimes free and curved, but more often straight t >

slightly bent, and attached by a narrow pedicel 1.8-6. 5 mm in diameter. Ci-3 usually slightly ridged. Upper thee i

and septal faces light beige; lower theca white or discoloured. Septa usually hexamerally arranged in 5 cycle t

(Si -3>S4>Ss), but some large coralla have up to 28 primary septa and a total of 1 12 septa. S5 narrower than S. .

but more exsert, fusing with their adjacent Si -3 to form highly exsert lancets. 24 P3, which are usually narrower

than the S3 they border, form a crown encircling a fascicular columella composed of broad, twisted lamellae.

Remarks. — Caryophyllia grandis is distinguished from C. ambrosia by its: attached, reinforced pedice :

highest cycle septa (i.e., S5) that are narrower than their S4; narrower pali; brownish theca; and usually straighte ,

less curved, corallum. In the Indonesian region C. grandis is also found at shallower depths than C. ambrosia: 25 1 -

567 m vs 468-1048 m, respectively.

DISTRIBUTION. — Indonesia: Arafura Sea (southeast of Tanimbar Islands); 251-567 m. Elsewhere: Malaysia

(Celebes Sea off Sabah); Indian Ocean from South Africa to western Sumatra; 183-595 m.

Subgenus CARYOPHYLLIA (ACANTHOCYATHUS) H. Milne Edwards & Haime, 1848

Key to the six species of Caryophyllia (Acanlhocyathus)

known from the Philippine/Indonesian region

1. Corallum bears thecal edge spines . 2

— Corallum bears 1 or 2 thecal edge crests (but no spines) . 4

2. Spines occur on both thecal edges, and sometimes even on thecal faces . 3

— Spines occur only on convex thecal edge . C. (A.) dentata

3. Corallum with 14 primary septa (>56 septa) and 14 pali; C1-2 rounded; 3 size classes of

septa (14: 14:28); thecal spines circular to elliptical in cross section . C. (A.) grayi

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

97

— Corallum with 12 primary septa (48 septa) and 12 pali; Ci-2 ridged; 4 size classes of septa

(6:6:12:24); thecal edge spines flattened in cross section . C. (A.) spinigera

4. Ci -2 ridged; S4^S3 in width . C. (A.) spinicarens

— Ci -2 rounded; S4^S3 in width . 5

5. Corallum with 14 primary septa (56 septa) and 14 pali; both thecal edges crested; up to

25 mm in GCD . C. (A.) karubarica

— Corallum with 12 primary septa (48 septa) and 12 pali; only convex thecal edge crested;

smaller, only up to 16 mm GCD . C. (A.) unicristata

Caryophyllia (A.) grayi ( H. Milne Edwards & Haime, 1848)

Figs 7 c, f, i

Acanthocyathus grayi H. Milne Edwards & Haime, 1848a: 293, pi. 9, figs 2, 2a. — ALCOCK, 1898: 15. — VAN DER HORST,

1931: 6. — Umbgrove, 1938: 264-265; 1950: 641-642, pi. 81, figs 27-32 (synonymy). — Wells, 1984: 209, pi. 2,

figs 5-9. — Zou, 1988: 76, figs 8-9.

Caryophyllia (A.) grayi - CAIRNS, 1994: 49, pi. 21, figs i-k (synonymy).

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5381, 1 (USNM 96843). — Stn 5593, 1 (USNM 96844)

"Hakuho Maru": stn KH72-1-50, I (USNM 96855).

Musorstom 1: stn 45, 2 (USNM 96851). — Stn 56, 10 (MNHN). — Stn 72, 5 (USNM 96847).

Musorstom 2: stn 29, 1 (MNHN). — Stn 47, 4 (USNM 96848).

Musorstom 3: stn 99. 1 (MNHN). — Stn 107, 1 (USNM 96850). — Stn 108, 1 (MNHN). — Stn 124, 3 (MNHN). —

Stn 131, 22: 1 1 (MNHN), 1 1 (USNM 96846). — Stn 140, 8 (MNHN).

Indonesia. "Siboga": stn 204, 14 (ZMA Coel. 1159). — Stn 289, 6 (ZMA Coel. 1168).

Deki: stn 4, 2 (NNM 22695). — Stn 6, 2 (NNM 22696). — Stn 25, 2 (NNM 22697). — Stn 44, 2 (NNM 22698). —

Stn 49, 2 (NNM 22699). — Stn 53, 7 (NNM 22700).

Mortensen's Java-S.A. Expedition: stn 1, 1 (ZMUC). — Stn 5, 21 (ZMUC). — Stn 6, II (ZMUC). — Stn 8,

37 (ZMUC). — Stn 9, 18 (ZMUC).

Corindon 2: stn 208, 2 (USNM 96852). — Stn 216, 4 (POL1PI). — Stn 251, 5 (MNHN). — Stn 261, 1 (MNHN).

Snellius 1: unnumbered station, 8.04.1929, 7°33’S, 1 14°36’E, 200 m, 2 (NNM 22701).

Japan. Syntype (BMNH 1840.9.29.42).

Type Locality. — Unknown.

Diagnosis. — Corallum ceratoid, compressed (GCD:LCD = 1.3- 1.5), and usually curved in plane of GCD.

Largest Philippine specimen (MUSORSTOM 1 stn 56) 16.7 x 21.4 mm in calicular diameter and 24.9 mm in

height. Corallum either attached to a small object through a slender pedicel 1.5- 1.8 mm in diameter or secondarily

unattached. Thecal edges rounded, the concave edge with 2 or 3 elongate spines, the convex edge bearing

3-5 elongate spines. Edge spines circular to elliptical in cross section. Costae rounded, not ridged. Septa usually

arranged in 14 sectors: 14:14:28 (56 septa), with 14 pali; however, a large specimen from Japan (BMNH

1840.9.29.42) with a GCD of 22.3 mm has 16 pali and a septal complement of 64. Primary septa highly exsert,

forming small (up to 3 mm) calicular lancets. Pairs of quaternary septa sometimes present in end half-systems.

A distinct crown of 14 pali occurs before secondary septa, encircling an elongate fascicular columella composed of

7-10 highly fused (basally), twisted elements.

Remarks. Caryophyllia (A.) grayi is distinguished from C. (A.) spinigera by having: more septa and pali

(usually 56 septa and 14 pali vs 48 septa and 12 pali for C. spinigera ); low, rounded Ct-2 (not ridged as in

' sP'n'8era)< a gently curved corallum in the plane of the GCD and thus an asymmetrical arrangement of thecal

e ge spines, cylindrical, not spatulate, thecal edge spines; the absence of lateral face thecal spines; a brownish

cora urn (corallum of C. spinigera is more gray); and equally developed primary septa (the 6 St of C. spinigera

are muc i more exsert than their 6 S2). C. (A.) grayi is more fully described and illustrated by Cairns (1994).

aryophyUia (A.) guangdongensis Zou, 1984, known only from the northern South China Sea (167-179 m), is

in erine iate between C. spinigera and C. grayi . It resembles C. spinigera in having spatulate edge spines and only

98

S. D. CAIRNS & H. ZIBROWIUS

48 septa with 12 pali, but differs from C. spinigera (and is thus similar to C. grayi) in having: convex, rounde I

Ci-2, equal-sized Si and S2, less exsert S1-2, and no thecal face spines.

Five more species are recognised in this subgenus: 4 having edge crests, not spines: C. (A.) spinicaren >

(Moseley, 1881), C. (A.) zanzibarensis Zou, 1984, C. (A.) karubarica sp. nov. and C. (A.) unicristata sp. nov .

and one having spines on only one thecal edge: C. (A.) dentata (Moseley, 1881). A 9th possible species, reportc 1

as Caryophyllia laoagana Smith, 1913, from the Pleistocene of the Philippines, may also belong to th

subgenus, but the single type specimen is very poorly preserved. The second author suggests that only the i

species having edge spines should be placed in the subgenus Acanthocyathus , the 4 species listed that have ed<

crests belonging more properly to the nominate subgenus.

DISTRIBUTION. — Philippines: Lubang Island; Verde Island Passage; Sibuyan Sea; Sulu Sea (Semirara Islam :

and west of Panay); 50-268 m. Indonesia: Makassar Strait; Banda Sea (Kai Islands and southeastern Sulawesi ;

Timor Sea (south of Timor); Bali Sea and Bali Strait; 50-150 m. Pleistocene of Talaud, Timor, and Vanua 1

(UMBGROVE, 1938, 1950). Elsewhere: South Africa; Bay of Bengal; Andaman Islands; South China Sea (Chariot :

Bank); Japan (Honshu and northern Ryukyu Islands); 37-490 m.

Caryophyllia (A.) dentata (Moseley, 1881)

Figs 8 a-d

Acanthocyathus sp. - MOSELEY, 1876: 550.

Acanthocyathus dentatus Moseley, 1881: 143, pi. 2, figs 7a-c.

MATERIAL EXAMINED. — Indonesia. "Challenger": stn 174, holotype (BMNH 1880.11.25.42).

Deki: stn 2, 1 (NNM 23077). — Stn 3, 5 (USNM 96858). — Stn 6, 4 (NNM 22752). — Stn 48, 2 (NNM 22753). --

Stn 63, 2 (NNM 22754). — Unnumbered station between Neira and Lontor, Banda Islands, 70-90 m, 1 (NNM 23076).

Karubar: stn 49, 1 (USNM 96856). — Stn 65, 5: 2 (MNHN), 3 (USNM 96857).

TYPE Locality. — "Challenger" stn 174D: 19°05'50"S, 178°I6'20"E (Fiji), 210 m.

DESCRIPTION. — Corallum compressed (GCD:LCD = 1.3- 1.5) and usually slightly curved in plane of GCD,

but rarely more than 90°. Holotype (BMNH 1880.11.25.42) 11 mm in GCD; largest known specimen (DEKI

stn 3) 9.4 x 12.3 mm in calicular diameter. Pedicel quite small (1.2- 1.5 mm in diameter) and circular in cro s

section. C1-2 of holotype highly ridged, but on all other specimens examined costae are low and rounded, covered

with small granules. Ci on convex thecal edge of holotype, as well as other specimens reported, ridged, bearing

4 or 5 prominent spines. Corallum white.

Septa of holotype and one specimen from the DEKI (between Neira and Lontor) hexamerally arranged in 4

cycles (Si>S2>S3>S4, 48 septa); however, in remaining 10 specimens reported herein the symmetry is decameral:

10:10:20, 40 septa. S1-2 (or 10 primary septa) highly exsert as much as 2.8 mm, forming rectangular lancets with

their adjacent S4 (or tertiaries). Inner edges of S immoderately sinuous. S3 (or secondaries) about 1/2 width of S 1-2,

having very sinuous inner edges. S4 (or tertiaries) 1/3 to 1/2 width of the S3, having moderately sinuous inner

edges. 12 (10 in the case of the decameral specimens) lamellar, highly sinuous pali occur in a crown before the S3

(or secondaries). Fossa of moderate depth containing a fascicular columella composed of 3-7 twisted elements.

Remarks. — As noted in the description, the holotype differs from all other specimens herein reported by

having distinctly ridged C1-2, but this is considered to be a variable character and thus within the range of variation

for the species. More disconcerting is the difference between the hexameral symmetry of the holotype and the

decameral symmetry of most of the other specimens reported. The decamerally symmetrical specimens may

represent a different species, but more specimens of both forms (especially the hexameral form) will have to be

studied to resolve this problem.

Caryophyllia (A.) dentata is similar to C. (A.) grayi, but distinguished by having spines on only one thecal

edge, having a smaller corallum, and having fewer septa (40-48 vs 56).

Source : MNHN , Paris

AZOOX ANTHELL ATE SCLERACTINIA

99

Distribution. — Indonesia-. Banda Sea (Kai and Banda Islands); Arafura Sea (southeast of Tanimbar Islands);

90-263 m. Elsewhere : Kandavu Island, Fiji; 384 m.

Caryophyllia (A.) spinigera ( Saville Kent, 1871)

Figs 7 e-f

Acanthocyathus spiniger Saville Kent, 1871: 275-276, pi. 23, figs 1 a-c.

Caryophyllia (A.) spiniger - Cairns, 1994: 49-50, pi. 21, fig. 1. pi. 22, figs a-d (synonymy).

Material EXAMINED. — Philippines. "Albatross"-, stn 5117, 1 (USNM 96859). — Stn 5273, 3 (USNM 96860)

— Stn 5278, 1 (USNM 96861). — Stn 5353, 1 (USNM 96862). — Stn 5369, 29 (USNM 92689). — Stn 5371, 66 (USNM

92690). — Stn 5372, 1 (USNM 96863). — Stn 5374, 3 (USNM 96864). — Stn 5376, 15 (USNM 96865) — Stn 5418

1 (USNM 96866).

Musorstom 1: stn 5, 1 (USNM 96869). — Stn 10, 1 (MNHN). — Stn 11, 1 (MNHN). — Stn 20, 3 (MNHN) —

Stn 24, 5 (MNHN). — Stn 25, 9: 6 (MNHN), 3 (BMNH 1992.8.11.2). — Stn 40, 1 (USNM 96874) — Stn 61

2 (MNHN). — Stn 71, 5 (MNHN). — Stn 72, 1 (MNHN).

Musorstom 2: stn 1, 3 (MNHN). — Stn 4, 1 (USNM 96878). — Stn 10, 4 (MNHN). — Stn 1 1, 4 (MNHN), — Stn 12,

17 (MNHN). — Stn 13, 9 (USNM 96887). — Stn 18, 1 (MNHN). — Stn 20, 2 (USNM 96881). — Stn 21, 1 (USNM

96882). — Stn 62, 1 (?). — Stn 63, 26 (USNM 96883). — Stn 64. 9 (USNM 96884). — Stn 66, 95' 91 (USNM 96885)

4 (BMNH 1992.8.1 1.3). — Stn 68, 4 (MNHN).

Musorstom 3: stn 86, 1 (MNHN). — Stn 87, 9 (MNHN). — Stn 88, 1 (MNHN). — Stn 90, 2: 1 (MNHN), 1 (USNM

96890). — Stn 91, 5 (MNHN). — Stn 92, 3 (MNHN). — Stn 96, 2 (MNHN). — Stn 97, 27 (USNM 96892) — Stn 98

27 (USNM 96893). — Stn 99, 27: 17 (MNHN), 10 (USNM 96894). — Stn 100, 8: 6 (MNHN), 2 (USNM 96895). —

Stn 101, 13 (USNM 96896). — Stn 102, 5 (MNHN). — Stn 103, 3 (USNM 96897). — Stn 108, 8 (USNM 96898). —

Stn 109, 6 (MNHN). — Stn 111, 23 (MNHN). — Stn 112, 14: 3 (MNHN), 11 (USNM 96899). — Stn 120, 6 (USNM

96900). — Stn 139, 4 (MNHN).

Indonesia. Deki: stn 42, 1 (NNM 22743).

Mortensews Java-S.A. Expedition: stn 2, 4 (ZMUC).

Karubar: stn 62, 1 (USNM 96867). — Stn 79, 2 (POLIPI).

Type Locality. — Japan, depth not given.

Diagnosis/Remarks. — This species was recently described and figured (Cairns, 1994) based on Philippine

specimens, however, the following observations can be added. The largest known specimen (MUSORSTOM 1

stn 61) is 16.7 x 20.9 mm in calicular diameter, 24.6 mm in height, containing 50 septa; a slightly smaller

specimen (Musorstom 2 stn 1) of GCD 20.4 mm has 52 septa and 14 pali. All other specimens examined have

48 septa and 12 pali. Many specimens originally attached to a small, conically-shaped bryozoan colony. The

coraHum is straight, compressed (GCD:LCD = 1 .3-1.4), and has ridged Ct-2. 2 to 4 pairs of elongate, spatulate

thecal edge spines occur on each corallum, the 4th pair present only on the largest coralla. In 1 5-20% of the coralla

examined, additional spines are also present on the lateral faces of the corallum, corresponding to each of the

ateral Ci. When present, the lateral thecal spines originate at a height intermediate between the 2nd and 3rd edge

spines. n rare cases, a 2nd set of lateral thecal spines may occur above the 1 st. Lateral thecal spines very irregular

m eve opment. sometimes present on one face and not the other; sometimes present on 1 lateral C| and not the

other on the same face; and sometimes 2 occurring on one lateral Ci and only one on the other C, on the same

a^eji resence °f lateral thecal spines is not necessarily related to corallum size, since many of the largest

I™ f aVC n°ne' Scf)tal formula: Si>S2>S3>S4, the Si being highly exsert (up to 4.5 mm) and forming tall

elements anC6tS' crown °f 12 planar P3 encircle a linear-fascicular columella composed of 3-12 broad, twisted

• Catyophylha (A .) spinigera is easily distinguished from its congeners by its elongate, spatulate edge spines; it

is compared to C. grayi in the account of that species.

Island^'!1’, ITu ~ ™lipPines- Lubang ^land; Verde Island Passage; Sibuyan Sea; Bohol; Suit

of Tan , ?a(BalabaC Island); 127-347 m- Indonesia: Banda Sea (Kai Islands); Arafun

ar s ands); Bali Sea; 200-245 m. Elsewhere: Japan (locality and depth unknown).

Sea (Semirara

Sea (southeast

100

S. D. CAIRNS & H. ZIBROWIUS

Caryophyllia (A.) spinicarens (Moseley, 1881)

Figs 8 g-i

Acanthocyathus spinicarens Moseley, 1881: 143-144, pi. 2, figs 6a-c.

Caryophyllia (Premocyathus) spinacarens (sic) - Cairns & KELLER, 1993: 237 (listed only).

Material EXAMINED. — Philippines. "Challenger": stn 210, holotype (BMNH 1880.11.25.43).

"Albatross": stn 5118, 1 (USNM 96901). — Stn 5198, 1 (USNM 96902). — Stn 5256, 8 (USNM 96903). —

Stn 5260, 1 (USNM 36477). — Stn 5374, 1 1 (USNM 96908). — Stn 5403, 1 (USNM 96909). — Stn 5408, 1 (USNM

96910). — Stn 541 1, 2 (USNM 9691 1). — Stn 5412, 1 (USNM 96912). — Stn 5417, 1 (USNM 96913). — Stn 5418,

2 (USNM 96914). — Stn 5505, 2 (USNM 96915). — Stn 5508, 3 (USNM 96916). — Stn 5527, 1 (USNM 96917). —

Stn 5535, I (USNM 96918). — Sin 5536, 1 (USNM 96919). — Stn 5537, 1 (USNM 96920). — Stn 5538, 3 (USNM

97021). — Stn 5541, 1 (USNM 96922). — Stn 5565, 2 (USNM 96923).

" Galatliea stn 423, 5 (ZMUC). — Stn 436, 2 (ZMUC).

Musorstom 1: stn 20, 2 (USNM 97024).

MUSORSTOM 2: stn 45, 2 (MNHN). — Stn 46. 16: 15 (MNHN), 1 (BMNH 1992.8.1 1 . 1). — Stn 63. 6 (USNM 97027).

— Stn 83, 2 (USNM 97028).

Musorstom 3: stn 92, 1 (MNHN). — Stn 119, 1 (MNHN). — Stn 125, 1 (MNHN). — Stn 138, 1 (USNM 97029). —

Stn 139, 3 (USNM 97030). — Sin 145, 1 (MNHN).

Indonesia. "Albatross": stn 5622, 4 (USNM 97034). — Stn 5625, 2 (USNM 97032). — Stn 5626, 3 (USNM

97033).

Deki: stn 62, 1 (NNM 22748).

South China Sea. " Albatross ": stn 5301, 6 (USNM 96905). — Stn 5314, 1 (USNM 96907).

"Hakuho Maru": stn KH72-1-52, 2: 1 (ORI), 1 (USNM 97031).

Type Locality. — "Challenger" stn 210: 9°26'N, 123°34'E (off Negros, Philippines), 686 m.

DESCRIPTION. — Corallum compressed (GCD:LCD = 1.28-1.67) and often slightly curved up to 45° in plane

of LCD. Angle of lateral edges 45°-76°; angle of thecal faces 22°-50°. Largest known specimen ("Albatross"

stn 5256) 14.1 x 19.6 mm in calicular diameter and 25.5 mm in height. Pedicel small (1.8-2. 5 mm in diameter),

unattached as an adult, and circular to elongate in cross section. Convex thecal faces meet in sharp thecal edges,

which are carinate from about 4 mm above base to calicular edge. Delicate edge crests, best preserved in small

coralla, up to 1.9 mm in height, sometimes convoluted and frilled, and occasionally notched with thin

discontinuities. This delicate structure usually reduced to a low, continuous ridge in larger coralla. Remaining Ci-2

on lateral faces highly ridged, up to 0.8 mm in height. Higher cycle costae (C3-4) low and convex, covered with

small, rounded granules. Corallum reddish-brown, the colour usually well preserved only near calice or on small

specimens, the theca of larger coralla often discoloured or partly black.

Septa usually hexamerally arranged in 4 complete cycles (Si-2>S4>S3), with 12 pali, but large coralla may

have 14 primary septa and 14 pali (56 septa), and coralla were also noted with 1 1 and 13 primary septa (44 and

52 septa, respectively). S1-2 highly exsert, having straight to slightly sinuous inner edges that extend 4/5 distance

to columella. S3 least exsert (1.1-1. 3 mm) and least wide septa (about 2/3 width of an Si -2), with sinuous inner

edges, each bordered by a slightly sinuous, lamellar P3 1.5- 1.7 mm in width. S4 about 2.2 mm exsert, fusing to

their adjacent Si or S2 in rectangular lancets. S4 equal to or slightly wider than S3, having slightly sinuous inner

edges. S4, especially of small coralla, spaced slightly closer to their adjacent Si or S2 than to their adjacent S3,

appearing to be angled inward toward their adjacent S3. Fossa relatively shallow, containing the palar crown

(12 P3) and a linear-fascicular columella composed of 7-20 closely-spaced twisted elements arranged in 1 or

2 rows.

Remarks. — As Moseley (1881) suggested and as followed herein, even though A. spinicarens bears thecal

edge crests instead of spines, it is similar to Cary’ophyllia (A.) grayi and was therefore placed in the same subgenus

(but see alternate opinion in discussion of C. (A.) grayi). C. (A.) spinicarens also differs from C. grayi in having:

S4-S3; highly exsert calicular lancets; and ridged Ci-2. It is more similar to C. zanzibarensis Zou, i984, known

only from Tanzania at 238-302 m (Cairns & Keller, 1993), differing only in having a less compressed corallum

(GCD:LCD of C. spinicarens , 1.28-1.67; C. zanzibarensis , 1.77-2.10).

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

101

DISTRIBUTION. — Philippines : from Lubang Island to Sulu Archipelago; 222-717 m. Indonesia : Molucca Sea

(Kayoa Island); Banda Sea (Kai Islands); Timor Sea (Timor); 290-750 m. Elsewhere: South China Sea (Pratas

Island. Hong Kong, and Vanguard Bank); 223-380 m.

Caryophyllia (A.) karubarica sp. nov.

Figs 9 a-c

MATERIAL EXAMINED/TYPES. — Indonesia. Karubar: stn 10, 2 paratypes (MNHN). — Stn 12, 2 paratypes

(POL1PI). — Stn 13, 1 paratype (MNHN). — Stn 39, 12 paratypes (USNM 97036). — Stn 58, holotype (MNHN). —

Stn 71. 1 paratype (POLIPI).

Type Locality. — Karubar stn 58: 8°22'S, 132°01'E (southeast of Tanimbar Islands), 457-461 m.

Etymology. — This species is named for the Karubar expedition.

Description. — Corallum compressed (GCD:LCD = 1.26-/. 36- 1.50) and usually slightly curved in plane of

LCD, but rarely more than 45°. Angle of lateral thecal edges 50°-61°; angle of thecal faces 38°-41°. One of the

larger specimens (the holotype) is 18.8 x 25.3 mm in calicular diameter and 24.5 mm in height. Pedicel small

(1.1 -1.6 mm in diameter), circular, and always unattached as an adult; 6 protosepta usually can be seen on the basal

disc. Thecal faces meet in sharply defined, carinate edges, but edge crests only 1.0- 1.7 mm in height, usually

discontinuous, and less developed on one side. Costae 0.8- 1.0 mm wide, slightly convex in shape, and separated by

narrow, shallow striae; every costa covered with low, rounded granules — 3 or 4 across the width of a costa.

Primary costae slightly more prominent than others, but no costae are ridged. Theca of proximal 4/5 of corallum

worn and white in colour, whereas uppermost theca adjacent to calice usually light reddish brown.

Septa of most specimens (GCD = 12-25 mm) 56 in number, arranged in 3 size classes and 14 sectors:

i.e., 14:14:28, and 24 pali; but, several of the larger specimens have 1 or 2 extra pairs of tertiary septa, resulting

in 58 or 60 septa and 15 or 16 pali. Primary septa highly exsert (up to 5 mm), having sinuous inner edges that

extend to the columella. Secondary septa only 2. 1-2.2 mm exsert, 2/3 width of a primary, also having sinuous

inner edges. Tertiary septa highly exsert (3. 5-3. 9 mm), each pair fused to its adjacent primary septum in a

prominent calicular lancet that produce a highly serrate margin. Tertiary septa about 4/5 width of a secondary,

having straight to only slightly sinuous inner edges. A ring of broad (up to 2.6 mm), sinuous pali occurs before

the secondary septa. Fossa shallow, containing a fascicular columella consisting of 1 or 2 parallel rows of large,

twisted elements.

Remarks. — Caryophyllia karubarica is most similar to C. spinicarens, but differs in having: convex

(rounded, not ridged) primary costae; less prominent edge crests; 14 primary septa, instead of a tendency toward 12

as in C. spinicarens ; a "fuller" corallum (i.e., a lower GCD:LCD), especially in lower half of corallum; and S4

that are less wide than the S3. Caryophyllia karubarica differs from C. unicristata sp. nov. in having a larger, less

curved corallum; 14 primary septa, instead of exclusively 12; two carinate thecal edges, instead of a ridge only on

the convex thecal edge; and more pronounced calicular lancets. These 2 species co-occur at 3 Karubar stations.

Distribution. — Indonesia: Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar Islands); 389-

477 m.

Caryophyllia (A.) unicristata sp. nov.

Figs 9 d-e

nic:\?\ATER,AL EXAM'NED/TYPES. — Indonesia. Karubar: stn 39, 5 paratypes (POLIPI). — Stn 40. 1 paratype

USNM 97038). — Stn 58, 2 paratypes (POLIPI). — Stn 59, 81: 6 (MNHN), 75 paratypes (USNM 97040). — Stn 62,

1/ paratypes (MNHN). — Stn 69, 1 paratype (POLIPI). — Stn 70, I paratype (MNHN). — Stn 71, 3 paratypes (USNM

y/U41). _ Stn 76, holotype (MNHN) and 4 paratypes (MNHN).

Source :

102

S. D. CAIRNS & H. ZIBROWIUS

Type Locality. — Karubar stn 76: 8°49'S, 131°36'E (south of Tanimbar Islands), 400 m.

ETYMOLOGY. — The species name unicristata (Latin unus, one + crista , crest) refers to the single edge crest of

this species.

DESCRIPTION. — Corallum relatively small, the largest specimen (KARUBAR stn 39) only 15.5 mm in GCD;

the holotype is 10.9 x 13.4 mm in calicular diameter and 16.0 mm in height. Corallum ceratoid, regularly curved

45°-90° usually in plane of GCD. Pedicel small (0.8-0.9 mm in diameter), unattached, and not reinforced. Calice

elliptical: GCD:LCD = 1.21-1.32. Base of pedicel often free, detached from substratum, its flat lower side

revealing the 6 protosepta; sometimes worn to a rounded tip; or occasionally overgrown by a bryozoan colony.

A low (0.9- 1.1 mm), sinuous crest occurs on convex edge of each corallum, the concave edge being evenly

rounded. Convex edge crest best developed in small coralla, often worn or lost in larger coralla. The other 1 1 Ci-2

are raised and slightly rounded (not ridged); C3-4 flat. All costae covered with low, rounded granules, 3 or 4 across

width of a costa. Upper 1-2 mm of theca and adjacent septa reddish-brown; remainder of corallum white.

Septa hexamerally arranged in 4 complete cycles: Si-2>S3>S4, and 12 pali. S 1-2 about 1.5 mm exsert, having

vertical, slightly sinuous inner edges that almost attain the columella. S3 about 0.4 mm exsert and 2/3 width of

the Si -2, having highly sinuous inner edges. S4 intermediate in exsertness, each pair of S4 fusing to its common

Si or S2 to form a low calicular lancet. S4 equal to or slightly less wide than S3, the widest S4 occurring in coralla

of greatest size. Inner edges of S4 slightly sinuous. 12 prominent P3 1.2- 1.4 mm wide, with sinuous edges.

Columella consists of 4-12 well-formed, twisted elements, 4-6 arranged in a line or a larger number in 2 parallel

rows.

Remarks. — Within the subgenus Acanthocyathus, A. unicristata needs to be compared only to the 3 species

that have ridged (nonspinose) thecal edges: C. spinicarens , C. zanzibarensis , and C. karubarica , from which it

differs in having: only one carinate thecal edge (on its convex side); a more circular calice (a relatively low

GCD:LCD of 1.21-1.32); a smaller pedicel diameter; consistently hexameral symmetry; and low calicular lancets.

Distribution. — Indonesia : Arafura Sea (southeast of Tanimbar Islands); 251-477 m.

Genus PREMOCYATHUS Yabe & Eguchi, 1942

Premocyathus dentiformis (Alcock, 1902) comb. nov.

Figs 9 f-j

Placotrochides dentiformis Alcock, 1902b: 121; 1902c: 33-34, pi. 4, figs 31, 31a.

Caryophyllia compressa Yabe & Eguchi, 1932a: 443 (nom. nud.).

Premocyathus compressus Yabe & Eguchi, 1942b: 121, 151-152, pi. 10, figs 13-14 (junior homonym).

Caryophyllia (P.) compressa - Mori, 1987: 21-30, 9 figs. — Cairns, 1994: 50-51, pi. 22, figs e-f (synonymy); 1995:

54-55, pi. 11, figs f-i.

? Caryophyllia (P.) ceratoconus Hu, 1987: 38-39, pi. 1, figs 16, 20.

Not Caryophyllia compressa Gardiner* Waugh. 1938: 180 [junior synonym = Caryophyllia (A.) zanzibarensis Zou,

1984, nom. nov.].

Not Caryophyllia (P.) compressa - Wells, 1956: F422, fig. 323, 3 [= Trochocyathus (T.) apertus sp. nov. herein).

Material EXAMINED. — Philippines. "Albatross": stn 5155, 1 (USNM 97042). — Stn 5217, 28 (USNM

62708).

Indonesia. "Siboga": stn 59, holotype (ZMA Coel. 1093).

"Galathea": stn 490, 4 (ZMUC).

Karubar: stn 15, 1 (MNHN). — Stn 18, 1 (USNM 97044).

Type Locality. — "Siboga" stn 59: 10°22.7'S, 123°16.5'E (off Timor), 390 m.

Diagnosis. — Corallum compressed (GCD:LCD = 1.2- 1.4) and curved 30°-45° in plane of GCD, the concave

thecal edge rounded, the convex thecal edge usually crested or even keeled. Base of corallum typically an open.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTIN1A

103

irregular scar 1.0- 1.5 mm in diameter. Only one specimen (Karubar stn 15) was found bearing an opposite bud at

this level, instead of having an open base. In fact, the usually open base is probably the result of breaking apart of

parent or opposite bud. Septal symmetry quite variable. Of the 57 septal/palar permutations inferred by MORI

(1987), the 4 well-preserved Philippine specimens displayed the following symmetries: 8/8/16, 8 (1 specimen),

10/10/20, 10 (1 specimen), and 10/10/18, 9 (2 specimens), which were the 13th, 4th, and 5th most common

arrangements, respectively, among specimens MORI examined from the type locality. Primary and secondary septa

and all pali have sinuous inner edges. Pali occur before secondary septa. Fascicular columella composed of

1-4 twisted elements.

REMARKS. — The holotype of Placotrochides dentiformis (ZMA Coel. 1093) is a poorly preserved specimen

4.2 x 6.9 mm in calicular diameter and 7.4 mm in height, having a septal complement of 8:8:10 (26 septa). This

septal arrangement is shared with only 7 of the 1090 specimens analyzed by MORI (1987). Due to poor preserva¬

tion, no pali are present, but a fascicular columella composed of twisted elements is recognizable. Although a poor

specimen, there is little doubt that it is conspeciftc with the species later reported as Premocyathus compressus.

Coralla of Trochocyathus apertus sp. nov. are similar in shape and size to P. dentiformis but differ in having a

papillose columella, P|-3 (not just P3), and in having a less compressed corallum.

Cairns & Keller (1993) and Cairns (1995) previously considered there to be 5 species in the subgenus

Caryophyllia (Premocyathus), but we now place 2 of them, Acantkocyathus spinicarens (Moseley, 1881) and

C. zanzibarensis Zou, 1984, in the subgenus Caryophyllia (Acanthocyathus)\ and the other 2, C. (Premocyathus)

burchae Cairns, 1984, and C. (P.) compressa sensu Wells (1956) (= Trochocyathus apertus sp. nov.) are herein

transferred to Trochocyathus. Only 1 species remains in Premocyathus, P. dentiformis. The genus Premocyathus is

thus defined as having: a curved, laterally compressed corallum with a carinate convex edge and an open base that

most likely is the result of separation from an opposite budded specimen; a variable septal symmetry; pali before

the penultimate septal cycle; and a fascicular columella.

Distribution. — Philippines : Ragay Gulf; Sulu Sea (Sulu Archipelago); 22-192 m. Indonesia: Banda Sea

(Kai Islands); Savu Sea (Timor); Java Sea; 221-545 m. Elsewhere: Japan (from Honshu to northern Ryukyu

Islands); Kermadec Islands; 1 15-757 m. Pleistocene of Japan. ? Plio-Pleistocene of Taiwan (as C. (P.) ceratoconus

Hu, 1987).

Genus CRISPATOTROCHUS Tenison Woods, 1878

Crispatotrochus rubescens (Moseley, 1881)

Figs 10 a-c

Cyathoceras rubescens Moseley, 1881: 157, pi. 2, figs 8a-c. — Cairns, 1984: 15.

Cyathoceras tydemani Alcock, 1902a: 93-94; 1902c: 14, pi. 1, figs 7, 7a (new synonym). — Faustino, 1927: 65,

pi. 9, figs 5-6.

Cyathoceras diomedeae Vaughan, 1907: 77-78, pi. 7, figs 1-2.

Crispatotrochus rubescens - Cairns, 1991: 15; 1994: 51, pi. 22, figs g-h (synonymy).

Material EXAMINED. — Philippines. "Albatross": stn 5519, 1 (USNM 60586).

Musorstom 2: stn 15, 4: 2 (MNHN), 2 (USNM 97049).

Indonesia. "Siboga": stn 105, syntype of C. tydemani (ZMA Coel. 579, Fig. 10c).

Deki: stn 3, 1 (NNM 22418). — Stn 48, 1 (NNM 22415). — Stn 59, 4 (NNM 22416).

Snellius 2: stn 4.066, 2 (NNM 22417).

Karubar: stn 16, 1 (USNM 97046). — Stn 67, 1 (USNM 97047). — Stn 86, 1 (MNHN).

South China Sea. "Hakuho Maru": stn KH73-2-44-2, 4: 2 (USNM 97050), 2 (ORI).

Type Locality. — "Challenger" stn 192: 5°49'15"S, 132°14’15"E (Kai Islands, Banda Sea), 236 m.

Diagnosis. — Corallum elongate-conical to trochoid, with a straight, flared calice and a robust pedicel about

5% diameter of GCD. Largest known specimen (" Albatross " stn 5519) 34 x 28 mm in calicular diameter, 39 mm

Source : MNHN , Paris

104

S. D. CAIRNS & H. ZIBROWIUS

in height, and 9.3 mm in pedicel diameter. Costae ridged near calice, but otherwise flat and granular. Septa

hexamerally arranged in 5 cycles (Si-2>S3>S4>Ss), the 4th cycle (48 septa stage) attained at a GCD of 7-9 mm.

Inner edges of Si -2 moderately sinuous. Fossa deep; pali absent; fascicular columella composed of numerous (15-

20), slender, twisted elements.

Remarks. — Crispatotrochus rubescens is more fully described and illustrated by Cairns (1994). Cyathoceras

tydemani is a juvenile specimen of C. rubescens , collected at a stage transitional between the 4th and 5th septal

cycles, i.e., 60 septa at a GCD of 8 mm.

DISTRIBUTION. — Philippines : Lubang Island; Bohol Sea (south of Negros); Sulu Sea (Sulu Archipelago);

275-522 m. Indonesia : Banda Sea (Kai Islands); Arafura Sea (south of Tanimbar Islands); Savu Sea (Sumba); 226-

315 m. Elsewhere: South China Sea (southern Formosa Strait); Japan (Honshu, Shikoku, and Kyushu); Hawaiian

and Christmas Islands; 1 10-634 m.

Crispatotrochus rugosus Cairns, 1995

Crispatotrochus rugosus Cairns, 1995: 57, pi. 13, figs a-b.

Material EXAMINED. — Indonesia. "Albatross": stn 5586, 1 (USNM 97052).

Type Locality. — NZOI stn Q70: 26°59.7'S, 159°18.9'E (Lord Howe Seamount Chain), 376 m.

Remarks. — Crispatotrochus rugosus was recently described based on specimens from the New Zealand

region; only a single, worn specimen of 8.7 mm GCD is reported herein. It is distinguished from C. rubescens by

having fine, transverse thecal ridges, and Si that are wider than S2. Fragments of a much larger specimen

(MUSORSTOM 2 stn 32, MNHN) — having an estimated GCD of 32 mm, 5 cycles of septa, and transverse thecal

costal ridges — may represent a large specimen of this species. Unfortunately, that specimen is missing its pedicel

and columellar region.

Distribution. — ? Philippines: Verde Island Passage; 192-220 m. Malaysia: Celebes Sea (Sabah); 616 m.

Elsewhere: Kermadec Islands; Lord Howe Seamount Chain; 142-508 m.

Genus LABYRINTHOCYATHUS Cairns, 1979

Labyrinthocyathus sp. A

Figs 10 f, i

Material EXAMINED. — Indonesia. Karubar: stn 36, 1 (MNHN).

Description. — Unique specimen 8.0 mm in calicular diameter, 15.5 mm in height, and 1.65 mm in pedicel

diameter. Theca thick (about 0.9 mm) and heavily encrusted with serpulid tubes, otherwise C1-2 slightly ridged.

Corallum white. Septa hexamerally arranged in 4 cycles: Si>S2>S3»S4. Si 1.7 mm exsert, having moderately

sinuous inner edges. S2 only slightly narrower than Si; S3 about 1/2 width of S2. S4 rudimentary: expressed only

as a short costoseptal ridge at calicular margin, absent from upper fossa, and present in lower fossa only as a

narrow lamella. Fossa of moderate depth; columella labyrinthiform.

Remarks. This specimen differs from the four Recent species of Labyrinthocyathus that have 4 cycles of

septa [L. dehcatus (Marenzeller, 1904); L. limatulus (Squires, 1964); L. langae Cairns, 1979; and L. facetus

Cairns, 1979] by having rudimentary S4. It may represent an undescribed species.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

1

Distribution. — Indonesia: Banda Sea (between Tanimbar and Aru Islands); 210-268 m.

Genus TROCHOCYATHUS H. Milne Edwards & Haime, 1848

Key to the 12 species of Trochocyathus known from the Philippine/Indonesian region

1. Corallum with 6 basal spines (Ci), 1 corresponding to each Ci . 2

— Corallum nonspinose, or, if spinose, the spines are limited to edge spines corresponding to

only 2 Ci . 3

2. Basal spines short (less than 5 mm); some S5 usually present . T. (A.) brevispina

— Basal spines longer (up to 10 mm); only 48 septa . T. (A.) longispina

3. Corallum reproduces predominantly by transverse division, resulting in a characteristic

basal scar on anthocyathus . 4

_ Corallum does not reproduce by transverse division, the base either being firmly attached

or having an open (broken) base . 6

4. Corallum discoidal or bowl-shaped; thecal spines not present . 5

— Corallum elongate-conical; flattened edge spines present . T. (T.) cooperi

5. Corallum discoidal: H:D about 0.5 . T. (T.) discus

— Corallum bowl-shaped: H:D 0.7-0.8 . T. (T.) gardineri

6. Base of corallum attached to a substratum; corallum straight . 7

— Base of corallum open, as though broken; corallum curved . 1 2

7. Corallum nonspinose . ®

— Corallum bears 1 or more pairs of thecal edge spines . 11

8. Theca transversely ridged . T. (T.) rhombocolumna

— Theca longitudinally costate . 9

9. Corallum (theca and septa) speckled with black pigment; pedicel robust (PD:GCD = 0.60-

0.80), composed of chambered concentric rings . T. (T.) maculatus

— Corallum not speckled; pedicel small to medium-sized (PD:GCD = 0.10-0.45), not

chambered . 1®

10. Pi -2 much smaller than P3; corallum usually contains 48 septa; columella a field of 6-20

medium-sized elements . T. (T.) philippinensis (nonspinose form)

— Pi -3 equal in width; corallum usually contains more than 48 septa; columella a field of up

to 40 slender elements . T. (T.) caryophylloides

11. Septa decamerally arranged in 3 size classes (40 septa); pedicel small (1.7- 1.9 mm

diameter); >2 pairs of delicate thecal edge spines usually present . T. (T.) semperi

— Septa hexamerally arranged (Si-2>S3>S4, 48 septa); pedicel larger (2. 9-6. 7 mm diameter);

usually only 1 pair of robust edge spines present .

. T. (T.) philippinensis (spinose form)

12. Crest on convex thecal edge continuous from base to calice; calice often brown-black near

calice . T. (T.) burchae

— Crest on convex thecal edge restricted to lower corallum; corallum white .

. T. (T.) apertus

106

S. D. CAIRNS & H. ZIBROWIUS

Subgenus TROCHOCYATHUS (TROCHOCYATHUS) H. Milne Edwards & Haime, 1848

Trochocyathus (T.) caryophylloides Alcock, 1902

Trochocyathus caryophylloides Alcock, 1902a: 94; 1902c; 14-15, pi. 2, figs 10, 10a. — Faustino. 1927; 80, pi. 7,

figs 5-6. — Yabe & Eguchi. 1942b: 123-124, pi. 10, fig. 21. — ? Zou et al„ 1988: 195. — Cairns, 1994: 52-53’

pi. 23, figs a-c, h.

Not Trochocyathus caryophylloides Zou, 1988: 76, pi. 5, fig. 5, 5a [= Trophocyathus (T.) Iphilippinensis sp. nov.

herein]..

Material EXAMINED. — Philippines. Musorstom 1: stn 63, 1 (USNM 97057).

Musorstom 2: stn 2, 1 (MNHN). — Stn 32, 1 (MNHN).

Musorstom 3: stn 88, 1 (MNHN).

Indonesia. Deki: stn 7, 2 (NNM 22763).

Karubar: stn 32, 1 (MNHN). — Stn 49. 1 (MNHN). — Stn 50, 2 (MNHN). — Stn 61, 1 (MNHN) — Stn 86

2 (USNM 97056).

T\ pe Locality. Siboga stns 96, 251, and 253: Celebes and Banda Seas, Indonesia, 1 15-304 m.

Diagnosis. — Corallum trochoid, firmly attached through a robust pedicel (PD:GCD = 0.1 1-0.41), up to

21 mm in GCD. Costae broad, flat to slightly convex, and covered with low, rounded granules. Corallum white to

light brown. Septal symmetry appears to change with size. Coralla 8-12 mm in GCD have 4 cycles of

48 hexamerally arranged septa (Si-2>S3>S4). Larger coralla over 16 mm GCD often have 16 primary septa and

3 size classes of septa, resulting in 64 septa, or hexameral symmetry but with an incomplete 4th cycle, also

resulting in 64 septa (MUSORSTOM 1 stn 63). Discrete pali arranged in 2 or 3 crowns before all but last cycle of

septa. Columella composed of an elliptical field on numerous (up to 45), slender rod-shaped papillae.

Remarks. — One aberrant specimen (Musorstom 2 stn 32) has undergone intratentacular budding, resulting

in a small colony of 3 massive corallites, each the result of equal, intratentacular division. T. caryophylloides is

more fully described and figured by Cairns (1994).

In our opinion, the species name was originally incorrectly formed, being derived from the root Caryophyllia ,

and thus should be caryophyllioides. However, according to the ICZN (Article 32dii) the original spelling cannot

be changed.

Distribution. — Philippines : Lubang Island; Verde Island Passage; 186-192 m. Indonesia : Banda Sea (Kai

Islands); Arafura Sea (southeast of Tanimbar Islands); 185-304 m. Elsewhere : Japan (Honshu and Fukue JimaV

1 15-344 m.

Trochocyathus (T.) rhombocolumna Alcock, 1902

Trochocyathus rhombocolumna Alcock, 1902a; 98; 1902c; 16, pi. 2, fig. 12. - Cairns, 1995; 60-61, pi 13 fig i

pi. 14, figs a-b (synonymy). y ’ 5 ’

Material EXAMINED. — Indonesia. Deki: stn 8, 1 (NNM 22755). — Stn 59, 1 (NNM 22756)

Karubar: stn 27, 1 (USNM 97061). — Stn 49, 2 (POLIPI). — Stn 86, 1 (MNHN).

Type Locality. — "Siboga" stn 95: 5°43.5'N, 1 19°40’E (Sulu Sea), 522 m.

fPnDrrGn-T^«TallU,m elon§ate-conical t0 trochoid- slight, firmly attached through a robust pedicel

(PD.GCD - 0.34-0.56), and up to 14 mm in GCD (Cairns, 1995). Theca covered with thin transverse ridges

Corallum white. Septa hexamerally arranged in 4 full cycles: S,>S2>S4>S3, the S, being highly exsert. Pali

arranged ,n 3 crowns before all but last septal cycle, each P2 and pair of P3 within a system forming a distinctive

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

107

triangular pattern. Columellar elements few in number and irregular in shape, sometimes rhomboidal in cross

section.

Remarks. _ Trochocyathus rhombocolumna is more fully described and illustrated by Cairns (1995) based

on New Zealand specimens. One specimen from KARUBAR stn 27 has a characteristic lenticular-shaped aperture of

an acrothoracican cirripede in the side of its corallum, but the symbiont no longer present.

DISTRIBUTION — Philippines: Sulu Sea (Sulu Archipelago); 522 m. Indonesia: Banda Sea (Kai Islands);

Arafura Sea (south and east of Tanimbar Islands); 209-385 m. Elsewhere: widespread in Indo-West Pac.f.c from

southwestern Indian Ocean to Hawaiian Islands, including ridges north of New Zealand; 1 10-530 m.

Trochocyathus (T.) maculatus Cairns, 1995

Trochocyathus (I) maculatus Cairns, 1995: 61, pi. 14, figs c-d.

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn 134, 2 (MNHN).

TYPE LOCALITY. — NZOI stn PI 15: 31°25.9’S, 159°02.2'E (off Lord Howe Island), 183 m.

DIAGNOSIS. - Corallum elongate-conical, straight, Oared distally, and attached by a broad pedicel 0.60-0.80

CCD. Pedicel increases in diameter by formation of thin exothecal dissepiments over raised basal costae, similar to

the process in Rhizosmilia. Largest Philippine specimen 8.6 x 10.3 mm in calicular diameter and 9.7 mm in

height; largest known specimen (CAIRNS, 1995) 13.8 mm in GCD. Costae granular, well-defined only in upper

part of corallum. Corallum white, but theca and all septa (not pali) speckled with dark brown-black pigmentation.

Septa hexamerally arranged in 5 cycles, the 5th never complete. The large Philippine specimen has only 1 pair of

S5 (50 septa) and a New Zealand specimen has as many as 64 septa. Si highly exsert, forming calicular lancets.

S2 smaller than Si but larger than S3. S4 variable in width, those adjacent to Si often wider than the S3, those

adjacent to S2 usually less wide than S3. Two crowns of pali present: 12 small Pi-2 occurring low in the fossa,

and 12 larger P3 rising much higher in fossa. P4 present on septa flanked by a pair of S5. Papillose columella a

field of 20-30 fine pillars.

Remarks. — Trochocyathus maculatus is more fully described and illustrated in the original account. It is

distinguished from its congeners by its distinctively speckled corallum and exothecal dissepiments.

Distribution. — Philippines: Sibuyan Sea (northeastern tip of Panay); 92-95 m. Elsewhere: Kermadec and

Lord Howe Islands; Taupo Seamount and Dampier Ridge, southeastern Australia; 100-183 m.

Trochocyathus (T.) philippinensis Semper, 1872

Figs lOd-e

Trochocyathus philippinensis Semper, 1872: 253, pi. 20, fig. 16. — FaUSTINO, 1927: 79-80, pi. 7. figs 3-4.

? Trochocyathus catyophylloides - Zou, 1988: 76, pi. 5, figs 5, 5a. [Not T. caryophylloides Alcock, 1902aJ.

Material EXAMINED. — Philippines. "Albatross": stn 5178, 8 (USNM 97065). — Stn 5213, 2 (USNM 97066).

— Stn 5217, 1 (USNM 97067). — Stn 5381, 3 (USNM 97070).

Musorstom 1: stn 13, 1 (USNM 97075). — Stn 14, 1 (MNHN). — Stn 35, 1 (USNM 97076). — Stn 64, 1 (MNHN).

Musorstom 2: stn 6, 1 (MNHN). — Stn 10, 1 (USNM 97079). — Stn 17, 1 (MNHN). — Stn 33, 32 (USNM 9708°)-

Musorstom 3: stn 88, 2 (USNM 97081). — Stn 96, 6 (USNM 97083). — Stn 98, 1 (USNM 97083). — Stn 102,

3 (MNHN). — Stn 108, 1 (MNHN). — Stn 124, 3. — Stn 131, 1 (USNM 97084). ^ „

Indonesia. Deki: stn 6, 2 (NNM 22772). — Stn 24, 2 (NNM ). — Stn 44, 8 (NNM 22773). — Stn 49, 7 (NNM

22774).

Source : MNHN , Paris

108

S. D. CAIRNS & H. ZIBROWIUS

Corindon 2: stn 248, 2 (MNHN).

Karubar: stn 2, 24 (POLIPI). — Stn 3, 8: 1 (MNHN), 7 (USNM 97071). — Stn 15, 1 (USNM 97072). — Stn 18

1 (USNM 97073). — Stn 32, 1 (POLIPI).

South China Sea. "Albatross": stn 5311, 2 (USNM 97068). — Stn 5314, 1 (USNM 97069).

Ryukyu Islands. "Tansei Marti stn KT93-09-AM6, 1 (USNM 93163). — Stn KT93-09-AM7, 1 (USNM 93160).

Type Locality. — Pandanon, west coast of Bohol, Philippines, 27-54 m.

DESCRIPTION. — Corallum ceratoid to trochoid and relatively small, the largest corallum (MUSORSTOM 3

stn 88) 12.3 x 17.6 mm in calicular diameter and 26.4 mm in height, but most specimens less than half this size;

one syntype measures 9.5 x 11.0 mm in calicular diameter and 19 mm in height (fide SEMPER, 1872). Calice

elliptical : GCD:LCD = 1.15-1.64. Coralla often maintain their attachment to substratum through a robust pedicel

2. 9-6.7 mm in diameter (PD:GCD = 0.20-0.44), the substratum often consisting of a small gastropod shell, an

echinoid spine, a pebble, or a corallum of a dead Scleractinia such as Flabellum or Balanophyllia. Corallum

usually straight, occasionally bent near base. About 1/3 of specimens examined bear slender thecal edge spines, the

remaining coralla having evenly rounded thecal edges. Of those having edge spines, size and symmetry are quite

variable. Some specimens have a pair of spines, but coralla bearing a single spine or just a nub or low crest on

one thecal edge are not uncommon. Only one specimen (MUSORSTOM 3 stn 96) has 3 spines: 2 on one edge and

1 on the other. Lower 1/3 to 1/2 of theca white and porcellaneous; however, upper 1/2 to 2/3 dark brown, bearing

well-formed, convex costae separated by deep intercostal furrows. Costae in upper corallum with minute granules.

Septa hexamerally arranged in 4 full cycles: Si-2>S3>S4I however, some large specimens have 1 or 2 pairs of

Ss resulting in 50-52 septa, and small coralla less than 9 mm GCD often lack several pairs of S4. S1-2 about

1.7 mm exsert, having vertical, slightly sinuous inner edges, each bordered by a small (0.3-0.5 mm) papillose to

lamellar palus. S3 about 1.3 mm exsert, with slightly more sinuous inner edges, and bordered by lamellar P3.

P3 each about 1.0 mm wide and forming a crown that rises higher in the fossa than the crown of P1-2. S4 equally

exsert as S3, in small to medium-sized coralla S4 slightly less wide than the S3, but S4 become proportionately

wider with growth of the corallum, until they sometimes slightly exceed the S3 in width. Fossa of moderate depth,

containing a papillose columella consisting of 6-20 interconnected papillae.

Remarks. — Among the 23 Recent species in the nominate subgenus of Trochocyathus, only 2 other have

thecal edge spines: T. semperi sp. nov. and T. cooperi (Gardiner, 1905). T. philippinensis is distinguished from

T. semperi by having: a larger corallum with a more robust pedicel (2.9-6.7 mm in diameter vs 1.7- 1.9 mm for

T. semperi ); hexameral septal symmetry resulting in 48 septa (not 40 septa); asymmetrical and erratic

development of thecal edge spines (vs usually two pairs for T. semperi)- and highest cycle septa (S4) that are

sometimes as large as the penultimate cycle (S3), whereas the tertiary septa of T. semperi are 1/2 the width of

their secondaries. Furthermore, T. philippinensis is more common in deeper water than T. semperi, i. e. , 1 GO-

268 m whereas most records of T. semperi are from less than 1 00 m.

Although similar in calicular features, T. cooperi differs from T. philippinensis by dividing transversely (basal

scar), and in having plate-like, lamellar thecal edge spines.

Distribution. — Philippines: Lubang Island; Verde Island Passage; Sibuyan and Samar Seas; Burias Pass;

Pandanon; Sulu Sea (Semirara Islands and west of Panay); 54-194 m. Indonesia: Makassar Strait; Banda Sea (Kai

Islands); 100-268 m. Elsewhere: South China Sea (north of Pratas Island); Japan (northern Ryukyu Islands);

108-223 m.

Trochocyathus (T.) semperi sp. nov.

Figs 10 g-h, 11 f

Philippines. "Albatross": stn 5133, 1 paratype (USNM 97085). — Stn 5142,

Material examined/Types.

5 paratypes (USNM).

MUSORSTOM 3: stn 140, 9 paratypes (MNHN),

Indonesia. Deki. stn 6, 3 paratypes (NNM 22784). — Stn 53, 2 paratypes (ZMUC), 2 paratypes (NNM 22785).

Corindon 2: stn 251, 59: holotype (MNHN), 10 paratypes (POLIPI), 48 paratypes (USNM 97086).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

109

TYPE LOCALITY. - Corindon 2 stn 251: 0°53.7'S, 1 19°29.6'E (Makassar Strait), 65 m.

ETYMOLOGY. - This species named for Carl Gottfried SEMPER, in recognition of his work on shallow water

azooxanthellate corals of the Philippine region (see Semper, 1872).

DESCRIPTION - Corallum ceratoid and relatively small, most specimens examined only 5-7 mm in GCD and

equally tall, the largest rMba,ross" stn 5142) being 7.7 x 9.7 mtr tin calicular dtameter anti 15 mn. .» i hetght.

Holotype 4 9 x 6.6 mm in calicular diameter and 8.4 mm in height. Calice elliptical. GCD. LCD - 1.25-1.37

Colalla often maintain their attachment to substratum through a robust pedicel 1.7- 1.9 mm >" diameter

(PD GCD = 0 18-0 30), the substratum often being a bivalve shell, large foramimferan, or other corals, sue

Heteropsammia. Pedicel next to basal disc often bent 30°-45°. Occasionally corallum detached from the substratum

revealing the basal disc and original 6-12 protosepta. Most coralla bear at least 2 pairs of delicate thecal

edge spines, the 1st pair beginning its development almost immediately above the basal disc as an ^tension of

thf 2 principal CSi; in 2 cases the lowermost spines were fused to the substratum Edge spines spatulate

proximalW cylindrical distally, and up to 3.6 mm in length, but. because of their delicate nature, only small

coralla be^ intact, elongate spines. All costae equally convex and granular. Upper 1/2 of theca blackish-brown,

lower 1/2 of theca and calicular elements white. „ , , ,

In early development, septa are hexamerally arranged in 3 cycles, but by a GCD of about ^ mm a decamer^

symmetry is adopted, adult coralla usually having 3 size classes of decamerally arranged septa (40 septa . Pnmary

septa 0.8 1.0 mm exsert, having vertical, slightly sinuous inner edges, each bordered by a small, Pooriy-formed

nalus Secondary septa 0.6 mm exsert and about 2/3 width of a primary, having sinuous inner edges each bordered

^ a naZ (about 0.5 mm), lamellar palus. Tertiary septa equally exsert, but only 1/2 width of a secondary.

Fossa o7moderate depth, containing a papillose columella composed of 5-9 slender elements, sometimes

indistinguishable in size and shape from the Pi.

Remarks - Only 3 of the approximately 23 Recent species of Trochocyathus (Trochocyathus) have thecal

edge spines: T. cooped (Gardiner, 1905); T. semperi ; and T. philippinensis , the last 2 species being compared

the previous account. Within the Scleractinia, edge spines are often associated with coralla i of th®[

transversely divide as a method of asexual propagation (e.g., Trochocyathus most spec es of

Truncatoflabellum)', however, in both T. philippinensis and T. semperi , no specimens show evidence of

division, many displaying their original basal disc.

DISTRIBUTION. - Philippines : Sibuyan Sea; Sulu Sea (Zamboanga Peninsula and Sulu Archipelago);

38-93 m. Indonesia: Makassar Strait; Banda Sea (Kai Islands); 65-245 m.

Trochocyathus (T.) apertus sp. nov.

Figs 1 1 a-d

Caryophyllia (. Premocyathus ) compressa - Wells, 1956: F422, fig. 323,3. [Not Premocyathus compressus Yabe &

Prvnocyaih wTcompressus - Ca.RNS, 1984: 14 (in part: " Albatross " specimens). [Not Premocyathus compressus Yabe &

Not ^Ca^ophyma (Premocyathus) compressa - Cairns, 1995: 54-55. [See discussion of Premocyathus dentiformis).

Material EXAMINED/TYPES. — Philippines. "Albatross": st"515(3, 1 19 paratypes (USNM 627(D). -

Stn 5164, 526+: holotype (USNM 97087), 500+ paratypes (USNM 62710), 26 (MNHN).

Indonesia. Deki: stn 10, 2 paratypes (NNM 22498).

MORTENSEN'S Java-S.A. Expedition: stn 9, 1 paratype (ZMUC).

Type Locality. — "Albatross" stn 5156: 5°01'40"N, 1 19°52'20"E (Sulu Archipelago), 33 m.

Etymology. — The species name (Latin apertus, open) alludes to the open base of all specimens.

Description. — Corallum relatively small, usually curved about 90° in plane of GCD, and somewhat

compressed (GCD:LCD = 1.19-1.37). Largest known specimen (ZMUC) 7.2 x 9.9 mm in calicular diameter anc

110

S. D. CAIRNS & H. ZIBROWIUS

14.3 mm in height; holotype 7.9 x 6.3 mm in calicular diameter and 11.8 mm in height. All coralla examined

unattached (free), with an open base 1.2-1. 8 mm in diameter that reveals the 12 septa: 6 thick and 6 thin. Each

corallum bears a short thecal edge crest on its lower convex edge, up to 5 mm long, up to 3 mm in height, and

about 0.5 mm thick. Crests best developed on small coralla, becoming worn or broken with age, not extending to

calicular edge. Crests absent from concave thecal edge, but occasionally the principal Ci of the concave edge is

slightly prominent or thickened. Costae low, convex, granular ridges separated by broad, shallow intercostal

furrows. Corallum white.

Septa hexamerally arranged in 4 cycles, but 4th cycle never complete. The most common septal complement is

40, arranged: 12:12:16, with 4 half-systems lacking pairs of S4 (e.g., the holotype). If the 12 half-systems are

numbered in a clockwise direction, starting with the half-system to the right of the principal septum aligned with

the convex edge, the 4 half-systems that lack S4 pairs are usually the 3rd, 5th, 8th, and 10th. Coralla having

44 septa, the largest number of septa observed, usually lack S4 pairs in the 5th and 8th half-systems. Si -2 up to

1.3 mm exsert, having vertical, slightly sinuous inner edges. S3 about 1.0 mm exsert, with slightly sinuous inner

edges, about 3/4 the width of an S2- S4 slightly less exsert and 3/4 width of an S3. Most, but not all, S1-2 bear a

small (0.3-0. 5 mm wide) papillose to lamellar palus, the papillae being indistinguishable from columellar

elements, except that they rise higher in the fossa. P3 always lamellar, about 1 mm wide, with straight edges, and

rising even higher in the fossa. The fossa contains a papillose columella consisting of 7-10 interconnected pillars,

each about 0.3 mm in diameter.

Remarks. — Cairns (1984) previously considered this species to be Premocyathus compressus

(= Premocyathus dentiformis herein) because of their similarity in corallum size and shape; because both species

have open bases; and because many specimens of T. apertus lacked Pi -2 and thus resembled a Caryophyllia ground

plan. In spite of these convergent characters, T. apertus differs significantly in having a papillose columella

composed of slender rods (not a fascicular composed of twisted lamellae), and in having often distinct Pi -2.

Furthermore, whereas the septal symmetry of P. dentiformis [= C. (P.) compressa sensu MORI, 1987] is quite

variable, the hexameral pattern that is most common in T. apertus (12:12:16) is quite rare (0.3%) in

P. dentiformis. Overall hexameral symmetry is the rule in T. apertus , but rare (only 3%) in P. dentiformis.

DISTRIBUTION. — Philippines : Sulu Sea (Sulu Archipelago); 33 m. Indonesia: Banda Sea (Kai Islands); Bali

Strait; 50-70 m.

Trochocyathus (T.) burchae (Cairns, 1984) comb. nov.

Premocyathus burchae Cairns, 1984: 14, pi. 2, figs G-H.

Caryophyllia (P.) burchae - Cairns, 1995: 54 (listed).

MATERIAL EXAMINED. — Philippines. " Albatross ": stn 5133, 6 (USNM 97064).

Indonesia. Deki: stn 10, 18 (NNM 22764). — Stn 82, 1 (NNM 23202).

MORTENSEN'S Java-S.A. Expedition: stn 5, 1 (ZMUC). — Stn 9, 4: 3 (ZMUC), 1 (USNM 97089).

Type Locality. — 20°43.7'N, 156°54.6'W (Lanai, Hawaiian Islands), 64 m.

Diagnosis/Remarks. — Little can be added to the original description based on Hawaiian specimens.

Although quite similar, T. burchae differs from T. apertus in having a smaller corallum with a smaller open base;

a brown-black pigmented upper theca; and a more distinctly developed thecal edge crest. The crest of the convex

side of T. burchae is up to 4.8 mm in height and usually continuous from base to calice, occasionally bilobate.

Furthermore, some coralla have a low crest on the lower concave thecal edge. The largest known specimen

(MORTENSEN stn 9) is 6.4 x 10.0 mm in calicular diameter.

Trochocyathus burchae is placed in this genus for the same reasons cited for T. apertus: its papillose columella

and the occasional presence of Pi-2 as well as P3.

DISTRIBUTION. — Philippines: Sulu Sea (Zamboanga Peninsula); 70 m. Indonesia: Banda Sea (Kai Islands);

Bali Strait; Sunda Strait (Java Sea); 35-70 m. Elsewhere: Lanai, Hawaiian Islands; 64 m.

Source :

AZOOXANTHELLATE SCLERACTINIA

111

Trochocyathus (T.) cooperi (Gardiner, 1905)

Fig. 1 1 e

? Trochocyathus weberi Alcock, 1902a: 95-96.

Trnnidocvathus cooperi Gardiner, 1905: 955, pi. 93, fig. 30. w

Trochocyathus sp. - Vaughan & WELLS, 1943: 47, fig. 20b (inverted view of specimen from Albatross stn 5142).

Trochocyathus cooperi - CAIRNS, 1994: 54, pi. 23, figs f-g.

MATERIAL EXAMINED - Philippines. " Albatross ": stn 5133, 1 (USNM 97090). - Stn 5136, 5 (USNM 97091).

S137 2 (USNM 97092) - Stn 5142, 9 (USNM 97093). - Stn 5143. 3 (USNM 97094). - Stn 5144, 1 (USNM

CT9S) - sm 5H6 5 (USNM 97«. - Sm 5147. 3 (USNM 97097). - St. 5151, 9 (USNM 970981. - Stn 5202.

1 (USNM 97099). — Stn 5355, 1 (USNM 97100).

MUSORSTOM 1: stn 57, 1 (MNHN).

Musorstom 2: stn 47, 1 (MNHN).

if,'”-’ Stn .7, 1 (NNM 22564). - Stn 24 4 (NNM 22565). - Stn 53, 4 (NNM

22566). — Stn 90, 1 (NNM 22569). — Stn 95, 1 (NNM). — Stn 103, 72 (NNM 22570).

SNELLIUS 2: stn 4.234, 1 (NNM 22571).

South Pacific. "Pele": stn TH1, 16 (USNM 73764).

Type locality. — Kolumadulu and Suvadiva, Maidive Islands, 64-70 m.

DIAGNOSIS. - Corallum (anthocyathus) compressed (GCD:LCD = 1.4-2.3). Angle of thecal edges, not

including crests, 19°-30°; angle of thecal faces, 20°-25°. One of the largest known specimens ( ^Watross

stn 5202) 12.1 x 17.3 mm in calicular diameter and 17.7 mm in height, whereas another, rejuvenescent coraUum

(MUSORSTOM 1 stn 57) is 15.9 x 17.6 mm in calicular diameter and 26.9 mm in height. Basal scar elliptical, up

to 4 0 mm in greater diameter. Base of each thecal edge bears a large (up to 4.5 mm), downward-projecting crest,

sometimes recurved toward the basal scar. Corallum reddish-brown, often with a more intense costal striping and/or

costal speckling near the calice. Septa hexamerally arranged in 4 full cycles (Sl-2>S3>S4). L^ge coral a may have

some additional pairs of S5 up to a total of 56-62 septa. Two crowns of pah occur, 12 small Pl-2 and 12 larger P3.

Papillose columella composed of 10-30 slender, cylindrical elements. , .

Anthocaulus stage rarely collected, attaining up to 6 mm in height, 7.7 mm in GCD, and having a robust

pedicel diameter of 3.4 mm. It may have 4 complete cycles of septa (Si-2>S3>S4) and prominent edge crests.

REMARKS. - Trochocyathus cooperi is one of 4 species in the subgenus that divides transversely. The other

3 species [T. gardineri (Vaughan, 1907); T. cepulla Cairns, 1995; and T. discus sp. nov.] are disco, dal (bowl¬

shaped), nonspinose, and nonnested. T. cooperi is more fully described and illustrated by CAIRNS ( 1994)

The description of Trochocyathus weberi Alcock, 1902a fits that of T. cooperi , except for the co our of .ts

corallum, which was stated to be "snow white". T. weberi was one of 8 species described by ALCOCK (1902a) in

his prelim, nary "Siboga" report but not included in his final report (ALCOCK, 1902c). It was never illustrated, a

station number was never cited, and it could not be found in the collections of the ZMA in 1994. Therefore, its

equivalence with T. cooperi remains questioned.

Distribution. — Philippines : Lubang Island; Sibuyan Sea; Sogod Bay, Leyte; Sulu Sea (Zamboanga

Peninsula, Sulu Archipelago, and Balabac Island); 34-96 m, with one assumed incorrect record at 918 m

(" Albatross " stn 5202). Indonesia: Banda Sea (Kai Islands); Flores Sea (Selayar Island, Sulawesi; Lintah S rai ),

Sunda Strait, Java Sea; 25-100 m. Elsewhere: Maidive Islands; northern Ryukyu Islands; Tahuata, Marquesas

Islands (reported herein); 70-80 m.

Trochocyathus (T.) gardineri (Vaughan, 1907)

Paracyathus gardineri Vaughan, 1907: 68-69, pi. 4, fig. 4.

Trochocyathus gardineri - CAIRNS, 1984: 16.

Source : MNHN Paris

1 12

S. D. CAIRNS & H. ZIBROWIUS

Not Paracyathus gardineri - Gardiner & Waugh, 1938: 183-184, pi. 3, fig. 5 [= Trochocyathus (T.) sp.].

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5567, 1 worn anthocyathus (USNM 97103).

Type Locality. — "Albatross" stn unknown: Hawaiian Islands (exact locality and depth unknown).

Diagnosis/Remarks. — Based on the single poorly-preserved specimen reported above, nothing can be added

to the original description of Vaughan, except for the range extension. T. gardineri reproduces by transverse

division, the anthocyathus having a Bat base with a central circular detachment scar about 5 mm in diameter.

At a calicular diameter ol 10-12 mm, the thecal walls abruptly turn upward, forming a cylindrical corallum. Septa

hexamcrally arranged in 4 complete cycles: Si-2>S4>S3. Three crowns of well-formed pali are present: Pi, Pi, and

P3. Columella papillose.

Distribution. — Philippines: Sulu Sea (Sulu Archipelago); 490 m. Elsewhere : Hawaiian Islands- 274-

470 m.

Trochocyathus (T.) discus sp. nov.

Figs 1 1 g-h, 12 a-c

0. ^JE1RI,AL exaM1NED/Types. — Indonesia. Karubar: stn 2, 1 dead corallum, paratype (USNM 97104). —

Stn 3, 20. holotype and 6 paratypes (MNHN), 3 paratypes (POLIPI), and 1 1 paratypes (USNM 97105).

Type Locality: Karubar stn 3: 5°48'S, 132°12'E (Kai Islands, Banda Sea), 278-300 m.

Etymology. — The species name (Latin discus , circular plate) refers to the shape of the anthocyathus.

Description. - Corallum (anthocyathus) discoidal and free, with a flat to slightly convex base. Central

region of base a circular detachment scar, measuring 3.6-4. 1 mm in diameter, with traces of 24 septa but no costae.

Instead, small (about 40 pm in width), pointed granules cover the scar region and surrounding base 4 or

- occurring across the width of a costa. Holotype 9.2 x 9.7 mm in diameter and 5.2 mm in height: largest known

specimen (Karubar stn 2) 12.2 mm in diameter. Toward calicular edge the costae are better defined by thin

-90 pm wide), deep intercostal furrows that become progressively wider toward the periphery. Costae convex

and about 0 4 mm w.de, bearing a coarser granulation than the base, only 3 or 4 blunt granules occurring across

“ f 3 C0S'a 12 a;b)- ^ ^ges and upper, outer regtons of septa reddish-brown in well-preserved

coral la, but base of colony and remaining septa, pah, and columella white. Anthocaulus unknown.

smalw!hheXrera,lly "T ?.ed m 4 CydeS’ 3 COmplete 4th cyc,e achieved at a GCD of about 8 mm. Specimens

verfical‘S T h T °f ^ h3ving 40'46 Septa>- S| high|y exsert (about > - 8 mm),

2/3 wfd,h o a ', g| y STrStTr gCS th3‘ CXtend 2/3 diStanCC 10 columella- S2 less exsert and about

3 width of an Si, also with slightly sinuous inner edges. S3 less exsert than S2 and about 2/3 width of an S2

m *”! m°dera‘ely jnuous inner edges. S4 dimorphic in width and exsertness: those adjacent to Si being slightly

more exsert and wider than an S3; those adjacent to S2 being slightly less exsert and less wide than an S3 Septal

P2 0 5 oTmmlid, ( T ^ gr3"UleS- lamellar’ but ->y 03-0.5 mm in 2h

nair wh'hin a 7 1 "T t ^ h'gher ,n foSSa; same size as * but recessed from the columella, each

of sen ta In T k y m l° US C°mm0n ?2 3 CheVr°n Pattern- Pali with granulation coarser than that

(fie Tie' h mnTTh5, n’r ?Ue Carinae' F0SSa Shal'°W’ containing 3 columella of 10-15 irregularly-shaped

( g- 1 1 g-h) papillae that are fused among themselves and to inner edges of Pi-2.

Remarks - Among the 4 species of Trochocyathus (Trochocyathus) that undergo transverse division this

ESa“!cvdeT''“ ^ I995- b“* «* ^ - -H-. bowUhaped XatZ wS

T clll hiin f ,Sma'Tr SIZC: *tS m°re regular ^ptal arrangement; and its dimorphic S4, those of

Distribution. — Indonesia-. Banda Sea (Kai Islands); 240-278 m.

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

113

Subgenus TROCHOCYATHUS (APLOCYATHUS) d'Orbigny, 1849

Trochocyathus (A.) brevispina sp. nov.

Figs 12 d-f

Odontocyathus ? sp. Alcock, 1902c: 24.

MATERIAL EXAMINED/TYPES. — Indonesia. "Siboga": stn 262, 1 paratype (ZMA Coel. 890).

Deki- stn 12 1 paratype (NNM 22499). — Stn 13, 1 paratype (NNM 22430). — Stn 41, 1 paratype (NNM 22500).

Karubar: stn 2, 1 paratype (POLIPI). — Stn 3, holotype (MNHN) and 8 paratypes (MNHN). — Stn 7, 19 paratypes

(USNM 97106).

Type LOCALITY. — Karubar stn 3: 5°47’40"S, 132°12’H"E (Kai Islands, Banda Sea), 278-300 m.

ETYMOLOGY. — The species name brevispina (Latin brevis , short + spina, spine) refers to the 6 short Ci

spines.

Description. — Corallum bowl-shaped and free, the base flat to evenly rounded, often with a imprint of

original substratum attachment or even an incorporated fragment of substratum at centre ot base. Holotype 20.3 x

22.1 mm in calicular diameter and 9.3 mm in height, containing 66 septa; largest specimen (Karubar stn 3)

22.3 x 26.1 mm in calicular diameter and 12.8 mm in height, containing 72 septa. Calice elliptical. GCD.LCD -

1.07-1.17. Costae well defined only near calice, where they are slightly convex and granular; remainder of base

usually worn and/or smooth, often with a hexagonal region of epitheca bounded by the 6 costal spines. All

specimens examined bear 6 short (not more than 5 mm in length) costal spines associated with the Ci, the spines

of some coralla being strongly compressed and ridged basally, these ridges extending to the centre of the base.

Well-preserved coralla black-brown in colour.

Septa hexamerally arranged in 5 cycles, several pairs of S5 present already in small coralla ol 12 mm GCD, but

even in large specimens 5th cycle never complete, the largest corallum having only 12 pairs of S5, or a total of

72 septa. Septal formula: Si-2>S4>S3>S5. Si-2 are 3.5-4.0 mm exsert, robust, having thick, straight inner edges

bordered by a well-defined palus. S3 slightly less exsert (2.5 mm) and about 3/4 width of an Si -2, each S3 also

bordered by a palus of equivalent size to a P1-2 but recessed slightly from the columella. S4 of equal exsertness to

S3, but slightly wider than S3, unless flanked by a pair of S5, in which case the S4 are narrower than an S3.

S4 flanked by S5 bear a palus of similar size to the others, but recessed even more from the columella than the P3.

When present, S5 are least exsert and least wide class of septa. All pali (Pi-4) thick, about 1.5 mm wide, having

straight, vertical inner edges; pali separated from their bordering septa by narrow (about 0.8 mm), deep notches.

Fossa shallow, containing a papillose columella composed of 9-20 small, irregularly-shaped pillars.

Remarks. — Three Recent species are now attributed to the subgenus Aplocyathus. Trochocyathus brevispina

differs from T. hastatus Bourne, 1903 (Figs 13 a-c), in always having 6 short costal spines, not 5 elongate ones. It

also differs in: having a larger corallum with more septa, the maximum known size of T. hastatus is 18 mm GCD

with 48 septa; having Si=S2, T. hastatus having S i>S2; and having equal-sized Pi-4, those ol T. hastatus being

unequal in size. T. brevispina is compared to T. longispina in the account of that species (see below).

Distribution. — Indonesia : Banda Sea (Kai Islands); 240-282 m (a dead corallum from 560 m).

Trochocyathus (A.) longispina sp. nov.

Figs 12 g-i

Material EXAMINED/TYPES. — Philippines. "Albatross": stn 5331, 1 paratype (USNM 97108). — Stn 5506,

1 paratype (USNM 97109). — Stn 5537, 1 paratype (USNM 97667).

Musorstom 1: stn 50, holotype (MNHN).

Musorstom 2: stn 44, 1 paratype (USNM 97111).

Indonesia. "Albatross": stn 5592, 1 paratype (USNM 97110).

Source : MNHN , Paris

114

S. D. CAIRNS & H. ZIBROWIUS

Type Locality. — Musorstom 1 stn 50: 13°49'N, 120°01'E (Lubang Island, Luzon), 415-510 m.

ETYMOLOGY. — The species name longispina (Latin longus, long + spina, spine) refers to the 6 elongate Ci

spines.

DESCRIPTION. — Corallum bowl-shaped and free, the base flat to slightly convex, often maintaining the scar of

original attachment. At a diameter of 9-1 1 mm the horizontal base is sharply inflected upward at 65-70°. Largest

specimen (holotype) 17.0 mm in calicular diameter and 12.7 mm in height. Calice circular. Costae well defined

only near calicular edge, otherwise lower theca and base uniformly granular. Six slender, elongate (up to 10 mm)

costal spines (Ci) project horizontally from the outer edge of base; spines circular in cross section. Corallum

white.

Septa hexamerally arranged in 4 complete cycles in all specimens reported (Si-2>S3>S4). Each Si -2 up to

3.0 mm exsert, with a slightly sinuous inner edge, bearing a small palus 0.5- 1.0 mm in width that contributes to

a crown of 12 Pi-2 located close to the columella. S3 less exsert (about 1.7 mm) and about 2/3 width of Si-2,

having moderately sinuous inner edges. Each S3 bordered by a large, lamellar P3 about 2.0 mm wide, the 12 P3

forming a crown that rises higher in the fossa and is slightly more recessed than the P1-2 crown. S4 about 0.8 mm

exsert, 1/2 the width of S3, without lobes. Fossa shallow, containing a prominent columella consisting of

irregularly-shaped papillae.

Remarks. — Trochocyathus longispina differs from T. brevispina sp. nov. in having a smaller corallum, less

septa, a circular (not elliptical) calice, a white (not black-brown) corallum, and longer costal spines that are circular

(not flattened) in cross section. T. hastatus Bourne, 1903 (Figs 13 a-c), the other Recent species in the subgenus,

differs from T. longispina in having S i>S2, a porcellaneous base, a brown-black corallum, and only 5 costal

spines. T. longispina also resembles Stephanocyathus (Acinocyathus) explanans (Marenzeller, 1904), both species

having 6 long costal spines, a white corallum, and a similar palar arrangement, but differs in having a smaller

corallum and correspondingly less septa and pali (48 vs 72 septa), and sinuous inner septal edges, especially

of the S3. The inner septal edges of S. explanans are straight, which is one of the few characters that differentiate

the subgenera Stephanocyathus (Acinocyathus) and Trochocyathus (Aplocyathus).

DISTRIBUTION. — Philippines: Zambalas, Luzon; Lubang Island; Sibuyan and Bohol Seas; 326-760 m.

Malaysia: Celebes Sea (Sabah); 558 m.

Genus TETHOCYATHUS Kuhn, 1933

Tethocyathus virgatus (Alcock, 1902)

Trochocyathus (Tethocyathus) virgatus Alcock, 1902a: 98-99; 1902c: 16-17, pi. 2, fig. 13. — Faustino, 1927: 82-83,

pi. 7, fig. 10.

Tethocyathus virgatus - Cairns, 1995: 65-66, pi. 16, figs c-f (synonymy).

MATERIAL EXAMINED. — Philippines. Musorstom 1: stn 32, 2 (MNHN). — Stn 63, 2 (MNHN).

Musorstom 2: stn 2, 1 (USNM 97113). — Stn 33, 2 (MNHN).

MUSORSTOM 3: stn 108, 4 (USNM 97114).

Indonesia. Mortensens Java-S.A. Expedition: stn 15, 4: 1 (NNM 23201), 3 (ZMUC).

Karubar: stn 16, 1 (USNM 97115). — Stn 27, 1 (POLIPI). — Stn 44, 2 (MNHN).

Type Locality. — "Siboga" stns 96 and 105: Sulu Archipelago, 275 m.

DIAGNOSIS. — Corallum elongate-conical to subcylindrical, firmly attached through a robust pedicel

(PD:GCD = 0.54-0.84). Largest Philippine specimen (MUSORSTOM 1 stn 63) 13.8 mm in GCD and 14.9 mm in

height, elsewhere coralla reported up to 16.3 mm in GCD (CAIRNS, 1995). Epitheca usually present, but variable

in development. CSi darkly pigmented, highlighting hexameral symmetry of corallum; however, in some

specimens CS2 are also pigmented, and in a few coralla upper edges of all septa are dark blackish-brown. Septa

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTIN1A

115

hexamerally arranged in 4 complete cycles: Si>S2>S4>S3, the Si usually quite thick and "swollen" appearing.

Three crowns of thick, ridged pali (Pi, P2, and P3), arranged in typical trochocyathid fashion. Columella of

moderate depth; papillose columella composed of 3-40 tuberculate pillars.

REMARKS. — Two specimens from MUSORSTOM 3 stn 108 show evidence of having been bored by an

acrothoracican cirripede crustacean, which is not uncommon in this genus (ZlBROWlUS, 1980; CAIRNS, 1995).

Tethocyathus virgatus is more fully described and illustrated by CAIRNS (1995).

DISTRIBUTION. — Philippines : Lubang Island; Verde Island Passage; Sulu Sea (Sulu Archipelago); 137-

275 m. Indonesia : Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar Islands); Flores Sea (Sumbawa);

240-315 m. Elsewhere : ridges north of New Zealand; 142-530 m.

Genus BOURNEOTROCHUS Wells, 1984

Bourneotrochus stellulatus (Cairns, 1984)

Deliocyathus stellulatus Cairns, 1984: 15-16, pi. 3, figs C-D.

Bourneotrochus veroni Wells, 1984: 213-214, pi. 3, figs 7-18.

Bourneotrochus stellulatus - CAIRNS, 1995: 71-72, pi. 18, figs f, i, pi. 19, figs a-c (synonymy).

MATERIAL EXAMINED. — Indonesia. Deki: stn 48, 3 (NNM 22760).

Snellrjs 2: stn 81.2, 2 (NNM 23075).

Karubar: stn 3, 2 (USNM 97118). — Stn 7, 2 (POLIPI). — Stn 18, 6 (MNHN).

TYPE Locality. — 19°48’N, 154°58'W (Hawaiian Islands), 337 m.

Diagnosis/Remarks. — This species was redescribed and illustrated by Cairns (1995) based on specimens

from ridges north of New Zealand. Six additional specimens from 3 stations are reported above. The anthocyathus

is relatively small (usually less than 6 mm GCD), discoidal to cylindrical corallum with a basal scar resulting

from transverse division. Each Ci bears a prominent costal spine. Septa are hexamerally arranged in 4 cycles, the

4th incomplete, usually resulting in 36 septa. Pali occur before all but the last cycle of septa and the columella is

papillose.

As previously discussed in this paper, at least 4 Recent species of Trochocyathus reproduce by transverse

division, and 5 species bear costal spines, which are present as edge spines in 3 species and as 5-1 1 C1-2 spines in

another 2 [i.e., Trochocyathus (Aplocyathus)]. One species, T. cooperi (Gardiner, 1905), has both edge spines and

transverse division, but no species of Trochocyathus has the character combination of transverse division and 6 Ci

spines, as does Bourneotrochus stellulatus. Nevertheless, a strong relationship to Trochocyathus is suggested,

perhaps even at the subgeneric level.

Distribution. — Indonesia : Banda Sea (Kai and Tanimbar Islands); 263-340 m. Elsewhere : Queensland; ridges

north of New Zealand; Chesterfield Islands; Funafuti and Tuvalu; Cook Islands; Hawaiian Islands; 274-476 m.

Pleistocene of Vanuatu (Wells, 1984).

Genus PARACYATHUS H. Milne Edwards & Haime, 1848

Paracyathus rotundatus Semper, 1872

Figs 13 d-e

Paracyathus rotundatus Semper, 1872: 253-254, pi. 20, figs 15a-b. — FAUSTINO, 1927: 72-73, pi. 5, figs 13-14.

Material EXAMINED. — Philippines. "Albatross": stn 5164, 5 (USNM 97120). Holotype (NMW 8177).

Musorstom 2: stn 9, 3 (MNHN).

Source :

116

S. D. CAIRNS & H. ZIBROWIUS

Indonesia. Deki: stn 68. 5 (NNM 22630). — Sin 71. 4 (NNM 22629). — Sin 82, 1 (NNM 22631).

South China Sea. Pelau Redang, Malaysia, 1 (BMNH).

Papua New Guinea. "Alpha Helix stn 79-M-21, 3 (USNM 80015).

Type Locality. — Lapinig Canal, Philippines, 11-18 m.

Description. — Corallum elongate-conical to trochoid, straight, and attached by a robust pedicel up to

0.5 GCD. SEMPER's type is slenderer than indicated in the original description: height 16 mm (vs 15 mm), calice

10.5 x 12.2 mm (vs 12 x 15.5 mm). Specimen illustrated herein 11.6 x 13.8 mm in calicular diameter, 20.2 mm

in height, and 5.7 mm in pedicel diameter. GCD:LCD = 1.14-1.63. Costae Oat and equal in width (about 0.4 mm),

covered with low, rounded granules — approximately 3 granules occurring across width of a costa. Septa, paliform

lobes, and columella pigmented a dark brown, purple-grey, blue, or blackish-brown. Theca usually white, but

occasionally also darkly pigmented at calicular edge.

Septa hexamerally arranged in 5 cycles, the last cycle never complete. Largest specimen with 84 septa

(5th cycle lacking 6 pairs of septa); a specimen 7.5 mm GCD with 66 septa. Si-2 about 1 mm exsert, having

vertical, straight inner edges that bear 1-3 paliform lobes, the uppermost lobe occurring about 1/2 distance to

columella. S3 only slightly less exsert than S1-2, each bearing 3-5 narrow paliform lobes, the uppermost lobe

reaching slightly higher than the P1-2. S4 less exsert and less wide than S3, their narrow paliform lobes mingling

with those of S3 low in fossa. S5 smallest septa, their paliform lobes fusing with those of S4 in a complex

tuberculate region. Fossa relatively deep. Columella trabecular to papillose, concave, and indistinguishable from

lowermost paliform lobes of all septa (nondiscrete).

REMARKS. — This is believed to be the first report of additional specimens of P. rotundatus since its original

description, which was based on one specimen.

Distribution. — Philippines : Lubang Island; Bohol; Sulu Sea (Sulu Archipelago); 18-66 m. Indonesia :

Sunda Strait, Java Sea; 35-54 m. Elsewhere : Gulf of Papua, Papua New Guinea; South China Sea (Malaysia);

55 m.

Paracyathus sp.

Figs 13 g-i

Paracyathus defilippi - MOSELEY, 1881: 144 (in part: "Challenger" stn 190). [Not Paracyathus defilippi Duchassaing &

Michelotti, I860].

Paracyathus agassizi - ALCOCK, 1902c: 18. [Not Paracyathus agassizi Duncan, 1873].

Material EXAMINED. — Indonesia. "Challenger": stn 190, 1 (BMNH).

"Siboga": stn 256, 1 (ZMA Coel. 1306).

Deki: unnumbered station, Ambon, unknown depth, 2 (NNM).

Diagnosis. — Only 4 specimens of this species are known, the largest ("Siboga" stn 256) 8.8 x 9.3 in

calicular diameter and 9.4 mm in height, and the "Challenger" specimen 3.9 x 4.5 mm in calicular diameter and

5.6 mm in height. Corallum elongate-conical and solidly attached through a broad, polycyclic pedicel and base.

Costae poorly developed, covered with low, rounded granules. Theca, septa, and pali all mottled with a brown

pigmentation, the columella being uniformly brown. Septa hexamerally arranged in 4 complete cycles according to

formula: Si-2>S4>S3. S 1 -2 highly exsert (up to 2.5 mm) and, along with their adjacent pairs of S4, form

12 rectangular lancets. Inner edges of S1-2 bear 1 or 2 narrow paliform lobes. S3 about 3/4 width of S1-2, each

bearing 1-3 slender paliform lobes. S4 usually slightly wider than S3, also bearing paliform lobes. Septal faces

bear prominent granules and all paliform lobes are highly sinuous and carinate. Columella papillose, consisting of

a field of numerous irregularly-shaped rods.

REMARKS. — The 4 specimens available for study of this species are not considered to be enough material to

properly characterise the species, but figures are provided to document this form.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

117

DISTRIBUTION. — Indonesia : Banda Sea (Kai and Ambon Islands); Arafura Sea; 90-397 m.

Genus STEPHANOCYATHUS Seguenza, 1864

Subgenus STEPHANOCYATHUS (STEPHANOCYATHUS) Seguenza, 1864

Stephanocyathus (S.) regius sp. nov.

Figs 14 a-c

Stephanocyathus nobilis - ZOU, 1988: 74-75 (in part: pi. 1, figs 4-7). [Not Ceratotrochus nobilis Moseley, 1873],

MATERIAL EXAMINED/TYPES. — Philippines. " Albatross stn 5444, 2 paratypes (USNM 97123). — Stn 5445,

16 paratypes (USNM 97124). — Stn 5447, 2 paratypes (USNM 97125).

Musorstom 1: stn 49, 7 paratypes (USNM 97129). — Stn 54, 2 paratypes (MNHN).

Indonesia. "Albatross": stn 5585, 1 paratype (USNM 97126). — Stn 5650, 3 paratypes (USNM 97127). —

Stn 5670, 1 paratype (USNM 97128).

"Hakuho Maru": stn KH72-1-26, holotype (USNM 97122) and 1 paratype (USNM 97130).

Corindon 2: stn 241, 2 paratypes (MNHN).

New Zealand Region. "Tangaroa": stn T243, 1 paratype (USNM 94362).

TYPE Locality. — "Hakuho Maru" stn KH72-1-26: 9°27'S, 127°58.6'E (Timor Sea, south of Leti Islands),

610-690 m.

ETYMOLOGY. — The shape of the corallum in this genus occasionally has been compared to a crown. This

theme is reiterated here (Latin regius, royal).

DESCRIPTION. — Corallum bowl-shaped, with a flat to slightly convex base. Holotype 29.0 mm in calicular

diameter and 12.5 mm in height; largest specimen ("Hakuho Maru" stn KH72-1-26) 32 mm in calicular diameter

and 11.5 mm in height. Centre of base often displays an irregularly-shaped imprint of detachment from

substratum; otherwise, base covered with low, convex, coarsely granular ridges, the Ci-2 sometimes slightly more

prominent than others. Corallum uniformly white.

Septa hexamerally arranged in 5 full cycles (Si>S2>S3>S4>Ss), some specimens as small as 18 mm GCD

having a full 5th cycle, whereas other larger specimens (e.g., 25-32 mm GCD) may lack several pairs of S5,

resulting in 86-94 septa. Si highly exsert (3. 9-4. 2 mm), having straight inner edges, each Si internally bordered

by a broad (up to 2.2 mm) notch and 2 or 3 paliform lobes, each lobe decreasing in size toward columella. S2

virtually the same as Si, but sometimes slightly less exsert. S3 less exsert (about 3.3 mm), only about 4/5 width

of an Si or S2, each S3 bordered by a narrow notch and 1-4 paliform lobes, the outermost lobe sometimes quite

wide (2.0 mm). Outermost lobes of P3 crown always stand higher in fossa than those of Pi -2 and are

more recessed from the columella. S4 less exsert (about 2.5 mm), about 4/5 width of an S3, with slightly sinuous

edges, and bordered by 3 or 4 paliform lobes, the outermost lobe being the largest and occurring higher in the fossa

and farther from the columella than P3. Innermost P4 often fuse with their common P3 near columella. S5 least

exsert septa (about 1.5 mm), with slightly sinuous inner edges, becoming rudimentary lower in fossa, bearing no

paliform lobes. Thus, only the Si and S5 are independent septa, the inner edges of the Pi reaching the columella

and those of the S5 diminishing in size, whereas the inner P3 merge with their common P2 and the inner P4

with their common P3. Fossa shallow, containing an elliptical field of 20-30 small densely fused papillae, the

innermost Pi -4 being indistinguishable in size and shape from the columellar elements.

Remarks. — Among the 7 species in the nominate subgenus, S. regius most closely resembles S. paliferus

Cairns, 1977 (known only from the western Atlantic at 229-715 m), both species having a similarly-shaped

corallum and well-developed paliform lobes. S. regius differs in having multiple paliform lobes and in lacking

small costal spines on the C1-2. S. regius also resembles Vaughanella multipalifera Cairns, 1995 in most

-haracters but differs in having an unattached corallum.

Source : MNHN , Paris

118

S. D. CAIRNS & H. ZIBROWIUS

ALCOCK (1902a) described 2 additional species of Stephanocyathus, each based on a small, worn corallum:

? Sabinotrochus flatiliseptis ("Siboga" stn 211, ZMA Coel. 1315, CD = 11.6 mm, Fig. 14 i) and

? Sabinotrochus bipatella ("Siboga" stn 52, ZMA Coel. 1314, CD = 8.1 mm. Fig. 14 f)- Given their small size

and poor state of preservation, they cannot be confidently identified as one of the better known species of

Stephanocyathus. Although they are illustrated herein, their names are not included in Table 1.

DISTRIBUTION. — Philippines: Lubang Island; Lagonoy Gulf; 563-975 m. Indonesia/Malaysia: Celebes Sea

(Sulu Archipelago); Makassar Strait; Timor Sea (south of Leti Islands); 690-2160 m. Elsewhere : South China

Sea; Malaysia (Celebes Sea off Sabah); Kermadec Islands (Macauley Island); 1035-1896 m.

Subgenus STEPHANOCYATHUS (ACINOCYATHUS) Wells, 1984

Stephanocyathus (A.) spiniger (Marenzeller, 1888)

Figs 13 f, 14 d

Stephanotrochus spiniger Marenzeller, 1888: 20-21.

Odontocyathus sexradiis Alcock, 1902a: 100-101.

Odontocyathus Stella Alcock, 1902b: 119-120.

? Odontocyathus coloradus Smith, 1913: 288, pi. 18, fig. 8.

Stephanocyathus (A.) spiniger - Wells, 1984: 209, pi. 2, figs 10-13. — Cairns & Parker, 1992: 26-27, pi. 7, figs g-i

(synonymy). — Cairns, 1994: 57, pi. 25, figs a-c (synonymy); 1995: 67-68, pi. 17, figs d-f, pi. 18, fig. c.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5273, 1 (USNM 97131). — Stn 5369, 15 (USNM

97132). — Stn 5371, 9 (USNM 97133). — Stn 5372, 1 (USNM 97134). — Stn 5440, 1 (USNM 97135). — Stn 5541,

2 (USNM 97136). — Stn 5545, 2 (USNM 97137).

MUSORSTOM 1: stn 2, 1 (MNHN). — Stn 25, 1 (USNM 97139). — Stn 61, 1 (MNHN). — Stn 63, 1 (MNHN). —

Stn 65, 1 (USNM 97140).

SlPHlLEXP: stn 78-SP40, 2 (USNM 80005).

MUSORSTOM 2: stn 12, 1 (USNM 97141). — Stn 18, 2 (MNHN). — Stn 21, 1 (USNM 97142). — Stn 26, 1 (USNM

97143). — Stn 62, 2: 1 (MNHN), 1 (POLIPI).

MUSORSTOM 3: stn 88, 1 (USNM 97144). — Stn 96, 1 (MNHN). — Stn 97, 1 (USNM 97145). — Stn 98, 1 (USNM

97146). _ Stn 99, 1 (USNM 97147). — Stn 102, 1 (MNHN). — Stn 103, 1 (MNHN). — Stn 109, 1 (MNHN). — Stn 120,

1 (MNHN). — Stn 143, 4 (MNHN).

Indonesia. Deki: stn 2, 12 (NNM 22514). — Stn 3. 1 (NNM 22519). — Stn 4, 1 (NNM 22520). — Stn 5, 3 (NNM

22515). — Stn 6, 3 (NNM 22516). — Stn 22, 3 (NNM 22522). — Stn 41, 1 (NNM 22523). — Stn 42, 1 (NNM 22517).

— Stn 49, 1 (NNM ). — Stn 50, 3 (NNM 22525). — Stn 52, 1 (NNM 22526). — Stn 63, 2 (NNM 22518).

Karubar: stn 61, 1 (MNHN). — Stn 86, 1 (MNHN).

South China Sea. "Hakuho Maru": stn KH72-1-50, 1 (USNM 97150).

Type Locality. — Sagami Bay, Honshu, Japan (depth not given).

Diagnosis/Remarks. — This common, relatively shallow-water Indo-West Pacific species was redescribed by

Cairns & Parker (1992) and Cairns (1994). It is characterised by having an unattached, bowl-shaped corallum

with 6 long (up to 25 mm), slender costal spines (Ci) that project horizontally from the edge of the base. Septa

hexamerally arranged in 5 cycles (Si»S2>S3>S4>Ss); however 1 large specimen (Karubar stn 86) of 40.6 mm

GCD has 6 additional pairs of S6, 1 S6 on each side of an Si, resulting in 1 10 septa. Largest Philippine specimen

("Albatross" stn 5541) 42 mm in calicular diameter, as measured from outer edge to outer edge of opposing exsert

Si. Si and S2 extremely exsert, each forming a rectangular calicular lancet with its adjacent S5 (or S6). Si, and

occasionally S2, pigmented a dark brown or black. Three crowns of broad paliform lobes (Pi-2, P3, and P4)

present, the Pi -2 often integrated into the columella.

DISTRIBUTION. — Philippines: Lingayen Gulf, Luzon; Lubang Island; Verde Island Passage; Sibuyan,

Visayan, and Bohol Seas; Sulu Sea (Semirara Islands and Sulu Archipelago); 152-401 m. ? Neogene of Masbate,

Philippines (Smith, 1913). Indonesia: Banda Sea (Kai Islands); Arafura Sea (south of Tanimbar Islands);

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

119

210-352 m. Elsewhere : widespread throughout Indo-West Pacific from southwestern Indian Ocean to Japan,

including South Australia and ridges north of New Zealand, and Charlotte Bank (South China Sea); 120-695 m.

Stephanocyathus (A.) explanans (Marenzeller, 1904)

Fig. 14 e

Stephanotrochus explanans Marenzeller, 1904a: 304-307, pi. 18, figs 19a-b.

Stephanocyathus nobilis - BOSHOFF, 1981: 39. [Not Ceratotrochus nobilis Moseley, 1873].

Stephanocyathus (A.) explanans - CAIRNS & KELLER, 1993: 243-244.

MATERIAL EXAMINED. — Indonesia. "Albatross": stn 5589, 4 (USNM 97151).

Karubar: stn 35, 1 (USNM 97152). — Stn 39, 10 (MNHN). — Stn 40, 9 (POLIP1). — Stn 58, 5 (USNM 97154). —

Stn 59, 14 (MNHN). — Stn 70, 2 (MNHN). — Stn 71, 1 1 (POLIPI). — Stn 75, 7 (POLIPI). — Stn 87, 1 (USNM 97156).

TYPE Locality. — "Valdivia" stns 194, 243, and 245: off Sumatra, Zanzibar Island, and Pemba, 245-614 m.

Description. — Corallum bowl-shaped, with a slightly rounded base, the thecal edges diverging at a 40°-45°

angle. Most coralla 21-27 mm in calicular diameter, the largest known corallum (KARUBAR stn 35) 27 mm in

calicular diameter and 20. 1 mm in height. All specimens bear 6 relatively short, slender, tapered, straight costal

spines (basal diameter 1.5-1. 7 mm) that project horizontally from the Ci at point of upward thecal inflection.

Spines best known from juveniles, where they are up to 9 mm long; in larger (older) coralla, Ci spines are either

broken or reduced to small nubs. In about 1/4 of specimens examined, 1-5 additional costal spines present on C2.

All costal spines circular in cross section. Lower corallum, to a level just above costal spines, usually eroded, as

though submerged in a soft substratum. Theca above this region costate, the costae slightly convex and granular.

Corallum white.

Septa hexamerally arranged in 5 cycles, the 5th cycle never complete: the most common septal complement is

72, achieved by having a pair of S5 in each of the 12 quarter systems adjacent to the 6 Si. S 1-2 about 3.3 mm

exsert, having straight, vertical inner edges, each bordered internally by a deep, broad (up to 1.6 mm) notch and a

rather narrow, lamellar paliform lobe only about 1 mm wide, which is often merged into the columella. S3 about

2 mm exsert, 3/4 width of an Si-2, and bordered by a narrow (about 0.5 mm) notch and a prominent, broad (2.0-

3.5 mm) paliform lobe. S4 that are flanked by S5 almost as wide and exsert as an S3, but bear a narrow paliform

lobe (about 0.9 mm), which is often fused into or subsumed by the P3. Unflanked S4 only 1.5 mm exsert, about

half width of an S3, becoming rudimentary lower in fossa. Because Pi -2 are directly adjacent to the columella, and

P4 are often fused into the P3 or poorly developed, many coralla appear to have only 1 crown of 12 massive P3.

S5 resemble unflanked S4. Inner edges of all septa and paliform lobes straight. Calicular margin serrate, but not

produced into lancets. Fossa relatively shallow. Columella composed of 8-10 granular papillae that are circular in

cross section, which distinguish them from the closely adjacent lamellar Pi-2-

Remarks. — Stephanocyathus explanans differs from the only other Recent species in this subgenus,

S. spiniger, in having a serrate (not lanceted) calicular edge; 48-72 (not 96) septa; 6-11 relatively short costal

spines that are circular in cross section (vs 6 elongate, basally compressed Cl); a completely white corallum;

equal-sized SI and S2; and wider and more developed P3.

Distribution. — Indonesia : Arafura Sea (southeast of Tanimbar Islands); 405-1016 m. Elsewhere: Malaysia

(Celebes Sea off Sabah); South Africa, Madagascar, and Tanzania; west of Sumatra; 183-614 m.

Subgenus STEPHANOCYATHUS (ODONTOCYATHUS) Moseley, 1881

Stephanocyathus (O.) weberianus (Alcock, 1902)

Figs 14 g-h

Stephanotrochus weberianus Alcock, 1902a: 101-102; 1902c: 25, pi. 3, figs 22, 22a.

Source : MNHN, Paris

120

S. D. CAIRNS & H. ZIBROWIUS

Stephanotrochus sibogae Alcock, 1902a: 102-103; 1902c: 25-26, pi. 3, figs 23, 23a.

Stephanotrochus sp. - Alcock, 1902c: 26.

Stephanocyathus (O.) ixine Squires, 1958: 54 (in part: "Albatross" stn 5545, pi. 8, figs 3-4)

Stephanocyathus nobilis - Zou, 1988: 74-75 (in part: pi. 1, figs 1-3). [Not Ceratotrochus nobtlts Moseley 1873]

Stephanocyathus (O.) weberianus - CAIRNS, 1994: 57-58, pi. 25, figs d-f (synonymy); 1995: 68-69, pi. 17, figs g-i

(synonymy).

MATERIAL EXAMINED. — Philippines. " Albatross stn 5348, 1 (USNM 97157). — Stn 5349, 1 (USNM 97158).

— Stn 5444, 2 (USNM 97159). — Stn 5445, 54 (USNM 46819).

Musorstom 1: stn 49, 4 (MNHN). — Stn 54, 1 (USNM 97166).

Indonesia "Albatross": stn 5586, 4 (USNM 97160). — Stn 5587, 1 (USNM 97161). — Stn 5601, 1 (USNM

97162) — Stn 5648, 1 (USNM 97163). — Stn 5650, 1 (USNM 97164). — Stn 5671, 1 (USNM 97165).

"Hakuho Man,": stn KH72-1-26, 6: 4 (USNM 97168), 2 (ORI).

Karubar: stn 20, 2 (POL1PI). — Stn 89. 1 (MNHN). — Stn 91, 33: 26 (USNM 97167), 7 (MNHN).

Type Locality. — "Siboga" stn 284: 8°43.1'S, 127°16.7'E (Timor Sea), 828 m.

DIAGNOSIS. — Corallum bowl-shaped, often with an eroded lower surface. Largest known specimen (Karubar

stn 91) 54 mm in calicular diameter and 30 mm in height. 12 to 18 costal tubercles correspond to the 12 Ci-2 and

the C3 of those half systems that possess 4 C5. In some specimens, tubercles are replaced by a thickened rim that

encircles the base of the corallum. Corallum white. Septa hexamerally arranged in 5 cycles (Si-2>S3>S4»Ss) —

the 5th cycle never complete, 72 being a common number of septa. Paliform lobes present before all but last

cycle of septa. Columella papillose to lamellar (Fig. 14h), the latter aspect a result of the intermingling of

lamellar Pi-2-

REMARKS. — This species is more fully described and illustrated by CAIRNS (1994).

DISTRIBUTION. — Philippines: Lubang Island; north of Samar; Sulu Sea (Zamboanga Peninsula); South

China Sea (Palawan); 563-1388 m. Indonesia: Makassar Strait; Molucca Sea; Ceram Sea; Banda Sea (Kai Islands;

Teluk Bone, Sulawesi); Timor Sea (southwest of Tanimbar Islands and south of Leti and Timor Islands); 567-

1756 m. Elsewhere: Malaysia (Celebes Sea off Sabah); South China Sea; Japan (Honshu, Kyushu, northern

Ryukyu Islands); Lord Howe Seamount Chain; Chesterfield Islands; 206-1302 m, although most records deeper

than 700 m.

Genus ERICIOCYATHUS nov.

Type Species. — Ericiocyathus echinatus , here designated.

Diagnosis. — Corallum solitary, bowl-shaped, and free. Septotheca costate, each C1-2 bearing a series of

elongate spines. Pali in a crown before penultimate septal cycle (P3). Columella papillose.

Remarks. — Among the caryophylliid genera, Ericiocyathus is most similar to Stephanocyathus

(Odontocyathus), both taxa having similarly shaped coralla of approximately the same size, and costal tubercles or

spines on their Ci -2. Ericiocyathus differs in having pali associated only with the S3, whereas S. ( Odontocyathus )

has 3 (or 4) cycles of paliform lobes. Also, S. ( Odontocyathus ) has only 12-18 costal tubercles (Ci-2, 3) relegated

to the lower, outer edges of the corallum, whereas Ericiocyathus has a series of elongate spines along each C1-2.

Stephanocyathus (Acinocyathus) and Trochocyathus (Aplocyathus) are also similar to Ericiocyathus in having

elongate costal spines, but differ in having a total of only 5 or 6 spines (one per Ci), and in having Pi -2 in

addition to P3.

ETYMOLOGY. — The genus name Ericiocyathus (Latin ericius, hedgehog + cyathus , cup) refers to the spiny

corallum. Gender: masculine.

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

121

Ericiocyathus echinatus sp. nov.

Figs 15 a-b

MATERIAL EXAMINED/TYPES. — Philippines. "Albatross": sin 5423, 1 paratype (USNM M238415). —

Stn 5425, holotype (USNM 97168). — Stn 5429, 1 paratype (USNM 97668). — Stn 5487, 8 paratypes (USNM 97170).

— Stn 5512, I paratype (USNM 97171). — Stn 5513, 3 paratypes (USNM 97172).

TYPE Locality. — "Albatross" stn 5425: 9°38'N, 121° 1 l'E (Cagayan Island, Sulu Sea), 907 m.

ETYMOLOGY. — The species name (Latin echinatus, covered with spines) refers to the many costal spines.

Description. — Corallum bowl-shaped and unattached, the base being evenly rounded and the thecal edge

angle 43°-53°. Small coralla bowl-shaped, but older coralla have high thecal edges, the largest specimen (the

holotype) 18.6 mm in calicular diameter and 16.8 mm in height. Each Ci-2 bears a row of long, tapered costal

spines that project perpendicularly to the theca. These spines are short (eroded) near the base, but on the sides of

the corallum costal spines up to 6.0 mm long with a basal diameter of 1.2 mm. Spines occur approximately every

2 mm, up to 6 aligned on each C1-2; the holotype has at least 60 spines, whereas smaller specimens have fewer.

Costae developed as low, convex, granulated ridges only near calicular edge. Corallum white.

Septa hexamerally arranged in 4 complete cycles (Si-2>S3>S4), septa of the 4th cycle beginning to be present

at a GCD of 5 mm. Si -2 about 3 mm exsert, having straight, vertical inner edges that extend about 2/3 distance to

columella. Si -2 do not have paliform lobes or pali. S3 about half as exsert as and 3/4 width of Si -2, with straight

to only slightly sinuous inner edges. S4 lowest septa, 2/3 width of an S3, having straight inner edges, becoming

rudimentary lower in fossa. A crown of 12 thin, lamellar, planar P3, each 1.5-2. 5 mm wide, encircles a columella

composed of 3-6 papillae or short lamellae. Fossa relatively shallow, the pali rising high in the fossa contrasting

with the columella, which is deeply seated.

DISTRIBUTION. — Philippines : Bohol Sea (Sogod Bay and Iligan Bay); Sulu Sea (Cagayan Island and Honda

Bay, Palawan); 814-1401 m.

Genus DELTOCYATHUS H. Milne Edwards & Haime, 1848

Key to the 7 species of Deltocyathus known from the Philippine-Indonesian region

1 . Adult corallum with 5 full cycles of septa (96) . 2

— Adult corallum with 4 cycles of septa or an incomplete 5th cycle (48-80 septa) . 4

2. P4 much larger than all other pali . D. rotulus

— P4 of similar size to other pali . 3

3. Base flat, no scar; all septa project an equal distance beyond calicular margin; S5 well

developed . D. magnificus

— Base convex (bowl-shaped), usually with a central scar or eroded region; S5 project much

less than S1-4 beyond calicular margin; S5 rudimentary, often absent near calicular edge ...

. D. suluensis

4. All septa project beyond calicular edge an equal distance; costae coarsely dentate .

. D. vaughani

— Septa of higher cycle less prominent than others; costae finely dentate or granular . 5

5. CS3 thick, projecting beyond calicular margin as 12 triangular lancet . D. Stella

— CS3 of normal thickness, although they may project as lancets . 6

Source : MNHN, Paris

122

S. D. CAIRNS & H. Z1BR0WIUS

6. Corallum often having several pairs of S5 (up to 80 septa); S4 fuse to P3 relatively high

in fossa; CS3 often hyperextended in a lobate structure . D. andamanicus

— Corallum always having only 4 cycles of septa (48 septa); S4 much smaller, fusing to S3

very low in fossa near columella; CS3 never hyperextended . D. philippinensis

Deltocyathus vaughani Yabe & Eguchi, 1932

Levipalifer orientalis Vaughan, 1900: 201-202, pi. 16, figs 3-7 (junior secondary homonym of Deltocyathus orientalis

Duncan, 1876).

Deltocyathus vaughani Yabe & Eguchi, 1932b: 388-389. — Zibrowius & Grygier, 1985: 121, fig. 12. — CAIRNS,

1994 : 54-55, pi. 23, figs i-j, pi. 24, figs a-c, f (synonymy).

Material EXAMINED. — Philippines. " Albatross ": stn 5110, 10 (USNM 40051). — Stn 5198, 8 (USNM

97173). — Stn 5529, 1 (USNM 97174). — Stn 5536, 2 (USNM 97175).

Indonesia. Deki: stn 62, 1 (NNM 22458).

Karubar: stn 28, 5 (MNHN). — Stn 39, 13 (USNM 97176). — Stn 40, 4 (POLIPI). — Stn 56. 1 (POL1P1). — Stn 59,

1 (MNHN). — Stn 71, 6 (MNHN). — Stn 76. 1 (MNHN).

Type Locality. — Bosyu (= Awa), Japan (depth not given).

Diagnosis. — Corallum discoidal, having a convex base with a basal angle of 130°- 170°. Largest known

specimen (Karubar stn 40) 26.8 mm in calicular diameter and 9.4 mm in height. Costae prominently ridged and

coarsely dentate. Corallum white. Septa hexamerally arranged in 4 complete cycles: Si>S2>S4>S3. One palus or

paliform lobe occurs on each septum (P1-4). Fossa shallow; columella papillose.

Remarks. — Deltocyathus vaughani is more fully described and illustrated by Cairns (1994). These are

the first records other than from Japan. It is distinguished from other species in the region by having

coarsely dentate costae, a patellate corallum, and costoseptae that project an equal distance beyond the calicular

edge (see key).

DISTRIBUTION. — Philippines: Lubang Island; Bohol Strait; 247-807 m. Indonesia : Banda Sea (Kai Islands);

Arafura Sea (south of Tanimbar Islands); 290-549 m. Elsewhere : Japan (Honshu and Kyushu); 88-1097 m.

Deltocyathus philippinensis sp. nov.

Figs 15 d-e

Deltocyathus italicus - ALCOCK, 1902c: 19 (in part: "Siboga" stn 95, 100). [Not Turbinolia italica Michelotti, 1838],

Material EXAMINED/TYPES. — Philippines. "Siboga": stn 95, 1 paratype (ZMA Coel. 5441). — Stn 100,

2 paratypes (ZMA Coel. 5443).

"Albatross": stn 5505, 4 paratypes (USNM 97179). — Stn 5506, holotype (USNM 97178). — Stn 5538, 1 paratype

(USNM 97180).

South China Sea. "Penguin": stn 32, Macclesfield Bank (ca. 16°N, 114°E), 342 m, 1 (BMNH 1892.10.17.123).

Type Locality. — "Albatross" stn 5506: 8°40'00"N, 124031'45"E (Macajalar Bay, Bohol Sea, Mindanao),

479 m.

Etymology. — Named for the region from which it was first collected.

DESCRIPTION. — Corallum bowl-shaped, the largest specimen (holotype) 15.0 mm in calicular diameter and

6.0 mm in height. Centre of base (up to 5-7 mm) usually eroded and discoloured to a chalky white. Beyond central

worn region radiate 24 ridged costae (C1-3), which become prominent near calicular edge. C1-3 covered with very

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

123

fine spines, both laterally and apically. C4 ridges present only near calicular edge, also bearing fine, elongate

spines. Intercostal spaces broad and smooth. Calicular margin adjacent to every CS1-3 produced into a short,

triangular apex, resulting in a moderately serrate calicular margin. Upper, outer edges of all septa light brown.

Septa hexamerally arranged in 4 complete cycles. Si extend up to 1.6 mm beyond calicular edge and reach

1/2 distance to columella, where each is bordered by a lamellar paliform lobe about 1.5 mm wide. Si are the only

independent septa. S2 as exsert as Si, about 4/5 width of an SI, bearing a palus of equal size, the P2 rising

slightly higher in the fossa and recessed slightly more from the columella than Pi. S3 as exsert as S1-2 but only

about 2/3 width of an S2. Each S3 bears a highly spinose palus (P3) 1.5- 1.7 mm in width, each pair of S3 within

a system fusing to its common P2 through several robust processes, the uppermost process at the level of

S2-P2 notch. P3 rise higher in fossa and are more recessed from the columella than are P2. S4 extend about 1.1 mm

beyond calicular edge and are about 2/3 width of an S3 in upper fossa. Deeper in the fossa, far below S3-P3 notch

and near columella, S4 are rudimentary, each pair of S4 joining its common S3 through 1-3 slender processes.

S4 do not bear paliform lobes. Fossa shallow to moderate in depth. Columella papillose, consisting of 15-20

fused granular elements, each about 0.25 mm in diameter.

Remarks. — The rudimentary nature of the S4 and the tenuous connection to their adjacent S3 distinguish this

species from other western Pacific Deltocyathus. D. philippinensis resembles D. eccentricus Cairns, 1979 (amphi-

Atlantic, 183-1000 m) in its S4 morphology, but differs in having only 48 septa (D. eccentricus often has pairs of

S5), an eroded central basal region, and ridged costae.

DISTRIBUTION. — Philippines : Bohol Sea (south of Siquijor and Macajalar Bay); Sulu Sea (Sulu Archipelago);

402-522 m. Elsewhere : South China Sea (Macclesfield Bank); 342 m.

Deltocyathus Stella sp. nov.

Figs 15 f-h

MATERIAL EXAMINED/TYPES. — Philippines. MUSORSTOM 2: stn 33, 1 paratype (MNHN).

Indonesia. Snellius 2: stn 4.034, 1 paratype (NNM 22777).

Karubar: stn 1, 35: holotype and 10 paratypes (MNHN), 10 paratypes (POLIPI), and 14 paratypes (USNM 97181).

— Stn 2, 9 paratypes (USNM 97182). — Stn 18, 15 paratypes (MNHN). — Stn 32, 1 paratype (POLIPI).

Type Locality. — Karubar stn 35: 5°46'45"S, 132°1 1T0"E (Banda Sea: Kai Islands), 156-305 m.

ETYMOLOGY. — The species name (Latin Stella, star) refers to the general shape of the corallum. It is treated

as a noun in apposition.

DESCRIPTION. — Corallum shaped as a shallow bowl, with a slightly convex base. Largest specimen examined

(Karubar stn 1) 12.3 mm in calicular diameter; holotype 10.6 mm in diameter and 3.9 mm in height. Centre of

base displays a flat to slightly concave attachment scar 1.3- 1.5 mm in diameter. Costae well defined and rounded,

separated by shallow, narrow intercostal striae. Costae covered with small, rounded granules. Most coralla

uniformly white, but several small fresh specimens from KARUBAR stn 18 are reddish-brown in colour.

Septa hexamerally arranged in 4 cycles, the 4th cycle complete at a GCD of about 6 mm; no specimens

examined have more than 48 septa. Si independent, extending about 0.5 mm beyond calicular edge, and reaching

about 1/2 distance to columella. Central most part of each Si bears a small Pi about 0.8 mm wide. S2 similar to

Si but slightly narrower, having pali (P2) that rise higher in the fossa and are more recessed from columella than

Pi. S3 highly exsert (1.5 mm beyond calicular edge) and quite thick (up to 0.45 mm), but only about 2/3 width ot

S2. P3 about 1.2 mm wide, bearing large, coarse granules up to 0.12 mm tall and fusing to their common P2 near

the columella. S4 about 0.75 mm exsert, each pair fusing to its common S3 in a short triangular lancet, producing

a highly serrate calicular margin. Inner edges of S4 fuse to adjacent P3 through 2 or 3 narrow processes below

S3-P3 notch. Inner edges of Si -3 slightly sinuous; edges of S4 dentate to laciniate. Fossa shallow. Columella

consists of 15-30 small (0.20 mm diameter) papillae, all fused at base level.

Source : MNHN , Paris

124

S. D. CAIRNS & H. ZIBROWIUS

Remarks. — Deltocyathus Stella is distinctive in having quite exsert and thickened S3. The 12 S3 are not

spinose as in D. ornatus Gardiner, 1899 or lobate as in D. andamanicus , but simply prominent and thickened.

Another spinose Deltocyathus, D. heteroclitus Wells, 1984 (Pleistocene of Vanuatu), differs in having spinose

(not thickened) CS3, an irregular number of costal spines, and an incomplete 4th cycle.

DISTRIBUTION. — Philippines : Verde Island Passage; 130-137 m. Indonesia ; Banda Sea (Kai Islands); Timor

Sea (southwestern Timor); 206-280 m.

Deltocyathus andamanicus Alcock, 1898

Fig. 15 c

Deltocyathus andamanicus Alcock, 1898: 16-17, pi. 1, figs 5, 5a. — Vaughan, 1907: 71-72, pi. 6, figs 4, 4a. —

Gardiner & Waugh, 1938: 196. — Pillai & Scheer, 1976: 16. — Cairns & Keller, 1993: 244-245, pi. 5, fig. F.

Deltocyathus sp. cf. D. andamanicus - Cairns, 1984: pi. 3, figs A-B.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5273, 1 (USNM 97183). — Stn 5403, 11 (USNM

82156). — Stn 5412, 3 (USNM 97184). — Stn 5417, 6 (USNM 97185).

MUSORSTOM 2: stn 63, 17: 10 (MNHN), 7 (USNM 97187).

Musorstom 3: stn 88, 1 (MNHN). — Stn 92, 16 (USNM 97188). — Stn 126, 6 (MNHN). — Stn 130, 16 (USNM

97189). — Stn 138, 2 (MNHN). — Stn 139, 32 (MNHN).

Indonesia. DEKi: stn 59, 2 (NNM 22460).

Snellius 2: stn 4.173, 2 (NNM 22461).

TYPE Locality. — "Investigator" stn 236a: Andaman Sea, 315-555 m.

Description. — Corallum shaped as a shallow bowl, with a slightly convex base. Largest known specimen

(" Albatross " stn 5417) 21.1 mm in calicular diameter and 5.5 mm in height. Centre of base often bears a circular

scar 1.1 -1.5 mm in diameter, which is usually surrounded by a larger irregularly-shaped region up to 10 mm in

diameter characterised by reduced costal development, increased erosion, and often a lighter colouration. Costae

consist of thin, finely dentate ridges. Corallum usually uniformly light brown, exceptionally white or even light

yellow, the latter characteristic of specimens dead when collected.

Septa hexamerally arranged in 5 cycles, but the last never complete, most coralla having 4-16 pairs of S5,

which results in a total of 56-80 septa. Si extend about 1.8 mm beyond calicular edge and have slightly sinuous

inner edges, each Si internally bordered by a lamellar P| 1.7- 1.9 mm in width. Si, not being joined to any other

septa, are the only independent septa. S2 similar to Si but less wide, their P2 rising higher in the fossa and being

more recessed from the columella than Pi. S3 highly exsert, extending up to 3.8 mm beyond the calicular edge. In

large, well-preserved coralla, the lower outer edge of the S3 is hyperextended as a small (about 1 mm) lobe, this

lobe being compressed in the septal plane. Each S3 bordered by a large palus 1 .5-2.4 mm wide that rises higher in

the fossa and is more recessed from the columella than P2. Inner edges of P3 fused to their common P2. S4 small

and low, extending only about 1.5 mm beyond calicular edge, each pair fused to their common S3 to form

rectangular lancets, resulting in a highly serrate calicular margin. Inner edges of S4 fuse to their adjacent P3

through a series of 3-5 slender processes. In half-systems having pairs of S5, S5 are equivalent to previously

described S4, and S4 are same size as previously described S3. Upper, outer edges of CS1-3 entire, whereas those of

the S4-5 are laciniate. Fossa shallow; columella composed of 10-20 small (about 0.3 mm diameter) granular pillars

that are fused basally.

Remarks. — Although the holotype was not directly examined (deposited at the Indian Museum and not

available for loan), photographs of it were obtained and, along with ALCOCK's (1898) figure and description, the

holotype of 1 8 mm GCD and 62 septa appears consistent with the specimens identified above.

Distribution. — Philippines: Lubang Island; Sibuyan and Visayan Seas; Bohol Strait; Sulu Sea (Semirara

Islands); 187-333 m. Indonesia: Banda Sea (Kai Islands); Flores Sea (Selayar Island, Sulawesi); 340-385 m.

Elsewhere: southwestern Indian Ocean; Andaman Sea; Hawaiian Islands; 238-397 m.

AZOOXANTHELLATE SCLERACT1N1A

125

Deltocyathus suluensis Alcock, 1902

Fig. 16 d

Deltocyathus italicus - Alcock, 1902c: 19 (in part: "Siboga" stn 251). [Not Turbinolia italica Michelotti, 1838].

Deltocyathus magnificus var. suluensis Alcock, 1902c: 20-21. — FAUSTINO, 1927: lb-11.

Deltocyathus formosus Cairns, 1995: 73-74, pi. 19, figs f-g (new synonym).

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn 93, 11 (MNHN).

Indonesia. "Siboga": stn 251, 1 (ZMA Coel. 1109).

Deki: stn 44, I (NNM 22462). — Stn 51, 4 (NNM 22463). — Stn 52, 5 (NNM 22464).

Stn 58, 3 (NNM 22466). — Stn 62, 4 (NNM 22467).

MORTENSEN'S Java-S.A. Expedition: stn 15, 1 (ZMUC).

— Stn 57, 1 (NNM 22465). —

"Galathea": stn 500, 6 (ZMUC).

Karubar: stn 2, 80 (USNM 97191). - Stn 3, 3 (USNM 97192). - Stn 7, 3 (MNHN). - Stn 18, 2 (MNHN). -

Stn 31, 5 (USNM 97193). — Stn 35, 4 (POL1PI). — Stn 36, 2 (POLIPI). — Stn 67, 4 (USNM 97196). — Stn 77,

1 (MNHN).

Type Locality. — "Siboga" stns 95 and 100: Sulu Archipelago, 450-522 m.

Diagnosis. — Corallum with a flat to slightly convex base and a small central basal scar; peripherally the

theca is thickened and abruptly upturned, resulting in a marked thecal edge. Largest known specimen (Karubar

stn 77) 21.8 mm in diameter. Costae low, wide rounded ridges covered with granules. Well-preserved specimens

show concentric bands of light brown pigmentation on the base and a light brown pigmentation on lower, outer

septal edges. Septa hexamerally arranged in 5 cycles, the 5th cycle complete (a total of 96 septa) at a GCD of

about 18 mm; however, pairs of S5 begin to appear in coralla as small as 4 mm GCD. In fact, S5 are usually more

visible in small, fiat-based coralla than in larger coralla where they are often rudimentary or even absent on the

peripheral, upturned calicular edge, even though their corresponding C5 are well developed. S1-4 extend equally

beyond calicular edge (1.4- 1.6 mm), the S5 being only 1/2 as exsert. Calicular margin serrate but not lanceted.

S 1-4 and Pi -4 arranged in typical deltocyathid fashion, but, as mentioned before, S5 rudimentary to absent,

especially near calicular edge of large coralla. S5 join their adjacent S4 very low in fossa near columella by a series

of 2-5 thin processes. Columella papillose.

Remarks. — Deltocyathus suluensis differs from D. magnificus Moseley, 1876, in having granular costae; a

central basal scar or eroded region; less exsert and less developed S5; and a smaller corallum. It differs Irom

D. rotulus (Alcock, 1898), another species with 5 cycles of septa, in having granular, convex (not serrate ridged)

costae; a serrate (not lanceted) calicular margin; smaller S5; and a papillose columella; and is usually lound in

shallower water (142-565 m vs 210-1986 m for D. rotulus).

DISTRIBUTION. — Philippines : Lubang Island; Sulu Sea (Sulu Archipelago); 450-540 m. Indonesia: Banda Sea

(Kai Islands); Arafura Sea (southeast of Tanimbar Islands); Bali Sea; 204-390 m. Elsewhere: southern Norfolk

Ridge; Kermadec Islands; 142-565 m.

Deltocyathus rotulus (Alcock, 1 898)

Figs 16 a-c

Trochocyathus rotulus Alcock, 1898: 16, pi. 2, figs 1 , la.

Deltocyathus fragilis Alcock, 1902a: 99-100; 1902c: 21, pi. 2, figs 15, 15a.

Deltocyathus rotulus - Cairns & Keller, 1992: 245, pi. 5, fig. I (synonymy). — CAIRNS, 1994: 55-56, pi. 24, figs j-k.

Material EXAMINED. — Philippines. "Albatross": stn 5423, 3 (USNM). — Stn 5425, 2 (USNM 97200).

Stn 5429, 2 (USNM 97201). — Stn 5438, 1 (USNM 97202). — Stn 5439, 1 (USNM 97203). — Stn 5445, 1 1 (USNM

97204). — Stn 5447, 1 (USNM 97205). — Stn 5527, 2 (USNM 97206).

Musorstom 1 : stn 54, I (MNHN).

Musorstom 3: stn 94, 1 (MNHN).

Source : MNHN , Paris

126

S. D. CAIRNS & H. ZIBROWIUS

Indonesia. "Albatross": sin 5582, 6 (USNM 92727). — Stn 5585. 1 (USNM 97207). — Sin 5586, 2 (USNM

97208). — Stn 5591, 1 (USNM 97209). — Stn 5601, 1 (USNM). — Stn 5648, 1 (USNM 97210). — Stn 5668, I (USNM

9721 1).

Deki: stn 6, 2 (NNM 22437). — Stn 58, 4 (NNM 22439).

"Galathea": stn 476, 1 (ZMUC).

Corindon 2: stn 241, 2 (MNHN). — Stn 286, 2 (POLIP1).

Snellius 2: stn 4.130. 1 (NNM 22440).

Karubar: stn 2. 1 (USNM 97213). — Stn 21, 6: 5 (MNHN), 1 (USNM 97214). — Stn 56, 3: 1 (MNHN), 2 (USNM

97215). — Stn 87, 7: 6 (MNHN), 6 (USNM 97216). — Stn 89, 9 (POLIPI). — Stn 91, 18 (POLIPI).

Type Locality. — North Maidive Atoll, 1408-1756 m.

Diagnosis. — Corallum discoidal, with a flat to slightly bowl-shaped base. Largest known specimen

("Albatross" stn 5582) 36 mm in calicular diameter. Centre of base always eroded and without costae; otherwise

Ci -5 thin, finely serrate ridges. Septa hexamerally arranged in 5 cycles (Si-2>S3>S4>S5), the 5th cycle complete at

a GCD of 30-32 mm. Each Si-2 and the adjacent pair S5 project as short rectangular lancets, producing a highly

serrate calicular margin. Pi -3 small, sometimes even absent (Fig. 16 b); however, P4 prominent (up to 4 mm

wide), forming a distinctive crown of 24 lobes. S5 rudimentary, each pair of S5 fused to their common S4 by

5-8 slender processes beginning near the S4-P4 notch and continuing to the columella. Fossa shallow, containing

a columella in the shape of a circular, undercut platform that supports numerous irregularly-shaped papillae.

Remarks. — This species is more fully described and illustrated by Cairns (1994). It is one of the few

species in the genus to attain 5 full cycles of septa and is unique in having such a well-developed P4 crown.

DISTRIBUTION. — Philippines: Dasol, Luzon; Lubang Island; Lagonoy Gulf; north of Samar; Bohol Strait;

Sulu Sea (Cagayan Islands and Honda Bay, Palawan); 543-1719 m. Indonesia: Makassar Strait; Molucca Sea;

Banda Sea (Kai Islands and Gulf of Bone, Sulawesi); Timor Sea (south of Tanimbar Islands); Flores Sea

(Sumbawa); Bali Strait; 210-1710 m. Elsewhere: Malaysia (Celebes Sea off Sabah); Indian Ocean from Durban to

Maidive Islands; Japan (Honshu); 510-1986 m.

Deltocyathus magnificus Moseley, 1876

Deltocyathus magnificus Moseley, 1876: 552-553; 1881: 147-148, pi. 4, fig. 10, pi. 13, figs 1-2. — ALCOCK, 1902c:

20. — Faustino, 1927: 76. — Cairns & Parker, 1992: 27-28, pi. 7, figs j-1, pi. 8, fig. a. — Cairns, 1994: 56,

pi. 24, figs d-e, g-h (synonymy).

Material EXAMINED. — Philippines. "Albatross": stn 5135, 1 (USNM 97219). — Stn 5198, 4 (USNM 97220).

— Stn 5314, 3 (USNM 97221). — Stn 5374, 2 (USNM 97222). — Stn 5444, 1 (USNM 62712). — Stn 5506, 1 (USNM

97223). — Stn 5508, 1 (USNM 97224). — Stn 5527, 1 (USNM 97225). — Stn 5536, 1 (USNM 97226).

"Galathea": stn 436, 2 (ZMUC).

"Hakuho Maru": stn KH72-1-20, 1 (USNM 97235).

Musorstom 2: stn 83, 2 (MNHN).

Musorstom 3: stn 120, 3 (USNM 97228).

Indonesia. "Albatross": stn 5567, 1 (USNM 97227).

"Hakuho Maru": stn KH72-1-28, 2: 1 (USNM 97236), 1 (ORI).

Deki: stn 3, 2 (NNM 22450). — Stn 6, 1 (NNM 22451). — Stn 12, 2 (NNM 22452). — Stn 41, 28 (NNM 22453). —

Stn 44, 1 (NNM 22454). — Stn 46, 2 (NNM 22455). — Stn 51, 1 (NNM 22456).

Karubar: stn 3, 5 (POLIPI). — Stn 7, 3 (MNHN). — Stn 31, 1 (MNHN). — Stn 59, 24 (USNM 97231). — Stn 69,

6 (MNHN). — Stn 70, 1 (USNM 97233). — Stn 71, 1 (POLIPI). — Stn 76, 1 (POLIPI).

Type Locality. — "Challenger" stn 192: 5°49'S, 132°14'E (Kai Islands, Banda Sea), 236 m.

DIAGNOSIS. — Corallum discoidal, with a flat to slightly concave base covered with straight, thin, finely

dentate costae. Largest known specimen (Karubar stn 69) a damaged corallum 47 mm in calicular diameter and

9 mm in height. All costoseptae project an equal distance (about 1.5 mm) beyond calicular margin. Septa

hexamerally arranged in 5 full cycles (Si>S2>S3>S4>Ss), the 5th cycle usually complete at a GCD of about

Source :

AZOOXANTHELLATE SCLERACTINIA

127

8 mm. Pairs of S5 join their common S4 by an elongate, porous lamella. All septa bear prominent pali or

paliform lobes, even S5 of larger coralla. Columella an elongate fusion of numerous papillae that rises as high as

the septa and thus there is no fossa.

REMARKS. — Deltocyathus magnificus, the largest species in the genus, is easily distinguished from

D. rotulus (Alcock, 1898) by its flat base, equally exsert septa, differently constructed columella, and smaller P4.

DISTRIBUTION. — Philippines : Verde Island Passage; Bohol Sea; Tanon Strait; north of Samar; Comotes Sea;

Sulu Sea (Semirara Islands and Sulu Archipelago); 220-717 m. Indonesia : Ceram Sea; Banda Sea (Kai Islands);

Arafura Sea (southeast of Tanimbar Islands); Timor Sea (south of Leti Islands); 118-477 m. Elsewhere: Malaysia

(Celebes Sea off Sabah); South China Sea (north of Pratas Island); southern Australia; Japan (Honshu, Kyushu,

and northern Ryukyu Islands); 88-1500 m.

Genus CONOTROCHUS Seguenza, 1864

Conotrochus funicolumna (Alcock, 1902)

Ceralotrochus (Conotrochus) funicolumna Alcock, 1902a: 93; 1902c: 11-12, pi. 1, figs 6, 6a. Faustino, 1927: 66,

pi. 9, figs 7-8. — Zou, 1988: 77, pi. 5, figs 1, la.

Conotrochus funicolumna - CAIRNS, 1984: 14, pi. 2, figs 1-J; 1994: 58-59, pi. 24, fig. i, pi. 25, figs g-1 (synonymy).

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5551. 1 (USNM 97238). — Stn 5567, 19 (USNM

97240).

"Hakuho Maru": stn KH72-1-20, 6 (USNM 97242).

Indonesia. "Albatross": stn 5586, 19 (USNM 97241).

Deki: stn 44, 1 (NNM 22551). — Stn 52, 4 (NNM 22552).

"Galathea": stn 500, 8 (ZMUC).

"Hakuho Maru": stn KH72-1-28, 2 (USNM 97243).

Karubar: stn 59, 1 (MNHN).

South China Sea. "Albatross": stn 5312, 1 (USNM 97237). — Stn 5318, 1 (USNM 97238).

Type Locality. — "Siboga" stns 95 and 100: Sulu Archipelago, 450-522 m.

DIAGNOSIS. — Corallum elongate-conical to trochoid, and usually free (often having a small-diameter, blunt

pedicel), but some coralla attached by a slender pedicel. Largest Indonesian specimen (Karubar stn 59) 14 mm in

calicular diameter and 24.0 mm in height. Theca thick, covered with coarse granules and a thin epitheca, the theca

projecting 0.5-0. 7 mm above outer, upper septal edges as a continuous rim. Septa hexamerally arranged in

4 complete cycles: Si>S2>S3>S4, the Si only slightly wider than S2. Fossa shallow, containing a prominent

columella composed of several short, twisted lamellae that are swirled in the typical clockwise direction.

Remarks. — Conotrochus funicolumna is more fully described and illustrated by Cairns (1994). It is

compared to C. brunneus (Moseley, 1881) in the following account.

Distribution. — Philippines: Sulu Sea (Sulu Archipelago); 353-522 m. Indonesia: Banda Sea (Kai Islands);

Arafura Sea (east of Tanimbar Islands); Timor Sea (south of Leti Islands); 268-616 m. Elsewhere: Malaysia

(Celebes Sea off Sabah); South China Sea (Pratas Island); Victoria, Australia; Japan (Honshu and Ryukyu Islands);

Hawaiian Islands; 88-600 m.

Conotrochus brunneus (Moseley, 1881)

Fig. 16 e

Pleurocyathus brunneus Moseley, 1881: 159-160, pi. 2, figs la-c.

Phloeocyathus hospes Alcock, 1902b: 116-117.

Ceratotrochus (Phloeocyathus) hospes - ALCOCK, 1902c: 12, pi. 2, figs 8, 8a.

Source : MNHN Paris

128

S. D. CAIRNS & H. ZIBROWIUS

Conotrochus brunneus - Cairns & Parker, 1992: 22. — Cairns & Keller, 1993: 246, pi. 4, figs F-G (synonymy). —

Cairns, 1995: 74-75, pi. 20, figs a-b.

Ceratotrochus (Conotrochus) brunneus - ZOU, 1988: 76-77, pi. 5, figs 2, 2a.

MATERIAL EXAMINED. — Philippines. " Albatross stn 5171, 1 (USNM M236517). — Stn 5217, 2 (USNM

97245). — Stn 5277, 1 (USNM 97246).

"Hakuho Maru"\ stn KH72-1-20, 1 (USNM 97260).

Musorstom 2: stn 32, 1 (USNM 97248). — Stn 33. 69 (MNHN). — Stn 63, 5 (USNM 97250).

MUSORSTOM 3: stn 92. 22: 21 (USNM 97251), 1 (BMNH 1992.8.11.4). — Stn 102, 2 (MNHN). — Stn 109, 1 (USNM

97252). — Stn 1 17, 1 (MNHN). — Stn 126, 2 (MNHN). — Stn 139, 2 (MNHN).

Indonesia. Deki: stn 3, 6 (NNM 22556). — Stn 4, 1 (ZMUC). — Stn 5, 1 (NNM). — Stn 6, 12 (NNM 22557). —

Sin 42, 1 (NNM 22559). — Stn 46, 1 (NNM 22560). — Stn 48. 18 (NNM 22561). — Stn 58, 3 (NNM 22628). — Stn 63,

8 (NNM 22563).

"Hakuho Maru ": stn KH85-1-A2, 1 (USNM 97247).

Karubar: stn 1, 1 (USNM 97253). — Stn 2, 31 (USNM 97254). — Stn 3, 7 (POLIPI). — Stn 7, 11 (MNHN). —

Stn 15, 6 (MNHN). — Stn 18, 1 (MNHN). — Stn 31, 1 (POLIPI). — Stn 32, 1 (POLIPI). — Stn 36, 2 (POLIPI). — Stn 69,

1 (USNM 97257). — Stn 71, 1 (MNHN). — Stn 77, 5 (MNHN).

South China Sea. "Albatross"', stn 5301, 1 (USNM 97247).

"Hakuho Maru": stn KH72-1-52, 10: 7 (USNM 97161), 3 (ORI).

Type Locality. — "Challenger" stn 194: 4°34'S, 129°57'30"E (Banda Island, Banda Sea), 366 m (see

Cairns, 1995).

DIAGNOSIS. — Corallum elongate-conical, originally attached by a slender pedicel that is usually augmented by

a broad lateral thecal adhesion near the pedicel, which together form a broad, solid attachment. Largest known spec¬

imen (KARUBAR stn 71) 12.1 mm in calicular diameter and 16.0 mm in height, but most coralla are much smaller,

having a GCD of 6-8 mm. Theca coarsely granular and glisteny, the theca of well-preserved coralla commonly

pigmented a reddish-brown arranged in longitudinal stripes or circumferential bands. Some specimens, however,

have a smooth, usually longitudinally pigmented, theca. Theca quite thick, reinforced internally with deposits of

stereome; upper thecal edge forms a smooth circular rim about 0.4 mm around the upper, outer edges of the septa.

Septa hexamerally arranged in 4 cycles (S i>S2>S3>S4), the 4th cycle rarely complete, most coralla having 36-44

septa. Fossa deep. Columella consists of short, twisted lamellae swirled in the typical clockwise direction.

Remarks. — Conotrochus brunneus is similar to C. funicolumna (Alcok, 1902) but differs in having a

smaller corallum, less septa (usually 36-44 vs 48 for C. funicolumna ), a secondarily attached pedicel, a pigmented

corallum, and internal stereome. Also, in general, C. brunneus is found in deeper water than C. funicolumna.

DISTRIBUTION. — Philippines: Lubang Island; Verde Island Passage; Sibuyan Sea; Mindoro Strait; Sulu Sea

(Sulu Archipelago); 97-460 m. Indonesia: Halmahera Sea; Banda Sea (Banda and Kai Islands); Arafura Sea (south

of Tanimbar Islands); Flores Sea (Sulawesi); 206-477 m, with 1 outlying doubtful record at 1089 m (Alcock,

1902c). Elsewhere: Indo-West Pacific from Madagascar to South China Sea (Vanguard Bank and Pratas Island),

including western Australia and ridges north of New Zealand; 237-1051 m.

Genus LOCHMAEOTROCHUS Alcock, 1902

Lochmaeotrochus oculeus Alcock, 1902

Figs 16 f-i

Lochmaeotrochus oculeus Alcock, 1902b: 117-118; 1902c: 13, pi. 2, figs 9, 9a.

Material EXAMINED. — Indonesia. "Siboga": stn 159, 5 coralla (syntypes, ZMA Coel. 814). — Stn 259,

2 clusters and 18 individual coralla (syntypes, ZMA Coel. 700).

"Albatross": stn 5586, 1 cluster consisting of 3 coralla (USNM 97263). — Stn 5592, 2 clusters consisting of

6 coralla (USNM 97264).

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

129

Deki: stn 52, 1 cluster (NNM 22554). — Stn 56, 2 clusters (NNM 22553).

MORTENSEN'S Java-S.A. EXPEDITION: stn 15, 3 clusters consisting of 22 coralla (ZMUC).

Karubar- stn 13, 15 clusters consisting of 89 coralla: 9 clusters (MNHN), 6 clusters (USNM 97265).

South China Sea. "Hakuho Maru stn KH73-2-44-2, 19 clusters and 12 coralla: 4 clusters (ORI), remainder

(USNM 97266).

TYPE Locality. — "Siboga" stns 159 and 259: (Kai Islands and Halmahera Sea), 411-487 m.

DESCRIPTION (emended). — Corallum conical to subcylindrical, having a circular calice up to 11 mm in

diameter and a corallum height up to 24 mm. Clusters of up to 30 coralla form pseudocolonies, each corallum

providing the substratum for the settlement of up to 6 additional coralla from its theca, resulting in a low, dense,

bushy agglomeration. The close proximity of multiple coralla often leads to fusion of their theca. Corallum

pedicel robust: PD:GCD up to 0.5. Theca covered with small, hollow granules and epithecal bands. Each corallum

internally reinforced with thick stereome. Base polycyclic. Transverse dissepiments sometimes present. Corallum

white. Theca extends up to 0.8 mm above upper, outer septal edges as a continuous rim, which is characteristic of

the genus.

Septa hexamerally arranged in 4 cycles, most coralla from Karubar stn 13 having 48 septa, although 36 septa

is also common in other specimens. Si 1.0- 1.2 mm exsert, having straight, vertical inner edges that border the

columella. S2 slightly less exsert and 3/4 to 4/5 width of an Si also having straight inner edges and occasionally

bordered by a narrow, lamellar to papillose paliform lobe. These P2 almost indistinguishable from columellar

elements and not always present. S3 slightly less exsert and 3/4 to 4/5 width of an S2, having slightly sinuous

inner edges, and sometimes bordered by a papillose or short lamellar paliform lobe, 2 of which sometimes unite

within a system forming a V-shaped structure before their common Si. In some coralla P3 are absent, rudimentary,

or irregular in development. S4 about 1/2 width of an S3 and have straight inner edges. Fossa of moderate depth,

containing a columella of granular papillae or short lamellae, but never swirled as in C. brunneus or

C. funicolumna.

Remarks. — ALCOCK (1902c: 13) stated that in this species "budding takes place near the calicular margin, is

fairly regular, and is a true gemmation." Based on re-examination of the syntypes and additional topotypic coralla,

it is clear that new corals occur over the entire theca in a very irregular manner (Fig. 16i) (not just the calicular

margin), and seem to prefer attachment to dead coralla or living corals not covered with an edge zone. This results

in corallum clusters of very irregular structure. Furthermore the bases of some coralla show a discrete margin at the

interface with the corallum on which it settled. Therefore, the clusters of coralla reported above are interpreted as

independent settlement of planulae, possibly clonemates, that form a pseudocolony much in ihe same way that

Desmophyllum dianthus (Esper, 1794), is known to form (CAIRNS, 1982).

ALCOCK (1902: 13) described L. oculeus as a colonial Conotrochus, very similar to C. hospes

(= C. brunneus). Even though L. oculeus is now interpreted as a solitary coral, it can be distinguished from

C. brunneus by having a tendency to form clusters or pseudocolonies, always having a white corallum, usually

having 48 septa, having a papillose columella (rarely sublamellar and never swirled), and often having either P2

and/or P3. This is believed to be the first report of this species and genus subsequent to their original descriptions.

Distribution. — Indonesia : Halmahera Sea; Banda Sea (Kai Islands); Bali Sea; 240-616 m. Elsewhere:

Malaysia (Celebes Sea off Sabah); South China Sea (southern Formosa Strait); 412-430 m.

Genus AULOCYATHUS Marenzeller, 1904

Aulocyathus recidivus (Dennant, 1906)

Ceratotrochus recidivus Dennant, 1906: 159-160, pi. 6, figs 1-2.

Aulocyathus recidivus - Cairns, 1982: 25-26, pi. 7, figs 7-9, pi. 8, fig. 1 (synonymy); 1994: 59-60, pi. 26. figs a-b;

1995: 75, pi. 20, figs c-f. — Cairns & Parker, 1992: 22-24, pi. 6, figs d-e, g-h (synonymy). — Cairns & Keller,

1993: 247, pi. 5, fig. C.

Source : MNHN. Paris

130

S. D. CAIRNS & H. ZIBROWIUS

MATERIAL EXAMINED. — Indonesia. "Albatross": stn 5585. I (USNM 97268). — Sin 5586. 1 (USNM 97269).

Type Locality. — Off Cape Jaffa and Neptune Island, South Australia, 165-190 m.

DIAGNOSIS. — Corallum elongate-conical, straight, and usually budded from fragment of the parent corallum.

The only well-preserved Celebes Sea specimen ("Albatross" stn 5585) is 9.5 mm in calicular diameter, 5.0 mm in

basal diameter, and 12.4 mm in height. Calicular margin circular and evenly serrate. Theca thin, with hollow,

conical spines. Corallum white. Septa hexamerally arranged in 4 to 5 cycles (S i>S2>S3>S4>Ss); however,

development of Sa and S5 quite irregular. Specimen from "Albatross" stn 5585 has 44 septa and includes systems

with 1 pair of S4, 2 pairs of S4, and even 1 system with 2 pairs of S4 and 1 pair of S5. Slender paliform lobes (P3)

usually present. Columella papillose.

Remarks. — Aulocyathus recidivus is more fully described and illustrated by Cairns (1982, 1994) and

Cairns & Parker (1992).

A small Aulocyathus specimen measuring only 2.5 mm in calicular diameter and 3.8 mm in height was

collected at MUSORSTOM 3 stn 87 (Lubang Island, 191-197 m, USNM 97653). It has a smooth, glistening theca

and 24 septa, and may represent a small specimen of A. juvenescens Marenzeller, 1904, heretofore known only

from Tanzania at 302-463 m (Cairns & Keller, 1993).

Distribution. — Malaysia: Celebes Sea off Sabah; 616-871 in. Elsewhere: Indo-West Pacific from

southwestern Indian Ocean to Japan, including South Australia and New Zealand region to Macquarie Island; 128-

1137 m.

Genus PARACONOTROCHUS C aims & Parker, 1992

Paraconotrochus zeidleri Cairns & Parker, 1992

Caryophyllia clavus var. epithecata - MOSELEY. 1881: 135. [Not Caryophyllia clavus var. epithecata Duncan, 1873].

Paraconotrochus zeidleri Caims & Parker, 1992: 21-22, pi. 5, fig. i, pi. 6, figs a-b.

MATERIAL EXAMINED. — Indonesia. "Albatross": stn 5589, 2 (USNM 97270). — Stn 5590, 1 (USNM 97271). —

Stn 5592, 1 (USNM 97272).

DEKi: stn 52, 8 (NNM 22751).

"Galathea": stn 500, 1 (ZMUC).

Karubar: stn 59, 1 (POLIP1). — Sin 69, 7: 6 (MNHN), 1 (USNM 97273). — Stn 77, 4: 3 (USNM 97274), 1 (MNHN).

Admiralty Islands. "Challenger": stn 219, 1 (BMNH).

Type Locality. — "Soela" stn 51: 41°15'S, 144°08’E (Tasmania), 520 m.

DIAGNOSIS. — Corallum turbinate, free or attached by a narrow pedicel. Calice elliptical: GCD:LCD = 1.15-

1.28; margin of calice finely serrate. Largest known specimen ("Albatross" stn 5592) 30.7 x 24.7 mm in calicular

diameter and 27.4 mm in height. Costae not well defined; white theca covered with low, coarse granules. Septa

hexamerally arranged in 5 cycles (Si-2>S3>S4>Ss), the lower, inner edges of S1-3 fusing to the columella deep

within fossa. In some specimens narrow paliform lobes (P3) are present, but in most specimens reported herein

they are absent. Columella very distinctive, composed of numerous short, swirled lamellae, the entire structure

well separated from inner septal edges.

Remarks. — This species is more fully described and illustrated by Cairns & Parker (1992).

Distribution. — Indonesia: Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar Islands); 35 1 -

558 m. Elsewhere: Malaysia (Celebes Sea off Sabah); New South Wales, western Tasmania, and Admiralty

Islands; 274-520 m.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

131

Genus DESMOPHYLLUM Ehrenberg, 1834

Desmophyllum dianthus (Esper, 1794)

Fig 17 g-h

Madrepora dianthus Esper, 1794: pi. 69, figs 1-3.

Desmophyllum dianthus - Cairns, 1994: 26-27, pi. 9. figs a-d (synonymy).

Desmophyllum cristagalli H. Milne Edwards & Haime. 1848a: 253. — ALCOCK, 1902c: 28. — Zibrowius, 1980: 1 17-

121, pi. 61, figs A-O. pi. 62, figs A-M (synonymy).

Not Desmophyllum sp. - ALCOCK, 1902c: 28 (= Madrepora oculata Linnaeus, 1758).

Material EXAMINED. — Indonesia. "Siboga": stn 259, 2 (ZMA Coel. 1242, see ALCOCK, 1902c).

"Captain Christiansen": 1°31'N, 124°47'E, 457 m, 12 March 1913, 6 (NNM 22800).

South China Sea. "Hakuho Maru": stn KH73-2-44-2, 9: 6 (USNM 97275), 3 (ORI).

Type Locality. — Sagami Bay (depth not given).

Remarks. — Desmophyllum dianthus is perhaps the most commonly collected deep-water coral and has a

virtually cosmopolitan distribution (see below), but, for unknown reasons, is rare in the Indonesian region. It is

known from only 2 localities: one reported by ALCOCK (1902c) and another reported herein. Many descriptions of

this species are available in the literature (e.g., ZlBROWIUS, 1974b. 1980; CAIRNS, 1979, 1982, 1994, 1995),

most under the name D. cristagalli.

DISTRIBUTION. — Indonesia: Celebes Sea (Manado, Sulawesi); Banda Sea (Kai Islands); 457-487 m.

Elsewhere: South China Sea (north of Pratas Islands); cosmopolitan, except for continental Antarctica and northern

boreal Pacific (see Cairns, 1994); 35-2460 m.

Genus DACTYLOTROCHUS Wells, 1954

Dactylotrochus cervicornis (Moseley, 1881)

Tridacophyllia cervicornis Moseley, 1881: 183-184, pi. 10, figs 2a-2d, 3a. — Bassett-Smith, 1890: 368.

Tridacophyllia primordialis Gardiner, 1899: 168, pi. 19, figs 7a-e.

Dactylotrochus cervicornis - Wells, 1954: 470-471, pi. 178, figs 1-3.

Material EXAMINED. — Indonesia. Karubar: stn 18, 4 (MNHN).

Type Locality. — Unknown.

Remarks. — This species is best described and illustrated by Wells (1954). The specimen he reported from

the Philippines ("Albatross" stn 5336) cannot be located at the USNM.

Distribution. — Philippines: north of Palawan; 84 m (Wells, 1954). Indonesia: Banda Sea (Kai Islands);

205-212 m. Elsewhere: South China Sea (Tizard Bank, Spratly Islands); Bikini, Marshall Islands; New Caledonia;

Loyalty Islands; 73-137 m, however yet unreported specimens from New Caledonia occur as deep as 400 m.

Genus ASTEROSMILIA Duncan, 1867

Asterosmilia marchadi (Chevalier, 1966)

Figs 17 a-b

Dasmosmilia marchadi Chevalier, 1966: 944-949, pi. 5, figs 3-4,

Source : MNHN, Paris

132

S. D. CAIRNS & H. ZIBROWIUS

Asterosmilia marchadi - Cairns, 1979: 140-142, pi. 26, figs 7, 9-10. - Zibrowius, 1980: 141-142, pi. 74. figs A-K

(synonymy). — Cairns & Keller, 1993: 249, pi. 6, figs A-B.

MATERIAL EXAMINED. — Philippines. " Albatross stn 5133, 10 (USNM 97278). — Stn 5152, 2 (USNM

97279). _ Sin 5156, 1 (USNM 97280). — Stn 5164, 55 (USNM 97282). — Stn 5178, 4 (USNM 97283). — Stn 5277,

1 (USNM 97284). — Stn 5357, 4 (USNM 97286).

MUSORSTOM 1: stn 32, 1 (USNM 97289). — Stn 57, 1 (USNM 97290). — Stn 73, 1 (USNM 97291).

Siphilexp: stn 78-T10, 1 (USNM 97287).

MUSORSTOM 2: stn 8, 1 (MNHN). — Stn 47, 1 (MNHN).

MUSORSTOM 3: stn 102, 2 (MNHN). - Stn 107, 3 (USNM 97293). - Stn 131, 7 (MNHN). - Stn 140, 10 (MNHN).

Indonesia. DEKI: stn 6, 1 (NNM 22537). — Stn 10, 2 (NNM 22538). — Stn 35, 2 (ZMUC), 3 (NNM 22539).

Mortensen'S Java-S.A. Expedition: stn 5, 42 (ZMUC). — Stn 6, 1 (ZMUC). — Stn 8, 6 (ZMUC). Stn 9,

2 (ZMUC).

Corindon 2: stn 216, 3 (MNHN). — Stn 251, 4 (MNHN).

— Stn 261, 4 (MNHN). — Stn 263, 1 (MNHN). —

Stn 266, 3 (USNM 97295).

Snellius 2: stn 4.235, 9 (NNM 22536).

South China Sea. "Albatross": stn 5311, 19 (USNM 97285).

Type Locality. — Senegal, 97-85 m.

DESCRIPTION. — Corallum ceratoid, usually curved about 45°. Approximately half of the coralla examined

originated by extratentacular budding from the edge zone of a parent corallum, their pedicels narrowing to a slender,

open base only 0.6-0.7 mm in diameter. Other coralla firmly attached to a substratum through a robust pedicel up

to 4.5 mm in diameter (PD:GCD = 0.18-0.30), these coralla assumed to have originated from planular settlement.

Largest known specimen (MUSORSTOM 3 stn 140) 14.9 x 18.0 mm in calicular diameter, 15.8 mm in height, and

4.5 mm in pedicel diameter; however, most coralla are less than 13 mm in GCD. All costae (Cm) consist of low,

rounded ridges, separated by equally wide, shallow furrows. Costae covered with tiny granules, 4 or 5 occurring

across the width of a costa. Theca quite thin. Many coralla display small scars of bud detachment on their theca,

some bearing as many as 25 shallow, circular (0.6-0. 7 mm in diameter) concavities. Although some coralla are

completely white, most coralla are reddish-brown, especially intense near calicular edge and fading to white at base.

Septa hexamerally arranged in 4 cycles, only larger coralla having pairs of S5, up to 66 septa. Si highly exsert

(about 2.9 mm), having straight, vertical inner edges that attain the columella. S2 slightly less exsert (about

2.5 mm), about 4/5 width of an Si. Both Si and S2 occasionally bear an irregularly-shaped paliform lobe that

mingles with the columella. S3 about 1.2 mm exsert and only 1/2 width of an S2- Each S3 bears a lamellar

paliform lobe about 1.2 mm wide, together forming a palar crown. S4 as exsert as S3 but only about 2/3

the width, with finely dentate inner edges, and attenuating in size low in fossa. If a pair of S5 flanks an S4, that

S4 also bears a paliform lobe, the P4 being larger than the P3 and occurring higher in the fossa and more

peripherally. Pairs of S4 flanking Si and S2 form low calicular lancets resulting in a serrate calicular margin.

Fossa of moderate depth, containing a crispate, twisted columella. Endothecal dissepiments often present.

Remarks. — Previous records of A. marchadi from the Atlantic and southwest Indian Ocean have been of

unattached specimens, presumably resulting from asexual budding. However, among the 28 lots reported above,

11 lots (mainly from MUSORSTOM 1-3 and Corindon 2) contained exclusively attached specimens; 11 lots

(mostly "Albatross" and ZMUC specimens) contained only free forms; and 6 lots (from all sources) contained

a mixture of both forms. Buds detach at a very early stage, probably because their attachment to the parent is so

small and tenuous. Examples of buds still attached to the parent corallum are uncommon, rarely exceeding 2 mm

in length; likewise, many already detached buds as small as 4 mm length were found in the study material.

We therefore conclude that the buds probably detach at a corallum length of 2-4 mm.

DISTRIBUTION. — Philippines: Lubang Island; Sibuyan and Visayan Seas; Sulu Sea (Panay; Zamboanga

Peninsula; Balabac Island, Palawan; Sulu Archipelago); 33-184 m. Indonesia: Makassar Strait; Banda Sea (Kai

Islands); Flores Sea (Selayar Island, Sulawesi); Bali Strait; 32-210 m. Elsewhere: tropical amphi-Atlantic;

southwestern Indian Ocean; Maidive Islands; South China Sea (Pratas Island); 32-229 m.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

133

Genus THALAMOPHYLLIA Duchassaing, 1870

Thalamophyllia tenuescens (Gardiner, 1899)

Figs 17 d-e

Desmophyllum tenuescens Gardiner, 1899: 161-162, pi. 19, figs la-b.

Thalamophyllia tenuescens - CAIRNS, 1995: 78, pi, 21, figs g-i.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5255, 2 corallites (USNM 97296).

Marigondon cave entrance, Mactan Island, Cebu, 22 m, coll. H. Schuhmacher, 28 May 1981, 1 colony (USNM

94142).

Musorstom 3: stn 131, 3 colonies (MNHN).

Indonesia. Deki: stn 54, 1 (NNM).

Corindon 2: stn 248, 1 colony (MNHN).

SNELLIUS 2: stn 4.226, 2 (NNM).

Karubar: stn 22, 1 colony (USNM 97298). — Stn 30, 1 (POLIPI).

Queensland. Steven's Reef, 12 m, 10 corallites (USNM 78584). — Reef D, Queensland, depth unknown. 3 corallites

(USNM 78585).

Type Locality. — Sandal Bay, Lifu, Loyalty Islands, 73 m.

DIAGNOSIS. — Corallum composed of a cluster of elongate-conical corallites, each originating from a common

basal coenosteum or the lower pedicel of a larger corallite. Largest known corallite (CORINDON 2 stn 248)

14.9 mm in GCD and 28.7 mm in height, but calices of most other corallites examined less than 5 mm in

diameter. Ci and often C2 highly ridged, producing a calice hexagonal in outline. Corallum white. Septa of most

corallites hexamerally arranged in 3 cycles (Si>S2»S3), only the larger corallites having a 4th cycle (48 septa).

Highest cycle septa (S3 or S4, depending on size) rudimentary, often absent from upper corallum. Fossa quite deep;

no columella.

REMARKS. — Previously reported specimens of T. tenuescens were small (GCD < 4.5 mm) and thus had only

3 cycles of septa. A colony from Corindon 2 stn 248, having a much larger calice (/.«., 14.9 mm GCD) and

4 cycles of septa, is included without hesitation because from its base bud two smaller corallites (GCD = 4.1 and

4.9 mm), each having only 24 septa and being similar to previously known specimens.

The colony from KARUBAR stn 22 has the C1-2 on the lower pedicel prominently ridged and extending outward

onto the common coenosteum. Exothecal dissepiments bridge adjacent Ci-2, forming chambers of tissue radiating

outward from the base of each corallite, each a potential bud.

This species is more fully described and illustrated by CAIRNS (1995).

Distribution. — Philippines: Sulu Sea (west of Panay); Bohol Strait; Davao Gulf; 22-183 m. Indonesia:

Makassar Strait: Banda Sea (Kai Islands); Flores Sea (Selayar Island); 85-288 m. Elsewhere: Queensland; Lord

Howe Seamount Chain; Kermadec Islands; Loyalty Islands; 8-315 m.

Genus RHIZOSMILIA Cairns, 1978

Rhizosmilia robusta Cairns, 1993

Rhizosmilia robusta Caims in CAIRNS & KELLER, 1993: 250-253, pi. 6, figs F-I.

Material EXAMINED. — Philippines. Musorstom 1: stn 65, 1 colony of 4 corallites (MNHN).

Type Locality. — "Anton Bruun" stn 373B: 26°00'S, 33°05'E (off southern Mozambique), 135 m.

Source : MNHN Paris

134

S. D. CAIRNS & H. ZIBROWIUS

Diagnosis — Phaceloid colonies composed of a few robust, trochoid corallites, the largest known corallite

29 8 x 36 7 mm in calicular diameter, 41.6 mm in height, and 18.1 mm in pedicel diameter. Base and lower

pedicel thickened with concentric rings of hollow chambers formed by layers of exothecal dissepiments that bridge

raised costae. Corallum white. Septa hexamerally arranged in 5 cycles (Si>S2>S3>S4>Ss), the largest corallite

also having 8 pairs of S& and even 1 pair of S7 in its end systems (/.<?., 1 16 septa). Small, lamellar paliform lobes

occur before S4, seemingly paired within each half system. Fossa deep; columella composed of granular rods that

are strongly fused into a massive structure.

REMARKS. — This species is more fully described and illustrated in its original description.

DISTRIBUTION. — Philippines : north of Lubang Island; 194-202 m. Elsewhere : southwestern Indian Ocean,

66-150 m.

Rhizosmilia sagamiensis (Eguchi, 1968)

Coenocyalhus sagamiensis Eguchi, 1968: C34. pi. CIO, figs 6-7.

Rhizosmilia sagamiensis - Cairns, 1994: 62-63. pi. 27, figs c-e.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5255, 1 (USNM 97300).

MUSORSTOM 3: stn 88. 1 (MNHN). - Sin 1 17, 4 (USNM 97301). - Stn 131, 3 (MNHN). - Stn 134, 1 (MNHN).

Indonesia. Corindon 2: stn 248, 10 (MNHN).

Type Locality. — Sagami Bay, 60-80 m.

DIAGNOSIS. — Corallum composed of a relatively small phaceloid clump of elongate-conical corallites, the

largest Philippine corallites (MUSORSTOM 3 stn 131) 8.8 x 10.1 mm in calicular diameter, 13.5 mm in height, and

4.5 mm in pedicel diameter. Corallite bases reinforced with concentric rings of hollow chambers (best seen in a

cross section of the pedicel), which is characteristic of the genus. Corallum primarily white; however, faces

of S1-2 usually bear reddish-brown crescent-shaped bands that parallel the curved septal edge. Septa hexamerally

arranged in 4 cycles (Si>S2>S3>S4), the Si being highly exsert. A crown of 12 P3 encircles the trabecular

columella.

REMARKS. — Rhizosmilia sagamiensis is more fully described and illustrated by Cairns (1995).

DISTRIBUTION. — Philippines: Lubang Island; Mindoro Strait; Sibuyan Sea (north of Panay); Sulu Sea (west

of Panay); Davao Gulf; 97-183 m. Indonesia: Makassar Strait; 170 m. Elsewhere: Japan (Sagami Bay to northern

Ryukyu Islands); 60-98 m.

Rhizosmilia elala sp. nov.

Figs 18 a-b

? Anomocora fecunda - EGUCHI, 1968: C42 (in part: pi. CIO, figs 1-5). [Not Coelosmilia fecunda Pourtales, 1871].

Material EXAMINED/TYPES. — Philippines. "Albatross": stn 5244, holotype (USNM 97304) and 15 paratype

colonies (USNM 97305).

Indonesia. Deki: stn 27, 3 corallites, paratypes (NNM 22497).

Karubar: stn 86, 1 paratype colony (MNHN).

Type Locality. — "Albatross" stn 5244: 6°52'N, 126°14'E (Pujada Bay, southeast coast of Mindanao),

313 m.

ETYMOLOGY. — The species name (Latin elatus , high) refers to the elongate growth form of this species.

DESCRIPTION. — Corallum consists of a phaceloid colony composed of elongate-ceratoid to cylindrical

corallites, the corallites often undergoing multiple rejuvenescence — one attaining the length of 12 cm, but only

Source : MNHN . Paris

AZOOXANTHELLATE SCLERACT1NIA

135

7.6 mm in distal calicular diameter. Main corallite of holotype colony 8.3 cm long and 11.8 mm in calicular

diameter, bearing 14 smaller corallites, most of which are assumed to be the result of independent settlement of

planulae. Corallites budded from edge zone of parent corallite as well as from common basal coenosteum. On

elongate corallites the edge zone covers only a small distal region of the corallum, whereas the remaining theca is

often encrusted with epifauna (serpulids, bivalves, etc.), including juvenile corallites of R. data. Thus, an elongate

corallite may bear numerous, irregularly spaced smaller corallites, the secondary corallites resulting either from true

budding or larval settlement. Well-preserved pedicels of this species indicate that the Ci-2 form prominent ridges

that radiate from the corallite base onto the adjacent coenosteum where exothecal dissepiments form over these

raised costae, characteristic of the genus Rhizosmilia. In the more distal parts, costae (Cm) are broad and convex,

covered with very fine granules, and separated from one another by narrow intercostal striae. Ci-2 usually slightly

more prominent than C3-4 and frequently pigmented a light reddish-brown, the remainder of the corallum being

white.

Septa hexamerally arranged in 4 complete cycles: Si>S2>S3>S4- Si highly exsert (up to 3.5 mm), having

straight, vertical inner edges that reach the columella. S2 slightly less exsert (up to 3.0 mm), about 2/3 width of

an Si. S3 about 1/2 width of an S2, and only about 2.3 mm exsert. Each S3 bordered by a thin lamellar paliform

lobe 1.3- 1.5 mm in width, the 12 P3 forming a distinct palar crown. S4 only about 1.8 mm exsert, about 1/2

width of an S3. Fossa of moderate depth, containing a crispate columella. Endothecal dissepiments common in

elongate coralla, usually visible only in longitudinal section.

REMARKS. — Five other species are known in Rhizosmilia: R. maculata (Pourtales, 1874); R. gigas (van der

Horst, 1931); R. sagamiensis (Eguchi, 1968); R. gerdae Cairns, 1978; and R. robusta Cairns, 1993. R. data is

distinguished in having elongate, subcylindrical corallites and pigmented Ci-2-

Rhizosmilia elata is similar to Anomocora in having elongate coralla with endothecal dissepiments and

irregular budding, but can be distinguished by its well-developed P3, the fact that its buds do not detach, and by

its characteristic basal structure of raised costae overlain with exothecal dissepiments.

Distribution. — Philippines: Pujada Bay, Mindanao; 313 m. Indonesia: Banda Sea (Kai Islands); Timor Sea

(south of Tanimbar Islands); 70-226 m. Elsewhere: ? Sagami Bay (EGUCHI, 1968); 100-380 m.

Genus PHYLLANGIA H. Milne Edwards & Haime, 1848

Phyllangia papuensis Studer, 1878

Figs 17 c, f, i

Phyllangia papuensis Studer, 1878: 642-643. — RIDLEY, 1884: 396-397, pi. 16, figs 5-10.

Blastomussa lawtoni Nemenzo, 1988: 216, fig. 1 (new synonym).

Phyllangia sp. - KROPP & MANNING, 1995: 535, fig. 3 a-d.

Material EXAMINED. — Philippines. Musorstom 2: stn 47, 9 colony fragments: 5 (MNHN), 4 (USNM 97307).

Ambon, coll. H. O. FORBES, 10 colony fragments (BMNH 1884.2.16.9-18; Ridley's, 1884 specimens).

Indonesia. Deki: stn 24, 1 colony (NNM 22496).

Solomon Islands. "Gazelle": stn 37, 10 colonies (syntypes), (ZMB 1849).

Indian Ocean. Madagascar. NW coast Nosy Be: 13°23.5'S, 47°58.5'E , 100 m. coll R. Plante, 29 October 1969,

1 large colony (MNHN).

Maldives Islands. Nilandu, 20 June 1900, depth unknown, 1 colony (NNM 22495).

Type Locality. — "Gazelle" stn 37: Solomon Islands, 88 m.

Description. — Colonies consist of many corallites that are budded from a thick dense basal coenosteum (up

to 1.5 mm) that envelopes a cylindrical substratum (i.e., gorgonian or antipatharian axis), such that corallites

point outward in all directions from the axis. Colony from Madagascar 17 cm long and 15 mm in diameter

(including the gorgonian axis), consisting of approximately 200 corallites. Philippine colony originally of similar

Source : MNHN , Paris

136

S. D. CAIRNS & H. ZIBROWIUS

size but encrusting a thinner antipatharian axis. Corallites elongate-conical to cylindrical and up to 7.9 mm in

diameter, although most are less than 5 mm in GCD. Corallites rise up to 7 mm above coenosteum, but most are

considerably lower. Coenosteum between corallites striate and white. Costae low and convex, bearing 3 or 4 small

rounded granules across the width of each costa. Upper theca and outer septal elements light reddish brown.

Septa hexamerally arranged in 3 to 4 cycles, a corallite less than 5 mm GCD usually having only 3 cycles

(Si»S2>S3), and larger corallites of 6.5 mm GCD having up to 44 septa (Si-2>S3>S4), but none having a full

4th cycle of 48 septa. Si highly exsert (up to 1.1 mm) and thick (0.2 mm), but not very wide, their inner edges

extending only about 1/2 distance to centre of calice. Lower, inner edges of Si sometimes bear a small (0.25 mm

wide), irregularly shaped paliform lobe. S2 much less exsert (0.6-0.7 mm) and only about 1/2 width of an Si:

S2 may also bear a larger paliform lobe up to 0.33 mm in width. S3 rudimentary, having laciniate inner edges. In

septal systems of corallites bearing pairs of S4, the S2 are accelerated to the same size of an Si, the S3 takes the

size of a normal S2 and has a paliform lobe, and the S4 are rudimentary, as previously described for the

unaccelerated S3. Fossa of moderate depth, containing a papillose columella, the rods often fused into a solid,

central structure.

REMARKS. — The 2 colonies from the Philippines and northwest Madagascar were the host to the same species

of hapalocarcinid crab (see K.ROPP & MANNING, 1995).

DISTRIBUTION. — Philippines: Sibuyan Sea; Balicasag Island, Bohol; 81-84 m. Indonesia : Banda Sea (Ambon

and Kai Islands); 100 m. Elsewhere: Bougainville Island (west of Solomon Islands); northwestern Madagascar;

Maidive Islands; 88-100 m.

Genus SYMPODANGIA nov.

Type SPECIES. — Sympodangia albatrossi, here designated.

ETYMOLOGY. — The genus name Sympodangia (Latin sympodium, structure resulting from alternate

branching + angia , a common coral generic suffix) refers to the growth mode of this coral. Gender: feminine.

DIAGNOSIS. — Colonial, small corallites closely packed in short sympodial branches. No individualized

columella; no endotheca. Axial edge of septa dentate.

REMARKS. — This genus is of uncertain affinities. The dentate septal edge is unlike typical rhizangiids.

Sympodial growth suggests affinities with Madrepora, but branches are stouter and less zigzag-shaped. Also, there

are more septa than in a Madrepora. Individual calices resemble those of Hoplangia, a monospecific genus, but

which has stronger Si.

Sympodangia albatrossi sp. nov.

Figs 18 c-g

Material EXAMINED. — Types: Philippines. "Albatross": stn 5398, holotype (USNM 97308); 400+ small

branch fragments, paratypes (USNM 97308), 28 branch fragments (MNHN).

Indonesia. "Albatross": stn 5586, 2 branch fragments, paratypes (USNM 97310).

Karubar: stn 18, 24 branch fragments, paratypes (USNM 97311).

Type Locality. — "Albatross" stn 5398: 11°35'N, 124°14'E (Biliran Island, Samar Sea, Philippines),

208 m.

Etymology. — This species is named for the RA ' Albatross.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

137

DESCRIPTION. — Small (rarely more than 4 cm in diameter), bushy colonies formed by closely spaced

sympodial budding, such that corallites from adjacent branches often fuse to one another. Adjacent corallites may

also be linked by slender (1 mm diameter), solid, coenosteal bridges, although this is far less common. Holotype a

branch fragment 35 mm long consisting of 20 corallites. Calices circular to slightly elliptical, ranging from 2.5 to

4.2 mm in diameter. No marked costae, the theca homogeneously covered with low, closely spaced, rounded

granules 90-120 pm in diameter, the larger-sized granules occurring only on coenosteum that encrusts a barnacle

valve substratum. Corallum light yellow-brown in colour.

Septa hexamerally arranged in 4 cycles, the 4th never complete. A full 3rd cycle of septa (a total of 24 septa)

present in calices of 2.4 mm GCD, and a maximum number of 36 septa (a pair of S4 in each system) is achieved

at GCD of about 3.4 mm. Si exsert (0.3-0.6 mm), having straight, vertical inner edges that bear 3-5 small

(85-100 pm diameter), cylindrical papillae on their lowermost edges adjacent to the columella. Accelerated S2

(those in systems having a pair of S4) about 0.2 mm exsert and 3/4 width of an Sr, lower, inner edges of S2

coarsely dentate, the teeth about 0.2 mm in width, beginning slightly higher in fossa than Si papillae. S3 slightly

less exsert than S3 but about equal in width, their inner edges bearing even larger teeth (up to 0.3 mm width) that

extend slightly higher in fossa than those of the S2. S4 less exsert and about 3/4 width of an S3, their inner edges

finely dentate for most of their length. Septal faces covered with tall (up to 90 pm), blunt granules. Fossa deep and

narrow. Columella papillose, but quite small, composed of 1-4 papillae of equal size to those on the lower edges

of the S 1 .

DISTRIBUTION. — Philippines : Samar Sea; 208 m. Indonesia : Banda Sea (Kai Islands); 212-616 m.

Elsewhere : Malaysia (Celebes Sea off Sabah).

Genus COLANGIA Pourtales, 1871

Colangia moseleyi (Faustino, 1927) comb. nov.

Cladocora conferta Moseley, 1881: 185, pi. 10, fig. 5, 5a (junior secondary homonym of Caryophyllia confer, a Dana,

1846: 380, pi. 27, fig. 6).

Cladocora moseleyi Faustino, 1927: 112-113, pi. 15, fig. 2-3 (nom. nov.).

Material EXAMINED. — Philippines. "Challenger", 1874, unnumbered stations off Zamboanga (Mindanao):

2 syntypes of Cladocora moseleyi: 30 fathoms (= 55 m), 1 colony, larger syntype (= lectotype) (BMNH, no register

number); 10 fathoms (=18 m), single corallite, smaller syntype (paralectotype), ? Polycyathus (BMNH no register

number).

Type Locality. — Zamboanga, Mindanao, Philippines, 55 m.

DESCRIPTION. — Lectotype collected alive, a compact colony of ca. 27 clustered corallites of various sizes,

some incomplete (damaged). Colony comprises some elongate parallel incomplete corallites, 10-20 mm high,

lower end broken. Endotheca can be seen inside the fractured corallites (parasmiliid character). In upper part colony

spreads laterally and short corallites overgrow the elongate ones, budding from a basal sheet as in Phyllangia.

Larger corallites up to 5 x 6 mm in calicular diameter. Septa little exsert (Si most exsert of all). Outer surface

quite smooth, locally with very attenuate costae covered by fine granules. Smaller calices with only 3 cycles ol

septa; larger calices with incomplete 4th cycle (for example 14 S4). Septa decreasing in width from Si to S4. Large

Si with straight vertical axial edge. One distinct large paliform lobe, with vertical axial edge, separated by notch

from corresponding septum, associated with S2 in smaller calices, or with S3 in larger calices (when S4 present in

same half system). Narrower septa of subsequent cycles with more sloping (less vertical) axial edge lrequently

bearing 2 or 3 narrow lobes, subhorizontally or obliquely upward directed. These narrow lobes very different trom

the main paliform lobes (but unlike septal dentation as in true Astrangia). Central area of fossa (up to ca. 1/3

calice diameter) occupied by columella composed of lobes similar to paliform lobes, but smaller.

Source : MNHN. Paris

138

S. D. CAIRNS & H. ZIBROWIUS

Remarks. — Moseley (1881) described material said to originate from 2 unnumbered shallow stations

(10 and 30 fathoms) off Zamboanga (Philippines), as a new species, Cladocora conferta. Cladocora moseleyi was

introduced by FAUSTINO (1927), who reproduced Moseley's description and illustration but with no new data as a

nomen novum in replacement for Moseley's Cladocora conferta, the latter then considered as a junior homonym

of Cladocora conferta (Dana, 1846). In fact, Dana’s (1846: 380-381, pi. 27, fig. 6) poorly described and illustrated

Caryophyllia conferta, of uncertain origin (possibly the West Indies), had been placed by later authors (starting

with H. Milne Edwards & Haime, 1849: 308) in the genus Cladocora, or even been assimilated (Vaughan,

1901: 298) as the West Indian Cladocora arbuscula (Lesueur, 1821), apparently without reexamining Dana's type.

Remaining of doubtful generic attribution, Dana's species may not be congeneric with Moseley's species.

Moseley's (1881) description of Cladocora conferta from the 30 fathoms station is based on a colony with many

clustered corallites (poorly illustrated). This syntype is here selected as the lectotype (BMNH, no register number).

It is not a Cladocora, but a Colangia, and closely resembles the West Indian Colangia immersa Pourtales, 1871,

type species of the genus. Colangia immersa should be restudied in detail. First described from Florida on the basis

of dead material from deep water, it more likely is a species of shelf depths. Moseley's second Philippine syntype

of Cladocora conferta, from the 10 fathoms station, (BMNH, no register number), is a different species, not a

Colangia: the elongate narrow corallite with only 3 cycles of septa and a lateral bud may be a Polycyathus. No

other specimen of Colangia has yet been recognized in the western Pacific. Elsewhere, the genus is known only

from the West Indies. This and the resemblance of Moseley's lectotype with the West Indian species is troubling.

Perhaps the origin is confused, the material not being from the Philippines. Confused origins of "Challenger"

material have been pointed out elsewhere.

DISTRIBUTION. — If indicated origin authentic (see Remarks), Colangia moseleyi is known only from

Mindanao (Philippines), 55 m (type locality).

Genus COENOSMILIA Pourtales, 1874

Coenosmilia arbuscula Pourtales, 1874

Figs 19 a-c

Coenosmilia arbuscula PourtaRs, 1874: 39-40, pi. 7, fig. 1. — Cairns, 1979: 130-131, pi. 24, figs 9-11 (synonymy).

Coenosmilia fecunda - Zibrowius, 1980: 131-133 (in part: pi. 68, figs A-F). [Not Coelosmilia fecunda Pourtalbs, 1871],

Coenosmilia sp. cf. C. arbuscula - Cairns, 1994: 61, pi. 27, figs a-b.

Not Parasmilia fecunda - Marenzeller, 1904a: 31 1. — Gardiner & Waugh. 1939: 229.

Material EXAMINED. — Philippines. "Albatross": stn 5543, 1 colony and 6 detached corallites (USNM

97312).

Type Locality. — "Hassler" stn unknown: off Barbados, 183 m.

DESCRIPTION (Specimen from "Albatross" stn 5543). — Corallum a bushy, irregularly branched colony of

about 30 corallites showing 5 generations of budding. Colony 9 cm in width, composed of elongate-conical

corallites, the largest measuring 12.1 x 12.5 mm in calicular diameter and 37 mm in length. Each corallite bears

up to 7 buds that originate from the edge zone just below calicular edge. C1-3 usually slightly ridged near calicular

edge, flat and granular, like the C4, on lower 2/3 of corallum. Corallum uniformly white.

Septa hexamerally arranged in 3 cycles (Si>S2>S3), S4 represented only as small exsert costosepta at calicular

margin. Si about 1.8 mm exsert, quite narrow (only about 2/5 calicular radius in upper corallite), having straight,

entire inner edges that thicken deep in fossa near columella. S2 about 1.1 mm exsert, 2/3 width of an Si, having

coarsely dentate inner edges near columella. S3 about 0.9 mm exsert, 2/3 width of an S2, having highly laciniate

inner edges. S4 about 0.4 mm exsert, occurring only at calicular edge. Fossa very deep and wide. Columella

variable, but most often composed of crispate lamellae that are attached to lower, inner edges of Si -2. Endothecal

dissepiments present.

AZOOXANTHELLATE SCLERACTINIA

139

REMARKS. — The Philippine specimen differs from most Atlantic specimens in having larger corallites,

a more robust colony, and a much deeper fossa. But, C. arbuscula is a variable species, and Atlantic specimens are

known to have even larger corallites, colonies of 4 corallite generations, and equally deep fossae. Since all other

characters are remarkably similar, this specimen, as well as one reported from Japan (Cairns, 1994), are identified

as C. arbuscula. The second author is dubious about the identification of the Philippine specimens with a species

known from the Atlantic, believing that there are probably some characters as yet not fully understood that

differentiate the two taxa.

DISTRIBUTION. — Philippines: Bohol Sea; 296 m. Elsewhere : tropical and warm temperate Amphi-Atlantic

(109-622 m); northern Ryukyu Islands (238-240 m).

Genus GONIOCORELLA Yabe & Eguchi, 1932

Goniocorella dumosa (Alcock, 1902)

Pourtalosmilia dumosa Alcock, 1902c: 36-37, pi. 5, fig. 33.

Goniocorella dumosa - CAIRNS, 1982: 31-34, pi. 9, figs 7-9, pi.

10, figs 1-2 (synonymy); 1994: 63-64, pi. 27, fig. j

(synonymy); 1995: 80-81, pi. 22, figs e-h (synonymy).

Goniocorella glanulosa (sic) Hu, 1987: 41, pi. 3. figs 3, 13-15, 17-20 (new synonym).

MATERIAL EXAMINED. — Indonesia. "Albatross": stn 5586, 9 dead fragments (USNM 97313).

South China Sea. "Hakuho Maru": stn KH73-2-44-2, 2 dead fragments (USNM 97314).

Type Locality. — "Siboga" stns 156 and 259: Banda Sea, 469-487 m.

DIAGNOSIS. — Bushy colonies formed by extratentacular, right-angled budding, the cylindrical coiallites

3-4 mm in diameter. Adjacent corallites often linked by slender, solid, tubular coenosteal bridges. Costae

inconspicuous. Septa hexamerally arranged in 3 cycles (Si>S2>S3). AH septa little exsert and narrow, resulting in

a vacuous fossa. No columella. Tabular endothecal dissepiments common.

Remarks. — Goniocorella dumosa is more fully described and figured by CAIRNS (1982, 1994).

DISTRIBUTION. — Indonesia: Halmahera Sea; Banda Sea (Kai Islands); 469-616 m. Elsewhere: Malaysia

(Celebes Sea off Sabah); throughout Indo-West Pacific from southwestern Indian Ocean to Japan, including South

China Sea (southern Formosa Strait) and New Zealand region; 88-1488 m. Plio-Pleistocene of Taiwan (Hu, 1987).

Genus CONFLUPHYLLIA nov.

Type Species. — Confluphyllia juncta, here designated.

DIAGNOSIS. — Colonial, extratentacular budding forming bushy colonies. Branch anastomosis common,

adjacent corallites also united by solid, cylindrical to sheet-like coenosteal bridges. No pali. Columella fascicular.

Tabular endothecal dissepiments present.

Remarks. — Confluphyllia is similar to Goniocorella , both genera having the same type of colony formation

as well as solid coenosteal bridges, but differs in having a robust, fascicular columella, Goniocorella having none.

Confluphyllia is further differentiated from Goniocorella by having more septa (up to 42 vs 24), which are more

exsert and wider; ridged C1-2; and larger-diameter corallites. Confluphyllia differs from Coenosmilia in having a

more robust colony that is reinforced with coenosteal bridges, and a fascicular columella ( Coenosmilia has a

crispate columella).

Source : MNHN, Paris

140

S. D. CAIRNS & H. ZIBROWIUS

ETYMOLOGY. — The genus name Confluphyllia (Latin confluus, which flows together + phyllia ,

a common coral suffix) refers to the confluent nature of the branches as linked by coenosteal bridges. Gender:

feminine.

Confluphyllia juncla sp. nov.

Figs 19 d-g

Material EXAMINED/TYPES. — Philippines. MUSORSTOM 3: stn 126. 2 dead colonies, paratypes (USNM

97315).

Indonesia. Deki: stn 59, 5 paratype colonies (NNM 22442).

Karubar: stn 25, holotype (MNHN) and 1 paratype colony (USNM 97316).

Type Locality. — Karubar stn 25: 5°25’S, 132°51'E (Banda Sea, Kai Islands), 318-352 m.

Etymology. — The species name (Latin jungere, to join, unite) refers to the connecting coenosteal bridges.

Description. — Corallum bushy, produced by closely-spaced extratentacular budding from the edge zone of

parent corallites, resulting in frequent branch anastomosis. Colonial structure reinforced by solid coenosteal bridges

that link adjacent corallites, as in Goniocorella. Coenosteal bridges circular in cross section (about 1 mm in

diameter), elliptical in cross section, or lamellar, the latter shape joining two adjacent corallites by a thin

coenosteal sheet several mm long. Holotype colony 7 cm wide, consisting of about 85 corallites, one (perhaps the

founder corallite) 6.5 cm in length. Ci slightly ridged, especially near calice, and usually white. C2 may also be

ridged, but only near calice. Otherwise, theca covered with low, rounded (0.2 mm diameter) granules and light

yellow-brown in colour. Edge zone on corallites extends about 7 mm from calice.

Septa hexamerally arranged in 4 cycles, the last cycle not complete. Corallites less than 4 mm in GCD have 24

or less septa, but larger corallites (up to 7.2 mm in GCD) have several pairs of S4, the most common

complements being 36 or 38 septa, although some corallites have as many as 42 septa. Si exsert (1.3- 1.5 mm),

having straight, vertical inner edges that extend to the columella. S2 less exsert (about 0.8 mm) and only 0.8 width

of an Si, also having straight inner edges that fuse with the columella deep in fossa. S3 about 3/4 exsertness and

width of an S3. S4 2/3 width of an S3. Fossa deep, containing a narrow fascicular columella consisting

of 1-5 thick, granular elements.

DISTRIBUTION. — Philippines : Sulu Sea (Semirara Islands); 266 m. Indonesia: Banda Sea (Kai Islands); 318-

385 m.

Family TURBINOLIIDAE H. Milne Edwards & Haime, 1848

Genus CONOCYATHUS d'Orbigny, 1849

Conocyathus zelandiae Duncan, 1876

Conocyathus zelandiae Duncan, 1876: 431, pi. 38, figs 1-3. — Wells, 1964: 113-1 14. — FlLKORN, 1994: 16. — CAIRNS,

1995: 83-84, pi. 23, figs f-i (synonymy).

Conocyathus sulcatus - TENISON Woods, 1878: 301-302. [Not Conocyathus sulcatus d'Orbigny in H. Milne Edwards &

Haime, 1851J.

Trematotrochus zelandiae - FOLKESON, 1919: 14.

Turbinolia australiensis Gardiner, 1939: 332-333, pi. 21, fig. 2.

Conocyathus - Wells, 1967: 355.

Material EXAMINED. — Indonesia. Snellius 2: stn 4.056, 1 1 (NNM 22637).

Karubar: stn 44, 4 (USNM 97320). — Stn 86, 1 (MNHN).

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

141

Type Locality. — Cook Strait, New Zealand (depth not given), but see CAIRNS (1995).

DIAGNOSIS. — Corallum conical, free, and quite small, the largest known corallum 3.4 mm in calicular

diameter and 6.1 mm in height. Ci-4 highly ridged, but C4 do not correspond to a septal cycle. Intercostal furrows

deep, each containing regularly spaced, circular pits 55-65 p.m in diameter. Corallum white. Septa hexamerally

arranged in 3 cycles: Si>S2>S3- Six large P2 form a palar ring that encircles a low, smooth, flat columella.

REMARKS. — 16 additional specimens of this tiny, rarely collected species are reported herein, the largest

Karubar specimen measuring 2.6 mm in calicular diameter and 4.5 mm in height. This species is more fully

described and figured by CAIRNS (1995) and GARDINER (1939, as Turbinolia australiensis).

Distribution. — Indonesia : Arafura Sea (south of Tanimbar Islands); Savu Sea (Sumba); 125-222 m.

Elsewhere : Persian Gulf; Western Australia; New South Wales; Arafura Sea (Arnhem Land, Australia); ? New

Zealand; 6-130 m.

Genus ENDOCYATHOPORA Cairns, 1989

Endocyathopora laticostata Cairns, 1989

Endocyathopora laticostata Cairns, 1989a: 39-40, pi. 21, figs a-e.

MATERIAL EXAMINED. — Indonesia. DEKl: unnumbered station in Bay of Ambon, 91 m, 5: 3 (ZMUC), 2 (USNM

97322). — Unnumbered station in Bay of Ambon, 100 m, 9 (NNM 22692). — Unnumbered station off Gelala. Ambon,

depth unknown, 73 (NNM 22691). — Unnumbered station between Neira and Lontor, Banda Islands, 75-90 m, I (NNM

22693).

MORTENSEN'S Java-S.A. Expedition: stn 9, 1 (ZMUC).

Type Locality. — "Albatross" stn 5136: 6°44'45"N, 121°48'E (Sulu Sea, Philippines), 46 m.

Diagnosis. — Corallum a narrow, regular cone with a blunt rounded base. Largest known specimen (Deki,

Ambon) 4.0 mm in calicular diameter and 8.6 mm in height. Costae broad and flat, covered with small (about

20 ^tm in diameter) granules arranged 5 or 6 across width of each costa. Intercostal furrows deep and narrow but

wider towards theca, undercutting adjacent costae. Inside corallum, aligned with each intercostal region, is a row of

small (110-120 |am) shallow depressions, which, by erosion following the death of the coral tend to become pores.

Corallum white. Septa hexamerally arranged in 3 complete cycles: S i>S2-3- Six P2 form a palar crown that

encircles a columella composed of 1 or 2 papillae.

Remarks. — This constitutes the second report of this turbinoliid, which is more fully described and

illustrated by CAIRNS (1989a).

Distribution. — Philippines : Sulu Sea (Mindanao and Basilan); 46-70 m. Indonesia : Banda Sea (Bay of

Ambon and Banda Islands); Bali Strait; 70-100 m.

Genus ALATOTROCHUS Cairns, 1994

Alatolrochus rubescens (Moseley, 1876)

Fig. 18 h

Platytrochus rubescens Moseley, 1876: 553.

Sphenotrochus rubescens - MOSELEY, 1881: 157-159, pi. 6, figs 8, 8a.

Alatotrochus rubescens - Cairns, 1994: 68-69, pi. 29, figs g-1 (synonymy); 1995: 84, pi. 24, figs a-b (synonymy).

Source : MNHN, Paris

142

S. D. CAIRNS & H. ZIBROWIUS

MATERIAL EXAMINED. — Philippines. "Hakuho Maru": stn KH72-1-20, 43 (USNM 97325), 10 (ORI).

MUSORSTOM 2: stn 2, i (MNHN).

Musorstom 3: stn 86, I (MNHN). — Stn 88, 1 (MNHN). — Stn 91, 1 (MNHN). — Stn 96, 2 (MNHN). — Stn 102,

1 (MNHN).

Indonesia. Deki: stn 44, 1 (NNM 22638). — Stn 49, 1 (NNM 22639).

Karubar: stn 2, 9 (USNM 97324). — Stn 3, 1 (POLIPI).

Type Locality. — "Challenger" stn 192: 5°49'15"S, 132°14'15"E (Kai Islands, Banda Sea), 136 m.

Diagnosis. — Corallum compressed, the calice elliptical (GCD:LCD = 1.15-1.25) and the thecal faces

meeting in sharp edges that, on the lower corallum, project as much as 4 mm as thick, solid edge crests. Largest

Philippine specimen (MUSORSTOM 3 stn 88) 1 1.8 x 18.3 mm in calicular diameter, which is close to the largest

recorded size. Four cycles of ridged costae (Cm) corresponding to Si -4, but C5 not corresponding to a septal cycle

(no S5). Edge crests also ridged, as though by costae, but these ridges oriented perpendicularly to thecal costae.

Septa hexamerally arranged in 4 complete cycles: Si>S2>S3»S4, the S1-2 being highly exsert and extending to

the columella. Columella papillose, consisting of 5-12 robust pillars; depending on the individual the columellar

pillars may be circular or lamellar in cross section.

Remarks. — Three relatively large specimens (MUSORSTOM 3 stns 88, 91, 102) differ in having their

4 lateral Ci swollen and protuberant in the upper 8-10 mm of the theca (Fig. 18h). This variation was previously

reported for a Japanese Pleistocene specimen (CAIRNS, 1994). Alatotrochus rubescens is more fully described and

illustrated by Cairns (1994).

DISTRIBUTION. — Philippines: Lubang Island; Sulu Archipelago; 187-460 m. Indonesia: Banda Sea (Kai

Islands); 136-278 m. Elsewhere: Japan (Kyushu and Ryukyu Islands); southern Norfolk Ridge; 193-751 m.

Genus CRYPTOTROCHUS C aims, 1988

"Cryptotrochus" venustus (Alcock, 1902)

Ceratotrochus venustus Alcock, 1902a: 92; 1902c: 10, pi. 1, figs 5, 5a.

Not Cryptotrochus venustus - Cairns, 1995: 88-89, pi. 26, figs g-i, pi. 27, figs a-b (= gen. nov., sp. nov., Cairns, in

press).

Material EXAMINED. — Indonesia. Deki: stn 57, 1 (NNM 22649). — Stn 58, 1 (NNM 22650).

Karubar: stn 2, 2 (POLIPI). — Stn 3, 44: 5 (MNHN), 36 (USNM 96906); Stn 7, 15 (MNHN).

Type Locality. — "Siboga" stn 256: 5°26.6'S, 132°32.5'E (Kai Islands, Banda Sea), 397 m.

DIAGNOSIS. — Corallum conical but slightly compressed, resulting in an elliptical calice with a GCD:LCD of

1.1 -1.2. Largest Indonesian specimen (Karubar stn 7) 10.1 x 11.2 mm in calicular diameter and 10.4 mm in

height. Costae ridged, often discontinuous, and separated by broad intercostal furrows. Five cycles of costae usually

present on coralla having only 4 cycles of septa, the C5 terminating at the calicular margin as exsert spurs without

septal continuation within calice. Thecal edges rounded in upper corallum but on lower corallum faces meet in an

acute, non-carinate, angle. Corallum white. Septa hexamerally arranged in 4 cycles (44-48 septa): Si>S2>S3>S4,

the Si being highly exsert. Six thin lamellar P2 encircle a papillose columella.

Remarks. — "Cryptotrochus" venustus differs from the type-species C. javanus Cairns, 1988, in having one

more cycle of costae; much wider and shallower intercostal regions; costae of independent origin; and a slightly

compressed corallum (that of C. javanus is conical). Because of these significant differences, C. venustus is not

confidently assigned to Cryptotrochus ; its systematic position is elaborated on in greater detail by Cairns (in

press).

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

143

DISTRIBUTION. — Indonesia: Banda Sea (Kai Islands); 200-397 m.

Genus NOTOCYATHUS Tenison Woods, 1880

Notocyalhus venustus (Alcock, 1902)

Citharocyathus venustus Alcock, 1902b: 119; 1902c: 22, pi. 3, figs 19, 19a.

Notocyathus venustus - Cairns, 1989a: 27-28, pi. 12, figs c-h (synonymy); 1994: 64. pi. 27, figs k-1.

MATERIAL EXAMINED. — Indonesia. Deki: stn 24, 10 (NNM 22682). — Stn 44, 1 (NNM 22683). — Stn 50, 17: 1

(USNM 97336), 16 (NNM 22685). — Stn 62, 1 (USNM 97338).

MORTENSEN'S Java-S.A. Expedition: stn 18, 1 (USNM 97339).

SNELLIUS 2: stn D2, 1 (NNM 23090). — Stn 81.2, 6 (NNM 22690).

KARUBAR: stn 2, 157 (MNHN). — Stn 3, 200+ (USNM 97327). — Stn 7, 1 18 (MNHN). — Stn 15, 1 (MNHN). —

Stn 22, 1 (USNM 97330). — Stn 31, 5 (POLIPI). — Stn 49, 3 (POLIPI). — Stn 50, 1 (USNM 97333).

Type Locality. — "Siboga" stn 59: 10°22.7'S, 123°16.5'E (Savu Sea), 390 m.

DIAGNOSIS. — Corallum conical, the largest Indonesian specimen (a syntype) 6.5 x 6.7 mm in calicular

diameter and 9.2 mm in height. Calice almost perfectly circular: GCD:LCD = 1.02-1.05. All costae bear a row of

outward projecting granules, as well as 2 rows of laterally oriented granules that project across the deep intercostal

furrow. Corallum white. Septa hexamerally arranged in 4 complete cycles: Si>S2>S3»S4- Si highly exsert, their

upper edges being coarsely dentate and inclined inward toward fossa. S4 considerably less exsert and less wide, but

as thick as an S3, their septal faces often lacking granules. Within each septal system a pair of lamellar P3 fuse in

V-shaped structures before their common P2. Fossa shallow if any; columella papillose.

Remarks. — As previously discussed (Cairns, 1989a, 1994), Notocyathus venustus is very similar to

N. conicus (Alcock, 1902), the coralla of small specimens being difficult or impossible to distinguish. To

reiterate, N. venustus differs in having a nearly circular calice and thus a lower GCD:LCD (1.02-1.05 vs 1 .01-1.22

for N. conicus), and more exsert S 1-2 and proportionally less exsert S4. In large specimens of N. venustus, S3

become almost as wide and exsert as the S2, but the S4 remain quite small and thin. In large specimens of

N. conicus, the S4 are only slightly less exsert and less wide than the S3. This species is more fully described and

illustrated by Cairns (1989a, 1994).

Distribution. — Philippines : Sibuyan Sea; Sulu Sea (Mindanao and Sulu Archipelago); 70-555 m.

Indonesia: Banda Sea (Kai and Tanimbar Islands); Savu Sea; Java Sea; 100-390 m. Elsewhere: Malaysia (Darvel

Bay, off Sabah, Celebes Sea); Japan (Kyushu and Ryukyu Islands); 193-422 m.

Notocyathus conicus (Alcock, 1902)

Citharocyathus conicus Alcock, 1902b: 118-119; 1902c: 22, pi. 3, figs 18, 18a.

Notocyathus conicus - CAIRNS, 1989a: 28, pi. 13, figs a-i (not "Albatross" stn 5586, a worn unidentified specimen)

(synonymy); 1994: 64-65, pi. 28, figs a-b; 1995: 91-92, pi. 27, figs c, g.

MATERIAL EXAMINED. — Philippines. "Hakuho Maru ": stn KH72-1-20, 1 (USNM 97343).

Musorstom 2: stn 11, 1 (MNHN). — Stn 33, 10 (USNM 81801).

MUSORSTOM 3: stn 102, 6 (MNHN). — Stn 106, 1 (MNHN).

Indonesia. Deki: stn 6, 20 (NNM 22761).

Corindon 2: stn 235, 3 (MNHN).

Karubar: stn 2, 2 (USNM 97341). — Stn 15, 4 (POLIPI). — Stn 18, 18 (MNHN). — Sin 32, 1 (USNM 97342).

Type Locality. — "Siboga" stn 95: 5°43.5'N, 1 19°40'E (Sulu Sea, Philippines), 522 m.

Source : MNHN , Paris

144

S. D. CAIRNS & H. ZIBROWIUS

DIAGNOSIS. — Corallum conical, largest Indonesian specimen (a syntype) 6.9 x 7.0 mm in calicular diameter

and 8.1 mm in height. Calice usually slightly elliptical: GCDiLCD = 1.01-1.22. Costae and costal granulation as

in C. venustus. Corallum white. Septa hexamerally arranged in 4 complete cycles: Si>S2>S3>S4. Difference

between exsertness and width of 4 septal cycles gradual, the S4 of large specimens sometimes almost as wide as

the S3. Pairs of lamellar P3 fuse in a V-shaped structure before their common P2. Fossa shallow; columella

papillose.

REMARKS. — This species is compared to N. venustus in the account of that species and is more fully

described and figured by Cairns (1989a, 1994).

Distribution. — Philippines: Lubang Island; Sibuyan Sea; Visayan Sea; Bohol Sea; Sulu Archipelago;

34-923 m. Indonesia: Makassar Strait; Banda Sea (Kai Islands); 206-210 m (depth of 1 1 10 m of CORINDON 2

stn 235 considered erroneous). Elsewhere: Japan (Kyushu, Bungo Strait, and Honshu); Norfolk and Kermadec

Ridges north of New Zealand; 70-710 m.

Genus DELTOCYATHOIDES Yabe & Eguchi, 1932

Deltocyathoides orientalis (Duncan, 1876) comb. nov.

Deltocyathus orientalis Duncan, 1876: 431, pi. 38, figs 4-7.

Peponocyathus australiensis - CAIRNS, 1989a: 29, 30-32, pi. 14, figs d-j, pi. 15, figs a-d (synonymy); 1994: 64-65,

pi. 28, figs c-f, pi. 41, fig. i (synonymy). — Cairns & Parker, 1992: 39-40, pi. 13, figs c-d.

Deltocyathus lens Alcock, 1902a: 99; 1902c: 19-20, pi. 2, figs 16, 16a. — Zou, 1988: 77-78, pi. 5, figs 6, 6a.

Not Peponocyathus orientalis - Yabe & EGUCHI, 1932a: 444-445 [= ? P. folliculus (PourtaRs, 1868)].

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5137 (confused origin, probably 5135), 1 (USNM

97344).

"Hakuho Maru ": stn KH72-1-20, 1 (USNM 97353).

Musorstom 2: stn 33, 142 (MNHN).

MUSORSTOM 3: stn 102, 1 (USNM 97345). — Stn 1 17, 1 (MNHN). — Stn 139, 1 (USNM 97346).

Indonesia. DEKI: stn 6, 1 1 (NNM 22640). — Stn 7, 8 (ZMUC). — Stn 24, 9 (NNM 22641). — Stn 26, 1 (ZMUC). —

Stn 44, 1 (NNM 22642). — Stn 46, 2 (ZMUC). — Stn 49, 18 (NNM 22643). — Stn 53, 19 (ZMUC).

"Galathea": stn 500, 1 (ZMUC).

CORINDON 2: stn 248, 3 (MNHN).

SNELLIUS 2: stn D2, 6 (NNM 23093). — Stn 81.2, 14 (NNM 22094).

Karubar: stn 2, 49 (MNHN). — Stn 3, 26 (POLIPI). — Stn 7, 40 (USNM 97349). — Stn 13, 2 (MNHN). — Stn 15,

35 (POLIPI). — Stn 18, 19 (USNM 97350). — Stn 22, 9 (MNHN). — Stn 28, 4 (USNM 97351). — Stn 29, 1 (MNHN). —

Stn 44. 22 (MNHN). — Stn 49, 18 (MNHN). — Stn 61, 35 (POLIPI).

Type Locality. — 34°12’N, 136°20'E (southeastern Honshu, Japan), 95 m.

DIAGNOSIS. — Corallum bowl-shaped, with a gently rounded base. Asexual reproduction through transverse

division not observed in this species. Coralla usually less than 7.5 mm in calicular diameter. Costae equal in

width, separated by deep intercostal furrows of about the same width. Each costa bears a row of outward-projecting,

blunt spines as well as smaller spines that project into intercostal furrows. Corallum white. Septa hexamerally

arranged in 4 complete cycles above a GCD of about 3.5 mm; 24 septa are present in smaller specimens, rarely

with any intermediate number. Septal formula: Si>S2>S4>S3, all septa being equally exsert. Si independent; pairs

of S3 join S2 and pairs of S4 to S3 within each septal system. P2-3, and occasionally Pi, present. Columella

papillose.

Remarks. — As implied by Cairns (1995), the genus Deltocyathoides should be employed for those

"peponocyathid" species that do not reproduce by transverse division, and the name Peponocyathus for those

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

145

species that do. This distinction is discussed in greater detail by Cairns (in press), and this species is more fully

described and illustrated by CAIRNS (1989a, 1994) as Peponocyathus australiensis.

Petrarcid ascothoracidan galls were found in the coralla of specimens from "Albatross" stations 5178, 5313,

5314, 5315, 5317, and 5403 (see Cairns, 1989a), the record from "Albatross" stn 5403 previously reported as

such by ZlBROWlUS and GRYGIER (1985).

DISTRIBUTION. — Philippines', throughout region from Luzon to Sulu Archipelago; 44-635 m. Indonesia:

Makassar Strait; Banda Sea (Kai, Tanimbar, and Tukangbesi Islands); Arafura Sea (east of Tanimbar Islands); Savu

Sea (Timor Sea); 85-393 m. Elsewhere: southwest Indian Ocean to Japan; 44-635 m.

Genus PEPONOCYATHUS Gravier, 1915

Peponocyathus minimus (Yabe & Eguchi, 1937)

Fig. 18 i

Discotrochus ( Cylindrophyllia ) minimus Yabe & Eguchi, 1937: 146-147, pi. 20, tigs 16-22; 1942b: 118.

Cylindrophyllia minimus - Squires, 1958: 58. — EGUCHI, 1965: 289, 2 figs. — Kikuchi, 1968: 8, figs 3a, b. —

Hu, 1988: 151, pi. 1, figs 9-12, 14-15.

Peponocyathus folliculus - Cairns, 1989a: 32-33 (in part: pi. 15, e-h, "Albatross": stns 5172, 5217, 531 1). [Not

Stephanophyllia folliculus Pourtalfes, 1868].

Cylindrophyllia orientalis - MORI & MlNOURA, 1983: 185-191, figs 1-6. [Not Peponocyathus orientals Yabe & Eguchi,

1932].'

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5172, 1 (USNM 81838). — Stn 5217, 1 (USNM 81839).

— Stn 5236, 36 (USNM 97354). — Stn 531 1, 6 (USNM 81841). — Stn 5312, 8 (USNM 97355). — Stn 5313, 13 (USNM

97356). _ Stn 5315, 3 (USNM 97357). — Stn 5424, 1 (USNM 97358). — Stn 5579, 1 (USNM 97359). — Stn 5586,

8 (USNM 97360).

"Galathea": stn 477, 3 (ZMUC).

Indonesia. Corindon 2: stn 210, 1 (MNHN). — Stn 235, 1 (MNHN).

Snellius 2: stn 81.2, 4 (NNM 22762, 23099).

Karubar: stn 2, 4 (POLIPI). — Stn 3, 195 (POLIPI). — Stn 7, 500+ (USNM 97362). — Stn 15. 16 (MNHN). —

Stn 18, 13 (MNHN). — Stn 22, 1 (MNHN). — Stn 44, 4 (MNHN). — Stn 49, 33 (MNHN). — Stn 61, 4 (MNHN).

Type Locality. — Neogene of Taiwan, Kyushu, and Honshu, and Recent of Toyama Bay, Japan.

DIAGNOSIS. — Anthocaulus not yet known for this species. Anthocyathus cylindrical (usually wider than

high), often with a flat base, the result of asexual transverse division. Anthocyathi in process of division rare (but

see Fig. 18i), the corallum base appearing "unfinished" or open (thickened closing deposit not yet developed),

characteristic of a recent fission from a parent corallum. Corallum rarely over 4.0 mm in calicular diameter. Costae

equal in width and covered with fine granules; costae separated by relatively wide intercostal furrows, often as wide

as the costae. If C4 are present, they originate on the horizontal flat base, not the vertical thecal walls. Corallum

white. Septa hexamerally arranged in 3 to 4 cycles, the 4th cycle never complete, most coralla having 24 or

36 septa: Si>S2-3. Small pali present before all but last septal cycle.

Remarks. — Peponocyathus minimus is similar to P. folliculus, both species being known from the

Philippine region. CAIRNS (1989a) mistakenly included 3 lots of P. minimus in a previous account of

P. folliculus (see synonymy). P. minimus differs in having a short, squat corallum (that of P. folliculus is

usually higher than wide); a flat base, not rounded; and intercostal furrows almost as wide as the costae (those ot

P. folliculus being very narrow and the costae quite wide). Furthermore, in P. minimus pairs of C4 originate on

the anthocyathus base, in P. folliculus on the thecal edges.

Peponocyathus minimus is described in greater detail by MORI & MlNOURA (1983) as Cylindrophyllia

orientalis.

Source : MNHN, Paris

146

S. D. CAIRNS & H. ZIBROWIUS

DISTRIBUTION. — Philippines : Sibuyan Sea; Sulu Sea (Cagayan Islands); eastern Mindanao; Sulu

Archipelago; 161-903 m. Indonesia : Celebes Sea (Borneo); Makassar Strait; Banda Sea (Tanimbar and Kai Islands);

Bali Strait; 124-616 m (depth of 1 1 10 m from CORINDON 2 stn 235 considered erroneous). Elsewhere : South

China Sea (Pratas Island); Japan (Honshu, Kyushu, and the Ryukyu Islands); 30-402 m. Pleistocene of Ryukyu

Islands.

Peponocyathus folliculus (Pourtal&s, 1868)

Siephanophyllia folliculus PourtaRs, 1868: 139.

Peponocyathus orientalis Yabe & Eguchi, 1932b: 444-445, unnumb. figs.

Peponocyathus folliculus - Zibrowius, 1980: 113-115, pi. 58, figs A-L, pi. 59, figs A-K (synonymy). — Cairns,

1989a : 32-33 (in part: pi. 16. figs a-c, "Albatross" stns 5277, 5577, 5584); 1994: 66-67, pi. 28, figs g-k.

MATERIAL EXAMINED. — Indonesia. Deki: stn 10, 3 (NNM 23097).

Snellius 2: stn 4.065 (station data possibly confused), 1 (NNM 23098).

Karubar: stn 15. 14: 8 (MNHN), 6 (USNM 97363).

Type Locality. — "Bibb" stn 51: 24°12'40"N, 81°19'25"W (Straits of Florida), 433 m.

DIAGNOSIS. — From Indonesia, known only from the anthocyathus stage. Corallum cylindrical, with a flat to

slightly rounded base that is the result of transverse division; small, rarely exceeding 4 mm in calicular diameter;

adult corallum usually higher than wide (i.e., H:D>1). Costae broad, convex, and finely granular. Corallum white.

Septa hexamerally arranged in 3 to 4 cycles, the 4th cycle never complete, 24-36 being the range of septal number.

Septa arranged: Si>S2>S3, each of the 6 S2 bearing a prominent paliform lobe. Fossa shallow; columella

papillose.

Remarks. — This species is more fully described and illustrated by Cairns (1994), and is compared to

P. minimus in the previous account.

DISTRIBUTION. — Philippines : Lubang Island; Sulu Sea (Sulu Archipelago); 146-534 m. Indonesia'. Banda Sea

(Kai Islands); Savu Sea (Sumba); 50-212 m. Elsewhere: Japan (Honshu, Kyushu, and northern Ryukyu Islands);

Atlantic; ? 30-582 m.

Genus TROPIDOCYATHUS H. Milne Edwards & Haime, 1848

Tropidocyathus lessonii (Michelin, 1842)

Flabellum lessonii Michelin. 1842: 119.

Trochocyathus (Tropidocyathus) lessoni - At.COCK, 1902c: 17, pi. 2, figs 14, 14a.

Tropidocyathus lessonii - Zou et ai. 1988: 195.

Tropidocyathus lessoni - Cairns, 1989a: 33-34, pi. 16, figs d-1 (synonymy); 1994: 67, pi. 29, figs a-b (synonymy). —

Cairns & Keller, 1993: 253, pi. 7, fig. C.

Material EXAMINED. — Philippines. Musorstom 1: stn 64, 1 (MNHN).

Musorstom 2: stn 29, 1 (USNM 97364). — Stn 33, 1 (MNHN).

Musorstom 3: stn 88, 1 (MNHN). — Stn 108, 1 (USNM 97365). — Stn 131, 9 (MNHN).

Indonesia. Deki: stn 49, 53 (NNM 22670). — Stn 53, 2 (NNM 22671).

Mortensen s Java-S.A. Expedition: stn 5, 3 (ZMUC). — Stn 8, I (ZMUC).

Karubar: stn 1, 1 (MNHN). — Stn 2, 8 (POLIPI).

Type Locality. — Unknown.

DIAGNOSIS. — Corallum cuneiform, with a rounded base and pronounced (up to 4 mm in height), thin, thecal

edge crests. Calice elliptical to rhombus-shaped, with a GCD:LCD = 1.1 -1.5. Largest Indonesian specimen

(Karubar stn 2) 13.2 x 15.7 mm in calicular diameter and 14.2 mm in height. Costae low and flat, covered with

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

147

low, blunt granules, 3 or 4 occurring across the width of each Ci-3- Costae do not extend to thecal edge crests;

however, crests are covered with low aligned granules and ridges. Theca pale orange; calicular elements white.

Septa hexamerally arranged in 4 cycles: Si>S2>S3>S4- Lamellar pali present before Si -3. Columella papillose.

Remarks. — Some Musorstom specimens were previously reported by Cairns (1989a). A more detailed

description and illustrations of this species are provided by Cairns (1989a, 1994).

DISTRIBUTION. — Philippines: Lubang Island; Verde Island Passage; Tablas Strait; Sulu Sea (Panay); Sulu

Archipelago; 68-421 m. Indonesia: Banda Sea (Kai Islands); Timor Sea; Savu Sea (Timor); Flores Sea; Bali Strait;

50-390 m. Elsewhere: South China Sea off Hong Kong; Japan (Kyushu and Ryukyu Islands); western Indian

Ocean (Natal, Somalia, and Kenya); 62-155 m.

"Tropidocyathus" pileus (Alcock, 1902)

Figs 19 h-i

Trochocyathus pileus Alcock, 1902a: 96-97; 1902c: 15-16, pi. 2, figs 11, 11a. — Faustino, 1927: 81, pi. 7, figs 7-8. —

Zou etal., 1988: 195.

Tropidocyathus pileus - Cairns, 1989a: 34-35, pi. 17, figs a-h (synonymy); 1994: 68, pi. 29, figs d-e; 1995: 91, pi. 28,

figs a-c.

MATERIAL EXAMINED. — Philippines. "Hakuho Maru": stn KH72-1-20, 47: 37 (USNM 97382), 10 (ORI).

Musorstom 1: stn 13, 1 (MNHN). — Stn 40, 7 (USNM 97367). — Stn 42, 2 (MNHN). — Stn 55, 1 (USNM 97369).

— Stn 58, 9 (USNM 97370). — Stn 62, 1 (MNHN).

Musorstom 2: stn 6, 3 (MNHN). — Stn 10, 1 (MNHN). — Stn 63, 30 (USNM 97372). — Stn 83, 19: 16 (MNHN),

3 (BMNH 1992.8.11.15).

Musorstom 3: stn 88, 1 (MNHN). — Stn 92, 322: 313 (USNM 97374), 9 (BMNH 1992.8.11.16). — Stn 96,

1 (USNM 97375). — Stn 99, 1 (MNHN). — Stn 108, 3 (USNM 97376). — Stn 109, 1 (MNHN). — Stn 124, 3 (MNHN).

Indonesia. DEKI: stn 49, 3 (NNM 22491).

Karubar: stn 2, 32 (POLIPI). — Stn 3, 3 (POLIP1). — Stn 7, 12 (MNHN). — Stn 35, 1 (USNM 97380). — Stn 36,

1 (MNHN). — Stn 62, 8 (USNM 97381). — Stn 67, 1 (MNHN). — Stn 85, 2 (POLIPI).

Type Locality. — "Siboga" stn 95: 5°43'N, 119°40'E (Sulu Archipelago, Philippines), 522 m.

Diagnosis. — Corallum variable in shape: viewed from the side, most coralla are trapezoidal, the basal

(shortest) edge often being straight or slightly curved. Other coralla triangular in profile or bowl-shaped. All coralla

laterally compressed, resulting in an elliptical calice with a GCD:LCD = 1.50-1.65. Largest specimen (Karubar

stn 2) 17.9 x 24.8 mm in calicular diameter and 20.8 mm in height. Thecal edges rounded and occasionally

slightly protuberant near corallum base. Highly ridged, vertically oriented costae continuous from calice to base,

extending onto edge protuberances. Costae bear 1 apical row of coarse teeth and smaller, laterally oriented granules.

Intercostal furrows deep. Corallum white. Septa hexamerally arranged in 4 to 5 cycles (see Remarks):

S|>S2>S4>S3 or Si>S2>S3>S5>S4- Lamellar pali occur before all but last septal cycle (P1-3 or P1-4), the P2 and

2 P3 within each system in a chevron-shaped arrangement. Columella linear-papillose.

Remarks. — Although "T." pileus is similar to the type species T. lessonii (Michelin, 1842) in palar

configuration, it differs in costal origin and ornamentation; depth of intercostal regions; columellar structure; and

in lacking thecal edge crests. Discussed in greater detail by Cairns (in press), it is doubtful that T. pileus should

remain in the genus Tropidocyathus.

All specimens obtained from the Indonesian region differ from typical "T. "pileus in having more than 4 cycles

of septa, often a full 5th cycle (96 septa) or pairs of S5 in the 4 systems adjacent to the principal septa, resulting

in 80 septa, some coralla even having a few pairs of S6- This higher number of septa does not seem to be related

to a larger corallum size, since specimens as small as 9.8 mm in GCD (e.g., Karubar stns 3, 7) have a lull

5th cycle. Coralla having 5 cycles of septa are often triangular (not trapezoidal) in profile and have finer (thinner)

costae, their C5 originating on the thecal faces 2-7 mm above the base.

This common Indo-West Pacific species is more fully described and illustrated by Cairns (1989a, 1994).

Source :

148

S. D. CAIRNS & H. ZIBROWIUS

Distribution. — Philippines : Verde Island Passage and Tablas Strait adjacent to Mindoro; Bohol Sea; Sulu

Archipelago; 143-522 m. Indonesia : Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar Islands); 240-

390 m. Elsewhere : widespread from southwest Indian Ocean to Japan, including Queensland, the Norfolk Ridge,

and the South China Sea off Hong Kong; 123-422 m.

"Tropidocyathus" labidus sp. nov.

Figs 20 a-g

MATERIAL EXAMINED/TYPES. — Indonesia. Deki: stn 52, 1 paratype (NNM 22775).

" Galaihea stn 500, 1 (ZMUC).

Karubar: stn 2, holotype (MNHN) and 5 paratypes (MNHN). — Stn 3, 38 paratypes (MNHN). — Stn 7,

40 paratypes (USNM 97384). — Stn 15, 1 paratype (MNHN). — Stn 32, 1 paratype (POLIPI).

Japan. "Tansei Maru stn KT93-09-AM8, 3 paratypes (USNM 97385).

Type Locality. — Karubar stn 2: 5°47'00"S, 132°1 1'35"E (Kai Islands, Banda Sea), 209-240 m.

Etymology. — The species name (Latin labidus , slippery) is an allusion to the smooth thecal surface.

Description. — Corallum compressed-conical, resulting in a GCD:LCD of 1.06-1.23. Holotype 6.4 x

7.9 mm in calicular diameter and 10.5 mm in height; largest specimen (DEKI stn 52) 9.8 x 11.4 mm in calicular

diameter and 13.2 mm in height. Tip of base pointed, covered by the slender lower distal edges of the 12 C3; C1-2

originate about 0.5 mm above basal tip; C4 originate 1. 5-3.0 mm above tip. C4 on centre of thecal faces originate

highest on face, whereas C4 near thecal edges originate lowest, such that the points of origin of the 6 pairs of C4

on either thecal face form a crescent (Fig. 20d), the arc of which encircles the basal tip. Each costa bears a row of

low, large (200-250 pm in diameter), smooth, rounded granules, producing a slippery texture. Intercostal furrows

deep and narrow (50 pm). Corallum theca pale orange, as in T. lessonii , but basal tip and all calicular elements

white. Dead specimens also uniformly white.

Septa hexamerally arranged in 4 complete cycles: Si>S2>S4>S3 (48 septa). Si highly exsert (up to 1.6 mm),

having highly sinuous inner edges. S2 slightly less exsert, about 4/5 width of an Si, also having sinuous inner

edges. S3 much less exsert (about 0.8 mm), about 2/3 width of an S2, having sinuous inner edges. S4 as exsert as

S3, but wider than the S3 (almost as wide as the S2), having straight, thin inner edges. A crown of 12 sinuous-

lamellar P 1 -2 encircles the columella, the Pi about 0.5 mm in width, the P2 about twice that width. A 2nd crown

of 12 P3, each P3 about the same width of a P2, stands higher in the fossa. Inner edges of P3 slightly recessed from

columella, each pair of P3 within a system forming a chevron arrangement with its common P2. All septa and

pali bear tall (up to 0.1 1 mm), slender, pointed spines. Fossa shallow. Columella consists of a line of 3-6 robust

papillae, fused at their bases to inner edges of Pi -2.

Remarks. — "Tropidocyathus" labidus differs from the other two Recent species of Tropidocyathus by having

a smaller adult corallum; a corallum that is only slightly compressed and bears no edge crests or protuberances;

a distinctive costal insertion pattern; and a smooth, coarse costal granulation, quite unlike that of the other

species. Like "T. " pileus, it probably should form the basis of a separate genus.

Distribution. — Indonesia: Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar Islands); 206-

390 m. Elsewhere : Amami 0 Shima, Ryukyu Islands; 422-425 m.

Genus IDIOTROCHUS Wells, 1935

Idiotrochus kikutii (Yabe & Eguchi, 1941)

Placotrochides kikutii Yabe & Eguchi, 1941a: 104; 1942b: 149, pi. 9, fig. 16a-c.

Idiotrochus kikutii - Cairns, 1989a: 36-37, pi. 18, figs a-b. d-h (synonymy); 1994: 69, pi. 30, figs a-d.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

149

MATERIAL EXAMINED. — Philippines: Musorstom 2: stn 32, 2 (MNHN). — Stn 33, 409: 33 (USNM 81911),

376 (MNHN).

Musorstom 3: stn 102, 1 (MNHN). — Stn 117, 2 (MNHN).

Indonesia. Deki: stn 46, 1 (ZMUC).

Snellius 2: stn D2, 1 (NNM 23088).

Karubar: stn 2, 17 (MNHN). — Stn 3, 30 (MNHN). — Stn 7, 32 (USNM 97387). — Stn 15, 111 (USNM 97388). —

Stn 18, 13 (MNHN). — Stn 22, 39 (POL1PI). — Stn 49, 23 (MNHN).

Type Locality. — Toyama Bay, Japan (depth not given).

DIAGNOSIS. — Corallum (anthocyathus) with an elliptical calice (GCD:LCD = 1.15-1.95) and a wedge-shaped

base, the lower edge often blade-like. Largest specimen examined (MUSORSTOM 2 stn 32) 3.6 x 4.6 mm in

calicular diameter and 7.9 mm in height. Theca porcellaneous; costae low, wide, and smooth, corresponding to the

interseptal regions. Intercostal striae narrow and shallow, corresponding in position to the septa. Septa hexamerally

arranged in 3 cycles: Si>S2^S3. A crown of 10-12 Pi -2 encircle a papillose columella. Pi aligned with principal

Si often reduced in size or absent.

Remarks. — Most of the Musorstom specimens were previously reported by CAIRNS (1989a), who also

provided a more detailed description and illustrations of this species.

DISTRIBUTION. — Philippines : Verde Island Passage; Sibuyan Sea; Sulu Archipelago; 97-581 m. Indonesia :

Banda Sea (Kai and Tanimbar Islands); 300-645 m. Elsewhere: Malaysia (Celebes Sea off Sabah); South China Sea

off Hong Kong (161 m); Honshu, Japan (depth unknown).

Genus THRYPTICOTROCHUS Cairns, 1989

Thrypticotrochus multilobatus Cairns, 1989

Thrypticotrochus multilobatus Cairns, 1989a: 37, pi. 19, figs b-g; 1995: 92, pi. 28, figs d-h. — Cairns & Keller, 1993:

254, pi. 7, figs F, I.

Material EXAMINED. — Philippines. "Siboga": stn 102, 2 (ZMA Coel. 710b).

Musorstom 2: stn 33, 1 (MNHN).

Musorstom 3: stn 100, 1 (MNHN).

Indonesia. Deki: stn 46, 1 (ZMUC).

Karubar: stn 2, 1 (MNHN). — Stn 3, 10 (POLIPI). — Stn 7, 24 (USNM 97391). — Stn 18, 1 (MNHN).

Type Locality. — "Albatross" stn 5576: 5°25'56"N, 120o03'39''E (Sulu Sea, Philippines), 507 m.

Diagnosis. — Corallum small and conical, rarely exceeding 4 mm in calicular diameter or 7 mm in height.

Base of corallum irregularly shaped as the result of regeneration from a parent fragment. Costae broad and serrate,

separated by thin, deep intercostal furrows. Corallum white. Septa hexamerally arranged in 4 cycles:

Si>S2>S3»S4. Inner edges of each Si-3 bear 1-3 narrow paliform lobes, the lowermost merging with

the papillose columella.

Remarks. — Since its original description, which was based on material from the Philippines, this species

has been reported from the southwestern Indian Ocean and north of New Zealand, and is probably widespread

throughout the Indo-West Pacific. It is more fully described by Cairns (1989a).

Distribution. — Philippines: Verde Island Passage; Ragay Gulf; Sulu Archipelago; 192-535 m. Indonesia:

Banda Sea (Kai Islands); 278-300 m. Elsewhere: South China Sea (Pratas Islands); southwest Indian Ocean

(Mozambique, Tanzania); Queensland; Norfolk and Kermadec Ridges; 95-925 m.

Source :

150

S. D. CAIRNS & H. ZIBROWIUS

Supcrfamily FLABELLOIDEA Bourne, 1905

Family GUYNIIDAE Hickson, 1910

Genus GUYNIA Duncan, 1872

Guynia annulata Duncan, 1872

Guynia annulata Duncan, 1872: 32, pi. 1, figs 1-8. — ZIBROWIUS, 1980: 161-162, pi. 83, figs A-Q (synonymy). —

Cairns, 1984: 23, pi. 5, figs A-B; 1989a: 42-43, pi. 21. fig. f, pi. 22, figs a-e. — Cairns & Wells, 1987: 42-43,

pi. 11, figs 8-9, 12-13. — Cairns & Parker. 1992: 42-43, pi. 14, figs g-h. — Cairns & Keller, 1993: 273, pi. 12,

figs H-l.

Material EXAMINED. — Philippines. Musorstom 2: stn 33. 10: 2 (MNHN), 8 (USNM 81892).

MUSORSTOM 3: stn 87. 1 (USNM 97382). — Stn 101, 1 (MNHN). — Stn 102, 7 (MNHN). — Stn 1 17, 1 (MNHN). —

Stn 131. 1 (MNHN). — Stn 140, 1 (USNM 97394).

Indonesia. Karubar: stn 2, 4 (1 octameral, 1 nonameral, 2 decameral) (POLIPI). — Stn 3, 6 (3 octameral and

3 decameral) (MNHN). — Stn 7, 2 (1 octameral, 1 decameral) (MNHN). — Stn 15, 24 (16 octameral. 8 decameral) (USNM

97396). — Stn 22, 2 (hexameral)(MNHN). — Stn 28. 2 (decameral) (POLIPI). — Stn 29, I (decameral) (USNM 97398).

Japan. "Tansei Maru": stn KT93-09-AM7, 3 (USNM 93159).

TYPE Locality. — Adventure Bank, Mediterranean, 168 m.

DIAGNOSIS. — Corallum cylindrical and quite small, rarely over 7 mm in length and 1.0- 1.3 mm in diameter,

straight to serpentine. Some coralla firmly attached along one side of the corallum, others attached at random

points of their theca to sand grains, pebbles, or foraminifera. One specimen (MUSORSTOM 3 stn 140) was laterally

attached to a dead specimen of Heteropsammia. Epitheca bears 2 sets of ridges: transverse, circumferential ridges,

and 16, 18, or 20 longitudinal costal ridges, which form a grid-like pattern of rectangles. Within each rectangle is a

chalky white thecal spot. Septa usually octamerally arranged in 2 cycles, the Si much thicker than the S2; among

the Karubar specimens there was an almost equal division between the typical octameral form (having 16 septa)

and decameral specimens (having 20 septa), and one specimen having 18 septa. These septal symmetries are

indicated in the Material Examined section. Furthermore, some small specimens have a hexameral symmetry. All

septa have highly sinuous inner edges. Columella a single twisted or trefoil ribbon.

Remarks. — This minute, cryptic species has one of the smallest calicular diameters of all scleractinian

corals. It is more fully described and illustrated by Zibrowius (1980), Cairns (1989a), and Cairns & Parker

(1992).

Distribution. — Philippines : Verde Island Passage; Sulu Sea (Mindoro and Panay); Sibuyan Sea; 97-194 m.

Indonesia: Halmahera Sea; Banda Sea (Kai Islands); 141-282 m. Elsewhere: nearly cosmopolitan in tropical and

warm temperate regions, including Amami o Shima, Ryukyu Islands (reported herein); not yet known from eastern

Pacific; 28-653 m.

Family FLABELLIDAE Bourne, 1905

Genus FLABELLUM Lesson, 1831

Subgenus FLABELLUM (FLABELLUM) Lesson, 1831

Flabellum (F.) pavoninum Lesson, 1831

Fig. 20 h

Flabellum pavoninum Lesson, 1831: 2. — Cairns, 1989a: 46-50, pi. 23, figs g-I, pi. 24, figs a-d,

1989b: 67; 1994: 70-71, pi. 30, figs g-i, pi. 31, figs a-e. — Cairns & Keller, 1993: 263.

g-h (synonymy);

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

151

Flabellum coalitum Marenzeller, 1888: 48-49. — Cairns, 1989a: 46, 50, pi. 24, figs e-f, i-1.

Flabellum sp. - CAIRNS, 1989a: pi. 24, figs e-f.

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn 131, 19: 8 (MNHN), 11 (USNM 97455).

Indonesia. CORINDON 2: stn 216, 18: 17 (MNHN), 1 (USNM 97456).

Karubar: stn 1, 9 (MNHN). — Stn 30, 5 (POL1PI).

Type Locality. — Hawaiian Islands (depth not given).

DIAGNOSIS. — The specimens reported herein are all relatively small coralla of the coalitum form, the largest

(CORINDON 2 stn 216) only 16.4 x 29.9 mm in calicular diameter and 29.2 mm in height. Angle of thecal edges

ranges from 7 1 °- 1 1 3°; angle of thecal faces, 33°-43°. Thecal faces slightly convex and coarse in texture, meeting in

acute thecal edges that often bear low, discontinuous crests. Calicular edge smooth. Theca of well-preserved

specimens reddish-brown, often more intensely striped along the Ci-3, as in F. politum Cairns, 1989.

Pedicel elongate and slender (1.1-1. 5 mm in diameter), having 6 protosepta in the basal disc. Septa

hexamerally arranged in 6 cycles: Si-3>S4>Ss>S6. Lower, inner edges of Si -3 moderately sinuous, whereas their

upper, outer margins (adjacent to theca) are notched, rising above calicular edge as exsert lobes. Columella

rudimentary.

Remarks. — Small specimens of F. pavoninum forma coalitum are similar to adult F. politum in many

characters, but can be distinguished by having coarse, slightly convex thecal faces; less acute thecal edges; and less

sinuous inner edges of the S1-3. F. pavoninum is more fully described by Cairns (1989a, 1994).

DISTRIBUTION. — Philippines : Sulu Sea (Panay); 120-122 m. Indonesia: Makassar Strait; Banda Sea

(Kai Islands); 96-156 m. Elsewhere: widespread throughout Indo-West Pacific from southwest Indian Ocean to

Hawaiian Islands (Cairns & Keller, 1993); 98-665 m. Forma coalitum is common off Japan (Honshu,

Kyushu); 73-658 m.

Flabellum (F.) magnificum Marenzeller, 1904

Flabellum magnificum Marenzeller, 1904a: 276-277, pi. 17, fig. 13. — Cairns, 1989a: 50-51, pi. 25, figs a-j

(synonymy); 1994: 72, pi. 31, figs j-1.

Flabellum pavoninum - Faustino, 1927: 46 (in part). [Not Flabellum pavoninum Lesson, 1831].

Flabellum pavoninum magnificum - Yabe & EGUCHI, 1942a: 89-90, pi. 5, figs la-c.

Material EXAMINED. — Philippines. Musorstom 2: stn 15, 1 (USNM 97476). — Stn 75, 4 (MNHN). —

Stn 78, 4 (MNHN).

Musorstom 3: stn 135, 1 (USNM 97477).

Indonesia. Deki: stn 3, 4 (NNM 22585). — Stn 12, 1 (NNM 22585). — Stn 41, 8 (NNM 22586). — Stn 42,

1 (NNM 22584).

KARUBAR: stn 12, 1 (MNHN). — Stn 36, 2 (MNHN). — Stn 59, 9 (MNHN). — Stn 67, 1 (USNM 97479). — Stn 69,

13 (USNM 97480). — Stn 70, 9: 7 (POLIP1), 2 (USNM 97481). — Stn 77, 1 1 (POLIPI). — Stn 85, 1 (POLIPI).

Type Locality. — "Valdivia" stn 199: 0°15.5'N, 98°04.8’E (western Sumatra), 470 m.

Diagnosis. — Angle of thecal edges 140°- 172°; angle of planar, slightly concave thecal faces, 44°-58°.

GCD:H = 1.29- 7.48- 1.71, all coralla being significantly wider than tall and having long thecal edges. Largest

specimen (KARUBAR stn 12) 57 x 90 mm in calicular diameter and 61 mm in height. Thecal faces meet in straight,

acute edges but only rarely are edge crests present. Calicular edge smooth. Coralla homogeneously reddish-brown

or white. Pedicel relatively short, 1.5-2. 3 mm in diameter, containing 6 protosepta at the basal disc. Septa

hexamerally arranged in 7 cycles: Si-4>Ss>S6>S7, only the larger coralla having a complete 7th cycle of

384 septa, and one (the largest specimen examined) also having one pair of S8- Si -4 quite narrow near calicular

edge but abruptly widening 10-12 mm lower in fossa, resulting in a very high septal concavity index. Lower, inner

edges of S1-4 highly sinuous. Columella as in F. lamellulosum.

Source : MNHN, Paris

152

S. D. CAIRNS & H. ZIBROWIUS

REMARKS. — Flabellum magnificum is one of the larger deep-water solitary scleractinians, second in size only

to F. impensum Squires, 1962 and perhaps certain gigantic Desmophyllum dianthus (Esper, 1794) and

Rhizotrochus typus H. Milne Edwards & Haime, 1848. It is easily confused with another large species,

F. lamellulosum Alcock, 1902, but can be distinguished by: its broad corallum (GCD:H = 1.48 average vs 1.05

average for F. lamellulosum)-, its long, uncrested thecal edges (LEL:H = 0.70 average vs 0.48 average for

F. lamellulosum)-, its sinuous Si -4 (SSI = 4.8 average vs 1.13 average for F. lamellulosum)-, and its wide Si -4

(SCI = 14.7 average vs 9.4 average for F. lamellulosum).

DISTRIBUTION. — Philippines-. Verde Island Passage; Sibuyan Sea; Bohol Strait; Suiu Archipelago; 29 1 -

486 m. Indonesia: Banda Sea (Kai Islands); Arafura Sea (south and southeast of Tanimbar Islands); 225-567 m.

Elsewhere: Malaysia (Celebes Sea off Sabah); west of Sumatra; Japan (Honshu and Shikoku); 307-700 m.

Flabellum (F.) patens Moseley, 1881

Fig. 20 i

Flabellum patens Moseley, 1881: 172 (in part: pi. 6, fig. 15). — Cairns, 1989a: 51-52, pi. 26, figs a-i (synonymy)-

1989b: 67; 1994: 71-72, pi. 31, figs g-i.

Flabellum australe - Alcock, 1902c: 30-31. [Not Flabellum australe Moseley, 1881].

Material EXAMINED. — Philippines. Musorstom 3: stn 126, 1 (MNHN),

Indonesia: "Siboga": stn 251, 1 (ZMA).

"Hakuho Maru": stn KH72-1-28, 2: 1 (USNM 97469), 1 (ORI).

Karubar: stn 5, 2 (MNHN). — Stn 31, 1 (MNHN).

Type Locality. — "Challenger" stn 192: 5°49'S, 132°14'E (Kai Islands, Banda Sea), 256 m.

Diagnosis. — Corallum highly compressed, resulting in a very low angle of thecal faces (20°-24°), a very

high GCD:LCD (2.0-2.5), and a very narrow fossa. Angle of thecal edges 76°-155°; GCD:H rather low (average =

1.04), indicating corallum to be approximately as tall as wide. Largest specimen known ("Albatross" stn 5313)

28.9 x 67.5 mm in calicular diameter and 54.7 in height, but most specimens examined less than 45 mm in

GCD. Calicular edge smooth. Thecal edges prominently crested, usually as one continuous ridge, but occasionally

forming spurs as in F. lamellulosum. Corallum white to grey, the theca near calicular edge usually discoloured and

slightly corroded as a band 3-5 mm in width that parallels the calicular edge, an indication of an association with a

commensal Lumbrineris polychaete (Zibrowius, SOUTHWARD, & Day, 1975). Pedicel elongate and slender

(1. 5-2.0 mm in diameter), containing 6 protosepta at the basal disc. Septa hexamerally arranged in 6 cycles:

Si-4>Ss>S6, only the largest specimens (GCD > 40 mm) having some S7. Si -4 relatively narrow, their inner

edges almost in contact with those from opposite side across a narrow fossa; columella rudimentary.

Remarks. — Flabellum patens is compared to F. lamellulosum in the account of the latter species and is

more fully described and illustrated by Cairns (1989a, 1994).

Distribution. — Philippines: Verde Island Passage; Mindoro Strait; Sulu Sea (Sulu Archipelago); 266-

439 m. Indonesia: Banda Sea (Kai Islands); Timor Sea (south of Sermata Islands); 204-295 m. Elsewhere: Japan

(Honshu, Shikoku, and Kyushu); Formosa Strait; South China Sea north of Pratas Island; 223-402 m.

Flabellum (F.) lamellulosum Alcock, 1902

Fig. 21 a

Flabellum lamellulosum Alcock, 1902a: 105-106; 1902c: 30, pi. 4, figs 28, 28a-b. — Cairns, 1989a: 52-53 pi 27 figs

a-1 (synonymy); 1989b: 67. v ' ’ e

^Haime ' ALC°CK’ 1902c: 30 (in part: "Sibo8a" stn 12). [Not Flabellum distinctum H. Milne Edwards &

Flabellum pavoninum - Faustino, 1927: 45-46 (in part). [Not Flabellum pavoninum Lesson, 1831].

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

153

MATERIAL EXAMINED. — Philippines. "Hakuho Maru": stn KH72-1-52, 3 (USNM 97454).

Musorstom 1: stn 9, 1 (MNHN). — Stn 11, 2 (MNHN). — Stn 12, 5 (MNHN). — Stn 13, 1 (USNM 97419). —

Stn 24, 6 (USNM 97420). — Stn 25, 1 (MNHN). — Stn 27, 1 (USNM 97422). — Stn 31, 2 (MNHN). — Stn 40

1 (MNHN). — Stn 55, 1 (MNHN). — Stn 61, 3 (MNHN). — Stn 68, 2: 1 (MNHN), 1 (BMNH 1992.8.11.10). — Stn 7L

1 (MNHN).

Musorstom 2: stn 1, 1 (USNM 97426). — Stn 10, 3 (USNM 97427). — Stn 12, 3: 1 (MNHN), 2 (USNM 97428). —

Stn 18, 1 (USNM 97429). — Stn 19, 1 (USNM 97430). — Stn 21, 5: 4 (MNHN), 1 (USNM 97431). — Stn 26, 1 (USNM

97432). — Stn 63, 7: 2 (MNHN), 5 (USNM 97433). — Stn 64, 2: 1 (MNHN), 1 (USNM 97434). — Stn 66, 2 (MNHN) —

Stn 68, 2: 2 (MNHN). — Stn 75, 1 (USNM 97437). — Stn 83, 1 (MNHN).

MUSORSTOM 3: stn 86, 2 (USNM 97433). — Stn 87, 4 (MNHN). — Stn 88, 10 (MNHN). — Stn 91, 2 (MNHN). —

Stn 92, 7 (MNHN). — Stn 96, 19: 17 (MNHN), 2 (BMNH 1992.8.11.9). — Stn 97, 2: 1 (MNHN), 1 (USNM 97440). —

Stn 98, 4 (MNHN). — Stn 99, 4 (MNHN). — Stn 100, 2: 1 (MNHN), 1 (USNM 97441). — Stn 101, 2 (MNHN). —

Stn 102, 1 (USNM 97442). — Stn 108, 7 (USNM 97443). — Stn 109, 6 (USNM 97444). — Stn 1 1 1, 2 (USNM 97445)

— Stn 1 12, 2 (USNM 97446). — Stn 130, 1 (MNHN). — Stn 135, 2 (USNM 97447). — Stn 143, 10.

Indonesia. Deki: stn 3, 7 (NNM 22574). — Stn 41, 1 (NNM 22575). — Stn 50, 1 (NNM 22576). — Stn 52,

7 (NNM 22577). — Stn 63, 2 (NNM 22578).

"Hakuho Maru": stn KH72-1-28, 2 (ORI).

Karubar: stn 12, 1 (MNHN). — Stn 36, 1 (MNHN). — Stn 59, 1 (USNM 97449). — Stn 67, 4 (POLIPI). — Stn 69,

4 (POLIPI). — Stn 70, 3 (MNHN). — Stn 77, 2: 1 (MNHN), 1 (USNM 97453).

Type Locality. — "Siboga" stn 251: 5°28.4'S, 132°02'E (Kai Islands, Banda Sea), 204 m.

Diagnosis. — Angle of thecal edges 130°-240°; angle of planar thecal faces 32°-51°. GCD:H = 0.79-7.05-

1.29, most coralla being higher than wide and having relatively short thecal edges. Largest specimen known

(Karubar stn 70) 49 x 68 mm in calicular diameter and 67 mm in height. Calicular edges smooth. Thecal edges

prominently crested, the crests beginning almost immediately adjacent to pedicel, formed by successive eversions

(spurs) of the calice associated with the 2 principal septa and costae and subsequent retrenchment of the calice edge.

Thecal edges of large specimens may bear 6 or 7 such spurs, some up to 8 mm in height. Well-preserved coralla

homogeneously reddish-brown or white with reddish-brown stripes corresponding to C1-4. Pedicel short and 1.5-

2.1 mm in diameter, containing 6 protosepta in the basal disc. Septa hexamerally arranged in 7 cycles, the last

cycle usually incomplete: Si-4>Ss>S6>S7. Septa relatively narrow, having thickened, straight, lower inner edges.

Columella a well-developed crescent of trabeculae 1.0- 1.5 mm in width.

Remarks. — A comparison of F. lamellulosum to F. magnificum Marenzeller, 1904, is given in the account

of the latter species. F. lamellulosum is also similar to F. patens Moseley, 1881, especially in having a high

corallum (GCD:H = 1.04-1.05 average for both species) and in having prominently crested thecal edges.

F. lamellulosum differs in having a much more open calice with a face angle of 32°-51° and a GCD:LCD of 1.43

(average), compared to 20°-24° and 2.25 (average) for F. patens. The more open fossa also accommodates a more

robust columella in F. lamellulosum. Furthermore, the corallum of F. lamellulosum is less dense and more fragile

than that of F. patens, and does not appear to have the lumbrinerid polychaete symbiont that is so common on the

theca of F. patens.

Petrarcid ascothoracidan galls are present in 2 coralla (MUSORSTOM 1 stn 71 and "Albatross" stn 5273),

evidenced as a characteristic columellar deformation (Fig. 21a).

This species is more fully described and illustrated by CAIRNS (1989a).

Distribution. — Philippines: Lubang Island and Verde Island Passage; Sulu Sea (Panay); Samar Sea; Bohol

Sea; 187-486 m. Indonesia: Banda Sea (Kai Islands); Arafura Sea (south of Tanimbar Islands); Timor Sea

(south of Leti Islands); Java Sea; 204-412 m. Elsewhere: South China Sea (Vanguard Bank, Spratly Islands);

265-286 m.

Flabellum (F.) politum Cairns, 1989

Flabellum politum Cairns, 1989a: 53-54, pi. 28, figs a-f (synonymy); 1989b: 67 ("species 1"); 1994: 73, pi. 32,

figs a-c.

154

S. D. CAIRNS & H. ZIBROWIUS

Material EXAMINED. — Philippines. Musorstom 2: stn 2, 3 (MNHN). — Stn 6, 4 (USNM 97459). — Stn 10,

3 (USNM 97460). — Stn 68, 2 (MNHN).

Musorstom 3: stn 86, I (MNHN). — Stn 88, 6 (USNM 97461). — Stn 96, 10 (MNHN). — Stn 98, 1 (USNM

97463).— Stn 100, 2: 1 (MNHN), 1 (USNM 97464). — Stn 102, 34 (USNM 97465). — Stn 107, 2 (MNHN). —

Stn 108, 10 (MNHN). — Stn 109, 1 (MNHN). — Stn 1 10, 1 (MNHN). — Stn 143, 6 (MNHN).

Indonesia. "Siboga": stn 204, 7 (ZMA Coel. 1233). — Stn 260, 50+ (ZMA Coel. 5435).

DEKi: stn 3, 4 (NNM 22579). — Stn 53, 3 (NNM 22580). — Stn 54, 4 (NNM 22581). — Stn 63, 1 (NNM 22582).

Karubar: stn 2, 2 (USNM 97466). — Stn 15, 2 (MNHN). — Stn 31. 1 (POLIPI). — Stn 65. 2 (MNHN).

Type Locality. — "Albatross" stn 5391: 12°13'15"N, 124°05'03"E (Samar Sea, Philippines), 216 m.

DIAGNOSIS. — Angle of thecal edges 90°-136°; angle of thecal faces 30°-45°. GCD:H = 0.98-1.30, indicating

that most specimens are slightly wider than high. Largest known specimen ("Albatross" stn 5392) 22.0 x

36.1 mm in calicular diameter and 29.7 mm in height. Calicular edges smooth. Thecal faces planar, smooth, and

porcellaneous, meeting in sharp, acute edges, which may bear a low thecal crest. Theca of well-preserved coralla

reddish-brown, with stripes of more intense pigmentation associated with C1-3. Pedicel elongate, 1. 0-1.4 mm in

cross section, and broken in all specimens examined, revealing the 6 protosepta. Septa hexamerally arranged in

6 complete cycles: Si-3>S4>Ss>S6 (192 septa). Lower, inner edges of S 1 -3 highly sinuous. Columella

rudimentary.

Remarks. — Flabellum politum is distinguished from other West Pacific species of Flabellum by having a

relatively small corallum; a smooth, lustrous theca; and S1-3 with highly sinuous inner edges. The species is more

fully described by Cairns (1989a).

DISTRIBUTION. — Philippines-. Lubang Island; Tablas Strait; Samar Sea; Ragay Gulf; Bohol Sea; Sulu Sea

(Sulu Archipelago); 40-280 m. Indonesia: Banda Sea (Kai Islands and southeastern Sulawesi); Arafura Sea

(southeast of Tanimbar Islands); 90-288 m. Elsewhere: South China Sea off Hong Kong; Japan (Korea Strait, East

China Sea, northern Ryukyu Islands); 70-402 m.

Subgenus FLABELLUM (ULOCYATHUS) Sars, 1851

Flabellum (U.) deludens Marenzeller, 1904

Flabellum deludens Marenzeller, 1904a: 269-272. pi. 17, figs 10, 10a. — Zibrowius & GRYGIER, 1985: 122, figs 16-17.

— Cairns, 1989a: 55-56, pi. 29, figs a-f (synonymy); 1994: 73, pi. 32, figs d-e.

Material EXAMINED. — Philippines. "Albatross": stn 5412, 1 (USNM 96653).

"Hakuho Maru": stn KH72-1-52, 7 (USNM 96666), 3 (ORI).

Musorstom 1: stn 5, 13 (MNHN). — Stn 9. 1 (MNHN). — Stn 10, 1 (USNM 96648). — Stn 11. 35 (USNM 96657).

— Stn 12. 2 (USNM 96924). — Stn 13, 2 (MNHN). — Stn 15, 1 (USNM 96646). — Stn 20, 5 (MNHN). — Stn 24,

12 (MNHN). — Stn 25, 14 (MNHN). — Stn 40, 4: 2 (MNHN), 2 (USNM 96925). — Stn 61, 3: 2 (MNHN), 1 (USNM

96922). — Stn 68, 3 (MNHN).

Musorstom 2: stn 1. 1 (MNHN). — Stn 10, 4 (USNM 96654). — Stn 11, 5: 1 (MNHN), 4 (USNM 96649). — Stn 12,

10 (MNHN). — Stn 13, 8 (MNHN). — Stn 15, 1 (USNM 96667). — Stn 20, 2 (USNM 96642). — Stn 21, 1 (MNHN). —

Stn 40, 7 (USNM 96647). — Stn 46, 2 (MNHN). — Stn 63, 7 (MNHN). — Stn 64, 19 (MNHN). — Stn 66, 83:

78 (MNHN), 5 (USNM 81925). — Stn 68, 6 (USNM 96640). — Stn 83, 1 (MNHN).

Musorstom 3: stn 87, 21: 20 (USNM 96662), 1 (MNHN). — Stn 88, 1 (MNHN). — Stn 91, 3 (MNHN). — Stn 92,

8 (USNM 96644), — Stn 96. 2 (MNHN). — Stn 97, 1 1 (USNM 97651). — Stn 98, 23 (MNHN). — Stn 99, 58 (USNM

96645). — Stn 100, 4 (USNM 96663). — Stn 101, 17 (MNHN). — Stn 102, 2 (MNHN). — Stn 103, 2: 1 (MNHN),

1 (USNM 96670). — Stn 108. 8 (USNM 96656). — Stn 109. 4: 1 (MNHN), 3 (USNM 96638). — Stn 1 1 1, 15 (MNHN)

— Stn 112, 6 (USNM 96659). — Stn 120, 55: 18 (MNHN), 37 (MNHN). — Stn 139, 2 (USNM 96926). — Stn 145 2~

1 (MNHN), 1 (USNM 96923).

Indonesia. DEKI: stn 46, 2 (NNM 22530). — Stn 49, 100 (NNM 22531).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

155

KARUBAR: stn 59, 1 (USNM 96669). — Stn 62, 8: 6 (MNHN), 2 (POLIPI). — Stn 63, 198: 33 (MNHN), 165 (USNM

96637). _ stn 65, 6 (USNM 96641). — Stn 76, 2 (USNM 96929). — Stn 79, 4 (POLIPI).

Type Locality. — "Valdivia" stns 185 and 203: west of Sumatra, 614-660 m.

DIAGNOSIS. — Corallum fragile. Angle of thecal edges 1 15°-150°; angle of thecal faces 64°-80°, resulting in a

very open corallum. Largest known specimen (Karubar stn 63) 39.9 x 53.7 mm in calicular diameter and

28.0 mm in height. Thecal edge crests low (rarely over 2 mm), thin, and continuous, extending from pedicel to

calice. Calicular edge deeply lacerate — a high (up to 6 mm) lancet corresponding to each S]-2 and adjacent pair of

S4 (or the 16 primary septa and adjacent tertiaries). Colour of corallum base white, but theca streaked with broad

reddish-brown stripes corresponding to each Ci-2, and narrower stripes corresponding to each C3. Upper, outer

Si -2 also pigmented. Septa usually hexamerally arranged in 5 cycles: Si-2>S3>S4»Ss, (he 5th cycle usually

incomplete (but see Remarks). Even in larger coralla the S5 are typically rudimentary, occurring in pairs flanking

the S3, not the Si -2, resulting in 72 septa. Columella elongate and rudimentary, formed by the fusion of lower,

inner edges of Si -2.

Remarks. — Cairns (1989a) divided the subgenus Flabellum (Ulocyathus) into 3 groups based on corallum

shape: laterally compressed, bowl-shaped, and constricted. Four similar species belonging to the laterally

compressed group are reported herein: F. deludens, F. marenzelleri, F. japonicum, and F. hoffmeisteri. In addition

to being laterally compressed, these 4 species also have reddish-brown striped theca; pigmented upper outer

margins of the major septa; small edge crests; and small pedicels. Their differences are summarized in Table 2, the

most useful discriminating characters being the nature of the calicular edge, septal symmetry, and the general shape

of the corallum. Among the 4 species, F. deludens has the largest edge angle and the most exsert septal lancets.

Table 2. — Comparison of the four laterally compressed Flabellum (Ulocyathus) known from the Philippine/Indonesian

region.

The Philippine and Indonesian specimens identified as F. deludens differ from a syntype of that species (ZMB)

and several other Indian Ocean specimens (i.e., "Marion Dufresne" cruise 27, station 4, CP6, west of Ceylon,

1035 m, USNM 82013) in having a smaller pedicel that is elongate, not circular, in cross section. The pedicel

dimensions of the Indian Ocean specimens average 2.34 x 2.46 mm (almost circular), whereas the pedicels of

coralla from the western Pacific are smaller and more elongate in cross section, averaging 1.28 x 2.14 mm

(although the pedicel of at least one specimen from Karubar stn 76 measures 2.15 x 2.40 mm, consistent with

Source :

156

S. D. CAIRNS & H. ZIBROWIUS

the Indian Ocean specimens). In both cases there are 12 protosepta at the basal disc. The pedicel size and shape

appears to be the only obvious morphological difference between specimens from the 2 regions, and, although this

character may distinguish 2 different species or subspecies, it is interpreted herein by the first author to reflect a

difference in substrate size availability. The second author disagrees with this interpretation, maintaining that the

western Pacific specimens are a separate species different from F. deludens. Support for that hypothesis is

strengthened by the fact that the western Pacific specimens also differ from the Indian Ocean specimens in having a

shallower depth range: mostly less than 350 m vs mostly deeper than 500 m for those from the Indian Ocean.

Cairns (1989a) reported one lot of F. deludens with octameral symmetry, but most of the specimens from the

Karubar stations reported herein have octameral (actually decahexameral) symmetry, consisting of 16 primary,

16 secondary, 32 tertiary, and 32 quaternary septa. The quaternaries occur in pairs flanking the secondary septa, not

the primary, resulting in 96 septa in large coralla. In these specimens the 4 calicular lancets adjacent to the princi¬

pal septa are smaller that the others. These decahexameral specimens are otherwise identical to typical F. deludens.

Flabellum deludens sensu CAIRNS is more fully described and illustrated by CAIRNS (1989a, 1994).

DISTRIBUTION. — Philippines : common throughout Philippines from Lubang Island to Moro Gulf; 187-

480 m, although most records are from less than 350 m. Indonesia: Banda Sea (Kai Islands); Arafura Sea (south of

Tanimbar Islands); 176-400 m. Elsewhere: Japan (Honshu, Shikoku, Kyushu); South China Sea (Spratly Islands);

west of Sumatra; northern Indian Ocean; 106-1035 m (but see Remarks).

Flabellum (U.) marenzelleri Cairns, 1989

Flabellum (U.) marenzelleri Cairns, 1989a: 57-58, pi. 30, figs a-e (synonymy).

Material EXAMINED. — Philippines. "Albatross": stn 5523, 1 (USNM 97410).

Musorstom 1: stn 40, 1 (MNHN).

Indonesia. Deki: stn 52, 1 (NNM 22757).

"Hakuho Maru": stn KH72-1-28, 3 (USNM 97407). — Stn KH85-1-A1, 3: 1 (USNM 97409), 2 (ORI). — Stn KH85-1-

A2, 1 (USNM 97408).

Karubar: stn 2, 4 (MNHN). — Stn 35, 2 (USNM 97406). — Stn 36, 2 (MNHN).

TYPE Locality. — "Albatross" stn 5289: 13°4r50"N, 120°58'30"E (Verde Island Passage, Philippines),

315 m.

DIAGNOSIS. — Angle of thecal edges 84°-l 12°; angle of planar thecal faces 39°-52°. Largest known specimen

(Karubar stn 35) 26.5 x 46.0 mm in calicular diameter and 28.1 mm in height. Thecal edge crests low (rarely

over 2 mm), thin, and continuous, extending from pedicel to calice. Calicular edge lacerate, a moderately high (up

to 4 mm) lancet corresponding to the 16 primary septa and their adjacent pairs of tertiary septa. Theca coarsely

granular. Theca purple-brown, with more intensely pigmented stripes corresponding to the 16 primary costae.

Pedicel elliptical to elongate in cross section (2. 1-2.4 in greater diameter), short, containing 12 protosepta at the

basal disc. Septa arranged in 4 size classes, larger coralla having 96 septa arranged accordingly: 16:16:32:32, as in

octamerally symmetrical F. deludens. Columella elongate and rudimentary.

Remarks. — Flabellum marenzelleri is similar to F. deludens, but can be distinguished (Table 2) by having a

denser and more compressed corallum, which is characterised by a lower edge angle and a lower face angle,

resulting in a higher GCD:LCD (1.65-1.75 vs 1.30-1.40 for F. deludens). F. marenzelleri also consistently has

a decahexameral symmetry, whereas only some specimens of F. deludens share this symmetry. F. marenzelleri

also has equal-sized, less exsert calicular lancets, those of F. deludens being dimorphic in size (alternately smaller

and larger) in octameral specimens and much more exsert.

Flabellum marenzelleri is more fully described and illustrated by CAIRNS (1989a).

Distribution. — Philippines: Verde Island Passage; Bohol Sea; 247-315 m. Indonesia: Banda Sea (Kai

Islands); Timor Sea (south of Leti Islands); Flores Sea (Sulawesi); 240-390 m.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTIN1A

157

Flabellum (U.) japonicum Moseley, 1881

Flabellum japonicum Moseley, 1881: 168-169, pi. 7, figs 4, 4a, pi. 16, fig. 12. — ALCOCK, 1902c: 32-33 (in part:

"Siboga" stn 17). — Faustino, 1927: 47-48, pi. 2, figs 5-6. — Cairns, 1989a: 56-57, pi. 29, figs g-i (synonymy);

1994: 73-74, pi. 32, figs g-h (synonymy).

MATERIAL EXAMINED. — Philippines. "Siboga": stn 95, 1 (ZMA).

"Galathea stn 436, 22 (ZMUC).

MUSORSTOM 2: stn 36, 2 (USNM 97402). — Stn 40, 1 (MNHN). — Stn 44, 2 (MNHN). — Stn 49, 10 (MNHN).

MUSORSTOM 3: stn 95, 1 (MNHN). — Stn 122, 15 (USNM 97403). — Stn 123, 10: 5 (MNHN), 5 (USNM 97404). —

Stn 128, 2: 1 (MNHN), 1 (USNM 97405).

TYPE Locality. — "Challenger" stn 232: 35° 1 l'N, 139°28'E (Sagami Bay, Japan), 631 m.

DIAGNOSIS. — Angle of thecal edges 90°-108°; angle of thecal faces 65°-88°. Largest known specimen

(MUSORSTOM 3 stn 123) 47 x 62 mm in calicular diameter and 33 mm in height. Corallum very fragile,

campanulate, and laterally compressed, having convex thecal faces that meet in sharp, crested thecal edges. Edge

crests low, usually not extending to calice. Calicular edge serrate, a small (up to 3.5 mm) equilaterally triangular

apex corresponding to each Si-2. Upper part of corallum with reddish-brown stripes corresponding to C1-2; lower

part often discoloured (superficially eroded). Pedicel circular to slightly elliptical in cross section (occasionally

elongate), 2. 1-2.7 mm in greater diameter, short, containing 12 protosepta at the basal disc. Septa hexamerally

arranged in 5 cycles: Si-2>S3>S4»Ss, a full 5th cycle present only in large coralla.

REMARKS. — Among the laterally compressed Flabellum ( Ulocyathus ), F. japonicum is very similar to

F. deludens (Table 2), but can be distinguished by having: smaller, triangular calicular apices resulting in a serrate

(not lacerate) calicular edge; less developed edge crests that do not extend to the calice edge; more convex thecal

faces; a more open calice characterised by a higher face angle; and a lessened thecal pigmentation, often discoloured

basally. F. japonicum is more fully described and illustrated by CAIRNS (1989a, 1994).

DISTRIBUTION. — Philippines : Lubang Island and Verde Island Passage; Tablas Strait; Sibuyan Sea; Bohol

Strait and Sea; Sulu Sea (Sulu Archipelago); 425-865 m. Indonesia : Bali Sea; 1060 m. Elsewhere: Japan (Honshu,

Shikoku, and Kyushu); 119-1141 m.

Flabellum (U.) hoffmeisteri Cairns & Parker, 1992

Flabellum japonicum - ALCOCK, 1902c: 32-33 (in part: "Siboga" stn 212). — HOFFMEISTER, 1933: 7, pi. 1, figs 1-2. [Not

Flabellum japonicum Moseley, 1881].

Flabellum n. sp. - Cairns, 1989a: 57 (in part: pi. 29, figs j-k).

Flabellum (U.) hoffmeisteri Cairns & Parker, 1992: 47-48, pi. 16, figs d-f (synonymy). — Cairns, 1995: 103-104,

pi. 33, figs g-h.

Material EXAMINED. — Indonesia. "Siboga": stn 212, 1 (ZMA).

Deki: stn 56, 3 (NNM 22541).

Karubar: stn 9, 1 (USNM 97411). — Stn 10. 13 (MNHN). — Stn 12, 83 (USNM 97412). — Stn 13, 1 (MNHN). —

Stn 35, 1 (POLIPI). — Stn 39, 2 (USNM 97413). — Stn 40, 16 (MNHN). — Stn 59, 76 (MNHN). — Stn 69, 19 (MNHN).

— Stn 70, 6 (MNHN). — Stn 71, 1 (USNM 97415). — Stn 75, 2 (POLIPI). — Stn 77, 23 (MNHN).

Type Locality. — "Soela" stn 27: 37°59'S, 150°05'E, Victoria, Australia, 452 m.

Diagnosis. — Angle of thecal edges variable, 63°-l 12°; angle of thecal faces 50°-67°. Largest known specimen

(Karubar stn 12) 41.5 x 62.5 mm in calicular diameter and 46.1 mm in height. Corallum campanulate, relatively

robust, and laterally compressed, having slightly convex thecal faces meeting in sharp thecal edges. Edge crests

absent or very low, in the latter case not extending to calicular edge. Calicular edges moderately serrate, a low (less

than 2 mm), triangular apex corresponding to the 16 primary septa. Upper corallum bears faint reddish-brown

Source :

158

S. D. CAIRNS & H. ZIBROWIUS

stripes associated with primary septa; lower part often discoloured and encrusted by other organisms

(i.e. , brachiopods). Pedicel elliptical in cross section (2. 3-2. 7 mm in greater diameter), short, containing

12 protosepta at the basal disc. Septa arranged in 4 size classes: 16 primary, 16 secondary, 32 tertiary, and a

variable number of quaternary septa. Pairs of quaternary septa first form adjacent to secondary septa and only in

large coralla do they occur adjacent to primary septa. Septa of a 5th size class occur in end-sectors of largest

coralla. Thus, coralla may have 64 septa (no quaternaries), 96 septa (1/2 of quaternaries present), 128 septa (a full

4th size class), or over 128 septa, if some septa of the 5th size class are present.

REMARKS. — Among the laterally compressed species of this subgenus, F. hoffmeisteri is similar to

F. japonicum but is distinguished by its decahexameral (x 1 6) septal symmetry; its moderately serrate calicular

edge and its lower face angle (angle between thecal faces) (Table 2). It differs from the other decahexamerally

symmetrical species, F. marenzelleri and some F. deludens, in having a much less jagged calicular edge, convex

thecal faces, virtually no edge crests, and a larger face angle.

DISTRIBUTION. — Indonesia: Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar); Flores Sea (Selayar

Island); 345-477 m. Elsewhere: Victoria and Tasmania; Kermadec and Colville Ridges; 1 10-646 m.

Flabellum (U.) messum Alcock, 1902

Flabellum laciniatum var. messum Alcock, 1902c: 31.

Flabellum (U.) messum - Cairns, 1989a: 58-59, pi. 30, figs f-i, k (synonymy); 1995: 101-102, pi. 33, figs a-c. —

Cairns & Keller, 1993: 263, pi. 10, figs G-H.

Material EXAMINED. — Indonesia. Karubar: stn 56, 8 (MNHN). — Stn 57, 14: 6 (MNHN), 8 (USNM 97474).

— Stn 72, 1 (POLIPI). — Stn 91, 2 (USNM 97475).

Type Locality. — "Siboga" stns 45, 284, and 314: Indonesia, 694-828 m.

Diagnosis. — Corallum highly compressed and constricted medially. Angle of straight, crested thecal edges

131°-210°; angle of concave thecal faces 36°-44°. Largest known specimen (Karubar stn 57) 31 x 56 mm in

calicular diameter and 42 mm in height. Thecal edge crests extend from pedicel to calicular edge. Calicular edge

highly lacerate, but rarely well preserved because of extreme fragility. Pedicel circular to slightly elliptical in cross

section, up to 3.4 mm in greater diameter, short, and containing 12 protosepta at the basal disc. Thecal faces rough

in texture. Corallum reddish-brown, except for pedicel and edge crests, which are white. Septa hexamerally arranged

in 5 cycles: Si-3>S4>Ss (96 septa). Fossa deep and narrow.

Remarks. — Flabellum messum is unique in this region within the subgenus in having a "constricted"

corallum (Cairns, 1989a). It is further distinguished by is reddish-brown theca (no stripes); short, stout pedicel;

and roughly textured theca. Flabellum messum is more fully described and illustrated by CAIRNS (1989a, 1995).

Distribution. — Philippines: Verde Island Passage; 368 m. Indonesia: Celebes Sea (south of Basilan);

Arafura Sea (southeast of Tanimbar Islands); Timor Sea (southeast of Timor); eastern Java Sea; 476-949 m.

Elsewhere: Malaysia (Darvel Bay, Celebes Sea); Mascarene Plateau, southwest Indian Ocean; Kermadec Ridge;

430-1035 m.

Flabellum (U.) sp.

Figs 21 d-f

Material EXAMINED. — Philippines. MUSORSTOM 2: stn 78, 1 (MNHN).

New Zealand Region: "Tangaroa": stn G3, 1 (USNM 94329). — Stn U582, 3 (USNM 94330).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

159

DESCRIPTION. — The single Philippine specimen measures 24.9 x 32.0 mm in calicular diameter, 21.8 mm in

height, and is curved 45° in plane of LCD. Its edge angle is 48° and its face angle is also approximately 48°, but

the lower 6 mm of the corallum has a much higher edge angle and lower face angle than the upper part. Thecal

faces convex, in lower half of corallum, meeting in an acute angle at thecal edges; in upper half thecal edges

rounded. Calicular edge lacerate, each Si -2 and adjacent pair of S4 forming a lancet about 3 mm in height. Pedicel

circular, 1.6- 1.7 mm in diameter, containing 6 protosepta at basal disc. Corallum white with very faint reddish-

brown stripes associated with C1-2 and upper, outer edges of S 1.2.

Septa hexamerally arranged in 5 cycles, the 5th incomplete: Si-2>S3>S4>Ss; pairs of S5 occur only adjacent to

S3 (not Si -2), resulting in 72 septa. Inner septal edges sinuous, the lower edges of Si -2 contributing to an elongate

columella.

A well-preserved specimen from the New Zealand region (“ Tangaroa " stn G3) is larger (29.8 x 33.6 mm in

calicular diameter and 31.1 mm in height), curved 90°, and has more septa, i.e., 90. It is otherwise similar to the

Philippine specimen.

Remarks. — The specimens reported above are similar to F. moseleyi Pourtales, 1880, which is the only

other species in the subgenus known to have a secondarily free, curved corallum. It differs in having fewer septa at

a corresponding GCD (the 5th cycle of F. moseleyi is complete at a GCD of about 30 mm) and in its method of

septal insertion, i.e., S4 pairs insert adjacent to S3 before adjacent to Si -2. F. moseleyi also has much more

prominent (up to 6 mm) and more slender, triangular calicular lancets. F. moseleyi is known only from the

Caribbean and eastern Gulf of Mexico, depth 216-1097 m (C.AIRNS, 1979). Although the specimens reported above

may represent an undescribed species, not enough specimens are available to properly describe it or definitively

distinguish it from F. moseleyi.

DISTRIBUTION. — Philippines: Verde Island Passage; 441-550 m. Elsewhere : Norfolk Ridge between Norfolk

Island and New Caledonia; Three Kings Ridge; 710-1058 m.

Flabellum (U.) sexcostatum Cairns, 1989

Flabellum (U.) sexcostatum Cairns, 1989a: 59, pi. 30, fig. j, pi. 31, figs a-b.

Material EXAMINED. — Philippines. Musorstom 1: stn 47, 4: 2 (MNHN), 2 (USNM 97470).

Type Locality. — "Albatross" stn 5284: 13°42’05"N, 120°30'45"E (Verde Island Passage, Philippines),

772 m.

DIAGNOSIS. — Corallum laterally compressed, the slightly convex thecal faces meeting at acute, carinate

edges; edge crests small and present only on lower half of corallum. Angle of thecal edges changes from 90°-130°

to 55°-75° 12-15 mm above the base; angle of thecal faces changes from 61°-73° to 29°-36°. Also at this height the

4 lateral Ci are prominently ridged. Largest known specimen (MUSORSTOM 1 stn 47) 32.3 x 50.7 mm in calicular

diameter and 42.2 mm in height. Calicular edges serrate, each Si -2 producing a tall (up to 4 mm) triangular lancet.

Pedicel elliptical in cross section (2.0-2. 1 mm in greater diameter), short, containing 12 protosepta at the basal

disc. Septa hexamerally arranged in 5 cycles: Si-2>S3>S4>Ss (96 septa).

Remarks. — This is the 2nd report of F. sexcostatum, the MUSORSTOM specimens being collected close to

the type locality.

Flabellum sexcostatum was not placed by CAIRNS (1989a) in a species group within the subgenus Flabellum

(Ulocyathus) because it is intermediate in shape between the laterally compressed and bowl-shaped groups. The

species is more fully described and illustrated by Cairns (1989a).

Distribution. — Philippines: known from only 12 specimens from the Verde Island Passage Luzon; 685-

772 m.

Source :

160

S. D. CAIRNS & H. ZIBROWIUS

Flabellum (U.) conuis Moseley, 1881

Figs 21 b-c

Flabellum conuis Moseley, 1881: 165-166, pi. 7, figs 6a-b.

Flabellum (U.) conuis - Cairns, 1989a: 59-60, pi. 31, figs c-g.

MATERIAL EXAMINED. — Philippines. "Hakuho Maru": stn KH72-1-8, 3 (USNM 97472).

Estase 2: stn 6, 1 (MNHN).

Indonesia. " Galathea stn 489, 2 (ZMUC).

Japan. "Tansei Maru": stn KT86-16 F, 1 (USNM 97473).

Type Locality. — "Challenger" stn 218: 2°33'S, 144°04'E (Admiralty Islands), 1994 m.

Diagnosis. — Angle of thecal edges 45°-65°; angle of thecal faces 35°-50°. Corallum campanulate. Largest

known specimen (" Hakuho Maru" stn KH72-1-8) 43.8 x 47.6 mm in calicular diameter and 37.3 mm in height.

Calice slightly elliptical: GCD:LCD = 1.05-1.30. Thecal edges and faces convex. No edge crests, but the 6 Ci are

slightly ridged on lower half of corallum. Calicular edge serrate, a short (up to 3.5 mm) equilaterally triangular

apex corresponding to each Si-2, but generally damaged because of high fragility. Pedicel circular in cross section

(2. 1-2.3 mm in diameter), short, containing 12 protosepta at the basal disc. Theca smooth, light grey in colour.

Septa hexamerally arranged in 5 cycles, the 5th cycle rudimentary and sometimes missing: Si-2>S3>S4»Ss.

Fossa deep and narrow.

Remarks. — Flabellum conuis is unique in this region within the subgenus in having a bowl-shaped or

campanulate corallum (Cairns, 1989a) and an almost circular corallum. Its deep range also distinguishes it from

other Flabellum in the region. It is more fully described by Cairns (1989a).

DISTRIBUTION. — Philippines: Sulu Sea (Palawan); Sulu Archipelago (Sibuto Passage); 2021-2570 m.

Indonesia: Bali Sea; 1 160 m. Elsewhere: Admiralty Islands; Bungo Strait, Japan; 1994-2603 m.

Genus POLYMYCES Cairns, 1979

Polymyces wellsi Cairns, 1991

Polymyces wellsi Cairns, 1991: 22, pi. 8, figs f, i, pi. 9, figs a-b; 1995: 108-109, pi. 35, figs d-f.

Material EXAMINED. — Philippines. Musorstom 3: stn 94, 1 (USNM 97484).

Indonesia. DEKI: stn 59, 22 (NNM 22721).

Karubar: stn 13, 3 (MNHN).

Type Locality. — "Johnson-Sea-Link" stn 1916: 1°18.7'S, 89°48.8’W (Espanola, Galapagos), 545-562 m.

Diagnosis. — Corallum elongate-conical, straight, with a slightly flared calice; corallum fragile. Philippine

specimen 14 mm in GCD and 45 mm in height. Calicular edge jagged, a high (up to 5 mm) triangular lancet

corresponding to each Si-2. Pedicel thickened by asymmetrical development of 4 contiguous, hollow rootlets,

2 flanking each side of the 2 principal septa. These rootlets grow downward, completely encircle the base, and

fuse, forming a V-shaped junction near the base of opposite thecal side. Theca smooth, streaked with reddish-brown

pigment corresponding to the Ci-2, although rootlets and pedicel remain white. Septa hexamerally arranged in

5 complete cycles: Si-2>S3>S4>S5-

Remarks. — The distinctive asymmetrically placed rootlets distinguish P. wellsi from all other flabellids in

the region. It is more fully described by Cairns (1991, 1995).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTIN1A

161

DISTRIBUTION. — Philippines: Lubang Island; 842 m. Indonesia: Banda Sea (Kai Islands); 385-417 m.

Elsewhere: Galapagos; northeastern New Zealand; Kermadec Islands; 355-1165 m.

Genus RHIZOTROCHUS H. Milne Edwards & Haime, 1848

Rhizolrochus typus H. Milne Edwards & Haime, 1848

Figs 22 d-e

Rhizolrochus typus H. Milne Edwards & Haime, 1848a: 282, pi. 8, fig. 16. — Cairns, 1989a: 79-81, pi. 41, figs f-j

(synonymy); 1994: 81, pi. 35, figs a-c, pi. 40, figs h-i (synonymy).

MATERIAL EXAMINED. — Philippines. Musorstom 3: stn 131, 9: 5 (MNHN), 4 (USNM 97486).

Indonesia. "Siboga": stn 260, 1 (ZMA).

Deki: stn 5, 2 (NNM 22426). — Stn 24, 4 (NNM 22420, 22422). — Stn 25, 1 (NNM 22423, 23096). — Sin 26,

2 (NNM 22419)'. — Stn 27, 3 (NNM 22427). — Stn 53, 2 (NNM 22428). — Stn 54, 1 (NNM 22421).

SNELLIUS 2: stn 4.106, 1 (NNM 22424). — Stn 4.115, 2 (NNM 22425).

"Cable Ship Telegraaf: northern coast of Sumatra (Segli), 549 m, 1 (ZMA).

South China Sea. Macclesfield Bank (cf. Bassett-Smith, 1890), 73-92 m, "dredge 22", 1 (BMNH 1893.9.1.213).

TYPE LOCALITY. — Singapore, South China Sea (depth not given).

Diagnosis. — Corallum conical (turbinate); calice elliptical (GCD:LCD = 1.20-1.45); calicular margin

smooth. Largest Philippine specimen ("Albatross" stn 5357) 40.1 x 57.2 mm in calicular diameter and 38.3 mm

in height. Pedicel narrow (1.0- 1.5 mm in diameter) and not reinforced; however, several cycles of discrete, hollow

rootlets (rootlet diameter 1. 0-2.5 mm), extend from lower corallum to substrate, firmly anchoring the corallum.

Corallum white. Septa hexamerally arranged in 6 cycles, the 6th complete only in large specimens:

Si-2>S3>S4>S5>S6. Upper, outer margin (near calice) of Si-3 quite narrow, but upper axial edge of same septa

project as broad lamellae into fossa.

Remarks. — Rhizotrochus typus is easily distinguished from all other flabellids in the Philippine/Indonesian

region by having numerous, discrete (i.e., free standing, not contiguous with corallum) rootlets. One large

specimen at the BMNH (1950.1.11.630) from Mauritius measures 108 mm in GCD. The species is more fully

described and figured by CAIRNS (1989a).

One unusual specimen of Rhizotrochus from DEKI stn 25 (NNM 23096) deserves special note. It is 13.3 x

19.1 mm in calicular diameter, 1 1.5 mm in height, and has 96 septa. It differs from other Rhizotrochus in having

a truncated base with a basal scar of 5.2 x 8.5 mm in diameter, which suggest that transverse division occurred. It

also has a polygonal calicular cross section, each C| being slightly ridged, rootlets occurring only in series on the

Ci. The presence of transverse division in Rhizotrochus is unexpected, since the upper part of the corallum is held

stationary by numerous rootlets, which should not even allow division to occur. Nonetheless, one specimen is

known with this character combination (Figs 22 d-e).

Distribution. — Philippines: Mindoro Strait; Sulu Sea (Balabac); 120-124 m. Indonesia: Banda Sea (Kai

Islands); Flores Sea (Lintah Strait); 70-296 m. Elsewhere: South China Sea (Macclesfield Bank); Malaysia (Darvel

Bay, Celebes Sea); Red Sea; Persian Gulf; Bay of Bengal; Singapore; Pelau; Japan (Honshu and Kyushu);

20-1048 m.

"Rhizotrochus1' flabelliformis Cairns, 1989

Flabellum latum - ALCOCK, 1902c: 31. [Not Flabellum latum Studer, 1878],

Rhizotrochus flabelliformis Cairns, 1989a: 81, pi. 41, figs k-1, pi. 42, figs b, d; 1995: 109-110, pi. 35, figs g-t, p .

figs b, d.

Source :

162

S. D. CAIRNS & H. ZIBROWIUS

Material EXAMINED. — Indonesia. Deki: stn 48, 1 (NNM 22412). — Stn 56, 1 (NNM 22413). — Stn 59

6 (NNM 22414).

"Galathea": stn 500, 1 (ZMUC).

TYPE Locality. — "Siboga" stn 105: 6°08'N, 121°19'E (Sulu Archipelago, Philippines), 275 m.

Diagnosis. — Corallum highly laterally compressed (GCD:LCD = 2.4-3.4); calicular edge smooth. Largest

known specimen (Deki stn 59) 25 x 73 mm in calicular diameter. Pedicel quite narrow (1.2- 1.5 mm in diameter),

but attachment reinforced by 2 compressed, massive (4-5 mm in diameter) rootlets, one originating from each

calicular edge and firmly anchored to the substratum. Corallum light reddish-brown, young ones having a more

intense Ci-2 pigmentation. Septa hexamerally arranged in 6 to 7 cycles (Si-4>Ss>S6>S7), but even largest

specimens with incomplete 7th cycle. Fossa deep and narrow; columella rudimentary.

Remarks. — The second author disagrees with the first in placing "R." flabelliformis in this genus. He

considers that it is not a true Rhizotrochus , not being attached by circles of adventitious, cylindrical rootlets. The

general morphology is closer to that of a species of Flabellum (Flabellum), the anchoring pair of compressed

opposite thecal edge eversions, by position and formation, being similar to irregular spurs on edge crests of

Flabellum ( Flabellum ) and to spines on edge crests of Truncatoflabellum. The use of these eversions for

attachment is a qualitative leap with respect to edge crest formation in those two other taxa.

This species is more fully described and illustrated by Cairns (1989a, 1994).

Distribution. — Philippines-. Sulu Archipelago; 275 m. Indonesia : Banda Sea (Kai Islands); Arafura Sea (east

of Tanimbar Islands); 263-390 m. Elsewhere : New Zealand region; 228-419 m.

Genus GARDINERIA Vaughan, 1907

Gardineria philippinensis Cairns, 1989

Gardineria philippinensis Cairns, 1989a: 82, pi. 42, fig. a.

Material EXAMINED. — Philippines. MUSORSTOM 1: stn 63, 1 (MNHN).

Indonesia. Karubar: stn 86, 1 (USNM 97488).

TYPE Locality. — "Albatross" stn 5217: 13°20'N, 123°14T5"E (Sibuyan Sea, Philippines), 192 m.

Description. — Corallum conical (turbinate), with a basal angle of 38°-42° and a circular calice. Largest

known specimen (MUSORSTOM 1 stn 63) 17.5 mm in calicular diameter and 19.6 mm in height, with a robust

pedicel 7.1 mm in diameter (PD:GCD = 0.41). Corallum attached exclusively through the base of its pedicel.

Theca usually heavily encrusted with bryozoans, foraminifera, serpulid tubes, and sponges. Unencrusted theca

white, bearing fine horizontal epithecal striae. Calicular margin smooth, rising as a thin lip as much as 1.5 mm

above upper, outer septal edges.

Septa hexamerally arranged in 4 complete cycles (48 septa): Si>S2>S3>S4. Si thick, their inner edges vertical

and straight, extending to columella; their peripheral edges meet the theca below the calicular margin, their upper

edges rise slightly above calicular edge. S2 similar to Si in shape, but slightly less exsert and narrower, also

attaining the columella. S3 about 1/2 width of S2, each bearing a small paliform lobe that merges with the

columella. S4 rudimentary. Fossa of moderate depth, containing a well-developed columella consisting of

12-17 papillose elements.

Remarks. — The original description of G. philippinensis was based on a type series consisting of the dead,

poorly-preserved holotype; a juvenile corallum; and a damaged pedicel of a 3rd specimen. The 2 specimens reported

above are larger and better preserved, permitting the observation that the species attains 4 full cycles of septa and

that it attaches exclusively through the base of its pedicel, not laterally.

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

163

Gardineria philippinensis is similar to G. hawaiiensis Vaughan, 1907, in shape and size but differs in attaining

a full 4th cycle of septa and in having a robust columella. It differs from Gardineria sp. A (reported by Cairns,

1995 from the Chesterfield Islands, Lord Howe Seamount Chain, and Norfolk Ridge, depth 291-378 m) by its

robust columella.

DISTRIBUTION. — Philippines : Lubang Island; Ragay Gulf, Luzon; Iligan Bay, Mindanao; 192-494 m.

Indonesia: Arafura Sea (southeast of Tanimbar Islands); 222-226 m.

Gardineria paradoxa (Pourtales, 1868)

Figs 21 g-h

Haplophyllia paradoxa PourtaRs, 1868: 140-141.

Duncania barhadensis PourtaRs, 1874: 45, pi. 9, figs 5-7.

Gardineria barhadensis - LEWIS, 1965: 1063.

Gardineria paradoxa - CAIRNS, 1979: 160-161, pi. 31, figs 4-6, 10 (synonymy).

MATERIAL EXAMINED. — Indonesia. Karubar: stn 5, 2: 1 (MNHN), 1 (USNM 97489).

Type Locality. — "Bibb" stn 22: 24°14’20"N, 80°59'40"W (Straits of Florida), 692 m.

DESCRIPTION (larger specimen). — Corallum elongate-conical: 20.2 mm in length and 11.2 mm in calicular

diameter. Corallum attached to substratum by pedicel as well as theca on one side all along from pedicel to calice.

Theca exteriorly eroded and encrusted with bryozoans. Although collected alive, specimen resembles a fossilized

corallum. Theca dense, up to 1.5 mm thick, extending as a robust lip about 1.3 mm above upper, outer septal

edges.

Septa decamerally arranged in 3 size classes: 10:10:20 (40 septa). Primary septa nonexsert, having straight,

vertical inner edges that join the columella low in fossa. Secondary septa similar in shape, about 2/3 width of a

primary, each secondary bearing a discrete paliform lobe about 0.8 mm in width. Tertiary septa narrow, about

1/2 width of a secondary, having finely dentate inner edges and extending only about 6 mm from calicular edge.

Columella papillose, composed of 13 cylindrical (0.5 mm in diameter), granular elements.

The smaller of the 2 specimens is a juvenile only 5. 1 mm in calicular diameter and contains only 30 septa.

Remarks. — This is first report of G. paradoxa outside the western Atlantic, where it is known from the

Greater and Lesser Antilles (Cairns, 1979). Comparison of the Karubar specimens to those from the Antilles

shows no significant differences; a specimen from Barbados (Fig. 21 g) is particularly similar to the large

Karubar specimen.

Gardineria paradoxa is distinguished from the other species in the genus by having decameral septal symmetry

and a strong secondary lateral attachment.

Distribution. — Indonesia: Banda Sea (Kai Islands); 285-323 m. Elsewhere: western Atlantic (Antilles);

91-700 m.

Genus JAVANIA Duncan, 1876

Javania insignis Duncan, 1876

Javania insignis Duncan, 1876: 435, pi. 39, figs 11-13. — Zibrowius, 1974c: 8-9, pi. 1, figs 1-6. — Cairns. 1989a:

77-78, pi. 40, figs d-e, g-h, j-k (synonymy); 1994: 80, pi. 34, figs i-k. — Cairns & Keller, 1993: 272.

Flabellum weberi Alcock, 1902a: 107.

Material EXAMINED. — Philippines. Musorstom 2: stn 32, 1 (MNHN). — Sin 33, 19 (USNM 97490).

Musorstom 3: stn 131, 4 (USNM 97491).

Source :

164

S. D. CAIRNS & H. ZIBROWIUS

Indonesia. Deki: stn 24, 2 (NNM 22435).

CORINDON 2: stn 248, 2 (MNHN).

Sneluus 2: stn 4.100, 1 (NNM 22434).

Karubar: stn 22, 1 (MNHN). — Stn 32, 2 (POLIPI).

Type Locality. — 34°13'N, 136°13'E (Honshu, Japan), 88 m.

Diagnosis. — Corallum elongate-conical, straight, robust, and slightly flared distally; calice usually highly

elliptical (GCD:LCD = 1.3-1. 7). Pedicel thickened with concentric layers of dense stereome, its diameter up to

55% of GCD. Corallum rarely over 25 mm in GCD or 43 mm in height; white. Septa hexamerally arranged in

5 cycles according to formula: Sl-2>S3»S4>Ss. S5 begin to appear at a GCD of 9-10 mm and the cycle is

usually complete at a GCD of 15-17 mm. Si -2 highly exsert, producing a lacerate calicular margin, but

S4-5 nonexsert and much smaller than lower cycle septa.

Remarks. — This species is more fully described and illustrated by Cairns (1989a, 1994).

Distribution. — Philippines : Verde Island Passage; Sulu Sea (west of Panay and Balabac); Davao Gulf,

Mindanao; 122-192 m. Indonesia : Makassar Strait; Banda Sea (Kai Islands); Flores Sea (Sumbawa); 73-296 m.

Elsewhere-, widespread from southwest Indian Ocean to Hawaiian Islands, including Celebes Sea (Darvel Bay) and

Japan; 46-825 m.

Javania lamprotichum (Moseley, 1880)

Desmophyllum lamprotichum Moseley, 1880: 41-42, figs 1-2.

? Desmophyllum alabastrum Alcock, 1902a: 105; 1902c: 28-29 (in part: "Siboga" stn 105, pi 4, fig 27 27a) —

Faustino. 1927: 64, pi. 5, figs 11-12.

Javania lamprotichum - Cairns, 1984: 21, pi. 4, figs D-E; 1995: 112, pi. 37, figs b-c.

Material EXAMINED. — Philippines. "Siboga": stn 105, 1 (ZMA, see Remarks).

Musorstom 1: stn 62, 1 (MNHN). — Stn 63, 1 (MNHN). — Stn 65, 1 (USNM 97492).

MUSORSTOM 2: stn 53, I (USNM 97493).

Musorstom 3: stn 88, 1 (MNHN). — Stn 94, 1 (USNM 97494).

Type Locality. — Unknown.

diagnosis. — Corallum elongate-conical and straight, having a pedicel thickened with layers of dense

stereome up to a diameter of 10 mm; theca of upper part of corallum relatively thin and delicate. Largest known

specimen (MUSORSTOM 1 stn 63) 29.5 x 44 mm in calicular diameter and 48 mm in height, with a pedicel

diameter of 9.0 mm. Corallum Oared distally, having an elliptical calice: GCD:LCD = 1.25-1.50. Upper corallum

usually light reddish-brown, occasionally white; lower corallum usually white. Septa hexamerally arranged in

5 complete cycles: Si-2>S3>S4>S5. All septa exsert to some degree, producing a serrate calicular edge.

REMARKS. — The figured syntype of Desmophyllum alabastrum Alcock, 1902 ("Siboga" stn 105) is missing

trom the ZMA; however, ALCOCK's (1902c) illustration appears to show the lower half of a Javania. In 1994, a

large, previously unidentified and well-preserved specimen of J. lamprotichum (35.8 x 27.6 mm in calicular

diameter, 8.3 mm pedicel diameter), was found at the ZMA, also from "Siboga" stn 105. Although not the

illustrated specimen and probably not even seen by ALCOCK, this is considered to be indirect evidence that

ALCOCK's (1902c) figured specimen of D. alabastrum might have been Javania lamprotichum. The other syntype

of D. alabastrum, from "Siboga " stn 95 (ZMA Coel. 1 252), appears to be a Thalamophyllia.

Javania lamprotichum differs from J. insignis Duncan, 1876, in having a larger, more delicate, flared corallum;

usually a pigmented, noncostate theca; and more prominent S4-5.

Coralla of 3 lots (MUSORSTOM 1 stns 63, 65, 2 stn 53) contain 1 or more borings of acrothoracican cirripedes.

Distribution. — Philippines: Lubang Island; Sulu Sea (Sulu Archipelago); 191-842 m. Elsewhere: Kermadec

Ridge; Johnston Atoll; Hawaiian Islands; 244-710 m.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

165

Javania pachytheca Cairns, 1995

Figs 21 i, 22 a

Javania pachytheca Cairns, 1995: 112-113, pi. 36, figs j-1, pi. 37, fig. a.

MATERIAL EXAMINED. — Indonesia. "Albatross": stn 5584, 1 (USNM 97495). — Stn 5634, 1 (USNM 97496).

Snellius 2: stn 81.2, 3 (NNM 23089).

TYPE Locality. — "Tangaroa" stn K846: 30° 1 3. l'S, 178°32.0'W (offMacauley Island, Kermadecs), 610 m.

DIAGNOSIS. — Corallum tall and slender, the specimen figured herein being 7.6 x 9.0 mm in calicular diame¬

ter, 20.2 mm in height, and 4.3 mm in pedicel diameter. Thecal wall quite thick (0.9-2.3 mm), covered with a fine

granulation. Corallum white or light brown. Septa hexamerally arranged in 4 complete cycles: Si-2>S3»S4, the

S4 being nonexsert and quite slender. Inner edges of all septa moderately sinuous. Fossa deep and narrow.

DISTRIBUTION. — Indonesia: Ceram Sea (Obi Islands); 534-601 m. Elsewhere: Malaysia (Celebes Sea off

Sabah); southwest Pacific from North Island, New Zealand to the Chesterfield Islands, including the Lord Howe

Seamount Chain; 360-1045 m.

Javania sp.

Figs 22 b-c

MATERIAL EXAMINED. — Indonesia. Karubar: stn 44. 1 (MNHN). — Stn 49, 1 fragment (USNM 97498). —

Stn 86, 1 (MNHN).

Diagnosis (specimen from Karubar stn 44). — Corallum straight, slightly flared distally: 13.7 x 15.6 mm

in calicular diameter and 33.8 mm in height, with a pedicel diameter of 7.2 mm. Upper part of corallum smooth

and porcellaneous, a light purple-grey in colour; lower corallum superficially eroded, discoloured, and encrusted.

Septa hexamerally arranged in 4 cycles: Si»S2>S3>S4- Si extremely exsert (up to 6.5 mm), having vertical,

straight inner edges. S2 1/2 as exsert but almost as wide as Si, having slightly sinuous inner edges. S3 only

0.5- 1.5 mm exsert, about 3/4 width of an S2, also having slightly sinuous inner edges. S4 not exsert (not even

reaching the top of the calice), about 1/3 width of an S3, having slightly sinuous inner edges. Fossa deep and

narrow.

Remarks. — The Karubar specimens reported above resemble the widespread species J. cailleti (Duchassaing

& Michelotti, 1864), but differ in having more exsert Si (much more exsert than their S2), and in having a purple-

grey pigmentation. Whereas some specimens of J. cailleti have highly exsert septa (e.g., holotype of D. nobile

Verrill, 1885; USNM 82016 from Lydonia Canyon, NW Atlantic), the Si of most specimens rarely exceed 4 mm

in exsertness, and in all cases S2 equal in size to SI. Furthermore, the corallum of J. cailleti is always white,

whereas the Karubar specimens are pigmented.

Descriptions and illustrations of J. cailleti are found in CAIRNS (1979, 1982, 1991) and ZIBROWIUS (1980).

It is a widely distributed species known from the Atlantic, Pacific, and Indian Oceans at depths of 86-2165 m.

Distribution. — Indonesia: Arafura Sea (south of Tanimbar Islands); 209-291 m.

Genus TRUNCATOFLABELLUM Cairns, 1989

Truncatoflabellum spheniscus (Dana, 1846)

Figs 23 a-b

Euphyllia spheniscus Dana, 1846: 160-161, pi. 6, figs la-e.

Truncatoflabellum spheniscus - CAIRNS, 1989a: 65-66, pi. 32, figs g-k (synonymy); 1994: 76, pi. 33, figs a-d

(synonymy).

Source :

166

S. D. CAIRNS & H. ZIBROWIUS

MATERIAL EXAMINED. — Indonesia. "Siboga": stn 299, 1 (ZMA Coel. 1229).

DEKi: stn 67, 15 (NNM 22616). — stn 71, 3 (NNM 22617). — Stn 74, 3 (NNM 22618). — Stn 90, 3 (NNM 22619).

— Stn 103, 100 (NNM 22620). — Stn 106, 13 (NNM 22621).

"Galathea stn 501, 1 (ZMUC).

"Hakuho Maru"\ stn KH72-1-29, 4 (USNM 97500). — Stn KH72-1-30, 7 (USNM 97501).

Karubar: stn 65, 99: 40 (MNHN), 29 (POLIPI), 30 (USNM 97499).

South China Sea. "Galathea": stn 330, 1 (ZMUC).

Singapore. 1, Phyletisches Museum, Jena, Germany (Coel. 922), coll. E. Haeckel, October 1900.

Australia. "Akaclemik Oparin": stn 18, 3 (USNM 93197).

Type Locality. — Singapore, South China Sea, 3-6 m.

Description. — Anthocyathus highly compressed (GCD:LCD up to 3.65), the planar thecal faces meeting in

rounded, but narrow, edges that bear 1 pair of edge spines about 4 mm above basal scar. Angle of thecal edges

57°-165°; angle of thecal faces quite low, 20°-31°. Largest known specimen (Karubar stn 65) 14.5 x 52.8 mm in

calicular diameter. Basal scar of most Indonesian specimens reported herein small, only 3. 2-5.0 mm in greater

diameter. Calicular margin strongly arched, smooth. Corallum white but often encrusted with bryozoa,

foraminifera, serpulids, and calcareous algae.

Large specimens (GCD > 45 mm) have septa hexamerally arranged in 6 cycles (Si-4>S5>S6, 192 septa), often

with additional pairs of S7, but smaller specimens have only 40, 42, 44, or 46 primary septa, a corresponding

number of secondaries, and twice that number of tertiary septa, resulting in coralla of 160, 168, 176, or 184 septa.

Si -4 (primary septa) narrow, notched near the calicular margin, and slightly concave midway down fossa. Lower,

inner edges of S1-4 quite thick and fused to columella. S5 (secondary septa) about 3/4 width of S1-4, not attaining

the columella. S6 (tertiary septa) about 1/2 width of S5, extending only a short distance from calicular edge. When

present, S7 (quaternary septa) are paired but are quite narrow and short. Fossa deep and elongate, containing a well-

developed columella about 1.4 mm in width.

Remarks. — Specimens from "Galathea" stns 330 and 501, and Haeckel's specimen are typical

T. spheniscus. with the large basal scar as in the type series. The other specimens reported above differ in having a

much smaller basal scar: 3. 2-5.0 mm in greater diameter vs 10.0-1 1.2 mm for the syntypes. The specimens appear

otherwise consistent in all characters, and thus scar diameter is considered by the first author to be variable in this

species. The second author is highly skeptical about identifying these small-scar specimens as T. spheniscus. His

general experience is that the size of the basal scar, reflecting the size at which transverse division occurred, is

standard in species of Truncatoflabellum and other transversely dividing species. T. spheniscus is distinguished

from most other species in the genus by its elongate, narrow calice and its distinctively shaped Si -4.

Distribution. — Indonesia: Arafura Sea (south of Tanimbar Islands); Timor Sea; Savu Sea; Sunda Strait,

Java Sea; 30-174 m. Elsewhere: Japan (Shikoku; Korea Strait; Honshu; northern Ryukyu Islands); Formosa Strait;

South China Sea (Singapore); Australia (Torres Strait, Gulf of Carpentaria, Western Australia); 2-106 m.

Truncatoflabellum aculeatum (H. Milne Edwards & Haime, 1848)

Flabellum aculeatum H. Milne Edwards & Haime, 1848a: 272, pi. 8, figs 3, 3a.

Truncatoflabellum aculeatum - Cairns, 1989a: 61, 64, pi. 31, figs h-l, pi. 32, figs a-c (synonymy).

Material EXAMINED. — Philippines. "Albatross": stn 5141, 3 (USNM 97502). — Stn 5253, 1 (USNM 97503)

Musorstom 3: stn 142, 3 (MNHN).

Indonesia. Deki: stn 14, 1 (NNM 22651). — Stn 60, 8 (NNM 22653). — Stn 82, 220 (NNM 22669). — Stn 84

2 (NNM 22655). — Stn 89, 1 (NNM 22656). — Stn 90, 1 (NNM 22657). — Stn 92, 2 (NNM 22658).

"Hakuho Maru": stn KH72-1-29, 4 (USNM 97506). — Stn KH72-1-30, 5 (USNM 97507).

Corindon 2: stn 260, 1 (USNM 97505). — Stn 292, 1 (MNHN).

Snellius 2: stn 4.099, 2 (NNM 22663). — Stn 4.134, 1 (NNM 22664). — Stn 4.228, 1 (NNM 22665) — Stn 4 232

2 (NNM 22666). — Stn 4.234, 4 (NNM 22667).

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACT1NIA

167

Type Locality. — Philippines (depth not given).

Diagnosis. — Anthocyathus compressed (GCD:LCD = 1. 8-2.6), the slightly convex thecal faces meeting in

rounded edges that usually bear 1 pair of edge spines directly adjacent to basal scar. Angle of thecal edges 45°-53°;

angle of thecal faces 28°-31°. Largest known specimen (KH72-1-29) 16.5 x 41.0 mm in calicular diameter and

25.5 mm in height, with a basal scar of 6.2 x 14.3 mm. Calicular margin of septal faces arched; corallum white.

Septa of most specimens hexamerally arranged in 5 complete cycles: Si-3>S4>Ss (96 septa); however, large

specimens have additional primary septa equal to Si -3 in size. Shape of septa as described for T. spheniscus , but

inner edges of Si -3 highly sinuous.

Remarks. — Although not noted by Cairns (1989a), T. aculeatum is similar to T. spheniscus (Dana, 1846),

especially in septal shape, and both species have been collected from the same stations. T. aculeatum is

distinguished by having a lower edge angle and higher face angle and thus a smaller GCDiLCD; a smaller

corallum; and less septa, most specimens having only 5 cycles (96 septa). The species is more fully described by

Cairns (1989a).

DISTRIBUTION. — Philippines : Visayan Sea; Bohol; Sulu Archipelago; 11-33 m. Indonesia: Makassar Strait;

Banda Sea (Kai Islands); Flores Sea (Lintah Strait and Selayar Island); Timor Sea; Sunda Strait, Java Sea;

18-81 m. Pleistocene of Talaud (UMBGROVE, 1938).

Truncatoflabellum candeanum (H. Milne Edwards & Haime, 1848)

Flabellum candeanum H. Milne Edwards & Haime, 1848a: 278, pi. 8, fig. 13.

Flabellum elegans H. Milne Edwards & Haime, 1848a: 277.

Truncatoflabellum candeanum - Cairns, 1989a: 70-71, pi. 36, figs d-h (synonymy); 1994: 76-77, pi. 33, figs e-f.

Material EXAMINED. — Philippines. Musorstom 1: stn 56, 8 (USNM 97508). — Stn 62, 1 (MNHN). —

Stn 64, 1 (MNHN). — Stn 72, 11: 10 (MNHN), 1 (USNM 97509).

Musorstom 2: stn 2, 1 (USNM 97510). — Stn 6, 5 (USNM 97511). — Stn 10, 2 (USNM 97512). — Stn 68,

1 (USNM 97513).

Musorstom 3: stn 88, 4 (USNM 97514). — Stn 90, 1 (MNHN). — Stn 91, 1 (MNHN). — Stn 92, 1 (USNM 97515).

— Stn 96, 3 (USNM 97516). — Stn 99, 1 (MNHN). — Stn 102, 25 (MNHN). — Stn 107, 2 (USNM 97517). — Stn 108,

5 (MNHN). — Stn 109, 1 (MNHN). — Stn 1 10, 1 (USNM 97518). — Stn 143, 20.

Indonesia. Deki: stn 54, 16 (NNM 22593). — Stn 58, 1 (NNM 22594).

Type Locality. — "Albatross" stn 5369: 13°48'N, 121°43'E (Luzon, Philippines), 194 m.

DIAGNOSIS. — Angle of acute thecal edges 40°-80°; angle of slightly convex thecal faces 30°-41°. GCD:LCD =

1.6- 1.9. Largest known specimen (holotype of F. elegans ) 16.5 x 32.3 mm in calicular diameter and 21.7 mm in

height. Most coralla bear 3 pairs of thecal edge spines: the lowest pair directly adjacent to basal scar and curved

downward; the middle pair directed horizontally; and the uppermost pair directed slightly upward. Spines often quite

long (up to 10 mm) and strongly compressed, having wide triangular bases. Basal scar up to 6 mm in greater

diameter. Calicular margin serrate, a small apex corresponding to each of the 20-24 primary septa. Upper,

peripheral edges of primary septa and corresponding costae, reddish-brown. Septa arranged in 3 size classes:

20-24:20-24:40-48, resulting in 80-96 septa. Columella well developed.

Remarks. — Truncatoflabellum candeanum is distinguished from other species by its multiple pairs of long,

flattened edge spines; and its scalloped calicular margin. It is more fully described by Cairns (1989a).

DISTRIBUTION. — Philippines: Lubang Island; Tayabas Bay; Samar Sea; Visayan Sea; 146-249 m. Indonesia:

Banda Sea (Kai Islands); 70-290 m. Elsewhere: Malaysia (Celebes Sea off Sabah); South China Sea off Hong

Kong; Japan (Korea Strait; Kyushu); 88-223 m.

Source :

168

S. D. CAIRNS & H. ZIBROWIUS

Truncatoflabellum incrustation Cairns, 1989

Truncatoflabellum incrustation Cairns, 1989a: 68-69, pi. 35, figs d-e.

Truncatoflabellum formosum Cairns, 1989a: 69-70 (in part: "Albatross" stns 5137, 5484).

MATERIAL EXAMINED. — Indonesia. "Siboga": stn 303, 1 (ZMA Coel. 1207).

Southwestern Sulawesi, 16.07.1985, 27-30 m, 1 (NNM 22595).

South China Sea. Macclesfield Bank, 64-82 m, 1 (BMNH).

Type Locality. — "Albatross" stn 5251: 7°05'12"N, 125°39'35"E (Mindanao, Philippines), 37 m.

DIAGNOSIS. — Angle of rounded thecal edges 23°-32°; angle of thecal faces 15°-19°. GCD:LCD = 1.65-2.10.

Largest known corallum (anthocyathus from "Albatross" stn 5253) 28 mm in GCD and 42 mm in height. One

pair of downward-projecting edge spines present near basal scar. Basal scar up to 6.0 mm in greater diameter. Theca

black-brown, but usually covered with a heavy encrustation of sessile organisms. Calicular edge smooth. Septa

hexamerally arranged in 5 complete cycles (Si-2>S3>S4>S5). Fossa deep and elongate, containing a rudimentary

trabecular columella.

Truncatoflabellum incrustatum is compared to T. irregulare (Semper, 1 872) in the following account, and is

described in greater detail by Cairns (1989a).

Distribution. — Philippines: Verde Island Passage; Leyte Gulf; Davao Gulf; Sulu Sea (Sulu Archipelago);

37-415 m. Indonesia: Savu Sea; Flores Sea (southwestern Sulawesi); 30-36 m. Elsewhere: South China Sea

(Macclesfield Bank); 64-82 m.

Truncatoflabellum irregulare (Semper, 1872)

Flabellum irregulare Semper, 1872: 242-245, figs 1-3, pi. 16, figs 7-17. — Cairns, 1989a: 67-68, pi. 34. figs i-k,

pi. 35, figs a-c (synonymy).

Not Flabellum irregulare - ALCOCK, 1902c: 32 (= Truncatoflabellum sp.).

MATERIAL EXAMINED. — Indonesia. "Siboga": stn 303, 3 (ZMA Coel. 1212).

Type Locality. — Lapinig Canal, Philippines, 11-18 m.

Diagnosis. — Angle of rounded thecal edges 36°-43°; angle of thecal faces 19°-21°. GCD:LCD = 1.6-2.0.

Largest known specimen (anthocyathus from "Albatross" stn 5145) 27.5 mm in GCD and 42.8 mm in height.

Usually one pair of downward-projecting thecal edge spines near basal scar. Basal scar usually 3x4 mm in

diameter. Theca white and often encrusted with sessile organisms. Calicular edge smooth. Septa arranged in

a variety of symmetries, 18:18:36 (72 septa) being the most common, but coralla with 16, 17, 19, and

12 primary septa are also known. Primary septa slightly exsert and notched near calicular edge. Fossa deep and

elongate, containing a rudimentary trabecular columella.

Remarks. Truncatoflabellum irregulare is distinguished from T. incrustatum by having irregular septal

symmetry and only 3 (not 4) size classes of septa. It also has a white (not dark) theca, a smaller basal scar, and

a septal notch on each primary septum.

The specimens reported as Flabellum irregulare by ALCOCK (1902c) ("Siboga" stns 49a, 253) were re-examined

and found to be juvenile specimens of an unknown species, not T. irregulare. T. irregulare is more fully described

and illustrated by Cairns (1989a).

Distribution. — Philippines: Bohol; Sulu Archipelago; 18-42 m. Indonesia: Savu Sea (Hainsisi, Samau

Island); 36 m.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

169

Truncatoflabellum paripavoninum (Alcock, 1894)

Fig. 22 f

Flabellum pari-pavoninum Alcock, 1894: 187; 1898: 21, pi. 2, figs 3a-b.

Truncatoflabellum paripavoninum - CAIRNS, 1989a: 72-73, pi. 37, figs j-1, pi. 38, fig. a (synonymy); 1995: 113-114, pi.

37, figs d-e.

MATERIAL EXAMINED. — Philippines. Musorstom 1: stn 44, 1 (MNHN). — Stn 47, 2 (MNHN).

MUSORSTOM 2: stn 25, 3 (MNHN). — Stn 77, 1 (MNHN).

Musorstom 3: stn 106, 2 (USNM 97548).

Indonesia. Karubar: stn 39, 1 (POLIP1). - Stn 56, 8 (USNM 97549). - Stn 57, 1 (MNHN). - Stn 70, 1 (USNM

97550). _ Stn 71, 9: 3 (MNHN), 6 (USNM 97551). — Stn 87, 2 (MNHN). — Stn 91, 1 (POL1P1).

Type Locality. — "Investigator" stn 177: 13°47'04"N, 73°07'E (Laccadive Sea), 1 163 m.

Diagnosis. — Angle of thecal edges 57°-138°; angle of thecal faces 31°-62°. Thecal faces virtually planar,

meeting in straight, nonspinose, noncrested, acute thecal edges. Largest known specimen (Karubar stn 71)

34.4 x 61.7 mm in calicular diameter and 48.2 mm in height. Basal scar elliptical but variable in size, greater

diameter 6.8 to 14.5 mm. Corallum white or uniformly reddish brown, the theca of most specimens being worn

and discoloured. Septa hexamerally arranged in 6 cycles (Si-3>S4>Ss>S6, 192 septa), the 6th cycle beginning to

appear at a GCD of 26-30 mm. Columella well developed, consisting of a robust fusion of the lower, inner edges

of S i-3, and being about 2.1 mm in width.

REMARKS. — Among the western Pacific species, T. paripavoninum is distinguished by having a large

corallum; nonspinose and noncrested thecal edges; and by occurring in relatively great depths. The diagnosis above

is based on the anthocyathus stage, only one specimen (Karubar stn 56) being an anthocaulus. The species is

more fully described and illustrated by CAIRNS (1989a, 1995).

Four specimens from Karubar stns 56 and 71 serve as the substratum for a stalked suberitid sponge

(? Rhizaxinella, identified by K. RUTZLER) (Fig. 22 f). The slender stalk (up to 60 mm long and 3 mm in

diameter) supports a "head" about 20 x 10 mm in size. The stem of the sponge is attached to the theca of the

living coral just below the calicular edge.

DISTRIBUTION. — Philippines : Lubang Island; Bohol Strait; Sulu Archipelago; 512-772 m. Indonesia: Arafura

Sea (southeast of Tanimbar Islands); Gulf of Bone, Sulawesi; Bali Sea; 411-1022 m. Elsewhere: Malaysia (Darvel

Bay, Celebes Sea); Laccadive Sea; Kermadec Islands; 1035-1450 m.

Truncatoflabellum formosum Cairns, 1989

Truncatoflabellum formosum Cairns, 1989a: 69-70 (in part: not "Albatross" stns 5137, 5484, 5162, and 5483, the first

2 stations being T. incrustatum, the latter 2 unidentified species of this genus), pi. 35, figs j-k, pi. 3b, tigs a-D

(synonymy); 1994: 77, pi. 33, figs g-h (synonymy).

Truncatoflabellum sp. nov. - Cairns, 1989a: 73, pi. 38, figs g-h.

? Truncatoflabellum formosum - CAIRNS & KELLER, 1993: 265, pi. 10, fig. I, pi. 11, fig- A.

MATERIAL EXAMINED. — Philippines. Musorstom 2: stn 2, 1 (MNHN). — Stn 32, 1 (MNHN).

Musorstom 3: stn 131, 5 (USNM 97540).

Indonesia. "Siboga": stn 274, 1 (ZMA Coel. 1208).

Corindon 2: stn 216, 6 (MNHN).

Karubar: stn 1, 4: 1 (POLIPI), 3 (USNM 97541). — Stn 3, 1 (POLIPI).

Type Locality. — "Albatross" stn 5249: 7°06’06"N, 125°40'08"E (Mindanao, Philippines), 42 m.

Source :

170

S. D. CAIRNS & H. ZIBROWIUS

DIAGNOSIS. — Angle of rounded thecal edges 37°-59°; angle of thecal faces 18°-34°. GCD:LCD = 1.4-1. 8.

Most anthocyathi have 2 pairs of thecal edge spines, the lowermost pair 3-4 mm above the basal scar and the upper

pair usually quite short, each of these upper spines with a broad, flat, triangular base. Largest Philippine specimen

(". Albatross " stn 5289) 13.4 x 23.0 mm in calicular diameter and 31.4 mm in height. Calicular margin arched and

smooth. Basal scar relatively small, 4-5 mm in greater diameter. Well-preserved specimens have reddish-brown

thecal stripes associated with the 20 primary septa. Septa arranged in 3 size classes (20:20:40 = 80 septa), some

pairs of tertiaries occasionally missing. Primary septa gracefully arched near calicular edge, having sinuous inner

edges. Ternary septa rudimentary, much smaller than the secondaries. All septa widely spaced.

Remarks. — This species is more fully described by Cairns (1989a, 1994).

Distribution. — Philippines: Lubang Island; Verde Island Passage; Sulu Sea (west of Panay and Sulu

Archipelago); Leyte Gulf; Davao Gulf; 42-315 m. Indonesia: Makassar Strait; Banda Sea (Kai Islands);

Gulf of Bone, Sulawesi; 57-933 m. Elsewhere: Japan (Honshu, Shikoku, Kyushu); ?southwest Indian Ocean;

106-230 m.

Truncatoflabellum pusillum Cairns, 1989

Truncatoflabellum pusillum Cairns, 1989a: 71-72, pi. 37, figs a-e. — Cairns & KELLER, 1993: 265, pi. 11, fig. E.

Material EXAMINED. — Philippines. Musorstom 2: stn 33, 10 (USNM 97532).

Musorstom 3: stn 87, 1 (MNHN). — Stn 96, 1 (MNHN). — Stn 102, 4 (MNHN). — Stn 109, 1 (MNHN).

Indonesia. Deki: stn 3, 2 (NNM 22588). — Stn 46, 4 (NNM 22589). — Stn 53, 1 (NNM 22590). — Stn 63

4 (NNM 22591).

CORINDON 2: stn 248, 4 (USNM 97536).

Karubar: stn 15, 20 (MNHN). — Stn 18, 5 (MNHN). — Stn 44, 2 (POLIPI).

Type Locality. — "Albatross" stn 5178: 12°43'N, 122°06'15"E (Sibuyan Sea, Philippines), 143 m.

Diagnosis. — Angle of thecal edges 14°-18°; angle of thecal faces 18°-20°. Thecal faces convex; thecal edges

rounded, each edge bearing 2-4 slender spines. Corallum slender and high, the largest specimen (Deki stn 3) 6.4 x

10.0 mm in calicular diameter, with a greater basal scar diameter of 2.7 mm. Theca smooth and porcellaneous,

bearing fine transverse striae; theca streaked with reddish-brown stripes, one corresponding to every interseptal

space adjacent to each Si -2. Septa hexamerally arranged in 3 cycles and usually 4 pairs of S4, 1 pair occurring in

the half-systems adjacent to each principal septum, resulting in 32 septa: Si-2>S3>S4. However, large specimens

may contain a full 4th cycle of septa. Inner edges of S 1.2 highly sinuous.

Remarks. — This species is more fully described and illustrated by Cairns (1989a).

Distribution. — Philippines: Lubang Island; Verde Island Passage; Sibuyan Sea; Sulu Sea (Sulu

Archipelago); 137-205 m. Indonesia: Makassar Strait; Banda Sea (Kai Islands); 85-300 m. Elsewhere:

Mozambique; 1 10-132 m.

Truncatoflabellum dens (Alcock, 1902)

Flabellum dens Alcock, 1902a: 106-107; 1902c: 32, pi. 4, figs 30, 30a. — Cairns, 1989a: 54, pi. 28 figs g-k

(synonymy).

Truncatoflabellum dens - CAIRNS, 1995: 114-115, pi. 37, figs f-h.

^ATERIAL EXAMINED. — Indonesia. "Siboga": stn 95, 9 syntypes: 8 (ZMA Coel. 1209, 1449), 1 (USNM

97 538).

Snellius 2: stn 4.032, 1 (NNM 22770).

Karubar: stn 2, 2 (MNHN).

Type Locality. "Siboga" stn 95: 5°43.5'N, 1 19°40'E (Sulu Archipelago, Philippines), 522 m.

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

171

DIAGNOSIS. — Angle of rounded thecal edges changes from 58°-80° at a height of about 6 mm to 21 -35 .

Corallum highly compressed, angle of thecal faces 14°-18° and GCD:LCD = 1.7-2.3. Corallum small, up to

12 7 mm in GCD and about 15 mm in height, having a small basal scar about 2x3 mm. Thecal edges usually

nonspinose. Theca bears reddish-brown stripes corresponding to each interseptal space. Calicular edge smooth.

Septa hexamerally arranged in 4 cycles (S i -2>S3»S4), larger coralla with up to 4 pairs of S5 (a total of 56 septa).

Si -2 have extremely sinuous inner edges. Fossa deep and elongate, containing a rudimentary columella.

REMARKS. — Truncatoflabellum dens is characterized by having a relatively small corallum with a bimodal

edge angle, no edge spines, and a very small basal scar diameter. It is more fully described and illustrated by

CAIRNS (1989a, 1995).

DISTRIBUTION. — Philippines : Sulu Sea (Sulu Archipelago); 522 m. Indonesia : Banda Sea (Tukangbesi and

Kai Islands); 300-385 m. Elsewhere: Kermadec and Norfolk Ridges; New Caledonia; 320-555 m.

Truncatoflabellum phoenix Cairns, 1995

Truncatoflabellum sp. B - Cairns, 1994: 79, pi. 33, figs i, 1.

Truncatoflabellum phoenix Cairns, 1995: 115-116, pi. 37, fig. i, pi. 38, figs a-f.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5146, 27 (USNM 97542). — Stn 5147. 7 (USNM

97543) - Stn 5159 21 (USNM 97544). - Stn 5162, 13 (USNM 97545). - Stn 5179, 21 (USNM 97546).

Musorstom 3: stn 137, 21 (MNHN).

Indonesia. SNELLIUS 2: stn D2, 3 (NNM 23203).

KarubaR: stn 7, 6 (MNHN). — Stn 22, 3 (MNHN). — Stn 44, 1 (POLIPI).

TYPE Locality. — "Tangaroa" stn C531: 29°14'40”S, 178°02'W (Raoul Island, Kermadecs), 179 m.

Diagnosis. - Corallum elongate and compressed (GCD:LCD = 1. 3-2.3), having nearly parallel thecal edges

and faces, which result in a basal scar of almost equal width to calice. Largest Philippine specimen ("Albatross

stn 5179) 2.9 x 3.9 mm in calicular diameter and 6.8 mm in height, with a greater scar diameter of 3.4 mm. In

addition to transverse division, this species also undergoes rejuvenescence, resulting in elongate coralla ol varying

diameters. Thecal faces convex; thecal edges rounded, each edge bearing 1-6 strongly downcurved spines. Theca

porcellanous, well-preserved specimens brown in colour, with more intense pigmentation in stripes adjacent to

each C,-2. Septa hexamerally arranged in 3 cycles (Si-2»S3, 24 septa), only rarely having additional pairs of S4

in end half-systems. Inner edges of Si-2 sinuous. Columella robust, about 0.4 mm wide.

Remarks. — Truncatoflabellum phoenix is distinguished from T. pusillum Cairns, 1989, and T. dens

(Alcock, 1902), species also having small coralla, by having: parallel thecal edges and faces that result in a basal

scar almost as large as the calice; usually only 24 septa; a more robust columella; an elongate corallum

characterized by multiple rejuvenescence events; and strongly downcurved thecal edge spines. The species is more

fully described and illustrated by CAIRNS (1994, 1995).

Distribution. — Philippines: Sibuyan Sea; Sulu Sea (Sulu Archipelago); 18-421 m, but most specimens

collected at 20-70 m. Indonesia: Banda Sea (Tanimbar Islands); Arafura Sea (southeast of Tammbar Islands); 291-

295 m. Elsewhere: Kermadec Islands; northern Ryukyu Islands; 80-179 m.

Truncatoflabellum mortenseni sp. nov.

Figs 22 g-h

Material examined/types. — Philippines. Musorstom 2: stn 33, 49 anthocauli and many juveniles (MNHN),

P Musorstom 3: stn 124, 1 anthocaulus and 1 anthocyathus, paratypes (MNHN). — Sin 131, 93 anthocauli and

18 anthocyathi, paratypes (MNHN and USNM 97521).

Source :

172

S. D. CAIRNS & H. ZIBROWIUS

Indonesia: Mortensen'S Java-S.A. Expedition: Stn 5, 29 + 81: 20 anthocauli and 76 anthocyathi, paratypes

(ZMUC). 1 anthocyathus, holotype (ZMUC), 9 anthocauli and 4 anthocyathi (USNM 97522). — Stn 6, 20 anthocauli and

73 anthocyathi, paratypes (ZMUC). — Stn 8, 7 anthocauli and 28 anthocyathi, paratypes (ZMUC). — Stn 9,

4 anthocyathi, paratypes (ZMUC). — Stn 18, 1 anthocaulus (ZMUC).

Karubar: stn 1,4 anthocauli, paratypes (MNHN). — Stn 30, 2 anthocauli and 2 anthocyathi, paratypes (MNHN).

Type Locality. — 11°36'N, 121°43'E (Sulu Sea west of Panay), 120-122 m.

ETYMOLOGY. — This species is named for Theodor MORTENSEN, who made many collections of Indo-West

Pacific fauna, including this species from the Java Sea.

DESCRIPTION. — Anthocaulus: Angle of rounded thecal edges 49°-61°; angle of convex thecal faces 23°-31°.

One pair of slender edge spines occurs 4-6 mm above the pedicel, these spines being elongate (up to 7.5 mm) only

on small specimens; on larger specimens these delicate structures are always broken. Base of thecal spines broad

and compressed. Pedicel circular and quite small (0.8- 1.1 mm in diameter), sometimes revealing the 6 protosepta.

Largest anthocaulus (MUSORSTOM 3 stn 131) 10.1 x 17.1 mm in calicular diameter and 19.5 mm in height.

Calicular margin of thecal faces slightly arched and smooth; GCD:LCD = 1.65-1.85. A thin reddish-brown thecal

stripe corresponds to each Si-3, a thinner stripe to each S4-5. Most anthocauli from MUSORSTOM 3 stn 131 tend

not to divide transversely after the edge spines are formed, continue to grow up to 18 mm in height, and have

64 septa; however, anthocauli from all other stations rarely exceed 7 mm in height before transversely dividing or

forming an incipient transverse fracture line.

Septa hexamerally arranged in 5 cycles (Si-3>S4>Ss), pairs of S5 present only in large anthocauli (up to

64 septa). S1-3 nonexsert, attenuate, gracefully concave near calicular edge, having highly sinuous lower, inner

edges. S4 about 1/2 width of S1-3, having straight inner edges. S5 1/3 to 1/2 width of the S4, having slightly

dentate inner edges. S5 originate in a progression from half-systems adjacent to the principal septa towards centre

of thecal face. Fossa deep and elongate; columella rudimentary.

Anthocyathus: One of the largest anthocyathi (holotype) measures 12.3 x 22.5 mm in calicular diameter and

16.9 mm in height, with a basal scar of 3.5 x 7.1 mm. Angle of thecal edges and faces similar to that of

anthocaulus. One, occasionally 2, pair(s) of edge spines, the lowermost pair occurring within 1 mm of basal scar.

Basal scar elliptical, 6.3-7. 3 mm in greater diameter. Theca pigmented as in anthocaulus.

Septa of anthocyathus hexamerally arranged in 5 cycles (Si-3>S4>Ss) as in anthocaulus, but in most

anthocyathi the 5th cycle is complete (96 septa). S 1-3 notched near theca, rising slightly above calicular margin

toward centre of fossa. Inner septal edges highly sinuous. S4 1/3 to 2/3 width of Si -3, but having less sinuous

inner edges. S5 rudimentary. Columella well developed, about 1 .3 mm in width.

Remarks. — In many ways the anthocaulus of T. mortenseni is similar to that of T. zuluense Cairns, 1993,

the corallum of both species often resisting transverse division after the basal pair of edge spines has formed.

T. mortenseni differs from T. zuluense in having: a larger edge angle; a smaller diameter pedicel; Si -3 that are

equal in size (versus Si-2>S3 in T. zuluense)-, and usually having more septa, i.e., 56-80 vs 48-56 for T. zuluense.

The anthocyathus of T. mortenseni is distinguished (Cairns, 1989a: table 6) by the combination of having a

septal formula of Si-3»S4>Ss and having 1 or 2 pairs of thecal edge spines.

Many live specimens of Truncatoflabellum mortenseni from Bali Strait, 50-70 m (MORTENSEN's stn 5, 6, 8)

were the substrate of the inarticulate disciniscid brachiopod Discradisca Stella (Gould) (det. A. LOGAN, 1993) and

the hipponicid prosobranch gastropod Malluvium sp. (det. A. Waren, 1993). Both epibionts were localized

generally near the calicular edge of live corals, Malluvium sp. exceptionally also on a dead coral.

Distribution. — Philippines-. Verde Island Passage; Tablas Strait; Sulu Sea west of Panay; 122-130 m.

Indonesia: Banda Sea (Kai Islands); Bali Sea and Bali Strait; 50-156 m.

Truncatoflabellum angustum sp. nov.

Figs 23 c-f

Truncatoflabellum dens - Cairns, 1995: 114 (in pan: pi. 37, figs f-h). [Not Flabellum dens Alcock, 1902],

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

173

MATERIAL EXAMINED/TYPES. — Philippines. " Albatross stn 5567, 11 paratypes (USNM 97524).

Musorstom 1: stn 64, 1 paratype (USNM 97525).

Musorstom 2: stn 63, 1 paratype (MNHN). t

Musorstom 3: stn 92, 2 paratypes (USNM 97526). — Stn 126, 4 paratypes (USNM 97527). — Stn 130, 1 paratype

(MNHN). — Stn 143, holotype and 12 paratypes (MNHN). ,IIOXIW

Indonesia. Karubar: stn 2, 19 paratypes (MNHN). - Stn 3. 2 paratypes (POLIPI) . - Stn 18, 2 paratypes (USNM

97530). _ Stn 31, 1 paratype (MNHN). — Stn 44, 20 paratypes (USNM 97531).

NONTYPES: Kermadecs. "Tangaroa": stn K858, 1 (ex USNM 94274, now USNM 97523).

"Acheron": stn BS441, 1 (USNM 94276).

TYPE Locality. — Musorstom 3 stn 143: 1 1°28.3'N, 124°1 1.6'E (Visayan Sea, Philippines), 205-214 m.

ETYMOLOGY. — The species name angustum (Latin angustus, slender, thin) alludes to the highly compressed

(slender) calice of this species.

DESCRIPTION. — Anthocyathus: Thecal faces flat to slightly convex, meeting in sharp edges that bear 3 or

4 pairs of slender, delicate edge spines. Angle of thecal edges 28°-52°; angle of thecal faces 17°-22 . Basal scar

relatively small: 1. 8-2.5 x 2.7-3. 3 mm. Largest specimen (holotype) 6.1 x 14.0 mm in calicular diameter and

10.8 mm in height, with a basal scar diameter of 2.1 x 3.1 mm. Calicular margin of septal faces slightly arched

and smooth; GCD:LCD = 1.85-2.31. Theca white to slightly reddish-brown.

Septa hexamerally arranged in 5 cycles, the last cycle never complete: Si-2>S3»S4>Ss. Most specimens have

only 4 pairs of Ss, 1 pair in each half-system adjacent to the 2 principal septa, resulting in a total of 56 septa.

Si-2 nonexsert, attenuate, having very sinuous inner edges that fuse to the columella lower in fossa. S3 7/10-9/10

width of S1-2 and usually less sinuous. S4 1/4- 1/3 width of the S3; S5 rudimentary, 1/2 width of S4. Fossa deep

and narrow, the columella being about 0.8 mm in width.

Anthocaulus: Only one unequivocal anthocaulus is known (MUSORSTOM 3 stn 130), the anthocyathus having

become detached during this study. This anthocaulus is 1.3 mm in pedicel diameter, 2.7 mm in height, and has a

GCD of 3.25 mm. It bears no thecal spines and has 24 septa arranged in 2 size classes: S i-2>S3-

Remarks. — Truncatoflabellum angustum has a septal number and arrangement similar to T. zuluense Cairns,

1993, and T. gardineri Cairns, 1993, both from the southwest Indian Ocean. T. angustum differs from both in

having a more elongate calice (higher GCD:LCD). It also differs from T. gardineri Cairns, 1993, in having a

smaller basal scar, and having thecal edge spines, not crests. It differs from T. zuluense in consistently severing its

anthocyathus from its anthocaulus.

Several specimens of T. angustum were reported and figured by CAIRNS (1995) as the anthocyathus stage of

T. dens, both species having been found at the same station and being similar in size and colouration. T. dens

(Alcock, 1902) is distinguished from T. angustum by having a bimodal edge angle and tending to have 32 septa vs

56 septa in T. angustum.

Distribution. — Philippines: Lubang Island; Sulu Sea (Semirara Islands, west of Panay, and Sulu

Archipelago); Visayan Sea; 195-490 m. Indonesia: Banda Sea (Kai Islands); 212-288 m. Elsewhere: Kermadec

Islands; 402-465 m.

Genus BLASTOTROCHUS H. Milne Edwards & Haime, 1848

Blastotrochus nutrix H. Milne Edwards & Haime, 1848

Blastotrochus nutrix H. Milne Edwards & Haime, 1848a: 284-285, pi. 8, fig. 14. — SEMPER 1872: 238-241, pi. 16,

figs 1-6. — Cairns, 1989a: 74-75, pi. 38, figs i-m, pi. 39, figs a-b (synonymy); 1989c: 643, figs a-b (upper).

Flabellum (Blastotrochus) nutrix - Faustino, 1927: 59-60, pi. 5, figs 1-6.

Material EXAMINED. — Indonesia. "Siboga": Stn 240. 1 (ZMA Coel. 1177); stn 315, 1 (ZMA CoeL 1 176). —

Unnumbered station. Banda Sea, 1 (USNM 97553, ex. ZMA Coel. 1 178). — Unnumbered station, Ambon, 1 (ZMA Coel.

1175).

Source :

174

S. D. CAIRNS & H. ZIBROWIUS

"Galathea": Stn 485, 4 (ZMUC).

Koedingarrang Keke, southwestern Sulawesi, 30-32 m, 17.07.1985, 8 (NNM 22431, 22433); Samalona,

southwestern Sulawesi, 23 m, 10.07.1985, 1 (NNM 22432).

Type Locality. — Philippines (depth not given).

DIAGNOSIS. — Corallum elongate and compressed, having rounded thecal edges and 1 pair of downward

projecting edge spines just above basal scar. Additional coralla (anthoblasts) bud from the thecal edges, up to

4 coralla occurring on each edge. Buds ultimately detach at a height of 4-5 mm. Largest Indonesian specimen

(USNM 97553, ex ZMA Coel. 1178) 8.9 x 1 1.2 mm in calicular diameter and 27.8 mm in height. Angle of thecal

edges 10°-16°; angle of thecal faces 10°-12°. Calice elliptical: GCDiLCD = 1.3-1. 6. Basal scar 2. 3-2. 5 x

2. 8-3. 3 mm in diameter. Theca white and usually encrusted. Septa hexamerally arranged in 4 cycles (Si-2>S3>S4),

although 4th cycle sometimes incomplete. Columella rudimentary, consisting of a fusion of lower, inner edges of

S 1 -2-

Remarks. — Although similar in shape to species of Truncatoflabellum, Blastotrochus differs in budding

corallites from its thecal edges. B. nutrix is more fully described and illustrated by Cairns (1989a, c).

Distribution. — Philippines : Lapinig Canal, north of Bohol; 1 1-18 m. Indonesia : Banda Sea (Ambon and

Banda Islands); Flores Sea (southwestern Sulawesi); Bali Strait; eastern Java Sea; 23-62 m.

Genus PLACOTROCHIDES Alcock, 1902

Placotrochides scaphula Alcock, 1902

Placotrochides scaphula Alcock, 1902b; 121-122; 1902c: 34, pi. 4, figs 32, 32a. — Cairns, 1989a: 78-79, pi. 40,

fig. 1, pi. 41, figs a-e (synonymy); 1994: 79-80, pi. 34, figs f-h; 1995: 116-117, pi. 38, fig. 38, pi. 39, fig. a. —

Cairns & Parker, 1992: 48-49, figs 15 h, i. — Cairns & Keller, 1993: 272-273, pi. 12, figs D, G.

Flabellum elongation Hu, 1987: 44, pi. 3, figs 4, 7-8 (new synonymy).

MATERIAL EXAMINED. — Indonesia. "Siboga": Stn 297, 1 (ZMA Coel. 1228).

Type Locality. — "Siboga" stn 212: 5°54.5'S, 120°19.2'E (Flores Sea), 462 m.

Diagnosis. — Corallum cylindrical, with virtually parallel thecal faces and parallel, rounded thecal edges. No

edge spines. Calice elliptical, the specimen reported herein 9.2 mm in GCD, with a basal scar of approximately

the same size as the calice. Shape of calice often asymmetrical, one thecal face being slightly less curved and

having more septa than the other face. Theca white, engraved with shallow, vertical striae that delimit wide, flat

costae. Septa hexamerally arranged in 4 cycles (Si-2>S3>S4), the 4th often incomplete, resulting in 40 septa. All

septa relatively thin and widely spaced. Fossa deep and elongate, containing a well-developed, elongate trabecular

columella that occupies medial 1/3 of fossa.

Remarks. — ALCOCK (1902b, c) chose a relatively small specimen ("Siboga" stn 212) as the holotype of this

species, not even mentioning the larger, better-preserved specimen reported herein from "Siboga" stn 297. It is

possible that Alcock did not have access to the entire "Siboga" collection when he wrote his report on the corals

from that expedition, as many other large, well-preserved "Siboga" specimens are reported herein for the first time.

Placotrochides scaphula is more fully described by Cairns (1989a, 1994) and Cairns & Parker (1992).

Distribution. — Philippines: Verde Island Passage; Sulu Archipelago; 520-522 m. Indonesia: Flores Sea

(southwestern Sulawesi); Timor Sea; 462-1628 m. Elsewhere: Japan (Honshu, Ryukyu Islands); southwest Indian

Ocean; Victoria, Australia; 80-1360 m. Plio-Pleistocene of Taiwan (Hu, 1987).

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

175

Genus PLACOTROCHUS H. Milne Edwards & Haime, 1848

Placotrochus laevis H. Milne Edwards & Haime, 1848

Placotrochus laevis H. Milne Edwards & Haime, 1848a: 283, pi. 8, figs 15, 15a. — Semper, 1872: 251-252, pi. 18,

figs 11-13. — Faustino, 1927: 61-62, pi. 5, figs 7-10. — Cairns, 1989a: 75-76, pi. 39, figs c-g (synonymy).

Placotrochus candeanus H. Milne Edwards & Haime, 1848a: 283-284. — Alcock, 1902c: 33.

MATERIAL EXAMINED. — Indonesia. "Siboga": stn 91, 1 (ZMA Coel. 1234). — Stn 116, 2 (ZMA Coel. 1310-

1311). - Stn 133, 1 (ZMA Coel. 1312). - Stn 273, 1 (ZMA). - Stn 279, 1 (ZMA Coel. 1313).

DEKr stn 10 9 (NNM 22599). — Stn 18, 1 (NNM 22600). — Stn 44. 1 (NNM 22601). — Stn 64, 3 (NNM 22602). —

Stn 65, 2 (NNM 22603). — Stn 66, 1 (NNM 22604). — Stn 69, 1 (NNM 22605). — Stn 89. 1 (NNM 22606). — Stn 91,

1 (NNM 22607). — Stn 110, 4 (NNM ). — Stn 116, 12 (NNM 22608).

SNELLIUS 2: stn 4.232, 1 (NNM 22596). — Stn 4.234, 1 (NNM 22597). — Stn 4.235, 1 (NNM 22598).

Karubar: stn 65, 5 anthocyathi (USNM 97554).

Type Locality. — Philippines (depth not given).

DIAGNOSIS. — Angle of thecal edges 40°-72°; angle of thecal faces 20°-33°. Thecal edges narrow, rounded (not

ridged or carinate); 1 pair of thecal edge spines near basal scar. Largest Indonesian specimen (Karubar stn 65)

10.3 x 19.7 mm in calicular diameter and 18.6 mm in height, with a basal scar diameter of 3.6 x 6.8 mm.

Ci -3 slightly ridged; corallum white. Septa hexamerally arranged in 5 complete cycles: Si-3>S4»Ss (96 septa).

Columella lamellar, the lamellae sometimes subdivided into many smaller elements.

Remarks. — This species is more fully described and illustrated by Cairns (1989a).

DISTRIBUTION. — Philippines : Bataan Peninsula; Sulu Sea (Basilan, Mindanao, and Sulu Archipelago),

35-69 m. Indonesia: Makassar Strait; Molucca Sea (Talaud Islands, Gulf of Tonuni); Banda Sea (Kai, Aru, Barat

Daya Islands); Arafura Sea (southeast of Tanimbar Islands); Flores Sea (Selayar Island); Bah Sea; Sunda Strait,

Java Sea; 12-289 m, but most records less than 100 m. Elsewhere: "South China Sea"; Australia (Arnhem Land;

Cape Jaubert, northwestern Australia; Queensland); India (Gulf of Mannar); 22-24 m.

Suborder DENDROPHYLLIINA

Family DENDROPHYLLIIDAE Gray, 1847

Genus BALANOPHYLLIA Searles Wood, 1844

Balanophyllia carinata (Semper, 1872)

Rhodopsammia carinata Semper, 1872: 257, pi. 19, figs 6a-b.

Rhodopsammia amoena Semper, 1872: 258, pi. 19, figs 5a-b.

Balanophyllia parallela - van DER Horst, 1922: 62. - Umbgrove, 1938: 272. [Not Rhodopsammia parallela Semper.

1 87?1

Balanophyllia carinata - ZiBROWiUS, 1985: 235-238, figs 15-24 (synonymy). — Zibrowius & GRYGIER, 1985: 127,

figs 30-35.

Material EXAMINED. — Philippines. "Albatross": stn 5134, 1 (USNM 97555). — Stn 5139. 5 (USNM 9^556),

— Stn 5151, 1 (USNM 97557). - Stn 5152, 4 (USNM 97558). — Stn 5156, 2 (USNM 97559). — Stn 5164, 17 (USNM

97560).

Siphilexp: stn 78-T14, 1 (USNM 77314).

Source :

176

S.D. CAIRNS & H. ZIBROWIUS

Indonesia "Siboga": stn 240, 5 (ZMA Coel. 567) (B. parallela of VAN DER HORST, 1922).

Deki- unnumbered station, Komkir, 75-90 m, 3 (ZMUC). - Stn 10, 20 (NNM 17352). — Stn 14, 2 (NNM 17357). —

Stn 18, 20 (NNM 17358). — Stn 20, 4 (NNM 17353). — Stn 24, 1 (NNM 17355). — Stn 31,2 (NNM 17354). — Stn 38,

1 (NNM 17351). — Stn 64, 20 (NNM 17345). — Stn 70, 7 (NNM 17346). — Stn 82, 20 (NNM 17347). — Stn 84, 4

(NNM 17364).

MORTENSEN'S Java-S.A. Expedition: stn 5, 82 (ZMUC). — Stn 6. 3 (ZMUC). — Stn 8, 6 (ZMUC).

CORINDON 2: stn 292, 1 (MNHN).

SNELL1US 2: stn 4.228, 1 (NNM 17363). — Stn 4.234, 4 (NNM 17361).

Type Locality. — Bohol near Pandonon, Philippines, 55 m.

Diagnosis. — Corallum unattached, straight to slightly curved, and compressed, the elliptical calice having a

GCD:LCD range of 1.15-1.55. Corallum edges sharp and slightly keeled, small buds asexually generating from the

edges. Once a bud detaches, a small irregularity may persist on thecal edge of parent. Because predominant mode of

reproduction appears to be by asexual budding, the base of most specimens is usually an open fracture about 1 mm

in diameter, revealing the 6 protosepta. Most specimens small, rarely over 10-11 mm in GCD, but ZIBROWIUS

(1985) reported a specimen 20 mm in GCD, and another listed above (SlPHlLEXP stn 78-T14) measures

14.2 x 21.6 mm in calicular diameter and 33.1 mm in height. Septa hexamerally arranged in 4 cycles in

a Pourtales plan, only larger specimens (GCD>19 mm) having pairs of S5, the largest specimen having 88 septa.

S 1-2 independent, slightly exsert, and relatively narrow, with smooth inner edges. Remaining septal cycles have

laciniate inner edges. Columella well developed and elongate, flat on top, and fused to lower, inner edges of the

S 1 -2-

Remarks. — Few specimens have been collected with attached buds, the buds apparently detaching at a

relatively small size.

This species has been found to host the petrarcid ascothoracidan Zibrowia auriculata Grygier, 1985, the gall of

which causes the columella to become larger and more porous than normal (ZIBROWIUS & Grygier, 1985).

B. carinata is more fully described and illustrated by ZIBROWIUS (1985).

Distribution. — Philippines : Visayan Sea; Bohol Sea; Sulu Sea (Basilan and Sulu Archipelago); 33-84 m.

Indonesia: Makassar Strait; Pleistocene of Talaud (Umbgrove, 1938); Banda Sea (Banda and Kai Islands); Flores

Sea (Selayar Island); Bali Strait; Sunda Strait, Java Sea; 45-100 m. Elsewhere: tropical Indo-West Pacific,

including Somalia, northern Indian Ocean, and Chesterfield Islands; 55-95 m.

Balanophyllia stimpsonii (Verrill, 1865)

Eupsammia stimpsonii Verrill, 1865: 150.

Eupsammia stimpsoniana Verrill, 1866: 29, pi. 2, figs 3, 3a.

Rhodopsammia socialis Semper, 1872: 260-261, pi. 20, figs 1-4. — FAUSTINO, 1927: 229, pi. 75, figs 9-12.

Rhodopsammia affinis Semper, 1872: 261-262, pi. 19, figs 7a-b.

Rhodopsammia incerta Semper, 1872: 264, pi. 19, figs 8a-b. — Faustino, 1927: 231, pi. 75, figs 3-4.

Leptopsammia conica van der Horst, 1922: 68-69, pi. 8, figs 14-15.

Balanophyllia affinis - Faustino, 1927: 228-232, pi. 75, figs 1-2. — van DER Horst, 1922: 62.

Balanophyllia stimpsonii - ZIBROWIUS, 1985: 234-235, figs 1-14 (synonymy). — Cairns & Keller, 1993: 274.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5133, 17 (USNM 97561). — Stn 5137, 1 (USNM

97562). — Stn 5143, 2 (USNM 97563). — Stn 5156, 1 (USNM 97564). — Stn 5164, 12 (USNM 97565).

Musorstom 2: stn 9, 1 (MNHN).

Indonesia. Deki: stn 10. 1 (NNM 17375). — Stn 14, 2 (NNM 17376). — Stn 18, 10 (NNM 17377). — Stn 30,

1 (ZMA Coel. 5479). — Stn 64, 4 (NNM 17369). — Stn 68, 1 (NNM 17368). — Stn 82, 100+ (NNM 17370). — Stn 84,

1 (NNM 17389). — Stn 100, 1 (NNM 17372).

Mortensens Java-S.A. Expedition: stn 5, 14 (ZMUC). — Stn 8, 2 (ZMUC).

"Te Vega": stn 1-54, 12 (USNM 97566).

Snellius 2: stn 4.228. 2 (NNM 17384). — Stn 4.232, 2 (NNM 17383). — Stn 4.234, 10 (NNM 17382). —

Stn 4.235, 4 (NNM 17381).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

177

Type Locality. — "North China Sea" (depth not given).

DIAGNOSIS. — Corallum unattached, straight to slightly curved, and ceratoid. Calice circular to elliptical:

CCD LCD =10-14 Thecal faces evenly rounded (corallum not highly compressed or keeled), bearing buds (some

upTo 20 mm in height), that originate randomly on theca. Tip of broken base 1.3-1.6 mm in dm meten M^st

specimens 9-10 mm in GCD, but ZiBROWlUS (1985) reported a specimen 15 mm in GCD. Septa hexamerally

arranged in 4 complete cycles, only larger specimens having some pairs of S5 i (uF 'to 3 l°tal of 66 _ septa .

Sl largest septa: slightly exsert and broad, having smooth, vertical inner edges. S2 about 3/4 width of an Si

having smooth inner edges. S3 only 1/4 width of an S2, having dentate to lacin.ate inner edges, each pair bending

toward their common S2. S4 rudimentary, each pair bending slightly toward their common S3. Fossa deep,

columella small, composed of several twisted laths.

REMARKS. - Balanophyllia stimpsonii differs from the other unattached western Pacific species,

(Se™r 1872, in having a less compressed corallum le„ a lower GCD:LCD and no thecal keels,; buddrng from

the'entire circumference of the corallum, no. jus, from keeled thecal edges; and & that are smaller than St.

R stimpsonii is more fully described and illustrated by ZiBROWlUS (1985).

ZZTcarLa, B. stimpsonii is also known to be a host for the pe, rated asco.hor.cdan genus Ztbrowta

Grygier, 1985, which produces a gall in the columella (ZiBROWlUS & GRYGIER, 1985).

DISTRIBUTION. — Philippines: Lubang Island; Bohol Sea; Sulu Sea (Mindanao and Sulu Archipelago);

18-70 m. Indonesia: Halmahera Sea (Gulf of Kau); Banda Sea (Kai Islands and southeastern

(Sumbawa and Selayar Island); Bali Strait; 35-75 m. Elsewhere: widespread throughout tropical Indo-West Pacific,

from southwestern Indian Ocean to Chesterfield Islands; 18-95 m.

Balanophyllia desmophyllioides Vaughan,

Figs 23 g-h

Balanophyllia desmophyllioides Vaughan, 1907: 149-150, pi. 45. fig. 1.

Balanophyllia destnophylloides - CAIRNS, 1984: 26.

1907

Philippines. MUSORSTOM 1: stn 3, 1 (MNHN). - Stn 63, 1 (MNHN). - Stn 65,

— Stn 32, 2 (MNHN). — Stn 49,

Material examined.

5 (UMusorstom82: stn 17, 1 (USNM 97569). - Stn 33, 10 (MNHN).

Indonesia. "Galathea": stn 488, 1 (ZMUC).

"Hakuho Mam": stn KH85-1-A2, 2 (USNM 97571).

CORINDON 2: stn 266, 2 (MNHN). msim

Karubar: stn 13, 1 (MNHN). — Stn 18, 3: 2 (MNHN). 1 (USNM 97570).

1 (POLIPI).

TYPE LOCALITY. - "Albatross" stn 4061: 20°16’ 10”N, 155°53'20”W (Hawaii), 44-152 m.

DESCRIPTION - Corallum elongate-conical to trochoid, the adult tending to become strongly compressed

(GCD LCD = 1 37- 1.90) on a robust, cylindrical pedicel (PD:GCD = 0.24-0.39). Largest known specimen

MUSORSTOM 1 stn 3) 18.1 x 35.1 mm in calicular diameter and 42.0 mm in height, with a pedicel diameter o

1 1 .0 mm. Profile of thecal face highly arched. Costae of lower corallum well defined tclturcNo

striae, but costal definition less clear in upper corallum, the theca being highly porous and granular texture.

eP1SeeCpm hexamerally arranged in 5 cycles, the 5th complete at a GCD of 15-19 mm.

independent, slightly exsert (about 2.2 mm), and quite thick and porous at the,r ^Per/ re2Uiar each

smooth in upper 1/2 of fossa, changing to coarsely dentate m lower 1/2 of

rectangular tooth about 0.3 mm wide. S4 smallest septa, about 1.3 mm exsert, 0.6 mm duck * ™

inner margin that attenuates about 1/2 distance down fossa. S5 and S4 equally exsert and thick, a pair of S5 fusing

Source :

178

S. D. CAIRNS & H. ZIBROWIUS

before each S4 high in fossa and extending to the columella as 1 septum. Lower inner edge of combined S5 dentate

like the Si -3. Fossa deep and spacious. Columella small in relation to size of corallum, composed of a narrow

(about 1.7 mm wide), elongate field of many, small (0.15 mm diameter) papillae that are weakly swirled in a

clockwise vortex. Columella low, but discrete and slightly convex.

Remarks. — Balanophyllia desmophyllioides is distinguished from other species by its highly arched calice;

its coarsely dentate S1-3 and S5; and the fusion of each pair of S5 high in the calice before their common S4. Some

of the specimens reported above are much larger than the holotype and subsequently reported specimens (Cairns,

1984), the holotype being only 15.0 mm in GCD and somewhat irregular in shape.

In our opinion, the species name was originally incorrectly formed, being derived from the root

Desmophyllum, and thus should be desmophylloides. However, according to the 1CZN (Article 32dii) the original

spelling cannot be changed.

Distribution. — Philippines: Lubang Island and Verde Island Passage, Luzon; 122-194 m. Indonesia:

Makassar Strait; Banda Sea (Kai Islands); Arafura Sea (east of Tanimbar Islands); Flores Sea (southwestern

Sulawesi); Bali Strait; 95-393 m. Elsewhere: Hawaiian Islands; 101-658 m.

Balanophyllia cornu Moseley, 1881

Figs 24 d-f

Balanophyllia cornu Moseley, 1881: 192-193, pi. 12, figs 11-15. — Alcock, 1902c: 41. — Cairns, 1994: 82-83

pi. 35, figs f-i (synonymy).

Not Balanophyllia cornu Sokolow, 1894: 88-91 (junior primary homonym).

Not Balanophyllia cornu - Cairns, 1984: 26 (= ? B. gigas Moseley, 1881).

Material EXAMINED. — Philippines. "Albatross": stn 5110, 5 (USNM 97573). — Stn 5268, 1 (USNM 97573).

— Stn 5280, 3 (USNM 97574). — Stn 5281, 1 (USNM M230138). — Stn 5367, 1 (USNM 97575) — Stn 5391

16 (USNM 97576). — Stn 5392, 100+ (USNM 97577). — Stn 5393, 1 (USNM 97578).

Musorstom 1: stn 3, 3 (USNM 97579). — Stn 14, 1 (MNHN). — Stn 61, 1 (MNHN). — Stn 63, 2 (USNM 97580)

Musorstom 2: stn 15, 4 (USNM 97582). — Stn 32, 7 (MNHN). — Stn 68, 1 (MNHN).

Musorstom 3: stn 88, 7 (MNHN). — Stn 96, 1 (MNHN). — Stn 100, 1 (MNHN). — Stn 102, 4 (MNHN) _ Stn 105

5 (MNHN). — Stn 108, 6: 1 (MNHN), 5 (USNM 97584). — Stn 120, 1 (USNM 97585). — Stn 126, 1 (USNM 97586). —

Stn 133, 1 (USNM 97587).

Indonesia. "Challenger": stn 192, 4 syntypes (BMNH 1880.11.25.143).

"Siboga": stn 297, 1 (ZMA 5482).

Deki: stn 7, 2 (ZMA Coel. 5484). — Stn 48, 4 (NNM 17332). — Stn 49, 30 (NNM 17532)

Mortensens Java-S.A. Expedition: stn 15, 1 (ZMUC).

"Galathea": stn 500, 8 (ZMUC).

"Hakuho Man,": stn KH72-1-28, 2 (USNM 97590). — Stn KH85-1-A2, 1 (USNM 97591)

Karubar: stn 7, 5 (POLIPI). - Stn 16, 1 (USNM 97588). — Stn 18, 4 (MNHN). - Stn 61, 3 (MNHN).

Type Locality. — "Challenger" stn 192: 5°49'15"S, 132°14'15"E (Kai Islands, Banda Sea), 256 m.

Description. — Corallum elongate-conical, with a slightly compressed calice (GCD:LCD = 1.1 -1.3). Two

growth forms occur: one having a straight, firmly attached (PD:GCD = 0.32-0.46) corallum, characteristic of the

type series; the other a curved (ceratoid), usually free corallum (PD:GCD = 0.12-0.21). The straight form has a

broadly encrusting base, whereas the curved form is either unattached or attached to a small object, such as a small

gastropod shell. The differences in growth form may simply be the result of the kind of substratum available on

which to settle, the costal and calicular characteristics being otherwise the same. Largest straight form

( usorstom 3 stn 105) 19.5 x 23.1 mm in calicular diameter and 34.6 mm in height, with a pedicel diameter of

T9 mm; largest specimen of ceratoid form (" Albatross " stn 5313) 22.9 x 27.5 mm in calicular diameter and

36.9 mm in height, with a pedicel diameter of 4.5 mm. Costae flat, 0.5-0.7 mm in width, and well defined by

deep, mow (about 0.1 mm) intercostal striae. Costae a reticulum covered by small, irregularly-shaped granules.

Lateral faces of costae also bear small granules that project into intercostal striae. Usually no epitheca, but when

present, restricted to lower pedicel.

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

179

Septa hexamerally arranged in up to 5 cycles: the 3rd (24 septa) complete at a GCD of about 8 mm; the 4th (48

septa) at 1 1-12 mm GCD; half of the 5th cycle (72 septa) at a GCD of 15-19 mm; and a full 5th cycle (96 septa)

at about 24 mm GCD. Si -2 essentially the same size, only slightly exsert, having smooth, vertical inner edges

that extend to the columella. S3 about 2/3 width of S1-2, also having smooth inner edges, which do not reach the

columella. In a corallum with 72 septa, which is the most common complement, the quarter-systems adjacent to

each S2 contain only 1 S4, whereas the quarter-systems adjacent to the Si contain 3 septa: 1 S4 and a pair of S5.

S4 adjacent to S2 are approximately the same size as the S5 adjacent to Si in the same half-system, the inner edges

of these 2 septa fusing near the columella. Inner edges of higher cycle septa also smooth. Fossa of moderate depth,

containing a well-developed, convex (discrete), elongate columella composed of many short lamellar elements

swirled in a clockwise direction. In some specimens the inner edges of the 4 lateral Si constrict the elongate

columella into 3 connected nodes.

DISTRIBUTION. — Philippines : Lubang Island; Verde Island Passage; Samar Sea; Sulu Sea (Semirara Islands);

185-368 m. Indonesia : Banda Sea (Kai Islands); Arafura Sea (east and southeast of Tanimbar Islands); Timor Sea

(south of Leti Islands and southwestern Timor); Flores Sea (southwestern Sulawesi); Bali Strait; 196-520 m.

Elsewhere: South China Sea (north of Pratas Islands); Formosa Strait; Japan (Honshu and Kyushu); 60-274 m.

Balanophyllia gemma (Moseley, 1881)

Figs 24 g-i

Thecopsammia gemma Moseley, 1881: 195, pi. 15, figs 8a-b.

Balanophyllia (Thecopsammia) gemma - ALCOCK, 1902c: 42. — Faustino, 1927: 223-224, pi. 73, figs 5-7.

Balanophyllia sp. - VAN DER HORST, 1922: 64 (in part: 1 of 3 specimens from "Siboga" stn 95).

Not Balanophyllia gemma - VAN DER HORST, 1926: 50, pi. 3, figs 12-13. — Cairns & KELLER, 1993: 221

(= Balanophyllia sp.).

Material EXAMINED. — Philippines. "Challenger": stn 201, holotype (BMNH 1880.11.25.147).

"Siboga": stn 95, 1 (ZMA Coel. 1166a).

"Albatross": stn 5135, 3 (USNM 97592).

Musorstom 2: stn 33, 3 (USNM 97593).

Indonesia. Deki: stn 46, 1 (NNM 17612). — Stn 59, 2 (NNM 17335).

Karubar: stn 49, 2 (POLIPI). — Stn 50, 8 (MNHN).

Type Locality. — "Challenger" stn 201: 7°03'N, 121°48'E (Sulu Sea, off Zamboanga Peninsula), 187 m.

DESCRIPTION. — Corallum cylindrical, straight, and firmly attached through a thick pedicel (PD:GCD =

0.70-0.75) and slightly expansive base. Largest known specimen (" Albatross " stn 5135) 8.7 x 9.6 mm in cahcular

diameter and 17.4 mm in height. A thick transversely corrugated epitheca extends to within 2-5 mm of calicular

edge. Costae equal in width (0.5-0.6 mm), flat, and highly porous, each costa uniformly covered with small

spines. Intercostal striae thin (0.05-0.10 mm), straight, and shallow.

Septa hexamerally arranged in 4 complete cycles, all 48 septa equally nonexsert, having smooth inner edges.

Si -2 equal in width, extending to columella. S3 about 1/2 width of S 1 -2- S4 that are adjacent to Si fuse to that Si

at calicular edge, their inner edges extending to the columella. Conversely, S4 that are adjacent to S2 also fuse to

that S2, each S4 bending toward the other S4 within its half-system, but not fusing with it and not quite reaching

the columella. Fossa shallow. Columella a discrete, elongate structure composed of short lamellae swirled in a

clockwise direction.

Remarks. — Balanophyllia gemma resembles a small, firmly attached specimen of B. cornu Moseley, 1881,

but can be distinguished by its well-developed epitheca, shallow fossa, and nonexsert septa.

Distribution. — Philippines: Verde Island Passage; Sulu Sea (Zamboanga Peninsula, Mindanao; Sulu

Archipelago); Basilan Strait; 137-294 m. Indonesia: Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar

Islands); Timor Sea (southwestern Timor); 185-522 m.

Source :

180

S. D. CAIRNS & H. ZIBROWIUS

Balanophyllia parvula Moseley, 1881

Figs 23 i, 24 a

Balanophyllia parvula Moseley, 1881: 194-195, pi. 15, figs 9, 9a. — ALCOCK, 1902c: 41. — Faustino, 1927' 234

pi. 73, figs 3-4.

MATERIAL EXAMINED. — Philippines. " Challenger stn 201, 2 synlypes (BMNH).

Musorstom 2: stn 32, 1 (MNHN).

Musorstom 3: stn 108, 6 (MNHN).

Indonesia. "Siboga": stn 251, 1 (NNM 22429).

Deki: stn 46, 1 (NNM 17334).

Karubar: stn 18, 1 (MNHN). — Stn 49, 9: 4 (MNHN), 5 (USNM 97596).

South China Sea. " Albatross stn 5310, 1 (USNM 97594).

Type Locality. — "Challenger" stn 201: 7°03'N, 121°48'E (Sulu Sea, off Zamboanga Peninsula), 187 m.

DESCRIPTION. — Corallum elongate-conical and straight, having a slightly compressed calice (GCD:LCD =

1.14-1.23) and attached by a robust pedicel (PD:GCD = 0.32-0.49). Largest known specimen (Karubar stn 49)

10.9 x 13.4 mm in calicular diameter and 18.4 mm in height, with a pedicel diameter of 4.4 mm. Costae quite

porous and well defined by narrow intercostal striae; however, lower 1/4 to 1/3 of corallum covered by a thin

epitheca.

Septa hexamerally arranged in 4 complete cycles in a strongly developed Pourtales plan. Si highly exsert (up to

3.0 mm), remarkably thick (up to 1.6 mm at calicular edge), having smooth, vertical inner edges that attain the

columella. S2 less exsert (up to 2.0 mm), only about 1/2 thickness of Si, and about 3/4 width of Si. Inner edges

of S2 also entire but do not quite attain the columella. S3 rudimentary, only about 0.7 mm exsert, 0.4 mm in

thickness, and about 1/4 width of an S2. S4 dimorphic in size: those adjacent to Si highly exsert (up to 2.8 mm),

fused to adjacent Si in robust triangular lancets; S4 adjacent to S2 slightly less exsert (up to 1.2 mm), also fused

to adjacent S2 but in less prominent lancets. Pairs of S4 unite before their common S3 high in fossa and extend to

the columella, sometimes fusing with inner edges of other S4 within their system near the columella. Inner edges

of S4 laciniate, bearing tall, regularly spaced teeth up to 0.6 mm tall. Fossa deep; columella rudimentary.

Remarks. — Balanophyllia parvula is distinguished from other Indo-West Pacific congeners by its remarkably

thick and exsert Si, which contribute to form tall, triangular calicular lancets. It is also distinguished in having a

strongly developed Pourtales plan, laciniate S4, and a rudimentary columella.

Moseley (1881) reported 3 specimens (syntypes) from "Challenger " stn 201 in his original description of

Balanophyllia parvula , 2 juveniles and 1 adult, the adult of GCD 8.0 mm being the figured specimen Both

juveniles were present at the BMNH in 1994, but the adult specimen could not be found. Judging from Moseley's

figure, B. parvula is a distinctive species represented by additional specimens reported above, but the 2 smaller

juvenile (GCD 4.4 and 4.8 mm) syntypes appear to be a different species, perhaps founder corallites of

a Rhizopsammia. Because of the possibility of 2 species being represented in Moseley’s type-series, the large

figured specimen is designated as the lectotype, even though it could not be found in 1994. ALCOCK (1902c)

reported 1 specimen of B. parvula from " Siboga " stn 251. A specimen at the NNM from this station fits

Moseley s figured B. parvula , but another Balanophyllia (species indet.) is also known from that station (ZMA).

Distribution. Philippines: Verde Island Passage; Sulu Sea (Zamboanga Peninsula); 192-195 m.

Indonesia : Banda Sea (Kai Islands); Arafura Sea (southeast of Tanimbar Islands); 206-300 m. Elsewhere • South

China Sea (north of Pratas Islands); 183 m.

Balanophyllia crassiseptum sp. nov.

Figs 25 a-c

MATERIAL EXAMINED/TYPES. — Philippines. Musorstom 1: stn 32, 3 paratypes (MNHN).

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

181

Indonesia. Deki: stn 7, 1 paratype (ZMA). — Stn 63, 1 paratype (NNM 17333).

Karubar: stn 49, 74 paratypes: 27 (MNHN), 3 (POLIPI), 44 (USNM 97654). — Stn 50, holotype (MNHN) and

32 paratypes (MNHN).

TYPE Locality. — Karubar stn 50: 7°59'09"S, 133°01'56"E (Arafura Sea southeast of Tanimbar Islands),

184-185 m.

ETYMOLOGY. — The species name crassiseptum (Latin crassus , thick + septum, literally fence or bar), alludes

to the thick septa of this species. The name is treated as a noun in apposition.

DESCRIPTION. — Corallum elongate-conical, straight, and compressed (GCD:LCD = 1.15-1.45). Pedicel

robust : PD:GCD = 0.45-0.55. Largest known specimen (holotype) 10.2 x 14.8 mm in calicular diameter,

17.0 mm in height, and 8.2 mm in pedicel diameter. Costae flat and porous, of variable width, and separated by

thin intercostal striae that sharply mark the costal boundaries. A thin epitheca covers basal 20-50% of corallum,

which is usually highly encrusted. Corallum white to light purple, the latter colour caused by an endolithic

microorganism.

Septa hexamerally arranged in 4 complete cycles. Si remarkably thick, as much as 1.5 mm at calicular edge.

Si quite exsert (up to 2.3 mm), porous, having a smooth inner edge that attains the columella. S2 also quite thick

(up to 0.9 mm) but only 1/2 as much as the Si. S2 up to 1.3 mm exsert, also porous, their inner edges also

attaining the columella. S3-4 0.5-0.6 mm thick. S3 rudimentary. S4 adjacent to Si more exsert than S2, but

S4 adjacent to S2 are as exsert as S3. Inner edges of S4 finely dentate. Fossa shallow, containing a well-developed,

discrete papillose columella.

Remarks. — Balanophyllia crassiseptum is very similar to B. parvula Moseley, 1881, both species having

very thick Si -2 and sometimes are found at the same stations. B. crassiseptum differs in having a larger columella,

less dentate inner edges of S4, and S2 that extend completely to the columella, not only 1/2 that distance as in

B. parvula.

The corallum of the holotype is bored by an acrothoracidan cirripede (Fig. 25 c) (see Grygier & Newman,

1985).

Distribution. — Philippines : Lubang Island; 183-193 m. Indonesia: Banda Sea (Kai Islands); Arafura Sea

(southeast of Tanimbar Islands); 185-250 m.

Balanophyllia rediviva Moseley, 1881

Figs 25 d-f

Balanophyllia rediviva Moseley, 1881: 193-194, pi. 15, figs 10-12. — van DER Horst, 1922: 59.

? Balanophyllia rediviva UMBGROVE, 1938: 273.

MATERIAL EXAMINED. — Philippines. Musorstom 1: stn 3, 1 (MNHN). — Stn 57, 1 (MNHN).

Musorstom 2: stn 33, 6 (MNHN).

Musorstom 3: stn 117, 3 (USNM 97598).

Indonesia. "Challenger": stn 192, syntypes (BMNH 1880.11.25.145).

Corindon 2: stn 248, 1 (MNHN).

Karubar: stn 22, 1 (MNHN).

Type Locality. — "Challenger" stn 192: 5°49T5"S, 132°14T5"E (Kai Islands, Banda Sea), 256 m.

Description. — Largest known corallum (Karubar stn 22) cylindrical, elongate and slightly bent: 8.3 x

9.5 mm in calicular diameter and 71 mm in length, having 3 episodes of rejuvenescence. Epitheca extends to

within 9-10 mm of calicular edge, but most of theca worn and covered with foraminifera and serpulid tubes. Theca

highly porous. C1-3 thin (about 0.3 mm) and slightly ridged, whereas C4 are much broader (about 0.7 mm) and

flat.

Source :

182

S. D. CAIRNS & H. ZIBROWIUS

Septa hexamerally arranged in 4 complete cycles. Si highly exsert (2.1 mm), having smooth inner edges that

extend to columella. S2 much less exsert (0.7 mm), about 4/5 width of an Si, also having entire inner edges.

S3 little exsert (0.3 mm), only about 1/3 width of an S2, having entire inner edges. S4 adjacent to Si highly

exsert (about 1.3 mm), at the calicular edge strongly fused to adjacent Si in triangular lancets. S4 adjacent to S2

considerably less exsert (0.3 mm) than those adjacent to Si, the inner edges of each pair of S4 within a half-system

fusing before its common S3 near the columella. Inner edges of S4 regularly dentate. Fossa of moderate depth,

containing a discrete, elongate, spongy columella that is sometimes constricted by the lower, inner edges of the

4 lateral Si.

Remarks. — Balanophyllia rediviva is similar to B. laysanensis Vaughan, 1907, in calicular features, but can

be distinguished by its elongate, cylindrical corallum; slightly ridged C1-3; and dentate S4.

Only 3 of Moseley's 4 syntypes could be located at the BMNH in 1994: the specimen illustrated

as Moseley's (1881) pi. 15, figs lOa-b is the only well-preserved corallum and is here designated as the lectotype.

DISTRIBUTION. — Philippines'- Lubang Island; Verde Island Passage; Mindoro Strait; 97-183 m. Indonesia :

Makassar Strait; Banda Sea (Kai Islands); 90-235 m. ? Holocene of Talaud Islands (Umbgrove, 1938).

Balanophyllia gigas Moseley, 1881

Balanophyllia gigas Moseley, 1881: 193. — VAN DER Horst, 1922: 58-59, pi. 8, fig. 22. — Cairns, 1994: 83, pi. 35,

figs j-1 (synonymy); 1995: 119-120, pi. 40, figs f-h (synonymy).

Material EXAMINED. — Philippines. MUSORSTOM 2: stn 1, 1 (MNHN). — Stn 27, 1 (USNM 97600).

Indonesia. Deki: stn 50, 1 (NNM).

Karubar: stn 13, 1 (POLIPI).

Type Locality. — Japan (depth not given).

Diagnosis. — Corallum ceratoid, often bent (up to 90°), and quite large. Largest Indonesian specimen (van

der Horst, 1922) 30 x 24 mm in calicular diameter, but some New Zealand specimens (Cairns, 1995) are larger

still. Pedicel robust: PD:GCD = 0.30-0.57. Basal 1/2 to 2/3 of theca epithecate and usually covered with

encrusting organisms. Costae flat, quite porous, and equal in width. Septa hexamerally arranged in 5 cycles in a

Pourtales plan, the 5th cycle rarely complete. S 1 -2 are 2-4 mm exsert, along with adjacent higher cycle septa

forming 12 triangular calicular lancets. S3 about 3/4 width of S1-2, the septa of all 3 cycles being independent,

smooth-edged, and reaching the columella; S4-5 have laciniate inner edges. Fossa deep. Columella discrete,

composed of short, swirled lamellar elements.

Remarks. — This species is characterised by attaining a large size (up to 79 mm in height and 33 mm in

calicular diameter), having epitheca, and usually having a bent corallum. It is more fully described and illustrated

by Cairns (1994, 1995).

Distribution. — Philippines : Lubang Island; Verde Island Passage; 100-188 m. Indonesia : Banda Sea

(Kai Islands); 90-393 m. Elsewhere : Japan (Honshu and Kyushu); Hawaiian Islands; New Zealand; 90-640 m.

Balanophyllia serrata sp. nov.

Figs 24 b-c

MATERIAL EXAMINED. — Philippines. Musorstom 1: stn 27, 2 paratypes (USNM 97601). — Stn 63, 1 paratvpe

(MNHN). — Stn 65, 4 paratypes (USNM 97602). — Stn 69, holotype (MNHN).

Musorstom 3: stn 108, 1 paratype (MNHN).

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

183

TYPE Locality. — Musorstom 1 stn 69: 13°58.8'N, 120°17.3'E (north of Lubang Island, Philippines),

187-199 m.

ETYMOLOGY. — The species name (Latin serratus, toothed like a saw) refers to the jagged calicular edge of

this species.

DESCRIPTION. — Corallum large, elongate-conical to trochoid, straight, and slightly flared distally. Largest

specimen (the holotype) 34.1 x 27.1 mm in calicular diameter and 45 mm in height, with a pedicel diameter of

17.3 mm. Pedicel robust: PD:GCD = 0.27-0.51; base encrusting. Calice elliptical: GCD:LCD = 1.17-1.37.

Costae 0.7-0. 9 mm wide, flat, and covered with small spines arranged 3 or 4 across the width of each costa.

Intercostal striae thin (0.2 mm) and deep, contributing to a highly porous theca. No epitheca, but lower pedicel of

large specimens covered with a solid stereome, obscuring the porous nature of the theca. No encrusting organisms

noted on stereome. , _ . . , . .

Septa hexamerally arranged in 5 cycles, a specimen of 27 mm GCD having a complete 5th cycle, but the

largest specimen of 34 mm GCD with only 88 septa (4 pairs of S5 not formed). Si -2 highly exsert:

Si 4.0-5. 5 mm exsert, S2 3-4 mm exsert. Both Si and S2 are independent septa, quite thick at the calicular edge

(up to 1.9 mm), having smooth inner edges that extend inward to the columella. S3 much less exsert, about

3/4 width of the S1-2, not reaching the columella. S4 approximately 1/2 width of an S3. S5 adjacent to lower order

septa highly exsert, fusing at the calicular margin to their adjacent lower order septa, and forming calicular lancets.

Lower inner edges of the 2 S5 within each half-system that are adjacent to Si -2 meet and fuse near columella.

Conversely, the 2 S5 within each half-system that are adjacent to S3 are the smallest septa (less wide than an S4)

and little exsert, their inner edges bending toward and often fusing to adjacent S4. S5 usually porous near the theca,

having coarsely dentate inner edges. Fossa of moderate depth. Columella rudimentary, elongate, and constricted by

lower, inner edges of lateral Si. Columella discrete, composed of short lamellae.

REMARKS. — This species has a large corallum and septal arrangement similar to that of B. gigas Moseley,

1881. It differs in having no epitheca, a slightly flared calice with more highly exsert Si -2, and a straight corallum.

Distribution. — Philippines: north of Lubang Island; 190-194 m.

Balanophyllia generatrix sp. nov.

Figs 25 g-i, 26 a-b

MATERIAL EXAMINED/TYPES. — Philippines. " Siboga stn 102, 1 corallum, paratype (ZMA Coel. 5483).

"Albatross": stn 5543, 1 quasicolony, paratype (USNM 97603). ,7U. . , ,.8Q,

Indonesia. "Siboga": stn 41, 1 quasicolony (ZMA Coel. 5538) and several isolated corallites (ZMA Coel. 5489),

ParDEKL stn 12, 1 quasicolony, paratype (NNM ). - Stn 58, 2 quasicolonies, paralyses (ZMA Coel. 5493)

Karubar: stn 16, 1 quasicolony, paratype (MNHN). - Stn 82, 1 quasicolony (holotype, MNHN) and 1 additional

quasicolony and several isolated coralla, paratypes (USNM 97604).

Type Locality. — Karubar stn 82: 9°30'00"S, 13r02'41"E (Arafura Sea south of Tanimbar Islands),

215-218 m.

Etymology. — The species name ( generatrix , Latin for "the one that produces offspring ) was the

unpublished museum (ZMA) name used by van DER HORST, who believed the species to be truly colonial. In fact,

the quasicolony results from contiguous independent planular settlement.

Description. — Corallum elongate-conical, usually occurring in a quasicolonial structure, i.e., individual

planulae settle close to one another, usually on the theca of an older or dead conspecific corallum, not as the result

of budding. The central individual may release the planulae that eventually colonize its theca, but in no case does

a corallum appear to bud from the theca of another living corallum. The bases of closely adjacent coralla often

Source :

184

S. D. CAIRNS & H. ZIBROWIUS

coalesce (fuse) appearing as though they have a common basal coenosteum, but this is not due to coloniality. The

holotype is such a quasicolony consisting of a dead central corallum 20.1 x 33.7 mm in calicular diameter and

56 mm in height, on which 28 individual coralla have settled, one as tall as 45 mm. Coralla may attain a height

of 70 mm (" Siboga " specimens) but never bud additional coralla. The calice is circular to elliptical in cross section

(GCD:LCD = 1.1-1.95), the more elliptical calices characteristic of larger coralla. Pedicel robust, the PD:GCD

being about 0.33. Costae poorly defined, most of the porous theca covered with a fine spination; no epitheca.

Septa hexamerally arranged in 5 full cycles, passing through the 4 cycle stage at a GCD of 6-9 mm. Si -2 equal

in size, only very slightly exsert (about 1 mm), having finely dentate lower, inner edges. S3 similar to S 1-2, also

being independent, but their inner edges do not reach quite as far into the fossa as the Si -2. S4 rudimentary. Pairs

of S5 fuse fairly high in the fossa, having coarsely dentate inner edges that extend as far toward the columella as do

those of the Si -2. Fossa quite deep, containing an elongate, discrete columella composed of many very small

papillae.

Remarks. — Balanophyllia generatrix is similar to B. gigas Moseley, 1881, both having large coralla and

5 cycles of septa, but B. generatrix is distinguished by its quasicolonial habit, very deep fossa, finer columella,

and coarse dentition of the S5.

One quasicolony from "Siboga" stn 41 and 2 from Deki stn 58 were found with van der Horst labels reading

Dendrophyllia generatrix , the specimen from the "Siboga" station labelled as the "type", the others as a "cotype".

Independently, specimens from the "Albatross" and Karubar expeditions had been segregated by the first author as

an unidentified species. Van der Horst's unpublished manuscript name is adopted for it herein. It is

understandable that van der Horst considered this species to be a colony resulting from budding and thus called it

a Dendrophyllia, but for reasons explained in the species description it is here described as a Balanophyllia.

Distribution. — Philippines : Bohol Sea; Sulu Archipelago; 296-535 m. Indonesia : Banda Sea (Kai Islands);

Arafura Sea (south of Tanimbar Islands); Flores Sea; 96-385 m.

Balanophyllia imperialis Saville Kent, 1871

Figs 26 c-f

Balanophyllia imperialis Saville Kent, 1871: 284, pi. 23, figs 5a-b. — van DER Horst, 1922: 60-61 (in part: "Siboga"

stn 153, pi. 8, fig. 25). — Faustino, 1927: 224. — Zou, 1988: 78. pi. 3, figs 1-14, pi. 4, figs 2a, 6a, 10a, 12a.

? Balanophyllia imperialis - HARRISON & POOLE, 1909: 905-906, pi. 86, figs 5a-c. — UMBGROVE, 1938: 272.

Balanophyllia sp. - Zibrowius & Grygier, 1985: 128, fig. 37.

Material EXAMINED. — Philippines. "Albatross": stn 5133, 1 (USNM 97605). — Stn 5146, 4 (USNM 97606)

— Stn 5174, 1 (USNM 97607).

Musorstom 3: stn 142, 5 (MNHN).

Indonesia. "Te Vega": stn 1-54, 7 (USNM 97609).

"Hakuho Maru"\ stn KH72-1-30, 8 (USNM 97608).

Corindon 2: stn 248, 1 (MNHN).

South China Sea. Holotype (BMNH 1984,4.27.3).

Type Locality. — Singapore, South China Sea (depth not given).

Description. — Corallum elongate-conical, straight to slightly curved, and attached by a slender pedicel:

PD:GCD = 0.09-0.18. Calice elliptical: GCD:LCD = 1.27-1.49. Largest known specimen (the holotype)

22.6 x 33.7 mm in calicular diameter and 40.7 mm in height, with a pedicel diameter of 3.3 mm. Costae well

defined by narrow, porous intercostal striae; C5 usually slightly wider than C1-4. Costae slightly convex and

covered with small spines. Usually no epitheca.

Septa hexamerally arranged in 5 cycles, but 5th cycle occasionally incomplete by a few to several pairs of S5,

even in large coralla. S1-2 independent, having smooth inner edges. Si about 2.1 mm exsert, S2 about 1.5 mm

exsert, neither forming calicular lancets. Septa arranged in a Pourtales plan, the inner edges of S3 smooth, those of

Source : MNHN . Paris

AZOOXANTHELLATE SCLERACTINIA

185

the S4-5 laciniate. Fossa deep, containing a discrete, elongate, robust (up to 2.8 mm in width) columella

consisting of tightly fused lamellar elements swirled in a clockwise direction.

REMARKS — The large size (GCD > 30 mm) and septal number of B. imperials invites comparison with

2 other species that have large coralla: B. gigas Moseley, 1881 and B. serrata sp. nov. B. imperials differs in

having a much narrower pedicel and in lacking calicular lancets. It is also characterised by having a very deep fossa

and a robust columella.

DISTRIBUTION. — Philippines : Visayan Sea; Sulu Sea (Zamboanga Peninsula and Sulu Archipelago),

27-70 m. Indonesia: Makassar Strait; Halmahera Sea; Timor Sea; 55-170 m; ? Holocene of Talaud Islands

(UMBGROVE, 1938). Elsewhere: South China Sea (Singapore); ? Mergui Archipelago; 18-38 m.

Genus ENDOPACHYS H. Milne Edwards & Haime, 1848

Endopachys grayi H. Milne Edwards & Haime, 1848

Endopachys grayi H. Milne Edwards & Haime, 1848b: 82-83, pi. 1 figs 2, 2a. - Semper 1872: 26T - van °j| Horst,

,922: 68. - FAUST, NO, 1927: 240-241, pi. 77, figs 1-2. - Umbgrove f f ^ P'- « q ^ 0; P ;

fig. 7. _ Zibrowius & Grygier, 1985: 128, figs 39-42. — Zou et al, 1988: 195 ~CA1™S, 1991: 24-25

figs i-j, pi. 11, figs a-b; 1994: 84-85, pi. 36, figs e, h, pi. 37, fig. i (synonymy); 1995: 121-122, pi. 41,

(synonymy). — Cairns & Keller, 1993: 276.

Endopachys weberi Alcock, 1902a: 109-110 [new synonym].

Endopachys sp. - van der Horst, 1922: 68, pi. 8, fig. 4 (same specimen as Alcock s E. weben).

pi. 10,

figs c-h

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5133, 6 (USNM 97610). —

97611). — Stn 5357, 1 (USNM 97612). — Stn 5593, 1 (USNM 97613).

Musorstom 1: stn 57, 2 (USNM 97614).

Musorstom 2: stn 29, 1 (MNHN).

Musorstom 3: stn 102, 1 (MNHN). — Stn 131, 3 (MNHN).

Indonesia. "Siboga": stn 51, 1 (holotype of E. weberi , ZMA Coel 7734)

Deki: stn 49, 2 (NNM 22714). — Unnumbered station, Ambon, 25-100 m. 3 (NNM 22715-1 /)

MORTENSEN'S Java-S.A. Expedition: stn 5, 6 (ZMUC). — Stn 6, 4 (ZMUC). — Stn 8, 1 (ZMUC).

Karubar: stn 2, 1 (USNM 97616).

Stn 5268, 1 (USNM

Type Locality. — Unknown.

Diagnosis. — Attached young stage rarely observed (holotype of E. weberi). Species best known trom the

unattached flabellate corallum, which results from budding. Corallum triangular in face view, the edge angle

(exclusive of lateral crests) 50°-80° and the face angle changing from a narrow 15°-28° basally to a more open

42°-57° distally. Largest known specimen 38.9 mm in GCD and 34 mm in height (Cairns & Keller, ),

largest Philippine corallum (MUSORSTOM 2 stn 29) 12.3 x 16.9 mm in calicular diameter and 15.2 mm in height

Edge crests delicate, about 0.5 mm in thickness and up to 4 mm in height, occasionally bearing 1 or more small

buds Costae and intercostal spaces not well defined; costae unridged. Septa hexamerally arranged in 5 cycles in

a Pourtales plan. Si-2 up to 2.5 mm exsert. Pairs of S5 within each quarter-system fuse before their common S4,

bearing a paliform lobe at this junction. Fossa deep; columella elongate, nondiscrete, and spongy.

Remarks. — The face angle of E. grayi is low initially and broadens with height, whereas that of E. bulbosa

is high as a juvenile and decreases with height. Other difference between the only two Recent species in this genus

are given in the Remarks of E. bulbosa. A more complete description and illustrations of E. grayi are found in

Cairns (1991, 1994, 1995).

Distribution. — Philippines: Lubang Island; Verde Island Passage; Sulu Sea (west of Panay, Zamboanga

Peninsula, and Balabac Island); 70-192 m. Indonesia: Halmahera Sea; Banda Sea (Kai and Ambon Islands); Bah Sea

Source :

186

S. D. CAIRNS & H. ZIBROWIUS

(Bali Strait and Madura Bay); 50-245 m. Elsewhere : Malaysia (Celebes Sea off Sabah); common in tropical and

warm temperate regions from southwest Indian Ocean to the Gulf of California, including South China Sea, Japan,

New Zealand, and the Hawaiian Islands; 37-386 m.

Endopachys bulbosa sp. nov.

Figs 27 a-g

Material EXAMINED/TYPES. — Indonesia. Karubar: stn 62, holotype and 1 paratype (MNHN) and 1 paratype

(USNM 97617). — Stn 67, 1 paratype (USNM 97618). — Stn 79, 1 paratype (MNHN).

Type Locality. — Karubar stn 62: 9°02'10"S, 132°43'05"E (Arafura Sea southeast of Tanimbar Islands),

239-250 m.

Etymology. — The species name (Latin bulbosa, bulbous or swollen) refers to the thick basal pads on the

corallum base.

Description. — Corallum unattached, flabellate (GCD:LCD = 1.32-1.41), and massive, the largest specimen

(the holotype) 33.7 x 45.7 mm in calicular diameter and 38.5 mm in height. Shape of corallum varies

characteristically with age, the theca of lower 1/3 to 1/2 of a large corallum being exceptionally thick (up to

4 mm) and having an edge angle of 142°-155° and a face angle of 90°-104°. However, 10-14 mm above base the

theca thins to about 1 mm, the edge angle decreases to 50°-55°, and the face angle to 40°-45°. Edge costae crested as

high as 7 mm, being 1.5 mm thick on lower corallum. All 12 Ci-2 highly ridged (up to 1.7 mm) in upper

corallum, but flat and very broad in region of basal thickening, the Ci being as much as 8 mm wide, the C2 as

much as 1.5 mm wide. C3-4 of thickened region only 0.3-0.5 mm wide; C3-5 of upper theca poorly defined.

Asexual budding not observed in type series.

Septa hexamerally arranged in 5 cycles. S1-2 highly exsert (up to 5 mm), with thick (2 mm) upper edges and

straight inner edges that attain the columella. S3 not exsert, about 2/3 width of an Si -2, their inner edges extending

almost to columella. S4 rudimentary, each S4 flanked by a pair of S5 that fuse before the S4 and extend to the

columella as 1 septum where each fuses with the other S5 within its half-system. No paliform lobes. Fossa of

moderate depth. Columella an elongate, discrete structure composed of many short lamellae swirled in a clockwise

direction.

Remarks. — The massive basal thecal thickening of this species is solid, not the exterior of an overly

enlarged internal cavity, and thus has the effect of weighting the corallum as though with ballast, which might

facilitate an upright or near-upright posture for the corallum. This basal thickening is not a characteristic of all

larger coralla of Endopachys, since large coralla of E. grayi H. Milne Edwards & Haime, 1848, of GCD 40 mm

do not have it. E. bulbosa also differs from E. grayi in having: well-defined, ridged C1-2; much more exsert S1-2;

a larger discrete, convex columella; and in lacking paliform lobes. E. maclurii (Lea, 1833), known from the

Eocene of southeastern U.S., has 6 strongly produced Ci, but nothing resembling the basal thickening of

E. bulbosa.

DISTRIBUTION. — Indonesia : Arafura Sea (south and southeast of Tanimbar Islands); 233-251 m.

Genus LEPTOPSAMMIA H. Milne Edwards & Haime, 1848

Leptopsammia slokesiana H. Milne Edwards & Haime, 1848

Figs 26 g-i

Leptopsammia stokesiana H. Milne Edwards & Haime, 1848b: 90, pi. 1, figs 4, 4a. — van der Horst, 1922: 68, pi. 8,

fig. 5. — Faustino, 1927: 242, pi. 77, figs 3-4.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

187

Balanophyllia stokesiana - Searles, 1956: 25, pi. 42, fig. c.

MATERIAL EXAMINED. — Philippines. MORTENSEN'S Pacific Expedition: Jolo Island, Sulu Archipelago, 46 m, 19

March 1914, 1 (NNM). Holotype, (BMNH 1855.12 27.1). ,, 3R im

Strait of Malacca. Malacca, depth unknown, Belcher collection, 1 (BMNH 1842.1 1.28. iu).

TYPE Locality. — Philippines (depth not given).

DESCRIPTION (Philippine specimen). — Corallum elongate-conical, straight, and firmly attached by a broad

pedicel: 7.3 x 9.1 mm in calicular diameter, 13.0 mm in height, and 4.6 mm in pedicel diameter. Porous theca

covered with low, rounded, granular costae; no epitheca. Septa hexamerally arranged in 5 cycles, the 5th

incomplete (a total of 54 septa): Si-2>S3>S4>S5. Si-2 nonexsert, having straight, smooth, vertical inner edges

that attain the columella. S3 about 3/4 width of Si-2, having straight, finely dentate inner edges S4 about 1/3

width of S3, having laciniate inner edges. S5 rudimentary. Columella discrete, composed of several tightly-tused

and twisted elements.

Remarks. — Only 5 specimens of this species are known: the holotype (BMNH 1855.12.27.1, GCD =

10 mm), a specimen reported by van DER HORST (1922) deposited at the ZMA (GCD = 17.6 mm), one reported

by SEARLE (1956) deposited at the USNM (78603), and the 2 specimens noted above. All 5 specimens are

remarkably similar in morphology.

DISTRIBUTION. — Philippines : Sulu Archipelago; 46 m. Indonesia: Flores Sea (Sumbawa); 69 m. Else¬

where : Malacca; depth unknown.

Leptopsammia crassa van der Horst, 1922

Figs 27 h-i

Leptopsammia crassa van der Horst, 1922: 69, pi. 8, figs 11-12.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5130, 1 (USNM 97621).

Indonesia. "Siboga": stn 258, holotype (ZMA Coel. 8462).

Type Locality. — "Siboga" stn 258: 5°26.6'S, 132°32.4'E (Kai Islands, Banda Sea), 22 m.

DIAGNOSIS (specimen from "Albatross" stn 5130). - Corallum 10.3 x 8.6 mm in calicular diameter, 9.7 mm

in height and 4.2 mm in pedicel diameter. Costae not well-defined. Porous theca 0.7 mm thick; no epitheca. Septa

hexamerally arranged in 4 cycles according to formula: S,-2>S3»S4. Si-2 nonexsert, having straight, vertical,

smooth inner edges. S3 2/3 width of Si-2, having laciniate inner edges. S4 rudimentary, having laciniate inner

edges that do not fuse with adjacent septa. Fossa of moderate depth, containing a small elongate columella

composed of several twisted elements.

Remarks _ The "Albatross" specimen was compared directly to the holotype (ZMA Coel. 8642) and

considered to be conspecific despite several differences. The " Albatross " specimen is slightly smaller, has thinner

theca, and has fewer septa, the holotype having a septal complement of: 15:15:30 (60 septa), which is thought to

be an aberration of a hexameral plan. Most other characters are the same.

Leptopsammia crassa differs from L. poculum (Alcock, 1902) (Figs 28 a-b), in havmgmdistmct costae (the

C 1-2 of L. poculum are slightly raised) and in having a stouter corallum. L. stokesiana H. Milne Edwards &

Haime, 1848, appears to differ from L. crassa in having some S5 and a costate theca. However, so few specimens

of Leptopsammia are known from the Indo-Pacific that a clear distinction among species is wanting.

DISTRIBUTION. — Philippines: Sulu Sea (Zamboanga Peninsula); 187 m. Indonesia: Banda Sea (Kai Islands);

22 m.

Source :

188

S. D. CAIRNS & H. ZIBROWIUS

Genus ENDOPSAMMIA H. Milne Edwards & Haime, 1848

Endopsammia philippensis H. Milne Edwards & Haime, 1848

Figs 28 c-e

Endopsammia philippensis H. Milne Edwards & Haime, 1848b: 91. pi. 1, figs 5, 5a; 1860: 108. — Faustino, 1927: 243-

244, pi. 77, figs 5-6. — Pillai & SCHEER, 1976: 71-72.

Thecopsammia regularis Gardiner, 1899: 169-170, pi. 19, fig. 8.

Balanophyllia regularis - van DER HORST, 1922: 63; 1926; 50, pi. 3, figs 10-11.

Endopsammia philippinensis (sic) - Wells, 1964: 118, pi. 2, figs 12-13. — CAIRNS, 1991: 26. — Cairns & KELLER,

1993: 221.

MATERIAL EXAMINED. — Philippines. Holotype, BMNH 1855.12.27.25.

Indonesia. Deki: unnumbered station, Damar Besar Island, Jakarta Bay, 4 (NNM 17518). — Unnumbered station,

Nyamuk Kecil Island, Jakarta Bay, 4 (NNM 17519). — Unnumbered station, Nuhucut Island, Kai Islands, 2-3 m, 5 (NNM

17520). — Unnumbered station, Sebesi Island, Lampung, Sumatra, Sunda Strait, 33 (NNM 17521).

Snellius 1: Potilyan Island, Pelokang, southwestern Sulawesi, 3 (NNM 17613).

Snellius 2: stn 4.070, 2 (NNM 17517).

Papua New Guinea: "Alpha Helix": stn M26, 1 (USNM 86818). — Stn M48, 2 (USNM 80018). — Stn M59,

I (USNM 88322).

Australia: Heron Island, Queensland, intertidal, 2 (USNM 83006).

Type Locality. — Philippines (depth not given).

Description. — Corallum subcylindrical and relatively small, the largest known specimen (USNM 83006)

8.4 x 9.3 mm in calicular diameter and 7.7 mm in height. Calice only slightly elliptical (GCD:LCD =

1.05-1.15); pedicel robust (PD:GCD = 0.68-0.77). Lower 3/4 of corallum usually covered with a thin epitheca,

which is often encrusted by algae or foraminifera. Theca near calicular edge highly porous and usually not costate.

Septa hexamerally arranged in 4 cycles but pairs of S4 often missing in smaller specimens. Si about 0.8 mm

exsert, having a smooth upper edge, which is dentate adjacent to columella. S2 less exsert, 1/2 to 2/3 width of an

Si, having slightly coarser inner edge dentition. S3 almost as wide as S2, having coarse to laciniate inner edges,

each pair of S3 within a half-system bending toward its common S2 and forming a loose fusion with that septum

adjacent to columella. S4 rudimentary, usually represented by only a row of tall spines that project from inner

theca. Fossa shallow to moderate in depth. Columella a well-developed, elliptical, spongy mass; nondiscrete, often

with a slightly concave upper surface.

Remarks. — H. Milne Edwards & Haime (1848b) undoubtedly intended to name this species

philippinensis (for the Philippine Islands), not philippensis (for Philippe), but according to the ICZN (Article

32cii), the original spelling cannot be changed.

Distribution. — Philippines: unspecified locality and depth (H. Milne Edwards & Haime, 1848b).

Indonesia: Banda Sea (Kai Islands); Flores Sea (Lintah Strait and Pelokang Island); Java Sea (Jakarta Bay); 2-10 m.

Elsewhere: tropical Indian Ocean; Loyalty Islands (Lifu); Queensland; Papua New Guinea (Bismarck, Solomon,

and Coral Seas); 0-73 m.

Genus RHIZOPSAMMIA Verrill, 1870

Rhizopsammia verrilli van der Horst, 1922

Figs 28 f-g

Rhizopsammia verrilli van der Horst, 1922: 64-65, pi. 8, figs 1-2. — ?Wells, 1983: 241-242. — Cairns, 1991: 25,

pi. 11, figs C-E (synonymy).

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

189

Dendrophyllia gracilis - Cairns, 1991: 23 (in part: USNM 78535). [Not Dendrophyllia gracilis H. Milne Edwards &

Haime, 1848].

MATERIAL EXAMINED. — Philippines: Cocos Island, 18 km east of Zamboanga, 6 m, 4 colonies (USNM 78534)

Indonesia. Snellius 1: unnumbered station, Binongko, Tukangbesi Islands, 6-10 m, 7 March 1930, 1 colony (NNM

22749).

~ West Pacific: Ngell Channel, Palau, 9 m, 3 colonies (USNM 78636).

Type Locality. — "Siboga" stns 220 and 282: Indonesia, 54-278 m.

DIAGNOSIS. — Colony reptoid, most corallites well separated from one another but interconnected by costate

stolons semi-circular in cross section, up to 6 mm in width, 1-3 stolons issuing from base of each attached

corallite. Additional corallites also bud from theca of erect corallites. Corallites up to 45 mm in height and 9 mm

in GCD. Costae equal and well defined; intercostal spaces highly porous. Septa hexamerally arranged in 5 cycles,

the 5th cycle always incomplete. Si -2 independent; septa arranged in a well-developed Pourtales plan. Inner edges

of highest cycle septa laciniate. Fossa deep; columella discrete and spongy.

Remarks. — The distinctive basal stolons distinguish this genus from other dendrophylliid genera, and the

large size of the corallites distinguish R. verrilli from the other Indonesian species, R. minuta van der Horst, 1922

and R. nuda van der Horst, 1926. It differs from the central Pacific R. chamissoi Wells, 1954, in having larger

corallites with more septa. Previously reported specimens of R. verrilli had not been known to have corallites

budding from the theca of other corallites, which led CAIRNS (1991) to identify at least one colony as

Dendrophyllia gracilis , the latter species having budding from both corallite edges as well as a common basal

coenosteum. _ . , . , ... . ,

As VAN DER Horst (1922) cautioned, a corallite of R. verrilli broken from its base could easily be mistaken

for a species of Balanophyllia or a Cladopsammia. Thus, complete coralla are required for accurate identification.

A more complete description of this species is given by Cairns (1991).

DISTRIBUTION. — Philippines : Sulu Sea (Zamboanga); 6 m. Indonesia: Banda Sea (Timor and Tukangbesi

Islands); 5-278 m. Elsewhere: Gulf of Thailand; Pelau; GaMpagos Islands; Cocos Island (eastern Pacific); 6-20 m.

Rhizopsammia nuda van der Horst, 1926

Rhizopsammia nuda van der Horst, 1926: 50-51, pi. 2, figs 10-12. „ 10791

Rhizopsammia (?) minuta- GARDINER & Waugh, 1939: 241. [Not Rhizopsammia minuta van der Horst, 1922],

Material EXAMINED. — Philippines. Marigondon Cave, Mactan Island, Cebu, 25-30 m, 1 colony (NNM

Indonesia. Dek.: stn 74, 1 colony (NNM 17503, 22779). - Stn 104, many corallites (NNM 17504, 22780).

Snellius 2: stn 4.100, 2 corallites (NNM 17511). - Stn 4.105, 1 colony (NNM 22783).

Type Locality. — Singapore, South China Sea (depth not given).

Diagnosis. — Colony reptoid, corallites united basally by thin (2.0-2.5 mm wide), flat stolons. Corallites

elongate-conical, up to 10 mm in height and 5 mm in GCD. Calice elliptical. Theca porous; no epithcca.

Septa hexamerally arranged in 4 cycles in a Pourtales plan. Inner edges of Si -2 dentate. No paliform lobes;

columella spongy.

Remarks. — Rhizopsammia nuda is similar to R. minuta van der Horst, 1922 (Fig. 28 h), the only

apparent difference being that the corallites of R. nuda are about twice as tall as those of R. minuta. Althoug

VAN DER HORST described both species, he did not compare them. A syntype of R. nuda is deposited at the ZM

(Coel. 5525) and 3 more colonies labelled as syntypes are deposited at the BMNH (1939.7.20.852, 853, and 8. i )•

This accounts for 4 of the 5 colonies mentioned in the original description.

Source :

190

S. D. CAIRNS & H. ZIBROWIUS

DISTRIBUTION. — Philippines'- Mactan Island, Cebu; 25-30 m. Indonesia : Lindah Strait (between Flores and

Sumbawa); Java Sea (Sunda Strait); 30-105 m. Elsewhere: Singapore (type locality, 9-22 m); Tanzania; 113-

220 m (Gardiner & Waugh, 1939).

Genus EGUCHIPSAMMIA Cairns, 1994

Eguchipsammia gaditana (Duncan, 1873)

Balanophyllia gaditana Duncan, 1873: 333.

Balanophyllia fistula - van DER HORST. 1922: 59 (in part, "Siboga": stn 310). [Not Balanophyllia fistula Alcock, 1902].

Dendrophyllia gaditana - Cairns, 1979: 181-182, pi. 36, figs 5-10 (synonymy). — ZIBROWIUS, 1980: 176-178, pi. 89,

figs A-N (synonymy). — Cairns & Keller, 1993: 279-280.

Eguchipsammia gaditana - Cairns, 1994: 85-86, pi. 37, figs d-f, h; 1995: 122-123, pi. 42, figs a-c.

MATERIAL EXAMINED. — Philippines. "Albatross": stn 5249, 4 branches (USNM 97622).

Indonesia. "Siboga": stn 310, 1 (ZMA).

Deki: stn 73, 4 (NNM 22750).

Karubar: stn 18, 10 branches (MNHN).

Type Locality. — "Porcupine" stn 29: 36°20’20"N, 6°47'W (Ibero-Moroccan Gulf), 417 m.

DIAGNOSIS. — Corallum consists of an elongate, cylindrical axial corallite from which secondary corallites bud

at right angle. Axial corallite unattached to substratum and often irregularly bent. Although some coralla achieve a

length of 53 mm (CAIRNS, 1994) and a GCD of 5.5 mm (ZIBROWIUS, 1980), the specimens reported above are

smaller, the longest only 15 mm and the calicular diameter ranging from 2-3 mm. Theca usually covered with a

thin epitheca giving lower corallum a porcellanous texture. Septa arranged in 3-4 cycles (depending on calicular

diameter) in a Pourtales plan. Si independent; each pair of S3 fuse before its common S2 high in the fossa.

Columella a small, nondiscrete, concave, spongy mass.

Remarks. — Eguchipsammia gaditana is described and figured in greater detail by ZIBROWIUS (1980) and

Cairns (1979, 1994, 1995). It is compared to E. wellsi in the account of that species.

DISTRIBUTION. — Philippines: Davao Gulf; 42 m. Indonesia: Banda Sea (Kai Islands); Flores Sea (Sumbawa);

Java Sea (Sunda Strait); 30-212 m. Elsewhere: widespread in tropical and temperate regions of world oceans, except

for eastern Pacific, but including north of New Zealand and Japan; 57-988 m.

Eguchipsammia wellsi (Eguchi, 1968)

Dendrophyllia (Alcockia) wellsi Eguchi, 1968: C63-64.

Eguchipsammia wellsi - Cairns, 1994: 86-87, pi. 37, figs a-c, g (synonymy).

Material EXAMINED. — Philippines. "Albatross": stn 5248, 5 (USNM 97624). — Sin 5249, 2 (USNM 97625)

— Stn 5357, 2 (USNM 97626).

Musorstom 3: stn 137, 3 (MNHN). — Stn 142, 1 (MNHN).

Type Locality. — "Soyo Maru" stn 210: 33°29'N, 135°28'E (Kii Peninsula, Honshu, Japan), 165 m.

Diagnosis. Corallum similar in shape to that of E. gaditana. Largest specimen reported herein ("Albatross"

stn 5248) 44 mm in length and 4.3 mm in GCD, which is typical for the species. Theca costate; no epitheca.

Septa hexamerally arranged in 3-4 cycles (up to 36 septa) in a Pourtales plan. Inner edges of S3-4 smooth.

Columella a discrete, convex structure composed of numerous small lamellae.

Source : MNHN, Paris

AZOOX ANTHELL ATE SCLERACTINIA

191

Remarks. — Although similar to E. gaditana (Duncan, 1873) in corallum shape, E. wellsi differs in having a

discrete, convex columella; having smooth inner septal edges that do fuse among themselves at a lower level in the

fossa; and in lacking epitheca. The species is more fully described and illustrated by Cairns (1994).

DISTRIBUTION. — Philippines: Sibuyan Sea; Visayan Sea; Davao Gulf; Sulu Sea (Balabac Island); 32-124 m.

Elsewhere : Japan (Honshu, Kyushu, and northern Ryukyu Islands); 1 10-196 m.

Genus CLADOPSAMMIA Lacaze-Duthiers, 1897

Cladopsammia echinata Cairns, 1984

Fig. 29 d

Cladopsammia echinata Cairns, 1984: 26-27, pi. 5, figs F-G.

MATERIAL EXAMINED. — Indonesia. Karubar: stn 86, 2 colonies: 1 (MNHN), 1 (USNM 97628).

TYPE Locality. — Sango 2 stn 4: 21°48'N, 160°09.1'W (Hawaiian Islands), 298-408 m.

Diagnosis. — Corallum irregularly and densely branched, resulting in a bushy clump of corallites. The

attachments of the colonies reported above are missing, all corallites budding from the theca of parent corallites and

occasionally merging with one another. Larger Karubar colony reported above 8 cm in height, 9 cm across, and

5 cm deep, consisting of about 120 corallites. Corallites elongate-conical to subcylindrical, and elliptical in cross

section, ranging from 3.2 to 8.4 mm in GCD. Ci-2 ridged near calice; otherwise theca uniformly covered with

small (0.15 mm height), slender, pointed spines, which are particularly well developed near pedicel and bases of

corallites. Septa hexamerally arranged in 4 complete cycles in a Pourtales plan, the Si -2 independent. Paliform

lobes absent. Columella spongy and relatively small.

Remarks. — Species previously known only from the Hawaiian Islands; original description more detailed.

DISTRIBUTION. — Indonesia: Arafura Sea (south of Tanimbar Islands); 222-226 m. Elsewhere: Hawaiian

Islands (Kauai, Nihoa, and Brooks Banks); 295-470 m.

Genus DENDROPHYLLIA Blainville, 1830

Dendrophyllia sp. cf. D. ijimai Yabe & Eguchi, 1934b

Fig. 29 e

Dendrophyllia minuscula - van DER HORST, 1922: 51-52, pi. 8, fig. 30. [Not Dendrophyllia minuscula Bourne. 1905J.

Dendrophyllia sp. - ZlBROWlUS & GRYGIER, 1985: 123, 126, figs 22-23.

Material EXAMINED. — Philippines. Musorstom 2: stn 33, 1 branch (MNHN).

Musorstom 3: stn 1 17, 2 branches (MNHN).

Indonesia. "Siboga": stn 49a, 1 (ZMA Coel. 5407, D. minuscula of van DER Horst, 1922).

Karubar: stn 30, 6 colonies: 3 (MNHN), 3 (USNM 97629).

Tasman Sea. "Tangaroa": stn Q47, 3 colonies (USNM 94236).

DESCRIPTION. — Coralla relatively small (up to 90 mm in height), consisting of a continuous, vertical,

slender (pedicel diameter 8 mm), founder axial corallite from which a variable number of secondary corallites bud at

right angle around circumference. Short tertiary buds occasionally form from the secondaries. Largest Indonesian

Source :

192

S. D. CAIRNS & H. ZIBROWIUS

colony (Karubar stn 30) 8 cm in height, bearing 16 corallites, the axial corallite tapering from 6.1 mm in basal

diameter to 3.4 mm distally. Corallites slightly elliptical in cross section and 3. 2-5. 8 mm in GCD, the axial

calices usually being the largest. Costae well developed, about 0.35 mm wide, and slightly convex; theca porous

near calicular edge.

Septa hexamerally arranged in 4 cycles, the 4th cycle complete usually only in axial corallites; other corallites

have 36-42 septa, pairs of S4 often missing from various half-systems. Si exsert (up to 0.7 mm), having smooth

lower, inner edges that reach the columella. S2 not exsert, 3/4 width of an Si, also having smooth inner edges.

S3 small, each flanked by a pair of S4 that meet, fusing before inner edge of their common S3 and extend as

1 septum to the columella. Inner edges of all septa entire, not laciniate or dentate. Fossa of moderate depth,

containing a large discrete columella composed of tightly fused, swirled lamellae.

Remarks. — This species belongs to a group (see Cairns, 1994) of about 5 species within the genus

Dendrophyllia characterised by having arborescent colonies with most budding occurring perpendicularly from

a central axial corallite: D. ramea (Linnaeus, 1758); D. minuscula Bourne, 1905; D. indica Pillai, 1967; D. velata

Crossland, 1952; and D. ijimai Yabe & Eguchi, 1934. Its growth form, calicular diameter, and septal number is

similar to D. ijimai , but because its corallites are uniformly 1-2 mm smaller in diameter and because the type of

D. ijimai is not available for comparison, only a tentative identification is suggested.

Distribution. — Philippines : Verde Island Passage; Mindoro Strait; 97-130 m. Indonesia: Banda Sea (Kai

Islands); Flores Sea; 69-111 m. Elsewhere: Taupo Seamount, Tasman Sea; 135 m. D. ijimai is known from Japan

and the western Indian Ocean; 10-366 m (Cairns & Keller, 1993).

Dendrophyllia arbuscula van der Horst, 1922

Figs 29 a-c

Dendrophyllia micranthus - van der Horst. 1922: 50 (in part: "Siboga" stn 277). [Not Oculina micranthus Ehrenberg,

1834],

Dendrophyllia arbuscula van der Horst, 1922: 53 (in part: "Siboga" stn 277; pi. 8, fig. 6). — EGUCHI, 1968: C55-56,

pi. C21 , figs 5, 13. — Pillai & Scheer, 1974: 462, fig. 7a. — Betterton, 1981: 242, figs 197-198. — Cairns,

1994: 90-91, pi. 38, figs i-1 (synonymy); 1995: 125-126, pi. 43, figs e-f.

Dendrophyllia subcornigera Eguchi, 1968: C64, pi. C32, figs 3-4.

Dendrophyllia subcornigera cylindrica Eguchi, 1968: C64-65, pi. C32, figs 1-2.

Dendrophyllia horsti Gardiner & Waugh, 1939: 237-238, pi. 2, figs 5-6.

? Dendrophyllia erecta Nemenzo, 1960: 19, pi. 10, fig. 1.

Dendrophyllia sp. cf. D. horsti - Cairns & KELLER, 1993: 278, pi. 13, figs F, I.

Not Dendrophyllia arbuscula - SCHEER & PILLAI, 1974: 64.

Material EXAMINED. — Philippines. "Albatross": sin 5279, 6 colonies (USNM 97630). — Stn 5280

2 colonies (USNM 97631).

Musorstom 3: stn 131, 7 (MNHN).

Indonesia. "Siboga": stn 277, 1 (ZMA).

Deki: stn 3, many colonies: (ZMA Coel. 7344), 2 (USNM 97633). — Stn 7, 2 colonies (NNM 22677). — Stn 24

1 (NNM 22678).

Karubar: stn 86. 6 colonies: 2 (MNHN), 4 (USNM 97632).

Type Locality. — "Siboga" stns 260 and 277: Banda Sea, 45-90 m.

Diagnosis. Corallum dendroid, attached by a thick base, which firmly anchors the primary axial corallite.

A variable number of secondary corallites bud at right angle to the axial, and tertiary corallites may also be

present. The early colony stage, represented by the axial and several secondary corallites, was described as D. horsti

by Gardiner & Waugh (1939), but the more fully developed colony containing secondary and tertiary branches

was first illustrated by van der Horst (1922: pi. 8, fig. 6) as the typical form. When secondary and tertiary

corallites are elongate (up to 5 cm) a more open corallum results, described as D. subcornigera cylindrica

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

193

by EGUCHI (1968). The largest Philippine specimen (" Albatross " stn 5279) is a highly branched corallum

(as illustrated by van der HORST, 1922) 7 cm wide, 5.5 cm in height, and 13.4 mm in pedicel diameter,

consisting of 20 corallites. Corallites elliptical in cross section: up to 12.2 mm in GCD. Costae broad

(0. 6-0.7 mm), flat, and quite porous, separated by narrow (0. 1 mm), shallow intercostal striae. Edge zone extends

only about 1 cm below calicular edge, below which the theca is usually encrusted. Septa hexamerally arranged in

4 cycles in a Pourtales plan, the septa of the first 2 cycles independent and exsert. Fossa shallow, containing

a well-developed, well-delimited, compact columella consisting of many small lamellae that are tightly fused

together in a clockwise swirl. Columella massive, up to 3 mm in width, and often constricted into a central and

2 narrower lateral parts by the 4 lateral S i .

REMARKS. — Van DER HORST's (1922, pi. 8, fig. 6) illustrated specimen from "Siboga" stn 277 (ZMA Coel.

5477) is a colony 80 mm in height bearing corallites 5-6 mm in GCD. One of the other 2 syntypes from "Siboga"

stn 260 (ZMA Coel. 1254) is Eguchipsammia fistula. We therefore designate the specimen from "Siboga" stn 277

as the lectotype, the 2 from "Siboga" stn 260 as paralectotypes.

In their description of D. horsti, Gardiner & Waugh (1939) noted its resemblance to D. arbuscula , assuming

that their colonies were "genetic dwarfs" in comparison. The growth series represented in "Albatross" stn 5279

suggests that it is more likely that D. horsti simply represents the early growth stage of a larger colony.

Dendrophyllia arbuscula belongs to Cairns' (1994) "second group" of Dendrophyllia species, i.e., species

having relatively small, bushy colonies with irregular branching from an axial corallite. The horsti-slage of

this species is more fully described by CAIRNS (1994).

DISTRIBUTION. — Philippines: Lubang Island; Sulu Sea (west of Panay; ? Luminusa Island); 122-353 m.

Indonesia : Banda Sea (Kai, Damar, and Barat Daya Islands); 45-245 m. Elsewhere: southwestern Indian Ocean to

Strait of Malacca; northern New Zealand region (Norfolk and Kermadec Islands); Japan (Honshu, East China Sea);

40-259 m.

Dendrophyllia alcocki (Wells, 1954)

Sclerhelia alcocki Wells, 1954: 465-466, pi. 177, figs 1-2.

Dendrophyllia palita Squires & Keyes, 1967: 28-29, pi. 6, figs 9-10.

Dendrophyllia alcocki- Zibrowius, 1974a: 570-573, figs 10-14. — Cairns, 1995: 126-127, pi. 43, figs g-i, pi. 44,

figs a-b (synonymy).

Material EXAMINED. — Indonesia. "Albatross": stn 5586, 1 fragment (USNM 97635).

Karubar: stn 18, 2 fragments (MNHN).

South China Sea. "Albatross": stn 5311, 1 fragment (USNM 97634).

"Hakuho Maru": stn KH73-2-44-2, 1 branch (USNM 97636).

Type Locality. — Bikini Atoll, Marshall Islands, 177-243 m.

Diagnosis. — Uniplanar or arborescent colonies formed by regular, extratentacular, sympodial branching.

Potential growth to at least a height of 11 cm (Cairns, 1995). Specimens reported above only small branch

fragments, each about 1 cm in length comprising only 4 or 5 corallites, with calices 4-5 mm in diameter.

Coenosteum dense and solid, slightly porous only near calicular edge. Theca covered with blunt spines that are

usually aligned on branch axis. Septa hexamerally arranged in 3 complete cycles: Si>S2^S3- Each pair of S3

meets before its common S2 to form a large palus (P2). Columella spongy.

Remarks. — According to the partial revision of the genus (Cairns, 1994), D. alcocki is one of at least

8 species in the "third group", characterised by having large, sympodially branched coralla. D. alcocki is more

fully described and figured by Cairns (1995).

Distribution. — Indonesia: Celebes Sea (Darvel Bay); Banda Sea (Kai Islands); 205-616 m. Elsewhere:

Maldives', throughout New Zealand region; Tasman Sea; New Caledonia region; Solomon Islands; Marshall

Islands; South China Sea (north of Pratas Islands); 118-570 m.

Source :

194

S. D. CAIRNS & H. ZIBROWIUS

Genus EN ALLOPS AMMI A Michelotti, 1871

Enallopsammia pusilla (Alcock, 1902)

Fig. 29 f

Dendrophyllia < Coenopsammia ) pusilla Alcock, 1902a: 113; 1902c: 44, pi. 5, figs 38, 38a.

Dendrophyllia (Coenopsammia) profunda - ALCOCK, 1902c: 43. [Not Diplolielia profunda Pourtales, 1867].

Coenopsammia profunda - Marenzeller, 1904a: 313-314, pi. 18, fig. 24. [Not Diplolielia profunda Pourtales, 1867].

Enallopsammia marenzelleri Zibrowius, 1973: 49-51 (in part: pi. 1, figs 1-7, only Indo-Pacific specimens, including

holotype); 1980: 204-205 (in pan: only Indo-Pacific specimens) [new synonym],

Enallopsammia sp. cf. E. marenzelleri - Cairns, 1982: 57-58, pi. 18, figs 5-6; 1995: 128-129, pi. 44, figs g-h.

MATERIAL EXAMINED. — Philippines. "Siboga": stn 95, holotype and paratype of D. pusilla (ZMA Coel. 1196,

589, respectively).

"Hakuho Maru": stn KH72-1-20, 4 branches (USNM 97638).

Indonesia. "Siboga": stn 266, holotype and paratype of E. marenzelleri (ZMA Coel. 6902, 588, respectively).

Deki: stn 1,4 branches (ZMUC), 6 branches (NNM 22735). — Stn 12, 1 (NNM 22739). — Stn 59, 2 (BMNH

1939.7.20.316), 10 (NNM 22736, 22740). — Stn 61, 1 (NNM 22737).

Snellius 2: Stn 4.144, 1 (NNM).

Karubar: stn 25, 7 colonies: 5 (MNHN), 1 (POLIP1), 1 (USNM 97640).

South China Sea: "Albatross": stn 5317, 3 branches (USNM 97637).

"Hakuho Maru": stn KH73-2-44-2, 8 branches: 6 (USNM 97639), 2 (ORI).

Type Locality. — "Siboga" stn 95 (5°43.5'N, 1 19°40'E (Sulu Archipelago, Philippines), 522 m.

Description. — Extratentacular budding (and branching) occurs in an irregular manner leading to a massive,

irregularly shaped colony with occasional branch anastomosis. Budding often sympodial on distal, small-diameter

branches, but buds may occur on all branch faces, producing a 3-dimensional corallum. Largest known colony

(Karubar stn 25) 160 mm in height and 290 mm in width, consisting of 7 main vertical branches originating

from a common base 27 mm in diameter. Corallites slightly elliptical (GCD:LCD = 1.02-1.24), up to 4.7 mm in

GCD, standing up to 3 mm above branch coenosteum. Costae convex and well developed over entire branch

coenosteum. On small-diameter branches, costae about 0.3 mm wide, bearing only 1 costal spine across their

width, and are flanked by deep, highly porous intercostal striae about 0.2 mm wide. On large-diameter branches,

costae are broader (up to 0.6 mm), bear 2 or 3 small spines across their width, and are flanked by narrower (about

0.1 mm), less porous intercostal striae. Intercostal pores of large-diameter branches gradually filled in with

stereome, resulting in a very dense corallum.

Septa hexamerally arranged in 3 cycles in an indistinct Pourtales plan. All septa nonexsert. Si independent and

quite narrow (about 0.5 mm), having smooth inner edges that reach the columella. S2 equal to Si in size and

shape. S3 narrower than S1-2, having dentate inner edges, each pair of S3 within a system bending toward its

common S2 and fusing with that septum near the columella. Fossa deep and spacious. Columella a circular,

concave, nondiscrete, spongy mass.

Remarks. — Enallopsammia pusilla was described on the basis of what now appear to be 2 relatively short

distal branches of a larger corallum - a total of 25 corallites (see ZIBROWIUS, 1973). Additional larger specimens

are now available from near to the type locality ("Hakuho Maru" KH72-1-20) as well as near the type locality of

E. marenzelleri (Karubar stn 25). These colonies strongly suggest that the holotype of E. marenzelleri is simply

a larger, more robust colony of E. pusilla.

Distribution. — Philippines: Sulu Sea (Sulu Archipelago); 514-522 m. Indonesia: Banda Sea (Kai and

Selayar Islands); 325-730 m. Elsewhere: Macquarie Ridge; Nicobar Islands, Bay of Bengal; South China Sea (north

of Pratas Island and southern Formosa Strait); 371-805 m.

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

195

Enallopsammia rostrata (Pourtal&s, 1878)

Amphihelia rostrata Pourtal&s, 1878: 204, pi. 1, figs 4-5.

Dendrophyllia (Coenopsammia) amphelioides Alcock, 1902a: 112-113; 1902c: 43-44, pi. 5, figs 37, 37a.

Anisopsamrnia rostrata - MARENZELLER, 1904a: 314-315, pi. 18, fig. 23.

Enallopsammia rostrata - Zibrowius, 1973: 44-45, pi. 2, figs 14-15. — Cairns, 1982: 57, pi. 18, figs 1-4 (synonymy);

1994: 92-93, pi. 39, figs d-f; 1995: 127-128, pi. 44, figs c-f. — Cairns & Parker, 1992: 52-53, pi. 18, figs e-i.

MATERIAL EXAMINED. — Philippines. " Albatross stn 5428, 3 branches (USNM 97642). — Stn 5499,

5 branches (USNM 97643).

Indonesia. Karubar: stn 13, 7 branches: 3 (MNHN), 1 (POLIPI), 3 (USNM 97644).

TYPE LOCALITY. — "Blake" stn 2: 23°14'N, 82°25'W (Straits of Florida), 1472 m.

REMARKS. — Enallopsammia rostrata differs from E. pusilla in having unifacial corallites (i.e., calices

confined to one face of the corallum, anterior by convention), arranged uniserially on small-diameter branches. The

corallites of most specimens of E. rostrata also bear a prominent septocostal rostrum; however, some specimens

lack this structure, these referred to as the "amphelioides" form by CAIRNS (1982). Of the 3 lots reported above,

specimens from "Albatross" stns 5428 and 5429 have septocostal rostra, whereas those from Karubar stn 13 do

not. The rostrate specimens have small corallites (3 mm GCD), consistent with the "delicate" specimens reported

by Cairns (1995) from New Zealand, whereas the nonrostrate specimens have massive coralla.

DISTRIBUTION. — Philippines: Sulu Sea (Palawan); Bohol Sea; 1013-2021 m. Indonesia: Halmahera Sea;

Ceram Sea; Banda Sea (Kai Islands); 417-1633 m. Elsewhere: cosmopolitan, except for eastern Pacific and

continental Antarctica; 1 10-2165 m.

Genus TUBASTRAEA Lesson, 1829

Tubastraea micranthus (Ehrenberg, 1834)

Oculina micranthus Ehrenberg, 1834: 304.

Dendrophyllia nigrescens Dana, 1846: 387. — Vaughan, 1918: 143-144, pi. 60, figs 1, la. — Searles, 1956: 24,

pi. 39A.

Coenopsammia viridis H. Milne Edwards & Haime, 1848b: 110.

Coenopsammia aequiserialis H. Milne Edwards & Haime, 1848b: 1 10-1 11. — Semper, 1872: 267.

Dendrophyllia micranthus - van DER Horst, 1922: 49-51 (in part: not "Siboga" sin 277, synonymy); 1926: 43-44,

pi. 2, figs 6-7. — Faustino, 1927: 218-220, pi. 72, figs 1-2. — Nemenzo, 1960: 16-17, pi. 8, fig. 2. — Scheer &

Pillai, 1974: 63, pi. 29, fig. 3. — BETTERTON, 1981: 242, figs 199-200.

Dendrophyllia micranthus var. grandis Crossland, 1952: 173, pi. 55, fig. 1, pi. 56, fig. 1.

Tubastraea micranthus - Zibrowius & Grygier, 1985: 130. — SCHUHMACHER, 1984: 94, figs la-b, 4.

Tubastrea micranthus - Latypov, 1990: 68, pi. 26, figs 1-2.

Tubastraea micrantha - CAIRNS & KELLER, 1993: 282.

Not Dendrophyllia micranthus - EGUCHI, 1968: C66 [= Dendrophyllia ijimai Yabe & Eguchi, 1934],

Material EXAMINED. — Philippines. "Albatross": stn 5554, 1 (USNM 97648).

Siphilexp: stn 78-CAC 189, 1 (USNM 97646).

Cocos Island, east of Zamboanga, 9 m, 1 colony (USNM 83685).

"Southern Philippines", depth unknown, 200+ branches (USNM 91088).

Indonesia. "Siboga": stn 240, 1 (ZMA Coel. 235).

Deki: stn 4, 1 (ZMA Coel. 5450). — Stn 10, several colonies (NNM).

"Alpha Helix": stn 1769, 1 (USNM 78551). — Stn 79-M122, 3 (USNM 97647).

Type Locality. — Unknown.

Source :

196

S. D. CAIRNS & H. ZIBROWIUS

Description. — Corallum dendroid, but more or less uniplanar, achieved by profuse extratentacular budding

from a relatively small number (2-8) of massive axial corallites. Coralla may attain a large size: e.g., 1 m in

height and 5 cm in basal diameter. Corallites of small-diameter distal branches generally occur only on branch

edges, but on larger-diameter branches corallites more uniformly distributed on all branch faces. Corallites usually

project upward at a 45° angle from axial branch and stand 5-7 mm above branch coenosteum. Corallites usually

6- 8 mm in GCD, but Nemenzo (1960) reported giant calices 10-12 mm in GCD. Costae well defined,

0. 4-0.5 mm in width, convex to ridged, bearing 1-3 low granules across their width at any point. Intercostal

furrows long and continuous, 0.15-0.20 mm wide, and occasionally punctuated with circular pores about 0.3 mm

in diameter that penetrate deeper into branch core. Branch porosity greatest in distal parts, the pores becoming

infilled and thus more dense with age (SCHUHMACHER, 1984). Corallum white; live tissues a striking dark green

or brown-black.

Septa hexamerally arranged in 3 cycles: Si>S2»S3. Si nonexsert, having straight inner edges that attain the

columella. S2 3/4 width of an Si, also having straight inner edges that attain the columella. S3 usually

rudimentary, represented by a very narrow dentate to laciniate lamella. Fossa deep, especially in axial corallites.

Columella rudimentary, composed of a solid, elongate fusion of lower, inner edges of the S1-2.

Remarks. — Tubastraea micranthus is the only species in the genus to have a tree-like, dendroid growth form.

Being a common, shallow-water species, it has received various names, including several alluding to its tissue

colour.

As defined by SCHUHMACHER & ZIBROWIUS (1985), T. micranthus belongs to a unique ecological category

among the Scleractima, i.e., azooxanthellate, yet constructional and hermatypic. In other words, although it lacks

zooxanthellae, it produces large colonies that contribute to a reef structure. According to SCHUHMACHER (1984),

T. micranthus does not grow as fast as other branching zooxanthellate corals, but because it reinforces its branch

strength through secondary calcification, it remains competitive with other reef corals.

Distribution. — Philippines : Negros and Bohol; Sulu Sea (Zamboanga Peninsula and Sulu Archipelago);

7- 46 m. Indonesia: Molucca Sea (Halmahera); Banda Sea (Ambon, Banda, Kai, and Damar Islands); 0.5-60 m.

Elsewhere: widespread in tropical Indo-West Pacific from southwestern Indian Ocean to Fiji (most corals referred to

this species from Japan are Dendrophyllia)\ 0-50 m.

Tubastraea diaphana (Dana, 1846)

Dendrophyllia diaphana Dana, 1846: 389, pi. 27, fig. 3. — Vaughan, 1918: 144-145, pi 60 figs 2-3

Dendrophyllia aequiserialis - Quelch, 1886: 147. [Not Coenopsammia aequiserialis H. Milne Edwards* Haime 18481

Dendrophyllia micranthus var. fruticosa Nemenzo, 1960: 17-18 pi 9 fig 1

Tubarigaia diaphana ’ SCHEER & PlLLAI' 1983: 174' pl‘ 41 ’ f,gs'M <synonymy)- - Cairns & Keller, 1993: 284, pi. 13,

Dendrophyllia sibogae van der Horst, 1922: 56-57, pi. 8, figs 18-19.

Tubastraea sp. - Guella et al. , 1988: 780.

Material examined. — Philippines. Santa Cruz, Zamboanga, 1 (USNM 78522)

GuE^rran/.Pia98a88e’ Palawan’ 3 m- 6 C0l0nieS (USNM 80822)' 4 C0l0nies (BMNH 1987.12.23.1-4) mentioned by

Siphilexp: stn 78SP-1-1, 5 (USNM 77162)

Type Locality. — Singapore, South China Sea (depth not given).

Diagnosis. - Colonies phaceloid, forming small, bushy clusters of corallites rarely more than 5 cm in height

or width. Colony results from closely spaced budding from a broad base. Corallites 6-1 1 mm in GCD and up to

22 mm m length. Costae well defined, as in T. micranthus. Theca thin and quite porous, especially near calicular

complete "T hexameral|y arra"ged in “P to 4 cycles, the 4th cycle rarely

• J1>S2»S3>S4. Si nonexsert, rather narrow near calicular edge, having smooth inner edges. S2 about

3/4 width of an S,, also having smooth inner edges. S3 usually present but represented only as low dentate ridges

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

197

In large coralla, traces of S4 occur in some half-systems near calicular margin. Fossa deep and spacious; columella

rudimentary.

Remarks. — This species is more fully described and illustrated by Vaughan (1918) and NEMENZO (1960),

by the latter as T. micranthus wax. fruticosa.

DISTRIBUTION. — Philippines : west coast of Mindoro; Sulu Sea (Dumaran Passage, northeast Palawan;

Zamboanga Peninsula); Negros; 1-5 m. Indonesia : Savu Sea; eastern Timor (VAN DER HORST, 1922); 27-54 m.

Elsewhere: widespread throughout tropical Indo-West Pacific from southwestern Indian Ocean to Fiji and Samoa;

1-15 m.

Tubastraea coccinea Lesson, 1829

Tubastraea coccinea Lesson, 1829: 93. — Wells, 1983: 243-244, pi. 18, figs 1-2 (synonymy). — Cairns, 1991:

26-27, pi. 12, figs c-e (synonymy); 1994: 93-94, pi. 39, figs g-i (synonymy). — Cairns & Keller, 1993: 282-284.

Lobophyllia aurea Quoy & Gaimard, 1833: 195, pi. 15, figs 7-11.

Coenopsammia willeyi Gardiner, 1899: 359, pi. 34.

Dendrophyllia willeyi - Vaughan, 1918: 143-144, pi. 60, figs 4, 4a.

Dendrophyllia aurea - VAN DER HORST, 1926: 46-48 (in part: pi. 2, figs 2-4, 8-9).

Tubastraea aurea - BOSCHMA, 1953: 111-118 (in part: pi. 10, figs 2, 6, pi. 11, figs 2, 4-6, pi. 12, figs 1-6). — Searles,

1956: 24, pi. 38B. — BETTERTON. 1981: 242-243, fig. 201. — SCHEER & Pillai, 1983: 173-174, pi. 40, fig. 8. —

SCHUHMACHER, 1984: 94-95. — LATYPOV, 1990: 65-66, pi. 27, fig. 4, pi. 32, fig. 5.

? Dendrophyllia turbinata Nemenzo, 1960: 18-19, pi. 9, fig. 2.

Tubastrea coccinea - Latypov, 1990: 66-67, pi. 27, fig. 1, pi. 32, fig. 3.

MATERIAL EXAMINED. — Philippines. Santa Cruz, Zamboanga Peninsula, 4 colonies (USNM 83645, 83652,

83667).

Siphilexp: stn 78-CAC194, 3 colonies (USNM 97645).

Indonesia. "Siboga": stn 231, 1 (ZMA Coel. 586).

Type Locality. — Bora Bora, Society Islands (depth not given).

Diagnosis. — Corallum plocoid, forming spherical to mound-shaped colonies up to 10 cm in diameter. Most

corallites originate from a common basal coenosteum, only rarely budding from the wall of another corallite.

Corallites short and squat: 10-13 mm in GCD and rarely over 10 mm in height. Costae similar to those of

T. diaphana. Tissue usually bright orange, but may also be yellow, pink, or purple. Septa hexamerally arranged in

4 cycles, the 4th usually incomplete: Si>S2»S3>S4- Septal arrangement also similar to that of T. diaphana.

Fossa of moderate depth, containing a rudimentary crispate columella. Endothecal dissepiments occasionally

present.

Remarks. — WELLS (1983) listed 18-19 junior synonymys of T. coccinea and recognised a total of 6 valid

species in the genus. The synonymy given above is therefore incomplete but gives most of the Indonesian records.

A more complete description of this species can be found in Cairns (1994).

Four species of Tubastraea occur in the Indonesian region, 3 of which have similar corallite characteristics but

are distinguished by their colony size and shape: T. micranthus, large and arborescent; T. diaphana, small and

phaceloid; and T. coccinea, medium-sized and plocoid. The 4th species, T. faulkneri Wells, 1982, is similar to

T. coccinea in growth form but has very widely-spaced corallites that are sunken in thick coenosteum, and S4 that

pair before their common S3. Wells (1982) reported T. faulkneri from Pelau, the Banda Sea (Banda and Ambon

Islands), Mindoro, and the Galapagos Islands at 3-8 m. According to Wells (1983), only 2 other species of

Tubastraea are valid: T. tagusensis Wells, 1982, and T.floreana Wells, 1982, both from the Galapagos Islands and

distinguished by their septal characteristics (Cairns, 1991 : table 4).

Distribution. — Philippines: Mindoro and Zamboanga Peninsula, Mindanao; 3-6 m. Indonesia: Banda Sea

(Kai and Ambon Islands); 40 m. Elsewhere: cosmopolitan in tropical shallow water, including warm temperate

region of Japan; 1 .5- 1 10 m.

Source :

198

S. D. CAIRNS & H. ZIBROWIUS

IN CERT AE SEDIS

Figs 29 g-i

Material EXAMINED. — Philippines. " Albatross stn 5179, 5 colony fragments, including SEM stub 813

(USNM 97651).

Indonesia. Deki: unnumbered station near Banda, 3 June 1922, 30 m, 1 colony (ZMUC).

DESCRIPTION. — Small colonies formed by sparse intratentacular budding. Corallites sometimes budded in

series, resulting in a unifacial arrangement or budded sympodially, resulting in a loose, bushy corallum with

anastomosing branches. Calice circular: 2. 1-3.3 mm in diameter. Corallum completely epithecate, only 60 pm

thick at calicular edge. Epitheca composed of narrow (20-45 pm wide) horizontal bands that encircle the corallites.

Septal symmetry and size classes not clearly defined: 20-24 septa per calice. Septa highly porous, the larger septa

composed of 4 or 5 trabecular spines, having clavate, tuberculate tips about 0.12 mm in diameter. Most septa are

connected to their adjacent septa by narrow synapticulae. Columella composed of 3 or 4 granular rods, each up to

0.17 mm in diameter.

Remarks. We can find no scleractinian family in which to convincingly place the species described above

Its epitheca and size are similar to those of Culicia (Rhizangiidae), but it differs in budding mode and septal

construction. Its septal structure is similar to that of the Micrabaciidae, but it differs in having a solid epitheca and

being colonial. Until more specimens are examined, we reserve judgment on the family affinities of this unusual

coral.

Distribution. — Philippines: Sibuyan Sea; 68 m. Indonesia-. Banda Sea (off Banda Island); 30 m.

ACKNOWLEDGEMENTS

We would like to thank Alain Crosnier, Philippe Bouchet AND Bertrand Richer de FORGES, who provided

many of the specimens used in this study and who provided encouragement for the completion of the study.

We also thank the following people, who generously loaned specimens used in the study and/or facilitated

collection: R. M. W. van Soest and J. J. Vermeulen (ZMA); M. Borel-Best and L. Ofwegen

( ), P. F. S. Cornelius (BMNH); Y. Shirayama (ORI); and K. Petersen and T. Wolff (ZMUC).

;,ooominary ldentiflcations of Balanophyllia carinata and B. stimpsonii from the DEKI were made by M. BRUNE

(1988) a student working at the NNM. Coral symbionts/epibionts have been identified by the following fellow

naturalists: subentid sponges by K. Rutzler, gastropods by A. WarEn, lumbrinerid polychaetes by T. Miura

ascothoracids by J.M. Grygier, brachiopods by A. Logan.

The scanni“g electron photomicrographs were taken in the SEM Laboratory, NMNH. The Department of

Photography of the NNM provided the negatives of the holotype of Madrepora minutiseptum.

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Yabe, H. & Sugiyama, T., 1941. — Recent reef-building corals from Japan and the south Sea Islands under the Japanese

mandate. Pt. 2. Science Reports of the Tdhoku Imperial University, ser. 2, Geology, special volume 2: 67-91,

pis 59-104.

ZlBROWius, H., 1973. — Revision des espfeces actuelles du genre Enallopsammia Michelotti, 1871, et description de

E. marenzelleri, nouvelle espfece bathyale h large distribution Ocdan Indien et Atlantique Central (Madreporarta,

Dendrophylliidae). Beaufortia, 21 (276): 37-54.

Zibrowius, H., 1974a. — Redescription of Sclerhelia hirtella from Saint Helena, South Atlantic, and remarks on Indo-

Pacific species erroneously referred to the same genus (Scleractinia). Journal of Natural History, 8 (5): 563-575.

ZIBROWIUS, H„ 1974b. — Scldractiniaires des lies Saint-Paul et Amsterdam (Sud de l'Ocdan Indien). Tethys, 5 (4)

["1973"]’: 747-778.

ZIBROWIUS, H„ 1974c. — Revision du genre Javania et considerations gdndrales sur les Flabellidae (Scldractmiaires).

Bulletin de I'Institut Oceanographique , Monaco, 71 (1429): 1-48.

Zibrowius, H., 1980. — Les scldractiniaires de la Mdditerranee et de l'Atlantique nord-oriental. Memoires de I Institut

Oceanographique , Monaco, 11: 1-284, 107 pis.

Zibrowius, H., 1982. — Deep-water scleractinian corals from the south-western Indian Ocean with crypts excavated by

crabs, presumably Hapalocarcinidae. Crustaceana, 43 (2): 113-120.

Zibrowius, H„ 1985. — Asexual reproduction by bud-shedding in shallow-water Balanophyllia of the tropical Indo-

Pacific (Cnidaria Scleractinia Dendrophylliidae). Proceedings of the Fifth International Coral Reef Symposium,

5: 233-238.

ZIBROWIUS, H. & GiLI, J.M., 1990. — Deep-water Scleractinia (Cnidaria Anthozoa) from Namibia, South Africa, and

Walvis Ridge, southeastern Atlantic. Scientia Marina, 54 (1): 19-46.

Zibrowius, H. & Grygier, M.J., 1985. — Diversity and range of scleractinian coral hosts of Ascothoracida (Crustacea

Maxillopoda). Annales de I'Institut Oceanographique, Paris, 61 (2): 115-138.

Zibrowius, H„ Southward, E.C. & Day, J.H., 1975. — New observations on a little-known species of Lumbrineris

(Polychaeta) living on various Cnidarians, with notes on its recent and fossil scleractinian hosts. Journal of the

Marine Biological Association of the United Kingdom, 55 (1): 83-108.

Zou, Renlin, 1984. — Studies on the deep-water Scleractinia from South China Sea. 1. A nomen novum and a new species

of Caryophyllia. Tropical Oceanology, 3 (3): 51-54, pis 1-2. [in Chinese, with English summary]

Source :

208

S. D. CAIRNS & H. ZIBROWIUS

Zou, Renlin, 1988. — Studies on the deep-water Scleractinia from South China Sea. 2. Record and narration of species as

well as time-spatial distributional characteristics. Tropical Oceanology, 7 (1): 74-83, pis 1-5. [in Chinese, with

English summary]

Zou, Renlin, Meng, Zhimin & Guan, Xilian, 1988. — Ecological analyses of deep sea scleractinian on the continental

shelf of the northern South China sea. Selected Oceanic Works, 1: 193-199. South China Sea Institute of Oceanology.

Academia Sinica.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

209

Fig. 1 a-d. — Madracis asanoi Yabe & Sugiyama, 1936: a. "Albatross" stn 5381 (USNM 96672), distal branch, x 1.9. —

b-d, "Albatross" stn 5277, USNM 96671: b. branch tip. x 1 1.9: c. unequal-sized corallites. x 17.3: d, calice. x 40.

Fig. 1 e-f. — Madracis sp. A, Corindon 2 stn 248 (USNM 96684): e, branch, x 12.3; f. calice, x 36.

Fig. 1 g-h. — Madracis sp. cf. M. pharensis, MUSORSTOM 3 stn 117 (USNM 96676): g. stereo view of a small encrusting

colony, x 13.5; h, calice, x 35.4.

210

S. D. CAIRNS & H. ZIBROWIUS

Fig. 2 a-d. — Fungiacyathus fissidiscus sp. nov., paratypes, Karubar stn 7 (USNM 96725): a, fracture showing septal

spines, x 21.8; b, calice, x 15; c, base showing regeneration from parent fragment, x 15.8; d. fracture showing septal

face granulation, x 24.8.

Fig. 2 e-f. — Leptopenus sp. A, Karubar stn 7 (MNHN): e-f, two views of the same corallum fragment, x 25.5, x 21.8,

respectively.

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

21 1

Fig. 3 a-b. — Culicia stellata Dana, 1846, "Alpha Helix" stn 79-M140 (USNM 80029): a, stereo calieular view, x 17.1;

b. enlargement of thecal lip and septal dentition, x 41.

Fig. 3 c-g. — Madrepora arbuscula (Moseley, 1881): c-d, f-g. MUSORSTOM 3 stn 126 (USNM 96750): c-d, corallum and

corallites of a massive specimen, x 0.25, x 0.62, respectively; f, stereo view of a fractured corallite illustrating lower

septa, columella, and dissepiment, x 15; g, calice and smooth coenosteum, x 12.8. — e, holotype, "Challenger"

stn 194 (BMNH 1880.11.25.96), x 1.2.

212

S. D. CAIRNS & H. ZIBROWIUS

F|C. 4 a-d. Madrepora minutiseptum sp. nov.: a-b, d, holotype, SNELLIUS 2 stn 4.196 (NNM 22734): a-b. anterior and

?oTCn-^MM OASny’ ^ ^ d' e"largement lowing anastomosing branches, x 0.91. - c p a ype

oil Japan (LtSNM 96754, ex ZMA 137). colony, x 0.38. H ’

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

213

Fig. 5 a-b. — Madrepora minutiseptum sp. nov., paratype, "off Japan" (USNM 96754): a. stereo view of a fractured

corallite showing septal dentition, x 23.5; b, two septa, x 105.

Fig. 5 c-e, g-h. — Neohelia sp. cf. N. porcellana : c. Snellius 2 stn 4.104 (USNM 96765), colony, x 1.05. — d, "Siboga"

stn 305 (ZMA), branch, x 14.5. — e, "Siboga" stn 289 (USNM 96766). calice, x 37. — g-h. Deki (Ambon. 91 m)

(ZMUC): g, coenosteal papillae, x 175; h. calice, x 58.

Fig. 5 f. — Neolielia porcellana Moseley, 1881, syntype, "Challenger" stn 177 (BMNH 1880.11.25.89). x 1.5.

214

S. D. CAIRNS & H. Z1BR0WIUS

FlG 6 a-b. — Anthemiphyllia frustum Cairns, 1994, Karubar sin 15 (USNM 96776): a. lateral view of 2 anthocyathi

that remained attached, x 5.4; b, calice of same specimen, x 6. 1.

Fig. 6 c-e. - Caryophyllia secla sp. nov., holotype. "Albatross" stn 5265 (USNM 96787): c-e. side, stereo calicular

view, and basal scar, x 2.9, x 3.4, x 14.3, respectively.

Fig. 6 t-h. — Caryophyllia transversalis Moseley, 1881: f. h. Karubar stn 79 (POL1PI): f, side view, x 2.2; h. edge

view with attached subentid sponge, x 1.8, — g, syntype, "Challenger" stn 192 (BMNH 1880.1 1.25.23), side view.

X z. / .

Source : MNHN , Paris

Fig. 7 a-b. — Caryophyllia octonaria sp. nov., holotype, Musorstom 1 sin 64 (MNHN), side and calicular views, x 4.5,

x 6.1, respectively.

Fig. 7 c, f, i. — Caryophyllia grayi (H. Milne Edwards & Haime. 1848): c. syniype. Japan (BMNH 1840.9.29.42), side

view, x 2.1. — f, i, Musorstom 1 stn 56 (MNHN): f, calicular view, x 2.5; i. side view showing edge spines, x 2.4.

Fig. 7 d-e. — Caryophyllia comulum sp. nov., holotype, CORINDON 2 stn 220 (MNHN), side and calicular views, x 2.7.

Fig. 7 g-h. — Caryophyllia grandis Gardiner & Waugh. 1938, Karubar stn 59 (MNHN), edge and calicular views of

corallum with 24 pali, x 1.1, x 1.3, respectively.

AZOOXANTHELLATE SCLERACTINIA

Source :

216

S. D. CAIRNS & H. ZIBROWIUS

Gv? *;dv o denla,“ (Moseley, 1881): a. hololypc, "Challenger" stn 174 (BMNH 1880.11.25.42), side

le , x 2.9. b, d, Deki sin 3 (USNM 96858), side and calicular views of a decameral specimen, x 3.4, x 4 3

respectively^— c, Deki (Neira) (NNM 23076), side view of hexameral specimen, x 2.8

Drominem (SaVi“e *mt' 187I) "Alba,ross" sln 5371 (USNM 92690): e, corallum with

prominent Ci spines, x 2; f, calicular view, x 2.

F|c. 8 g-i. — Cary ophy Ilia spinicarens (Moseley, 1881): g-h, holotype, "Challenger" stn 210 (BMNH 1880 II ^5 43)

ddLa.e edgeCcUreasr,.sV'exW3S.6X ’ X 3'3' reSpeC,ively' ~ 1 "Alb«'ross" stn 5256 (USNM 96903), small corallum with

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

217

Fig. 9 a-c. — Caryophyllia karubarica sp. nov.. holotype. Karubar stn 58 (MNHN), side. edge, and calicular views,

x 2.0, x 2.15, x 2.15, respectively.

Fig. 9 d-e. — Caryophyllia unicristata sp. nov., holotype. Karubar stn 7 ( MNHN). side and calicular views, x 3.1,

x4. 1, respectively.

Fig. 9 f-j. — Premocyathus dentiformis (Alcock, 1902): f. h, "Albatross" stn 5217 (USNM 62708), side and calicular

views, x 6.4, x 7.8, respectively. — g. i, holotype, "Siboga" stn 59 (ZMA Coel. 1093), calicular and basal views, x

9.4, x 8.4, respectively. — j, Karubar stn 15 (MNHN), specimen showing regeneration from basal region, x 7.1.

Source :

218

S. D. CAIRNS & H. ZIBROWIUS

MUS0RST“ 1 “ 72 <MN™>. *• - views of

f!g !o * h Z^rbr,in,h0T,hUS Sp- A- KARUBAR s,n 36 <MNHN), side and calicular views, x 4.4, x 5 6 respectively

x 7.5 respecUvd^ SP' n°V" h°l0,ype' CoR,NDON 2 SIn 251 (MNHN). side and calicular views, x 6.4,

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

219

Fig. 1 1 a-d. — Trochocyathus apertus sp. nov.: a-c. holotype. "Albatross" stn 5164 (USNM 97087). side, open base, and

calicular views, x 5.2, x 13, x 7.2, respectively. — d, paratype, Mortensen'S Java Exp., stn 9 (ZMUC). calice, x 5.3.

Fig. lie. — Trochocyathus cooperi (Gardiner, 1905). Musorstom 2 stn 47 (MNHN), side view of an anthocyathus. x 4.

Fig. 11 f. — Trochocyathus semperi sp. nov., CORINDON 2 stn 251 (POLIPI), 2 small coralla with costal spines, x 5.6.

Fig. 11 g-h. — Trochocyathus discus sp. nov.: g, paratype. Karubar stn 3 (USNM 97105). stereo calicular view, x 7.5.

— h, holotype, Karubar stn 3 (MNHN), calice, x 5.

220

S. D. CAIRNS & H. ZIBROWIUS

Fig 12 a-c. — Trochocyathus discus sp. nov.: a, holotype, Karubar stn 3 (MNHN), side view, x 5.2. — b-c. paratyoe

Karubar sin 3 (USNM 97105): b. costal granulation at calicular edge, x 25; c, basal scar, x 14.

FlG' '2,d'f- — Trocl‘ocy“ihus brevispina sp. nov., holotype, Karubar stn 3 (MNHN), calicular, basal, and side views

x 2. 1 , x 2.1, x 2.5, respectively.

FIG. 12 g-i. — Trochocyathus longispina sp. nov., holotype, MUSORSTOM I stn 50 (MNHN), calicular. basal, and edge

views, x 2.6. x 2.1, x 2.8, respectively.

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

221

Fig. 13 a-c. — Trochocyathus hasiatus Bourne, 1903, synlype, Tutanga (BMNH 1903.12.1.2), calicular, basal, and edge

views, all x 3.4.

Fig. 13 d-e. — Paracyathus rotundatus Semper, 1872: d, "Alpha Helix" stn 79-M21 (USNM 80015), side view, x 2.6. —

e, holotype, Philippines (NMW 8177), calice, x 4.2.

Fig. 13 f. — Stephanocyathus spiniger (Marenzeller, 1888), "Siboga" sin 159 (ZMA Coel. 1305), hololype of

Odontocyathus Stella, x 3.2.

Fig. 13 g-i. — Paracyathus sp.: g. "Challenger" stn 190 (BMNH), side view, x 9.2. — h. Deki (Ambon) (NNM), calice,

x 13. — i, "Siboga" stn 256 (ZMA Coel. 1306), calice, x 5.3.

Source :

222

S. D. CAIRNS & H. ZIBROWIUS

Fig. 14 a-c. — Stephanocyathus regius sp. nov., holotype, "Hakuho Maru" sin KH 72-1-26 (USNM 97122) stereo

calicular and basal views, both x 1.7.

FlG„|f d' ~ Stephanocyathus spiniger (Marenzeller, 1888), "Siboga" stn 156 (ZMA Coel. 1304), base of syntvpe of

Udontocyathus sexradu.x 1.

5°' \\ %' ~ Stephanocyathus explanans (Marenzeller, 1904), Karubar stn 40 (MNHN), oblique calicular view, x 2.

HG. 14 t. Sabinotrochus bi patella Alcock, 1902, holotype, "Siboga" stn 284 (ZMA Coel. 1322) x 6

14 g-h. - Stephanocyathus weberianus (Alcock, 1902): g, syntype. "Siboga" stn 284 (ZMA Coel. 1322), side

x h, Karubar stn 91 (MNHN), labyrinthiform columella, x 2.

Fig. 14 i. — "Sabinotrochus" Jlatiliseptis Alcock. 1902, holotype. "Siboga" stn 21 1 (ZMA Coel. 1315), x 4.4.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

223

Fig. 15 a-b. — Ericiocyathus echinatus sp. nov., holotype, "Albatross" stn 5425 (USNM 97169), side and calicuiar

views, both x 2.4.

Fig. 15 c. — Deltocyathus andamanicus Alcock, 1898, "Albatross" stn 5417 (USNM 97185). calicuiar view, x 2.5.

Fig. 15 d-e. — Deltocyathus philippinensis sp. nov., holotype. "Albatross" stn 5506 (USNM 97178), stereo calicuiar

and basal views, both x 3.6.

Fig. 15 f-h. — Deltocyathus Stella sp. nov.: f-g. holotype, Karubar stn 1 (MNHN), calicuiar and basal views, both

x 4.2. — h. paratypes, Karubar stn I (MNHN), 4 coralla showing growth stages, x 3.

Source :

224

S. D. CAIRNS & H. ZIBROWIUS

Fig. 16 a-c. — Deltocyathus rolulus (Alcock, 1898): a, "Albatross" stn 5582 (USNM 92727), corallum having Pi. 4,

x 1.4. — b. "Albatross" stn 5585, corallum with very small P|.3 (USNM 97207), x 1.6. — c. "Siboga" stn 45 (ZMA),

a worn syntype of D. fragilis, x 2.8.

FIG. 16 d. — Deltocyathus suluensis, Alcock, 1902, syntype, "Siboga" stn 95 (ZMA Coel. 5442), x 3.6.

FIG. 16 e. — Conotrochus brunneus Moseley, 1881. holotype. "Challenger" stn 194 (BMNH 1880.1 1.25.62), x 10.1.

Fig. 16 f-i. — Lochmaeotrochus oculeus Alcock, 1902: f, h-i, Karubar stn 13 (MNHN): calice and coralliles from same

colony, x 5.6. x 2.5, x 2.5, respectively. — g, syntype, "Siboga" stn 259 (ZMA Coel. 700), x 3.4.

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

225

Fig. 17 a-b. — Asterosmilia marchadi (Chevalier. 1966). Mortensen's Java Exp., stn 8 (ZMUC): a. side of corallum

showing several detachment scars, x 3.4; b, oblique calicular view, x 3.4.

Fig. 17 c, f. i. — Phyllangia papuensis Studer, 1878. Musorstom 2 stn 47 (MNHN), colony, calice, and costal

granulation, x 0.75, x 11.3, x 12.1, respectively.

Fig. 17 d-e. — Thalamophyllia tenuescens (Gardiner. 1899). COR1NDON 2 stn 248 (MNHN), side and calicular views of

a colony, x 1.6, x 2.7, respectively.

Fig. 17 g-h. — Desmophyllum dianthus (Esper. 1794), "Siboga" stn 259 (ZMA Coel. 1242), side and calicular views,

x 1.2, x 2, respectively.

226

S. D. CAIRNS & H. ZIBROWIUS

Fig. 18 a-b. — Rhizosmilia elata sp. nov„ holotype. "Albatross" stn 5244 (USNM 97304), corallum and calice x 0 83

x 4, respectively.

FIG. 18 c-g. — Sympodangia albatross i sp. nov.: c. g. paratypes, Karubar stn 18 (USNM 97311): c. longitudinal

sInC5398S(USNM 0™ , “!'!°"' X 8' C°S'al granulaIion over a barnacle, x 43. - d. holotype, "Albatross"

stn 5398 (USNM 97308), x 1.8. - e-f, paratypes. ’Albatross" stn 5398 (USNM 97309), branch and calice. x 10 1

x zj, respectively.

FlGx’ 2 h5 — Alatotrochus rubescens (Moseley, 1876), MUSORSTOM 3 stn 102 (MNHN), side view showing enlarged Ci,

F'G incipient SSScitT^ * Eg"C“' ”37>' “ 5586 <USNM ™“>. view showing

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

227

Fig. 19 a-c. — Coenosmilia arbuscula Pourtates, 1874, "Albatross" stn 5543 (USNM 97312), colony, corallite with small

buds, and calice, x 0.8, x 2.1, x 3.2, respectively.

Fig. 19 d-g. — Confluphyllia juncta sp. nov.: d-f, holotype, Karubar stn 25 (MNHN): d. colony, x 0.8; e, corallites

united by coenosteal bridges, x 2.4; f. distal branch, x 2.4. — g, paratype, Karubar stn 25 (USNM 97316), calice

showing columella, x 7.5.

Fig. 19 h-i. — "Tropidocyathus" pileus (Alcock, 1902), Karubar stn 2 (POL1PI), side and calicular views of corallum

with S5, x 2, x 2.3, respectively.

228

S. D. CAIRNS & H. ZIBROWIUS

Fig. 20 a-g. — "Tropidocyaihus" labidus sp. nov.: a-b. holotype, Karubar stn 2 (MNHN), side and calicular views,

x 5.4, x 6.6. respectively. — c-g. paratypes. Karubar stn 2 (MNHN): c. costal granulation near calice, x 30.5!

d-e, side and calicular views, x 9.1, x 9.8, respectively; f, higher cycle costal origins near base, x 30.5;

g, enlargement of pali and columella, x 20.

Fig. 20 h. — Flabellum pavoninum Lesson, 1831, Musorstom 3 stn 131 (USNM 97455), side view, x 2.1.

Fig. 20 i. — Flabellum patens Moseley, 1881, Karubar stn 31 (MNHN), side of corallum showing band of discolouration

(erosion) caused by Lumbrineris polychaete, x 1.

Source : MNHN Paris

AZOOXANTHELLATE SCLERACTINIA

229

Fig. 21 a. — Flabellum lamellulosum Alcock, 1902, Musorstom 1 stn 71 (MNHN), columellar gall of a petrarcid

ascothoracidan crustacean, x 3.

Fig. 21 b-c. — Flabellum conuis Moseley, 1881, "Hakuho Maru" stn KH72-1-8 (USNM 97 472), side and calicular views,

x 1, x 1.1, respectively.

Fig. 21 d-f. — Flabellum sp.. Musorstom 2 stn 78 (MNHN), edge, side, and calicular views, x 1.7, x 1.6. x 1.8,

respectively.

Fig. 21 g-h. — Gardineria paradoxa (Pourtalfes, 1868): g, Barbados (USNM 80893), calice, x 4.1. — h, Karubar stn 5

(MNHN), calice, x 4.1.

Fig. 21 i. — Javania pachy theca Cairns, 1995, "Albatross" stn 5634 (USNM 97496), calicular view, x 6.3.

Source :

230

S. D. CAIRNS & H. ZIBROWIUS

Fig. 22 a. —Javania pachytheca Cairns, 1995, "Albatross" stn 5634 (USNM 97486), side view, x 3.1.

Fig. 22 b-c. Javania sp.. Karubar stn 44 (MNHN), side and calicular views, x 1.9, x 3.4, respectively.

Fig. 22 d-e. — Rhizotrochus "typus" H. Milne Edwards & Haime, 1848, Deki stn 25 (NNM 23096): d, oblique view of base

showing scar of transverse division, x 2.9; e. calice, x 2.9.

Fig. 22 f — Truncatoflabellum paripavoninum (Alcock, 1894), Karubar stn 71 (MNHN), corallum with suberitid sponge

attached, x 0.52. 6

Fig. 22 §-h. — Truncatoflabellum mortenseni sp. nov.: g. holotype, MORTENSEN'S Java Exp. stn 5 (ZMUC), calice x 3 3

— h-. ParatyPes- MORTENSEN'S Java Exp. stn 5 (USNM 97522), 5 small coralla illustrating stages of ihc transverse

division process, x 1.7. 66

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

231

Fig. 23 a-b. — Truncatoflabellum spheniscus (Dana, 1846), "Hakuho Maru" sin KH72-1-30 (USNM 97501): a. side view,

x 1.1; b, 2 anthocauli, one in process of dividing, x 1.8.

Fig. 23 c-f. — Truncatoflabellum angustum sp. nov.: c-e. hololype. Musorstom 3 sin 143 (MNHN). edge, side, and

calicular views, x 4.5, x 3.3, x 3.7, respectively. — f, paratype, Musorstom 3 stn 130 (MNHN), anthocyathus and

anthocaulus of same specimen, x 5.8.

Fig. 23 g-h. — Balanophyllia desmophyllioides Vaughan, 1907, Musorstom 1 stn 3 (MNHN), side and calicular views,

x 1,5, x 1.7, respectively.

Fig. 23 i. — Balanophyllia parvula Moseley, 1881, Karubar stn 49 (MNHN), side view, x 3.3.

Source : MNHN , Paris

232

S. D. CAIRNS & H. Z1BR0WIUS

Fig. 24 a. — Balanophyllia parvula Moseley. 1881. Karubar stn 49 (MNHN), calice, x 4.7.

Fig. 24 b-c. — Balanophyllia serrata sp. nov., holotype. MUSORSTOM 1 sin 69 (MNHN). side and calicular views, x 1 1

x 1.6, respectively.

FIG. 24 d-f — Balanophyllia cornu Moseley, 1881: d-e, "Albatross" sin 5280 (USNM 97574). side and calicular views of

attached form, x2.1, x 3.1, respectively. — f, "Albatross" sin 5313 (USNM 92880), side view of curved form,

Fig. 24 g-i. Balanophyllia gemma (Moseley. 1881): g, i, holotype, "Challenger" stn 201 (BMNH 1880.11.25.147).

epUheca C3l‘CUlar vieWS' X 5'2' X 6'6‘ resPectively- — *>, "Albatross" stn 5135 (USNM 97592), side view showing

Fig. 25 a-c. — Balanophyllia crassiseptum sp. nov., holotype, Karubar stn 50 (MNHN): a-b. side and calicular views,

x 3.2, x 3.8, respectively; c, other side showing boring of an acrothoracican cirripede. x 2.9.

FlG. 25 d-f, — Balanophyllia rediviva Moseley, 1881, Karubar stn 22 (MNHN), corallum, costae, and calice of same

specimen, x 1.1, x 3.2, x 6.5, respectively.

Fig. 25 g-i. — Balanophyllia generatrix sp. nov.: g, i, paralype, "Siboga" stn 41 (ZMA 5538), top and side view of same

pseudocolony, both x 0.6. — h, holotype, Karubar stn 82 (MNHN), x 0.6.

AZOOX ANTHELLATE SCLERACTINIA

233

Source :

234

S. D. CAIRNS & H. ZIBROWIUS

Fig. 26 a-b. — Balanophyllia generatrix sp. nov„ paratype, "Siboga" sin 41 (ZMA Coel. 5538), 2 calicular views, both

x 3.4.

FIG. 26 c-f. — Balanophyllia imperials Saville Kent, 1871: c, f, "Hakuho Man," stn KH72-1-30 (USNM 97608), side and

calicular views, x 2, x 3.8, respectively. — d-e, holotype, Singapore (BMNH 1984.4.27.3), side and calicular views,

x 1.4, x 1.7, respectively.

Fig. 26 g-i. Leptopsammia stokesiana H. Milne Edwards & Haime, 1848: g, Mortensen's Pacific Exp (Jolo Island)

(NNM), side view, x 3.8. — h-i, holotype, Philippines (BMNH 1855,12.27.1), side and calicular views, x 2.9, x 5 5

respectively.

Source : MNHN. Paris

AZOOXANTHELLATE SCLERACTINIA

235

Fig. 27 a-g. — Endopachys bulbosa sp. nov.: a-d. holotype. Karubar stn 62 (MNHN), side. edge, calicular, and basal

views, x 1.1, x 1.2, x 1.15, x 1.4, respectively; e-g, paratypes, Karubar sin 62 (MNHN): e-g, side and 2 basal

views showing basal thickening and broad C1-2, x 1.3, x 1.8, x 1.9, respectively.

Fig. 27 h-i. — Leptopsammia crassa van der Horst. 1922, holotype, "Siboga" stn 258 (ZMA Coel. 8462), side and

calicular views, x 2.8, x 4.5, respectively.

Source : MNHN, Paris

236

S. D. CAIRNS & H. ZIBROWIUS

Fig. 28 a-b. — Leptopsammia poculum (Alcock. 1902). holotype, "Siboga" stn 260. side and calicular views x 2 6 x 7

respectively.

F'G 28 c-e. — Endopsammia philippensis H. Milne Edwards & Haime. 1848. "Siboga" stn 213 (ZMA Coel 568)

(BalanophyU.a regulans of van der Horst. 1922), side and calicular views, x 5.8. x 6. x 6.8. respectively

remoidg'rolo^' vTrmia T ^ H°rSt’ 1 9 22> "Siboga" stn 282 (ZMA Coel. 5478). calice and

reptoia colony, x 6.1, x 1, respectively.

F,Gcoloiy,^ R{“1Z°pSamm,a "unula van der Horsl- 1922- holotype. "Siboga" stn 279, ZMA Coel. 6896, stereo view of

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

237

Fig. 29 a-c. — Dendrophyllia arbuscula van der Horst, 1922; a. syntype, "Siboga" stn 277 (ZMA Coel. 5477), x 0.75. —

b-c, "Albatross" stn 5279 (USNM 97630), colony and calice, x 0.81, x 4.4, respectively.

Fig. 29 d. — Cladopsammia echinata Cairns 1984, Karubar stn 86 (USNM 97628), colony, x 0.55.

Fig. 29 c. — Dendrophyllia sp. cf. D. ijimai, "Siboga" stn 49a (ZMA Coel. 5467), branch, x I.

Fig. 29 f. — Enallopsammia pusilla (Alcock, 1902), Karubar stn 25 (MNHN), colony fragment, x 0.36.

Fig. 29 g-i. — Incertae Sedis, "Albatross" stn 5179: g, 3 corallites and epitheca, x 9.7; h, calice, x 22; i, enlargement of

septa and columella, x 42.

Source :

238

S. D. CAIRNS & H. ZIBROWIUS

INDEX

Only the "SYSTEMATIC ACCOUNT" (p. 66-198) and the photographs (p. 209-237) are indexed.

Generic (and subgeneric) names used herein in combination with specific (or subspecific) names, or in citation,

are given between brackets. / separates variant spellings; spelling retained herein in first position.

Taxa of generic and specific level that receive full taxonomic treatment herein are in bold.

Page numbers in bold refer to full taxonomic treatment; numbers in italics refer to illustrations.

Acanthocyathus 96-102, 103

Acinocyathus 114, 118-119

Acropora 80, 8 1

aculeatum ( Flabellum , Truncatoflabellum)

166-167

aequiserialis (Coenopsammia, Dendrophyllia ) 195,

196

affinis ( Balanophyllia , Rhodopsammia ) 1 76

agassizi ( Paracyathus ) 1 16

alabastrum (Desmophyllum) 164

Alatotrochus 141-142, 226

albatrossi ( Sympodangia ) 136-137, 226

alcocki ( Dendrophyllia , Sclerhelia) 193

alcocki ( Madrepora ) 79

Alcockia 190

ambrosia ( Caryophyllia ) 88, 95-96

amoena (Rhodopsammia) 1 75

A mphel ia/A mphihelia 79, 80, 83, 195

amphelioides (Coenopsammia, Dendrophyllia) 195

andamanicus (Deltocyathus) 86, 122, 124,

223

angustum ( Truncatoflabellum ) 172-173 ,231

Anisopsammia 195

annulata (Guynia) 150

Anomocora 134, 135

Anthemiphyllia 86-87, 214

apertus (Trochocyathus) 102, 103, 105, 109-

110. 219

Aplocyathus 113-114, 115, 120

arbuscula ( Amphihelia , Lophohelia, Madrepora)

79,80-81,82,2/7

arbuscula (Cladocora) 1 38

arbuscula (Coenosmilia) 138-139, 227

arbuscula (Dendrophyllia) 192-193, 237

asanoi (Madracis) 66-67. 209

asperula ( Madracis ) 66, 67

Asterosmilia 131-132, 225

Aulocyathus 129-130

a urea ( Dendrophyllia , Lobophyllia, Tubastraea) 197

australe (Flabellum) 152

australiensis (Peponocyathus) 144, 145

australiensis (Turbinolia) 140, 141

axillaris ( Cyathelia/Cyathohelia , Madrepora) 84

Balanophyllia 108, 175-185, 189, 190 231-

234

barbadensis (Caryophyllia) 92

barbadensis ( Duncania , Gardineria) 163

Bathyactis 68. 69, 70-73

bipatella (Sabinotrochus) 1 18, 222

Blastomussa 135

Blastotrochus 173-174

brevispina (Aplocyathus , Trochocyathus ),

105, 113, 220

brunneus (Conotrochus , Phloeocyathus) 127,

129, 224

bulbosa ( Endopachys ) 185, 186, 235

burchae (Caryophyllia, Premocyathus,

Trochocyathus) 103, 105, 110

cailleti (Javania) 165

candeanum (Flabellum, Truncatoflabellum)

167

candeanus (Placotrochus) 175

carinata (Balanophyllia, Rhodopsammia) 175-

176, 177

Caryophyllia 87-102, 103, 109, 110, 130, 138,

214-217

caryophylloide s/cary ophyllioides

( Trochocyathus ) 105, 106, 107

cepulla (Trochocyathus) 111, 112

ceratoconus (Caryophyllia, Premocyathus) 102, 103

Ceratotrochus 117, 119, 120, 127, 128, 129, 142

cervicornis ( Dactylotrochus , Tridacophyllia)

131

chamissoi ( Rhizopsammia ) 189

Citharocyathus 1 43

Cladocora 137, 138

Cladopsammia 189, 191, 237

davits (Caryophyllia) 89, 90, 91, 130

coalition (Flabellum) 151

coccinea (Tubastraea) 197

Coelosmilia 138

Coenocyathus 134

Coenopsammia 194, 195, 197

Coenosmilia 138-139, 227

Colangia 137-138

coloradus ( Odontocyathus ) 1 18

complicata (Stephanophyllia) 76, 77-78

compressa/compressus ( Caryophyllia,

Premocyathus) 102, 103, 109, 110

conferta (Cladocora) 137, 138

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

239

Confluphyllia 139-140, 22 7

conica (Leptopsammia) 176

conicus ( Notocyathus ) 143-144

Conocyathus 140-141

Conotrochus 127-128, 129, 224

conuis ( Flabellum , Ulocyathus ) 160, 229

cooperi (Trochocyalhus, Tropidocyathus) 105,

108, 109, 111, 1 15, 219

cornu ( Balanophyllia ) 178-179, 232

cornulum (Caryophyllia) 88, 94-95, 215

crassa ( Leptopsammia ) 187, 235

crassiseptum ( Balanophyllia ) 180-181, 233

Crispatolrochus 103-104,2/8

crispus ( Fungiacyathus ) 73

crist agalli ( Desmophyllum ) 131

crosnieri ( Caryophyllia ) 87. 89

Cryptotrochus 142-143

Culicia 78-79, 198, 211

cultrifera ( Caryophyllia ) 94

Cyathelia/Cyaihohelia 84

Cyathoceras 103, 218

cylindrica ( Dendrophyllia ) 1 92

Dactylotrochus 131

Dasmosmilia 131

decactis (Madracis) 66

defilippi (Paracyathus) 1 16

delicatulus ( Labyrinthocyathus ) 1 04

Deltocyathoides 144-145

Deltocyathus 86, 115, 121-127, 144, 223-224

deludens ( Flabellum , Ulocyathus) 154-156

157, 158

Dendrophyllia 184, 189, 190, 191-193, 194,

195, 196, 197, 237

dennanti (Fungiacyathus) 73

dens ( Flabellum , Truncatoflabellum) 170-

171, 172

dentata/dentatus ( Anthemiphyllia ,

Discotrochus) 86

dentata/dentatus ( Acanthocyathus ,

Caryophyllia) 96, 98-99, 216

dentiformis (Placotrochides, Premocyathus)

102-103, 110, 2/7

desmophyllioides/desmophylloides

( Balanophyllia ) 177-178, 231

Desmophyllum 79, 131, 133, 152, 164, 225

diaphana ( Dendrophyllia , Tubastraea) 196-197

dianthus (Desmophyllum, Madrepora) 131,

152, 225

diomedeae (Caryophyllia) 87, 88

dioniedeae (Cyathoceras) 103

Diplohelia 194

Discotrochus 86, 145

discus (Trochocyathus) 105, 111, 112, 219-220

distinction (Flabellum) 1 52

dumosa (Goniocorella, Pourtalosmilia) 139

eccentricus (Deltocyathus) 123

echinata (Cladopsammia) 191 ,237

echinatus (Ericiocyathus) 120-121, 223

Eguchipsammia 190-191

elata (Rhizosmilia) 134-135. 226

elegans (Flabellum) 167

elongata ( Caryophyllia ) 89

elongatum (Flabellum) 174

Enallopsammia 194-195. 237

Endocyathopora 141

Endopachys 185-186, 235

Endopsammia 188. 236

ephyala (Caryophyllia) 88

epithecata ( Caryophyllia ) 130

erecta (Dendrophyllia) 192

Ericiocyathus 120-121, 223

Eupsammia 176

explanans (Acinocyathus , Stephanocyathus ,

Stephanotrochus) 1 14, 119. 222

facetus (Labyrinthocyathus) 1 04

faulkneri (Tubastraea) 197

fecunda (Anomocora, Coelosmilia, Coenosmilia,

Parasmilia) 134, 138

fissidiscus ( Bathyactis . Fungiacyathus) 72-

73, 210

fissilis (Fungiacyathus) 73

fistula (Balanophyllia) 1 90

flabelliformis (Rhizotrochus) 161-162

Flabellum 70, 108, 146, 150-160, 161, 162,

163, 166, 167, 168. 169, 170, 174, 228-229

flatiliseptis (Sabinotrochus) 118, 222

floreana (Tubastraea) 197

folliculus (Peponocyathus, Stephanophyllia)

144, 145, 146

formosa (Madrepora, Sclerohelia) 79

formosissima (Letepsammia, Stephanophyllia)

73-75, 76

formosum (Truncatoflabellum) 169-170

frag il is (Bathyactis, Fungiacyathus) 71

fragilis (Deltocyathus) 125

franki (Letepsammia) 74

frustum (Anthemiphyllia) 86-87. 214

fruticosa (Dendrophyllia) 196

Fungia 68, 69, 70. 71, 77

Fungiacyathus 68-73, 87, 210

fun gulus (Stephanophyllia) 76-77. 78

funicolumna (Ceratotrochus, Conotrochus)

127, 128

gaditana (Dendrophyllia, Eguchipsammia),

190, 191, 193

240

S. D. CAIRNS & H. ZIBROWIUS

gardineri ( Paracyathus , Trochocyathus ) 105,

111-112

gardineri (' Truncatoflabellum ) 173

Gardineria 162-163, 229

gemma (. Balanophyllia , Thecopsammia ) 179,

232

generatrix ( Balanophyllia ) 183-184, 233-234

gerdae ( Rhizosmilia ) 135

gigas ( Balanophyllia ) 178, 182, 184, 185

gigas ( Rhizosmilia ) 1 35

Goniocorella 115, 140

gracilis ( Dendrophyllia ) 189

grandis (Cary ophy Ilia) 88, 96, 215

grandis (Dendrophyllia) 195

granulosa/glanulosa ( Goniocorella ) 1 39

granulosus (Fungiacyathus , Bathyactis) 71

grayi (Endopachys) 185-186

grayi (Acanthocyathus , Caryophyllia) 96,

97-98. 99. 100, 2/5

guangdongensis (Acanthocyathus, Caryophyllia) 97

Guy nia 150

Haplophyllia 163

hastatus (Aplocyathus, Trochocyathus) 113, 114,

221

liawaiiensis (Caryophyllia) 88, 9 3

hawaiiensis (Gardineria) 163

heteroclitus (Deltocyathus) 124

Heteropsammia 1 50

hoffmeisteri (Flabellum , Ulocyathus) 155,

157-158

Hoplangia 136

horsti (Dendrophyllia) 192, 193

hospes (Ceratotrochus. Phloecyathus) 127, 129

Idiotrochus 148-149

ijimai (Dendrophyllia) 191-192, 195. 237

immersa (Colangia) 138

imme rsa (Oculina) 83

impensum ( Flabellum ) 152

imperials ( Balanophyllia ) 184-185,254

incerta (Caryophyllia) 92

incerta (Rhodopsammia) 176

incertae sedis 198, 237

incrustatum (Truncatoflabellum) 168, 169

indica (Dendrophyllia) 192

infundibulifera ( Amphelia/Amphihelia , Madrepora,

Oculina) 82, 83

insignis (Javania) 163-164

irregular e ( Flabellum , Truncatoflabellum)

168

interjecta (Madracis) 66

italica (Turbinolia) 122, 125

ixine (Odontocyathus, Stephanocyathus) 120

japonica (Culicia) 78, 79

japonica (Letepsammia, Stephanophyllia) 75, 77

japonicum ( Flabellum , Ulocyathus) 70, 155,

157. 158

Javania 163-164, 229-230

javanus (Cryptotrochus) 142

juncta (Confluphyllia) 139-140. 227

juvenescens ( Aulocyathus ) 130

kauaiensis (Madracis) 66

karubarica (Acanthocyathus , Caryophyllia)

97, 217

kikaiensis (Bathyactis) 69

kikutii (Idiotrochus, Placotrochides) 148-149

kirbyi (Madracis) 66

labidus (Tropidocyathus) 148, 228

Labyrinthocyathus 104-105. 218

laciniatum (Flabellum) 158

laevis (Placotrochus) 175

lamellifera (Caryophyllia) 87, 88, 90

lamellulosum ( Flabellum ) 151, 229

lamprotichum (Javania) 164

langae (Labyrinthocyathus) 1 04

laoagana ( Acanthocyathus , Caryophyllia) 98

laticostata (Endocyathopora) 141

latum ( Flabellum ) 161

lawtoni (Blastomussa) 135

laysanensis (Balanophyllia) 182

lens (Deltocyathus) 1 44

leptoclados (Oculina) 83

Leptopenus 73. 210

Leptopsammia 176, 186-188, 234-236

lessonii/lessoni ( Trochocyathus ,

Tropidocyathus) 146-147

Letepsammia 73-75, 76

Levipalifer 122

limatulus (Labyrinthocyathus) 104

Lobophyllia 197

Lochmaeotrochus 128-129, 224

longispina (Aplocyathus, Trochocyathus)

105. 113-114, 219

Lophohelia 79, 80

maclurii (Endopachys) 186

maculata ( Rhizosmilia ) 135

maculatus (Trochocyathus) 105, 107

Madracis/Madrasis 66-68. 209

Madrepora 77. 79-84. 85, 131, 136. 211-213

magnificus (Deltocyathus) 121, 125

magnificum ( Flabellum ) 151-152, 153

marchadi ( Asterosmilia , Dasmosmilia) 131-

132, 225

marenzelleri ( Enallopsammia ) 194

Source : MNHN, Paris

AZOOXANTHELLATE SCLERACTINIA

241

marenzelleri ( Flabellum , Ulocyathus), 155,

156

marmorea ( Caryophyllia ) 92

messum ( Flabellum , Ulocyathus) 158

micranthus ( Dendrophyllia , Oculina,

Tubastraea) 192, 195-196, 197

minimus ( Cylindrophyllia , Discotrochus,

Peponocyathus) 145-146, 226

minuscula (Dendrophyllia) 191, 192

minuta ( Rhizopsammia ) 189, 236

minutiseptum ( Madrepora ) 82-84, 211-212

mortenseni (Truncato flabellum) 171-172,

230

moseleyi ( Cladocora , Colangia) 137-138

moseleyi (Flabellum) 159

multilobatus ( Thrypticotrochus ) 149

multipalifera (Vaughanella) 118

neglecta (Stephanophyllia) 75, 77

Neohelia 84-85, 2/3

niphada (Rhombopsammia) 75-76

nobile (Desmophyllum) 165

nobilis (Ceratotrochus, Stephanotrochus) 117, 119,

120

Notocyathus 143-144

nuda ( Rhizopsammia ) 189-190

nutrix ( Blastotrochus , Flabellum) 173-174

octonaria ( Caryophyllia ) 87, 92, 2/5

octopali ( Caryophyllia ) 92

oculata ( Amphihelia , Madrepora) 79-80, 81,

82, 83

oculeus ( Lochmaeotrochus ) 128-129, 224

Oculina 82, 83, 195

Odontocyathus 113, 118, 1 19-120, 221-222

orientalis ( Cylindrophyllia , Deltocyathoides ,

Deltocyathus, Peponocyathus) 144-145, 146

orientalis (Levipalifer) 122

omatus ( Deltocyathus ) 124

pachyclados ( Oculina ) 83

pachytheca ( Javania ) 165, 229-230

palaoensis (Madracis) 66

paliferus/palifera ( Bathyactis , Fungiacyathus)

69-70

paliferus ( Stephanocyathus ) 1 17

palita ( Dendrophyllia ) 193

panda (Caryophyllia) 87

papuensis (Phyllangia) 135-136, 225

Paraconotrochus 130

Paracyathus 112, 115-117, 22/

paradoxa (Gardineria, Haplophyllia) 163, 229

parallela (Balanophyllia, Rhodopsammia) 175, 176

Parasmilia 138

paripavoninum ( Flabellum ,

Truncatoflabellum) 169, 230

parvula (Balanophyllia) 180, 231-232

patella (Fungia, Madrepora) 77

patens (Flabellum) 152, 153, 228

pavoninum (Flabellum) 150-151, 152, 228

Peponocyathus 144, 145-147, 226

pharensis (Madracis) 67, 209

philippensis/philippinensis (Endopsammia)

188, 236

philippinensis (Deltocyathus) 122-123, 223

philippinensis ( Gardineria ) 162-163

philippinensis (Trochocyathus) 105, 106,

107-108, 109, 218

phoenix (Truncatoflabellum) 171

Phyllangia 135-136, 225

pileus (Trochocyathus, Tropidocyathus) 147-

148, 227

Placotrochides 102, 103, 148, 174

Placotrochus 175

Pleurocyathus 111

poculum (Leptopsammia) 1 87, 236

politum (Flabellum) 153

Polycyathus 137, 138

Polymyces 160-161

porcellana (Neohelia) 84-85, 2/3

Pourtalosmilia 139

Premocyathus 100, 102-103, 109, 110, 2/7

primordialis (Tridacophyllia) 131

profunda (Coenopsammia, Dendrophyllia,

Diplohelia) 194

profunda (Madracis) 68

pusilla (Coenopsammia, Dendrophyllia,

Enallopsammia) 194, 195, 237

pusillum (Truncatoflabellum) 170

quadragenaria (Caryophyllia) 88, 93

ramea (Amphihelia, Madrepora) 79

ramea (Dendrophyllia) 192

recidivus (Aulocyathus, Ceratotrochus) 129-

130

rediviva (Balanophyllia) 181-182, 233

regius (Stephanocyathus) 117-118, 222

regularis (Balanophyllia, Thecopsammia) 188

Rhizopsammia 188-190, 236

Rhizosmilia 133-135, 226

Rhizotrochus 152, 161-162

Rhodopsammia 175, 176

rhombocolumna (Trochocyathus) 105, 106-

107

Rhombopsammia 74, 75-76

robusta (Rhizosmilia) 133-134

rostrata (Amphihelia, Anisopsammia,

Enallopsammia) 195

242

S. D. CAIRNS & H. ZIBROWIUS

rotulus (Deltocyathus, Trochocyathus) 121,

125-126, 224

rotundatus ( Paracyathus ) 115-116, 221

rubescens ( Alatotrochus , Platytrochus,

Sphenotrochus) 141-142, 226

rubescens ( Crispatotrochus , Cyathoceras)

103-104, 218

rugosa ( Caryophyllia ) 87, 91-92

rugosus (Crispatotrochus) 104

Sabinotrochus 1 18, 222

sagamiensis ( Coenocyathus , Rhizosmilia) 134

scaphula ( Placotrochides ) 174

Sclerhelia/Sclerohelia 79, 191

scobinosa ( Caryophyllia ) 88, 94, 95

secfa ( Caryophyllia ) 87, 89-90, 91, 2/4

semperi ( Trochocyathus ) 105, 108-109, 2/5-

219

serrata ( Balanophyllia ) 182-183, 185, 252

sexcostatum ( Flabellum , Ulocyathus) 159

sexradii (Odontocyathus) 222

sibogae ( Bathyactis , Fungiacyathus) 69, 70,

71

sibogae ( Dendrophyllia ) 196

sibogae ( Stephanotrochus ) 120

socialis ( Rhodopsammia ) 1 76

spheniscus ( Euphyllia , Truncato flabellum)

165-166, 167, 23/

Sphenotrochus 141

spinicarens/spinacarens ( Acanthocyathus ,

Caryophyllia , Premocyathus) 97, 98, 100-

101, 102, 103, 216

spiniger (Acinocyathus , Stephanocyathus ,

Stephanotrochus) 118-119, 22/

spinigera/spiniger (Acanthocyathus,

Caryophyllia) 97, 98 99,216

squiresi (Rhombopsammia) 76

stabilis (Bathyactis) 70

s/e//a (Deltocyathus) 121, 123-124, 225

(Odontocyathus) 221

stellata (Culicia) 78-79, 2/ /

stellulatus (Bourneotrochus, Deltocyathus)

115

Stephanocyathus 114, 117-120, 221-222

Stephanophyllia 73, 74, 75, 76-78, 145, 146

Stephanotrochus 118, 119, 120

stephanus (Bathyactis, Fungiacyathus) 68-69,

stimpsonii/stimpsoniana (Balanophyllia,

Eupsammia) 176-177

stokesiana (Balanophyllia, Leptopsammia)

186-187, 234

subcomigera ( Dendrophyllia ) 192

sulcatus (Conocyathus) 140

suluensis (Deltocyathus) 121, 125, 224

superstes (Letepsammia, Stephanophyllia) 74,

75

symmetrica ( Bathyactis , Fungia) 68, 69, 70, 7 1

Sympodangia 136-137, 226

tagusensis (Tubastraea) 197

tenuescens (Desmophyllum, Thalamophyllia)

133, 225

tenuis/tenui (Amphihelia, Lophohelia, Madrepora)

79

Tethocyathus 114-115

Thalamophyllia 133, 164, 225

Thecopsammia 179, 188

Thrypticotrochus 149

transversalis (Caryophyllia) 88, 90-91, 2/4

Trematotrochus 140

Trochocyathus 84, 102, 103, 105-114, 115,

120, 146, 147, 218-221

Tropidocyathus Ill, 146-148, 227-228

truncata (Culicia) 78, 79

Truncato flabellum 162, 165-173, 174, 230-

231

Tubastraea/Tubastrea 195-197

tubulifera (Oculina) 83

turbinata (Dendrophyllia) 197

Turbinolia 122, 125

turbinolioides (Bathyactis , Fungiacyathus)

72, 87

tydemani (Cyathoceras) 103, 104, 218

typus (Rhizotrochus) 152, 161

Ulocyathus 154-160

unicristata (Acanthocyathus, Caryophyllia)

97, 98, 101-102, 2/7

variegatus (Bathyactis, Fungiacyathus) 71-

72

Vaughanella 1 1 8

vaughani (Deltocyathus) 121, 122

velata (Dendrophyllia) 192

venustus (Ceratotrochus, Cryptotrochus) 142-

143

venustus ( Citharocyathus , Notocyathus) 143

144

veroni (Bourneotrochus) 1 15

verrilli (Rhizopsammia) 188-189, 236

virgatus (Tethocyathus, Trochocyathus) 114-

115

virginea (Madrepora, Oculina) 83

viridis (Coenopsammia) 195

weberi (Endopachys) 185

Source : MNHN , Paris

AZOOXANTHELLATE SCLERACTINIA

243

weberi ( Flabellum ) 163

weberi ( Trochocyathus ) 1 1 1

weberianus ( Odontocyathus ,

Stephanocyathus, Stephanotrochus) 119-

120, 222

wellsi ( Eguchipsammia ) 190-191

wellsi ( Polymyces ) 160

willey i ( Coenopsammia , Dendrophyllia) 197

zeidleri (Paraconotrochus) 130

zelandiae ( Conotrochus , Trematotrochus ) 140-

141

zanzibarensis (Acanthocyathus, Caryophyllia) 100

102, 103

zuluense ( Truncatoflabellum ) 172, 173

Source : MNHN, Paris

SUIT rs DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS E

Mollusca Bivalvia: Pectinoidea (Propeamussiidae and

Pectinidae) from eastern Indonesia

Henk H. DIJKSTRA

Institute of Systematics and Population Biology

(Zoological Museum)

University of Amsterdam

P.O. Box 94766, 1090 GT Amsterdam, The Netherlands

&

Woro W. KASTORO

Puslitbang Oseanologi - LIPI

P.O. Box 4801/JKTF, Jakarta 11048, Indonesia

ABSTRACT

Thirty species of Pectinoidea are reported from off the Kai and Tanimbar Islands, Arafura Sea, at shelf to bathyal

depths (111-1244 m). Of these, eight are new records for the Indonesian Archipelago and two are new species:

Parvamussium conspecium sp. nov. (Propeamussiidae) and Veprichlamys versipellis sp. nov, (Pectinidae). On average,

the bathymetric occurence of species is shallower in the study area than in the South-West Pacific. Amussium

margaritiferum Dautzenberg & Bavay, 1912, is synonymized with Propeamussium investigatoris (E.A. Smith, 1906), and

Parvamussium crypticum Hayami & Kase, 1993, is synonymized with P. carbaseum Dijkstra, 1991. Lectotypes are

designated for five nominal taxa.

RESUME

Mollusca Bivalvia : Pectinoidea (Propeamussiidae et Pectinidae) de l'lndonesie orientale.

Au cours de la campagne oceanographique franco-indondsienne Karubar en mer d'Arafura, trente espdces de

Pectinoidea ont dtd rdcoltdes au voisinage des Ties Kai et Tanimbar, sur le plateau et la pente continentals entre 111 m et

1244 m de profondeur. Sur ce total, huit sont signalees pour la premidre fois dans l'archipel indondsien et deux espdces

nouvelles sont ddcrites : Parvamussium conspecium sp. nov. (Propeamussiidae) and Veprichlamys versipellis sp. nov.

(Pectinidae). Dune manidre gdndrale, les espdces rdcoltdes, dejd connues du Sud-Ouest Pacifique, ont dtd trouvdes d des

profondeurs moindres qu'ailleurs. Amussium margaritiferum Dautzenberg & Bavay, 1912, est mis en synonymie avec

Propeamussium investigatoris (E.A. Smith, 1906), et Parvamussium crypticum Hayami & Kase, 1993, avec P. carbaseum

Dijkstra, 1991. Des lectotypes sont designes pour cinq espdces nominales.

Dijkstra, H. H. & Kastoro, W. W., 1997. — Mollusca Bivalvia: Pectinoidea (Propeamussiidae and Pectinidae) from

eastern Indonesia. In: A. Crosnier & P. Bouchet (eds), Resultats des Campagnes Musorstom, Volume 16. Mem. Mus.

natn. Hist. not.. 172: 245-285. Paris ISBN: 2-85653-506-2.

Source : MNHN, Paris

246

H. H. DIJKSTRA & W. W. KASTORO

INTRODUCTION

The fauna of the eastern seas of the Indonesian archipelago has remained little known to this date. The Siboga

Expedition carried out nineteen stations (sins 250-268) near the Kai [= Kei] Islands in December 1899 (Tydeman,

1902: 14). At this occasion, seven species of Pectinoidea were collected in nearshore waters (Dautzenberg &

Bavay, 1912). Later, in 1922, during the Danish Expedition to the Kai Islands (Mortensen, 1923),

Dr Th. Mortensen made 63 dredge and trawl hauls in sublittoral to bathyal depths. The Pectinoidea were never

reported on, and this material (now in ZMUC) will be studied and treated elsewhere by the senior author. Finally,

the Indonesian-Dutch SNELLIUS-II Expedition (1984-1985) did some sampling in the northwest of Banda Sea, but

the investigations did not touch the Kai or Tanimbar Islands. The Pectinoidea of that expedition were described by

DIJKSTRA (1991).

The present paper reports on the Propeamussiidae and Pectinidae collected during the Indonesian-French

Karubar cruise. For a narrative of the cruise and complete station list, see CROSNIER, RICHER DE FORGES &

Bouchet (1997: this volume page 9). In addition, a few odd samples collected in 1980 during the Corindon

cruise in the strait ot Makassar have also been considered. We follow the style and presentation of an earlier paper

on deep-water Pectinoidea from the New Caledonia region (Dijkstra, 1995b), and extensive reference is made to

that paper: lists of synonyms and references, diagnosis and descriptions are not repeated for species already

discussed in the context of the New Caledonia fauna. The material is deposited in MNHN, Paris and POLIPI,

Jakarta, with voucher specimens in the private collection of the senior author. Comparative material from

Indonesia and type material was studied from various museum collections, namely AMS, BMNH MNHN

RMNH, WAM, ZMA, ZMUC, ZSI.

ABBREVIATIONS AND TEXT CONVENTIONS

Repositories

AMS

BMNH

HD

IGPS

IOAS

KBIN

POLIPI

MNHN

NMNZ

NMW

NSMT

RMNH

UMUT

USNM

WAM

ZMA

ZMUC

ZSI

Other abbreviations

OD

SD

Australian Museum, Sydney

The Natural History Museum, London

H.H. Dijkstra collection, Sneek

Institute of Geology and Palaeontology, Sendai

Institute of Oceanology, Academia Sinica, Qingdao

Institut Royal des Sciences Naturelles de Belgique, Bruxelles

Puslitbang Oseanologi-LIPI [Research and Development Centre for Oceanology -

Indonesian Institute of Sciences], Jakarta

Museum national d'Histoire naturelle, Paris

Museum of New Zealand Te Papa Tongarewa, Wellington

National Museum of Wales, Cardiff

National Science Museum, Tokyo

Nationaal Natuurhistorisch Museum, Leiden

University Museum, University of Tokyo, Tokyo

National Museum of Natural History, Washington, DC

Western Australian Museum, Perth

Zoologisch Museum, Amsterdam

Zoologisk Museum, Copenhagen

Zoological Survey of India, Calcutta.

Original designation

Subsequent designation

Source : MNHN, Paris

PECTINOIDEA FROM EASTERN INDONESIA

247

SYSTEMATIC ACCOUNT

Class BIVALVIA Linnaeus, 1758

Subclass PTERIOMORPHIA Beurlen, 1944 [emend.. Boss, 1982]

Superorder EUPTERIOMORPHIA Boss, 1982

Order OSTREOIDA Waller, 1978

Suborder PECTININA Waller, 1978

Superfamily PECTINOIDEA Wilkes, 1810 [emend., Waller, 1978]

Family PROPEAMUSS1IDAE Abbott, 1954

Genus PROPEAMUSSIUM de Gregorio, 1884

Propeamussium alcocki (E.A. Smith, 1894)

Figs 1-4

Amussium alcocki E.A. Smith, 1894: 172, pi. 5, figs 15-16.

Propeamussium alcocki - DlJKSTRA, 1995b: 13, figs 1-4, 133-137 [references, description, discussion],

MATERIAL EXAMINED. — The type material (see Dijkstra, 1995b: 13).

Indonesia. Karubar, Tanimbar Islands : stn CP 52, 08°03'S, 131°48’E, 1244-1266 m, 3 spms. — Stn CP 53,

08°18'S, 131°41'E, 1026-1053 m, 2 spms. — Stn CP 89, 08°39'S, 131°08'E, 1058-1084 m, 3 spms. — Stn CP 91,

08°44'S, 131°05'E, 884-891 m, 17 spms, 1 lv.

Distribution. — Gulf of Aden, Laccadive Sea, Bay of Bengal, Coral Sea, New Caledonia, and Loyalty Islands

(Dijkstra, 1995b: 13). Now the Arafura Sea. Present material alive in 891-1244 m.

Propeamussium caducum (E.A. Smith, 1885)

Figs 5-8

Amussium caducum E.A. Smith, 1885: 309, pi. 23, figs 1-lc.

Propeamussium caducum - Dijkstra, 1995b: 15, figs 9-10, 129-132 [synonymy, references, description],

MATERIAL EXAMINED. — The type material (see Dijkstra, 1995b: 17).

Indonesia. Corindon, Makassar Strait: stn B 247, 00°55'S, 119°26'E, 520 m, 1 rv.

Karubar, Kai Islands : stn CP 35, 06°08'S, 132°45'E, 390-502 m, 1 lv. — Stn CP 39, 07°47'S, 132°26'E, 466-

477 m, 17 spms.

Tanimbar Islands: stn CP 54, 08°21'S, 131°43'E, 836-869 m, 4 spms, 2 lv, 1 rv. — Stn CC 56, 08°16'S, 131°59’E,

549-552 m, 29 spms. — Stn CC 57, 08°19'S, 131°53'E, 603-620 m, 6 spms. — Stn CC 58, 08°I9’S, 132°02'E, 457-

248

H. H. D1JKSTRA & W. W. KASTORO

461 in, 3 spms, 3 lv, 4 rv. — Stn CP 71, 08°38’S, 131°44'E, 477-480 m, >50 spms, 7 lv, 3 rv. — Stn CP 72, 08°36'S,

131°33'E, 676-699 in, >50 spms, 7 lv, 4 rv. — Sin CP 73, 08°29’S. 131°33'E, 840-855 m, 2 spms. — Sin CP 75,

08°46'S, 131°36'E, 451-452 m, 17 spms. 3 lv, 1 rv.

Distribution. — Japan, Philippines, Indonesia (Dautzenberg & Bavay, 1912, Thiele & Jaeckel, 1931,

Dijkstra, 1991), Bay of Bengal, Arabian Sea, Gulf of Aden, Tanzania, New Caledonia. Present material alive in

452-840 m.

Remarks. — The present specimens are similar to the type material, but the growth lines are weaker, there are

no radial striae on the left valve, internal ribs number 9 (instead of 10 in the type material), and the colour is cream

instead of whitish transparent.

Propeamussium ina (Dautzenberg & Bavay, 1912)

Figs 9-10

Amussiwn ina Daulzenberg & Bavay, 1912: 32, pi. 28, figs 18-21.

Propeamussium (Parvamussium) ina - DUKSTRA, 1990: 9, 10.

Parvamussium ina - ROMBOUTS, 1991: 68.

MATERIAL EXAMINED. — The type material (see below).

Indonesia. CORINDON, Makassar Siraii: stn B 268, 01°57'S, 119°16'E, 200 m, 2 lv.

Karubar, Kai Islands: stn DW 31. 05°40'S, 132°51'E, 288-289 m, I rv (see Remarks).

Type Material. — Lectotype (H 14, L 14, D 6.5 mm) ZMA Moll. 3.12.01 1, here designated, live taken.

Two paralectotypes: ZMA Moll. 3.12.012. DlJKSTRA (1990: 10) noticed the existence of three complete

specimens in ZMA, instead of 4 valves mentioned by DAUTZENBERG & Bavay (1912: 32). Bavay's manuscript

in KBIN mentions these specimens.

Type Locality. — "Siboga", stn 312, 08°19'S, 1 17°41’E, Saleh Bay, North coast of Sumbawa, Indonesia,

274 m.

Distribution. — Indonesia, shells in 200-288 m, live record in 274 m.

DESCRIPTION. — Shell small, up to ca. 14 mm high, fragile, transparent, slightly orbicular, equivalve,

inequilateral, slightly convex, auricles unequal, umbonal angle ca. 105°. Prodissoconch ca. 200 pm in height.

Both valves smooth, fine concentric striae on right valve. Auricles unequal with some more striae. Marginal apron

very fragile and mostly broken off. Internal ribs 6 and 2 small auricular ribs. Hinge line straight. Resilifer

triangular. No byssal fasciole, no byssal notch. Ctenolium absent. Colour pale brown with numerous small

reddish maculations on left valve, right valve paler without maculations, visible internal ribs whitish.

Remarks. — The present specimen from the Kai Island is somewhat similar to the type material of P. ina,

although more elongate (umbonal angle ca. 90°) and whitish transparent. P. steindachneri (Sturany, 1901), from

the northeastern Indian Ocean and the Red Sea, differs somewhat by having a more elongate shape and its colour

with larger reddish maculations and small whitish streaks. P. rubrotinctum (Oyama, 1951) from southern Japan to

New Caledonia differs from P . ina by having more numerous internal ribs (commonly 10) and the maculations are

somewhat larger.

Propeamussium invesligaloris (E.A. Smith, 1906)

Figs 11-15

Amussium invesligatoris E.A. Smith, 1906: 255.

Amussium margaritiferum Dautzenberg & Bavay, 1912: 36, pi. 27, figs 15-18. Syn. nov.

Source : MNHN, Paris

PECTIN01DEA FROM EASTERN INDONESIA

249

FIGS 1-4. — Propeamussium alcocki (E.A. Smith, 1894), Karubar, stn CP 91, 30.8 x 27.8 mm (db): 1, left valve,

exterior; 2, left valve, interior; 3, right valve, exterior; 4, right valve, interior.

Figs 5-8. — P. caducum (E.A. Smith, 1885), Karubar, stn CC 56, 17.0 x 13.9 mm (db): 5. left valve, exterior; 6. left

valve, interior; 7, right valve, exterior; 8, right valve, interior.

Source : MNHN, Paris

250

H. H. DIJKSTRA & W. W. KASTORO

MATERIAL EXAMINED. — The type material (see below).

Indonesia. KARUBAR, Kai Islands : stn CP 09. 05°23'S. 132°29'E. 368-389 m, 10 spms. 4 lv. — Stn CC 10.

05°21'S, 132°30'E, 329-389 m. 1 spm. — Stn CP 12, 05°23'S, 132°37'E, 413-436 m, 6 spms. 1 lv. — Stn DW 13,

05°26'S, 132°38’E, 417-425 m. 4 spms, 3 lv, 5 rv. — Stn CP 17, 05°15'S, 133°01'E, 439-459 m, 7 spms. — Stn CP 39,

07°47'S, 132°26'E, 466-477 m, 13 spms.

Tanimbar Islands: stn CP 65, 09°14'S, 132°27’E. 174-176 m, >50 spms. — Stn CP 69, 08°42'S, 131°53'E, 356-

368 m, 46 spms, 3 lv, 2 rv. — Stn CP 77, 08°57'S, 131°27'E, 346-352 m, 23 spms, 1 lv. — Stn CP 78, 09°06'S,

131°24'E, 284-295 m, 4 spms, 1 lv.

TYPE Material. — Amussium investigatoris : lectotype (H 25.0, L 23.0, D 4.5 mm) ZSI M835/1, here

designated, live taken. Three paralectotypes: ZSI M836-838/1 and two paralectotypes: BMNH 1906.10.12.99-100.

Although the largest syntype (H 26.0, L 25.1, D 5.4 mm) is preserved at the BMNH, and closest to Smith' s

measurements, a syntype from the ZSI is selected as lectotype in accordance with Smith (1894: 158). The type

specimens are extremely fragile and the marginal apron often broken off. — A. margaritiferum : holotype ZMA

Moll. 3.12.021.

Type Locality. — A. investigatoris : "Investigator", stn 248, 08°37'N, 75°37'E, W of Travancore, 410-

519 m. — A. margaritiferum: "Siboga", stn 137, 0°23.8'N, 127°29'E, channel between Makjan and Halmahcira,

Moluccas, 472 m.

Distribution. — Northern Indian Ocean and eastern Indonesia. Present material alive in 176-466 m.

Description. — Shell relative small, fragile and semi-transparent, up to ca. 25 mm high, suborbicular,

equilateral, inequivalve, slightly convex, left valve somewhat more so than right, auricles equal, umbonal angle ca.

100-1 10°. Proclissoconch ca. 220 |im. Left valve sculptured with many irregularly spaced radial riblets, which are

squamous near ventral margin. Latticed microsculpture only in early growth stage (ca. 10 mm in height). Anterior

and posterior auricle with ca. 7-10 fine squamous radial riblets, weaker on right valve. Right valve sculptured with

fine regularly spaced concentric lamellae. Internal riblets 10 with 2 auricular riblets, and a few (2-4) intercostal

rudimentary riblets near the periphery. Hinge line straight. Resilifer triangular. No byssal fasciole, or byssal notch,

no ctenolium. Colour of left valve orange-tinted, of right whitish.

Remarks. — The present specimens correspond very well with the type material of P. investigatoris, although

the latticed sculpture is somewhat less developed. P.jeffreysii (E.A. Smith, 1885), from the Philippines, differs

by having a liner and more irregularly spaced radial sculpture, developed into concentric lamellae near the periphery

of the left valve. P . maorium (Dell, 1956), from the SW Pacific, differs by having a smooth surface, lacking the

latticed microsculpture, in early growth stage, and a finer radial sculpture on the left valve.

Propeamussium rubrotinctum (Oyama, 1951)

Figs 16-19

Parvamussium (Parvamussium) rubrotinctum Oyama, 1951: 81, pi. 13, figs 8-10.

Propeamussium rubrotinctum - Dijkstra, 1995b: 21, figs 23-26 [synonymy, reference, description].

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 32, 05°47'S, 132°38'E 307-311 m I lv -

Stn CP 36, 06°05'S, 132°44'E, 210-268 m, 11 spms, 1 lv.

Distribution. — Japan, South China Sea, New Caledonia, and the Loyalty Islands (Dijkstra, 1995b: 21);

new record for Indonesia. Present material alive in 210-268 m.

Remarks. — The present specimens are similar to Oyama's description, but the figured specimen is

somewhat more orbicular and has intercalated rudimentary riblets. Oyama reported other specimens lacking

intercalated riblets, just as in the present material.

Source : MNHN, Paris

PECTINOIDEA FROM EASTERN INDONESIA

251

Figs 9-10. — Propeamussium sp. cf. ina (Dautzenberg & Bavay, 1912). Karubar, stn DW 31, 10.0 x 9.2 mm (rv) :

9, right valve, exterior; 10, right valve, interior.

Figs 11-15. — P. investigatoris (E.A. Smith, 1906), Karubar, stn CP 09, 24.8 x 23.4 mm (db): 11. left valve, exterior;

12, left valve, interior; 13, right valve, exterior; 14, right valve, interior; 15, left valve, exterior, ventral marginal

detail, scale bar 1 mm.

Source :

252

H. H. DIJKSTRA & W. W. KASTORO

Figs 16-19. — Propeamussium rubroiincium (Oyama, 1951), Karubar, stn CP 36, 17.9 x 17.0 mm (db): 16, left valve,

exterior; 17, left valve, interior; 18, right valve, exterior; 19, right valve, interior.

Figs 20-23. — P. siratama (Oyama, 1951), Karubar, stn DW 13, 12.1 x 12.0 mm (lv), 11.3 x 10.9 (rv): 20 left valve

exterior; 21, left valve, interior; 22, right valve, exterior; 23, right valve, interior.

Source MNHN, Paris

PECT1N01DEA FROM EASTERN INDONESIA

253

Propeamussium sibogai (Dautzenberg & Bavay, 1904)

Figs 24-29

Amussium sibogai Dautzenberg & Bavay, 1904: 207, figs 1-4.

Propeamussium sibogai - Dijkstra, 1995b: 23, figs 19-22 [synonymy, references, description).

Material EXAMINED. — The type material (see Dijkstra, 1995b: 23).

Indonesia. Karubar, Kai Islands: stn DW 07, 05°46'S, 132°21'E, 283-285 m, 1 rv (fragment). — Stn CP 36,

06°05'S, 132°44’E, 210-268 m, 5 spms.

DISTRIBUTION. — South Africa, Japan, Philippines, Indonesia, NW Australia, New Caledonia, Loyalty Islands

(Dijkstra, 1995b: 23). Present material alive in 210-268 m.

REMARKS. — The present material is very similar to the holotype from the Bali Sea. P. watsoni (E.A. Smith,

1894) differs by having a fine radiating sculpture in early growth stage and fine, closely spaced concentric lamellae

on the left valve, more numerous (10-14) and not so broad internal ribs. P. alcocki (E.A. Smith, 1894) differs by

having a more oval shape, a more fragile shell, and 11-12 narrower internal ribs. P. andamanicum (E.A. Smith,

1894) differs in its more oval shape, and its 9-10 shorter and narrower internal ribs. P. sewelli (Knudsen, 1967),

from eastern Africa, differs by having a fine radiating and concentric sculpture on the left valve, and 1 1 narrower

internal ribs. All these species are whitish transparent.

Propeamussium siratama (Oyama, 1951)

Figs 20-23

Ctenamusium (Micramussium) siratama Oyama, 1951: 80, pi. 13, figs 5-7.

Propeamussium (Propeamussium) siratama - Dijkstra, 1990: 2, pi. 1, figs 3-4.

Parvamussium siratama - ROMBOUTS, 1991: 70.

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn CP 09, 05°46'S, 132°21'E, 283-285 m, 2 spms.

— Stn DW 13, 05°26'S, 132°38'E, 417-425 m, 2 lv, 5 rv. — Stn DW 28, 05°31'S, 132°54'E, 448-467 m, 1 lv.

Type Material. — Holotype (H 7.4, L 7.5, D 2.8 mm) possible in the Oyama collection at Toba

Aquarium, Japan [not seen).

Type Locality. — Off Manazuruzaki, Sagami Sea, Japan, 234-291 m.

DISTRIBUTION. — Western Pacific from Japan to Indonesia, 234-467 m; present specimens alive in 283-

285 m.

DESCRIPTION. — Shell small, up to ca. 12 mm high, fragile, orbicular, nearly equivalve, equilateral, slightly

convex, auricles equal, umbonal angle 1 10-1 15°. Prodissoconch ca. 200 pm in height. Left valve sculptured with

fine, irregularly spaced radial lirae from early growth stage until beyond central part of disc, with microscopic

regularly spaced concentric lamellae, smooth central part and periphery. Auricles frequently smooth, sometimes

with fine concentric lirae. Right valve sculptured with fine, regularly spaced concentric lirae, more prominent near

ventral margin. Inner surface with 9-10 ribs, sometimes with one rudimentary intercostal one. Hinge line straight.

No byssal fasciole, or byssal notch. Ctenolium absent. Colour whitish or pale-brown transparent.

Remarks. — The present specimens fit the original description, but they reach a larger size, have fewer (9)

internal ribs and lack the very fine concentric lamellae on the left valve. A somewhat similar species is

P. malpelonium (Dali, 1908), from tropical eastern Pacific off Colombia, which differs by a coarser and more

developed microsculpture on the left valve, and more numerous (11) internal ribs. P. malpelonium is only recorded

from abyssal depths, 2690-4505 m (Grau, 1959: 14).

Source :

254

H. H. DIJKSTRA & W. W. KASTORO

FIGS 24-29. — Propeamussium sibogai (Dautzenberg & Bavay, 1904), Karubar, stn CP 36, 49.1 x 52.1 mm (db) :

24, left valve, exterior; 25, left valve, interior; 26, right valve, exterior; 27, right valve, interior; 28, left valve,

detail internal costa, scale bar 1 mm; 29, right valve, detail internal costa, scale bar 1 mm.

Source : MNHN. Paris

PECTINOIDEA FROM EASTERN INDONESIA

255

Genus PARVAMUSSIUM Sacco, 1897

Parvamussium araneum Dijkstra, 1991

Figs 30-38

Parvamussium araneum Dijkstra, 1991: 8, figs 3-10.

Material EXAMINED. — The type material (see below).

Indonesia. Karubar. Kai Islands: stn DW 18, 05°18'S, 133°01'E, 205-212 m, 5 lv, 2 rv. — Stn DW 31, 05°40'S,

1 32°5 1 'E, 288-289 m, 1 rv. — Stn CP 34, 06°09'S, 132°41'E, 435-445 m, 2 lv.

Tanimbar Islands: stn DW 49, 08°00'S, 132°59'E, 206-210 m, 1 lv, 1 rv.

Type Material. — Holotype RMNH 56531, live taken.

Type Locality. — Snellius-II, stn 4.060, 9°51.8'S, 120°46.4'E, NE coast of Sumba, E of Melolo, 240 m.

Distribution. — Indonesia, shells in 155-435 m, alive in 240-300 m.

Remarks. — The present specimens are alike the type material, but on the left valve the commarginal

lamellae are more irregularly spaced.

Parvamussium carbaseum Dijkstra, 1991

Figs 39-43

Parvamussium carbaseum Dijkstra, 1991: 9, figs 11-21.

Parvamussium sp. Kase & Hayami, 1992: 448. — Hayami & Kase, 1993: 3, fig. 5.

Parvamussium crypticum Hayami & Kase, 1993: 54, figs 173-181. Syn. nov.

MATERIAL EXAMINED. — The type material (see below).

Indonesia. Karubar, Kai Islands: stn DW 29, 05°36'S, 132°56'E, 181-184 m, 2 rv.

TYPE Material. — Parvamussium carbaseum: holotype RMNH 56534. — P. crypticum: holotype UMUT

RM 19451a.

TYPE Locality. — Parvamussium carbaseum: SNELLIUS-II, stn 4.031, 5°54'S, 123°58.4'E, Tukang Besi

Islands, Banda Sea, Indonesia, 390 m. — P. crypticum: "Shodokutsu" (- small cave) of Ie Islet, 26°42.9'N,

127°50.1'E, Okinawa, Ryukyu Islands, Japan, alive, 7-20 m.

DISTRIBUTION. — Okinawa, alive in 7-20 m; eastern Indonesia, shells in 155-495 m.

REMARKS. — The present specimens are similar to the type material of P. carbaseum, and differ by their larger

size, up to ca. 5 mm high, and by having a few more rudimentary intercostal riblets. HAYAMI & Kase (1993: 56)

stated that P. crypticum differs from by "the byssal notch decidedly deeper, and the ratio of length/height a little

larger...". However, similar features are observed in the present material and P. crypticum is here synonymized.

P. araneum Dijkstra, 1991, differs from P. carbaseum by having a larger size, a latticed sculpture, and by having

fewer (only a few rudimentary) internal ribs. P. texturatum (Dautzenberg & Bavay, 1912) differs in external

sculpture, which is more prominent with lamellose radial costae and more widely spaced fine commarginal

lamellae.

Parvamussium cassium Dijkstra, 1991

Figs 44-47

Parvamussium cassium Dijkstra, 1991: 11, figs 22-27.

Source :

256

H. H D1JKSTRA & W. W. KASTORO

Figs 30-38. — Parvamussium araneum Dijkstra, 1991, Karubar, stn DW 18: 30, left valve, exterior, scale bar 1 mm;

31, left valve, interior, scale bar 1 mm; 32, right valve, exterior, scale bar 1 mm; 33, right valve, interior, scale bar

1 mm; 34, left valve, prodissoconch, dissoconch, preradial stage, scale bar 10 pm; 35, left valve, exterior, central

detail, scale bar 100 pm; 36, left valve, exterior, central detail, scale bar 100 pm; 37, right valve, exterior, anterior

and posterior auricles, scale bar 1 mm; 38, right valve, interior, dorsal marginal detail, scale bar 1 mm.

Figs 39-43. — P. carbaseum Dijkstra, 1991, Karubar, stn DW 29: 39, right valve, exterior, scale bar 1 mm; 40, right

valve, interior, scale bar 1 mm; 41, right valve, central detail, scale bar 100 pm; 42, right valve, anterior and

posterior auricles, scale bar 1 mm; 43, right valve, interior, dorsal marginal detail, scale bar 1 mm.

Source : MNHN Paris

PECTINOIDEA FROM EASTERN INDONESIA

257

Figs 44-47. — Parvamussium cassium Dijkstra. 1991, Karubar. sin CP 05, 8.0 x 7.8 mm (db): 44. left valve, exierior,

45, left valve, interior; 46. right valve, exterior; 47, right valve, interior.

Figs 48-56. — P. conspecium sp. nov., Karubar, stn DW 15: 48-51, holotype, 5.1 x 4.7 mm (db): 48. left valve,

exterior; 49, left valve, interior; 50, right valve, exterior; 51. right valve, interior. — 52-56. paratype: 52. left

valve, exterior, scale bar 1 mm; 53, left valve, antero-marginal detail, scale bar 100 pm; 54. left valve, interior,

scale bar 1 mm; 55, left valve, exterior, preradial stage, scale bar 100 pm; 56, left valve, exterior, dissoconch.

preradial stage, scale bar 100 pm.

Source :

258

H. H. D1JKSTRA & W. W. KASTORO

MATERIAL EXAMINED. — The lype material (see below).

Indonesia. Karubar. Kai Islands: stn CP 05. 05°49'S. 132°18'E. 296-299 m, 1 spm.

Type Material. — Holotype RMNH 56549, live taken.

TYPE Locality. — Snellius-II. stn 4.142, 6°29.7'S, 121°10.8'E, NE Taka Bone Rate (Tiger Island),

E of Tarupa Kecil, 450-600 m.

DISTRIBUTION. — Flores Sea and Banda Sea, Indonesia; alive in 299-450 m.

Remarks. — The present specimen is similar to the type material, although more orbicular in shape, with

fewer secondary radial riblets on the left valve and the small scales on the intersections are slightly more

prominent. Other conchological features are identical.

Parvamussium conspectum sp. nov.

Figs 48-61

Parvamussium sp. cf. texturatum - DlJKSTRA, 1991: 17, figs 44-52.

Material EXAMINED. — Indonesia. Karubar. Kai Islands: stn DW 02, 05°47'S, 132°13’E, 209-240 m. 1 lv

(paratype POLIPI). 2 rv (paratypes HD. POLIPI). — Stn DW 15, 05°17'S. 132°41'E, 212-221 m, 1 spm. (holotype), 2 lv,

2 rv (paratypes MNHN) — Stn DW 18, 05°18'S, 133°01'E, 205-212 m, 1 lv (paratype HD). — Sin DW 24. 05°32'S,

132°5 1 'E. 230-243 m, 2 lv (paratypes MNHN).

Type Material. — Holotype. live taken, MNHN. Paratypes: 2 HD, 2 POLIPI, 5 MNHN.

Type Locality. — Kai Islands, E Indonesia, Karubar, stn DW 15, 050°17'S, 132°41'E, 212-221 m.

Distribution. — Eastern Indonesia; shells in 100-290 m, alive in 212-250 m.

Description. — Shell small, up to ca. 5 mm high, fragile, semi-transparent, valves equally convex, elongate,

inequivalve, slightly inequilateral, auricles unequal in size, umbonal angle ca. 90°. Prodissoconch ca. 190 pm in

height. Left valve sculptured with ca. 20 primary and secondary irregularly spaced radial riblets, ca. 10 closely set

commarginal lamellae in early growth stage (before pre-radial stage), widely spaced on central part of disc and more

closely so towards ventral margin. Intersections of radial riblets and commarginal lamellae strongly squamous.

Anterior auricle larger than posterior and provided with closely spaced, strongly irregularly developed concentric

lamellae, more prominent anteriorly. A row of small lamellae produced near disc Hank. Right valve sculptured

with ca. 20 widely spaced concentric lamellose lirae. Sculpture of auricles similar to that of left valve. A radial lira

on anterior auricle near byssal fasciole. Internal ribs 9, commencing 1 mm below resilifer and developed towards

0.5 mm above periphery, with a small auricular riblet on each side. Hinge line straight. Resilifer triangular.

Byssal fasciole and notch small. Ctenolium absent. Left valve creamy-orange, right valve transparent white. Inner

side of both valves glossy. Dimensions (holotype): H 5.1, L 4.7, D 1.1 mm.

Remarks.— Parvamussium sp. cf. texturatum of DlJKSTRA (1991: 17, figs 44-52) differs from the present

material only in the internal ribs being shorter and slightly more numerous (10 instead of 9). P. texturatum

(Dautzenberg & Bavay, 1912) differs by having a more orbicular shape, a weaker sculpture of commarginal

lamellae on the left valve, and in having more numerous internal ribs (15). P. vesiculatum Dijkstra, 1995, differs

by having a more nodose sculpture at intersections on the left valve, more numerous closely spaced concentric

lamellae on the right valve, and less developed internal riblets (only a few rudimentary near the anterior and

posterior margins). P. undisonum Dijkstra, 1995 differs in size (larger, up to ca. 14 mm), is more oblique, and

more weakly sculptured with more close-set commarginal lamellae on the left valve.

Etymology. — Lat. conspectus, adj. = conspicuous.

Source : MNHN Paris

PECTINOIDEA FROM EASTERN INDONESIA

259

Figs 57-61. — Parvamussium conspectum sp. nov., Karubar, sin DW 15, paratype (MNHN): 57, right valve, exierior,

scale bar 1 mm; 58, right valve, exterior, central detail, scale bar 100 pm; 59, right valve, interior, scale bar 1 mm;

60, right valve, exterior, prodissoconch. dissoconch, preradial stage, scale bar 10 pm; 61. right valve, exterior,

dissoconch, preradial stage, scale bar 10 pm.

Figs 62-67. — P. cristatellum Dautzenberg & Bavay, 1912, Karubar. stn DW 13, 8.4 x 8.9 mm (lv, typical), 8.3 x

8.1 mm (lv, atypical), 7.8 x 7.8 mm (rv); 62. left valve, exterior; 63. left valve, interior; 64. right valve, exterior;

65, right valve, interior; 66. left valve (atypical), exterior; 67, left valve (atypical), interior.

Source :

260

H. H. DIJKSTRA & W. W. KASTORO

Figs 68-72, — Parvamussium cristatellum Dautzenberg & Bavay, 1912. Karubar, stn DW 13: 68, left valve, exterior,

central detail, scale bar 1 mm; 69, left valve, exterior, antero-ventral detail, scale bar 100 urn; 70. left valve,

exterior, prodissoconch, dissoconch, preradial stage, scale bar 10 pm; 71. right valve, exterior, central detail, scale

bar 100 pm; 72, right valve, interior, antero-dorsal detail, scale bar 100 pm.

Figs 73-74. — P. pauciliratum (E.A. Smith, 1903), Karubar, stn DW 28, 7.0 x 7.5 mm (rv): 73. right valve, exterior;

74, right valve, interior.

Figs 75-78. — P. scitulum (E.A. Smith, 1885), Karubar: stn DW 22, 9.1 x 9.8 mm (lv), 7.8 x 8.0 mm (rv): 75. left

valve, exterior; 76, left valve, interior; 77, right valve, exterior; 78, right valve, interior.

Source : MNHN, Paris

PECTIN'OIDEA FROM EASTERN INDONESIA

261

Parvamussium cristatellum (Dautzenberg & Bavay, 1912)

Figs 62-72

Pecten (Amussium) cristatum (sic) Bavay. 1905b: 187. pi. 17. figs 2a-c (non Pecien cristatus Bronn, 1828).

Amussium cristatellum Dautzenberg & Bavay, 1912: 36. pi. 28, figs 5-8 [nom. nov. for Pecten (Amussium) cristatus

Bavay].

Parvamussium cristatellum - DlJKSTRA, 1991: 13. figs 28-32 [synonymy, references, description, discussion].

Material EXAMINED. — The type material (see below).

Indonesia. Karubar. Kai Islands : stn DW 13, 05°26'S, 132°38'E, 417-425 m, 20 lv, 20 rv, — Stn DW 14, 05°18'S,

132°38'E, 245-246 m, 1 lv. — Stn DW 18, 05°18'S, 133°01'E, 205-212 m, 3 lv, 7 rv. — Stn DW 28, 05°33'S. 132°51'E,

304-314 m, 4 lv, 5 rv. — Stn DW 29, 05°36'S, 132°56'E, 181-184 m, 4 lv. — Stn DW 32, 05°47'S, 132°51'E. 170-

206 m. 3 lv, 1 rv. — Stn CP 34, 06°09'S, 132°41’E, 435-445 m. 4 lv.

Tanimbar Islands: stn CP 65, 09°14'S, 132°27'E. 174-176 m, 4 lv, 5 rv. — Stn CP 69, 08°42'S. 131°53'E, 356-

368 m, 8 lv, 8 rv.

Type Material. — Lectotype (lv illustrated by Bavay, 1905b. pi. 17, fig. 2a; H 7.1, L 7.2 mm) ZSI

M3360/1, here designated. Two paralectotypes (Bavay, 1905b, pi. 17, figs 2b-c) ZSI M3360/2-3.

Type Locality. — "Masandam insulam" [= Andaman Islands, India], depth unknown.

Distribution. — Northeastern Indian Ocean and Indonesia.

Remarks. — The present specimens are similar to the type material. The internal ribbing and sculpture

are variable, from weak to more prominent, sometimes lacking sculpture or commarginal lamellae on the left

valve. P. thetidis somewhat differs by having a weaker sculpture and more close-set commarginal lamellae on

the left valve. P. siebenrocki (Sturany, 1901) from the northwestern Indian Ocean is nearly identical to

P. cristatellum , and differs only in the radial costae on the left valve being somewhat weaker; however,

intermediates seem to exist and the two may be synonyms. P. formosum (Melvill, 1907) from the western Indian

Ocean is quite smooth and the auricles are very finely sculptured and more similar to those in P. torresi

(E.A. Smith, 1885).

Parvamussium pauciliratum (E.A. Smith, 1903)

Figs 73-74

Amussium paucilirata (sic) E.A. Smith, 1903: 622, pi. 36, figs 23-24.

Parvamussium pauciliratum - DlJKSTRA, 1995b: 26. figs 107-110, 151-152 [references, description],

MATERIAL EXAMINED. — The type material (see Dijkstra, 1995b: 28).

Indonesia. Karubar, Kai Islands: stn DW 28, 05°31'S, 132°54'E, 448-467 m, 1 rv.

Distribution. — Northern Indian Ocean, Indonesia to New Caledonia; 27-448 m, alive in 27-45 m.

REMARKS. — The present specimen is very similar to the type material, although the internal ribs are

somewhat more prominent. However, development of internal ribs is variable (very weak and small to strongly

developed). Juveniles often lack the internal ribs.

Parvamussium scitulum (E.A. Smith, 1885)

Figs 75-78

Amussium scitulum E.A. Smith, 1885: 312, pi. 23, figs 4-4b.

Parvamussium scitulum - DlJKSTRA, 1995b: 31, figs 43-46. 153-154 [synonymy, references, description, discussion].

Source :

262

H. H. D1JKSTRA & W. W. KASTORO

Material examined. — The type material (see Dijkstra. 1995b: 31).

Indonesia. KARUBAR, Kai Islands: stn CP 05, 05°49'S, 132°18'E, 296-299 m, 1 lv. — Stn DW 22, 05°22'S,

1 33°01'E. 124-850 m, 7 lv, 16 rv. — Stn DW 30, 05°39'S, 132°56’E, 111-118 m, 3 lv, 9 rv. — Stn DW 31. 05°40'S,

132°51'E, 288-289 m, 1 lv.

DISTRIBUTION. — Western and southwestern Pacific from Japan to New Caledonia; 50-300 m, with shells

occasionally carried deeper, present material (shells only) in 1 18-296 m.

Remarks. — The present specimens are similar to the type material, and reach a larger size, up to 10 mm. In

P. dautzenbergi Dijkstra, 1990, also from Indonesia, the left valve is more prominently reticulated.

Parvamussium squalidulum Dijkstra, 1995

Figs 79-82

Parvamussium squalidulum Dijkstra, 1995b: 32, figs 47-50.

Material examined. — The type material (see Dijkstra, 1995b: 32).

Indonesia. KARUBAR, Kai Islands : stn DW 02, 05°47'S, 132°13’E, 209-240 m, 3 lv. 6 rv. — Stn DW 22, 05°22'S,

133°01'E. 124-850 m, 1 lv.

Distribution. — Coral Sea to New Hebrides Arc, alive in 260-400 m; new record for Indonesia.

Remarks. — The material from stn DW 02 differs slightly in sculpture. Concentric lamellae are somewhat

more widely spaced on both valves and scales on riblets are weaker.

Parvamussium thetidis (Hedley, 1902)

Figs 83-89

Amusium thetidis Hedley, 1902: 304, fig. 49.

Parvamussium thetidis - Dijkstra, 1995b: 35, figs 99-102 (synonymy, references, description, discussion).

MATERIAL EXAMINED. — The type material (see Dijkstra, 1995b: 35).

Indonesia. Karubar, Kai Islands: stn DW 24, 05°32'S. 132°51'E, 230-243 m, 3 lv, 2 rv. — Stn DW 28, 05°3 1 S

132°54’E, 448-467 m, 1 lv. — Stn DW 31. 05°40'S, 132°51'E, 288-289 m, 1 lv, 8 rv.

Tanimbar Islands: stn CP 77. 08°57’S, 131°27'E, 346-352 m, 1 spm.

Distribution. — Eastern Australia, Coral Sea to New Hebrides Arc; new record for Indonesia. Present

material alive in 346-352 m, shells in 243-448 m.

Remarks. — The left valve of the specimen from stn CP 77 is slightly more prominently sculptured, with

more numerous radial riblets and stronger commarginal lamellae.

Parvamussium torresi (E.A. Smith, 1885)

Figs 90-94

Amussium torresi E.A. Smith, 1885: 311, pi. 23, figs 3-3b.

Parvamussium torresi - Dijkstra. 1995b: 36, figs 51-54, 125-128 [references, description).

Material EXAMINED. — The type material (see Dijkstra. 1995b: 36).

Indonesia. Karubar, Kai Islands: stn DW 01. 05°46'S, 1 32°1 0'E, 156-305 m, 2 lv, 1 rv. — Stn DW 02 05°47’S

1 32° 1 3'E, 209-240 m, 13 lv. 15 rv. — Stn DW 03. 05°48'S, 132°13'E, 278-301 m. 6 lv, 8 rv. — Stn DW 01, 05°46's'

132°21'E, 283-285 m, 2 lv, 7 rv. — Sin DW 15. 05°17'S, 132°41'E, 212-221 m. 5 lv, 1 rv. — Stn DW 18, 05°18's’

133°01'E, 205-212 m, 6 spms, 2 lv, 16 rv, — Stn DW 29, 05°36'S, 132°56'E, 181-184 m. 4 lv, 12 rv. — Stn DW 31

05°40’S, 132°51'E, 288-289 m, 10 lv, 10 rv. — Stn DW 32, 05°47'S, 1 32°51'E, 170-206 m. 2 spms.

Source : MNHN, Paris

PECTINOIDEA FROM EASTERN INDONESIA

263

Figs 79-82. — Parvamussium squalidulum Dijkstra, 1995, Karubar, stn DW 02. 8.2 x 8.9 mm (lv), 7.2 x 7.8 mm (rv):

79, left valve, exterior; 80, left valve, interior; 81, right valve, exterior; 82, right valve, interior.

Figs 83-89. — P. thetidis (Hedley, 1902), Karubar. stn DW 31; 83, left valve, exterior, scale bar 1 mm; 84. left valve,

interior, scale bar 1 mm; 85. right valve, exterior, scale bar 1 mm; 86. right valve, interior, scale bar 1 mm; 87, left

valve, exterior, prodissoconch, dissoconch, preradial stage, scale bar 10 pm; 88. left valve, exterior, central detail,

scale bar 100 pm; 89, right valve, exterior, central detail, scale bar 100 pm.

Source :

264

H. H. DIJKSTRA & W. W. KASTORO

FlGSJ0;9!' Parvamussium torresi (E.A. Smith, 1885). Karubar, sin DW 02, 6.9 x 7.1 mm (lv), 6.3 x 6.3 mm (rvV

9°, left valve, exterior; 91, left valve, interior; 92, right valve, exterior; 93, right valve, interior; 94 left valve

exterior, prodissoconch, scale bar 100 pm.

FlGS0,95:9f8- — P yesiculalum Dijkstra. 1995; 95-96. Karubar, stn DW 32, 7.9 x 8.1 mm (lv); 95, left valve, exterior

Z enft valv„e' 1"t®nor; ~ 97‘98’ Karubar, stn DW 18: 97. left valve, exterior, dissoconch, preradial stage, scale

bar 10 pm; 98. left valve, exterior, antero-ventral detail, scale bar 100 pm.

FlGiftn9'|1°f!)' ~,P' virga,um Dijkstra- I991- Karubar, stn DW 32, 10.0 x 10.2 mm (lv): 99. left valve, exterior;

iuu. lert valve, interior.

Source MNHN . Paris

PECTINOIDEA FROM EASTERN INDONESIA

265

Distribution. — Southern Philippines, Indonesia, Coral Sea to the Loyalty Islands; 205-600 m, alive in

205-355 m.

Remarks. — The present specimens are similar to the type material; a few specimens (stn DW 01 and

DW 32) have colour maculations. P. formosum from the Arabian Sea is closely related but differs in having

a more prominent sculpture of radial striae on auricles and near the anterior and posterior margins of the left valve.

P. scitulum (E.A. Smith, 1885) differs from P. torresi by having a more compressed shell, more prominent

sculpture of radial striae, and by being more brightly coloured. The right valve of P. torresi is covered with

regularly spaced concentric lirae, which are generally lacking on P. scitulum.

Parvamussium vesiculatum Dijkstra, 1995

Figs 95-98

Parvamussium vesiculatum Dijkstra, 1995b: 37, figs 59-62, 93-96.

MATERIAL EXAMINED. — The type material (see Dijkstra. 1995b: 37).

Indonesia. Karubar, Kai Islands: stn DW 18, 05°18'S, 133°01’E. 205-212 m. 3 spnis. — Stn DW 29. 05°36'S,

132°56'E, 181-184 m, I Iv, I rv. — Stn DW 32, 05°47'S, 132°51'E, 170-206 m, 1 lv.

DISTRIBUTION. — New Caledonia and Loyalty Islands; new record for Indonesia. Present material alive in 205-

212 m.

Parvamussium virgatum Dijkstra, 1991

Figs 99-100

Parvamussium virgatum Dijkstra, 1991: 20. figs 62-65.

MATERIAL EXAMINED. — The type material (see below).

Indonesia. KARUBAR, Kai Islands: stn DW 32. 05°47'S, 1 32°5 I E, 170-206 m, I Iv,

Type Material. — Holotype RMNH 56556.

Type Locality. — Banda Sea, Tukang Besi Islands, NW of Binongko, SNELLIUS-II, stn 4.033. 05°52.5'S,

123°58.5’E, 250-290 m.

Distribution. — Eastern Indonesia; shells in 206-305 m.

Remarks. — The present specimen is similar to the type material. P. scitulum (E.A. Smith, 1885) differs by

having a more compressed shell, and a stronger sculpture with radial and concentric lirae on the left valve; the

auricles are also more prominent sculptured.

Family PECTINIDAE Wilkes, 1810

Subfamily CAMPTONECTINAE Habe, 1977

Genus DELECTOPECTEN Stewart, 1930

Delectopecten alcocki (E.A. Smith, 1904)

Figs 101-108

Pecten alcocki E.A. Smith, 1904: 13.

Delectopecten alcocki - Dijkstra, 1995b: 50, figs 111-114, 147-150 [references, description].

Source

266

H. H. DIJKSTRA & W. W. KASTORO

Material EXAMINED. — The type material (see Dijkstra, 1995b: 50).

Indonesia. Karubar. Kai Islands: stn CP 05. 05°49'S, 132°I8'E, 296-299 m, 1 spm. — Stn DW 08. 05°20'S.

I32°31’E. 358-360 m, 10 Iv, 5 rv. — Stn CP 09. 05°23'S, 132°29'E, 68-389 m, 18 spms. 2 Iv, 10 rv. — Stn CC 10.

05°21’S, 132°30'E, 329-389 m, 25 spms, 9 lv. 6 rv. — Stn DW 13, 05°26'S, 132°38'E, 417-425 m. 1 spm, 25 lv, 23 rv.

— Stn CP 16. 05°17’S, 132°50'E, 315-349 m, 3 spms. — Stn CP 17. 05°15’S. 133°01'E. 439-459 m, 1 spm. —

Stn DW 18, 05°18'S, 133°01'E. 205-212 m. 2 rv. — Stn CP 20. 05°15'S, 132°59'E. 769-809 m. 1 rv. — Stn CC 21.

05°14'S, 133°00'E, 688-694 m, 4 spms, 1 lv. 1 rv. — Stn DW 28, 05°31'S. 132°54'E, 448-467 m, 1 Iv, 1 rv. —

Stn DW 31, 05°40'S, 132°51'E, 288-289 m, 1 spm. — Sin CP 35, 06°08'S, 132°45’E, 390-502 m, 2 spms, 1 rv, —

Stn CP 38, 07°40'S. 132°27’E, 620-666 m, 4 spms.

Tanimbar Islands: stn DW 44, 07°52'S, 132°48’E, 291-295 m, 1 lv. 3 rv. — Stn CC 56. 08°16'S. 131°59’E, 549-

552 m. 1 spm. — Stn CC 58, 08°19'S, 132°02'E, 457-461 m. 2 spms, 1 lv. — Stn CP 59, 08°20'S, 132°H'E, 399-

405 m, 3 spms, 4 lv, 5 rv, — Stn DW 60, 08°21'S, 132°14'E, 387-389 m. 5 lv, 2 rv. — Sin CP 63, 08°00’S, 132°58'E,

214-215 m. 1 rv. — Stn CP 65. 09°14'S, 132°27'E, 174-176 m. 8 spms, 22 Iv, 23 rv. — Stn CP 69, 08°42'S, I31°53'E,

399-405 m, many spms. — Stn CP 71, 08°38'S, 131°44'E. 477-480 m, 1 lv.

Distribution. — East Africa, Gulf of Aden, Bay of Bengal, Philippines, Indonesia, Coral Sea (Dijkstra,

1995b: 50). Present material alive in 176-688 m.

REMARKS. — The present specimens are similar to the type material, although somewhat more variable in

sculpture (diverging radial striae very weak to more prominent and radially aligned scales absent to strongly

developed). In the type material the diverging radial striae are almost lacking and the scales are weak.

Delectopecten fluctualus (Bavay, 1905)

Figs 109-113

Pecten (Chlamys) fluctualus Bavay, 1905b: 188, pi. 17, figs 3a-b.

Delectopecten fluctuatus - Dijkstra, 1995b: 51, figs 83-86 [references, description].

Material examined. — The type material (see Dijkstra, 1995b: 51).

Indonesia. Karubar, Kai Islands: stn DW 13. 05°26'S. 132°38'E, 417-425 m, 1 iv. — Stn DW 31, 05°40'S.

132°51'E, 288-289 m, 3 spms.

Tanimbar Islands: stn CP 86, 09°26'S. 13I°I3'E, 223-225 m, 2 spms.

Distribution. — Andaman Sea, Indonesia. Loyalty Islands (Dijkstra, 1995b: 51). Present specimens alive

in 225-288 m.

Remarks. — Present specimens differ from the holotype in being semi-transparent white instead of opaque

cream, and by having more numerous radial riblets, up to ca. 40-70, near the ventral margin.

Subfamily CHLAMYDINAE Teppner, 1922

Tribe CHLAMYDINI Tcppner, 1922

Genus LAEVICHLAMYS Waller, 1993

Laevichlamys aliae (Dijkstra, 1988) (comb, nov.)

Figs 132-133

Chlamys aliae Dijkstra, 1988: 17-18, unumbered figs.

Chlamys aliae - DIJKSTRA, 1990: 7, 9. — ROMBOUTS, 1991: 90.

MATERIAL EXAMINED. — The type material (see below).

Indonesia. Karubar, Kai Islands: stn DW 29. 05°36’S, 132°56'E, 181-184 m, 1 Iv. — Stn DW 30 05°39'S

132°56'E, 111-118 m. 7 lv, 3rv.

Source : MNHN, Paris

PECTINOIDEA FROM EASTERN INDONESIA

267

Figs 101-108. — Delectopecten alcocki (E.A. Smith, 1904), Karubar, stn CP09: 101, left valve, exterior, scale bar

1 mm; 102, left valve, exterior, anterior auricular detail, scale bar 10 pm; 103, left valve, exterior, antero-ventral

detail, scale bar 100 pm; 104. left valve, exterior, prodissoconch, scale bar 10 pm; 105, right valve, exterior, scale

bar 1 mm; 106, right valve, exterior, anterior auricle, scale bar 100 pm; 107, right valve, exterior, antero-ventral

detail, scale bar 100 pm; 108, right valve, exterior, vesicular detail, scale bar 10 pm.

Figs 109-1 10. — D. fluctuatus (Bavay, 1905), Karubar, stn DW 31, 10.0 x 9.1 mm (db); 109. left valve, exterior; 110,

left valve, interior.

Source :

268

H. H. DIJKSTRA & W. W. KASTORO

Type Material. — Holotype ZMA Moll. 3. 88.046. live taken.

Type Locality. — Off Punta Engano, Mactan, Cebu. Philippines, 1 10 in.

Distribution. — Philippines and Indonesia; living in 30-200 m (Dijkstra, unpubl. data).

Remarks. — The present specimens are similar to the type material. The shagreen microsculpture and

strongly developed radial costae suggest a placement in Laevichlamys rather than Chlamys Roding, 1798.

Laevichlamys deliciosa (Iredale, 1939) (comb, nov.)

Fig. 134

Mimachlamys deliciosa Iredale. 1939: 350-351, pi. 5, figs 22-22a.

Clilamys deliciosa - Dijkstra, 1991: 30 [synonymy, references].

Material EXAMINED. — The type material (see below).

Indonesia. Karubar. Kai Islands: stn DW 02, 05°47’S, 132°13'E, 209-240 m, 1 lv. — Stn DW 18. 05°18'S.

133°01'E, 205-212 m. 1 spm. — Stn DW 22, 05°22'S, 133°01'E, 124-850 m, 3 lv.

Tanimbar Islands: stn DW 50. 07°59'S, 133°02'E. 184-212 m, 1 spm.

Type Material. — Holotype AMS C89669.

Type Locality. — Low Isles, SE of Lizard Island, N Queensland, Great Barrier Reef Exped. stn 14,

14°41'S, 145°29'E, 35 m.

Distribution. — Western and southwestern Pacific, alive in 80-205 m.

DESCRIPTION. — Shell small, commonly 15 mm high, occasionally to ca. 25 mm, equivalve, somewhat

equilateral, valves convex, auricles very unequal, umbonal angle ca. 80-85°. Prodissoconch height ca. 240 Jim.

Both valves sculptured with numerous fine radial riblets commencing in early growth stage (ca. 1 mm) and

increasing to ca. 45-50 by intercalating riblets towards ventral margin. Radial riblets bear small erect prickly

scales. Interspaces microscopically granulated or reticulated, and smooth near ventral margin. Anterior auricles have

9-14 fine radial riblets; posterior auricles have fewer (6-10) and weaker riblets. Postero-dorsal margin of hinge line

somewhat declined. Byssal gap small. Active ctenolium present with 4-6 teeth. Colour uniformly cream, pink,

orange, red or purple, sometimes also stained.

Remarks. — Present specimens are almost identical to the type material. Specimens from stations DW 18 and

DW 50 reach 22 mm and 24 mm in height.

Genus SEMIPALLIUM Jousseaume, 1928

Semipallium Jousseaume in Lamy, 1928: 169. Type species (OD): Pecien tigris Lamarck, 1819. Recent, Indo-West

Pacific.

DIAGNOSIS. — A byssate Chlamydini with shagreen (reticulated) microsculpture, sculptured with regularly

spaced primary radial costae, fine secondary radial riblets commonly present at least in late growth stage; auricles

very unequal; inner surface undulated; byssal notch moderately deep, ctenolium well developed.

Distribution. — Miocene-Recent. Tropical Indo-West Pacific; littoral to sublittoral depths.

Remarks. Hertlein (1969: N365) considered Semipallium as a valid Indo- Pacific genus, placed in the

Decaiopecten group. Waller (1993: 202) treated it as a genus of Chlamydini.

Source : MNHN. Paris

Figs 111-113. — Deleciopecien fluctuatus (Bavay, 1905), Karubar, stn DW 31: 111. left valve, exierior. antero- ventral

detail, scale bar 100 pm; 112, left valve, exterior, prodissoconch, preradial stage, scale bar 100 pm; 113, right

valve, exterior, anterior auricle, ctenolium, scale bar 1 mm.

Figs 114-117. — Veprichlamys versipellis sp. nov., holotype, 23.3 x 19.5 mm (db): 114, left valve, exterior;

115, left valve, interior; 116, right valve, exterior; 117, right valve, interior.

PECTINOIDEA FROM EASTERN INDONESIA

269

270

H. H. DIJKSTRA & W. W. KASTORO

Semipallium dianae (Crandall, 1979)

Figs 135-137

Chlamys dianae Crandall. 1979: 114, figs 3-8.

Chlamys dianae - Matsukuma, Okutani & Habe, 1991: 137, 185, pi. 135, fig. 9. — LAN, 1993: 161, 219, fig.

Semipallium dianae - DiJKSTRA, 1991: 38. — ROMBOUTS, 1991: 59. pi. 5, figs 3-3a-b.

MATERIAL EXAMINED. — The type material (see below).

Indonesia. Karubar, Kai Islands : stn DW 22. 05°22'S. 133°01'E. 124-850 m, 2 Iv, 1 rv.

Type Material. — Holotype in the Taiwan Museum. Taipei, Taiwan 7911.

TYPE Locality. — Ryukyu Islands, S Japan, alive, ca. 30 m.

Distribution. — Western and southwestern Pacific, from southern Japan, the Philippines, Indonesia and the

Solomon Islands; alive in 20-55 m (DiJKSTRA, unpubl. data).

Description. — Shell commonly ca. 35 mm high, occasionally up to ca. 50 mm, elongated, slightly convex,

equivalve, subequilateral, auricles very unequal, umbonal angle ca. 80-90°. Prodissoconch height ca. 280 pm. Both

valves covered with shagreen (reticulated) microsculpture and sculptured with 8-10 (commonly 9) regularly spaced,

rounded radial costae. Fine radial riblets developed near ventral margin, most prominent on right valve. Anterior

auricle with 5-6, posterior auricle with 2-4 radial ribs. Hinge line straight on anterior, somewhat declined on

posterior auricle. Inner surface plicated, sometimes striated near ventral margin. Resilifer triangularly oblong,

elongated. Byssal fasciole broad, byssal notch relative deep. Ctenolium well developed with 5-7 teeth. Colour very

variable, occuring in brown, orange, red, purple and yellow, commonly creamy-yellowish with brown and milky

white dots and streaks.

Remarks. — The present valves from the Kai Islands are similar to the type material, although more

monochrome yellow and orange.

Genus VEPR1CHLAMYS Iredale, 1929

Veprichlamys versipellis sp. nov.

Figs 114-131

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 13, 05°26'S, 132°38'E, 417-425 m 5 v

(MNHN). — Stn CP 33, 06°05'S, 132°38'E, 307-311 m. 1 v (MNHN).

Tanimbar Islands: stn DW 44, 07°52'S, 132°48 E. 291-295 m, 5 v (3 MNHN, 2 L1PI). — Stn CP 46 08°01'S

1 32°5 1 ’E, 271-273 m. 1 fresh db (holotype). — Stn DW 61, 09°05'S, 132°44'S, 235-236 m. 14 v (10 MNHN. 4 LIPI). —

Stn CP 79, 09°16'S, 131°22'E, 239-250 m, 3 v (1 MNHN, 2 HD). — Stn CP 86. 09°26'S. 131°13'E, 223-225 m 2 v

(MNHN).

Type Material. — Holotype MNHN. Paratypes: 22 MNHN. 6 LIPI, 2 HD.

Type Locality. — Arafura Sea, E of Tanimbar Islands, Karubar, stn CP 46 08°01’S 132°51'E 271-

273 m.

Distribution. — Banda and Arafura Sea, shells only in 225-417 m.

Description. — Shell fragile, up to 35 mm high, somewhat obliquely ovate, compressed, equilateral,

equivalve, auricles tnequivalve, umbonal angle ca. 90°. Prodissoconch height ca. 280 |im. Both valves sculptured

with numerous (ca. 40) irregularly spaced, primary radial costae, commencing at 1 mm shell height, and ca. 12,

Source : MNHN Paris

PECTINOIDEA FROM EASTERN INDONESIA

271

Figs 118-127. — Veprichlamys versipellis sp. nov.: 118-119, holotype: 118, left valve, interior, dorsal detail, scale

bar 1 mm; 119, right valve, interior, dorsal detail, scale bar 1 mm. — 120-121, Karubar, stn DW 44, paratypes

(Iv, rv): 120, left valve, exterior, central detail, scale bar 1 mm; 121, right valve, exterior, ventral marginal detail,

scale bar 1 mm. — 122-127, Karubar, stn CP 86, paratype (db): 122, left valve, exterior, scale bar 1 mm;

123, left valve, exterior, prodissoconch, radial stage, scale bar 100 pm; 124, left valve, exterior, anterior auricle,

scale bar 100 pm; 125, right valve, exterior, scale bar 1 mm; 126, left valve, exterior, ventral marginal detail, scale

bar 100 pm; 127, right valve, exterior, anterior auricle, ctenolium, scale bar 100 pm.

Source :

272

H. H. DIJKSTRA & W. W. KASTORO

Figs 128-131. — Veprichlamys versipellis sp. nov., Karubar, stn CP 86, paratype (db): 128, right valve, exterior,

prodissoconch, preradial stage, scale bar 100 pm; 129, left valve, exterior, central detail, scale bar 100 pm;

130, left valve, exterior, ventral marginal detail, scale bar 100 pm; 131, right valve, exterior, postero-marginal

detail, scale bar 100 pm.

Figs 132-133. — Laevichlamys aliae (Dijkstra, 1988), Karubar, stn DW 30, 23.8 x 20.6 mm (lv): 132. left valve,

exterior. — 133, left valve, interior.

Fig 134. — L. deUciosa (Iredale, 1939), Karubar. stn DW 22, 9.0 x 7.6 mm, left valve, exterior.

Figs 135-137. — Semipallium dianae (Crandall, 1979), Karubar, stn DW 22, 25.1 x 21.9 mm (lv): 135, left valve,

exterior; 136, left valve, interior; 137, left valve, exterior, central detail, scale bar 1 mm.

Source : MNHN, Paris

PECTINOIDEA FROM EASTERN INDONESIA

273

secondary radial riblets commencing at central part of disc, and increasing towards ventral margin. Costae

squamose. Microsculpture of interspaces variable: early growth stage with diverging striae near anterior and

posterior margin, shagreen microsculture above central part of disc and near ventral margin, in between radial striae

and diverging to the anterior and posterior margins. Anterior auricle of left valve larger than posterior, sculptured

with 10 weakly developed squamous radial lirae; posterior auricle with 6 fine squamous radial lirae, interspaces of

early growth stage striae microscopically reticulated. Auricles of right valve sculptured with more prominent radial

costae, fewer in number (anterior 6, posterior 5). Anterior hinge line straight, posterior somewhat suppressed.

Byssal fasciole broad, byssal notch rather deep. Inactive and active ctenolium beside ledge of suture, with 4 teeth.

Postero-lateral margins of disc scarcely gaping. Resilifer elongate triangular. Inner surface of both valves plicate

near periphery. Colour creamy with pink-reddish dots and scales. Dimensions (holotype): H 23.4, L 20.0,

D 6. 1 mm.

REMARKS. — Juvenile specimens resemble V. jousseaumei (Bavay, 1904), from the western Pacific, which

differs by the absence of intercalating ribs and of the shagreen microsculpture. Adult specimens of V. versipellis

differ by having shagreen microsculpture on the anterior auricles, on the central part of the disc and near the ventral

margin, whereas V. jousseaumei has radial striae. V. versipellis has many (ca. 50), irregularly spaced, radial

riblets, V. jousseaumei fewer (ca. 20), regularly spaced, radial ribs. V. perillustris (Iredale, 1925) from southern

and southeastern Australia differs having by a more oblique shape, fewer (ca. 20) radial ribs, more prominent scales

on the ribs, and lack of a shagreen (reticulated) microsculpture. V. kiwaensis (Powell, 1933) resembles

V. perillustris and differs from V. versipellis by having fewer (ca. 20) regularly spaced radial ribs and more

prominent microscopic striae; shagreen microsculpture is absent. V. incantata (Hertlein, 1972) from the Galapagos

Islands differs by attaining a larger size (ca. 45 mm high), having a more elongate shape, a larger convexity of the

valves and lacking the shagreen microsculpture.

Etymology. — From the Latin versipellis , adj. = metamorphosis, with regard to the inconstancy of shell

microsculpture.

Tribe AEQUIPECTININI Nordsieck, 1969

Genus CRYPTOPECTEN Dali, Bartsch & Rehder, 1938

Cryptopecten Dali, Bartsch & Rehder, 1938: 93. Type species (OD): Cryptopecten alii Dali, Bartsch & Rehder, 1938.

Recent, Hawaii Islands, 95-435 m.

Synonymy and Diagnosis: see DlJKSTRA (1995b: 60).

Cryptopecten bullatus (Dautzenberg & Bavay, 1912)

Figs 138-145

Pecten ( Chlamys ) bullatus Dautzenberg & Bavay, 1912: 17, pi. 27, figs 1-2,

Cryptopecten bullatus - Dukstra, 1995b: 60, figs 115-118 [synonymy, references, description, discussion].

Material EXAMINED. — The type material (see DlJKSTRA, 1995b: 60).

Indonesia. CORINDON .Makassar Strait: stn B 248, 00°54'S, 119°29'E, 170 m, 3 v.

Karubar, Kai Islands: stn DW 02, 05°47'S, 132°13'E, 209-240 m, 2 lv. — Stn DW 15, 05°17'S, 132°41’E, 212-

221 m, 5 lv, 6 rv. — Stn DW 18, 05°18'S, 133°01'E, 205-212 m, 2 spms, 2 lv, 2 rv. — Stn DW 24, 05°32'S, 132°51E.

230-243 m, 1 lv, 3 rv. — Stn DW 29. 05°36'S, 132°56'E, 181-184 m, 1 rv. — Stn DW 30, 05°39'S, 132°56'E. 111-

118 m, 4 lv, 1 rv. — Stn DW 32, 05°47'S, 132°51E. 170-206 m, 1 lv.

Tanimbar Islands: stn DW 44, 07°52'S, 132°48'E, 291-295 m, 4 lv. — Stn CP 47, 08°01'S, 132°55'E, 235-246 m,

1 rv. — Stn DW 49, 08°00'S, 132°59'E. 206-210 m, 1 lv, 2 rv. — Stn DW 50, 07°59'S, 133°02'E. 184-186 m, 1 lv, —

Stn DW 61, 09°05'S, 132°44’E, 235-236 m, 2 lv, 1 rv. — Stn CP 79, 09°16’S, 131°22'E, 239-250 m, 1 rv. —

Stn DW 80, 09°37'S, 131°02'E, 199-201 m, 1 lv. — Stn CP 86, 09°26'S, 131°13'E, 223-225 m, 2 spms, 1 lv, 1 rv.

Source :

274

H. H. DIJKSTRA & W. W. KASTORO

Figs 138-145. — Cryptopecten bullatus (Dautzenberg & Bavay, 1912): 138-141, Karubar, stn DW 18, 19.4 x

19.9 mm (db): 138, left valve, exterior; 139, left valve, interior; 140, right valve, exterior; 141, right valve

- 142‘145- Karubar, stn DW 24: 142, right valve, exterior, central detail, scale bar 100 urn;

143 right valve, exterior, prodissoconch, scale bar 100 |im; 144, left valve, exterior, ventral marginal detail, scale

bar 1 mm; 145, right valve, exterior, ventral marginal detail, scale bar 1 mm.

Source : MNHN, Paris

PECTIN01DEA FROM EASTERN INDONESIA

275

DISTRIBUTION. — Throughout the western, southwestern and Central Pacific; also known from the western

Indian Ocean; present specimens from 1 1 1-295 m, alive in 212-223 m.

Cryptopecten nux (Reeve, 1853)

Figs 146-149

Peciert coruscans Reeve, 1853: sp. 143, pi. 32, fig. 143 (non Hinds, 1845).

Pecten nux Reeve, 1853: errata.

Cryptopecten nux nux - Hayami, 1984: 100, pi. 2, fig. 4, pi. 3, figs 1-2, pi. 9, figs 2-5, pi. 12, figs 1-2 [synonymy,

references, description, discussion],

MATERIAL EXAMINED. — The type material (see below).

Indonesia. Corindon, Makassar Strait: stn B 248, 00°54'S, 119°29'E, 170 m, 3 v.

Karubar, Kai Islands: stn DW 01, 05°46'S, 132°10'E, 156-305 m, 5 lv, 2 rv. — Stn DW 02, 05°47'S, 132°13'E, 209-

240 m, 3 lv, 2 rv. — Stn DW 22, 05°22'S, 133°01'E, 124-850 m, 7 lv, 18 rv.

Type Material. — Lectotype BMNH 1950.11.14.52, designated by Wagner (1989).

Type Locality. — Panglao, Bohol, Philippines (restricted by Wagner, 1989).

DISTRIBUTION. — Western and northwestern Indian Ocean, western and southwestern Pacifc, alive in 30-200 m

(Dukstra, unpubl. data).

Genus HAUMEA Dali, Bartsch & Rehder, 1938

Haumea Dali, Bartsch & Rehder, 1938: 86. Type species (OD): Haumea juddi Dali, Bartsch & Rehder, 1938 (= Pecten

loxoides G.B. Sowerby II, 1882). Recent, Hawaiian Islands, 7-15 m.

Diagnosis. — Free swimming Aequipectinini, slightly obliquely suborbicular, right valve more convex than

left, auricles subequal, valves sculptured with 18-20 radial costae, interspaces with fine concentric lamellae,

auricles weakly sculptured with 4-6 radial riblets and fine close-set concentric lamellae, byssal notch moderately

deep, ctenolium present.

DISTRIBUTION. — Pliocene-Recent. Western Indian Ocean, western and southwestern Pacific; littoral to

sublittoral depths.

Remarks. — Hertlein (1969: N357) treated Haumea as a synonym of Argopecten Monterosato, 1899, a

subgenus of Chlamys. Waller (1991 : 32) placed Argopecten in the Aequipecten group, or Aequipectinini (1993 :

198). Haumea differs from Argopecten by having a more compressed, and somewhat obliquely orbicular left valve

(in Argopecten more convex and elongate, especially in young specimens) and smaller, subequal auricles (in

Argopecten larger, inequal with a prominent anterior auricle in the right valve). Microsculpture of Haumea is finer

with more closely spaced concentric lamellae. The genus includes H. inaequivalvis (G.B. Sowerby II, 1842),

H. loxoides and H. rehderi (Grau, 1960). Argopecten is not known in the Indo-Pacific region.

Haumea inaequivalvis (G.B. Sowerby II, 1842)

Fig. 150

Pecten inaequivalvis Sowerby, 1842: 50, pi. 19, figs 193, 194, 195.

Pecten inaequivalvis - REEVE, 1852: sp. 1, pi. 1, figs 1, 6. — DESHAYES, 1863: 31. — MARTENS, 1880: 138. — Melvill

& Standen, 1898: 46.

Vola inaequivalvis - H. & A. Adams, 1858: 554. — DUNKER, 1882: 244.

276

H. H. DIJKSTRA & W. W. KASTORO

Pecten (Vola) inaequivalvis - KOster & Kobelt, 1888: 236, pi. 62, figs 5-8.

Chlamys ( Aequipecten ) inaequivalvis - Dautzenberg & Bouge, 1933: 426.

Chlamys inaequivalvis - Kira, 1962: 137, pi. 49, fig. 13.

Haumea inaequivalvis - Dijkstra, 1984b: 28, 4 figs. — ROMBOUTS, 1991: 43, pi. 25, fig. 1.

Crypiopecien inaequivalvis - Bernard, Cai& Morton, 1993: 50.

MATERIAL EXAMINED. — The type material (see below).

Indonesia. Karubar, Kai Islands: stn DW 30, 05°39'S, 132°56'E, 111-118 m, 1 rv.

Type Material. — Lectotype, here designated, the shell figured by Reeve (1852: pi. 1, fig. 6; H 30.8,

L 33.5, D 13.5 mm) BMNH 1994126/1. Two paralectotypes: BMNH 1994126/2-3.

Type Locality. — Philippine Islands.

DISTRIBUTION. — Western and northeastern Indian Ocean, western, southwestern and central South Pacific;

alive in 3-46 m (Dijkstra, unpubl. data).

Description. — Shell rather small, usually 20 mm high, occasionally up to ca. 30 mm, suborbicular,

inequivalve, right valve more convex than left, auricles unequal, umbonal angle ca. 100-1 10°. Prodissoconch

height ca. 220 ftm. Both valves sculptured with 18-20 regularly spaced, prominent radial costae. Radial ribs near

anterior and posterior margin weakly concentrically striated. Interspaces between radial costae with concentric

lamellae. Auricles sculptured with 2-4 weak radial riblets. Inner surface strongly plicated near ventral margin.

Hinge line straight. Byssal notch moderately deep. Resilifer obliquely triangular. Active ctenolium weak, with

4-5 teeth. Colour blackish grey or brownish marked with a few white spots and black streaks, right valve whitish

or light brown.

Remarks. — The present young specimen is similar to the type material. Young specimens could be easily

confused with the closely related species Haumea rehderi (Grau, 1960) from the same region, which differs by

having more prominent and widely spaced concentric lamellae between the radial costae, more prominent radial

riblets on the auricles, and a deeper byssal notch. Haumea loxoides differs by having a more fragile shell, a more

oblique shape, and a creamy mottled with red colour.

Genus VOLACHLAMYS Iredale, 1939

Volachlamys Iredale, 1939: 356. Type species (OD): Pecten cumingii Reeve, 1853. Recent, Queensland, Australia.

Diagnosis. — Non-cemented, free swimming Aequipectinini, orbicular, equilateral, subequivalve, auricles

subequal, valves sculptured with 14-24 radial costae, interspaces with concentric lamellae, byssal notch well-

developed and moderately deep, ctenolium present.

Distribution. — ?Miocene-Recent (Hayami, 1989: 16). Indo-West Pacific; intertidal to sublittoral depths.

Volachlamys singaporina (G.B. Sowerby II, 1842)

Figs 151-155

Pecten singaporinus G.B. Sowerby II, 1842: 74, pi. 13, fig. 55, pi. 14, fig. 71.

Pecten pica Reeve, 1853: sp. 115, pi. 27, figs 115 a-b.

Pecten singaporinus - Reeve, 1853: sp, 74, pi. 20, fig. 74. — KOster & Kobelt. 1888: 94, pi. 25, figs 2-4

Pecten pica - KOSTER & Kobelt, 1888: 255-256, pi. 67, figs 1-2.

Chlamys ( Argopecten ) singaporina - DIJKSTRA, 1990: 8.

Volachlamys singaporina - ROMBOUTS, 1991: 62, pi. 22, fig. 6. — Dharma, 1992: 84, pi. 20, figs 7-7a-d

Chlamys singaporina - Bernard, Cai & Morton, 1993: 49,

Source : MNHN. Paris

PECTINOIDEA FROM EASTERN INDONESIA

277

Figs 146-149. — Cryptopecten nux (Reeve. 1853), Karubar, stn DW 22, 6.9 x 6.9 mm (lv), 6.1 x 6.2 mm (rv):

146, left valve, exterior; 147, left valve, interior; 148, right valve, exterior; 149, right valve, interior.

Fig. 150, — Haumea inaequivalvis (G.B. Sowerby II, 1842), Karubar, stn DW 30, 6.0 x 6.8 mm, right valve, exterior.

Figs 151-153. — Volachlamys singaporina (G.B. Sowerby II, 1842), Karubar, stn CP 65, 36.5 x 37.1 mm (lv):

151, left valve, exterior; 152, left valve, interior; 153, left valve, exterior, antero-central detail, scale bar 1 mm.

Source :

278

H. H. DIJKSTRA & W. W. KASTORO

FlGS 154-155. — Volachlamys singaporina (G.B, Sowerby II, 1842), Karubar, stn CP 65, 32.8 x 33.9 mm (rv):

154, right valve, exterior; 155, right valve, interior.

FlG® 156-159. — Anguipecien cf. picturatus Dijkstra, 1995, Karubar, stn DW 30, 26.6 x 24 5 mm (lv) 21 4 x

20.0 mm (rv): 156, left valve, exterior; 157. left valve, interior; 158, right valve, exterior; 159. right valve

intp.nnr & *

Source : MNHN. Paris

PECTINOIDEA FROM EASTERN INDONESIA

279

MATERIAL EXAMINED. — The type material (see below).

Indonesia. Karubar, Tanimbar Islands : stn DW 64, 09°13'S, 132°31'E, 179-180 m. 3 lv. — Stn CP 65, 09°14'S,

132°27'E, 174-176 m, 10 lv, 7 rv.

Type Material. — P. singaporinus: lectotype (H 54.2, L 54.7, D 15.9 mm) BMNH 1994127/1, here

designated. Two paralectotypes: BMNH 1994127/2-3. — P. pica : three syntypes BMNH 1994139.

TYPE Locality. — P. singaporinus: Singapore. — P. pica: "New Zealand" [incorrect locality].

DISTRIBUTION. — Throughout the southwestern Pacific to northern Australia; alive from intertidal to subtidal

depths. Present specimens probably washed into deep water.

DESCRIPTION. — Shell up to ca. 45 mm high, suborbicular, equilateral, equivalve, very compressed, auricles

unequal, umbonal angle ca. 90°. Prodissoconch height ca. 260 |im. Both valves sculptured with 18 to 24

(commonly 20 to 22) regularly spaced, smooth rounded radial costae. Interspaces between radial costae with fine

concentric lamellae. Anterior auricle of left valve sculptured with 6-8 weak radial riblets and very fine close-set

concentric lamellae; riblets nearly absent on posterior auricle. Hinge line straight. Byssal notch moderately deep.

Ctenolium well-developed with 4-6 teeth. Resilifer triangular oblong. Inner side with prominent plicae near ventral

margin. Colour greyish or creamy with brown and/or whitish maculations and streaks.

REMARKS. — Present specimens are similar to the original and subsequent descriptions and illustrations by

Reeve (1853), although with fewer radial costae (18-20). Sowerby (1842) mentioned 24, Reeve (1853) ca. 22

and KUSTER & Kobelt (1888) 20-22 radial costae. Material examined from throughout the southwestern Pacific,

South China Sea, Malaysia and Indonesia (BMNH, HD, MNHN, RMNH, ZMA) shows variation in number of

radial costae, decreasing to eastern Indonesia and northeastern Australia. It is possible that V. cumingii (Reeve,

1853) from Queensland is only a geographical variant (under study).

Subfamily PECTININAE Wilkes, 1810

Tribe DECATOPECTININI Waller, 1986

Genus ANGUIPECTEN Dali, Bartsch & Rehder, 1938

Anguipecten Dali, Bartsch & Rehder, 1938: 92. Type species (OD): Anguipecten gregoryi Dali, Bartsch & Rehder, 1938

(= Pecten lamberti Souverbie in Souverbie & MONTROUZIER, 1874). Recent, Hawaiian Islands, 470-571 m .

DIAGNOSIS. — Free swimming Decatopectinini, suborbicular, laterally compressed, with 9-40 rounded radial

costae, sculptured with very closely spaced commarginal lamellae, auricles subequal to equal, byssal notch nearly

absent, no byssal fasciole, ctenolium weakly developed.

Distribution. — Miocene-Recent. Indo-West Pacific; littoral to sublittoral depths.

Anguipecten cf. picturatus Dijkstra, 1995

Figs 156-159

Pecten aurantiacus A. Adams & Reeve in A. ADAMS, 1850: 74, pi. 21, fig. 12 (non Roding, 1798, nec J. Sowerby. 1820,

nec Defiance, 1825).

Anguipecten picturatus Dijkstra, 1 995a: 1 7 ( nom . nov. for P. aurantiacus Adams & Reeve).

Pecten aurantiacus - REEVE, 1853: sp. 105, pi. 26, fig. 105. — K0STER & KOBELT, 1888: 171, pi. 47, fig. 7.

Gloripallium aurantiacum - MASUDA, 1962: 197.

Anguipecten lamberti - Abbott & Dance, 1982: 312, fig. (non Souverbie in Souverbie & Montrouzier, 1874).

Anguipecten aurantiacus - DIJKSTRA, 1984a: 9, figs; 1991: 41.

280

H. H. DIJKSTRA & W. W. KASTORO

Bractaechlamys (sic) aurantiaca - Matsukuma, Okutani & Habe, 1991: 185, pi. 134, fig. 4.

Decadopecten (Anguipecten) aurantiacus - ROMBOUTS, 1991: 38, pi. 13, fig. 12.

Bractechlamys aurantiaca - Bernard, Cai & Morton, 1993: 50. — Lan, 1993: 161, fig,

MATERIAL EXAMINED. — The type material (see below).

Indonesia. Karubar, Kai Islands : stn DW 30, 05°39'S, 132°56’E, 111-118 m, 2 lv, 2 rv.

Type Material. — Holotype BMNH 1950. 1 1 .14.8.

Type Locality. — "China Sea" [= South China Sea].

Distribution. — Western Indian Ocean, western and southwestern Pacific; alive in subtidal to sublittoral

waters (2-90 m) (DIJKSTRA, unpubl. data).

Description. — Present specimens of the Kai Islands are up to 25 mm high, somewhat triangularly

elongated, nearly equilateral and equivalve, right valve slightly more convex than left valve, auricles subequal,

umbonal angle 85°. Prodissoconch heigh ca. 250 pm. Both valves undulated and sculptured with 7 rounded radial

costae. Microsculpture consisting of very fine and closely set, commarginal lamellae. Faint secondary radial riblets

developed on radial costae near ventral margin. Auricles weakly sculptured with 4-5 radial riblets. Hinge line

straight, somewhat raised on right valve. Inner surface plicated, more prominently so near periphery. Resilifer

triangular, erected. Byssal notch very small, byssal fasciole absent. Ctenolium weakly developed with 3 teeth.

Colour creamy, maculated with dots and streaks, right valve more uniform and paler.

Remarks. — The present valves differ strongly from the type material of P. aurantiacus by having a more

orbicular shape (typical specimens are more elongated), fewer 7 vs 9-15) radial costae and are nearly lacking

secondary radial riblets on costae (strongly developed and scabrous in typical specimens). The shape is somewhat

similar to Mirapecten rastellum (Lamarck, 1819), but the latter is sculptured with strongly erected scales on the

radial costae and dorsal margin of right valve. The byssal notch is also wider than in M. rastellum and the auricles

are larger.

DISCUSSION

A total of 19 species ol Propeamussiidae and 11 species of Pectinidae are present in the material from the

Karubar cruise in the Arafura Sea. Twenty-eight species (18 propeamussiids and 10 pectinids) were collected near

the Kai Islands, and twelve species (6 each of propeamussiids and pectinids) near the Tanimbar Islands. Two

species ( Parvamussium conspectum and Veprichlamys versipellis) are new to science, and 8 are new records for

Indonesia: Propeamussium alcocki, P. investigators, P. rubrotinctum, Parvamussium squalidulum, P. thetidis,

P. vesiculatum, Delectopecten fluctuatus, and Haumea inaequivalvis.

There are 33 stations without pectinoids and 26 stations have only one species. Five stations (02, 13, 29, 31,

32) have six species and station 18 has seven species (3 of them live-taken). Half of the species are represented by

at least one live-taken sample, others only by empty shells. Haumea inaequivalvis and Volachlamys singaporina

are intertidal to subtidal species and the Karubar records represent empty shells carried downslope to water depths

where the species is apparently not normally living. Other than these two species, the depth range indicated by

empty shells appears compatible with what is known for the relevant species elsewhere in the Indo-Pacific.

The Kai Islands were selected as the site of the Danish 1922 and Karubar 1991 Expeditions because

Mortensen had hypothesized that deep-sea species occured there at considerably shallower depths than elsewhere

in the world. Ten species of Pectinoidea were collected alive during both the Karubar and New Caledonia

dredging programs (DIJKSTRA, 1995b) and in this respect it may be of interest to compare their bathymetric range

in the two regions (Table 1). Propeamussium alcocki appears to range deeper in the Arafura Sea than in New

Caledonia, but the first occurence of 8 species is shallower or considerably shallower in the Arafura Sea than in

New Caledonia. The limited evidence available thus appears to support MORTENSEN's hypothesis.

Source : MNHN Paris

PECTINOIDEA FROM EASTERN INDONESIA

281

100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 m

Table 1. — Bathymetric range of Pectinoidea in the Arafura Sea (thick bar) and New Caledonia (thin line).

The Indonesian archipelago has been touched by several major expeditions ( Challenger , Valdivia , Galathea ), and

was the main focus of several specific ones ( Siboga , Snellius-II. Karubar). A total of 43 deep-water pectinoids

are now recorded from the archipelago (Table 2, next page), with most of present knowledge based on the last three

mentioned expeditions. Despite this collecting effort, the deep-sea fauna of Indonesia is still probably very

incompletely known:

(a) Seventeen species (40%) were first discovered or recorded during the recent SNELLIUS-II (1984-85) and

Karubar (1991) expeditions.

(b) Nine species (21%) taken during the historical Challenger, Valdivia and Siboga expeditions have never been

taken again since then in Indonesian waters.

(c) In the Karubar material, on average, each species is present at 4.4 stations, but there is considerable

variation in occurence patterns, with Delectopecten alcocki present at 23 stations and 9 species present at single

stations. The new species and species representing new records for Indonesia are present on average at 3.9 stations:

it thus seems that they are not significantly rarer than the species already known from Indonesia, which are present

on average at 4.7 stations.

This indicates that rather many more species of Pectinoidea will probably be discovered when new regions of

the Indonesian archipelago will be properly sampled.

ACKNOWLEDGEMENTS

We are very grateful to Dr P. BOUCHET (MNHN) for his advice and critical comments and for suggesting the

data in the discussion. Prof. A. MATSUKUMA kindly translated Japanese text into English. We also express our

gratitude to several staff members of the following institutions, who allowed us to study the type and systematic

collections, for loaning type material or donating specimens for comparative taxonomic study: Ms K. Way

(BMNH), Dr K.V. Surya Rao (ZSI), Dr Z. Wang (IOAS). Prof E. Gittenberger and Mr J. Goud (RMNH),

Dr W. PONDER and Mr I. Loch (AMS), Dr P.G. Oliver and Ms A. Trew (NMW), Prof. I. Hayami of the

Department of Applied Biology, Kanagawa University at Hiratsuka and Mr T.C. Lan (Taipei). Thanks are also due

to Mr R.G. MOOLENBEEK (ZMA) for his help, to Mr D. PLATVOET and Mr H. HOENSELAAR (ZMA) for their

technical assistance in preparing SEM-micrographs and to Dr Rudo VON COSEL (MNHN) for taking most of the

photographs of the shells.

Source

282

H. H. DIJKSTRA & W. W. KASTORO

CH VA SI GA SN KA

Table 2. Pectinoidea from Indonesia normally occuring in depths deeper than 100 m, as collected by

the Challenger (CH), Valdivia (VA), Siboga (SI), Galathea (GA), Snellius-II (SN) and Karubar (KA) Expeditions.

PECTINOIDEA FROM EASTERN INDONESIA

283

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ESUL VIS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS E

Mollusca, Gastropoda: The Muricidae collected

during the KARUBAR Cruise in eastern Indonesia

Roland HOUART

Scientific Associate

Institut Royal des Sciences Naturelles de Belgique

Departement des Invertebres Recents

Rue Vautier, 29

B-1000 Brussels, Belgium

ABSTRACT

Sixteen species of Muricidae were collected during the French-Indonesian Karubar cruise. Most of them are new

records for the region. Leptotrophon kastoroae sp. nov. is described and compared to three similar species from

New Caledonia.

RESUME

Mollusca Gastropoda : Les Muricidae recoltes lors de la campagne Karubar en Indonesie

orientale.

Seize espbces de Muricidae ont dtd rdcoltees lors de la campagne franco-indonesienne Karubar. La plupart de celles-ci

n’dtaient pas connues dans la region. Leptotrophon kastoroae sp. nov. est d£crite et comparde it trois espbces similaires

de Nouvelle-Calddonie.

INTRODUCTION

Rather few species of Muricidae were collected during the French-Indonesian Karubar cruise on board of the

Indonesian R. V. "Baruna Jaya 1 Indonesia and Papua New Guinea have a very diverse muricid fauna (DHARMA,

1988; Hinton, 1972, 1979), and it was thus unexpected that as few as 16 species are represented in the expedition

material. Furthermore one third (6 species) occurred only as empty shells. Eleven species (69%) are represented by

a single sample and none has been collected at more than three stations. Most probably the bottom types surveyed

during KARUBAR were not favourable to muricids. Most hauls sampled fauna on bottoms of mud or sandy mud,

whereas a majority of muricids favour hard substrates. Only 15 stations (16% of all stations) yielded muricids, and

in 10 of these the family Muricidae is represented by a single species. Three stations (stn 15, 22, 50) yielded two

species, and two (stn 18, 30) gave three species each. Despite the small size of the collection, as many

Houart, R., 1997. — Mollusca Gastropoda: The Muricidae collected during the Karubar Cruise in eastern Indonesia.

In: A. Crosnier & P. Bouchet (eds), Rdsultats des campagnes Musorstom. Volume 16. Mem. Mus. natn. Hist, nat.,

172 : 287-294. Paris ISBN 2-85653-506-2.

Source : MNHN, Paris

288

R. HOUART

as 1 1 species (69%) are new records for the Indonesian archipelago. All this points out to a diverse, but still poorly

recorded, fauna, at least in the 100-250 m depth range, where most of the findings have been made.

ABBREVIATIONS AND TEXT CONVENTIONS

Repositories

AMS

MNHN

NSMT

POLIPI

RH

Australian Museum, Sydney, Australia

Museum national d'Histoire naturelle, Paris, France

National Science Museum, Tokyo, Japan

Puslitbang Oseanologi-LIPI [Research and Development Centre for Oceanology - Indonesian

Institute of Sciences], Jakarta

Author's collection

Other abbreviations

dd empty shell

lv collected alive

SYSTEMATIC ACCOUNT

Family MURICIDAE Rafinesque, 1815

Subfamily MURICINAE Rafinesque, 1815

Genus HAUSTELLUM Schumacher, 1817

Haustellum multiplication (Sowerby, 1895)

Figs 1-2

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: sin CP 65, 09°14'S, 132°27'E, 174-176 m,

1 dd.

Remarks. — The species is already known from neighbouring localities, on the Australian side of the Arafura

Sea (Ponder & Vokes, 1988). Only one dead and damaged specimen was collected during the expedition.

Genus CHICOREUS Montfort, 1810

Chicoreus (Triplex) axicornis (Lamarck, 1822)

Material EXAMINED. —Indonesia. Karubar, Kai Islands: stn DW 30, 05°39'S, 132°56'E, 111-118 m, 1 lv.

Chicoreus (Siratus) cf. pliciferoides Kuroda, 1942

Figs 3-4

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn CP 25, 05°30'S, 132°52’E, 336-346 m, 1 dd.

Remarks. — The identity of the single collected specimen is uncertain. It is probably not C. pliciferoides, but

this is the most similar species. The present material is lighter, and relatively smaller, with narrower varices.

Nevertheless, the shell of C. pliciferoides being very variable morphologically, an examination of more numerous

specimens from the same region is necessary before a decision can be reached.

Chicoreus (Chicopinnatus) orchidiflorus (Shikama, 1973)

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 30, 05°39'S, 132°56'E, 111-118 m, 1 dd.

Source : MNHN, Paris

MOLLUSCA GASTROPODA MURIC1DAE FROM EASTERN INDONESIA

289

Figs 1-2. — Haustellum multiplication (Sowerby, 1895). Indonesia, Tanimbar Islands, 32.8 mm.

Figs 3-4. — Chicoreus ( Siratus ) cf. pliciferoides Kuroda, 1942. Indonesia, Kai Islands, 50.1 mm.

Figs 5-6. — Poirieria (Flexopteron) poppei Houart, 1993. Indonesia, Kai Islands, 33.3 mm.

Source : MNHN, Paris

290

R. HOUART

Genus POIRIERIA Jousseaume, 1880

Poirieria (Flexopteron) poppei Houart, 1993

Figs 5-6

Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn CC 10, 05°21'S, 132°30'E, 229-289 m, 1 Iv.

Remarks. — New record for Indonesia.

Genus DERMOMUREX Montcrosato, 1890

Dermomurex (Takia) infrons Vokes, 1974

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn DW 49. 08°00'S, 132°59'E, 206-210 m,

1 Iv. — Stn DW 50, 07°59'S, 133°02'E, 184-186 m, 1 Iv, 1 dd.

REMARKS. — New record for Indonesia. Dermomurex infrons was originally described from Japan (as Murex

inermis Sowerby, 1841, non M. inermis Philippi, 1836). The type locality is confirmed by specimens that were

recently collected off Tosa Bay and Sagami Bay (NSMT). Specimens are also known from Transkei, South Africa

(material in Natal Museum, Pietermaritzburg).

Subfamily MURICOPSINAE Radwin & D'Attilio, 1971

Only three species (3 specimens) were collected during the Karubar expedition. All have been originally

described from the Philippine Islands and are new records for Indonesia.

Genus FAVARTIA JOUSSEAUME, 1880

Favartia jeanae Bertsch & D'Attilio, 1980

Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 18. 05°I8'S, 133°OI'E, 205-212 m, 1 lv.

Genus MU REX! ELLA Clench & Perez Farfante, 1945

Murexiella judithae (D'Attilio & Bertsch, 1980)

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 30, 05°39'S. 132°56'E, 111-118 m, 1 dd.

Murexiella peregrina Olivera, 1980

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 22, 05°22'S, 133°01'E. 85-124 m. I lv.

Subfamily ERGALATAXINAE Kuroda & Habe, 1971

Genus ERGALATAX Iredale, 1931

Ergalatax tokugawai Kuroda & Habe, 1971

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn DW 50, 07°59'S, 133°02'E, 184-196 m

2 lv, 1 dd.

Remarks. New record for Indonesia. The species is known from Japan and the Philippine Islands.

Source : MNHN, Paris

MOLLUSCA GASTROPODA MURICIDAE FROM EASTERN INDONESIA

291

Genus CYTHAROMORULA Kuroda, 1953

Cytharomorula springsteeni Houart, 1995

Material EXAMINED. — Indonesia. Karubar, Kai Islands: sin DW 18, 05°18'S, I33°01'E, 205-212 m, 1 dd. —

Sin DW 32, 05°47'S, 132°51'E, 170-206 m. 3 dd.

Remarks. — New record for Indonesia. The species was only recently described from the Philippine Islands

(HOUART, 1995a: 255).

Genus ORANIA Pallary, 1900

Orania archaea Houart, 1995

Material EXAMINED. — Indonesia, karubar, Kai Islands: stn DW 22, 05°22'S, 133°01'E, 85-124 m, 1 Iv.

Remarks. — Although only recently described (Houart, 1995a: 267), it is present in many collections from

many localities in the Indo-West Pacific, but was misidentified.

Subfamily TYPHINAE Cossmann, 1903

Genus SIPHONOCHELUS Jousseaume, 1880

Siphonochelus japonicus (A. Adams, 1863)

MATERIAL EXAMINED. — Indonesia. Karubar. Kai Islands : stn DW 13, 05°26'S, 132°38'E, 417-425 m, 2 lv,

1 dd. — Stn DW 15, 05°17'S, 132°41'E, 212-221 m, I lv, 5 dd.

Tanimbar Islands: stn DW 49, 08°00'S, 132°59'E, 206-210 m, ldd.

REMARKS. — New record for Indonesia. Originally described from Japan, its presence there is confirmed by

material from Sagami Bay (NSMT 44066). The species is also known from off Queensland, Australia (AMS

Cl 69094).

Subfamily TROPHONINAE Cossmann, 1903

Genus TROPHONOPSIS Bucquoy & Dautzenberg, 1882

Trophonopsis carduelis (Watson, 1883)

Figs 7-8

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn CP 20, 05°15'S, 132°59'E. 769-809 m, 2 Iv,

1 dd. — Stn CC 21, 05°14'S, 133°00’E, 688-694 m, 2 dd.

REMARKS. — New record for Indonesia. Trophonopsis carduelis was described from off Sydney (Australia). The

holotype, although having one teleoconch whorl more, is very similar to the specimen illustrated here.

Trophonopsis plicilaminatus (Verco, 1909)

Figs 9-10

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 69, 08°42'S, 131°53'E, 356-368 m,

1 lv, 6 dd.

REMARKS. — New record for Indonesia, The species is known from the type locality, off Beachport, South

Australia, and from New Caledonia (HOUART, 1995b).

Source :

292

R. HOUART

Figs 7-8. — Trophonopsis carduelis (Watson, 1883). Indonesia, Kai Islands, 22.8 mm.

Figs 9-10. — Trophonopsis plicilaminaius (Verco, 1909). Indonesia, Kai Islands, 13 mm.

Figs 11-12. — Leptotrophon kastoroae sp. nov. Indonesia, Kai Islands, holotype, 11.2 mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA MUR1CIDAE FROM EASTERN INDONESIA

293

Genus LEPTOTROPHON Houart, 1995

Leptotrophon kastoroae sp. nov.

Figs 11-12

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 14, 05°18'S, 132°38'E, 245-246 m, 2 lv,

1 dd. — Stn DW 15, 05°17'S, 132°41'E, 212-221 m, 1 lv (holotype), 12 dd (paratypes: 2 MNHN, 2 POLIPI, 1 RH). —

Stn DW 18, 05°18’S, 133°01E, 205-212 m, 1 lv, 4 dd.

Type Material. — Holotype MNHN. Paratypes: 2 MNHN, 2 POLIPI, 1 RH.

Type Locality. — Indonesia. Kai Islands, Karubar, stn DW 15, 05°17'S, 132°41'E, 212-221 m.

DISTRIBUTION. — Indonesia, Kai Islands, alive in 212-245 m.

DESCRIPTION. — Shell up to 11.2 mm (holotype), slender, spinose, lightly built. Spire high, with

1.5 protoconch whorls, and up to 5 weakly convex, spinose teleoconch whorls. Suture impressed. Protoconch

broad, globose, smooth, glossy. Terminal varix unknown (eroded). Axial sculpture of teleoconch whorls

consisting of numerous, weak growth striae and high, rounded varices, each with long, narrow, sharp primary and

secondary spines. Shoulder spine longest, weakly adapically bent. First and second whorls with 9 varices, third

with 9 or 10, fourth with 10 or 11, last whorl with 10 varices. Spiral sculpture of strong, smooth, primary and

secondary cords: first to third whorl with 2 primary cords, fourth with 2 primary and 1 secondary cords, shoulder

with 1 secondary cord, last whorl with 5 or 6 primary and secondary cords, shoulder with 1 secondary cord.

Occasionally with numerous, narrow spiral striae. Aperture small, rounded. Columellar lip flaring, smooth. Lip

erect, adherent adapically. Anal notch indistinct. Outer lip smooth, with 4-6 more or less visible denticles within,

very variable in strength. Siphonal canal long, narrow, straight, open, smooth. White.

Remarks. — The genus Leptotrophon Houart, 1995 was described to include numerous small species of

Trophoninae with a spinose, delicate, and small shell, all from the New Caledonia region (Houart, 1995b). The

occurrence of another, new species in Indonesia is a very interesting range extension for the genus. Leptotrophon

kastoroae is similar to three other spinose species of Leptotrophon-. L. spinacutus Houart, 1986; L. bernadettae

Houart, 1995, and L. rigidus Houart, 1995. From L. spinacutus, L. kastoroae differs in having relatively longer

spines, a narrower siphonal canal, and a broader aperture with a narrower apertural varix. From L. bernadettae, it

differs in its relatively broader and rounded protoconch, narrower columellar lip, and more slender shell.

L. kastoroae is relatively smaller than L. rigidus, with more numerous, longer spines, a twice larger protoconch,

and a narrower siphonal canal.

ETYMOLOGY. — Named for Mrs W. KASTORO (Indonesian Institute of Sciences Research and Development

Centre for Oceanology, Jakarta), one of the member of the Karubar cruise.

REFERENCES

Dharma, B., 1988. — Indonesian Shells. Vol. 1. PT. Sarana Graha, Jakarta: i-xvi + 1-111.

Hinton, A., 1972. — Shells of New Guinea and the Central Indo-Pacific. Robert Brown & Associates, Port Moresby and

Jacaranda Press, Milton, 94 pp.

Hinton, A., 1977. — Guide to Shells of Papua New Guinea. R. Brown & Assoc., Port Moresby , 74 pp.

Houart, R., 1995a. — The Ergalataxinae (Gastropoda, Muricidae) from the New Caledonia region with some comments

on the subfamily and the description of thirteen new species from the Indo-West Pacific. Bulletin du Museum national

d’Histoire naturelle, Paris, 4e s6r„ 16, section A, n° 2-4, 1994 (July 1995): 245-197.

Houart, R., 1995b. — The Trophoninae (Gastropoda, Muricidae) of the New Caledonian region. In: P. BouCHET (ed.),

Resultats des Campagnes MUSORSTOM, Volume 14. Memoires du Museum national d'Histoire naturelle, 164:

459-498.

Source :

294

R. HOUART

Ponder. W.F. & E.H. Vokes, 1988. — Revision of the Indo-West Pacific fossil and Recent species of Murex s.

Haustellum (Mollusca: Gastropoda: Muricidae). Records of the Australian Museum, suppl. 8: 1-160.

Source :

;ULT\TS DES CAMPAGNES MUSORSTOM. VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DI

Mollusca Gastropoda: Arafura Sea Cancellariidae

collected during the KARUBAR Cruise

Andre VERHECKEN

Scientific Associate, Afdeling Malacologie,

Koninklijk Belgisch Instituut voor Natuurwetenschappen,

Vautierstraat, 29, B-1000 Brussels, Belgium

ABSTRACT

The deep-water Cancellariidae collected during the Karubar cruise near the Kai and Tanimbar Islands are represented

by 20 species (9 new), only two of which were recorded earlier from the Arafura Sea. As many as 14 species (70% of the

total) are represented by single specimens, and 17 (85%) have been collected at one station only: this points to a still

more diverse cancellariid fauna. New species of Axelella, Perplicaria, and Solatia represent the first occurence of these

genera in the Indo-West Pacific. Admete aethiopica Thiele, 1925, recently suspected to be a species of Turridae, is

confirmed as a cancellariid.

RESUME

Mollusca Gastropoda : Cancellariidae de la mer d'Arafura recoltes durant la campagne Karubar.

Les Cancellariidae de mer profonde, recoltes pendant la campagne Karubar prbs des lies Kai et Tanimbar

(mer d’Arafura) sont representes par 20 espbces, dont 9 nouvelles. Deux de ces esp&ces, seulement, 6taient d6j<l connues

de la mer d’Arafura. Pas moins de 14 espbces (70% du total) sont repr6sentees par des specimens uniques, et 17 (85%) ont

dtd rficoltees uniquement a une station : ceci indique que la faune locale de cancellaires est encore plus diversifide que ne le

laissent apparaltre les rdsultats de cette expedition. Des espfeces nouvelles d 'Axelella, Perplicaria et Solatia permettent de

mentionner, pour la premiere fois, ces genres dans l'lndo-Ouest Pacifique. 11 est par ailleurs confirme qu Admete aethiopica

Thiele, 1925, supposee recemment etre un Turridae, est bien un Cancellariidae.

INTRODUCTION

The malacofauna of the Indonesian seas has not yet been adequately inventoried. A review of the Cancellariidae

from that area has been published (VERHECKEN, 1986), based on the rather poorly documented material collected

mainly early this century, furthermore mostly from shallow water. By contrast, the material studied here is

Verhecken, A., 1997. — Mollusca Gastropoda: Arafura Sea Cancellariidae collected during the Karubar Cruise. In:

A. CROSNtER & P. Bouchet (eds), Resultats des Campagnes MUSORSTOM, Volume 16. Mem. Mus. natti. Hist. nat.. 172:

295-323. Paris ISBN 2-85653-506-2.

Source : MNHN Paris

296

A. VERHECKEN

excellently documented and originates from depths from the continental shelf down to about 800 m. Despite that

only a limited geographical area of the northern Arafura Sea was covered, it is unexpectedly rich in specimens (72)

and species (20, nine of which are here described as new to science) of Cancellariidae.

Supraspecific taxonomy of Cancellariidae has not yet been worked out satisfactorily and generic allocation of

several species discussed here proved to be difficult. Genera of small deep-water cancellariids have been named based

on fossil and Recent Australian and New Zealand species. A great confusion prevails, which will be difficult to

unravel without reevaluation of the relevant type material and examination of representative collections. Hence,

generic names used here may be open to discussion and their usage does not necessarily imply their recognition as

the most appropriate name. Therefore, in this paper all genus-level names are treated as genera (no subgenera), and

no new generic names are introduced.

ABBREVIATIONS AND TEXT CONVENTIONS

Repositories

AMS Australian Museum, Sydney

AMNH American Museum of Natural History, New York

AV Author's collection, Mortsel

BMNH The Natural History Museum, London

DMNH Delaware Museum of Natural History, Greenville

NZGS Institute of Geological and Nuclear Sciences, Lower Hutt

KBIN Koninklijk Belgisch Instituut voor Natuurwetenschappen. Brussels

MNHN Museum national d'Histoire naturelle, Paris

NMW National Museum of Wales, Cardiff

NSMT National Science Museum, Tokyo

POLIPI Puslitbang Oseanologi-LIPI [Research and Development Centre for Oceanology - Indonesian

Institute of Sciences], Jakarta

USNM National Museum of Natural History, Smithsonian Institution, Washington DC

WAM Western Australian Museum, Perth

ZMA Zoologisch Museum, Amsterdam

ZMHU Museum fiir Naturkunde, Berlin

Other abbreviations

dd empty shell

lv collected alive

sPm specimen (condition at the time of collecting unknown)

OD original designation

SD subsequent designation

Counting of protoconch whorls follows Verduin (1984). Full references to taxa mentioned in comparisons are

listed in Petit & Harasewych (1990) and are not repeated here.

Paratypes and other reference material are in POLIPI as noted under each species. All other material is in

MNHN.

Source : MNHN , Paris

MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

297

SYSTEMATIC ACCOUNT

Family CANCELLARIIDAE Forbes & Hanley, 1851

Subfamily CANCELLARIINAE

Genus AXELELLA Petit, 1988

Axelella Petit, 1988: 130 (nom. nov. for Olssonella Petit, 1970, non Glibert & Van de Poel, 1967). Type species (OD):

Cancellaria smithii Dali, 1888. Recent, Western Atlantic.

Species of Axelella are found almost exclusively in central American seas, both Caribbean and Panamic-

Paciftc. The two species treated here are certainly not typical representatives and are placed in the genus only by

lack of a more appropriate genus.

Axelella kastoroae sp. nov.

Figs 1-4

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 31, 05°40'S, 132°51'E, 288-289 m, 1 dd.

Type Material. — Holotype MNHN.

Type Locality. — Indonesia. Off Kai Islands, Karubar, stn DW 31, 05°40'S, 132°51'E, 288-289 m.

DESCRIPTION. — Shell biconical, with convex whorls and a large aperture. Protoconch (Figs 2-3) smooth,

paucispiral with 1.0 whorl, maximum diameter 1.0 mm, exposed height 0.7 mm. Transition to teleoconch marked

only by start of teleoconch sculpture. Teleoconch with 4 1/4 convex whorls. Axial sculpture of narrow rounded

ribs: 12, 12, 13 and 12 on first to fourth whorl respectively, 10 on last whorl. Spiral sculpture of broad flat bands,

6 on first teleoconch whorl. From the second whorl on, between the broader bands up to 4 narrower ones occur, all

bands fitting closely together. On third and fourth whorl 8 primary bands, plus secondary and tertiary ones. On last

whorl up to 60 spirals bands, divided axially by sharp, raised incremental riblets (Fig. 4). Suture impressed, very

slightly canaliculate. Aperture semicircular, tapering anteriorly into a short siphonal canal oriented slightly

abaxially. Columella straight, parallel to shell axis, with three strongly oblique folds, adapical one strongest, weak

abapical fold near rim of siphonal canal. Outer lip with crenulated edge, 14 lirae inside. Umbilical slit narrow,

almost completely closed by columellar callus. Siphonal fasciole present but not strong.

Dimensions: 15.5 x 10.4 mm.

REMARKS. — Axelella kastoroae is rather similar to Cancellaria quasilla Petit, 1987 (= Cancellaria cretacea

E. A. Smith, 1899, non Nyst, 1881) from off Southern India, Burma, and off Northern Somalia. The apical

whorls of the holotype of C. quasilla (material examined in the Zoological Survey of India, Calcutta) are strongly

eroded, but specimens from off Somalia (AV) have a multispiral protoconch. In spite of a large distance in time

and space, A. kastoroae also resembles Cancellaria paraguanensis H. K. Hodson in HODSON & HODSON, 1931,

from the Miocene of Venezuela, as figured by JUNG (1965, pi. 75, figs 15-16). The latter species grows up to

36 mm, has a stronger siphonal fasciole and fewer axials on the last whorl as compared to A. kastoroae. HODSON

(1931: 44) and JUNG (1965: 555) did not place C. paraguanensis in a subgenus and Petit (1987: 154) compared

C. quasilla with species of Merica, stating that a determination of subgeneric placement was not yet possible.

These species, which in my opinion are congeneric, differ from typical Cancellaria (s. 5.) species in lacking the

bifid posterior columellar fold. Further, Axelella kastoroae differs from species of Merica in its much more convex

whorls and its broad axial sculpture. The microsculpture of A. kastoroae (Fig. 4) resembles that of Nipponaphera

habei Petit, 1972 (Japan and the Philippines) but is finer; the latter species has a multispiral protoconch and

the teleoconch whorls are strongly angulate. In general form, the new species also bears some resemblance to

Source :

298

A. VERHECKEN

Figs 1-4. — Axelella kasioroae sp. nov.: 1. holotype, 15.5 mm; 2-3, protoconch. Scale bar: 1 mm; 4, detail of

teleoconch sculpture. Scale bar: 0.4 mm.

Figs 5-7. — Axelella cf. nodosivaricosa (Petuch): 5, 14.9 mm, stn CP 67, 146-233 m; 6-7, protoconch. Scale bar

1 mm.

Figs 8-10. — Bonellitia atopodonta (Petit & Harasewych): 8, 22.6 mm, stn CP 83, 285-297 m; 9-10, protoconch of

Fig. 8. Scale bar: 1 mm.

Figs 11-13. — Bonellitia garrardi (Petit): 11, 7.4 mm, stn DW 49, 206-210 m; 12-13, protoconch of same. Scale bar

0.5 mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

299

Axelella funiculata (Hinds, 1843) from western Central America, but that species is more elongated, it has a much

more impressed suture and stronger columellar folds, it lacks the sculpture of sharp, raised incremental riblets and

it has a multispiral protoconch.

ETYMOLOGY. — Named in honour of Mrs Widana KASTORO (POLIPI).

Axelella cf. nodosivaricosa (Petuch, 1979)

Figs 5-7

Agairix (Olssonella) nodosivaricosa Petuch, 1979: 11, 15, figs 26-27.

Scalpiia nodosivaricosa - Springsteen & Leobrera, 1986: 334, pi. 95, fig. 10.

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 67, 08°58'S, 132°06'E, 146-233 m,

I dd.

Type Material. — Holotype (11x9 mm), DMNH 126397.

Type Locality. — Philippines. Off Balicasag, Bohol Island, 300 m.

DISTRIBUTION. — A. nodosivaricosa is known from the Philippines and Japan (AV, KBIN), now possibly

extended to the Arafura Sea.

REMARKS. — The rather eroded shell (14.9 x 9.8 mm) is very close to specimens of A. nodosivaricosa from

the Philippines and Japan (AV, KBIN) but differs from them in having the whorls more inflated, fewer axial ribs

on spire whorls (8, 9, 9 vs 14, 13, 9 on second to fourth whorl respectively), and spiral sculpture only weakly

indicated. The last adult whorl has 7 axial ribs in both the KARUBAR specimen and typical A. nodosivaricosa. The

present specimen has a paucispiral protoconch with 1 whorl, maximum diameter 0.9 mm, exposed height 0.7 mm,

nucleus diameter 0.4 mm. Typical -4. nodosivaricosa (KBIN. AV, Figs 60-62) have a protoconch with 1 1/4 to

1 3/4 whorl, maximum diameter 0.95 mm, exposed height 0.8 mm, nucleus diameter 0.3 mm, and this results in

rather different general appearance (Figs 6-7 vs Figs 61-62). The main difference resides in the regularly squared

cancellation of the earlier whorls in A. nodosivaricosa, versus the strong axial sculpture in the present specimen.

Since only one rather eroded shell is at hand, it is hard to judge the value of cited differences; therefore this shell is

here provisionally identified as conforming A. nodosivaricosa.

Genus BONELL1TIA Jousseaume, 1887

Bonellilia Jousseaume, 1887: 223. Type species (OD): Cancellaria bonellii Bellardi, 1841. Miocene-Pliocene Italy.

Admeiula Cossmann, 1889 has been used for species in this genus; for a discussion on these names, see Verhecken,

1986 : 33.

Bonellilia atopodonta (Petit & Harasewych, 1986)

Figs 8-10

Cancellaria atopodonta Petit & Harasewych, 1986: 440, figs 5-6.

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn CP 20, 05°15'S, 132°59' E, 769-809 m, 1 dd. —

Stn CC 21, 05°14'S, 133°00'E, 688-694 m, 1 lv. — Stn DW 24, 05°32'S, 132°51'E, 230-243 m, 1 dd.

Tanimbar Islands: stn CP 38, 07°40’S, 132°27'E, 620-666 m, 1 lv (POLIPI). — Stn CP 59. 08°20'S. 132°H'E,

399-405m, 1 dd (POLIPI). — Stn CP 69, 08°42'S, 131°53'E, 356-368 m, 7 dd, 1 lv. — Stn CP 70. 08°41'S, 131°47E.

410-413 m, 2 dd (POLIPI). — Stn CP 72, 08°36'S, 131°33'E, 676-699 m. 1 lv. — Stn CP 77, 08°57’S, 131°27'E, 346-

352 m, 1 lv (POLIPI). — Stn CP 78, 09°06'S, 131°24'E, 284-295 m. 1 dd (POLIPI). — Stn CP 83, 09°23'S, 131°00'E,

285-297 m, 1 dd.

Total: 19 specimens; dimensions up to 24.8 x 15.4 mm.

Source :

300

A. VERHECKEN

Type Material. — Holotype (21.5 x 13.6 mm) and 2 paratypes MNHN.

Type Locality. — Philippines. SSW of Batangas, Luzon, Musorstom 2, stn CP 78, 13°49’N, 120°28'E,

441-510 m.

Distribution. — Philippines, now extended to the Arafura Sea, alive in 352-688 m, shells in 243-809 m.

Remarks. — The protoconch of most specimens is heavily corroded. Fresh shells have a pale fawn perios-

tracum with strong solid hairs, 0.5 mm long, flattened and broadened at base. This species is nearest to Bonellitia

garrardi , from which it differs by its larger size, sharper sculpture, and paucispiral protoconch (Figs 9-10).

B. garrardi has more rounded whorls, and lacks the deep suture and distinct shoulder of B. atopodonta.

Bonellitia garrardi (Petit, 1974)

Figs 11-13

Cancellaria (Merica) nassoides Schepman, 1911: 263, pi. 18, fig. 9 (non von Koenen, 1889).

Admetula garrardi Petit, 1974: 104 (nom. nov. for C. nassoides Schepman). — Garrard 1975: 33, fig. 2(10).

Neadmete nassoides - Habe, 1961: 435, pi. 23, fig. 5.

Bonellitia garrardi - VERHECKEN, 1986: 34, figs 1-2.

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn DW 49, 08°00'S, 132°59'E, 206-210 m

1 dd.

Type Material. — Holotype (16.2 x 10.1 mm) ZMA.

Type Locality. — Indonesia. Near Kai Islands, "Siboga", stn 256, 05°26.6'S, 132°32.5'E, 397 m.

Distribution. — Japan, Indonesia, and Queensland, Australia (VERHECKEN, 1986: 34); Philippines (USNM,

several lots, unpublished).

Remarks. — The present specimen is small (7.4 x 4.6 mm), slightly worn, and has part of the outer lip

broken. Its large, blunt-tipped multispiral protoconch (1 7/8 whorls; maximum diameter 1.0 mm; exposed height

0.6 mm; Figs 12-13) differentiates it from the sympatric B. atopodonta and from B. superstes Finlay, 1930, from

New Zealand, both having a relatively high paucispiral protoconch.

Genus BROCCHINIA Jousseaume, 1887

Brocchinia Jousseaume, 1887: 221. Type species (SD by Sacco, 1894: 68): Valuta mitraeformis Brocchi, 1814 non

Lamarck, 1811 [= Brocchinia tauroparva Sacco, 1894], Pliocene, Italy.

Inglisella Finlay, 1924 [type species (OD): Ptychatractus pukeuriensis Suter, 1917, Miocene, New Zealand]

may be a synonym. Finlay (1924: 501) First placed P. pukeuriensis provisionally in Brocchina [sic], but later

proposed the new genus Inglisella, differing "in the thin shell, different form of growth, discrepant sculpture, much

straighter columella, and rather pronounced posterior notch in the outer lip just at the keel" (FINLAY, 1924: 513).

The Karubar species have a solid shell and a columella with rather strong folds; consequently they are here

placed in Brocchinia.

Brocchinia fischeri (A. Adams, 1860)

Figs 14-18, 22-24

Cancellaria (Merica) fischeri A. Adams, 1860: 411.

ISolutosveltia abyssicola Habe, 1961: 438, pi. 23, fig. 4.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

301

Merica fischeri - A. ADAMS, 1868: 368.

"Cancellaria" fischeri - HABE, 1961: 437.

Inglisella fischeri - Garrard, 1975: 39, figs 4 (8)-(ll).

Cancellaria (Merica) fischeri - Habe, 1985: 10, pi. 2, fig. 4.

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 02, 05°47'S, 132°13'E, 209-240 m, 1 dd. —

Stn DW 14, 05°18'S, 132°38'E, 245-246 m, 1 lv (POLIPI). — Stn DW 15, 05°17'S, 132°41'E, 212-221 m, 1 lv, 2 dd. —

Stn DW 28, 05°31'S, 132°54'E, 448-467 m, 8 dd.

Tanimbar Islands: stn DW 49, 08°00'S, 132°59'E, 206-210 m, 1 lv, 2 dd. — Stn DW 50, 07°59'S, 133°02'E,

184-186 m, 1 lv (POLIPI).

Total 16 specimens, dimensions up to 12.8 x 5.4 mm, with up to 6 teleoconch whorls.

Type Material. — C. fischeri: Not located. Three shells from the Cuming collection, BMNH 1968419,

were considered possible syntypes and figured as such by GARRARD (1975) and Habe (1985). However, the

locality "was written in pencil on the reverse of the board, together with the citation for the original description;

the front of the board also bears the locality Korea. The only original writing on the board is the label on the front

with the name ' Fischeri , A. Ad' and the initials M.C. on the reverse denoting the origin of the specimens with

Hugh Cuming" (K. Way, in lilt, viii-1995). This lot is now in the general collection, BMNH, with the label

"These specimens had been separated as the types; this is unlikely".

S. abyssicola: whereabouts of holotype (7.9 x 4.8 mm) and paratype (6.8 x 4.2 mm) not mentioned by Habe

(1961).

Type Locality. — C. fischeri: "Strait of Corea", 114 m. — 5. abyssicola: Japan, off Kochi Prefecture,

Shikoku, about 150 m.

Distribution. — Korea; formerly recorded from Japan [A. Adams, 1868: 368; and a single specimen each in

KB IN, ZMHU, NMW (1955.158.1906), and BMNH], but such records have not been confirmed recently (Habe,

1985: 10); China Sea, off Pratas Island, 234 m (USNM 284939); Philippines (USNM 288372, 289413; said to be

from 40 and 1017 m !); Borneo, 622 m (USNM 289966); off South and Western Australia, 79-147 m (GARRARD,

1975: 39).

Remarks. — Brocchinia fischeri has remained little known in the literature. It was omitted by LOEBBECKE

(1881-1886), and TRYON (1885: 84) simply mentioned it under the "unfigured and unidentified species of

Cancellaria". It has only been illustrated by Garrard (1975) and Habe (1985). B. fischeri proves to be rather

variable (Figs 14-18), spire angle 36°-41°. Most Karubar specimens as well as 8 specimens in AMS show a

pitted surface at magnification x 30-60, and SEM pictures show a complex surface with irregularly shaped cavities

(Fig. 24). This may represent an intritacalx, an outer shell layer known in Muricidae and several other families,

including Cancellariidae (D'ATTILIO & Radwin, 1971). This microsculpture is practically absent on exposed parts

of the shell, such as the spiral cords, and this observation agrees with the softness of intritacalx layers. The fact

that a similar pitted layer is also present in related species ( B . kaiensis. B. spec. A, B. spec. B) might indicate that

this is indeed an intritacalx with possibly taxonomic value, rather than an accidental corrosion present on the

teleoconch but not on the protoconch. This type of intritacalx structure has not yet been described in

Cancellariidae.

Measurements of intact protoconchs (mean and standard deviation, n = 14): number of whorls, 1.17, 0.24;

maximum diameter, 0.74, 0.08 mm; exposed height, 0.59, 0.10 mm.

The name Solutosveltia abyssicola is based on shells with an aberrant aperture and may well be a synonym of

B. fischeri. A shell from KARUBAR: stn DW 28 (Fig. 17) seems to be intermediate between B. fischeri and

5. abyssicola (Fig. 64). The genus Solutosveltia is based only on the presence of a semidetached aperture, a

character of doubtful taxonomic value.

The elongated form of Brocchinia fischeri is similar to that of B. clenchi Petit, 1986, from the Atlantic, which

has a much weaker sculpture and no pitted shell surface. B. exigua (Smith, 1891) from off southeastern Australia

has a less constricted suture, lacks the pitted shell surface, has 2 spiral cords per whorl, and only one weak

columellar fold.

Source :

302

A. VERHECKEN

Figs 14-18. Brocchinia fischeri (A. Adams), illustration of variability: 14, 12.8 mm, stn DW 28 448-467 m

ioo o'!"1"1’ S'n °W 28; *6' 8 2 mm' stn DW 15' 2I2'221 m; 17 ■ 10 3 mm’ stn DW 28; 18, 9.8 mm. stn DW 02

Fig. 19. — Brocchinia kaiensis sp. nov., holotype, 4.4 mm.

Fig. 20. — Brocchinia spec. A, 9.0 mm, stn DW 15, 212-221 m.

Fig. 21. — Brocchinia spec. B, 5.7 mm, stn DW 49, 206-210 m.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CANCELLARI1DAE FROM THE ARAFURA SEA

303

Brocchinia fischeri belongs to a group of closely related species, occurring both as fossil and Recent in

the Pacific from Japan to New Zealand, and also in the Atlantic including Europe, and which have been classified

under various generic names. Brocchinia was introduced for a European tertiary fossil; Inglisella Finlay, 1924,

Anapepta Finlay, 1930, Gergovia Cossmann, 1899, Microsveltia Iredale, 1925, Solutosveltia Habe, 1961, etc.

have been proposed for Indo-Pacific, Australian and New Zealand species.

FIGS 22-24. — Brocchinia fischeri (A. Adams), stn DW 49, 206-210 m: 22-23, protoconch; 24, detail of

(intritacalx ?) sculpture on last teleoconch whorl. Scale bar: 1 mm.

Figs 25-26. — Brocchinia kaiensis sp. nov., holotype, protoconch. Scale bar: 0.5 mm for both protoconchs.

Brocchinia kaiensis sp. nov.

Figs 19, 25-26

Material examined. — Indonesia. Karubar, Kai Islands: stn DW 28, 05°31'S, 132°54'E, 446-467 m:

1 lv.

Type Material. — Holotype lv, MNHN.

Type Locality. — Indonesia. Off Kai Islands. Karubar, stn DW 28, 05°31'S, 132°54'E, 446-467 m.

Description. — Shell small, globose, spire angle 65°. Protoconch paucispiral, somewhat eroded, with about

1.5 whorls with impressed suture, maximum diameter 0.8 mm, exposed height 0.7 mm (Figs 25-26). Transition

to teleoconch eroded. Teleoconch of about 2 3/4, rapidly expanding whorls. Axial sculpture practically absent on

first whorl; 9 indistinct broad rounded ribs on second and last whorls. Spiral sculpture consisting of 3 spiral bands

on first two whorls, width about 0.1 mm, remaining equal in strength when crossing over the axial ribs,

almost no nodules formed at intersections. Whorls convex, suture lined by a very narrow and shallow canal.

Source

304

A. VERHECKEN

Aperture semicircular; outer lip crenulated, no inner lirae, only broad grooves corresponding to spiral bands on

outer surface of last whorl. Columella straight, short, with a moderately strong fold and a fainter one forming the

rim of the wide siphonal canal. Narrow zone of thin columellar callus; no umbilicus, no siphonal fasciole.

Dimensions: 4.4 x 4.3 mm.

REMARKS. — B. kaiensis also has a pitted shell surface between the spiral bands. The general outline is

intermediate between B. tanimbarensis and B. fischeri. From B. tanimbarensis, B. kaiensis differs by having

distinct spiral bands, also in interspaces between axials, no sutural ramp and practically no nodules at intersection

of axial and spiral sculpture. B. fischeri is much more slender than B. kaiensis, its protoconch is more depressed,

its columellar folds and callus are much stronger, and it has a parietal tooth.

ETYMOLOGY. — Named after the Indonesian archipelago of the Kai Islands.

Brocchinia spec. A

Figs 20, 27-28

Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 15, 05°17'S, 132041'E, 212-221 m, 1 dd.

Description. — Shell high, conical, spire angle 27°, with straight sides and small aperture. Protoconch

paucispiral with 1 1/4 whorl, nucleus relatively large, whorls smooth and rounded (Figs 27-28), maximum

diameter 0.8 mm, exposed height 0.6 mm. Protoconch wider than adapical part of first teleoconch whorl.

Transition to teleoconch clearly marked. Teleoconch with 5 whorls, the First two with 7 weak, broad axial ribs,

subsequent whorls without axial sculpture apart from strongly prosocline growth lines. Spiral sculpture of

3 narrow bands on spire whorls, grouped in abapical two thirds of whorl. Younger whorls flush, suture very

slightly grooved. Base with 4 spiral bands getting weaker abapically. Last whorl with 13 spiral bands. Teleoconch

surface pitted. Aperture rounded, triangular. Outer lip expanded abapically, with 8 strong lirae inside at some

distance from edge. Columella short, inclined adaxially, with 2 strong folds and a weaker one at rim of short

siphonal canal. One parietal tooth. Columellar callus expanded into a solid but thin-edged columellar collar, half

covering a deeply placed umbilical slit.

Dimensions: 9.0 x 3.9 mm.

Remarks. — This single shell is characterised by its elongated, almost perfectly conical shape, differentiating

it from B. fischeri with its rather strongly constricted suture. Already the first teleoconch whorl has the

characteristic flush shape, versus the more rounded whorls of B. fischeri. However, the possibility cannot be

excluded that this shell is only an aberrant form of B. fischeri: it has columellar folds, callus, and parietal tooth

much like it, and the early whorls similarly do have axial sculpture. The loss of axial sculpture might have been

caused by some trauma to the mantle. Protoconch dimensions do not allow a distinction to be made. More material

will be needed to assess whether or not it is a separate species.

Brocchinia spec. B

Figs 21, 31-32

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands : stn DW 49, 08°00’S, 132°59'E 206-210 m

1 lv. ‘

Description. Shell small, elongated, spire angle 53°. Protoconch paucispiral, 1 1/4 smooth whorl,

maximum diameter 0.8 mm, exposed height 0.6 mm (Figs 29-30). Teleoconch with 3.5 whorls, suture impressed

with rather narrow groove. Axial sculpture of broadly rounded ribs, 8, 7, 9 on 1st to 3rd whorl. Spiral sculpture

consisting on first two whorls of 3 cords, packed together in abapical 2/3 of whorl; on 3rd whorl 4 well-defined

spiral cords and a smoothly indicated one in adapical part of whorl; on last half whorl, secondary spirals are

formed between the main cords. Fifteen spirals on last whorl behind outer lip. Teleoconch surface pitted.

Source : MNHN Paris

MOLLUSCA GASTROPODA CANCELLARI1DAE FROM THE ARAFURA SEA

305

Aperture elongated oval, outer lip thin, possibly not fully developed. No umbilicus, no siphonal fasciole. Only a

narrow zone with thin columellar callus. Columella straight, one faint fold and a very faint one forming the rim of

a wide siphonal canal.

Dimensions: 6.1 x 3.5 mm.

Figs 27-28. — Brocchinia spec. A, protoconch.

Figs 29-30. — Brocchinia spec. B, protoconch.

FIGS 31-32. — Brocchinia tanimbarensis sp. nov., protoconch of juvenile shell from stn CP 71, 477-480 m.

Scale bar: 0.5 mm for all figures.

Remarks. — This single shell differs from B. fischeri in its broader spire angle, relatively higher aperture, and

somewhat less convex whorls. The columellar folds of B. fischeri are much stronger and are placed more

transversely. This shell can be considered to be subadult, with incompletely formed aperture, but even then the

difference in strength of columellar folds still holds. In a specimen of B. fischeri (Fig. 18) the columellar folds can

be observed through a boring hole in the last whorl and, although not as bold as in a normally formed aperture,

they are much sharper than in Brocchinia sp. B, where they are very much like those of B. kaiensis. The

protoconch and the sculpture of the teleoconch, however, are very near that of B. fischeri. Therefore, recognition of

this single shell as a different species is not straightforward, and more specimens will be needed to confirm it as a

separate, undescribed species.

Brocchinia tanimbarensis sp. nov.

Figs 31-35

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: sin DW 60, 08°21'S, 132°14’E, 387-389 m,

1 dd (paratype POLIPI). — Stn CP 69. 08°42' S, 131°53' E, 356-368 m, 1 lv (paratype MNHN). — Stn CP 70, 08°41'S,

306

A. VERHECKEN

131°47'E, 410-413 m. 2 lv (paratypes POLIP1). — Sin CP 71. 08°38'S, 131°44'E. 477-480 m. 10 lv, 1 dd (paratypes:

3 POLIPI. 8 MNHN). — Stn CP 77, 08°57'S, 131°27'E, 346-352 m. 1 lv (holotype).

Type Material. — Holotype MNHN. Paratypes: 6 POLIPI, 9 MNHN.

Figs 33-35. — Brocchinia tanimbarensis sp. nov.: 33, holotype, 6.5 mm; 34, 7.4 mm, stn CP 69, 356-368 nr

35. 7.0 mm, stn CP 71, 477-480 m.

FIG. 36. — Merica elegans (Sowerby), 30.4 mm, stn CP 65, 174-176 m.

Fig. 37. — Merica oblonga (Sowerby), 38.6 mm, stn CP 65, 174-176 m.

FIGS 38-39. — Microsveltia karubar sp. nov.: 38, holotype, 9.9 mm; 39, paralype. 7.6 mm, stn CP 71, 477-480 m.

Fig. 40. — Microsveltia metivieri sp. nov., holotype, 6.3 mm.

Source : MNHN , Paris

MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

307

Type LOCALITY. — Indonesia. Off Tanimbar Islands, Karubar, stn CP 77, 08°57'S, 131°27’E, 346-352 m.

DISTRIBUTION. — Known only from off the Tanimbar Islands.

DESCRIPTION. — Shell small, solid, white, with short spire, spire angle 60-80°. Protoconch (strongly eroded

on most shells, preserved only on the smallest one but already with corroded surface, Figs 31-32) paucispiral with

3/4 whorl, surface sculpture etched, maximum diameter 0.9 mm, exposed height 0.6 mm, transition to teleoconch

indistinct. Possibly, a quarter whorl with only smooth sculpture of 2 spiral lines may also be part of the

protoconch. Teleoconch with up to about 3 1/8 whorls. Sculpture starts on the first teleoconch whorl with

3 smoothly indicated spiral lines, undulating to form 8 gradually stronger nodules. Three spirals on spire whorls,

6 on last whorl. Axial sculpture of broad rounded ribs, 10 and 1 1 on second and third whorl, 13 on last whorl.

Shell sculpture prominent at intersection of axials and spirals, forming strong raised rounded nodules. Whorls with

a flat narrow sutural shelf. Aperture semicircular, columella parallel to shell axis, with two rather strong folds, the

weaker abapical fold on the rim of broad short siphonal canal. Outer lip thin, wavy because of nodulose sculpture,

no inner lirae. Thin white columellar callus; no umbilicus nor siphonal fasciole.

Dimensions: holotype 6.5 x 5.5 mm; paratypes up to 7.4 mm.

Remarks. — None of these specimens has a pitted shell surface like B. fischeri, but most of them have spo¬

radic remains of a soft white layer, possibly an intritacalx. The Recent Oamaruia deleta Finlay, 1930, of which

only the juvenile holotype has been figured, reaches 1 1 mm and has 3 distinct columellar folds, the lowest being

stronger (POWELL, 1979: 224). It may be the closest living relative of B. tanimbarensis. Oamaruia major

Marwick, 1965, from the Opoitian (Pliocene, Piacenzan) of New Zealand is also rather close to B. tanimbarensis

but differs by its size up to 11.3 mm, its sculpture with only 2 spirals on spire, its more elongate aperture and

3 columellar folds. The New Zealand Miocene Admete suteri Marshall & Murdoch, 1920, type species (OD) of

Oamaruia Finlay, 1924, has a protoconch of two whorls, "the apex obliquely disposed", 17-21 axials on the last

whorl, and also has 3 columellar folds. Waipaoa marwicki Dell, 1956, from off New Zealand, depth about 600 m,

in general form resembles small specimens of B. tanimbarensis, but lacks the columellar folds and the nodulose

sculpture. B. tanimbarensis is rather different from B. fischeri and the closely related B. exigua from New South

Wales, that share with it the general form of the aperture and columella, but lack the strong nodulose sculpture and

are much more elongate. B. tanimbarensis resembles closely the Atlantic species B. nodosa (Verrill & Smith

in VERRILL, 1885) and B. azorica (Bouchet & Waren, 1985). B. nodosa is much larger (up to 16 mm: MNHN

specimen from Bay of Biscay), and also B. azorica may grow somewhat larger. B. nodosa and B. azorica differ from

B. tanimbarensis in having no distinct sutural ramp, the nodules more pointed, and the columellar and parietal

callus thicker so that the underlying sculpture is barely visible. B. pustulosa Verhecken, 1991a, from off Brazil,

637 m, has a sutural area comparable to that of the new species, grows up to 1 1.6 mm high, and has a much

higher spire.

Etymology. — Named after the Indonesian archipelago of the Tanimbar Islands.

Genus MERIC A H. & A. Adams, 1854

Merica H. & A. Adams, 1854: 277. Type species (SD by COSSMANN, 1899: 13): Cancellaria melanosioma Sowerby, 1849.

North-western Indian Ocean.

The two species of Merica from the Karubar expedition were taken dead at the same station; they may have

been carried down from more shallow depths.

Merica elegans (Sowerby, 1822)

Fig. 36

Cancellaria elegans Sowerby, 1822: fig. 3.

Source :

308

A. VERHECKEN

Cancellaria reeveana Crosse, 1861: 237 (unnecessary name change: VERHECKEN, 1986: 39). — LOEBBECKE, 1881: 12,

pi. 2, figs 1-2, 4-6.

Merica elegans - VERHECKEN, 1986: 40, fig. 9.

Not Cancellaria ( Merica ) elegans - Garrard, 1975: 3, fig. 1(1).

Material EXAMINED. — Indonesia. Karubar, Tanimbar Island: stn CP 65, 09°14'S, 132°27’E, 174-176 m,

1 dd.

Type Material. — A confusion about the "type lot" of this species asks for some explanation. The shell

figured by SOWERBY was "in Mrs. Mawe's collection", but no shell in BMNH has a label indicating that origin.

Garrard (1975: 4) mentioned "Holotype unknown. ... Location of type: BMNH. Not located at present" and

I (VERHECKEN, 1986: 40) myself stated "there is no indication that this specimen is, or should be, in BMNH".

However, PETIT & Harasewych (1986: 442) mention a type lot BMNH 1968387 of 3 specimens, "labeled

C. elegans, Baclayon, Bohol Id., Philippines on the back of an old board", stating that "it is possible that two

specimens with locality data were added later, as only one specimen has an old label with the number '4', the same

sort as used by SOWERBY, glued inside the aperture". About 15 years ago one of these 3 shells was labelled

"Awaiting neotype selection by Petit", a designation which has never been published. Furthermore, SOWERBY's

figure has the number "3", not "4". This material has now been checked again (vi- 1995) and discussed with

K. Way (BMNH). Of the 3 shells, the one (33.6 x 22.7 mm) once intended for neotype selection resembles best

but is broader than SOWERBY's figure (33.5 x 20.8 mm), and shows no traces of a label having been glued inside

the aperture, although there is an old label with the species name in SOWERBY's handwriting (fide K. Way). That

shell now has a label "Although there is no evidence that this specimen reached Cuming via Mrs. Mawe; its very

close similarity to the type figure and Sowerby's own hand on the label must make it eligible to be considered as a

possible type. K. Way, 1986". Since there is no solid proof that this is the figured shell, its designation as

lectotype would pose a problem; and because it has no reliable locality data, it does not qualify for selection as a

neotype. I suggest the best solution would be to select as neotype a well documented shell resembling as closely

as possible this BMNH shell, when such specimen can be found.

Type Locality. — Unknown to SOWERBY (I). Ticao, Philippines, according to G. B. Sowerby (II) (1849:

447), possibly based on 4 shells ex Mrs De Burgh coll. (BMNH).

Distribution. — Philippines; Indonesia.

Remarks. — Within the genus Merica, M. elegans is characterised by its numerous weak prosocline ribs,

only very slightly canaliculate suture, and slightly inflated fusiform outline. The present specimen (30.4 x

18.8 mm) is more slender than Merica melanostoma Sowerby, 1849, and M. subsinensis Loebbecke. 1881. Its

suture is less canaliculate, and its sculpture is dominantly spiral, much more like that of Merica sinensis Reeve,

1856, which however has a strongly oblique protoconch. The species described and illustrated by Garrard [1975:

3, fig. 1(1)] as Cancellaria (Merica) elegans and occurring subtidally down to 49 m off NE Australia (including

Gulf of Carpenteria, near the Arafura Sea) to Queensland, differs markedly from the present material.

Merica oblonga (Sowerby, 1825)

Fig. 37

Cancellaria oblonga Sowerby, 1825: Appendix p. xv. — SOWERBY 1832: fig. 19.

Cancellaria bifasciaia Deshayes, 1830: 181. — Loebbecke, 1885: 30, pi. 9, figs 1-2.

Merica bifasciata - Habe, 1961: 434, pi. 24, fig. 27.

Merica oblonga - Chenu, 1859: 277, fig. 1847. — Petit, 1974: 112, fig. 5. — Verhecken, 1986: 41, figs 7-8.

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 65, 09°14'S, 132°27'E 174-176 m

2 dd (38.6 x 19.2 mm; 35.1 x 17.2 mm).

Type Material. C. oblonga: Sowerby’s shell from the Tankerville auction has not been located (not in

BMNH). — C. bifasciata: holotype (21.4 x 11.8 mm) in MNHN.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

309

TYPE Locality. — C. oblonga: Unknown. KlENER (1841: 6) indicated "l’ocean equinoxial, les cotes de

Panama", which is erroneous (KEEN, 1971: 649). SOWERBY (II) (1849: 447) first mentioned a correct locality:

"Straits of Macassar". — C. bifasciata: Unknown.

Distribution. — Japan to Indonesia; Northern Indian Ocean to Aden; Eastern South Africa (?) (Verhecken,

1986: 41; VERHECKEN & WRANIK, 1991: 60).

REMARKS. — Specimens in collections are usually smaller in size [heights of 43 mm (Loebbecke Museum

und Aquarium, Diisseldorf) and 37 mm (VERHECKEN, 1986: 42) are exceptional] and have a much smoother

sculpture. The coarse sculpture of present specimens resembles that of M erica laddi Petit, 1987 [= Cancellaria

(Merica) petiti Ladd, 1982, non Olsson, 1967], from the Pliocene of Fiji, which is smaller and has a more

acuminate spire.

Genus MICROSVELTIA Iredale, 1925

Microsveltia Iredale, 1925: 265. Type species (OD): Microsvellia recessa Iredale, 1925. Recent, New South Wales,

Australia.

Garrard (1975: 35) considers Microsveltia a synonym of Gergovia Cossmann, 1899, because Cancellaria

platypleura Tate, 1898, type species (OD) of Gergovia "was almost certainly ancestral [to M. recessa], and the

introduction of a new genus was unwarranted". However, COSSMANN originally included Gergovia in Merica as a

new section; and M. recessa has very little resemblance to any species of Merica. Hence, this synonymy is not

accepted here, and Microsveltia is considered valid. FINLAY (1930: 241) considered Microsveltia to be "an absolute

synonym" of Inglisella; but the Brocchinia- like species included in Inglisella do not seem closely related to

M. recessa. Inclusion of the following four species in Microsveltia is only provisional, awaiting a thorough

revision of the cancellariid genera.

Microsveltia karubar sp. nov.

Figs 38-39

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: sin CP 71, 08°38'S, 131°44’E, 477-480 m,

3 Iv.

Type Material. — Holotype lv, MNHN. Paratypes: 1 Iv MNHN, 1 lv POLIPI.

Type Locality. — Indonesia. Off Tanimbar Islands, Karubar, stn CP 71, 08°38'S, 131°44'E, 477-480 m.

DESCRIPTION. — Shell small, with relatively high, conical spire and small aperture, whorls bicarinate, suture

well impressed. Protoconch (severely corroded in all shells) apparently paucispiral and rather high, not flattened.

Teleoconch with up to about 5 whorls. Axial sculpture of rather narrow rounded ribs, constricted near suture,

numbering 7-10, 9-11, 11-13, 11-15 on 2nd to 5th whorl respectively; base without axial sculpture. Spiral

sculpture of narrow cords numbering 2, 3-4, 3-4 on second to fourth whorl, 10-12 on last whorl, including

secondary spirals. Whorls rounded, angular at main spirals, constricted near deeply impressed suture. Aperture

small, oval, ending abapically in short but distinct siphonal canal. Columella short, with one strong fold and a

very weak one at the start of the siphonal canal, which is slightly inclined adaxially. Outer lip crenulate; no lirae

inside. Siphonal fasciole slightly developed. No umbilicus, only a slight, completely closed, depression. Almost

no columellar callus. Thin pale fawn periostracum.

Dimensions: holotype 9.9 x 5.8 mm. Paratypes 7.6 x 4.6 mm (MNHN); 5.6 x 3.6 mm (POLIPI).

Remarks. — The constricted profile of the whorls resembles that of M. metivieri, but the latter species has

much broader ribs. The sculpture somewhat resembles that of Cancellaria patricia Thiele, 1925, from East Africa

(type locality) and south-eastern Australia (Verhecken, 1991b), but that species has much weaker columellar

310

A. VERHECKEN

folds and a shorter spire. M. recessa has a reticulated sculpture much stronger than in M. karubar , moreover it has

two columellar folds of about the same strength, a small and narrow umbilicus, partly covered by callus, and a fine

bristly greenish-brown periostracum (Garrard, 1975: 37).

ETYMOLOGY. — This species is named after the French-Indonesian scientific expedition that discovered it.

Microsveltia metivieri sp. nov.

Figs 40-42

Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 28, 05°31'S, 132°54'E, 448-467 m, I dd.

Type Material. — Holotype dd, MNHN.

Type Locality. — Indonesia. Off Kai Islands, Karubar, stn DW 28, 05°31'S, 132°54'E, 448-467 m.

Description. — Shell small, white, with acuminated spire, spire angle 55°, and strongly constricted suture.

Protoconch (Figs 41-42) white, paucispiral with one whorl, slightly deviating from teleoconch axis, nucleus

relatively large, diameter 0.3 mm, suture slightly impressed. Maximum diameter 0.6 mm, exposed height

0.5 mm. Transition to teleoconch rather indistinct. Teleoconch with 4 3/4 whorls. Axial sculpture consisting of

strong, broad rounded ribs, far less prominent near suture, numbering 8, 8, 9, 11 on 1st to 4th whorl respectively,

12 on last whorl. Growth lines microscopic, hardly visible. Spiral sculpture of well-marked cords, width up to

0.1 mm, numbering 2, 2, 3, 5 on 1st to 4th teleoconch whorl respectively, 11 on last whorl. The spirals keep

their profile when crossing over the axials, without nodular intersection. Spiral cords more closely set in adapical

two-third of whorls, thus giving the impression of a slightly excavated suture, but in younger whorls a much

narrower spiral cord appears just above abapical suture. Last whorl regularly convex; axial sculpture disappearing

towards umbilicus. Aperture rounded, columella straight, inclined adaxially, with two weak folds, anterior one

weaker. Siphonal fasciole well developed, enclosing a narrow umbilicus which is almost closed by a thin

columellar callus. No lirae inside outer lip. Short, broad siphonal canal.

Dimensions: 6.3 x 3.8 mm.

Remarks. The sculpture of strong ribs of M. metivieri superficially resembles the much stronger sculpture

ol Scalptia mercadoi Old, 1968, from the Philippines, but that species has a multispiral protoconch and grows up

to 34 mm (AV) for 6 teleoconch whorls, or 18 mm for 4.5 whorls. M. metivieri is rather close to, but different

Irom M. recessa Iredale, 1925, from New South Wales, Australia, which has bicarinate whorls with nodules on

intersections of coarse spiral and axial sculpture, "microscopic hair-like growth lines", and 8 lirations inside outer

lip (Garrard, 1975: 37).

Etymology. — This species is named in honour of Bernard Metivier (MNHN).

Microsveltia procerula sp. nov.

Figs 43-46

2 ( Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 71, 08°38'S, 131°44'E, 477-480 m.

Type Material. — Holotype lv and 1 paratype lv, MNHN.

Type Locality. — Indonesia. Off Tanimbar Islands, Karubar , stn CP 71, 08°38'S, 131°44'E, 477-480 m.

Description. Shell small, high-spired, thin-walled, younger whorls translucent. Protoconch (Figs 43-44)

paucispiral, bulbous, with about one smooth slightly oblique whorl, suture impressed, maximum diameter

Source : MNHN , Paris

MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

311

0.7 mm, exposed height 0.6 mm. Transition to teleoconch not discernible because of shell damage in that area.

Teleoconch with about 4 whorls. Axial sculpture consists of rather narrow, slightly opisthocline rounded ribs, 12,

1 1, 10 and 1 1 on 1st to 4th whorl respectively. Spiral sculpture of narrow threads, 2, 3, 3 + 1 secondary, on 1st to

3rd whorl, 10 on last whorl, crossing over axials without nodules. Aperture rounded, trapeziform, with short, wide

siphonal canal; outer lip thin, without inner lirae. Columella short, parallel to shell axis. One very weak

columellar fold in holotype (Fig. 45); paratype (Fig. 46) with a relatively strong fold, and a weak one at rim of

siphonal canal which deviates adaxially from the columellar direction. No umbilicus, no columellar callus.

Siphonal fasciole not developed. Shell covered by thin pale-beige periostracum, forming microscopic axial folds

which apparently do not reflect the presence of a similar sculpture on shell surface.

Dimensions: holotype 5.0 x 2.6 mm; paratype 6.0 x 3.0 mm.

REMARKS. — Microsveltia procerula is close to M. cf. sagamiensis, but differs by its thin shell, its less

constricted suture, its sutural area sloping down towards shoulder of whorl, and its stronger columellar folds.

It differs from M. metivieri in its less conical spire, and the strength and form of its axial ribs. M. procerula

closely resembles Cancellaria turriculata Tate, 1889, from the upper Eocene of South Australia, judging from the

original description and illustration (TATE, 1889: 156, pi. X, fig. 14), but conspecificity can be excluded because

of the very large gap in time.

ETYMOLOGY. — The specific name is from the Latin adjective procerulus, diminutive of procerus , meaning

"rather long, elongate", and refers to the slender form of this small shell.

FiGS 41-42. — Microsveltia metivieri sp. nov., holotype, protoconch.

FIGS 43-44. — Microsveltia procerula, holotype, protoconch.

Scale bar: 0.5 mm for all figures.

Source :

312

A. VERHECKEN

l™^5'46 ~ Micr°svel,ia Penile sp. nov.: 45, holotype, 5.0 mm; 46, paratype, 6.0 mm, stn CP 71. 477-480 m.

HG. 47. Microsveltia cf. sagamiensis (Kuroda & Habe), 4.3 mm, stn DW 44, 291-295 m.

Figs 48-50. — Perplicaria boucheti sp. nov.: 48, holotype, 9.0 mm; 49-50, protoconch. Scale bar: 1 mm.

MG. 51. — Solatia arafurensts sp. nov., holotype, 36.0 mm.

Fig. 52. — lAdmete aethiopica Thiele, 5.2 mm, stn CP 71, 477-480 m.

Fig. 53. — Plesiotriton vivus Habe & Okutani, 40.3 mm, stn DW 44, 291-295 m.

Fig. 54. — Tritonoharpa beui sp. nov., holotype, 16.5 mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CANCELLARI1DAE FROM THE ARAFURA SEA

313

Microsveltia cf. sagamiensis (Kuroda & Habe, 1971)

Figs 47, 55-57

Neadmete sagamiensis Kuroda & Habe in KURODA, Habe & Oyama, 1971: 204, pi. 109, fig. 24.

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands : stn DW 44, 07°52'S, 132°48'E, 291-295 m,

1 dd.

Type Material. — The illustrated syntype of Neadmete sagamiensis (6.3 x 3.3 mm) is here designated

lectotype; it is in the collection of the Emperor of Japan, together with at least 3 paralectotypes (not seen).

TYPE Locality. — West of Jogashima, Sagami Bay, Japan, 1 10-150 m.

DISTRIBUTION. — Japan: from Sagami Bay, Honshu and Tosa Bay, Shikoku, sandy bottom, 50-200 m.

Description. — Shell minute, solid, elongated, spire angle 50°, constricted near suture, aperture small.

Protoconch bulbous, paucispiral with about 1 whorl, suture impressed (Figs 55-56). Maximum diameter about

0.6 mm, exposed height 0.5 mm. Remains of a finely granular microsculpture still present (Fig. 57). Transition

to teleoconch not discernible because of shell damage in that area. Teleoconch with about 3.5 shouldered whorls.

Axial sculpture of rounded ribs, narrow on first, broadly rounded on later whorls, 10, 9 and 10 on 1st to

3rd teleoconch whorl, 7 on last whorl. Spiral sculpture of narrow (up to 0.1 mm wide) cords running over the

axial ribs without forming nodules, 2 cords on 1st whorl, 2 + 1 secondary spiral and 1 on shoulder plane on

2nd whorl, 3 + 2 on shoulder plane on 3rd whorl. The zone between suture and the first adapical spiral cord is

wider than the spiral interspaces. Last whorl with 1 1 narrow, well-marked spiral cords. Suture strongly constricted,

sutural area almost horizontal and slightly wavy because of vanishing axial ribs; shoulder of axial ribs produced in

a rounded angle. Aperture small, rounded triangular. Outer lip with thin edge (partly broken) and 5 widely spaced

rather indistinct inner lirae continuing deep inside aperture. Columella straight, with 2 very weak folds deep inside

aperture. Short, broad siphonal canal. Columellar callus narrow and thin, but covering most of the umbilical slit.

Almost no siphonal fasciole. No periostracum.

Dimensions (Karubar specimen): 4.25 x 2.5 mm.

REMARKS. — Petit (1974: 1 1 1) considered Neadmete sagamiensis to be a synonym of Admete cancellata

Kobelt, 1887, described from Japan, and said to have 'narrow high radial ribs', narrower than their interspaces, and

crossed by spirals of about the same strength, forming a neatly quadratic cancellation (KOBELT, 1887a: 12;

1887b : 105). Kobelt's illustration (1887b: pi. 24, fig. 14) is so small that it can hardly be used for

identification, but the sculpture described is very obvious on the type [holotype by monotypy, ICZN art. 73a(ii),

see KOBELT, 1887b; 10.2 x 6.2 mm, ZMHU 101677, Fig. 63]. The dimensions of the holotype and of its

paucispiral protoconch (1 1/8 whorl, max. diameter 0.95 mm, exposed height 0.8 mm) are significantly larger than

in the present material; and the sculpture is quite different, so that identification of the Karubar shell as

A. cancellata is rejected. Both the Karubar shell and the lectotype of Neadmete sagamiensis (according to the

original illustration) have axial ribs broader than their interspaces and much stronger than the spirals, and the

whorls are more shouldered than the regularly rounded whorls of A. cancellata. Therefore, PETIT's identification of

N. sagamiensis as Admete cancellata does not seem completely warranted, but variability within these taxa is as

yet unknown. Since the present specimen is certainly different from the holotype of A. cancellata and closest to

M. sagamiensis, the latter name is used here. The Karubar specimen differs from the lectotype of

N. sagamiensis in having the whorls more shouldered and the aperture more triangular. The protoconch of the

lectotype, according to KURODA & Habe's description and illustration, is larger and has more whorls. Because of

these differences, identification of the present material is only tentative. M. cf. sagamiensis is quite similar to

M. metivieri but differs in having the whorls more shouldered, a wider and flatter sutural area, the axial ribs less

pronounced, the whorls less convex, and the columellar folds much weaker.

314

A. VERHECKEN

55~57- Microsvehia cf. sagamiensis (Kuroda & Habe): 55-56, proloconch, stn DW 44. 291-295 m Scale bar

.5 mm; 57, detail of protoconch sculpture. Scale bar: 0.1

Published illustrations of M. recessa from New South Wales, Australia are somewhat contradictory, drawings

of the holotype (Iredale, 1925: pi. 43, fig. 16; LASERON, 1955: 268, fig. 11) resembling the present shell

except lor the 2 strong columellar folds, but photographs of the holotype [Garrard, 1975: fig. 3(8)1 and other

specimens [Garrard, 1975: fig. 3(7); Ka.cher, 1978: card 1919] showing a very coarse quadratic sculpture.

. recessa has two bold and narrow spiral cords, forming prominent lateral nodules at junction with ribs, and

strong hrat'ons mside the outer lip (Garrard, 1975: 37). In general form and in the sculpture of broad axial ribs

overridden by narrow spiral bands, M. sagamiensis seems also to be close to Cancellaria japonica Smith, 1879

but extensive damage to the latter's holotype (1 1.9 x 5.8 mm, BMNH 1878.1 1.7.90, figured by Petit, 1974: 1 lo’

text-fig. 3) prevents clear conclusions. Subsequent to the original description, no additional specimens of

^japonica have been reported; it could not be recognised as any known Japanese species by Habe and by

Okutani (Petit, 1974: 1 10). Hence, its real relation to M. sagamiensis remains unclear.

Kuroda & Habe (1971) placed sagamiensis in the genus Neadmete Habe, 1961. Its type species (ICZN

Opinion 1370), N. okutanu Petit, 1974, has a dominantly spiral sculpture (see Habe, 1961: pi 23 fig 11

incorrectly identified as Neadmete japonica, later renamed N. okutanii). The genus has been used for several rather

arge, cold-water species from the northeastern Pacific (Kanakoff & McLean, 1966; ABBOTT, 1974- 248) with a

dominT ^ ^ SCUIptUre' Hence' 1 Prefer not t0 use ,his g^us for the present species in which the axials

Genus PERPLICARIA Dali, 1 890

Mi' i89°- piiocene’ Fiorwa' usa-

Miocene, France. ' ’ g' ' Typ sPec,es (by monotypy): Dagumia vigneauxi Magne, 1966.

Source : MNHN Paris

MOLLUSCA GASTROPODA CANCELLAR1IDAE FROM THE ARAFURA SEA

315

Perplicaria has been known only from five fossil species in the Caribbean and France, and one Recent species

(Perplicaria clarki M. Smith, 1947) from West Central America. The genus ranges from the early Miocene

(Perplicaria prior Maury, 1910) to Recent.

Perplicaria boucheti sp. nov.

Figs 48-50

MATERIAL EXAMINED. — Indonesia. KARUBAR, Tanimbar Islands: stn CP 79, 09°16'S, 131°22'E, 239-250 m,

1 dd.

Type Material. — Holotype dd, MNHN.

TYPE Locality. — Indonesia. Tanimbar Islands, Karubar, stn CP 79, 09°16'S, 131°22'E, 239-250 m.

DESCRIPTION. — Shell outline typical for a Perplicaria. Protoconch whitish, paucispiral with one rapidly

expanding whorl (Figs 49-50), maximum diameter 0.9 mm, exposed height 0.8 mm. Surface smooth and shiny,

last 1/16 whorl showing a smoothly indicated start of spiral sculpture. Transition to teleoconch clearly marked by

an expansion of shell width and the beginning of strong teleoconch sculpture. Teleoconch with 2 3/4 whorls,

rapidly expanding in height. On first half whorl only spiral sculpture of 7 low Hat bands. On second whorl

36 gently indicated axial ribs, and 1 1 primary spiral cords, with one narrower second order spiral in between. Last

whorl with 19 broad spiral cords, second and third order spirals in between. On last whorl, axial ribs have

practically disappeared. Axial ribs and growth lines opisthocyrt. Whorls rounded, suture impressed. Inclination of

whorls steeper than inclination of spiral bands. Aperture oblong, 54 % of total shell height, slightly expanded

abapically. Outer lip solid, with narrow and slightly crenulated edge, 15 lirae inside. Columella straight, with

3 folds, adapical one strongest, abapical fold forming rim of short siphonal canal. No umbilicus, only a very

narrow slit is present. Columellar callus thin, almost completely covering the umbilical slit. Colour pale fawn,

inside of aperture white, with an indistinct colabral pale orange band near base of lirae.

Dimensions: 9.0 x 4.5 mm.

REMARKS. — P. boucheti represents the first Perplicaria known from the Eastern hemisphere, and the second

known living species of the genus. Perplicaria clarki differs in having a multispiral protoconch (as ascertained

based on a shell from Gobernadora Island, Panama, AMNH 253840), and adult size reaching 33 mm (Keen, 1971 :

656). Based on analogy with the other species of this genus, the holotype of P. boucheti may not be fully grown.

ETYMOLOGY. — This species is named in honour of Philippe BOUCHET (MNHN).

Genus SOLATIA Jousseaume, 1887

Solatia Jousseaume, 1887: 222. Type species (OD): Solatia solat Jousseaume, 1887 (junior subjective synonym of

Solatia piscatoria Gmelin, 1789. See Verhecken, 1988: 665), North-west Africa.

This genus groups mainly European fossil and two Recent species in the eastern Atlantic (VERHECKEN, 1988).

Only Solatia buccinoides (Sowerby, 1832) from tropical West America has also been placed in Solatia (KEEN,

1971: 654), but this placement is questionable.

Solatia arafurensis sp. nov.

Fig. 51

Cancellaria sp. - Wilson. 1994: 175. pi. 37, tig. 19.

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 72, 08°36'S, 1 3 1 °33'E, 676-699 m,

1 lv.

316

A. VERHECKEN

Type Material. — Holotype lv, MNHN.

Type Locality. — Indonesia. Off Tanimbar Islands, Karubar, stn CP 72, 08°36'S, 131°33'E, 676-699 m.

Distribution. — Type locality, and off Port Hedland, north Western Australia (Wilson, 1994; material not

examined).

Description. — Shell large, solid, oblong, strongly sculptured; last whorl elongated. Protoconch missing,

early teleoconch whorls strongly corroded. Teleoconch axial sculpture of strong, broad rounded ribs, interspaces

about as wide as ribs, 13, 15, 16 on spire whorls, 16 on last whorl. Spiral sculpture of broad rounded cords,

smoothly indicated, 4 per whorl, plus one on shoulder of whorl, fused onto nearest cord. Intersection with axials

strongly nodular. Suture impressed, forming a narrow concave sutural area, bordered by strong nodules on shoulder

of ribs and obliquely crossed by strong growth lines. Aperture white, oblong, ending abapically in a well-defined

siphonal canal. Columella straight, parallel to shell axis, two rather strong columellar folds placed near half

height. Columellar callus thin, transparent, covering almost completely narrow umbilical slit. Outer lip slightly

crenulated, no lirae inside.

Dimensions: 36.0 x 20.1 mm.

Remarks. — Except for two recently formed sections of the last whorl, the shell surface is heavily corroded

and chalky. In spite of this, aperture interior is intact and smooth, and the specimen was live-taken. Large parts of

the shell surface were covered by a black layer (about 0.5 mm thick, still partly visible on Fig. 51) with the aspect

of bitumen, which however does not dissolve in acetone or dichloromethane, usual solvents for bituminous

substances. It might be periostracum, or remains of an epibiont. The last section of the last whorl, where shell

dissolution must have been minimal, had very little of this layer: this seems to point out an epibiont rather than

periostracum.

This species is close only to Solatia buccinoides (Sowerby, 1832) from tropical West America, which grows

up to 40 mm, has a brown shell, much stronger spiral sculpture, and sutural area sloping down towards shoulder.

S. buccinoides also has a wider aperture, the outer lip expanded near half height with 1 1 lirae inside, and a posterior

canal. The two columellar folds are stronger than in S. arafurensis. Placement of this new species in Solatia is by

reference to S. buccinoides, since there seems to be no better fitting genus.

This is clearly the species mentioned and illustrated by WILSON (1994), who gives the following information:

"4 cm. North West Shelf. This unidentified species has been trawled recently on the scampi grounds on the outer

edge of the continental shelf off Port Hedland".

Etymology. — Named after the Arafura Sea, where the holotype was collected.

Subfamily ADMETINAE Troschel, 1856

Genus ADMETE Kr0yer in Moller, 1842

Admete Kroyer in Moller, 1842: 88. Type species (by monotypy): Admete crispa Moller, 1842 (= ITritonium viridulum

Fabricius, 1780). North Atlantic.

Species of Admete typically occur at high latitudes or in deep water.

lAdmete aethiopica Thiele, 1925

Figs 52. 58-59

Admete aethiopica Thiele, 1925: 201, pi. 22, fig. 23.

MATERIAL EXAMINED. — Somalia. The type material (see hereafter).

Indonesia. Karubar, Tanimbar Islands: stn CP 71, 08°38'S, 131°44'E, 477-480 m, 1 lv.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CANCELLAR1IDAE FROM THE ARAFURA SEA

317

Type Material. — Lectotype, here designated, the shell figured by Thiele, 3.7 x 2.4 mm. Paralectotypes:

10 specimens, "Valdivia", stn 251, 01°40.6'S, 41°47.1'E, 693 m, and 2 specimens, "Valdivia", stn 256, 01°49’N,

45°29.5'S, 1 134 m. All types are in ZMHU; no registration number. Paralectotype 10 (3.5 x 2.0 mm, lv) is quite

different and resembles the turrid Propebela exarata (Moller, 1842) from Greenland, as illustrated by SNELI &

Stokland (1986: 122, fig. 2).

TYPE Locality. — Eastern Africa. Off Somalia, "Valdivia", stn 251, 01°40.6'S, 41°47.1'E, 693 m.

Distribution. — Off Somalia, 693-1 134 m; now the Arafura Sea, 480 m.

DESCRIPTION. — [KARUBAR specimen) Shell small, thin-walled, semi-translucent, whitish; spire short.

Protoconch smooth, paucispiral, with about 1 1/8 whorl, maximum diameter 0.7 mm, exposed height 0.5 mm

(Figs 58-59). Nucleus relatively large, diameter 0.26 mm. Transition to teleoconch only marked by start of axial

teleoconch sculpture. Teleoconch with 3 slightly inflated whorls, suture impressed, last whorl large, height

4.0 mm (77 % of total shell height). Axial sculpture on 1st to 3rd whorl consisting of respectively 13, 12,

12 smoothly indicated, slightly sigmoid rounded ribs, disappearing near shell base. One rather strong spiral cord

on shoulder, forming nodular intersection with axials, adjacent to somewhat concave spiral depression. Smooth

spiral bands, width 0.1 mm, numbering 6 on 2nd, 14 on last whorl, separated by narrow groove, crowded towards

shell periphery and base. Shoulder area flat, slightly sloping down towards shoulder cord, axials obliquely crossing

over it. Aperture oval, slightly square-cut adapically. Columella straight, almost parallel to shell axis, with 2 very

weak folds near half height; siphonal canal wide and short. Outer lip thin, translucent, no inner lirae. No

umbilicus.

Dimensions: 5.2 x 3.4 mm.

Figs 58-59. — lAdmete aethiopica Thiele, stn CP 71, 477-480 m, protoconch. Scale bar: 0.5 m.

REMARKS. — The lectotype of Admete aethiopica has a dome-shaped paucispiral protoconch of 1 whorl,

maximum diameter 0.7 mm, exposed height 0.6 mm, nucleus large, diameter 0.33 mm. The teleoconch has

2.5 whorls, axial ribs 15 and 16 on 1st and 2nd whorl, 13 on last whorl; 13 spirals on apertural side of last whorl.

Height of aperture 1.9 mm, height of last whorl 3.0 mm. Paralectotypes measure up to 4.4 x 2.7 mm (Thiele

gave 4.25 x 2.5 mm), average 3.8 x 2.3 mm. The double row of nodules near shoulder, as figured by Thiele, is

only clearly present in the five largest shells, and can be even stronger than in the lectotype. It may be a

characteristic of fully grown specimens. The Indonesian specimen, although larger than any of the types, has the

second row only vaguely indicated and has fewer axial ribs than the lectotype. The number of axial ribs in the type

material (excluding paralectotype 10) is 14-16 and 15-16 on the 1st and 2nd teleoconch whorls respectively. The

obtusely angled truncated columella figured by Thiele occurs in most, but not all of the paralectotypes. Also the

318

A. VERHECKEN

strength of the spiral striae is quite variable. Considering all this, and despite the geographic distance, there seems

to be no reason to separate the East African material and the present Indonesian material.

Subsequent to its original description, Admete aethiopica has been mentioned only by PETIT & HARASEWYCH

(1990: 9), stating that Admete aethiopica is "not a cancellariid". This statement was based on a verbal

communication by P. BOUCHET in 1984 (Petit, in litt., June 1995), who considered Thiele's species to be a

juvenile Gymnobela at that time. BOUCHET (in litt., July 1995) now admits that his 1984 opinion was erroneous

and that Admete aethiopica is a cancellariid. Placement of the species in Admete is not certain, but seems the most

appropriate for the time being. THIELE found no radula, and I equally found neither radula nor tubular jaw. Two

perfectly spherical glassy statocysts, diameter 0.1 mm, could be seen by transparency after rehydrating the animal

prior to dissection.

Figs 60-64. — Japanese cancellariids. 60-62, Axelella nodosivaricosa (Petuch): 60, 15.9 mm, Yaku Island, 130 m;

61-62, protoconch, scale bar: 1 mm. — 63, Admete cancellata Kobelt, holotype (ZMHU 101677), 10.2 mm. —

64, Solutosveltia abyssicola Habe, enlarged reproduction of Habe, 1961: pi. 23, fig. 4.

Subfamily PLESIOTRITONINAE Beu & Maxwell, 1987

Genus PLESIOTRITON Fischer, 1884

Plesiotriton Fischer, 1884: 654. Type species (OD): Cancellaria volutella Lamarck, 1803. Eocene, France.

Range of the genus: Upper Cretaceous to Recent. Only two Recent species are known: the Indo-Pacific

P. vivus Habe & Okutani, 1981, and the Pacific P. mirabilis Beu & Maxwell, 1987.

Plesiotriton vivus Habe & Okutani, 1981

Fig. 53

Plesiotriton vivus Habe & Okutani, 1981: 144, figs 2-3. — Beu & Maxwell, 1987: 28, fig 2D pi 4 f i m

Ptsanella viva - Springsteen & Leobrera, 1986: 98, pi. 18, fig. 25.

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn DW 44. 07°52'S, 132°48'E, 291-295 m.

Source : MNHN Paris

MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

319

Type Material. — Holotype (39.0 x 15.2 mm), NSMT-Mo 58602. Two paratypes NSMT-Mo 58603-4.

Type Locality. — Philippines. Off Panglao, Bohol Island, deep water.

Distribution. — Philippines, Indonesia, Zanzibar (Beu & Maxwell, 1987: 28).

Remarks. — The placement of this species in Plesiotriton was maintained with doubt by Beu & Maxwell,

who figured the protoconch (1987: 10, fig. 2D).

Genus TRITONOHARPA Dali, 1908

Tritonoharpa Dali, 1908: 319. Type species (OD): Tritonoliarpa vexillata Dali, 1908. Recent. Panamic Western America.

Synonyms: Nivitriton Iredale, 1929, Esbelta Sarasua, 1975 (fide Beu & Maxwell, 1987: 33).

Tritonoharpa differs from Plesiotriton in lacking the prominent columellar folds and (as far as presently known)

a radula. Beu & Maxwell (1987: 33) include 19 Recent species, most of them from the tropical Indo-West Pacific.

Six species of Tritonoharpa are currently known from Indonesia (Bf.u & Maxwell 1987), three of them still

unnamed.

Tritonoharpa beui sp. nov.

Fig. 54

Tritonoharpa n. sp.?C, aff. T. angasi (Brazier) - BEU & MAXWELL 1987: 35, pis 12i-j, 1-p.

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 30, 05°39'S, 132°56'E, 111-118 m, 1 dd

(holotype).

TYPE Material. — Holotype dd, MNHN. Paratype 22.4 x 7.4 mm, WAM 3043.83: Mariel King Memorial

Expedition, Tanimbar Islands: stn TSIII: 1-7, 6 miles west of Labuan Olendir, Selaru. 08°07'S, 130°51'E, 25-vi-1970,

46-64 m; (Beu & Maxwell, 1987: 35; not seen).

Type Locality. — Indonesia. Off Kai Islands, Karubar, stn DW 30, 05°39'S, 132°56'E, 1 1 1-1 18 m.

Distribution. — Kai and Tanimbar Islands, southern Moluccas, Indonesia.

DESCRIPTION. — Shell elongate, axial sculpture slightly more prominent than spiral sculpture; aperture long

(45% of total shell length) and rather narrow. Protoconch relatively large, slightly deviating from teleoconch axis,

paucispiral with 1.5 swollen smooth whorls, not planorboid; maximum diameter 1.2 mm, exposed height

1.0 mm. Transition to teleoconch shell clearly marked by sudden start of teleoconch spiral sculpture. Teleoconch

with 5.5 weakly convex whorls. Axial sculpture of narrow, low, opisthocline non-collabral ribs, 16, 17, 18, 25 on

1st to 4th teleoconch whorl, 25 on last whorl. Varices formed at irregular intervals (150°-360°) on spire whorls,

spaced about 240° on adult whorls, not parallel to axial ribs, raised rather high above suture, which is coronated by

axial ribs of the subsequent whorl. Varices reflected and concave on abapertural face. On base, fine growth lines

present between axials. Spiral sculpture of flattened narrow cords, 0.1 mm wide, 8, 8, 8, 10, 10 on 1st to

5th whorl, 34 on last whorl. Three to 6 microscopic spiral lines in spiral interspaces. Spirals form tiny nodules

when crossing axial ribs. Aperture forming a rounded elongated parallellogram, ending adapically in a small sinus.

Outer lip smooth inside, edge crenulated with 16 small nodules, bordered at a distance of 0.5 mm by a strong

varix. The same arrangement can also be seen on older varices, where further shell growth occurred not at the

curled-up edge of outer lip, but somewhat inside it. Inner lip erect, extended into a well developed columellar

collar. Columella almost parallel to shell axis, slightly swollen near centre but without a distinct fold. Siphonal

canal well developed, open and strongly twisted dorsally. Umbilical chink partly hidden by anterior end of

columellar collar. Background colour pale beige; a large brown blotch at mid-distance between two varices, and a

Source :

320

A. VERHECKEN

narrow brown band extending over the width of 3 spiral cords on varices, brown elements paler and indistinct on

apical whorls.

Dimensions (holotype): 16.5 x 6.0 mm.

Remarks. — A photograph of the Karubar specimen was submitted to A. BEU (NZGS), who advised that

its presumed identity with Tritonoharpa n. sp. ?C, aff. T. angasi appears to be correct. Beu & MAXWELL (1987:

35) state that it "lacks the interstitial cords of T. angasi". This refers to the secondary spiral cords (BEU, in litt.)

and these are indeed lacking on the holotype. The only difference between holotype and paratype is the slightly

greater inflation of the whorls of the latter. The protoconch of the paratype was figured by Beu & Maxwell

(1987: pi. 12, fig. o).

Distinction of species within Tritonoharpa is not easy. T. beui differs from some of its congeners by its nearly

straight-sided whorls and general shell outline, from others by its microscopic spiral sculpture or by the

protoconch retained in adult specimens (several species are normally decollate at this size). For further

differentiation from other species of Tritonoharpa , see BEU & Maxwell (1987: 35).

etymology. — This species is named in honour of A. G. Beu (NZGS), senior author of the important study

on Plesiotritoninae where this species was figured for the first time and was recognised as a probably new species.

DISCUSSION

The present collection is remarkable in the number of new records and new taxa collected in the somewhat

restricted geographical area covered by the expedition. From the Arafura Sea, only 7 cancellariids had been recorded

in the literature and/or are represented in museum collections: the holotype of Cancellaria nassoides Schepman,

1911 [= Bonellitia garrardi ] from the Kai Islands (ZMA); a specimen of Trigonostoma bicolor (Hinds, 1 843) from

"Samarang" stn 258, Tual, Kai Islands, 22 m (ZMA); a damaged specimen of T. antiquatum (Hinds, 1843) from

approximately 100 miles North of Croker Island, Arafura Sea, 09°30' S, 132°34' E, 124 m (AMS); a specimen of

Neadmete okutanii (identification by T. Garrard) from off W. Aru Island, 54-65 m (USNM 747371); and three

species of Tritonoharpa: T. pseudangasi Beu & Maxwell, T. brunnea Beu & Maxwell and T. aff. angasi, together

represented by 5 specimens. The present material adds 18 additional species, only 3 of which are here well

represented in number of specimens. The total number of cancellariid species known to occur in Indonesian seas is

now 41 (VERHECKEN, 1986; Beu & MAXWELL, 1987; the present study). This compares favourably to other,

much more thoroughly studied, (Indo-) Pacific areas: Australia 48, Philippines 16, Japan 36 species. Among the

new species, some appear to have their closest relative in west central America and the Atlantic Ocean. Faunal

affinities between Plio-Pleistocene molluscs from the latter areas and from western Pacific islands had already been

reported (Ladd, 1982: 19-20).

Another result of the present study is the presence of a shell layer, possibly an intritacalx, which has been

noted in some, but not all species here placed in Brocchinia. Surface layers of this type had not yet been described

in Cancellariidae. Further work will be necessary to evaluate the significance of this character at genus level.

ACKNOWLEDGEMENTS

The material reported in this paper was collected by P. BOUCHET, W. KASTORO and B. Metivier, between

23 October and 3 November 1991, working on board the R/V "Baruna Jaya 1 " during the Karubar expedition.

Thanks are due to P. BOUCHET (MNHN) for making this material available for study. J. ClLLlS (KB IN) made the

SEM-photographs. A. G. Beu (NZGS), R. Kilias (ZMHU), R. G. Moolenbeek (ZMA), R. E. Petit (North

Myrtle Beach, South Carolina, USA) and K. M. Way (BMNH) provided helpful information or sent material on

loan. T. Backeuau (KB IN) read a draft of the manuscript.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

321

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Kaicher, S. D., 1978. — Card Catalogue of world-wide shells. Pack 19, Cancellariidae. St Petersburg, Florida.

Kanakoff, G. P.. & McLean, J. H., 1966. — Recognition of the cancellariid genus Neadmete Habe, 1961 in the West

American fauna, with description of a new species from the Lomita Marl of Los Angeles County, California.

Contributions in Science, Los Angeles County Museum of Natural History, 117: 1-6.

Keen, A. M., 1971. — Sea Shells of tropical West America. Stanford. 2nd ed., xi + 624 pp.

KlENER, L. C., 1841. — Species general et Iconographie des Coquilles vivantes. Genre Cancellaire, 44 pp., 9 pis. Paris.

KOBELT, W„ 1887a. — Eine neue Admete. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 19: 12.

KOBELT, W., 1887b. — Genus Admete Kroyer. Systematisches Conchylien-Cabinet, 2nd ed., Band 4, Abt. 4: 97-108,

pi. 24. Niirnberg.

Kuroda, T., Habe, T. & Oyama, K.,1971. — Sea Shells of Sagami Bay. Tokyo, xvi + 741 + 489 + 51 pp., 121 pis.

Ladd, H. S., 1982. — Cenozoic fossil mollusks from western Pacific islands; Gastropods (Eulimidae and Volutidae

through Terebridae). United States Geological Survey Professional Paper, 1171: 1-100, pis 1-41.

Laseron, C. F., 1955. — The New South Wales Cancellariidae. Records of the Australian Museum, 23: 267-272.

Loebbecke, T., 1881-86 [in 1881-1887], — Das Genus Cancellaria. Systematisches Conchylien-Cabinet, 2nd ed.,

Band 4, Abt. 4: 1-96, pis 1-23. [Part 309: 1-16, pis 1-5 (1881); part 335: 17-32, pis 6-10 (1885); part 340: 33-56,

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MaGNE, A., 1966. — Daguinia vigneauxi n. gen., n, sp. Journal de Conchyliologie, 105: 127-128.

MOLLER, H. P. C., 1842. — Index Molluscorum Groenlandiae. Naturhistorisk Tidsskrift, 4: 76-97.

Petit, R. E., 1974. — Notes on Japanese Cancellariidae. Venus , 33 (3): 109-1 15.

Petit, R. E„ 1987. — New names for two species of Cancellariidae. The Nautilus, 101 (3): 154.

Petit, R. E., & Harasewych, M. G., 1986. — New Philippine Cancellariidae (Gastropoda: Cancellariacea), with notes on

the fine structure and function of the nematoglossan radula. The Veliger, 28 (4): 436-443.

Petit, R. E. & Harasewych, M. G., 1990. — Catalogue of the superfamily Cancellarioidea Forbes and Hanley, 1851

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Petuch, E„ 1979. — Twelve new Indo-Pacific gastropods. Nemouria, 23: 1-21.

Powell, A. W. B., 1979. — New Zealand Mollusca. Collins, Auckland, xiii + 500 pp., pis 1-82.

Sacco, F„ 1894. — / Molluschi dei Terreni Terziarii del Piemonte e della Liguria. Pt, 16, Cancellariidae, 78 pp., 3 pis.

Torino.

Schepman, M. M., 1911. — Prosobranchs of the Siboga Expedition, Part 4. Siboga-Expeditie, 49 (1): 247-363,

pis 18-24.

Sneli, J.-A. & Stokland, O., 1986. — On the taxonomical status of Tritonium viridulum Fabricius, 1780 (Gastropoda:

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Sowerby, G. B. (II), 1825. — A Catalogue of the Shells contained in the collection of the late Earl of Tankerville.

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Sowerby, G. B. (II), 1832-33. — Cancellaria. In: The Conchological Illustrations, parts 9-13. 5 pis, lextpages

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MOLLUSCA GASTROPODA CANCELLARIIDAE FROM THE ARAFURA SEA

323

Thiele, J., 1925. — Gastropoda der Deutschen Tiefsee-Expedition. II Teil. Wissenschaftliche Ergebnisse der Deutschen

Tiefsee-Expedition auf dem Dampfer "Valdivia" 1898-1899, 17 (2): 35-382, pis 13-46.

Tryon, G. W., 1885. — Family Cancellariidae. Manual of Conchology, 7: 65-98, pis 1-7. Philadelphia.

Verduin, A., 1984. — On the taxonomy of some recent European marine species of the genus Cingula s. 1. (Gastropoda:

Prosobranchia). Basteria, 48: 37-87.

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Verhecken, A., 1988. — Notes sur la nomenclature, la taxonomie et la biometrie de Solatia piscatoria (Gmelin, 1791)

(Gasteropodes, Cancellariidae). Bulletin du Museum national d'Histoire naturelle, ser. 4, 10 (A, 4): 661-673.

Verhecken, A., 1991a. — Description of two new species of bathyal Cancellariidae (Mollusca, Gastropoda) from off

Brazil. Bulletin du Museum national d'Histoire naturelle, ser. 4, 12 (A, 3-4): 547-553.

Verhecken, A., 1991b. — Occurrence of Cancellaria patricia Thiele off South-east Australia; with notes on three

Australian taxa of Cancellariidae (Neogastropoda: Cancellarioidea). Journal of the Malacological Society of Australia,

12: 69-76.

Verhecken, A. & Wranik, W„ 1991. — Additional data on the Cancellariidae of the Gulf of Aden. Gloria Maris, 30 (4):

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Kallaroo.

Source : MNHN, Paris

SUL VTS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS D

Mollusca Gastropoda:

turrid gastropods (Conoidea)

New deep-water

from eastern Indonesia

Alexander SYSOEV

Zoological Museum, Moscow State University.

Hertzen st. 6, Moscow 113009, Russia

ABSTRACT

Nineteen new species are described from the bathyal zone of the Arafura Sea at depths between 146 and 1084 m. The

genus Lusitanops is recorded for the first time from the Indo-Pacific and Clinura vitrea sp. nov, is the first Recent

representative of this hitherto Cenozoic fossil genus. Based on shell and radula morphology, the classification of

Heteroturris in the Clathurellinae is confirmed. Including new species described here, there are now 92 turrid species

recorded from Indonesia at depths greater than 200 m.

RESUME

Mollusca Gastropoda : Nouveaux Turridae bathyaux (Conoidea) de Test de l'Indonesie.

Dix-neuf esp&ces nouvelles de Turridae sensu lato (= Turridae s. s. + Conidae) sont decrites de la mer d' Arafura, il des

profondeurs comprises entre 146 et 1084 m. Le genre Lusitanops est signale pour la premiere fois de l'Indo-Pacifique, et

Clinura vitrea sp. nov. represente la premiere occurrence dans les faunes modernes de ce genre, jusqu'ici connu comme

fossile du Cenozoique. L'attribution du genre Heteroturris & la sous-famille Clathurellinae est confirmee par la

morphologie de sa radula. Certaines esp&ces nouvelles atteignent de grandes dimensions (par exemple Comitas rex

sp. nov., 87 mm el Nihonia maxima sp. nov., 128 mm), ce qui suggbre que la faune bathyale de Turridae d'Indon6sie

(92 espfeces actuellement recens6es & des profondeurs sup6rieures it 200 m) est encore loin d'etre inventoriee de fa9on

satisfaisante.

INTRODUCTION

Conoidea, or Turridae sensu lato as they used to be known, represent a significant component of deep-water

gastropod assemblages worldwide. Because of the sheer size of the family, and also because many species are

extremely scarce, Indo-Pacific turrids are poorly known. Despite the rich harvest of turrids obtained there by

the landmark “Siboga" expedition one hundred years ago, the fauna of Indonesian waters is no exception.

Sysoev, A., 1997. — Mollusca Gastropoda: New deep-water turrid gastropods (Conoidea) from eastern Indonesia. In:

A. Crosnier & P. BOUCHET (eds), Resultats des Campagnes Musorstom, Volume 16. Mem. Mus. natn. Hist. not.. 172:

325-355. Paris ISBN: 2-85653-506-2.

Source : MNHN Paris

326

A. SYSOEV

SCHEPMAN (1913) recorded 52 species, most of them new, from water depths exceeding 200 m in the Indonesian

archipelago. The Karubar expedition, which worked in eastern Indonesia during October and November 1991,

obtained a very rich material of Conoidea, comprising over 100 species. Work on this collection is in progress and

the results will greatly extend our knowledge of the deep-water Indonesian fauna. The purpose of the present paper,

which is a preliminary report on this turrid fauna, is to record and describe some of the more spectacular new

species collected during the expedition. Twenty new species are described from 19 deeper- water stations, at depths

between 146 and 1084 m. They belong to 15 genera.

Type material is housed in Musdum national d'Histoire naturelle, Paris (MNHN) and Pusat Penelitian dan

Pengembangan Oseanologi LIPI, Jakarta (POLIPI).

SYSTEMATIC ACCOUNT

Superfamily CONOIDEA Fleming, 1822

Family TURRIDAE H. & A, Adams, 1853

Subfamily TURRINAE H. & A. Adams, 1853

Genus GEMMULA Weinkauff, 1875

Type Species: Pleurotoma gemmata Reeve, 1843 (= Gemmula hindsiana Berry, 1958) [Non Pleurotoma

gemmata Conrad, 1835].

Gemmula closterion sp. nov.

Figs 11-12, 16

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn DW 61, 09°05'S. 132°44'E, 235-236 m.

1 dd (paratype MNHN). — Stn CP 67, 08°58'S, 132°06'E, 146-233 m, 1 Iv (holotype), 2 dd (paratypes MNHN and

POLIPI). — Stn CP 79, 09°16'S, 131°22'E, 239-250 m, 2 lv (paratypes MNHN and POLIPI).

Type Material. — Holotype and 3 paratypes MNHN, 2 paratypes POLIPI.

DIAGNOSIS. — Shell small, up to 22 mm high, very narrow, spire height about 40% of shell height,

periostracum rather thick, light-brown. Protoconch non-planktotrophic, of 1.5 whorls, smooth except for a few

arcuate axial ribs near transition to teleoconch. Teleoconch whorls with strong bicarinate subsutural fold, narrow

and concave subsutural ramp, and prominent peripheral keel with rounded gemmae. Suture deeply channeled. Base

with three very strong spiral cords separated by wide and usually smooth interspaces. Canal long and straight. Anal

sinus deep, wide, U-shaped.

Description (holotype). — Shell, slender, very narrow (diameter/height 0.27), spire height comprising 40% of

shell height. Color white under rather thick light-brown periostracum. Protoconch consisting of 1.5 smooth

whorls, diameter is 0.85 mm, with 5 narrow arcuate axial ribs near transition to teleoconch. Teleoconch consisting

of 8.75 whorls with a narrow, concave subsutural ramp, a strong, gemmate peripheral keel and another strong cord

between keel and abapical suture. Keel bearing 16 rounded and slightly longitudinally elongate gemmae on last two

whorls, bordered by 2 narrow cords on either side. Subsutural fold strong, bicarinate, with 1 wavy cord running

along its edge and 1 more prominent, weakly granulate cord below it. Suture deeply channeled and covered by edge

of subsutural fold. Base with 3 very strong cords separated by wide smooth interspaces, 3 weaker and rather widely

spaced cords near base of canal, which is covered by 15 weak cords. Aperture small, ovate, inner lip with straight

columellar side covered by white callus. Outer lip chipped, but growth lines define rather deep and broad, U-shaped,

peripheral anal sinus.

Source : MNHN , Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA 327

FIGS 1-8. — Radulae. 1, Borsonia jay a, paratype. — 2, Heteroturris gemmuloides, paratype. — 3. Gymnobela ioessa,

paratype. — 4, Gymnobela muricata , paratype. — 5, Gymnobela milrodeta, holotype. — 6, Gymnobela baruna,

holotype. — 7, Clinura vitrea, holotype. — 8, Xanthodaphne cladara, paratype. Scale bar 50 pm.

Dimensions: height 18.0 mm, last whorl height 10.8 mm, aperture height 8.2 mm, shell diameter 4.8 mm.

Largest paratype 22.2 x 5.7 mm. Diameter/height ratio of intact shells 0.26-0.30, mean 0.28.

Paratypes very similar to holotype, differing only in details of spiral sculpture. Subsutural fold on last whorl

occasionally with 1 additional thin thread in its lower part, cords below keel may number up to 4, and two

paratypes have a thin cord in 1 interspace between main cords on shell base. Also, in some paratypes, the

subsutural ramp may have numerous, rather strong, oblique growth lines.

REMARKS. — Gemmula closterion is one of the smallest species in the genus. I cannot ascertain that the

specimens examined are adult, but this is likely since shells from 3 different stations are of about the same size.

328

A. SYSOEV

Even if the type specimens are subadult, the species is nevertheless easily recognizable by its paucispiral

protoconch, a character so far unique in Gemmula s. str., which indicates that the species has non-planktotrophic

development (probably lecithotrophic). The most similar species is G. graeffei (Weinkauff, 1875) (- G. hombroni

Hedley, 1922), which differs in having a uniformly brown shell, a non-channeled suture, spiral ribs on the entire

subsutural ramp area, and a typical multispiral protoconch. A similar, paucispiral protoconch occurs in the

subgenus Kuroshioturris Shuto, 1961, which differs from the nominotypical subgenus by numerous conchological

characters.

Etymology. — From the Greek kloster , spindle; klosterion is a diminutive. It is treated as a noun in

apposition.

Subfamily COCHLESPIRINAE Powell, 1942

Genus COMITAS Finlay, 1926

Type SPECIES: Surcula oamarutica Suter. 1917 (= Drillia fusifomiis Hutton, 1877).

Comitas rex sp. nov.

Figs 19-20

Material EXAMINED. — Indonesia. KaRUBAR, Tanimbar Islands : stn CP 84. 09°23'S, 13I°09'E, 246-275 m,

i dd (holotype).

Type Material. — Holotype MNHN.

DIAGNOSIS. — Shell large, 87 mm high, fusiform, spire height comprising 40% of shell height, with long

and broad canal. Color of periostracum light brown, with abapically to whorl periphery a band of dark reddish-

brown rectangular blotches on axial ribs. Whorls strongly angulate, with wide concave subsutural ramp covered by

very fine spiral threads. Suture shallow. Axial ribs very strong, widely spaced, oblique, gradually weakening

towards base ol last whorl. Spiral sculpture of strong, rounded and widely spaced cords, occasionally with

additional secondary cord in interspaces. Interspaces covered by 2-4 fine spiral threads. Cords equally strong on

axial ribs and between them. Anal sinus deep, wide and rounded, deepest point just below middle of subsutural

ramp. Outer lip projecting forward below sinus.

Description (holotype). — Shell solid, elongate fusiform, with a high turreted spire consisting of 13.5

whorls strongly angulated somewhat above periphery, with a wide, concave subsutural ramp. Protoconch and first

2-3 teleoconch whorls eroded, but protoconch probably paucispiral. Suture well defined, shallow, straight. Axial

ribs very strong, rather sharp, widely spaced, opisthocline, produced adapically without forming a knob, entirely

traversing spire whorls, gradually weakening abapically, abruptly stopping on shell base; 10 on penultimate whorl

and 8 on last whorl. Subsutural ramp covered by very fine and closely set spiral threads. Spiral cords below ramp

coarse, rounded, equally strong on axial ribs and between them; interspaces broader than cord, with 2-4 coarse

threads and occasionally an additional secondary cord. Base almost flat, smoothly continuous with canal. Canal

long, proportionally very broad, and slightly curved adaxially. Inner lip of aperture weakly and rather evenly

concave, with thin, longitudinally rugose callus. Outer lip projecting moderately forward below sinus. Anal sinus

deep, wide and rounded, deepest point just below middle of subsutural ramp. Shell covered by thin, firmly attached

light brown periostracum, with abapically to whorl periphery a reddish-brown band forming distinctly darker

rectangular blotches on axial ribs, most clearly visible on wet shell.

Dimensions: height 87.1 mm, last whorl height 52.3 mm, aperture height 42.7 mm, diameter 27.5 mm.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CONOrDEA FROM EASTERN INDONESIA

329

Remarks. — The sculpture and general colour pattern resemble those of C. ilariae Bozzetti, 1991, but

Comitas rex clearly differs by having angulate whorls with stronger axial ribs and concave subsutural ramp, and

much broader canal. Another similar species is C. kaderlyi (Lischke, 1872), which, however, is characterized by

more numerous and weaker axial ribs and finer spiral sculpture.

Etymology. — From the Latin rex, king, with reference to the large size of the species. It is treated as a

noun in apposition.

Genus NIHONIA MacNeil, 1961

Type SPECIES: Nihonia shimajiriensis MacNeil, 1961.

Nihonia maxima sp. nov.

Figs 13-15

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: sin CP 84, 09°23'S, 131°09'E, 246-275 m,

1 dd (holotype).

Type Material. — Holotype MNHN.

Diagnosis. — Shell large for genus, over 120 mm high, slender, fusiform, spire rather high, comprising 35%

of shell height. Suture shallow. Spiral sculpture of narrow, granular cords in the slightly concave substural ramp,

and rather weak and widely spaced primary cords below the shoulder. Interspaces with thinner secondary cords and

thin, wavy, and closely set tertiary ones. Aperture narrow, canal long and straight. Anal sinus deep, asymmetrical,

outer lip strongly projecting forward below sinus. Colour yellowish-white, with reddish-brown primary spiral

cords.

DESCRIPTION (holotype). — Shell solid, slender, fusiform, with relatively high spire comprising 35% of shell

height. Protoconch and tip of teleoconch missing. Teleoconch consisting of 10.5 whorls, adapical 3 whorls

styloid, with very slow increase in diameter. Suture tightly adpressed, sometimes hardly distinguishable, clearly

lined on spire whorls by a narrow, strongly granular subsutural cord, which becomes rather obsolete on last

2 whorls. Subsutural ramp concave, especially on juvenile whorls, less so on adult whorls. Spiral sculpture of

spire whorls consisting of thin cords on the subsutural ramp, a broader cord bordering abapically the subsutural

ramp, and 3 (later 2) strong cords below whorl periphery. On subsequent whorls sculpture becoming more

complex. Cords in subsutural ramp fading out until only three indistinct ones remain in the middle of subsutural

ramp of last adult whorl. Secondary and wavy, closely set tertiary spiral cords appearing gradually in interspaces

between main cords. On periphery and base of last adult whorl, sculpture consisting of 4 single and 2 twinned

flattened, rather weak primary cords. No axial sculpture except numerous strong incremental lines intersecting

spiral cords, spiral cords sometimes interrupted at intersections, especially on canal. Aperture rather narrow, ovate,

smoothly continuous with long, straight canal. Inner lip covered by thin callus. Outer lip thin, strongly projecting

forward below anal sinus. Sinus deep, asymmetrical, deepest part in abapical half of subsutural ramp. Colour

yellowish-white, with reddish-brown primary spiral cords.

Dimensions: height 128.2 mm, last whorl height 83.8 mm, aperture height 71.2 mm, diameter 32.1 mm.

Remarks. — In whorl profile and type of sculpture, Nihonia maxima is most similar to the type species of

the genus, N. shimajiriensis, from the Pliocene of Okinawa. However, N. shimajiriensis is much smaller

(the incomplete holotype is 26 mm high at about 9 whorls, including 1.5 whorls of the protoconch, but the last

whorl is partly broken), lacks the styloid apical whorls with slowly increasing diameter present in N. maxima, and

has strongly angulate whorls bearing more pronounced cords. N. maxima can be easily distinguished from the two

330

A. SYSOEV

common Recent species of the genus, N. australis (Roissy, 1805) and N. mirabilis (Sowerby, 1914), by its

complex and much weaker spiral sculpture.

ETYMOLOGY. — From the Latin maximus, largest, with reference to the very large adult size.

Genus CLAVOSURCULA Schepman, 1913

Type Species: Clavosurcula sibogae Schepman, 1913.

Clavosurcula schepmani sp. nov.

Figs 17-18

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn CC 21, 05°14'S, 133°00'E, 688-694 m, 1 lv

(holotype).

Tanimbar Islands: stn CP 69, 08°42'S, 131°53'E, 356-368 m, 1 dd (paratype).

Type Material. — Holotype and paratype MNHN.

DIAGNOSIS. — Shell biconical with cyrtoconoid spire. Protoconch of 1.5 smooth light-brown whorls,

indicating non-planktotrophic development. First teleoconch whorls angulate, with subsutural row of tubercles,

and strong, oblique, axial knobs at periphery. Last whorls with abapical portion of suture at level of undulating

peripheral keel, leaving only subsutural ramp exposed. Base of last whorl with broad Battened spiral cords separated

by narrow grooves.

Description (holotype). — Shell thin, biconical, spire cyrtoconoid. Protoconch light brown, teleoconch

white. Protoconch of 1.5 evenly convex whorls, diameter 850 pm. Teleoconch of 7.3 rapidly expanding whorls.

First 3 teleoconch whorls with broad, smooth, concave ramp with a subsutural row of pointed tubercles, and

strong peripheral keel overhanging abapical part of whorl, sculptured by broad, opisthocline axial ribs, forming

pointed knobs on periphery, and extending to abapical suture, crossed by rather indistinct, narrow, spiral cords.

After third whorl, exposed whorl height abapically of peripheral keel becomes covered by successive whorl,

gradually leaving only concave ramp exposed. On subadult and adult whorls, subsutural row of tubercles coalesced

to a wavy subsutural fold, and adapical margin of peripheral keel forming spirally striated suprasutural undulating

fold, suture impressed, undulating. Subsutural ramp broad, rather flat, with weak indistinct spiral cords, on

penultimate whorl sculptured in median area by two more distinct cords. Base of last whorl sculptured by 35 rather

strong, wide, Battened cords separated by narrow interspaces, interspaces broader on canal. Base slightly convex,

smoothly connecting to a long and straight canal. Aperture narrow, subrectangular. Inner lip covered by very thin

callus. Anal sinus deep, with rounded outline, deepest point above middle of subsutural ramp. Outer lip chipped

but, judging from growth lines, greatly projecting forward below sinus.

Dimensions: height 25.5 mm, diameter 12.2 mm, last whorl height 20.0 mm, aperture height 17.9 mm.

Remarks. — The paratype measures 36.0 x 16.0 mm for 8.5 whorls. The early whorls are decorticated, but

probably only part of the protoconch is missing. Spire whorls have the same sculpture as on the holotype In the

last two whorls there is a distinct spiral striation between abapical edge of sinus zone and peripheral keel, which is

straight, not undulating.

Clavosurcula schepmani is very similar to the type and only species of the genus, C. sibogae , from the

Mores Sea in 794 m. The latter species is, however, larger (38 mm at 8 whorls; SHUTO's [1970] measurements of

the holotype are erroneous), broader, and without peripheral knobs and subsutural tubercles, even on first

teleoconch whorls.

Etymology^ — The species is named in honor of M. M. Schepman for his pioneering work on the deep-sea

turrids of the Siboga expedition.

Source : MNHN , Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

331

Family CONIDAE Fleming, 1822

Subfamily CLATHURELLINAE H. & A. Adams, 1858

Genus BORSONIA Bellardi, 1839

Type Species: Borsonia prima Bellardi, 1839.

Borsonia jaya sp. nov.

Figs 1, 9, 39-44

Material EXAMINED. — Indonesia. Karubar. Tanimbar Islands: sin CP 72, 08°36'S, 131°33'E, 676-699 m,

1 lv, 1 dd (paratypes MNHN). — Stn CP 87, 08°47'S, 130°49'E, 1017-1024 m. 2 lv (paratypes MNHN), 3 dd (2 paratypes

MNHN and 1 paratype POLIPI). — Stn CP 89, 08°39'S, 131°08'E, 1058-1084 m, 4 lv (holotype and 1 paratype MNHN,

2 paratypes POLIPI), 2 dd (paratypes MNHN). — Stn CP 91, 08°44'S, 131°05'E, 884-891 m, 1 lv (paratype POLIPI), 1 dd

(paratype MNHN).

Type Material. — Holotype and 10 paratypes MNHN, 4 paratypes POLIPI.

DIAGNOSIS. — Shell fusiform, solid, up to 70 mm high,

with high spire, periphery angulate, subsutural ramp concave

in abapical part and devoid of axial sculpture. Axial ribs

oblique, narrow, short, weakening abapically, restricted to

periphery on last adult whorl. Spiral cords rather strong and

closely set, narrower on ramp, covering the whole shell

surface. Aperture pyriform with moderately long canal, inner

lip with low pleat in adapical part of columellar edge. Anal

sinus broad, moderately deep. Shell chalky, covered with dark-

brownish grey periostracum, aperture of adult, live collected

specimens light orange inside. Operculum small, pyriform,

with terminal nucleus. Radular teeth large, straight, with short

rounded basal part.

DESCRIPTION (holotype). — Shell slender, fusiform, solid,

with high spire forming 41.5% of shell height. Protoconch

and early teleoconch whorls corroded. Remaining part of

teleoconch consisting of 8 convex whorls, angulate at

periphery, suture shallow, slightly channeled. Subsutural

ramp weakly convex adapically, concave abapically. Axial ribs

short, opisthocline, narrow, separated by interspaces wider

than ribs, abapically extending from angulation to suture,

evanescent on outer base; 16 ribs on penultimate whorl, 18 on

last whorl, where their vertical extension is restricted to periphery. Spiral cords rather strong, flattened, sometimes

with a narrower cord between primary ones, narrower on subsutural ramp. Interspaces between cords narrow, not

exceeding half of cord width, except on canal and adjacent portion of base, where interspaces exceed cord width.

Last whorl moderately convex, base smoothly continuous with rather long and straight canal. Aperture broad,

pyriform, evenly curved inner lip with broad callus and low pleat in adapical part of columellar edge. Outer lip

evenly curved, projecting forward below anal sinus. Sinus moderately deep, as broad as subsutural ramp, deepest

point in the middle of ramp. Shell chalky, covered with dark-brownish grey periostracum, aperture light

orange inside.

FIGS 9-10. — Opercula.

9, Borsonia jaya , paratype. Scale bar 0.5 mm. —

10, Heieroturris gemmuloides, paratype. Scale

bar 0.25 mm.

Source :

332

A. SYSOEV

Dimensions: height 60.9 mm, last whorl height 35.6 mm, aperture height 28.7 mm, diameter 19.8 mm.

Radula of a paratype (stn CP 87, height 35.5 mm) with large, straight, narrow teeth with rounded basal part,

length 490 pm. Operculum small, pyriform, with slightly curved axis and terminal nucleus.

Remarks. — The paratypes are 35.5 to 70.5 mm high, with some variation in proportions of the last whorl

(e.g. Figs 41-42). Younger specimens (Fig. 43) have a relatively broader body whorl with axial ribs proportionally

higher, extending to shell base. In such small specimens the aperture is greyish-white inside. The importance of

columellar pleat may vary slightly, it is present in all but one specimen. Axial ribs on last whorl number 17-19.

Bor sonia jay a is very similar to B. epigona Martens, 1901 from west of Sumatra, 646-676 m, but the latter is

much smaller (up to 28 mm high), with distinctly angulate shell base, straight axial ribs, very prominent

columellar pleat, and, judging from the original illustration, without spiral sculpture in the subsutural ramp.

B-jaya also shows some similarity to the type-species of Buridrillia Olsson, 1942, Clathrodrillia (Buridrillia)

panarica Olsson, 1942 from the Pliocene of Central America. However, it has been recently shown (Emerson &

McLean, 1992) that Buridrillia has radular teeth of modified wishbone type and therefore belongs to the subfamily

Crassispirinae.

Etymology. — From the Indonesian jaya, meaning large, beautiful.

Genus HETEROTURRIS Powell, 1967

Type Species: Heteroturris sola Powell, 1967.

Remarks. Heteroturris was originally described in the subfamily Turrinae because POWELL (1967)

considered it closely allied to Lophiotoma Casey, 1904. Powell also noticed that the sculpture of the subsutural

ramp resembles that ot Microdrillia Casey, 1903, which he classified in subfamily Clavinae. Taylor et al. (1993)

have included Microdrillia in the “tomopleurid” group of genera of the subfamily Clathurellinae (Conidae). The

style and position of anal sinus in Heteroturris is rather different from the character states of Turrinae, and the

radular morphology of H. gemmuloides confirms the classification of Heteroturris in Clathurellinae.

Heteroturris gemmuloides sp. nov.

Figs 2, 10, 21-23

Material EXAMINED. — Indonesia. KARUBAR, Tanimbar Islands : stn CP 59, 08°20'S 132°11'E 399-405 m

1 I v (paratype MNHN). - Stn CP 69. 08°42’S. 131°53'E. 356-368 m, 1 Iv (holotype), 1 dd (paratype POLIPI)

Philippines. Musorstom 2: stn CP 75, 13°5rN, 120°30’E. 300-330 m. 1 Iv (paratype MNHN).

Type Material. — Holotype and 2 paratypes MNHN, 1 paratype POLIPI.

Diagnosis. — Shell large for the genus, height up to 40 mm, solid, narrowly fusiform, with high spire and

long canal. Whorls with rather prominent bicarinate subsutural fold, concave subsutural ramp, and prominent,

tuberculate, peripheral keel. Subsutural ramp bearing numerous, regular, curved folds formed by thickened scars of

anal sinus. Abapically of keel, spiral cords strong and widely spaced, sometimes with a thinner cord in interspaces

two on exposed part of spire whorls, over 20 on base and canal of last whorl. Anal sinus deep, U-shaped, with

parallel sides. Shell chalky white under rather thick light-brown periostracum. Operculum small, vestigial, with

large central nucleus. Radula long, with more than 50 transverse rows. Radular teeth small, inflated at the base

with proximal opening.

Description (holotype). — Shell solid, narrowly fusiform, with tall spire comprising 40% of total shell

eight. Protoconch and apical teleoconch whorls eroded. Remaining part of teleoconch consisting of 10.5 angulate

Source : MNHN . Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

333

whorls. Subsutural ramp wide, concave with rather massive, broad, flat-topped, bicarinate subsutural fold and

2 weak spiral threads in the middle, ramp bearing numerous, regular, curved folds formed by thickened scars of

anal sinus. On last whorl, cords of subsutural fold becoming strongly and irregularly granulate, and appearing like

a spirally twisted rope, due to intersection with growth lines. Peripheral keel moderately prominent, bearing

numerous rounded tubercles truncated adapically at edge of ramp, 24 tubercles on penultimate whorl, 27 on last

whorl. Spiral sculpture consisting of one abapical cord bordering suture, and one additional strong cord (missing on

apical whorls) below peripheral keel. On last whorl, periphery with 4 widely interspaced strong cords, additional

thinner cords appearing in between. Base and canal covered by ca. 26 cords, sometimes with thinner cord in

interspaces; adapical basal cords of almost same strength as those at periphery, but prominence of cords gradually

decreases towards tip of canal, where cords are obsolete. Axial sculpture consisting of strong incremental lines,

intersection with spiral cords granular. Aperture rather narrow, ovate, gradually continued by long and straight

canal. Inner lip covered by thick callus, parietal wall weakly and evenly curved. Anal sinus deep, U-shaped, with

parallel sides, occupying entire width between subsutural fold and peripheral keel. Colour chalky white under thick

light-brown periostracum.

Dimensions: height 40.5 mm, last whorl height 24.4 mm, aperture height 21.1 mm, diameter 1 1.8 mm.

Operculum (of paratype) small, 1.05 x 0.6 mm, vestigial, very thin, with large central nucleus. Radula long,

narrow, with over 50 transverse rows. Radular teeth rather small, mean length 270 pm (in specimen 38.5 mm

high), short, base strongly inflated, with proximal opening.

Remarks. — Paratypes are smaller than holotype: 38.5 x 11.3 mm and 33.0 x 9.7 mm. Specimens from

Indonesia are very similar to the holotype, except for minor variation in the strength and number of peripheral

tubercles, and width of subsutural fold, which may be slightly concave between two marginal cords. The specimen

from the Philippines (Fig. 23) has a stronger subsutural fold, and a slightly narrower subsutural ramp, especially

on last whorl, with only one thin thread in its middle. Heteroturris gemmuloides is very similar to H. sola , but

differs in attaining twice the size, with a broad, bicarinate subsutural fold, and a nodulose peripheral keel. The

shape of radular teeth is very similar to that of Typhlomangelia polythele Barnard, 1963 and T. adenica Sysoev,

1996 (Barnard, 1963, fig. 3f; Sysoev, 1996, fig. 4).

ETYMOLOGY. — Named gemmuloides because of its superficial resemblance to species of Gemmula.

Distribution. — Tanimbar Islands, Indonesia, and southwest of Luzon Island, Philippines, taken alive at

330-399 m.

Heteroturris serta sp. nov.

Fig. 28

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 28, 05°3 1 'S, 132°54'E. 448-467 m, 1 dd

(holotype).

Type Material. — Holotype MNHN.

DIAGNOSIS. — Shell solid, small, less than 10 mm high, narrowly fusiform, with high spire and long canal.

Protoconch with 3.25 whorls, initial 1.5 whorls smooth, subsequent whorls sculptured by strong, closely spaced

axial ribs. Suture channeled. Whorls with very strong, smooth, rounded peripheral keel and thick subsutural fold,

crenulated adapically. Subsutural ramp narrow, concave, with one spiral cord in the middle, and regular arcuate

folds formed by thickened scars of anal sinus. Spiral cords very strong and widely spaced. Last whorl with three

cords on shell base below keel. Anal sinus broad and shallow. Colour light yellowish.

DESCRIPTION. — Shell solid, thick-walled, small, narrowly fusiform with tall spire comprising 37% of shell

height. Protoconch multispiral, consisting of 3.25 whorls; first 1.5 whorls smooth, subsequent whorls covered

with numerous arcuate axial ribs, which gradually become stronger towards abrupt protoconch/teleoconch

334

A. SYSOEV

discontinuity. Teleoconch consisting of 6.25 whorls. Spiral sculpture on spire whorls consisting of a thick,

overhanging subsutural fold with flattened and steeply descending surface, a strong, smooth peripheral keel, and

another cord bordering abapical suture (on last 2 spire whorls only). Axial sculpture consisting of strong

incremental folds that do not extend over main spiral cords. Subsutural fold consisting of a strong, smooth spiral

cord abapically and a much weaker crenulated adapical cord, separated by area covered by oblique incremental

wrinkles. Subsutural ramp rather narrow, concave, covered by distinct, crisp, arcuate incremental folds formed by

scars of anal sinus, and on last 3 whorls bearing a narrow cord running in the middle of ramp and forming tubercles

at intersection with axial folds. Periphery and base of last whorl with 4 strong cords with wide interspaces,

followed by 13 gradually weakening cords on canal. Canal long and straight. Aperture rather narrow, pyriform,

with a strong lira inside. Inner lip covered by thick callus, parietal side evenly curved. Outer lip chipped, but

growth lines indicating broad and shallow anal sinus. Colour yellowish-white, with a very thin periostracum.

Dimensions: height 9.5 mm, last whorl height 6.0 mm, aperture height 4.8 mm, diameter 3.4 mm.

Remarks. — Heteroturris serta differs from H. sola in having a solid and broad subsutural fold, stronger spiral

sculpture, and a protoconch with fewer whorls. It also differs from H. gemmuloides in its stronger spiral sculpture

and smooth peripheral keel. The holotype of H. serta may not be fully adult, but it is well preserved and the

species should be easily recognizable, either as a juvenile with protoconch or as a mature specimen with adult

sculpture.

Etymology. — From the Latin sertus, plaited, with reference to the very regular incremental sculpture.

Subfamily RAPHITOMINAE Bellardi, 1875

Genus CRYPTODAPHNE Powell, 1942

Type SPECIES: Cryptodaphne pseudodrillia Powell, 1942.

Cryptodaphne rugosa sp. nov.

Figs 24-27

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 13, 05°26'S, 132°38'E 417-425 m 1 dd

(holotype). — Stn DW 24, 05°32,S, 132°51’E, 230-243 m, 1 dd (paratype POLIPI).

Tanimbar Islands : stn DW 44, 07°52'S, 132°48'E, 291-295 m. 2 Iv (paratypes MNHN).

Type Material. — Holotype and 2 paratypes MNHN, 1 paratype POLIPI.

Diagnosis. — Shell rather small, up to 13 mm high, thin, teleoconch shell surface finely granulate.

Protoconch multispiral, consisting of ca. 3 diagonally cancellated whorls. Teleoconch whorls with deep, slightly

channeled suture, and very broad, slightly concave subsutural ramp, occupying most of height on spire whorls.

Strong spiral keel adjacent to abapical suture in early whorls and well below periphery in penultimate whorl. Last

whorl with 10 fine spiral cords and numerous closely set sigmoid incremental lines in subsutural ramp, and several

strong and narrow cords below keel, thin secondary and tertiary cords in interspaces. Siphonal canal short and

curved. Anal sinus moderately deep, asymmetrical, with deepest point in lower half of ramp. Colour of protoconch

light brown, teleoconch off-white.

Description (holotype). — Shell rather small, thin but solid, narrowly biconical, with tall spire comprising

40% of shell height. Protoconch multispiral, protoconch I and initial part of protoconch II missing, remaining part

consisting of about 2 whorls with diagonally cancellated sculpture, diameter 650 pm. Protoconch/teleoconch

boundary sharp, with deeply opisthocyrt protoconch outer lip. Teleoconch consisting of 6.75 whorls separated by

deep, slightly channeled suture, shell surface minutely granular. Most of whorl height occupied by very broad,

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

335

slightly concave subsutural ramp. Primary sculpture consisting of spiral cords, only the strongest adapical one

exposed on early whorls, forming a suprasutural keel; penultimate whorl with one additional strong cord exposed

below keel; last whorl with 8 strong, widely spaced primary cords exposed on base below peripheral keel,

1-3 thinner secondary cords in interspaces; 9 weaker cords on canal. Subsutural ramp smooth on early whorls, on

later whorls bearing narrow spiral cords, 8 on penultimate, 10 on last whorls, the second from adapical margin and

the most abapical ones strongest. Axial sculpture of numerous, narrow sigmoid folds formed by thickened

incremental lines. Axial folds and spiral cords forming reticulate pattern in ramp, small nodules at points of

intersection. Canal rather short and curved. Aperture oval, columella thick, forming a distinct angle with parietal

part of inner lip. Anal sinus asymmetrical, moderately deep, steeply descending adapically, deepest point rounded in

abapical half of ramp. Protoconch light brown, teleoconch off-white.

Dimensions: height 1 1.8 mm, diameter 4.8 mm, body whorl height 7.0 mm, aperture height 5.2 mm.

Remarks. — In the paratypes (dimensions 12.8 x 5.1 mm, 9.3 x 3.7 mm, and 7.7 x 3.5 mm), the strength

and relative position of the main spiral keel vary a little, as do also the concavity of the subsutural ramp and the

number of cords on the ramp and in interspaces of primary spirals. One paratype has a protoconch of about

3 whorls with etched surface, but even in that specimen the earliest part is dissolved.

Cryptodaphne rugosa resembles C. affinis (Schepman, 1913), but can be easily distinguished by its

proportionally larger body whorl, differently shaped anal sinus, and details of sculpture. In fact, it is much more

similar to the fossil type-species than to any of the Recent species assigned to the genus (Powell, 1966;

Shuto, 1971). The New Zealand type species C. pseudodrillia was originally recorded from the Lower Miocene

(Powell, 1942) and later (Powell, 1966) the Upper Oligoccne, but Beu & Maxwell (1990) list it in the Early

Miocene faunal assemblage. C. rugosa is almost twice the size of the holotype of C. pseudodrillia and differs

mainly in the shape of the anal sinus, in C. pseudodrillia “descending almost vertically and then abruptly produced

forward” (Powell, 1942: 165), as well in the twisted canal, absence of “close-spaced spiral threads” covering the

entire shell surface, and presence of minute granulation.

The diagonally cancellate protoconch sculpture clearly indicates a position in Raphitominae, however C. rugosa

possesses features more frequently encountered in Clathurellinae, such as the closely spaced axial folds, formed by

thickened growth lines, well developed over the entire subsutural ramp, and the presence of minute granulation on

the shell surface.

Etymology. — Rugosus, Latin adjective meaning rough, with reference to granular microsculpture.

Genus GYMNOBELA Verrill, 1884

TYPE Species: Gymnobela engonia Verrill, 1884.

Gymnobela ioessa sp. nov.

Figs 3, 29-33

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 54, 08°2I'S, 131°43'E, 836-869 m,

5 Iv (holotype and 3 paratypes MNHN. 1 paratype POLIPI). 5 dd (3 paratypes MNHN and 2 paratypes POLIPI).

Type Material. — Holotype lv and 6 paratypes MNHN, 3 paratypes POLIPI.

DIAGNOSIS. — Shell exceeding 45 mm in height, fusiform, solid, with high spire and short canal. Whorls

angulate below periphery, angulation obsolete on last adult whorl. Subsutural ramp broad, moderately to weakly

concave, without axial sculpture and with fainter spiral cords, sometimes with thickened scars of anal sinus

adapically. Axial ribs oblique, rather short, reaching abapical suture, only occupying periphery on adult whorls,

obsolete on last whorl of large adults. Spiral cords flat, separated by narrow interspaces and, sometimes, additional

336

A. SYSOEV

narrower cords. Anal sinus broad, moderately deep, somewhat angulate. Colour reddish-violet, with lighter

subsutural band, aperture brown inside. Radular teeth straight, narrow, barbed, with narrow trilobate basal part.

Description (holotype). — Shell fusiform, thin but solid, with tall spire comprising 41.5% of shell height.

Protoconch and outer layers of apical teleoconch whorls dissolved, only traces of brown protoconch columella

remaining. Teleoconch consisting of 10 whorls, suture shallow, slightly channeled. Apical whorls angulate at

periphery, angulation weaker and situated a little below periphery in subaduit whorls, last whorl rather evenly

rounded. Whorl profile concave above angulation, slightly convex below it. Axial sculpture consisting of ribs that

extend abapically from whorl angulation to outer base, oblique, weakening abapically, 14 on penultimate whorl,

obsolete on last whorl. Spiral sculpture consisting of narrow grooves delimiting low, flat spiral bands, grooves

weaker on subsutural ramp, deeper below periphery. Base evenly convex, smoothly continuous with rather short,

straight, moderately broad canal. Aperture broad, oval, parietal side weakly concave, separated by obtuse angle from

straight columella. Inner lip a thin, broad callus, outer lip evenly curved. Anal sinus moderately deep, broad,

slightly angulate, deepest point just below middle of subsutural ramp. Colour dull reddish-violet, with a lighter

band subsuturally in part of ramp, aperture and columella orange brown. Periostracum very thin, transparent.

Dimensions: height 46.3 mm, last whorl height 27.1 mm, aperture height 21.2 mm. diameter 16.0 mm.

Last 1-1.5 protoconch whorls preserved on 2 paratypes, brown, with sculpture of spiral cords interrupted by

close set, curved axial riblets. Radular teeth of 34.2 mm high paratype straight, narrow, barbed, with narrow

trilobate basal part, mean length 170 pm (Fig. 3). No operculum.

Remarks. — The paratypes are smaller than the holotype (largest 45.5 x 16.3 mm). In smaller specimens, the

axial ribs number 14 or 15 on the last whorl and do not extend onto shell base. The adapical part of the subsutural

ramp may bear numerous, regular, thickened scars of the anal sinus.

Gymnobela ioessa resembles G. muricata , occurring sympatrically, but differs in having a more slender,

reddish-violet shell, higher axial ribs reaching suture abapically, and smaller radular teeth with narrow, trilobate

basal part. G. ioessa also attains a smaller size.

G. ioessa and G. muricata form a distinct group within the genus, which combines characters of two different

subfamilies of Conidae. The general shell outline and colouration, and the character of anal sinus are similar to

those of Clathurellinae (e.g., Borsonia and Typhlosyrinx Thiele, 1925), whereas the protoconch sculpture and the

structure of radular teeth indicate a position in the Raphitominae. These species, however, can be placed in

Gymnobela taken in a broad sense, as is frequently adopted in the current literature.

Etymology. — From the Greek ioeis (feminine ioessa ), meaning violet or dark brown, based on the ground

colour of the shell.

Gymnobela muricata sp. nov.

Figs 4, 34-38

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands : stn CP 54, 08°21'S, 131°43’E, 836-869

0 Iv (2 paratypes MNHN, 1 paratype POLIPI), 2 dd (holotype and paratype MNHN). — Stn CP 73 08°29'S 131°33

840-855 m, 1 Iv (paratype POLIPI). - Stn CP 91, 08°44'S, 131°05'E, 884-891 m, 1 dd (paratypes MNHN)!"

Type Material. — Holotype (dd) and 4 paratypes MNHN, 2 paratypes POLIPI.

m,

'E,

Diagnosis. — Fully adult shell very large, up to 70 mm high, broad, solid, with high, regularly conical spire,

short, broad siphonal canal. Whorls angulate just below middle of whorl, angulation obsolete in last adult whorls.

Subsutural ramp wide, weakly concave, without axial sculpture. Axial ribs oblique, short, not reaching abapical

suture, only occupying periphery in last whorls, obsolete on last whorl of large adults. Spiral grooves narrow,

ehmiting flat, unevenly broad cords, weaker on ramp than on periphery and base. Anal sinus broad, moderately

deep somewhat angulate. Colour white to light-brown. Radular teeth straight, long, weakly barbed, with bifurcate

Source : MNHN Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

337

Description (holotype). — Shell broadly fusiform, thin but solid, with tall spire comprising 39% of total

shell height. Protoconch and apical whorls dissolved. Teleoconch consisting of 10 whorls, suture shallow. Earliest

teleoconch whorls angulate just below middle of whorl, angulation weaker on subsequent whorls, last whorl

almost evenly rounded. Whorl profile weakly concave above angulation, almost flat below. Axial ribs very short,

oblique, extending abapically from whorl angulation on earliest teleoconch whorls, forming rounded knobs just

below periphery on subadult whorls (18 ribs on penultimate whorl), obsolete on last adult whorl. Spiral sculpture

consisting of narrow, shallow, unevenly spaced grooves separating low, flat cords. Cords weaker but more regular

on subsutural ramp, stronger but uneven below whorl angulation. Last whorl rather inflated, with evenly convex

base, clearly separated from short, very broad, slightly twisted siphonal canal. Aperture broad, oval. Inner lip

covered by thin, broad, longitudinally wrinkled callus, concave parietal and columellar sides forming obtuse angle.

Outer lip evenly convex. Anal sinus moderately deep, broad, slightly angulate, deepest point in abapical part of

subsutural ramp. Colour chalky light yellowish brown.

Dimensions: height 69.1 mm, last whorl height 42.4 mm, aperture height 33.3 mm, diameter 25.7 mm.

Radular teeth of 33.3 mm high paratype weakly barbed with strong basal spur and bifurcate basal part, mean

length 275 pm (Fig. 4). No operculum.

Remarks. — The paratypes are smaller than the holotype (the largest is 52.1 x 20.6 mm). Small specimens

are dull greyish-white, with a very thin transparent periostracum, and proportionally longer and narrower canal. The

protoconch is dissolved in all specimens, but there are traces of a brown layer, suggesting that the protoconch was

of the planktotrophic type.

For differences with G. ioessa, see that species.

ETYMOLOGY. — From the Latin muricatus (adj.), meaning pointed or spiny like a murex, with reference to the

high spire and short, knob-like axial ribs.

Gymnobela mitrodeta sp. nov.

Figs 5, 45-46

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn CP 12, 05°23'S, 132°37'E, 413-436 m, 1 lv

(holotype).

Type Material. — Holotype MNHN.

Diagnosis. — Shell of medium size, 24 mm high, thin but solid, buccinoid. Protoconch multispiral, small,

with 2+ diagonally cancellated whorls. Teleoconch whorls angulate at periphery, suture deep, slightly channeled.

Subsutural ramp occupying about half of exposed whorl height between subsutural row of blunt tubercles and

crowned periphery, ramp concave on both sides of convex median part. Spiral sculpture of fine cords in ramp,

stronger but narrow cords at periphery and broad rounded cords abapically, secondary and tertiary cords in

interspaces. Axial ribs oblique, numerous, very short on last whorls. Canal moderately long, slightly curved and

obliquely truncated. Anal sinus asymmetrical, with apex in lower half of subsutural ramp. Protoconch brown,

teleoconch white, with two light reddish-brown bands at periphery and on canal. Radular teeth small with well-

developed basal part.

Description (holotype). — Shell thin but solid, buccinoid, with angulated shoulder at whorl periphery.

Protoconch I and initial part of protoconch II missing, remaining part consisting of about 2 whorls with

diagonally cancellated sculpture, diameter 550 pm. Protoconch/teleoconch boundary sharp. Teleoconch consisting

of 7.8 convex, strongly angulated whorls, suture deep, narrowly channeled by subsutural ridge. Subsutural ramp

occupying about half of spire whorls, concave on early whorls, but becomes progressively convex in the middle

part on three last whorls; smooth on first 2 whorls, on subsequent whorls sculptured by spiral cords, very weak

below subsutural ridge, increasing in strength on abapical half of subsutural ramp, and becoming very narrow near

338

A. SYSOEV

shoulder. Below shoulder, spiral sculpture of strong, rounded and widely spaced primary cords, 2 on spire whorls,

4 on periphery of last adult whorl, interspaces with densely packed thinner secondary and tertiary cords. Spiral

cords weaker on base and siphonal canal. Axial sculpture consisting of strong ribs, which are interrupted by ramp,

and form blunt, elongated tubercules on subsutural ridge and at periphery. On early whorls, ribs almost straight,

extending over all exposed part of whorl below periphery, on two last whorls ribs more numerous (28 on

penultimate, 34 on last whorl), oblique, very short, occupying only periphery. Base moderately constricted to a

rather short, slightly twisted, and obliquely truncated canal. Aperture oval, siphonal canal poorly demarcated. Inner

lip covered by thin and glossy callus, parietal area weakly convex, columellar area almost straight. Anal sinus

strongly asymmetrical, deepest part in abapical half of subsutural ramp, abapical edge almost horizontal, outer lip

strongly projecting forward below sinus. Protoconch brown, teleoconch ground color white, with two pale reddish-

brown bands, one encircling peripheral angulation on last two whorls, the other obliquely encircling canal.

Dimensions: height 24.2 mm, diameter 1 1.2 mm, last whorl height 16.3 mm, aperture height 13.4 mm.

Radular teeth small, 130 (xm long, with well-developed basal part, subquadrate in front view (Fig. 5).

REMARKS. — Gymnobela mitrodeta can be easily distinguished from other species of the genus by its complex

spiral sculpture, short and numerous axial ribs, tuberculated subsutural fold, strongly asymmetrical anal sinus, and

peculiar color pattern.

ETYMOLOGY. — From the Greek mitrodetos (adjective), crowned by a turban, with reference to the orange

spiral band encircling the periphery of subadult and adult whorls.

Gymnobela micraulax sp. nov.

Figs 47-48

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 91. 08°44'S, 131°05'E, 884-891 m,

2 lv (holotype and paratype).

Type Material. — Holotype and paratype MNHN.

Diagnosis. — Shell of medium size, up to 27 mm, thin, semi-transparent, narrow, with high spire.

Teleoconch whorls angulate at periphery or above it, with steep, smooth subsutural ramp. Suture shallow.

Spiral sculpture of narrow grooves. Axial ribs strong, oblique, vanishing rapidly below periphery before abapical

suture. Canal moderately long, inner lip without callus. Outer lip strongly projecting below deep anal sinus.

Colour light-brown.

DESCRIPTION (holotype). — Shell fusiform, thin, semi-transparent, narrow, with high spire forming about

40% of total height. Protoconch corroded apically, remaining 2 whorls sculpture etched, but traces of oblique

reticulation present. Teleoconch consisting of 8.5 shouldered whorls, whorl angulation above periphery, suture

shallow, slightly adpressed, steeply descending subsutural ramp almost flat adapically and concave near whorl

angulation. Axial sculpture of strong, broad and opisthocline ribs, forming axially elongated nodule below

shoulder, vanishing abapically before reaching suture. 14 axial ribs on penultimate whorl, 13 on last whorl.

Incremental lines forming raised wrinkles in adapical part of subsutural ramp. Spiral sculpture consisting of

narrow, rather evenly spaced grooves, 9 on exposed part of penultimate whorl, 35 on last whorl, more crowded on

canal. Base weakly convex, smoothly connected to moderately long, obliquely truncated canal. Aperture narrow,

poorly demarcated from canal, inner lip weakly and evenly concave, without a callus, outer lip thin, fragile,

strongly projecting forward below anal sinus. Anal sinus deep, deepest point just below suture. Colour of

protoconch with traces of brown, teleoconch glassy light-brown.

Dimensions: shell height 26.6 mm, diameter 8.1 mm, body whorl height 15.8 mm, aperture height 12.8 mm.

Remarks. — The paratype measures 22.0 x 7.1 mm and has 7.5 teleoconch whorls (protoconch corroded,

surface etched). Whorl angulation is approximately at periphery, which also results in a slightly broader subsutural

ramp. It also differs slightly from the holotype in the ribs being slightly more axially elongated.

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

339

Gymnobela micraulax can be easily distinguished from other Indo-Pacific species of Gymnobela by its high

spire and sculpture of narrow grooves.

ETYMOLOGY. — Micraulax, Greek (adj.) meaning with small furrows, with reference to the spiral sculpture.

Gymnobela baruna sp. nov.

Figs 6, 49-50

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn CC 21, 05°14'S, 133°00'E, 688-694 m, 1 Iv

(holotype), 1 dd (paratype).

Type Material. — Holotype and paratype MNHN.

Diagnosis. — Shell thin, up to 36 mm high, with high spire occupying 45% of shell height. Proto¬

conch multispiral, with diagonally cancellated sculpture. First teleoconch whorls with angular periphery,

weakly concave anal sinus occupying ca. 60% of subsural ramp, periphery with very short oblique axial ribs,

last whorls almost evenly convex. Spiral sculpture of fine spiral cords and strongly sigmoid incremental

lines, intersection irregularly reticulate. Colour white, adapical half of last whorls with very pale yellowish-white

band.

Description (holotype). — Shell consisting of 1+ protoconch and 9.5 teleoconch whorls, slender, thin but

rather solid, high, fusiform, tall spire occupying 45% of shell height, suture rather deep, slightly channeled. Tip of

protoconch dissolved, remaining whorl surface etched but remnants of sculpture typical of multispiral larval shell

with diagonally cancellated sculpture, indicating planktotrophic development. First 6 teleoconch whorls distinctly

shouldered, weakly concave sinus zone occupying about 60% of broad subsutural ramp, whorl profile almost flat

above and below strong peripheral angulation. Subsequent whorls almost evenly convex, subsutural ramp flat with

poorly defined abapical border. Axial sculpture consisting of strongly sigmoid incremental lines, forming raised

opisthocyrt riblets in sinus zone, and strongly prosocline threads below sinus; very short, broad, opisthocline ribs

on periphery of spire whorls, ca. 15 per whorl, vanishing on 6th and subsequent whorls. Spiral sculpture

consisting of cords, weakly defined and widely spaced in sinus zone, sharply defined, thin, rounded and closely

spaced on rest of the whorl, a few stronger below periphery. Intersection of incremental lines and spiral cords rather

regularly reticulate on spire whorls, more irregularly so on last whorl, due to unevenness of lines and cords.

Aperture rather broad, oval, widely open siphonal canal not distinctly set off. Inner lip with thin, polished callus

extending over convex parietal and concave columellar areas. Outer lip chipped, anal sinus (from shape of growth)

rather deep, its deepest point in middle of subsutural ramp. Colour chalky white, adapical half of last whorls with

very pale yellowish-white band.

Dimensions: height 36.2 mm, diameter 12.8 mm, last whorl height 21.3 mm, aperture height 16.7 mm.

Radular teeth (Fig. 6) straight, unbarbed, basal part subquadrate in frontal view, length 250 pm.

Remarks. — The paratype (21.6 x 8.5 mm at 2+ protoconch and 7.5 teleoconch whorls) is a damaged and

slightly eroded shell. Whorl profile and sculpture correspond to the characters of holotype at comparable size, but

whorl angulation is more pronounced than on corresponding (7th) whorl of the holotype; also, spiral cords and

growth lines are slightly coarser.

Gymnobela baruna is rather different from most other representatives of the genus. The general shell outline,

the absence of prominent axial sculpture and fine spiral cords are more characteristic of Xanthodaphne . However,

the high spire and clearly defined broad subsutural ramp, delimited by the whorl angulation in early whorls, are

features that do not fit that genus. G. baruna is most similar to the species illustrated by Matsumoto (1979,

pi. 17. fig. 3) as Daphnella proximo Kuroda. The latter, however, is a manuscript name and a nomen nudum

[ICZN Art. 13a(i)].

ETYMOLOGY. — From the name of the Indonesian research vessel "Baruna Jaya 1" which collected the

material. Baruna is used as a noun in apposition.

340

A. SYSOEV

Genus MIOA WATERIA Vella. 1954

Type Species: Awateria personate! Powell, 1954.

Mioawateria asarotum sp. nov.

Fig. 55

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: sin DW 13, 05°26'S. 132°38'E, 417-425 m, I dd

(holotype).

Type Material. — Holotype MNHN.

Diagnosis. — Shell small, about 7 mm high, short, wide, solid, with few rapidly expanding whorls.

Protoconch multispiral with diagonally cancellated sculpture. Teleoconch whorls angulate. Last whorl almost as

high as wide, completely covered by sculpture of low, wide cords, intersecting numerous, stronger but narrower

axial ribs, intersection forming axially elongated nodules. Aperture broad. Canal very short and poorly

differentiated from aperture. Anal sinus broad and shallow.

Description. — Shell small, solid, broadly biconical, with low spire and very large, inflated last whorl,

occupying 74% of total shell height. Protoconch consisting of 2+ convex whorls, protoconch I and probably about

0.5 whorl of protoconch II missing, sculpture of opisthocyrt, opisthocline axial riblets, forming diagonal

cancellation with oblique threads in abapical two-thirds. Axial riblets nearly orthocline near clearly defined

protoconch/teleoconch discontinuity. Teleoconch consisting of 3.8 rapidly expanding whorls, with sharp

angulation above periphery, subsutural ramp weakly concave. On first teleoconch whorl, two spiral cords and

a third, poorly defined one on shoulder, intersecting thinner axial riblets, intersection forming axially elongate

nodules. On 2nd and third whorls, an additional spiral cord exposed above suture, shoulder cord more distinct,

intersecting axial riblets of similar strength, intersection forming rounded beads. On last whorl, 19 spiral cords and

35 ribs; subsutural cord indistinct on broad subsutural fold, peripheral cords broad, low, separated by narrow

groove, cords on base and canal higher and separated by interspaces occupying equal width; axial ribs sharp,

stronger than cords, forming raised, prosocline, sigmoid wrinkles in subsutural zone, rather orthocline and

extending over whole whorl height below angulation, forming checkerboard pattern with spiral grooves. Aperture

broad, parietal wall convex with rather thin callus, columella almost straight with thicker callus. Canal short and

broad. Outer lip broken, but growth lines indicating very shallow sinus, with deepest point at whorl angulation.

Protoconch brown, teleoconch beige-white.

Dimensions: height 6.9 mm, diameter 4.8 mm, last whorl height 5.1 mm, aperture height 3.9 mm.

Remarks. Mioawateria asarotum is similar to the type species, M. personata, but differs in having much

lower spire and inflated body whorl. In shell outline, it is similar to M. rhomboidea (Thiele, 1925) from West

Africa, but differs in its smaller shell with curved axial ribs and very wide spiral cords. M. extensaeformis

(Schepman, 1913), which is rather common in the Indo-Pacific bathyal, has a narrower shell and much weaker

spiral sculpture.

ETYMOLOGY. Latin asarotum (noun in apposition), a floor laid in mosaic, with reference to the elongated

squares formed by intersection of axial and spiral sculpture.

Genus CLINURA Bellardi, 1875

Type Species: Murex (Pleurotoma) calliope Brocchi, 1814 (Neotype figured by ROSSI RONCHETTI 1955

fig. 163).

Source MNHN Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

341

Clinura vitrea sp. nov.

Figs 7, 54

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 91, 08°44'S, 131°05'E, 884-891 m,

1 Iv (holotype).

Type Material. — Holotype MNHN.

Diagnosis. — Shell of medium size, 21 mm high, thin, porcellaneous, with high pagodiform spire.

Teleoconch whorls very convex, with keeled periphery. Axial ribs numerous, very short, restricted to whorl

periphery, forming narrow oblique knobs on keel. Spiral sculpture of strong cords below periphery, interspaces

rather wide. Last whorl attenuated towards moderately long, straight canal. Subsutural ramp broad, almost devoid

of sculpture, sinus rounded, deepest point in middle of ramp. Radular teeth short, straight, with bilobate basal part.

Description (holotype). — Shell thin, porcellaneous, broadly fusiform, consisting of 8.5+ very convex

teleoconch whorls (protoconch and apical teleoconch whorls dissolved) with high pagodiform spire occupying 42%

of total shell height. Suture impressed, periphery angulate, subsutural ramp broad, convex, whorl profile concave

abapically of slightly overhanging peripheral keel, base regularly convex. Outer shell layer etched on spire whorls,

details of sculpture preserved on last two whorls only. Axial sculpture consisting of numerous (28 on penultimate

and last whorls), short, opisthocline riblets, forming oblique nodules on peripheral keel and extending to abapical

concavity only; incremental lines very distinct in ramp, occasionally thickened as opisthocyrt wrinkles. Spiral

sculpture in subsutural ramp consisting of 6-8 poorly defined cords, on and below peripheral keel consisting of

strong, sharply defined cords, interspaces broader than cords, 2 stronger cords just above suture in exposed part of

spire whorls. Base convex, canal distinctly set off, long, straight, narrow. Aperture ovate, inner lip a thin glaze

over weakly convex parietal area and almost straight columella. Outer lip chipped, based from incremental lines

anal sinus occupies whole ramp, rounded, deepest point on abapical side of middle part of ramp. Colour por¬

cellaneous white, etched shell surface and interspaces between cords opaque, chalky white, ramp and cords vitreous.

Dimensions: height 21.3 mm, diameter 10.5 mm, last whorl height 13.4 mm, aperture height 11.1 mm.

Radular teeth 250 |4m long, straight, short, with a bilobed and turned out basal part (Fig. 7).

REMARKS. — Until now the genus Clinura was known only from Oligocene to Pliocene deposits in Europe,

Indonesia and New Zealand (POWELL, 1966), and C. vitrea is the first Recent record. Regrettably, however, the

protoconch of the new species is unknown, and this makes the generic placement only provisional. It is similar to

the type species from the Pliocene of Italy. Characters shared by fossil species of Clinura and C. vitrea are the

fusiform-biconic shell profile with a pagodiform spire, narrow canal, crenulate peripheral keel, and strong spiral

sculpture below whorl periphery. In C. calliope the anal sinus is rather shallow and has the apex in the upper part

of subsutural ramp but the form and position of anal sinus vary rather greatly within the genus (see BEETS, 1942)

and in some species are similar to those of C. vitrea.

ETYMOLOGY. — Latin vitreus (adjective), like glass, with reference to both the fragility of the shell and the

semi-transparent appearance.

Genus VEPRECULA Melvill, 1917

Type Species: Clathurella sykesi Melvill & Standen, 1903.

Veprecula bandensis sp. nov.

Fig. 53

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: sin CC 21, 05°14'S, I33°00'E. 688-694 m, I lv

(holotype).

342

A. SYSOEV

Type Material. — Holotype MNHN.

DIAGNOSIS. — Shell large for genus, 21 mm high, very thin and fragile, slender, with high spire. Protoconch

multispiral with oblique axial ribs. Teleoconch whorls weakly angulate at shoulder, narrow, weakly concave

subsutural ramp with regular fold-like scars of anal sinus. Sculpture finely reticulate, of numerous thin axial

riblets and thin strong spiral cords, interspaces spirally elongate. Canal moderately long, slightly twisted and

obliquely truncated. Anal sinus deep, deepest point in adapical part of ramp. Colour white.

DESCRIPTION. — Shell very thin and fragile, slender, fusiform, with high spire comprising 37% of total shell

height. Protoconch I and tip of multispiral protoconch II dissolved, only last whorl remaining, convex, sculptured

by strong, coarse, prosocline axial ribs; protoconch / teleoconch discontinuity sharp. Teleoconch consisting of

7.0 whorls, suture tightly impressed, whorls weakly angulate at shoulder, evenly convex below shoulder.

Subsutural ramp narrow, weakly concave, without spiral sculpture, with numerous regular, raised, opisthocyrt

wrinkles corresponding to scars of anal sinus. Sides finely reticulately sculptured by numerous (42 on last whorl),

prosocyrt, sharply defined, narrow axial riblets overriden by almost equally strong but narrower spiral cords,

interspaces spirally very elongate, intersection of cords and riblets spirally elongate; 3 spiral cords on first

teleoconch whorl, additional cords added at shoulder and between cords, reaching 20 on penultimate whorl, 35 on

periphery and base of last whorl, plus 13, slightly stronger, on canal. Base evenly connected to moderately long,

twisted, and obliquely truncated canal. Aperture pyriform. Inner lip strongly concave on abapical part of parietal

side, covered by thin, translucent, axially wrinkled callus. Outer lip damaged, anal sinus (based on incremental

scars) deep, deepest point in adapical part of ramp. Colour of protoconch deep brown, of teleoconch semi¬

transparent white with thin, tightly adhering beige periostracum.

Dimensions: height 21.1 mm, diameter 7.7 mm, last whorl height 13.2 mm, aperture height 10.2 mm.

Remarks. — Veprecula bandensis can be included in Veprecula on the basis of shell outline, reticulate

sculpture, and characteristic sculpture of protoconch. It is most similar to the type species, V. sykesi , but differs in

having a much larger shell with lower spire, fainter sculpture, twisted canal, and poorly prominent intersection of

axial and spiral sculpture. Other known species of the genus have a much stronger sculpture, consisting of widely

spaced ribs and cords.

The new species is also similar to Daphnella thia Melvill & Standen, 1903 (the original figure of that species

is rather inadequate, but it was recently illustrated by BOSCH et al., 1995: 167) in shell outline and sculpture, but

differs in having more curved ribs, a well differentiated subsutural ramp, and different protoconch sculpture.

Etymology. — Bandensis , an adjective based on the stem Banda (Sea), the type locality.

Genus XANTHODAPHNE Powell, 1942

Type Species: Pleurotoma (Thesbia) membranacea Watson, 1886.

Xanthodaphne cladara sp. nov.

Figs 8, 51-52

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn CC 21, 05°14'S, 133°00'E 688-694 m 2 Iv

(holotype and paratype).

Type Material. — Holotype and paratype MNHN.

Diagnosis. — Shell large for genus, about 32 mm high, buccinoid, very thin, fragile, light-brown.

Protoconch brown, of about 3 diagonally cancellated whorls. Teleoconch spiral sculpture of very fine cords,

appearing on third whorl and becoming stronger towards last whorl. Subsutural zone of first whorls with regular!

Source : MNHN, Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

343

opisthocyrt folds formed by thickened growth lines, becoming obsolete on last whorls. Canal rather long,

columella slightly twisted.

DESCRIPTION (holotype). — Shell buccinoid, very thin, fragile, consisting of 2.1+ protoconch and

6 teleoconch whorls. Tip of protoconch broken, probably consisting of protoconch I and less than 0.5 whorl of

protoconch 11. Remaining whorls forming a rather low spire, diameter 1 .05 mm, sculptured over abapical two-

thirds by oblique reticulation, and opisthocyrt riblets on adapical third. Protoconch/teleoconch boundary sharp.

Teleoconch whorls very convex, with rather deep, impressed suture, slightly concave subsutural ramp defined only

on first 4 whorls, last two whorls very regularly convex. Sculpture fine and delicate, consisting of sigmoid

incremental lines, obsolete in last 2 whorls, strong in first 4 whorls, forming raised opisthocyrt wrinkles in

subsutural zone, and low, indistinct, prosocyrt lines below shoulder; numerous, fine, flattened spiral cords,

interspaces narrow, indistinct on first two whorls, stronger on subsequent whorls, also present in subsutural zone.

Last adult whorl strongly convex, with weak incremental lines and thin, but strong spiral cords, 4 or 5 per mm on

periphery, a little more crowded on base, more spaced on canal. Canal rather long, broad, straight. Aperture broad,

columella slightly twisted, without inner lip callus. Based on growth lines, anal sinus broad, very shallow. Colour

light-brown, protoconch brown.

Dimensions: height 31.7 mm, diameter 13.8 mm, last whorl height 22.3 mm, aperture height 17.1 mm.

Remarks. — The paratype (30.4 x 12.4 mm at 6 teleoconch whorls) has a less inflated last whorl:

diameter/height 0.41 vs. 0.435 in the holotype. Such a difference could possibly represent sexual dimorphism.

Growth lines on the last whorl are more distinctly sigmoid than in the holotype, due to more prosocyrt profile on

periphery. Radular teeth 280 pm, straight, with short but strongly expanded basal part (Fig. 8).

Xanthodaphne cladara is similar to X. subrosea (Barnard, 1963) from off Cape Point, South Africa, 2524-

2780 m, but differs in having a more slender shell, with more slowly expanding whorls, a smaller protoconch

(diameter 1.45 mm in subrosea ), and fainter spiral sculpture.

ETYMOLOGY. — From the Greek kladaros (adj.; feminine kladara), easily broken, with reference to the fragility

of the shell.

Genus LUSITANOPS Nordsieck, 1968

Type Species: Pleurotomella lusitanica Sykes, 1906.

Lusitanops dictyota sp. nov.

Fig. 56

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands : sin CP 38, 07°40'S, 132°27'E, 620-666 m,

1 lv (holotype).

New Caledonia. Biocal: stn. CP 60. 24°01'S, 167°08'E, 1480-1530 m, 1 lv.

Type Material. — Holotype MNHN.

DIAGNOSIS. — Shell large for genus, about 17 mm high, thin, light-brown. Protoconch of 3+ diagonally

cancellated brown whorls. Teleoconch sculpture of thin spiral cords covering the entire shell surface, and

numerous, regularly and closely set incremental riblets, intersection finely reticulate. Aperture broad. No anal

sinus. No radula, no operculum.

Description (holotype). — Shell buccinoid, thin, consisting of 3 + protoconch and 4.3 teleoconch whorls.

Tip of protoconch (probably only protoconch I) damaged, remaining whorls convex, diameter 1.05 mm, sculptured

on abapical two-thirds by opisthocyrt riblets intersected by equally strong oblique cords, on adapical third by much

Source :

344

A. SYSOEV

fainter spiral threads in axial interspaces. Protoconch/teleoconch boundary sharp. Teleoconch whorls evenly

convex, suture impressed, rather deep. Spiral sculpture consisting of fine, flattened cords, occasionally in last

2 whorls with thinner additional cord in interspaces, interspaces equal to or narrower than cord width, 17 on

penultimate whorl, about 75 on last whorl. Axial sculpture consisting of numerous, evenly spaced, orthocline,

thickened incremental lines, interspaces narrow. On first half teleoconch whorl, spiral cords stronger than

incremental ribs; on subsequent 2 whorls, cords and ribs of equal strength, cords a little rugose at intersection with

ribs, producing a finely reticulate sculpture; on last 2 whorls, spiral cords stronger than incremental riblets. Base

regularly convex, connected rather abruptly to rather short, straight canal. Aperture broad, columellar and parietal

areas of inner lip forming an obtuse angle, callus very narrow, outer lip partly broken, but apparently regularly

convex. No distinct anal sinus. Colour light-brown, protoconch brown. No operculum. No radula.

Dimensions: height 16.9 mm, diameter 9.4 mm, last whorl height 13.2 mm, aperture height 10.4 mm.

REMARKS. — Lusitanops dictyota is similar to L. cingulata Bouchet & Waren, 1980 from the upper abyssal of

the North-Eastern Atlantic, but differs in having a much larger shell with fainter and closely set spiral cords. This

is the first record of Lusitanops outside the North-Eastern Atlantic.

The specimen from New Caledonia is very similar to the holotype, except for being much smaller (8.6 x

5.2 mm at 3+ adult whorls), with slightly broader last whorl. Its protoconch consists of 2.5 remaining whorls,

upper ca. 1.25 whorls missing.

Etymology. — From the Greek dictyotos (adj.), reticulate, with reference to the sculpture of the teleoconch.

Distribution. — Tanimbar Islands, Indonesia, and southward of New Caledonia, taken alive at 620-1530 m.

CONOEDEA incertae sedis

Genus THELECYTHARELLA Shuto, 1969

Type Species: Agladrillia oyamai Shuto, 1965.

Among existing genera of Conoidea, the new species described below conforms rather well with

Metaclathurella Shuto, 1983 [type species: Austropusilla (Metaclathurella) crokerensis Shuto, 1983), originally

classified by Shuto as a subgenus of Austropusilla Laseron, 1954. KlLBURN (1995) has synonymized

Metaclathurella and Lioglyphostomella Shuto, 1970 with Thelycytharella [sic!], and stated that Austropusilla is

not related. This synonymization seems reasonable, with Metaclathurella only characterized by a complete absence

of axial sculpture. Within this complex, T. vitrea (Reeve, 1845) and T. kecil sp. nov., form a distinct group

characterized by a very small (5.5 mm high) pupoid shell without axial sculpture, and further research may prove

that they belong to a separate genus.

The taxonomic position of Thelecytharella is unclear. It was originally placed in the subfamily Mangeliinae, an

opinion followed by KlLBURN (1995), without an indication of reasons for such placement. However,

conchological characters alone are insufficient to allocate it to a subfamily, and even to a family. Examination of

anatomy is badly needed, but the present material is represented by a single empty shell.

Thelecytharella kecil sp. nov.

Fig. 58

Material EXAMINED, — Indonesia. Karubar, Kai Islands: stn DW 28, 05°31'S. 132°54'E 448-467 m 1 dd

(holotype). ’

Type Material. — Holotype MNHN.

Source : MNHN, Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

345

DIAGNOSIS. — Shell small, 5.5 mm, but solid, subcylindrical, few-whorled, white. Protoconch multispiral,

smooth. Teleoconch whorls very weakly but evenly convex, covered by strong spiral cords, 5 on first whorl, 22 on

last whorl. Aperture narrow, canal very short. Anal sinus moderately deep, rounded, directed somewhat adapically.

DESCRIPTION (holotype). — Shell very solid, subcylindrical, consisting of 2+ protoconch and 3.1 teleoconch

whorls. Tip of protoconch dissolved, plugged by secondary callus, remaining whorls smooth, polished, with

strongly opisthocyrt incremental lines and strong basal keel partly covered by successive whorl; protoconch

diameter 850 |um, i.e. half diameter of first teleoconch whorl. Protoconch/teleoconch boundary sharp. Teleoconch

whorls rapidly growing but slowly expanding, weakly and evenly convex, suture shallow. Sculpture primarily

spiral, consisting of strong rounded cords, 5, 7 and 22 on first to third whorls respectively, two adapical cords more

widely spaced at level of sinus. Very fine spiral microsculpture in interspaces between cords. No axial sculpture

other than fine, sigmoid incremental lines. Aperture ovate, narrow, siphonal canal very short, indistinctly set off.

Inner lip evenly curved, covered by thick callus with free edge. Outer lip with thin edge but strengthened behind by

low varix, deep stromboid notch at base of siphonal canal. Anal sinus moderately deep, semi-enclosed, directed

slightly adapically. Colour white.

Dimensions: height 5.5 mm, diameter 1.6 mm, last whorl height 3.4 mm, aperture height 2.4 mm.

REMARKS. — In shell shape, the new species is very similar to T. vitrea from the Philippines (Iectotype

figured by KlLBURN, 1995, fig. 12), but clearly differs in having strong spiral sculpture.

Etymology. — From the Indonesian kecil (adj.), small, with reference to the small adult size. It is used here

as an invariable noun in apposition.

Genus ALICEIA Dautzenberg & Fischer, 1 897

TYPE Species: Aliceia aenigmatica Dautzenberg & Fischer, 1897.

Remarks. — The taxonomic position of Aliceia is rather enigmatic, and the species described below offers no

help. The anal sinus of the type species is shallow, whereas in A. simplicissima (Thiele, 1925) it is deep, rather

narrow, and distinctly peripheral. Such style of anal sinus suggests that the genus may belong to the family

Turridae, subfamily Turrinae, or to the bathytomine group of Clathurellinae (Conidae). At the same time, the

protoconch is more similar to some genera of the subfamily Raphitominae (Conidae), e.g., Pleurotomoides Bronn,

1831 and Famelica Bouchet & Waren, 1980. In shell outline, sinus character, and the absence of sculpture,

A. simplicissima resembles the turrine genus Lucerapex Iredale, 1936. However, Lucerapex has a paucispiral

globular smooth protoconch and nodules or scales on the peripheral keel.

Aliceia presently includes three species: the North Atlantic A. aenigmatica , the Indo-West Pacific

A. simplicissima , and the Hawaiian species figured by Kay (1979: 364, fig. 115 N) as “ Thatcheriasyrinx sp." and

suggested by Bouchet & Waren (1980) to be also a member of Aliceia. This genus thus appears to have a

world-wide distribution. Whereas A. aenigmatica and “ Thatcheriasyrinx sp.” are very similar to each other in shell

outline, and possess a broad and shallow anal sinus, a false umbilicus, and semi-tubular processes on the

periphery, A. simplicissima differs in having a slender shell lacking sculpture, with a rather deep and narrow anal

sinus. Nevertheless, the thin small shell without spiral cords or axial ribs, the more or less peripheral position of

the sinus, and the peculiar protoconch with axial pillars below peripheral keel are characters shared by all three

species, and it seems reasonable to include the species of Thiele in Aliceia, at least until more material, with soft

parts, comes to hand.

Aliceia simplicissima (Thiele, 1925)

Fig. 57

Pleurotoma (Gemmula?) simplicissima Thiele, 1925: 175 (209), pi. 23 (35), fig. 3-3a.

Source :

346

A. SYSOEV

MATERIAL EXAMINED. — Indonesia. "Valdivia" : stn 199, 0°15,5'N, 98°04'E, 470 m: 1 dd 4.8 x 2.0 mm.

Karubar, Tanimbar Islands: sin CP 69, 08°42'S, 131°53'E, 356-369 m, 1 dd 5.75 x 2.2 mm

Zanzibar. "Valdivia": stn 245, 05°27.9'S, 39°18.8'E, 463 m: 1 dd 3.8 x 1.75 mm.

Type Material. — The shell collected by the "Valdivia" at the station 245 is here designated as the lectotype.

The other shell is a paralectotype. Both are deposited at the Zoologisches Museum, Berlin, no registration number.

DESCRIPTION (Karubar specimen). — Shell slender, thin, semi-transparent, consisting of 3+ protoconch and

4.7 teleoconch whorls, suture impressed, deep. Tip of protoconch (probably protoconch I and less than 0.5 whorl

of protoconch II) broken, remaining whorls smooth above periphery, sculptured by short straight axial ribs below

periphery, indistinct near protoconch/teleoconch boundary. Teleoconch whorls obtusely angulate at periphery,

angulation stronger towards adult whorls, periphery occupied by anal sinus forming a band with sharply defined

edges. Incremental lines thin, indistinct, strongly prosocyrt above and below periphery, strongly opisthocyrt in

sinus zone. No spiral sculpture. Aperture broadly subtriangular, canal moderately long and straight. Shell colour

translucent white, with one light-brown spiral band at base of exposed whorl height, encircling base of last whorl,

and a second narrower band encircling base of canal.

Remarks. — None of the shells of the Zoologisches Museum, Berlin, corresponds to measurements given by

THIELE (5x1.8 mm), but the shell from Zanzibar (apparently figured by THIELE) is better preserved and is for this

reason designated as lectotype. It has 3.5 teleoconch and 3+ protoconch whorls.

ACKNOWLEDGEMENTS

The work was carried out during my stay as visiting curator in MNHN. I am greatly indebted to Philippe

BOUCHET who contributed from collecting the material in the field to revising the manuscript. Bruce Marshall

kindly polished the language and improved the descriptions in many ways.

APPENDIX

Check-list of deep-water turrid gastropods from Indonesia

To estimate the richness of the turrid fauna of Indonesian waters, and to provide a basis for further research, it

seems useful to give here a check-list of deep-water turrid gastropods presently known from Indonesia. This list

includes species recorded at depths greater than 200 m, in the area within the political borders of Indonesia. A total

of 92 species are listed.

The taxonomic position of species has been re-evaluated, when possible, but brief descriptions and poor

illustrations of some species allow to allocate them only tentatively to genera, awaiting the examination of

respective type material.

Source MNHN Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

347

Source : MNHN, Paris

348

A. SYSOEV

_ Species _

"Drillia" audax Melvill & Standen, 1903

Paradrillia celebensis (Schepman, 1913)

"Crassispira" aequatorialis Thiele, 1925

CONIDAE/CLATHURELLINAE

Bathytoma alractoides (Watson, 1881)

Borsonia smithi Schepman, 1913

Borsonia symbiotes (Wood-Mason & Alcock, 1891)

Borsonia timorensis (Schepman, 1913)

Borsonia epigona Martens, 1901

Borsonia jaya Sysoev, 1997

Maoritomella batjanensis (Schepman, 1913)

Typhlosyrinx supracostata (Schepman, 1913)

Glyphostoma cara (Thiele, 1925)

Heteroturris gemmuloides Sysoev, 1997

Heteroturris serta Sysoev, 1997

MANGEUINAE

Benthomangelia trophonoidea (Schepman, 1913)

Benthomangelia gracilispira (Powell, 1969)

Anticlinura biconica (Schepman, 1913)

"Mangelia terpnisma" abyssicola Schepman, 1913

Guraleus? verhoeffeni (Martens, 1904)

Guraleus halmahericus (Schepman, 1913)

Guraleus (Euguraleus) savuensis (Schepman, 1913)

Stellatoma rufostrigata (Schepman, 1913)

Heterocithara sibogae Shuto, 1970

RAPH1TOMINAE

Neopleurotomoides rufoapicatus (Schepman, 1913)

Isodaphne perfragilis (Schepman, 1913)

Pagodidaphne gradata (Schepman, 1913)

Pagodidaphne schepmani (Thiele, 1925)

Cryptodaphne affinis (Schepman, 1913)

Cryptodaphne abbreviata (Schepman, 1913)

Cryptodaphne ( Acamptodaphne) biconica (Schepman, 1913)

Source : MNHN, Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

349

REFERENCES

Barnard, K.H., 1963. — Deep sea Mollusca from west of Cape Point, South Africa. Annals of the South African Museum,

46 (17): 407-452.

Beets, C., 1942. — Notizen iiber Thatcheria Angas, Clinura Bellardi und Clinuropsis Vincent. Leidsche Geologische

Mededeelingen, 13 (1): 356-367.

Beu, A.G. & Maxwell, P.A., 1990. — Cenozoic Mollusca of New Zealand. New Zealand Geological Survey

Paleontological Bulletin, 58: 518 pp.

BOSCH. D.T., Dance, S.P., Moolenbeek, R.G., & Oliver, P.G.. 1995. — Seashells of Eastern Arabia. Dubai: Motivate

Publishing. 296 pp.

Bouchet, P. & War£n, A., 1980. — Revision of the North-East Atlantic bathyal and abyssal Turridae (Mollusca,

Gastropoda). Journal of Molluscan Studies, suppl. 8: 119 pp.

Emerson, W.K. & McLean, J.H., 1992. — Buridrillia deroyorum, new species from the Galapagos Islands, a living record

of a Neogene turrid genus. Nautilus, 106 (1): 39-42.

Kay, E.A., 1979. — Hawaiian marine shells. Bernice P. Bishop Museum Special Publication 64(4). Bishop Museum

Press, Honolulu. 655 pp.

Kilburn, R.N., 1995. — Turridae (s.l.) of southern Africa and Mozambique (Mollusca: Gastropoda, Conoidea). Part 8.

Conidae: subfamily Mangeliinae, section 3. Annals of the Natal Museum, 36: 261-269.

Source : MNHN Paris

350

A. SYSOEV

Matsumoto, Y., 1979. — Molluscan shells of Mie Prefecture, Japan. Toba Aquarium, 179 pp.

Powell, A.W.B., 1942. — The New Zealand Recent and fossil Mollusca of the family Turridae. Bulletin of the Auckland

Institute and Museum, 2: 192 pp.

Powell, A.W.B., 1964. — The family Turridae in the Indo-Paciftc. Part 1. The subfamily Turrinae. Indo-Pacific Mollusca,

1 (5): 227-346.

Powell, A.W.B., 1966. — The molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute and

Museum, 5 : 184 pp.

Powell, A.W.B., 1967. — The family Turridae in the Indo-Pacific. Part la. The subfamily Turrinae concluded. Indo-

Pacific Mollusca, 1 (7): 409-431.

Powell, A.W.B., 1969. — The family Turridae in the Indo-Pacific. Part 2. The subfamily Turriculinae. Indo-Pacific

Mollusca, 2 (10): 207-415.

Rossi Ronchetti, C., 1955. — I tipi della “Conchiologia fossile subapennina" di G. Brocchi. II. Gastropodi, Scafopodi.

Rivista Italiana di Paleontologia e Stratigrafta, 5 (2): 91-343.

Schepman, M.M., 1913. — The Prosobranchia of the Siboga-Expedition. Part 5. Toxoglossa. Siboga-Expeditie

Monograph, 49: 365-452.

Shuto, T., 1970a. — Taxonomic notes on the turrids of the Siboga-collection originally described by M. M. Schepman

1913 (Part II). Venus. 28 (4): 161-178.

Shuto, T., 1970b. — Taxonomic notes on the turrids of the Siboga-collection originally described by M, M. Schepman

1913 (Part II). Venus, 29 (2): 37-54.

Shuto, T„ 1971. — Taxonomic notes on the turrids of the Siboga-collection originally described by M. M. Schepman

1913 (Part III). Venus, 30 (1): 5-22.

Shuto, T„ 1983. — New turnd taxa from the Australian waters. Memoirs of the Faculty of Science, University - of Kyushu

ser. D, Geology, 25 (1): 1-26.

Sysoev, A.V., 1996. — Deep-sea conoidean gastropods collected by the John Murray Expedition, 1933-34. Bulletin of

the Natural History Museum, London, (Zoology), 62 (1): 1-30.

Taylor. J.D., Kantor, Yu. 1. & Sysoev, A.V., 1993. — Foregut anatomy, feeding mechanisms, relationships and

classification of the Conoidea (= Toxoglossa) (Gastropoda). Bulletin of the Natural History Museum, London

(Zoology), 59 (2): 125-170.

Thiele, J„ 1925. — Gastropoda der deutschen Tiefsee-Expedition. II Teil. Wissenschaftliclie Ergebnisse der deutschen

Ttef see-Expedition auf dem Dampfer "Valdivia" 1898-1899, 17 (2): 1-348 [35-382],

Source : MNHN, Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

351

Figs 11-18. — Genera Gemmula, Nihonia, and Clavosurcula. 11-12, 16, Gemmula closterion: 11, paratype,

Karubar stn CP 79, 22.1 x 5.8 mm; 12, holotype; 16, paratype, Karubar stn CP 67, 14.9 x 4.3 mm. —

13-15, Nihonia maxima, holotype. — 17-18 .Clavosurcula schepmani, holotype.

Source : MNHN, Paris

352

A. SYSOEV

Figs 19-28 — Genera Comilas, Heteroturris, and Cryptodaphne. 19-20, Comitas rex, hololype. — 21-23 Heteroturris

fmrP 7S 97« QfralyPe' ^R,U,B*R S,n CP 59‘ 385 x 111 mm- 22- h0l0'ypc; 23. paratype, Musorstom 2

f" J ,* 27 8 x 9-> mm- — ,24’27- Cryptodaphne rugosa: 24-25, hololype; 26, paratype, Karubar sin DW 24,

12.8 x 5.1 mm; 27, paratype, Karubar sin DW 44. 7.7 x 3.4 mm. - 28, Heteroturris serta, hololype.

Source : MNHN Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

353

Figs 29-38. — Genus Gymnobela. 29-33, G. ioessa: 29, holotype; 30-33, paratypes, Karubar stn CP 54,

46.6 x 16.4, 37.2 x 13.9, 35.9 x 14.2, and 30.3 x 12.9 mm, respectively. — 34-38, G. muricata: 34, holotype;

35-36, paratype, Karubar stn CP 73, 52.1 x 20.6 mm; 37-38, paratypes, Karubar stn CP 54, 29.4 x 12.8 and

43.3 x 17.5 mm, respectively.

Source : MNHN, Paris

354

A. SYSOEV

sm cp 91 ~f>?TT™YSOn,aJ?d Gymnobe'a-39-44-Borsonia Java: 39, holotype; 40, paratype, Karubar

nCP 77 so * 521 1 A' paraIype' Karubar stn CP 87, 58.0 x 20.8 mm; 42, paratype, Karubar

2 Cp 89' 70 5 x 22 2 mm' K\RU,BAR s,n GP 87’ 438 x >6.0 44. paratype. Karubar

holotype ' ^ 22 2 ~ 45'46' Gymnobela mitr<>deta , holotype. — 47-48, Gymnobela micraulax ,

Source : MNHN. Paris

MOLLUSCA GASTROPODA CONOIDEA FROM EASTERN INDONESIA

355

«3~

stn CP 69, 5.75 x 2.2 mm. — 58. Thelecytharella kecil, holotype.

Source : MNHN, Paris

ATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 16

RESULTATS Dt

Mollusca Cephalopoda: Mid-depth octopuses

(200-1000 m) of the Banda and Arafura Seas

(Octopodidae and Alloposidae)

Mark D. NORMAN * F.G. HOCHBERG**

& C.C. LU ***

* Department of Zoology, University of Melbourne

Parkville, VIC 3052. Australia

-“Invertebrate Zoology. Santa Barbara Museum of Natural History

2559 Puesta del Sol Road.. Santa Barbara, CA 93105. USA

-“‘Invertebrate Zoology, Museum of Victoria, 328 Swanston Walk

Melbourne, VIC 3000, Australia

ABSTRACT

»»«• - - tte -'»■ ™

depth distributions and phylogenetic affinities of this fauna are discussed.

RESUME

Mollusca Cephalopoda i Pl.uvr.s baibyales <200-11100 m) de. mere de Band. «t d'Aralur.

(Octopodidae et Alloposidae). . . , , •

x, ,, TV unrUpppr FG * I ti C C 1997 — Mollusca Cephalopoda: Mid-depth octopuses (200-1000 m) of

Norman, M.D., Hochberg, F.G. & Lu. (-^ - • prosnier & P Bouchet (eds), Resultats des Campagnes

the Banda and Arafura Seas (Octopodidae and Alloposidae) /«. A Cros N.er < 1SBN 2-85653-506-2.

MUSORSTOM, Volume 16. Mem. Mus. naln. Hist, nat., 172 . 357

Source : MNHN Paris

358

M.D. NORMAN, F.G. HOCHBERG & C.C, LU

199 et 869 metres de profondeur. Deux especes nouvelles sont decrites : Benthoctopus karubar sp. nov. et Octopus pyrum

sp. nov. Une autre espfece d Octopus, distincte d’O. pyrum, est prdsente, mais reste inddterminee. Le genre Pteroctopus est

signald pour la premifere fois du domaine indo-pacifique sur la base d’une femelle rdcoltde pendant la campagne Karubar,

et ce materiel est conspecifique avec des individus males rdcoltes en Nouvelle-Caledonie et au Vanuatu. La presence

d'Eledone palari en mer d’Arafura dtend un peu au nord l'aire de distribution connue de cette espbce, rdcemment decrite

d'Australie. Enfin, l'octopode pelagique Haliphron atlanticus (plus connu sous le nom Alloposus mollis) est egalement

reprdsentd par un seul individu, une femelle prd-mature. La repartition bathymdtrique et les affmites phylogdn<5tiques de

cette faune sont discutees par rapport ii celles d'Australie et du reste de l'lndo-Pacifique.

INTRODUCTION

The octopod fauna of the tropical Indo-West Pacific region is still poorly known, despite many species having

high profiles in commercial and subsistence fisheries. Recent research into shallow-water octopuses of this region,

particularly the Indo-Malayan region, has encountered a diverse, largely undescribed fauna (NORMAN, 1992a,

1992b, 1993a, 1993b, 1993c, 1993d; Norman & Hochberg, 1994; Norman & Sweeney, 1997; Norman, in

press).

Octopuses from beyond the continental shelves (> 200 m) in the Indo-West Pacific have received even less

attention than shallow taxa. This is a product of the few deep-water surveys undertaken in this region and/or the

limited retention of cephalopod material on such cruises. The expeditions of the "Dana" (THORE, 1949) and

"Siboga" (Adam, 1954) are two of the few exceptions. ROBSON (1925, 1932), Voss (1967, 1988a, 1988b) and

NESIS (1987) provided reviews of deeper sea octopods of the world, collating available information on animals

found at such depths. These authors recognised that the majority of described species are based on limited (often

poorly preserved) material, from few stations. Few works have provided detailed morphological comparison of

species occurring in deeper waters [exceptions being Voss & Pearcy (1990) and Lu & Stranks (1994)].

In 1991, French-Indonesian collaboration resulted in the KARUBAR research cruise to the Banda and

Arafura Seas. A series of 91 benthic trawl stations were carried out employing the Indonesian research vessel,

"Banina Jaya /”, at depths between 100 and 1250 metres. Material collected through this cruise is lodged in the

collections of the Museum National d’Histoire Naturelle, Paris (MNHN) and the Puslitbang Oseanologi - LIPI

Jakarta (POLIPI).

In a visit to the Paris museum by the first two authors in November 1995, octopod material collected through

the Karubar cruise was examined. Benthic (incirrate) octopuses originated from 18 stations at depths ranging

between 199 and 869 metres. Finned octopods (Cirroctopoda) were also encountered at a number of stations and

this material will be treated elsewhere.

MATERIAL AND METHODOLOGY

Trawls were carried out on board the “ Baruna Jaya 1 ” from October 22 to November 5, 1 99 1 . Four gear types

were employed of which 3 captured octopods: Waren dredge, (Drague Waren, station code: DW), Beam trawl (chalut

a perche, station code: CP) and shrimp trawl (chalut a panneaux [crevettes], station code: CC). All molluscs

collected in these trawls (including cephalopods) were separated on board by P. BOUCHET, W Kastoro and

B. MEtivier.

Octopod I material was collected from 18 Karubar stations. Locality details for these stations are provided in

lable (p 379). Type material for the 2 new species described here are lodged in the cephalopod collections of the

Museum National d’Histoire Naturelle (MNHN), Paris, the Puslitbang Oseanologi - LIPI (POLIPI), Jakarta and

the Museum of Victoria (MV), Melbourne.

Morphological characters and measurements used in the descriptions below are illustrated in Figure 1 Gill

counts refer to the number of lamellae on each side of each gill (= per demibranch , an inner and outer demibranch

on each gill) Count per demibranch excludes central terminal (anterior) lamella, e.g., animals illustrated in

Figures le-f have a gill count of 10.

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

359

BO. 1. _ Orientation, terminology and meat., entente. L . left; R = right. - ortawion reMve « ~

b. whole animal, dorsal view. Arms numbered from dorsal to ventral as I to 4. AC - arm crown. DMWS dorsal

mantle white spots (sensu Packard & Sanders. 1971); FWS = frontal wh.te spots ®"P; D *S,AN” ;

1971); H = typical right-hand position of hectocotylised arm in males (left hand '" ce_na n e er ' ML - mant e

length; OC = position of ocellus in ocellate octopuses; ST = position ot stylets, TL - iota >eng^ •_

animal ventral view — d, oral view of arms and webs in males: AL = arm length; H = hectocotylised arm. LG - ligula

SCR- spermatophore groove; SGU = sperma.ophore guide; WD = web depth. Webs designated from dorsal to ventral

sectors by Ers A to E - elf, mantle cavity contents: A = anus; AF = anal flaps; BH = branchial hearts; F = funnel,

FO = funnel organ (W shape shown on male, UU shape shown on female); G = = gills; ! 3 = o gan

— e mantle cavity contents of mature male: SS = spermatophore storage sac T - testis, IO - terminal organ

("penis") — f mantle cavity contents of mature female: DO = distal oviducts; LE = mature ovary as in large-egg

sDedes- SE = mature ovary as in "small-egg" species. - g, components of hectocotylised arm tip of mature male.

CL = calamus length- LL^ ligula length; SG = spermatophore groove. — h, components of spermatophores.

CB = cement body; CT = cap thread; EA = ejaculatory apparatus; SR = sperm reservoir. - 1, midsection of ejacula ory

apparatus in intact "armed" spermatophore of 0. aegma showing inward pointing teeth ( ).

Source : MNHN, Paris

360

M.D. NORMAN. F.G. HOCHBERG & C.C. LU

State of maturity is divided into 3 stages: Immature (or juvenile), Submature and Mature. In immature material

reproductive organs are not visible or tiny. Submature specimens have developed reproductive ducts (visible as

distinct terminal organ or oviducts), but lack spermatophores or a swollen ovary. Mature females possess a large

ovary, which occupies one-third or more of the mantle cavity and contains distinct individual eggs (shown for

large- or small-egg species in Fig. If).

Diagnoses and descriptions presented here are based on submature and mature specimens. Data for juvenile

material is not included as counts and relative measurements (such as sucker counts and arm lengths versus mantle

length) undergo considerable ontogenetic change in the early growth stages and can cause overlap in otherwise valid

diagnostic characters. Weights are presented for specimens preserved in 80% ethanol.

RESULTS

A total of 37 specimens of 6 species of octopods were collected from between 200 and 900 metres. Of these

species, sufficient material for 5 species was available to enable description and treatment here, including 2 new

species.

A single submature male (21.8 mm ML) of an additional species of the genus Octopus was encountered in

material from Karubar station CP 84, off the Tanimbar Islands between 246 and 275 metres. It is distinct from

the new species, Octopus pyrum, in possessing significantly shorter arms (-2 times mantle length) and a

distinctive large pink leucophore on the posterior tip of the mantle. It lacks the diagnostic band of founder

chromatophores found on the ventral mantle of O. pyrum. The submature state and poor condition of this

specimen, however, prevents identification.

Additional material of 2 species collected from the same or adjacent regions were encountered in the collections

of two Australian museums: the Northern Territory Museum in Darwin (NTM) and the Museum of Victoria

Melbourne (MV).

Species collected in the Karubar cruises and housed in the Museum National d'Histoire Naturelle, Paris and

the Puhtsbang Oseanologi, Jakarta, are presented below, with depth ranges:

Family Octopodidae

Benthoctopus karubar sp. nov.

Octopus pyrum sp. nov.

Octopus sp. indeterminate

Pteroctopus sp.

Eledone palari Lu & Stranks, 1992

410-869 m

329-511 m

246-275 m

205-620 m

200-300 m

Family Alloposidae

Haliphron atlanticus Steenstrup, 1859

284-295 m

SYSTEMATIC ACCOUNT

Family OCTOPODIDAE

Genus BENTHOCTOPUS Grimpe, 1921

Benthoctopus karubar sp. nov.

Figs 2-4, 1 la

? Benthoctopus sp. C - Adam, 1954: 186, fig. 37, pi. Ill, fig. 3.

m: 1 juv., 18.6 mm ML (MNHN

Source : MNHN. Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

361

Tanimbar Islands : stn CP 54. 836-869 m: 1 9 , 70.9 mm ML (paratype MNHN 2038 _ - Stn CC 57 603-620 m.

1 $, 96.8 mm ML (paratype MNHN 2049). - Stn CP 70. 410-413 m: 1 <3 59 6 mm ML fho^type MNHN 2026), 1 <3,

51 0 mm ML (paratype POLIPI). — Stn CP 71. 477-480 m: 1 <3. 46.0 mm ML (paratype MNHN 2027).

TYPE MATERIAL. — Holotype MNHN. Paratypes: 3 MNHN, 1 POLIPI. See above listing.

Type Locality. — Off Tanimbar Islands, Arafura Sea, 08°41’S, 131°47 E, 410-413 m.

Flr 2 _ Benthoctopus karubar sp. nov. — a, dorsal whole animal, 59.6 mm ML male holotype (MNHN 2026) Scale bar

= 20 mm. - b, stylet, 70.9 mm ML female (MNHN 2038). Scale bar = 5 mm. - c, funnel organ, 51.0 mm ML male,

paratype (POLIPI). Scale bar = 5 mm. — d, ligula, holotype. Scale bar - 5 mm.

Source : MNHN, Paris

362

M.D. NORMAN, F.G. HOCHBERG & C.C. LU

Diagnosis. — Large species, ML to at least 100 mm. Arms short to moderate, around 2-3 times ML. Dorsal

arms slightly longer than other arms. Webs deep, >33% of longest arm, approximately equal in length, ventral

web slightly shallower. Suckers moderate sized, 7-10% of ML, forming two rows. Sucker counts to around 100 in

males, 150 in females on normal arms, 47-55 on hectocotylised arm of males. Enlarged suckers absent in both

sexes. Funnel organ UU-shaped. Gill count 8-9 lamellae per demibranch. Ink sac and anal flaps absent.

Hectocotylus on right third arm. Ligula large (>10% of arm length in mature males), pointed with an open

s 2 stoiS Seated - \ommS“nehM= °eSOphagOUs; P = Phreatic tissue; PSG = posterior salivary gland;'

Source : MNHN, Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

363

shallow groove. Calamus small and pointed (around 25% of ligula length). Spermatophores of moderate length

(40 5mm 70% of ML), produced in moderate numbers (26 in storage sac of holotype). Eggs large. Colour pattern:

Pink to purple on all surfaces. Oral web dark purple. Skin sculpture: scattered with small low round papillae over

dorsal head and some of mantle.

DESCRIPTION — Counts and measurements are presented in Table 2 (p. 380). Data presented below is

presented as ranges and means (latter in italics) for four KARUBAR specimens (1 mature and 1 submature male,

2 submature females).

Large species (Figs 2a, 1 la); mantle length to at least 100 mm, total length to at least 400 mm; weight to at

least 750 g. Mantle round to ovoid, longer than wide (width 48.0-74.4-97.7% of ML). Head wide (52.1 -66.2-

74.5% of ML, 72.1-97.9-108.6% of mantle width). Skin soft, semi-gelatinous in majority of specimens

examined. Eyes large, slightly pronounced. Stylets present as thin and clear non-mineralized rods (Fig. 2b) length

around 40% of ML. Mantle opening moderately wide, approximately 50% of circumference of body at leve o

opening Funnel of moderate length, approximately 35% (30.8-34.7-40.6%) of mantle length, free portion

approximately one-third (24.3-52.5-38.5%) of funnel length. Funnel organ UU-shaped (Fig. 2c) outer limbs

approximately equal in length to median ones (outer limbs 96.8-98.7-102.2% of median limbs). Funnel organ

occupies approximately 55% (54.1-56.7-59.2%) of funnel length. .

Arms moderate length, around 2-3 times (2.2-2. 7-3.1) mantle length. Arms robust, sub-cylindrical along

length, tapering in distal third. Arm autotomy at base of arms absent. Arms approximately equal in length dorsal

lateral arms slightly longer. Suckers forming 2 rows and small to moderate sized, 7-10% of mantle length, slig y

larger in female specimens (M: 7.6, 7.2; F: 9.9, 10.7% of ML). Enlarged suckers absent in both sexes. Up to

100 suckers on intact normal arms of males, up to 150 in females (maximums in males: 96, 102; females 124

146). Webs deep (deepest webs 33.1-35.4-37.7% of longest arm). Webs approximately equal in depth, ventral web

shallower than other sectors. , .

Third right arm of males hectocotylised. Modified arm slightly shorter than opposite arm, approximately tw ee

(17 16 times) mantle length and around 80% (83.7, 72.7%) length of opposite arm. Ligula large (6.4 [in

submature male], 13.3 [in holotype] % of arm length) and sharp (Fig. 2d). Liguh groove open. Hoor of groove

with medial rib lacking distinct transverse ridges. Calamus small and sharp, around 25% of ligula length (26.6 >% of

ligula in holotype). Spermatophore groove well developed, wide and thin with fine transverse ridges.

Spermatophore guide distinct, bordered by small ridges or digits of skin. Approximately 50 suckers on

hectocotylised arm (47, 53 and 55 in three males).

Gills with 8-9 lamellae on both inner and outer demibranchs, plus terminal lamella.

Digestive tract illustrated in Fig. 3a. Anterior salivary glands large, longest dimension over 50 k ot !engdi o

buccal mass. Posterior salivary glands moderate sized (equal in length with buccal mass, approximately 40 /.of

digestive gland length). Crop diverticulum well developed. Stomach bipartite. Caecum coiled to form more than

1 5 whorls, distinctly striated. Digestive gland approximately ovoid. Ink sac and anal flaps absent Buccal mass,

digestive gland and intestine covered in large purple chromatophores, potentially used as a means of masking l.gh^

produced by bioluminescen. prey. Beaks illustrated in Figs 3b-d. Upper beak with a short, shght * ^

and narrow hood (Fig. 3b). Lower beak with distinct rostrum, narrow hood and relatively parallel lateral

separated in posterior 20% (Figs 3c-d). Radula with seven teeth and two marginal plates in each ^ transverse row

(Fig 3e). Rhachidian tooth with 1-2 lateral cusps, typically 2, on each side of large medial cone. Lateral cusps n

symmetrical to slightly asymmetrical sedation, migrating from lateral to medial position over approximate y

6 7 Ma'le'gerdtalia'illustrated in Fig. 4a. Terminal organ (“penis”) in mature males T-shaped with diverJ,^lu"1

distinctly longer than distal portion of organ. Spermatophores (Fig. 4b) of moderate length W.5^6T#*tf

ML), produced in moderate numbers (26 in storage sac). Sperm reservoir under half spermatophore length ( 7^ )

containing thin sperm cord coiled in around 42 coils. Ejaculatory apparatus of spermatophore with 6-10 thin

rCg Female genTtsd^a illustrated in Fig. 4c. The largest female examined (96.8 mm ML, MNHN 2049) was almost

mature and contained around 150 large eggs, reaching 14 mm long. One of the larger eggs showing follicular folds

is illustrated in Fig. 4d.

364

M.D. NORMAN. F.G. HOCHBERG & C.C. LU

ASG

F,G reproductive°fracF appendix” A^5G m3le repr°dUCtiVe SyStem of 59 6 mm ML holotype (-MNHN 202

96.8 mm ML female!* paralyse °MNHN 2M9r-Sc!7vLSCDOb=rd\i5 i^'cT, t'ft|'_fenlaie reProducIlve system

PO = proximal oviduct. Scale bar = 10 mm. - d, submature eggs showing3 follicular folds (FFXsSfe

Source : MNHN Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

365

Colour pattern of pink to dark purple colour produced by tiny crimson to purple chromatophores. Oral web dark

purple in most specimens. Multiple (-8) irregular rows of subdermal founder chromatophores on arms. Little skin

sculpture. Dorsal head and some of mantle scattered with small low rounded papillae (Fig. 2a). Lateral ridge absent.

REMARKS. — Adam (1954) reported four specimens of Benthoctopus from the “ Siboga ” Expedition in

Indonesian waters (in the region covered by the 1991 KARUBAR cruise), which he treated as 4 distinct undescribed

taxa (species A-D). Collection depths ranged from 304 to 1886 m. Only Adam’s Benthoctopus sp. C, a female,

shows similarities with the species described here, matching gill counts and radula dentition, and sharing similar

arm lengths (longest 2.9 times mantle length), web depths (deepest 36% of longest arm) and dark skin colouration.

Adam provides no details of funnel organ shape.

Adam’s remaining species are distinct from B. karubar. Benthoctopus sp. A (female from 304 m) possesses a

W-shaped funnel organ. Benthoctopus sp. B (male from 1886 m) has a distinctive radula with very large first

lateral teeth (ADAM, fig. 36). Benthoctopus sp. D (female from 1158 m) possesses a much lower gill count, 5-6

lamellae per demibranch. Benthoctopus sp. C, collected from 794 m at 7°24’S, 118°15.2 E, was also the only

specimen of the 4 to fall within the depth range of 400-800 m reported here for B. karubar.

As noted by NESIS (1987), the genus Benthoctopus is poorly studied. Most species are ill-defined with the pro¬

posed diagnostic characters often overlapping between taxa. Of the described species of Benthoctopus, only six

species share with B. karubar a UU-shaped funnel organ and the absence of distinctly enlarged suckers in either sex.

These taxa are compared to B. karubar in Table 3 (p. 380). The only described species close to the new species

treated here is B. levis, described from the Antarctic waters surrounding Heard Island in the southern Indian Ocean.

Benthoctopus karubar females lay large eggs (at least 14 mm long) and hence hatchlings would be benthic

with limited capacity for dispersal. The distance between records for these 2 taxa, coupled with temperature

differences between Antarctic waters and those of the Banda and Arafura Seas, are sufficient to consider these

SPCV0SS & PEARCY (1990) reviewed members of the genus Benthoctopus and described 5 new species from

the north-east Pacific Ocean. All are distinct from Benthoctopus karubar, as all possess a W-shaped tunnel organ.

The systematics of the genus Benthoctopus and its relationships with members of the genus Bathypolypus

Grimpe, 1921 require significant revision. Members of the genus Benthoctopus , as it currently stands, are

primarily grouped on the basis of a single character, the absence of a functional ink sac. It is likely that ancestral

octopuses of different shallow-water lineages have been convergent in loss of the ink sac as their ancestors shifted

to lightless depths (where an ink sac offers no selective advantage). It is possible that this genus, as it currently

stands, will prove to be polyphyletic.

ETYMOLOGY. - This species derives its names from the 1991 Karubar cruise, during which the known

specimens were collected.

DISTRIBUTION. - Benthoctopus karubar is reported here from the Arafura Sea, Indonesia (near Kai and

Tanimbar Islands). Material examined here was collected between 400 and 800 m.

Genus OCTOPUS Lamarck, 1798

Octopus pyrunt sp. nov.

Figs 5-7, lib

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn CC 10, 329-389 m. 1 6, 31.8 mm ML (holotype

MNHN 2029): 1 c3 , 34.8; 2 2 , 22.4, 29.9 mm ML (paratypes POLIPI).

Tanimbar Islands: stn CP 69, 356-368 m: 1 2, 29.1 mm ML, (para.ype MNHN 2033), 1 2, 30.1 mm ML ( M In H in

^Australia. RV "Soela": stn S06/85/27, off Townsville coast, 20°25.8’S, 152°57.7'E - 20°23.6’S, 152°57.7 E,

51 1 m, coll. CS1RO: 1 <3, 22.9 mm ML (paratype MV F78818).

Type Material. — Holotype MNHN. Paratypes: 1 MNHN, 3 POLIPI, 1 MV.

Source :

366

M.D. NORMAN, F.G. HOCHBERG & C.C. LU

Type Locality. — Off Kai Islands, Banda Sea, 05°21 ’S, 1 32°30'E, 329-389 m.

diagnosis, - Small to moderate size, ML to 35 mm. Arms moderately long, around 3-4 times ML Arms

deepes^oto^oTl 'englh‘ S°metimeS slighl|y lon§er than °^r a™s. Webs moderately deep.

Suckers smaH 5 5 M f mT’ 7 'dpP!°Xmale]y eClual in dePlh' lalera> webs sometimes slightly deeper.

hectocotvS’ f of ML formmg 2 rows. Sucker counts to around 120 on normal arms, 46-55 on

hectocotylized arm of males. Enlarged suckers absent in both sexes. Funnel organ W-shaped. Gill count

Source : MNHN, Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

367

7-8 lamellae per demibranch. Ink sac present. Hectocotylus on right third arm. Ligula large (8-10% of arm length)

robust and pear-shaped with thickened sides to the ligula groove at the base. Calamus large and pointed (around

50% of ligula length). Spermatophores of moderate length (15-26 mm, 67-75% of ML), produced m low numbers

(1-3 in storage sacs). Egg size unknown. Colour pattern: Orange to crimson base colour on dorsal surfaces, cream

on ventral surfaces. Dorsal surfaces scattered with numerous raised leucophores, cream to gold in colour.

Distinctive transverse band of around 50 founder chromatophores across midsection of ventra mantle. Large

chromatophores on ventral digestive gland, visible within mantle cavity adjacent to gills. Skin sculpture, scattered

with numerous raised round papillae over dorsal surfaces, including many over eyes.

DESCRIPTION. — Counts and measurements are presented in Table 4 (p. 381). The description below is based

on 6 Karubar specimens (2 mature males and 4 submature females). Data is presented as ranges and means (latter

in italics). Raw data for the Australian male specimen is also presented in Table 4 (see Remarks below )

Small to moderate sized species (Figs 5a, 1 lb); mantle length (ML) to it least 35 mm total 1 length to at least

170 mm; weight to at least 19 g. Mantle ovoid, longer than wide (width 55.2-7/. 7-87.9% of ML; 50.2%

slightly squashed Australian male). Head of moderate width (47.5-54.7-63.8% of ML. 67.0-76.7-86.9 % of mantle

width 52 8% and 109.0% respectively for Australian male). Eyes large, slightly pronounced. Stylets present as

thin and clear non-mineralized rods (Fig. 5c), length around 25% of ML. Mantle opening moderately wide

approximately 50% of circumference of body at level of opening. Funnel of moderate length, approximately

40% of mantle length (36.5-47.7-45.1% of ML, 36.7% in Australian male) : free portion ^ '

funnel length (42 2-47.5-52.3% of funnel length, 41.7% in Australian male). Funnel organ W-shaped (Fig. 5d)

luter limbs shorter than median ones (outer limbs 71.4-70-82.5% of median limbs. 833)% m Australian male).

Funnel organ occupies approximately 50% of funnel length (45.0-57.7-57.9%, 55.9% in Australian male) _

Arms of moderate length, around 3-4 times (2.8-J.2-3.9 x ML, 2.8 in Australian male) mantle length. Arms

sub-cylindrical along length, tapering evenly along length to narrow tips. Ann autotomy at base of Ttrms absent

Arms approximately equal in length, lateral arms sometimes slightly longer Suckers sma L le s tha 8% f

mantle length in both sexes (5.5-0-7.6% of ML, 6.1% in Australian male), forming 2 rows. Enlarged suckers

absent in both sexes. Up to 126 suckers on intact normal arms in submature and mature specimens (maxim

106 to 126 immaterial examined). Webs of moderate depth (deepest webs 19.4-22.6-27.8% of longest arm, 25.4%

in Australian male). Webs approximately equal in depth, lateral webs slightly deeper in some specimen^

Third right ann of males hectocotylized. Modified ann short, approximately twice mantle length ( .8-2 0 times

ML 1 7 in Australian male) and around 65% length of opposite arm. Ligula large (8.0-103)% of arm length, 0%

m Australian male), robust and pear-shaped (Fig. 5e). Ligula groove deep with

near shape. Floor of groove with medial rib and transverse ridges. Calamus large, narrow and sharp, around 50 h of

Hgula length (45.3 and 55.8% of ligula in Karubar males, 51.4% in Australian male). Spermatophore groove

wd. devefped. wide and thin with fine transverse ridges. Spermatophore guide distinct, bordere by flattened

papillae or digits of skin. Approximately 50 suckers on hectocotylized arm (46 and 55 in KARUBAR males,

Australian male).

Gills with 7-8 lamellae on both inner and outer demibranchs, plus terminal lamella.

Digestive tract illustrated in Fig. 6a. Anterior salivary gland length approximately one third length of

buccaf mass Posterior salivary glands large (almost as large as digestive gland). Crop diverticulum well devel¬

op dS^ach bipartite. Caecum coiled ,o§ form single whorl, distinctly striated. Digestive gland approximately

ovoid with no evidence of pancreatic tissue. Ink sac present but small, partially embedded m ventral surface of

digestive aland. Anal flaps present. Beaks illustrated in Figs 6b-d. Upper beak with a short slightly hooked

rostrum and narrow hood (Fig. 6b). Lower beak with worn rostrum in both d,ssecte^ d 2

hood, widely spread wings and flared lateral walls separated in posterior third Fig. 6d). Radula with 7 teeth a

marginal plates in each transverse row (Fig. 6e). Rhachid.an tooth with -2 la, era cusp on each **£££

medial cone. Lateral cusps in asymmetrical seriation, migrating from lateral to med.al position over 5-6 transverse

'^Male genitalia illustrated in Fig. 7a. Terminal organ (“penis”) in mature males T-shaped with diverticu^m

equal in size to distal portion of organ. Holotype w.th spermatophore within terminal organ ,cau^g

diverticulum to stretch forming a plate-like portion (Fig. 7c). Spermatophores (F,g. 7b) of moderate length

368

M.D. NORMAN, F.G. HOCHBERG & C.C, LU

(15-26 mm. 67-75% of ML), produced in low numbers (1-3 in spermatophore storage sac). Sperm reservoir under

half spermatophore length (38.5%), containing fine to robust sperm cord (coiled in around 17 coils in partially

discharged spermatophore of Karubar paratype, around 32 coils in better condition spermatophore of Australian

m®.le); E^CuIat°ry apparatus of spermatophore with irregularly placed segments, followed by a distinctive while

cylindrical portion towards oral end.

''^Symbo^asTFig.Ta! plus"lS = ink’sac^SbaT- ^ loTm’ IT T "T Para‘ype (P0LIPI): a' digestive tract.

2 mm. - c, lateral view’ of lower'^.-d.'^nTraT viewTf ^ S-Tr JST ^ ^ ^ ** a" beakS =

Source : MNHN. Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

369

No mature females were encountered in this study. Egg size and number unknown.

Basal colour pattern of orange to crimson on dorsal surfaces, cream on ventral surfaces. Dorsal surfaces covered

in numerous round papillae, cream to gold in colour, colouration probably produced by fixed leucophores (Fig. 5af

Blue-black subdermal colour often visible over each eye. Ventral mantle with diagnostic transverse band of around

50 founder chromatophores at anterior third of mantle (Fig. 5b). Single row of founder chromatophores down

midline of ventral arms. Large chromatophores also visible on ventral visceral envelope within mantle cavity,

^SkTn sculpture of loose granular texture formed by raised leucophore papillae over dorsal mantle, head, arm

crown and arm bases. Lateral ridge absent.

holotype: BH = branchial heart; G = gill; RP = renal papilla. Scale bar = 5 mm.

Source : MNHN. Paris

370

M.D. NORMAN, F.G. HOCHBERG & C.C. LU

Remarks. — The mature male specimen from off north-east Australia matched the Indonesian material in all

characters and is accordingly included here. This specimen is slightly distorted but still possesses the diagnostic

ligula, ventral mantle founder chromatophores, leucophores and matching sucker and gill counts.

There is only one recognised taxon which shows similarities with Octopus pyrum. The distinctive pear-shaped

ligula with the swollen basal edges (Fig. 5e) shows some similarities with that of Bathypolypus valdiviae

described by Thiele (in Chun, 1915, pi. LXXX) from 500 m on the Agulhas Banks, off South Africa. These two

taxa, however, are easily distinguished by the following characters in B. valdiviae : absence of an ink sac and crop

diverticulum; a VV-shaped funnel organ; shorter arms (twice mantle length); deeper webs (33-40% of longest arm);

larger ligula (13-18% of hectocotylized arm) and a radula with a rhachidian tooth which lacks lateral cusps.

ETYMOLOGY. — Derived from the Latin pyrum meaning "pear", which refers to the distinctive pear-shaped

ligula of this species.

DISTRIBUTION. — Octopus pyrum is reported here from the waters of the Banda and Arafura Seas, Indonesia

(from off the Kai and Tanimbar Islands) and north-east Australia (off Townsville, Queensland). Indonesian material

was trawled from between 329-389 m, while the male specimen from off Townsville, Australia was collected from

511m.

Genus PTEROCTOPUS P. Fischer, 1882

Pteroctopus sp.

Figs 8-10, 11c

Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn CC 10, 329-389 m: 1 9 , 50.3 mm ML (MNHN

2030). — Stn DW 18, 205-212 m: 1 9, 43.0 mm ML (MNHN 2019).

Tanimbar Islands: stn CC 57, 603-620 m: 1 9 , 30.2 mm ML (MNHN 2064).

diagnosis. — Moderate sized species, ML to at least 50 mm. Small eye openings, broad head. Narrow mantle

opening, less than 40% of circumference of body at level of opening. Arm lengths approximately 3 times mantle

length. Arms approximately equal in length. Webs deep, >30% of longest arm length. Webs approximately equal

in depth, lateral webs slightly deeper. Webs extend as membranous flared margins along entire length of arms, very

well developed at arm tips. Suckers small, around 7% of ML, forming 2 rows. Sucker counts to around 150 in

females on normal arms. Enlarged suckers absent in females. Large VV-shaped funnel organ. Gill count 8-9

lamellae per demibranch. Ink sac and anal flaps present. Eggs appear large-type in submature ovary. Colour

pattern: crimson brown dorsally, pink ventrally. Skin sculpture: scattered with regular small round papillae over all

dorsal surfaces. Two narrow elongate papillae over each eye.

Description. — Counts and measurements are presented in Table 5 (p. 381). Data presented below are raw

data for the 2 larger Karubar females, both submature.

Moderate sized species (Figs 8a, 11c); mantle length to at least 50 mm, total length to at least 230 mm;

weight to at least 1 10 g. Mantle ovoid, slightly flattened dorso- ventrally and longer than wide (width 77.4, 89.9%

of ML). Head wide (64.7, 80.7% of ML, 83.5, 89.8% of mantle width). Skin soft, semi-gelatinous. Eyes moderate

sized with small openings (Figs 8a, 8e). Stylets present as thin and clear non-mineralized rods (Fig. 8c), length

around 40% of ML. Mantle opening narrow, less than 40% of circumference of body at level of opening (Fig. 8b).

Funnel of moderate length, approximately 40% of mantle length (43.9, 35.1% of ML), free portion short

approximately 20% (22.6, 20.5%) of funnel length. Funnel organ VV-shaped (Fig. 8d), outer limbs approximately

^ T^ia?f°neS (°Uter 'imbS 8?1’ ,08 0% 0f mcdian limbs)' Funnel organ occupies approximately

4U-5U/c (39.4, 56.3%) of funnel length.

Arms of moderate length, around 3 times (2.9, 3.3 x ML) mantle length. Arms moderately robust, sub-

cylindr.cal along length, tapering in distal 20%. Arm autotomy at base of arms absent. Arms approximately equal

in length. Suckers small, around 7% (6.3, 7.0%) of mantle length, forming 2 rows. Enlarged suckers absent in

Source : MNHN, Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

371

F|G. 8. - Pteroctopus sp. - a. dorsal whole animal, 50.3 mm ML female (MNHN 2030)_ Scale bar - 20 mrm

h ventral mantle 43.0 mm ML female (MNHN 2019). Scale bar = 20 mm. — c, stylet, 50.3 mm ML female (MNHN

2030) Scale bar = 5 mm. — d. funnel organ, 50.3 mm ML female. (MNHN 2030). Scale bar - 5 mm. e. lateral view

of head of 43.0 mm ML female (MNHN 2019), showing pair of papillae over each eye. Scale bar - 5 mm. f, flared

web margin on distal arms, 43.0 mm ML female (MNHN 2019). Scale bar - 5 mm.

Source : MNHN, Pahs

372

M.D. NORMAN, F.G. HOCHBERG & C.C. LU

females. Up to 150 suckers on intact arms of females (maximums in females: 144, 148). Webs deep (deepest webs

31.7, 36.3% of longest arm). Webs approximately equal in depth, laterals slightly deeper than dorsal and ventral

webs. Webs extend as membranous flared margins along entire length of arms, very well developed at arm tips

(Fig. 8f).

Gills with 8-9 lamellae on both inner and outer demibranchs, plus terminal lamella.

FK\vmh«u « °C,^“S-Sp- 7 a7 di.*?tive tract components of 43.0 mm ML female (MNHN 2019): a, digestive tract.

Symbol^as in Fig. 3a plus IS = ink sac. Scale bar = 10 mm. - b, upper beak, lateral view. Scale bar for all beaks =

2 mm. c, lower beak, lateral view. — d. lower beak, ventral view. — e, radula.

Source : MNHN Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

373

Digestive tract illustrated in Fig. 9a. Anterior salivary glands small, less than 20% of length of buccal mass.

Posterior salivary glands moderate sized (longest dimension almost equal in length with buccal mass,

approximately 40% of digestive gland length). Crop swollen, but not distinctly branched to form a diverticulum.

Stomach bipartite. Caecum coiled in single whorl, distinctly striated. Digestive gland approximately ovoid. Ink sac

and anal flaps present. Beaks illustrated in Figs 9b-d. Upper beak with a short, slightly hooked rostrum and

moderate hood (Fig. 9b). Lower beak with rounded rostrum, narrow hood and relatively parallel lateral walls

separated in posterior 15% (Figs 9c-d). Radula with 7 teeth and 2 marginal plates in each transverse row (Fig. 9e).

Rhachidian tooth with 1-2 lateral cusps, typically 1, on each side of large medial cone. Lateral cusps in

symmetrical seriation, migrating from lateral to medial position over approximately 4 transverse rows.

No males encountered in the KARUBAR cruise. A full description of male material from the Coral Sea and New

Caledonia is in preparation.

Submature female genitalia illustrated in Fig. 10a. Ihe

larger female (50.3 mm ML, MNHN 2030) was submature

with a small ovary, but possessed well-developed muscular

distal oviducts. Eggs were forming in clusters throughout the

submature ovary (Fig. 10b). These eggs are already 4 mm

long in this small ovary and would thus be large-type eggs

when mature (>10% of mantle length).

Basal colour crimson brown dorsally and pink ventrally,

formed by tiny crimson chromatophores. Skin sculptured in

numerous small rounded papillae in regular texture over all

dorsal surfaces (Figs 8a, 11c). Few slightly larger bumps

scattered over dorsal mantle. Two long papillae over each eye

(Figs 8a, 8e). Lateral ridge absent.

Remarks. — Considerable confusion surrounds a

distinctive group of mid-depth octopuses, coined here as the

Pteroctopus/Danoctopus complex, identified by the following

diagnostic characters:

- muscular animals with loose, semi-gelatinous skin

sculptured in small and regular low patches,

- distinctive narrow mantle aperture,

- VV-shapcd funnel organ with narrow limbs,

- two narrow elongate papillae over each eye,

- flared web membranes extending to the arm tips, and

- a wide head with small eye apertures.

Members of this group occur in mid-depth habitats from

around 100 to 1000 m in the Atlantic, Indian and Pacific

Oceans. Within this group are two separate subgroups, (1) those species treated under the genus Danoctopus

Joubin, 1933 (and its synonyms Berry a Hoyle, 1939 and Hapaloctopus Taki, 1962) in which the hectocotylus o

males is the third right arm; and (2) others treated under the genus Pteroctopus, where the third left arm ot males is

hectocotylised. A fifth nominal genus within this group, Sasakinella Taki, 1964 is based on a single female

specimen from this group. As no males have been connected with Taki’s type specimen, the status of this generic

name remains unresolved. NESIS (1987) recognised Danoctopus, Pteroctopus and Sasakinella as valid, though ill-

defined, genera containing equally ill-defined species.

The 3 female KARUBAR specimens reported here clearly belong within this complex but were difficult to

identify at both generic and species levels. The absence of male material prevented identification at the generic

level, using the existing generic character of the side of the hectocotylised arm. Identification of these specimens

was resolved on examination of material from comparable depths in the adjacent waters of the Coral Sea and New

Caledonia, housed in MNHN. Amongst this material were numerous specimens from this group, including

Fig. 10 — Pteroctopus sp.

a-b, reproductive system of 50.3 mm ML

submature female (MNHN 2030): a, ovary.

Symbols as in Fig. 4c. Scale bar = 10 mm. —

b, cluster of immature ovarian eggs. Scale bar =

2 mm.

Source :

374

M.D. NORMAN, F.G, HOCHBERG & C.C. LU

4 males, all with the third left arm hectocotylised. This attribute clearly identifies them as members of the genus

Pteroctopus and constitutes the first record of this genus from tropical Indo-West Pacific waters. This

material matched the Karubar females in counts, dimensions, skin sculpture and colour, and we consider them

conspeciftc.

Only 2 species have been coined in the genus Pteroctopus , P. tetracirrhus delle Chiaje, 1830 (described from

the Mediterranean Sea) and P. witjazi Akimushkin, 1963 (described from the Sea of Okhotsk, Russia, far northern

Pacific). NESIS (1987) considers the differences distinguishing the latter species as minor and ill-defined. As a

consequence of this lack of resolution, we have chosen to treat the taxon reported here as an indeterminate species

of the genus Pteroctopus. Material from the Coral Sea and New Caledonia is currently being worked up and species

level resolution for this taxon will be undertaken in that work.

Th^ P'eroctopus/Danoctopus complex requires extensive revision, particularly development of characters which

may aid to identify female specimens of all genera, as well as distinguish species within each member genus.

Distribution. - Karubar material was collected off the Kai Islands, Banda Sea and Tanimbar Islands

Arafura Sea, between 205 and 620 m. Additional material discussed above from the Coral Sea and New Caledonia

was collected between 383 and 600 m.

Genus ELEDONE Leach, 1817

Eledone palari Lu & Stranks, 1992

Fig. lid

,uk5™L examined. — Indonesia. Karubar, Tanimbar Islands: stn CP 62, 246-253 nr 1 juv 21 8 mm MI

44.6-54.2; T 9 " 8 ML (Snh'nmI, 1 ' “ 1 ^ • 51 9 ML Stn CP 86, 223-225 .3 3,

p ^Australia. Arafura Sea, Northern Territory, 9°46’S. 130°14'E, 270-300 m, 15 Sept. 1987: 1 2, 23.0 mm ML (NTM

I matureCR+IP7TIObN ~ ^ f°ll0Win8 ^ description » based on Karubar specimens-

i^ed m S r6(pa3e82)and ' “ + 2 — ** ™ Serial

Moderate sized, robust octopuses (Fig. 1 Id), ML to 63 mm, we. weigh, to 120 g. Arms short 1 5-2 0 times

dTp" fromt lT L2'3A WebS ^ 5°'60 % «f longest arm length Webs decrease in

* males K in SmlVsenH W fUC5erS Smal1’ 5'7% °f M1" f°rming 3 SinglC r0W' Sucker COUnts’ UP 10 50

in eidier'scx UU \ T T* nT °f malC 3rm) with 44‘45 suckers- No enlarged suckers

r °rgaKni C°Unt 5- Ink S3C Present' Ink ducl °Pens anterior opening of anus.

Anal flaps absent. Ligula robust with large calamus. Tips of non-hectocotylized arms in males with spondform

““‘“Bed portions »p 20% of a™ ,«„gth. 4,e spe™,t„phtr„f JZt™

large «p1„T5mr )4rf mm”8 'h ejaCU'at0,7 appara,US' Spem> COrd with ~15 Eggs

dorsallv Whimwhhc?4? ML);Produced >" low "umbers. Colour pattern: pink cream-brown, to red-brown

Danillie nn h h sca tered sma11 cnmson chromatophores on ventral surfaces. Skin sculpture: pairs of large

(Fig! lid)" ma" and ^ Cr°Wn’ SCt With'n nng °f Sma'ler paP"lae- Later., ridge we", develop

mnr!h!|ARI(S TV ,The onginal description of the species by Lu & Stranks (1992) provided details of the

d's,,w're — p» — - £

Source : MNHN, Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOS1DAE OF EASTERN INDONESIA

375

Stylets, the shell vestige of octopodids, are well developed in Eledone palari. They are keratin-like (non-

mineralised) and approximately one-third of mantle length (21 mm in 62.8 mm ML female).

The inner margin of the mantle aperture is distinct from the form common to other members of the family

Octopodidae. A well-developed flange on the posterior margin of the funnel corresponds with a matching deep fold

inside the anterior edge of the ventral mantle. This structure is reminiscent of the flap-type mantle locking

apparatus found in the pelagic octopods, Tremoctopus violaceus delle Chiaje, 1830 (Family Tremoctopodidae) and

Haliphron atlanticus (Family Alloposidae).

Sucker counts on normal arms are low. Males possess up to 50 suckers proximal to the modified glandular

tips. Females possess up to 76 suckers on the normal arms, which lack the glandular tissue found in the males.

Spermatophores are produced in low numbers (1-5 in storage sac) and are ‘•armed”. The inside of the ejaculatory

apparatus is armed with sharp inward pointing teeth (as for example in Fig. li), which would splay out on the

outer surfaces of the spermatophore on eversion. Mangold (1989) illustrated such armature in the intact and

everted spermatophores of Eledone cirrhosa (Lamarck, 1798). .

The generic status of this octopus requires review as it has a suite of morphological characters distinct from

remaining species of the genus Eledone. All other members of the genus Eledone are restricted to the Atlantic

Ocean and Mediterranean Sea.

DISTRIBUTION — Karubar material was collected from east of the Tanimbar Islands in the Arafura Sea,

between 8°S and 10°S, at depths of around 200 to 300 m. LU & STRANKS (1992) report the distribution of this

species as the continental slope of Australia, from the Great Australian Bight, South Australia north to the

southern Arafura Sea (~10°S), at depths between 1 10 and 620 m. All depth records reported by Lu & Stranks

from shallower than 200 m are from cooler latitudes, south of 26°S.

Family ALLOPOSIDAE

Genus HALIPHRON Steenstrup, 1859

Haliphron atlanticus Steenstrup, 1859

Fig. lie

Synonyms: Alloposus mollis Verrill. 1880; Heptapus danai Joubin. 1929. (See full synonymy in THORE, 1949).

MATERIAL EXAMINED. — Karubar, Tanimbar Islands : stn CP 78. 284-295: 1 2, 73.2 mm ML (MNHN 20461.

DESCRIPTION. — Counts and measurements are provided in Table 7 (p. 382). Mantle length 73.2 ram, total

length approximately 300 mm, wet weight 165 g. Mantle round and semi-gelatinous with no constriction between

the mantle and the head (Fig. 1 le). Head broad, as wide as mantle. Eyes almost ventro-lateral m aspect. Mantle

opening very wide, >60% of circumference at level of opening (Fig. 1 le). Funnel-locking apparatus present as

transverse flap on funnel, locking into corresponding crease on mantle wall. Arms short, longest 1.7 ' tunes .

Arm formula unclear due to damaged arms in this specimen. Webs deep, >40% of longest arm length. Undamaged

webs of similar depths. Suckers small, 6.0% of ML, first 6 suckers forming a single row from mouth to web

margin, remaining suckers in 2 rows. At least 50 suckers on third arms, with 10-12 tiny suckers on tips. No

enlarged suckers in single specimen. Large W-shaped funnel organ, lateral limbs equal in length with medial ones

(Fig. lie). Gill count 10 in outer demibranch, 9 in smaller inner demibranch. Ink sac and rudimentary anal flaps

present with ink duct emerging anterior to opening of anus. Single female immature, reproductive tract not

developed. Colour pattern: large pink chromatophores on smooth, loose and torn skin. Specimen coated in

coagulated mucous and/or fine mud.

REMARKS. — We follow the works of KRISTENSEN & KNUDSEN (1983) and WiLLASSEN (1986) in adopting

the name Haliphron atlanticus Steenstrup, 1859 as senior synonym over the name Alloposus mollis Verrill. 1880.

376

M.D. NORMAN, F.G. HOCHBERG & C.C. LU

ORE (1949) treated this taxon under the name A. mollis, reporting a large submature female (400 mm total

length) from the Banda Sea (05°52’S, 131°14’E), close to the collection site for the Karubar specimen

Our record is based on a single immature female, 73.2 mm in mantle length, -300 mm in total length and

weighing 165 grams. WlLLASSEN (1986) reported two much larger females from off the coast of Norway the

largest being mature at a mantle length over 450 mm and weighing over 4. 1 kg.

Little is known of the biology of this semi-gelatinous pelagic octopus. Thore discussed the funnel locking

apparauis and muscular septa within the mantle cavity of this taxon, stating that “these septa give the impression

at the power of expulsion of water from the mantle cavity must be great, thus making the animal a rather fast

comnared "TTh Ff ^ Pr°PU,Si°n for >bc — ,ty of materia, captured in JawL

IXXle, 1885er C3t ma °lher’ POtentia"y Sl0Wer' semi-Selalinous octopods such as Japetella

™enhabTf th!S Speci“ are n0t known- THORE (I949> caPtured specimens of this species between 20

and 350 m where the sea floor was over 2400 m deep. Other specimens have been collected in ben

ic dredges (Verrill, 1882). Our specimen was collected in a beam trawl around 290 m, however

,hHe anima‘aS thG net travelled through the water column. Alvarino

LTlhLd of L aV0V 6 k l SPeCICS 35 °bserva,ions ^ a young animal in surface waters

idendficalTon § ^ ** "CL ^ Specimen was eventually captured enabling

rPnnDr!SHR,hUKI0Ni The.lKARUBAR female was collecled off the Tanimbar Islands, Arafura Sea. THORE (1949)

d'ian an Pa T n °f 'hiS (Under lhe "a™ MoP°™ «*«*) “ circumgloba, m Atllmic

Indian and Pacific Oceans, between latitudes 40°N and 40°S.

UliOUSSION

depths are more gradual, dropping to around 10°C at 500 m and 8°C at 1000 m ^ ^ angCS 31

sPeckrhlavUenabL^ret°,n arHUfnd T " “ V'SibIC “ ** °Ct°PUSeS °f northe™ Australia. Over 20 shallow-water

None of .he! b 8 d from the waters of the Great Barrier Reef and northern Australia (NORMAN 1993c)

sssssrssssssss!?

Australia (see discussion of these species groups in Norman & Sween^v ^99") ? g° ‘nClUd,ng

faunas is reflected in a single Ldm ToltcTT T C°mp°S,t,0n and affinities >*ween shallow and deeper

ssssrsSSS

encountered at greater depths. ’ ' 1 Wov- 1991)- This species was not

Source : MNHN , Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

377

The phylogenetic affinities of the mid-depth KARUBAR species are less clear. Members of the genus

Benthoctopus are reported from throughout the world’s oceans (NESIS, 1987). Benthoctopus requires extensive

revision with most species placed in this genus solely on the basis of absence of an ink sac. As discussed above, it

is premature to discuss phylogenetic affinities of Benthoctopus species.

The new species described here, Octopus pyrum , shows no affinities with any known shallow-water taxa. The

distinctive pear-shaped ligula shows superficial similarities with that of Bathypolypus valdiviae , however the two

species are distinct in a suite of other characters (see above).

As discussed above, the Pteroctopus species reported here belongs in the Pteroctopus/Danoctopus group,

with members occurring in mid-depths (100-1000 m) throughout temperate and tropical latitudes of all oceans.

The wide distribution of this group suggests an older lineage, potentially dating back to times of the circumglobal

and equatorial Tethys Sea, potentially accounting for representation of this group in all oceans at these latitudes.

The anomalous Eledone palari shares a single row of suckers with the remaining members of the genus

Eledone, all of which are restricted to the temperate and tropical waters of the Atlantic Ocean and Mediterranean

Sea. It is distinct, however, in a number of morphological characters, including a different floorplan to the male

reproductive tract, normal octopodid ligula and calamus, spongiform tissue on the modified arm tips of mature

males, deep webs and distinct skin sculpture (regular paired papillae). The generic status of this species requires

revision as it is likely to be distinct from the genuine Eledone , as represented in the Atlantic Ocean.

Eledone palari is restricted to the continental slopes surrounding the entire Australian land mass between 1 10

and 620 m. It produces large eggs (up to 15 mm) and as a consequence its hatchlings would be benthic in habit

with limited capacity for dispersal. The geological stability and isolation of the Australian land mass over the last

350 million years (as it has travelled north from the fragmenting continent of Gondwana) may explain the

distribution and unique nature of this taxon. It is possible that the ancestors of E. palari were carried north on this

migrating land mass from southern polar waters, over sufficient time that direct ancestors of this unique octopus

no longer exist. Conversely, the nearest relatives may prove to be the south polar genera Bentheledone Robson,

1932, Thaumeledone Robson, 1930 and Megaleledone Taki, 1961, all of which share a hectocotylus with well-

developed calamus. All of these genera possess a single row of suckers and occur in deep and cold polar waters

between 800 and 5300 m.

The reported circumglobal distribution of Haliphron atlanticus may reflect its pelagic habit. Nothing is known

of the origins and phytogeny of this distinct animal.

Limited octopod material has been collected from greater depths (>1000 m) in tropical latitudes of the Indo-

West Pacific region. The limited material collected from these depths includes benthic octopuses placed in the

genus Benthoctopus Grimpe, 1921 (e.g., ADAM, 1954), pelagic gelatinous octopuses such as species of Japetella

Hoyle, 1885, Eledonella Verrill, 1884 and Vitreledonella Joubin, 1918 (see THORE, 1949), and finned cirrate

octopuses, such as Grimpoteuthis pacifica (Hoyle, 1885, collected from 4465 m).

The benthic octopod faunas which exist above and below 200 m in this tropical region hence appear

phylogenetically distinct. These differences suggest that there have been no successful regional or recent descents of

shallow-water lines into these darker cooler depths, at least within this region of the Indo-Malayan Archipelago. In

order to further our knowledge of the nature and origins of these little known animals, further sampling, detailed

morphological descriptions and appropriate molecular analyses are all required.

ACKNOWLEDGMENTS

The authors would like to express their gratitude to Renata BOUCHER-RODONI, and Guy, Julian and Fabrice

BOUCHER for their hospitality and support during a research visit by the first two authors to Paris in November

1995. These visits were funded through the visiting curatorship scheme of the Museum National d’Histoire

Naturelle. We would also like to thank Philippe BOUCHET and Philippe MAESTRATI (MNHN). along with

Dr BOUCHER-RODONl’s staff, for considerable assistance during this visit. Thanks to Richard WlLLAN for

assistance with material from the Northern Territory Museum, Darwin.

378

M.D. NORMAN. F.G. HOCHBERG & C.C. LU

REFERENCES

^Sib^a' Expedite, ScmS-?^" 3' ^ Cdphal°P°deS k ''exclusion d« genres Sepia, Sepiella et Sepia, euthis.

ALrZnco&J jgr records 0{Alloposus mo,lis Verrm (Cephaiopoda; oc*°p°da>

°c,opoda- *■

H° Part |WThe nTnnl^T °j IT’ SpeC'CS "f CePha,0Poda collected during the cruise of H.M.S. " Challenger

Part 1. Ihe Octopoda. Annals and Magazine of Natural History, ser. 5, 15; 222-236.

KRIfhl7«,u,-T'|KM& KNUI?fEN- l983- — A catalogue of the type specimens of Cephalopoda (Mollusca) in

the Zoological Museum. University of Copenhagen. Steenstrupia, 9. 217-227.

LU\Cn?J»SlRr5' Ju" ]"l - Eleelone Palari’ a new species of octopus (Cephalopoda: Octopodidae) from

Australia. Bulletin of Marine Science, 49 ( 1 -2)[" 1 99 1 "]: 73-87. ;

LU\n5.&fSTRABNKS'AT'N'' ‘994o_ Syn°Psis of Pareledone and Megaleledone species, with description of two new

species from East Antarctica (Cephalopoda: Octopodidae). Memoirs of the Museum of Victoria. 54: 221-242

K' <ed)- Trai,i *

,rnrPr“ - ,h“r amts — »

No«~ Oc,opodid“)' * “ ““

N°Sw u. Z T S,TeL°f ,he °T,r macr°p- *r”1’ (“^oPKto: Octopodidae) front the Great

carrier Keel Australia. Memoirs of the Museum of Victoria, 53 (2)(" 1 992"]: 267-308.

N ° desrH nr inn^f I993bt ~ °cellate octopuses (Cephalopoda: Octopodidae) of the Great Barrier Reef Australia-

MSPSw-rsawS0” 3 redeSCripti0n 0f 0c,°PUS P°ly*™ Gray, 1849. Memoirs of Museum of Victoria,

— p~.

"TeSs

Norman, M.D. & Sweeney m i (iqq7\ -n.„

the Philippine Islands. In press invertebrate Tnonomy (CephaloPoda: Octopodinae, of

P^^ttufaame^Animal^Behaviour, 19^ *780-790.^ Pa"ernS °f °C‘°PUS and ma,urallon of the response to

Robson. G.C., 1926. - The deep sea Octopoda. Proceedings of the Zoological Society of London, (1925): 1323-1356.

R°^ridsh^N^semn^(NaturarHistory^London.r359,,p|x'eP^£,^OP0^a ** " ^ °Ct0p°da the Octopodinae).

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

379

Thore, S., 1949. — Investigations on the "Dana" Octopoda, I. Dana Report, 33: 1-85.

Voss, G.L., 1967. — The biology and bathymetric distribution of deep-sea cephalopods. Studies in Tropical

Oceanography, 5: 511-535.

Voss GL 1988a. — Evolution and phylogenetic relationships of deep-sea octopods (Cirrata and Incirrata). In:

m'r. Clarke, & E.R. Trueman (eds.). The Mollusca, Volume 12. Palaeontology and Neontology of cephalopods:

253-276. Academic Press, London.

Voss, G.L., 1988b. — The biogeography of the deep-sea Octopoda. Malacologia, 29 (1): 295-307.

Voss, G.L. & Pearcy, W.G., 1990. — Deep-water octopods (Mollusca: Cephalopoda) of the northeastern Pacific.

Proceedings of the California Academy of Sciences, 47 (3): 47-94.

Willassen, E., 1986. — Haliphron atlanticus Steenstrup (Cephalopoda: Octopoda) from the Coast of Norway. Sarsia,

71: 35-40.

TABLES

Table 1. — Summary of Karubar station data referred to in this paper.

Source MNHN, Paris

380

M.D. NORMAN, F.G. HOCHBERG & C.C. LU

Table 2. — Counts and measurements (raw data, measurements in mm) for 2 males and 2 females of Benthoctopus karubar

sp. nov. D = damaged, H = hectocotylized arm.

Registration number

Karubar station:

Status

Sex

Maturity

Mantle length

Total length

Weight (g)

Mantle width

Head width

Shallowest web depth

Deepest web depth

Arm lengths (L/R): 1

2

3

4

Sucker diameter

Sucker count: R3

L3

Ligula length

Calamus length

Spermatophore number

Spermatophore length

Spermatophore width

Sperm reservoir length

Sperm cord whorls

Egg number

Egg length

Egg width

26

40.5

1.1

15.0

42

submature

-150

14

4

C°mpar,isoIn of Benthoctopus karubar sp. nov. with similar described species. AL1 = Arm Leneth Index

SDI = Su^rD"ametenrglide ^ = L!?glh Hgula iengIh/h«tocotylised arm length. - M = mature -

depth” of deepest web/Tengdi^f fonges^arm61^ d'ame,er/mamle len*Ih- ’ Sub = — ' WDI = web depth index,

Source : MNHN, Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

381

Table 4. _ Counts and measurements (raw data, measurements in mm) for material of Octopus pyrum sp. nov.

D = damaged. - H = hectocotylized arm. - pd = sperm reservoir partially discharged.

Table 5. — Counts and measurements (raw data, measurements in mm) for two females of Pteroctopus sp. collected in

Karubar cruise. D = damaged.

Source : MNHN, Paris

382

M.D. NORMAN, F.G. HOCHBERG & C.C. LU

Table 6. - Counts and measurements (raw data, measurements in mm) for 3 males and 3 females of Eledone patari

D = damaged. - H = hectocotylized arm. '

T*BS[n CpVs^MNH^O^^D^'damaged."^ ™> “ °t H^ron CanUcu,. Karubar

Source : MNHN, Paris

CEPHALOPODA OCTOPODIDAE AND ALLOPOSIDAE OF EASTERN INDONESIA

383

Fig. 1 1. — a, Benthoctopus karubar sp. nov. Dorsal view of 96.8 mm ML female, paratype (MNHN 2049). — b. Octopus

pyrum sp. nov. From left: dorsal view of male paratype (34.8 mm ML, POLIPI) and ventral views of female paratype

(22.4 mm ML, POLIPI) and male holotype (31.8 mm ML, MNHN 2029). — c. Pteroctopus sp. Dorsal view of 50.3 mm

ML female (MNHN 2030). — d. Eledone palari. From left: dorsal view of 45.5 mm ML female, lateral view of

35.8 mm ML male, ventral view of 19.7 mm ML juvenile, (all MNHN 2020). — e, Haliphron atlanticus. Ventral view

of 73.2 mm ML female (MNHN 2046). Mantle and funnel dissected open to display gills and funnel organ

respectively.

Source : MNHN. Paris

Source : MNHN, Paris

ATS DES CAMPAGNES MUsbRSTOM. VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS D

Crustacea Decapoda : Stylodactylidae recoltes en

Indonesie, aux lies Wallis et Futuna et au Vanuatu

(campagnes KARUBAR, MUSORSTOM 7 et 8)

Donnees complementaires sur les Stylodactylidae

de Nouvelle-Caledonie

Regis CLEVA

Museum national d'Histoire naturelle,

Laboratoire de Zoologie (Arthropodes),

61 rue de Buffon, 75231 Paris Cedex 05, France

RESUME

De nombreux specimens de Stylodactylidae ont etd rdcoltes, en 1991. aux ties Kai et Tanimbar (Moluques) lors

de la campagne franco-indon6sienne Karubar. Ce materiel renferme huit especes, dont une nouvelle du genre

Parastylodactylus. P. moluccensis. Deux especes sont signalees pour la premiere fois de la region : Parastylodactylus

richeri Cleva, 1 990, et Neostylodactylus affinis Hayashi & Miyake. 1968. Les cinq autres etaient deja connues de la faune

carcinologique indondsienne : Stylodactylus tokarensis Zarenkov, 1968, S. multidentatus Kubo, 1942, 5. libratus Chace,

1983. Parastylodactylus bimaxillaris (Bate, 1888), et Stylodactylus licinus Chace, 1983, cette demure signalec tout

rdcemment par TaKEDA et Hanamura (1994). Par ailleurs, certains specimens, tout d’abord identifies & S. libratus avec

doute et presentant des caractferes qui les rapprochent de Stylodactylus pubescens Burukovsky, 1990, nous posent un

problfeme qui n'a pas trouve de solution satisfaisante ; ils sont cites ici comme Stylodactylus sp. Stylodactylus

brevidactxlus Cleva, 1990, compte tenu de la variability observ6e chez 49 specimens de S. multidentatus Kubo, recoltes

lors de cette campagne, est mis en synonymie avec cette espfece.

Nous avons ajoutd au materiel indonesien, pour chacune des differentes especes, les spdcimens rapportes recemment

de Wallis et Futuna, du Vanuatu et de Nouvelle-Caledonie. Les espbces de ces trois demises regions qui ne figurent pas

parmi celles de la campagne Karubar, sont mentionnees & la fin de cette etude.

ABSTRACT

Crustacea Decapoda: Stylodactylidae collected in Indonesia, the Wallis and Futuna Islands

and the Vanuatu (Karubar, MUSORSTOM 7 and 8 cruises). Additional information on the

Stylodactylidae from New Caledonia.

During the French-Indonesian expedition Karubar off Kai and Tanimbar Islands (Moluccas) in 1991. eight species ot

Stylodactylidae were collected. One of these species, Parastylodactylus moluccensis was new. Two other species.

Cleva, R., 1997. — Crustacea Decapoda : Stylodactylidae r6coltes en Indondsie, aux lies Wallis et Futuna et

au Vanuatu (campagnes Karubar. MUSORSTOM 7 et 8). DonmSes compldmentaires sur les Stylodactylidae recoltds

en Nouvelle-Caledonie. In : A. Crosnier & P. Bouchet (eds), Resultats des Campagnes MUSORSTOM. Volume 16.

Mem. Mus. natn. Hist. nat.. 172 : 385-407. Paris ISBN: 2-85653-506-2.

Source : MNHN, Paris

386

R. CLEVA

Parastylodactylus richeri Cleva, 1990, and Neostylodactylus affinis Hayashi & Miyake, 1968, are recorded from the

region for the first time and the remaining five species, Stylodactylus tokarensis Zarenkov, 1968, S. multidentatus Kubo,

1942, S. libratus Chace, 1983, Parastylodactylus bimaxillaris (Bate, 1888), and Stylodactylus licinus Chace, 1983, are

already known from the Indonesian area, the last one having been recorded recently by Takeda and Hanamura (1994)

On the other hand, some specimens, at first identified doubtfully as Stylodactylus libratus, and related to Stylodactylus

pubescens Burukovsky, 1990, have been causing trouble to us, and we have not find till now a satisfying solution : they

are mentionned here as Stylodactylus sp. Stylodactylus brevidactylus Cleva, 1990, considering the variability observed

through 49 specimens of S. multidentatus Kubo collected during this cruise, is synonymised with this species.

We added to the indonesian material, for each different species, the specimens collected recently from Wallis and

Futuna, the Vanuatu and New-Caledonia. The species from these three countries which have not been collected during the

Karubar expedition are mentionned at the end of this study.

INTRODUCTION

La campagne franco-indonesienne Karubar, effectuee au large des ties Kai et Tanimbar (Moluques) par le

navire oceanographique Baruna Jaya I ", entre 1 50 et 1 250 m de profondeur, du 2 1 octobre au 6 novembre 1 99 1 , a

rapporte de nombreux echantillons de Stylodactylidae (Crosnier, Richer de Forges & Bouchet, 1997).

D'autres recoltes de specimens appartenant a cette famille ont ete faites, depuis, au large des ties Wallis et

Futuna, lors de la campagne MUSORSTOM 7, effectuee par le navire de FORSTOM, “Alls”, jusqu’a des profondeurs

de 1300 m. du 5 mai au 4 juin 1992 (Richer de Forges & Menou, 1993) et au large de la Nouvelle-Caledonie

lors de di verses campagnes en eaux profondes, toutes faites avec 1’ “Alis” : Beryx 1 1, du 13 au 23 octobre 1992

(Lehodey et all., 1992), Smib 8, du 27 janvier au 2 fevrier 1993, Bathus l,du 10 au 19 mars 1993, Bathus 2,

du 10 au 18 mai 1993, Bathus 3, du 23 novembre au ler decembre 1993, HALIPRO 1, du 19 au 31 mars 1994,

Bathus 4, du ler au II aout 1994 (Richer de Forges & Chevillon, 1996).

A cela s’ajoutent une recolte faite, toujours en Nouvelle-Caledonie, lors de (’expedition MONTROUZIER, a une

profondeur de 52 m (BOUCHET, 1994) et, enfin, des recoltes faites au Vanuatu, lors de la campagne MUSORSTOM 8

eifectuee a bord de 1' "Alis", du 19 septembre au 14 octobre 1994, entre 100 et 1600 m de profondeur (RICHER DE

Forges etal., 1996).

Ce sont toutes ces recoltes qui sont etudiees dans la presente note.

Les specimens etudies sont deposes dans Ies collections du Museum national d’Histoire naturelle a Paris

(MNHN), au Puslitbang Oseanologi - LIPI, a Jakarta (POLIPI) et au National Museum of Natural History a

Washington (USNM).

Les abreviations utilisees dans les pages qui suivent sont : LC : longueur de la carapace mesuree du fond de

I orbite au milieu du bord posteneur de la carapace ; LR : longueur du rostre ; MxP3, P3, P4, P5 : troisieme paire

de maxtllipedes, troisieme, quatrieme et cinquieme paires de pereiopodes.

LISTE DES STATIONS

Abreviations utilisees : CP : Chalut a Perche ; CC : Chalut a Crevettes ; DW : Drague Warren.

Karubar. Indonesie

St. CP 05. — 22.10.1991,

St. CP 09. — 23.10.1991,

St. CC 10. — 23.10.1991,

St. CP 16. — 24.10.1991

moluccensis.

St. CP 17. — 24.10.1991,

St. DW 18. — 24.10.1991

St. CP 19. — 25.10.1991,

St. CP 20. — 25.10.1991,

05°46 39 S- 1 32°20’4"E, 285-323 m : Stylodactylus tokarensis.

05 19 21 S-132°30'35"E, 361-389 m : Parastylodactylus bimaxillaris.

05 26 1 1 S-132°27'37"E, 329-389 m : Parastylodactylus bimaxillaris.

, 05° 17 06 S- 132°51'19"E, 315-348 m : Stylodactylus libratus. Parastylodactylus

05 17 03 S-133°00'24"E, 459-439 m : Stylodactylus tokarensis. S. libratus. S sp

, 05° 1 7'49"S- 1 33°00'5 1 "E, 205-212 m : Neostylodactylus affinis.

05 15 52 S-133°00'01"E, 604-576 m : Stylodactylus licinus.

05 16 30 S-132°58'20"E, 768-810 m : Stylodactylus licinus.

Source : MNHN , Paris

STYLODACTYLIDAE INDO-OUEST PACtFIQUES

387

St. CC 21. — 25.10.1991, 05°16'25"S-132°59’03"E, 688-694 m : Stylodactylus licinus.

St. DW 24. — 26.10.1991, 05o31,35"S-132°51'15"E, 243-240 m : Stylodactylus multidentatus multidentatus.

St. CP 27. _ 26.10.1991, 05°34'22"S-132°5r29"E, 304-314 m : Stylodactylus tokarensis, S. multidentatus

multidentatus, Parastylodactylus richeri, P. moluccensis.

St. DW 32. — 26.10.1991, 05°46,31"S-132o50l42"E, 170-206 m : Neostylodactylus affinis.

St. CP 33. —27.10.1991, 06o02'10"S-132o38'21"E, 281-311 m : Parastylodactylus bimaxillaris.

St. CP 35. — 27.10.1991, 06°07,22"S-132°43,45"E, 390-502 m : Stylodactylus licinus.

St. CP 36. — 27.10.1991, 06°05'50"S-132o44'29"E, 268-210 m : Parastylodactylus bimaxillaris.

St. CP 38. — 28.10.1991, 07o38,41"S-132°29,22"E. 666-620 m : Stylodactylus licinus.

St. CC 41. _ 28.10.1991, 07°47’28"S-132°39'04"E, 401-393 m : Parastylodactylus bimaxillaris.

St. CP 45. — 29.10.1991, 07°53' 11"S- 132°47'20"E, 301-305 m : Parastylodactylus bimaxillaris.

St. DW 50. — 29.10.1991, 07°59,09"S-133°01,56"E, 184-185 m : Neostylodactylus affinis.

St. CC 56. — 31.10.1991, 08o12'39"S-132°0ri5"E, 552-549 m : Stylodactylus licinus.

St. CC 57. — 31.10.1991, 08°15’48"S-131°56'38"E, 603-622 m : Stylodactylus licinus.

St. CP 66. — 01.1 1.1991, 09°02,19"S-132o10'49"E, 211-217 m : Stylodactylus multidentatus multidentatus.

St. CP 67. — 01.1 1.1991, 08°58'59"S- 1 32°07'20"E, 233-246 m : Stylodactylus multidentatus multidentatus.

St. CP 69. — 02.11.1991, 08°45' 1 7"S- 1 3 1 °5 1 '35"E, 356-367 m : Parastylodactylus bimaxillaris.

St. CP 70. — 02.1 1.1991, 08o39'14"S-131°49'16"E, 411-410 m : Parastylodactylus bimaxillaris.

St. CP 71. — 02.1 1.1991, 08°39'39"S- 1 3 1 °42'29"E, 477-480 m : Stylodactylus licinus.

St. CP 76. — 03.1 1.1991, 08°49'08"S- 13 1°35'36"E, 400 m : Parastylodactylus bimaxillaris.

St. CP 79. — 03.1 1.1991, 09° 1 3'34"S- 1 3 1 °22'35"E, 250-239 m : Stylodactylus multidentatus multidentatus.

St. CP 83. — 04.11.1991, 09o24'28"S-130o59'48"E, 285-298 m : Stylodactylus multidentatus multidentatus,

Parastylodactylus bimaxillaris.

St. CP 84. — 04.1 1.1991, 09°22'4 1 "S- 1 3 1 °07' 1 7"E, 275-246 m : Stylodactylus multidentatus multidentatus.

$t Cp 85. _ 04.1 1.1991, 09o22’51"S-131o12'04"E, 244-239 m : Stylodactylus multidentatus multidentatus.

St. CP 86. — 04.11.1991, 09°23'59"S- 1 3 1° 1 4'29"E, 226-222 m : Stylodactylus multidentatus multidentatus,

Parastylodactylus bimaxillaris.

MUSORSTOM 7. lies Wallis et Futuna.

St. CP 531. — 16.05.1992, 12°32'S-176°39’W, 580-600 m: Stylodactylus licinus.

St. CP 550. — 18.05.1992, 12°15'S-177°28'W, 800-810 m : Stylodactylus licinus.

St. CC 554. — 18.05.1992, 12°14'S-177°28'W, 795-820 m : Stylodactylus licinus.

St. CP 565. — 20.05.1992, 1 l047'S-178o25’W, 900 m : Stylodactylus licinus.

St. CP 638. — 30.05.1992, 13°37'S-179056'E, 820-840 m : Stylodactylus brucei.

Sans numero de station, 05.1992, drague, 500-610 m : Parastylodactylus semblatae.

BERYX 11. Nouvelle-Caledonie.

St. CP 08. — 15.10.1992, 24°54'S-168021'E, 540-570 m : Parastylodactylus tranterae.

St. DW 09. — 26.10.1991, 24°44'S-170°07’E, 790-825 m : Stylodactylus sp.

St. DW 27. — 18.10.1992, 23°37'S-167°4rE, 460-470 m : Stylodactylus laurentae.

St. CP 53. — 21.10.1992, 23°48'S-168°17'E, 540-950 m : Stylodactylus sp.

St. CP 58. — 22.10.1992, 23°19’S-167°59'E, 850-920 m : Stylodactylus licinus.

St. CP 59. — 22.10.1992, 23° 19'S- 1 67°59'E, 750-800 m : Stylodactylus licinus.

St. CP 60. — 22.10.1992, 23°19’S-168°00'E, 580-600 m : Stylodactylus licinus. S. sp.

SMIB 8. Nouvelle-Caledonie.

St. DW 146. — 27.01.1993, 24°55,2'S- 168°2 1 ,7’E, 514-522 m : Stylodactylus laurentae.

St. DW 150. — 27.01.1993, 24°54,3’S-168022,2’E, 519-530 m : Parastylodactylus tranterae.

St. DW 160. — 28.01.1993, 24°46,1,S-168°08,1'E, 280-282 m : Stylodactylus libratus.

St. DW 180. — 30.01.1993, 23°47,7'S-168°18,1’E, 460-525 m : Stylodactyloides crosnieri.

St. DW 183. — 31.01.1993, 23°18,3’S-168o04,9'E, 330-367 m : Stylodactyloides crosnieri.

Source :

388

R. CLEVA

St. DW 185. — 31.01.1993, 23°16'S-168°04,3'E, 305-355 m : Stylodactylus laurentae.

St. DW 187. — 31.01.1993, 23° 1 7,7'S- 1 68°05,6'E, 390-540 m : Stylodacty loides crosnieri.

St. DW 1 9 1 . - 0 1 .02. 1 993, 22°57,0'S- 1 68° 1 9,2'E, 564-580 m : Parastylodactylus semblatae

St. DW 193. - 01.02.1993, 22°58,7'S-168°20,rE. 500-508 m : Stylodactylus laurentae

St. DW 194. — 01.02.1993, 22°59,6’S-168°22,5'E, 491 m : Stylodactylus laurentae

St. DW 195. - 01.02.1993, 22°58,9'S-167°20,2'E, 508-514 m : Stylodactylus laurentae.

St. DW 201. - 02.02.1993, 22°58,6'S-167°20,3'E, 500-504 m : Stylodactylus laurentae,

semblatae.

Parastylodactylus

Bathus

St. CP 657. -

St. CP 660.

St. CP 668.

St. CP 670. -

St. CP 671. -

St. DW 687.

St. CP 701. •

St. CP 709.

Bathus

St. DW 721.

St. DW 731.

St. CP 737. -

St. CP 738.-

Sl. CP 742. -

St. DW 758.

St. CP 760. -

1. Nouvelle-Caledonie.

- 12.03.1993, 21°14'S-165°54'E, 490-530 m :

- 13.03.1993, 21o10'S-165°53'E, 786-800 m

- 14.03.1993, 20°57'S-165°34'E, 205-219 m

- 14.03.1993, 20°54'S-165°53'E, 394-397 m

- 14.03.1993, 20o51'S-165o28'E, 450-470 m :

- 16.03.1993, 20°34'S-165°07'E, 408-440 m

- 18.03.1993, 20°57'S-165°35’E, 302-335 m

- 19.03.1993, 21°4rS-166°37'E, 650-800 m

Parastylodactylus tranterae.

: Stylodactylus licinus.

: Stylodactylus multidentatus multidentatus.

Parastylodactylus richeri.

Parastylodactylus tranterae.

: Parastylodactylus richeri.

: Stylodactylus multidentatus multidentatus.

: Stylodactylus licinus.

2. Nouvelle-Caledonie.

1 1.05.1993, 22°53 S-167°I7'E. 525-547 m : Parastylodactylus semblatae.

13.05.1993, 22°49S-166°44'E, 300-370 m : Stylodactylus multidentatus multidentatus

— 13.05.1993, 23°03'S-167°00'E, 350-400 m : Stylodactylus laurentae.

— 13.05.1993, 23°02'S-166°56'E, 558-647 m : Parastylodactylus tranterae.

— 14.05.1993, 22°33 S-166°25'E, 340-370 m : Stylodactylus multidentatus multidentatus.

16.05.1993, 22 18 S-166°10'E, 377-386 m : Stylodactylus multidentatus multidentatus.

— 16.05.1993, 22° 18 S-166°10'E, 455 m : Stylodactylus multidentatus multidentatus.

Operation Montrouzier. Nouvelle-Caledonie.

Chenal de Touho, 04.09.1993, 52 m : Neostylodactylus amarynthis.

Bathus

St. DW 776.

St. DW 786.

St. DW 794.

St. DW 830.

St. CP 831.

St. CP 832. -

St. CP 833. -

St. DW 836.

St. CC 841.

St. CP 842.

St. CP 844.

St. CP 846.

St. CC 848.

3. Nouvelle-Caledonie.

- 24.1 1.1993, 24°44'S-170°08'E, 770-830 m : Stylodac^lus sp.

- 25.11.1993, 23°54'S-169°49'E, 699-715 m : Stylodacty’lus sp

- 26.1 1.1993, 23°48'S-169°49'E, 751-755 m : Sty’lodactylus sp.

29.1 1.1993, 23° 19 S- 1 68°0 1 'E, 361-365 m : Stylodactyloides crosnieri

- 30.11.1993, 23°04'S-166°55'E, 650-658 m : Stylodactylus licinus.

Z in! mcS’ 650'669 m 1 Sty‘odactylus teinus, Parastylodactylus tranterae.

0.1 1.1993, 23 03 S-166 58 E, 441-444 m : Parastylodactylus tranterae.

30.1 1.1993, 23°02 S-166°59'E, 295-306 m : Stylodactyloides crosnieri

- 30.1 1.1993, 23°02'S-166°53'E, 640-680 m : Stylodactylus licinus.

01.12.1993, 23°05 S-166°48'E, 830 m : Stylodactylus licinus.

01.12.1993, 23°06'S-166°45'E, 908 m : Stylodactylus licinus.

- 01.12.1 993, 23°02'S-166°57'E, 500-514 m : Parastylodactylus tranterae

- 01.12.1993, 23°02'S-166°52'E, 680-700 m : Stylodactylus licinus.

Halipro 1. Nouvelle-Caledonie.

I" pp «ao7' ~SnM94’ 21°26'S-166018'E. 720-950 m : Stylodactylus licinus.

St CH 874 30 03 1004 \ aa^55'^ 450~490 m : St)lodactylus multidentatus multidentatus.

St. CH 874. - 30.03.1994, 23°05'S-166°48'E, 708-830 m : Stylodactylus licinus.

STYLODACTYL1DAE INDO-OUEST PACIFIQUES

389

St. CP 877. — 31.03.1994. 23°03’S-166o59'E, 464-480 m : Stylodactylus laurentae.

BATHUS 4. Nouvelle-Caledonie.

St. CP 892. — 02.08.1994, 21°0rS-164°27'E, 580-600 m : Parastylodactylus tranterae.

St. CP 897. — 03.08.1994, 20o15'S-l63°5rE, 305-350 m : Stylodactylus multidentatus multidentatus.

St. CP 900. — 03.08.1994, 20°16'S-163°50'E, 580 m : Stylodactylus multidentatus multidentatus.

St. CP 905. — 04.08.1994, 19°02'S-163°15'E, 294-296 m : Stylodactylus multidentatus multidentatus.

St. CP 910. — 05.08.1994, 18°59'S-163°08'E, 560-608 m : Parastylodactylus bimaxillaris. P. semblatae.

St. CP 91 1. — 05.08.1994, 18°57'S-163°08’E, 566-558 m : Parastylodactylus semblatae.

St. CP 912. — 05.08. 1994, 18°55'S-163°07'E, 702-690 m : Parastylodactylus tranterae. P. semblatae.

St. CP 921. — 06.08.1994, 18°46'S-163°17'E, 613-610 m : Parastylodactylus semblatae.

St. CP 922. — 06.08.1994, 1 8°48'S- 1 63° 1 8'E, 600 m : Parastylodactylus semblatae.

St. DW 923. — 06.08.1994, 18°51'S-163°24'E, 502-470 m : Stylodactylus laurentae.

St. DW 924. — 07.08.1994, 18o54’S-163024'E, 344-360 m : Stylodactylus laurentae.

St. DW 929. — 07.08.1994, 1 8°5 1 'S- 163°23'E, 502-516 m : Parastylodactylus semblatae.

St. CP 930. — 07.08. 1994, 1 8°5 l'S-163°23'E, 530-520 m : Parastylodactylus semblatae.

St. DW 931. — 07.08.1994, 1 8°55’S-163024'E, 360-377 m : Stylodactylus laurentae.

St. DW 934. — 08.08.1994, 19o05'S-163°28'E, 231-240 m : Neostylodactylus affinis.

St. CP 953. — 11.08.1994, 2r45'S-166°36'E, 220-234 m : Stylodactylus multidentatus multidentatus.

RFO. Nouvelle-Caledonie, 12.09.1994, 22°33,4rS-166°25,74'E, 300 m : Parastylodactylus richeri.

MUSORSTOM 8. Vanuatu.

St. CP 974. — 22.09.1994, 19°2rS-169°28'E, 492-520 m : Parastylodactylus semblatae.

St. CP 980. — 22.09.1994, 1 9°2 I S- 1 69°25'E, 450-433 m : Parastylodactylus richeri.

St. CP 990. — 24.09.1994, 18°5rS-168°50'E, 980-990 m : Stylodactylus licinus.

St. CP 991. — 24.09.1994, 18°51'S-168°52'E, 936-910 m : Stylodactylus licinus.

St. CP 992. — 24.09.1994, 18052'S-168o55'E, 775-748 m : Stylodactylus licinus. S. macropus.

St. CP 1007. — 25.09.1994, 18°51'S-168°55'E, 720-830 m : Stylodactylus licinus. S. multidentatus

multidentatus.

St. CP 1024. — 28.09.1994, 17048'S-168038'E, 335-370 m : Parastylodactylus richeri.

St. CP 1025. — 28.09.1994, 17049'S-168°39'E, 385-410 m : Parastylodactylus richeri.

St. CP 1027. — 28.09.1994, 17°53'S-168039'E, 550-571 m : Parastylodactylus semblatae.

St. CC 1034. — 29.09.1994, 17°54'S-168°42'E, 690-750 m : Stylodactylus licinus.

St. CP 1035. — 29.09.1994, 17°56'S- 168°44'E, 765-780 m : Stylodactylus licinus.

St. CP 1050. — 01.10.1994, 16°39'S-168°0rE, 541-577 m : Stylodactylus licinus.

St. CP 1055. — 01.10.1994, 16°30’S-167°55'E, 572-580 m : Stylodactylus licinus.

St. CC 1056. — 01.10.1994. 16°33'S-167°55'E, 602-620 m : Stylodactylus licinus.

St. DW 1061. — 02.10.1994, 16°14'S-167°20'E, 458-512 m : Stylodactylus multidentatus multidentatus.

St. DW 1065. — 02.10.1994, 16°16'S-167°21'E, 360-419 m : Stylodactylus multidentatus multidentatus.

St. CP 1074. —04.10.1994, 15°48'S-167024'E, 775-798 m : Stylodactylus licinus.

St. CP 1075. — 04.10.1994, 1 5°53'S- 1 67°27'E, 956-944 m : Stylodactylus licinus.

St. CP 1080. —05.10.1994, 15°57'S-167027'E, 799-850 m : Stylodactylus licinus.

St. CP 1087. — 06.10.1994, 15°10'S-167°14,E, 394-421 m : Stylodactylus multidentatus multidentatus.

St. CP 1088. — 06.10.1994, I5°09'S-167°15'E, 425-455 m : Stylodactylus multidentatus multidentatus.

St. CP 1091. — 06.10.1994, I5°10'S-167°13'E, 344-350 m : Parastylodactylus bimaxillaris.

St. CP 1092. — 06.10.1994, 1 5° 1 0'S- 1 67° 1 2'E, 314-321 m : Stylodactylus multidentatus multidentatus.

St. CP 1 1 14. — 08.10.1994, 14°52'S-167°03'E, 647 m : Stylodactylus licinus.

St. CP 1 135. — 1 1.10.1994, 15°40'S-167°02'E, 282-375 m : Neostylodactylus affinis.

St. CP 1 136. — 1 1.10.1994, 15°40'S-167°0rE, 398-400 m : Stylodactylus multidentatus multidentatus.

390

R. CLEVA

St. CP 1137. — 11.10.1994, 15°4rS-167°02'E, 360-371 m : Stylodactylus multidentatus multidentatus,

Parastylodactylus richeri.

ETUDE SYSTEM ATIQUE

Genre STYLODACTYLUS A. Milne Edwards, 1881

Stylodactylus licinus Chace, 1983

Fig. 4 A-D

Stylodactylus licinus Chace, 1983 : 14, fig. 6. — Hayashi, 1986 : 93, fig. 52 (photo couleurs) ; 1991 : 41. — Cleva.

1990 : 87, fig. 3 a-j, 18 f-g ; 1994 : 58. — Takeda & Hanamura, 1994 : 17, fig. 8 a.

Stylodactylus tokarensis Zarenkov, 1968 : 58 (pro parte), fig. 2 (perdiopode 3) et fig. 3 (non autres dessins de la fig. 2 =

Stylodactylus tokarensis Zarenkov, 1968).

Stylodactylus stebbingi - Toriyama & Hayashi. 1982 : 90, 92, 95. 105 (non Hayashi & Miyake, 1968, fide Hayashi in

Baba, Hayashi & Toriyama, 1986 : 93). — King, 1984 : 178, 179 (fig.), 181 ; 1986 : 12, fig. 9 (non Hayashi &

Miyake, 1968).

Materiel EXAMINE. — Indonesie Karubar : St. CP 19, 604-576 m : 8 8 6 a 12,5 mm ; 7 $ 7 H 15 mm (4 ov )

(USNM) ; 1 2 avec bopyre (MNHN-Na 12150). — St. CP 20, 768-810 m : 10 6 6 a 13 mm ; 21 2 (9 ov et 1 non ov

avec bopyre) 6 a 15 mm (MNHN-Na 12134). — St. CC 21, 688-694 m : 9 <3 7,5 a 12,5 mm ; 22 2 (1 1 ov.) 6 & 15 mm

(POLIP1). — St. CP 35, 390-502 m : I 2 9 mm (MNHN-Na 12136). — St. CP 38, 666-620 m : 9 2 (5 ov.) 6 & 17 mm

(MNHN-Na. 12135). — St. CC 56, 552-549 m : 1 2 ov. 16,5 mm (POLIPI). — St. CC 57, 603-622 m : 1 <3 avec bopyre

16 mm (MNHN-Na 12151); 1 2 ov. 14,5 mm (POLIPI). — St. CP 71 , 477-480 m : 1 2 ov. 13,5 mm (MNHN-Na 12137)

Wallis et Futuna. Musorstom 7 : st. CP 531, 580-600 m : 1 6 10.5 mm (MNHN-Na. 14454) — St CP 550 800-

810 m : 1 2 ov. 16,5 mm (MNHN-Na 14455). — St. CC 554, 795-820 m : I 2 13 mm (MNHN-Na 14457) —

St. CP 565, 900 m ; 1 <3 14,5 mm (MNHN-Na 14456).

Nouvelle-Caledonie, Beryx 1 1 ; st. CP 58, 850-920 m : 1 6 12,5mm (MNHN-Na 14625). — St. CP 59, 750-

80°m : 1 <3 8,5mm, 3 2 10 it 12 mm (MNHN-Na 14619) ; 1 6 9 mm, 1 2 ov. 15,5 mm (MNHN-Na 14624) —

St. CP 60, 580-600 m : 1 2 1 1 mm (MNHN-Na 14621).

Bathus 1 : st. CP 660, 786-800 m : 1 S 6,5 mm, 1 2 ov. 12 mm (MNHN-Na 14617). — St. CP 709 650-800 m •

1 2 8,5 mm (MNHN-Na 14623).

3 : st CP 83 '■ 65°-658 m : 1 2 9,5 mm (MNHN-Na 14484). — St. CP 832, 650-669 m ; 1 2 7 mm (MNHN-

m3 — Sn S? 841, 64°-680 m : 1 2 12 mm (MNHN-Na 14476). — St. CP 842, 830 m : I 2 ov. 15 mm (MNHN-

N* 3 “"St. CC 844, 908 m : 1 <3 11 mm (MNHN-Na 14462). — St. CC 848, 680-700 m : I 2 ov, 12,5 mm

(MNHN-Na 14483).

( M NHN * Na°i 3 1 9 2) CP ^ ' 72°'95° m : 1 6 9 mm (MNHN-Na 14479). — St. CH 874, 708-830 m : 1 2 ov. 16 mm

mnVan!!a!U' Musorstom 8 : st CP 990, 980-990 m : 1 6 8 mm, 1 2 10,5 mm (MNHN-Na 13194) — St CP 991 936-

910m; 2 c3 12 *13 mm ,1 2 9,5 mm (MNHN-Na 13195). - St. CP 992, 775-748 m : 1 ,3 11,5 mm (MNHN-Na 13196),

I* ov' ,13 (MNHN-Na 14458). - St. CP 1007, 720-830 m : 1 2 15 mm (MNHN-Na 13197), - St. CC 1034 690-

T’A.oo!0,5 T ‘^NHN-Na l3198>- - St. CP 1035, 765-780 m : 5 <3 7,5 k 15 mm, 3 2 (I ov.) 10 &

' " '<T ™ Cn °50- 541-577 m : 2 <3 10 et 11,5 mm, 12 10 mm (MNHN-Na 13200) ; 1 2 13

r I m “JV. CP '°55:. 572'580 m : 1 6 15 mm (MNHN-Na 14459), 1 c3 16 mm (MNHN-Na 13201), 1

d 16,5 mm (MNHN-Na 14461). — St. CC 1056, 602-620 m : 2 6 9 et 9,5 mm (MNHN-Na 13202). — St CP 1074 775-

2 «sn , omm ^NHN;?a 132°3)' - SL CP i075’ 956-944 m = 1 * 13 mm (MNHN-Na 13204) - St CP 1080

799-850 m : 1 2 ov. 15 mm (MNHN-Na 13205). - St. CP 1 1 14, 647 m : 1 <3 10 mm (MNHN-Na 13206).

Le,gra,nd nombre de specimens identifies ici a Stylodactylus licinus (146 au total, dont 92 indonesiens) permet

d ctendre les variations individuelles relevees lors de notre etude de 1990 (Cleva, 1990 : 88), principalement en ce

qui concerne la longueur relative du rostre, ainsi que le nombre de ses epines. C'est ainsi que, sur les 77 specimens

dont le rostre est intact, le rapport des longueurs du rostre a la carapace (LR/LC) varie de 1 ,2 a 2, 1 (moyenne =

environ 1.65), alors qu'en 1990 nous avions releve des variations comprises entre 1,5 et 2,0. On compte de 32 a

6 epines dorsales (moyenne = 38), et de 14 a 29 epines ventrales (moyenne = 20), contre respectivement 34 a 45

et 18 a 27 releves alors (un male d'Australie, etudie en 1994, presente 12 epines ventrales seulement).

Source : MNHN, Paris

STYLODACTYLIDAE INDO-OUEST PACIFIQUES

391

Ces chiffres nous permettent de lever l'incertitude concernant l'identification d'un specimen japonais de la baie

de Tosa, prete par K.-I. HayaSHI en 1990, dont le rostre, plus court que celui des specimens que nous avions alors

examines, nous faisait douter du statut exact de cet individu (Cleva, 1990 : 90). Hayashi (1991 : 42) a de

nouveau signale S. licinus du sud de Kyushu, ou une femelle ovig^re a 6t6 capturee a 808-826 m de profondeur.

Les autres differences individuelles relevdes sont les suivantes : absence d'epines supra-orbitaires chez une jeune

femelle indonesienne de 6 mm (station CC 21), et chez un male du Vanuatu de 9 mm (station CC 1056) ; 4£me

pleuron abdominal non termine en pointe (d'un seul cote ou des deux cotes) chez une quinzaine de specimens. Ces

variations ne sont apparemment pas en relation avec la taille des animaux.

La courbure du rostre, plus ou moins accentuee, ne permet pas d'indiquer avec certitude le sexe des specimens :

on peut cependant remarquer que chez les males, et notamment les plus grands, le rostre est presque horizontal,

tandis que sa courbure vers le haut est plus prononcee chez les femelles et les jeunes males.

REMARQUES. — Takeda et Hanamura (1994) ont tout recemment signale Stylodactylus licinus de

l'lndonesie (Mer de Flores, 280 m, 1 9 ov.).

Dans le materiel qu'ils ont dtudie figure egalement un specimen male, identifie Stylodactylus sp. (1994 : 18,

fig. b-c), capture entre 558 et 593 m. II y a tout lieu de penser, a la lecture de leur description et de leurs

remarques, qu’il s'agit en fait d'un exemplaire de Stylodactylus licinus : le rostre plutot droit et plus fin doit

corresponds au fait qu’il s'agit d’un male, de taille moyenne. Un examen de ce specimen s'av&re toutefois

indispensable pour en confirmer l'identification.

Coloration. — La couleur de fond des specimens varie assez sensiblement du rose au rouge (Fig. 4 A-D).

Distribution. — Stylodactylus licinus est connu des Philippines (550-970 m), de la Nouvelle Caledonie

(580-950 m), des lies Chesterfield (650-970 m), du Vanuatu (541-990 m), des lies Fidji (494 m), des lies Wallis et

Futuna (580-900 m), du Japon (432-826 m), de l'Australie (222-1000 m) et de l'lndonesie (280-810 m). La

distribution bathymetrique de l'espece semble concentree entre 500 et 1000 m, les recoltes a des profondeurs

moindres (Australie, 222 m et Indonesie, 280 m) semblant assez exceptionnelles.

Stylodactylus tokarensis Zarenkov, 1968

Stylodactylus tokarensis Zarenkov, 1968 : 58 (pro parte ), fig. 2 (non fig. 2, dessin du troisifeme p6reiopode. et fig. 3 =

Stylodactylus licinus Chace, 1983). — Cleva. 1990 : 91, fig. 3 k-p, 4-5.

MATERIEL EXAMINE. — Indonesie. Karubar : st. CP 05, 285-323 mild 7.5 mm (POL1PI). — St. CP 17, 459-

439 m : 1 6 8 mm (MNHN-Na 12138). — St. CP 27, 304-314 m:li ires abime et incomplet (POLIPI).

Chez les deux males en bon etat, le rapport des longueurs LR/LC est egal ou a peine superieur a 1. Les

formules rostrales s'ecrivent 30(7)/7 ; 28(7)/6 ; 30(6)/6, nombres qui s'inscrivent dans les limites des variations

observees dans notre travail de 1 990.

DISTRIBUTION. — Nous avions deja signale Sty-lodactylus tokarensis de l’lndonesie (Detroit de Macassar, 595-

592 m, Cleva, 1990 : 91) ; les recoltes de 1991 ont ete realisees entre 285 et 459 m. Cette espece, decrite a partir

d'un specimen male capture en mer de Chine orientale, a 820 m de profondeur, est egalement presente en

Nouvelle-Caledonie (485-850 m) et aux ties Chesterfield (500-570 m).

Stylodactylus multidentatus multidentatus Kubo, 1942

Stylodactylus multidentatus Kubo, 1942 : 34, fig. 4-5. — Hayashi & Miyake, 1968 : 586, fig. 1. — Miyake, 1982 : 26,

pi. 9, fig. 5 (photo couleur). — Chace, 1983 : 11 (cl6), 20, fig. 8 a-o. — Chan & Yu, 1985 : 290, pi. 1 E-F (photos

couleurs). — Hayashi, 1986 : 93, fig. 53 (photo couleur). — Kensley, Tranter & Griffin, 1987 : 293.

Stylodactylus multidentatus multidentatus - Cleva, 1990 : 100, fig. 7, 8 h-m ; 1994 : 59.

392

R. CLEVA

Stylodactylus discissipes - Balss, 1933 : 84 (non Bate, 1888).

Stylodactylus bimaxillaris - Miyake, 1982, pi. 9. fig. 4 (non Bate, 1888).

Stylodactylus brevidactylus - CLEVA, 1990 : 106, fig. 8 a-g.

Materiel EXAMINE. — Indonesie. Karubar : st. DW 24, 243-240 m : cdphalothorax seulement — St CP 27

304-314 m : 3 <3 9,5 £ 16,5 mm (MNHN-Na 12139). — St. CP 66, 211-217 m : 1 2 14,5 mm (POLIPI). — St. CP 61

233-246 m : 9 <3 14,5 k 18 mm ; 4 9 (3 ov.) 16,5 & 19,5 mm (MNHN-Na 12140). — St. CP 79 250-239 m • 8 c3 15 i

l1 ™ „\9,?J8 0V') l0'5 * 18'5 mm (P0L1PI>' - St- cp 83, 285-298 m : 3 <3 14,5 It 18,5 mm (MNHN-Na 12156) -

St. CP 84, 275-246 m : 3 <3 16 h 19 mm ; 1 2 ov. 13,5 mm (USNM). — St. CP 85, 244-239 m : 1 6 16.5 mm (POLIPI)

— St. CP 86. 226-222 m : 3 <3 16,5 it 18,5 mm ; 4 2 (3 ov.) 9 a 20,5 mm (MNHN-Na 12141)

Vanuatu. Musorstom 8 : st. CP 1007, 720-830 m : I 2 13 mm (MNHN-Na 13210). — St DW 1061 458-5P m

1 <3 14 mm (MNHN-Na 1321 1). - St. DW 1065, 360-419 m:13 12mm (MNHN-Na 13212), 1 2 12 mm (MNHN-Na

14472). - St. CP 1087. 394-421 m : 4 2 (1 ov.) 8,5 a 14 mm (MNHN-Na 13213). - St. CP 1088, 425-455 m l 6

*MNHN"Na 13214). — St. CP 1092, 314-321 m : 1 2 8,5 mm (MNHN-Na 13215). — St. CP 1 136 398-400 m •

25 c3 10,5 a 15,5 mm, 9 2 (6 ov.) 10,5 a 15 mm (MNHN-Na 13216). — St. CP 1137, 360-371 m : 29 <3 9 5 a 14 5 mm

(1 sp. avec bopyre), 20 2 (8 ov.) 8 & 14 mm (MNHN-Na 13217).

Nouvelle-Caiedonie. Bathus 1 : st. CP 668, 205-219 m : I <3 8,5 mm, 3 2 5 & 6.5 mm (MNHN-Na 14641) -

St. CP 701, 302-335 m : 1 2 6 mm (MNHN-Na 14639).

Bathus 2 : st. DW 731, 300-370 m : 1 c3 14 mm (MNHN-Na 14637). — St. CP 742, 340-470 m : 1 <3 1 1 mm 12

8 mmm(MN^NHNa^463466)29) ~ ^ °W ?58’ 3?7~386 m : 1 8.5 mm (MNHN-Na 14638). — St. CP 760. 455 m : 1 8

Halipro 1 : st. CP 869, 450-490 m : 1 6 14,5 mm (MNHN-Na 14478).

, . Pa™us,4,: st- CP 897 • 305-350 m : 1 6 1 1 mm (MNHN-Na 14380). — St. CP 900, 580 m : 1 c3 12 mm (MNHN-Na

“TO, H (MNHN-Na 13207). - * CP 905, 294-296 m : 1 9 8,5 mm (MNHN-Na mi) - s" CP 953

220-234 m : 1 <3 11,5 mm (MNHN-Na 13209).

L'observation de ce materiel complementaire confirme. la encore, la variability intraspecifique importante de

cette espece, et nous a conduit a reconsidercr le statut de Stylodactylus brevidactylus , decrit lors de notre etude de

1990 a Part.r d'un specimen unique, aux dactyles des P3 a P5 particulierement courts par rapport a ce que I'on

observe habituellement chez Stylodactylus multidentatus. Dans les collections etudiees, deux specimens

indonesiens de la station CP 27 montrent cn effet des dactyles courts, mais dont les longueurs relatives par rapport

au propode sent mtermedia.res entre celles notees chez S. multidentatus (CLEVA, 1990 : 102) et celles indiquees

pour S. brevidactylus (Cleva, 1990 : 106) ; ce dernier peut alors etre consid6re comme le terme extreme, observe

jusqu a present pour ce caracterc chez 5. multidentatus et, avec cette hypothese. doit etre mis en synonymie avec

ui. Ceci semble dautant plus justifte que les autres caracteres retenus pour definir 5. brevidactylus (taille des

spinules de ecaille antennaire, longueur relative des pereiopodes) sont, eux aussi, sujet a des variations notables

meme s. une tendance moyenne semble se degager (un releve precis de chaque caractere conduirait de toute evidence

a une combe de Gauss). C est amsi, par exemple, que dans la tres grande majorite des cas, les spinules qui ornent le

bord de 1 ecaille antennaire sont peu nombreuses et de petite taille.

s,in!^“ fb!T'eS dr la l0ngUeUr rClatiVe dU r0Stre el de SOn arma,ure- Chez les specimens indonesiens,

smscrivent tout a fail dans celles notees en 1990 pour les exemplaires des Philippines : le rapport LR/LC varie de

7\ , l’2 ‘m°yenne ^ P°ur 39 specimens) ; on compte de 40 a 63 epines rostrales dorsales (moyenne = 48) et

15 a 28 epines ventrales (moyenne = 21). Par ailleurs, le rapport LR/LC des specimens de Nouvelle-Caiedonie

est sens.blement plus grand que chez les specimens indonesiens, comme nous Pavions deja note en ce qui concerne

plus'Se Us"' P PPP'nS (CLEVA’ 1990 : 104) : CheZ 11 SP6dmens au ros,rc en bon etat, il varie entre 1.20 et

varSriTefrrfmov ^ aVOnS/elf d 9ue. sur 70 specimens au rostre complet, le rapport LR/LC

. ; . ’4 <m°ycnne - 1,3). Le nombre d’epines rostrales dorsales varie entre 34 et 52 (moyenne = 44)

P4 vin'( mSrpePinCL Ve,Ura ! Cr;C ‘ 6 Ct 3 1 (m°yenne = 22)' Les rapP°rls des longueurs propode/dacty le des P3 ei

3 1 W T,Ct,VrDtde 1 • ^ 3'2 61 d£ 2'4 4 3’7 (raPPels : ^imens phi.ippins : P3 1 .8 a 2,2 ; P4 • 2 2 l

; ” i ' P4 : : ,mPOr,am' “ conform, la mise en

synonymie ae 5. brevidactylus avec S. multidentatus.

366“T,°“gement r'PandUe d3nS ''Ind°-0^st Pacifique : Japon (225-300 m), Philippines (152-

366 m), Taiwan (150-250 m), Austra.ie (237-412 m), Nouvelle-Caiedonie (205-580 m), Vanuatu (314 830 m),

STYLODACTYLIDAE INDO-OUEST PACIFIQUES

393

Indonesie (146-314 m). La repartition bathymetrique semble concentrde entre 150 et 400-450 m environ. Un

specimen a ete rdcoltd entre 720 et 830 m lors de la station CP 1007 de MUSORSTOM 8, ce qui semble

exceptionnel, pour ne pas dire douteux. Si Ton fait abstraction de ce cas particulier, la profondeur maximale

observee est de 580 m.

Stylodactylus libratus Chace, 1983

Fig. 1 A. C, E ; 2 A, C

Stylodactylus libratus Chace, 1983 : 12, fig. 5. — Kensley, Tranter & Griffin, 1987 : 292 . — Cleva, 1990 : 108,

fig. 9 b, 18 a-b. — Hayashi, 1991 : 40.

MATERIEL EXAMINE. — Indonesie. "Albatross" Expedition 1907-1910 : st. 5645. Celebes, Selal Butung,

5°29'06"S- 1 22°36’06"E, 377 m, 16.12.1909 : 1 6. holotype, 14 mm (USNM 196081).

Karubar : st. CP 16, 315-348 m:ld 11,5 mm (MNHN-Na 12158). — St. CP 17, 459-439 m : 1 6 12 mm (MNHN-

Na 12967).

Nouvelle-Caledonie. Biocal : st. CP 105, 21°30’S-166°2I ’E, 335-330 m, 08.09.1985 : 1 <$ 8,5 mm, I 9 ov.

9 mm (MNHN-Na 10658).

Smib 8 : st. DW 160, 280-282 m : 1 9 ov. 8 mm (MNHN-Na 14797).

lies Loyaute. Musorstom 6 : st. DW 391, 20o47,35'S-167°05,70'E, 390 m, 13.02.1989 : 1 9 7,5 mm (MNHN-Na

11368). — St. CP 401, 20°42, 1 5'S- 1 67°00,35'E, 270 m, 14.02.1989 : I 6 7.5 mm (MNHN-Na 11899).

lies Chesterfield. Musorstom 5 : st.299. 22°47,70'S-159°23,70,E, 360-390 m. 11.10.1986 : 1 6 5,5 mm

(MNHN-Na 10919). — St. DW 301, 22°06,90'S- 159°24.60'E. 487-610 m, 12.10.1986 : 1 8 4,5 mm, 2 9 5,5 et 7 mm

(MNHN-Na 10918).

Les deux specimens recoltds en Indonesie peuvent etre rapportds avec une assez bonne approximation il l’espece

de Chace, decrite d’apres un specimen unique provenant de la meme region, meme si existent entre eux quelques

differences comme la taille relative du rostre et celle de l’ceil par exemple (Fig. I E et 2 A). Les neuf specimens de

Nouvelle-Caledonie et des lies Loyaute et Chesterfield, deja cites dans notre travail de 1990, beaucoup plus petits,

ont egalement ete identifies a S. libratus. Tous ont en commun un rostre au moins aussi long que la carapace

(LR/LC compris entre 1 et 1,3 ), arme ventralement d'au moins 5 epines (entre 5 et 13).

Un second “groupe" de specimens, identifies dans un premier temps a S. libratus avec des reserves, continue de

nous interroger. II est etudie ci-apres, sous la denomination Stylodactylus sp.

Stylodactylus sp.

Fig. 1 B. D, F ; 2 B. D, F

MATERIEL examine. — Indonesie. Karubar : st. CP 17, 459-439 m : 2 9 13 et 16 mm (MNHN-Na 12142).

Nouvelle-Caledonie. Biocal : st. DW 51. 23°05,S-167°44,E. 680-700 m, 31.08.1985 : 1 9 8,5 mm (MNHN-Na

10917) (identifid h S. libratus par Cleva, 1990).

Chalcal 2 : st. DW 72, 24°54,50'S-168°22,30'E, 527 m, 28.10.1986 : 1 9 10.5 mm (MNHN-Na 10920) (identifie ti

S. libratus par Cleva, 1990).

Beryx 11 : st. DW 09. 790-825 m : I 8 12,5 mm (MNHN-Na 12159). — St. CP 53, 540-950 mil 9 11,5 mm

(MNHN-Na 14649). — St. CP 60, 580-600 m : 1 <3 13 mm (MNHN-Na 14647) ; 1 6 10 mm, 1 9 carapace abimee

(MNHN-Na 14648).

Bathus 3 : st. DW 776, 770-830 m : 1 6 9 mm. 2 9 5,5 et 10,5 mm (MNHN-Na 14480). — St. DW 786, 699-

715 m : 1 9 9 mm (MNHN-Na 12164). — St. DW 794, 751-755 m : 1 9 7,5 mm (MNHN-Na 12163).

Outre le fait que ces specimens possedent un rostre court (LR/LC compris entre 0,7 et 0,85), arme

ventralement de 5 epines au plus (2 a 5), d’autres differences apparaissent nettement lorsque l’on compare, deux a

deux, des specimens de ce type avec des specimens identifies a S. libratus de meme taille et, si possible, de meme

sexe (Na 11368 9 S. libratus et Na 12163 9 S. sp., fig. 1A et IB ; Na 10658 6 et Na 10917 9 ; Na 10658 9

et Na 12164 9, fig. 1C et ID ; Na 12158 S et Na 14649 9, fig. IE et IF). Ces differences semblent bien

indiquer, & priori, que Ton est en presence de deux “lignees” : on peut noter en effet que, chez Stylodactylus sp.,

Source : MNHN, Paris

394

R. CLEVA

Fig

r \ZA‘:Sty!?d?CctyluS UbJ.TS- Nouvelle-Cal6donie, $ 7,5 mm (MNHN-Na 11368) ; B, Stylodactylus sp„ Nouvelle-

Cal6donie, 2 7,5 mm (MNHN-Na 12163) ; C, S. libraius, Nouvelle-Caledonie, 2 9 mm (MNHN-Na 10658) •

DKiSlr^o?'VN°Urel,le-Cal6d0nie’ 2 9 mm (MNHN-Na 12164) ; E, S. libra, us, Indonesie, 6 11,5 mm

(MNHN-Na 12158) ; F , Stylodactylus sp„ Nouvelle-Calddonie, 2 11,5 mm (MNHN-Na 14649). Echelles = 1 mm.

Source : MNHN, Paris

STYLODACTYLIDAE INDO-OUEST PACIFIQUES

395

l’oeil est plus volumineux, les epines rostrales plus longues et plus robustes, les epines antennaires et

branchiosteges plus longues. Les spinules qui ornent les ecailles antennaires sont egalement plus longues.

Toutefois, lorsqu’il s’agit de specimens plus grands, les differences sont plus difficiles a apprehender : ainsi le

male 12159 Stylodactylus sp. possede des epines rostrales plus courtes que celles du male 12142 S. libratus

(Fig. 2A et 2B), ce qui contredit ce qui precede. De meme, le volume de la cornee est assez comparable chez les

plus grands specimens (Fig. 2A et 2B, 2C et 2D).

Ces specimens a rostre court ont des affinit^s certaines avec Stylodactylus pubescens Burukovsky, espece

recoltee sur les rides sous-marines de Sala-y-Gomez et de Nasca, au large du Chili, dans l’Est Pacifique, par

545-800 m de profondeur. L’auteur nous a aimablement fait don de deux specimens, un male de LC 9,5 mm et

une femelle ovigere de LC 8,5 mm. Cette espece, tout comme Stylodactylus sp., possede un rostre plus court que

la carapace, arme ventralement de 2 a 4 epines, et des epines antennaires et branchiosteges longues. Compares a

des specimens de Stylodactylus sp. de taille equivalente, ces 2 specimens montrent des epines rostrales

sensiblement plus Fines, un rostre plus recourbe vers le bas, des pereiopodes sensiblement plus longs et plus greles

(Fig. 2E et 2F).

On comprendra dans ces conditions nos hesitations a decider du statut de Stylodactylus sp.

1) Les differences observees avec certains des specimens identifies a S. libratus , les differences de profondeur de

recolte (de 439 a 950 m pour Stylodactylus sp., contre 270 it 610 m pour S. libratus ), les quelques differences

relevdes avec S. pubescens ainsi que l’eloignement des zones de recoltes, font pencher pour l'hypothese d’une

espece nouvelle.

2) Les caracteres intermediaires (et parfois contradictoires) presentes par certains specimens font pencher pour

une seconde hypothese, celle d’une seule espece, 5. libratus , qui presenterait une grande variability intraspecifique,

comme on a deja pu l’observer chez d’autres especes, a laquelle s’ajoute ici une probable variability geographique,

notre matyriel provenant d’lndonesie et de Nouvelle-Caledonie.

La seconde interrogation trouve d'autant plus sa justification que, dans notre travail de 1990, un specimen

recolte a Madagascar par 400 m de fond, identifie S. aff. libratus, n'est peut-etre qu'un exemplaire de l'espece de

CHACE, au rostre sensiblement plus long, et avec des dpines surnumeraires sous l’epine branchiostyge (Cleva,

1990 : 110).

Des informations sur la coloration des specimens, malheureusement absentes actuellement, nous permettraient

sans aucun doute de prendre une decision finale quant au statut reel de ces differents specimens.

Signalons enfin que HAYASHl (1991 : 41) a recemment fait mention d'un specimen male de Stylodactylus

libratus recolty au Japon (Kyushu-Palau Ridge).

Genre NEOSTYLODACTYLUS Hayashi & Miyake, 1968

Neostylodactylus affinis Hayashi & Miyake, 1968

Fig. 4 E

Neostylodactylus affinis Hayashi & Miyake, 1968 : 605, fig. 7. — Chace, 1983 : 4 (cl6). — Cleva. 1990 : 112 (cle),

1 13.

Materiel EXAMINE. — Indonesie. Karubar : st. DW 18, 205-212 m : 1 2 3 mm (POLIPI). — St. DW 32. 170-

206 m : 1 2 3,5 mm environ (carapace abimye) (MNHN-Na 12143). — St. DW 50, 184-185 m : 1 2 ov. 3,5 mm

(POLIPI).

Nouvelle-Caledonie. Bathus 4 : st. DW 934, 231-240 m : 1 2 3,3 mm (MNHN-Na 14466).

Vanuatu. Musorstom 8 : st. CP 1135, 282-375 m : 1 2 3 mm (MNHN-Na 13218).

Les trois specimens indonesiens, incomplets et pas en tres bon etat, ont cependant ete identifies avec une bonne

approximation a N. affinis. Nous avons note que, sur le specimen de la station DW 32 et le specimen neo-

caledonien, dont les quatriemes pereiopodes droits sont encore en place, ainsi que sur celui du Vanuatu dont les

deux P4 sont prysents, l'ischion et le meats sont fusionnes (pas de ligne de suture visible). Une ligne de suture

Source :

396

R. CLEVA

Source : MNHN, Paris

STYLODACTYLIDAE INDO-OUEST PACIFIQUES

397

bien visible s'observe, par contre, au niveau des cinquiemes pereiopodes du specimen de la station DW 50 et

des specimen de Nouvelle-Caledonie et du Vanuatu, ce qui confirme nos observations anterieures (Cleva, 1990 :

112).

COLORATION. — La mention portee sur Petiquette accompagnant le specimen photographic indique

“uniformement verdatre”, ce qui correspond mal a la photo que nous reproduisons (Fig. 4 E). II est vraisemblable

que cette photo est surexposee car l’animal apparait plutot jaune.

Distribution. — Detroit de Coree, 120 m ; Nouvelle Caledonie et lies Chesterfield, 231-440 m ; Indonesie,

premier signalement, 170-212 m ; Vanuatu, premier signalement, 282-375 m.

Genre PARASTYLODACTYLUS Figueira, 1971

Parastylodactylus bimaxillaris (Bate, 1888)

Fig. 4 F

Stylodactylus bimaxillaris Bate, 1888 : 855, pi. 138, fig. 3. — Calman. 1939 : 188. — Hayashi & Miyake, 1968 : 599,

fig. 5. — Miyake, 1982 : 25 (non pi. 9, fig. 4 = Stylodactylus multidentatus Kubo, 1942).

Parastylodactylus bimaxillaris - Chace, 1983 : 8, fig. 4. — Chan & Yu, 1985 : 289, pi. I A-D (photos couleurs). —

Cleva, 1990 : 115, fig. 1 1 a, 12 a ; 1994 : 62.

Non Stylodactylus bimaxillaris - Calman, 1925 : 16. — Barnard, 1950 : 652, fig. 1 22 f-h (= Stylodactylus stebbingi

Hayashi & Miyake, 1968).

Non Stylodactylus bimaxillaris - Miyake, 1982, pi. 9, fig. 4 (= Stylodactylus multidentatus Kubo, 1942).

MATERIEL EXAMINE. — Indonesie. Karubar : st. CP 09, 361-389 m : 9 d 6 <1 8 mm ; 14 2 (8 ov.) 4,5 & 8 mm

(POLIPI) ; 1 3,22 (1 ov.) avec bopyres (MNHN-Na 12152). — St. CC 10, 329-389 m : 1 $ ov. 7 mm (POLIPI). —

St. CP 33, 281-31 1 m : 1 2 ov. 6,5 mm (USNM). — St. CP 36. 268-210 m : 1 6 6 mm, 1 2 ov. 7 mm (MNHN-Na

12144). — St. CC 41, 401-393 m : 1 2 ov. 8,5 mm (MNHN-Na 12145). — St. CP 45, 301-305 m : 2 $ 5.5 mm (USNM).

— St. CP 69, 356-367 m : 1 $ ov. 6,5 mm (MNHN-Na 12146). — St. CP 70, 411-410 m : 1 2 ov. 6.5 mm (POLIPI). —

St. CP. 76, 400 m : 1 2 6 mm (MNHN-Na 12147). — St. CP 83, 285-298 m : 1 2 ov. 6,5 mm (MNHN-Na 12148). —

St. CP 86, 226-222 m : I 2 4,5 mm (MNHN-Na 12149).

Nouvelle-Caledonie. Bathus 4 : st. CP 910, 560-608 m : 1 2 ov. 7 mm (MNHN-Na 13219).

Vanuatu. Musorstom 8 : st. CP 1091, 344-350 m : 1 <S 4 mm (MNHN-Na 13220), 1 2 ov. 5 mm (MNHN-Na

14473).

Cet abondant materiel (40 specimens) apporte quelques donnees nouvelles sur la variability de l'espece : le

rostre, qui mesure de 1,6 a 2 fois la longueur de la carapace, porte de 18 a 35 epines dorsales (dont 6 a 8 post-

rostrales), et de 5 a 9 epines ventrales (moyennes sur les 29 specimens au rostre intact : epines dorsales : 25 ;

epines ventrales : 6). Les chiffres les plus bas releves jusqu'ici semblent etre ceux d'un specimen australien chez

qui Ton compte 17 epines dorsales et 4 ventrales (CLEVA, 1994 : 62).

II semble se confirmer, par ailleurs, que le rostre est plus incurve vers le haut chez les femelles que chez les

males.

Distribution. — Largement repandu dans l'lndo-Pacifique (Afrique du Sud, Mozambique, Madagascar,

golfe d'Aden, Philippines, Taiwan, detroit de Coree et mer de Chine orientale, Japon, Australie, Nouvelle-

Caledonie, Vanuatu), Parastylodactylus bimaxillaris etait deja connu de la region indonesienne, puisque les types

de Bate ont ete recoltes au nord de la Nouvelle-Guinee. La distribution bathymetrique de l'espece s'etend entre 106

et 608 m.

Parastylodactylus richeri Cleva, 1990

Fig. 4 G-H

Parastylodactylus richeri Cleva, 1990 : 127, fig. 15 c, 16 e-h.

Source :

398

R. CLEVA

MATERIEL EXAMINE. — Indonesie. Karubar : st. CP 27, 304-314 m : 1 9 moins de 3,5 mm (POLIPI).

Vanuatu. Musorstom 8 : st. CP 980, 450-433 m : I 9 ov. 5 mm (MNHN-Na 14468). — St. CP 1024, 335-370 m :

1 9 ov. 4,5 mm (MNHN-Na 14470). — St. CP 1025, 385-410 m : 1 9 ov. 6 mm (MNHN-Na 14469). — St. CP 1137,

360-371 m : 1 9 ov. 5,5 mm (MNHN-Na 13222).

Nouvelle-Caledonie. Bathus 1 : st. CP 670, 394-397 m : 1 9 ov. 5 mm avec bopyre (MNHN-Na 14631). —

St. DW 687, 408-440 m : 1 8 3 mm (MNHN-Na 14645).

RFO, 300 m : 2 9 (1 ov.) 3,5 et 4,5 mm (MNHN-Na 13221).

Le specimen indonesien, dont la carapace est abimee et le rostre incomplet, a pu toutefois Stre iden¬

tify a Parastylodactylus richeri, espece decrite de la Nouvelle-Caledonie, et dont 4 autres specimens ont et 6

rapportds de cette region. Quelques differences ont ete relevees par rapport au materiel type chez ces

9 nouveaux exemplaires. Elies sont indiquees ci-apres. Entre parentheses sont rapelles les chiffres notes pour le

materiel type.

Le merus des P3 porte 3 ou 4 epines laterales (4), et son dactyle est orne de 4 a 7 spinules (4 a 6) ; celui des P4

porte 1 ou 2 epines (2 ou 3) et son dactyle 3 a 7 spinules (4 & 6) ; celui des P5 porte 2 a 4 epines (2 a 4) et son

dactyle 4^6 spinules (6). Le rapport des longueurs propode/dactyle est respectivement de : P3 : 1,8 a 3,1 (2,6) ;

P4 : 2,1 a 3,7 (3,0 a 3,2) ; P5 : 4,7 a 7,4 (6,1 a 6,8).

Le rostre porte 21 a 26 epines dorsales (22 a 25 pour le materiel type) et 3 ou 4 epines ventrales (3), et le

rapport des longueurs du rostre a la carapace varie entre 1,2 et 1,55 (1,35).

COLORATION. — Elle n'est pas tres caracteristique. La teinte generale est rosatre, avec quelques taches plus

rouge ou orange. Des anneaux de couleur rougeatre s'observent sur les pereiopodes (Fig. 4 G-H).

Remarque. — Parastylodactylus richeri est tres proche d'une autre espece aux dactyles des P3 et P4

particulierement longs (et inermes), Parastslodactylus longidactylus Cleva, 1990, decrite des Philippines et non

retrouvee depuis.

Distribution. — Nouvelle-Caledonie (300-440 m), Vanuatu (335-450 m), Indonesie, premier signalement,

304-314 m.

Parastylodactylus moluccensis sp. nov.

Fig. 3

Materiel EXAMINE. — Indonesie. Karubar : st. CP 16. 315-348 m : 1 9 6,5 mm (MNHN-Na 12157). —

St. CP 27, 304-314 m : 1 6 7 mm environ (carapace abimee) (MNHN-Na 12122).

Types. — La fcmelle recoltee a la station CP 16 dont la carapace mesure 6,5 mm est l'holotype (MNHN-Na

12157), le male recoltd lors de la station CP 27 est un paratype.

Description (d apres 1 holotype) — Rostre mesurant environ 1,7 fois la longueur de la carapace. Sa moitie

distale est assez fortement redressee vers le haul. Son extremite manque. II porte 16 epines dorsales (dont

5 implantees sur le bord dorsal de la carapace, en arriere du niveau du fond de l'orbite), de taille variable et

d implantation irreguliere, et 6 fortes epines ventrales, de taille plus homogene, sensiblemcnt plus longues et plus

robustes que les epines dorsales les plus fortes. Quelques soies plumeuses s’observent a la base ou entre certaines

epines, et d'autres ont probablement du se detacher.

Carapace avec epines supra-orbitaire, antennaire (emoussee), et branchiostege.

Cinquieme pleuron abdominal arrondi, avec unc forte epine sur sa marge posterieure.

Telson avec 2 rangees de 4 epines dorsales.

(Eil a cornee bien developpec ; une cornee secondaire (ocelle) bien individualistic ; pedoncule oculaire orne de

3 rangees de 2 a 4 soies plumeuses disposees comme suit : deux sont situees en position dorsale (1'une a la limite

de la cornee principale et l'autre legerement en arriere de la premiere, au niveau de 1'ocelle), la troisieme est placee

lateralement, a la limite egalement de la cornee principale.

Source : MNHN, Paris

STYLODACTYLIDAE INDO-OUEST PAC1FIQUES

399

Fig. 3. — Parasiylodactylus moluccensis sp. nov., 9 holotype 6.5 mm.. Indonesie (Karubar, st. CP 16, MNHN-Na

12157) : A, cdphalothorax ; B. abdomen et telson ; C. D. E . troisieme, quatribme et cinquieme pereiopodes droits.

Echelles = 1 mm.

Source : MNHN, Paris

400

R. CLEVA

Stylocerite terming en pointe aigiie, atteignant le 2eme tiers du premier article du pedoncule antennulaire.

Ecaille antennaire au bord exteme inerme, de longueur egale a celle de la carapace.

Longueur des troisiemes maxillipedes el des pereiopodes par rapport a l'extremite de l'ecaille antennaire : des

MxP3 ne subsiste que le premier article ; chez le paratype, les MxP3 depassent cette extremite de la longueur du

dernier article ; PI d’un peu plus de la longueur du propode ; P2 des 2/3 de la longueur du propode ; P3 n'atteint

pas tout a fait l'extremite de l'ecaille antennaire ; P4 atteint le milieu de l'ecaille et P5 depasse le niveau du premier

tiers de l'ecaille.

De longues soies plumeuses ornent les articles de tous les pereiopodes.

Ischion-merus des troisiemes pereiopodes un peu plus de 2 fois plus long que le carpe (lobe distal compris),

avec une epine ventrale pres de sa base, une epine laterale sub-distale, et une petite epine distale dorsale ;

propode environ 2 fois plus long que le carpe et 4,5 fois plus long que le dactyle ; dactyle avec 3 ou 4 spinules

ventrales.

Ischion et merus des quatriemes pereiopodes completement fusionnes (pas de ligne de suture visible), plus de

2 fois plus longs que le carpe ; merus avec une epine laterale sub-distale et une petite epine dorsale distale ;

propode environ deux fois plus long que le carpe, et 4,5 fois plus long que le dactyle ; dactyle avec 4 spinules

ventrales (des 2 cotes).

Ischion et merus des cinquiemes pereiopodes completement fusionnes, environ 2 fois plus longs que le carpe ;

spinulation du merus identique & celle des quatriemes pereiopodes ; propode environ 2 fois plus long que le carpe et

4,5 fois plus long que le dactyle ; dactyle avec 5 spinules ventrales (des 2 cotes).

Le paratype differe de I'holotype par quelques points de detail :

— Telson avec 2 rangees de 4 et 5 epines dorsales.

— Pereiopodes un peu plus courts (par rapport a l’extremite de l'ecaille antennaire).

— Rapport des longueurs ischion-merus/carpe des trois dernieres paires de pereiopodes sensiblement plus

grand.

ETYMOLOGtE. — Le terme moluccensis fait reference au lieu de recolte de cette nouvelle espece.

Discussion. L association de certains des caracteres de ces deux specimens : presence d’une epine supra-

orbitaire, cinquieme pleuron abdominal arrondi mais avec une forte epine, fusion complete dc l'ischion et du merus

des P4 et P5, dactyles des trois dernieres paires de pereiopodes courts et robustes en font indeniablement un taxon

nouveau, le sixieme du genre Parastylodactylus.

II differe :

Pm l°n8‘dactylus Cleva, 1990, et P . richeri Cleva, 1990, par la presence d'une epine supra-orbitaire et

d'une forte epine sur le 5eme pleuron abdominal ; le rostre plus long (LR/LC = 1,7 contre 1,2 a 1,55), aux epines

plus longues et robustes, et avec des epines ventrales plus nombreuses (6 au lieu de 4, 3 ou 2) ; les dactyles des P3

a P5 courts et robustes ; la fusion de l'ischion et du merus des P4 et P5 ; l’ornementation du telson (4 ou 5 epines

par rangee dorsale au lieu de 3).

de P. bimaxillaris (Bate, 1888), par la fusion de l'ischion et du merus des P4 et P5 ; la presence d'une

epine proximo-ventrale sur l'ischion-merus des P3 ; la position de l'epine sur le cinquieme pleuron abdominal ;

I ornementation du pedoncule oculaire (3 rangees de soies au lieu d'une seule).

, ~ de P' tranterae Cleva’ 1990, par le nombre d'epines dorsales du rostre (16 dont 5 post-rostrales, contre 42

a 50 dont 1 1 a 16 post-rostrales) ; la presence d'une epine supra-orbitaire ; la fusion de l'ischion et du merus des P4

et P5 ; 1 ornementation du telson (3 paires d’epines chez P. tranterae).

,n , ~ de Pse™blatae C|eva, 1990, par le nombre d'epines dorsales du rostre (16 dont 5 post-rostrales, contre

30 a 39 dont 8 a 1 2 post-rostrales) ; la fusion de l'ischion et du merus des P4 et P5 ; la presence d'une epine sur le

cinquieme pleuron abdominal.

Coloration. — Inconnue.

Distribution. — Indonesie, 304-348 m

Source : MNHN, Paris

STYL0DACTYL1DAE INDO-OUEST PACIFIQUES

401

AUTRES ESPECES RECOLTEES A WALLIS ET FUTUNA (MUSORSTOM 7)

ET AU VANUATU (MUSORSTOM 8)

Stylodactylus brucei Cleva, 1994

Stylodactylus brucei Cleva. 1994 : 54, fig. 1A, C-F.

MATERIEL EXAMINE. — lie Wallis. MUSORSTOM 7 : st. CP 638, 820-840 m : 1 6 holotype, 37 mm (MNHN-Na

12121).

Cette espece, la plus grande de la famille (longueur totale 220 mm), a ete recemment decrite lors de I'etude

d'une collection de Stylodactylidae d'Australie (Cleva, 1994). L'exemplaire recolte a Wallis, en meilleur dtat que

les specimens australiens, a etc choisi comme holotype.

Distribution. — Australie (900-1000 m), tie Wallis (820-840 m).

Stylodactylus macropus Chace, 1983

Stylodactylus macropus Chace, 1983 : 16, fig. 7. — Cleva, 1990 : 95.

MATERIEL EXAMINE. — Vanuatu. MUSORSTOM 8 : st. CP 992. 775-748 m : 1 2 15 mm (MNHN-Na 13223).

Distribution. — Philippines (700-925 m), Nouvelle-Caledonie (800-825 m), lies Chesterfield (745-825 m),

Vanuatu, premier signalement (775-748 m).

Parastylodactylus semblatae Cleva, 1990

Parastylodactylus semblatae Cleva, 1990 : 122, fig. 12 c, 14, 18 c; 1994 : 62.

Materiel EXAMINE. — iles Wallis et Futuna. MUSORSTOM 7 : sans numero de station, 500-610 m : I 2

5.5 mm (MNHN-Na 14464).

Vanuatu. Musorstom 8 : st. CP 974, 492-520 m : 1 $ ov. 7 mm (MNHN-Na 14471). — St. CP 1027, 550-571 m :

1 $ 6 mm (MNHN-Na 14467).

Distribution. — Australie (458-500 m), Nouvelle-Caledonie et iles Chesterfield (260-702 m), Vanuatu (492-

571 m), Wallis et Futuna (500-610 m).

AUTRES ESPECES RECOLTEES EN NOUVELLE-CALEDONIE

Stylodactylus laurentae Cleva, 1990

Stylodactylus laurentae Cleva, 1990 : 96, fig. 6, 19.

MATERIEL EXAMINE. — Nouvelle-Caledonie. Beryx 1 1 : st. DW 27. 460-470 m : 2 $ 4,5 et 6 mm (MNHN-Na

14646).

Smib 8 : st. DW 146, 514-522 m : 1 2 ov. 8.5 mm (MNHN-Na 12133), 1 2 8 mm (MNHN-Na 14633). —

St. DW 185, 305-355 m : 1 6 6 mm (MNHN-Na 14643). — St. DW 193, 500-508 m : 1 8 8,5 mm, 1 2 ov. 7,5 mm

(MNHN-Na 14618). — St. DW 194, 491 m : 1 8 7,5 mm (MNHN-Na 14634). — St. DW 195. 508-514 m : 2 2 5,5 et

7 mm (MNHN-Na 14627). — St. DW 201, 500-504 m : 1 2 ov. 7,5 mm (MNHN-Na 14644).

Bathus 2 : st. CP 737, 350-400 m : 1 2 ov. 7.5 mm (MNHN-Na 14632).

Halipro 1 : st. CP 877, 464-480 m : 1 2 4,5 mm (MNHN-Na 13224).

Bathus 4 : st. DW 923, 502-470 m : 1 2 8.5 mm (MNHN-Na 13225). — St. DW 924, 344-360 m : 2 2 6 mm

(MNHN-Na 13226). — St. DW 931, 360-377 m : 1 2 ov. 7 mm (MNHN-Na 13227).

Source :

402

R. CLEVA

Distribution. — Nouvelle-Caledonie et ties Chesterfield (300-610 m).

Neostylodactylus amarynthis (de Man, 1902)

Stylodactylus sp.(amarynthis) de Man, 1902 : 897, pi. 27, fig. 64 a-b.

Stylodactylus Amarynthis - DE Man, 1920 : 32, pi. 5, fig. 9, 9 a-h.

Stylodactylus amarynthis - Kemp, 1925 : 258.

Neostylodactylus amarynthis - Hayashi & Miyake, 1968 : 603, fig. 6. — Chace, 1983 : 4, fie 1-3 — Cleva 1990 ■

112 ; 1994 : 60, fig. 3.

Materiel EXAMINE. — Nouvelle-Caledonie. Montrouzier : chenal de Touho, 52 m : 1 5 ov. 3 mm (MNHN-

Na 12160).

DISTRIBUTION. — La Reunion, ties Andaman, Indonesie, Philippines, Japon, Australie, Nouvelle-Caledonie

(premier signalement), cntre 9 et 120 m (Cleva, 1990 : 1 13).

Parastylodactylus tranterae Cleva, 1990

Parastylodactylus tranterae Cleva, 1990 : 1 19. fig. 1 1 b, 12 b, 13.

Nouvelle-Caledonie. Beryx II : st. CP 08, 540-570 m : 1 9 ov. 8.5 mm (MNHN-Na

Materiel examine.

14626).

Smib 8 : st. DW 150, 519-530 m : 1 2 8,5 mm (MNHN-Na 14635).

' Lst’ CP 657- 490-530 m : 1 d 5 mm environ. 1 2 7 mm avec bopyre (MNHN-Na 14622). — St CP 671

450-470 m : 1 2 ov. 9 mm (MNHN-Na 14620),

Bathus 2 : st. CP 738, 558-647 m : 3 6 4,5 h 7,5 mm (MNHN-Na 14628).

Bathus 3 : st. CP 832, 650-669 m : 1 2 9,5 mm (MNHN-Na 14474). — St. CP 833. 441-444 m : 2 2 7 5 et 8 5 mm

avec bopyre (MNHN-Na 12161). — St CP 846, 500-514 m : 1 2 10,5 mm avec bopyre (MNHN-Na 14477)

Na Sf 4 1 St' CP 892‘ 580‘6°° m : ' 9 8 mm (MNHN-Na 14382). — St. CP 912, 702-690 m : 1 2 ov.9 mm (MNHN-

DISTRIBUTION. — Australie (540 m), ties Chesterfield (650-700 m), Nouvelle-Caledonie (441-702 m).

Parastylodactylus semblatae C leva, 1990

Parastylodactylus semblatae Cleva, 1990 : 122, fig. 12 c, 14. 18 c; 1994 : 62.

s^nlv 2mLsEnnAsniN£' -.N°uvelle-Caledonie. Sm.b 8 : st. DW 191, 564-580 m : 1 2 6 mm (MNHN-Na 14465).

St. DW 201, 500-504 m ; 1 d 5 mm, 1 2 6 mm (MNHN-Na 14642).

Bathus 2 ; st. DW 721, 525-547 m : 1 <5 6,5 mm (MNHN-Na 14640).

Bathus 4 : st. CP 910 560-608 m : 1 d 6,5 mm, 2 2 6 et 6,5 mm (MNHN-Na 13228). — St. CP 911 566-558 m ■

613 6l'o T'r 9 sJT ^mu^m32239)' _ S‘ CP 9'2’ 702-690 m : 1 9 6 mm (MNHN-Na 13230). - St. CP 92fi

St DW 9'>o so? sir,6 "T 'i4^^3 l3231>- — St. CP 922, 600 m : 1 d 6,5 mm. 1 2 6 mm (MNHN-Na 13232). —

1373^ VI' s°s2'5 6 T 5 °uV'.6'5 mm (MNHN-Na 13233)- — St. CP 930, 530-520 m : 3 d 5 a 5,5 mm (MNHN-Na

13234), 1 2 5,5 mm photographiee (MNHN-Na 13235).

II est mteressant de noter que, chez quelques specimens, on peut observer une petite epine sur le bord posterieur

MncnTc P eT" abd°minal d’un seul c6t6 (c’est le cas pour 3 femelles de Bathus 4 el pour celle de

Musorstom 7) ou meme des deux cotes (male de 5 mm recolte lors de la station CP 930 de Bathus 4) ■ ce

caractere qui peut done s'observer quelquefois, est a prendre en consideration lors de l'utilisation de la cle des

especes du genre Parastylodactylus. Cette remarque est en mettre en parallele avec celle faite tt propos de

Stylodactylus lianas , ou certains spectmens n’ont pas le pleuron abdominal 4 termine en pointe aigiie (cf. supra).

S7|D‘S,TR,IBU™D,; _ ^ustralie (458-500 m), Nouvelle-Caledonie et ties Chesterfield (260-702 m), Vanuatu (492-

571 m), ties Wallis et Futuna (500-610 m).

Source : MNHN, Paris

STYL0DACTYL1DAE INDO-OUEST PAC1FIQUES

403

Stylodactyloides crosnieri Clcva, 1990

Stylodactyloides crosnieri Cleva, 1990 : 129, fig. 17, 18 d-e ; 1994 : 62.

MATERIEL EXAMINE. — Nouvelle-Caledonie. Smib 8 : st. DW 180, 460-525 m : I ? 13.5 mm (MNHN-Na

12132). — St. DW 183, 330-367 m : 1 6 15,5mm (MNHN-Na 14616). — St. DW 187, 390-540 m : 1 $ 14,5 mm

(MNHN-Na 14630).

Bathus 3 : st. DW 830, 361-365 m : 2 6 15,5 et 16,5 mm, 1 9 1 1,5 mm (MNHN-Na 14482). — St. DW 836, 295-

306 m : 1 $ ov. 15,5 mm (MNHN-Na 12162).

Distribution. — Nouvelle-Caledonie et ties Chesterfield (200-540 m), Australie (357 in), Nouvelle-Zelandc

(Yaldwyn, 1991, in litt.).

CONCLUSIONS

Le grand nombre de specimens de Stylodactylidae recoltes lors de ces campagnes, permettent d'apporter une

interessante contribution a la connaissance de cette famille, tout en laissant apparattre la complexity de la definition

de certains taxons. Ainsi :

— Une espece nouvelle, Parastylodactylus moluccensis, de I'lndonesie, est decrite.

— La distribution geographique de plusieurs autres est etendue ou leur presence, supposee, confirmee :

Stylodactylus licinus a Wallis et Futuna et au Vanuatu, S. multidentatus au Vanuatu, Stylodactylus macropus au

Vanuatu, Neostylodactylus affinis en Indonesie et au Vanuatu, N. amarynthis en Nouvelle-Caledonie,

Parastylodactylus bimaxillaris au Vanuatu, P. richeri en Indonesie et au Vanuatu, P. sembiatae a Wallis et Futuna

et au Vanuatu.

— La variability intraspecifique de plusieurs especes (Stylodactylus licinus, S. multidentatus, Parastylodactylus

bimaxillaris, notamment) est dysormais mieux connue, grace aux importantes series de specimens recoltees.

Cette variability apparait parfois tres importante : il suffit de voir dans quelles proportions varient, pour une

meme espece, la longueur relative du rostre ou le nombre de ses epines (pour ne citer que cet exemple facile a

observer) pour s'en convaincre. Elle nous a ainsi conduit a mettre en synonymie Stylodactylus brevidactylus

Cleva, 1990, decrit a partir d'un seul specimen aux dactyles notablement courts, avec S. multidentatus. Elle nous

conduira peut-etre, aussi, a mettre en synonymie avec 5. libratus, dont le materiel type est reduit a un specimen

unique, ou avec S. pubescens, les exemplaires identifies dans un premier temps a 5. libratus avec des reserves, et

separes provisoirement sous l'appellation Stylodactylus sp.

Dans le meme ordre d'idees, des series suffisamment importantes de specimens de differentes regions s'aveirent

indispensables pour mieux evaluer une probable variability geographique souvent seulement pressentie, et qui nous

conduira peut-etre h reconsiderer le statut de certains specimens.

— La coloration de deux especes, Neostylodactylus affinis et Parastylodactylus richeri, inconnue jusqu'a

present, est indiquee. II est vraisemblable que, compte tenu de ce qui a ete dit plus haut, beaucoup d'incertitudes et

de questions restant en suspens pourraient etre rysolues si Ton disposait d'informations sur la couleur des animaux

recoltes, particuliyrement dans les cas ou apparait un dessin caracteristique.

REMERCIEMENTS

B. Richer de forges et A. Crosnier, tous deux de 1’ORSTOM, ont mis h notre disposition le materiel

etudie ici ; J.-P. ROCROI (BIMM, Museum national d'Histoire naturelle), nous a traduit le texte russe de la

description de Stylodactylus pubescens Burukovsky et K. Baba ceux des recentes publications de K.-I. Hayashi ;

F. A. CHACE et A. CROSNIER ont effectue une lecture critique de notre manuscrit ; M. DE SAINT LAURENT nous a

conseille a plusieurs reprises ; J. Rebi£re a repasse a l'encre nos dessins et J.-L. Menou est l'auteur des

photographies. Nous sommes heureux de pouvoir les remercier tous ici.

Source :

404

R. CLEVA

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Zarenkov, N.A., 1968. — New data on rare shrimps (Thalassocaridae, Rhynchocinetidae, Stylodactylidae,

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Source :

406

R. CLEVA

FIGURE 4

A-D Stylodactylus licinus, Vanuatu, Musorstom 8 : A. $ ov. 13 mm (MNHN-Na 14458) ; B, 2 13 mm (Na 14460) ■

C cS 16,5 mm (MNHN-Na 14461) ; D. S 15 mm (MNHN-Na 14459) ; E , Neostylodactylus affinis Nouvelle’

' ¥\Parasylodac,ylus bimaxiUaris , Vanuatu. Musorstom 8, 2 ov. 5 mm

H, 2^ov. 6^ l,MNW^a\<M<&)0<*aCty US MUS0RST0M 8 : G’ ? ov' 4’5 «™> (MNHN-Na 14470) ;

Source : MNHN, Paris

STYLODACTYLIDAE INDO-OUEST PACIFIQUES

407

Source : MNHN . Paris

Source : MNHN, Paris

SUL ATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DE

Crustacea Decapoda: Palinuridae, Scyllaridae

and Nephropidae collected in Indonesia

by the KARUBAR Cruise, with an identification key

for the species of Metanephrops

Tin -Yam CHAN

Institute of Marine Biology

National Taiwan Ocean University

Keelung, Taiwan, R.O.C.

ABSTRACT

The lobsters collected by the Karubar cruise from Indonesia are examined. Twenty-one species are identified, ten of

which being newly recorded in Indonesia. Moreover, the Karubar cruise obtained an intact specimen of Metanephrops

arafurensis (de Man, 1905) and a re-description is given for this poorly known species. The four species groups in

Metanephrops are redefined and a revised key to the species of Metanephrops is provided.

RESUME

Crustacea Decapoda : Palinuridae, Scyllaridae et Nephropsidae recoltes en Indonesie lors de la

campagne Karubar. Cle d'identification des especes du genre Metanephrops.

Les especes appartenant aux families des Palinuridae, Scyllaridae et Nephropidae, recolt6es lors de la campagne

Karubar en Indondsie, ont ete etudides: 21 especes ont 6te trouvees, dont 10 n'avaient pas encore ete signalees en

Indonesie. Un specimen de Metanephrops arafurensis (de Man, 1905) ayant etd recolte, cette espece, mal connue, est

reddcrite. Les quatre groupes d'espfeces du genre Metanephrops sont redefinis et une cle d'identification de toutes les

especes de ce genre est propostSe.

INTRODUCTION

The Karubar cruise in 1991 collected a number of deep-sea lobster specimens from Indonesia. The material is

found to contain five species of palinurids, five species of scyllarids and 1 1 species ot nephropids. Although no

new species were found, the Karubar material extends the known distributions for many species, such as

Linuparus trigonus (von Siebold, 1824), Palinustus unicornutus Berry, 1979, Ibacus pubescens Holthuis, 1960,

I. novemdentatus Gibbes, 1850, Nephropsis acanthura Macpherson, 1990, N. holthuisi Macpherson, 1993,

Chan, T.-Y., 1997, — Crustacea Decapoda: Palinuridae, Scyllaridae and Nephropidae collected in Indonesia by the

Karubar Cruise, with an identification key for the species of Metanephrops. In: A. Crosnier & P. Bouchet (eds),

Rdsultats des Campagnes MUSORSTOM, Volume 16. Mem. Mus. natn. Hist. nat.. 172: 409-431. Paris ISBN: 2-85653-

506-2.

Source : MNHN Paris

410

T.-Y. CHAN

N. serrata Macpherson, 1993. N. stewarti Wood-Mason, 1872, N. sulcata Macpherson, 1990, and Metanephrops

australiensis (Bruce, 1966). The most interesting finding is a complete specimen of Metanephrops arafurensis (de

Man, 1905), which was previously known only from a mutilated type. Together with the additional knowledge

gained of the characteristics of the other Metanephrops species, their relationships are discussed and a revised key

to the species of this genus is provided.

MATERIAL AND METHODS

The specimens used in the present study are deposited at the Museum national d'Histoire naturelle, Paris

(MNHN), Puslitbang Oseanologi-LIPI, Indonesia (POLiPI), National Taiwan Ocean University (NTOU), National

Museum of Natural History, Washington, D.C. (USNM), Nationaal Natuurhistorisch Museum. Leiden (RMNH)

and Zoologisch Museum, University of Amsterdam, Amsterdam (ZMA). Furthermore, the collections of the

MNHN and NTOU contain 15 of the 18 known species of Metanephrops. Only three species, namely M.

binghami from the Caribbean, M. motunauensis and M. challenger i from New Zealand have not been examined.

Their characteristics are mainly by referring to Yaldwyn (1954), HOLTHUIS (1964, 1991), JENKINS (1972) and

Takeda (1990).

The terminology used for the body parts of Nephropsis mainly follows MACPHERSON (1990), while that for

Metanephrops follows CHAN & Yu (1991). The measurements are of carapace length (cl) which is measured along

the dorsal midline from the postorbital margin to the posterior margin of the carapace. Only restricted synonymies

of Indonesian records and important works on the species are given.

In the lists of material, CP = beam trawl, CC = shrimp otter trawl, DW = Warren dredge.

SYSTEMATIC ACCOUNT

Family PALINURIDAE

Genus L1NUPARUS White, 1847

Linuparus trigonus (\ on Siebold, 1824)

Palinurus Trigonus von Siebold, 1824: 15 (type-locality: Japan).

Linuparus trigonus - HOLTHUIS, 1991: 114, figs 215-216.

-?i i ^i7TER!Ao examinED-— Indonesia. Karubar. Tanimbar Islands : stn CP 66, 09°01'S, 132°09’E, 1,11 1991

21 1-217 m: 1 2 91.2 mm (MNHN). — Stn CP 79, 09°16'S, 131°22'E, 250-239 m, 3 1 1 1991- 1 2 72 3 mm (POI IPO '

Stn CP 83, 09°23'S, 131°00'E, 285-297 m, 4.11.1991: I 6 69.1 mm (POLIPI) (POLIPI). -

Remarks. — Although this species is often reported in the western Pacific (Japan, Korea, China, Taiwan, the

Philippines and Australia, at depths of 30-3 1 8 m), it is recorded here for the first time in Indonesia.

Genus PALINUSTUS A. Milne Edwards, 1880

Palinustus unicornutus Berry, 1979

— A,rica)- ■ ««= •* <*

24 ?7“Z<™TN)nd0nC!i!i' K™B*R lCai s“ Dw 1*. 05-18-S. 133‘01’E, 205.312 m.

Source : MNHN, Paris

PALINURIDAE, SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

411

Remarks. — P. unicornutus was previously thought to be restricted to the eastern coast of South Africa.

Recently, however, it was found that this species actually has a wide distribution in the Indo-West-Pacific (Chan

& YU, 1995).

Palinustus waguensis Kubo, 1963

Palinustus waguensis Kubo, 1963: 63, figs 1-3 (type-locality: Japan). — Holthuis, 1991: 126, figs 237-238. — Chan &

Yu, 1995: 389, fig. 7, 8D, 9E, 10E.

MATERIAL EXAMINED. — Indonesia. Karubar. Kai Islands: stn DW 30, 05°39'S, 132°56'E, 118-111 m,

26.10.1991: 1 <3 31.4 mm (MNHN).

Remarks. — Similar to P. unicornutus, recent finds show that this species is also widely distributed in the

Indo-West-Pacific (Chan & Yu, 1995). Nevertheless, its occurrence in Indonesia had already been suggested by

Holthuis (1991).

Genus PUERULUS Ortmann, 1897

Puerulus angulatus (Bate, 1888)

Panulirus angulatus Bate, 1888: 81, pi. 11 -figs 1-4 (type-locality: New Guinea).

Puerulus angulatus - Holthuis, 1991: 162, figs 301-302.

Not Puerulus angulatus - DE Man, 1916: 36, pi. 2-fig. 5 (= Puerulus velutinus Holthuis, 1963).

MATERIAL EXAMINED. — Indonesia. Karubar. Kai Islands: stn DW 2, 05°47'S, 132°13'E, 209-240 m.

22.10.1991: 1 juv. 14.3 mm (POL1P1). — Stn DW 18, 05°18'S, 133°01'E, 205-212 m, 24.10.1991: 1 6 50.3 mm

(POLIPI). — Stn CP 36, 06°05'S, 132°44'E, 268-210 m, 27.10.1991: 4 juv. 14.8-17.9 mm (MNHN).

Tanimbar Islands: stn CP 85, 09°22'S. 131°14'E, 245-240 m, 4.11.1991: 1 juv. 13.4 mm (MNHN). — Stn CP 88.

08°45'S, 130°47'E, 1188-1178 m, 5.11.1991: 1 juv. 13.0 mm (MNHN).

REMARKS. — This species is widely distributed in the Indo-West-Pacific (from 274 to 536 m deep) and has

been reported from Indonesia by HOLTHUIS (1991). The specimen from Stn CP 88 has the body spines and

pleopods exceptionally long and was collected from great depth (1 188-1178 m). Since the arrangement of the

spines on body in this specimen is very similar to those of the other juveniles of this species, it is probably still

in the very early puerulus stage.

Puerulus velutinus Holthuis, 1963

Puerulus angulatus - DeMan, 1916: 36, pi. 2-fig. 5 ( non Bate, 1888).

Puerulus velutinus Holthuis, 1963: 55 (type-locality: Indonesia); 1966: 273; 1991: 165, figs 307-308.

MATERIAL EXAMINED. — Indonesia. Karubar Kai Islands: stn CP 35, 06°08'S, 132°45'E, 390-502 m, 27.10.

1991 : 1 ovig. $ 58.0 mm (MNHN).

REMARKS. — P. velutinus is unique in the genus, in bearing a large postorbital spine. It is known from the

southern Philippines down to NW Australia (Wadley & Evans, 1991), at depths of 485-683 m.

Family SCYLLARIDAE

Genus IBACUS Leach, 1815

Ibacus brevipes Bate, 1888

Ibacus brevipes Bate, 1888: 62, pi. 9-fig. 1 (type-locality: Indonesia). — HOLTHUIS, 1985: 47, figs 13-14; 1991: 201

figs 384-385.

Source

412

T.-Y. CHAN

Material EXAMINED. — Indonesia. Karubar. Kai Islands: stn CP 36, 06°05'S, 132°44'E 268-210 m 27 10

1991: 6 6 20.6-38.5 mm, 2 9 21.7-40.3 mm (POLIP1).

Remarks. — The Karubar specimens were taken from almost the same locality and depth as the type

collected by the "Challenger" (i.e. 05°49.15’E, 132°14.14'E, 256 m) near the Kai Islands. In the Karubar

material the number of posterolateral teeth on the carapace may be as high as 18 (12-17 in HOLTHUIS, 1985). As

mentioned by HOLTHUIS (1985), the teeth on the anterior margin of the distal antennal segment are sometimes

strongly reduced in the males. This species has been reported from the South China Sea, the Philippines,

Indonesia and New Caledonia (at depths of 186-457 m).

Ibacus pubescens Holthuis, 1960, comb. nov.

Ibacus ciliatus pubescens Holthuis, 1960: 147 (type-locality: the Philippines); 1985: 33, fig. 8; 1991- 203 fig 338

right.

Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn CP 36, 06°5'S, 132°44’E 268-210 m 27 10

1991: 1 2 24.4 mm (POLIP1).

Tanimbar Islands: stn CP 82, 09°32'S, 131°02'E 219-215 m. 4.11.1991: I 2 35.3 mm

09°22'S, 1 3 1 0 1 4'E, 245-240 m, 4.11.1991: 1 2 21.8 mm (POL1PI),

(MNHN). — Stn CP 85,

Remarks. — This present form differs from the typical I. ciliatus (von Siebold, 1824) by having the entire

body distinctly pubescent and slightly more posterolateral teeth on the carapace (one Karubar specimen even has

15 posterolateral teeth on one side). The typical form has a northern distribution, from Japan to the South China

Sea and northern Philippines. The pubescent form has been found in the southern range of the distribution of the

species (i.e. southern Philippines and NW Australia), but is only reported for the first time from Indonesia here

Similar to the figure provided by HOLTHUIS (1985, 1991), the posterior margin of abdominal tergite V is evenly

serrated in the Karubar material. However, all the specimens from Taiwan have the posterior margin of abdomi¬

nal tergite V provided with only three or four distinct tubercles near the lateral ends (Chan & Yu, 1993: 187-

upper photo). The figures of/, ciliatus ciliatus given by HOLTHUIS (1985, 1991) also show a similar arrangement

ol tubercles on abdominal tergite V. Thus, it seems justifiable to treat the pubescent, southern form as a distinct

species, rather than subspecies, as in the comparable situation of Metanephrops velutinus and M. andamanicus It

is interesting that material from the Philippines to Australia often has the body more pubescent.

The depth range of this species is from 151-391 m (HOLTHUIS, 1985).

Ibacus novemdentatus Gibbes, 1850

'baC(igs 39cf 39^f”,a,M* Gibbes, 1850: 19 (type-locality: unknown). — Holthuis, 1985: 52, figs 15-17; 1991: 204.

199^VfroLmmX(POLIPII)'“Ind0neSia' KaRUBAR ls,a"ds: s,n 35- 06°08'S. 132°45'E, 390-502 m, 27.10.

Tanimbar Islands: stn CP 81, 09°35'S, 131°02'E, 200-207 m, 4.11.1991: I <J 52.0 mm (POL1P1).

.mRE“AfS' — Although I. novemdentatus is widely distributed in the Indo-West-Pacific (at depths of 37-

400 m), it has nevertheless not been reported from Indonesia before. This species generally bears 8 posterolateral

teeth on the carapace, but the male from the Karubar cruise has 7, while the female has 9 posterolateral teeth

Genus SCYLLARUS Fabricius, 1775

Scyllarus sp.

??mfomRl.AL E*A.MINED- — Indonesia. KARUBAR. Kai Islands: stn CP 5, 05°49'S 132°18'E 269-299 m

SST JUV- mm <MNHN)- ~ S,n °W 32* 05°47'S- l32°5''E- >70-106 m. 26 io H991 fjuv 6 6 mm

Source :

PALINURIDAE, SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

413

REMARKS. — These two specimens are very young juveniles (probably at the very early postlarval stages). No

trace of gonopores is found and their pleopods are extremely long. The thoracic sternum has the anterior process

undeveloped (i.e. anterior end of the thoracic sternum truncate and not anteriorly protruded at all) and bears a pair of

elongate posterolateral spines. The juvenile of S. cultrifer collected by the Karubar cruise (see below) has a

similar size but the pleopods are rudimentary and the anterior extremity of the thoracic sternum already shows the

characteristic shape of the species. Thus, it is highly likely that these two small specimens are the juveniles of a

large Scyllarus species. They show some similarities to S. crenatus (WHITELEGGE, 1900), described from East

Australia, in possessing a distinct rostrum, the arborescent markings on the abdomen being indistinct (almost

absent in one specimen) and only the posterior margins of abdominal somites I to III medially incised. S. crenatus

is still known only from the types and it is also likely that WHlTELEGGE's (1900) specimens (about cl 5.2 mm)

are juveniles of other species. This situation is further complicated because the taxonomic status of many species

of Scyllarus is still unclear.

Scyllarus cultrifer (Ortmann, 1897)

Arctus cultrifer Ortmann. 1897: 272 (type-locality: Japan).

Arctus sordidus - Bate, 1888: 66, pi. 9-fig. 3 ( non Stimpson, 1860).

Scyllarus cultrifer meridionalis Holthuis, 1960: 150 (type-locality: the Philippines).

MATERIAL EXAMINED. — Indonesia. Karubar. Kai Islands: stn DW 24, 05°32'S. 132°51'E, 243-230 m,

26.10.1991: 1 9 8.0 mm (MNHN).

REMARKS. — S. cultrifer is unique in the genus by having both the pereiopods III and IV subchelate. This

species is widely distributed in the Indo- West- Pacific (from littoral to about 290 m deep). HOLTHUIS (1960)

assigned the southern material (i.e. from the Philippines, Indonesia and east of South Africa) to a distinct

subspecies, S. cultrifer meridionalis Holthuis, 1960. However, the KARUBAR specimen shows intermediate

characters between the typical and southern forms. Its posterior margin of abdominal tergite IV lacks a median

incision but the other characteristics all conform to the typical form. As suggested by Harada (1962) and Chan

and Yu (1993), it may not be necessary to divide this species into two subspecies. S. cultrifer has already been

reported by Bate (1888, under the name "Aractus sordidus ") from the Kai Islands.

Family NEPHROPIDAE

Genus NEPHROPSIS Wood-Mason, 1873

Nephropsis acanthura Macpherson, 1990

Nephropsis acanthura Macpherson, 1990: 311, figs 5d, 9d-f, lla-b, 16d (type-locality: the Philippines). — Holthuis,

1991: 35, fig. 61-62. — Griffin & Stoddart, 1995: 234.

MATERIAL EXAMINED. — Indonesia. Karubar. Tanimbar Islands: stn CP 87, 08°47'S, 130°49’E, 1017-1024 m,

5.11.1991: 1 6 11.1 mm (POLIPI). — Stn CP 89, 08°39'S, 131°08’E, 1084-1058 m, 5.11.1991: 1 ovig. 9 18.9 mm

(MNHN).

Remarks. — Although this species is widely distributed in the Indo-West-Pacific (at depths of 720-1305 m),

it is here recorded for the first time from Indonesia. N. acanthura is distinct in having an erect basal spine on the

telson and can be distinguished from the closely related species N. occidentalis Faxon. 1893, from the eastern

Pacific by the rostrum being distinctly longer than one-half the carapace length. The size of N. acanthura is also

smaller than its eastern Pacific counterpart. However, the carapace is not granulate in the Karubar specimens and

that of the female is rather heavily pubescent. Furthermore, the carapace of the male has some post-supraorbital

spinules which are lacking in the female (see also GRIFFIN & Stoddart, 1995).

Source :

414

T.-Y. CHAN

Nephropsis ensirostris Alcock, 1901

Nephropsis ensirostris Alcock. 1901: 162, pi. 1-fig. 2 (type-locality: Arabian Sea). — deMan, 1916: 113

— Macpherson, 1990: 303, figs 5a, 6. 8a-b, 16a. — Holthuis, 1991: 41. figs 71-72.

Material EXAMINED. — Indonesia. Karubar. Tanimbar Islands: stn CP 70, 08o4]'S, 131°47'E, 413-410 m

2.11.1991: 1 6 19.5 mm (MNHN). — Stn CP 72, 08°36’S, 13 1°33'E. 699-676 m, 2.1 1.1991: 1 2 16.8 mm (POLIPI).

Remarks. — The present species is unique in the genus in lacking teeth on the rostrum. N. ensirostris is

widely distributed in the Indo-West-Pacific (at depths of 315-1300 m) and its occurrence in Indonesia has been

recorded by de Man (1916) and Macpherson (1990). The two Karubar males agree well with the description of

Macpherson (1990), except that the outer spine on the terminal half of the carpus of large chelipcd is absent in

one specimen and poorly developed on one side in the other. The smaller male also has the left anterior spine of

the subdorsal carina missing.

Nephropsis holthuisi Macpherson, 1993

Nephropsis holthuisi Macpherson, 1993: 55, figs 1-3 (type-locality: NW Australia). — Griffin & Stoddart, 1995: 234.

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn CP 38. 07°40’S, 132°27'E 620-666 m

28.10.1991: 1 2 9.0 mm (MNHN).

Tanimbar Islands: stn CC 57. 08°19'S, 131°53'E. 603-620 m. 31.10.1991: 1 6 32.1 mm, I ovig 2 28 1 mm

(MNHN),

Remarks. — This species is extremely similar to N. rosea Bate, 1888, from the western Atlantic.

Macpherson (1993) only used the relative position of the gastric tubercle on the carapace to separate them. Other

than being smaller in size, the Indo-West-Pacific material appears to have the anterior margin of the abdominal

pleuron II slightly more convex and the large chelae less granulate [three specimens of N. rosea from French

Guyana (MNHN-AS 574) were compared]. Since, at present, only six specimens are known from

north-western and eastern Australia and Indonesia (at depths of 603-1105 m), more material is necessary to

determine the taxonomic status of this Indo-West-Pacific form.

Nephropsis serrata Macpherson, 1993

Nephropsis Steward - DE Man, 1916: 112, pi, 3-fig. 17. (non Wood-Mason. 1872).

Nephropsis serrata Macpherson, 1993: 59, figs 4-6 (type-locality: NW Australia).

? Nephropsis hamadai Watabe & Ikeda, 1994: 102 (type-locality: Japan).

,^fJrfRIAL EXAMINED- — ,ndonesia. Karubar. Kai Islands: stn CC 21. 05°14'S. 133°00'E, 688-694

n ,oo,': ,3AJi'4'26'fiI^m' 2 2 22-5-32-3 mm (MNHN). — Stn CP 38, 07°40’S, 132°27'E,' 620-666

28.10.1991: 1 <J 26.3 mm, 12 2 13.0-30.1 mm, 1 juv. 9.4 mm (POLIPI)

Tanimbar Islands: stn CP 59, 08°20'S. 132°H'E, 405-399 m, 31.10.1991: 1 2 23.1 mm (MNHN).

m,

m.

Remarks. The present species is very similar to N. steward and they often occur together. Nevertheless,

M serrata differs constantly from the latter species in having some additional spines on the subdorsal carina Thus

the Stboga" specimen reported by DE Man (1916) from the Kai Islands has the subdorsal carina denticulate and

should belong to the present species instead of N. steward. The size of N. serrata is also much smaller than that of

N. steward. However, the differences in the rostral length and large chela between these two species, mentioned by

Macpherson, (1993), appear to be rather variable in N. steward.

N. hamadai Watabe & Ikeda, 1994, recently described from Japan, is very similar to N. serrata. The characters,

such as the shape of the coxae of the pereiopods and the relative distances of the orbital margin, cervical groove

and posterior margin of the carapace, used by Watabe and Ikeda to distinguish N. hamadai from N. serrata are

found to be very vanable in the Karubar and North Western Australian specimens (10 specimens in NTOU,

CSIRO in exchange). It is highly likely that N. hamadai belongs to the same species as N. serrata.

Source : MNHN , Paris

PALINURIDAE, SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

415

N. serrata was previously known from western Australia only. The present report extends its range northward

to Indonesia, at depths of 300-694 m.

Nephropsis stewarti Wood-Mason, 1873

Nephropsis stewarti Wood-Mason, 1873: 60 (type-locality: Andaman Sea).

Nephropsis stewarti - Macpherson, 1990: 312, figs 5e, 10, llc-d, 16. — Holthuis, 1991: 45, figs 80-81.

Not Nephropsis Stewarti- DEMan, 1916: 112, pi. 3-fig. 17 (= N. serrata Macpherson. 1993).

MATERIAL EXAMINED. — Indonesia. Karubar. Kai Islands: stn CP 12. 05°23'S, 132°37’E. 436-413 m,

23 10 1991: 2 3 13.8-24.6 mm, 1 $ 25.3 mm (MNHN). — Stn CP 35, 06°08'S. 132°45'E, 390-502 m, 27.10.1991:

1 S 53 9 mm (MNHN). — Stn CP 39, 07°47'S. 132°26'E. 477-466 m, 28.10.1991: 1 9 51.8 mm (MNHN).

Tanimbar Islands: stn CC 56, 08°16’S, 131°59'E, 552-549 m, 31.10.1991: 1 spec. 58.0 mm (POL1PI). — Stn CP 59,

08°20'S, 132°1 l'E, 405-399 m, 31.10.1991: 1 3 64.1 mm (MNHN). — Stn CP 69, 08°42'S, I31°53'E, 356-368 m,

2.11.1991: 1 3 32.5 mm (MNHN).

REMARKS. — This species is widely distributed in the Indo-West-Pacihc (at depths ot 170 to over 1060 m).

However, the previous record in Indonesia by DE MAN (1916) actually represented N. serrata. Thus, it can be

considered that N. stewarti is correctly reported from Indonesia only now. N. stewarti is probably the largest

species of the genus, the body length (excluding the rostrum) of a male collected by the Karubar cruise reaching

18.2 cm.

Nephropsis sulcata Macpherson, 1990

Nephropsis sulcata Macpherson, 1990: 319, figs 13e-g, 14a-b. 15a-b, 16g (type-locality: the Philippines). — Holthuis,

1991: 47, figs 84-85.

MATERIAL EXAMINED. — Indonesia. Karubar. Tanimbar Islands: stn CP 81, 09°35’S. 131°02'E, 200-207 m,

4 11 199L 2 ovig. 9 both 21.0 mm. 4 9 15.7-21.5 mm (MNHN). - Stn CP 87. 08°47'S, 130°49'E, 1017-1024 m,

5.11.1991: I <J 22.1 mm. 2 9 12.2-22.5 mm (POLIPI). — Stn CP 89. 08°39'S, 131°08'E. 1084-1058 m. 5.11.1991:

Id 15.6 mm (rostrum missing, tentatively identified as the present species) (MNHN).

Remarks. — N. sulcata is widely distributed in the Indo- West-Pacific but has not been recorded in Indonesia

before. Moreover, one lot of the Karubar specimens (Stn CP 81) was collected at a depth of only 200-207 m

which is much shallower than previously thought for N. sulcata (415-1 1 1 5 m deep in MACPHERSON, 1990, 1993).

It is found that some characteristics used by MACPHERSON (1990) to separate N. sulcata from the closely

related Atlantic species N. atlantica Norman, 1882, are not very satisfactory [specimens of N. sulcata and

N. atlantica in MNHN, mentioned by MACPHERSON (1990), have been compared]. The dorsal carina on the

abdomen is also very distinct and the median groove of the rostrum may sometimes overreach the distal rostral

teeth in the material from the Atlantic. On the other hand, the distance between the post-supraorbital spine and the

gastric tubercle is often more than 0.5 (to about 0.6) times the distance between the gastric tubercle and the post-

cervical groove in the Indo-West-Pacific material. Nevertheless, the size of Indo- West-Pacific specimens appears to

be much smaller than that of the Atlantic material and the carpus of pereiopod II is always shorter than the palm.

Furthermore, the posterior border of the abdominal somite V often bears a distinct spine in the Indo-West-Pacific

specimens, but this spine is usually absent in the material from the Atlantic.

Genus METANEPHROPS Jenkins, 1972

Metanephrops arafurensis (de Man, 1905)

Figs 1, 2 a-c, 3, 4 b, 5 b

Nephrops arafurensis de Man. 1905: 587; 1916: 107, pi. 3-fig. 16 (type-locality: Indonesia). — Yaldwyn, 1954: 730.

Metanephrops arafurensis - JENKINS. 1972: 171. — CHAN & Yu, 1987: 184. - HOLTHUIS, 1991: 67, figs 130-131.

Source :

416

T.-Y. CHAN

Fig. 1 . — Metanephrops arafurensis (de Man. 1905).

<S 50.7 mm (MNHN): a, dorsal view; b, lateral view;

Indonesia, Tanimbar Islands, 7°46'S,

c, dorsal view of carapace.

132°31'E, 443-468 m.

Source : MNHN . Paris

PALINURIDAE. SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

417

MATERIAL EXAMINED. — Indonesia. "Siboga": stn 262, 05°53.8'S, I32°48.8'E, 560 m, 12.18.1899: 1 8

34.0 mm, type (ZMA).

Karubar. Tanimbar Islands: stn CC 40, 07°46'S, 132°31'E, 443-468 m, 28.10.1991: I 8 50.7 mm (MNHN).

Description. — Carapace spinulose. 4-5 post-rostral teeth present. Region between post-rostral carinae only

having some spinules on posterior part. Posterior margin of hepatic groove spinulose, but that of cervical groove

lacking spines except those at anterior ends of post-cervical ridges. Three lateral, post-cervical ridges present.

Antennal spine wing-like and with outer margin somewhat crenulated. Distal segment of maxilliped III rather long

and slender. Large cheliped weakly ridged but covered with sharp tubercles, with inner margins bearing some large

spines, and outer margin of chela angular.

Abdomen with distinct dorsal carina (that of somite I with only anterior end present); raised parts naked but

depressed parts heavily pubescent; tergite I with deep, but short, lateral transverse furrows; tergites II and III with

broad but medially interrupted transverse furrows (as wide as or wider than main facade and having intermediate,

narrow, eroded, transverse carina between) and dorsally arched, lateral, longitudinal furrows, submedian notches

rudimentary but some pits may be present on raised parts near dorsal carina; sculpture on tergite IV similar, but

with main facade broader and having more (and larger) pits, while ventral end of lateral furrow recurved

anterodorsally; non-articular surface of tergite V with raised parts less than depressed parts and mainly composed of

an eroded cross at middle; tergite VI with median ridge bearing a pair of submedian spines, as well as strong spines

at both anterior and posterior ends, posterolateral spine well-developed, lateral lobe terminated posteriorly by a

strong spine and outer margin having 2-3 spines. Abdominal pleura ventrally pointed and with margins crenulated.

Telson bearing 3 spinules at middle, with pair of elongate, submedian, basal spines and 2-3 large spines on

dorsolateral ridges. Endopod of uropods covered with spinules and having a strong basal spine, exopod only with

outer part spinulose. Posterolateral spines of telson and uropods all very well-developed.

Coloration. — Body generally orange pink. Eyes dark brown. Anterior carapace somewhat pink. Large

cheliped slightly banded with pale and deep orange bands. Distal 2/3 ol fingers of large chelae and hinges of

abdominal somites whitish.

SIZE. — The two males known are cl 34 mm and 50 mm.

Distribution. — Only known near the Kai Islands in Indonesia, at depths of 443-560 m.

REMARKS. — M. arafurensis was originally described from a multilatcd specimen and no other material had

been obtained since the original description. The finding of a complete specimen (although with left large cheliped

smaller and probably regenerated) of this species by the Karubar cruise is therefore of particular importance.

The Karubar male is almost identical with the type (condition good except for original damage), except that

the right post-rostral carina bears only four teeth (actually the fifth post-rostral tooth is rather small and situated

more medially than the anterior teeth). Aside from lacking pits on the raised parts near the dorsal carina, the

sculpture of the anterior three abdominal somites of the Karubar male is very similar to that of the type.

Although no illustration was given by DE MAN (1916) of the posterior abdomen, a broken abdominal somite IV

and the left third of somite V are present in the type. They are also very similar to those of the Karubar male and

only differ in the arrangement of pits on the raised parts.

Many authors (eg. BRUCE, 1966; JENKINS, 1972; Chan & Yu, 1987; HOLTHUIS, 1991) suspected that

M. arafurensis is most similar to M. auslraliensis. Indeed the spinulation of the carapace is almost identical

between these two species, with only the spines being relatively more well-developed and the post-rostral carina

sometimes bearing one more, small tooth in M. arafurensis. However, the large cheliped and abdominal sculpture

of M. arafurensis are very different to those of M. australiensis. The presence of spinules on the exopods of the

uropods and the middle of the telson also separate the former species from the latter.

The spinulose large cheliped and complicated abdominal sculpture of M. arafurensis show close resemblance

with the fossil species M. motunauensis Jenkins, 1972, from the late Pliocene of New Zealand. The large

chelipeds appear to be almost identical in these two species. The spinulation ol the carapace and ol the tail fan are

also very similar, except that the entire posterior margin of the cervical groove is spinulose in A7. motunauensis.

Source :

418

T.-Y. CHAN

Although M. motunauensis seems to have fewer spinules on the posterodorsal carapace and tail fan, these spinules

may well be eroded and become indistinct in fossils. The clearest difference between M. arafurensis and

M. motunauensis is of the abdominal sculpture. Although the basic patterns of sculpture are similar [although

JENKINS (1972) mentioned a difference in the shape of the lateral longitudinal furrows actually similarly arched

dorsally in both species], the transverse furrows of M. motunauensis are narrow, as in most of the other species of

the genus. However, the transverse furrows are very broad in M. arafurensis with the abdominal tergites IV and V

having many pits and that of the latter even largely depressed. Furthermore, the median ridge of abdominal tergite

VI bears three pairs of submedian spines in M. motunauensis, but the two anterior pairs of spines are represented

by a single, large spine in M. arafurensis.

M. arafurensis seems to be very rare as only one specimen was obtained by the intensive samplings of the

Karubar cruise and the specimen was collected from almost the same location and depth as the type.

Metanephrops australiensis (Bruce, 1966)

Fig. 2 d-e, 3, 4 b

Neplirops australiensis Bruce, 1966b: 245; pis 25-27 (type-locality: NW Australia).

Metanephrops australiensis - Jenkins, 1972: 171. — Chan & Yu, 1987: 184. — Holthuis, 1991: 68, figs 134-135,

Material EXAMINED. — Australia. "Umitaka Maru": N.E. of Port Hedland, 17°05’S, 119°48'E, 434 m,

26.11.1964: 1 ovig, $ 51.8 mm paratype (RMNH-D21 151).

N.W. Shelf. 1 8° 19S, 117°49'E, 414 m, 25.2.1985: 1 8 50.6 mm (NTOU, CSIRO in exchange). — I6°31'S,

120°16'E, 440 m, 28.8.1986: 4 juv. 19.4-21.5 mm (NTOU, CSIRO in exchange). — 16°32'S, 120°25'E, 440 m!

30.8.1986: 1 8 57.2 mm (NTOU, CSIRO in exchange).

Indonesia. Karubar. Kai Islands: stn CP 35. 06°08’S, I32°45'E, 390-502 m, 27.10.1991: 1 juv. 11.1 mm

(MNHN).

Philippines. "Albatross": stn D5290, I3°40'09"N, 120°55'30"E, 391 m. 22.2.1908: 2 2 31.8-35.3 mm (USNM

170461).

Remarks. — It is interesting that the present species was supposed to be endemic to NW Australia (at depths

of 418-500 m), but is now found in Indonesia. Although the specimen collected by the Karubar cruise is a small

juvenile, it has the typical abdominal sculpture of the species, except for lacking distinct pits. However, its

posteriormost post-rostral teeth are situated more medially and the posterior region between the post-rostral carinae

bears more spinules. Moreover, the exopods of the uropods, as well as the middle and lateral parts of the telson,

bear some spinules. These spinules are always absent in larger specimens (i.e. from cl 19.1 mm onwards). The

fingers are also slightly longer than the palm (1.1 times) in the juvenile. Nevertheless, all these different characters

in juveniles may merely represent the ancestral relationships between M. australiensis and M. neptunus.

The northward distribution of M. australiensis actually reaches as far as the Philippines. The Metanephrops

collection in the USNM has a lot of two females obtained by the "Albatross" from the Philippines (USNM

170461) very similar to the present species. The only difference between the two Philippine specimens and the

material of Australia (NTOU and RMNH) is that the lateral abdomen possess rudimentary longitudinal furrows.

These longitudinal furrows on abdominal tergites II and III are quite distinct (though very shallow) in the smaller

female (fig. 4e). Furthermore, the intermediate, eroded carinae within the transverse furrows of abdominal tergite II

are very distinct in this female and the large chelae are slightly more granulate in the Philippine specimens.

Nevertheless, since specimens from Australia sometimes also have rudimentary longitudinal furrows laterally on

the abdomen (fig. 4d) and since these furrows are intermediately developed in the other female from the

Philippines, it does not seems necessary to treat the Philippine material as a different species.

M. australiensis nonetheless mainly occurs in NW Australia. The extensive sampling in the Philippine-

Indonesian region by many large expeditions has so far only obtained three specimens of this species (i.e. one

juvenile by the Karubar cruise and two females by the "Albatross").

Source :

PALINUR1DAE. SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

419

Fig. 2 a-c. — Metanephrops arafurensis (de Man, 1905), Indonesia. Tanimbar Islands, 7°46'S, 132°31'E. 443-468 m,

6 50.7 mm (MNHN): a, lateral view of abdomen; b, dorsal view of abdomen; c, dorsal view of posterior abdominal

somites and tail fan.

FIG. 2 d-e. - Metanephrops australiensis (Bruce, 1966); d, NW Australia, 18°19'S, 1 17°49'E. 414 m, 6 50.6 mm NTOU

CSIRO in exchange); dorsal view of abdominal tergites II and III. — e, The Philippines, 13°40.09N, 120 55.3 E, 392

m, 2 31.8 mm (USNM 170461): dorsal view of abdominal tergites I-IV.

Source : MNHN. Paris

420

T.-Y. CHAN

Metanephrops neptunus (Bruce, 1965)

Fig. 3, 4 b, 5 c-d

Nephrops neptunus Bruce, 1965: 274, pis 13-15 (type-locality: South China Sea).

Metanephrops neptunus - Jenkins, 1972: 171. — Chan & Yu, 1987: 184. — Macpherson, 1990: 299 — Holthuis

1991: 76, figs 148-149.

Material EXAMINED. — South China Sea. "Cape St. Man’": stn 26, 19°25 5'-19°22'N 1 14°07 5’-l 14°l 1'F

732-795 m, 7.1.1964: 1 «J 59.0 mm allotype (RMNH-D21 152). ‘ H4 11 h.

Tungsha Tao (or Pratas), coll. Taiwan Fisheries Research Institute, Keelung: 1 spec

Philippines. "Albatross": stn 5423, 09°38.3'N, 121°H'E, 929 m, 31.3.1909: 1 9 42.0 mm (USNM 170451)

Indonesia. Karubar. Kai Islands: stn CP 20. 05°15'S, 132°59'E, 769-809 m, 25.10.1991' 1 6 87 1 mm (MNHN)

Tammbar Islands, stn CP 38, 07°40'S, 132°27'E, 620-666 m, 28.10.1991: 1 juv 15 3 mm (MNHN)

CORINDON 2: stn 214. Makassar Strait, 00°31.4'N, 117°50.1'E, 595 m. 1.11.1980: I spec (MNHN-AS 257)

CR.pn ' tralia„N W: Shelf’ l8°19'S' ,17°49'E' 414 ™ 2 6 26.0-69.0 mm. 1 ovig. 9 71.6 mm. I 9 36.5 mm (NTOU

CSIRO in exchange). '

Remarks. M. neptunus has been reported in Indonesia by Macpherson (1990), from Makassar. As

mentioned by BRUCE (1965), the anterior transverse furrows of the abdominal tergites II and III are rather variable

in this species. These furrows can be as long as the posterior transverse furrows (i.e. in the two large specimens of

the Karubar and the Makassar specimen) or have the submedian parts variably raised and fused with the main

facades [,.e. in the types, the specimen from Tungsha Tao, the specimen from the Philippines, the small specimen

trom Karubar and some specimens from Australia (Wadley & Evens, 1991 and specimens in NTOU)]. The red

colour of the body also varies a lot in different individuals. Some have the body nearly all red (i.e. the female from

Karubar, which has only the abdominal somites I to IV whitish, fig. 5 c) while some others are mainly white

i.e. the Australian specimen in Wadley & EVANS, 1991). Half red and half white individuals can also be found

(eg. the types, specimens from Tungsha Tao, Makassar, and the Karubar male, fig. 5 d). Since no correlation is

found between colour patterns, shape of the anterior transverse furrows on the abdomen, localities, depth and sexes

they probably represent natural variations of the species.

Apart from its huge size and presence in deeper waters, the general appearance of M. neptunus rather differs

rom all the other species of the genus [eg. smaller eyes, only two lateral post-cervical ridges, large chelae with

fingers considerably longer (1.3- 1.5 times) than palm, each of abdominal tergites II to V bearing Two transverse

furrows etc.] The small juvenile obtained by the Karubar cruise already has all these "aberrant" characteristics

hus, although juveniles of M. austrahensts show some resemblances with M. neptunus. they can be readily

distinguished from each other. y

Although M neptunus ris widely distributed from the South China Sea to western Australia (at depths of 300-

m and usually more than 500 m) and has a very large size (body length reaching 25 cm in Holthuis, 1991)

SZttSSZSZZ perhaps be d“e “ ,Us '"habi“* — - — «• m*

Metanephrops sibogae (de Man, 1916)

Fig. 3, 4 b

Nephrops sibogae de Man, 1916: 102. pi. 4-fig. 18

Metanephrops sibogae - Jenkins, 1972: 171. _

Griffin & Stoddart, 1995: 232

(type-locality: Indonesia).

Chan & Yu, 1987: 184.

— Yaldwyn, 1954: 730.

— Holthuis, 1991: 79, figs 154-155. —

Source :

PALINURIDAE, SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

421

REMARKS. — M. sibogae is very similar to M. boschmai (Holthuis, 1964) but is generally considered to differ

in the inner margin of the merus of the large cheliped lacking large spines (HOLTHUIS, 1964, 1991; Chan & Yu,

1987). However, 1-3 large spines are found on the merus of the large cheliped in 23 of the 32 specimens collected

by the Karubar cruise. Therefore, the use of the spinulation on the large chelipeds to separate these two species

is not at all satisfactory. Nevertheless, a direct comparison of some M. boschmai specimens from western

Australia (NTOU and 6 paratypes at RMNH) shows that several constant differences can be found between the two

species (see also Griffin & Stoddart, 1995).

The posterior margin of the hepatic groove always bears 4-6 spinules in M. boschmai, while these spinules are

lacking in all the M. sibogae specimens examined. Furthermore, there are 1-5 additional spinules present around

the dorsal postorbital spines in M. boschmai and these spinules are usually absent in M. sibogae (only two

specimens have an additional spinule on one side of the carapace). The paired spines on the dorsal post-cervical

ridge are well-separated in M. boschmai, but abutting in M. sibogae. The median ridge of abdominal tergite VI is

generally armed with a pair of submedian spines in M. boschmai, but only a single median spine in M. sibogae.

Very rarely (2/32) an additional pair of posterior submedian spinules may be present in M. sibogae (as shown in

DE MAN, 1916: pi. 6-fig. 18) though these are always smaller than the median spine. On the other hand, the

median ridge of abdominal tergite VI may occasionally lack distinct spines in both species. As mentioned by

Holthuis (1946) the distal segment of maxilliped III is more oval in M. boschmai (1.8-2.04 times as long as

broad in M. boschmai and 2. 6-3.2 times as long as broad in M. sibogae). The shape of the scaphocerite, however,

is not very different between the two species as described by HOLTHUIS (1964). The large chela is weakly ridged

and granular (sometimes rather sharp) in adults of M. sibogae, but always rounded and very finely granular in

M. boschmai. Other than inhabiting different regions (i.e. M. sibogae occurs from southern Indonesia to northern

Australia at depths of 246-320 m, while M. boschmai is restricted to western Australia at similar depths), the two

species have very different coloration (see Wadley & Evans, 1991). M. sibogae is rather uniformly orange-pink.

The colour of M. boschmai is much attractive: Its body is also orange-pink but the posterior margins of each

abdominal tergite and tailfan are whitish, while the large chela has the fixed finger red, but the movable finger

whitish. Furthermore, the postorbital margin is deep red in some small specimens of M. boschmai.

The abdomen is generally smooth in the present species. However, sometimes rudimentary, transverse, as well

as longitudinal, grooves, or rows of shallow pits, may be present on the abdomen (also present in M. boschmai,

but usually even less distinct).

Metanephrops velutinus Chan & Yu, 1991

Fig. 3, 4 b, 5 a

Metanephrops velutinus Chan & Yu, 1991: 35, pis 2b. 4b. 6c, 8a, c-d (type-locality: the Philippines). — HOLTHUIS,

1991: 82. figs 160-161. — Griffin & Stoddart, 1995: 233.

Material EXAMINED. — Indonesia. Karubar. Kai Islands: stn CC 10, 05°21'S. 132°30’E, 329-389 m,

23.10.199: 2 <3 35.7-48.6 mm, 1 ovig. 2 51.9 mm, 1 2 37.2 mm (MNHN). — Stn CP 12, 05°23'S, 132°37'E, 436-413

m. 23.10.1991. 1 <3 44.8 mm (MNHN). — Stn CP 34. 06°09'S, 132°41'E, 435-445 m, 27.10.199: 4 <3 26.7-49.1 mm, 1

ovig. 2 47.9 mm, 2 2 32.5-56.4 mm (MNHN).

Tanimbar Islands: stn CC 40, 07°46’S, 132°31'E. 443-468 m, 28.10.1991: 2 <3 43.7-56.4 mm (POLIPI). —

Stn CC 41 , 07°45'S, 132°42'E, 401-393 m, 28.10.1991: 5 6 18.1-55.3 mm. 1 ovig. 2 47.5 mm. 1 2 55.1 mm

(MNHN). — Stn CC 42, 07°53'S, 132°42'E. 354-350 m, 28.10.1991: 3 <3 36.0-40.8 mm. 2 ovig. 2 49.1-50.8 mm,

2 2 39.4-55.3 mm (MNHN). — Stn 59. 08°20'S. 132°H'E. 405-399 m, 31.10.1991: 1 <3 41.9 mm (POLIPI). —

Stn CP 69, 08°42'S, 131°53'E, 356-368 m, 2.11.1991: 3 <3 35.5-46.5 mm, 2 2 38.6-39.9 mm (POLIPI).

Remarks. — The present species was only recently described by Chan and Yu (1991) from the Philippines

and western Australia, where it was previously mistaken for M. andamanicus (Wood-Mason, 1891). The

34 specimens collected by the Karubar cruise in southeastern Indonesia all have the raised parts of the abdomen

coarse and pubescent. The record of M. andamanicus by DE MAN (1916) from the other side of Indonesia was

suspected to represent the present species (Chan & Yu, 1991). However, a re-examination of DE Man's (1916)

specimen {"Siboga": stn 12, Java Sea, 07°15'S, 115°15.6'E, 289 m, 14.3.1899: 1 S 44.9 mm (ZMA)] revealed

Source

422

T.-Y. CHAN

that the raised parts of its abdomen are naked and smooth. The Metanephrops collection of the USNM also

contains two M. andamanicus specimens from the Java Sea [Cruise 684, Java Sea, 08°34’S, 1 14°36'E, 322 m,

22.6.1981: 1 S 34.0 mm, 1 2 50.0 mm (USNM 456448)]. Thus, it is possible that M. andamanicus has a

distribution from the Indian Ocean to the western part of Indonesia, while M. velutinus occurs from the

Philippines down to the eastern part of Indonesia and western Australia (at depths of 238-702 m). Material from

intermediate localities, such as the Flores and Banda Seas, will probably provide more insights to the exact

distribution of these two, closely related species. This species was found recently in the Torres Strait.

NOTES ON THE RELATIONSHIPS OF THE SPECIES OF METANEPHROPS

The discovery that the poorly known species M. arafurensis is very similar to the single fossil species of the

genus, M. motunauensis , along with improved knowledge of the species of Metanephrops in recent years (CHAN

& Yu, 1987, 1991, MACPHERSON, 1990), allow a brief revision of the relationships between the species of this

genus.

The number of species (including the fossil species) of Metanephrops has been increased from 14 to 18 since

its erection in 1972. Generally, members of this genus are divided into four groups (DE Man, 1916; Yaldwyn,

1954; JENKINS, 1972; Chan & Yu. 1987; Holthuis, 1991). The "japonicus" group has the carapace smooth,

abdomen conspicuously sculptured and the large chela bearing prominent, spinulated ridges. The "binghami" group

has the carapace finely granulate, abdomen smooth but the large chela with prominent spinulate ridges. The

" arafurensis " group has the carapace spinulose, abdomen with conspicuous furrows and the large chela variably

ridged. The "thornsoni" group has the carapace smooth, abdomen weakly sculptured or smooth, and the large chela

also weakly ridged to smooth.

The division of the genus into these four groups appears to be valid for the 18 species known at present:

The "japonicus" group. — M. japonicus (Tapparone-Canefri, 1873), M. andamanicus (Wood-Mason.

1891), M. sagamiensis (Parisi, 1917), M. formosanus Chan & Yu, 1987, M. mozambicus Macpherson, 1990,

M. armatus Chan & Yu, 1991, and M. velutinus Chan & Yu, 1991.

The "binghami" group. — M. rubellus (Moreira, 1903) and M. binghami (Boone, 1927).

The "arafurensis" group. — M. arafurensis (de Man, 1905), M. neptunus (Bruce, 1965), M. australiensis

(Bruce, 1966) and M. motunauensis Jenkins, 1972.

The "thornsoni" group. — M. thornsoni (Bate, 1888), M. challengeri (Balss, 1914), M. sibogae (de Man,

1916), M. boschmai (Holthuis, 1964) and M. sinensis (Bruce, 1966).

However, the spines and ridges on the large chela are rather small and weak in some species of the "japonicus"

group, such as M. velutinus and M. formosanus. The abdomens of M. mozambicus and M. formosanus lack a

distinct dorsal carina and have the sculpture rather simple (Chan & Yu. 1987, 1991). On the other hand, the large

chela in some species of the "thornsoni" group is somewhat granulate-ridged (e.g. M. sibogae) or has large spines

(e.g. M. thornsoni). Moreover, distinct transverse furrows are present on the abdomen of M. sinensis and

M. thornsoni. Within the ” arafurensis " group, the abdominal sculpture can be very complicated (e.g. M. neptunus

and M. arafurensis) or rather simple (e.g. M. australiensis) while the large chela may be heavily spinulose (e.g.

M. neptunus) or nearly smooth (e.g. M. australiensis).

The spinulation of the carapace, however, appears to be quite constant in separating the species groups of

Metanephrops. The "arafurensis" group has the carapace uniformly spinulose. The "binghami" group has the

carapace rather smooth but the entire posterior margins of both the hepatic and cervical grooves are spinulose. The

carapaces ol the "japonicus" and" thornsoni" groups are smooth, with the posterior margins of the cervical grooves

not serrated. Moreover, the ridging of the large chela appears to have two, somewhat different, forms regardless of

the degree of development of spines on it. In the "japonicus" and "binghami" groups, both the dorsolateral and

ventrolateral margins of the large chela are strongly ridged, making the outer border of the chela a flat surface (that

of the japonicus" group also has a longitudinal, medial depression). On the other hand, the outer border of the

arge che a in the "arafurensis" and "thornsoni" groups is always angular. Although there are large differences in

the complexity of the abdominal sculpture amongst the species of the ' japonicus " and "arafurensis" groups the

Source :

PALINURIDAE, SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

423

furrows on the abdomen are always distinctive in these two groups. Moreover, distinct longitudinal furrows are

present in the "japonicus" group, while the "arafurensis" group usually has some large pits on the abdomen (not

clear in the fossils of M. motunauensis). The abdomens of some species of the "thomsoni" group are sculptured,

but only narrow transverse furrows are present, without large pits on tergites. It should also be noted that the

uropods of the "arafurensis" group are unique in being spinulose. Thus, the definitions of the four species groups

are modified as follows:

"Arafurensis" group. — Carapace uniformly spinulose. Large chela moderately to weakly ridged and from

spinulose to finely granulate, outer border always angular. Abdomen conspicuously sculptured, with or without

deep longitudinal furrows but always bearing some large pits at least in adults (not clear in the fossil species

M. motunauensis). Dorsal surface of uropods spinulose.

"Japonicus" group. — Carapace generally smooth, only bearing some spines on anterior and dorsal parts,

posterior margin of cervical groove not serrated. Large chela ridged and sharply tuberculate to spinulose, outer

border somewhat flat, with a longitudinal medial depression. Abdomen conspicuously sculptured and with deep

longitudinal furrows. Uropods unarmed dorsally.

"Binghami" group. — Carapace generally smooth, but entire posterior margins of hepatic and cervical

grooves spinulose. Large chela ridged and spinulose, with outer border flat. Abdomen not sculptured and uropods

unarmed dorsally.

"Thomsoni" group. — Carapace generally smooth except for some spines on anterior and dorsal parts.

Large chela smooth to weakly ridged, bearing a few large spines; outer border always angular. Abdomen smooth or

bearing only narrow, transverse furrows. Uropods unarmed dorsally.

Since the distinguishing characters previously used for some species are also found to be unsatisfactory (e.g.

M. sibogae vs. M. boschmai and M. thomsoni vs. M. sinensis, etc.), a revised key to the species of

Metanephrops is provided:

1. Carapace rather uniformly spinulose; dorsal surface of uropods covered with spinules .

. 2 ( "arafitrensis " group)

— Carapace smooth between ridges and large spines; uropods unarmed dorsally . 5

2. Region between post-rostral carinae heavily spinulose; abdominal tergites II to V each

with two transverse furrows; large chelae with fingers distinctly longer than palm .

. M. neptunus

— Region between post-rostral carinae only bearing some spinules on posterior part;

abdominal tergites II to V each with one transverse furrow; large chelae with fingers

shorter than palm . 3

3. Abdomen lacking deep longitudinal furrows; large chelae finely granular or nearly smooth.

. M. australiensis

— Abdomen with deep longitudinal furrows; large chelae sharply tuberculate . 4

4. Posterior margin of cervical groove not serrated; transverse furrows on abdominal tergites

more or less as wide as main facades . M. arafurensis

— Posterior margin of cervical groove entirely spinulose; transverse furrows on abdominal

tergites much narrower than main facades . M. motunauensis (fossil species)

5. Posterior margin of cervical groove entirely spinulose . 6 ("binghami" group)

— Posterior margin of cervical groove not serrated . 7

6. Spinules present between post-rostral carinae; abdominal tergites III to V bearing distinct

lateral spines; dorsolateral post-cervical ridge nearly smooth . M. rubellus

— Spinules absent between post-rostral carinae; abdominal tergites III to V without lateral

spines; dorsolateral post-cervical ridge spinulose . M. binghami

Source :

424

T.-Y. CHAN

7. Abdomen bearing distinct transverse and longitudinal furrows; large chelae distinctly

ridged, with outer borders flat . 8 ("japonicus" group)

— Abdomen smooth or having only narrow transverse furrows; large chelae smooth or

weakly ridged, with outer border angular . 14 (" thomsoni " group)

8. Abdominal tergite V bearing distinct lateral spines; median ridge of tergite VI dorsally

armed with paired spines . 9

— Abdominal tergite V without distinct lateral spines; median ridge of tergite VI unarmed

dorsally . 1 q

9. Raised parts of abdomen subdivided; abdominal tergite I bearing well-developed dorsal

car'na . . japonicus

— Raised parts of abdomen smooth; abdominal tergite I lacking distinct dorsal carina .

. . armatus

10. Large spines present on large chela; abdomen without dorsal carina . M. formosanUs

— Large spines absent on large chela; abdomen bearing dorsal carina . 1 l

1 1 . Post-rostral carinae usually with at least one side having 4-5 teeth; spine on lateral lobe

of abdominal tergite VI long and nearly reaching posterolateral groove .... M. sagamiensis

— Post-rostral carinae never bearing more than 3 teeth; spine on lateral lobe of abdominal

tergite VI short and with tip far from posterolateral groove . 12

1 2. Raised parts of abdomen coarse and pubescent . M. velutinus

— Raised parts of abdomen smooth and naked . 13

13. Abdomen with dorsal carina well-developed; main facades of abdominal tergites IV and V

well-separated from dorsal carina, those of tergites II and III bearing posterior submedian

notches . M andamanicus

— Abdomen with dorsal carina almost leveled; main facades of abdominal tergites IV and V

more or less fused with dorsal carina, those of tergites II and III with posterior submedian

notches minute or absent . . nwzambicus

14. Abdomen bearing transverse furrows

— Abdomen smooth .

15. Three postorbital spines present; large chela without large spines along inner margin but

lateral margin of movable finger bearing bush of setae; abdominal tergite I usually having

short, lateral, transverse furrows . sinensis

2 postorbital spines present; large chela generally bearing some large spines along inner

margin, lateral margin of movable fingers naked; abdominal tergite I generally lacking

distinct transverse furrows . . thomsoni

1 6. Dorsal post-cervical ridge unarmed, except at anterior end . M. challenged

— Dorsal post-cervical ridge spinulose . I 7

17. Posterior margin of hepatic groove armed with spinules; large chela rounded; median

ridge of abdominal tergite VI generally bearing a pair of submedian spines ... M. boschmai

- Posterior margin of hepatic groove devoid of spinules; large chela weakly ridged; median

ridge of abdominal tergite VI generally bearing a median spine . M. sibogae

DEPTH AND GEOGRAPHICAL DISTRIBUTIONS OF THE SPECIES OF METANEPHROPS

Figures 3 and 4 show the known vertical and geographical distributions for the species of Metanephrops

embers of the japonicus group have the widest distribution in the Indo-West-Pacific. The "binghami" group is

PALINURIDAE, SCYLLAR1DAE AND NEPHROPIDAE FROM INDONESIA

425

restricted to the western Atlantic. Species of the "thomsoni" group mainly occur along the western periphery of

the Pacific, from Japan to New Zealand. The "arafurensis" group has a similar, but more restricted, distribution in

the Philippine-Australian region (with only the fossil species found in New Zealand). Since most ot the species

(1 1 out of 18) are present in the Indo-Malay region, the genus probably originated there. JENKINS (1972) is likely

right in suggesting that the "binghami" group originated from the Indo-Malay region and migrated to the Atlantic

through the Tethys, instead of reaching the Atlantic via southern Africa.

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pIG 3. — Vertical disiribution of the extant species of Metanephrops. Thick bars represent main distribution ol the

species.

Amongst the four species groups, the "japonicus" group seems to be the most successful in terms of having

the most species and spread to the other parts of the Indo- West-Pacific. As suggested by JENKINS (1972), the

" thomsoni " group is probably the youngest, and therefore still has a limited distribution. However. I disagree with

JENKINS (1972) when he suggests that members of the "thomsoni" group, by being rather abundant, are actively

displacing species of the other groups north- and southwardly. It is now known that species of different groups are

often found in the same region, and M. australiensis, as well as all the members of the "japonicus group, also

occur in large numbers, being exploited commercially (Chan & Yu, 1987, 1991; Wadley & Evans. 1991,

HOLTHUIS, 1991). On the other hand, species of Metanephrops have rather clear vertical zonations. In the same

Source :

426

T.-Y. CHAN

region, members of the "thomsoni" group are usually found in shallower waters and those of the "arafurensis"

group in deeper waters, while species of the "japonicus" group occur in between the two (CARTER et al. , 1983;

Anon., 1984; Chan & Yu, 1987, 1993; Wallner & Phillips, 1988).

Fig. 4 a. Geographical distribution of the species of Meianephrops of the "binghami" group:A: M. binghami

• : M. rubellus,

ACKNOWLEDGEMENTS

Grateful acknowledgement is extended to A. CROSNIER and B. RICHER DE FORGES, of the Institut framjais de

Recherche scientifique pour le Developpement en Cooperation (ORSTOM), and K. Moosa, of the Puslitbang

Oseanologi-LIPI, Jakarta, for providing the Karubar material for this study; F.A. Chace and R.B. MANNING of

the National Museum of Natural History, Washington D.C., L.B. Holthuis and C.H. Fransen of the Nationaal

Natuurhistorisch Museum, Leiden, for kindly allowing the author to examine the collections of their museums.

I sincerely thank S. Pinkster of the Zoologisch Museum, University of Amsterdam, for sending me on loan

the type of M. arafurensis and the "Siboga" M. andamanicus , D. Evans of the CSIRO Marine Laboratories,

Western Australia, for donating a set of nephropid specimens from Western Australia to the NTOU; D.A. Lee of

the Taiwan Fisheries Research Institute, Keelung, for providing me with informaion on Tungsha Tao

Metanephrops, and the Museum national d'Histoire naturelle, Paris, for providing a short-term research grant to the

author to work in the Museum, thereby making the present study possible. This study was also a contribution

from a research grant on the decapod crustaceans of Taiwan (NSC 84-261 l-B-01 9-006), supported by the National

Science Council, R.O.C.

Source :

PALINURIDAE, SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

427

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Source :

430

T.-Y. CHAN

Fig. 5 a. — Metanephrops velulinus Chan & Yu. 1991. Indonesia, Tanimbar Islands, 7°46'S, 132°31'E, 443-468 m, 8

(POLIPI).

Fig. 5 b. — Metanephrops arafurensis (de Man, 1905). Indonesia, Tanimbar Islands, 7°46'S, 132°31'E, 443-468 m 8

50.7 mm (MNHN).

Fig. 5 c-d. — Metanephrops neptunus (Bruce, 1965): c, Indonesia. Kai or Tanimbar Islands, 1 2 79.2 mm (POLIPI). — d.

Indonesia. Kai Islands, 05°15'S, 132°59'E, 769-809 m, <3 87.1 mm (MNHN).

Source :

PALINURIDAE. SCYLLARIDAE AND NEPHROPIDAE FROM INDONESIA

431

Source : MNHN, Paris

ATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DE

Crustacea Decapoda: Hermit crabs

of the family Paguridae

from the KARUBAR Cruise in Indonesia

Patsy a. McLaughlin

Shannon Point Marine Center

Western Washington University

1900 Shannon Point Road

Anacortes, Washington, 98221-4042, U.S.A.

ABSTRACT

The French-Indonesian 1991 campagne to the islands of Kai, Aru, and Tanimbar, part of the Maluku region of

Indonesia, revealed an unexpected wealth of hermit crabs of the family Paguridae. Although only 295 specimens were

collected in depths ranging from 85 to 1024 meters, an incredible 19 genera and 36 species are represented, of which

seven genera and 26 species are described for the first time. Included are the monotypic Alainopaguroides gen. nov„

Enneopagurus gen. nov„ Enneophyllus gen. nov., lcelopagurus gen. nov., and Tarrasopagurus gen. nov., and their

respective new species. The genus Michelopagurus , gen. nov., is established for "Pagurodes" limatulus Henderson, 1888,

and one additional new species, and the genus Pseudopagurus is created for "Pagurodes" piliferus Henderson, 1888, the

last of the original trio of species initially assigned to the heterogeneous Pagurodes. A lectotype for Pagurodes inarmatus

Henderson, 1888, the type species of the now monotypic Pagurodes , is also designated. The genus Turleania is proposed

as a replacement name for Laurentia McLaughlin & Haig.

Of the new genera, three are particularly noteworthy. Not only are Enneopagurus and Enneophyllus just the second and

third genera of the Paguridae to be characterized, in part, by the absence of gills on the third maxillipeds. the latter genus

is unique, at least for the present. Its type species, E. spinirosiris sp. nov., is the first pagurid known to have a well

developed epi-rosiral spine. Alainopaguroides joins that very specialized group of genera distinguished by marked

reduction in the abdomen, accompanied by total loss of male pleopods and reduction in the number of female pleopods.

Two additional genera of this group, Solitariopagurus and Porcellanopagurus, are also represented in the Karubar

collection, each by a new species.

In addition to the new genera, new species are described in several of the less commonly reported genera, e.g.,

Caiapaguroides, Decaphyllus, Catapagurus, and Tomopaguropsis. Although Pagurus is widely represented in the colder

waters, particularly of the northern hemisphere, the discovery of three new species from the restricted geographic region

of the Karubar campagne was unexpected. A third species has been added to, and extends the distributional range of, the

recently described Bathypaguropsis from Australian and New Zealand waters. A new species described in Australerenius

has provided continuity to the heretofore disjunct distribution of this genus. Only one genus, Pylopaguropsis, was

represented entirely by known species.

McLaughlin, P. A., 1997. — Crustacea Decapoda: Hermits crabs of the family Paguridae from the Karubar Cruise in

Indonesia. In: A. Crosnier & P. Bouchet (eds), Rdsultats des Campagnes MUSORSTOM, Volume 16. Mem. Mus. nain.

Hist. not.. 172: 433-572. Paris ISBN 2-85653-506-2.

Source : MNHN Paris

434

P. A. MCLAUGHLIN

The Karubar collection is also significant for its number of highly evolved genera. Specifically, development of the

male sexual tube(s) is uncommonly prevalent. In the 19 genera included in the collection, males of 13 develop a sexual

tube on one or both coxae of the fifth pereopods, or nearly two-thirds of the total genera.

All species are fully illustrated and detailed descriptions or diagnoses provided. Keys are provided for the regional

genera and species, including those reported from the Maluku area, but not included in the Karubar collection.

RESUME

Crustacea Decapoda : Pagures de la famille des Paguridae recoltes lors de la campagne Karubar

en Indonesie.

La campagne franco-indonesienne Karubar faite aux Moluques, en 1991, dans la region des ties Kai Aru et Tanimbar

a reveld une nchesse inattendue en bemard-l'ermite de la famille des Paguridae. Bien que 295 specimens seulement de cette

famille aient ete rdcoltes k des profondeurs comprises entre 85 el 1024 mdtres, ils foment un ensemble incroyable de 19

genres et 36 espdces, parmi lesquels sept genres et 26 espdces sont nouveaux. On y trouve les genres monotypiques

Alainopagurus gen. nov., Enneopagurus gen. nov., Enneophyllus gen. nov., Icelopagurus gen. nov, et Tarrasopagurus

gen. nov et les espdces nouvelles qui leur correspondent. Le genre Michelopagurus , gen. nov., est dtabli pour

Pagurodes hmatulus Henderson, 1888, et une espdce nouvelle additionnelle, tandis que le genre Pseudopagurus est cree

pour Pagurodes' piliferus Henderson, 1888, la dernidre des trois espdces assignees, h l'origine, au genre hdtdrogdne

Pagurodes. Un lectotype pour Pagurodes inarmatus Henderson, 1888, l'espece type du genre Pagurodes, maintenant

monotypique, est designd. Le genre Turleania est propose en remplacement de Laurentia McLaughlin & Haig, preemploye.

Parmi les nouveaux genres, trois sont particulierement interessants. Non seulement Enneopagurus et Enneophyllus

sont es second et troisidme genres de Paguridae it etre caractdrises, en partie, par l’absence de branchics sur les troisidmes

maxilhpddes, mais en outre, le dernier citd est unique, au moins pour le moment, son espdce type, E. spinirostris, etant le

prem'er pagunde connu d posseder une epine epirostrale bien developpde. Alainopagurus fait partie du groupe trds

specialist de genres se distinguant par une reduction marquee de I'abdomen. accompagnde par la perte totale des pleopodes

males et la reduction en nombre des pleopodes femclles. Deux genres appartenant d ce groupe, Soliiariopagurus et

Porcellanopagurus, sont dgalement reprdsentes dans les rdcoltes de Karubar, chacun par une espdce nouvelle,

Des especes nouvelles sont dgalement decrites dans plusieurs autres genres peu communs, it savoir Catapaguroides,

Decaphylius, Caiapagurus el Tomopaguropsis. Bien que Pagurus soil largement reprdsente dans les eaux froides, en

HIemiSphere n°^’ 3 d,§C0l!verte de tr0ls nouvelles espdces appartenant d ce genre dans la region restreintc

prospectee par la campagne Karubar eta.t inattendue. Une troisidme espdce a ete ajoutee au genre Bailiypaguropsis

rdcemment decrit des eaux australiennes et ndo-zelandaises. La description d'une nouvelle espdce dans'le genre

Ausualeremus permet de rendre coherente la distribution de ce genre jusqu'd present discontinue. Seul un genre

Pvlopaguropsis, dtait reprdsentd dans la collection par des espdces toutes connues.

La collection Karubar est dgalement significative par le nombre de genres trds evolues qu'elle renferme En

da^ ?a co\'lectiirenPrTen‘ n’a'eS CS' anormalemenI predominant. Parmi les 19 genres representes

“• m“es a,ec un "be se"ei d<i,e,oppb sur d™ c»“

r^„rT,0Ule:! !eS esp6ces s°nl 'purees et ddcrites en detail ou des diagnoses sont fournies. Des cles d'identification

proposdes. §CnreS ^ CSP T*C°MS ^ KARUBAR et ceux et celIes d6Ja signalds de la rdgion dtudiee sont

INTRODUCTION

Prior to the 1991 French-Indonesian campaign, Karubar (named for the islands of Kai, Aru and Tanimbar)

frnm?i° ma")ne herna,t^'rab fauaa of the Maluku (formerly Moluccas) region of Indonesia was known primarily

I aurInt iSS KnSJ t J :MlERS’ I884)’ "ChallenSer" (HENDERSON, 1888), "Siboga" ( DE Saint

NT, 1968a, b McLaughlin & Haig, 1996), "Snellius" (Buitendijk, 1937) and "Alpha Helix" (Humes

, Forest, 1984, Haig & Ball, 1988), and the shallow water collections of the Indonesian Institute of

nl r/T °Rr- ’r93' ‘"5)' Eigh‘ SpedeS Were rep0rted in the combincd collections of the

A/m and Challenger and one from the "Snellius". Five species from the "Siboga" collections were descr.bed

by“ScAINT Lavrent <1968a- b); however, in subsequent publications Forest and de Saint Laurent (1968)

mSTTS ,ndlCated thal nUmer0US SPeCieS fr0m lhat expedili0n ~d - described

,1 HA,Gf(1i"6> have just recently described three of those. Rahayu and FOREST (1993 1995)

although To, !.|PeCieS d'°fnid gCnUS Di°geneS 3nd 20 SpedeS 0f Clibanarius from Indonesian waters,

although not all were represented ,n the Maluku region. Isolated species reports have also come from severa

PAGURIDAE FROM THE KARUBAR EXPEDITION

435

sources, particularly those concerning associations with rhizocephalans, e.g., the "Siboga" (van Kampen &

BOSCHMA, 1925; BOSCHMA, 1931b); MORTENSEN's Pacific Expedition (BOSCHMA, 1931a), and the Danish

Expedition to the Kei Islands (van Baal, 1937), as well as museum collections (DE Man, 1881; Lewinsohn,

1969); however, the most comprehensive report is that of Haig and Ball (1988) from the "Alpha Helix"

expedition. These authors documented the occurrence of 46 marine species (including four new and four left

undescribed); however, most were collected at depths of 20 meters or less; seven genera of the Paguridae were

represented.

It has long been postulated that the most diverse marine faunas are to be found in the tropical oceans, especially

the Indo-West Pacific, but at depths generally less than 200 meters (e.g., Ekman. 1953; BRIGGS, 1974). Within

the Paguridea (sensu FOREST, 1987), most hermit crabs inhabiting these tropical environs were thought to belong

to the family Diogenidae. The hermit crabs of the family Paguridae that were collected during the Karubar

expedition consisted of 295 specimens, all coming exclusively from depths ranging from 85 to 1024 meters. The

assemblage includes an astonishing 19 genera, of which seven are proposed herein, and 36 species, 29 of which are

reported for the first time. More than 65 percent of the species covered by this report were collected from depths in

excess of 200 meters, whereas only four species appeared restricted to more shallow depths.

Of the new genera, Michelopagurus gen. nov., with type species Pagurodes limatulus Henderson, 1888, has

been erected for one of two species originally described in the heterogeneous genus Pagurodes Henderson, 1 888,

but subsequently restricted from that genus by designation of P. inarmatus Henderson, 1888, as the type species

(DE Saint Laurent, 1969). Although the second species, Pagurodes piliferus Henderson, 1888, is not present in

the KARUBAR collection, it is in the interest of stability in nomenclature that a new genus, Pseudopagurodes gen.

nov., also be established for it.

The general terminology used in the species descriptions is that of McLaughlin (1974), with exception of the

fourth pereopods. A distinction is made herein between subehelate fourth pereopods, in which the pereopod is

developed as a prehensile structure by the folding back of the dactyl against the propodus, and semichelate fourth

pereopods, where the ventral margin of the propodus is produced beneath the dactyl to such an extent that flexion

of the dactyl becomes much more akin to the action of a dactyl against a fixed finger of a chelate appendage. Terms

pertaining to regions of the carapace follow those proposed by Pilgrim ( 1973) and Morgan and Forest (1991).

Gill structure, i.e., trichobranchiate, intermediate, or phyllobranchiate follow the definitions provided by

LEMAITRE (1989). The lists of specimens examined follow the station data provided by CROSNIER el al. (1997).

The station abbreviations DW, CP, and CC refer to Waren dredge, beam trawl, and shrimp trawl respectively.

Shield lengths (to the nearest 0. 1 mm) of the specimens examined are indicated in parentheses, and measured from

the tip or midpoint of the rostrum to the midpoint of the posterior margin of the shield. In keys provided for the

Maluku regional taxa, those not encountered during the Karubar expedition are indicated by an asterisk (*). The

majority of the specimens reported herein are shared between the Museum national d'Histoire naturelle, Paris

(MNHN) and the Puslitbang Oseanologi - LIPI, Jakarta (POLIPI). Supplemental materiels of selected species are

deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM) and

the Swedish Natural History Museum, Stockholm (SNHM). Comparative materials used in the study have come

from the Museum national d’Histoire naturelle (MNHN). Museums and Art Galleries of the Northern Territory.

Darwin, Australia (MNT), National Museum of Natural History, (USNM), Natural History Museum. London

(NHM), Natural History Museum and Institute, Chiba, Japan (CBM-ZC), Rosenstiel School of Marine and

Atmospheric Sciences, University of Miami (UMML), and the author's personal collection. Photographs were

taken with a Nikon 35 mm camera and a professional camera 4x5 inches.

LIST OF GENERA AND SPECIES

Genus ALAINOPAGUROIDES gen. nov.

A. lemaitrei sp. nov.

Genus ANAPA GRIDES de Saint Laurent-Dechance, 1966.

? Anapagrides sp.

436

P. A. MCLAUGHLIN

Genus ANAPAGURUS Henderson, 1886 (key).

Genus AUSTRALEREMUS McLaughlin, 1981.

A. indonesiensis sp. nov.

A. triserratus (Ortmann, 1 892).

Genus BATHYPAGUROPSIS McLaughlin, 1994.

B. rahayuae sp. nov.

Genus CATAPAGUROIDES A. Milne Edwards & Bouvier, 1892.

C. cristimanus de Saint Laurent, 1968 (key).

C. declivis sp. nov.

C. inermis de Saint Laurent, 1968 (key).

C. karubar sp. nov.

C. melini de Saint Laurent, 1968 (key).

C. mortenseni de Saint Laurent, 1968 (key).

C. spinulimanus de Saint Laurent, 1968 (key).

Genus CATAPAGURUS A. Milne Edwards, 1880.

C. ensifer Henderson, 1 893 (key).

C. holthuisi sp. nov.

C. oculocrassus sp. nov.

Catapagurus sp. of Haig & Ball, 1988 (key).

C. tanimbarensis sp. nov.

Genus DECAPHYLLUS de Saint Laurent, 1968.

D. barunajaya sp. nov.

D. junquai de Saint Laurent, 1 968 (key).

D. maci sp. nov.

D. similis de Saint Laurent, 1968 (key).

Genus ENNEOPAGURUS gen. nov.

E. garciagomezi sp. nov.

Genus ENNEOPHYLLUS gen. nov.

E. spinirostris sp. nov.

Genus ICELOPAGURUS gen. nov.

I. crosnieri sp. nov.

Genus MICHELOPAGURUS gen. nov.

M. chacei sp. nov.

M. limaiulus (Henderson, 1888).

Genus MICROPAGURUS McLaughlin, 1986 (key).

Genus NEMATOPAGURUS A. Milne Edwards & Bouvier, 1892.

N. alcocki sp. nov.

N. australis (Henderson, 1888) (key).

N. cf. indicus Alcock, 1905.

N. ostlingochirus sp. nov.

N. scutelliformis sp. nov.

Nematopagurus sp.

N. spinulosensoris McLaughlin & Brock, 1974.

Genus PAGURODES Henderson, 1888.

P. inarmatus Henderson, 1888 (photos).

Genus PAGURUS Fabricius, 1775.

P. capsularis sp. nov.

PAGURIDAE FROM THE KARUBAR EXPEDITION

437

P. compressipes (Miers, 1884) (key, photos).

P. haigae sp. nov.

P. hedleyi Grant & McCulloch, 1906 (key).

P. hirtimanus (Miers, 1880) (key).

P. kaiensis sp. nov.

P. moluccensis Haig & Ball, 1988 (key).

P. pergranulatus (Henderson, 1 896) (key).

IPagurus sp.

Genus PORCELLANOPAGURUS Filhol, 1885.

P. jacquesi sp. nov.

Genus PSEUDOPAGURODES gen. nov.

P. piliferus (Henderson, 1888) (photos).

Genus PYLOPAGUROPSIS Alcock, 1905.

P.fimbriata McLaughlin & Haig, 1989 (key).

P. laevispinosa McLaughlin & Haig, 1989.

P. lewinsohni McLaughlin & Haig, 1989 (key).

P. zebra (Henderson, 1893).

Genus SOL1TARIOPAGURUS Tiirkay, 1986.

S. tuerkayi sp. nov.

Genus SPIROPAGURUS Stimpson, 1858 (key).

Genus TARRASOPAGURUS gen. nov.

T. rostrodenticulatus sp. nov.

Genus TOMOPAGUROPSIS Alcock, 1905.

T. crinita sp. nov.

T. miyakei sp. nov.

Genus TURLEANIA nom. nov.

T. albatrossae (McLaughlin & Haig. 1996) (key).

T. balli (McLaughlin & Haig. 1996) (key).

T. multispina sp. nov.

T. senticosa (McLaughlin & Haig, 1996).

T. sibogae (McLaughlin & Haig, 1996) (key).

SYSTEMATIC ACCOUNT

Family PAGURIDAE Latreille, 1803

Key to the genera of the Maluku Paguridae

1 . Crista dentata of third maxilliped with accessory tooth . 6

— Crista dentata of third maxilliped without accessory tooth . 2

2. Rostrum strongly deflected downward over ocular lobes; with epi-rostral spine .

. Enneophyllus gen. nov.

— Rostrum not strongly deflected downward over ocular lobes; without epi-rostral spine ... 3

3. Pleurobranch present above fourth pereopod; males with elongate left sexual tube; females

with paired gonopores . . 4

— Pleurobranch absent above fourth pereopod; males with elongate right sexual tube; females

with single left gonopore . 5

Source : MNHN Paris

438

P. A. MCLAUGHLIN

4. Male sexual tube with terminal tuft of sparse setae; paired arthrobranchs on third

maxillipeds . Turleania nom. nov.

— Male sexual tube with terminal fringe of dense stiff setae; no paired arthrobranchs on third

maxillipeds . Enneopagurus gen. nov.

5. Males with 3 unpaired left pleopods; fourth pereopod semichelate . Catapaguroides

— Males with 4 unpaired left pleopods; fourth pereopod simple, not semichelate .

. Decaphyllus

6. Males with sexual tube developed on one or both coxae of fifth pereopods . 7

— Males without sexual tube developed on one or both coxae of fifth pereopods . 17

7. Abdomen well developed; males with 2 to 4 unpaired left pleopods . 9

— Abdomen reduced; males without unpaired left pleopods . 8

8. Pleurobranch present above fourth pereopod; lateral margins of shield rounded .

. Alainopaguroides gen. nov.

— Pleurobranch absent above fourth pereopod; lateral margins of shield drawn out into

spinose projections . Solitariopagurus

9. Females with paired first pleopods modified as gonopods . 1 0

— Females without paired first pleopods modified as gonopods . 1 2

10. Chelipeds subequal; rostrum smoothly rounded; males with very short to very long sexual

tube on coxa of right fifth pereopod; left sometimes also with very short sexual tube . 1 1

— Chelipeds markedly unequal; rostrum denticulate; males with moderately short sexual tube

on coxa of left fifth pereopod, right sometimes also with short tube .

. Tarrasopagurus gen. nov.

11. Males with elongate right sexual tube; ocular acicles very broadly separated; gills

phyllobranchiate . Nematopagurus

— Males with very short right sexual tube; ocular acicles separated by approximately basal

width of 1 acicle; gills trichobranchiate . Michelopagurus gen. nov.

12. Males with well developed right sexual tube . 13

— Males with well developed left sexual tube . 1 5

13. Right sexual tube elongate, curving up over dorsal surface of body from right to left;

telson subtriangular . Catapagurus

— Right sexual tube short, not curving up over dorsal surface of body; telson not

subtriangular . 1 4

14. Ocular acicles elongate; chelipeds subequal; fourth pereopods with prominent tubular

preungual process at base of claw . Icelopagurus gen. nov.

— Ocular acicles short; chelipeds markedly unequal; fourth pereopods without prominent

preungual process . Anapagrides

15. Telson with median cleft; ocular acicles simple . 16

— Telson without median cleft; ocular acicles multifid . Micropagurus*

16. Chelipeds grossly unequal . Anapagurus*

— Chelipeds subequal . Spiropagurus*

17. Pleurobranchs above second, third and fourth pereopods . 20

— Pleurobranchs above fourth pereopod only . 18

18. Abdomen reduced; lateral margins of shield drawn out into strongly calcified projections;

males without unpaired left pleopods . Porcellanopagurus

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

439

— Abdomen not reduced; lateral margins of shield not drawn out into strongly calcified

projections; males with some unpaired left pleopods . 1 9

19. Females with first pleopods paired and modified as gonopods; right chela circumscribed by

row of spines . Australeremus

— Females without first pleopods paired and modified as gonopods; right chela not

circumscribed by row of spines . Pagurus

20. Chelipeds subequal . Tomopaguropsis

— Chelipeds grossly unequal, right markedly larger than left . 21

21. Females with first pleopods paired and modified as gonopods; males with 3 unpaired left

pleopods . Pylopaguropsis

— Females without first pleopods paired and modified as gonopods; males with four unpaired

left pleopods . Bathypaguropsis

Genus ENNEOPHYLLUS nov.

DIAGNOSIS. — Nine pairs of phyllobranchiate gills; arthrobranchs absent from third maxillipeds. Rostrum well

developed, strongly depressed. Ocular acicles simple. Antennal peduncle with supernumerary segmentation not

clearly evident. Third maxilliped with 1 spine on basis; ischium with crista dentata somewhat reduced, without

accessory tooth (Fig. la). Sternite of third maxillipeds unarmed. Chelipeds unequal, right appreciably larger.

Dactyls of ambulatory legs with spinose ventral margins. Sternite of third pereopods with small anterior lobe.

Fourth pereopods semichelate, with single row of scales in propodal rasp. Fifth pereopods weakly semichelate.

Sternite of fifth pereopods developed as single small lobe.

Coxa of left fifth pereopod in males with elongate, basally stout sexual tube directed exteriorly and curved

dorsally across abdomen from left to right, with few terminal setae; right fifth coxa with gonopore, no apparent

sexual tube; 3 unequally biramous unpaired left pleopods. Females not known.

Abdomen straight; uropods (Fig. Ik) only slightly asymmetrical. Telson with transverse suture very weakly

indicated; terminal margins oblique.

TYPE SPECIES. — Enneophyllus spinirostris sp. nov. Gender masculine.

ETYMOLOGY. — From the Greek ennea meaning nine, and phyllon meaning leaf, and referring to the nine

pairs of phyllobranchiate gills in this genus.

AFFINITIES. — Enneophyllus very closely resembles the other two genera of pagurids now recognized that have

only nine pairs of gills, Enneobranchus Garcia-Gomez, 1988, and Enneopagurus gen. nov. All three are

characterized by the absence of an accessory tooth on the crista dentata of the third maxilliped, an elongate left male

sexual tube, and a subtriangular telson that has the transverse suture faintly, if at all, indicated. However, the three

genera differ fundamentally in gill structure: phyllobranchiate in Enneophyllus, intermediate in Enneobranchus, and

trichobranchiate in Enneopagurus. Additional characters that distinguish the three taxa include: 1) the termination

of the left sexual tube - apparently lacking any terminal setae in Enneobranchus (cf. GARCIA-G6MEZ, 1988), with

a dense fringe of setae in Enneopagurus, but only a very few setae in Enneophyllus-, 2) a distinctive preungual

process at the base of the dactylar claw on the fourth pereopod in Enneobranchus, but lacking in both

Enneopagurus and Enneophyllus-, and 3) subequal chelipeds in both Enneobranchus and Enneopagurus, but

conspicuously unequal in Enneophyllus.

Remarks. — Enneophyllus at present is known only from a single representative of the type species,

E. spinirostris sp. nov. For this reason, no attempt has been made at this time to determine the precise structure

440

p.a. McLaughlin

of the mouthparts in this genus; however, a flagellum on the first maxilliped can be observed without dissection.

The supernumerary segment of the antennal peduncle that is usually calcified and readily apparent is, in this

specimen, represented only as a chitinous area between the third and fourth segments. A character that sets

Enneophyllus apart, not only from Enneobranchus and Enneopagurus , but all other pagurids, is its remarkable

rostral structure. The rostrum is exceptionally well developed, lobe-like, and bent downward over the ocular plate

between the ocular acicles (Fig. lb-c). At least in E. spinirostris a prominent epi-rostral spine is developed.

Enneophyllus spinirostris sp. nov.

Figs la-k, 33a-b

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands : stn DW 49. 08°00'S, 132°59'E, 210-206 m,

29.10.1991: 1 <J (1.5 mm) (MNHN-Pg 5250).

TYPES. — The unique specimen is the holotype.

DESCRIPTION. — Shield (Fig. lb) considerably longer than broad, but with lateral portions distinctly rounded;

anterior margin between rostrum and lateral projections concave; anterolateral margins terraced; posterior margin

truncate; dorsal surface glabrous, but with small areas of decalcification anterolaterally. Rostrum strongly produced

as rounded lobe, deflected downward over ocular lobes, and provided with extremely prominent epi-rostral spine

(Fig. lb-c). Lateral projections well developed, acutely triangular, with marginal or submarginal spine.

Ocular peduncles (including corneae) approximately 0.60 shield length; peduncles appearing somewhat laterally

compressed; corneae slightly dilated. Ocular acicles moderately small, roundly triangular, with submarginal spine;

separated basally by width of rostrum or equivalent to basal width of 1 acicle.

Antennular peduncles, when fully extended, overreaching ocular peduncles by 0.80 length of ultimate segment.

Ultimate segment with 1 long seta dorsodistally. Penultimate segment with few short setae. Basal segment with

statocyst region expanded laterally, with small spine on dorsolateral margin distally.

Antennal peduncles overreaching ocular peduncles by approximately half length of ultimate segment. Fifth and

fourth segments with scattered setae. Third segment with small ventrodistal spinule. Second segment with

dorsolateral distal angle produced, terminating in acute simple spine and with slightly smaller spine on mesial

margin; dorsomesial distal angle with prominent acute spine. First segment with spine at dorsolateral distal angle;

1 small spine on ventrolateral margin distally. Antennal acicle reaching to proximal margin of fifth peduncular

segment; straight, terminating in acute spine and with few very short setae. Antennal flagellum short, not reaching

tip of dactyl of left cheliped, with 1 or 2 short (1 or 2 article length) setae on each article. Crista dentata with

7 relatively evenly-sized teeth (Fig. la).

Chelipeds unequal; right appreciably larger. Right cheliped (Fig. Id) moderately elongate, chela (Fig. 33a)

operculate; propodal-carpal articulation perpendicular. Dactyl 0.80 length of palm, broad; cutting edge with

2 distinct calcareous teeth; terminating in small calcareous claw; slightly overlapped by fixed finger; dorsal surface

flattened, smooth, dorsomesial margin drawn out into subacute obliquely elevated unarmed ridge; mesial and

ventral surfaces with few scattered setae. Palm approximately equaling length of carpus; dorsomesial margin

rounded proximally, strongly elevated into prominent acute crest in distal half, with 1 spinule basally on distal

margin; dorsal surface weakly convex on palm, but weakly concave on fixed finger, dorsolateral margin not

delimited in proximal third of palm, but produced as obliquely elevated crest distally and extending length of fixed

finger; mesial face of palm and ventral surfaces of palm and fixed finger with scattered setae; cutting edge of fixed

finger with 2 prominent calcareous teeth, terminating in calcareous claw. Carpus slightly shorter than merus;

dorsomesial distal angle with small subacute spine, dorsomesial margin with few low protuberances and long

setae, all surfaces with sparse tufts of setae, most numerous and longest on mesial and ventral surfaces. Merus

subtnangular; dorsodistal margin with small spine; dorsal surface with row of very small protuberances and few

setae; ventromesial margin with prominent spine at distal angle and 1 smaller spine in distal third; ventrolateral

margin with 1 spinule at distal angle. Ischium with row of minute spinules and few setae on ventromesial margin.

PAGURIDAE FROM THE KARUBAR EXPEDITION

441

Fig. 1. — Enneophyllus spinirostris sp. nov., holotype 8 (1.5 mm) from Karubar Stn DW 49: a, basis and ischium ol

left third maxilliped; b. shield and cephalic appendages; c, rostrum and right ocular acicle (lateral view); d. right

cheliped (mesial view); e. left cheliped (mesial view); f. right second pereopod (lateral view); g. left third pereopod

(lateral view); h. anterior lobe of sternite of third pereopods; i, coxae and sternite of fifth pereopods; j, left sexual

tube (dorsal view); k, uropods and telson. Scales equal 0.5 mm (a, c, h. k) and 1.0 mm (b. d-g. i-j).

Left cheliped (Figs le, 33b) with propodal-carpal articulation nearly perpendicular; chela somewhat

dorsoventrally flattened. Dactyl only slightly longer than palm, unarmed but with scattered setae, longer and more

Source :

442

P. A. MCLAUGHLIN

numerous ventrally; cutting edge with row of small corneous teeth, terminating in small corneous claw. Palm

approximately 0.65 length of carpus; row of few low protuberances and tufts of long setae on rounded dorsomesial

margin, dorsal surface of palm and fixed finger smooth, with few scattered short setae, dorsolateral margin rounded

on palm, minutely spinulose on fixed finger; cutting edge of fixed finger with jagged row of very small calcareous

teeth, terminating in very small corneous claw. Carpus approximately equal to length of merus, dorsal surface

broadened disially; dorsomesial distal angle with small spine, dorsomesial margin with row of spinulose

protuberances and tufts of long setae, dorsal midline with spine at distal margin and longitudinal row of minute

protuberances and sparse tufts of setae, dorsolateral margin only weakly delimited by few minute protuberances and

few setae. Merus subtriangular; dorsal margin with row of low spinulose protuberances and setae, 1 spine at distal

margin; ventrolateral margin with 1 very small spine at distal angle; ventromesial margin with 1 prominent spine

at distal margin. Ischium with row of small spines on ventromesial margin.

Second and third pereopods (Figs lf-g) moderately short, not overreaching outstretched right cheliped; generally

similar from left to right. Dactyls approximately 0.25 longer than propodi (left second and third broken in

holotype); in dorsal view, generally straight; in lateral view slightly curved ventrally; terminating in slender

corneous claws; dorsal margins each with row of widely-spaced long moderately stiff setae; ventral, lateral and

mesial faces with few scattered setae; ventral margins with 5 or 6 corneous spines. Propodi with 1 corneous spine

on ventrolateral distal margin, dorsal and ventral surfaces each with row of widely-spaced low protuberances and

long moderately stiff setae. Carpi each with 1 spine at dorsodistal angle, row of low protuberances and long

moderately stiff setae on dorsal surface and 1 additional small spine. Mcri each with low protuberances and sparse

tufts of setae dorsally and ventrally; ventrodistal margins each with small spine (second) or unarmed (third). Ischia

unarmed, but with sparse tufts of setae dorsally and ventrally. Fourth pcreopod with small spine at dorsodistal

margin of carpus. Sternite of third pereopod with roundly rectangular anterior lobe (Fig. lh), unarmed but with few

long setae. Sternite of fifth pereopods (Fig. li) narrow, with few distal setae.

Left male sexual tube (Figs li-j) very stout basally; right gonopore almost completely obscured by circle of

short setae. Abdomen elongate, nearly twice length of cephalothorax, straight; membraneous, with no indication of

segmentation of somites 2-5; tergite of somite 6 well calcified. Telson (Fig. Ik) with posterior lobes nearly

symmetrical, each with prominent spine at outer angle and I or 2 tiny corneous spinules on oblique terminal

margins, median cleft moderately deep; lateral margins each with chitinous plate.

Color (in preservative). — Calcified integument somewhat iridescent; tint of orange on chelae and proximal

portions of fixed fingers and dactyls, distal portions white; ambulatory legs with tint of orange on dactyls and

distal halves of propodi.

Habitat. — Scaphopod shell.

Distribution. — Known only from the type locality in the Tanimbar Islands, Indonesia; 206-210 m.

Etymology. — From the Latin spina , meaning spine and rostrum , and indicating the presence of the unusual

epi-rostral spine.

Affinities. — While clearly not closely related to any known pagurid taxa, Enneophyllus spinirostris does

possess a number of convergent characters. The shared generic characters have already been discussed. At the

’Jooo C 'eVe1, the °Perculate r‘8hl chela is reminiscent of species of Catapaguroides A. Milne Edwards & Bouvier

1892 and Pylopagurus A. Milne Edwards & Bouvier, 1891. The use of a scaphopod microhabitat is also seen in

some species of Pylopagurus, Orthopagurus Stevens, 1927, occasionally Pagurus species (cf. McLaughlin &

KONISHI, 1994), and the parapagurid genus Tsunogaipagurus Osawa, 1995. The left sexual tube curving over the

abdomen is suggestive of species of Catapagurus A. Milne Edwards, 1880.

Remarks. — The phylogenetic position of this extraordinary taxon cannot be even speculated upon until the

morphology of the female is known, and until sufficient specimens become available to permit a more detailed

study of the mouthparts and "skeletal" anatomy. Nonetheless, in having both reduced gill number and well devel¬

oped sexual tube, it can be hypothesized that the rostral development reflects yet another apomorphic character

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

443

Genus ENNEOPAGURUS nov.

DIAGNOSIS. — Nine pairs of trichobranchiate gills; arthrobranchs absent from third maxillipeds. Rostrum

moderately well developed, triangular, not deflected. Ocular acicles simple. Antennal peduncle with supernumerary

segmentation. Maxillule (Fig. 2a) with external lobe of endopod somewhat produced, not recurved, but provided

with 2 or 3 moderately long setae. Third maxilliped with at least 1 prominent spine on basis; ischium with crista

dentata moderately well developed but without accessory tooth (Fig. 2b). Chelipeds subequal, right more robust.

Dactyls of ambulatory legs with unarmed or very weakly spinulose ventral margins. Sternite of third pereopods

with subquadrate anterior lobe. Fourth pereopods semichelate; with single row of scales in propodal rasp. Fifth

pereopods semichelate. Sternite of fifth pereopods developed as single small subovate or subcircular lobe.

Coxa of left fifth pereopod in males with well developed, rather stout sexual tube (Figs 3a-b) directed exteriorly

and upward, terminally somewhat spatulate and with fringe of dense curved setae; right fifth coxa with gonopore

and occasionally vas deferens slightly protruded; 3 unequally biramous unpaired left pleopods. Females with paired

gonopores; no paired pleopods, unpaired left pleopods on somites 2 to 5.

Uropods asymmetrical. Telson with transverse suture weakly indicated; terminal margins oblique.

Type Species. — Enneopagurus garciagomezi sp. nov. Gender masculine.

ETYMOLOGY. — From the Greek ennea meaning nine, and pagouros meaning crab, and referring to the

presence on only nine pairs of gills in this genus.

Affinities. — As previously noted, Enneopagurus , Enneophyllus gen. nov. and the Atlantic genus

Enneobranchus share several generic characters, however, none are mutually exclusive. Despite our limited

knowledge of its morphology, Enneophyllus is so markedly different from the other two genera that its shared

characters must be considered convergent.

Differences in gill morphology, terminal setation of the male sexual tube, and the absence of a distinctive

preungual process at the base of the dactyl are sufficient to distinguish Enneopagurus from Enneobranchus ;

however, their overall morphological similarities suggest a generally close relationship. Nevertheless, if gill

number is not considered, Enneopagurus appears even more closely allied to the Indo-Pacific genus, Turleania

nom. nov. Species of both of these genera have trichobranchiate gills, lack an accessory tooth on the crista dentata

of the third maxilliped, and have the well developed male left sexual tube terminating in a tuft of setae (Figs 3a-d).

The two genera are separated, not only by the presence of arthrobranchs on the third maxilliped in Turleania nom.

nov., but by differences in the armature of the dactyls of the ambulatory legs and in setal development of the male

sexual tube. In all known species of Turleania nom. nov. the ventral margins of the dactyls of the ambulatory legs

are each provided with a row of well developed spines; the sexual tube is an elongate, usually somewhat spiraled

structure with rounded tip and sparse terminal tuft of straight setae. The dactyls of the ambulatory legs in

Enneopagurus, however, are either unarmed or have only a very few tiny corneous spinules; the left sexual tube is

moderately short, curved toward the exterior, with a broad, spatulate tip practically obscured by a dense fringe of

curved setae.

REMARKS. — Despite their differences, Enneopagurus and the Atlantic Enneobranchus might be considered

analog genera, set apart from most other pagurid genera chiefly by the absence of gills on the third maxillipeds.

Two additional characters shared by these genera, i.e., lack of an accessory tooth on the crista dentata, and left male

sexual tube closely align them with a second analog pair of Indo-Pacific/ Atlantic genera, Turleania nom. nov. and

Iridopagurus de Saint Laurent-Dechance, 1966a.

Garcia-G6mez (1988) related Enneobranchus to Anapagrides de Saint Laurent-Dechanc6, 1966b (sensu lato)

and Iridopagurus de Saint Laurent- Dechancd, 1966a, noting that all three genera shared the diagnostic characters of

1) a long left sexual tube and, 2) absence of an accessory tooth on the crista dentata. He also believed that they all

possessed "intermediate" type branchiae, defining his term "intermediate" by reference to LEMAlTRE's subsequently

published PhD dissertation (Lemaitre, 1989, Fig. 2I-K). Recently McLaughlin and SANDBERG (1995) showed

444

P. A. MCLAUGHLIN

that DE Saint Laurent- Dechance's (1966b) Anapagrides , as defined by its type species, A. facetus (Melin,

1939), did not correspond to the diagnosis given by its author, nor by DE Saint Laurent (1968b), and emended

Anapagrides to reflect the characters of A. facetus. Subsequently, McLaughlin and Haig (1996) proposed the

genus Laurentia, a name now found to be a junior homonym (see, p. 477) for DE Saint Laurent-Dechance's

(1966b) taxon. Garcia-GCmez's (1988) remarks now apply to Turleania nom. nov.

Enneopagurus, Turleania nom. nov., Enneobranchus, and Iridopagurus appear to form a very cohesive

phylogenetic unit in being four of only five taxa presently known that have a left male sexual tube, but lack an

accessory tooth on the crista dentata. Additionally, species of all four share, albeit not exclusively, elongate

ambulatory dactyls, propodal rasps of the fourth pereopods which consist of a single row of scales, and generally

subtriangular telsons. However, some interesting divergent pathways, both morphological and geographic, should

be noted: 1) gills in species of Turleania nom. nov. are trichobranchiate, as are those of Enneopagurus,, species of

the Atlantic Iridopagurus and Enneobranchus have intermediate gills; 2) the external lobe of the endopod of the

maxillule is well developed, but not recurved, in species of Enneopagurus and Turleania nom. nov., but obsolete

in Enneobranchus (cf. GARCIA-GOMEZ, 1988) and Iridopagurus (cf. Garcia-Gomez, 1983); 3) the rostrum is

triangular and well developed in Enneopagurus and Turleania species, in contrast to the reduced and broadly

rounded rostra of species of both Atlantic genera; 4) the transverse suture dividing the telson into anterior and

posterior lobes is weakly indicated in the Indo-Pacific genera, but apparently absent in Enneobranchus and

Iridopagurus. Although there is a suggested trend toward simplification in morphological structure from Indo-

Pacific to Atlantic in these analog genera, all are still highly complex taxa about which we still have only

fragmented knowledge.

Enneopagurus garciagomezi sp. nov.

Figs 2a-j, 3a-b, 33 c-f

” CP °7‘47'S' 132°“'E' 4“'477 m'

nJ^n?L‘Slands- Stn CP 56' 08°16'S- 13i°59'E, 552-548 m, 31.10.1991: 3 «J . 2 9 , 1 ov. 9 (2.4-3.3 mm)

(LSNM 276005). Stn CP 59, 08°20'S, 132°11'E, 405-399 m, 31.10.1991: 2 3, 2 5 (2 7-3 3 mm) (SNHM 4808)

Stn CP 70. 08°41'S, 131°47'E, 413-410 m, 2,11.1991: 1 <J (3.3 mm) (MNHN-Pg 525 1) - S tn CP 71

! 3 1 °47'E ' 413 ~4\0 m ' 2 \ \\ qqi ' 7 ' !? *8 * 5 <'•«»•* S3 525^ - Sm CP 7fr OS'llt]

08“42S l3|O5rt0 fsk 5 i ;3n mm) (MNHN‘PS 5253>- 1 5 (2.8 mm) (POLIPI). - Stn CP 69,

131°36'E 452 451 min ?q o. J ^ 1 ov‘ 9 (2-3‘3-6 mm) <MNHN-pg 5254). - Stn CP 75, 08°46'S,

3.1 1.1991: 6^^(I.8-3./mni)^JSNM^ ^ ~ ** " °8°5TS- 13 352’346

ai,VPESu ~ The male (3-3 mm- MNHN-Pg 5251) collected at the station 70 of Karubar cruise is the holotype

All tne other specimens are paratypes.

Description. — Shield (Fig. 2c) slightly broader than long or length approximately equal to width; anterior

margin between rostrum and lateral projections concave; anterolateral margins sloping or terraced; posterior margin

truncate, dorsal surface with several tufts of setae. Rostrum triangular, well developed, reaching beyond bases of

ocular ac.cles, terminating subacutely and with tiny spinule. Lateral projections well developed, acutely or

obtusely triangular, with submarginal spine.

Ocular peduncles (including comeae) approximately 0.75 shield length; corneae strongly dilated. Ocular acicles

narrowly triangular, with submarginal spine; separated basally by less than 0.50 to 0.75 basal width of 1 acicle

Antennular peduncles, when fully extended, overreaching ocular peduncles by 0.75 to nearly entire length of

ultimate segment. U timate segment with 5 to 8 long setae forming arc on dorsodistal margin and irregular row of

exoLdedir ^.0rSal,S,Urface- Penultimate segment with few short setae. Basal segment with sta.ocyst region

expanded la eral y and dorsoventrally flattened, with small spine near dorsolateral margin in distal half

Antenna] peduncles overreaching ocular peduncles by 0.50 to 0.75 length of ultimate segment. Fifth and fourth

segments with scattered long setae. Third segment unarmed or with small ventrod.stal spLle. Second segme

with dorsolateral distal angle produced, terminating in acute simple or more frequently bifid spine; dorsomesia!

Source :

PAGURIDAE FROM THE KARUBAR EXPEDITION

445

distal angle with prominent acute spine. First segment with small spine at dorsolateral distal angle; 1 or 2 very

small spines on ventrolateral margin distally. Antennal acicle frequently reaching to distal margin of cornea or

beyond; terminating in acute spine and with long setae on mesial margin. Antennal flagellum long, overreaching

outstretched chelipeds, with 1 to 3 long and sometimes 1 or 2 shorter setae every 1 to 3 articles, at least in

proximal third. Crista dentata with 7 to 14 regularly or irregularly-sized teeth, decreasing in size distally.

Fig 2. — Enneopagurus gareiagomezi sp. nov., a-b, paratype 2 (3.6 mm) from Karubar Stn CP 69; c-i, holotype 6

(3.3 mm) from Stn DW 70; j, paratype (2.9 mm) from Stn CP 39: a. left maxillule; b. basis and ischium of left third

maxilliped; c, shield and cephalic appendages; d, right cheliped (dorsal view); e, left cheliped (dorsal view); f, right

second pereopod (lateral view); g, left third pereopod (lateral view); h, anterior lobe of sternile of third pereopods;

i-j, telson. Scales equal 0.5 mm (a-b, h-j), 2.0 mm (c), and 3.0 mm (d-g).

Source : MNHN, Paris

446

P. A. MCLAUGHLIN

Chelipeds subequal; left frequently longer, but not as robust. Right cheliped (Figs 2d, 33c, e) elongate, quite

slender. Dactyl slightly shorter to approximately as long as palm; cutting edge with blunt or sometimes

denticulate calcareous margin and 2 or 3 more prominent calcareous teeth; terminating in very small corneous

claw; dorsomesial margin not delimited; dorsal surface convex, elevated proximally into slender ridge, all surfaces

unarmed but with numerous fine setae. Palm 0.60 to 0.80 length of carpus; dorsomesial and dorsolateral margins

not delimited, dorsal surface convex, surfaces of palm and fixed finger unarmed, but with many fine setae; cutting

edge of fixed finger with several small and 2 or 3 somewhat larger calcareous teeth proximally and row of tiny

calcareous teeth distally, terminating in very small corneous claw. Carpus slightly longer than merus; dorsomesial

margin sometimes with row of small spines, or more often only 1 to 3 spines and several low tubercles; all

surfaces with very short transverse rows of moderate to long setae; ventrolateral distal margin frequently with small

spine. Merus with short transverse rows of setae on all surfaces; ventrolateral margin with 1 prominent spine at

ventrodistal angle, ventromesial margin with somewhat smaller spine at distal angle. Ischium with dorsal and

ventral rows of setae. Coxa with prominent spine on ventrolateral distal angle and smaller spine on ventromesial

distal angle.

Left cheliped (Figs 2e, 33d, f) slender, rarely shorter than right and frequently overreaching right by distal third

of chela; fingers in males usually obliquely curved ventrolaterally. Dactyl 1.25 to 1.50 length of palm; cutting

edge with row of very small corneous teeth, terminating in corneous claw; surfaces unarmed, but with numerous

short setae. Palm 0.60 to 0.75 length of carpus, dorsomesial and dorsolateral margins not delimited, all surfaces of

palm and fixed finger unarmed, but with numerous short to moderately long setae; cutting edge of fixed finger with

row of very small widely-spaced calcareous teeth interspersed with minute calcareous or corneous teeth, terminating

in tiny corneous claw. Carpus slightly longer than merus; dorsomesial margin with 1 or 2 distal spines and row of

spinulose protuberances, sometimes, at least partial row of small spines, dorsolateral margin not delimited; all

surfaces with numerous very short transverse rows of long setae; ventrolateral distal angle usually with small

spine. Merus with very short transverse rows of long setae on all surfaces; ventrolateral margin with I prominent

acute spine at distal angle; ventromesial margin with 1 smaller spine at distal angle. Ischium with long setae

dorsally and ventrally. Coxa with acute spine at each ventrodistal angle.

Second and third pereopods (Figs 2f-g) very long, overreaching outstretched chelipeds at least 0.25 length of

dactyls; generally similar from left to right. Dactyls long and slender, usually at least half again length of propodi;

in dorsal view, twisted in distal half; in lateral view, curved ventrally; terminating in slender corneous claws; dorsal

margins each with row of very long stiff setae, ventral, lateral and mesial faces with few scattered setae,

ventromesial margins with row of long, fine setae, at least in distal half, ventral margins each usually also with

1 to 5 tiny corneous spines. Propodi often with 1 corneous spine on ventrolateral distal margin, few low

protuberances and numerous setae dorsally; few scattered setae ventrally. Carpi each with 1 spine on dorsally

adjacent to dorsodistal angle and row of low protuberances on dorsal surface; second pereopods, and rarely third,

also with 1 additional small spine or spinulose protuberance on dorsal surface proximally. Meri and ischia

unarmed, but with tufts of setae dorsally and ventrally. Sternite of third pereopod with subquadrate anterior

lobe (Fig. 2h) armed marginally with 2 to 6 small spines and numerous long setae.

Telson (Figs 2i-j) with posterior lobes slightly asymmetrical, each usually with prominent spine at outer angle

and 2 to 4 additional spines on oblique terminal margin, occasionally margins nearly straight, median cleft

obsolete; lateral margins each with corneous plate.

Color (in preservative). — Calcified integument somewhat iridescent.

Habitat. — Variety of gastropod shells.

Distribution. — Kai and Tanimbar Islands; 356-552 m.

Etymology. — The species is named for Dr Julio Garci'a-GOmez, Miami-Dade Community College,

mm!, Florida, not only a friend and colleague, but the first carcinologist to document gill loss on the third

maxillipeds in pagurids.

Aff|N|TIES — Among Karubar species, Enneopagurus garciagomezi bears considerable resemblance to

species of Turleama nom. nov. However, in addition to the characters separating the two genera, E. garciagomezi

Source

PAGURIDAE FROM THE KARUBAR EXPEDITION

447

is characterized by setose, but unarmed chelae, and ambulatory legs that have little or no armature on the ventral

margins of the dactyls.

Remarks. — The sexual tube in males of this species (Figs 3a-b) tends to be moderately short to moderately

long, thick, curved outwardly and directed dorsally; the tip is spatulate and provided with a dense fringe of curved

setae. Males obviously infected with rhizocephalans still exhibit a well developed sexual tube.

Fig. 3. — Male secondary sexual characters of Enneopagurus and Turleania nom. nov. a-b, Enneopagurus garciagomezi

sp nov., paratype 6 (3.2 mm) from Karubar Stn CP 69. — c-d, Turleania multispina sp. nov., paratype 6 (1.9 mm)

from Stn DW 03: a, c, sternite and coxa of left fifth pereopods (ventral view); b, d, terminal portion of left sexual

tube. Scale equals 0.5 mm (c, d) and 1.0 mm (a, b).

Sexual dimorphism is seen in the left cheliped and in the setation of the sternite of the fifth pereopods.

Although the dactyl and fixed finger of the left chela are long, slender, and venlrally curved in both sexes, the

curvature is usually more pronounced in males where there is also a tendency for the chela to become laterally

twisted. Females have a considerably greater amount of setae on the sternite and coxae of the filth pereopods.

Genus DECAPHYLLUS de Saint Laurent, 1968

Decaphyllus de Saint Laurent, 1968a: 925; 1968b: 1100.

EMENDED Diagnosis. — Shield well calcified or with varying amount of decalcification particularly medially,

lateral margins strongly convex, well calcified. Posterior carapace weakly calcified or entirely membraneous. Eight

to 10 pairs of phyllobranchiate gills (no pleurobranchs; arthrobranchs of third maxilliped small, vestigial or

absent). Ocular acicles simple. Antennal peduncle with supernumerary segmentation. Maxillule with external

endopodal lobe obsolete or absent. Third maxilliped with crista dentata of ischium reduced, no accessory tooth;

merus with very strong dorsodistal spine. Sternite of third maxillipeds unarmed. Fourth pereopods simple, neither

semi- nor subchelate; without propodal rasp. Sternite of fifth pereopods entire, not divided into 2 lobes by median

groove.

Males with very long sexual tube developed on coxa of right fifth pereopod, directed from right to left across

ventral body surface and curved anteriorly. Coxa of left fifth pereopod with short or moderately short sexual tube

directed from left to right. Females with single gonopore on coxa of left third pereopod.

Abdomen with unpaired pleopods on somites 2 to 5 in both sexes, uniramous or very unequally biramous in

males. Telson without transverse suture; terminal margin entire or with minute median cleft; posterior lobes

strongly asymmetrical, each with 2 to 4 spines.

REMARKS. — In her revision of the genera Catapaguroides A. Milne Edwards & Bouvier, 1892 and

Cestopagurus Bouvier, 1897, DE SAINT LAURENT (1968a) established the genus Decaphyllus for three previously

unknown species. In that publication, de Saint Laurent presented only a brief generic diagnosis and a slightly

Source :

448

P. A. MCLAUGHLIN

more detailed diagnosis of its type species, D. spinicornis de Saint Laurent. In a subsequent paper she (DE SAINT

Laurent, 1968b) provided a detailed description of the general characters of Decaphyllus , gave full descriptions of

the three species, D. spinicornis, D. similis de Saint Laurent and D. junquai de Saint Laurent, and compared this

rather distinctive genus with its closest cohort, Catapaguroides. Decaphyllus spinicornis is known only from

Japan, whereas D. similis and I), junquai are reported from New Guinea and Indonesia.

Two additional species have been discovered during the course of this study, D. barunajaya sp. nov. and

D. maci sp. nov. Both of these species differ from DE SAINT Laurent's (1968a, b) taxa in having a very

prominent spine on the ventral margin of the first antennal segment. Decaphyllus barunajaya sp. nov. also differs

more notably in lacking arthrobranchs on the third maxillipeds. Gill loss in the Paguridea typically is manifest by

loss of pleurobranchs, from four being present in the Pylochelidae and some diogenids to none in some genera of

the Paguridae. Decaphyllus is only one of four pagurid genera where this total loss of pleurobranchs has been

observed. The tendency toward reduction and/or disappearance in the paired arthrobranchs of the third maxilliped,

without complete pleurobranch loss, has been reported in coenobitids and some pylochelids, but until this present

report was known in the Paguridae, only in Enneobranchus.

Although other characters can be enlisted to justify the establishment of the genera Enneobranchus,

Enneophyllus, and Enneopagurus, loss of arthrobranchs on the third maxilliped is the fundamental character. That

rationale might similarly suggest that a distinct genus be established for D. barunajaya sp. nov., were it not for the

overwhelming number of supplemental characters it shares with the other four species of Decaphyllus. These

include the broadly rounded rostrum; characteristic shape of the ocular acicles; lower ramus of antennular flagellum

with only three segments; distinctive profile of the coxal endite of the maxillule; reduced and irregular dentition of

the crista dentata; absence of an accessory tooth on the crista dentata; loss of the pleurobranch above the fourth

pereopod; uniform armature of the dorsal margins of the carpi of the second pereopods; very specialized develop¬

ment of the dactyl and propodus of the fourth pereopod; length and orientation of both the right and left sexual

tubes; male pleopod development; single female left gonopore; and unusual telson shape and armature. In view of

all these shared characters, clearly a simple emendation of Decaphyllus to accommodate D. barunajaya is the

appropriate action.

Key to the Indonesian species of Decaphyllus

1. Gill lamellae of third maxillipeds vestigial or absent; dactyl of right cheliped unarmed .

. — . D. barunajaya sp. nov.

— Gill lamellae of third maxillipeds normally developed; dactyl of right cheliped armed at

least with few spinules . 2

2. Dactyl of right cheliped with numerous spines or spinules on dorsal surface; telson with

nearly symmetrical posterior lobes, each with 2 small spines . D. similis*

— Dactyl of right cheliped with only 1 or 2 spines or spinules on dorsal surface; telson with

asymmetrical posterior lobes, terminal margins each with 2 to 4 spines . 3

3. Palm of right cheliped with prominent median longitudinal row of spines on dorsal

surface; telson with 3 or 4 spines on each terminal margin .

. . Decaphyllus maci sp. nov.

— Palm of right cheliped without prominent median longitudinal row of spines on dorsal

surface; telson with 2 spines on each terminal margin . Decaphyllus junquai*

Decaphyllus barunajaya sp. nov.

Figs 4a-l

Material EXAMINED. — Indonesia. Karubar. Kai Islands : stn DW 31, 05°40'S, 132°51'E, 288-289 m, 26. 10

1991: I 6 (1.8 mm) (MNHN-Pg 5255).

Tanimbar Islands: stn DW 50, 07°59'S, 133°02'E, 184-186 m, 29. 10. 1991: 1 ov. $ (1.9 mm) (MNHN-Pg 5256).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

449

TYPES. — The ovigerous female (MNHN-Pg 5256) from the Karubar station DW 50 is the holotype. The

other specimen is a paratype.

DESCRIPTION. — Shield (Fig. 4a) longer than broad; anterior margin between rostral region and lateral

projections very weakly concave; anterolateral margins sloping; posterior margin roundly truncate; surface with

median regions poorly calcified. Rostrum very broadly rounded or nearly obsolete. Lateral projections well

developed, with terminal spine or spinule.

Ocular peduncles approximately equaling length of shield, dorsal surface with row of tufts of fine setae; corneae

very slightly dilated. Ocular acicles drawn out distally into acute spine, mesial margin with several long setae;

separated basally by slightly more than half basal width of 1 acicle.

Antennular peduncles overreaching ocular peduncles by 0.20 to 0.50 length of ultimate segment. Basal segment

with prominent spine on dorsolateral margin medially. Penultimate and ultimate segments unarmed.

Antennal peduncle reaching nearly to base of cornea. Fifth and fourth segment with few scattered setae. Third

segment with acute spine on ventrodistal margin. Second segment with dorsolateral distal angle strongly produced,

terminating in bifid spine, dorsomesial distal angle with small spine. First segment with strong spine on

ventrodistal margin. Antennal acicle long, usually reaching to distal margin of ultimate peduncular segment or

slightly beyond; terminating in small spine; mesial surface with numerous long setae. Antennal flagellum with 2

to 4 short setae every 1 or 2 articles.

Crista dentata of third maxilliped with 5 or 6 irregularly-spaced and sized teeth; merus with very strong spine

on dorsodistal angle. Arthrobranchs of third maxillipeds vestigial or absent.

Chelipeds subequal in length, right only slightly longer, but appreciably stronger. Right cheliped (Fig. 4b)

with dactyl set at slightly oblique angle to palm; cutting edges of dactyl and fixed finger each with row of small

calcareous teeth. Dactyl slightly shorter than palm; unarmed, but with moderately dense long setae dorsally and

ventrally. Palm slightly shorter than carpus; dorsomesial margin with row of small spines not reaching to distal

margin, dorsal midline with row of small spines not extending onto fixed finger, dorsolateral margin row of small

spines and dorsal surface laterad of midline with numerous small spines, lateral face with faint transverse slriations,

some minutely spinulose and long setae; fixed finger unarmed but with moderately dense long setae particularly

dorsally and ventrally. Carpus slightly longer than merus; dorsomesial margin with row of spines, smallest

proximally and distally, dorsolateral surface with row of small spines, surfaces all with long setae. Merus with

1 acute spine on dorsodistal margin laterally; dorsal surface with transverse rows of long setae; ventromesial and

ventrolateral margins each with small spine distally, partially obscured by long setae, ventrolateral margin with

additional small spine in proximal half. Ischium with 1 anteriorly directed and 1 posteriorly directed spine and long

setae on ventral margin.

Left cheliped (Figs 4c-d) with longitudinal hiatus between dactyl and fixed finger; cutting edges each with row

of small calcareous teeth. Dactyl slightly to considerably longer than palm, unarmed but with long setae

particularly mesially and ventrally. Palm 0.50 to 0.70 length of carpus; dorsal surface with 2 rows of small spines,

dorsolateral margin with irregular single or double row of small spines, not extending on to fixed finger; latter

unarmed but with long moderately dense setae particularly laterally and ventrally; surfaces of palm also with

scattered long setae. Carpus with row of 4 to 6 spines on dorsomesial margin and 2 to 4 spines on dorsolateral

margin; all surfaces with scattered setae. Merus with transverse rows of setae on dorsal surface, 1 prominent spine

on dorsodistal margin mesially; ventrolateral margin with 2 rather widely-spaced spines, ventromesial margin with

3 spines; ventral surface with scattered long setae. Ischium with 1 anteriorly directed and 1 posteriorly directed

spine and scattered setae.

Ambulatory legs (Figs 4e-f) overreaching tip of right cheliped. Dactyls 1.50 to nearly twice length of propodi;

slightly curved ventrally; all surfaces unarmed, but with numerous setae, particularly longer and stronger on dorsal

margins. Propodi slightly longer than carpi, unarmed but with numerous moderately long setae, particularly

dorsally and ventrally. Carpi each with dorsodistal spine and 2 (second) or 1 (third) additional spines on dorsal

margin in proximal half. Meri each with 1 spine in proximal half of dorsal margin, also usually 1 spine

proximally in distal half, at least on second pereopods; ventral margins with numerous setae. Ischia unarmed but

with numerous long setae. Fourth pereopods with claw of dactyl (Figs 4g-h) almost entirely masked by tufts of

450

P. A. MCLAUGHLIN

dense, pectinate setae. Fifth pereopods semichelate. Anterior lobe of sternite (Fig. 4i) of third pereopods

subsemicircular, with long marginal setae.

Fig. 4. — Decaphyllus barunajaya sp. nov., a-c, e-i, k, holotype V (1.9 mm), from Karubar Stn DW 50;

d, j, 1, paratype <J (1.8 mm) from Stn DW 31: a, shield and cephalic appendages; b, right cheliped (dorsal view);

c, left cheliped (dorsal view); d, left cheliped (mesial view); e, right second pereopod (lateral view); f, left third

pereopod (lateral view); g, dactyl and propodus of right fourth pereopod (lateral view; setae omitted); h, dactyl and

propodus of left fourth pereopod (lateral view); i, anterior lobe of sternite of third pereopods; j, coxae and sternite of

fifth pereopods; k-1, telson. Scales equal 0.5 mm (g, h, i, k, 1) and 1.0 mm (a-f, j).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

451

Male with unequally biramous pleopod on abdominal somite 2, pleopods 3 to 5 uniramous. Right sexual tube

(Fig. 4j) reaching to level of coxa of left third pereopod; left sexual tube reaching to base of coxa of right fifth.

Telson (Figs 4k-l) with median cleft not apparent; terminal margin with prominently produced left exterior

angle separated by irregularly-spaced series of 3 or 4 spines from weakly developed right exterior angle; lateral

margins each with narrow chitinous plate.

COLOR (in preservative). — Shield and cephalic appendages with faint orange hue. Chelipeds and ambulatory

legs mottled orange with white distally on most segments. Calcified integument somewhat iridescent.

Habitat. — Unknown.

Distribution. — Tanimbar and Kai Islands, Indonesia; 184 to 289 m.

ETYMOLOGY. — The species is named for the vessel "Baruna Jaya from which the specimens were

collected.

AFFINITIES. — Decaphyllus barunajaya differs from the other species of the genus in the absence of

arthrobranchs on the third maxillipeds, in having an unarmed dactyl of the right cheliped, two proximal spines on

the dorsal surfaces of the carpi of the second pereopods, and shorter right sexual tube. The slightly dilated corneae,

unarmed ventral margin of the carpus of the right cheliped, and distinctly different telson armature are additional

characters that set D. barunajaya apart from D. spinicornis. Among Indonesian species, it resembles D. similis in

having moderately long ocular peduncles with slightly dilated corneae, but differs in having shorter antennular

peduncles, a bifid dorsolateral distal angle of the second antennal segment, and two rows of spines on the carpus of

the left cheliped. Decaphyllus barunajaya is also readily distinguished from D. junquai by its longer ocular, but

shorter antennular peduncles and armature of the right cheliped. Additional characters that serve to distinguish

Decaphyllus barunajaya from D. maci sp. nov. include the anterior lobe of the sternite of the third pereopods,

which is subsemicircular in the former species, but subtriangular in the latter. The carpus of the left cheliped has

only two to four spines on the dorsolateral margin in D. barunajaya, but five or six in D. maci, and the left chela

exhibits a pronounced hiatus between the dactyl and fixed finger in the former species that is lacking in the latter.

Decaphyllus maci sp. nov.

Figs 5a-i

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 30, 05°39'S, 132°56'E, 118-111 m,

26.10.1991: 1 <S (1.8 mm) (MNHN-Pg 5257), 1 ov. 9 (1.8 mm) (MNHN-Pg 5258).

Types. — The ovigerous female (MNHN-Pg 5258) from Karubar station DW 30 is the holotype. The male

is a paratype.

DESCRIPTION. — Shield (Fig. 5a) longer than broad; anterior margin between rostral region and lateral

projections very weakly concave; anterolateral margins sloping; posterior margin roundly truncate; surface with

median regions poorly calcified proximally and distally. Rostrum very broadly rounded or nearly obsolete. Lateral

projections well developed, with terminal spine.

Ocular peduncles approximately equaling length of shield, dorsal surface with row of tufts of long setae;

corneae not dilated. Ocular acicles each drawn out distally into long, acute spine, mesial margin with several long

setae; separated basally by approximately 0.75 basal width of an acicle.

Antennular peduncles overreaching ocular peduncles by 0.50 to 0.80 length of ultimate segment. Basal segment

with prominent spine on dorsolateral margin medially. Penultimate and ultimate segments unarmed.

Antennal peduncle reaching to distal half of ocular peduncle but not to base of cornea. Fifth and fourth segment

with few scattered setae. Third segment with acute spine on ventrodistal margin. Second segment with dorsolateral

distal angle strongly produced, terminating in unequally bifid spine; dorsomesial distal angle with prominent spine.

First segment with strong spine on ventrodistal margin. Antennal acicle longer than peduncle, usually reaching to

452

p. a. McLaughlin

base of cornea; terminating in small spine; mesial surface with numerous long setae. Antennal flagellum with 2 to

4 short and 1 or 2 somewhat longer setae every 1 or 2 articles.

Crista dentata of ischium of third maxilliped with 3 to 5 irregularly-spaced and sized teeth; merus with very

strong spine on dorsodistal angle. Arthrobranchs of third maxillipeds normally developed.

Fig. 5. — Decaphyllus maci sp. nov„ a-c, e-h, holotype 2 (1.8 mm) from Karubar Stn DW 30; d, i, paratype 6

(1.8 mm) from Stn DW 30: a, shield and cephalic appendages; b, right cheliped (dorsal view); c, left cheliped (dorsal

view); d. left cheliped (mesial view); e. right second pereopod (lateral view); f, left third pereopod (lateral view);

g. anterior lobe of sternite of third pereopods; h-i, telson. Scale equals 0.5 mm (g-i) and 1.0 mm (a-f).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

453

Chelipeds subequal in length, right only slightly longer, but appreciably stronger. Right cheliped (Fig. 5b)

with dactyl generally straight; cutting edges of dactyl and fixed finger each with row of small calcareous teeth.

Dactyl approximately 0.80 length of palm; with 1 spine on dorsal surface proximally and relatively dense long

setae, particularly laterally and ventrally. Palm approximately equal to length of carpus; dorsomesial margin with

row of spines not reaching to distal angle, dorsal midline with row of small spines not extending full length of

palm, dorsolateral margin with row of moderately widely-spaced small spines, few additional spinules on dorsal

surface laterad of midline; lateral and mesial faces with scattered long setae; fixed finger unarmed but with relatively

dense long setae particularly laterally and ventrally. Carpus slightly longer than merus; dorsomesial margin with

row of moderately small spines, largest proximally, dorsolateral margin with row of slightly larger spines; surfaces

all with scattered long setae. Merus with 1 acute spine on dorsodistal margin laterally; dorsal surface with

transverse rows of long setae; ventromesial and ventrolateral margins each with small distal and proximal spine and

scattered long setae, ventral surface with median spinule. Ischium with 1 anteriorly directed and 1 posteriorly

directed spine and long setae on ventromesial margin, ventrolateral distal angle with minute spinule.

Left cheliped (Figs 5c-d) without longitudinal hiatus between dactyl and fixed finger but tips crossing; cutting

edge of dactyl with row of sharp spine-like teeth, cutting edge of fixed finger with row of small, blunt calcareous

teeth. Dactyl approximately equal to length of palm, unarmed but with long setae particularly mesially and

ventrally. Palm 0.60 to 0.80 length of palm; dorsal surface with row of small spines in midline not extending

onto fixed finger and 1 or 2 small spines dorsolaterally, dorsolateral margin with row of small spines, extending

only slightly, if at all, onto fixed finger; dorsomesial margin with 2 or 3 acute spines; fixed finger unarmed but

with long moderately dense setae particularly laterally and ventrally; surfaces of palm also with scattered long

setae. Carpus with row of 4 or 5 spines on dorsomesial margin and 5 or 6 spines on dorsolateral margin;

ventrolateral distal angle with small acute spine; all surfaces with scattered setae. Merus with transverse rows of

setae on dorsal surface, 1 acute spine on dorsodistal margin; ventrolateral margin with 2 or 3 rather widely-spaced

spines, ventromesial margin with 3 spines, ventral surface with I or 2 acute spines and scattered long setae.

Ischium with 1 anteriorly directed and 1 posteriorly directed spine and scattered setae on ventromesial margin.

Ambulatory legs (Figs 5e-f) overreaching tip of right cheliped. Dactyls 1.50 to nearly twice length of propodi;

slightly curved ventrally; all surfaces unarmed but with numerous setae, particularly longer and stronger on dorsal

margins. Propodi 0.25 to 0.40 longer than carpi, unarmed but with numerous moderately long setae, particularly

dorsally and ventrally. Carpi each with dorsodistal spine and 1 additional spine on dorsal margin in proximal half.

Meri each with 1 spine on dorsal surface at mid-length and 1 additional spine in proximal half; ventral margins

with numerous setae. Ischia unarmed but with numerous long setae. Fourth pereopods with claw of dactyl almost

entirely masked by tufts of dense, pectinate setae. Fifth pereopods semichelate. Anterior lobe of sternite (Fig. 5g)

of third pereopods bluntly subtriangular, with long submarginal setae.

Male with uniramous pleopods on abdominal somites 2, 4 and 5, pleopod of somite 3 unequally biramous.

Right sexual tube reaching to level of coxa of left cheliped; left sexual tube not quite reaching to base of coxa of

right fifth.

Telson (Figs 5h-i) sometimes with very small median cleft; terminal margin with prominently produced left

exterior angle separated by irregularly-spaced series of 5 or 6 spines from more weakly developed right exterior

angle; lateral margins each with narrow chitinous plate.

COLOR (in preservative). — Shield and cephalic appendages with faint orange hue. Chelipeds and ambulatory

legs also with faint orange hue, appearing as indistinct bands on most segments. Calcified integument somewhat

iridescent.

Habitat. — Unknown.

Distribution. — Kai Islands, Indonesia; 111-118 m.

Etymology. — This species is named for the late E.J. "Mac" McGEORGE, whose photographs have enhanced

many pagurid reports.

AFFINITIES. — As previously indicated, all known species of Decaphyllus share a considerable number of

morphological characters. In proportions and armature of the cephalic appendages, and armature of the right

454

P. A. MCLAUGHLIN

cheliped, D. maci most closely resembles D. spinicornis. Both species have relatively long ocular peduncles with

corneae showing no dilation; bifid termination of the dorsolateral distal angles of the second segments of the

antennal peduncles; and antennular peduncles that overreach the ocular peduncles by 0.50 to 0.80 length of the

ultimate segment. Similarly, the right chelipeds of both species have a single small spine on the dorsoproximal

surface of the dactyl and both dorsomesial and median longitudinal rows of spines. However, D. maci also has a

row of small spines on the clearly delimited dorsolateral margin, whereas D. spinicornis is described as having the

dorsolateral margin only weakly delimited and with few scattered spinules. Although both species also have two

rows of spines on the dorsal surface of the carpus, the ventrolateral margin is armed with a few spines in

D. spinicornis, but unarmed in D. maci. Differences are also found in the development of the telson. In D. maci a

small median cleft is sometimes apparent, but even when obscured, there is a distinct separation of right and left

lobes, each terminal margin armed with two or occasionally three small spines in addition to the spine of the

external angle. By contrast, D. spinicornis has only two spines on the terminal margin between the spinose

external angles.

From the remaining species of Decaphyllus , all occurring in Indonesian waters, D. maci is best distinguished

from D. similis by its shorter antennular peduncles, dactyl of the right cheliped having only a single spine on the

dorsal surface, and by having two rows of spines on the dorsal surface of the carpus of the left cheliped. From

D. junquai, this new species is also readily separated by the relative shortness of its antennular peduncles, less

strongly armed right chela, and more strongly armed telson. Like D. spinicornis, the telsons of both D. similis and

D. junquai have only two spines on the terminal margin between the spinose external angles. Decaphyllus maci

bears a superficial resemblance to D. barunajaya sp. nov. in having relatively long ocular and short antennular

peduncles; bifid termination of the dorsolateral distal angles of the second segments of the antennal peduncles;

strong spine on the ventrodistal margin of the first segment of the antennal peduncle; and weak median

calcification of the shield. However, the two species are easily distinguished by the armature of the chelipeds and

the carpi of the second pereopods.

Genus CATAPAGUROIDES A. Milne Edwards & Bouvier, 1892

Catapaguroides A. Milne Edwards & Bouvier, 1892: 211 (in part). — Bouvier, 1922: 26 (in part). — de Saint Laurent,

1968a: 927.

DIAGNOSIS. — Ten pairs of phyllobranchiate gills (no pleurobranch above arthrobranchs of fourth pereopod).

Antennal peduncle with supernumerary segmentation. Ischium of third maxilliped with crista dentata more or less

reduced, without accessory tooth. Chelipeds unequal, right appreciably stronger. Carpi of ambulatory legs with

dorsodistal spine. Fourth pereopods semichelate; propodal rasp with single row of corneous scales.

Males with sexual tube developed on coxa of right fifth pereopod, directed from right to left under thorax and

recurved anteriorly. Short tube developed on coxa of left fifth pereopod and concealed between 2 thick tufts of setae

on sternite; unpaired biramous left third to fifth pleopods. Females with single gonopore on coxa of left third

pereopod. No paired pleopods and 4 unpaired, left second to fifth biramous pleopods.

Telson with transverse suture only weakly delineated; posterior lobes not markedly asymmetrical.

Remarks. — Prior to the revisionary work of DE Saint Laurent-Dechance (1966b) and de Saint

LAURENT (1968a), the position of the male sexual tube was often the only criterion used in generic assignments.

Use of the collective possession of a right sexual tube had resulted a heterogeneous conglomerate of taxa assigned

to the genera Catapaguroides, Cestopagurus, and Catapagurus that otherwise shared few characters of phylogenetic

significance. With her redefinition of Catapaguroides (DE Saint LAURENT, 1968a), the genus was left with only

three of its formerly assigned taxa, all with Atlantic distributions. She reassigned three Indo-Pacific species, two

previously included in Cestopagurus and one in Catapagurus to Catapaguroides and described six new species. Five

of these, C. inermis de Saint Laurent, 1968a; C. cristimanus de Saint Laurent, 1968a; C. mortenseni

de Saint Laurent, 1968a; C. melini de Saint Laurent, 1968a; and C. spinulimanus de Saint Laurent. 1968a, occur

in Indonesian waters, although none were collected during the Karubar expedition.

PAGURIDAE FROM THE KARUBAR EXPEDITION

455

Key to the Indonesian species of Catapaguroides

1. Right cheliped with subcircular chela and subtriangular carpus (dorsal view) . 2

— Right cheliped with subrectangular or subovate chela and subrectangular carpus (dorsal

view) . 3

2. Right cheliped with row of spines on dorsomesial margin of palm, carpus with row of

spines in dorsal midline; palm of left chela with 2 spinulose tubercles .

. C. cristimanus*

— Right cheliped with 1 prominent spine on dorsomesial margin of palm; carpus few spines

on dorsomesial margin and 2 spines in dorsal midline distally; palm of left chela unarmed.

. C. karubar sp. nov.

3. Palm of right cheliped armed with numerous spines, at least on dorsomesial margin . 4

— Palm of right cheliped unarmed or with only 1 small tubercle at dorsomesial distal angle ..

. 6

4. Dorsal surface of palm of right cheliped with 3 irregular rows of fine spines or spinules ...

. C. spinulimanus*

— Dorsal surface right cheliped unarmed or with only I spine or tubercle proximally,

dorsomesial margin with row of spines . 5

5. Carpus of right cheliped distinctly shorter than palm; dactyl with convex dorsal surface ....

. C. melini*

— Carpus of right cheliped distinctly longer than palm; dactyl with median ridge or crest in

distal half . C. declivis sp. nov.

6. Antennular peduncles overreaching distal margins of corneae by nearly half length of

penultimate segment; ocular peduncles moderately slender . C. inermis*

— Antennular peduncles overreaching distal margins of corneae by length of ultimate

segment; ocular peduncles moderately short and stout . C. mortenseni*

Catapaguroides declivis sp. nov.

Figs 6a-j, 33g

MATERIAL EXAMINED. — Indonesia. Karubar. Tanimbar Islands: sin DW 49, 08°00'S, I32°59'E. 210-206 m,

29.10.1991: 1 6 (1.5 mm) (MNHN-Pg 5259); 1 ov. $ (1.1 mm) (MNHN-Pg 5260); 1 6 (0.9 mm) (POLIP1). —

Stn DW 50, 07°59'S, 133°02'E, 184-186 m, 29.10.1991: 1 ov. 9 (1.2 mm) (USNM 276009).

Types. — The male (MNHN-Pg 5259) from the Karubar station DW 49 is the holotype. The other

specimens are paratypes.

DESCRIPTION. — Shield (Fig. 6a) longer than broad; anterior margin between rostrum and lateral projections

weakly concave; anterolateral margins sloping, posterior margin truncate, but with shallow median concavity;

dorsal surface with few setae. Rostrum broadly rounded. Lateral projections triangular, produced little if at all

beyond level of rostrum; with small marginal spinule.

Ocular peduncles moderately long and moderately stout, 0.70 to 0.80 shield length; dorsomesial margins each

with row of sparse tufts of long setae; corneae slightly dilated. Ocular acicles narrowly triangular, with few

marginal setae and submarginal distal spine.

Antennular peduncles overreaching ocular peduncles by at least full length of ultimate peduncular segment.

Basal segment elongate, with statocyst lobe produced laterally and with acute spine on distolateral margin, distal

margin with several long setae. Penultimate segment with few scattered short setae. Ultimate segment with 1 or

2 very long, distally plumose setae.

Source :

456

P. A. MCLAUGHLIN

Fig. 6. — Catapaguroides declivis sp. nov., a-b, e-j, paratype 6 (1.1 mm) from Karubar Stn DW 49; c-d, holotype <J

(1-5 mm) from Sin DW 49: a, shield and cephalic appendages; b, right cheliped; c, carpus and chela of right cheiiped;

d carpus and chela of left cheliped; e, right second pereopod (lateral view); f. left third pereopod (lateral view);

g, dactyl, propodus and carpus of left fourth pereopod (lateral view); h, anterior lobe of sternite of third pereopods;

i, coxae and sternite of fifth pereopods; j, telson. Scales equal 0.5 mm (g-j) and 1.0 mm (a-f).

Antennal peduncle moderately short, reaching to distal margin of cornea or slightly beyond. Fifth segment with

tew scattered setae and 2 or 3 longer at distal margin. Fourth segment glabrous. Third segment with few setae and

acute spine at ventrodistal margin. Second segment with dorsolateral distal angle produced, terminating in strong

simple or bifid spine; dorsomesial distal angle with small spine. First segment with small spine on ventrolateral

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

457

margin. Antennal acicle slender, arcuate, reaching approximately to middle of ultimate peduncular segment;

terminating in small spine and with several marginal long setae. Antennal flagella (missing in holotype)

overreaching right cheliped; with 1 or 2 moderately long and also 1 or 2 very short setae every 1 or 2 articles

proximally, only very short setae distally.

Right cheliped (Figs 6b-c, 33g) with dactyl articulating obliquely with palm; propodal-carpal articulation

rotated clockwise 15° to 45° from perpendicular. Dactyl 0.70 to approximately equaling length of palm; cutting

edge with 2 prominent calcareous teeth; terminating in small calcareous claw and slightly overlapped by fixed

finger; dorsal surface with distinct but very faintly granular or smooth crest in proximal half, dorsomesial margin

drawn out into very weak subacute ridge, armed proximally with 2 to several very small spinulose tubercles;

mesial and ventral surfaces with few scattered moderately long setae. Palm 0.65 to 0.80 length of carpus;

dorsomesial margin with single or irregularly double row of 5 to 1 1 small to moderately strong acute or subacute

spines, 1 or 2 well-developed tubercles at dorsoproximal margin in mesial half, dorsal surface of palm and fixed

finger convex, dorsolateral margin not delimited; mesial face and ventral surface of palm and fixed finger with

scattered setae; cutting edge of fixed finger with 2 prominent calcareous teeth, terminating in corneous or

calcareous claw. Carpus 1.25 to 1.50 times longer than merus; dorsomesial distal angle with acute spine, usually

also with row of 3 to several small spinules on dorsomesial margin, long setae along entire dorsomesial margin

and on mesial face; dorsal surface laterad of midline with row of 5 to 18 spines, dorsolateral surface strongly

sloping ventrally; lateral and ventral surfaces with scattered long setae, most numerous ventrally, ventrolateral

distal angle with acute spine. Merus subtriangular; dorsodistal margin with small spine or spinule; dorsal surface

somewhat flattened but with double row of long setae sometimes arising from low protuberances; ventromesial

margin with 2 to 4 prominent spines distally and row of long setae; ventrolateral margin with 1 or 2 prominent

spines near distal angle and row of long setae. Ischium with long setae dorsally and ventrally.

Left cheliped (Fig. 6d) (missing in male paratype) not reaching to proximal margin of dactyl of right; with

propodal-carpal articulation twisted 30° to 45° counterclockwise from perpendicular; dactyl and fixed finger curved

ventrally and with slender hiatus. Dactyl 1.25 to 1.50 longer than palm, unarmed but with scattered setae; cutting

edge with row of small corneous teeth. Palm approximately 0.60 length of carpus; dorsomesial margin with 2 or

3 spines and sparse tufts of long setae, dorsal surface of palm unarmed or with 1 minute spinule proximally in

midline, fixed finger unarmed; both with few long setae, particularly on dorsomesial margin of palm; dorsolateral

margin not delimited. Carpus approximately 1.25 length of merus; dorsomesial distal angle with strong slender

spine, margin with 2 or 3 spines and numerous long setae; dorsolateral distal angle with 1 or 2 tiny spinules,

dorsolateral margin with 3 to 6 spines; ventrolateral distal angle with prominent spine; mesial and ventral surfaces

with few scattered long setae. Merus with tufts of setae on dorsal margin; ventrolateral margin with 2 slender

spines in distal half; ventromesial margin with 1 or 2 spines in distal half. Ischium with small spine at

ventromesial distal angle. Coxa with acute spine at distolateral angle.

Ambulatory legs (Figs 6e-f) elongate, overreaching right cheliped by nearly entire length of dactyls;

terminating in long, slender claws. Dactyls of right pereopods slightly longer than propodi; dactyls of left second

(regenerating in holotype) from 1.25 to 1.50 and dactyls of left third from 1.50 to nearly twice length of propodi;

slightly curved and twisted distally, surfaces with few long and short setae most numerous dorsally; ventral

margins with 2 or 3 short, fine corneous spinules. Propodi of second pereopods each with single long and 1 or

2 pairs of short or moderately short stiff articulated bristles at ventrodistal margin (males) or unarmed (females);

propodi of third with ventrodistal margins unarmed or with 1 short corneous spinule; dorsal and ventral surfaces all

with sparse tufts of setae most numerous on third. Carpi each with spinule at dorsodistal angle and few setae

dorsally. Meri and ischia each with few setae. Fourth pereopods with 3 or 4 denticles on ventral margin of dactyl;

propodal rasp consisting of 6 to 8 corneous scales (Fig. 6g). Fifth pereopod semichelate. Sternite of third

pereopods with roundly rectangular anterior lobe (Fig. 6h).

Male with long sexual tube developed on coxa of right fifth pereopod (Fig. 6i), directed from right to left and

reaching beyond coxa of left fifth pereopod; coxa of left with short sexual tube directed from left to right and

partially obscured by setae arising from sternal surface.

Telson (Fig. 6j) with asymmetrical posterior lobes separated by deep median cleft; terminal margins each with

3 or 4 very strong and often 1 smaller acute spines; lateral margins each delimited by chitinous plate.

Source :

458

P. A. MCLAUGHLIN

Color. — Unknown.

Habitat. — Unknown.

Distribution. — At present known only from the Tanimbar Islands, Indonesia; 206-209 m.

ETYMOLOGY. — From the Latin declivis meaning downhill or sloping, referring to the strongly sloping

dorsolateral face of the carpus of the right chcliped.

Affinities. — Catapaguroides declivis bears some resemblance to both C. melini and C. cristimanus. From

C. melini, C. declivis differs in having shorter ocular peduncles with more dilated corneae; shorter antennal

peduncles; distinctly different armature of the right cheliped; shorter male right sexual tube; and more strongly

armed telson. Catapaguroides declivis is readily distinguished from C. cristimanus by the longer, more slender

shield of the former species and most specifically, by its narrower and differently armed right chela, which has one

or two dorsoproximal tubercles on the palm and lacks sharply crested dorsomesial and dorsolateral margins.

Catapaguroides declivis is immediately distinguished from the second species of Catapaguroides found during the

Karubar expedition by shape and armature of the carpus of the right cheliped.

REMARKS. — The spines on the dorsomesial margin of the palm of the right chela are stronger, but fewer in

number in the very small, but mature, female (0.92 mm) than in either of the larger males. In the holotype, the

right chela is more strongly twisted, and the row of spines on the dorsal surface of the carpus is composed of

numerous, but irregular, more serrate appearing spines. The characteristic proximomedial tuberculation of the right

chela is a single tubercle in both male and female paratypes, and double in the holotype.

Catapaguroides karubar sp. nov.

Figs 7a-h

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 18, 05°17’49"S, 133°00’51"E, 205-212 m

24.10.1991: 1 $ (1.1 mm) (MNHN-Pg 5261).

TYPES. — The single specimen is the holotype.

DESCRIPTION. — Shield (Fig. 7a) approximately as long as broad; anterior margin between rostrum and lateral

projections weakly concave; anterolateral margins sloping, posterior margin truncate. Rostrum broadly rounded.

Lateral projections triangular, produced only slightly beyond level of rostrum, with small marginal spine.

Ocular peduncles moderately long and moderately stout, approximately 0.80 shield length; corneae

approximately 0.30 length of peduncle, not dilated. Ocular acicles triangular, with small distal spine; separated

basally by basal width of an acicle.

Antennular peduncles when extended overreaching ocular peduncles by slightly less than length of ultimate

peduncular segment. Basal segment elongate, with statocyst lobe produced laterally and with acute spine on

dorsolateral margin. Penultimate segment with few scattered short setae. Ultimate segment with 2 or 3 long,

distally plumose setae.

Antennal peduncle moderately short, reaching to proximal margin of cornea. Fifth segment with 1 or 2 long

setae at distal margin. Fourth segment glabrous. Third segment with very prominent acute spine at ventrodistal

margin. Second segment with dorsolateral distal angle strongly produced as elongate spine, with accessory spinule

on mesial margin; dorsomesial distal angle with prominent spine. First segment with acute spine on ventrolateral

margin. Antennal acicle slender, arcuate, reaching distal margin of ultimate peduncular segment; terminating in

small spine, and with few setae distally. Antennal flagella short; each article with 2 or 3 moderately short and I or

2 somewhat longer setae. Crista dentata with 5 or 6 irregularly-spaced and sized teeth on ischium; basis unarmed.

Right chcliped (Fig. 7b) with operculatc chela. Dactyl articulating somewhat obliquely with palm; approximately

1.30 times length of palm; cutting edge with 3 widely separated calcareous teeth; terminating in

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

459

Fig. 7. — Catapaguroides karubar sp. nov.. holotype 2 (1.1 mm) from Karubar Stn DW 18: a. shield and cephalic

appendages; b, right cheliped; c, left cheliped; d, right second pereopod (lateral view); e, dactyl of right second

pereopod (mesial view); f. left third pereopod (lateral view); g. anterior lobe of sternite of third pereopods; h. telson.

Scale equals 0.5 mm (g, h) and 1.0 mm (a-f).

small corneous claw, slightly overlapped by fixed finger; dorsal surface convex, with weak ridge in proximal half,

dorsomesial margin distinctly lamellar; ventral surface with distinct median ridge. Palm approximately 0.65 length

of carpus; dorsomesial margin not delimited, but with 1 well-developed spine in distal third, dorsal surface convex.

460

P. A. MCLAUGHLIN

with 2 small spines adjacent to proximal margin; fixed finger convex, but with distinct depression laterally,

dorsolateral margin lamellar; ventral surface of palm and fixed finger with scattered short setae; cutting edge of

fixed finger with 3 widely-spaced calcareous teeth, terminating in corneous claw. Carpus only slightly longer than

mcrus; dorsomesial distal angle with prominent acute spine and 2 small spinules near rounded dorsomesial margin

proximally; dorsal midline with 2 spines in distal half, dorsolateral surface strongly sloping ventrally, dorsolateral

margin not delimited; lateral and ventral surfaces with few fine setae. Merus subtriangular; dorsal surface somewhat

rounded, distal margin with strong spine mesially; ventromesial margin with 1 prominent spine distally;

ventrolateral margin with 1 prominent and 1 smaller spine near distal angle. Ischium unarmed.

Left cheliped (Fig. 7c) shorter and appreciably less robust than right. Dactyl longer than palm, unarmed but

with few setae of moderate length; cutting edge with row of corneous teeth, terminating in small corneous claw.

Propodal-carpal articulation rotated counterclockwise approximately 75°; palm and fixed finger unarmed and with

only few scattered setae; cutting edge of fixed finger with row of very small calcareous teeth, terminating in small

corneous claw. Carpus approximately 1.40 times longer than palm but shorter than merus; dorsomesial margin

with row of 3 widely-spaced spines, 1 spine distally on strongly sloping dorsolateral margin. Merus roundly

triangular; ventromesial margin with strong spine distally; ventrolateral margin with 2 spines in distal half.

Ischium unarmed, but with few fine setae on ventromesial margin.

Ambulatory legs (Figs 7d-f) elongate, overreaching right cheliped; terminating in long, slender claws. Dactyls

1.25 to 1.35 times longer than propodi, slightly curved and twisted distally, dorsal surfaces each with row of stiff

setae; ventromesial margins each with row of 5 or 6 long corneous spines. Propodi each with row of widely-

spaced, sparse tufts of setae on dorsal margins, ventrodistal angles with 2 or 3 stiff setae. Carpi each with spinule

at dorsodistal angle and few setae dorsally. Meri and ischia each with few setae. Fourth pereopods with 4 denticles

on ventral margin of dactyl; propodal rasp consisting of 5 corneous scales. Fifth pereopods semichelate. Sternite of

third pereopods with roundly rectangular anterior lobe (Fig. 7g).

Male unknown.

Uropods strongly asymmetrical. Telson (Fig. 7h) with weakly delimited transverse suture; slightly asymmetri¬

cal posterior lobes separated by deep median cleft; terminal margins strongly oblique, each with 1 small acute

spine at midlength and spinose outer angle; lateral margins each delimited by chitinous plate.

Color. — Unknown.

Habitat. — Unknown.

Distribution. — At present known only from the type locality in the Kai Islands, Indonesia; 205-212 m.

Etymology. — The acronym for the 1991 French-Indonesian expedition to Indonesia.

Affinities. — The operculate shape of the right cheliped of C. karubar suggests a close relationship with

C. cristimanus, another species known only from the Kai Islands region of Indonesia; however the quite different

armature of both chelipeds will immediately separate to two species, as will the armature of the telsons.

Catapaguroides cristimanus has been described as having a longitudinal row of spine on the dorsal midline of the

palm ol the right cheliped and two tubercles on the dorsomesial margin of the palm of the left. The right chela of

C. karubar has only two proximal spines and one mesial marginal spine on the dorsal surface; the left chela is

entirely unarmed. The telson of C. cristimanus is reported to have three long spines on each terminal telsonal

margin, whereas, C. karubar has a single small spine on each margin. As previously noted, the shape and armature

of the carpi of the right chelipeds will immediately separate C. karubar from C. declivis. In the former, the carpus

is subtriangular and armed with very few spines, whereas in the latter species, the carpus is subrectangular and

armed with a complete row of spines on the dorsolateral margin.

Remarks. — Although this species is known from a single female, its assignment to Catapaguroides is

reasonably certain. It shares with other members of the genus, 10 pairs of phyllobranchiate gills, reduced crista

dentata lacking an accessory tooth, and single female left gonopore.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

461

Genus SOLITARIOPAGURUS Tiirkay, 1986

Solitariopagurus Tiirkay, 1986: 139.

Diagnosis. — Ten pairs of phyllobranchiate gills. Anterior carapace vaulted and strongly calcified; lateral

margins of shield developed into blunt or spiniform projections. Rostrum and lateral projections widely separated.

Antennal peduncle with supernumerary segmentation. Maxillule with external lobe of endopod obsolete or absent.

Ischium of third maxilliped with well developed crista dentata and one accessory tooth. Stemite of third pereopods

with moderately slender transverse anterior lobe and nearly perpendicular posterior plate. Stemite of fifth pereopods

widely separated from preceding sternal plates. Fourth and fifth pereopods subchelate.

Males with stout, moderately long, unequal sexual tubes on coxae of both fifth pereopods, right approximately

twice length of left; each with long setae terminally; no paired or unpaired pleopods. Female with single gonopore

opening posteriorly on coxa of left third pereopod; no paired pleopods; unpaired left uniramous pleopods on

abdominal somites 2 to 4.

Abdomen reduced; tergal plate of first somite chitinous or very faintly calcified; tergal plate of second only

weakly delineated; tergal plates of somites 3 to 5 clearly defined, chitinous or very weakly calcified; tergite of sixth

somite weakly calcified. Uropods symmetrical; protopods each with very prominent, posteriorly directed spine.

Telson with transverse suture at least indicated; terminal margin entire.

REMARKS. — Solitariopagurus was initially thought to be another of the rather exceptional genera of deep

water pagurids. The type species, S. profundus Tiirkay, 1986, was described from depths between 1300 and

2000 meters in the Red Sea. However, both the recently described, S. triprobolus Poupin & McLaughlin, 1996,

and S. tuerkayi sp. nov. have been collected in relatively shallow depths between 1 1 1 and 400 meters.

Solitariopagurus tuerkayi sp. nov.

Figs 8a-n, 34a

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 18, 05°18'S, 133°01'E, 205-212 m,

24.10.1991: 1 6 (2.9 mm) (MNHN-Pg 5262); 2 $ (2.9, 3.0 mm) (MNHN-Pg 5263). — Stn DW 30, 05°39'S, 132°56'E.

118-111 m, 26.10.1991: 1 6 (3.0 mm) (USNM 276002).

Tanimbar Islands: stn DW 50, 07°59'S, 133°02'E, 184-186 m, 29.10.1991: 1 <3 (3.3 mm) (POLIPI).

Types. — The male (2.9 mm) (MNHN-Pg 5262) from the Karubar station DW 18 is the holotype. The

other specimens are paratypes.

Description. — Anterior carapace (Figs 8a, 34a) strongly vaulted; shield length consistently shorter than

breadth, total carapace length usually slightly longer; anterior margin between rostrum and lateral projections

straight or very slightly concave; lateral margins each with relatively short, but prominent spine at anterolateral

angle, broad and usually weakly bilobed subacute spine at midlength, and strong broad or moderately slender spine

adjacent to cervical groove; dorsal surface strongly calcified, with transverse row of 4 prominent, narrow to broad

tubercles proximal to anterior margin, sometimes with few scattered very small spinulose tubercles laterally;

posterolateral region weakly delineated and usually somewhat globular; posterior margin broadly rounded. Linea

transversalis present as well calcified, broad rod. Rostrum elongate, usually reaching to proximal half of ocular

peduncles; broad, slightly to strongly upturned, triangular, with elevated lateral margins and median keel; usually

terminating bluntly. Lateral projections triangular; elongate, as long or slightly longer than rostrum. Posterior

carapace with anterolateral regions developed distolaterally as pair of unarmed calcified plates, sometimes projecting

laterally as short blunt or spinose process; posterolateral regions distomesially and posteromedian plate distally

with very weakly calcified transverse rod-like area; remainder of posterior carapace membraneous (damaged in

illustrated male paratype).

Source :

462

p.a. McLaughlin

Fig-8-- Soliumopagurus tuerkayi sp. nov„ a-k, m-n, paratype 6 (3.3 mm) from Karubar Sin DW 50; 1. holotype 6

(-.9 mm) from Sin DW 18; a. shield and cephalic appendages; b, thoracic stemites; c, right cheliped (dorsal view)-

d, carpus and merus of right cheliped (mesial view); e, left cheliped (dorsal view); f. carpus and merus of left cheliped

(mesial view); g, right second pereopod (lateral view); h, merus of second right pereopod (mesial view); i left third

pereopo (lateral view); j. dactyl, propodus and carpus of left fourth pereopod (lateral view); k, propodus and dactyl of

right fifth pereopod (lateral view); 1-m. stemite and coxae of fifth pereopods; n, abdomen, uropods and telson (dorsal

view). Scales equal 0.5 mm (1, m), 1.0 mm (b, j, k), 2.0 mm (a, c-i, n).

Ocular peduncles short, slightly less than half length of shield, with prominent ventral swelling proximally and

slight submedian constriction; cornea somewhat dilated, diameter 0.25 to 0.35 length of peduncle. Ocular acicles

small, acutely triangular, obscured from dorsal view by base of rostrum; separated basally by more than basal

width of one acicle.

Source MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

463

Antennular peduncles elongate, overreaching ocular peduncles by full length of ultimate segment or slightly

more. Ultimate segment with 2 or 3 long plumose setae on dorsodistal margin. Penultimate segment with few

scattered short setae. Basal segment with small spinule on distolateral margin. Epistomial plate well calcified,

broad.

Antennal peduncles overreaching ocular peduncles by 0.33 to 0.50 length of ultimate segment, but appreciably

shorter than antennular peduncles. Fifth and fourth segments with few scattered, very short setae. Third segment

unarmed. Second segment with dorsolateral distal angle produced, terminating subacutely; dorsomesial distal angle

rounded. First segment with row of small or very small tubercles on ventrolateral margin. Antennal acicle

moderately short, not reaching distal margin of fourth peduncular segment, subtriangular and terminating

subacutely, with 2 or 3 minute spinules on lateral margin. Antennal flagellum long, overreaching outstretched

chelipeds; every 1 to 4 articles with 1 or 2 very short setae.

Sternite of third maxillipeds (Fig. 8b) produced into blunt or acute spinose process on either side of midline.

Sternite of chelipeds (Fig. 8b) moderately broad, elongate and subtriangular with surface concave, apex blunt or

with shallow median groove. Sternite of second pereopods (Fig. 8b) broad, plate-like, with median longitudinal

groove. Sternite of third pereopods with narrowly subrectangular anterior lobe.

Right cheliped (Figs 8c-d, 34a) elongate; considerably stronger than left; propodal-carpal articulation

perpendicular. Dactyl equal to or slightly longer than palm; articulating obliquely; cutting edge smooth or serrate

and with 2 prominent calcareous teeth; terminating in small calcareous claw, slightly overlapped by fixed finger;

dorsal surface convex, smooth or minutely granular, dorsomesial margin minutely serrate; ventral surface smooth

or microscopically granular. Palm equal to or slightly longer than carpus; somewhat dorsoventrally compressed;

dorsal surface convex, smooth or minutely granular; spinulose or serrate dorsomesial and dorsolateral margins

slightly elevated; fixed finger also with smooth or minutely granular dorsal surface; cutting edge smooth or serrate,

with large teeth; ventral surface of fixed finger with obliquely longitudinal and microscopically tuberculate ridge

extending onto palm in distal half. Carpus equal to or slightly longer than merus; trapezoidal (in dorsal view),

spinulose dorsomesial and dorsolateral margins slightly elevated; dorsal surface with scattered small tubercles or

spinules, also with longitudinal median ridge or crest, most distinct distally and armed with spinules or tuberculate

spines, one strongest in proximal half; mesial and lateral faces minutely granular or spinulose; ventral surface

"hourglass" in shape, ventromesial and ventrolateral margins each with row of very small tubercles or blunted

spines. Merus broadly subtriangular; dorsal margin spinose or spinulose; lateral and mesial faces spinulose or

granular; ventrolateral margin with row of small spines, strongest proximally; ventromesial margin with row of

small spines distally, 3 or 4 very prominent spines proximally. Ischium with row of small tubercles or spinules

on ventromesial margin.

Left cheliped (Figs 8e-f, 34a) dorsoventrally compressed; not reaching to base of dactyl of right; dactyl and fixed

finger curved ventrally. Dactyl 1 .50 to twice length of palm; cutting edge with row of corneous teeth; terminating

in corneous claw and very slightly overlapped by fixed finger; dorsal and ventral surfaces unarmed; dorsomesial

margin serrate, at least in proximal half; mesial face with few scattered very short setae. Palm 0.60 to 0.75 length

of carpus; dorsal surface smooth or minutely granular, dorsomesial and dorsolateral margins serrate and slightly

elevated; ventral surface granular or minutely tuberculate, with short distal longitudinal row of small tubercles

extending onto proximal half of fixed finger; cutting edge of fixed finger with row of small calcareous teeth

interspersed with corneous teeth; terminating in corneous claw. Carpus slightly shorter to slightly longer than

merus; dorsal surface trapezoidal (dorsal view), with few scattered very small tubercles or spinules; dorsomesial and

dorsolateral margins raised and serrate, midline elevated into prominent crest armed with row of simple or

multidenticulate spinulose tubercles or small spines, strongest proximally, and with I very prominent tuberculate

spine at midlength; mesial, lateral, and ventral surfaces spinulose or tuberculate, ventrodistal margin with row of

tubercles. Merus broadly subtriangular; dorsal surface spinose or spinulose, mesial and lateral faces spinulose or

tuberculate particularly near ventral margins; ventromesial margin with row of small spinules and 3 strong spines

proximally; ventrolateral margin minutely spinulose distally becoming row of stronger spines in proximal half.

Ischium with row of small tubercles or spines on ventromesial margin.

Ambulatory legs (Figs 8g-i) moderately long and slender, but not overreaching extended right cheliped;

generally similar. Dactyls approximately equal to length of propodi; laterally compressed, slightly curved

464

P. A. MCLAUGHLIN

ventrally; dorsal margins with few scattered short setae; ventral margins each with row of 9 to 12 corneous spines.

Propodi 1.75 to nearly twice length of carpi; dorsal margins serrate; mesial and lateral faces minutely spinulose,

particularly ventrally; ventral margins each with I or 2 corneous spines at distal angle and 2 additional widely-

spaced corneous spines. Carpi approximately half or slightly less than half length of mcri; dorsal margins

minutely serrate, no distinct spine at distal angle; lateral faces each with dorsal and median longitudinal ridges

separated by concavity, most distinct on third, and with numerous microscopic spinules in ventral halves. Meri

with serrate or spinulose dorsal margins; ventral surfaces oblique (Fig. 8h), ventromesial and ventrolateral margins

spinulose. Fourth pereopods strongly subchelate; propodal rasp (Fig. 8j) consisting of single row of often distally

bulbous corneous scales. Fifth pereopods weakly subchelate; dactyl and propodus (Fig. 8k) each with small rasp of

corneous scales dorsally.

Males with subequal or markedly unequal sexual tubes on coxae of fifth pereopods (Figs 81-m); each with

subterminal and terminal long setae. Females with uniramous, unpaired left pleopods on somites 2 to 4.

Abdomen (Fig. 8n) markedly reduced, segmentation clearly delineated dorsally. Tergite of first abdominal

somite usually chitinous, rod-shaped. Tergites of somites 2 to 5 moderately broad, weakly chitinous plates,

occasionally showing slight tendency toward calcification. Tergite of sixth somite with moderately well calcified

anterior rod-like and posterior paired rectangular plates. Uropods symmetrical; protopods each with very strong,

posteriorly directed subacute spine armed dorsally with longitudinal row of 5 or 6 small spinules; exopods

subcircular, each with large circular rasp of corneous scales; endopods appreciably smaller, ovate, each with small

oval rasp. Telson elongate; transverse suture clearly delineated; terminal margin entire, with rounded external

angles.

Color (in preservative). — After four years in alcohol only faint orange tint remains on chelae and dactyls of

ambulatory legs; propodi of ambulatory legs with one or two very faint bands.

Habitat. — Unknown.

distribution. — Presently known only from the Kai and Tanimbar Islands, Indonesia; 1 1 1-212 m.

Etymology. - This species is named for Dr Michael Turkay, Forschungsinstitut Scnckenberg, who first

recognized the distinctiveness of this genus.

Affinities. — Sohtariopagurus tuerkayi bears a strong resemblance to 5. triprobolus from French Polynesia

in the elongation of the rostrum and lateral projections and development of the lateral carapace spines, as well as

the structure of the abdomen, uropods and telson. However, the lateral projections of S. tuerkayi are generally

shorter and stouter than ,n 5. triprobolus ; the antennular peduncles of the former species are appreciably shorter.

t first of the lateral carapace projections also are much smaller in 5. tuerkayi than those of S. triprobolus There

are clear similarities in the shape of the chelipeds of the two species; however, the dorsodistal angles of the carpi

are not developed into wing-like projections in 5. tuerkayi as they are in 5. triprobolus. The dorsal surfaces of

these segments in 5. tuerkayi each have a raised median crest armed with a row of spines or tubercles set off by

one particularly prominent, tuberculate spine in midlength, whereas the median elevations of these surfaces in

5. triprobolus are slight and very weakly armed. The dactyls and propodi of the ambulatory legs of 5. tuerkayi are

approximately equal in length, in contrast to the much longer propodi of 5. triprobolus.

Genus PORCELLANOPAGURUS Filhol, 1885

188S: «. <*> M. - Borradaile, 1916: 1.1, _ Bennett,

ozu. FOREST, 1951a. 82, 1951b: 182. - Wolff. 1961: 28. - Miyake. 1978: 117,-TOrkay, 1986: 140.

mareinToTshil'lH^ phy'lobranchiale gilIs- Anterior carapace vaulted and well calcified; lateral

nargins of shie d developed into blunt or spiniform projections. Rostrum and lateral projections widely separated

Posterolateral plates calcified anteriorly and usually drawn out into projecting lobes; remamder of

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

465

membraneous or with areas of slight calcification. Ocular acicles reduced, simple. Antennal peduncle with

supernumerary segmentation. Maxillule with external lobe of endopod slightly produced, not recurved. Third

maxilliped with well developed crista dentata and 1 accessory tooth.

Chelipeds unequal. Ambulatory legs generally similar. Fourth pereopods usually semichelate. Fifth pereopods

chelate.

Males without sexual tube on coxa of one or both fifth pereopods; without paired or unpaired pleopods.

Females with paired gonopores; no paired pleopods, unpaired left pleopods on somites 2 to 4.

Abdomen reduced, usually globular, membraneous, but with tergites at least faintly delineated. Uropods

symmetrical or slightly asymmetrical.

REMARKS. — In the original description of Porcellanopagurus tridentatus Whitelegge, 1900, the author

overlooked the first segment of the antennal peduncle, i.e., the segment upon which the antennal gland opens, and

attributed the armature of the second peduncular segment to the first. Takeda (1981, 1985) and Suzuki and

TAKEDA (1987) may, in part, have followed WHlTELEGGE's (1900) interpretation of the first segment in their

inaccurate descriptions of the segmentation and armature of the antenna of their Porcellanopagurus species. These

authors described the basal segment as having a spinule at its outer distal angle and its inner angle developed

forward as a long lobe to grasp the inner margin of second segment, when in fact it is the second segment that is

so armed, and the third segment that is produced. In P. belauensis Suzuki & Takeda, 1987, the authors further

described the "third segment twice as long as the antennal acicula." Neither the true third nor fourth segments are

illustrated as longer than the acicle (SUZUKI & TAKEDA, 1987, figs 2b-c); only the fifth (ultimate) segment agrees

with that description.

Poupin and McLaughlin (1996) noted that, among other characters, the subchelate fourth pereopods found in

species of Solitariopagurus suggested a closer relationship with Alainopagurus Lemaitre & McLaughlin, 1995,

than with Porcellanopagurus, in which the fourth pereopods are semichelate. The fourth pereopods of

P. truncatifrons Takeda, 1981, were described by the author as having "propodus widened and spatulated; its lower

border fringed with a row of horny curved spines, its distal end being prolonged only slight beyond articulation

with dactylus which is talon-like and curved dorsally." From his figure [Takeda, 1981, fig. 3 (3)], it would

appear that this appendage is inverted, thereby giving the impression of a dorsally directed dactyl; however, the

appendage appears distinctly subchelate.

Reference to the telson has been intentionally omitted in the generic diagnosis of Porcellanopagurus given here.

In most species, a typical pagurid-like telson is developed. However, in two recently described Japanese species,

P. truncatifrons and P. nihonkaiensis Takeda, 1985, telsons apparently are lacking. For P. truncatifrons, Takeda

(1981: 12) commented "It is remarkable that I failed to find the telson, but the presence of marginal hairs along the

posterior border of the penultimate segment may justify the absence of the telson." Takeda (1985) illustrated

only the sixth abdominal somite and uropods, but made no comment about the missing telson in

P. nihonkaiensis. Both of these species are known only from their respective holotypes; therefore, it is not clear

whether this ostensible telson loss reflects evolutionary change or simply injury.

Porcellanopagurus jacquesi sp. nov.

Figs 9a-l, 34b

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 18, 05°18'S, 133°01'E, 205-212 m,

24.10.1991: 1 6 (2.8 mm) (MNHN-Pg 5264); 1 <3 (3.2 mm) (MNHN-Pg 5265).

Types. — The male (2.8 mm) (MNHN-Pg 5264) from the Karubar station DW 18 is the holotype. The

other male is a paratype.

DESCRIPTION. — Anterior carapace (Figs 9a, 34b) vaulted; shield length slightly shorter than maximum

breadth, total carapace length usually slightly longer; anterior margin between rostrum and lateral projections

466

P. A. MCLAUGHLIN

F 7 u°,C.elan7aiU^US jace*ues‘ SP' nov- a-b- e, g-1, paratype <J (3.2 mm) from Karubar Stn DW 18

c-d hootype 6 (2.8 mm) from Stn DW 18: a, cephalothorax with cephalic appendages and abdomen

f’uLTT Peduncle (lateral view); c, thoracic sternites; d, right cheliped; e. left chcliped (dorsal view)

' lefl cheliped (mesial view); g, right second pereopod (lateral view); h. left third pereopod (lateral view); i. dactyl

viPw^L aI'd CfPUS ,°f Cft f°U/r Pere°P°d (,ateral Vlew»; J- carpus, propodus and dactyl of left fifth pereopod (lateral

view), k stemite and coxae of fifth pereopods; 1, telson. Scales equal 1.0 mm (b-c, i-1), 2.0 mm (d, 0. and 3.0 mm

straight or very slightly concave; lateral margins each with short acute spine at basally broadened anterolateral

angle, broad multtsp.nose process at midlength, and strong broad or moderately slender spinose process posterior to

cervical groove; dorsal surface strongly calcified, with faintly marked postrostral furrows and weakly delineated

usually somewhat globular posterolateral regions; posterior margin roundly truncate. Cardiac sulci and

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

467

posteromedian plate chitinous or weakly calcified. Rostrum broadly triangular, usually reaching to proximal fourth

of ocular peduncles; terminating subacutely or bluntly. Lateral projections triangular; produced, but not in advance

of rostrum.

Ocular peduncles short, slightly less than half length of shield, with distinct submedian constriction; cornea

slightly dilated, diameter approximately 0.33 length of peduncle. Ocular acicles small, acutely triangular, obscured

from dorsal view by base of rostrum.

Antennular peduncles moderately short, but overreaching ocular peduncles by nearly full length of ultimate

segment. Ultimate segment with 2 or 3 long plumose setae on dorsodistal margin. Penultimate and basal segments

unarmed.

Antennal peduncles (Fig. 9b) overreaching ocular peduncles by 0.25 to 0.50 length of ultimate segment. Fifth

and fourth segments with few scattered, very short setae. Third segment unarmed. Second segment with dorsolateral

distal angle produced, terminating in acute spine; dorsomesial distal angle rounded. First segment with cluster of

spines on lateral surface ventrally. Antennal acicle moderately short, but sometimes reaching beyond distal margin

of fourth peduncular segment, terminating acutely. Antennal flagellum long, slightly overreaching outstretched

chelipeds; every 1 to 4 articles with 1 or 2 very short setae, at least in proximal half.

Sternite of third maxillipeds (Fig. 9c) produced into rounded or acute spinose process on either side of midline.

Sternite of chelipeds (Fig. 9c) slender, elongate with asymmetrical posterior lobe. Sternite of second pereopods

(Fig. 9c) broad, plate-like, with median longitudinal groove. Sternite of third pereopods with roundly

subrectangular anterior lobe.

Right cheliped (Figs 9d, 34b) (missing in paratype) stout; not much longer, but considerably stronger than left;

propodal-carpal articulation twisted clockwise approximately 45° from perpendicular. Dactyl slightly longer than

palm; articulating obliquely; cutting edge with 4 prominent teeth; terminating in small calcareous claw, slightly

overlapped by fixed finger; dorsal surface convex, armed with few tiny spinules, dorsomesial margin minutely

serrate; ventral surface with few widely-spaced tufts of setae. Palm approximately equal to length of carpus;

somewhat swollen dorsoventrally; dorsal surface convex, dorsomesial and dorsolateral margins slightly elevated,

tuberculate or spinulose; fixed finger with few tufts of short setae on dorsal surface; cutting edge with 2 large teeth;

ventral surface of palm and fixed finger microscopically spinulose and with scattered tufts of setae. Carpus slightly

longer than merus; trapezoidal (in dorsal view), dorsomesial and dorsolateral margins also slightly elevated, at least

distally, crenulate and with few low transverse ridges; all surfaces with scattered short low transverse ridges

providing rough-textured appearance. Merus broadly subtriangular; dorsal margin with several transverse ridges and

few fine setae; lateral and mesial faces minutely granular in ventral halves; ventromesial margin with row of small

spines; ventrolateral margin with row of somewhat stronger spines distally and few widely-spaced spinules

proximally. Ischium with row of small spinules on ventromesial margin.

Left cheliped (Figs 9e-f, 34b) with dactyl and fixed finger curved ventrally. Dactyl approximately 1.25 length of

palm; cutting edge with row of corneous teeth; terminating in corneous claw and very slightly overlapped by fixed

finger; dorsal surface unarmed, dorsomesial margin not delimited; mesial and ventral surfaces with scattered setae.

Palm nearly 0.75 length of carpus; dorsal surface smooth or microscopically granular, dorsomesial and dorsolateral

margins serrate and slightly elevated; ventral surface weakly granular; cutting edge of fixed finger with row of

small calcareous teeth, terminating in corneous claw. Carpus 0.65 to 0.75 length of merus; with low transverse

ridges mesially and laterally; dorsolateral margin raised proximally and distally; mesial, lateral and ventral surfaces

each with few short transverse low ridges and sparse setae. Merus subtriangular; dorsal surface with few short

transverse ridges and short setae; lateral face minutely spinulose or tuberculate; ventromesial margin with row of

small spines, ventrolateral margin with row of stronger spines. Ischium with row of small tubercles on

ventrolateral margin; row of small spines on ventromesial margin.

Ambulatory legs (Figs 9g-h) moderately long and slender, overreaching extended right cheliped; generally

similar. Dactyls approximately equal to or slightly shorter than propodi; in dorsal and lateral views, nearly

straight; dorsal margins with short setae; mesial and lateral faces with few short setae; ventral margins each with

row of 8 to 1 1 corneous spines. Propodi 1.10 to 1.25 length of carpi; dorsal margins slightly tuberculate; ventral

margins each with 1 or 2 corneous spines at distal angle and 2 to 6 additional widely-spaced corneous spines. Carpi

approximately half or slightly more than half length of meri; dorsal margins slightly spinulose, no distinct spine

468

P. A. MCLAUGHLIN

at distal angle; lateral faces each with longitudinal ridge separated by concavity from dorsal margin. Meri with

minutely protuberant or serrate dorsal margins; ventral surfaces oblique (mesial view), ventromesial and

ventrolateral margins spinulose. Fourth pereopods strongly semichelate; propodal rasp (Fig. 9i) consisting of

single row of corneous scales. Propodus of fifth pereopods with small rasp of corneous scales (Fig. 9j).

Males with coxae of fifth pereopods (Fig. 9k) drawn out posteromedially, gonopores each masked by long

setae. Females unknown.

Abdomen (Fig. 9a) somewhat reduced, globular. Tergite of first abdominal somite rod-shaped, membraneous

and only faintly delineated. Tergites of somites 2 to 5 also membraneous, moderately broad, weakly indicated.

Tergite of sixth somite moderately well calcified, divided into anterior and posterior lobes by transverse suture,

each with median furrow. Uropods symmetrical; protopods with weak posteriorly directed subacute protuberance on

right side only. Telson (Fig. 91) (missing in holotype) with transverse suture weakly delineated; posterior lobes

separated by very small median cleft, terminal margins rounded, unarmed.

COLOR (in preservative). — Both specimens, after four years in alcohol, showed faint light orange banding at

proximal margins of dactyls and medianly on propodi of ambulatory legs; carpi each with nearly imperceptible

longitudinal stripe on lateral ridge.

FlABITAT. — Unknown.

DISTRIBUTION. — At present known only from the type locality in the Kai Islands. Indonesia; 205-212 m.

Etymology. — This species is named for Professor Jacques FOREST of the Museum national d'Histoire

Naturelle, Paris, in recognition, not only of his landmark studies of Porcellanopagurus , but his many

contributions to hermit crab systematics.

Affinities. — Porcellanopagurus jacquesi most closely resembles P. japonicus Balss, 1913, P. nihonkaiensis

Takeda, 1985, and P . belauensis Suzuki & Takeda, 1987, in having a broadly triangular rostrum, somewhat broad

and spinulose or denticulate lateral carapace lobes, and additionally with the latter two species, the laterally ridged

carpi of the ambulatory legs. Porcellanopagurus jacquesi shares with P. japonicus a small, spiniform posterior

carapace lobe; in both P. nihonkaiensis and P. belauensis this lobe is quite blunt. Like P. jacquesi, a postrostral

furrow is present in both P. japonicus and P. belauensis that is lacking in the only known specimen of

P. nihonkaiensis. The presence of a strongly acute, spiniform rostrum and absence of laterally ridged carpi of the

ambulatory legs immediately distinguishes P. japonicus from the other three species. The development, in

P. nihonkaiensis and P. belauensis, of a triangular distal lobe on the rostrum also sets these species apart from

P. jacquesi, where the rostrum terminally is subacute or rounded and slightly depressed. The width of the shield is

appreciably greater in P. belauensis than in the other three species.

In P. nihonkaiensis the ventral margins of the propodi of the ambulatory legs are described as having five

corneous spines, the dactyls seven; P. belauensis is reported to have three corneous spines on the ventrodistal angle

and a row of seven additional spines on the ventral margins of the propodi of the second pereopods, while the

spines on the third pereopods number two distally and five marginally. The dactyls of the second pereopods have

nine spines, the third, 1 1 in this species. Porcellanopagurus jacquesi is intermediate in having one or two corneous

spines on the ventrodistal angles of the propodi and three to six on the ventral margins; the ventral spines of the

dactyls range from eight to 1 1.

Iakeda (1985, figs 1 A-C) illustrated, but did not discuss the tergites of the sixth abdominal somite and

uropods in P. japonicus, P. nihonkaiensis and P. truncatifrons. For P. japonicus the sixth tergite is represented by

a rounded anterior lobe with shallow, incomplete median furrow and rectangular posterior lobe with deeper but still

incomplete median furrow. This tergite in P. nihonkaiensis and P. truncatifrons is illustrated as having a triangular

anterior lobe with central median depression; the rectangular posterior lobe has a well developed and nearly

complete longitudinal groove. As described above and illustrated in Fig. 9a, P. jacquesi has two subequal

rectangular lobes, each with a longitudinal median furrow, incomplete only on the posterior lobe. Like P. jacquesi,

the uropods of P. japonicus show only a small protuberance posteriorly from the protopod of the right uropod. In

P. nihonkaiensis the protopods of both uropods show a well developed posterior protuberance, and in

PAGUR1DAE FROM THE KARUBAR EXPEDITION

469

P. truncatifrons an even more prominent spiniform protuberance. Depicted by SUZUKI and TAKEDA (1987,

fig. 3p), but not described is a similar strong spine on the protopod of each uropod of P. belauensis ; the sixth

abdominal tergite in this species is clearly divided into four distinct, well separated lobes.

Balss (1913) did not describe the telson of P. japonicus but his figure (fig. 40) shows a reduced and

subtriangular structure. As previously mentioned, TAKEDA (1985, fig. 1B-C) illustrates P. rtihonkaiensis and

P. truncatifrons without telsons. His specimen of P. japonicus is depicted with an elongate telson, lacking both

transverse suture and median cleft. The telson of P. belauensis, as illustrated by SUZUKI and TAKEDA (1987), has

the anterior and posterior lobes divided by a complete transverse suture, and the posterior lobes by a longitudinal

median groove over their entire length. Porcellanopagurus jacquesi has a well developed telson with transverse

suture weakly marked and with a small median cleft separating the posterior lobes.

Rather abundant setation was illustrated by BALSS (1913, fig. 40) for P. japonicus. Both in the description and

figures (SUZUKI & TAKEDA, 1987) P. belauensis is characterized as having abundant setation on the carapace and

all appendages. In contrast, setation was described by TAKEDA (1985) as being sparse in P. nihonkaiensis. Very

sparse setation is also characteristic of P. jacquesi.

Remarks. — Although the specimen selected as the holotype is the smaller of the two, it is the only

specimen with all appendages still attached. The carapaces of both specimens were damaged in collection; however,

that of the paratype had been severed in such a way that it has been possible to reconstruct it accurately (Fig. 9a).

The abdomen of this specimen is in excellent condition, whereas that of the holotype is badly shriveled and the

telson is missing.

WHITELEGGE (1900), in his original description of Porcellanopagurus tridentatus, described the coxae of the

fifth pereopods as having a "tubular prolongation, directed inwards and downwards, and their margins are fringed

with long setae". His statement could apply equally well to the coxae of P. jacquesi, although there is no doubt

that the two are distinct taxa. When discussing the relationship of Solitariopagurus with Porcellanopagurus,

TURKAY (1986), called attention to WHITELEGGE’s (1900) report, but commented that he could detect no sexual

tubes in the male syntype of this species that he examined. Close inspection of the coxae of both males of

P. jacquesi similarly does not support the supposition that these elongations represent sexual tubes.

Genus ALAINOPAG UR OIDES nov.

DIAGNOSIS. — Eleven pairs of phyllobranchiate gills. Anterior carapace vaulted and generally well calcified,

with anterolateral regions slightly depressed. Linea transversalis as calcified rod, posterior carapace membraneous or

with slight calcification. Ocular acicles simple. Antennal peduncle with supernumerary segmentation. Maxillule

with external endopodal lobe obsolete or absent. Third maxilliped with crista dentata of ischium (Fig. 10a)

somewhat reduced, but with 1 accessory tooth. Sternite of third maxillipeds with anterior margin rounded on either

side of median concavity. Sternite of second pereopods subdivided into broad lateral lobes by deep longitudinal

median groove. Sternite of third pereopods (Fig. 10b) with narrow, transverse anterior lobe and broad posterior lobe

divided by deep median groove. Fourth pereopods weakly semichelate, propodal rasp rudimentary; with prominent

tubular preungual process (Fig. 10c) at base of claw.

Abdomen reduced; tergal plate of first somite chitinous; tergal plates of second through fifth somites

sometimes very faintly delineated; tergite of sixth somite (Fig. lOd) weakly calcified, subdivided into narrow to

moderately broad anterior lobe, and posterior pair of broad plates separated by distinct median groove. Uropods

generally symmetrical. Telson with transverse suture; posterior lobes usually separated by shallow median cleft.

Males with moderately long and stout sexual tube on coxa of right fifth pereopod (Fig. lOe), left often with

very short tube; no paired or unpaired pleopods. Female with paired gonopores; no paired pleopods; unpaired left

biramous pleopods on abdominal somites 2 to 4 (Fig. lOf).

Type Species. — Alainopaguroides lemaitrei, sp. nov. Gender masculine.

Etymology. — This genus is named for Alain CROSNIER, marine biologist of ORSTOM, in recognition of

his many contributions to our knowledge of the decapod fauna of the Indo-Pacific.

470

p. a. McLaughlin

AFFINITIES. — Although Alainopaguroides shares a number of important characters with several other genera,

no precise phylogenetic relationships can yet be determined. In reduction of the abdomen, total absence of paired or

unpaired pleopods in males and lack of the left fifth pleopod in females, Alainopaguroides, Porcellanopagurus,

Ostraconotus A. Milne Edwards, 1 880, Solitariopagurus, and Alainopagurus conceivably might all be considered

sister taxa. However, 11 pairs of phyllobranchiate gills are common to species of Porcellanopagurus,

Alainopagurus and Alainopaguroides, while Ostraconotus and Solitariopagurus species have only 10. The rostrum

is very strongly produced and broadly triangular in species of Solitariopagurus and some species of

Porcellanopagurus, developed as a prominent slender spiniform projection in Alainopagurus, but broad and bluntly

truncated in other species of Porcellanopagurus or upturned in Ostraconotus and Alainopaguroides. The lateral

regions of the anterior carapace are drawn out into spinose lobes in Porcellanopagurus and Solitariopagurus, or

simply globular in Alainopagurus, whereas only spinulose margins are seen in Ostraconotus, and simply rounded

margins in Alainopaguroides. Varying areas of weak calcification are observed on the posterior carapaces of species

of four of the genera; yet, only in Ostraconotus is there nearly complete calcification of the posterior carapace. The

second through fifth abdominal tergites of Solitariopagurus and Alainopagurus are chitinized or calcified to some

extent, while those of the other three genera are only faintly indicated.

Quite different specializations in the fourth pereopods are seen among the five genera. With the one exception

previously mentioned, in Porcellanopagurus species the fourth pereopod is semichelate; the propodal rasp consists

of a single row of very small spinules. The semichelae of Alainopagurus and Alainopaguroides are similar in

having the ventrodistal portion of the propodus very weakly produced; the propodal rasp consists of a single row of

small spines in Alainopagurus, but only two or three distal scales or corneous spines in Alainopaguroides. The

fourth pereopods of Solitariopagurus species are distinctly subchelate, allowing the dactyl to articulate against the

ventral margin of the propodus; the propodal rasp consists of a row of bulbous scales. Major structural differences

and sexual dimorphism are seen in the fourth pereopods of Ostraconotus. Females of the only known species,

Ostraconotus spatulipes A. Milne Edwards, 1880, have an extremely paddle-shaped propodus and simple

articulating dactyl; the propodus of the male is approximately half the breadth of the female, but the dactyl is

similar. Rather typical pagurid type, minutely chelate, fifth pereopods are found in Ostraconotus,

Porcellanopagurus and Alainopaguroides, but subchelate in Solitariopagurus and in Alainopagurus. Species of

Porcellanopagurus, Alainopagurus and one species of Solitariopagurus are known to utilize bivalve shells as their

microhabitat; nothing is known for the others.

Extensions of the vas deferens as sexual tubes are present in all of the aforementioned genera except

Porcellanopagurus. Species of this latter genus occasionally show a posterior elongation of one or both coxae, but

no tubular structure arises from a gonopore. In contrast, males of Alainopagurus and Solitariopagurus have short,

equal or unequal sexual tubes produced from the gonopores of both fifth pereopods. Males of Ostraconotus have a

single right sexual tube, whereas Alainopaguroides has an elongate right sexual tube and usually a very short left,

although it may only resemble a small protuberance in smaller specimens. While females of all five genera have

no paired pleopods and only left unpaired egg-bearing pleopods (left second through fourth), paired gonopores occur

in Porcellanopagurus, Ostraconotus, and Alainopaguroides, but only single left gonopores are found in females of

Solitariopagurus and Alainopagurus.

Until Turkay's (1986) account of Solitariopagurus, Ostraconotus and Porcellanopagurus had been considered

the prime examples of carcinization in the Paguridae (e.g., BORRADAILE, 1916; Wolff. 1961). The recent

discovery of Alainopagurus, and now Alainopaguroides, brings to five the candidates for this distinction. As may

be seen from the foregoing brief review of major characters in these genera, no clearly defined evolutionary

pathway can yet be observed.

Alainopaguroides lemaitrei sp. nov.

Figs lOa-m, 34c-e, 35a

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 13, 05°26'S, 132°38'E, 417-

415 m, 24.10.1991: 1 6 , 4 $ (1. 3-2.6 mm) (MNHN-Pg 5266). — Stn DW 31, 05° 40'S, 132°51'e' 288-

289 m, 26.10.1991: 1 <J (2.1 mm) (POLIPI), — Stn CP 35, 06°08’S, 132°45'E, 390-502 m, 27.10.1991: 1 ov. $

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

471

(4.9 mm) (MNHN-Pg 5267). — Stn CP 36. 06°05'S, 132°44'E, 268-210 m, 27.10.1991: 15,19 (2.6. 3.6 mm) (USNM

276007).

Tanimbar Islands: stn CP 59. 08°20'S. 132°11'E. 405-399 m, 31.10.1991: 1 3 (6.2 mm) (MNHN-Pg 5268 A), 3 2,

2 ov. 2 (4. 7-5. 2 mm) (MNHN-Pg 5268 B); 1 3, 2 2, 1 ov. 2 (4.5-S.6 mm) (USNM 276006). — Stn CP 62, 09°01'S,

132°42'E, 246-253 m, 1.11.1991: 1 3 (5.4 mm) (SMNH 4813). — Stn CP 70, 08°41'S. 131°47'E, 413-410 m.

2.11.1991: 1 3 (3.9 mm) (POLIPI).

Types. — The male (6.2 mm) (MNHN-Pg 5268 A) from Karubar station CP 59 is the holotype. The other

specimens are paratypes.

Description. — Shield (Figs lOf, 34c, 35a) usually considerably broader than long, dorsal surface swollen,

anterolateral region somewhat depressed; anterior margin between rostrum and lateral projections slightly concave;

posterior margin roundly truncate; dorsal surface with light covering of short to moderately long setae, longest and

most abundant laterally; frequently with moderately deep transverse postrostral depression. Rostrum broad, blunt or

rarely subacute, unarmed, upturned. Lateral projections broadly rounded or obtusely triangular, with small marginal

or submarginal spine or spinule.

Ocular peduncles (including corneae) very short, slightly more than half shield length, slender basally but

enlarged distally; with corneae strongly dilated. Ocular acicles moderately short to moderately long, 0.35 to

0.50 length of peduncle (excluding cornea), slender, triangular; terminating acutely and with dorsal surface

longitudinally grooved; mesial margins each with row of moderately long setae and few additional setae laterally;

separated basally by more than basal width of one acicle.

Antennular peduncles (left broken in holotype) overreach ocular peduncles by approximately half length of

penultimate segment. Ultimate segment with 2 widely separated rows of long setae on dorsal surface. Penultimate

segment with few setae. Basal segment unarmed, but with statocyst enlarged.

Antennal peduncles overreaching ocular peduncles (including corneae) by 0.25 to 0.50 length of ultimate seg¬

ment. Ultimate and penultimate segments with few to several moderately long setae. Third segment with

small spinule on ventrodistal margin. Second segment with dorsolateral distal angle produced, terminating

in slender spine and frequently with accessory spine on lateral margin, with long setae dorsally, mesially and later¬

ally; dorsal surface with numerous long setae and prominent longitudinal furrow, mesial and lateral margins with

low, spinulose protuberances and long setae; dorsomesial distal angle with small spine. First segment with strong

spine on laterodistal margin; ventrally produced and with 1 small spine distolaterally. Antennal acicle reaching

nearly to distal margin of ultimate peduncular segment; terminating in small spine; mesial, lateral and dorsal sur¬

faces all with long setae, margins sometimes also with spinulose protuberances. Antennal flagellum longer than

outstretched ambulatory legs; each article naked or proximal articles with 1 or 2 minute bristles.

Right cheliped (Figs lOg, 34d) stronger; frequently but not always, shorter than left in large males; palm, fixed

finger and dactyl somewhat dorsoventrally compressed. Dactyl 0.65 to 0.75 length of palm; cutting edge with

2 broad calcareous teeth and often numerous small calcareous denticles; terminating in small corneous claw,

slightly overlapped by fixed finger; dorsomesial margin not delimited, dorsal surface with few tufts of short setae;

mesial and ventral surfaces with moderate to dense long setae. Palm 1.35 to 1.50 length of carpus; dorsal surface

very slightly convex, unarmed; dorsomesial and dorsolateral margins not delimited, mesial and lateral faces

spinulose, sometimes almost imperceptibly so; fixed finger with scattered short and long setae on weakly convex

dorsal surface; cutting edge with 1 broad sometimes denticulate calcareous tooth, few to many much smaller

calcareous teeth proximally and distally, terminating in small corneous claw; ventral surfaces of palm and fixed

finger with moderate to dense long setae. Carpus 0.65 to 0.75 length of merus; dorsomesial and dorsolateral

margins each with row of irregularly-sized slender spines, strongest distally, dorsal surface with scattered spinules;

lateral, mesial and ventral surfaces also spinose, ventromesial and ventrolateral margins with slightly stronger

serrations; all surfaces with scattered fine setae. Merus subtriangular; dorsodistal margin with 1 strong sometimes

nearly erect spine; slightly flattened dorsal margin with very short transverse rows of spines or spinules; lateral

face spinulose, ventrolateral margin with irregular single or double row of small spines, 1 more prominent spine at

distal angle; mesial and ventral surfaces spinulose, ventromesial margin spinulose or with distinct row of small

spines, 1 stronger spine at distal angle. Ischium with row of fine setae on ventral margin.

472

P. A. MCLAUGHLIN

Fig 10. — Alainopaguroides lemaitrei sp. nov., a, c-d, g-1, paratype $ (5.6 mm) from Karubar Sin CP 59;

b. e, m, holotype 6 (6.2 mm) from Stn CP 59: a, basis and ischium of left third maxilliped with crista dentata and

accessory tooth; b, stemite of third pereopods; c, tip of dactyl and preungual process of left fourth pereopod; d. sixth

abdominal somite and telson (dorsal view); e, sternite and coxae of Fifth pereopods; f. cephalothorax with cephalic

appendages and abdomen; g. right cheliped (lateral view); h. left cheliped (lateral view); i, right second pereopod

(lateral view); j. left third pereopod (lateral view); k. dactyl and propodus left fourth pereopod (lateral view);

1. propodus and dactyl of right fifth pereopod (lateral view); m, telson. Scales equal 0.1 mm (c), 1.0 mm (a, k-1),

2.0 mm (b, d, e, m) and 5.0 mm (f-j).

Left cheliped (Figs lOh, 34e) in large males sometimes overreaching right cheliped by as much as half length

of dactyl; chela dorsoventrally compressed; dactyl and fixed finger slightly arched. Dactyl long, approximately

twice length of palm; dorsomesial margin microscopically serrate, at least proximally; dorsal and ventral surfaces

with scattered long and short setae; cutting edge with row of tiny corneous teeth, terminating in small corneous

claw and slightly overlapped by fixed finger. Palm 0.50 to 0.65 length of carpus; with nearly flat dorsal surface,

dorsomesial and dorsolateral margins usually minutely serrate, latter also often with row of widely-spaced tiny

bristles extending almost entire length of fixed finger; mesial, lateral and ventral surfaces with scattered usually

PAGURIDAE FROM THE KARUBAR EXPEDITION

473

short setae; cutting edge of fixed finger with few widely-spaced very small calcareous teeth, interspaces occupied

by short rows of corneous teeth, terminating in small simple or bifid claw; dorsal and ventral surfaces with

scattered long setae. Carpus about 0.65 length of merus, subrectangular; dorsomesial and dorsolateral margins

weakly elevated, each with row of small slender spines, strongest at distal angles, dorsal surface slightly concave,

unarmed; mesial and lateral faces nearly perpendicular, unarmed; ventromesial margin with row of widely-spaced

short setae, ventrolateral margin with row of very small spines. Merus subtriangular; dorsodistal margin with

spine, slightly flattened dorsal margin with small spinules and moderately short setae; lateral face spinulose,

ventrolateral margin with row of small spines; ventromesial margin with row of spinules, ventral surface with

scattered small spines and spinules. Ischium with row of fine setae on ventromesial margin.

Ambulatory legs (Figs lOi-j) overreaching right chelipcd by full length of dactyls; generally similar. Dactyls

slightly blade-shaped, particularly third, usually somewhat shorter than propodi; laterally compressed; in dorsal

view, slightly twisted in distal half; in lateral view, curved venlrally; terminating in small corneous claws; dorsal

surfaces each with row of regularly-spaced stiff long setae; lateral faces each with faint broad longitudinal sulcus

proximally, mesial faces glabrous; ventral margins each with row of shorter and finer but also regularly-spaced

setae. Propodi 1.75 to twice length of carpi, somewhat laterally compressed; dorsal and ventral surfaces each with

row of long setae; mesial and lateral faces unarmed. Carpi 0.45 to 0.65 length of meri; dorsal surfaces each with

row of tiny spinules, 1 stronger spine at distal margin; lateral faces each with median longitudinal row of short

fine setae sometimes accompanied by tiny spinules; mesial and ventral surfaces unarmed. Meri each usually with

small spine at dorsodistal margin, at least on second, dorsal surfaces often spinulose or spinose and with abundant

short fine setae; mesial faces glabrous; lateral faces minutely spinulose, each also with irregular rows of short fine

setae; ventromesial margins unarmed or with few minute spinules proximally, ventrolateral margins, and

occasionally also broadened ventral surfaces, each with irregularly double or triple rows of spinules or small

spines, 1 slightly larger spine at distal angle. Ischia unarmed. Fourth pereopods each with 2 or 3 ovate corneous

scales or spines at ventrodistal angle of propodus (Fig. 10k), lateral face often abundantly setose; dactyl with

prominent preungual process at base of claw (Fig. lOd). Fifth pereopods chelate; propodus with diffuse rasp (Fig.

101).

Uropods (Fig. lOf) with elongate moderately narrow rasps of corneous scales on both exopods and endopods;

protopods not produced posteriorly. Telson (Figs lOd, m) with roundly subtriangular posterior lobes usually

separated by small median cleft, sometimes cleft nearly obsolete; narrowly or broadly oblique terminal margins

each usually with 1 or 2 spinules and very short bristles.

COLOR. — Unknown.

Habitat. — One specimen found occupying gastropod shell well covered by anemone.

Distribution. — Presently known only from the Kai and Tanimbar Islands, Indonesia; 210 - 502 m.

Etymology. — This species is named for Dr. Rafael Lemaitre, National Museum of Natural History,

Smithsonian Institution in recognition of his continuing contributions to pagurid systematics.

AFFINITIES. — The generic discussion previously presented pertains directly to A. lemaitrei as it is the only

known representative of the genus. However, the elongation of the right sexual tube and its right-to-left direction

over the dorsal part of the abdomen in this genus is more reminiscent of sexual tube development in Catapagurus

than in any of the other previously discussed genera. Alainopaguroides also shares with Catapagurus such charac¬

ters as short and stout ocular peduncles with dilated corneae and moderate to long ocular acicles, although these are

also seen in Ostraconotus spatulipes. The general structure and armature of the chelipeds and ambulatory legs, as

well as the specialized preungual process on the fourth pereopod, also resemble those of Catapagurus species.

REMARKS. — Contrary to the typical relationship of shield length to animal size in the majority of pagurids,

this is not the case in A. lemaitrei. In this species, there is a marked increase in shield breadth in proportion to

length with increased animal size. However, as width increases, so does the general convexity of the shield,

making accurate linear width measurements impossible.

474

P. A. MCLAUGHLIN

As indicated in the description, there is a tendency in some males for the length of the left cheliped to

noticeably exceed that of the right; however, this phenomenon does not appear to necessarily be a function of size.

In the holotype, which was the largest male examined, the left cheliped was shorter than the right, as it was in all

females. In two of the slightly smaller males, the left cheliped overreached the right by a quarter to a half the

length of the left dactyl.

Genus ANAPAGRIDES de Saint Laurent-Dechance, 1966

Anapagrides de Saint Laurent-Dechance, 1966b: 262 (in part). — Miyake, 1978: 141 (in part). — McLaughlin &

Sandberg, 1995: 580.

Nanopagurus McLaughlin, 1986: 797.

Not Anapagrides - de Saint Laurent, 1968b: 1 1 15. — Haig & Ball, 1988: 177 (= Turleania nom. nov.).

DIAGNOSIS. — Eleven pairs of phyllobranchiate gills. Shield with well developed rostrum. Ocular acicles

triangular. Antennal peduncle with supernumerary segmentation. Third maxilliped with well developed crista

dentata and 1 accessory tooth. Chelipeds unequal, right appreciably larger. Ambulatory legs similar from left to

right; carpi with or without dorsodistal spinule. Fourth pereopod semichelate; with single row of corneous scales

in propodal rasp.

Males with coxae of fifth pereopods slightly asymmetrical; right larger and with short sexual tube directed

posteriorly. No paired pleopods, 3 unpaired left pleopods. Females with single gonopore on coxa of third left

pereopod. No paired pleopods, 4 unpaired pleopods; second to fourth with both rami well developed, fifth reduced.

Telson with transverse suture; posterior lobes separated by median cleft; terminal margins with few small

spines.

Remarks. — The inclusion of Anapagrides in the Karubar material is based upon a single ovigerous female

of uncertain specific identity, and must therefore be considered tentative.

? Anapagrides sp.

Figs lla-g

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 18, 05°18'S, 133°01'E 205-212 m 24 10

1991: 1 ov. 9 (1.1 mm) (MNHN-Pg 5269). ’

Diagnosis. Shield (Fig. 11a) slightly longer than broad. Rostrum broadly triangular, not produced in

advance of lateral projections. Lateral projections triangular, with small marginal or submarginal spine. Ocular

peduncles moderately short and stout, much shorter than antennular peduncles. Ocular acicles narrowly triangular,

with submarginal spine. Antennal peduncles reaching slightly beyond distal margin of corneae; shorter than

antennular peduncles.

Right cheliped (Fig. 1 lb) with moderately long fine setae particularly laterally on carpus and ventrally on

merus. Dorsal surface of dactyl elevated in midline and armed with row of tiny spinules, dorsomesial margin with

row of very small spines. Palm with row of small spines on dorsomesial and dorsolateral margins and slightly

shorter row adjacent to dorsomesial margin, dorsal surface of palm and fixed finger with scattered spinules. Carpus

with row of spines on dorsomesial margin, 1 spine on dorsodistal margin and row of slightly smaller spines on

dorsal surface mesially; dorsolateral margin not distinctly delimited, but with row of tiny spinules; ventrolateral

margin with row of minute tubercles. Merus with row of spines on ventrolateral margin.

Left cheliped (Fig. 1 lc) with long fine setae, particularly mesially on carpus and ventrally on merus. Dorsal

surface of dactyl with few tiny spinulose protuberances. Palm strongly elevated in midline and armed with row of

small spines extending to distal half of fixed finger. Carpus with short row of spines on dorsolateral margin,

dorsomesial margin with 1 small spine distally and row of minutely spinulose protuberances; distal margin with

1 spine dorsally and 2 laterally. Merus with 2 strong spines on ventrolateral margin distally.

Ambulatory legs (Figs lld-e) similar. Dactyls each with row of 7 or 8 corneous spines on ventral margins;

mesial faces of third pereopods each with row of widely-spaced corneous spines dorsally. Propodi with 1 or

PAGURIDAE FROM THE KARUBAR EXPEDITION

475

Fig. 11. — lAnapagrides sp., a-g, ov. 9 (1.1 mm) from Karubar Sin DW 18. — Turleania senticosa McLaughlin & Haig,

1996. h, 6 (2.1 mm) from Stn DW 22; i, 6 (2.3 mm) from Stn DW 22; j, 6 (1.6 mm) from Sin DW 50;

k, 9 (1.6 mm) from Sin DW 50: a, h, shield and cephalic appendages; b, chela and carpus of right cheliped (dorsal

view); c, chela and carpus of left cheliped (dorsal view); d, right second pereopod (lateral view); e, dactyl of left third

pereopod (mesial view); f, anterior lobe of sternite of third pereopods; g, i-k, telson. Scales equal 0.5 mm (f-g, i-k)

and 1.0 mm (a-e, h).

Source :

476

P. A. MCLAUGHLIN

2 spines on ventrodistal margins and 2 or 3 widely-spaced and smaller spinules on ventral surface. Carpi each with

minute spinule on dorsodistal margin. Sternite of third pereopods with subsemicircular unarmed anterior lobe

(Fig. Ilf).

Telson (Fig. 1 1 g) with terminal margins of posterior lobes horizontal to slightly oblique, armed with 4 and

7 small spines, lateral margins with serrate marginal plate.

Color (in preservative). — Only remaining color: left cheliped faint orange with tips of dactyl and fixed finger

white, distal margin of carpus darker orange and light orange band distally on merus.

Habitat. — Unknown.

Distribution. — Known only from one locality in the Kai Islands, Indonesia; 205-212 m.

Remarks. — This specimen has been referred to Anapagrides because of its considerable similarity to ,4. reesei

(McLaughlin, 1986), a species known only from subtidal Pocillopora rubble at Kahe Point, Oahu, in the Hawaiian

Islands. Anapagrides reesei , like the Karubar specimen, is very small, with females ovigerous in very much the

same size range (1.2- 1.4 mm). The armature of the left cheliped is nearly identical in the two species, as is that of

the right chela. Like A. reesei, the anterior lobe of the sternite of the KARUBAR specimen is subsemicircular; the

terminal margins of the telsons of both taxa are slightly oblique and armed with a few spines, the lateral margins

carry a serrate chitinous plate. The latter species differs from the Hawaiian taxon in having slightly longer antennal

peduncles, a row of tiny spines on the dorsolateral margin of the carpus of the right cheliped, and minute spinules

on the dorsodistal margins of each of the ambulatory legs.

There is also an affinity, albeit less striking, between the Karubar specimen and A. facetus (Melin, 1939),

particularly in the relative proportions of the cephalic appendages; however the rostrum of A facetus is stronger

than the lateral projections. In A. facetus at least the dactyl of the third left pereopod has a row of corneous spines

on the mesial face (cf. McLaughlin & Sandberg, 1995). Similar armature of the mesial faces of the dactyls of

the third pereopods can be observed in the Karubar specimen. Differences in the female of /l. facetus include

having a pair of strong spines on the dorsal surface of the carpus of the right cheliped, whereas the Karubar

specimen has a lateral row of small spines. The anterior lobe of the telson of A. facetus is subrectangular.

In the absence of male characters, a female specimen having a single left gonopore and no paired first pleopods

modified as gonopods also conceivably might represent a species of Trichopagurus de Saint Laurent, 1970b or

Pagurixus. However, the gills are intermediate in the monotypic Trichopagurus, but unquestionably phyllobranch

in the Karubar specimen. The occurrence of an unpaired left gonopore in species of Pagurixus is variable; but in

all species, the rostrum is usually well developed and the lateral projections weak or obsolete. A reversed condition

is seen in the Karubar specimen. Additionally, all of the 13 species of Pagurixus now recognized (cf. Morgan,

1993; Komai & Asakura, 1995) are described as having a subrectangular or subquadrate anterior lobe developed

on the sternite of the third pereopods.

Tan asopagurus rostrodenticulatus gen. nov., sp. nov., and 1 Anapagrides sp. also share common characters such

as the armature of the chelipeds and spination of the ambulatory legs. Females of the two taxa are readily

distinguished, T. rostrodenticulatus by the presence of paired gonopores and first pleopods, ? Anapagrides sp. by a

single left gonopore and the absence of paired first pleopods. If assignment of the Karubar specimen to

Anapagrides is correct, males would correspondingly be easily distinguished by sexual tube development, i.e., left

in T. rostrodenticulatus, right in Anapagrides sp. Although these two taxa occurred sympatrically at Stn DW 18,

the intermediate type gills of the two presumably juvenile male specimens assigned to T. rostrodenticulatus

immediately distinguished them from l Anapagrides sp., despite their lack of sexual tubes and denticulate rostra.

Genus TURLEANIA nom. nov.

Anapagrides de Saint Laurent-Dechanctf 1966b: 262 (in pan). - Miyake, 1978: 142 (in part)

\a966bS' °E SAINT LAURENT' 1%8b: ' ' 15' “ HA,G & BALLl l988: ' 77; not AliaPaSr^ de Sain. Laurent-Dechanc6,

Laurentia McLaughlin & Haig, 1996: 76; not Laurentia Ragonot, 1888: 49.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

477

DIAGNOSIS. — Eleven pairs of trichobranchiate gills. Rostrum narrowly triangular. Ocular acicles simple or

multifid. Antennal peduncle with supernumerary segmentation. Maxillule with external lobe of endopod somewhat

produced, not recurved. Crista dentata of ischium of third maxilliped without accessory tooth. Dactyls of

ambulatory legs with armed ventral margins. Chelipeds unequal or subequal, right appreciably stouter. Fourth

pereopods semichelate; with single row of scales in propodal rasp. Sternite of fifth pereopods developed as single

small subovate or subquadrate lobe.

Coxa of left fifth pereopod in males (Fig. 3c) with moderately long or long, often weakly spiraled, sexual tube

provided with sparse terminal tuft of stiff setae (Fig. 3d); right fifth coxa with gonopore, occasionally vas deferens

slightly produced, but not developed as distinct sexual tube; 3 uniramous or unequally biramous unpaired left

pleopods. Females with paired gonopores; no paired pleopods, unpaired left pleopods on somites 2 to 5.

Telson with transverse suture only weakly indicated; terminal margins oblique.

Remarks. — Laurentia McLaughlin & Haig, 1996 has proved to be a junior homonym of Laurentia Ragonot,

1888, a lepidopteran genus (cf. Fletcher & Nye, 1984: 80), and as such must be replaced. McLaughlin

and HAIG chose the name Laurentia in recognition of the work of M. DE SAINT LAURENT. In propos¬

ing the replacement name Turleania, an anagram (from Laurentia), the dedication remains unchanged; gender

feminine.

All four originally described species, type species T. albatrossae McLaughlin & Haig, 1966, T. balli

McLaughlin & Haig, 1996, T. sibogae McLaughlin & Haig, 1996, and T. senticosa McLaughlin & Haig, 1996,

were collected in Indonesian waters; however, only T. senticosa is represented in the KARUBAR material.

The generic diagnosis has been broadened to include a fifth species. Turleania multispina sp. nov. is the first

species in the genus with multifid ocular acicles.

Key to the species of Turleania

1. Ocular acicles simple . . . . . 2

— Ocular acicles multifid . T. multispina sp. nov.

2. Right chela with dorsal surface unarmed or with only few scattered spinules . 3

— Right chela with dorsal surface armed with numerous spines or spinules . 4

3. Dactyl of right cheliped with row of spines on dorsomesial margin, dorsodistal margin of

carpus with median spine; an-terior lobe of sternite of third pereopod with 4 marginal

spines . T. balli *

— Dactyl of right cheliped with only few low protuberances on dorsomesial margin,

dorsodistal margin of carpus unarmed; anterior lobe of sternite of third pereopod with

1 marginal spine . T. sibogae*

4. Dorsomesial margin of right chela with row of strong spines; telson with asymmetrical

posterior lobes, each with strong spine at outer angle and 1 or 2 spines on terminal

margins . T. albatrossae*

— Dorsomesial margin of right chela with spinules or irregular rows of small spines; telson

with only slightly asymmetrical posterior lobes, each with acute outer angle and 1 or

2 low protuberances and sparse tufts of setae on terminal margins . T. senticosa

Turleania senticosa (McLaughlin & Haig, 1996)

Figs llh-k

Laurentia senticosa McLaughlin & Haig, 1996: 87, figs 3E, 6

MATERIAL EXAMINED. — Indonesia. KarubaR. Kai Islands: stn DW 18, 05°18'S, 133°01'E, 205-212 m.

24.10.1991: 1 ov. 9 (1.5 mm) (POLIPI). — Stn DW 22, 05°22'S, 133°01'E. 85-124 m, 25.10.1991: 2 <3 (2.1, 2.3 mm)

(MNHN-Pg 5270).

Source :

478

P. A. MCLAUGHLIN

Tanimbar Islands: stn DW 50, 07°59'S, !33°02'E, 184-186 m, 29.10,1991: 1 <J, 2 9, 1 ov. 2 (1.6- 1.8 mm) (USNM

275999, 276000).

DIAGNOSIS. — Shield (Fig. 1 lh) slightly to considerably longer than broad. Rostrum triangular, terminating

bluntly or subacutely. Lateral projections well developed, triangular, with strong marginal or submarginal spine.

Ocular peduncles subcylindrical, 0.65 to 0.80 shield length; overreached by both antennular and antennal peduncles;

corneae dilated. Ocular acicles subtriangular, with small submarginal spine.

Second segment of antennal peduncle with dorsolateral distal angle produced, terminating in acute spine and

sometimes with accessory spine; dorsomesial distal angle with very prominent spine. Antennal acicle reaching to

base of cornea or slightly beyond; terminating in acute spine and with long setae mesially and terminally.

Antennal flagellum moderately short, with 1 or 2 short or long setae every 1 to 4 articles.

Right cheliped moderately long and stout; sometimes with hiatus between dactyl and fixed finger. Dactyl 0.60

to 0.90 length of palm; dorsomesial margin and dorsal midline each with row(s) of very small spines or spinules

and long setae. Palm with dorsomesial margin not clearly delimited, but with irregular rows of spinules or small

spines; dorsal surface convex, armed with small spines or spinules particularly in lateral half and on Fixed finger;

surfaces all with long, but not particularly dense setae. Carpus with row of moderately slender, acute spines on

dorsomesial margin, dorsal surface with 1 row of slightly smaller spines laterad of midline and scattered

small spines or spinules laterally, dorsolateral margin not delimited; surfaces all with long setae. Merus with 1 or

2 prominent spines on ventrolateral margin distally, ventromesial margin with 1 prominent spine at distal angle,

sometimes also 1 additional spine at midlength, and occasionally smaller spine proximally.

Left cheliped not appreciably shorter than right but much less robust. Dactyl unarmed or with few spinules and

tufts of long setae in dorsal midline, low occasionally spinulose protuberances and long setae on dorsomesial

margin. Palm with convex dorsal surface armed with tiny spines or spinules, particularly laterally, and long setae.

Carpus with row of slender spines on dorsolateral and dorsomesial margins, both rows partially obscured by long

setae; ventrolateral margin with acute spine distally. Merus with long setae on all surfaces; ventrolateral margin

with 1 or 2 prominent acute spines distally, and frequently irregular row of smaller spinules on lateral face

ventrally; ventromesial margin with 1 spine near distal angle.

Ambulatory legs similar from left to right. Dactyls slightly to considerably longer than propodi, slender; dorsal

and ventral margins each with row of stiff setae, latter also with 5-8 long corneous spines. Carpi each with

1 spine on dorsal surface adjacent to dorsodistal angle, 1 additional spine on dorsal surface proximally (second).

Mcri each with 1 spine on ventral margin in distal third (second) or unarmed (third). Sternite of third pereopods

with small, subovatc or subtriangular anterior lobe usually with 1 to 4 marginal spines and row of long setae.

Telson (Figs lli-k) with posterior lobes slightly asymmetrical, each outer angle acutely produced or with

terminal spine, 0 to 3 spines on oblique terminal margins.

Color (in preservative). Ocular acicles retain faint orange tint. Chclipeds with some iridescence. Right

cheliped with faint spot of orange near the tip of fixed finger and larger patch proximally. Left cheliped with band

ol light orange proximally on dorsal surfaces of dactyl and fixed finger. Band of light orange present distally on

dactyls of ambulatory legs.

Habitat. — Unknown.

Distribution. — Seram, Kai and Tanimbar Islands, Indonesia; 85-186 m.

Affinities. — McLaughlin and Haig (1996), discussed the similarities between T. senticosa and

T albalrossae- Turleania multispina similarly resembles T. senticosa in having the dorsal surfaces of the chelae

armed with small spines or spinules. The other species, T. balli and T. sibogae both have unarmed chelae. The

specimens of T. senticosa collected during the Karubar expedition all have the spines of the dorsomesial region

of the palm of the right cheliped more strongly developed than in the type series. With that exception, two males

r°m lhe Kai Islands agree exceptionally well with the holotype; the specimens from Tanimbar Islands do not

exhibit the hiatus between the dactyl and fixed finger.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

479

Remarks. — Antennal flagella were missing from the type specimens described by McLaughlin and Haig

(1996). They have been included in the diagnosis presented based upon the present material. The Karubar speci¬

mens of T. senticosa , while agreeing with the type material from Scram Island in most characters, do exhibit some

variations not noted by McLaughlin and Haig. As previously indicated not all of the present specimens have a

hiatus between the dactyl and fixed finger of the right cheliped; in one specimen, the spine on the ventromesial

distal angle of the merus is absent. Similarly, the left cheliped of one specimen has one rather than two spines on

the ventrolateral distal angle. The most noteworthy variation is seen in the telson. In all of present specimens the

posterior lobes are clearly asymmetrical; the oblique margins are armed with two or three spines (Figs 1 Ib-d).

Turleania multispina sp. nov.

Figs 12a-j

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 02, 05°47'S, 132°13'E, 209-240 m.

22.10.1991: I ov. 9 (2.1 mm) (USNM 276003). — Stn DW 03, 05°48'S, 1 32°13'E, 278-301 m. 22.10.1991:

1 6 (1.8 mm) (MNHN-Pg 5271) ; 1 9 (1.9 mm) (POLIPD. — Stn DW 31, 05° 40'S, 132°51'E, 288-289 m. 26.10.1991 :

1 ov. 9 (2.1mm) (MNHN-Pg 5272). —Stn CP 35, 06°08'S, 132°45'E. 390-508 m, 27.10.1991:

1 ov. 9 (2.3 mm) (MNHN-Pg 5273).

Types. — The ovigerous female (2.3 mm) (MNHN-Pg 5273) from Karubar station CP 35 is the holotype.

The other specimens are paratypes.

Description. — Shield (Fig. 12a) longer than broad; anterior margin between rostrum and lateral projections

somewhat concave; anterolateral margins sloping; posterior margin truncate; dorsal surface with few tufts of setae.

Rostrum triangular, well developed, reaching beyond bases of ocular acicles, terminating acutely. Lateral

projections well developed, acutely or obtusely triangular, with submarginal spine.

Ocular peduncles (including corneae) approximately 0.80 shield length; corneae dilated. Ocular acicles

subrectangular, with 3 to 6 terminal spines; separated basally by approximately half basal width of one acicle.

Antennular peduncles, when fully extended, overreaching ocular peduncles (including corneae) by 0.75 to nearly

entire length of ultimate segment. Ultimate segment with 2 to 6 long setae on dorsodistal margin and scattered

shorter setae on dorsal and ventral surfaces. Penultimate segment with few short setae. Basal segment with

statocyst region expanded laterally and dorsoventrally flattened; with 0-3 small spines on protuberances of

dorsodistal margin mesially and 1 stronger spine dorsodistal margin laterally.

Antennal peduncles overreaching ocular peduncles by 0.35 to 0.50 length of ultimate segment. Fifth and fourth

segments with scattered setae. Third segment unarmed or with small ventrodistal spinule. Second segment with

dorsolateral distal angle produced, terminating in acute simple or bifid spine; dorsomesial distal angle with

prominent acute spine. First segment often with small spine at dorsolateral distal angle; 1 to 3 small spines on

ventrolateral distal margin. Antennal acicle reaching beyond base of cornea, but not to distal margin; terminating

in acute spine and with long setae on mesial margin. Antennal flagellum long, overreaching outstretched chelipeds;

with 2 to 4 short (< 2 articles length) setae every 1 to 3 articles in proximal half, fewer longer and irregularly-

spaced setae in distal half.

Right cheliped (Fig. 12b) moderately long and stout. Dactyl slightly less than length of palm; cutting edge

with 2 large calcareous teeth in proximal half, row of very small calcareous teeth distally; terminating in small

corneous claw; dorsomesial margin not delimited, dorsal surface convex, with long setae and scattered small spines

or spinules in proximal third; ventral and mesial surfaces also with scattered long setae. Palm 0.60 to 0.75 length

of carpus; dorsomesial margin not delimited, but sometimes with prominent tubercle at proximal angle; dorsal

surface convex, with scattered small spines and spinules, not extending onto dorsolateral surface or fixed finger,

armature partially to entirely obscured by long simple or plumose setae; dorsal surface of fixed finger also with

numerous long setae; cutting edge with 2 large rather sharp and several small calcareous teeth, terminating in small

corneous claw. Carpus slightly longer than merus; dorsomesial distal angle with acute spine and usually I or

2 smaller spines on dorsomesial margin, dorsal surface with short transverse rows of long setae,

480

P. A. MCLAUGHLIN

Fig. 12. — Turleania multispina sp. nov., holotype 9 (2.3 mm) from Karubar Stn CP 35: a, shield and cephalic

appendages; b, right cheliped; c, left cheliped; d. right second pereopod (lateral view); e, dactyl of left second

pereopod (mesial view); f, left third pereopod (lateral view); g, dactyl of left third pereopod (mesial view); h, dactyl

and propodus of right fourth pereopod (lateral view); i, anterior lobe of sternite of third pereopods; j, telson Scale

equals 0.5 mm (h-j) and 1.0 mm (a-g).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

481

dorsolateral margin not delimited; mesial, lateral and ventral surfaces also with scattered long setae. Merus with

numerous long setae on dorsal margin and mesial and lateral faces; ventrolateral and ventromesial distal angles each

with 1 acute spine, or occasionally only blunt protuberance at ventromesial angle. Ischium with setae mesially and

ventrally. Coxa with spine at ventrolateral distal angle.

Left cheliped (Fig. 12c) slender; usually equaling right in length but less robust. Dactyl approximately equal to

or slightly longer than palm; cutting edge with row of very small corneous teeth, terminating in corneous claw;

dorsal surface convex, unarmed but with numerous long setae. Palm 0.50 to 0.60 length of carpus; dorsal surface

convex, armed with scattered small spines and spinules laterally and on proximal half of fixed finger, both with

numerous long setae; cutting edge of fixed finger with row of small calcareous teeth, terminating in small

corneous claw; ventral surfaces also with numerous long setae. Carpus approximately as long as merus; dorsodistal

margin with 1 mesial acute spine and second in midline, dorsolateral and dorsomesial margins unarmed but with

tufts of long setae; mesial, lateral and ventral faces also with long setae. Merus with long setae on dorsal, lateral

and ventral surfaces; ventrolateral margin with 2 acute spines distally, ventromesial margin with I spine near distal

angle. Ischium with long setae on ventral margin. Coxa with spine at ventrolateral distal angle.

Ambulatory legs (Figs 12d-g) similar from left to right. Dactyls 1.20 to 1.50 length of propodi, slender; in

dorsal view, straight; in lateral view, curved ventrally; terminating in long, slender corneous claws; dorsal margins

each with row of long stiff setae, mesial faces with scattered long setae and 1 to 3 widely-spaced corneous spines

near ventral margin. Propodi with long setae dorsally and ventrally, often arising from low protuberances,

particularly on dorsal surfaces. Carpi each with 1 spine on dorsal surface adjacent to dorsodistal angle, often

1 additional spine on dorsal surface proximally (second). Meri unarmed but with several tufts of moderately long

setae on dorsal and ventral margins. Ischia unarmed. Sternite of third pereopods with small, subquadrate or

subcircular anterior lobe (Fig. 12i) unarmed or with 1 or 2 marginal spinules. Fourth pereopod with 8 or 9 clearly

separated, sharp corneous scales in propodal rasp (Fig. 12h). Fifth pereopods semichelate.

Telson (Fig. 12j) with posterior lobes only slightly asymmetrical, each outer angle prominent, blunt or

subacute; terminal margins oblique and armed with 2 to 4 acute spines.

COLOR (in preservative). — Calcified integument somewhat iridescent. Ventral surfaces of carpi of chelipeds

retaining faint orange tint.

Habitat. — Two of the specimens were inhabiting shells covered by an unidentified bryozoan.

Distribution. — Kai Islands, Indonesia; 209-502 m.

Etymology. — From the Latin spina meaning spined, and refers to the multispined ocular acicles of this

species.

AFFINITIES. — In having the dorsal surfaces of the chelae armed with small spines or spinules, T. multispina

resembles both T. albatrossae and T. senticosa ; however, this new species is immediately distinguished from all

known species of the genus by its multispined ocular acicles.

Genus MICHELOPAGURUS nov.

Pagurodes Henderson, 1888: 94 (in part). — ALCOCK, 1901: 224; 1905b: 106 (in part). — Gordan. 1956: 324 (in pan;

lit.). — de Saint Laurent, 1969: 740 (in part).

1 Pagurodes - Bouvier, 1922: 22. — INGLE, 1993: 102; not Pagurodes Henderson, 1888.

DIAGNOSIS. — Eleven pairs of trichobranchiate gills. Rostrum broadly rounded or obtusely and bluntly

triangular. Ocular acicles simple. Antennal peduncle with supernumerary segmentation. Maxillule (Fig. 13a) with

external lobe of endopod well developed, not recurved. Ischium of third maxilliped with accessory tooth on crista

dcntata. Chelipeds elongate, subequal, right appreciably stouter. Fourth pereopods semichelate; with single row of

scales or rarely incomplete double row in propodal rasp; no distinctive preungual process.

482

P. A. MCLAUGHLIN

Coxae of fifth pereopods in males (Fig. 13b) symmetrical; right, left, or both with short sexual tube;

3 unequally biramous unpaired left pleopods. Females with paired gonopores; paired first pleopods modified as

gonopods, unpaired left pleopods on somites 2 to 5.

Telson with transverse suture; posterior lobes separated by distinct median cleft; terminal margins rounded;

lateral margins, at least left, with corneous plate.

ETYMOLOGY. — The genus is named for Michele de Saint Laurent, who first recognized that

Henderson's (1888) Pagurodes consisted of three distinct taxa.

Type Species. — Pagurodes limatulus Henderson, 1888. Gender masculine.

Remarks. — Henderson (1888) described the genus Pagurodes for three superficially similar Indo-Pacific

species collected during the "Challenger" expedition. Pagurodes inarmatus was based upon eight syntypes;

P. piliferus and P. limatulus were each represented by single specimens, although a second small, poorly preserved

specimen was "doubtfully" referred to P. piliferus. HENDERSON indicated that while the trichobranchiate gills,

presumably, possessed by all three species suggested a relationship with Parapagurus Smith, 1879, Pagurodes was

distinct because males had sexual tubes rather than paired first and second pleopods, modified as gonopods.

When DE Saint Laurent (1969) erected the genus Acanthopagurus de Saint Laurent, she expressed her belief

that HENDERSON'S (1888) species of Pagurodes actually represented three distinct genera. She designated

Pagurodes inarmatus as the type species of HENDERSON'S taxon, thus restricting Pagurodes, and indicated that

P. piliferus and P. limatulus would be separated in a later publication. That separation was never formalized.

Having now reexamined five of HENDERSON'S syntypes of P. inarmatus, including the male specimen from

"Challenger" station 168 upon which he based his description, as well as all "syntypes" of the two other species, it

is clear that DE Saint Laurent (1969) was correct in her evaluation. Pagurodes, as defined by the characters of

the examined syntypes of P. inarmatus, is a plainly identifiable taxon. Although P. inarmatus is not represented in

the Karubar material, it is in the interest of stability in nomenclature that Henderson's (1888) described male

(NHM 88.33) (Figs 15d, 35b-e) be designated as the lectotype of P. inarmatus.

Alcock (1905b) redescribed P. limatulus, pointing out that his female specimen(s) possessed paired first

pleopods modified as gonopods; however, he did not propose a distinct genus for this species. Instead he simply

noted in his generic description that paired first pleopods were present in females of at least one species. When

BouviER (1922) assigned two new abyssal Atlantic species to Pagurodes sensu lato, he remarked that since his

species were both represented by only single specimens, he could not be sure that all characters of the genus were

present. As previously indicated, DE SAINT LAURENT (1969) noted that neither P. limatulus nor P. piliferus were

congeners of P. inarmatus. Of P. limatulus she commented that this species would be assigned to a new taxon,

together with BOUVIER'S (1922) species, Pagurodes richardi Bouvier and Pagurodes atlanticus Bouvier. That

publication was never completed. In his comprehensive treatment of Atlantic pagurids, INGLE (1993) retained

P. richardi and P. atlanticus in Pagurodes (sensu Bouvier, 1922), noting that until DE Saint Laurent's (1969)

opinions could be reevaluated through a thorough study of Henderson's (1888) Indo-Pacific species, the generic

placement of the Atlantic species was uncertain.

McLaughlin (1988) considered the possible relationship of "Pagurodes" limatulus with the Atlantic

Pagurus" piercei Wass, 1963, unlikely, when she proposed the genus Goreopagurus for the latter taxon. The

recent discovery of a second species of Goreopagurus from the eastern Pacific (McLaughlin & Haig, 1995)

confirms the distinctive characters of this genus that set it apart from Henderson's (1888) species.

Pagurodes limatulus is herein reassigned to Michelopagurus gen. nov. as its type species; Pagurodes richardi and

Pagurodes atlanticus are also, provisionally, reassigned to this genus.

Michelopagurus limatulus (Henderson, 1888) new combination

Figs 13a-d, 36a-f

Pagurodes limatulus Henderson, 1888: 97, pi. 10, fig. 6. — Alcock, 1905b: 107, pi. 12, fig. 6. — Estampador. 1937:

507 (list). — Gordan, 1956: 325 (lit.). — de Saint Laurent, 1969: 740. — McLaughlin, 1988: 262.

Pagurodes sp. ? limatulus: Alcock, 1901: 225.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

483

Material EXAMINED. — South of Philippines. " Challenger ", Stn 214, 1414 m: 1 8 holotype (3.5 mm)

(NHM 88.33).

Indonesia. Karubar, Kai Islands: stn CP 20, 05°15'S, 132°59'E, 769-809 m, 25.10.1991: 5 cJ.ll $ , 3 ov. 2

(2. 1-3.6 mm) (MNHN-Pg 5274). — Stn CC 21, 05°14'S, 133°00'E, 688-694 m, 25.10.1991: 5 8,62 (1. 9-3.6 mm)

(USNM 276001). — Stn CP 38, 07°40'S, 132°27'E, 620-666 m, 28.10.1991: 2 8,32 (2.1-4.0 mm) (MNHN-Pg 5275);

2 2 (USNM 275997); I 2 (1.7) (POLIPI); 18,12 (2.9-3.2) (SNHM 4809). — Stn CP 91, 08°44'S, 131°05'E, 884-891

m, 5.11.1991: 1 2 (2.4 mm) (POLIPI).

Diagnosis. — Shield (Figs 13c, 36a, d) broader than long. Rostrum produced beyond level of lateral

projections, broadly rounded. Lateral projections somewhat produced, broadly rounded. Ocular peduncles (including

comeae) very short and stout, less than half length of shield; comeae 0.25 to 0.33 length of peduncles, dilated little

if at all. Ocular acicles triangular, with submarginal spine.

Antennular peduncle with very prominent spine on dorsolateral margin of basal segment. Third segment of an¬

tennal peduncle with very strong spine at ventrodistal angle; second segment with dorsolateral distal angle strongly

produced, terminating in simple or bifid spine and frequently with small secondary spine on mesial margin, dor-

somesial distal angle with acute spine. Sternite of third maxillipeds with prominent spine on either side of

midline.

Right cheliped (Figs 36b, e) moderately elongate. Dactyl slightly shorter than palm; surface with scattered

setae, occasionally unarmed but more frequently with row of tubercles on dorsomesial margin. Palm narrow,

somewhat compressed dorsoventrally; dorsomesial margin tuberculate and with 2 or 3 prominent spines at

proximal margin, convex dorsal surface with short transverse sometimes tuberculate ridges and rows of setae,

usually 1 spine or tubercle in midline at proximal margin. Carpus with row spines on dorsomesial margin, dorsal

surface often with short transverse rows of tubercles and slightly oblique row of small spines, dorsolateral margin

with irregular row of small spines not extending to proximal or distal margins. Merus with irregular single or

double row of spines or spinulose protuberances on ventrolateral margin, most distal usually strongest;

ventromesial margin with row of small spines.

Left cheliped (Figs 36c, 0 with elongate slender unarmed or weakly tuberculate dactyl and fixed finger, tending

to curve ventrally. Palm with midline elevated and armed 1 or 2 double rows of small spinules or tubercles, dorsal

surface spinulose or tuberculate. Carpus with row of spines on both dorsolateral and dorsomesial margins; ventro¬

lateral margin with row of tuberculate spines or spinules. Merus with small spines on distal half of ventromesial

margin; 1 to 3 spines distally and transverse rows of spinules or tubercles in proximal half of ventrolateral margin.

Second and third pereopods similar from left to right. Dactyls 1.20 to 1.35 length of propodi; dorsal margins

each with row of stiff setae, mesial and lateral faces each with longitudinal sulcus, flanked above on mesial faces

by 1 or 2 rows of corneous spinules; ventral margins each with row of 1 1 to 19 corneous spines. Propodi with

low protuberances and tufts of setae on dorsal surfaces, 1 or 2 corneous spinules at ventrodistal margins. Carpi

each with dorsodistal spine and row of low protuberances with tufts of setae on dorsal surface. Meri each with 1 to

3 spines at ventrolateral distal angle and row, sometimes double, of spinules or tubercles on ventral surface

(second) or unarmed (third). Fifth pereopods weakly semichelate. Anterior lobe of sternite of third pereopods

subrectangular, subdivided by median longitudinal groove into two sub-lobes, each with tuft of setae.

Males usually with vas deferens produced as short, almost transparent, sexual tube from both right and left coxa

of fifth pereopods, sometimes from only one, and occasionally hardly produced at all.

Telson (Fig. 13d) with prominent median cleft separating slightly asymmetrical posterior lobes; rounded

terminal margins each with 3 to 7 prominent spines often interspersed with smaller spines.

COLOR (in preservative). — Overall reddish orange.

Habitat. — Gastropod shells sometimes encased in bryozoan.

Distribution. — South of Philippine Islands, Indonesia; ?Travancore coast of India; 620 - 1414 m.

AFFINITIES. — Michelopagurus limatulus shows greater affinity to M. chacei sp. nov. than to either of its

Atlantic congeners, particularly in the armature of the telson and lack of armature of the anterior lobe of the

484

pa. McLaughlin

sternite of the third pereopods. The two Indo-Pacific species are separated by the shorter and more stout ocular

peduncles of M. limatulus and its more elongate and slender chelipeds.

Remarks. — In addition to the species of Henderson (1888) and Bouvier (1922) now assigned to

Michelopagurus , de Saint Laurent (1969) indicated that she had seen three undescribed species among the

Albatross and Siboga collections. As indicated above, one additional species assignable to this genus is present

in the Karubar material, but it is unknown whether it is one of the three seen by DE Saint Laurent.

Michelopagurus chacei sp. nov.

Figs 14a-h, 37c-e

oh >^AJERIAL EXAMINED- — Ind°nesia. Karubar, Kai Islands: stn DW 13, 05°26'S, 132°38'E, 417-425 m

24 10.1991: 1 ov. $ (2.5 mm) (MNHN-Pg 5276). — Stn CP 26, 05°34'S, 132°52'E, 265-302 m, 26.10 199L 1 6

(2.1 mm) (MNHN-Pg 5277).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

485

Types. — The ovigerous female (2.5 mm) (MNHN-Pg 5276) from Karubar station DW 13 is the holotype.

The other specimen is a paratype.

Description. — Shield (Fig. 14a, 37c) as broad or slightly broader than long; anterolateral margins slightly

terraced; posterior margin truncate; dorsal surface with scattered tufts of setae. Rostrum broadly rounded, reaching

approximately to level of lateral projections. Lateral projections strongly produced, obtusely triangular, unarmed.

Ocular peduncles short and stout, dorsal surface with tufts of setae; corneae large, occupying 0.25 to

0.33 length of peduncle, but only weakly dilated. Ocular acicles triangular, terminating subacutely and with strong

submarginal spine; separated basally by approximately 0.75 basal width of one acicle.

Antennular peduncles, when fully extended, overreaching ocular peduncles (including corneae) by 0.25 to

0.50 length of ultimate peduncular segment. Ultimate segment with prominent tuft of long setae on dorsal surface

distally and with row of short setae at least in distal half. Penultimate segment glabrous. Basal segment with acute

spine on dorsolateral margin.

Antennal peduncle overreaching ocular peduncles by nearly full length of ultimate segment. Fifth and fourth

segments with few scattered setae. Third segment with spine at ventrodistal margin. Second segment with

dorsolateral distal angle produced, terminating in strong simple or bifid spine and sometimes with small accessory

spine on mesial margin; dorsomesial distal angle with prominent spine. First segment with spine at laterodistal

margin and 1 spine on ventrodistal margin. Antennal acicle quite long, reaching to distal half of fifth peduncular

segment; terminating in acute spine and tuft of setae; mesial margin with few setae. Antennal flagellum long but

usually not overreaching tips of dactyls of outstretched ambulatory legs; with 1 or 2 very few short setae every

2 to 4 articles.

Chelipeds subequal; right usually slightly longer and stouter. Right cheliped (Figs 14b, 37d) elongate,

moderately slender. Dactyl 1.10 to 1.20 length of palm; dorsomesial margin with row of very small spines, dorsal

surface unarmed but with few scattered setae; mesial and ventral surfaces with scattered of setae; cutting edge with

2 widely-spaced calcareous teeth in proximal half, row of corneous teeth distally. Palm slightly compressed

dorsoventrally; 0.75 length of carpus; dorsomesial margin with irregular row of small spines, decreasing in size

distally, dorsolateral margin not delimited; dorsal, mesial and lateral surfaces unarmed but with scattered short

setae; fixed finger unarmed; cutting edge with 2 large and distal row of small calcareous teeth. Carpus slightly

broadened distally; dorsomesial margin with row of prominent spines, dorsolateral margin distinct proximally and

armed with irregular row of very small spinulose tubercles; dorsal surface with double longitudinal row of very

small spines laterad of midline and numerous very small spines forming short quasi-transverse rows in proximal

half; mesial and lateral faces with sparse tufts of setae; ventrolateral distal angle with small spine. Merus with

transverse rows of setae dorsally; ventromesial distal angle with 1 spine and few marginal spinules; ventrolateral

margin with 2 spines distally and spinules proximally. Ischium with ventromesial margin spinulose proximally.

Left cheliped (Figs 14c, 37e) only slightly shorter than right. Dactyl and fixed finger long and slender,

somewhat dorsoventrally compressed, with tips deflected ventrally ; margins of dactyl and fixed finger not delimited,

rounded surfaces smooth. Palm markedly shorter than both dactyl and carpus; dorsal surface elevated in midline and

armed with row of small spines not extending onto fixed finger, dorsomesial margin with row of small spines on

spinulose tubercles; dorsolateral margin not delimited, lateral surface with few small spinules dorsally. Carpus

with dorsolateral and dorsomesial margins each with row of spines, 1 spine on dorsodistal margin; ventrolateral

distal angle with 1 or 2 small spines, ventral surface spinulose. Merus with transverse rows of setae on dorsal

margin; ventrolateral margin with row of spines, ventromesial margin with row of very small spines or spinules.

Ischium unarmed.

Ambulatory legs (Figs 14d-e) similar from left to right; moderately long and slender; overreaching outstretched

chelipeds by nearly entire length of dactyls. Dactyls relatively long, 1.10 to 1.25 length of propodi; nearly

straight; dorsal surfaces each with row of stiff setae; mesial and lateral faces with few setae; ventral margins each

with 9 to 1 1 corneous spines. Propodi each with irregular row of setae on dorsal surface; mesial and lateral faces

unarmed; ventral surfaces with few setae and 1 or 2 corneous spines at distal angle. Carpi each with small

dorsodistal spine and dorsal row of sparse tufts of setae. Meri usually with 1 or 2 spinules distally on ventral

margins of second, third unarmed but with few dorsal and ventral setae. Ischia unarmed. Sternite of third pereopods

486

P. A. MCLAUGHLIN

Fig. 14. — Michelopagurus chacei sp. nov. a, f, h. holotype 6 (2.5 mm) from Karubar Sin DW 13; b-e, g, paratype 6

(2.1 mm) from Stn CP 26; a, shield and cephalic appendages; b, right cheliped; c, left cheliped; d, second right

pereopod (lateral view); e, third left pereopod (lateral view); f. anterior lobe of sternite of third pereopods; g. coxae

and stemite of fifth pereopods; h, telson. Scales equal 0.5 mm (f, h), 1.0 mm (g), and 2.0 (a-e).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

487

with anterior lobe (Fig. 14f) broadly subrectangular, concave as result of median depression, unarmed but with

submarginal tuft of setae adjacent to each lateral angle. Fifth pereopods chelate.

Coxae of fifth pereopods in males (Fig. 14g) with short right sexual tube and even shorter left.

Telson (Fig. 14h) with well defined transverse suture; posterior lobes subcircular to subtriangular, median cleft

prominent; terminal margins slightly oblique or rounded, armed with 3 to 5 spines interspersed with much smaller

slender spines or spinules.

Color. — Unknown.

Habitat. — Unknown.

Distribution. — Kai Islands, Indonesia; 264-425 m.

ETYMOLOGY. — Dedicated to Dr Fenner A. CHACE, Zoologist Emeritus of the Division of Crustacea,

National Museum of Natural History, Smithsonian Institution, who, during a continuing long and active career

has contributed so much to decapod systematics.

Affinities. — As previously noted, Michelopagurus chacei bears a considerable resemblance to M. limatulus,

particularly in the structure of the anterior lobe of the sternite of the third pereopods and development of the telson

armature. In addition to the distinguishing characters of M. limatulus mentioned above, the shield of M. chacei

tends to be slightly broader and the antennular flagella longer than seen in M. limatulus ; however, these characters

may well be subject to intraspecific variation when large samples are available for examination. The ambulatory

legs appear to provide the best characters for separating the two species. A row of spinules or tubercles is present

on the ventral margins of the meri of the second pereopods of M. limatulus, and the ventral margins of the dactyls

are provided with 11 to 19 corneous spines. In M. chacei , the meri of the second pereopods have only one or two

distal spinules on the ventral margins; the ventral margins of the dactyls are armed with nine to 1 1 corneous

spines.

Genus PSEUDOPAGURODES nov.

Pagurodes Henderson, 1888: 94 (in part). — ALCOCK, 1901: 224; 1905b: 106 (in part). — Gordan, 1956: 324 (in part;

lit.). — de Saint Laurent, 1969: 740 (in part).

DIAGNOSIS. — Eleven pairs of intermediate type gills. Rostrum reduced and rounded. Ocular acicles small,

widely separated. Maxillule with external lobe of endopod well developed, not recurved. Third maxilliped with well

developed crista dentata and 1 accessory tooth. Chelipeds subequal. Dactyls of ambulatory legs without corneous

spinules on ventral margins. Fourth pereopods semichelate; propodal rasp with single row of scales; no preungual

process at base of dactylar claw. Fifth pereopods chelate.

Male unknown. Female with paired gonopores, no paired pleopods on first abdominal somite, and 4 unequally

biramous left pleopods on somites 2 to 5.

Asymmetrical uropods. Telson with transverse suture, subtriangular posterior lobes separated by median cleft.

ETYMOLOGY. — From the Greek pseudes meaning false, and pagouros meaning crab, reflecting the deceptive

similarities between this genus and Pagurodes sensu stricto.

Type Species. — Pagurodes piliferus Henderson, 1888. Gender masculine.

REMARKS. — With the restriction of Pagurodes to taxa exhibiting the characters manifest by P. inarmatus and

the establishment of Michelopagurus gen. nov. for P. limatulus , Pagurodes piliferus is left in a systematic limbo.

The damaged male specimen from the Arafura Sea that HENDERSON (1888) doubtfully assigned to P. piliferus was

subsequently redetermined to be Pagurus compressipes Miers, 1884, by M. de Saint Laurent (unpublished).

I concur with her identification. The remaining, and true type specimen of Pagurodes piliferus (NHM 88.33) is a

female (Figs 13e-f, 37a-b) with intermediate gills, thus clearly not allied to Pagurodes sensu stricto, nor to

488

P. A. MCLAUGHLIN

Michelopagurus gen. nov. In gill structure Pseudopagurodes approaches Tarrasopagurus gen. nov., but differs from

that genus in having subcqual, elongate chelipeds and ambulatory legs, while lacking paired first pleopods. No

specimens of Pseudopagurodes were found during the KARUBAR expedition.

Genus ICELOPAGURUS nov.

Diagnosis. — Eleven pairs of phyllobranchiate gills. Rostrum triangular. Ocular acicles triangular, elongate.

Antennal peduncle with supernumerary segmentation. Maxillule (Fig. 15a) with external lobe of endopod

rudimentary or vestigial. Crista dentata of third maxilliped somewhat reduced, but with accessory tooth. Chelipeds

elongate, subequal, right stouter. Sternal plate of third pereopods broad, with weak longitudinal groove. Fourth

pereopods semichelate; with single row of spiniform scales in propodal rasp; dactyl with tubular preungual process

(Fig. 15b).

Coxae of fifth pereopods in males (Fig. 15c) symmetrical, right with stout, relatively short sexual tube directed

posteriorly and externally, left usually with very short sexual tube; 3 unequally biramous unpaired left pleopods.

Females with paired gonopores; without paired first pleopods modified as gonopods, unpaired left pleopods on

somites 2 to 5.

Telson with transverse suture; rounded posterior lobes separated by distinct median cleft with nearly

perpendicular margins, terminal margins armed; lateral margins with very narrow chitinous plate.

Type Species. — Icelopagurus crosnieri sp. nov. Gender masculine.

Etymology. — From the Greek ikelos meaning like or resembling, and pagouros meaning crab, and referring

to the similarities shared with another pagurid genus, namely, Catapagurus.

Remarks. — As indicated in the derivation ol its name, Icelopagurus is superficially very similar to

Catapagurus. Both genera are characterized by unusually elongate ocular acicles, rudimentary or vestigial external

endopodal lobe on the maxillule, more or less reduced crista dentata with one accessory tooth, distinctive tubular

preungual process on the dactyl of the fourth pereopod, well developed right male sexual tube, and females lacking

specialized secondary sexual characters. However, Icelopagurus is readily separated from Catapagurus by the

shortness of the male sexual tube that does not curve up over the dorsal surface of the body and by the very

distinctive development of the telson.

Icelopagurus is also ostensibly quite similar to Pagurodes sensu stricto. As presently known, both taxa are

monotypic, and their type species share such immediately observable characters as elongate chelipeds and

ambulatory legs, well calcified shields that tend to be somewhat vaulted, long antennal acicles, triangular rostra,

very short, stout ocular peduncles provided with low protuberances and tufts of setae, and males with a short sexual

tube arising from the coxa of the right fifth pereopod and an even shorter tube from the left (Figs 15c-d). However,

upon closer inspection, the ocular acicles of Pagurodes inarmatus (Fig. 35b) are short, reaching only to the basal

portion of the peduncles; the external lobe of the maxillulary endopod is completely absent; the gills are

trichobranchiate; the dactyl of the fourth pereopod lacks a preungual process; and the posterior lobes of the telson

are acutely subtriangular with strongly oblique terminal margins armed with 3 or 4 small spines. In contrast,

/. crosnieri sp. nov. has very long ocular acicles (fig. 38b), reaching to or beyond the bases of the corneae; the

external lobe of the maxillulary endopod is vestigial; the gills are phyllobranchiate; the dactyl of the fourth

pereopod is provided with a large tubular preungual process; and the posterior lobes of the telson are broadly

rounded, with convex terminal margins armed with 4 or 5 widely-spaced long corneous spines.

Icelopagurus crosnieri sp. nov.

Figs 15a-c, e-j, 38a-d

, . MATERIAL EXAMINED. — Indonesia. Karubar. Tanimbar Islands: stn CP 87, 08°49'S, 130°49'E, 1017-1024 m

?mmhn p «™2lT!1<nU; SNM 276004)' ~ Stn cp 91, 08°44'S, 131°05'E, 884-891 m. 5.11.1991: 1 ov. 9 (4.3 mmj

(MNHN-Pg 5278) ; 1 8,2 9, 1 ov. 2 (3. 1-4.3 mm) (MNHN-Pg 5279); 1 2 (3.2 mm) (POLIPI).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

489

Types. — The ovigerous female (4.3 mm) (MNHN-Pg 5278) from Karubar station CP 91 is the holotype.

The other specimens are paratypes.

Description. — Shield (Figs 15e; 38a-b) broader than long, well calcified, slightly vaulted; cervical groove

deep; anterior margin between rostrum and lateral projections concave; anterolateral margins sloping; posterior

margin truncate; dorsal surface with numerous tufts of setae. Posterior carapace with patches of calcification

medianly posterior to cervical groove, remainder membraneous. Rostrum triangular, terminally subacute or acute,

usually not reaching level of lateral projections. Lateral projections strongly produced, acutely or obtusely

triangular, with prominent terminal marginal or submarginal spine.

Ocular peduncles very short and stout, dorsal surface with tufts of setae; corneae large, occupying 0.25 to

0.33 length of peduncle, but not dilated. Ocular acicles slender, elongate, length variable (extending from

0.25 length of peduncle to nearly base of cornea), terminating acutely or with distinct spine; separated basally by

entire to 1 .25 basal width of one acicle.

Antennular peduncles, when fully extended, overreaching ocular peduncles by slightly less to slightly more

than entire combined lengths of ultimate and penultimate peduncular segments. Ultimate segment with prominent

tuft of long setae on dorsal surface distally and with additional row of short setae. Penultimate segment with sparse

row of short setae dorsally. Basal segment unarmed.

Antennal peduncle overreaching ocular peduncles by at least 0.25 length of penultimate segment. Fifth and

fourth segments with few scattered setae. Third segment with small spine at ventrodistal margin. Second segment

with dorsolateral distal angle produced, terminating in simple or bifid spine and with small accessory spine on

lateral margin; dorsomesial distal angle with small spine. First segment with spine at laterodistal margin and

1 spine on ventrodistal margin. Antennal acicle quite long, reaching to or beyond distal margin of Fifth peduncular

segment; terminating in acute spine; mesial margin with row of long setae. Antennal flagellum long but usually

not overreaching tips of dactyls of outstretched ambulatory legs, naked or with very few short setae on proximal

articles. Sternite of third maxillipeds with median concavity, unarmed.

Chelipeds subequal; right usually slightly longer and stouter. Right cheliped (Figs 15f, 38c) with dactyl 0.65

to 0.80 length of palm; dorsomesial margin rounded, unarmed or minutely spinulose, dorsal surface with scattered

moderately long setae and row of shorter setae adjacent to cutting edge; ventral surface also with few tufts of setae;

cutting edge calcareous, with 2 prominent teeth. Palm somewhat dorsoventrally compressed; slightly shorter to

approximately equal to length of carpus; dorsomesial and dorsolateral margins not well defined; dorsal, mesial and

lateral surfaces covered with very small spinules or tubercles; Fixed finger minutely spinulose on proximal portion

of dorsal surface; cutting edge with 2 or 3 large and distal row of small calcareous teeth. Carpus appreciably

broadened distally; dorsomesial and dorsolateral margins distinct distally, each armed with irregular double row of

small spines or spinulose tubercles; dorsal surface with numerous small spines; mesial and lateral faces with

spinules or tubercles dorsally, minutely spinulose or granular ventrally. Merus with all surfaces uniformly

spinulose or tuberculate. Ischium with minutely spinulose ventromesial margin.

Left cheliped (Fig. 38d) only slightly shorter than right. Dactyl and fixed Finger long and slender, somewhat

dorsoventrally compressed, with tips deflected ventrally; margins of neither dactyl nor fixed finger delimited,

rounded surfaces smooth or minutely spinulose. Palm markedly shorter than both dactyl and carpus; dorsal surface

weakly convex, dorsomesial and dorsolateral margins not delimited; all surfaces spinulose or tuberculate, ventral

surface minutely so. Carpus with dorsal surface relatively flat, covered with small spines; mesial, lateral and

ventral surfaces spinulose. Merus with dorsal, lateral and ventral surfaces uniformly spinulose; mesial face nearly

smooth. Ischium with minutely spinulose ventral margin.

Ambulatory legs (Figs 15g-h) similar from left to right; slender and very long, overreaching outstretched

chelipeds by half to entire length of dactyls. Dactyls long, very slender, ventrally curved, and slightly twisted

distally; approximately 1.25 length of propodi; dorsal surfaces each with row of low protuberances and long stiff

setae; mesial faces each with longitudinal row of shorter setae. Propodi each with irregular double row of small

spines and short setae on dorsal surface; mesial, lateral and ventral surfaces usually spinulose. Carpi with single or

double row of small spines on dorsal surface; mesial and lateral faces spinulose, at least dorsally. Meri with

spinulose dorsal, lateral and ventral surfaces, spinules strongest ventrally; mesial faces smooth or minutely

spinulose. Ischia with minutely spinulose ventral margins. Fifth pereopods weakly chelate.

490

P. A. MCLAUGHLIN

Fig. 15. — Icelopagurus crosnieri sp. nov, a, c, g-j, paratype 8 (3.1 mm) from Karubar Stn CP 91; b. paratype 2

(5.2 mm) from Stn CP 87; e, holotype 8 (4.3 mm) from Stn CP 91; f, paratype 2 (4.3 mm) from Stn CP 91

— d, Pagurodes inarmatus, leciotype 8 (7.0 mm) from "Challenger" Stn 168: a. maxillule; b. distal portion of

dactyl of fourth pereopod; c-d, sternite and coxae of fifth pereopods; e. shield and cephalic appendages; f. right

cheliped (mesial view); g, dactyl of right second pereopod (mesial view); h, left third pereopod (lateral view)-

i. sternite of third pereopods; j, telson. Scales equal 0.25 mm (b), 0.5 mm (j), 1.0 mm (a. c, i), and 3.0 mm (d-h).

Sternite of third pereopods (Fig. 15i) broad; anterior lobe subrectangular, unarmed but with marginal row of

setae.

Coxae of fifth pereopods in males (Fig. 15c) with right sexual tube short and moderately thick basally, curving

posteriorly and externally; small protuberance of left vas deferens.

Telson (Fig. 15j) with prominent transverse suture; posterior lobes obliquely rounded and chitinous lateral plate

armed with 4 to 6 long corneous spines; median cleft prominent, margins slightly oblique and armed with 0 to

2 small spines and often 1 more prominent at outer angle.

COLOR. — Unknown.

Habitat. — Gastropod shells.

Source : MNHN Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

491

Distribution. — Tanimbar Islands, Indonesia; 884-891 m.

Etymology. — Dedicated to Alain Crosnier, marine biologist of ORSTOM, who made the collection

available for study.

AFFINITIES. — Icelopagurus crosnieri most closely resembles Catapagurus oculocrassus sp. nov. in overall

morphology, but is immediately distinguished from that species by the major differences in the telsons of the

two species. As noted in the remarks for the genus, /. crosnieri also shares several general characters with

Pagurodes inarmatus , but differs significantly from the latter species in the spination of the chelipeds, as well as

in the numerous characters cited above.

Genus TARRASOPAGURUS nov.

DIAGNOSIS. — Eleven pairs of intermediate gills. Rostrum obtusely triangular or broadly rounded, with 1 or

more marginal spinules. Ocular acicles triangular. Antennal peduncle with supernumerary segmentation. Maxillule

(Fig. 16a) with external lobe of endopod moderately well developed, not recurved. Crista dentata well developed,

1 accessory tooth. Chelipeds markedly unequal, right considerably longer and stronger. Fourth pereopods

semichelate; with single row of scales in propodal rasp.

Coxae of fifth pereopods in males (Fig. 16b) generally symmetrical, left with short sexual tube directed

anteriorly or posteriorly, right sometimes also with short tube developed, sometimes with vas deferens only

slightly protruded; 3 unpaired, unequally biramous pleopods. Females with paired gonopores; paired first pleopods

modified as gonopods, unpaired left pleopods on somites 2 to 5.

Telson with transverse suture; posterior lobes subequal, terminal margins oblique.

ETYMOLOGY. — From the Greek tarraso meaning confused, and pagouros meaning crab, and referring to the

characters of this genus shared with several other pagurid genera.

Type Species. — Tarrasopagurus rostrodenticulatus sp. nov. Gender masculine.

REMARKS. — As its etymology indicates, Tarrasopagurus shares a number of characters with several other

pagurid genera. In having intermediate type gills (Fig. 16c), markedly unequal chelipeds and relatively short

ambulatory legs, it agrees with Cestopagurus as redefined by DE Saint Laurent (1968c). It is distinguished from

species of Cestopagurus by the presence in the latter genus of an elongate right sexual tube, which is directed from

right to left across the ventral part of the body and a very short left sexual tube that may or may not be developed.

Intermediate gills are also a character that Tarrasopagurus shares with Pseudopagurodes, but females of that genus

lack the paired first pleopods found in Tarrasopagurus species; males of Pseudopagurus are not presently known.

Short paired male sexual tubes is a character that Tarrasopagurus shares with Parapagurodes McLaughlin & Haig,

1973; but again, females of this latter genus lack paired first pleopods modified as gonopods. A character that

Tarrasopagurus shares with Michelopagurus nov. gen., in addition to short or very short male sexual tubes, is

paired female first pleopods. However, the gills in Michelopagurus are clearly trichobranchiate in structure,

whereas those of Tarrasopagurus are intermediate. The chelipeds in this genus are grossly unequal; the ambulatory

legs are relatively short, overreaching the outstretched right cheliped little if at all. In contrast, the chelipeds of

Michelopagurus species are elongate and subequal; the ambulatory legs are long and slender, and considerably

overreach the outstretched chelipeds.

Tarrasopagurus rostrodenticulatus sp. nov.

Figs 16a-l

Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn CP 05, 05°49'S, 132°18'E. 296-299 m.

22.10.1991: 1 6 (2.7 mm) (MNHN-Pg 5280). — Stn DW 13, 05°26'S. 132°38'E. 417-425 m. 24.10.1991: 4 2 (0.9-

492

p.a. McLaughlin

1.2 mm) (MNHN-Pg 5281). — Sin DW 18, 05°17'49"S, 133°00'51"E, 205-212 m, 24.10.1991: 4 <3, 4 2 (1. 6-1.8 mm)

(MNHN-Pg 5282); 1 <3,2 $ (1.0-1. 1 mm) (POLIPI); 2 <3,4 2, 1 ov, 2 (1.3-1. 7 mm).(USNM 276011). — Stn DW 31,

05°40'S, 132°51'E, 288-289 m, 26.10.1991: 1 <3 (1.6 mm) (MNHN-Pg 5283). — Stn CP 36, 06°05'S, 132°44'E, 268-

210 m, 27.10.1991: 1 2 (1.9 mm) (USNM 276010).

Types. — The male (2.7 mm) (MNHN-Pg 5280) from Karubar station CP 05 is the holotype. The other

specimens are paratypes.

DESCRIPTION. — Shield (Fig. 16d) longer than broad; anterior margin between rostrum and lateral projections

weakly concave; anterolateral margins terraced; posterior margin truncate; dorsal surface with few tufts of setae.

Rostrum broadly rounded or weakly subtriangular, usually not reaching level of lateral projections, armed with 1 to

several small marginal spines or spinules. Lateral projections broadly rounded or obtusely triangular, usually with

small terminal marginal or submarginal spine.

Ocular peduncles short and stout; corneae occupying approximately 0.15 to 0.25 length of peduncle, not

dilated. Ocular acicles small, triangular, with submarginal spine; separated basally by 0.75 to entire basal width of

one acicle.

Antennular peduncles, when fully extended, overreaching ocular peduncles (including corneae) by slightly less

to slightly more than entire length of ultimate peduncular segment. Ultimate segment with 1 long seta on dorsal

surface in distal third, and with additional row of sparse short setae. Penultimate segment with few short setae

dorsally. Basal segment with strong spine on dorsolateral margin.

Antennal peduncle overreaching ocular peduncles by 0.50 to 0.75 length of ultimate segment. Fifth and fourth

segments with few scattered setae. Third segment with small spine at ventrodistal margin. Second segment with

dorsolateral distal angle produced, terminating in bi- or trifid spine and usually with 1 or 2 smaller accessory spines

on both lateral and mesial margins; dorsomesial distal angle with small spine. First segment with spine at

laterodistal margin and 1 or 2 spines on ventrolateral margin. Antennal acicle long, reaching to or beyond distal

margin of cornea; terminating in acute spine and tufts of moderately long setae. Antennal flagellum long but

usually not overreaching tips of dactyls of outstretched ambulatory legs; with 1-3 moderately long and I or 2 very

short setae every, or every other, article. Sternite of third maxillipeds with slight median concavity, unarmed.

Chelipeds markedly unequal; right longer and stronger. Right cheliped (Figs 16e-f) with dactyl 0.65 to 0.80

length of palm; armature variable: dorsomesial margin rounded and unarmed, delimited but unarmed, or delimited

by row of small spines; dorsal surface convex or with distinct median elevation, unarmed or with scattered small

spinules or tubercles; ventral surface with few tufts of setae; cutting edge calcareous, usually with 2 more

prominent teeth. Palm somewhat dorsoventrally swollen; slightly to considerably shorter than carpus; dorsomesial

and dorsolateral margins frequently not well defined, often armed with 1 or 2 irregular rows of low tubercles,

strongest mesially, less commonly with irregular, almost double row of distinct spines on dorsomesial margin and

row of blunt spines or spinulose tubercles on dorsolateral margin extending nearly to tip of fixed finger; dorsal

surface with few very small tubercles, with scattered small tubercles and spinules, or with numerous small spines;

mesial, lateral and ventral surfaces with few scattered setae; fixed finger sometimes with few spinules or low

tubercles on dorsal surface; cutting edge calcareous, also usually with 2 or 3 larger teeth. Carpus somewhat longer

than merus; with double or triple row of small spines on and/or adjacent to dorsomesial margin; longitudinal row

of small spines or spinules on dorsal surface laterad of midline, dorsolateral margin not delimited, but dorsal

surface laterally and extending onto lateral faces usually armed with scattered spinules or tubercles; ventrolateral

margin usually with few spinules; mesial surface generally glabrous; ventral surface with few long setae. Merus

with few very short setae on dorsal surface; ventrolateral margin with row of 4 or 5 slender spines, ventromesial

margin with 1 or 2 spines distally. Ischium unarmed.

Left cheliped (Figs 16g-h) reaching to or slightly beyond base of dactyl of right; with or without hiatus

etween dactyl and fixed finger. Dactyl 0.25 to 0.33 longer than palm; dorsomesial margin not delimited dorsal

mesial and ventral surfaces with few scattered setae. Palm 0.50 to 0.75 length of carpus; dorsal surface elevated in

midhne and armed with row of spines usually extending at least to proximal half of fixed finger; dorsomesial

surface weakly to strongly sloping, with longitudinal row of spines not clearly marginal; dorsolateral surface very

strongly sloping, usually with 1 longitudinal row of small spinules and scattered spinules or tubercles; ventral

PAGURIDAE FROM THE KARUBAR EXPEDITION

493

Fig. 16. — Tarrasopagurus rostrodenticulatus sp. nov., a, c, f. h, paratype 6 (1.5 mm) from KARUBAR Stn DW 18;

d, holotype 6 (2.7 mm) from Stn CP 05; b, e, g, i-1, paratype 6 (1.5 mm) paratype, from Stn DW 18: a, right

maxillule; b, coxae and sternite of fifth pereopods; c, intermediate arthrobranchiate gill lamella; d, shield and

cephalic appendages; e, right cheliped (dorsal view); f, carpus and chela of right cheliped (dorsal view); g. left

cheliped (dorsal view); h. carpus and chela of left cheliped (dorsal view); i, second right pereopod (lateral view);

j, third left pereopod (lateral view); k, propodus and dactyl of left fourth pereopod (lateral view); 1, telson. Scales

equal 0.25 mm (a, c), 0.5 mm (b, k, 1), and 1.0 mm (d-j).

494

P. A. MCLAUGHLIN

surface with scattered setae. Carpus approximately as long as merus; dorsomesial and dorsolateral margins each

with row of spines; ventrolateral margin with 2 or 3 spines distally. Merus with row of acute spines on

ventrolateral margin, ventromesial margin with 3 or 4 widely-spaced small spines or spinules. Ischium unarmed.

Ambulatory legs (Figs 1 6i-j) similar from left to right. Dactyls equal to or only very slightly longer than

propodi; very slightly curved, not twisted; dorsal surfaces each with row of sparse tufts of setae; ventral margins

each with row of 6 to 9 strong corneous spines. Propodi with tufts of short setae on dorsal surfaces; ventrodistal

angles each with corneous spine, ventral surfaces each with row of widely-spaced corneous spinules. Carpi each

with tiny spinule at dorsodistal angle, dorsal surfaces with few tufts of setae. Meri and ischia unarmed. Sternite of

3rd pereopods with large semicircular anterior lobe, unarmed but with marginal row of setae. Fourth pereopod with

moderately elongate dactyl (Fig. 16k). Fifth pereopods chelate.

Coxae of fifth pereopods in males (Fig. 16b) with short left and frequently also short right sexual tube.

Telson (Fig. 161) with prominent transverse suture; posterior lobes with chitinous lateral margins, median cleft

prominent; terminal margins oblique, with 4 or 5 spines, largest usually at outer angle.

COLOR (in preservative). — Ocular peduncles with orange tint, darker basally. Ambulatory legs with distal

halves of dactyls and propodi white, proximal halves orange; carpi each with longitudinal orange stripe dorsally and

laterally; meri each with faint orange band on lateral face distally.

Habitat. — Gastropod shells.

Distribution. — Kai Islands, Indonesia; 205-425 m.

Etymology. — The specific epithet reflects the unusual denticulate rostral margin of this species.

Affinities. — At first glance, the small size and general shape and armature of the chelipeds and ambulatory

legs of Tarrasopagurus rostrodenticulatus might suggest a relationship with species of Pagurixus Melin, 1939.

However, where males of Pagurixus species have the right gonopore masked by a mesially directed tuft of stiff

setae, males of Tarrasopagurus have developed sexual tubes, albeit quite short; females possess paired first

pleopods modified as gonopods. Females of Pagurixus have no special sexual modifications, although not

uncommonly only one gonopore develops. The denticulate rostral margin of T. rostrodenticulatus immediately sets

it apart from all other known species.

Remarks. — Two very small specimens (0.8, 1.0 mm) (POLIPI) from Stn DW 18, appeared to be juvenile

males ot this species, but neither had either sexual tube developed, nor could any denticulations be seen on the

rounded rostra. Neither are considered paratypes. Within the size range of 1.0 to 1.5 mm (shield length),

denticulations on the rostral margin are very tiny and difficult to observe; however, with increased animal size

these denticulations become quite obvious.

Genus CATAPAGURUS A. Milne Edwards, 1880

Catapagurus A. Milne Edwards, 1880: 46. — Smith, 1882: 14. — Henderson, 1888: 75. — A. Milne Edwards &

Bouvier, 1893: 125. — Alcock, 1905b: 1 14. — Forest & de Saint Laurent, 1968: 151. — de Saint Laurent.

1970a: 1456. — Miyake, 1978: 141.

Hemipagurus Smith, 1881: 422.

Diagnosis. — Eleven pairs of phyllobranchiate gills. Shield with rostrum weakly developed. Ocular acicles

simple, slender, elongate. Antennal peduncle with supernumerary segmentation. Maxillule with external lobe of

endopod rudimentary. Ischium of third maxilliped with crista dentata more or less reduced, 1 accessory tooth.

Sternite of third maxillipeds unarmed.

Chelipeds long and slender; right stronger. Ambulatory legs very long and slender; dactyls and propodi similar.

Fourth pereopods semichelate; propodal rasp consisting of 1 row of scales; distinctive preungual process present.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

495

Males with right sexual tube orientated toward exterior then recurved over anterior part of abdomen; 2 (pleopods

3 and 4) or 3 (pleopods 3-5) unpaired, usually uniramous pleopods. Females with paired gonopores; without paired

first pleopods modified as gonopods, with pleopods 2 to 4 biramous, pleopod 5 similarly present or absent.

Telson with transverse suture; posterior lobes subtriangular; terminal margins oblique.

Remarks. — In her revision of Catapaguroides and Cestopagurus, DE Saint Laurent (1968a) transferred

three species from Catapagurus to other existing genera, including C. vallatus Melin, 1939, to Nematppagurus.

However, she did not consider Catapagurus australis Henderson, 1888. Despite HENDERSON'S correct diagnosis of

the generic characters of Catapagurus, a recent rexamination of the syntypes of C. australis (NHM 1888.33) have

shown that the structure of the sexual tubes and the weak development of the ocular acicles provide convincing

evidence that this species too should have been assigned to Nematopagurus.

Even with the transfer of the three species aforementioned from Catapagurus, Forest and DE Saint Laurent

(1968) and de Saint Laurent (1970a) indicated that the genus still included about ten Indo-Pacific species,

of which several remained to be described. It is unfortunate that the revision promised by de Saint Laurent

(1970a) was never completed. Not only there are serious inconsistencies in the diagnoses that have been presented

by several authors cited in the above synonymy, but major problems in species interpretations prevail as well,

which are beyond the scope of the KARUBAR study. HAIG and BALL (1988) reported the occurrence of a

species they believed to represent Catapagurus ensifer Henderson, 1893, and a second "probably ... undescribed"

species from the Maluku area, noting that the lack of an up-to-date revision of the genus made identification

difficult. The new species described herein are assigned to Catapagurus as defined by Forest and DE Saint

Laurent (1968) and DE Saint Laurent (1970a), until such time as a thorough review of the genus can be

completed.

Key to the Indonesian species of Catapagurus

1 . Dactyls of ambulatory legs spatulate or blade-shaped . 2

— Dactyls of ambulatory legs not spatulate or blade-shaped . 3

2. Telson with roundly triangular posterior lobes separated by very broad median cleft .

. C. ensifer*

— Telson with acutely triangular posterior lobes separated by moderately narrow median cleft

. Catapagurus sp. of Haig & Ball*

3. Telson with broad median cleft, terminal margins of posterior lobes oblique and armed

with 1 to 4 small spines; ocular peduncles very short and stout, ocular acicles usually

reaching beyond midpoint of peduncle . C. oculocrassus sp. nov.

— Telson with narrow median cleft, terminal margins of posterior lobes perpendicular or

nearly so, unarmed or with only few very short bristles; ocular peduncles not extremely

short, ocular acicles not reaching beyond midpoint of peduncle . 4

4. Mesial faces of dactyls of ambulatory legs each with ventral row of corneous spines .

. C. tanimbarensis sp. nov.

— Mesial faces of dactyls of ambulatory legs without ventral row of corneous spines .

. C. holthuisi sp. nov.

Catapagurus oculocrassus sp. nov.

Figs 17a-k, 39a-b

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands : stn CP 20. 05°15'S, 132°59'E. 769-809 m,

25.10.1991: 2 <3 , 3 $ , 2 ov. 9 (2.7-3.7 mm) (MNHN-Pg 5284). — Stn CP 21, 05°14'S, 133°00'E, 688-694 m.

25.10.1991: 2 6 , 6 9 , 4 ov. 9 (1. 5-2.7 mm) (MNHN-Pg 5285). — Stn CP 38, 07°40'S, 132°27'E, 620-666 m.

28.10.1991: 1 6 (3.1 mm) (MNHN-Pg 5286). — Stn CP 39, 07°47'S, 132°26’E, 477-466 m, 28.10.1991: 1 ov. 9

(2.4 mm) (POLIPI).

Source :

496

P. A. MCLAUGHLIN

Tanimbar Islands: sin CC 56. 08°16'S, 131°59'E. 552-549 m. 31.10.1991: 1 <3, 1 9 (2.1. 2.3 mm) (SNHM 4810). —

SinCC 57, 08°19'S, 131°53'E. 603-620 m, 31.10.1991: 2 6 , 2 ov. $ (1.9-2.2 mm) (MNHN-Pg 5287);

1 6 (1.8 mm) (POLIPI); 2 <3, 2 9. 1 ov. 9 (1.7-2.3 mm) (USNM 276016).

TYPES. — The male (3.1 mm) (MNHN-Pg 5286) from Karubar Station CP 38 is the holotype. The other

specimens are paratypes.

DESCRIPTION. — Shield (Figs 17a-b) usually broader than long, occasionally longer than broad in some

females; anterior margin between rostrum and lateral projections slightly concave; anterolateral margins rounded;

posterior margin roundly truncate; surface with numerous tufts of setae. Rostrum broadly rounded, produced to or

slightly beyond level of lateral projections. Lateral projections very obtusely triangular, subacute or acute, with

terminal spine or spinule.

Ocular peduncles very short and stout, dorsal surfaces each with 2 or 3 short transverse rows of setae; corneae

usually dilated (not noticeably so in illustrated paratype); equal to slightly less than half length of peduncle. Ocular

acicles triangular, moderately to very slender, reaching to or slightly beyond bases of corneae; terminating in acute

spine, mesial margins each with row of moderately long setae; separated basally by 1.00 to 1.35 basal width of

one acicle.

Antennular peduncles overreach distal margin of corneae by half to entire length of penultimate segment.

Ultimate segment with short row of long setae or bristles on dorsal surface mesially and longer row laterally,

usually several additional setae at dorsodistal margin. Penultimate and basal segments unarmed, but with few

scattered setae.

Antennal peduncles overreach distal margin of cornea by 0.75 to 1.25 length of ultimate segment. Fifth and

fourth segments with few scattered setae. Third segment with small spine or spinule at ventrodistal angle. Second

segment with dorsolateral distal angle produced, terminating in acute spine, lateral margin usually with 1 or

2 spinules distally; dorsomesial distal angle with smaller spine, mesial margin with few setae. First segment with

small spine on laterodistal margin; ventral margin also with 1 small spine distolaterally. Antennal acicle long,

reaching to distal half or third of fifth segment, with small terminal spinule; mesial margin with numerous long

setae. Antennal flagellum long, overreaching outstretched chelipeds; I or 2 very short setae every 1 to 4 articles, at

least in proximal third.

Right cheliped (Figs 17c, 39a) long, moderately slender, somewhat dorsoventrally compressed. Dactyl 0.50 to

0.80 length of palm; dorsal surface minutely granular, at least in mesial half and with few scattered setae; rounded

dorsomesial margin and mesial face granular or spinulose; cutting edge with 1 or 2 broad calcareous teeth,

terminating in small corneous claw. Palm equal to or slightly longer than carpus; dorsomesial margin not

delimited, dorsolateral margin weakly indicated by slightly elevated granular or spinulose ridge sometimes

extending nearly to tip of fixed finger; dorsal surface smooth, granular or minutely spinulose; mesial and lateral

surfaces minutely spinulose or granular; fixed finger with scattered moderately long setae, cutting edge with 1 or

2 distinct teeth, terminating in small corneous claw; ventral surfaces smooth or microscopically granular and with

scattered long setae. Carpus slightly shorter to slightly longer than merus; dorsomesial margin with irregular

sometimes double or triple row of small to very small spines and spinules, dorsal surface spinulose or granular,

dorsolateral margin spinulose but not distinctly delimited; lateral mesial and ventral surfaces spinulose or granular.

Merus subtriangular; dorsal surface with few short transverse setal ridges at least in distal half, occasionally 1 or

2 small spines at distal margin; mesial, lateral and ventral surfaces granular or spinulose; ventromesial margin

sometimes with larger spines in distal half; ventrolateral margin rounded, granular or spinulose. Ischium unarmed

or with row of minute spinules on ventromesial margin.

Left cheliped (Figs 17d, 39b) slender, somewhat dorsoventrally compressed; dactyl and fixed finger 1.00 to

1.75 length of palm; in lateral view, straight or ventrally curved; terminating in corneous claws. Dactyl with

dorsomesial margin and mesial half of dorsal surface spinulose in proximal half, numerous long setae distally.

Palm and proximal half of fixed finger with minutely spinulose dorsal surfaces; rounded dorsomesial margin

spinulose, dorsolateral margin with row of very small spines on faintly raised ridge; mesial and lateral faces

spinulose; ventral surfaces smooth or microscopically spinulose, distal halves of dactyl and fixed finger with

numerous tufts of long setae. Carpus slightly shorter or equal to length of merus; dorsomesial margin with

irregular single or nearly double row of small spines; dorsolateral margin also with single or double row of smaller

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

497

spines; all surfaces at least partially covered with very small spinules and sparse short to moderately long setae,

particularly dorsally. Merus with few short occasionally spinulose transverse ridges or low protuberances and tufts

of setae; lateral, mesial and ventral surfaces spinulose, slightly stronger spines on ventromesial and ventrolateral

margins distally. Ischium unarmed or with microscopically spinulose ventromesial margin.

Ambulatory legs (Figs 17e-f) elongate, overreaching outstretched right cheliped. Dactyls not blade-shaped; in

dorsal view, twisted; in lateral view, slightly curved ventrally in distal halves; exceeding length of propodi usually

Fig. 17. — Catapagurus oculocrassus sp. nov., a, c, g-k, holotype 6 (3.1 mm) from Karubar Stn CP 38; b. e-f, paratype

6 (3.7 mm) from Karubar Stn CP 20; d, paratype $ (3.7 mm) from Stn CP 20: a-b. shield and cephalic appendages;

c, right cheliped (mesial view); d, left cheliped (dorsal view); e, second right pereopod (lateral view); f. third left

pereopod (lateral view); g, sternite of third pereopods; h, tip of dactyl and preungual process of left right fourth

pereopod (lateral view); i, coxae and sternite of fifth pereopods; j, tip of right sexual tube; k, telson. Scales equal

0.25 mm (h), 0.5 mm (j), 1.0 mm (k), 2.0 mm (a-b, g, i), 3.0 mm (d), and 5.0 mm (c, e-f)-

498

P. A. MCLAUGHLIN

by more than 0.35 own length, third pair usually slightly longer than second; dorsal margins each with row of

long rather stiff setae; mesial faces each with ventral row of long fine setae; lateral faces and ventral margins with

few setae. Propodi 1.5 to nearly twice length of carpi; dorsal surfaces each with row of small spines interspersed

with short often stiff setae; mesial faces and sometimes also lateral faces spinulose, at least in dorsal halves; ven¬

tral margins each usually with row of minute setae and 1 stiff bristle at distal angle. Carpi short, frequently less

than half length of meri; dorsal surfaces each with sparse setae and numerous small spinules, largest distally;

lateral faces, and mesial faces dorsally, also spinulose. Meri with short transverse rows of moderately long setae on

dorsal surfaces, tending to be spinulose or spinose in distal third, usually 1 or 2 slightly stronger spines near distal

margin; ventromesial and ventrolateral margins delimited only distally, usually unarmed, ventral surfaces

spinulose, spinules strongest on second. Ischia with few tufts of setae dorsally and ventrally. Sternite of third

pereopods (Fig. 17g) with roundly rectangular anterior lobe. Carpus of fourth pereopods unarmed or with small

blunt spine at dorsodistal angle; preungual process of dactyl (Fig. 17h) approximately as long as claw. Fifth

pereopods chelate.

Male with moderately long right sexual tube (Figs 17i-j) directed toward exterior and upward over dorsal surface

of abdomen. Left coxa with slight protuberance of vas deferens. Pleopods of third and fourth somites very

unequally biramous, fifth uniramous. Females also with uniramous fifth pleopod.

Uropods with protopod of right (only) produced posteriorly into small spine. Telson (Fig. 17k) with posterior

lobes acutely triangular, terminating in corneous spine; oblique terminal margins each with 1 to 4 small spines;

lateral margins each with narrow chitinous marginal plate.

COLOR (in preservative). — Some specimens in alcohol for four years retain an orange tint on chelipeds,

particularly carpi and meri.

HABITAT. — Gastropod shells frequently covered by bryozoans.

Distribution. — Known only from the Kai and Tanimbar Islands, Indonesia; 466-809 m.

ETYMOLOGY. — From the Latin oculus meaning eye, and crassus meaning stout, indicating the stout ocular

peduncles characteristic of this species.

Affinities. — At first glance, C. oculocrassus bears a more striking similarity to Icelopagurus crosnieri than

to other species of Catapagurus. Both C. oculocrassus and I. crosnieri have noticeably setose shields, very short,

stout ocular peduncles, long and moderately broad ocular acicles, elongate antennular peduncles, relatively slender

subequal chelipeds, comparable tubular preungual process at the base of the claw of the dactyl of the fourth

pereopod, and a right male sexual tube. Differences of cheliped armature are species distinctive, but it is the

subtriangular, relatively weakly armed posterior telsonal lobes of C. oculocrassus that provides the most

convincing generic distinction.

The species of Catapagurus described herein are morphologically very closely allied, in that none have the

blade-like ambulatory dactyls seen in Catapagurus ensifer and the figured, but unnamed species reported by Haig

and Ball (1988) from the Banda and Arafura regions of Indonesia. Catapagurus oculocrassus is most reliably

distinguished from the following new species by the posterior lobes of the telson which have oblique terminal

margins armed with one to four small spines. In all of the other known species, including C. ensifer and Haig and

Ball's species, the perpendicular terminal margins are unarmed or provided only with very short stiff bristles; the

posterior lobes are separated by a very broad and deep median cleft. Until further study of Catapagurus is

completed, its actual generic limits remain uncertain.

Catapagurus tanimbarensis sp. nov.

Figs 18a-m, 39c-d

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn 49, 08°00’S, 132°59'E, 210-206 m,

29.10.1991: 1 ov. 2 (2.1 mm) (MNHN-Pg 5288); 1 <5, 1 ov. $ (1.9-2.1 mm) (MNHN-Pg 5289).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

499

TYPES. — The ovigerous female (2.1 mm) (MNHN-Pg 5288) is the holotype. The other specimens are

paratypes.

DESCRIPTION. — Shield (Figs 18a-b) slightly longer than broad; anterior margin between rostrum and lateral

projections concave; anterolateral margins sloping; posterior margin roundly truncate; surface with several tufts of

setae. Rostrum broadly rounded, produced to or slightly beyond level of lateral projections. Lateral projections

obtusely triangular, with terminal spine.

Ocular peduncles moderately short and stout, approximately 0.75 length of shield, dorsal surfaces with 1 or

2 tufts of setae; corneae strongly dilated. Ocular acicles narrowly triangular, slender, reaching to or slightly beyond

mid-length of peduncle; terminating acutely, mesial margins with few moderately short setae; separated basally by

nearly entire basal width of one acicle.

Antennular peduncles overreach distal margin of corneae by entire length of ultimate segment. Ultimate

segment with short oblique row of long setae at dorsodistal margin, 2 rows of widely-spaced short setae on dorsal

surface. Penultimate segment with few setae. Basal segment with spine on produced ventromesial distal angle.

Antennal peduncles overreach distal margin of cornea by half length of ultimate segment. Fifth and fourth

segments with few scattered setae. Third segment with small spine or spinule at ventrodistal angle. Second

segment with dorsolateral distal angle produced, terminating in acute spine, lateral margin usually with 1 accessory

spinule distally; dorsomesial distal angle with relatively long spine, mesial margin with few setae. First segment

with small spine on laterodistal margin, ventral margin also with 1 small spine distolaterally. Antennal acicle

moderately long, reaching to proximal third of fifth segment, but not overreaching distal margin of cornea; with

small terminal spinule, mesial margin with few long setae. Antennal flagellum long, overreaching outstretched

chelipeds; 1 or 2 very short setae every 1 or 2 articles.

Right chelipcd (Fig. 39c) long, moderately slender, somewhat dorsoventrally compressed. Dactyl approximately

equal to length of palm; dorsal surface spinulose, at least in proximal half and with scattered setae; dorsomesial

margin with row of spinules, mesial face weakly spinulose; cutting edge with 2 widely-spaced calcareous teeth

interspersed with smaller calcareous denticles; terminating in small corneous claw and slightly overlapped by fixed

finger. Palm equaling length of carpus; dorsomesial margin weakly delimited by multiple rows of very small

spinules, dorsolateral margin with slightly elevated spinulose ridge extending nearly half length of fixed linger;

dorsal surface covered by minute spinules except for smooth distal third of fixed finger, mesial and lateral surfaces

minutely spinulose or granular; fixed finger with moderately long setae distally; cutting edge with 3 widely-spaced

large and few smaller calcareous teeth, terminating in small corneous claw; ventral surfaces smooth or

microscopically granular and with scattered very short setae. Carpus approximately equal to merus in length;

dorsomesial margin with row of small spines becoming double or triple row distally, dorsodistal margin with few

small spines, dorsal surface minutely spinulose, dorsolateral margin with double row of spinules; lateral, mesial

and ventral surfaces spinulose. Merus subtriangular; dorsodistal margin with 2 or 3 small spines, dorsal surface

with few short transverse spinose and setose ridges at least in distal half; mesial, lateral and ventral surfaces

microscopically spinulose; ventrolateral margin with 1 spine at distal angle and minute spinules or tubercles

proximally; ventromesial margin rounded, granular or spinulose. Ischium with small spine at ventrolateral distal

angle.

Left chelipcd (Figs 18c, 39d) slender, reaching to base of dactyl of right; somewhat dorsoventrally compressed;

dactyl and fixed finger 1.25 to nearly twice length of palm; in lateral view, straight or slightly curved ventrally;

terminating in corneous claws. Dactyl with dorsomesial margin faintly serrate in proximal 0.25 to 0.50, several

moderately long setae marginally and on dorsal surface. Palm and proximal 0.50 to 0.75 of fixed finger with

minutely spinulose dorsal surfaces; rounded dorsomesial margin spinulose, dorsolateral margin faintly marked by

row of very small spines forming weak ridge proximally; mesial and lateral faces spinulose; ventral surfaces

smooth or microscopically spinulose, distal halves of dactyl and fixed finger with few long setae. Carpus equal to

or slightly longer than merus; dorsomesial and dorsolateral margins each with row of small spines; all surfaces

with very small spinules and few short to moderately long setae. Merus with short, occasionally spinulose,

transverse ridges and stiff setae; lateral and mesial faces minutely spinulose; ventral surface with scattered very

small spines or tubercles, ventromesial and ventrolateral margins spinulose, each with 1 stronger spine at distal

angle. Ischium unarmed.

500

P. A. MCLAUGHLIN

Fig. 18. — Catapagurus tanimbarensis sp. nov., a, d-f, k, holotype ov. 9 (2.1 mm) from Karubar Stn DW 49;

b-c. g-j, 1, paratype 6 (2.0 mm) from Stn DW 49; m, paratype 9 (1.9 mm) from Stn DW 49: a-b shield and cephalic

appendages; c, carpus and chela of left cheliped (mesial view); d right second pereopod (lateral view); e, dactyl of

right second pereopod (mesial view); f, third left pereopod (lateral view); g, anterior lobe of sternite of third

pereopods; h, tip of dactyl and preungual process of left right fourth pereopod (lateral view); i. coxae and sternite of

fifth pereopods with sexual tubes; j, tip of right sexual tube; k-m. telson. Scales equal 0.25 mm (h), 0.5 mm (j-m),

1.0 mm (a-b, g, i), and 2.0 mm (c-f).

Ambulatory legs (Figs 18d-f) elongate, overreaching outstretched right cheliped. Dactyls not blade-shaped; in

dorsal view, straight; in lateral view, slightly curved ventrally in distal half; equaling or exceeding length of

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDmON

501

propodi usually by approximately 0.25 own length, third pair usually slightly longer than second; dorsal margins

each with row of setae, longer and more bristle-like distally (most broken off in holotype); mesial faces each with

ventral row of 12 to 14 corneous spinules; lateral faces with few setae; ventral margins glabrous. Propodi 1.75 to

twice length of carpi; dorsal surfaces each with row of low protuberances and short stiff setae or bristles; mesial

and ventral faces unarmed; ventral margins occasionally with few short setae and 1 stiff bristle at distal angle.

Carpi short, 0.50 to 0.75 length of meri; dorsal surfaces each with sparse setae and row of small spines, strongest

on second pereopods; lateral, mesial and ventral surfaces glabrous. Meri with 4 short transverse ridges dorsally,

each usually with 1 small spine and 2 or 3 short bristles; ventromesial and ventrolateral distal angles each with

very small spine, ventromesial and ventrolateral margins of second pereopods minutely spinulose or granular, third

unarmed. Ischia unarmed. Sternite of third pereopods with narrowly subrectangular anterior lobe (Fig. 18g).

Preungual process of fourth pereopods (Fig. 18h) elongate and setose. Fifth pereopods chelate.

Male with moderately long right sexual tube (Figs 18i-j) directed toward exterior and upward over dorsal surface

of abdomen. Left coxa with slight protuberance of vas deferens encircled by short stiff setae. Pleopods of third and

fourth somites uniramous and rudimentary, fifth uniramous and moderately well developed. Females also with

uniramous fifth pleopod.

Uropods with protopod of right (only) produced posteriorly and armed with small spinule. Telson (Figs 1 8k-m)

with posterior lobes separated by moderately deep V-shaped median cleft, triangular; terminating acutely or

subacutely and sometimes with small corneous spine at tip; oblique terminal margins each with 2 to several very

short stiff bristles and rarely small protuberance or spinule; lateral margins each with narrow chitinous marginal

plate.

Color. — Unknown.

Habitat. — One specimen occupied a shell of Natica sp.

DISTRIBUTION. — Known from the Kai and Tanimbar Islands, Indonesia; 184-301 m.

ETYMOLOGY. — Named for the Tanimbar Islands of Indonesia.

AFFINITIES. — Catapagurus tanimbarensis and the following new species share many general morphological

attributes, but can be distinguished by the shapes of their telsons, as is the case of other Indo-Pacific species of the

genus (Haig & Ball, 1988). Additionally, neither of the other new species described herein have the mesial faces

of the ambulatory dactyls armed with a ventral row of spines. The shorter antennular peduncles, but longer

antennal acicles distinguish C. tanimbarensis from C. holthuisi sp. nov., while the more strongly dilated corneae

and more slender ocular acicles immediately separate C. tanimbarensis from C. oculocrassus.

Remarks. — Particularly in the holotype, but also to a lesser extent in both paratypes, the setae on the dorsal

margins of the dactyls of the ambulatory legs have been broken off. All that remains to indicate their original

presence is a row of distinct sockets (Fig. 18e).

Two additional ovigerous females, both with shield lengths of 1.5 mm, are assigned to C. tanimbarensis based

on the shape of the telson and armature of the ambulatory dactyls; but because of their conditions, neither are

considered paratypes. The specimen from station DW 50 (POLIPI), collected at a depth of 184 to 186 m, is

missing both third pereopods and chelipeds; the specimen from station DW 03 (MNHN), collected at a depth of

278-301 m, is missing both chelipeds. However, both are included in the geographical and depth distributions of

the species.

Catapagurus holthuisi sp. nov.

Figs 19a-j, 39e-f

Material

26.10.1991: 1 6

EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 27, 05°33'S,

(1.5 mm) (MNHN-Pg 5290).

1 32°5 1 E, 304-314 m.

Source :

502

P. A. MCLAUGHLIN

Tanimbar Islands: sin CP 77, 08°57'S, 131°27'E, 352-346 m, 03.11.1991: 1 ov. $ (2.8 mm) (MNHN-Pg 5291).

— Sm DW 84, 09°23’S, 131°09'E, 275-246 m. 04.11.1991: 2 ov. 9 (1.8-1 .8 mm) (USNM 276017). — Sin 86, 09°26'S,

131°13'E, 225-223 m, 04.1 1.1991: 1 ov. 2 (1.5 mm) (POLIPI).

TYPES. — The ovigerous female (2.8 mm) (MNHN-Pg 5291) from Karubar station CP 77 is the holotype.

The other specimens are paratypes.

DESCRIPTION. — Shield (Figs 19a-b) slightly to considerably broader than long; anterior margin between

rostrum and lateral projections concave; anterolateral margins sloping; posterior margin roundly truncate; surface

with numerous tufts of setae. Rostrum usually broadly sublriangular, occasionally rounded, not produced to level

of lateral projections. Lateral projections triangular, with marginal spine.

Ocular peduncles moderately short and stout, approximately 0.65 length of shield, dorsal surfaces each with

1 or 2 tufts of setae; corneae strongly dilated. Ocular acicles narrowly triangular, slender, reaching beyond

mid-length of peduncle; terminating acutely, sometimes with distinct simple or minutely bifid spine, mesial

margins each with few moderately short setae; separated basally by more than basal width of one acicle.

Antennular peduncles overreach distal margin of corneae by 0.25 to 0.40 length of penultimate segment.

Ultimate segment with 3 or 4 long setae at dorsodistal margin, row of long setae on dorsal surface. Penultimate

segment with very few setae. Basal segment unarmed.

Antennal peduncles overreaching distal margin of cornea by 0.50 to 0.75 length of ultimate segment. Fifth and

fourth segments with scattered setae. Third segment with spine at ventrodistal angle. Second segment with

dorsolateral distal angle produced, terminating in acute spine; dorsomesial distal angle with well developed spine,

mesial margin with few setae. First segment with small spine on laterodistal margin, ventral margin also with

1 small spine distolaterally. Antennal acicle moderately short, usually reaching little, if any, beyond proximal

margin of Fifth segment or distal margin of cornea; with small terminal spinule, mesial margin with few long

setae. Antennal flagellum long, overreaching outstretched chelipeds; 1 or 2 very short setae every 1 to several

articles.

Right cheliped long, moderately slender; chela (Fig. 39e) somewhat dorsoventrally compressed. Dactyl

approximately equal to length of palm; dorsal surface minutely spinulose, at least in mesial half, and with scattered

setae; dorsomesial margin with row of spinules and sparse long setae, mesial face weakly spinulose; cutting edge

with 2 broad calcareous teeth; terminating in small corneous claw and slightly overlapped by fixed finger. Palm

equaling length of carpus; dorsomesial margin rounded and armed with multiple rows of very small spinules or

tubercles, dorsolateral margin usually with slightly elevated spinulose ridge extending nearly entire length of fixed

finger; dorsal surface covered by minute spinules or granules except for smooth distal half of fixed finger; mesial

and lateral surfaces minutely spinulose or granular; Fixed Finger with moderately long setae distally; cutting edge

with 1 prominent calcareous tooth proximally and 2 widely-spaced broad smaller teeth and few calcareous denticles

distally, terminating in small corneous claw; ventral surfaces smooth or microscopically granular and with sparse

setae. Carpus slightly shorter than merus; dorsomesial margin with single or irregularly double row of small

spines, extending onto mesiodistal margin dorsally, rounded or very weakly ridged dorsolateral margin armed with

multiple rows of spinules or small tubercles; lateral, mesial and ventral surfaces spinulose. Merus subtriangular;

dorsodistal margin with 2 or 3 small spines, dorsal surface with few short transverse spinose and setose ridges at

least in distal half; mesial, lateral and ventral surfaces microscopically spinulose; ventrolateral margin with 1 spine

at distal angle and minute spinules or tubercles proximally; venlromesial margin rounded, granular or spinulose.

Ischium with spine at ventrolateral distal angle, ventrolateral and ventromesial margins sometimes spinulose.

Left cheliped (Figs 19c, 39f) slender, reaching well beyond base of dactyl of right; somewhat dorsoventrally

compressed; dactyl and fixed Finger 1.25 to twice length of palm; in lateral view, straight or very slightly curved

ventrally; terminating in corneous claws. Dactyl with row of spinules on dorsomesial margin in proximal half and

row of long setae along entire length of margin. Palm 0.65 to 0.75 length of carpus; dorsal surface of palm and

Fixed finger spinulose and with scattered long setae; rounded dorsomesial margin spinulose, dorsolateral margin

with row of very small spines forming weak ridge extending almost entire length of Fixed Finger; mesial and lateral

faces spinulose; ventral surfaces smooth or microscopically spinulose, dactyl and Fixed finger with numerous long

setae. Carpus equal to length of merus; dorsomesial and dorsolateral margins each with row of small spines;

Source : MNHN Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

503

of right sexual tube; j, telson. Scales equal 0.10 mm (h), 0.5 mm (i), 1.0 mm (b. g, j), 2.0 mm (a), and 3.0 mm (c-f)-

distomesial and distolateral margins each with 2 or 3 small spines; all surfaces with small spinules or tubercles and

few moderately long setae. Merus with spine at dorsodistal margin, 1 or 2 additional spines on dorsal surface in

distal half and double row of stiff setae; lateral and mesial faces minutely spinulose at least in ventral halves;

ventral surface with scattered small spines or tubercles, ventromesial and ventrolateral margins each with row of

small spines, strongest at distal angle. Ischium with spine on ventrolateral margin distally.

Ambulatory legs (Figs 19d-f) elongate, overreaching outstretched right cheliped by more than half length of

dactyls. Dactyls not blade-shaped; in dorsal view, straight or slightly twisted; in lateral view, slightly curved

Source :

504

P. A. MCLAUGHLIN

ventrally in distal half; equaling or exceeding length of propodi usually by 0.25 to 0.35 own length, third pair

commonly slightly longer than second; dorsal margins each with row of long moderately stiff setae; mesial faces

each with ventral row of equally long setae; lateral faces occasionally with row of widely-spaced short setae, third

often with faint longitudinal sulcus dorsally; ventral margins glabrous. Propodi 1.65 to twice length of carpi;

dorsal surfaces minutely spinulose, each also with row of long setae; mesial and lateral faces also minutely

spinulose; ventral margins each with few spinules and row of very fine, moderately short setae. Carpi 0.65 to 0.75

length of meri; dorsal surfaces each with sparse stiff setae and irregular almost double row of small spines or

spinules, strongest on second pereopods; lateral faces minutely spinulose; mesial and ventral surfaces smooth.

Meri each with 1 spine at dorsodistal margin, row of low sometimes spinulose protuberances and stiff setae on

dorsal surface, first 3 or 4 often also with small spine; ventromesial and ventrolateral margins of second pereopods

each with row of small spines, strongest ventrolateral ly; third with spine at ventrolateral distal angle, ventrolateral

margins usually minutely tuberculate or spinulose. Ischia unarmed. Sternite of third pereopods (Fig. 19g) with

narrowly subrectangular anterior lobe and broad posterior lobe subdivided by weak longitudinal furrow. Preungual

process of fourth pereopods (Fig. 19h) elongate and setose. Fifth pereopods semichelate.

Males with long right sexual tube directed toward exterior and upward over dorsal surface of abdomen from left

to right; with partially protruded tip (Fig. 19i); left coxa with small almost transparent sexual tube partially

obscured by circle of short setae; moderately short uniramous left pleopods on abdominal somites three to five.

Females with well developed biramous second to fourth pleopods, fifth uniramous.

Uropods with protopods usually not noticeably produced, unarmed. Telson (Fig. 19j) with triangular posterior

lobes separated by broad subrectangular or rectangular median cleft, terminating in strong corneous spine; generally

perpendicular margins usually with row of very short bristles; lateral margins each with narrow chitinous marginal

plate frequently armed with 1 or 2 moderately long stiff bristles.

COLOR. — Unknown.

Habitat. — One specimen occupied shell of Natica sp.

Distribution. — Known from the Kai and Tanimbar Islands, Indonesia; 223-352 m.

ETYMOLOGY. — This species is dedicated to Dr Lipke B. Holthuis, Nationaal Natuurhistorisch Museum,

Leiden, one of the world’s foremost carcinologists and always a willing advisor.

AFFINITIES. — As previously indicated, C. holthuisi appears quite closely allied to C. tatiimbarensis

particularly in the shape and armature of the chelipeds. The two species are immediately distinguished by the

armature of the ambulatory dactyls; fringed with long setae dorsally and ventromesially in C. holthuisi, but

provided with a mesioventral row of corneous spines in C. tanimbarensis. In the absence of the ambulatory legs,

or in instances where the setae have been broken off, the broad subrectangular median cleft and perpendicularly-

sided terminal margins of the posterior telsonal lobes of C. holthuisi readily separate this species from both

C. tanimbarensis and C. oculocrassus in which the median cleft is V-shaped and the terminal margins oblique.

Genus NEMATOPAGURUS A. Milne Edwards & Bouvier, 1892

Nematopagurus A. Milne Edwards & Bouvier, 1892: 209; 1899: 59; 1900: 200. — Alcock. 1905b: 108. — Miyake,

1978: 128.

Diagnosis. Eleven pairs of phyllobranchiate gills. Shield with rostrum weakly and obtusely sublriangular,

broadly rounded or obsolete. Ocular peduncles relatively stout; corneae often prominently dilated. Ocular acicles

triangular or ovate, with strong submarginal spine. Antennal peduncle with supernumerary segmentation.

Maxillule with external lobe of endopod usually rudimentary, sometimes somewhat developed, not recurved.

Ischium of third maxilliped with crista dentata well developed, 1 accessory tooth.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

505

Chclipeds moderately long and slender; subequal, with right generally slightly longer and/or more robust.

Ambulatory legs moderately long; dactyls and propodi usually similar. Fourth pereopods semichelate; propodal

rasp with 1 row of scales; dactyls without prominent preungual process.

Males with moderate to long usually filamentous sexual tube emanating from coxa of right fifth pereopod and

orientated from left to right across ventral body surface; coxa of left with papilla or short sexual tube; 3 unpaired

unequally biramous pleopods. Females with paired gonopores, paired first pleopods modified as gonopods,

4 unpaired pleopods.

Telson with transverse suture; slightly asymmetrical posterior lobes divided by distinct median cleft; terminal

margins rounded or somewhat oblique.

REMARKS. — As previously mentioned, personal examination of the type specimens of Calapagurus australis

Henderson, 1888, has shown that this species correctly belongs in Nematopagurus and is herein formally

transferred. The specimen upon which HENDERSON (1888) based his description was from "Challenger" station 188

in the Arufura Sea. He stated that the abdomen was missing from that particular specimen, but that the carapace

measured 4.8 mm and the ocular peduncle 2.5 mm. The second specimen, from the reefs at Levuku, Fiji Islands,

was reported to be a "very imperfect specimen”. Catapagurus australis is represented by two syntypes (NHM

1888.33), a male with a shield length of 1.5 mm and carapace length of 2.8 mm, lacking an attached abdomen,

although a somewhat damaged abdomen is present, and a second slightly larger male with attached abdomen, and

shield and carapace lengths of 2.1 and 3.5 mm respectively. The ocular peduncles of neither specimen approach the

2.5 mm given by HENDERSON. The smaller specimen, from "Challenger" station 188 is better preserved, as

Henderson noted, and is accompanied by detached right and left chclipeds and right third pereopod. The syntype

from the Fiji Islands has only a right cheliped and both third pereopods, all detached and poorly calcified. Since

both specimens agree well with HENDERSON'S description, there is no need to designate a lectotype at this time.

With the inclusion of HENDERSON'S taxon, 12 species are currently assigned to Nematopagurus , of which

1 1 are found in the Indo-Pacific region. Three additional new species are described herein. Two others species are

represented in the KARUBAR collection, but their identities both are questionable. One is tentatively assigned to

Nematopagurus indicus Alcock, 1905b; the other most closely resembles N. gardineri Alcock, 1905a, b, but its

conspecificity with this taxon is doubtful. When the host of additional new species of this genus awaiting

description (J. FOREST, personal communication; personal observations) have been fully studied. Nematopagurus

will be one of the most speciose genera in the entire region. In the interim, specimen assignments to known taxa,

particularly of those species exhibiting a median longitudinal row of spines on the dorsal surface of each chela,

must be considered conditional, at best.

Key to the Maluku species of Nematopagurus

1. Chelae of chclipeds marked by transverse scutes . N. scutelliformis sp. nov.

— Chelae of chelipeds not marked by transverse scutes . 2

2. Chelipeds with dorsal surfaces armed with spines modified by tear-drop shaped sensory

structures . /V. spinulosensoris

— Chelipeds without dorsal surfaces armed with spines modified by tear-drop shaped sensory

structures . 3

3. Mesial and lateral faces of carpi of chelipeds each with short transverse rows of long stiff

iridescent bristle-like setae . 4

— Mesial and lateral faces of carpi of chelipeds without short transverse rows of long stiff

iridescent bristle-like setae . 5

4. Dactyls of chelipeds each with dorsomesial row of spines; dorsal surface of fixed finger of

right chela spinulose . N. australis*

— Dactyls of chelipeds each without dorsomesial row of spines; dorsal surface of fixed finger

of right chela unarmed . N. alcocki sp. nov.

Source :

506

P. A. MCLAUGHLIN

5. Telson with subtriangular posterior lobes; terminal margins oblique . 6

— Telson with rounded posterior lobes; terminal margins convex and armed with marginal

and submarginal spines . N. cf. indicus

6. Chelae with setae of dorsal surfaces curved or curled; dactyls of ambulatory legs with 5 or

fewer corneous spines on ventral margins . N. ostlingochirus sp. nov.

— Chelae with setae of dorsal surfaces long and straight; dactyls of ambulatory legs with

more than 5 corneous spines on ventral margins .

. Nematopagurus sp. (? = N. gardineri sensu Haig & Ball, 1988)

Nematopagurus cf. indicus Alcock, 1905

Figs 20a. e-f, 40a-c

Nematopagurus indicus Alcock, 1905b: 109, pi. 12, fig. 4. — Kemp & Sewell, 1912: 26. — BALSS, 1912: 110.

Not Nematopagurus indicus - Miyake, 1961: 12 (list); 1978: 30 (list). — Miyake, Sakai & Nishikawa, 1962: 126 (list). =

Nematopagurus valiants (Melin, 1939).

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn DW 49, 08°00'S, 132°59'E, 210-226 m,

29.10.1991: 1 6 (5.2 mm) (MNHN-Pg 5292).

DIAGNOSIS. — Shield (Fig. 20a) longer than broad, with tufts of setae anteriorly and laterally. Rostrum

broadly rounded, not reaching level of lateral projections. Lateral projections with 1 or 2 marginal spines. Ocular

peduncles moderately stout; approximately as long as antennal peduncles but reaching only to middle of ultimate

segment of fully extended antennular peduncles and with sparse setae dorsally; corneae dilated. Ocular acicles

moderately small, terminating subacutely and with strong submarginal spine. Antennal acicle slightly arcuate;

with terminal spine and setose margins, reaching distal half of ultimate peduncular segment. Antennal flagellum

long, nude.

Chelipeds similar in form and ornamentation; left slightly shorter; right (Fig. 40a) stouter; copiously setose,

but not obscuring surface armature. Palm of right cheliped longer than broad and approximately equal to length of

dactyl; dorsomesial margin with row of spines, dorsal surface with median row of spines not extending onto fixed

finger; dorsolateral margin with row of spines extending only to proximal half of fixed finger; ventral surfaces of

dactyl, palm and fixed linger with few long setae. Carpus about as long as merus and longer than palm;

dorsomesial margin with row of spines, dorsal surface with irregular longitudinal row of somewhat smaller spines

laterally; 1 spine on dorsodistal margin. Merus with 3 strong spines on ventrolateral margin; ventromesial margin

with 1 distal and 1 proximal spine, ventral surface with scattered spines.

Left cheliped (Fig. 40b) with palm slightly shorter than dactyl; similar in armature and setation to that of right.

Carpus with dorsomesial and dorsolateral row of spines; 1 spine at dorsodistal margin. Merus with ventromesial

and ventrolateral row of spines; ventral surface with few spinulose protuberances.

Ambulatory legs overreaching chelipeds by 0.50 to 0.75 length of dactyls (second shorter). Dactyl of left third

straight, almost peg-like (Fig. 40c), with row of 14 corneous spines on ventral margin; second and right third with

slight ventral curvature, ventral margins with 8 to 12 corneous spines; all with abundant long setae randomly

arranged. Propodi with short transverse rows of setae on dorsal surfaces. Carpi each with spine at dorsodistal angle;

second right with 2 and second and third left each with 1 additional spine on dorsal surface posteriorly. Meri of

second pereopods each with row of spines on ventral margin; ventral margins of third only with protuberances.

Right sexual tube (Fig. 20e) short, directed from right to left across ventral surface of body, but not reaching

coxa of left fifth pereopod. No development of left sexual tube. Telson (Fig. 20f) with asymmetrical posterior

lobes separated by prominent median cleft; terminal margins with several strong curved spines and few additional

smaller spines on dorsal surface marginally, most numerous on left.

Color (in preservative). — Only faint reddish tint on the tips of spines of chelae remains after four years in

alcohol.

Habitat. — Unknown.

Source : MNHN Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

507

Fig. 20. — Nematopagurus cf. indicus Alcock, 1905, a, e-f, 6 (5.2 mm) from Karubar Stn DW 49. —

Nematopagurus sp., b-c, g, $ (1.7 mm) from Stn DW 18. — Nematopagurus spinulosensoris McLaughlin & Brock,

1974, d, h, ? (5.6 mm) from Stn CP 86: a-b, d, shield and cephalic appendages; c, right ocular acicle; e, coxae and

sternite of fifth pereopods; f-h, telson. Scales equal 0.5 mm (c, g), 1.0 mm (b, 0. 2.0 mm (e, h), 3.0 mm (a), and

5.0 mm (d).

Source :

508

P. A. MCLAUGHLIN

Distribution. — Tanimbar Islands, Indonesia; 210-226 m. ?Off Nicobar Islands and Travancore and Cochin

coasts of India; approximately 185-433 m.

Remarks. — Despite the short right sexual tube and lack of a left, the Karubar specimen certainly appears

to belong to Nematopagurus. Whether it is correctly assigned to N. indicus is less certain. At present only

N. indicus and N. gardineri Alcock, 1905a (repeated practically verbatim by ALCOCK, 1905b with the same

illustration), have been described with a median longitudinal row of spines on the palms of the chelae; however,

one new species reported herein is also similarly armed. Neither of the former species is described in much detail,

although ALCOCK (1905a, b) stated that N. gardineri was the more pilose of the two. Except for its pilosity and

armature of the ambulatory legs, the present Karubar specimen agrees more closely with N. indicus than with

N. gardineri. The diagnosis given above is based on the KARUBAR specimen; differences with ALCOCK's

description are discussed below.

The only reports of N. indicus are those of ALCOCK (1905b), BalSS (1912), and KEMP and SEWELL (1912).

Alcock (1905b)'s description was rather general, dealing more with length ratios than specific diagnostic

characters; his figure, if accurate, provides some additional information. Balss (1912) gave no information on his

specimen other than to note that its capture southwest of Great Nicobar in the Nicobar Islands represented an

extension of the range of the species. KEMP and SEWELL (1912) remarked that their male and female specimens

agreed with ALCOCK's (1905b) description and type specimens in all characters except the eyes, which were

distinctly stouter and reached only to the proximal third of the ultimate peduncular segment of the antennule. In

this regard, the KARUBAR specimen agrees more closely with those reported by KEMP and SEWELL (1912).

ALCOCK (1905b) contrasted N. indicus with N. gardineri by "regularly disposed tufts of setae on the hepatic and

gastric regions" of the carapace in the former species and a "smooth carapace" in the latter. In his figure (ALCOCK,

1905b, pi. 12, fig. 4), the carapace setae are shown in two longitudinal rows on the shield. As noted in the

diagnosis, tufts of setae in the single Karubar specimen occur laterally and anteriorly on the shield. The rostrum

of N. indicus was described by ALCOCK as being "very broadly triangular"; it is broadly rounded in the present

specimen. He specified that the ocular peduncles reached beyond the middle of the ultimate segment of the

antennular peduncle. As previously indicated, the ocular peduncles of the Karubar specimen reach only to the

middle; however, minor variations in this length ratio are to be expected. Similarly, the length of the dactyl of the

right cheliped was reported to be shorter than the palm in the Indian specimens, whereas the ratio would be

approximately 1 : 1 if the tip of the dactyl were not broken in the Karubar specimen.

Alcock (1905b) remarked that the dactyls of the ambulatory legs were stout, compressed and curved, with the

third pair slightly longer than the second. In the Karubar specimen, the third pereopods also are longer than the

second, but whether the peg-like development of the dactyl of the left third (Fig. 40c) is characteristic of the

Karubar taxon or is attributable to injury cannot be determined from this single specimen. Only a spine at the

dorsodistal margin of the carpus of each ambulatory leg was reported in the description, but his figure (ALCOCK,

1905b, pi. 12, fig. 4) shows a proximal spine on the left second. Two proximal spines are present on the second

pereopods and one on the third in the Karubar specimen, as were illustrated, but not described, by ALCOCK

(1905a, pi. 68, fig. 3; 1905b, pi. 12, fig. 2) for N. gardineri.

The right sexual tube was described for N. indicus as a stout tube ending in a very long, lax, curly filament, the

left was short and blunt. In the Karubar specimen, the right is not particularly stout, but quite short, not

reaching to the coxa of the left fifth pereopod. However, it is possible that the more filamentous terminal portion

had been broken off. Variation in the development of the left sexual tube in species of Nematopagurus is not

uncommon, thus the absence of a left tube in the Karubar specimen would not, of itself, exclude the specimen

from N. indicus.

Nematopagurus sp.

Figs 20b-c, g, 40d-e

1 Nematopagurus gardineri - Haig & Ball, 1988: 185; ?not Nematopagurus gardineri Alcock, 1905a, b.

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 18, 05°18'S. 133°01'E, 205-212 m,

24.10.1991: 1 9 (1.7 mm) (MNHN-Pg 5293).

Source : MNHN. Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

509

DIAGNOSIS. — Shield (Fig. 20b) approximately as long as broad, smooth, but with few sparse tufts of setae.

Rostrum broadly rounded, reaching level of lateral projections. Lateral projections each with marginal spinule.

Ocular peduncles moderately stout; slightly longer than antennal peduncles but reaching only to distal 0.75 of

ultimate segment of fully extended antennular peduncles, with tuft of stiff setae dorsally at base of dilated corneae.

Ocular acicles moderately small, terminating subacutely and with strong laterally produced submarginal spine,

giving bifid appearance (Fig. 20c). Antennal acicle somewhat arcuate, with terminal spine and stiff setae mesially,

reaching distal half of ultimate peduncular segment; flagellum long, with 1 or 2 short setae every 2 to 4 articles.

Chelipeds similar in form, sculpture and length; right (Fig. 40d) stouter; with numerous long fine setae,

particularly dorsally but not obscuring surface armature. Palm of right cheliped longer than broad and

approximately equal to length of dactyl; dorsomesial margin with row of spines, dorsal surface with median row of

spines not extending onto fixed finger; dorsolateral margin with row of spines extending only to proximal half of

fixed finger; ventral surfaces of dactyl, palm and fixed finger with moderately long setae. Carpus about as long as

merus and palm; dorsomesial margin with row of spines, dorsal surface with second longitudinal row of somewhat

smaller spines laterally; 1 spine on dorsodistal margin. Merus with 1 small spine on each ventral margin distally;

ventral surface with scattered low protuberances and few setae.

Left cheliped (Fig. 40e) with palm slightly shorter than dactyl; similar in armature and setation to that of right.

Carpus with dorsomesial and dorsolateral row of spines; 1 spine at dorsodistal margin. Merus with 3 widely-spaced

spinules on ventromesial margin; ventrolateral margin with 1 small spine distally.

Ambulatory legs with dactyls of second pair shorter than third; each with row of 8 corneous spines on ventral

margin; all with sparse tufts of moderately long setae dorsally. Propodi with tufts of long stiff setae on dorsal

surfaces. Carpi each with spine at dorsodistal angle and tuft of stiff setae on dorsal surface posteriorly. Meri

unarmed, but with widely-spaced tufts of stiff setae dorsally and ventrally.

Telson with slightly asymmetrical posterior lobes separated by prominent median cleft; terminal margins with

several small spines, strongest near outer angles.

COLOR (based on N. gardineri sensu Haig & Ball, 1988). — Carapace with scattered red and white

chromatophores on semi-transparent background. Antennules light reddish brown distally. Antennae reddish brown.

Meri and carpi of chelipeds with large red spots on semi-transparent background. Carpi, propodi and dactyls of

ambulatory legs with reddish brown longitudinal stripes on semi-transparent background.

Habitat. — Questionably, gastropod shells with associated anemones.

Distribution. — Kai Islands and ? Arafura Sea, Indonesia; 758-212 m.

REMARKS. — Although the KARUBAR specimen is a small female, lacking paired first pleopods, it agrees well

with the general morphology of species of Nematopagurus. The sternal region of the fifth pereopods and anterior

portion of the abdomen are damaged; however, upon close inspection, two openings in the ventral integument,

corresponding in position to those of first pleopods can be seen. There is little doubt that it is correctly assigned to

this genus, and it bears certain similarities to ALCOCK's (1905a, b) N. gardineri , particularly in the seemingly bifid

structure of the ocular acicles. The acicles are not truly bifid, but the well developed submarginal spine is mesial in

position and clearly visible dorsally (Fig. 20b, c) giving the impression of a bifid acicle. Additionally, ALCOCK

specified that the ocular peduncles were equal in length to the anterior margin of the carapace, longer than the

antennal peduncles and reaching to the distal fourth of the ultimate segment of the antennular peduncles. The

KARUBAR specimen agrees in the ratios of ocular peduncle length to anterior shield margin and antennal peduncle,

but not antennular. In contrast, ALCOCK described and illustrated the corneae as not dilated. The ocular peduncles of

the Karubar specimen arc much stouter than ALCOCK illustrated for N. gardineri, and the corneae are clearly

dilated. His description of N. gardineri specifies a single spine at the dorsodistal marginal of the carpi of the

ambulatory legs, but his figure (ALCOCK, 1905a, pi. 68, fig. 3; 1905b, pi. 12, fig. 2) shows two and one

proximal spines on the denuded left second and third pereopods respectively; only a dorsodistal spine is present in

the Karubar specimen. Although the chelipeds are provided with an abundance of long fine setae in the KARUBAR

specimen, this pilosity does not begin to approach the density of the other Karubar specimen tentatively

assigned to N. indicus. This disparity is in marked disagreement with ALCOCK's (1905a, b) observations.

510

p. a. McLaughlin

Very few additional accounts of N. gardineri have been published. The first appears to have been KENSLEY's

(1969) report of the species from the International Indian Ocean Expedition's (IIOE) "Anton Briiun" Stn 390

(29°35'S, 31°42'E). KENSLEY gave only a cryptic account of his male specimen, but his figure (KENSLEY, 1969,

fig. 6e-h) illustrated a species agreeing, at least in armature of the chelipcds, with ALCOCK's description. However,

KENSLEY illustrated a specimen with ocular peduncles broadly expanded distally and with dilated comeae. Although

the type locality for N. gardineri in the Maidive Islands is in relatively close geographic proximity to KENSLEY's

IIOE station, conspecificity of his and ALCOCK's (1905b) specimens can not be confirmed at this point.

MlYAKE (1978) subsequently identified a species from Japanese waters as N. gardineri. While his single male

specimen from Sagami Bay agreed generally with most of the segmental ratios given by ALCOCK (1905a, b), it

apparently differed substantively in having unequal chelipeds. Miyake (1978) reported that the right was "vastly

larger" than the left. The ambulatory legs of MlYAKE's specimen were reported to have three spines "basally"

(presumably intended to mean proximally) on the carpi of the second pair, and 11 or 12 spines on the ventral

margins of the dactyls. It is very unlikely that MlYAKE's and ALCOCK's specimens are assignable to the same

taxon, nor that either is conspecific with the Karubar specimen.

Although Haig and Ball (1988) indicated that their three specimens, collected in the Arafura Sea during the

"Alpha Helix" expedition, agreed with the original ALCOCK (1905a, b) descriptions in most characters, they too

found exception in the shape of the ocular peduncles. Like the KARUBAR specimen, HAIG and BALL (1988) de¬

scribed their specimens as having peduncles "markedly expanded at the cornea." These authors also commented on

Miyake's (1978) record, noting that he did not mention the form of the ocular peduncles. Apparently they over¬

looked his reference to grossly unequal chelipeds. Since Haig and Ball's (1988) description was limited to notes

on living color and the comments on the ocular peduncles referred to previously, it is not possible to ascertain

whether or not their specimens and the KARUBAR female belong to the same taxon but, given the close proximity

of the collection sites, it is probable. It is doubtful that either are conspecific with ALCOCK’s (1905a, b) species.

Nematopagurus spinulosensoris McLaughlin & Brock, 1974

Figs 20d, h, 4Ia-b

Nematopagurus spinulosensoris McLaughlin & Brock, 1974: 246, figs 1-3. — McLaughlin & Lane, 1975: 520,

pis 1-3.

Nematopagurus spinulosensorius - TOrkay, 1986: 139 (misspelling).

Nematopagurus muricatus -Thompson, 1943: 424. — Miyake, 1978: 129; not Nematopagurus muricatus Henderson,

1888.

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 45, 07°54'S, 132°47'E, 302-305 m.

29.10.1991: 1 2 (5.0 mm) (MNHN-Pg 5294). — Stn DW 49, 08°00'S, 132°59'E, 210-206 m, 29.10.1991: I <J

(3.0 mm) (POLIPI). — Stn CP 86, 09°26'S, 131°13 E, 225-223 m, 04.11.1991: 1 2 (5.6 mm) (MNHN-Pg 5295).

DIAGNOSIS. — Shield (Fig. 20d) longer than broad. Rostrum usually obtusely rounded, occasionally obtusely

triangular. Ocular peduncles overreached by both antennular and antennal peduncles; corneae usually strongly

dilated. Ocular acicles acutely triangular, moderately slender, with prominent longitudinal furrow and very strong

submarginal spine.

Chelipeds subequal, right usually somewhat larger; chelae and carpi of both chelipeds with numerous sensory-

modified spines on dorsal surfaces. Right cheliped (Fig. 41a) with dorsal surface of dactyl generally flattened, dor-

somesial margin, or dorsal surface mesially, usually with irregular longitudinal row of unmodified small spines or

tubercles. Palm with irregular single or double row of usually unmodified moderately strong spines on dorsomesial

margin; dorsal surface with several irregular rows of customarily modified spines, extending onto fixed finger prox¬

imally; dorsolateral margin with single or double row of moderately strong, usually modified spines, extending

onto fixed finger as single row of blunt unmodified spines or tubercles. Carpus with row of strong unmodified

spines on dorsomesial margin; dorsal surface with irregular rows of moderately strong, generally modified spines;

laterodistal margin with spine. Distal margin of merus usually with 2 or 3 strong spines; ventrolateral margin

with row of strong spines mesiodistal margin and ventromesial face distally with few small spines.

Source : MNHN. Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

511

Left cheliped (Fig. 41b) with short row of small unmodified spines or spinulose tubercles usually in dorsal

midline of dactyl. Palm with single or double row of frequently modified spines on dorsomesial margin; dorsal

midline with 2 or 3 irregular rows of usually modified spines extending onto fixed finger; dorsolateral margin with

double or triple row of small modified spines proximally becoming single row of small unmodified spines or

tubercles on fixed finger. Carpus with row of frequently unmodified spines on dorsomesial margin; dorsal surface

with 2 or 3 irregular rows of modified spines proximally, tending to cluster distally, distal margin occasionally

with 1 or 2 spines; dorsolateral margin with single or double row of commonly modified spines. Merus with

1-3 spines on distal margin; ventromesial and ventrolateral margins each with row of spines.

Second and third pereopods generally similar. Dactyls long, slender; ventral surfaces each with row of 10 to

13 strong corneous spines. Carpi each with row of strong spines on dorsal surfaces. Sternite of third pereopods

with subsemicircular anterior lobe, anterior margin with long stiff setae.

Coxa of left fifth pereopod with vas deferens usually slightly protruded. Telson (Fig. 20h) with posterior lobes

subtriangular or subquadrate, left usually slightly larger; separated by very shallow median cleft; terminal and

usually also lateral margins weakly calcified, terminal margins rounded or somewhat oblique, each with numerous

small calcareous spines marginally and several stronger calcareous acute or blunt spines submarginally; lateral

margins unarmed or occasionally each with row of small calcareous spines or spinules.

COLOR. — In life: chelipeds and ambulatory legs generally vivid salmon-pink, bordering on iridescent;

antennal flagella bright yellow.

In preservative: Shield pale orange or straw-colored; ocular peduncles light orange with dark orange ring

proximally. Chelipeds very pale orange with white spines; carpi with darker red-orange proximally and ventrally.

Ambulatory legs pale orange with lighter longitudinal stripes on dactyls and propodi; carpi pale orange with darker

red-orange proximally; meri pale orange and white (McLaughlin & Brock, 1974).

Habitat. — Unknown.

Distribution. — Hawaiian Islands, Japan, Maldives, Indonesia; east coast of South Africa; 180 to 250 m.

AFFINITIES. — In having the dorsal surfaces of the chelae armed with numerous well-developed spines,

N. spinulosensoris superficially resembles N. muricatus. However, neither N. muricatus nor any other known

species of this genus are provided with the tear-drop sensory structures so characteristic of the spines of

N. spinulosensoris.

Remarks. — Turkay (1986) reported that he had examined the specimens identified as N. muricatus by

Thompson (1943) from the "John Murray" Expedition and by Miyake (1978) from Tosa Bay, Japan, and had

found these to actually represent N. spinulosensoris. Apparently neither of these latter authors noticed the

distinctive spines apparently unique to N. spinulosensoris.

Nematopagurus scutelliformis sp. nov.

Figs 21a-h, 41c-f

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn CP 05, 05°49'S, 1 32° 1 8’E, 296-299 m.

22.10.1991: 1 6 (4.4 mm) (SNHM 4811). — Stn CP 06, 05°49'S, I32°2I'E. 298-287 m, 22.10.1991: I 2 (6.1 mm)

(MNHN-Pg 5296). — Stn CP 16, 05°I7’S. 132°50'E, 315-349 m. 24.10.1991: I 6 (5.1 mm) (MNHN-Pg 5297).

Tanimbar Islands: stn CP 46. 08°01'S, 132°51'E, 271-293 m. 29.10.1991: 1 6 . 1 9 (2.3, 3.3 mm) (MNHN-

Pg 5298). — Stn CP 83, 09°23’S, 131°00'E, 285-297 m. 4.11.1991: 1 ov. 2 (6.9 mm) (USNM 276008). — Stn CP 84,

09°23'S, 131°09’E, 275-296 m, 4.11.1991: I ov. 2 (3.6 mm) (POLIP1).

Types. — The male (5.1 mm) (MNHN-Pg 5297) from Karubar station CP 16 is the holotype. The other

specimens are paratypes.

DESCRIPTION. — Shield (Fig. 2 la) as broad as long or broader; anterolateral margins sloping; anterior margin

between rostrum and lateral projections concave; posterior margin truncate; dorsal surface with few tufts of setae.

512

P. A. MCLAUGHLIN

Rostrum broadly rounded, not exceeding lateral projections. Lateral projections prominent, roundly triangular or

subquadrate, each usually with small submarginal spine laterally.

Ocular peduncles short, 0.75 to 0.80 length of shield; dorsal surfaces each with median tuft of stiff setae at base

of cornea, dorsomesial surface with few setae; corneae strongly dilated. Ocular acicles small, triangular; terminating

subacutely, with deeply concave dorsal surface and prominent submarginal spine; separated basally by entire basal

width of one acicle.

Antennular peduncles moderately short, exceeding distal margin of corneae by 0.25 to 0.75 length of ultimate

segment. Ultimate segment with 2 or 3 tufts of setae on dorsolateral surface in distal half. Penultimate segment

with few scattered setae. Basal segment with small spine on lateral face.

Antennal peduncles moderately short, overreaching distal margin of cornea by 0.20 to 0.25 length of ultimate

segment. Fifth and fourth segments with few tufts of stiff setae. Third segment with small spine at ventrodistal

angle. Second segment with dorsolateral distal angle produced, terminating in simple spine, lateral and mesial

margins with few stiff setae; dorsomesial distal angle with small spine. First segment frequently with tiny spinule

on laterodistal margin. Antennal acicle moderately long, reaching to or beyond proximal half of ultimate

peduncular segment; arcuate, terminating in acute spine; mesial margin with tufts of long stiff setae. Antennal

flagella long, overreaching tip of right cheliped; occasionally few articles each with 1 or 2 very short setae or

bristles, at least in proximal half.

Chelipeds suhequal; right (Figs 21b. 41c, e) approximately as long, but slightly stronger than left, moderately

elongate. Dactyl slightly shorter than palm; cutting edge with 3 strong calcareous teeth proximally, few corneous

teeth distally, terminating in small corneous claw and slightly overlapped by fixed finger; dorsal surface with

several low transverse scutes mesially and extending onto rounded dorsomesial margin, each with marginal row of

short stiff setae, few tufts of longer setae adjacent to cutting edge; mesial face dorsally and ventral surface also with

tufts of longer setae. Palm approximately as long as carpus; dorsomesial margin not delimited, but with 1 or

2 spines proximally; dorsal surface with 7 to 9 rows of partially to nearly complete transverse scutes continued

onto lateral face proximally, each with marginal row of short stiff setae, 3 distal-most scutes interrupted at

dorsolateral margin by distinct calcareous spine; proximal 0.75 to 0.80 of dorsal surface of fixed finger with

transverse rows of scutes provided with marginal short stiff setae, each interrupted at dorsolateral margin by strong

spine; distal 0.20 to 0.25 of dorsal surface nearly smooth, with only scattered tufts of setae; cutting edge with row

of strong calcareous teeth in proximal 0.65, small calcareous teeth interspersed with corneous teeth distally,

terminating in small corneous claw; ventral surfaces of palm and fixed finger smooth, with few scattered fine setae.

Carpus slightly shorter than merus; dorsodistal margin with 1 small spine and row of uniformly short stiff setae;

dorsomesial margin with row of strong spines; dorsal surface with transverse rows, each consisting of 2 or 3 scutes

extending onto dorsal half of lateral face, and provided marginally with short stiff setae; dorsolateral margin not

delimited; lateral, mesial and ventral surfaces with scattered tufts of setae. Merus subtriangular; dorsal margin with

few transverse ridges and long stiff setae; lateral and mesial faces tufts of stiff setae; ventrolateral margin unarmed

or with 1 small spine distally, frequently few transverse ridges and stiff setae in proximal 0.65; ventromesial

margin 3 or 4 widely-spaced small spines, ventral surface with few low protuberances or ridges and tufts of stiff

setae. Ischium with few stiff setae, occasionally also with 1 small spinulose tubercle on ventromesial margin near

proximal angle.

Left cheliped (Figs 4 Id, 0 moderately long, usually reaching nearly to tip of dactyl of right; moderately slender.

Dactyl slightly longer than palm; cutting edge with row of small corneous teeth, terminating in small corneous

claw and slightly overlapped by fixed finger; dorsal surface with tufts of long setae adjacent to cutting edge, few

marginally setiferous scutes mesially, extending onto rounded dorsomesial margin; mesial face dorsally and ventral

surface each with few long setae. Palm 0.65 to 0.80 length of carpus; dorsomesial margin not delimited, but with

1 or 2 spines proximally; dorsal surface, like that of right, with 7 to 9 transverse rows of nearly complete single

scutes, each extending onto lateral face and provided with marginal row of short stiff setae, 3 distal-most

interrupted at dorsolateral margin by spine; dorsal surface of fixed finger with several transverse scutes, each with

marginal fringe of short stiff setae and terminating in spine at dorsolateral margin, distal quarter to third nearly

smooth, but with scattered tufts of setae; cutting edge with row of small calcareous teeth interspersed with

corneous teeth. Carpus slightly longer than merus; dorsodistal margin with 1 small spine and uniform row of short

PAGURIDAE FROM THE KARUBAR EXPEDITION

513

Fig. 21. — Nemaiopagurus sculelliformis sp. nov., a-g, holotype S (5.1 mm) from Karubar Stn CP 16; h, paratype

ov. 2 (7.0 mm) from Stn CP 83: a, shield and cephalic appendages; b, right cheliped (lateral view); c, right second

pereopod (lateral view); d. dactyl of right second pereopod (mesial view); e, left third pereopod (lateral view); f, coxae

and stemite of fifth pereopods; g-h, telson. Scale equals 1.0 mm (f-g), 2.0 mm (h) and 5.0 mm (a-e).

stiff setae, dorsomesial margin with row of prominent spines, dorsolateral margin not delimited; dorsal surface

with transverse rows each consisting of 2 or 3 individual marginally setiferous scutes and extending onto lateral

face; mesial and ventral surfaces with tufts of stiff setae. Merus subtriangular; dorsal surface with transverse ridges

and stiff setae, distal margin with row of moderately long stiff setae; lateral and mesial faces with tufts of stiff

setae; ventral surface with few low protuberances and tufts of setae; ventromesial margin with row of 4 widely-

spaced spines, decreasing in size proximally, ventrolateral margin with 1 small distal spine and few short

sometimes spinulose ridges in proximal half. Ischium with scattered tufts of setae, and occasionally 1 spinulose

tubercle on ventromesial margin near proximal angle.

Source :

514

P. A. MCLAUGHLIN

Ambulatory legs (Figs 21c-e) overreaching right cheliped by approximately half length of dactyls. Dactyls 0.25

to 0.35 longer than propodi; in lateral view, slightly curved ventrally; in dorsal view, twisted (at least in large

specimens); each terminating in strong corneous claw; dorsal surfaces each with 1 or 2 rows of short corneous

spines often obscured by 2 or 3 rows of long stiff setae; mesial faces each with row of spiniform bristles or

corneous spines; lateral faces each with few scattered setae; ventral surfaces each with row of 9 to 17 strong

corneous spines increasing in size distally. Propodi 0.25 to 0.35 longer than carpi; dorsal surfaces each with row of

short transverse ridges extending onto lateral faces and set with short to moderately long stiff spiniform bristles;

mesial and lateral faces with scattered setae; ventral surfaces usually with 2 or 3 widely-spaced small corneous

spinules and fine setae, 1 or 2 corneous spines at ventrodistal angle. Carpi 0.65 to 0.75 length of meri; dorsal

surfaces each with spine at dorsodistal angle, occasionally 1 additional spine in proximal half at least on right

second pereopod, and all with row of tufts of stiff setae; mesial and ventral surfaces each with few scattered tufts of

long setae; lateral faces each with I to 3 rows of stiff setae, longest medially. Meri laterally compressed; dorsal

surfaces each with row of transverse ridges and stiff setae; lateral and mesial faces usually with few tufts of setae;

ventral surface usually also with tufts of setae. Ischia each with setae dorsally and ventrally. Anterior lobe of

sternite of third pereopods subsemicircular or subrectangular, with long stiff setae medially and/or on anterior

margin. Fifth pereopods chelate.

Males with well developed, elongate, filiform sexual tube on coxa of right fifth pereopod (Fig. 2 IQ. left with

very short sexual tube; unpaired pleopods 3-5 with exopods well developed, endopods substantially reduced. Telson

(Figs 21g-h) with posterior lobes slightly asymmetrical, nearly subsemicircular; separated by deep median cleft;

terminal margins rounded, each with 1 or more rows of acute spines; lateral margins oblique, each (or at least left)

with row of small subacute spines increasing in size toward outer angle; dorsal surface frequently spinulose near

terminal margins.

COLOR (in preservative). — Spines and margins of scutes of chelipeds with faint pinkish orange tint; scutal

setae usually weakly iridescent.

FIabitat. — Not known.

Distribution. — Kai and Tanimbar Islands, Indonesia; 271-349 m.

ETYMOLOGY. — From the Latin scutella meaning small flat dish, and forma meaning shape, and referring to

the scute-like form of the ornamentation of the chelipeds.

AFFINITIES. — Nematopagurus scutelliformis sp. nov. shares several morphological peculiarities with

N. scutellichelis Alcock, 1905b, most notably in having the chelae and carpi ornamented with broad, Hat,

imbricating scutes; however, the two species are readily distinguished by the spination of the chelae and

ambulatory legs. Each chela of Nematopagurus scutelliformis has a row of spines on dorsolateral margin of the

palm distally and fixed finger proximally, that is absent in N. scutellichelis. Additionally, the dorsal surfaces of the

carpi of both ambulatory legs of N. scutellichelis are provided with a row of spines; the propodi are "ringed or

scutellated, the squames and scutes being nude and polished" (ALCOCK, 1905b: 113). The carpi of these appendages

in N. scutelliformis , although having numerous tufts of stiff setae, have only a dorsodistal spine and occasionally

one proximal spine on the second right pereopod; the propodi have short transverse setiferous ridges dorsally that

extend onto the lateral surfaces of the segments only slightly.

Remarks. Although N. scutelliformis is ostensibly very similar in appearance to N. scutellichelis, a

species reported only from very deep water (in excess of 1500 m) off the Maidive Islands in the Indian Ocean, there

is no doubt that the two taxa are distinct. Despite the fact that in some of the larger KARUBAR specimens, the

spines of the right cheliped tend to be obscured by stiff setae, those of the left are very distinct in individuals of

any size. It is improbable that ALCOCK (1905b) would have failed to observe such spines on the left chela in his

specimen(s), which, from the carapace measurement given, appear to be in the size range of the Karubar

specimens. However, even if he had, the differences in armature and ornamentation of the ambulatory legs provide

additional differentiating characters.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

515

Nematopagurus ostlingochirus sp. nov.

Figs 22a-i

MATERIAL EXAMINED. — Indonesia. Karubar, Tanimbar Islands: sin DW 49, 08°00'S, I32°59'E, 210-206 m,

29.10.1991: 1 <$ (1.2 mm) (MNHN-Pg 5299).

Type. — The single specimen collected is the holotype.

DESCRIPTION. — Shield (Fig. 22a) slightly longer than broad; anterolateral margins sloping; anterior margin

between rostrum and lateral projections concave; posterior margin truncate; dorsal surface with very sparse setae.

Rostrum broadly rounded, not reaching distal margins of lateral projections. Lateral projections prominent, roundly

subquadrate, each with small submarginal spine laterally, left with additional accessory spinule.

Ocular peduncles moderately short, but only slightly shorter than length of shield; dorsal surfaces each with

median tuft of stiff setae at base of cornea, 1 additional tuft on dorsal surface and second tuft mesially; corneae

dilated. Ocular acicles small, triangular; terminating acutely, with concave dorsal surface and prominent simple or

bifid submarginal spine; separated basally by slightly less than basal width of one acicle.

Antennular peduncles short, overreaching distal margins of corneae by 0.20 length of ultimate segment or less.

Ultimate segment with 1 or 2 fine setae. Penultimate segment with few scattered setae. Basal segment with small

spine on dorsolateral distal margin.

Antennal peduncles short, not overreaching distal margins of corneae. Fifth and fourth segments with few setae.

Third segment with small spine at ventrodistal angle. Second segment with dorsolateral distal angle produced,

terminating in strong spine, lateral margin with 1 or 2 accessory spines; dorsomesial distal angle with small spine.

First segment with 1 or 2 small spines on ventrolateral margin distally. Antennal acicle moderately long, reaching

proximal half of ultimate peduncular segment; arcuate, terminating in acute spine; mesial margin with few

moderately long stiff setae. Antennal flagella slightly longer than outstretched right cheliped; every 1 or 2 articles

with 1 or 2 very short setae and occasionally 1 long seta.

Chelipeds subequal; right (Figs 22b-c) slightly longer and stronger. Dactyl slightly shorter than palm; cutting

edge with 5 calcareous teeth, terminating in small corneous claw and overlapped by fixed finger; dorsal surface with

few spinules proximally and several moderately short, usually plumose and distally curved or curled setae, longer

simple setae distally, dorsomesial margin with row of spinules in proximal half; mesial and ventral surfaces with

scattered longer setae. Palm slightly longer than carpus; dorsomesial margin with row of small spines; dorsal

midline with longitudinal row of spines extending to distal half of fixed finger, dorsal surface covered with

moderately short usually plumose curved or curled setae and scattered longer simple setae; dorsolateral margin with

row of spines extending approximately half length of fixed finger; dorsal surface of fixed finger with similarly

plumose curved or curled setae proximally and scattered longer simple setae distally; cutting edge with 2 large

calcareous teeth proximally, 1 large and several small calcareous teeth distally, terminating in small corneous claw;

lateral and ventral surfaces of palm and fixed finger with scattered simple setae. Carpus approximately equal to

length of merus; dorsodistal margin with 1 small spine, dorsomesial margin with row of spines; dorsal surface

with scattered moderately long stiff setae and row of smaller spines approximating dorsolateral margin; lateral and

mesial faces with few short transverse rows of stiff setae. Merus subtriangular; dorsodistal margin with row of stiff

setae; dorsal margin and mesial and lateral faces with few long stiff setae; ventrolateral margin with 2 small spines

distally; ventromesial margin 2 slightly stronger spines in distal half. Ischium with few setae dorsally and

ventrally.

Left cheliped (Fig. 22d) long, reaching almost to tip of dactyl of right; moderately slender. Dactyl slightly

shorter than palm; cutting edge with row of small corneous teeth, terminating in small corneous claw and slightly

overlapped by fixed finger; dorsal surface with several small spines and few short plumose and curled setae in

proximal half, scattered longer simple setae distally; mesial and ventral surfaces with few moderately long simple

setae. Palm approximately 0.65 length of carpus; dorsomesial margin with row of spines; dorsal surface with

longitudinal row of small spines in slightly raised midline, extending half length of fixed finger and partially

obscured by short plumose curved or curled setae; dorsolateral margin with row of spines, extending to distal half

516

P. A. MCLAUGHLIN

Fl° 22u ~ Nema,°Pa8ur“s ostlingochirus sp. nov., holotype S (1.2 mm) from Karubar Stn DW 49: a. shield and

cephalic appendages; b, chela and carpus of right cheliped (dorsal view; setae omitted); c, right cheliped (lateral

view); d, left cheliped (dorsal view; setae omitted); e, left third pereopod (lateral view); f. dactyl of left third pereopod

(mesial view); g, anterior lobe of sternite of third pereopods; h, coxae and stemite of fifth pereopods; i, telson. Scale

equals 0.5 mm (g-i) and 1.0 mm (a-f).

Source : MNHN} Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

517

of fixed finger and numerous longer but similarly plumose curled setae; cutting edge of fixed finger with row of

small calcareous teeth. Carpus slightly longer than merus; dorsodistal margin with 1 small spine; dorsomesial and

dorsolateral margins each with row of spines strongest mesially ; mesial, lateral and ventral surfaces with few long

stiff setae; ventrolateral distal angle with small spine. Merus subtriangular; dorsal surface with few stiff setae,

particularly at distal margin; ventromesial and ventrolateral margins each with 2 strong spines in distal half.

Ischium with few setae ventrally.

Only detached left third pereopod (Figs 22e-f) remains with holotype. Dactyl approximately 1.25 length of

propodus; dorsal surface with few short setae; mesial face with row of 8 corneous spines dorsally; lateral face with

few scattered setae; ventral margin with row of 5 corneous spines. Propodus somewhat longer than carpus; surfaces

each with few setae; 2 small corneous spinules at ventrodistal angle. Carpus approximately 0.65 length of merus;

dorsal and ventral surfaces with few setae, dorsodistal angle with small spine. Merus with few setae dorsally and

ventrally. Ischium unarmed. Anterior lobe of sternite of third pereopods (Fig. 22g) narrowly subsemicircular, with

few marginal setae. Fifth pereopods chelate.

Male with long right sexual tube (Fig. 22h) not noticeably filamentous terminally; left tube stout, moderately

short, directed toward right coxa. Telson (Fig. 22i) with posterior lobes practically symmetrical, nearly

subquadrate; separated by moderately deep median cleft; terminal margins straight, both with 3 large spines, left

with additional smaller spine; lateral margins rounded, each with distinct chitinous plate.

COLOR. — Unknown.

Habitat. — Unknown.

Distribution. — Known only from type locality in the Tanimbar Islands, Indonesia; 206-210 m.

ETYMOLOGY. — From the Greek ostlingos meaning curled hair, and cheir meaning hand, denoting the curling

setae on the dorsal surfaces of the chelae of this species.

Affinities. — In having a median row of spines on the dorsal surfaces of both chelae, N. ostlingochirus bears

some similarity to both N. indicus and N. gardineri', however, the new species is easily distinguished by the

distinctive setation of the chelae.

Nematopagurus alcocki sp. nov.

Figs 23a-i

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 22, 05°22'S, 133°01'E, 124-85 m,

25.10.1991: 1 $ (1.3 mm) (MNHN-Pg 5300); 1 ov. 2 (1.7 mm) (MNHN-Pg 5301).

Types. — The ovigerous female (1.7 mm) (MNHN-Pg 5300) from Karubar station DW 22 is the holotype.

The other female is a paratype.

DESCRIPTION. — Shield (Fig. 23a) as broad as long; anterolateral margins sloping; anterior margin between

rostrum and lateral projections concave; posterior margin roundly truncate; dorsal surface with very few tufts of

setae. Rostrum broadly rounded, approximately equaling level of lateral projections. Lateral projections prominent,

roundly triangular, each with small submarginal spine laterally.

Ocular peduncles moderately short, 0.80 to nearly entire length of shield; dorsal surfaces each usually with

median tuft of stiff setae at base of cornea and 1 additional on dorsomesial surface; corneae strongly dilated. Ocular

acicles small, triangular; terminating subacutely, with deeply concave dorsal surface and prominent submarginal

spine; separated basally by slightly less to slightly more than basal width of one acicle.

Antennular peduncles short, overreaching distal margin of corneae by 0.10 to 0.25 length of ultimate segment.

Ultimate segment with 1 or 2 setae on dorsolateral distal margin. Penultimate segment with few scattered setae.

Basal segment with small spine on lateral face dorsally.

Source :

518

P. A. MCLAUGHLIN

Fig. 23. Nematopagurus alcocki sp. nov„ holotype 2 (1.7 mm) from Karubar Sin DW 22: a. shield and cephalic

appendages; b, right cheliped (dorsal view); c. left cheliped (dorsal view); d, right second pereopod (lateral view);

e, dactyl ol right second pereopod (mesial view); f, left third pereopod (lateral view); g. dactyl of left third pereopod

(mesial view); h. anterior lobe of stemite of third pereopods; i, telson. Scale equals 0.5 mm (h-i) and 1 .0 mm (a-g).

Source : MNHN Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

519

Antennal peduncles short, not overreaching distal margins of corneae. Fifth and fourth segments with few stiff

setae. Third segment with small spine at ventrodistal angle. Second segment with dorsolateral distal angle

produced, terminating in simple spine, lateral and mesial margins with few stiff setae; dorsomesial distal angle

with small spine. First segment with 2 small spines on ventrolateral margin distally. Antennal acicle moderately

long, reaching beyond proximal half of ultimate peduncular segment; arcuate, terminating in acute spine; mesial

margin with moderately long stiff setae. Antennal flagella long; occasionally few articles with 1 or 2 very short

setae or bristles, at least in proximal half.

Chelipeds subequal; right (Fig. 23b) slightly longer and stranger. Dactyl somewhat shorter than palm; cutting

edge with 3 strong calcareous teeth proximally and few calcareous teeth distally, terminating in small corneous

claw and overlapped by fixed finger; dorsal surface with scattered short setae and longer setae on rounded

dorsomesial margin, occasionally 1 small spinule proximally; mesial and ventral surfaces also with tufts of longer

setae. Palm slightly shorter than carpus; dorsomesial margin with row of small or very small spines; dorsal mid¬

line with longitudinal row of small spines in proximal third, sometimes extending distally as row of tiny spinules,

dorsal surface with numerous short setae, most abundant distally and laterally, dorsolateral margin with row of very

small spines extending approximately half length of fixed finger; dorsal surface of fixed finger with short setae

proximally and scattered longer setae distally; cutting edge with row of strong calcareous teeth in proximal 0.75,

small calcareous teeth distally, terminating in small corneous claw; lateral and ventral surfaces of palm and fixed

finger with scattered setae. Carpus slightly longer than merus; dorsodistal margin with 1 small spine, dorsomesial

margin with row of spines; dorsal surface with scattered setae, dorsolateral margin with row of somewhat smaller

spines; lateral and mesial faces with long stiff bristle-like and often somewhat iridescent setae. Merus

subtriangular; dorsodistal margin with row of bristle-like setae, dorsal margin and mesial and lateral faces with

short transverse ridges of long stiff bristle-like setae; ventrolateral margin with 1 small spine and 2 tiny spinules

distally, stiff setae in proximal 0.65; ventromesial margin 1 or 2 small spines distally and 1 or 2 tuberculate

protuberances proximally, ventral surface with few low protuberances or ridges and stiff setae. Ischium with few

stiff setae.

Left cheliped (Fig. 23c) moderately long, but not reaching to tip of dactyl of right; moderately slender. Dactyl

slightly longer than palm; cutting edge with row of small corneous teeth, terminating in small corneous claw and

slightly overlapped by fixed finger; dorsal surface with few scattered setae; mesial and ventral surfaces with few

long setae. Palm approximately 0.65 length of carpus; dorsomesial margin with 1 spine at proximal angle, few

transverse ridges and stiff setae, and adjacent row of tiny spinules; dorsal surface with row of very small spines

laterad of midline and extending onto fixed finger proximally and numerous short setae, dorsolateral margin with

row of tiny spines, extending to distal half of fixed finger; cutting edge with row of small calcareous teeth. Carpus

somewhat longer than merus; dorsodistal margin with 1 small spine, dorsomesial and dorsolateral margins each

with row of moderately small spines; mesial and ventral surfaces with numerous short transverse ridges and bristle¬

like long setae; ventrolateral distal angle with small tubercle, ventral surface with few low tubercles and long setae.

Merus subtriangular; dorsal surface with few transverse ridges and bristle-like setae, distal margin with row of

moderately long similar setae; lateral and mesial faces with short transverse ridges and bristle-like setae, ventral

surface with few low protuberances and tufts of setae; ventromesial and ventrolateral margins each with few small

spines distally and low protuberances proximally. Ischium with few tufts of setae.

Ambulatory legs (Figs 23d-g) similar; dactyl of third left (third right missing in holotype) approximately

0.25 longer than dactyls of second; all terminating in small corneous claws; dorsal surfaces each with row of

sparse tufts of moderately short setae and few corneous spines; mesial faces each with row of corneous spines

dorsally; lateral faces each with few scattered setae; ventral surfaces each with row of 6 to 9 corneous spines

increasing in size distally. Propodi 0.30 to 0.45 longer than carpi; dorsal surfaces each with row of tufts of long

bristle-like setae and few corneous spinules; mesial and lateral faces with few scattered setae; ventral surfaces with

1 or 2 small corneous spinules at ventrodistal angle. Carpi 0.65 to 0.75 length of meri; dorsal surfaces each with

spine at dorsodistal angle and 1 or 2 much smaller spines in proximal half, partially obscured by tuft of stiff

bristle-like setae; lateral faces each with 1 or 3 short rows of bristle-like setae. Meri laterally compressed; dorsal

surfaces each with row of short transverse ridges and bristle-like setae; lateral and mesial faces with few setae;

ventral surfaces each with 1 small spine in distal third (second) or unarmed (third) and tufts of setae. Ischia each

520

P. A. MCLAUGHLIN

with few setae dorsally and ventrally. Anterior lobe of sternite of third pereopods (Fig. 23h) roundly

subrectangular, with long marginal setae. Fifth pereopods chelate.

Males unknown. Telson (Fig. 23i) with posterior lobes slightly asymmetrical, nearly subsemicircular;

separated by moderately deep median cleft; terminal margins rounded or slightly oblique, each with 2 to 4 large and

2 smaller spines; lateral margins slightly oblique.

COLOR (in preservative). — Segments of chelipeds and few spines retain faint small spots or patches of light

orange; bristle-like setae with yellowish tint.

DISTRIBUTION. — Known only from the type locality in Kai Islands, Indonesia; 85-124 m.

Etymology. — This species is named for A. Alcock, noted carcinologist of the Indian Museum, whose

monographic work on Indian Paguridea is still the cornerstone for regional studies.

AFFINITIES. — Nematopagurus alcocki appears most closely allied with N. australis , particularly in having

long, stiff irridescent setae on the mesial and lateral faces of the carpi of both chelae, and a covering of short setae

on the dorsal surfaces of the palms and fixed fingers. The ocular peduncles of Henderson's (1888) specimen from

the Arafura Sea are a little more slender than seen in the specimens of N. alcocki , and the corneae only slightly

dilated; however the peduncles of syntype from the Fiji Islands more closely approach those of the Karubar

species. Nonetheless, the two species are immediately distinguished by the dactyls of the chelipeds. The

dorsomesial margins of both dactyls are provided with a row of spines in N. australis, whereas they are completely

unarmed or may have one very small spine proximally in N. alcocki. Other differences include the spination of the

dorsal surface of the palm of N. australis that is lacking in N. alcocki, and the longer ocular peduncles of the

former species. HENDERSON described the carpi of the ambulatory legs as slightly spinulose. Only a dorsodistal

spine is actually present on each of these appendages, whereas the carpi of N. alcocki have one or two small spines

proximally on the dorsal margins, in addition to the dorsodistal spine.

Remarks. — Although this species is described from only female specimens, there can be little doubt as to

the accuracy of its assignment to Nematopagurus. In addition to well developed paired first pleopods modified as

gonopods in the holotype, N. alcocki has such additional generic characters as the small, deeply grooved ocular

acicles, reduced, rounded rostrum and clearly exposed interocular lobes, subequal chelipeds, and elongate ambulatory

legs with ventrally armed dactyls.

The paratype is an immature female, as the gonopores are quite small and the gonopods minute; only a

rudimentary left second pleopod is apparent. The chelipeds of this specimen have similar patterns of spination to

that described for the holotype; however, the spines are appreciably stronger. In contrast, the dorsal surfaces of the

chelae are not as abundantly provided with short setae and the characteristic bristle-like longer setae are weaker and

more sparse on both the chelipeds and ambulatory legs.

One additional female specimen (1.7 mm) (POLIPI) of Nematopagurus was found at stn DW 22, and probably

also represents N. alcocki; however, it is missing the chelipeds, left ambulatory legs, including the coxa of the

third, the coxae and sternite of the fifth pereopods, and the abdomen. This specimen is not considered a paratype,

even though the carpi of both right ambulatory legs have a very small proximal spine on the dorsal margin, and

the second has a tiny spine on the ventral margin of the merus.

Genus AUSTRALEREMUS McLaughlin 1981

Pylopagurus - Forest & DE Saint Laurent, 1968: 145 (in part), not Pylopagurus A. Milne Edwards & Bouvier, 1891. _

Miyake, 1978: 119 (in part). — McLaughlin, 1981a: 2 (in part).

Australeremus McLaughlin, 1981a: 4. — McLaughlin & GUNN, 1992: 68.

DIAGNOSIS. Eleven pairs phyllobranchiate gills. Ocular acicles triangular. Basal antennular segment with

strong lateral spine; ventrodistal margin produced into elongate lobe. Antennal peduncle with supernumerary

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

521

segmentation. Maxillule with external lobe of endopod well developed, not recurved. Third maxilliped with well

developed crista dentata and very strong accessory tooth.

Right cheliped often not appreciably larger than left. Chela of right subrectangular to subtriangular; dorsal

surface of palm circumscribed by row of dorsomesial, dorsoproximal and dorsolateral marginal spines; angle of

propodal-carpal articulation approximately 15° from horizontal plane. Left chela with variable propodal-carpal angle

of articulation; dactyl elongate and considerably narrower than fixed finger; dorsolateral margin of chela elevated, at

least proximally, and frequently expanded. Fourth pereopods semichelate; propodal rasp consisting of single row of

corneous scales.

Males without paired pleopods or sexual tubes; with 3 unpaired, unequally biramous left pleopods. Females

with paired first pleopods modified as gonopods, with 4 unpaired biramous pleopods, second to fourth with both

rami well developed, fifth with endopod reduced.

Abdomen frequently straight or only weakly flexed. Uropods symmetrical or asymmetrical. Telson with

transverse suture, sometimes only weakly indicated; posterior lobes symmetrical or subequal, terminal margins

straight, oblique or rounded, armed; lateral margins each with undifferentiated, usually weakly calcified or chitinous

plate.

REMARKS. — McLaughlin (1981a) chose the generic epithet in the belief that species of the genus all were

endemic to the Southern Ocean. In their subsequent review of Australeremus, McLaughlin and GUNN (1992)

reassigned Eupagurus triserratus Ortmann, 1892, and Pylopagurus serpulophilus Miyake, 1978 (as a junior

synonym) from Japan to the genus, thus giving it a broad, but disjunct distribution. The discovery, not only of

A. triserratus, but of an additional new species in the Maluku region of Indonesia suggests that Australeremus

may be far more ubiquitous than previously believed and simply overlooked by collectors. At least several species

occupy habitats not routinely sampled, such as bryozoan and serpulid worm tubes.

Australeremus triserratus (Ortmann, 1892)

Figs 24a, c

Eupagurus triserratus Ortmann, 1892: 308, pi. 12, fig. 15.

? Eupagurus tricarinatus - Balss, 1913: 58 (?not Eupagurus tricarinatus Stimpson, 1858).

Eupagurus triserratus (?) - SHIINO, 1936: 184.

Eupagurus (Eupagurus) triserratus - MELIN, 1939: 29, figs 9, 10.

Pagurus triserratus - Kim, 1964: 5, pi. 1, fig. 6; 1970: 8; 1973: 225, 599, fig. 50, pi. 65, fig. 30.

Pylopagurus serpulophilus Miyake, 1978: 120, pi. 4, fig. 4; 1982: 120, pi. 40, fig. 5. — McLaughlin. 1981a: 3.

Australeremus triserratus - McLaughlin & Gunn, 1992: 87, fig, 14, pi. 1.

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands : stn DW 50, 07°59'S, 133°02'E, 184-186 m.

29.10.1991: 1 6 (0.9 mm) (MNHN-Pg 5302).

Diagnosis. — Shield (Fig. 24a) slightly to considerably longer than broad. Rostrum acutely triangular. Ocular

peduncles slightly shorter than antennular peduncles. Ocular acicles acutely triangular, with strong submarginal

spine.

Right cheliped (Fig. 24c) with row of spines on dorsomesial margin of dactyl, dorsal surface with scattered low

or spinulose tubercles. Dorsomesial, dorsoproximal and dorsolateral margins of palm each with row of acute spines

entirely circumscribing palm and fixed finger; dorsal surface slightly convex and with numerous blunt or spinulose

tubercles, midline with single or occasionally double row of larger spines. Carpus often with appreciably concave

mesial face; dorsomesial margin with row of spines at least in distal half, second row of smaller spines laterally.

Merus with row of spines on ventrolateral margin distally, ventromesial margin with 3 or 4 spines proximally.

Left cheliped (missing in Karubar specimen) with propodal-carpal articulation 45° to 50° from horizontal

plane. Dactyl dorsoventrally flattened, unarmed. Palm and fixed finger with row of strong spines on dorsolateral

margin, dorsal surfaces generally flattened, armed with 2 irregular rows of tubercles or spinules. Carpus with row

of acute spines on dorsal margin.

522

P. A. MCLAUGHLIN

Ambulatory legs generally similar. Dactyls each with 7 to 1 1 corneous spines on ventral margin; propodi with

low protuberances on dorsal surfaces. Carpi each with 1 spine at dorsodistal angle, second often also with

1 additional spine on dorsal surface proximally. Sternite of third pereopods with subsemiovate anterior lobe.

Uropods symmetrical. Telson (Fig. 24g) with terminal margins of posterior lobes straight or slightly oblique,

armed with 2 to 4 strong spines and occasionally 1 or 2 small spines; lateral plates reduced.

Color (in preservative). — Ground color of body and legs light red-brown. Carapace with pair of dark

red-brown spots in front and behind cervical groove. Antennal flagellum with light and dark red-brown segments

alternatively. Chela and carpus dark red-brown; mcrus light red-brown with three dark colored cross-bands. Walking

legs light red-brown; meri and dactyls each with two dark colored cross-bands; carpi and propodi each with one dark

colored cross-band (after MIYAKE, 1978, pi. 4, fig. 4).

Habitat. — Serpulid worm tubes, at least in part.

Distribution. — Sagami Bay, Tanabe Bay, Amakusa, Japan; East China Sea; Bonin Islands; South China

Sea; Tanimbar Islands, Indonesia; 60 - 400 m.

Affinities. — Based upon the armature of the chelipeds and carpi of the ambulatory legs, McLaughlin and

Gunn (1992) suggested a possible relationship among the species, A. triserratus, A. steward (Filhol, 1883), and

A. eltaninae McLaughlin & Gunn, 1992. However, a much closer kinship now appears to exist with

A. indonesiensis sp. nov., as is discussed under that taxon.

Remarks. — The diagnosis given is here for the species in general, not just as characterized by the very small

Karubar specimen. This report of A. triserratus represents a considerable extension of the southern range of this

species. McLaughlin and Gunn (1992) earlier had documented its occurrence only as far south as the South

China Sea southwest of Kaohsiuhg, Taiwan. Despite its small size and the absence of the left cheliped, the single

specimen in the Karubar collection unquestionably represents A. triserratus. The right palm has the marginal

encirclement of spines characteristic of Australeremus and the median row of spines on the dorsal surface indicative

of A. triserratus.

Australeremus indonesiensis sp. nov

Figs 24 b, d-f, h, 42a-b

_ Mater,al EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 18, 05°18'S, I33°0I'E 205-212

n n TLlu^i’lr1 (MNHN-Pg 5303). - Stn DW 31, 05°40'S. 132°51'E, 288-289 m, 26.10.1991: 1

(2.U mm) (MNHN-Pg 5304).

m,

$

Types. — The female (1.9 mm) (MNHN-Pg 5303) from Karubar station DW 18 is the holotype. The other

female is a paratype.

Description. — Shield (Fig. 24b) longer than broad; anterior margin between rostrum and lateral projections

concave; posterior margin truncate; dorsal surface with scattered tufts of moderately long setae. Rostrum triangular,

with small terminal spine; reaching beyond bases of ocular acicles. Lateral projections obtusely or acutely

triangular, with marginal or submarginal spine. Posterior carapace with few tufts of long setae mesially adjacent to

cervical groove.

Ocular peduncles 0.75 to 0.85 length of shield, moderately slender, dorsal surfaces with few sparse tufts of

setae; corneae slightly, if at all, dilated. Ocular acicles triangular, terminating subacutely and with strong

submarginal spine; separated basally by approximate basal width of one acicle.

Antennular peduncles (when extended) overreaching ocular peduncles by third to half length of ultimate

segment. Basal segment with spine at dorsolateral distal margin. Penultimate segment with scattered short setae.

Ultimate segment usually with 1 long seta at dorsodistal margin and few much shorter setae on dorsal surface.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

523

Fig. 24. — Australeremus triserratus (Ortmann, 1892), a, c, g, 6 (0.9 mm) from Karubar Sin DW 50. — Australeremus

indonesiensis sp. nov., b, d-f, h, holotype 2 (1.9 mm) from Stn DW 18: a-b, shield and cephalic appendages;

c-d, right chela (lateral view); e, right second pereopod (lateral view); f, anterior lobe of stemite of third pereopods;

g-h, telson. Scales equal 0.5 mm (f-h) and 1.0 mm (a-e).

Source :

524

P. A. MCLAUGHLIN

Antennal peduncles overreach ocular peduncles by 0.50 to 0.75 length of ultimate segment. Fifth and fourth

segments with scattered setae dorsally and ventrally. Third segment with unarmed ventrodistal angle. Second

segment with dorsolateral distal angle produced, terminating in acute bifid spine, mesial and lateral margins each

with 1 to 3 accessory spinules; dorsomesial distal angle with acute spine. First segment sometimes with spine at

dorsolateral distal angle, 1 spine on ventrolateral margin distally. Antennal acicle reaching to proximal third of

ultimate segment, terminating in small spine and with 1 to 4 quite long setae; mesial face with few moderately

short setae. Antennal flagellum overreaching right cheliped; every, or every other, article with 1 to 3 long

(4-6 article length) and 1 or 4 shorter setae.

Right cheliped (Figs 24d, 42a) with dactyl from slightly shorter to equaling length of palm; cutting edge with

3 or 4 calcareous teeth proximally, short row of corneous teeth in distal third, terminating in corneous claw and

slightly overlapped by fixed finger; dorsomesial margin with row of spines, dorsal surface with longitudinal row of

spinules or low spinulose tubercles and few setae, 1 or 2 additional spinules or tubercles laterally and adjacent to

cutting edge; mesial and ventral surfaces with scattered long setae. Palm 0.65 to 0.80 length of carpus;

dorsomesial, dorsoproximal and dorsolateral margins each with row of acute spines entirely circumscribing palm

and fixed finger, dorsal surfaces of palm and fixed finger slightly convex and with numerous small spinules, 1 or

2 somewhat stronger adjacent to dorsomesial margin; cutting edge of fixed finger with 1 large calcareous tooth

medianly, smaller teeth proximally and distally, terminating in corneous claw; surfaces with few scattered setae.

Carpus slightly longer than merus; dorsomesial margin with row of spines at least in distal half, only few setae

proximally in paratype, 1 spine at dorsodistal margin and row of spines on dorsolateral margin; all surfaces with

scattered long setae. Merus subtriangular; dorsal margin with few setae, particularly distally; ventromesial margin

with row of small spines distally, ventrolateral margin with few spinulose protuberances. Ischium unarmed.

Left cheliped (Fig. 42b) with propodal-carpal articulation 35° to 45° clockwise from horizontal plane. Dactyl

1.25 length of palm; cutting edge with row of corneous teeth, terminating in small corneous claw; dorsomesial

margin with row of low sometimes spinulose protuberances and few setae, dorsal surface with few scattered setae.

Palm half length of carpus; dorsolateral margin of palm and fixed finger with row of strong spines, dorsal surfaces

generally flattened, armed with few scattered tiny spinules, dorsomesial margin with few spinulose protuberances

and setae. Carpus acutely triangular in cross-section; dorsal margin with row of strong spines, distal margin with

2 smaller spines mesially; 1 or 2 blunt spines on or adjacent to ventromesial margin distally; lateral and ventral

surfaces with scattered long setae. Merus slightly shorter than carpus; dorsal margin with few setae; ventromesial

margin with row of small spines or protuberances and long setae, ventrolateral margin unarmed or with few low

protuberances and long setae. Ischium with few spinules on ventromesial margin.

Ambulatory legs (Fig. 24e) generally similar (third left missing in both holotype and paratype). Dactyls

slightly shorter to approximately equaling length of propodi; dorsal margins each with row of sparse tufts of long

setae; mesial and lateral faces with scattered shorter setae; ventral margins each with 7 or 8 corneous spines and

numerous long setae. Propodi 1.50 to 1.75 length of carpi; dorsal and ventral surfaces with low protuberances and

sparse tufts of long setae. Carpi 0.50 to 0.65 length of meri; dorsal margins each with 1 tiny spinule at distal

angle, dorsal and particularly ventral surfaces with sparse tufts of long setae, second pereopods also with 1 slightly

more prominent protuberance in proximal half. Dorsal and ventral surfaces of meri with low protuberances and

sparse tufts of long setae. Ischia unarmed, but with tufts of setae. Sternite of 3rd pereopods with small

subsemiovate anterior lobe (Fig. 24h) and tuft of long setae.

Dorsal surface of sixth abdominal somite with several long setae; uropods symmetrical. Telson (Fig. 24h) with

transverse suture very faintly indicated; terminal margins of posterior lobes slightly oblique, armed with 2 or

3 spines and 1 or 2 spinules on either side of median cleft, outer angles blunt or with corneous spinule; lateral

margins rounded, chitinous plates limited to very small area adjacent to outer angle.

Color (in preservative). — Only a general faint orange tint remains on the chelae and ambulatory legs after

four years in alcohol.

Habitat. — Unknown.

Distribution. — Presently known only from the Kai Islands, Indonesia; 205-289 m.

Source : MNHN, Pahs

PAGURIDAE FROM THE KARUBAR EXPEDITION

525

ETYMOLOGY. — The specific name, indonesiensis, reflects the general geographic environs of this species,

only the second to be described from outside the "Southern" Ocean.

AFFINITIES. — As previously indicated, A. indonesiensis is morphologically most similar to A. triserratus.

Both species are characterized by the lack of armature on the dorsal margins of the segments of the ambulatory

legs. The lack of a median row of large spines on the dorsal surface of the right chela will immediately distinguish

A. indonesiensis from A. triserratus.

Genus PAGURUS Fabricius, 1775

Cancer Linnaeus, 1758: 625 (in part).

Pagurus Fabricius, 1775: 410 (in part).

Eupagurus Brandt, 1851: 105 (in part).

Bernhardus Dana, 1851: 267 (in part).

Not Pagurus Berthold, 1827: 255 (nomen nudum).

Not Pagurus Fabricius sensu Dana, 1851: 267 (= Dardanus Paul'son, 1875).

DIAGNOSIS. — Eleven pairs of phyllobranchiate gills. Rostrum variable. Ocular acicles simple, bifid or

multifid. Crista dentata well developed, with 1 or more accessory teeth. Sternite of third maxillipeds unarmed or

armed. Chelipeds generally grossly unequal, right usually appreciably larger. Dactyls of ambulatory legs

commonly with spinose ventral margins. Sternite of third pereopods variable. Fourth pereopods usually

semichelate, with 1 to several rows of scales in propodal rasp. Sternite of fifth pereopods variable.

Coxae of fifth pereopods generally symmetrical in both sexes. Males with paired gonopores; no paired

pleopods, usually with 3 or 4 unpaired left pleopods, rarely without unpaired pleopods. Females usually with

paired gonopores; no paired pleopods, usually 4 unpaired left pleopods, rarely only 3.

Abdomen usually spirally twisted, rarely straight. Uropods most commonly asymmetrical, occasionally

symmetrical. Telson typically with transverse suture; posterior lobes frequently separated by well developed median

cleft; terminal margins rounded, straight or oblique.

Remarks. — Pagurus is the "catch-all" genus for any hermit crab having 1 1 pairs of phyllobranchiate gills,

but lacking secondary sexual modifications or similar exclusive characters, and as such is the most specious, albeit

heterogeneous, of all pagurid genera. However, despite the widespread occurrence of more than 150 species

throughout the world's oceans, fewer than one quarter of those have been reported from the tropical Indo-Pacific.

In addition to the KARUBAR specimens, five species of Pagurus are known specifically from the Maluku area.

A small male of Pagurus cornpressipes (Miers, 1884) was the specimen from the Arafura Sea that HENDERSON

(1888) tentatively assigned to his "Pagurodes" piliferus. Although BuiTENDlJK (1937) published only on the

Diogenidae of the "Snellius" expedition, she did identify some of the Paguridae from that collection, among them a

specimen of P. hirtirnanus (Miers, 1880) from Wotap Island ("Tenimber Islands"), now part of the collection of

the National Museum of Natural History (USNM 122050). Haig and Ball (1988) reported P. hirtirnanus and

P. moluccensis Haig & Ball, 1988, from the Banda Islands, P. hedleyi Grant & McCulloch, 1906, from the

Arafura Sea, and P. pergranulatus (Henderson, 1896) from Seram.

Key to the Maluku species of Pagurus

1 . Interocular lobes developed as pair of spinose processes . Pagurus moluccensis*

— Interocular lobes not developed as pair of spinose processes . 2

2. Ventromesial margins of carpus and/or merus of right cheliped strongly produced into

ventrally directed "wing-like" lobe . 3

— Ventromesial margins of carpus and/or merus of right cheliped not produced into ventrally

directed "wing-like" lobe . 4

Source :

526

P. A. MCLAUGHLIN

3. Dorsomesial distal angle of right chela produced into distinct spinose or tuberculate lobe;

dactyls of ambulatory legs longer than propodi; second pereopods each with dorsodistal and

dorsoproximal spine . P. pergranulatus*

— Dorsomesial distal angle of right chela not produced into distinct spinose or tuberculate

lobe; dactyls of ambulatory legs shorter than propodi; second pereopods each with only

dorsodistal spine . P. hedleyi*

4. Dorsal surfaces of chelae armed with numerous very strong spines . 5

— Dorsal surfaces of chela unarmed or armed only with granules, tubercles or small spines....

. 6

5. Dactyl of fourth pereopod with multiple rows of corneous scales in propodal rasp; lateral

face of propodus of left third pereopod with numerous short transverse rows of setae .

. . P. haigae sp. nov.

— Dactyls of fourth pereopods with single row of corneous scales in propodal rasp; lateral

face of propodus of left third pereopod without numerous short transverse rows of setae . . .

. TPagurus sp.

6. Right cheliped not appreciably longer than left; dorsomesial and dorsolateral margins of

chelae each forming low, but distinct ridge, dorsal surfaces armed with transverse ridges

and setae . p. compressipes*

— Right cheliped appreciably longer than left; dorsomesial and dorsolateral margins of chelae

not forming low, but distinct ridges, dorsal surfaces not armed with transverse ridges and

setae . 7

7. Dorsal surfaces of chelae with covering of dense short setae practically obscuring armature

. P. hirtimanus*

— Dorsal surfaces of chelae without covering of dense short setae practically obscuring

armature . 8

8. Dorsal surface of right chela smooth, granular, minutely spinulose or with small simple

tubercles; dactyls of ambulatory legs nearly twice length of propodi .

. P. kaiensis sp. nov.

— Dorsal surface of right chela with numerous small spines and tubercles, latter usually with

central corneous capsule; dactyls of ambulatory legs much less than twice length of

Propodi . P. capsularis sp. nov.

Pagurus kaiensis sp. nov.

Figs 25a-l, 42e-f

Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 14, 05°18'S, 132°38'E, 245-246 m,

24.10.1991: 1 8 (2.6 mm) (MNHN-Pg 5305). — Stn DW 15, 05°17'S, 132°4I'E, 212-221 m 24 10 1991

1 9 (2.8 mm) (POLIPI). — Stn DW 18. 05°18’S, 133°01'E, 205-212 m. 24.10.1991: I 8 (8.4 mm) (MNHN-Pg 5306)-

I 9 (3.0 mm) (MNHN-Pg 5307). — Stn CP 36, 06°05'S, 132°44'E. 268-210 m, 27.10.1991: 1 9 (3.9 mm) (USNM

276015).

Types. — The male (8.4 mm) (MNHN-Pg 5306) from the Karubar station DW 18 is the holotype. The

other specimens are paratypes.

Description. Shield (Fig. 25a) as long as broad or slightly broader; anterior margin between rostrum and

lateral projections concave; anterolateral margins terraced; posterior margin truncate; surface frequently with few

scattered setae. Rostrum broadly triangular, terminating acutely or bluntly and often with terminal spine (but

absent in holotype). Lateral projections obtusely triangular, with strong submarginal spine.

Source :

PAGURIDAE FROM THE KARUBAR EXPEDITION

527

Ocular peduncles short, approximately 0.70 to 0.80 length of shield, appreciably broadened distally; corneae

dilated. Ocular acicles narrowly triangular, terminating acutely and with very small submarginal spine; separated

basally by basal width of one acicle.

Antennular peduncles overreaching corneae by 0.75 to entire length of ultimate segment. Ultimate segment

with 1 long and few shorter setae on dorsal surface. Penultimate segment with few short setae ventrally. Basal

segment with strong spine on laterodistal margin dorsally.

Antennal peduncle overreaching ocular peduncle (including cornea) by 0.50 to 0.65 length of ultimate segment;

with supernumerary segment. Fifth and fourth segments with few scattered setae. Third segment with strong spine

at ventrodistal angle. Second segment with dorsolateral distal angle very prominently produced, reaching to mid or

distal portion of fourth segment and terminating in very strong spine, usually with 1 or 2 accessory spines on both

mesial and lateral margins; dorsomesial distal angle with prominent, slender, acute spine. First segment frequently

with very small spine on lateral margin distally, ventrolateral margin with 1 prominent spine. Antennal acicle

long, reaching to distal half of ultimate peduncular segment, arcuate, terminating in acute spine, mesial margin

with sparse row of setae. Antennal flagellum usually slightly shorter than outstretched right cheliped, with 1 to

3 short setae every 2 to 6 articles.

Maxillule with external lobe of endopod moderately well developed, not recurved. Ischium of third maxilliped

with 1 accessory tooth on crista dentata. Sternite of third maxillipeds unarmed or with very small median spinule.

Chelipeds grossly unequal. Right cheliped (Fig. 42e) with dactyl somewhat shorter to approximately equaling

length of palm; cutting edge with row of calcareous teeth, terminating in small calcareous claw; dorsal surface

angularly convex, with scattered small spinules and few setae, midline with row of very small spines extending

nearly to tip; dorsomesial margin with row of small spines; ventral surface with few setae. Palm 0.75 to

approximately equaling length of carpus; dorsomesial margin with row of spinules and 1 more prominent tubercle

at dorsomesial proximal angle, convex dorsal surface generally smooth, faintly granular, or minutely spinulose,

but with weak longitudinal depression setting off distinctly spinulose or tuberculate mesial portion, dorsolateral

margin with row of small spines, obsolete in proximal fourth or third but increasing slightly in size distally and

extending almost to tip of fixed finger; mesial, lateral and ventral surfaces minutely spinulose or granular; cutting

edge of fixed finger with row of small calcareous teeth and 1 more prominent tooth medially, terminating in small

calcareous claw. Carpus approximately as long as merus; dorsomesial margin with irregular row of small to

moderately strong spines; dorsal surface with 3 irregular rows of smaller spines; dorsolateral margin not delimited,

dorsolateral surface with numerous short transverse spinulose ridges; distomesial and distolateral margins each with

row of small spines; ventral surface tuberculate. Merus triangular; 1 spine on dorsodistal margin, dorsal surface

with rows of short transverse spinulose ridges and short setae; ventromesial and ventrolateral margins each with

row of small spines, usually more acute laterally; ventral surface spinulose or tuberculate. Ischium unarmed or

with row of widely-spaced very small spinulose protuberances and few moderately long setae on ventral margin.

Left cheliped (Fig. 42f) with dactyl exceeding length of palm by 0.10 to 0.35 own length; cutting edge with

row of very fine corneous teeth, terminating in small corneous claw; dorsomesial margin with row of spinules at

least in proximal half, dorsal midline usually slightly elevated and often with row of spinules in proximal half;

dorsal, mesial and ventral surfaces with scattered moderately long setae, most numerous distally and ventrally.

Palm 0.40 to 0.65 length of carpus; triangular in cross-section, dorsal surface elevated in midline, but not forming

distinct ridge or crest, with irregular double row of very small spines and tubercles, extending onto proximal half

of fixed finger, dorsal surface of fixed finger frequently spinulose or weakly tuberculate; dorsolateral margin with

row of small or very small spines, usually not extending to tip of fixed finger; dorsolateral and dorsomesial

surfaces spinulose and strongly sloping ventrally, dorsomesial margin with irregular row of spinulose

protuberances or small spines. Carpus approximately as long as merus; dorsolateral margin with row of acute

spines, laterodistal margin with few spines or spinules dorsally; dorsodistal margin with strong usually double

spine, dorsomesial margin with row of smaller spines or spinulose protuberances; dorsal, mesial and lateral faces

often tuberculate and with scattered setae; ventromesial margin with row of widely-spaced low blunt or spinulose

protuberances, ventrolateral margin with row of very small spinulose tubercles, ventral surface tuberculate. Merus

with short transverse rows of setae on dorsal margin; ventromesial margin with row of small blunt spines.

528

p. a. McLaughlin

ventrolateral margin with row of more acute spines; mesial, lateral and ventral surfaces with scattered setae.

Ischium sometimes with row of small spinulose tubercles on ventromesial margin.

Ambulatory legs (Figs 25b-e) similar from left to right. Dactyls 1.65 to 1.85 longer than propodi; in dorsal

view, straight or very slightly twisted; in lateral view, slightly arched; terminating in small corneous claws; dorsal

F|C. 25. — Pagurus kaiensis sp. nov., a-f, h-j, holotype 6 (8.4 mm) from Karubar Stn DW 18; g, k, paratypc

V (4.0ram) from Stn CP 36. — 1, Pagurus compressipes syntype 6 (3.9 mm) (NHM 1881.31): a, shield and

cephalic appendages; b, right second pereopod (lateral view); c, dactyl of right second pereopod (mesial view);

d, left third pereopod (lateral view); e, dactyl of left third pereopod (mesial view); f, dactyl and propodus of left fourth

pereopod (lateral view); g, tip of dactyl and preungual process of right fourth pereopod (lateral view); h. anterior lobe

of sternite of third pereopods; i, coxae and sternite of fifth pereopods; j-1, telson. Scales equal 0.1 mm (e)

1.0 mm (f, h, k-1), 2.0 mm (j), 3.0 mm (i), and 5.0 mm (a-e).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

529

margins each with row of low protuberances and stiff bristles, increasing in length distally; lateral and mesial faces

each with shallow longitudinal sulcus at least in proximal half, lateral faces often also with dorsal and ventral row

of fine setae, mesial faces each with short row of small corneous spinules dorsally and few setae ventrally;

mesioventral or ventral surfaces each with 1 1 to 14 corneous spines, increasing in length distally. Propodi 1.30 to

1.50 longer than carpi; dorsal surfaces each with irregular row of spinulose protuberances or spinules, frequently

accompanied by very short spiniform bristles; ventrodistal margins each usually with 1 or 2 corneous spinules,

ventral surface with row of quite small corneous spinules, more numerous and closely-spaced on second pereopods

of holotype. Carpi 0.65 to 0.75 length of meri; each with row of small spines or spinulose tubercles becoming

strongest at distal angles and accompanied by very short, spiniform bristles and sparse short setae. Meri each with

series of transverse rows of short setae on dorsal surfaces; second pereopods each with 2 or 3 small spines on

ventromesial and ventrolateral distal margins, narrow ventral surface with irregular double row of small spines,

third unarmed. Ischia with few setae most abundant dorsally. Propodal rasp of fourth pereopods with single row of

corneous scales; dactyl with prominent preungual process (Figs 25f-g) at base of claw. Sternite of third pereopods

with few long setae on subsemiovate or subsemicircular anterior lobe (Fig. 25h).

Sternite of fifth pereopods (Fig. 25i) developed anteriorly as two subovate lobes separated by shallow median

groove, anterior margins each with tuft of moderately dense setae. Males with paired gonopores partially obscured

by row of moderately long setae; 3 unequally biramous unpaired pleopods, pleopods 3 and 4 with exopods

moderately well developed, endopods reduced, pleopod 5 with exopod moderately well developed, endopod

rudimentary or vestigial. Females with paired gonopores, 4 unpaired pleopods, pleopods 2 to 4 with both rami

well developed, pleopod 5 with exopod well developed, endopod markedly reduced.

Uropods asymmetrical; exopods and endopods both with well developed rasps. Telson (Figs 25j-k) with

transverse suture; posterior lobes strongly asymmetrical, separated by indistinct or very small median cleft;

terminal margins oblique, with marginal and submarginal spines, strongest at outer angles; lateral margins

sometimes with row of small spines, or at least left with short marginal chitinous plate.

COLOR (in preservative). — Distal halves of ocular peduncles and penultimate segments of antennular

peduncles orange. Chelipeds with overall faint orange tint, appreciably faded on chelae, but darker on carpi and

meri; meri also with splotches of white. Ambulatory legs each with longitudinal stripe of orange on lateral face of

carpus and on lateral, mesial, and ventral surfaces of dactyl.

Habitat. — Unknown.

Distribution. — Kai Islands, Indonesia; 210-268 m.

ETYMOLOGY. — The specific epithet is derived from the Kai Islands, the type locality of this species.

AFFINITIES. — Pagurus kaiensis does not appear closely allied to any of the other regional Pagurus species. In

small specimens, the general shape of the left chela approaches that of P. hirtimanus, but without the dense

setation of that species. In having long ambulatory dactyls and a single row of scales in the propodal rasp,

P. kaiensis resembles P. compressipes, but the two are immediately distinguished by the markedly different

armatures of the chelipeds (Figs 42g-h) and telson (Fig. 251) of the latter species.

Pagurus capsularis sp. nov.

Figs 26a-k

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn DW 49. 08°00'S. I32°59'E, 210-206 m,

29.10.1991: 1 6 (6.6 mm) (MNHN-Pg 5308). — Stn DW 50. 07°59'S, 133°02'E, 184-186 m, 29.10.1991: 1 6

(3.2 mm) (MNHN-Pg 5309).

Types. — The male (6.6 mm) (MNHN-Pg 5308) from Karubar station DW 49 is the holotype. The other

male is a paratype.

530

P. A. MCLAUGHLIN

DESCRIPTION. — Shield (Fig. 26a) longer than broad; anterior margin between rostrum and lateral projections

concave; anterolateral margins sloping; posterior margin truncate; surface with few tufts setae. Rostrum broadly

triangular, terminating acutely or with terminal spine. Lateral projections obtusely triangular, with submarginal

spine or spinule.

Ocular peduncles short, approximately 0.65 length of shield, slightly broader distally; corneae weakly dilated.

Ocular acicles narrowly triangular, terminating acutely and with very small submarginal spine; separated basally by

slightly more than half basal width of one acicle.

Antennular peduncles overreaching corneae by approximately 0.50 length of ultimate segment. Ultimate

segment with 1 long and few shorter setae on dorsal surface. Penultimate segment with few short setae. Basal

segment with numerous long stiff setae and strong spine on latcrodistal margin dorsally.

Antennal peduncle reaching nearly to distal margin of cornea. Fifth and fourth segments with few scattered

setae. Third segment with strong spine at ventrodistal angle obscured by tufts of long stiff setae. Second segment

with dorsolateral distal angle very prominently produced, reaching to distal half of fourth segment, terminating in

simple or bifid spine partially obscured by stiff setae; dorsomesial distal angle with acute spine, several long stiff

setae on mesial margin. First segment with small spine on lateral margin distally, ventrolateral margin with

1 prominent spine. Antennal acicle moderately long, reaching to distal half of ultimate peduncular segment, but

not overreaching distal margin of cornea; arcuate, terminating in acute spine, mesial margin with row of tufts of

stiff setae. Antennal flagellum longer than outstretched right cheliped, with 1 to 3 short or long setae every 4 to

8 articles.

Maxillule with external lobe of endopod obsolete. Ischium of third maxilliped with 1 accessory tooth on crista

dentata. Stemite of third maxillipeds unarmed.

Chelipeds grossly unequal. Right cheliped (Fig. 26b) (regenerating in paratype) with dactyl shorter than palm

and overlapped by fixed finger; cutting edge with row of low broad calcareous teeth and adjacent row of tufts of

short stiff setae; dorsal surface convex, with closely-spaced spinulose tubercles on mesial side of midline

proximally, distally only with few tufts of setae; dorsomesial margin with row of small tuberculate spines, mesial

surface with several low broad tubercles. Palm approximately equaling carpus in length; dorsomesial margin with

row of small spines, convex dorsal surface with irregular rows of small spines and tubercles, 1 more prominent

row of spines in midline, tubercles frequently with central corneous somewhat flask-shaped capsules (Fig. 26c);

dorsal surface of fixed finger with row of spinules and double row of tufts of setae adjacent to cutting edge, latter

with row of small calcareous teeth, terminating in strong calcareous claw; mesial face and lateral face dorsally with

low unmodified tubercles or protuberances; all surfaces with scattered short setae. Carpus approximately as long as

merus; dorsomesial margin with row of moderately strong spines and numerous long setae, dorsal surface with

scattered setae, 1 irregular row of few spines adjacent to dorsomesial margin, 1 short row of quite small spines in

midline and additional row adjacent to weakly delimited dorsolateral margin, distal margin with few spinules;

mesial and lateral faces with scattered setae. Merus with few short transverse rows of setae; ventromesial margin

with few small blunt spinules, but not ventrally produced into wing-like expansion; ventral surface with

unmodified spinulose tubercles, ventrolateral margin with row of acute spines. Ischium unarmed.

Left cheliped (Fig. 26d) with dactyl exceeding length of palm by approximately third own length; cutting edge

with row of corneous teeth, terminating in large corneous claw; dorsomesial margin not delimited; dorsal, mesial

and ventral surfaces with rows of tufts of moderately long stiff setae. Palm approximately half length of carpus;

triangular in cross-section, dorsal surface elevated in midline but not forming distinct ridge or crest, with row of

tuberculate spines extending onto proximal half of fixed finger; dorsolateral margin with row of spines, dorsolateral

and dorsomesial surfaces strongly sloping ventrally, each with covering of tubercles, most of which provided with

central corneous somewhat flask-shaped structure, more numerous laterally but not encompassing entire dorsal

surface of fixed finger, dorsomesial margin not delimited. Carpus approximately as long as merus; dorsolateral

margin with row of acute spines, laterodistal margin with row of spines extending onto ventrolateral margin

distally; dorsodistal margin with pair of strong spines, dorsomesial margin with row of equally strong spines;

mesial, lateral and ventral surfaces with numerous low sometimes spinulose or spinose protuberances and tufts of

long stiff setae. Merus with short transverse rows of stiff setae on dorsal margin; ventromesial margin with 1

prominent spine and few spinulose tubercles, partially masked by long stiff setae; ventrolateral margin with

PAGURIDAE FROM THE KARUBAR EXPEDITION

531

row of very strong acute spines and long stiff setae, ventral surface with spinulose tubercles and tufts of long stiff

setae; mesial and lateral with low protuberances and setae. Ischium unarmed.

Fig. 26. — Pagurus capsularis sp. nov„ a, f-h, k, paratype 8 (6.6 mm) from Karubar Stn DW 49; b-e, i, j, holotype 8

(3.2 mm) from Stn DW 50: a, shield and cephalic appendages; b, right cheliped (dorsal view); c, capsulate tubercle;

d, chela and carpus of left cheliped (dorsal view); e, right second pereopod (lateral view); f, left third pereopod

(lateral view); g, dactyl of left third pereopod (mesial view); h, dactyl and propodus of right fourth pereopod (lateral

view); i, anterior lobe of sternite of third pereopods; j-k, telson. Scales equal 0.5 mm (c, i), 1.0 mm (h, j), 3.0 mm

(b, d-e, k), and 5.0 mm (a, f-g).

Source :

532

P. A. MCLAUGHLIN

Ambulatory legs (Figs 26e-g) similar from left to right. Dactyls 1.35 to 1.50 longer than propodi; in dorsal

view, straight; in lateral view, slightly curved ventrally; terminating in moderately long corneous claws; dorsal

margins each with setae proximally and row of corneous spines, increasing in length distally; lateral faces with

dorsal and ventral row of fine setae; mesial faces each with row of corneous spines dorsally and setae ventrally;

ventral surfaces each with 7 to 12 strong corneous spines, increasing in length distally. Propodi 1.10 to

1.20 longer than carpi; dorsal surfaces each with row of low protuberances and tufts of stiff setae; ventrodistal

margins each with 1 or 2 corneous spinules, ventral surface with row of widely-spaced quite small corneous

spinules. Carpi 0.75 to 0.85 length of meri; each with dorsodistal spine and sparse row of tufts of setae; second

pereopods also with small spine on dorsal surface in proximal half. Meri and ischia with dorsal and ventral tufts of

setae. Propodal rasp of fourth pereopods (Fig. 26h) with multiple rows of sharp corneous scales; dactyl with

prominent preungual process at base of claw. Anterior lobe of sternite of third pereopods (Fig. 26i) subsemicircular

or roundly subrectangular, unarmed or with marginal spine and several long setae. Sternite of fifth pereopods

developed anteriorly as 2 somewhat flattened semirectangular lobes separated by shallow median depression,

anterior margins each with row of fine setae.

Females unknown. Males with 4 unpaired left pleopods; second small and uniramous in holotype, all 4

unequally biramous in paratype; exopods moderately well developed, endopods absent or rudimentary. Uropods

asymmetrical; exopods and endopods both with well developed rasps. Telson (Figs 26j-k) with transverse suture;

posterior lobes asymmetrical, separated by very small or faintly indicated median cleft; terminal margins oblique or

weakly concave, each with row of moderately slender spines, not reaching to, or not stronger at outer angles, nor

extending onto weakly calcified lateral margins.

COLOR (in preservative). — Ocular peduncles with patch or band of orange on dorsal surface proximally and

partial orange stripe on ventral surface. Ambulatory legs with orange at distal margins of meri, proximal and distal

margins of carpi and propodi and proximal margins of dactyls; distal tips of dactyls white.

Habitat. — Unknown.

Distribution. — Collected only in Tanimbar Islands, 184-210 m.

Etymology. — From the Latin capsula, meaning a case or box, and reflecting the presence of central

capsulate structures on many of the tubercles and spines on the dorsal surfaces of both chelae.

Affinities. — The shape of left chela in P. capsularis and the presence of four unpaired male pleopods is very

reminiscent of two other species found in the Maluku region, P. hirtimanus and P. pergramilatus (Haig & Ball,

1988). Pagurus capsularis is readily differentiated from P. hirtimanus by the lack of a dense covering of setae on

the chelae that is seen in the latter species. This new species is immediately distinguished from P. pergranulatus

by marked differences in the ventromesial margin of the merus of the right cheliped. This margin in

P. pergranulatus is developed into a distinct somewhat wing-like ventrally produced lobe and provided with a dense

tuft of setae; no such development of this margin occurs in P. capsularis. Differences in the shape and armature of

the right chelae in the two species, while also distinct, are less dramatic. In P. capsularis , the dorsodistal angle is

armed with spines, but not noticeably produced as a subacute lobe; the dorsal surface is armed with numerous

small spines and tubercles, with a somewhat irregular, more prominent row in the longitudinal midline. In

contrast, P. pergranulatus has the dorsodistal angle of the palm produced as a subacute lobe (cf. ALCOCK &

ANDERSON, 1897, pi. 31, fig. 1); the dorsal surface is provided with a series of granules each having what has

been described as an anterior concavity or central depression (HENDERSON, 1896). No reference has been made by

previous authors (e.g., Henderson, 1896; Alcock, 1905b; Haig & Ball, 1988) to distinctive capsulate

structures on the tubercles of the chelae of P. pergranulatus-, however, in the single specimen available for

examination (" Soela " station T 17/43, 19°47.4'S, 1 16°31.4'E, 56-57 m; MNT) structures similar to those described

for P. capsularis were observed. While the residual color patterns of the "Soela" specimen agree with those given

by Haig and Ball (1988) for P. pergranulatus, both clearly differ from the remnants of color in the specimens of

P. capsularis.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

533

The presence of a median row of spines on the dorsal surface of the right chela and three irregular rows of

spines on the carpus of P. capsularis suggests a similarity with P. samoensis (Ortmann, 1892), but the Karubar

species differs from ORTMANN's in having the dorsal surface of the right chela strongly convex, and having much

shorter antennal acicles, but longer dactyls of the ambulatory legs.

REMARKS. — The capsules occupying the central surface of the tubercles of the chelae in both P. capsularis

and P. pergranulatus are reminiscent of the structures seen in Nematopagurus spinulosensoris. However, those of

the former two species have a more ball-like base and elongate, slender and drawn-out distal portion, whereas the

capsules of N. spinulosensoris are stouter and have a more tear-drop appearance. Since P. capsularis is known only

from the holotype and single paratype, no attempt has been made to determine the internal structure of these

capsules; nevertheless, a subterminal opening, similar to that observed in N. spinulosensoris, does appear to be

present. The capsules of that species were shown to be highly complex structures (MCLAUGHLIN & LANE, 1975).

Pagurus haigae sp. nov.

Figs 27a-h. 43a-d

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands : stn CP 16, 05°17’S, 132°50'E, 315-349 m,

24.10.1991: 1 8 (18.6 mm) (MNHN-Pg 5310); 1 2 (12.1 mm) (MNHN-Pg 5310 bis). — Stn CP 26, 05°34'S, 132°52'E,

265-302 m, 26.10.1991: 1 8 (10.1 mm) (MNHN-Pg 5311); 1 8 (7.3 mm) (SNHM 4812). — Stn CP 27, 05°33'S,

132°51'E, 304-314 m, 26.10.1991: 1 8 (14.1 mm, molt) (POLIPI).

Tanimbar Islands : stn CC 41, 07°45'S, 132°42'E, 401-393 m, 28.10.1991: 1 8 (11.5 mm) (USNM 276014).

TYPES. — The male (18.6 mm) (MNHN-Pg 5310) from KARUBAR station DW 16 is the holotype. The other

specimens are paratypes.

DESCRIPTION. — Shield (Fig. 27a) longer than broad; anterior margin between rostrum and lateral projections

concave; anterolateral margins terraced; posterior margin truncate; surface with few tufts setae. Rostrum triangular,

terminating acutely, with or without terminal spine. Lateral projections triangular, with strong marginal or

submarginal spine.

Ocular peduncles moderately short, 0.50 to 0.75 length of shield, slightly broader distally; corneae weakly

dilated; dorsal surface frequently with row of sparse setae. Ocular acicles roundly triangular, terminating subacutely

and with strong submarginal spine; separated basally by 0.50 to nearly entire basal width of one acicle.

Antennular peduncles overreaching corneae by 0.25 to nearly entire length of ultimate segment. Ultimate

segment usually with 1 or 2 setae at dorsodistal margin and also often with row of short setae on dorsal surface.

Penultimate segment with very few short setae. Basal segment with hooked spine on laterodistal margin dorsally

and few setae mesially.

Antennal peduncle overreaching distal margin of cornea by 0.25 to 0.50 length of ultimate segment; with

supernumerary segment. Fifth and fourth segments with few scattered setae. Third segment with strong spine at

ventrodistal angle sometimes partially obscured by tufts of long stiff setae. Second segment with dorsolateral distal

angle prominently produced, reaching to or beyond distal half of fourth segment, terminating in simple or bifid

spine and with 3 to 5 spines on mesial margin sometimes partially obscured by thick setae; dorsomesial distal

angle with acute spine. First segment with spine on lateral margin distally, ventrolateral margin with 4 to 6 small

spines laterally and distally. Antennal acicle long, reaching to distal half of ultimate peduncular segment, and

considerably beyond distal margin of cornea; slightly sinuous, terminating in acute spine, mesial margin with row

of tufts of stiff setae. Antennal flagellum shorter than outstretched right cheliped, with 2 very short setae every

article and 1 to 3 longer setae every 4 to 8 articles.

Maxillule with external lobe of endopod vestigial. Ischium of third maxilliped with 1 accessory tooth on crista

dentata. Sternite of third maxillipeds with strong spine on either side of midline.

Chelipeds grossly unequal; spines of chelae and carpi usually with tiny corneous tips and most practically

obscured by tufts of long thick setae. Right cheliped (Figs 43a, c) with dactyl equal to length of palm; cutting edge

Source :

534

P. A. MCLAUGHLIN

FlG „27- -7 pagurus haigae sp. nov., a-g, paratype 5 (12.1 mm) from Karubar Stn CP 16; h. holotype 6 (18.6 mm) from

S'" CP 16' _ ’’ Pa8“rus y°k°yai Makarov, 1938, 6 (8.5 mm) from Uchibo coast of Boso Peninsula. Japan,

(CBM-ZC 1668): a, shield and cephalic appendages; b. right second pereopod (lateral view); c, dactyl of right second

pereopod (mesial view); d, left third pereopod (lateral view); e, dactyl of left third pereopod (mesial view); f, anterior

lobe of stermte of third pereopods; g-i, telson. Scales equal 2.0 mm (i), 3.0 mm (f-g), 5.0 mm (a-e. h).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

535

with row of 4 strong calcareous teeth and short distal row of corneous teeth, terminating in corneous claw and

slightly overlapped by fixed finger; dorsal surface convex, with median row of small spines decreasing in size

distally but extending nearly to tip; dorsomesial margin with row of spines also decreasing in size distally; mesial

and ventral surfaces with tufts of long thick setae. Palm 0.60 to 0.75 length of carpus; dorsomesial margin usually

only weakly delimited by quasi double row of strong spines, frequently 1 more prominent spine or tubercle at

dorsoproximal angle, convex dorsal surface with 8 or 9 irregular rows of strong spines; dorsolateral margin not

distinctly delimited except on fixed finger; dorsal surface of fixed finger with median row of spines decreasing in

size distally, cutting edge with few calcareous teeth, terminating in strong corneous claw; mesial, lateral and

ventral surfaces with low sometimes spinulose protuberances and tufts of long thick setae. Carpus slightly shorter

to as long as merus; dorsomesial margin with row of moderately strong spines, dorsal surface with few to

numerous small spines or low sometimes spinulose protuberances and tufts of long thick setae, distal margin with

row of spinules and few slightly larger spines; dorsolateral margin not delimited, lateral face with low sometimes

spinulose protuberances and tufts of long setae, laterodistal margin with row of small spines; mesial face with few

spines dorsally, scattered low protuberances and setae ventrally, mesiodistal margin with row of very small blunt

spines; ventral surface with scattered setae. Merus subtriangular; dorsodistal margin with 2 to 4 slender spines,

dorsal surface with few short transverse rows of setae; lateral face microscopically spinulose and with scattered fine

setae, ventrolateral margin with row of small spines not extending to proximal margin but frequently terminating

proximally in 1 or 2 larger blunt spines or tubercles; mesial face with numerous low protuberances and tufts of

long setae, ventromesial margin with few small spines, sometimes 1 or 2 larger tubercles at proximal angle;

ventral surface with scattered tufts of setae. Ischium unarmed, but with row of setae on distolateral and

ventromesial margins.

Left cheliped (Figs 43b, d) with dactyl 1.75 to more than twice length of palm; cutting edge with row of

corneous teeth, terminating in large corneous claw; dorsomesial margin not delimited or with 2 or 3 small spines

proximally, dorsal midline unarmed or with few spinules or spinulose tubercles in proximal half; dorsal, mesial

and ventral surfaces with rows of tufts of long thick setae. Palm slightly less than half length of carpus; triangular

in cross-section, dorsal surface with row of strong spines decreasing in size distally, usually extending to distal

half, occasionally nearly to tip, of fixed finger; dorsolateral margin with double row of strong spines, decreasing in

size and becoming single row on fixed finger; dorsolateral and dorsomesial surfaces strongly sloping ventrally,

lateral face with numerous strong spines; mesial face more frequently with smaller spines or spinulose tubercles,

dorsomesial margin with row of 3 or 4 spines or tubercles; ventral surfaces of palm, fixed finger and dactyl all with

tufts of long setae. Carpus approximately as long as merus; dorsolateral margin with row of acute spines,

dorsodistal margin with 1 strong spine, dorsomesial margin with row of smaller spines, strongest proximally;

laterodistal margin with few spines dorsally, lateral face with low frequently spinulose protuberances and long

setae, ventrolateral margin with row of spines and moderately dense row of long setae; mesial faces with numerous

tufts of long setae, ventromesial margin with 2 to 4 small, often blunt spines distally. Merus with short transverse

rows of long setae on dorsal margin; mesial face with few low protuberances and setae, ventromesial margin with

few small spines; lateral face spinulose, particularly ventrally, ventrolateral margin with row of very strong acute

spines sometimes interspersed with shorter spines and row of long setae, frequently 1 or pair of stronger acute

blunt spines on each margin proximally. Ischium with row of small spines on ventromesial margin.

Ambulatory legs (Figs 27b-e) with dactyls 1.20 to 1.45 longer than propodi; in dorsal view, very slightly

twisted; in lateral view, slightly curved ventrally; terminating in moderately long corneous claws; dorsal margins

each with 2 rows of long thick setae; lateral faces each with weak to prominent longitudinal sulcus and few setae

(second and third right), moderately dense but randomly placed long setae on third left; mesial faces each with faint

longitudinal sulcus (not shown in Figs 27c, e), second pair flanked dorsally and ventrally by long setae, sometimes

also with dorsal row of corneous spines, third pair with row of corneous spines often interspersed with tufts of

setae dorsally and medially; ventromesial surfaces each with 8 to 17 strong corneous spines, increasing in length

distally, but partially obscured by thick setae. Propodi 1.10 to 1.35 longer than carpi; dorsal surfaces each with

row of low transverse protuberances and tufts of setae; lateral faces each frequently with small tubercle at proximal

margin medially or dorsally, second and third right pereopods each with 2 or 3 longitudinal rows of sparse tufts of

setae, left third with entire surface covered (but not extremely densely) by short transverse rows of moderately short

536

P. A. MCLAUGHLIN

stiff setae; mesial faces with few scattered setae; ventrodistal margins each with 1 or 2 small corneous spinules,

ventral surfaces with row of tufts of setae, more numerous on left and much denser on third left. Carpi 0.75 to

nearly equaling length of meri; second pereopods each with row of strong spines partially obscured by long setae

on dorsal surface, third with dorsodistal spine, dorsal surface unarmed or often with 1 to several smaller spines

partially obscured by row of tufts of setae; lateral faces also with 2 or 3 longitudinal rows of sparse tufts of setae.

Meri each with several transverse rows of long setae dorsally and ventrally, second also with single or double row

of small spines on ventral margin. Ischia with dorsal and ventral tufts of setae, second each frequently with row of

small spines on ventral margin. Propodal rasp of fourth pereopods with triple row of corneous scales. Sternite of

third pereopods with few long setae on subsemicircular anterior lobe (Fig. 21V). Sternite of fifth pereopods

developed anteriorly as two somewhat flattened semirectangular lobes separated by shallow median depression,

anterior margins each with row of fine setae.

Males with 3 unequally biramous unpaired pleopods, all with exopods moderately well developed, endopods

rudimentary. Females with paired gonopores, 4 unpaired biramous pleopods. Uropods asymmetrical; exopods and

endopods both with well developed rasps. Telson (Figs 27g-h) with transverse suture; posterior lobes somewhat

asymmetrical, separated by small median cleft; terminal margins slightly to strongly oblique, each with row of

2 to 5 strong calcareous spines interspersed with smaller calcareous or corneous spines; lateral margins usually

with few to numerous corneous spinules and occasionally calcareous spines.

Color (in preservative). — General overall orange tint; somewhat mottled on shield. Antennal flagella

alternating series of 8 to 10 transparent articles followed by similar number of burnt-orange. Meri of chelipeds and

ambulatory legs with darker orange, but with white band on distal margin dorsally and laterally.

Habitat. — One specimen occupying Natica shell.

Distribution. — Collected in Kai and Tanimbar Islands, 265 to 401 m.

Etymology. — This species is dedicated to the late Janet Haig, who contributed so much to our knowledge

of anomuran fauna of the Indo-Pacific.

Affinities. — Pagurus haigae appears most similar to P. brachiomastus (Thallwitz, 1892) and P. yokoyai

Makarov, 1938. Like P. haigae, both of these species have strongly spinose and setose chelae; the carpi of the

ambulatory legs each carry a row of spines on the dorsal surface. Other characters shared by P. haigae and

P. brachiomastus include a pair of large tubercles on the ventral surface of the merus of the right cheliped, usually

densely setose lateral faces of the dactyl and propodus of the third left pereopod, and light or white colored band on

the distal portion of the meri of the chelipeds and ambulatory legs. However, the shortness of the dactyls and

spinose upper margins of the propodi of the ambulatory legs of P. brachiomastus, as well as its longer more

slender ocular peduncles, will immediately distinguish this species from P. haigae. As indicated by Miyake

(1978 : 98, text-fig. 37c) the dense setation on the lateral faces of the dactyl and propodus of the third left pereopod

is a character restricted to females of P. brachiomastus. No sexual dimorphism in this character was noted in the

Karubar specimens of P. haigae.

It is with P. yokoyai that P. haigae shares the greatest morphological likeness, and it would be impossible to

distinguish the two taxa based on the descriptions of Yokoya (1933) (as Eupagurus gracilipes Yokoya not

E gracilipes Stimpson, 1858) or Makarov (1938, 1962). Miyake's (1978) description provides much greater

detail, but only on the basis of color patterns and the illustrated telson (Text-fig. 45b) would differentiation be

teasible. Having compared the Karubar specimens of P. haigae with three male specimens of P. yokoyai there is

no doubt that the taxa are distinct. In addition to differences in the telson armature of the two species (Figs 27g-i),

the right cheliped of the latter species lacks the strong tubercuiate spines on the ventromesial margin of the merus,

the carpus is longer and narrower, and the spines of the dorsal surface of the chela are stronger. The left chela of

P. yokoyai is usually narrower and has fewer but stronger spines; the carpus is also narrower and the dorsomesial

margin more weakly armed. The second pereopods have five to eight corneous spines on the distal halves of the

ventral margins of the dactyls, and a double row of corneous spines on the mesial face dorsally.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

537

Remarks. — Yokoya's (1933) description of Eupagurus gracilipes consisted of little more than a few

illustrations and remarks on its differences from Pagurus pectinatus (Stimpson, 1858). Makarov (1938, 1962)

noted, in a footnote to his diagnosis of Stimpson's (1858) P. gracilipes that, Yokoya's (1933) name was

preoccupied, and proposed that the species be called P. yokoyai. Makarov (1938, 1962) did suggest that

P. yokoyai was probably more closely related to P. brachiomastus than to P. pectinatus, which, in fact is the

case. Pagurus haigae, like P. brachiomastus and P. yokoyai, shares numerous characters with P. pectinatus, such

as strongly spinose and setose chelae; however, the much more elongate ocular peduncles and stronger rostrum of

the latter species clearly set it apart from the former three species.

Miyake's (1978, Text-fig. 44) illustration of P. yokoyai is indicated as a female, but with only three unpaired

left pleopods. This is either an illustrator's error or the sex of the illustrated specimen is incorrect, as it is most

unlikely that the second left pleopod would be absent in this Pagurus species. One of the three specimens

examined [d (8.5 mm) from Kushimoto, Wakayama, Japan] had well developed male gonopores, but four unpaired

pleopods. However, there was no indication of rhizocephalan infestation that might have attributed to this apparent

feminization.

IPagurus sp.

Fig. 28a-d

MATERIAL EXAMINED. — Indonesia. KARUBAR, Kai Islands : stn DW 22, 05°22'S, 133°01'E, 124-85 m,

25.10.1991: 1 <3 (1.2 mm) (MNHN-Pg 5312).

DIAGNOSIS. — Shield (Fig. 28a) longer than broad. Rostrum obtusely triangular, reaching slightly beyond

level of lateral projections. Lateral projections each with tiny submarginal spinule. Ocular peduncles moderately

long; slightly shorter than extended antennal peduncles and reaching only to proximal half of ultimate segment of

fully extended antennular peduncles, dorsal and mesial surfaces with few tufts of stiff setae; corneae not dilated.

Ocular acicles moderately small, terminating subacutely and with strong submarginal spine. Antennal acicle

reaching to base of cornea or slightly beyond; arcuate; with terminal spine and few tufts of stiff setae. Antennal

flagella broken, but still reaching beyond distal margins of propodi of ambulatory legs, with 2 or 3 moderately

long setae every 2 to 4 articles. Crista dentata well developed, with 1 accessory tooth. Sternite of third pereopods

with slight median protuberance.

Right cheliped missing.

Left cheliped (Figs 28b-c) with palm slightly more than half length of dactyl. Dorsal surface of dactyl with row

of widely-spaced small spines in midline. Palm subtriangular in cross-section; dorsal midline with row of spines

extending to distal half of fixed finger; surface with numerous tufts of long setae, but not obscuring spines. Carpus

with dorsomesial and dorsolateral row of few strong spines; ventrolateral distal angle with acute spine; tufts of

long setae dorsally and ventrally. Merus with 2 strong spines on ventrolateral margin, 1 on ventromesial margin.

Ambulatory legs (Fig. 28d) (only second and third left) with dactyls only slightly longer than propodi; dorsal

surfaces with tufts of setae; ventral margins each with row of 6 or 8 corneous spines. Propodi each with

widely-spaced low protuberances and tufts of setae on dorsal surfaces; ventrodistal angles each with corneous spine,

ventral surfaces each with row of widely-spaced very small corneous spinules. Carpi each with dorsodistal spine

and few low protuberances with tufts on setae on dorsal surface, second also with 1 spine proximally, partially

obscured by setae. Meri with low protuberances and lufts of setae dorsally and ventrally; second also with spine on

ventral margin in distal half. Fourth pereopods semichelate; propodal rasp with single row of corneous scales.

Male with 3 unequally biramous left pleopods (third to fifth). Telson (Fig. 28e) with distinct transverse suture;

slightly asymmetrical posterior lobes separated by prominent median cleft; terminal margins somewhat oblique,

each with 4 moderately strong spines; lateral margins each with weakly serrated chitinous marginal band.

COLOR. — Unknown.

Habitat. — Unknown.

538

P. A. MCLAUGHLIN

Fig. 28. — IPagurus sp. <J (1.2 mm) from Karubar Stn DW 22: a, shield and cephalic appendages; b, left chela (dorsal

view); c, left cheliped (mesial view); d, left second pereopod (lateral view); e, telson. Scale equals 0.5 mm (e) and

1.0 mm (a-d).

Distribution. — Known only from one station (DW 22) in Kai Islands, Indonesia; 124-85 m.

Remarks. — This specimen is tentatively assigned to Pagurus as it lacks any secondary male sexual

characters. The armature of the left cheliped is consistent with that seen in many Pagurus species, as is the

structure and armature of the telson.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

539

Genus BATHYPAGUROPSIS McLaughlin, 1994

Bathypaguropsis McLaughlin, 1994: 469.

DIAGNOSIS. — Thirteen pairs of trichobranchiate gills. Shield with central dorsal surface sometimes only

weakly calcified; rostrum well developed. Ocular acicles triangular, dorsal surface flattened or slightly convex.

Antennal peduncle with supernumerary segmentation. Maxillule with external lobe of endopod articulated, not

recurved. Ischium of third maxilliped with well developed crista dentata and 1 accessory tooth.

Right cheliped massive, chela operculate or nearly so; propodal-carpal articulation approximately 30° from

perpendicular; dactyl articulating obliquely with palm. Left cheliped moderately elongate, slender; propodal-carpal

articulation approximately 30° to 60° counterclockwise from perpendicular; dactyl and fixed finger opening

obliquely. Ambulatory legs with dactyls and propodi similar. Fourth pereopods semichelate, propodal rasp

consisting of 1 or more, sometimes incomplete, rows of scales.

Males with paired gonopores, each partially masked by tuft of stiff setae; no paired pleopods or sexual tubes.

Four unpaired pleopods on left, with exopods only moderately well developed, endopods markedly reduced. Females

with paired gonopores. No paired pleopods; left second to fifth unpaired, second to fourth with both rami well

developed and egg-carrying, fifth reduced as in males.

Telson with transverse suture; posterior lobes subtriangular; terminal margins oblique, unarmed or spinulose.

Bathypaguropsis rahayuae sp. nov.

Figs 29a-h, 42c-d

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 14. 05°18'S, 132°38'E, 245-246 m.

24.10.1991: 1 6 (3.2 mm) (MNHN-Pg 5313).

TYPE. — The single specimen is the holotype.

DESCRIPTION. — Shield (Fig. 29a) longer than broad; anterior margin between rostrum and lateral projections

concave; anterolateral margins sloping; posterior margin truncate. Rostrum prominent, triangular, acute, with

terminal spinule. Lateral projections triangular, with small marginal spine. Posterior carapace with area between

cardiac sulci and sulci cardiobranchalis weakly calcified.

Ocular peduncles slightly more than half shield length, dorsomesial surface with row of setae; corneae not

dilated. Ocular acicles simple, triangular, with small marginal terminal spine; separated basally by width of

rostrum, or approximately half basal width of one acicle.

Antennular peduncles moderately long, overreaching ocular peduncles by slightly less than entire length of

ultimate segment. Basal segment with small acute spine on lateral surface dorsally. Penultimate segment with few

scattered setae. Ultimate segment with few long setae on dorsal surface.

Antennal peduncles exceeding ocular peduncles by half length of ultimate segment, but reaching only to distal

half of ultimate segment of antennular peduncle. Fifth and fourth segments with few scattered setae. Third segment

with very strong spine at ventrodistal margin. Second segment with dorsolateral distal angle produced into broad,

triangular process, terminating in acute spine, mesial margin unarmed, lateral margin with 1 or 2 spines;

dorsomesial distal angle with acute spine. First segment with small spine at laterodistal margin; ventral margin

produced, with 1 spine laterally. Antennal acicle reaching beyond proximal margin of ultimate peduncular segment;

slightly arcuate, with row of sparse tufts of setae on mesial margin and terminating in small spine. Antennal

flagellum long, nearly overreaching outstretched right cheliped; every 1 or 2 articles usually with 2 or 3 very short

(< 1 article length) setae, and often additional 1 or 2 longer setae every 2 to 5 articles.

Right cheliped (Figs 29b, 42c) massive, operculate. Dactyl broad, slightly shorter than palm; cutting edge with

1 very broad, faintly cusped, calcareous tooth, terminating in very small corneous claw; dorsal surface very slightly

elevated in the midline proximally, with scattered low tubercles and few short transverse setal ridges in proximal

Source :

540

P. A. MCLAUGHLIN

g-h

Fig. 29. — Bathypaguropsis rahayuae sp. nov., holotype 6 (3.2 mm) from Karubar Sin DW 14: a. shield and cephalic

appendages; b, carpus and chela of right cheliped (mesial view); c, right second pereopod (lateral view); d, dactyl of

second left pereopod (mesial view); e. third left pereopod (lateral view); f, dactyl of third right pereopod (mesial

view); g. propodus and dactyl of left fourth pereopod (lateral view); h, telson. Scales equal 1 .0 mm (g-h) and 2.0 mm

(a-f).

half, slightly pitted and with few tufts of setae distally; dorsomesial margin rounded, mesial face with low broad

tubercles, 1 large blunt tubercle proximally; ventromesial margin with corneous-capped blunt tubercles. Palm with

maximum breadth equal to length, nearly twice length of carpus; dorsomesial distal angle with prominent blunt

spine, dorsomesial margin with row of rather widely-spaced blunt or subacute tuberculate spines; dorsal surface

convex and covered with flattened granules and very low short ridges, with 1 rather inconspicuous tubercle at

proximal margin; dorsolateral margin only faintly delimited by row of widely-spaced tiny spinules, slightly larger

distally and on fixed finger; mesial face with scattered low spinules and spinulose tubercles or granules, particularly

Source : MNHN Paris

PAGUR1DAE FROM THE KARUBAR EXPEDITION

541

in distal half, distal margin with 2 blunt corneous-capped spines dorsally and 1 at ventral angle; lateral face with

scattered granules, low tubercles and/or short transverse ridges; ventral surface with several large, flattened, blister¬

like tubercles, often with corneous caps, few similar tubercles on ventral surface of fixed finger; cutting edge of

fixed finger with 3 calcareous teeth, terminating in very small corneous claw. Carpus slightly shorter than merus,

subquadrate when viewed dorsally; dorsomesial distal angle somewhat depressed but with adjacent large blunt spine,

dorsomesial margin with 1 moderately strong and 2 smaller spines in slightly irregular row, 1 additional strong

spine on dorsal surface mesially, remainder of dorsal surface with scattered spinules or low blunt or spinulose

tubercles, primarily in distal half; distal margin with 4 small spines extending onto lateral face; dorsolateral margin

not delimited, lateral and mesial surfaces with scattered minute spinulose tubercles or granules, ventral surface with

rounded unarmed distal ridge. Merus broadly subtriangular, armed on ventral surface by short row of blunt tubercles

on ventrolateral proximal margin and 2 tubercles in midline proximally. Ischium unarmed.

Left cheliped (Fig. 42d) not reaching to base of dactyl of right, slender; propodal carpal articulation

approximately 35° counterclockwise from perpendicular. Dactyl slightly longer than palm; surfaces unarmed but

with scattered tufts of short setae; cutting edges of dactyl and fixed finger each with row of small corneous teeth in

distal 0.50 to 0.65; terminating in small corneous claws. Palm little more than half length of carpus; all surfaces

unarmed, but with few scattered setae, particularly on fixed finger. Carpus slightly shorter than merus; dorsomesial

margin with 1 small spine at distal angle and 2 widely-spaced very small spines marginally; 1 tiny spinule at

dorsolateral distal angle, dorsolateral margin not delimited. Merus entirely unarmed. Ischium with 2 minute

spinules on ventromesial margin distally.

Ambulatory legs (Figs 29c-f) similar, but with right slightly longer than left. Dactyls 1.20 to 1.25 length of

propodi; in dorsal view, straight; dorsal margins each with row of tufts of long setae; mesial faces each with dorsal

and ventral sparse row of moderately long setae; lateral faces each with 1 row of very sparse tufts of short setae;

ventral margins each with row of 7 to 10 corneous spines and few tufts of setae. Propodi slightly longer than

carpi; each with few scattered setae on dorsal surface; ventrodistal angles each with 1 corneous spinule; mesial,

lateral, and ventral surfaces with scattered setae. Carpi 0.75 to 0.80 length of meri; dorsodistal angles of right

(only) each with very small spine, dorsal surfaces of all with few setae. Meri and ischia with scattered setae

particularly on ventral margins. Fourth pereopods with propodal rasp (Fig. 29g) consisting of row of long

corneous scales. Anterior lobe of sternite of third pereopods subrectangular, with central semicircle fringed with

short setae.

Telson (Fig. 29h) with posterior lobes slightly asymmetrical, left larger; separated by moderate median cleft;

terminal margins each with 5 very small spinules.

Color. — Unknown.

Habitat. — Unknown.

DISTRIBUTION. — Known only from the type locality in the Kai Islands, Indonesia; 245-246 m.

ETYMOLOGY. — This species is named for Dr D. L. RAHAYU, Indonesian Institute of Sciences, in recognition

of her contributions to our knowledge of the regional hermit crab fauna.

AFFINITIES. — Bathypaguropsis rahayuae sp. nov. is the third species of this genus to be recognized, and

despite the overall morphological similarities among the three taxa, it is most closely allied to B. yaldwyni

McLaughlin, 1994. Although the ocular peduncles are shorter and the antennal acicles longer in B. yaldwyni , the

general armature of the chelae and carpi of the chelipeds would be comparable if intraspecific variability in

B. rahayuae was to approach that of B. yaldwyni. The terminal margins of the telson of B. rahayuae are armed

with spinules as they are in B. yaldwyni , although the number in the single specimen of B. rahayuae is smaller.

Nevertheless, the two species can be easily distinguished by the spines on the ventral margins of the ambulatory

dactyls, 15 to 31 in B. yaldwyni , but only 7 to 10 in B. rahayuae. The smaller number of dactylar spines is similar

to that of B. marionensis McLaughlin, 1994 (8 to 14); however, the strongly produced dorsomesial distal angle and

more numerous marginal spines of the right chela, as well as the armed ventromesial margin of the left, clearly

distinguish this species from B. rahayuae.

542

P. A. MCLAUGHLIN

Remarks. — McLaughlin (1994) noted that one adult specimen of B. yaldwyni was parasitized by an

unidentified rhizocephalan, and while female appearing pleopods were present, neither male nor female gonopores

could be detected. The holotype of B. rahayuae similarly had been infected by an unidentified rhizocephalan, but no

evidence of structural alteration nor feminization has been observed. Only normal male gonopores are present.

Genus PYLOPAGUROPSIS Alcock, 1905

Pylopaguropsis Mcock. 1905b: 133. — de Saint Laurent-Dechanc6, 1966b: 259. — McLaughlin & Haig. 1989: 125.

Galapagurus Boone, 1932: 12.

DIAGNOSIS. — Thirteen pairs of trichobranchiate gills. Shield with well developed rostrum. Ocular acicles

triangular, sometimes slender. Antennal peduncles with supernumerary segmentation. Maxillule with external lobe

of endopod very weakly to moderately well developed. Ischium of third maxilliped with 1 accessory tooth on well

developed crista dentata.

Right cheliped usually massive, chela operculate or semioperculate; dactyl frequently articulating obliquely

with palm.

Left cheliped moderately elongate, slender; propodal-carpal articulation usually twisted counterclockwise 30-70°

from perpendicular when viewed dorsally; dactyl and fixed finger opening obliquely. Ambulatory legs with dactyls

and propodi of second pair (third pereopods) frequently dissimilar. Fourth pereopods semichelate; propodal rasp of

1 to 4 rows of corneous scales.

Females with paired gonopores; paired first pleopods modified as gonopods, second to fifth unpaired. Males

with paired gonopores, no paired pleopods or sexual tubes, 3 unequally biramous left pleopods. Telson with

transverse suture; posterior lobes often asymmetrical; terminal margins oblique, concave or horizontal, usually

armed with 1 to many spines; lateral margins unarmed or with 1 to 3, or sometimes row of small spines.

Remarks. — Haig and Ball (1988) reported two undescribed species of Pylopaguropsis from the

Alpha Helix expedition. Although both species, P. lewinsohni McLaughlin & Haig, 1989 and P. fimbriata

McLaughlin & Haig, 1989, had been recognized much earlier, their actual publication had been unavoidably

delayed. Neither species is represented in the Karubar material.

Key to the Indonesian species of Pylopaguropsis

1. Palm of right chela fringed with spines and long setae; carpus of fourth pereopod with

dorsodistal spine . P. fimbriata *

— Palm of right chela not fringed with spines and long setae; carpus of fourth pereopod

without dorsodistal spine . 2

2. Left chela with 1 or more rows of spines on dorsal surface . P. laevispinosa

— Left chela unarmed or with few scattered spinules or spinulose tubercles on dorsal surface . .

. 3

3. Propodus of right third pereopod with 1 longitudinal sulcus on lateral face . P. zebra

— Propodus of right third pereopod with 3 longitudinal sulci on lateral face .

. P. lewinsohni*

Pylopaguropsis zebra (Henderson, 1893)

Figs 30a, c, 43e-f

Eupagurus zebra Henderson, 1893: 425, pi. 39, figs 12-15.

Pagurus zebra - MIYAKE, 1975: 260, pi. 116, fig. 2; 1978: (in part) 108, fig. 43; 1982: 225.

Pylopagurus zebra - McLaughlin & Haig, 1989: 143, figs 3b, 5b, 7b, 9b, 1 lb, 13b (for complete synonymy).

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

543

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 15, 05°17'S, 1 32°4I'E, 212-221 m,

24.10.1991: 1 $ (2.7 mm) (POLIPI). — Stn DW 18, 05°18'S, 133°01'E, 205-212 m, 24.10.1991: 1 6 (1.8 mm) (USNM

275998).

Tanimbar Islands: stn DW 49, 08°00'S, 132°59'E, 206-209 m, 29.10.1991: 2 <? , 1 2 (3.3-3.S mm) (MNHN-Pg

5314).

DIAGNOSIS. — Shield (Fig. 30a) approximately as broad as long or slightly longer than broad. Rostrum

prominent, acute, terminating in small spinule. Ocular peduncles overreached by antennular peduncles. Ocular

acicles triangular, acute; separated basally by breath of rostrum.

Right chela (Fig. 43e) with dactyl compressed dorsoventrally; dorsomesial margin expanded and armed with row

of strong spines, dorsal surface with several spines at dorsomesial proximal angle, few also on dorsal surface. Palm

with irregular single or double row of low spines or spinulose tubercles on dorsomesial margin, dorsolateral

margin with row of strong spines; dorsal surface with few irregular rows of small spines or spinulose tubercles,

extending onto fixed finger. Carpus trapezoidal (dorsal view), with regular or irregular rows of spines or spinulose

tubercles on dorsal surface, few strong spines on dorsodistal margin; mesial face strongly produced ventrally,

ventromesial margin tuberculate. Merus with row of strong conical spines on produced ventromesial margin;

mesial, lateral and ventral surfaces usually tuberculate.

Left cheliped (Fig. 430 long and slender, reaching almost to middle of palm of right. Palm and fingers twisted

counterclockwise approximately 45° from perpendicular. Dactyl unarmed. Palm with scattered low protuberances,

some occasionally slightly spinulose. Carpus with row of small to moderately strong spines on dorsomesial

margin, 1 prominent spine on dorsolateral distal angle and 1 or more spines, spinules or low protuberances on

dorsolateral margin. Merus with row of small spines on ventromesial margin; ventrolateral margin with row of

acute spines.

Ambulatory legs with second pair (third pereopods) dissimilar. Dactyls each with row of corneous spines on

mesial face close to dorsal margin; ventral margins each with row of 8-10 strong corneous spines. Propodi each

with 1 or 2 corneous spinules at ventrodistal margin. Third right pereopod with dactyl longer than left, 1.50 to

twice as deep; laterally compressed; lateral face with prominent longitudinal sulcus; ventral margin with row of

13 or 14 strong corneous spines. Propodus with distinct dorsolateral margin, lateral face with shallow

longitudinal sulcus dorsally, surface broad and flattened medially. Carpi ol both second and third pereopods each

with small spine at dorsodistal margin. Meri unarmed or with small spine on ventrolateral distal margin. Sternite

of third pereopods with long narrow anterior lobe slightly protruded medially. Propodal rasp of fourth pereopods

with 1 row of sharp corneous scales.

Telson (Fig. 30c) with posterior lobes separated by broad median cleft; terminal margins oblique, each with

3 or 4 strong spines, lateral margins with narrow corneous plate.

COLOR (in preservative). — Ocular peduncles with 2 longitudinal red stripes dorsally and 1 ventrally on white

base color. Right cheliped with base color of pinkish-white on palm and carpus, red on merus; palm and carpus

with scattered red spots, usually associated with tufts of setae, carpus also with broad longitudinal red stripe on

mesial face; merus with several narrow longitudinal white stripes. Left cheliped with base color of pinkish-white;

palm with two longitudinal red stripes and I prominent red spot on dorsal surface; carpus and merus each with

dorsal, lateral and mesial longitudinal red stripes. Ambulatory legs each with longitudinal red stripes on lateral and

mesial faces of all segments, carpi also with dorsal red stripe.

Habitat. — Unknown.

Distribution. — Northwestern Australia, Indonesia; South Africa, Japan; 102-125 meters.

Remarks. — The Karubar specimens differ from the lectotype and other specimens described by

McLaughlin and Haig (1989) in having the carpus of the left cheliped less strongly armed. In the present

specimens, the telsons all have strongly oblique terminal margins. The telson of the lectotype was reported

missing by McLaughlin and Haig (1989); their illustrated specimen from Japan had horizontal terminal

margins.

Source :

544

P. A. MCLAUGHLIN

Fig. 30. — Pylopaguropsis zebra (Henderson, 1893), a, c, $ (3.3 mm) from Karubar Stn DW 49. — Pylopaguropsis

laevispinosa McLaughlin & Haig, 1989, b, d, 6 (3.2 mm) from Stn DW 22: a-b, shield and cephalic appendages;

c-d, telson. Scales equal 1.0 mm (c-d) and 2.0 mm (a-b).

Pylopaguropsis laevispinosa McLaughlin & Haig, 1989

Figs 30b, d, 44a-b

Pylopaguropsis laevispinosa McLaughlin & Haig, 1989; 166, figs 4e, 6e, 8f, lOe, 12e, 13j, 21.

PAGURIDAE FROM THE KARUBAR EXPEDITION

545

MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 22, 05°22'S,133° Ol'E, 85-124 m,

25.10.1991: 3 6 (1. 2-3.3 mm) (MNHN-Pg 5315). — Stn DW 30, 05°39'S, 132°56'E, 111-118 m, 26.10.1991: 1 9

(2.3 mm) (MNHN-Pg 5316).

Tanimbar Islands: stn DW 50, 07°59'S, 133°02’E, 184-186 m, 29.10.1991: I ov. 9 (1.2 mm) (POLIPI).

DIAGNOSIS. — Shield (Fig. 30b) longer than broad. Rostrum obtusely triangular; terminating in small

spinule. Ocular peduncles overreached by antennular peduncles. Ocular acicles triangular, acute; separated by

slightly more than basal width of one acicle.

Right chela (Fig. 44a) somewhat dorsoventrally compressed; dactyl as long as palm or slightly shorter,

articulation only slightly oblique; dorsomesial margin with row of strong spines, dorsal surface with scattered

spines; ventromesial margin with row of spinulose tubercles. Palm with irregular single or double row of

moderately strong spines on dorsomesial margin, dorsal surface of palm and fixed finger with numerous spines,

sometimes forming irregular rows; dorsolateral margin with row of spines, strongest on fixed finger; ventral

surface convex, tuberculate laterally and granular medially. Carpus with row of strong spines mesially and laterally

on dorsal surface and irregular transverse row of strong spines on distal margin, extending onto mesial and lateral

faces. Merus with row of acute spines on ventrolateral margin; ventromesial margin with row of acute spines,

ventroproximal margin with prominent blunt tubercle.

Left cheliped (Fig. 44b) long, slender; chela twisted counterclockwise 30° to 40° from perpendicular. Dactyl

with row of small spines on dorsal surface; dorsomesial margin with row of strong spines, not extending to tip.

Palm with 2 irregular rows of strong spines in midline of sloping dorsal surface, 1 extending onto fixed finger as

small spinulose tubercles, dorsal surface mesially and laterally each with row of moderately strong spines; ventral

surface with 1 row of small spines laterally in distal portion of palm and proximal portion of fixed finger. Carpus

with 2 rows of corneous-tipped spines on dorsal surface, strongest distally. Merus with row of strong acute spines

on ventrolateral margin; ventromesial margin with row of smaller subacute spines, prominent spinulose tubercle at

ventromesial proximal angle.

Ambulatory legs generally similar. Dactyls long and slender; ventral margins each with row of strong,

corneous spinules. Propodi with lateral faces evenly convex, unarmed. Carpi each with dorsodistal spine and

at least 2 or 3 (male) or row (female) of small spines (second) or with only dorsodistal spine (third). Sternite

of third pereopods with anterior lobe subsemicircular. Propodal rasps of 4th pereopods with 1 row of corneous

scales.

Telson (Fig. 30d) with asymmetrical posterior lobes separated by shallow median cleft; terminal margins

oblique, each with few to several small spines, sometimes extending onto lateral margins, particularly on left.

COLOR (in preservative). — Shield orange tinged, rostral margin accentuated in dark orange. Ocular

peduncles cream; acicles with margins accentuated in orange. Antennular peduncles cream, with faint

yellowish brown in distal third of ultimate segment. Antennal peduncles with orange stripe on ultimate

segment dorsally and on mesial face ventrally; fourth segment orange and white striped dorsally and white

ventrally; acicle with broad orange longitudinal stripe on dorsal surface. Right cheliped with chela faint orange or

cream-colored; carpus generally with faint orange hue, mesial, lateral and dorsal surfaces orange with white stripe

proximally. Merus orange and white striped dorsally and on dorsal halves of mesial and lateral faces, ventral surface

with faint orange hue. Left cheliped with chela very faint orange; carpus and merus with orange and white

longitudinal stripes. Second and third pereopods with orange and white longitudinal stripes (McLaughlin &

Haig, 1989).

Habitat. — Unknown.

Distribution. — Okinawa and Indonesia; 3-125 m.

Remarks. — Pylopaguropsis laevispinosa was described from two females collected in Okinawa, Ryukyu

Islands. The present diagnosis has been modified to reflect the variations exhibited by large males of this species.

The presence of P. laevispinosa in the Kai and Tanimbar Islands of Indonesia represents a major southern extension

of the range of this species.

Source :

546

P. A. MCLAUGHLIN

Genus TOMOPAGUROPSIS Alcock, 1905

Tomopaguropsis Alcock, 1905b: 136.

DIAGNOSIS. — Thirteen pairs of trichobranchiate gills. Shield with well developed rostrum. Ocular acicles

triangular. Antennal peduncle with supernumerary segmentation. Maxillule with external lobe of endopod well

developed, recurved. Third maxilliped with well developed crista dentata and 1 accessory tooth. Sternite of third

maxillipeds with prominent spine on either side of midline.

Chelipeds subequal; right usually somewhat more robust. Ambulatory legs similar from left to right; carpus

with or without dorsodistal spine. Fourth pereopod semichelate; propodal rasp with several rows of corneous

scales. Fifth pereopods chelate.

Males with coxae of Fifth pereopods symmetrical; paired gonopores; with or without paired first pleopods

modified as gonopods, 4 unpaired unequally biramous left pleopods. Females with paired gonopores; no paired

pleopods, 4 unpaired pleopods, second to fourth usually with both rami well developed, fifth reduced. Telson with

transverse suture; posterior lobes separated by median cleft; terminal margins spinose.

TYPE Species herein selected. — Tomopaguropsis lantana Alcock, 1905b.

REMARKS. — ALCOCK (1905b) established Tomopaguropsis to accommodate his new species Tomopaguropsis

lantana , and Eupagurus Iproblematicus A. Milne Edwards & Bouvier, 1893, an Atlantic species. In his original

diagnosis of Tomopaguropsis , ALCOCK (1905b) reported that males were provided with a small pair of pleopods on

the first abdominal somite. Although no mention was made of paired male pleopods in the original description of

E. Iproblematicus , ALCOCK's assignment of this taxon to Tomopaguropsis presumably was based on

MILNE Edwards and Bouvier's (1893: 153) subsequent remark: "Cette espece se distingue des Eupagurus par

ses panes anterieures, dont les doigts sont mobiles dans un plan oblique, et par ses lamelles branchiales qui sont

profondement bifides a l'extremite. Ces deux caracteres, et la plupart des autres, la rapprochent des Parapagurus, et

Ton trouve meme chez le male deux fausses pattes sexuelles anterieures, reduites a l'etat de bourgeons tres courts,

ainsi que la trace d'une fausse patte sexuelle de la 2^me paire".

The presence of paired first pleopods modified as gonopods in Tomopaguropsis was the character that

Alcock (1905b) likened to the condition reported by Milne Edwards and Bouvier (1893) for their genus

Tomopagurus A. Milne Edwards & Bouvier, 1893. However, PROVENZANO (in FOREST & DE SAINT LAURENT,

1968) noted that normal males of Tomopagurus lacked paired first pleopods, and proposed that the presence of

paired first pleopods in the holotype of Tomopagurus rubropunctatus A. Milne Edwards & Bouvier, 1 893, was due

to the feminizing effect of a rhizocephalan of the genus Peltogaster; females of Tomopagurus were known to have

paired first pleopods. DE Saint Laurent (1970b) suggested that variation in the presence of paired pleopods in

both sexes probably occurred not only in Tomopagurus but in Tomopaguropsis as well. To demonstrate this

variation in Tomopaguropsis , she referred to an undescribed species which lacked male gonopods; in Tomopagurus

she cited the notation by Provenzano (in Forest & de Saint Laurent, 1968). McLaughlin's (1981a)

description of the new species, Tomopagurus wassi McLaughlin, a species lacking female paired first pleopods,

and her (MCLAUGHLIN, 1981b) description of feminization in a male of Rhodochirus McLaughlin, 1981a,

supported DE SAINT Laurent's suggestion for Tomopagurus. I have not reviewed the holotype of

Tomopaguropsis problematica , but I have examined four male specimens (UMML 4975, 4978-80); all lack paired

first pleopods. Similarly, males of T. crinita sp. nov. lack paired pleopods. In all other respects, males of both

species clearly are assignable to Tomopaguropsis.

ALCOCK (1905b) did not designate a type species for Tomopaguropsis , and the only other report of T. lantana

is that of Kemp and Sewell (1912) who noted the collection of one additional specimen during the 1910-1911

cruise of R.I.M.S.S. "Investigator". Although T. problematica was listed first in ALCOCK's (1905b) remarks, and

has been reported more frequently (e.g., Williams, 1984; Abele & KlM, 1986) it is probable that his generic-

diagnosis was based principally on the Indian taxon. Tomopaguropsis lantana, therefore, is herein selected as the

type species. Whether or not paired first pleopods in males of Tomopaguropsis is a normally variable condition

will only be determined when several males of T. lantana have been examined.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

547

Tomopaguropsis crinita sp. nov.

Figs 31a-g, 44c-d

MATERIAL EXAMINED. — Indonesia. KARUBAR , Kai Islands : stn CC 10, 05°21'S, 132°30'E, 329-389 m,

23.10.91: 1 <3 (5.4 mm) (USNM 276012). — Stn DW 30, 05°39'S, 132°56’E, 111-118 m, 26.10.1991: I <3 (3.7 mm)

(USNM 276030). — Stn CP 59, 08°20'S, 132°11'E, 405-399 m, 31.10.1991: 1 2 (4.7 mm) (MNHN-Pg 5317).

Tanimbar Islands: stn CP 69, 08°42'S, 131°53'E, 356-368 m, 2.11.1991: 1 <3 (4.1 mm) (MNHN-Pg 5318). —

Stn DW 77, 08°57'S, 131°27'E, 346-352 m, 3.11.1991: 1 <3 (4.3 mm) (MNHN-Pg 5319); 1 <3 (2.4 mm) (POLIPI).

Types. — The male (4.3 mm) (MNHN-Pg 5319) from Karubar station DW 77 is the holotype. The other

specimens are paratypes.

DESCRIPTION. — Shield (Fig. 31a) subtriangular, longer than broad; anterior margin between rostrum and

lateral projections weakly concave; anterior margins sloping; posterior margin truncate; dorsal surface frequently

only weakly calcified in midline, somewhat rugose laterally and with few tufts of setae. Rostrum broadly

triangular, produced slightly beyond bases of ocular acicles; with very small terminal spinule. Lateral projections

triangular, terminating subacutely, unarmed or with tiny spinule.

Ocular peduncles short and moderately stout; dorsomesial surface with row of long setae; corneae small, not

dilated. Ocular acicles elongate and subtriangular, dorsally rounded, with tuberculate terminal spine and few setae

distally; approximate, or separated basally by less than 0.30 basal width of one acicle.

Antennular peduncles overreaching distal margins of corneae by 0.70 to entire length of ultimate segment.

Ultimate segment with row of long setae on dorsodistal margin. Penultimate segment glabrous or with very few

scattered setae. Basal segment with small spine on dorsolateral margin medially.

Antennal peduncles overreaching distal margins of corneae by 0.25 to 0.60 length of ultimate segment.

Fifth segment with few setae. Fourth segment with spinule at dorsodistal margin. Third segment with strong

ventrodistal spine and long setae. Second segment with dorsolateral distal angle produced, terminating in bifid or

simple spine, dorsal surface and lateral margin with long setae; dorsomesial distal angle with small spine.

First segment usually with small spine on dorsolateral distal margin; ventral margin produced, unarmed. Antennal

acicle moderately long, reaching nearly to distal margin of ultimate peduncular segment or sometimes well beyond;

slightly arcuate; mesial and lateral margins and dorsal surface distally with very long setae; terminating in simple

spine. Antennal flagellum moderately long, frequently reaching beyond tips of outstretched chelipeds; each article

with several very long (> 6 articles length) setae.

Chelipeds subequal, left occasionally longer, but right slightly more robust; dactyls and fixed fingers horizontal

or arched ventrally. Right cheliped (Figs 31b, 44c) with dactyl slightly shorter to slightly longer than palm;

dorsomesial margin not delimited; dorsal, mesial and ventral surfaces all with numerous tufts of long setae; cutting

edge with 1 or 2 large calcareous teeth proximally, row of corneous teeth in distal half, terminating in corneous

claw. Palm slightly shorter than carpus; usually with row of small spines on proximal 0.60 of dorsomesial

margin, occasionally with only single spine at proximal angle; dorsal surfaces of palm and fixed finger with

numerous rows of tufts of long setae and rarely 1 or 2 very small spinules in proximal third, dorsolateral margins

not delimited; lateral and ventral surfaces with numerous tufts of long setae; cutting edge of fixed finger with

calcareous teeth proximally and row of fused or individual corneous teeth distally. Carpus approximately equaling

length of merus; dorsomesial margin with row of spines, strongest distally, dorsodistal margin usually with 1 or

2 spines or spinulose tubercles, dorsolateral margin not delimited; surfaces all with long or short transverse

sometimes slightly protuberant rows of long setae. Merus with short transverse rows of long setae on dorsal

margin; dorsodistal margin with 2 or 3 spines; lateral face usually with scattered low, sometimes spinulose

protuberances and tufts of setae; ventromesial and ventrolateral distal angles each with 1 distal spine and

occasionally second spinule or spinulose protuberance on margin distally. Ischium with tuberculate ridge on

lateroproximal margin, laterodistal margin usually with 2 or 3 tiny spinules; ventromesial margin with few low

protuberances and tufts of short setae.

Left cheliped (Figs 31c, 44d) with dactyl slightly shorter to slightly longer than palm; often short hiatus

between dactyl and fixed finger proximally; dorsomesial margin of dactyl not delimited; dorsal, mesial and ventral

Source :

548

P. A. MCLAUGHLIN

Fig. 31. — Tomopaguropsis crinita sp. nov., holotype 6 (4.3 mm) from Karubar Stn DW 77: a, shield and cephalic

appendages; b. right cheliped (dorsal view); c. left cheliped (dorsal view); d, right second pereopod (lateral view);

e, left third pereopod (lateral view); f, dactyl and propodus of left fourth pereopod; g, telson. Scales equal

1.0 mm (f-g) and 3.0 mm (a-e).

surfaces all with numerous tufts of long setae; cutting edge with row of corneous teeth, terminating in corneous

claw. Palm slightly shorter than carpus, dorsomesial margin with 1 spine at proximal angle and sometimes few

Source : MNHN Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

549

additional small spines on margin in proximal half, dorsal surfaces of palm and fixed finger with numerous

irregular rows of long setae, dorsolateral margins not delimited; lateral and ventral surfaces with numerous tufts of

long setae; cutting edge of fixed finger with row of very small calcareous teeth proximally, corneous teeth distally.

Carpus approximately equaling length of merus; dorsodistal margin with 1 or 2 spines or spinules, dorsomesial

distal angle with strong spine and 1 to 6 spines on dorsomesial margin, dorsolateral margin not delimited; surfaces

with short transverse, sometimes slightly protuberant rows of long setae. Merus with short transverse, somewhat

protuberant or spinulose rows of long setae on dorsal margin; dorsodistal margin with 1 or 2 small spines;

ventromesial and ventrolateral distal angles each with 1 distal spine or spinule and occasionally second spinule or

spinulose protuberance on margin distally; lateral surface frequently with low, sometimes spinulose protuberances

in ventral half. Ischium with tuberculate ridge on lateroproximal margin, ventrolateral distal margin with 2 or

3 spinules; ventromesial margin with 1 or 2 spinulose protuberances and tufts of setae.

Ambulatory legs (Figs 3 ld-e) with dactyls 1.25 to nearly twice length of propodi; in dorsal view, slightly

twisted; in lateral view, curved ventrally; all surfaces, but particularly dorsal and ventrolateral margins, with rows

of long setae; mesial faces each with row of tufts of long setae; ventral margins also with 9 to 25 short, fine

corneous spines. Propodi with transverse rows of long setae dorsally; scattered setae on mesial, lateral and ventral

surfaces; ventromesial distal angles usually with 1 or 2 corneous spines. Carpi each frequently with spine at

dorsodistal angle; surfaces, particularly dorsal and lateral, with numerous tufts of setae. Meri and ischia each with

short transverse sometimes slightly protuberant rows of long setae particularly dorsally. Fourth pereopods without

prominent preungual process at base of claw (Fig. 310- Anterior lobe of sternite of third pereopods subquadrate,

with convex median, marginally setose, elevation.

Abdomen of males with 4 unpaired, unequally biramous pleopods; endopods well developed, exopods

approximately half length of endopods. Posterior margin of sixth abdominal somite with lateral angles each drawn

out into prominent simple, bi- or trifid spine. Telson (Fig. 3 1 g) with deep transverse suture; posterior lobes nearly

symmetrical, separated by narrow to moderately broad median cleft; terminal margins rounded, armed with several

blunt or acute, sometimes corneous-tipped spines, extending onto lateral margins.

Color. — Unknown.

Habitat. — Unknown.

Distribution. — At present known only from the islands of Kai and Tanimbar, Indonesia; 1 1 1 - 346 m.

ETYMOLOGY. — From the Latin crinitus meaning long-haired, and referring to the very long setae of the

antennal acicles, ocular peduncles, chelipeds and ambulatory legs of this species.

AFFINITIES. — In having quite dense setation on the chelipeds and ambulatory legs, as well as elongate and

very prominent ocular acicles, T. crinita agrees with the description given by ALCOCK (1905b) for T. lantana.

However, T. crinita lacks the rows of spinules on the chelae reported for T. lantana. ALCOCK's (1905b, pi. 13,

fig. 4) depicts a dorsodistal spine on the carpus of each ambulatory leg of T. lantana-, T. crinita lacks any carpal

spine. In his generic diagnosis, ALCOCK attributed paired first pleopods modified as gonopods to males of

Tomopaguropsis, and these are depicted for T. lantana (ALCOCK, 1905b, pi. 13, fig. 4a). As previously noted,

male gonopods are not present in T. crinita, nor in the Atlantic species, T. problematica.

Tomopaguropsis miyakei sp. nov.

Figs 32a-h, 44e-f

MATERIAL EXAMINED. — Indonesia. KARUBAR. Kai Islands: stn DW 35, 06°08'S, 132°45'E, 390-502 m.

27.10.1991: 1 9 (2.2 mm) (MNHN-Pg 5320).

Type. — The single specimen is the holotype.

550

P. A. MCLAUGHLIN

DESCRIPTION. — Shield (Fig. 32a) approximately as long as broad; anterior margin between rostrum and lateral

projections weakly concave; anterior margins sloping; posterior margin truncate; dorsal surface only weakly

calcified centrally, with few setae laterally. Rostrum triangular, produced slightly beyond bases of ocular acicles;

with small terminal spine. Lateral projections broadly triangular, terminating subacutely, with spine or spinule.

Ocular peduncles moderately short; dorsomesial surface with few long setae; corneae not dilated. Ocular acicles

rather short, subtriangular, dorsally rounded, with terminal spine and few setae distally; separated basally by more

than half basal width of one acicle.

Antennular peduncles overreaching distal margins of corneae by approximately 0.75 length of ultimate

segment. Ultimate segment with 1 or 2 long setae on dorsodistal margin and 2 or 3 on dorsal margin. Penultimate

segment with few scattered setae. Basal segment with prominent spine on dorsolateral margin medially.

Antennal peduncles slightly overreaching distal margins of corneae. Fifth segment with few setae. Fourth

segment with transverse row of long setae adjacent to dorsodistal margin. Third segment with strong ventrodistal

spine and long setae. Second segment with dorsolateral distal angle produced, terminating in simple spine, mesial

margin with small spine, dorsal surface and lateral margin with long setae; dorsomesial distal angle with small

spine. First segment with spine on dorsolateral distal margin; ventral margin produced and with 1 spine

ventrolaterally. Antennal acicle moderately long, reaching nearly to distal margin of ultimate peduncular segment,

slightly arcuate, mesial and lateral margin distally each with very long setae; terminating in simple spine.

Antennal flagellum moderately long, reaching about to tips of outstretched chelipeds; each article with several very

long setae, shortest proximally.

Chelipeds subequal, right somewhat longer and more robust; dactyls and fixed fingers weakly arched ventrally.

Right cheliped (Figs 32b, 44e) with dactyl approximately equal to length of palm; dorsomesial margin not

delimited; dorsal, mesial and ventral surfaces all with numerous tufts of long setae arising from small surface pits;

cutting edge with 1 calcareous tooth proximally and row of corneous teeth in distal half, terminating in corneous

claw. Palm about 0.75 length of carpus, dorsomesial margin with row of rather widely-spaced small spines on

proximal 0.80 and second adjacent row, 1 distinct tubercle and 2 very small spinules in dorsal midline proximally;

dorsal surfaces of palm and fixed finger with numerous tufts of long setae; dorsolateral margin rounded and with

few tiny spinules on palm, but with row of small spines on proximal 0.75 of fixed finger; lateral and ventral

surfaces with numerous tufts of long setae; cutting edge of fixed finger with 1 large calcareous tooth proximally

and row of slender calcareous teeth interspersed with corneous teeth distally; 1 very prominent calcareous-tipped

tubercle at proximal angle. Carpus approximately equaling length of merus; distal margin with few small spines

dorsally and laterally; dorsomesial margin with row of moderately strong spines and adjacent mesial and lateral row

of smaller spines and spinules; dorsolateral margin not delimited but dorsal surface laterally and lateral surface

dorsally each with irregular transverse rows of spinulose protuberances and long setae; mesial and ventral surfaces

with scattered long setae. Merus with sparse short setae on all surfaces; ventromesial margin with 2 very small

spines distally and 2 larger spines proximally; ventrolateral margin with 1 spinule distally. Ischium with small

spinulose tubercle on ventromesial margin.

Left cheliped (Figs 32c, 44f) with dactyl slightly longer than palm; dorsomesial margin of dactyl not delimited;

surfaces all with few sparse tufts of long setae; cutting edge with row of corneous teeth; terminating in corneous

claw and slightly overlapped by fixed finger. Palm approximately 0.75 length of carpus; widely-spaced double row

of small spines on rounded dorsomesial margin; dorsal surfaces of palm and fixed finger with few small widely-

spaced spines laterad of midline, strongest on fixed finger, and row of tiny spinules in region of rounded

dorsolateral margin; surfaces all with scattered long setae; cutting edge of fixed finger with row of very small

calcareous teeth proximally, corneous teeth distally. Carpus slightly shorter than merus; dorsomesial margin with

irregular row of spines and few long setae, dorsolateral margin with irregular row of smaller spines; lateral face

with few small spines at distal margin and 1 small spine centrally; 1 tiny spinule at ventrolateral distal angle;

mesial and ventral surfaces with sparse tufts of setae. Merus with sparsely scattered setae on dorsal surface, lateral

and mesial faces each with few low minutely spinulose protuberances and setae in ventral half; ventromesial

margin with 1 small spine distally, 1 at midlength and cluster of 3 slightly larger spines proximally; ventrolateral

margin with 1 distal spine. Ischium with 1 proximal and 1 distal small spine on ventromesial margin, distolateral

margin with spinule dorsally.

PAGURIDAE FROM THE KARUBAR EXPEDITION

551

Fig. 32. — Tomopaguropsis miyakei sp. nov., holotype $ (2.2 min) from Karubar Stn CP 35: a, shield and cephalic

appendages; b, right cheliped (mesial view); c. carpus and chela of left cheliped (lateral view); d, right second

pereopod (lateral view); e, left third pereopod (lateral view); f. dactyl and propodus of left fourth pereopod; g, tip of

dactyl and preungual process of left fourth pereopod (lateral view); h, telson. Scale equal 0.1 mm (g), 0.5 mm (f), and

1.0 mm (a-e, h).

Ambulatory legs (Figs 32d-e) with dactyls 1.25 to 1.35 length of propodi; in dorsal view, slightly twisted; in

lateral view, curved ventrally; all surfaces with sparse tufts of long rather stiff setae, most numerous dorsally;

ventral margins few setae and 8 or 9 corneous spines. Propodi with numerous sparse tufts of long moderately stiff

setae also most numerous dorsally; ventromesial distal angles each with 1 or 2 corneous spines. Carpi with spine

at dorsodistal angle, second also with row of widely-spaced spinules on dorsal surface, both pairs with sparse tufts

of setae dorsally and laterally. Meri and ischia each with dorsal and ventral row of low, sometimes spinulose

protuberances and sparse setae. Anterior lobe of sternite of third pereopods subrectangular, with marginal setae.

Fourth pereopods with moderately prominent preungual process at base of claw (Figs 32f-g).

Source :

552

p.a. McLaughlin

Male not known. Females with very unequally biramous pleopods. Posterior margin of sixth abdominal somite

with lateral angles rounded. Telson (Fig. 32h) with prominent transverse suture; posterior lobes nearly

symmetrical, separated by V-shaped median cleft; terminal margins rounded, each armed with 5 acute, corneous-

tipped and 2 very small spines on left, 5 on right, lateral margins not distinctly delimited.

Color. — Unknown.

Habitat. — Unknown.

DISTRIBUTION. — At present known only from the type locality in the Kai Islands of Indonesia; 390 - 502 in.

ETYMOLOGY. — This species is dedicated to the eminent Japanese carcinologist, Dr. Sadayoshi MIYAKE, in

recognition of his many contributions to our knowledge of the pagurid fauna of Japan.

Affinities. — Tomopaguropsis miyakei appears more comparable to T. lantana than to T. crinita. The dorsal

surfaces of the chelae of both T. miyakei and T. lantana are spinulose, while those of T. crinita are marked by

transverse rows of setae. However, there are very few spinules on the right chela of T. miyakei , in contrast to the

rows of spinules described by ALCOCK (1905b) for T. lantana. Two other differences between the two species are

apparent. For T. lantana the ocular acicles are reported to be long and stout, perhaps similar to the ocular acicles of

T. crinita. The ocular acicles of T. miyakei are neither long nor particularly stout. ALCOCK (1905b: 137) described

the chelipeds and ambulatory legs as being "very hairy, but not so thickly so as to entirely conceal the surface

sculpture". Although the chelipeds and ambulatory legs of T. miyakei are furnished with numerous sparse tufts of

long setae, the setation could not be considered analogous to that described and illustrated for T. lantana (ALCOCK,

1905b, pi. 13, fig. 4).

AC KNO WLEDGEMENTS

I am deeply indebted to Alain CROSNIER and Bertrand RICHER DE FORGES, both from ORSTOM, for providing

me with the opportunity to study this extremely significant collection. It was fortunate that two scientists more

specially involved in hermit crabs, Jacques FOREST of the Museum national d'Histoire naturelle, Paris, and Dwi

Listyo Rahayu of the Pulitsbang Oseanologi-LIPI from Ambon, were on board during the Karubar Cruise. It is

a testiment to their expertise that I have had such an excellent collection with which to work. Type materials from

the "Challenger" and "Alert" were graciously loaned by Paul CLARK, Natural History Museum, London. Valuable

comparative materials were provided by A. J. BRUCE, formerly of the Museums and Art Galleries of the Northern

Territory, Darwin, Australia, Michio IMAFUKU, Kyoto University, Kyoto, and Tomoyuki Komai, Natural History

Museum & Institute, Chiba, Japan, Rafael LEMAITRE, National Museum of Natural History, Smithsonian

Institution, and Nancy VOSS, Rosenstiel School of Marine and Atmospheric Sciences, University of Miami,

Miami, Florida. Technical assistance was provided by Janet ARMBRUST, Mt. Vernon, Washington. Photos are the

work of E. J. McGEORGE and Denis SERRETTE, Museum national d'Histoire naturelle, Paris. Mrs M. Joan

Thorne, Editorial Manager, Zoological Record, called my attention to a possible case of homonymy. To all I am

most grateful. This is a scientific contribution from the Shannon Point Marine Center, Western Washington

University.

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PAGURIDAE FROM THE KARUBAR EXPEDITION

559

FiG. 33 a-b. — Enneophyllus spinirostris sp. nov., holotype 8 (1.6 mm), Karubar Stn DW 49: a, carpus and chela of

right cheliped; b, merus, carpus and chela of left cheliped.

Fig. 33 c-f. — Enneopagurus garciagomeii sp. nov., c-d, paratype <J (3.4 mm), Karubar Stn CP 71. —

e-f, paratype $ (2.7 mm), Karubar Stn CP 75: c, e, carpus and chela of right cheliped; d, f, carpus and chela of left

cheliped.

Fig. 33 g. — Catapaguroides declivis sp. nov., holotype 8 (1.5 mm), Karubar Stn DW 49: chela of right cheliped.

Source :

560

P. A. MCLAUGHLIN

Fig. 34 a. — Soliiariopagurus tuerkayi sp. nov., paratype 6 (3.3 mm), Karubar Stn DW 50: whole animal.

Fig. 34 b. — Porcellcmopagurus jacquesi sp. nov., holotype 6 (2.8 mm), Karubar Stn DW 18: whole animal.

Fig. 34 c-e. — Alainopaguroides lemaitrei sp. nov., holotype 6 (6.2 mm), Karubar Stn CP 59: c, whole animal;

d, right chela; e, left chela and carpus.

Source : MNHN, Paris

Fig. 35 a. — Alainopaguroides lemaitrei sp. nov., holotype S (6.2 mm). Karubar Stn CP 59, carapace and cephalic

appendages.

Fig, 35 b-e. — Pagurodes inarmatus Henderson, 1888, lectotype <5 (7.0 mm), "Challenger" Sin 168: b, shield and

cephalic appendages; c, right chela and carpus; d, left chela and carpus; e, dactyls and propodi of left second and third

pereopods.

PAGUR1DAE FROM THE KARUBAR EXPEDITION

561

Source :

562

P. A. MCLAUGHLIN

FIG. 36. — Michelopagurus limatulus (Henderson, 1888), new combination: a-c, holotype 8 (3.1 mm), "Challenger" Stn

214 (NHM 88.33). — d-f, 8 (2.8 mm), Karubar Stn CP 38: a, d, carapace and cephalic appendages; b, e, carpus (or

part only) and chela of right cheliped; c, f, carpus (or part only) and chela of left cheliped.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

563

FlG. 37 a-b. — Pseudopagurodes piliferus (Henderson, 1888), new combination, a-b, holotype 9 (4.5 mm). Challenger

Stn Tablas Islands (NHM 88.33): a, carpus and chela of right cheliped; b, carpus and chela of left cheliped.

Fig. 37 c-e. — Michelopagurus chacei sp. nov., c-e, holotype 6 (2.5 mm) from Karubar Stn DW 13: c, shield and

cephalic appendages; d, carpus and chela of right cheliped; e, carpus and chela of left cheliped.

Source : MNHN, Paris

564

p.a. McLaughlin

F‘G u8',7 lcfloPcSurus crosnieri sp. nov., holotype ov. 2 (4.3 mm). Karubar Stn CP 91: a, whole animal; b. enlarged

shield and cephalic appendages; c, carpus and chela of right cheliped; d, carpus and chela of left cheliped.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

565

Fig. 39 a-b. — Catapagurus oculocrassus sp. nov., paratype ov. 2 (2.7 mm), Karubar Stn CP 21: a, chela of right

cheliped; b, chela of left cheliped.

Fig. 39 c-d. — Catapagurus tanimbarensis sp. nov., holotype 2 (2.1 mm), Karubar Stn DW 49: c, chela of right

cheliped; d, chela of left cheliped.

Fig. 39 e-f. — Catapagurus holthuisi sp. nov., holotype ov. 2 (2.8 mm), Karubar Stn CP 77: e, chela of right cheliped;

f, chela of left cheliped.

Source .

566

P. A. MCLAUGHLIN

Fig. 40 a-c. — Nematopagurus cf. indicus Alcock, 1905, 6 (5.2 mm), Karubar Stn DW 49: a, carpus and chela of right

cheliped; b, carpus and chela of left cheliped; c, dactyls and propodi of left second and third pereopods.

Fig. 40 d-e. — Nematopagurus sp., 9 (1.7 mm), Karubar Stn 18: d, chela of right cheliped; e, carpus and chela of left

cheliped.

Source : MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

567

FlG. 41 a-b. — Nematopagurus spinulosensoris McLaughlin & Brock, 1974, 2 (5.6 mm), KaRUBAR Sin CP 86 a, carpus

and chela of right cheliped; b, carpus and chela of left cheliped.

Fig. 41 c-f. — Nematopagurus scutelliformis sp. nov., c-d, paratype 2 (7.0 mm), Karubar Stn CP 83; e-f, paratype,

2 (6.9 mm), Karubar, st. CP 101: c, e, carpus and chela of right cheliped; d, f, carpus and chela of left cheliped.

Source :

568

P. A. MCLAUGHLIN

Fig. 42 a-b. — Australeremus indonesiensis sp. nov., holotype $ (1.9 mm), Karubar Stn DW 18: a, carpus and chela of

right cheliped; b, carpus and chela of left cheliped.

Fig 42 c-d. — Bathypaguropsis rahayuae sp. nov., holotype 6 (3.2 mm), Karubar Stn DW 14: c, carpus and chela of

right cheliped; d, carpus and chela of left cheliped.

Fig. 42 e-f. — Pagurus kaiensis sp. nov., a-b, holotype 6 (8.4 mm), Karubar Stn DW 18: e, chela of right cheliped;

f, carpus and chela of left cheliped.

Fig. 42 g-h. — Pagurus compressipes (Miers, 1881), syntype 6 (3.9 mm) NHM 1881.31: g, chela of right cheliped;

h, chela of left cheliped.

Source . MNHN, Paris

PAGURIDAE FROM THE KARUBAR EXPEDITION

569

Fig. 43 a-d. — Pagurus haigae sp. nov,, a-b, paratype 2 (12.1 mm), Karubar Stn CP 16. — e-d, paratype 6 (7.3 mm)

from Stn CP 26: a, c, chela of right cheliped; b, d, chela of left cheliped.

Fig. 43 e-f. — Pylopaguropsis zebra (Henderson. 1893), 2 (2.7 mm), Karubar Stn DW 15: e, chela of right cheliped;

f, chela of left cheliped.

Source : MNHN, Paris

570

P. A. MCLAUGHLIN

Fig. 44 a-b. — Pylopaguropsis laevispinosa McLaughlin & Haig, 1989, 6 (3.2 mm), Karubar Stn DW 22: a, carpus and

chela of right cheliped; b, carpus and chela of left cheliped.

Fig. 44 c-d. — Tomopaguropsis crinita sp. nov., paratype 6 (5.4 mm), Karubar Stn CC 10: c, carpus and chela of right

cheliped; d, carpus and chela of left cheliped.

Fig. 44 e-f. — Tomopaguropsis miyakei sp. nov., holotype 2 (2.2 mm), Karubar Stn CP 35: e, carpus and chela of right

cheliped; f, carpus and chela of left cheliped.

PAGUR1DAE FROM THE KARUBAR EXPEDITION

571

INDEX

Page numbers in bold indicates full taxonomic treatments, numbers in italics refer to illustrations.

Acanthopagurus 482

Alainopaguroides 435, 438, 469, 470, 473

lemaitrei 435, 469. 470, 472, 473, 560, 561

Alainopagurus 465, 470

Anapagrides 435, 438, 443, 444, 474, 476

facetus 444, 476

reesei 476

sp. 474, 475, 476

Anapagrides sp. 435

Anapagurus 436, 438

Australeremus 436, 439, 520, 521

eltaninae 522

indonesiensis 436, 522, 523, 525, 568

steward 522

triserratus 436, 521, 522, 523, 525

Bathypaguropsis 436, 439, 539

marionensis 541

rahayuae 436, 539, 540, 541, 542, 568

yaldwyni 541, 542

Bemhardus 525

Cancer 525

Catapaguroides 436, 438, 442, 447, 448, 454, 455,

460, 495

cristimanus 436, 454, 455, 458, 460

declivis 436, 455, 456, 458, 460, 559

inermis 436, 454, 455

karubar 436, 455, 458. 459, 460

melini 436, 454, 455, 458

mortenseni 436, 454, 455

spinulimanus 436, 454, 455

Catapagurus 436, 438, 442, 454, 473, 488, 494, 495,

498

australis 495, 505

ensifer 436, 495, 498

holthuisi 436, 495, 501, 503, 504, 565

oculocrassus 436, 491. 495, 497, 498, 501, 504,

565

tanimbarensis 436, 495. 498, 500, 501, 504, 565

vallatus 495

Cestopagurus 447, 454, 491, 495

Clibanarius 434

Dardanus 525

Decaphyllus 436, 438, 447. 448, 453, 454

barunajaya 436, 448, 450, 451

junquai 436, 448, 454

maci 436, 448, 451 ,452, 454

similis 436, 448, 451, 454

spinicornis 448, 451, 454

Diogenes 434

Diogenidae 435

Enneobranchus 439, 440, 443, 444, 448

Enneopagurus 436, 438, 439, 440, 443, 444, 448

garciagomezi 436, 443, 444, 445, 446, 447, 559

Enneophyllus 436, 437, 439, 440, 443, 448

spinirostris 436, 439, 440, 441, 559

Eupagurus 525, 546

gracilipes 536, 537

problematicus 546

triserratus 52 1

zebra 542

Galapagurus 542

Goreopagurus 482

Hemipagurus 494

Icelopagurus 436, 438, 488

crosnieri 436, 488, 490, 491, 498, 564

Iridopagurus 443, 444

Laurentia 444, 476, 477

senticosa 477

Michelopagurus 435, 436, 438. 481, 482, 484, 487,

488, 491

chacei 436, 483, 484, 486, 487, 563

limatulus 436, 482, 483, 484, 487. 562

Micropagurus 436, 438

Nanopagurus 474

Nematopagurus 436, 438, 495, 504, 505, 508, 509, 520

alcocki 436, 505. 517, 518, 520

australis 436, 505, 520

cf. indicus 436, 506, 507, 566

gardineri 505, 508, 509, 510, 517

indicus 505, 506, 508, 509. 517

muricatus 510, 51 1

ostlingochirus 436, 506, 515, 516, 517

scutellichelis 514

scutelliformis 436, 505, 511, 513, 514, 567

sp. 436, 506, 507, 508, 566

spinulosensoris 436, 505, 507, 510, 51 1, 533, 567

spinulosensorius 510

vallatus 506

Orthopagurus 442

Ostraconotus 470

spatulipes 470, 473

Ostraconotus, 470

Paguridae 435, 437

Paguridea 435

Pagurixus 476, 494

Pagurodes 435, 436, 481, 482, 487, 488

atlandcus 482

inarmatus 435, 436, 482, 487, 488, 491, 561

limatulus 435, 482, 487

piercei 482

Source :

572

P. A. MCLAUGHLIN

piliferus 435, 482, 487, 525

richardi 482

Pagurus 436, 439, 442, 525, 529, 537, 538

brachiomastus 536, 537

capsularis 436, 526, 529, 531, 532, 533

compressipes 437, 487, 525, 526, 529, 568

gracilipes 537

haigae 437, 526, 533, 534, 536, 537, 569

hedleyi 437, 525, 526

hirtimanus 437, 525, 526, 529, 532

kaiensis 437, 526, 528, 529, 568

moluccensis 437, 525

pectinatus 537

pergranulatus 437, 525, 526, 532, 533

samoensis 533

sp. 437, 537, 538

triserratus 521

yokoyai 534, 536, 537

zebra 542

Parapagurodes 49 1

Parapagurus 482, 546

Pocillopora 476

Porcellanopagurus 437, 438, 464, 465, 468, 469, 470

belauensis 465, 468, 469

jacquesi 437, 465, 466, 468, 469, 560

japonicus 468, 469

nihonkaiensis 465, 468, 469

tridentatus 465, 469

truncatifrons 465, 468, 469

Pseudopagurodes 435, 437, 487, 488, 491

piliferus 437, 484, 563

Pseudopagurus 49 1

Pylopaguropsis 437, 439, 542

fimbriata 437, 542

laevispinosa 437, 542, 544, 545, 570

lewinsohni 437, 542

zebra 437, 542, 544, 569

Pylopagurus 442, 520

serpulophilus 521

zebra 542

Rhodochirus 546

Solitariopagurus 437, 438, 461, 465, 469, 470

profundus 46 1

triprobolus 461, 464

tuerkayi 437, 461, 462, 464, 560

Spiropagurus 437, 438

Tarrasopagurus 437, 438, 488, 491, 494

rostrodenticulatus 437, 476, 491, 493, 494

Tomopaguropsis 437, 439, 546, 549

crinita 437, 546, 547, 548, 549, 552, 570

lantana 546, 549, 552

miyakei 437, 549, 551, 552, 570

problematica 546, 549

Tomopagurus 546

wassi 546

Trichopagurus 476

Tsunogaipagurus 442

Turleania 437, 438, 443, 444, 446, 474, 476, 477

albatrossae 437, 477, 478, 481

balli 437, 477, 478

multispina 437, 447, 477, 478, 479, 480

senticosa 437, 475, 477, 481

sibogae 437, 477, 478

!ILTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DE

Crustacea Decapoda: Parapaguridae

from the KARUBAR Cruise in Indonesia,

with descriptions of two new species

Rafael LEMAITRE

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

Washington, D.C. 20560

ABSTRACT

During the French-Indonesian Karubar campaign, ten species and a megalopal stage of deep-water hermit crabs of

the family Parapaguridae, were collected. Two of the species found in the collection are undescribed, Oncopagurus

glebosus sp. nov., and Paragiopagurus insolitus sp. nov.. and are characterized by several unusual or unique characters.

One previously described species, Oncopagurus orientalis (de Saint Laurent, 1972), was found to be insufficiently defined.

These three species are described or diagnosed, and illustrated. Another species, Parapagurus latimanus Henderson. 1888.

is reported for the first time from Indonesia. Two megalopal stage specimens of a parapagurid species cannot be assigned

with certainty based on current knowledge, to any species; they are also illustrated and discussed. A list of all

15 parapagurid species currently known from Indonesian waters is presented, including references where diagnoses and

illustrations can be found.

RESUME

Crustacea Decapoda: Parapaguridae recoltes lors de la campagne Karubar en Indonesie.

Descriptions de deux especes nouvelles.

Durant la campagne franco-indonesienne Karubar, 10 especes et un stade megalope de pagures d’eau profonde de la

famille des Parapaguridae ont 6t6 recoils. Deux de ces especes sont nouvelles: Oncopagurus glebosus et Paragiopagurus

insolitus et se distinguent par plusieurs caractires inhabituels ou uniques. Une espece decrite precedemment, Oncopagurus

orientalis (de Saint Laurent, 1972) s'est reveUe mal definie. Ces trois especes sont done l'objet d'une description ou d'une

diagnose et sont illustrees. Une autre espece, Parapagurus latimanus Henderson, 1888. est signalee pour la premiere fois

d' Indonesie. Deux megalopes de Parapaguridae ne peuvent etre rattachees avec certitude h aucune espece connue; elles sont

egalement illustrees et discuses. Une lisle des 15 especes de Parapaguridae, connues des eaux indonesiennes, est etablie,

accompagnee de references oil des diagnoses et des illustrations les concemant peuvent etre trouvdes.

Lemaitre, R., 1997. — Crustacea Decapoda: Parapaguridae from the Karubar Cruise in Indonesia, with

descriptions of two new species. In: A. Crosnier & P. Bouchet (eds), Resultats des Campagnes Musorstom, Volume 16.

Mem. Mus. natn. Hist, nat., 172: 573-596. Paris ISBN: 2-85653-506-2.

Source : MNHN . Paris

574

R. LEMAITRE

INTRODUCTION

The family Parapaguridae Smith, 1882, redefined by DE Saint Laurent (1972), in recent years has been the

subject of revisionary studies that have established new generic boundaries and proposed a number of new genera

(Lemaitre, 1989, 1993, 1996; Osawa, 1995). As result, ten genera are now recognized, five of which are

monotypic: Probeebei Boone, 1926, Tylaspis Henderson, 1885, Typhlopagurus de Saint Laurent, 1972,

Bivalvopagurus Lemaitre, 1993, and Tsunogaipagurus Osawa, 1995; five others contain the remainder of the

species: Parapagurus Smith, 1879, Sympagurus Smith, 1883, Strobopagurus Lemaitre, 1989, Oncopagurus

Lemaitre, 1996, and Paragiopagunis Lemaitre, 1996.

The parapagurid fauna from Indonesia is known largely from collections obtained in earlier expeditions such as

the U.S. “Albatross" (late 1800's to early 1900's), Dutch Siboga Expedition (1899-1900), the Danish

Th. Mortensen’s Pacific Expedition (1914-1916), and the Danish Galathea (1950-1952). Based on these

collections, 12 species of parapagurids have been reported from various Indonesian localities (DE SAINT LAURENT,

1972; Lemaitre, 1994, 1996).

During the recent French-Indonesian deep-water sampling campaign known as Karubar, conducted from

October to November of 1991 on board the " Baruna Jaya I", seven of the 12 species known to occur in the

Indonesian region were obtained. In addition, the Karubar material was found to contain two distinctive new

species, Oncopagurus glebosus sp. nov., and Paragiopagunis insolitus sp. nov., described herein. One species,

Parapagurus latimanus Henderson, 1888, had not been previously reported from this region. Another species in

this material, previously included in the genus Sympagurus, S. orientalis (de Saint Laurent, 1972), but recently

assigned by Lemaitre (1996) to the genus Oncopagurus, was found to be insufficiently defined. It is illustrated

and diagnosed. Also reported and discussed are two megalopal stage specimens of an undetermined species of

parapagurid. Although based on current knowledge these postlarvae cannot be assigned to any species, they are of

interest because of the paucity of information on the larval development of parapagurids (Lemaitre &

McLaughlin, 1992).

The Karubar material remains deposited in the Museum national d’Histoire naturelle, Paris (MNHN), except

lor some duplicates deposited in the National Museum of Natural History, Smithsonian Institution, Washington

D C- (USNM). For comparative purposes, paratypic material of Oncopagurus orientalis (de Saint Laurent),

borrowed from the Zoologisch Museum, Amsterdam (ZMA), and Zoologisk Museum, Copenhagen (ZMK), was

examined.

In the material examined section, the length of the shield (to the nearest 0.1 mm), indicated in parentheses, is

measured from the tip ol the rostrum to the midpoint of the posterior margin of the shield. Measurements included

for the megalopae are carapace length (CL), measured from the tip of the rostrum to the posterior midpoint of the

carapace; and total length (TL), measured from the tip of the rostrum to the midpoint of the telson, excluding the

telsonal setae. Abbreviations used are: unmat. , immature (sex indetermined); ovig., ovigerous; Stn, station.

The Karubar campaign was named for the islands of Kai, Aru, and Tanimbar.

The general terminology employed follows McLaughlin (1974). In the descriptive text, the term

semichelate to describe the condition of the fourth and fifth pereopods, is used following McLaughlin’s (1997:

435) definition, i.e. "where the ventral margin of the propodus is produced beneath the dactyl to such an extent that

flexion of the dactyl becomes more akin to the action of a dactyl against a fixed finger of a chelate appendage”.

This is in contrast with the "subchelate” condition, "in which the pereopod is developed as a prehensile structure

by the folding back of the dactyl against the propodus”.

Source :

PARAPAGURIDAE FROM EASTERN INDONESIA

575

SYSTEMATIC ACCOUNT

Family PARAPAGURIDAE Smith, 1882

Genus STROBOPAGURUS Lemaitre, 1989

Strobopagurus sibogae (de Saint Laurent, 1972)

Parapagurus sibogae de Saint Laurent, 1972: 116, figs 10, 23 (type locality: Indonesia, Siboga Exp. Stn 12).

Strobopagurus sibogae - LEMAITRE, 1989: 36; 1996: 167, fig. 1.

Material EXAMINED. — Indonesia. KARUBAR, Kai Islands: stn CP 36, 06°05'S. 132°44'E. 268-210 m,

27.10.1991: 2 2 3.3, 4.7 mm (USNM 276034).

Tanimbar Islands: stn CP 83, 09°23'S. 131°00'E. 285-297 m, 4.11.1991: 1 2 3.3 mm (MNHN-Pg 5380). —

Stn CP 84, 09°23'S, 131°09'E, 275-246 m, 4.11.1991: 3 <3 3.4-4 .4 mm. 1 2 3.5 mm (MNHN-Pg 5381).

DISTRIBUTION. — Western Pacific: Indonesia; China Sea; Japan; and Australia. Depth: 40 to 550 m.

Genus PARAPAGURUS Smith, 1879

Parapagurus latimanus Henderson, 1888

Parapagurus latimanus Henderson. 1888: 91, pi. 9, fig. 2 (type locality: "Challenger'' Stn 167A, New Zealand). —

Murray, 1895: 597. — Gordan, 1956: 338. — Lemaitre, 1986: 526; 1989; 11. — Lemaitre & McLaughlin. 1992:

762, fig. 9.

Parapagurus pilosimanus latimanus - de Saint Laurent, 1972: 103, pi. 1, fig. 5.

MATERIAL EXAMINED. — Indonesia. Karubar. Tanimbar Islands: stn CP 52, 08°03'S, 131°48'E. 1244-1266 m,

30.10.1991: 4 8 4.2-8.9 mm, 2 2 4.5, 4.8 mm (MNHN-Pg 5365). — Stn CP 87, 08°47'S, 130°49'E. 1017-1024 m,

5.11.1991: 1 8 5.2 mm (USNM 276026).

Diagnosis. — (See Lemaitre & McLaughlin, 1992).

Distribution. — Southern Australia (DE Saint Laurent, 1972); New Zealand; and now Indonesia. Depth:

909 to 1995 m.

Genus SYMPAGURUS Smith. 1883

Sympagurus brevipes (de Saint Laurent, 1972)

Parapagurus brevipes de Saint Laurent. 1972: 105, figs 2, 14 (type locality: Indonesia, Siboga Exp. Stn 12).

Sympagurus brevipes - Lemaitre, 1989: 37; 1994: 412; 1996; 170, figs 2, 3a-b, 4, 5a. 6.

Material EXAMINED. — Indonesia. Karubar. Kai Islands: stn CC 05, 05°49'S, 132°18'E, 296-299 m,

22.10.1991: I 8 18.5 mm. 1 2 ovig. 16.0 mm (MNHN-Pg 5373). — Stn CP 09, 05°23'S. 132°29‘E. 368-389 m,

23.10.1991: 1 2 ovig. 14.4 mm (MNHN-Pg 5374). — Sin CC 10, 05°21'S, 132°30’E, 329-389 m. 23.10.1991: 1 <3

23.2 mm, 2 2 9.6, 14.0 mm, 1 2 ovig. 14.4 mm (USNM 276027). — Stn CP 16, 05°17'S, 132°50'E. 315-349 m,

24,10.1991: 1 2 ovig. 13.6 mm (MNHN-Pg 5367). — Stn CP 26. 05°34’S, 132°52’E, 265-302 m, 26.10.1991: 1 <3

8.5 mm (MNHN-Pg 5370). — Stn CP 36, 06°05'S, 132°44'E, 268-210 m. 27.10.1991: 1 2 15.8 mm, 1 2 ovig.

12.9 mm (MNHN-Pg 5366). — Stn CP 37. 06°07'S, 132°42'E. 363-241 m. 27.10.1991: 1 <3 16.1 mm (MNHN-Pg

5375).

Tanimbar Islands: stn CP 39, 07°47’S, 132°26'E, 477-466 m, 28.10.1991: 1 8 21.1 mm (MNHN-Pg 5377). —

Stn CP 45, 07°54'S, 132°47’E, 302-305 m, 29.10.1991: 1 2 15.8 mm (MNHN-Pg 5376). — Stn CC 57, 08°19'S,

131 °53'E, 603-620 m, 31.10.1991: 1 immat. 3.7 mm (MNHN-Pg 5371). — Stn CP 69, 08°42'S, 131°53'E, 356-368 m,

2.11.1991: 1 <3 5.5 mm, 1 2 4.3 mm (MNHN-Pg 5372). — Stn CP 77, 08°57'S, I31°27'E. 352-346 m, 3.11.1991: 1 2

5.4 mm (MNHN-Pg 5369).

Source :

576

R. LEMA1TRE

Karubar. 1991. (No station data): 1 3 22.5 mm (MNHN-Pg 5368).

Diagnosis. — (See Lemaitre, 1996).

Distribution. — Indo-Pacific: Zanzibar; Indonesia; Philippines; and Australia. Depth: 210 to 794 m.

Sympagurus papposus Lemaitre, 1996

Sympagurus papposus Lemaitre, 1996: 180, figs 3c-d, 5 b. 8-10 (type locality: E of Broken Bay, New South Wales,

Australia. FRV Kapala Stn K75-01-02).

Material examined. — Indonesia. Karubar, Tanimbar Islands: stn CC 40. 07°46'S. 132°31’E, 443-468 m,

28.10.1991: 1 3 13.4 mm, 2 9 ovig. 121.1. 12.3 mm (MNHN-Pg 5379). — Stn CC 41, 07°45'S, 132°42'E, 401-

393 m, 28.10.1991: 1 3 14.6 mm (USNM 276033). — Stn CP 91, 08°44'S, 131°05'E, 884-891 m, 5.11.1991: 1 3

7.0 mm (MNHN-Pg 5378).

Diagnosis. — (See Lemaitre, 1996).

Distribution. — Indo-Pacific: Madagascar; Indonesia: and Australia. Depth: 205 to 960 m.

Remarks. — As pointed out by Lemaitre (1996), this species is very similar to S. dofleini (Balss, 1912).

Females of S. papposus can be separated from males or females of S. dofleini by the armature of the anterior lobes

of the telson. The left anterior lobe of the telson, and sometimes also the right anterior lobe, are armed

ventrolaterally with a fringe or cluster of slender corneous spines mixed with bristle-like setae in S. papposus. The

anterior lobes have in both sexes at most a row of setae in S. dofleini. Males of the two species, however, can

only be separated using a number of subtle differences. The anterolateral projections of the shield are broadly

rounded, often obsolete, in 5. papposus-, the projections are broadly triangular, often terminating acutely in

S. dofleini. The spines on the antennal scales are stronger, and more broadly spaced in S. papposus than in

5. dofleini. The distal lobe of the male first gonopod is broader in S. papposus than in S. dofleini. The two

species also seem to utilize different habitats or symbiotic associations. S. papposus has been found living

exclusively in large zoanthids ( Epizoanthus sp.) whereas 5. dofleini is most frequently found living in large

actinians ( Stylobates sp.).

Genus ONCOPAGURUS Lemaitre, 1996

Oncopagurus minulus (Henderson, 1896)

Parapagurus minulus Henderson, 1896: 531 (type locality: off the north Maidive Atoll. "Investigator" Stn 150). —

Alcock & Anderson, 1897: pi. 32, fig. 3, 3a. — Alcock. 1901: 222; 1905: 101, pi. 10, fig. 3. — Gordan, 1956:

338 (lit). — de Saint Laurent, 1972: 108.

Sympagurus minulus - Lemaitre, 1989: 37; 1994: 412.

Oncopagurus minulus - LEMAITRE, 1996: 201, fig. 21.

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands: stn CP 54, 08°21'S, 131°43'E, 836-869 m,

30.10.1991: 1 $ ovig. 3.1 mm (MNHN-Pg 5345). — Stn CP 87. 08°47'S, 130°49'E, 1017-1024 m. 5.11.1991: 1 3

3.2 mm (MNHN-Pg 5346). — Stn CP 91, 08°44’S, 131°05'E, 884-891 m. 5.11.1991: 1 3 3.0 mm (USNM 276030).

Diagnosis. — (See Lemaitre, 1996).

Distribution. — Indo-Pacific: Maldives; Indonesia; and Australia. Depth: 800 to 2308 m.

Oncopagurus monstrosus (Alcock, 1894)

? Parapagurus monstrosus" Alcock, 1894: 243 [type locality, by lectotype designation (Lemaitre, 1996: 199): Bay of

Bengal].

Source :

PARAPAGURIDAE FROM EASTERN INDONESIA

577

Sympagurus monsirosus - Henderson, 1896: 533, — Alcock & Anderson, 1897: pi. 32, fig. 4. — ALCOCK, 1901: 223.

— Lemaitre, 1989: 37; 1994: 412.

Sympagurus arcuaius var. monsirosus - Alcock, 1905: 104, pi. 10, fig. 5. — Gordan, 1956: 341. — Kemp & Sewell,

1912: 26.

Parapagurus monsirosus - de Saint Laurent, 1972: 108. — Miyake, 1978: 72; 1982: 119, pi. 40, fig. 1. — Baba et al. ,

1986: 302, fig. 146. — Imafuku, 1992: 234, unnumbered fig.

Oncopagurus monsirosus - LEMAITRE, 1996: 199, figs 19, 20.

Not Parapagurus arcuaius var. monsirosus - BALSS, 1912: 99, pi. 10, fig. 3. [= Sympagurus brevipes (de Saint Laurent,

1972)].

MATERIAL EXAMINED. — Indonesia. Karubar. Kai Islands: stn CC 10, 05°2 1 'S, 132°30’E, 329-389 m,

23 10 199L 1 6 6.1 mm (MNHN-Pg 5348), 1 2 3.1 mm (USNM 276028). — Stn CP 12, 05°23'S, 132°37'E. 436-

413 m 23 10 1991: I 6 4.9 mm (MNHN-Pg 5351). — Stn CP 26, 05°34'S, 132°52'E. 265-302 m, 26.10.1991: 1 6

5.1 mm (USNM 276029). — Stn CP 35, 06°08'S, 132°45'E, 390-502 m, 27.10.1991: 3 6 3.2-4.3 mm, 2 2 2.4,

2.5 mm (MNHN-Pg 5352).

Tanimbar Islands: stn CP 39, 07°47'S, 132°26'E, 477-466 m, 28.10.1991: 1 6 3.4 mm (MNHN-Pg 5350). —

Stn CP 69, 08°42'S, 131°53E, 356-368 m, 2.11.1991: 1 6 5.1 mm (MNHN-Pg 5349). — Stn CP 70, 08°41'S, 13I°47'E,

413-410 m, 2.11.1991: 1 2 4.5 mm (MNHN-Pg 5347).

Diagnosis. — (See Lemaitre, 1996).

DISTRIBUTION. — Indo-Pacific: Gulf of Aden; Bay of Bengal; Japan; Philippines; Indonesia; and Australia.

Depth: 202 to 1000 m.

Oncopagurus orientalis (de Saint Laurent, 1972)

Figs 1-2

Parapagurus orientalis de Saint Laurent, 1972: 114, figs. 8, 16.

Sympagurus orientalis - LEMAITRE, 1989: 37; 1994: 412.

TYPE Material. — Hololype: Philippines Islands. “ Albatross ”: stn 5289, southern Luzon, 13°41'50"N,

120°58'30"E, 314 m, 22 07.1908: 6 2.9 mm (USNM 168311).

Paralypes: Philippines Islands. "Albatross": stn 5268, Batangas Bay, 13°42'N, 120°57 15 E, 170 fms (311 m),

8.06.1908: 3 6 2. 0-2.4 mm (USNM 168320). — Th. Mortensen's Pacific Exp. 1914-16, 3 mi SW of Tucuran. 550 m.

10.03.1914: 1 2 1.7 mm (ZMK).

Paralypes: Indonesia. "Siboga": stn 137, 00°23.8'N, 127°29'E, 472 m. 3. VIII. 1899. coll. M. WEBER: 2 6 1.7.

1.9 mm (ZMA De 103.108). — "Galathea" Exp. 1950-52: stn 490. Bali Sea, 05°25'S, 117°03'E, 545-570 m,

14.09.1951: 1 2 ovig. 1.8 mm (ZMK).

ADDITIONAL Material EXAMINED. — Indonesia. Karubar. Kai Islands: stn CP 35, 06°08'S, 132°45'E, 390-

502 m, 27.10.1991: 1 6 2.5 mm, 1 2 2.2 mm (MNHN-Pg 5353).

DIAGNOSIS. — Shield (Fig. la) as long as broad; dorsal surface weakly calcified medially; rostrum broadly

rounded, with low dorsal ridge; anterior margins weakly concave; lateral projections broadly subtriangular, usually

terminating in small spine; ventrolateral margin armed with small spine; posterior margin broadly rounded. Ocular

peduncles more than half length of shield; ocular acicles subtriangular, terminating in strong bifid or occasionally

multifid spine; corneae weakly dilated. Sternite of third maxillipeds with small spine on each side of midline.

Antennular peduncle, when fully extended (not shown extended in Fig. la), exceeding distal margin ot cornea by

full length of ultimate segment. Antennal peduncle (Fig. lb) not exceeding distal margin of cornea; third segment

with strong ventromesial distal spine; second segment with dorsolateral distal angle produced, terminating in

strong spine; first segment with small lateral spine; acicle not exceeding distal margin of cornea, mesial margin

armed with 8 to 12 spines; flagellum with series of short setae (<1 article in length) and long setae (>3 articles in

length) every 4 to 8 articles. Chelipeds markedly dissimilar, with some iridescence and moderately dense setae.

Right cheliped (Fig. 2a-e) with chela longer than broad; fingers moderately curved ventromesially; dactyl with

concave ventromesial face; palm with scattered small spines on dorsal face, dorsolateral and dorsomesial margins

each well delimited by row of spines; mesial face of palm rounded, with small spines or tubercles. Left cheliped

(Fig. 20 with carpus weakly calcified on dorsal surface; carpus with dorsodistal spine. Ambulatory legs (Fig. lc-d)

Source :

578

R. LEMAITRE

FlG, ■ — 0"coPa8urus orientals (de Saint Laurent, 1972), Karubar Stn CP 35 (MNHN-Pg 5353). a-g, $ (2 2 mm)-

, d 2 5 mmJ: a: ;’hleld and cephalic appendages; b, right antennal peduncle, lateral view; c, right second pereopod!

lateral view, d, nght third pereopod, lateral view; e, propodus and dactyl of right fourth pereopod, lateral view; f. left

(on left) and nght (on nght) uropods, dorsal view; g, telson, dorsal view; h, male second right (upper) and left (lower)

pleopods, lateral view. Scales equal 1 mm (a, c, d), and 0.5 mm (b, e-h).

Source : MNHN, Paris

PARAPAGURIDAE FROM EASTERN INDONESIA

579

slender; dactyl with 1 to 4 minute spinules (usually not visible in lateral view) on ventromesial margin, and

dorsal and dorsomesial rows of long setae; carpus with small dorsodistal spine; merus of right third pereopod with

row of small spines on dorsal margin; meri of left second and third pereopods with dorsal margins unarmed.

Anterior lobe of sternite of third pereopods with small marginal spine, setose. Fourth pereopod (Fig. le) with

dactyl terminating in short, corneous claw in both sexes; propodal rasp consisting of ovate scales. Uropods and

telson (Fig. lf-g) markedly asymmetrical. Telson lacking transverse suture separating anterior and posterior lobes;

Fig. 2. — Oncopagurus orientalis (de Saint Laurent, 1972), a-b. holotype, 6 (2.9 mm), "Albatross Stn 5289,

Philippines (USNM 168311); c-f, 2 (2.2 mm), Karubar Stn CP 35 (MNHN-Pg 5353): a, carpus and chela of right

cheliped (from de Saint Laurent, 1972); b, chela of same, ventral view; c, right cheliped, dorsal view;

d-e, chela of same in ventral (d) and mesial (e) views; f, left cheliped, dorsal view. Scales equal 1 mm.

Source :

580

R. LEMAITRE

posterior lobes separated by shallow U-shaped median cleft, right lobe weakly developed (frequently obsolete),

terminal margins armed with often strongly curved corneous spines. Male lacking first gonopods; second gonopods

(Fig. lh) vestigial or rudimentary, unsegmented, usually paired, asymmetrical, or sometimes with unpaired left.

Females with vestigial right second pleopod.

Habitat and symbiotic Associations. — The Karubar specimens were found living in coarse-textured

zoanthids. Other specimens have been found in gastropod shells.

Distribution. — Indo-Pacific: Philippines; Moluccas; Indonesia. Depth: 300 to 575 m.

Remarks. — Oncopagurus orientalis is one of four Oncopagurus species in which males lack first gonopods

and the second gonopods are vestigial or rudimentary. The others are O. haigae de Saint Laurent, 1972, O. tuamotu

Lemaitre, 1994, and O. cidaris Lemaitre, 1996. The multifid condition of the ocular acicles in O. orientalis is the

most obvious character that immediately distinguishes this species from the other three. Although the four species

are otherwise superficially similar, they differ in such characters as the shape and armature of the right cheliped;

relative length of dactyls of ambulatory legs; presence or absence of sexual dimorphism in the dactyl of the fourth

pereopod; and in males, degree of development of the second gonopods.

The right palm of O. orientalis shows some degree of variability in length, and also in armature of the ventral

face. As expected, such variabilty is related to size and sex of the individuals. The palm of the male holotype, for

example, is longer than broad (Fig. 2a), whereas in Karubar females the palm is broader than long (Fig. 2c). The

armature of the ventral face can consist of scattered small tubercles (Fig. 2b), or scattered small tubercles and

moderately large tubercles often arranged in an oblique row (Fig. 2d).

Oncopagurus glebosus sp. nov.

Figs 3-6

Materul EXAMINED. — Holotype : Indonesia. Karubar, Tanimbar Islands : stn CP 86, 09°26'S 13ri3'E 225-

223 m, 4.11.1991: 6 1.8 mm (MNHN-Pg 5342).

Paratypes: Indonesia. Karubar, Tanimbar Islands : stn DW 49, 08°00’S, 132°59'E, 210-206 m, 29.10.1991: 15 6

!,,;6OBn’ml 22 °/[s-2-2' 23 mm' 1 juv. 1.2 mm (MNHN-Pg 5344). — Stn DW 80, 09°37'S, 13r02'E. 199-201 m,

C,, TO A T r?*2 5 L2' 15 mm- 1 2 ov'g- 18 mm (USNM 276035). — Stn CP 86. 09°26'S, 131°13E,

2760363) 9 3 3' 17 mm <MNHN-pg 5343>. 1 3 2 3 mm, 2 2 1.3, 2.7 mm. 1 2 ovig. 2.5 mm (USNM

Description. Shield (Fig. 3a) as broad as long; dorsal surface weakly calcified on usually more than half of

surface, with scattered tufts of short setae; rostrum broadly rounded, weakly produced, with short mid-dorsal rid«e;

anterior margins concave; lateral projections subtriangular, terminating in small spine; anterolateral margfns

s opmgj postenor margin broadly rounded; ventrolateral margins of shield with small spine on one or both sides.

Anterodistal margin of branchiostegite rounded, unarmed, setose.

Ocular peduncles more than half length of shield, with dorsal row of long setae. Cornea moderately dilated.

Ocular acicles subtriangular, terminating bluntly or subacutely; with strong submarginal spine; separated basally

by less than basal width of 1 acicle. *

Antennular peduncle long, slender; when fully extended (not shown extended in Fig. 3a), exceeding distal

margin of cornea by entire length of ultimate segment. Ultimate segment twice as long as penultimate segment,

with scattered setae. Basal segment with strong ventromesial spine; lateral face with distal subrectangular lobe

armed with 1 small spine, and strong spine proximally. Ventral flagellum usually with 5 or 6 articles

Antennal peduncle (Fig. 3b) reaching distal margin of cornea. Fifth segment unarmed, but with scattered setae

Fourth segment with strong dorsodistal spine. Third segment with strong ventromesial distal spine. Second

segment with dorsolateral distal angle produced, terminating in strong, simple spine; mesial margin with spine on

dorsodistal angle. F.rst segment with 1 small spine on lateral face; ventromesial angle produced, with row of 3 or

4 small spines laterally. Antennal acicle slightly curved outward (in dorsal view), not reaching distal margin of

cornea, terminating in strong spine (rarely bifid); mesial margin armed with row of 8 to 1 1 spines. Flagellum

Source :

PARAPAGURIDAE FROM EASTERN INDONESIA

581

Fig. 3. — Oncopagurus glebosus sp. nov., paratype 3 (2.3 mm), Karubar Sin CP 86 (USNM 276036): a, shield and

cephalic appendages; b, right antennal peduncle, lateral view; c, right cheliped, lateral view (setae omitted);

d-e, chela of same in ventral (d) and lateral (e) views (setae ommited in e); f, left cheliped, dorsal view. Scales equal

1 mm (a, c-f), and 0.5 mm (b).

Source : MNHN. Paris

582

R. LEMAITRE

long, exceeding extended right cheliped and ambulatory legs; with serial arrangement of short (<1 article in length)

and long (3-5 articles in length) setae every 2-4 articles.

Mandible (Fig 4a) as figured. Maxillule (Fig. 4b) with external lobe of endopod weakly developed, internal lobe

with 1 long seta. Maxilla (Fig. 4c) with endopod exceeding distal margin of scaphognathite. First maxilliped

(Fig. 4d) with endopod exceeding exopod in distal extension. Second maxilliped (Fig. 4e) with exopod about

4 times as long as broad. Third maxilliped (Fig. 4f) with exopod about 6.5 times as long as broad; crista dentata

consisting of about 8 calcareous or corneous-tipped teeth; basis with 1 tooth mesially; coxa unarmed or with small

tooth mesially. Stemite of third maxillipeds with small spine on each side of midline.

KSSKKSSS

Source : MNHN , Paris

PARAPAGURIDAE FROM EASTERN INDONESIA

583

Fig. 5. — Oncopagurus glebosus sp. nov., paratype 3 (2.3 mm), Karubar Stn CP 86 (USNM 276036): a, second right

pereopod, lateral view; b, dactyl of same, mesial view; c, right third pereopod, lateral view; d, dactyl of same, mesial

view; e, stemite of third pereopods, ventral view; f, propodus and dactyl of left pereopod, lateral view; g, propodus

and dactyl of left fifth pereopod, lateral view. Scale equals 1 mm (a-d), and 0.5 mm (e-g).

584

R. LEMAITRE

Chelipeds markedly dissimilar. Right cheliped (Fig. 3c-e) massive; dorsal surfaces of merus, carpus and chela

each with moderately dense setae (not shown in Fig. 3c, e); chela with dense fringe of long setae on lateral and

mesial margins. Fingers curved ventromesially, terminating in small, usually blunt corneous claws; cutting edges

with irregularly-sized calcareous teeth. Dactyl about as long as mesial margin of palm, set at strongly oblique

angle to longitudinal axis of palm; mesial margin broadly curved, well delimited by row of strong spines

diminishing in size distally; dorsal face with scattered small tubercles; ventral face with longitudinal ridge covered

with irregular rows of tubercles; ventromesial face concave. Fixed finger broad at base, dorsal face with scattered

small tubercles, lateral margin well delimited by row of spines; ventrolateral face often strongly concave, ventral

face with median longitudinal ridge covered with irregular rows of tubercles. Palm longer than broad, dorsolateral

margin well delimited by row of strong spines; dorsomesial margin delimited by row of spines; mesial face

rounded, with scattered tubercles; dorsal surface with irregular rows of spines medially; ventral surface (Fig. 3d-e)

with irregularly arranged tubercles or blunt spines, and raised frequently very prominent cluster of tubercles

medially. Carpus with dorsolateral margin usually well delimited by row of spines distally, rounded proximally;

dorsodistal margin with row of spines; dorsal face with numerous small spines; ventromesial margin with row of

spines; ventral face with scattered small tubercles. Merus with scattered tubercles on dorsal face; ventromesial

margin with row of spines. Ischium with ventromesial row of spines. Coxa with ventromesial and ventrolateral

margins each with small distal spine.

Left cheliped (Fig. 3f) usually weakly calcified on dorsolateral face of carpus and on lateral face of merus.

Fingers terminating in small corneous claws; dorsal and ventral surfaces unarmed except for scattered tufts of setae;

cutting edge of dactyl with row of minute, fused corneous teeth; cutting edge of fixed finger with row of regularly

spaced, small, evenly-sized calcareous teeth. Dactyl slightly longer than length of mesial margin of palm. Palm

unarmed except for scattered setae and proximomedial row of blunt spines on dorsal face. Carpus with strong

dorsodistal spine, and smaller spine laterally on dorsodistal margin; dorsal margin with long setae; ventral face

smooth. Merus with long setae on dorsal margin; with ventrolateral row of spines, and small ventromesial spine

distally. Ischium and coxa each with 1 small spine on ventromesial and ventrolateral margins distally.

Ambulatory legs (Fig. 5a-d) similar, exceeding extended right cheliped by approximately 0.25 length of dactyl.

Dactyls broadly curved, about 1.6 times as long as propodi, and terminating in sharp corneous claws; each with

dorsal and dorsomesial rows of long setae, and 1-5 minute spinules on ventromesial margin. Propodi each with

row of setae on dorsal margin. Carpi each with small dorsodistal spine, and setae dorsally. Meri unarmed except for

1 or 2 small ventrodistal spines (second pereopod) or with row of small spines on dorsal margin (third pereopod).

Ischia with small dorsodistal and ventrodistal spine (second) or unarmed (third). Coxae with 1 small spine on

ventromesial and ventrolateral margins distally (second) or unarmed (third). Anterior lobe of sternite of third

pereopods (Fig. 5e) rounded, setose, unarmed or with small subdistal spine.

Fourth pereopod (Fig. 5f) semichelate. Dactyl terminating in sharp corneous claw; with ventrolateral row of

small corneous spinules. Propodus longer than broad, rasp formed of 1 row of rounded scales. Carpus with long

setae on dorsal margin. Merus with rows of long setae on dorsal and ventral margins.

Fifth pereopod (Fig. 5g) semichelate. Propodal rasp extending to mid-length of segment.

Uropods and telson (Fig. 6a-c) markedly asymmetrical. Telson lacking transverse suture; dorsal surface with

scattered setae; posterior lobes separated by shallow unarmed cleft, terminal margin of lobes armed with long, often

strongly curved corneous spines.

Males with paired first and second gonopods; first gonopods not yet appearing or not fully developed in

juveniles (SL <1.5 mm). First gonopods (Fig. 6d) each with nearly flat distal lobe and long marginal setae.

Second gonopods (Fig. 6e) each with distal segment flat; distal half of distal segment with long setae marginally

and on anterior face; basal segments each with row of setae laterally. Females with vestigial second right pleopod.

Habitat. — Gastropod shells.

Distribution. — Known so far only from Tanimbar Islands, Indonesia. Depth: 199 to 225 m.

ETYMOLOGY. The specific name is from the Latin glebosus, meaning lumpy, and is in reference to the

lumpy appearance given by tubercles to the ventral face of the right chela.

Source :

PARAPAGURIDAE FROM EASTERN INDONESIA

585

Remarks. — Oncopagurus glebosus sp. nov. is distinguished not only from all its congeners but from all

other parapagurids, by the unique condition of the ocular acicles. This is the only species in the family known to

have ocular acicles with a submarginal spine. The ocular acicles in all other parapagurids (except the highly

specialized Tylaspis anomala Henderson, 1885, which lacks acicles), terminate in a simple to multifid marginal

spine.

The distinct armature of the ventral surface of the right palm also distinguishes O. glebosus sp. nov. from

other Oncopagurus species. The ventral surface has numerous irregularly arranged tubercles, some of which

frequently form a prominent cluster medially (Fig. 3d). The cluster is usually markedly raised above the surface; in

586

R. LEMAITRE

small individuals (SL < 1 .5 mm) it is not as prominent, and the tubercles may be smaller and wider apart than in

large individuals. The presence of prominent armature in the form of spines or variously shaped tubercles on the

ventral face of the right chela is a condition present in three other parapagurid species, Paragiopagurus boletifer (de

Saint Laurent, 1972), P. rugosus (de Saint Laurent, 1972), and Tsunogaipagurus chuni (Balss, 191 1).

Genus PARAGIOPAGURUS Lemaitre, 1996

Paragiopagurus acutus (de Saint Laurent, 1972)

Parapagurus acutus acutus de Saint Laurent, 1972: 113, figs 7, 18 (type locality: Philippines, "Albatross" Stn 5222).

Sympagurus acutus acutus - Lemaitre, 1989: 37; 1994: 412.

Paragiopagurus acutus - Lemaitre, 1996: 211, figs 25-26.

Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 28. 05°31'S, 132°54'E, 448-467 m,

26.10.1991: 1 2 immat. 1.5 mm (MNHN-Pg 5362). — Stn CP 27, 05°33'S, 132°51'E, 304-314 m, 26 10 199L 1 2

4.7 mm (MNHN-Pg 5359). — Stn CP 36, 06°05'S, 132°44'E, 268-210 m, 27.10.1991: 1 6 6.4 mm (MNHN-Pg 5360).

Tanimbar Islands: stn CC 56, 08°16’S, 131°59'E, 552-549 m, 31.10.1991: 2 6 4.8, 5.2 mm, 1 2 3 7 mm (USNM

276032). — Stn CP 67, 08°58'S, 132°06'E, 233-146 m, 1.11.1991: 3 2 4.3-4.6 mm, 1 2 ovig. 3.7 mm (MNHN-Pg

5355). — Stn CP 77, 08°57’S, 131°27’E, 352-346 m, 3.11.1991: 4 6 5. 2-6.6 mm (MNHN-Pg 5358). — Stn CP 79,

09°16'S, 131°22'E, 250-239 m, 3.11.1991: 9 3 2.2-4.5 mm. 6 2 3.7-4.5 mm, 2 2 ovig. 4.0, 4.3 mm (USNM 276031).

— Stn CP 83, 09°23'S, 131°00'E. 285-297 m. 4.1 1.1991: 2 <3 4.1, 6.0 mm (MNHN-Pg 5356). — Stn CP 84, 09°23'S,

131°09'E, 275-246 m, 4.11.1991: 26 3 1. 7-4.6 mm, 12 2 1. 6-4.0 mm, 12 2 ovig, 3.0-3.9 mm (MNHN-Pg 5361). —

Stn CP 85, 09°22'S, 131°14'E, 245-240 m, 4.11.1991: 8 3 3. 5-5.0 mm (MNHN-Pg 5357). — Stn CP 86 09°26’S

13 1°I3'E, 225-223 m. 4.11.1991: 3 3 2.2-3J mm (MNHN-Pg 5354).

Diagnosis. — (See Lemaitre, 1996).

COLOR. — The following coloration is based on a male (6.4 mm, Stn CP 36, MNHN-Pg 5360) after

preservation in alcohol for four years. Overall straw white to yellowish. Shield with small light orange area on

each side of anterior half. Ocular peduncles dark orange on ventral face, dorsal face faded orange. Right cheliped

with dorsal surface of chela having small light orange area proximally; carpus with three wide orange stripes (one

dorsal, one mesial, one lateral); merus with two stripes (one lateral, one mesial). Carpus of left cheliped with

orange stripe on lateral face, and light orange tint on mesial face. Ambulatory legs with orange tint on lateral faces

of carpi and propodi (more strongly colored on carpi); with orange area on lateral and mesial faces of meri.

Distribution. — Western Pacific: Philippines, China Sea, Indonesia, Japan, and Australia. Depth: 146-

558 m.

Paragiopagurus insolitus sp. nov.

Figs 7-10

MATERIAL EXAMINED. - Holotype: Indonesia. Karubar, Kai Islands: stn DW 28, 05°31'S 132°54'E 448-

467 m, 26.10.1991: <3 2.2 mm (MNHN-Pg 5363).

Paratypes: Indonesia. Karubar, Kai Islands: stn DW 28, 05°31'S, 132°54'E, 448-467 m. 26.10.1991: 3 3 1.3-

1.8 mm, 1 2 ovig, 1.7 mm (MNHN-Pg 5364); 1 <3 2.0 mm, 1 2 ovig. 1.6 mm (USNM 276037).

Description. Shield (Fig. 7a) slightly longer than broad; dorsal surface weakly calcified on usually more

than half of surface, with scattered short setae; rostrum broadly rounded, weakly produced, with short mid-dorsal

ridge; anterior margins weakly concave; lateral projections subtriangular, terminating in small spine; anterolateral

margins sloping; posterior margin broadly rounded; ventrolateral margins with small spine on one or both sides

Anterodistal margin of branchiostegite rounded, unarmed, setose.

0cu'ar peduncles more than half length of shield, each with dorsal row of long setae. Cornea at most weakly

dilated. Ocular acicles subtriangular, terminating in long slender spine reaching nearly to mid-length of ocular

peduncles; separated basally by less than basal width of 1 acicle.

Source :

PARAPAGURIDAE FROM EASTERN INDONESIA

587

Fig. 7. — Paragiopagurus insolitus sp. nov., Karubar Stn DW 28. a-b, d-g, holotype 6 (2.2 mm) (MNHN-Pg 5363);

c, paratype <5 (2.0 mm) (USNM 276037); a, shield and cephalic appendages; b. right antennal peduncle, lateral view;

c, epistome, ocular peduncles and acicles, anterior view (es, epistomial spine; Is, labral spine); d. right cheliped,

dorsal view; e-f, chela of same in mesial (e) and lateral (f) views; g, left cheliped, dorsal view. Scales equal 0.5 mm (a.

d-g), and 0.25 mm (b, c).

Source : MNHN, Paris

588

R. LEMAITRE

FlGm8„,.h P°[ragl0Pa8“rus Msohtus sp. nov„ paratype 6 (2.0 mm), Karubar Stn DW 28 (USNM 276037) Left

in,fmaJ r'T: ,a- mandlble; b’ maxiilule (proximal endile not shown); c, maxilla; d first maxilliped

e, second maxilliped; f, third maxilliped. Scales equal 0.5 mm (a), 1 mm (b-d), and 0.25 mm (e, 0- ?

Source :

PARAPAGURIDAE FROM EASTERN INDONESIA

589

Antennular peduncle long, slender, exceeding distal margin of cornea by about one-fifth of penultimate

segment. Ultimate segment twice as long as penultimate segment, with scattered setae. Basal segment with strong

ventromesial spine; lateral face with distal subrectangular lobe armed with small spine, and strong spine

proximally. Ventral flagellum usually with 5 or 6 articles.

Antennal peduncle (Fig. 7b) exceeding distal margin of cornea by about one-fifth length of fifth segment. Fifth

segment unarmed, but with scattered setae. Fourth segment with strong dorsodistal spine. Third segment with

strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong,

simple spine; mesial margin with spine on dorsodistal angle. First segment with lateral face unarmed;

ventromesial angle produced, with row of 3 or 4 small spines laterally. Antennal acicles slightly curved outward

(in dorsal view), slightly exceeding distal margins of corneae, terminating in strong spine; mesial margins armed

with row of 10 to 13 spines. Flagellum long, exceeding extended right cheliped and ambulatory legs; with short

setae 1 flagellar article in length or less.

Mandible (Fig. 8a) with incisor process consisting of several irregularly-shaped teeth. Maxillulc (Fig. 8b) with

external lobe of endopod obsolete, internal lobe with 1 long seta. Maxilla (Fig. 8c) with endopod exceeding distal

margin of scaphognathite. First maxilliped (Fig. 8d) with endopod exceeding exopod in distal extension. Second

maxilliped (Fig. 8e) with long, slender exopod about 10 times as long as broad. Third maxilliped (Fig. 80 with

long, slender exopod about 10 times as long as broad; merus with dorsodistal spine; crista dentata consisting of

about 12 calcareous or corneous-tipped teeth; basis with 1 tooth mesially; coxa unarmed. Sternite of third maxil-

lipeds with spine on each side of midline. Epistome (Fig. 7c) with median region strongly produced anteriorly

(somewhat pyramid-shaped with ventral face concave); terminating in small, often inconspicuous blunt spine.

Chelipeds markedly dissimilar. Right cheliped (Fig. 7d-f) with sparse setae; surfaces of merus, carpus, and chela

with some iridescence. Fingers weakly curved ventromesially, terminating in small, usually blunt corneous claws;

cutting edges with irregularly-sized calcareous teeth. Dactyl shorter than length of mesial margin of palm, set at

moderately oblique angle to longitudinal axis of palm; mesial margin broadly curved, weakly delimited proximally

by irregular row of few small tubercles; dorsal face with scattered small tubercles and tufts of setae; ventral face

elevated along midline forming longitudinal ridge. Fixed finger broad at base, dorsal face with scattered small

tubercles and tufts of setae; lateral margin well delimited proximally by row of spines. Palm longer than broad,

dorsolateral margin at most weakly delimited by row of small spines or tubercles; dorsomesial margin weakly

delimited by row of small well-spaced spines; dorsal surface smooth or at most with scattered small spines or

tubercles laterally and mesially; mesial face rounded, with scattered tubercles; ventral surface smooth. Carpus with

rounded lateral and mesial faces; dorsal surface with irregular rows of small spines or tubercles; dorsomesial margin

delimited in distal half by row of small spines; dorsodistal margin with 3 or 4 median spines; ventromesial margin

with row of spines; ventral face with scattered small tubercles. Merus with scattered tubercles on dorsal and ventral

faces; dorsal surface with longitudinal row of bristle-like setae, and row of setae on dorsodistal margin;

ventromesial margin with row of spines. Ischium unarmed. Coxa with ventromesial and ventrolateral margins each

with small distal spine.

Left cheliped (Fig. 7g) usually weakly calcified laterally on carpus and merus. Fingers terminating in small

corneous claws; dorsal and ventral surfaces unarmed except for scattered tufts of setae; cutting edge of dactyl with

row of minute, fused corneous teeth; cutting edge of fixed finger with row of low small teeth. Dactyl about as long

as length of mesial margin of palm. Palm unarmed; with scattered setae. Carpus with small dorsodistal spine;

dorsal margin with bristle-like setae; ventral face smooth. Merus with bristle-like setae on dorsal margin; usually

with small spine on ventrolateral margin, and 1 to 3 spines on ventromesial margin. Ischium unarmed. Coxa with

ventromesial and ventrolateral margins each with small spine distally.

Ambulatory legs (Fig. 9a-d) similar right from left, exceeding extended right cheliped by approximately

0.25 length of dactyls. Dactyls broadly curved, each about 1.5 times as long as propodus, and terminating in sharp

corneous claw; with dorsal and dorsomesial rows of long setae, and 5 to 8 minute spinules on ventromesial

margin. Propodi each with row of short setae on dorsal margin. Carpi each with small dorsodistal spine, and short

setae dorsally. Meri unarmed. Ischia unarmed or each with small spine on ventrolateral margin distally. Coxae of

second pereopods each with 1 small spine distally and 1 small spine proximally on ventromesial margin; coxae of

third pereopods unarmed. Anterior lobe of stemite of third pereopods (Fig. 9e) sub-semicircular, unarmed, setose.

Source :

590

R. LEMAITRE

FIG. 9. — Paragiopagurus insolitus sp. nov., Karubar Stn DW 28. a-e, holotype <J (2.2 mm) (MNHN-Pg 5363V

c 8ripPhTa.h PH 6 (2'° H ^ (U^NM 276.037): a' r'Sht second pereopod, lateral view; b, dactyl of same, mesial view!

f'ufTf pereop°d; >ateral view; d, dactyl of same, mesial view; e, sternite of third pereopods, ventral view;

f, left fourth pereopod, lateral view; g, left fifth pereopod, lateral view. Scale equals 0.5 mm (a-d), and 0.25 mm (e-g)

Source :

PARAPAGURIDAE FROM EASTERN INDONESIA

591

Fourth pereopod (Fig. 9f, 10a) semichelate. Dactyl terminating in blunt corneous claw almost entirely masked

by long, stiff plumose setae arising near base of claw; with ventrolateral row of small corneous spinules. Propodus

shorter than greatest height; rasp formed of 1 row of rounded or ovate scales. Carpus with long setae on dorsal

margin. Merus with rows of long plumose setae on dorsal and ventral margins.

Fifth pereopod (Fig. 9g) semichelate. Dactyl with about 4 or 5 fused corneous spines distally. Propodal rasp

extending to mid-length of segment.

Gills phyllobranchiate.

Uropods and telson (Fig. lOb-c) markedly asymmetrical. Telson lacking transverse suture; dorsal surface with

scattered setae; posterior lobes separated by wide and shallow unarmed cleft, terminal margins of lobes armed with

long, often strongly curved corneous spines.

Males lacking first gonopods, and with paired second gonopods. Second gonopods (Fig. lOd) each with distal

segment flat; distal segment with long setae marginally and on anterodistal face; basal segments each with scattered

setae, with or without rudimentary exopod. Females lacking vestigial second right pleopod; fifth left pleopod not

egg carrying.

Fig. 10. — Paragiopagurus insolitus sp. nov., Karubar Stn DW 28. a, d, paratype 6 (2.0 mm) (USNM 276037); b-c,

holotype 6 (2.2 mm): a, dactyl and distal end of propodus of left fourth pereopod, lateral view; b, left (b) and right (c)

uropods, dorsal view; c, telson, dorsal view; d, left second gonopod, anterior view. Scales equal 0.2 mm (a), 0.5 mm

(b, c), and 0.25 mm (d).

Source

592

R. LEMAITRE

Habitat. — Unknown (probably gastropod shells).

Distribution. — Known so far only from the Kai Islands, Indonesia. Depth: 448 to 467 m.

Etymology. — The specific name is from the Latin insolitus, unusual. The name is given for the unusual

condition of the ocular acicles, epistome, maxillipeds, and fourth pereopods.

REMARKS. — Paragiopagurus insolitus sp. nov. is a singularly distinctive species. The shape or conformation

of the ocular acicles, mouthparts, epistome, and dactyl of the fourth pereopod, are unusual or unique among

parapagurids. The relative length of the terminal spine of the ocular acicles is the longest known for any

parapagurid, and usually reaches nearly to midlength of the ocular peduncles. The incisor process of the mandible is

unusual in that it consists of several irregularly-shaped teeth (Fig, 8a), rather than having a single small median

tooth as in all other parapagurids for which the mandibles have been described. The second and third maxillipeds

each have a very long and slender exopod that is nearly 10 times as long as broad (Fig. 8e-f); in other parapagurids

the exopod is at most six times as long as broad. The epistome is strongly produced anteriorly, forming a

somewhat pyramid-shaped process with a concave ventral face (Fig. 7c); in other parapagurids the epistome is

evenly rounded.

The specialized setal arrangement seen on the dactyl of the fourth pereopod of P. insolitus sp. nov., is unique

among parapagurids (Fig. 10a). The claw of the dactyl is almost entirely masked by long plumose setae that arise

near the base and all around the claw. Although the function of this setal arrangement is unknown, it bears some

similarity with that reported by DE Saint Laurent (1968a, b) and McLaughlin (1997) for species of the

pagurid genus Decaphyllus de Saint Laurent, 1968.

Megalopa

Fig 11

Material EXAMINED. — Indonesia. Karubar, Tanimbar Islands : stn CP 59. 08°20'S, I32°U'E, 405-399 m,

31. 10.1991: 1 specimen (CL 5.5 mm, TL 23.4 mm), in gastropod shell (USNM 276038). — Stn CP 87, 08°47'S,

130°49'E. 1017-1024 m, 5.11.1991: 1 specimen (CL 5.6 mm, TL 22.1 mm) (MNHN-Pg 5382).

Remarks. — Although the two megalopal stage specimens reported herein evidently belong to the

Parapaguridae, they cannot be assigned with certainty at the present time to any particular species. The complete

larval rearing for any parapagurid species has not been achieved beyond the second zoeal stage (WILLIAMSON &

von Levetzow, 1967; Provenzano unpublished). The two Karubar specimens are identical morphologically,

and appear to be of the same species.

Of the two Karubar stations where parapagurid megalopae were obtained, no adult parapagurid stages were

found at Stn CP 59. Adult specimens of Parapagurus latimanus and Oncopagurus minutus were obtained at

Stn CP 87; conceivably the megalopae might be of one of these two species. However, O. minutus is a species

whose adult individuals are small, growing rarely to the size of the Karubar megalopae. Based on station co¬

occurrence and size, it is more likely that the megalopae are of P. latimanus, a species whose individuals can grow

to a size considerably larger than these postlarvae.

The Karubar specimens show similarities with the megalopae assigned by Lemaitre & McLaughlin

(1992: 754, Figs 6-7) to Sympagurus dimorphus (Sluder, 1883), but clearly differ at least in the cephalic shield,

development of the rostrum, and armature of the chelipeds. The shield in the Karubar megalopae (Fig. 1 la) has

a more pronounced and longer median longitudinal ridge on the anterior half than in S. dimorplius. The rostrum is

terminally rounded in both the Karubar and S. dimorphus megalopae; however, in the former, the rostrum has

a membranous ventral extension (Fig. 1 la-b) that is more developed than in the latter. The right and left chelipeds

of the Karubar specimens have spines or small tubercles on the dorsomesial margins of the palm and dactyl, and

on the dorsal margin of the carpus (Fig. 1 lc-d); the chelipeds are unarmed in the megalopae assigned to

S. dimorphus.

Source :

PARAPAGUR ID AE FROM EASTERN INDONESIA

593

FIG. 11. — Megalopa (CL 5.6 mm, TL 22.1 mm), Karubar Stn CP 87 (MNHN-Pg 5382): a, shield and ocular peduncles;

b, shield, left lateral view; c, right cheliped, dorsal view; d. left cheliped, dorsal view; e, abdominal somites 1-3, left

lateral view; f, telson, dorsal view. Scales equal 2 mm.

Source : MNHN, Paris

594

R. LEMAITRE

GENERAL REMARKS AND LIST OF INDONESIAN PARAPAGURIDS

Recent studies based largely on specimens obtained during the Karubar campaign indicate that the Indonesian

region harbors a previously unrecognized impressive diversity of hermit crabs of the family Paguridae

(McLaughlin, 1997). The Indonesian parapagurid fauna contains a total of 15 species, or about 25% of the

59 species of the family currently known worldwide, and evidently does not approach the species richness of the

Paguridae from the region. However, the discovery of two new species, Oncopagurus glebosus and Paragiopagurus

insolitus, characterized by unusual or unique conditions previously unknown in the family, are indicative of the

broad range of morphological diversity that exists in the Parapaguridae.

The following is a list of the 15 species and megalopa of Parapaguridae currently known from Indonesian

waters, including references where diagnoses and illustrations can be found. An asterisk indicates that the species

was not obtained during the Karubar campaign.

Parapagurus latimanus Henderson, 1888 — de Saint Laurent (1972, as P. pilosimanus latimanus)',

Lemaitre & McLaughlin (1992).

Strobopagurus sibogae (de Saint Laurent, 1972) — DE SAINT LAURENT (1972, as Parapagurus sibogae)',

Lemaitre (1996).

*Sympagurus affinis (Henderson, 1888) — DE Saint LAURENT (1972, as Parapagurus affinis)', Lemaitre

(1994).

Sympagurus brevipes (de Saint Laurent, 1972) — DE Saint LAURENT (1972, as Parapagurus brevipes)',

Lemaitre (1996).

*Sympagurus dofleini (Balss, 1912) — LEMAITRE (1994).

Sympagurus papposus Lemaitre, 1996 — Lemaitre (1996).

*Sympagurus planimanus (de Saint Laurent, 1972) — DE SAINT Laurent (1972, as Parapagurus

planimanus)', Lemaitre (1996).

*Sympagurus trispinosus (Balss. 1911) — LEMAITRE (1996).

*Oncopagurus indicus (Alcock, 1905) — LEMAITRE (1996).

Oncopagurus minutus (Henderson, 1 896) — LEMAITRE (1996).

Oncopagurus monstrosus (Alcock, 1894) — LEMAITRE (1996).

Oncopagurus orientalis (de Saint Laurent, 1972) — DE Saint LAURENT (1972, as Parapagurus oriental is)',

this report.

Oncopagurus glebosus sp. nov.

Paragiopagurus acutus (de Saint Laurent, 1972) — DE Saint LAURENT (1972, as Parapagurus acutus acutus );

Lemaitre (1996).

Paragiopagurus insolitus sp. nov.

Megalopa (sp. indet.) — this report.

ACKNOWLEDGMENTS

This study was possible thanks to Alain CROSNIER, who provided me with the opportunity to examine the

rich hermit crab collections that he and other French colleagues, especially Bertrand RICHER DE FORGES, have

obtained during recent expeditions to various Indo-Pacific regions. His editorial work and patience is also

gratefully acknowledged. I thank Tomoyuki Komai (Natural History Museum & Institute, Chiba, Japan), and

Patsy A. McLaughlin (Shannon Point Marine Center, Western Washington University, USA), for their useful

criticisms to the manuscript.

REFERENCES

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from H.M. Indian Survey Steamer 'Investigator1, Commander R.F. Hoskyn, R.N., commanding. Ser. 2, No. 1. Annals

and Magazine of Natural History , (6) 13: 225-245.

Source :

PARA PAGURIDAE FROM EASTERN INDONESIA

595

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 decapod species collected by the Royal Indian Marine Survey Ship

Investigator. Calcutta, iv + 286 pp., 3 pis.

ALCOCK, A., 1905. — Catalogue of the Indian Decapod Crustacea in the Collection of the Indian Museum. Part II.

Anomura. Fascicle I, Pagurides. Calcutta: Indian Museum, xi + 197 pp., pis 1-16.

ALCOCK, A, & ANDERSON, A.R.S., 1897. — Illustrations of the Zoology of the Royal Marine Surveying Steamer

Investigator. Crustacea, 5, pis 28-32, Calcutta.

Baba, K., Hayashi, K.-I. & TORIYAMA, M„ 1986. — Decapod crustaceans from continental shelf and slope around Japan.

Japan Fisheries Resource Conservation Association, Tosho Printing Co., Ltd., Tokyo, 336 pp.

Balss, H., 1911. — Neue Paguriden aus den Ausbeuten der Tiefsee-Expedition "Valdivia" und der japanischen Expedition

Prof. Dofleins. Zoologischer Anzeiger, 38: 1-9.

BaLSS, H„ 1912. — Paguriden. In: C. CHUN (ed.), Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition auf dem

Dampfer "Valdivia" 1898-1899. Jena, Verlag von Gustav Fischer, 20 (2): 85-124, pis 7-11.

Boone, L., 1926. — Unusual deep-sea Crustacea — Some forms secured by the Arcturus Oceanographic Expedition. A New

family of Crustacea. New York Zoological Society Bulletin. 29 (2): 69-73.

Gordan, J., 1956. — A bibliography of pagurid crabs, exclusive of Alcock, 1905. Bulletin of the American Museum of

Natural History, 108: 253-352.

Henderson, J.R., 1885. — In: T.H. Tizard et a!., Narrative of the cruise of the H.M.S. Challenger with a general account

of the scientific results of the expedition. Report on the Scientific Results of the Voyage of H.M.S. Challenger,

during the years 1873-76, 1 (2): 511-1110.

Henderson, J.R., 1888. — Report on the Anomura collected by H.M.S. Challenger during the years 1873-76. Report on

the Scientific Results of the Voyage of H. M. S. Challenger during the years 1873-76, (Zoology), 27: xi + 221 pp.,

pis 1-21.

Henderson, J.R., 1896. — Natural history notes from H.M. Indian Marine Survey Steamer "investigator", Commander

C.F. Oldham, R.N., commanding, Ser. 2, No. 24. Report on the Paguridae collected during the season 1893-94.

Journal of the Asiatic Society • of Bengal, 65 (3): 516-536.

Imafuku, M„ 1992. — Anomuran members. In: M. Takeda (ed.), Invertebrates. 8, Macrura, Anomura, and others. The

earth for animals, 68: 234-235. Asahi Shimbun Press, Tokyo. (In Japanese).

Kemp, S. & Sewell, R.B.S., 1912. — II. Notes on Decapoda in the Indian Museum. III. The species obtained by

R.I.M.S.S. "Investigator" during the survey season 1910-1 1. Records of the Indian Museum, 7 (1): 15-32, pi. 1.

Lemaitre, R., 1986. — Western Atlantic species of the Parapagurus pilosimanus complex (Anomura: Paguroidea:

Parapaguridae): description of a new species and morphological variations. Journal of Crustacean Biology, 6 (3):

525-542.

Lemaitre, R., 1989. — Revision of the genus Parapagurus (Anomura: Paguroidea: Parapaguridae), including

redescriptions of the western Atlantic species. Zoologische Verhandelingen, (253): 1-106.

Lemaitre, R„ 1993. — A new genus of Parapaguridae (Decapoda, Anomura). Crustacean Research. 22: 11-20.

Lemaitre, R., 1994. — Crustacea Decapoda: Deep-water hermit crabs (Parapaguridae) from French Polynesia with

descriptions of four new species. In: A. Crosnier (ed.), R6sultats des Campagnes Musorstom, Volume 12. Memoires

du Museum national d'Histoire naturelle, 161: 375-419.

Lemaitre, R., 1996. — Hermit crabs of the family Parapaguridae (Crustacea: Decapoda: Anomura) from Australia: species

of Strobopagurus Lemaitre, 1989, Sympagurus Smith, 1883, and two new genera. Records of the Australian Museum.

48 (2): 163-221.

Lemaitre, R. & McLaughlin, P.A., 1992. — Descriptions of megalopa and juveniles of Sympagurus dimorphus (Studer,

1883), with an account of the Parapaguridae (Crustacea: Anomura: Paguroidea) from Antarctic and Subantarctic waters.

Journal of Natural History, 26: 745-768.

McLaughlin, P.A., 1974. — The hermit crabs (Crustacea Decapoda. Paguridae) of northwestern North America.

Zoologische Verhandelingen, (130): 1-396.

McLaughlin, P.A., 1997. — Crustacea Decapoda: hermit crabs of the family Paguridae from the Karubar Expedition in

Indonesia. In: A. Crosnier & P. BouCHET (eds), Resultats des Campagnes Musorstom, Volume 16. Memoires du

Museum national d'Histoire naturelle, 172: 433-572.

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Miyake, S„ 1978. — The crustacean Anomura of Sagami Bay. Biological Laboratory, Imperial Household. 161 pp.

Miyake, S., 1982. — Japanese crustacean decapods and stomatopods in color. Vol. I, Macrura, Anomura, and

Stomatopoda. Hoikusha Publishing Co., Ltd., Osaka, 261 pp.

Murray, J„ 1895. — Report on the scientific results of the voyage of H.M.S. Challenger during the years 1873-76.

A summary of the scientific results. Report on the scientific results of the voyage of H.M.S. Challenger, during the

years 1873-76, (Zoology), 1-2: 1-1608.

Osawa, M., 1995. — A new parapagurid genus, Tsunogaipagurus, for Sympagurus chuni (Balss, 1911) (Crustacea:

Decapoda: Anomura). Proceedings of the Japanese Society of Systematic Zoology, (53): 62-70.

Saint Laurent, M. de, 1968a. — Revision des genres Catapaguroides et Cestopagurus et description de quatre genres

nouveaux. I. Catapaguroides A. Milne Edwards et Bouvier et Decapltyllus nov. gen, (Crustaces Decapodes Paguridae).

Bulletin du Museum national d'Histoire naturelle, (2) 39 (5) [1967]: 923-954.

Saint Laurent, M. de, 1968b. — Revision des genres Catapaguroides et Cestopagurus et description de quatre genres

nouveaux. I. Catapaguroides A. Milne Edwards et Bouvier et Decapltyllus nov. gen. (Crustaces Decapodes Paguridae)

(suite). Bulletin du Museum national d’Histoire naturelle, (2) 39 (6) [1967]: 1100-11 19.

Saint Laurent, M. de, 1972. — Sur la famille des Parapaguridae Smith. 1882. Description de Typhlopagurus foresti gen.

nov., et de quinze espbces ou sous-especes nouvelles de Parapagurus Smith (Crustacea, Decapoda). Biidraeen tot de

Dierkunde, 42 (2): 97-123.

Smith, S.I., 1879. — The stalked-eyed crustaceans of the Atlantic coast of North America north of Cape Cod. Transactions

of the Connecticut Academy of Arts and Sciences, 5 (1): 27-136, pis 8-12.

Smith. S.I.. 1882. — XVII. Report on the Crustacea. Part I. Decapoda. Reports on the dredging, under the supervision of

Alexander Agassiz, on the east coast of the United States, during the summer of 1880, by the U.S. Coast Survey

Steamer "Blake", commander J.R. Bartlett U.S.N., commanding. Bulletin of the Museum of Comparative Zoology,

Harvard College, 10 (1): 1-108, pis 1-16.

Smith, S.I., 1883. — Preliminary report on the Brachyura and Anomura dredged in deep water off the south coast of New

England by the United States Fish Commission in 1880, 1881, and 1882. Proceedings of the United States National

Museum, 6 (1): 1-57, pis 1-6.

Studer, T„ 1883. — Verzeichniss der Crustaceen, welche wahrend der Reise S.M.S. Gazelle an der Weskuste von Africa,

Ascension und dem Cap der guten Hoffnung gesammelt wurden. Abandlungen der Preussischen Akademie der

Wissenschaften, 2 (1882-1883): 1-32, pis 1-2.

Wiluamson, D.I. & von Levetzow, K.G., 1967. — Larvae of Parapagurus diogenes (Whitelegge) and some related species

(Decapoda, Anomura). Crustaceana, 12 (2): 179-192.

Source : MNHN, Paris

RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS

Crustacea Decapoda: Species of the genera

Agononida Baba & de Saint Laurent, 1996

and Munida Leach, 1820 (Galatheidae)

from the KARUBAR Cruise

Enrique MACPHERSON

Centro de Estudios Avanzados de Blanes (CSIC)

Cami de Santa Barbara s/n

17300 Blanes (Girona). Spain.

ABSTRACT

Twenty six species of galaiheid crustaceans belonging to the genera Agononida Baba & de Saint Laurent, 1995 and

Munida Leach. 1820, were caught off the Molucca archipelago, during the Karubar Cruise (October-November, 1991).

Three species are described as new: A. emphereia, M. compacta and M. punctata.

RESUME

Crustacea Decapoda : Especes des genres Agononida Baba & de Saint Laurent, 1995 et Munida

Leach, 1820 (Galatheidae) recoltees lors de la campagne KARUBAR.

Les espfeces des genres Agononida Baba & de Saint Laurent. 1995, et Munida Leach, 1820, recoltdes dans 1 archipel

des Moluques sont au nombre de 26. Trois espbces (A. emphereia, M. compacta et M. punctata) sont nouvelles.

A. emphereia, proche de A. soelae (Baba, 1986) et de A. incerta (Henderson, 1888) se differencie facilement par

l'armature de la carapace. M. compacta est proche de M. rhodonia Macpherson, 1994, mais se differencie par la forme et

l'armature des chelipcdes. M. punctata se distingue de M. rubrodigitalis Baba, 1994, par la forme du rostre.

INTRODUCTION

The genera Agononida Baba & de Saint Laurent, 1995 and Munida Leach, 1820 are represented in Indonesian

waters by more than 15 species (Baba, 1988; Baba & de Saint Laurent, 1995; Macpherson, 1993, 1994;

MACPHERSON & Baba, 1993). During a recent cruise (Karubar) carried out in October-November

Macpherson, E., 1997. — Crustacea Decapoda: Species of the genera Agononida Baba & de Saint Laurent, 1995 and

Munida Leach, 1820 (Galatheidae) from the Karubar Cruise. In: A. Crosnier & P. Bouchet (eds), Resultats des

Campagnes Musorstom, Volume 16. Mem. Mus. natn. Hist, nat., 172: 597-612. Paris ISBN 2-85653-506-2,

Source : MNHN, Paris

598

E. MACPHERSON

1991 to the Molucca Archipelago (Kai and Tanimbar islands) (see CROSNIER et all., 1997), numerous

representatives of these genera were collected. The aim of this paper is to study this interesting collection.

The types of the new species and other material have been deposited in the collections of the Museum national

d'Histoire naturelle, Paris (MNHN), the Puslitbang Oseanologi-LIPI, Jakarta (POLIPI) and the National Museum

of Natural History, Washington. When no information on the depository is provided, the material is held by the

MNHN, Paris. Measurements given are of carapace length excluding rostrum, and the terminology used mainly

follows previous papers (MACPHERSON & de SAINT Laurent, 1991). In order to avoid repetitious descriptions,

only distinctive characters have been included in the text.

LIST OF STATIONS

The abbreviations of the gears used arc: DW = Waren dredge, CP = Beam trawl, CC = Otter trawl.

Station DW 1. — 22.10.1991, 05°46’S, 132°10'E, 156-305 m: M. agave. M. caesura. Munida sp.

Station DW 2. — 22.10.1991, 05°47'S, 132°13’E, 209-240 m: M. leptitis, M. philippinensis, Munida sp.

Station CP 5. — 22.10.1991, 05°49'S, 132°18'E, 296-299 m: A. incerta, A. squamosa, M. caesura, M. leptitis,

Munida sp.

Station CP 6. — 22.10.1991, 05°49'S, 132°21'E, 287-298 m: A. incerta, M. agave, M. striola, Munida sp.

Station CP 9. — 23.10.1991, 05°23'S, 132°29'E, 368-389 m: A. incerta, M. compacta, M. punctata,

M. striola.

Station CC 10. — 23.10.1991, 05°21'S, 132°30'E, 329-389 m: A. incerta, A. similis, M. compacta.

Station CP 12. — 23.10.1991, 05°23'S, 132°37'E, 413-436 m: A. incerta, A. similis, M. compacta.

Station DW 13. — 24.10.1991, 05°26'S, 132°38'E, 417-425 m: M. compacta.

Station DW 14. — 24.10.1991, 05°18'S, 132°38'E, 245-246 m: M. caesura. M. leptitis.

Station CP 15. — 24.10.1991, 05° 17'S, 132°41'E, 212-221 m: M. agave.

Station CP 16. — 24.10.1991, 05°17'S, 132°50'E, 315-349 m: A. incerta, M. caesura, M. striola.

Station CP 17. — 24.10.1991, 05°15’S, 133°01'E, 439-459 m: M. compressa.

Station DW 18. — 24.10.1991, 05°18'S, 133°01'E, 205-212 m: M. agave, M. caesura, M. hyalina,

M. rufiantennulata.

Station CP 19. — 25.10.1991, 05°15'S, 133°0I'E, 576-605 m: /\. soelae, M. compacta.

Station CP 20. — 25.10.1991, 05°15'S, 132°59'E, 769-809 m: M. curvirostris, M. microps.

Station CC 21. 25.10.1991, 05°14'S, 133°00'E, 688-694 m: A. emphereia, M. compacta, M. curvirostris

Station DW 22. — 25.10.1991, 05°22'S, I33°01’E, 85-124 m: M. clinata, M. minuta.

Station CP 25. 26.10.1991, 05°30'S, 132°52'E, 336-346 m: A. incerta, A. squamosa, M. caesura,

M. leviantennata, M. punctata, M. rubrodigitalis.

Station CP 26. — 26.10.1991, 05°34'S, I32°52'E, 265-302 m: A. incerta.

Station CP 27. 26.10.1991, 05°33'S, 132°51'E, 304-314 m: A. incerta, A. squamosa, M. caesura.

Station DW 32. — 26.10.1991, 05°47'S, 132°51'E, 170-206 m: M. japonica.

Station CP 33. — 27.10.1991, 06°05’S, 132°38'E, 307-311 m: A. incerta, M. striola.

Station CP 35. 27.10.1991, 06°08'S, 132°45'E, 390-502 m: A. incerta, M. compacta, M. leviantennata,

M. punctata.

Station CP 36. - 27.10.1991, 06°05'S, 132°44'E, 210-268 m: A. squamosa, M. caesura, M. striola.

Station CP 39. — 28.10.1991, 07°47'S, 132°26'E, 466-477 m: M. compacta

Station CC 40. - 28.10.1991, 07°46'S, 132°31'E, 443-468 m: A. incerta, M. compacta.

Station CC 41. - 28.10.1991, 07°45'S, 132°42'E, 393-401 m: A. incerta, M. compacta.

Station CC 42. - 28.10.1991, 07°53'S, 132°42'E, 350-354 m: A. incerta, A. similis, M. caesura,

M. compacta, M. striola.

Station DW 44. — 29.10.1991, 07°52'S, 132°48'E, 291-295 m: M. caesura.

Station CP 45. - 29.10.1991, 07°54'S, 132°47'E, 302-305 m: A. incerta., M. caesura.

Station CP 46. - 29.10.1991, 08°01'S, 132°51'E, 271-273 m: A. incerta, A. similis, M. caesura

Source :

SPECIES OF THE GENERA AGONONIDA AND MUNIDA FROM THE KARUBAR EXPEDITION

599

Station CP 47. — 29.10.1991, 08°01'S, 132°55'E, 235-246 m: A. incerta, A. similis, M. striola.

Station DW 49. — 29.10.1991, 08°00'S, 132°59'E, 206-210 m: M. japonica, M. leptitis, M. philippinensis.

Station CC 56. — 31.10.1991, 08°16’S, 131°59’E, 549-552 m: M. compacta.

Station CC 57. — 31.10.1991, 08°19'S, 131°53'E, 603-620 m: 4. soelae, M. compacta.

Station CC 58. — 31.10.1991, 08°19'S, 132°02'E, 457-461 m: M. compacta.

Station CP 59. — 31.10.1991, 08°20'S, 132°1 l'E, 399-405 m: A. incerta, M. compacta.

Station CP 62. — 01.1 1.1991, 09°01'S, 132°42’E, 246-253 m: A. similis., M. compacta.

Station CP 63. — 01.1 1.1991, 08°00’S, 132°58'E, 213-214 m: A. similis, M. armata.

Station CP 65. — 01.1 1.1991, 09°14’S, 132°27’E, 174-176 m: M. armata, M. philippinensis.

Station CP 66. — 01.1 1.1991, 09°01'S, 132°09'E, 211-217 m: A. similis.

Station CP 67. — 01.1 1.1991, 08°58'S, 132°06'E, 146-233 m: A. similis, M. philippinensis, M. striola.

Station CP 69. — 02.11.1991, 08°42'S, 131°53'E, 356-368 m: A. incerta, M. compacta, M. leviantennata,

M. striola.

Station CP 70. — 02.1 1.1991, 08°41'S, 131°47'E, 410-413 m: A. incerta, M. compacta.

Station CP 71. — 02.1 1.1991, 08°38'S, 131°44'E, 477-480 m: A. incerta, M. compacta.

Station CP 72. — 02.1 1.1991, 08°36'S, 131°33'E, 676-699 m: 4. eminens.

Station CP 75. — 03.1 1.1991, 08°46'S, 131°36'E, 451 m: M. compacta.

Station CP 76. — 03.1 1.1991, 08°50'S, 131°33TE, 400-401 m: 4. incerta, M. compacta.

Station CP 77. — 03.1 1.1991, 08°57'S, 131°27'E, 346-352 m: 4. incerta, M. compacta, M. leviantennata.

Station CP 78. — 03.11.1991, 09°06'S, 131°24'E, 284-295 m: 4. similis.

Station CP 79. — 03.1 1.1991, 09°16'S, 131°22'E, 239-250 m: 4. similis, M. kuboi, M. striola.

Station CP 82. — 04.1 1.1991, 09°32'S, 131°02'E, 215-219 m: 4. similis.

Station CP 83. — 04.11.1991, 09°23'S, 131°00’E, 285-297 m: 4. incerta, 4. similis, M. kuboi, M. philip¬

pinensis, M. striola.

Station CP 84. — 04.11.1991, 09°23'S, 131°09'E, 246-275 m: 4. incerta, M. kuboi, M. philippinensis,

M. striola.

Station CP 85. — 04.11.1991, 09°22'S, 131°14'E, 240-245 m: 4. incerta, A. similis, M. rubrodigitalis,

M. striola.

Station CP 86. — 04.1 1.1991, 09°26'S, 131°13'E, 223-225 m: 4. similis.

Station CP 87. — 05.1 1.1991, 08°47'S, 130°49'E, 1017-1024 m: M. microps.

SYSTEMATIC ACCOUNT

Genus AGONONIDA Baba & de Saint Laurent, 1995

Agononida emphereia sp. nov.

Fig. 1

Material EXAMINED. — Indonesia. Karubar: stn 21, 688-694 m: 1 $ 18,5 mm (MNHN-Ga 3947); 3d 18.2 to

21.4 mm; 1 ov. 2 20.5 mm (MNHN-Ga 3948).

TYPES. — The male of 18.5 mm from the Stn 21 (MNHN-Ga 3947) has been selected as holotype, the other

specimens are paratypes.

ETYMOLOGY. — From the Greek, emphereia, likeness (considered here as a substantive in apposition), in

reference to the similarity between the new species and other closely related species.

DESCRIPTION. — Carapace with numerous secondary striae and scales. One protogastric spine behind each

epigastric spine (practically absent in one specimen). One small spine on hepatic region, usually on each side.

Three or four spines in a row on each branchiocardiac boundary, anteriormost postcervical, larger than remainder.

Source : MNHN, Paris

600

E. MACPHERSON

Cardiac region with 1-2 spines (absent in one specimen). Posterior border of carapace unarmed. Anterolateral spine

pronounced, situated at anterolateral angle of carapace. Branchial margins with 4 spines. Fourth to seventh thoracic

sternites with numerous short arcuate striae. Second, third and fourth abdominal segments each with 4 spines on

anterior transverse ridge; posterior ridge of fourth segment with strong median spine. Eye moderately large,

maximum corneal diameter about 1/3 length of anterior border of carapace between bases of external orbital spines.

Basal antennular segment (distal spines excluded) not exceeding eye, distolateral spine shorter than distomesial.

Distomesial prolongation of first antennal segment well developed, reaching rostral tip; second segment with

2 strong distal spines, distomesial spine slightly overreaching third segment, 1-2 spines on mesial border; third

segment unarmed. Flexor margin of merus of third maxilliped with median spine, extensor border with distal

spine. Fingers of cheliped unarmed, fixed finger bifid distally. First walking leg about 3 times carapace length.

Dactylus of walking legs about 2/3 propodus length, armed with small spine-like setae on median third of ventral

border.

Remarks. — The new species is closely related to A. soelae (Baba, 1986) from North-West Australia,

New Caledonia and Indonesia and A. incerta (Henderson, 1888) (see below) in the general spination of the

carapace. The three species have spines on the anterior and posterior ridges of the fourth abdominal. The new

species differs from A. soelae in the following aspects:

— The carapace and sternal plastron are less rugose in A. soelae than in the new species.

— The distomesial prolongation of the first antennal segment is short in A. soelae , reaching the end of the

second segment of the antennal peduncle; this prolongation is very long in the new species, clearly overreaching

the antennal peduncle and reaching rostral tip.

— The posterior border of the carapace is unarmed in the new species, whereas A. soelae has 4-6 spines.

The extensor border of the merus of the third maxilliped is armed with one distal spine in A. emphereia,

unarmed in A. soelae.

The new species is distinguished from A. incerta by the following characters:

— Two protogastric spines in the new species, absent in A. incerta.

One small hepatic spine, usually on each side, is present in A. emphereia , absent in A. incerta.

The cardiac spines are always absent in A. incerta , whereas in A. emphereia they are usually present.

Distribution. — Indonesia, between 688 and 694 m.

Agononida eminens (Baba, 1988)

Munida eminens Baba, 1988: 95, fig. 35; 1994: II. — Macpherson, 1994: 456, fig. 72; 1995: 392.

Material EXAMINED. — Indonesia. Karubar: stn 72, 676-699 m: 1 8 15.8 mm.

»IS™BUT10N- ~ ThC species was describcd from specimens collected in the Philippines between 564 and

686 m (Baba, 1988). Since then the species has been cited in eastern Australia, New Caledonia, Loyalty Islands,

Chesterfield Islands, Wallis and Futuna area and Indonesia, between 650 and 970 m (Baba 1 994

Macpherson, 1994; 1995). The present material has been collected from 676-699 m.

Agononida incerta (Henderson, 1888)

"“fig a74”;C'l995He3n9d4erSOn 18881 * ,3’ ^ 4a‘ ~ Baba' l988; '<* 1994: .2. - Macpherson, .994: 478,

Material EXAMINED. — Indonesia. Karubar: stn 5, 296-299 m: 6 8 17 4 to 27 0 mm- 12 9 15 8 tn

389 m-^ «J 1?8 » 20 09im- l 9 n n '° 3 °V‘ 9 216 t0 25 4 mm; 162 115 t0 27 0 mm- - Stn 9, 368-

, Z o 20;0 mm> 8 2 12 0 t0 17 6 mm; 1 JUV- 5.0 mm (POL1PI). — Stn 10, 329-389 nr 8 <3 13 3 to

30.7 mm; 5 ov. 9 21.3 to 23.8 mm; 3 9 22.4 to 23.6 mm. - Stn 12, 4.3-436 m: 3 <3 12 4 to 29" mm; 3 o v 9

Source :

SPECIES OF THE GENERA AGONONIDA AND MUNIDA FROM THE KARUBAR EXPEDITION

601

b, sternal plastron; c. ventral view of cephalic region, showing antennular and antennal peduncles; d, right third

maxilliped, proximal segments of endopod, lateral view; e, left chela, dorsal view; f, right first walking leg, lateral

view.

Source : MNHN, Paris

602

E. MACPHERSON

20.0 to 24.0 mm; 4 2 12.5 to 30.7 mm; 1 juv. 8.0 mm (POLIPI). — Stn 16, 315-349 m: 5 3 17.6 to 24.6 mm;

2 2 12.0 and 18.0 mm. — Stn 25, 336-346 m: 3 3 11.8 to 27.0 mm; 5 ov. 2 11.0 to 23.0 mm; 1 juv. 5.6 mm.

Stn 26, 265-302 m: 1 juv. 5.3 mm. — Stn 27, 304-314 m: 1 3 19.5 mm. — Stn 33, 307-311 m: 2 3 14.8 and

15.0 mm; I 2 7.0 mm. — Stn 35, 390-502 m: 9 3 11.5 to 27.0 mm; 3 2 11.0 to 20.0 mm. — Stn 40, 443-468 m:

7 3 18.3 to 35.8 mm; 3 ov. 2 25.2 to 36.8 mm; 12 2 13.4 to 30.6 mm (POLIPI). — Stn 41, 393-401 m: 1 1 3 13.8 to

40.4 mm; 4 ov. 2 21.3 to 30.5 mm; 5 2 10.6 to 25.2 mm; 1 juv. 10.4 mm. — Stn 42, 350-354 m: 4 3 15,0 to

20.6 mm; 2 ov. 2 24.1 and 24.4 mm; 2 2 15.0 and 19.7 mm. — Stn 45. 302-305 m: 1 2 9.5 mm. — Stn 46, 217-

273 m: 2 3 28.4 and 31.5. — Stn 47, 235-246 m; 3 3 27.4 to 29.8 mm; 1 ov. 2 24.6 mm; 2 2 17.5 to 26.4 mm. _

Stn 59, 399-405 m:6d 9.7 to 15.6 mm; 1 ov. 2 22.4 mm; 6 2 14.7 to 15.0 mm (POLIPI). — Stn 69, 356-368 m:

2 3 23.5 and 25.7 mm; 3 ov. 2 22.0 to 25.6 mm; 1 2 16.8 mm. — Stn 70, 410-413 m: 3 3 14.5 to 16.4 mm;

10 2 15.0 to 18.4 mm (POLIPI). — Stn 71, 477-480 m: 3 2 11.4 to 14.4 mm. — Stn 76, 400-401 m: 3 3 9.7 to

18.5 mm; 4 2 15.8 to 25.0 mm. — Stn 77, 346-358 m: 1 3 23.0 mm; 1 ov. 2 25.8 mm (POLIPI). — Stn 83, 285-

297 m: 1 3 32.4 mm; 2 ov. 2 23.6 and 27.8 mm; 1 2 16.3 mm. — Stn 84, 246-275 m: 2 3 13.5 and 17 6 mm-

1 2 6.8 mm. — Stn 85, 240-245 m: 8 3 16.0 to 30.5 mm; 1 2 25.0 mm.

Distribution. — Known from east African coast, Japan, Philippines, Indonesia, eastern Australia, New

Caledonia, Loyalty Islands, Chesterfield Islands, Wallis and Futuna, and Kiribati, between 17 and 720 m. The

specimens examined were collected in 217-502 m.

Agononida similis (Baba, 1988)

Munida similis Baba, 1988: 129, figs 49-50.

Material EXAMINED. — Indonesia. KARUBAR: stn 10. 329-389 m: 10 3 16.0 to 23.6 mm; 2 ov. 2 20.0 and

20.7 mm; 6 2 14.3 to 21.0 mm. — Stn 12, 413-436 m: 1 2 12.6 mm. — Stn 42, 350-354 m; 1 3 14.7 mm; 5 2 1 1 6 to

i3n3 .Ti' ~ S,n 46‘ 27 1-273 m: 1 2 16 5 mm- — Stn 47- 235-246 m: 4 3 14.0 to 21.0 mm; 3 ov. 2 19.0 to 22.4 mm-

7 2 14.0 to 19.1 mm. — Stn 62, 246-253 m: 8 3 11.8 to 22.0 mm; 1 ov. 2 19.1 mm; 6 2 5.7 to 17.8 mm (POLIPI). —

Stn 63. 214-215 m: 2 3 18.0 and 19.1 mm; 1 2 14.8 mm. — Stn 66, 211-217 m: 1 3 13 6 mm-

2 2 15.4 and 16.0 mm (POLIPI). - Stn 67. 146-233 m: 7 <J 7.8 to 20.4 mm; 5 2 9.0 to 18.5 mm. - Stn 78 284-

tPnMPtt I-6 S'n l9' 239~25° m: 9 13 15 0 10 16 6 mm; 1 ov- 9 17'2 mm; 3 9 8 0 10 l7-5 mm

P0^PI ar, n82’213'219 m: 3 20.6 mm; 1 ov. 2 20.5 mm. — Stn 83, 285-297 m: 7 2 5.6 to 8.0 mm. —

Stn 84 246-275 m : 2 2 6.8 and 7.3 mm. - Stn 85. 240-245 m: 11 3 12.5 to 22.0 mm; 4 ov. 2 17.7 to 18.0 mnv

(POLIPI) l° mm; ' jUV' 9 0 mm (P0LIPI)- ~ Sln86' 223-225 m: 7 13 15 4 10 21-7 mm; 4 2 15.1 to 16.5 mm

Remarks. — Agononida similis is closely related to A. squamosa Henderson. 1885 (see below). Both species

can be distinguishable by the shape of the rostrum and the mesiodistal spine of the first segment of the antennal

peduncle (Baba, 1988). The specimens examined here always have the posterior border of the carapace without

spines, whereas in all other specimens of A. squamosa examined two spines are present. Although Baba (1988)

showed certain variations in the presence of these spines, the constancy here observed suggests that this character

may have a specific value.

distribution. - Philippines and Indonesia in 263-494 m (Baba, 1988). The present material has been

collected between 146 and 436 m.

Agononida soelae (Baba, 1986)

Munida soelae Baba. 1986: 2, fig. 3; 1988: 82 (key). — Macpherson, 1994: 530.

i ■*N?cfrTERIAL EXAMINED- — Indonesia. Karubar: stn 19, 576-605 m: 1 3 13.4 mm

i o iv. u mm.

Stn 57, 603-620 m:

Distribution. - North-West Australia and New Caledonia, between 450 and 600 m (Baba 1986

Macpherson, 1994). Indonesia, between 576 and 620 m.

Source :

SPECIES OF THE GENERA AGONONIDA AND MUNIDA FROM THE KARUBAR EXPEDITION

603

Agononida squamosa (Henderson, 1885)

Munida squamosa Henderson, 1885: 409; 1888, 131, pi. 13, fig. 1. — MACPHERSON, 1993: 425, fig. 1 h-i; 1994: 537,

fig. 96; 1995: 406. — Baba, 1994: 16.

Material EXAMINED. — Indonesia. Karubar: sin 5, 296-299 m: 2 5 11.6 and 12.0 mm; 1 ov. 2 16.2 mm;

1 2 10.0 mm. — Stn 25, 336-346 m: 1 ov. 2 14.0 mm (USNM). — Stn 27, 304-314 m: 1 2 13.5 mm; 1 ov. 2

14.7 mm; 1 2 10.6 mm (POLIPI). — Sin 36, 210-268 m: 2 2 7.7 and 9.4 mm.

Distribution. — Japan, Indonesia, Admiralty Islands, northeastern Australia, New Caledonia, Loyalty Islands

and Wallis Island, between 176 and 752 m (MACPHERSON, 1994, 1995). The material examined here was collected

in 210-346 m.

Genus MUNIDA Leach, 1820

Munida agave Macpherson & Baba, 1993

Munida agave Macpherson & Baba, 1993: 387, figs 1-2.

MATERIAL EXAMINED. — Indonesia. Karubar: stn 1, 156-305 m: 5 d 4.0 to 8.0 mm; 5 2 5.1 to 9.3 mm. —

Stn 6, 289-298 m: 1 d 10.7 mm. — Stn 15, 212-221 m: 2 2 5.0 and 5.7 mm. — Stn 18. 212-221 m: 4 d 5.0 to

6.3 mm; 2 2 4.0 and 5.1 mm.

Remarks. — The specimens examined are not well preserved and the antennular peduncles of many specimens

are broken. Therefore, one of the important specific characters (the length of the distal spines) is often unknown.

Most of the morphological characters agree quite well with the description of M. agave. However, the number of

spines along the anterior border of the second abdominal segment is variable (0 to 6) and in one specimen the

thoracic sternites are squamate (smooth in M. agave). As it was pointed out in the original description,

probably more than one species are included in this complex. Therefore, in view of the difficulties in the

correct identification, the present material is provisionally referred to M. agave until more specimens become

available.

Distribution. — Japan and Philippines in 89-197 m (Macpherson & Baba, 1993), Indonesia, between 156

and 305 m.

Munida armata Baba, 1988

Munida armata Baba, 1988: 86, fig. 31. — Macpherson, 1993: 427.

Material EXAMINED. — Indonesia. Karubar: stn 63, 214-215 m: 5 d 12.7 to 16.0 mm. — Stn 65, 174-

176 m: 2 d 16.0 and 16.2 mm; 2 2 7.7 to 13.4 mm (POLIPI).

Distribution. — Philippines in 182-216 m (Baba, 1988; Macpherson, 1993), Indonesia, between 174 and

215 m.

Munida caesura Macpherson & Baba, 1993

Munida caesura Macpherson & Baba, 1993: 388, fig. 3.

Source : MNHN, Paris

604

E. MACPHERSON

"isssiiss sss

Source : MNHN, Paris

SPECIES OF THE GENERA ACONONIDA AND M UN IDA FROM THE KARUBAR EXPEDITION

605

MATERIAL EXAMINED. — Indonesia. Karubar: stn 1, 156-305 m: 5 8 4.7 to 8.6 mm; 6 9 3.6 to 9.3 mm. —

Stn 5, 296-299 m: 15 8 7.6 mm to 17.5 mm; 20 9 9.7 to 15.3 mm. — Stn 14, 245-246 m; 2 8 9.0 and 12.4 mm;

3 9 10.0 to 14.8 mm (USNM). — Stn 16, 315-349 m: 3 8 9.0 to 10.5 mm (POLIPI). — Stn 18, 205-212 m: 10 8 3.1 to

9.3 mm; 4 9 4.0 to 8.5 mm. — Stn 25, 336-346 m: 1 8 8.1 mm; 3 9 9.0 to 14.4 mm. — Stn 27, 304-314 m:

2 8 8.3 and 11.0 ram; 1 9 12.5 mm. — Stn 36, 210-268 m: 1 ov. 9 15.2 mm; 2 9 12.6 and 17.7 mm. — Stn 42,

170-206 m; 1 8 10.8 mm. — Stn 44, 291-295 m: 3 9 3.4 to 9.6 mm. — Stn 45, 302-305 m: 1 9 9.3 mm. — Stn 46,

271-273 m: 4 8 8.4 to 10.3 mm; 2 ov, 9 12.3 and 12.5 mm; 1 9 8.9 mm (POLIPI).

Distribution. — Japan, Philippines, Indonesia in 250-390m (Macpherson & Baba, 1993). The present

material was collected in 156-346 m.

Munida clinata Macpherson, 1994

Munida clinata Macpherson, 1994: 457, fig. 11.

MATERIAL EXAMINED. — Indonesia. Karubar: stn 22, 85-124 m: 4 8 4.0 to 5.4 mm.

DISTRIBUTION. — Munida clinata was described from specimens collected in the Philippines, New Caledonia

and Chesterfield Islands, between 28 and 245 m (MACPHERSON, 1994). The material from Indonesia was caught in

85-124 m.

Munida compacta sp. nov.

Fig. 2

Munida curvirostris Macpherson, 1993: 428 (in part).

MATERIAL EXAMINED. — Indonesia. Karubar: stn 9, 368-389 m: 23 8 8.3 to 13.3 mm; 39 9 10.1 to 17.4 mm

(USNM). — Stn 10, 329-389 m: 17 8 11.4 to 19.7 mm; 34 9 9.0 to 17.3 mm. — Stn 12, 413-436 m: 6 8 10.0 to

16.4 mm; 5 9 11.8 to 17.5 mm (POLIPI). — Stn 13, 417-425 m: 1 8 11.6 mm . — Stn 19, 576-605 m: 1 9 6.2 mm. —

Stn 21, 688-694 m:6J 6.3 to 18.6 mm; 1 1 9 5.7 to 16.5 mm. — Stn 35, 390-502 m:7J 9.0 to 16.0 mm; 8 9 12.7

to 19.0 mm. — Stn 39, 466-477 m: 9 8 8.2 to 16.8 mm; 1 ov. 9 16.0 mm; 8 9 7.4 to 15.4 mm. — Stn 40, 443-

468 m: 12 8 11.5 to 17.4 mm; 10 9 12.2 to 21.0 mm (USNM). — Stn 41, 393-401 m:6<5 14.0 to 18.3 mm; 5 9 9.0

to 18.1 mm. — Stn 42, 350-354 m: 6 8 13.6 to 20.0 mm; 4 ov. 9 17.4 to 19.4 mm. — Stn 56. 549-552 m: 4 8 12.7 to

15.0 mm; 1 9 16.2 mm. — Stn 57, 603-620 m: 1 9 8.7 mm. — Stn 58, 457-461 m: 1 8 18.8 mm. — Stn 59, 399-

405 m: 10 8 8.8 to 19.4 mm; 2 ov. 9 16.6 and 17.5 mm; 13 9 8.3 to 18.2 mm (POLIPI). — Stn 62, 246-253 m: 1 9

12.0 mm, — Stn 69, 356-368 m: 13 8 8.6 to 20.3 mm; 1 ov. 9 16.6 mm (MNHN-Ga 3949); 4 ov. 9 16.8 to 20.6 mm;

16 9 8.7 to 22.8 mm. — Stn 70, 410-413 m: 1 8 14.7 mm; 5 9 9.8 to 18.4 mm. — Stn 71. 477, 480 m: 1 8 9.7 mm;

6 9 9.4 to 15.2 mm. — Stn 75, 451-452 m: 2 8 14.0 and 15.3 mm; 3 9 9.4 to 14.1 mm. — Stn 76. 400-401 m: 5 8

10.0 to 15.0 mm; 3 9 10.0 to 1 1.4 mm. — Stn 77, 346-352 m: 3 9 18.3 to 19.0 mm (POLIPI).

Types. — The ovigerous female of 16.6 mm from Stn 69 (MNHN Ga 3949) has been selected as the holotype,

the other specimens are paratypes.

ETYMOLOGY. — From the Latin, compactus , thick, in reference to the shape of the carapace.

Description. — Carapace with numerous secondary striae. Intestinal region with numerous scales.

Anterolateral spine well developed, situated at anterolateral angle. Branchial margins with 5 well developed

spines. Fourth thoracic sternite with a few short arcuate striae; fifth to seventh without striae. Abdominal

segments with numerous transverse striae; second segment with 9-11 spines on anterior ridge. Eye large,

maximum corneal diameter about 2/5 length of anterior border of carapace between bases of anterolateral spines of

carapace. Basal segment of antennule (distal spines excluded) ending at same level of cornea, distal spines

subequal. First segment of antennal peduncle with distomesial spine nearly reaching end of second segment;

distomesial spine of second segment exceeding third segment, usually with accompanying small spine proximally.

Extensor margin of merus of third maxilliped unarmed. Distomesial spine of cheliped merus well developed,

reaching midpoint of carpus. Fixed finger with one distal spine; movable finger with basal spine. First walking leg

Source :

606

E. MACPHERSON

about twice carapace length; dactylus as long as propodus, with spine-like setae reaching distal part of ventral

margin.

Remarks. — Munida compacta is closely related to M. curvirostris Henderson, 1885 (see below). Both species

have five spines on the lateral margins of the carapace behind the cervical groove, the front margins are transverse,

the eyes are large, the abdominal segments have spines along the anterior ridge of the second tergite, the lateral

portions of the posterior thoracic sternites have no granules, the distal spines on the basal antennular segment are

subequal and the extensor margin of the merus of the third maxilliped is unarmed. However, the species are easily

differentiable in several regards. The new species has numerous secondary striae and scales on the carapace and

abdominal segments (more than 5 secondary striae on the second abdominal segment). These striae and scales are

practically absent in M. curvirostris (less than 5 secondary striae on the second abdominal segment). Furthermore,

in M. compacta the dactylus of the walking legs is as long as the propodus, whereas in M. curvirostris the

dactylus is clearly shorter than the propodus.

The new species also resembles Munida rhodonia Macpherson, 1994, from New Caledonia, Loyalty and

Chesterfield Islands (Macpherson, 1994).

However, they can be distinguished by several differences:

— The chelipeds are clearly different. They are more massive in M. rhodonia than in the new species and

the spines on the dorsal side of the palm are very small in M. rhodonia , whereas they are well developed in the

new species. Moreover, in the new species, the distomesial spine of the merus of the cheliped reaches or

overreaches the midlength of the carpus; in M. rhodonia , this spine usually does not reach the midlength of this

article.

— In M. rhodonia the dactylus of the first walking leg is clearly shorter than the propodus. In the new species

the dactylus is as long as the propodus.

Distribution. — Indonesia, between 246 and 694 m.

Munida compressa Baba, 1988

Munida compressa Baba. 1988: 91, figs 33-34. — Macpherson, 1993: 427.

Material EXAMINED. — Indonesia. Karubar: stn 17, 439-459 m: 1 2 14.4 mm.

Distribution. — Japan, Southern China Sea, Philippines, Arafura and Molucca Seas in 180-668 m (Baba,

1988; Macpherson, 1994). The present specimen was collected between 439 and 459 m.

Munida curvirostris Henderson, 1885

Munida curvirostris Henderson, 1885: 412, — Macpherson, 1993: 428 (in part).

MATERIAL EXAMINED. — Indonesia. Karubar: stn 20, 769-809 m: 14 6 7.2 to 15.6 mm- 5 ov 2 13 5 to

18.7 mm; 20 2 8.3 to 14.8 mm. — Stn 21, 688-694 m: 4 6 5.6 to 18.3 mm; 2 2 13.8 and 14.1 mm.

Remarks. — A revision of M. curvirostris Henderson, 1885 is presently being prepared by Baba. Several

samples from MUSORSTOM 1 (Station 50), MUSORSTOM 2 (Station 75) and CORINDON (Stations 209 and 276)

cruises (see Macpherson, 1993) were incorrectly identified as M. curvirostris , belonging instead to M. compacta

(see above).

Distribution. — East coast of Africa, Arabian Sea, Maldives, Andaman Sea, Japan, Philippines, Indonesia

eastern Australia, between 141 and 1360 m (Baba, 1988; Macpherson, 1994). The present material was collected

from 688-809 m.

Source :

SPECIES OF THE GENERA AGONONIDA AND MUNIDA FROM THE KARUBAR EXPEDITION

607

Munida hyalina Macpherson, 1994

Munida hyalina Macpherson, 1994: 477, fig. 22.

MATERIAL EXAMINED. — Indonesia. Karubar: stn 18, 205-212 m:2i 3.6 and 4.1 mm.

Remarks. — The specimens from Indonesia arc similar to the type material. However, one specimen has

4 small spines on the branchial margins (3 spines in the types) and one transverse stria posterior to the median

ridge on the second abdominal segment (absent in the types).

Distribution. — New Caledonia and Chesterfield Islands, between 310 and 720 m (Macpherson, 1994) and

Indonesia, between 205 and 212.

Munida japonica Stimpson, 1858

Munida japonica Stimpson, 1858: 252. — Macpherson & Baba, 1993: 399, fig. 9 (references and synonymy).

Material EXAMINED. — Indonesia. Karubar: stn 32, 170-206 m: 5 6 3.2 to 9.5 mm; 1 ov. 2 9.6 mm;

3 2 4.3 to 4.5 mm. — Stn 49, 206-209 m: 4 6 4.3 to 14.2 mm; 1 2 6.5 mm.

DISTRIBUTION. — Munida japonica has been previously cited in Japan and the Philippines in 102-220 m

(Macpherson & Baba, 1993). The present material has been collected between 170 and 209 m.

Munida kuboi Yanagita, 1943

Munida kuboi Yanagita, 1943: 20, figs 5-6. — Baba. 1988: 109, fig. 40; 1990: 964. — Macpherson, 1993: 431.

Material EXAMINED. — Indonesia. Karubar: stn 79, 239-250 m: 2 6 1 1.0 and 1 1.8 mm; 1 2 10.8 mm. —

Stn 83, 285-297 m: 3 6 10.6 to 14.0 mm; 6 2 8.6 to 15.5 mm (POLIPI). — Stn 84, 246—250 m: 5 6 10.1 to

13.2 mm; 1 2 10.8 mm.

DISTRIBUTION. — South Africa, Madagascar, Japan, Philippines and Indonesia, between 78 and 405 m (Baba,

1988, 1990; MACPHERSON, 1994). The specimens examined here were collected between 239 and 297 m.

Munida leptitis Macpherson, 1994

Munida leptilis Macpherson, 1994: 487, fig. 27; 1995: 394, fig. 14.

Material EXAMINED. — Indonesia. Karubar: stn 2, 209-240 m: 3 6 4.8 to 5.3 mm; 2 2 3.6 and 4.3 mm;

1 juv. 2.4 mm. — Stn 5, 296-299 m: 1 6 4.0 mm. — Stn 14, 245-246 m: 3 6 4.7 to 5.1 mm; 7 2 4.0 to 5.1 mm. —

Stn 49, 206-210 m: 4 6 3.3 to 4.4 mm; 1 ov. 2 3.6 mm; 4 2 3.0 to 4.0 mm.

Remarks. — The specimens examined agree quite well with the type material. However, in some the movable

finger of the chelipeds has some spines along the mesial border (only one basal spine in the types). Furthermore,

the dactylus of the walking legs in the specimens from Stn 14 are unarmed on the tetminal third ot the ventral

margin (armed with movable spines along the whole border in the types). These differences suggest the existence

of several forms or species (see also MACPHERSON, 1995). Munida leptitis was described from only two

specimens collected in New Caledonia and Loyalty Islands, therefore the discovery of additional topotypic

specimens would be desirable in order to determine the variability of the species and to confirm the identity of the

present material.

Distribution. — New Caledonia and Loyalty Islands, between 21 and 440 m (Macpherson, 1994), Wallis

and Futuna area (MACPHERSON, 1995). The specimens from Indonesia were collected between 206 and 299 m.

Source : MNHN Paris

608

E. MACPHERSON

Munida leviantennata Baba, 1988

Munida leviantennata Baba, 1988: 1 11, figs 41. 42; 1994: 12, fig. 5. — MaCPHERSON, 1994: 491; 1995: 395.

Material EXAMINED. — Indonesia. Karubar: sin 25, 336-346 m: 1 8 11.5 mm. — Sin 35, 390-502 m: 2 <3

16.3 and 16.7 mm; I 2 14.8 mm (POLIPI). — Sin 69. 356-368 m: 1 <3 7.7 mm. — Stn 77, 346-352 m: 1 8 17.7 mm.

Distribution. — Philippines, Indonesia, eastern Australia, New Caledonia, Chesterfield Islands and Wallis

Island, between 300 and 660 m (Baba, 1988, 1994; MaCPHERSON, 1994). The present material was obtained

between 336 and 502 m.

Munida microps Alcock, 1894

Munida microps Alcock, 1894: 326. — Baba, 1988: 122; 1994: 13. — MaCPHERSON, 1994: 496, fig. 32; 1995: 397.

Material EXAMINED. — Indonesia. Karubar: stn 20, 769-809 m: 1 2 15.0 mm. — Stn 87, 1017-1024 nr

1 ov, 9 13.7 mm.

Distribution. — Previously known from Arabian Sea, Maldives Islands, Philippines, Indonesia, southeastern

Australia, New Caledonia, Chesterfield Islands, Wallis and Futuna Islands, between 970 and 1240 m

(MaCPHERSON, 1994). The present material was collected between 686 and 1024 m.

Munida minuta Macpherson, 1993

Munida minuta Macpherson, 1993: 432, fig 3.

Material EXAMINED. — Indonesia. Karubar: stn 22, 85-124 m: 2 8 3.6 and 4.0 mm.

DISTRIBUTION. — Philippines in 92-97 m (MACPHERSON, 1993). The specimens from Indonesia were obtained

in 85-124 m.

Munida philippinensis Macpherson & Baba, 1993

Munida philippinensis Macpherson & Baba, 1993: 410, fig. 16.

ooa ™aTER',A1; ^MINED. — Indonesia. Karubar: stn 2, 209-240 m: 1 <3 6.3 mm; 2 2 4.0 and 5.3 mm. — Stn 49

206- 09 m: 1 8 4.4 mm, 1 ov, 2 4.9 mm (USNM). - Stn 65, 174-176 m: 1 <3 5.0 mm. - Stn 67, 146-233 m: 2 <3 6 2

6 6 mm ?POLIP<I)' $ 8 mm' ~ S'" 83> 285’297 m: ' °V' 2 70 mm <P0LIP1>- — Stn 84, 246-275 m: 1 6

Distributor — Philippines, in 170-220 m (Macpherson & Baba, 1993). Indonesia, between 146 and

297 m.

Munida punctata sp. nov.

Fig. 3

(MN^Ga 3A952fXAMSl,nE2s' ~ ,I"^nes*a /A«UBAR- «n 9, 368-389 m: 3 <J 8.2 to 11.0 mm; 2 2 8.0 and 8.3 mm

MNHN-Ga 3950 • 7,3 in 5 I f ‘° ”m (MNHN-Ga 3953). - Stn 35, 390-502 m: 1 <3 12.7 mm

tMiNHiN ua M 50), 2 8 10.5 to 12.4 mm; 2 2 10.0 and 12.0 mm (MNHN-Ga 3951).

Types. - The male of 12.7 mm from Stn 35 (MNHN-Ga 3950) has been selected as holotype, the other

specimens are paratypes.

Source :

SPECIES OF THE GENERA ACONONIDA AND MUNIDA FROM THE KARUBAR EXPEDITION

609

Fig. 3. — Munida punctata sp. nov., 6 12.7 mm. holotype from Stn 35: a. carapace and abdomen, dorsal view;

b. rostrum, lateral view; c. sternal plastron; d. ventral view of cephalic region, showing antennular and antennal

peduncles; e, right third maxilliped. proximal segments of endopod, lateral view; f. right cheliped. dorsal view;

g, right first walking leg, lateral view; h. dactylus of right first walking leg, lateral view.

ETYMOLOGY. — From the Latin, puncta , point, puncture, in reference to the red point near the rostrum lip.

DESCRIPTION. — Carapace with numerous secondary striae. Intestinal region with some median scales.

Rostrum spiniform, not laterally compressed. Anterolateral spine moderately short, situated at anterolateral angle.

Source : MNHN, Paris

610

E. MACPHERSON

not reaching level of sinus between rostrum and supraocular spine. Branchial margins with 5 small spines.

Thoracic sternites with numerous short arcuate striae. Second abdominal segment with 8-9 spines along anterior

transverse ridge. Second and third segments each with some transverse striae. Males with two gonopods on first

and second abdominal segments. Eye large, maximum corneal diameter about 1/2 length of anterior border of

carapace between bases of external orbital spines. Basal segment of antennule (distal spines excluded) ending at

level of cornea, distal spines subequal. First segment of antennal peduncle with short distomesial spine, clearly'not

reaching end of second segment; distomesial spine of second segment long, barely exceeding antennal peduncle.

Extensor margin of merus of third maxilliped unarmed. Distomesial spine of cheliped merus well developed,

though not reaching midpoint of carpus. Movable finger of chelipeds with basal spine, fixed finger with terminal

spine. First walking leg about twice carapace length. Dactyli of walking legs slightly shorter or as long as

propodus, with spinules along entire ventral margin.

Remarks. — The new species resembles Munida rubrodigitalis Baba. 1994, from northeastern Australia, New

Caledonia, Loyalty Islands and Indonesia (Baba, 1994; Macpherson, 1994, see also below). Both species have

five spines on the branchial margin, the second abdominal segment armed with spines along the anterior ridge, the

distal spines on the basal antennular segment subequal, short chelipeds and a red, distal mark on the rostrum

(specimens in preservative). However, they can be easily distinguished by the shape of the rostrum, which is

laterally compressed in M. rubrodigitalis, whereas in the new species it is clearly spiniform.

Distribution. — Indonesia, between 336 and 502 m.

Munida rubrodigitalis Baba, 1994

Munida rubrodigitalis Baba, 1994: 13, fig. 6.

Munida sp. — MACPHERSON, 1994: 558, figs 13b, 90.

Material EXAMINED. — Indonesia. Karubar: stn 25, 336-346 m:

285-297 m: 1 6 13.3 mm; 2 2 11.6 and 12,5 mm.

2 9.0 mm (MNHN-Ga 3660). — Stn 83,

Distribution. — Eastern Australia, between 497 and 503 m (Baba, 1994), New Caledonia and Loyalty

Islands in 466-650 m (Macpherson, 1994). Indonesia, between 285 and 346 m.

Munida rufiantennulata Baba, 1969

Munida rufiantennulata Baba, 1969: 23, fig. 7; 1988: 128; 1989: 131. - Macpherson, 1994: 523. figs. 46, 83.

Material examined. — Indonesia. Karubar: stn 18, 205-212 m: 3 6 4.3 to 9.4 mm; 7 2 3.0 to 7.7 mm.

DISTR'BUTION - Japan, Philippines. Indonesia, New Caledonia, Loyalty Islands, Mattew and Hunter Islands

and Chesterfield Islands, between 379 and 610 m (Baba, 1969, 1988, 1989; Macpherson, 1994) The material

examined here was collected in 205-212 m.

Munida striola Macpherson & Baba, 1993

Munida striola Macpherson & Baba, 1993: 416, fig. 20.

Material EXAMINED. — Indonesia. Karubar: stn 6. 287-298. I ov. 2 10 3 mm — Stn 9 368 389 m I ^

ZgSK ? UJ mm- - s" » 3»7-3‘"“ * ”.a3 1. Val

Source :

SPECIES OF THE GENERA AGONONIDA AND MUNIDA FROM THE KARUBAR EXPEDITION

61 I

15.4 mm; 4 ov. 2 12.0 lo 17.0 mm; 4 2 7.7 to 13.6 mm. — Stn 84. 246-275 m: 5 6 12.4 to 17.5 mm; 1 ov. 2

16.5 mm; 3 2 12.1 to 14.0 mm. — Stn 85, 240-245 m:2i 11.6 and 15.0 mm; 1 ov. 2 13.5 mm.

Distribution. — M. striola has been cited in Japan, Philippines and Indonesia in 215-300 m (Macpherson

& Baba, 1993). The material examined here was collected between 146 and 389 m.

Munida sp.

MATERIAL EXAMINED. — Indonesia. Karubar: stn 1, 156-305 m: 3 ov. 2 7.1 to 8.0 mm. — Stn 2. 209-

240 m : 3 8 8.5 to 10.7 mm. — Stn 5. 296-299 m: 7 <5 10.3 to 10.6 mm; 1 ov. 2 9.2 mm; 3 2 8.3 to 8.5 mm. —

Stn 6. 289-298 m: 1 1 6 10.5 to 12.0 mm; 4 ov. 2 8.3 to 10.3 mm; 3 2 10.3 to 1 1.5 mm.

Remarks. — This species is actually being studied by K. Baba (Kumamoto University, Japan) and probably

it belongs to a new genus (K. Baba, personnal communication).

Distribution. — Indonesia, between 156 and 305 m.

ACKNOWLEDGEMENTS

I am very grateful to A. CROSNIER of ORSTOM and M. K. Moosa from the Puslitbang Oseanologi-LIPI for

their support and help and for making this interesting material available to me. Thanks are also due to K. Baba

(Kumamoto University) for his comments, suggestions and improvements of ihe manuscript.

REFERENCES

Baba, K., 1969. — Four new genera with their representatives and six new species of the Galatheidae in the collection of

the Zoological Laboratory, Kyushu University, with redefinition of the genus Galatliea. Ohmu (Occasional Papers

from ilie Zoological Laboratory, Faculty of Agriculture, Kyushu University), 2 (1): 1-32.

Baba, K., 1986. — Two new anomuran Crustacea (Decapoda: Anomura) from North-West Australia. The Beagle, 3 (1):

1-5.

Baba. K., 1988. — Chirostylid and Galatheid Crustaceans (Decapoda; Anomura) of the "Albatross" Philippine

Expedition, 1907-1910. Researches on Crustacea, Special Number 2; v + 203 pp.

Baba, K., 1989. — Anomuran Crustacean obtained by dredging from Oshima Strait, Amami-Oshima of the Ryukyu

Islands. Memoirs of the National Science Museum, Tokyo, 22: 127-134.

Baba, K., 1990. — Chirostylid and Galatheid Crustaceans of Madagascar (Decapoda, Anomura). Bulletin du Museum

National d'Histoire Naturelle, ser. 4, 11, sect. A, (4): 921-975.

Baba, K.. 1994. — Deep-sea Galatheid crustaceans (Anomura: Galatheidae) collected by the 'Cidaris 1' expedition off

central Queensland, Australia. Memoirs of the Queensland Museum, 35: 1-21.

Baba. K.. & de Saint Laurent, M., 1996. — Crustacea Decapoda: Revision of the genus Bathymunida Balss, 1914, and

description of six new related genera (Galatheidae). In: A. Crosnier (ed.), R6sultats des Campagnes Musorstom,

Vol. 15. Memoires du Museum National d'Histoire Naturelle, 168: 433-502.

Crosnier, A., Richer de Forges, B. & BOUCHET, P., 1997. — La campagne Karubar en Indon6sie. au large des Ties Kai

etTanimbar. In: A. Crosnier & P. Bouchet (eds), Resultats des Campagnes Musorstom, Vol. 16. Memoires du

Museum National d'Histoire Naturelle. 172: 9-26.

Henderson, J. R„ 1885. — Diagnoses of the new species of Galatheidea collected during the "Challenger" Expedition.

Annals and Magazine of Natural History, ser. 5, 16: 407-421.

HENDERSON, j. R., 1888. — Report on the Anomura Collected by H.M.S. Challenger During the Years 1873-76.

Challenger Reports, Zoology, 27: vi + 221 pp., 21 pis.

Source : MNHN, Paris

612

E. MACPHERSON

Macpherson, E.. 1993. — Crustacea Decapoda: Species of the genus Munida Leach, 1820 (Galatheidae) collected during

the Musorstom and Corindon cruises in the Philippines and Indonesia, In: A. Crosnier (ed.). Resultats des

Campagnes Musorstom, Vol. 10. Memoires du Museum National d’Histoire Naturelle, 156: 421-442.

Macpherson, E., 1994. — Crustacea Decapoda: Studies on the genus Munida Leach. 1820 (Galatheidae) in

New Caledonian and adjacents waters with descriptions of 56 new species. In: A. Crosnier (ed.). Resultats des

Campagnes MUSORSTOM, Vol. 12. Memoires du Museum National d’Histoire Naturelle, 161: 421-569.

Macpherson, E„ 1996. — Crustacea Decapoda: Species of the genus Munida Leach, 1820 and Paramunida Baba, 1988

(Galatheidae) from Wallis and Futuna. In: A. Crosnier (ed.). Rdsultats des Campagnes Musorstom, Vol. 15.

Memoires du Museum National d’Histoire Naturelle, 168: 387-421.

Macpherson, E.. & Baba. K.. 1993. — Crustacea Decapoda: Munida japonica Stimpson, 1858, and related species

(Galatheidae). In: A. Crosnier (ed.), Resultats des Campagnes Musorstom, Vol. 10. Memoires du Museum National

d'Histoire Naturelle, 156: 381-420.

Macpherson. E.. & de Saint Laurent, M.. 1991. — Galatheid crustaceans of the genus Munida Leach, 1818, from

French Polynesia. Bulletin du Museum National d'Histoire Naturelle, ser. 4, section A, Zoologie, 13 (3-4): 373-422.

Source :

;SULTATS DES CAMPAGNES MUSORSTOM. VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 16 — RESULTATS

Crustacea Decapoda: Ethusinae (Dorippidae),

mainly from the KARUBAR Cruise

CHEN Huilian (H. L. CHEN)

Institute of Oceanology, Academia Sinica,

7 Nan-Hai Road, Qinngdao 266071, China

ABSTRACT

Material of Ethusinae collected by a French-lndonesian expedition in Indonesia (KARUBAR, 1991), and two French

expeditions to Wallis and Futuna Islands (MUSORSTOM 7. 1992), and off New Caledonia (Bathus 3. 1993) yielded a total

of 1 1 species belonging to three genera. One genus and five species are new and three species are recorded lor the first

time from Indonesia.

RESUME

Crustacea Decapoda : Ethusinae (Dorippidae) recoltes principalement lors de la campagne

KARUBAR.

Les Ethusinae recoltees par une expedition t'ranco-indonesienne en Indon6sie (Karubar, 1991) et deux autres

expeditions frangaises aux ties Wallis et Futuna (Musorstom 7, 1992) et en Nouvelle-Caledome (Bathus 3. 1993),

comprennent 1 1 espfeces appartenant h 3 genres. Un genre et cinq espfices sont nouveaux pour la science et trois espdees

n’avaient jamais 6t6 encore signalees en Indonesie.

INTRODUCTION

This report is based on the collections taken during a Franco-Indonesian expedition in Indonesia (Karubar,

1991), and two French expeditions at Wallis and Futuna Islands (MUSORSTOM 7, 1992) and off New Caledonia

(Bathus 3, 1993).

Eleven species, belonging to 3 genera, have been identified, of which one genus and 5 species are described as

new, and 3 species (marked with an asterisk in the accompanying list) are reported for the first time from

Indonesia.

Chen Huilian (H. L. Chen), 1997. — Crustacea Decapoda: Ethusinae (Dorippidae), mainly from the Karubar Cruise.

In. A. CROSN1ER & P. Bouchet (eds), Resultats des Campagnes MUSORSTOM. Volume 16. Mem. Mus. nain. Hist. nat..

172: 613-625. Paris ISBN: 2-85653-506-2.

Contribution no. 2557, Institute of Oceanology, Academia Sinica, Qingdao, China.

Source : MNHN, Paris

614

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

LIST OF SPECIES

Subfamily ETHUSINAE Guinot, 1977

Parethusa glabra gen. et sp. nov.

Ethusa dilalidens sp. nov.

Ethusa indica Alcock, 1 894

Ethusa indonesiensis sp. nov.

Etlmsa longidentata sp. nov.

Ethusa sexdentata (Stimpson. 1 858)

Ethusa bicornuta sp. nov.

* Ethusa brevidentata Chen, 1993

Ethusa desciscens Alcock, 1 896

* Ethusa investigatoris Alcock, 1 896

* Ethusa pubescens Chen, 1993

SYSTEMATIC ACCOUNT

Subfamily ETHUSINAE Guinol, 1977

Genus PARETHUSA nov.

DIAGNOSIS. — Carapace longer than broad, swollen, dorsal and ventral surfaces smooth. Grooves and regions

very indistinct. Front thin, separated into 2 broadly triangular teeth (the external side of the two median frontal;

teeth is angled, suggestive of the beginning of a tooth) by a large, V-shaped notch. Basal segment of antennules

not swollen, eyes located on ventral surface, eye-stalks short, stout and movable. Last two legs short and small,

dactyli talon shaped.

TYPE species. — Parethusa glabra sp. nov. by present designation.

Gender. — Feminine.

ETYMOLOGY. — The name is formed by a combination of the Greek word par (near) and the feminine name

Ethusa.

Remarks. — This new genus is closely related to Ethusa and Ethusina. but may be distinguished from them

by the following:

The new genus is characterized by the presence of talon-shaped dactyli on the last two legs, the long and slender

second pleopods and the teeth of the front (2 broadly triangular ones). I am inclined to consider Parethusa as a

primitive member of the Ethusinae.

Parethusa glabra sp. nov.

Figs 1-2

Material

27.10.1991: 1 6

EXAMINED. — Indonesia. Kai Islands. Karubar: st. CP 35. 06°08'S,

holotype 8.3 x 8.0 mm (MNHN-B 22886).

I 32°45'E. 390-502 m.

Source

ETHUS1NAE MAINLY FROM THE KARUBAR EXPEDITION

615

Description. — Carapace inflated, slightly longer than broad. Dorsal and ventral surfaces smooth. Grooves

and regions very indistinct, only urogastric and cardiac-intestinal regions slightly defined. Front thin, separated into

two broadly-triangular teeth by a large V-shaped notch. Orbit very small, with a small V-shaped notch between

frontal and exorbital teeth, dorsal surface with a broad, shallow groove. Exorbital teeth blunt, short, with outer

border converging inwardly. Base of antennules not swollen. Eyes located on ventral surface, eyestalks short, stout

and movable.

Fig. 1. — Parethusa glabra gen. et sp. nov., <J holotype (MNHN-B 22886): a. carapace; b, anterior portion of carapace

(ventral view); c, male abdomen. Scales = 1 mm.

Chelipeds nearly symmetrical, surface smooth. Merus 4.5 times longer than high, palm twice as long as high.

Fingers as long as propodus, cutting edges without teeth.

Second and third pereiopods long and smooth, third longest and fourth shortest of all pereiopods. Merus ol third

7.5 times longer than high and propodus 5 times as long as high. Dactylus longer than propodus. Fourth

pereiopods short and stout, 3 times longer than high, carpus longer than propodus, propodus 2.5 times longer than

high, distal part with some spines.

Male abdomen moderately broad, consisting of five segments (3rd-5th fused): first stout, 3 times as long as

second. Base of fused segments strongly convex on each side, depressed in middle. Sixth segment 2.7 times longer

than broad, anterior border depressed. Telson bluntly triangular.

Male first pleopods stout, basal half twice as broad as distal half, with a small notch distally. Second pleopods

longer than first, distal part curved and thin.

Genus ETHUSA Roux, 1830

Ethusa dilatidens sp. nov.

Fig. 3

MATERIAL EXAMINED. — Indonesia. Tanimbar Islands. Karubar: st. CP 83, 09°23’S. 131°00E, 285-297 m,

04.11.1991: 1 S holotype 12.0 x 12.2 mm (MNHN-B 22887).

Source : MNHN, Paris

616

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

FIG. 2. — Parethusa glabra gen. et sp. nov., 6 holotype (MNHN-B 22886): a, cheliped; b. third pereiopod; c. fourth

pereiopod; d. fifth pereiopod; e-g. male first pereiopod; h-i, second male pleopod. Scales: a-f, h = 1 mm; e i =

0. 1 mm.

Source : MNHN, Paris

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

617

FIG. 3. - Ethusa dilatidens sp. nov., <J holotype (MNHN-B 22887): a. carapace; b. cheliped; c second pereiopod;

d. third pereiopod; e. fourth pereiopod; f. male abdomen; g-h. male first pleopod; l-j. male second pleopod.

Scales : a-g = 1 mm. h. j = 0.1 mm.

Source : MNHN, Paris

618

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

Description. — Carapace slightly broader than long, dorsal surface with very fine granules and short

pubescence. Cervical and branchial grooves and regions distinct. Branchial regions strongly convex, mesogastric

and cardiac-intestinal regions slightly convex. Front divided into 4 short teeth by one V-shaped and 2 U-shaped

notches. Exorbital teeih broad at base, slightly broader than long, outer borders converging inwards.

Chelipeds symmetrical, palm twice as long as high and slightly longer than fingers, cutting edges of fingers

without teeth.

Second and third pereiopods smooth and hairless. Third pereiopod (left only) the longest: merus 5.6 times

longer than high and propodus 4 times longer than high ; merus of second pereiopod only 4.5 times and propodus

only 3.2 limes longer than high. Right fourth pereiopod short, with fine granules and short hairs near the tip.

Carpus as long as propodus.

Male abdomen consisting of 5 segments (3rd-5th fused): first 1.5 times as long as second. Sixth segment 1.15

times longer than broad. Telson bluntly triangular, broader than long.

Male first plcopods moderately stout, basal half thicker than distal, gradually narrowed distally, slightly

produced at distal 1/7, and both borders of distal end with some spines. Male second pleopods slender, tip sharp.

Etymology. — The name is formed by a combination of the Latin dilatus (expanded) and dens (tooth), in

reference to the shape of the exorbital teeth.

Remarks. — The new species is similar to Etlwsa sexdentata (Stimpson, 1858) in ihe shape of the carapace,

but can easily be distinguished from it by the exorbital teeth being broader than long, its outer border converging

inwards, the symmetrical male chelipeds, and the form of ihe male first pleopod. Ethusa dilatidens is also similar

10 E- obliquidens Chen, 1993, bul differs from the latter in having the exorbital teeth longer than broad ; the telson

of male abdomen broadly triangular or semi-circular ; the first segment 1.5 times longer than the second; and the

distal end of first pleopods blunty rounded.

Ethusa indica Alcock, 1894

Ethusa indica Alcock, 1894: 405 ; 1896: 283. — Alcock & Anderson, 1895, pi. 14, fig. 2. — Chen 1993: 324

Nagai, 1995: 60, pi. 1, fig. 6.

MATERIAL EXAMINED. — Indonesia. Kai Islands. Karubar: si. CP 12,05°23'S. 132°37'E. 436-413 m

mo so\"1; It™,* I2'? To ; \ 9 90 X 90 ‘ 130 x 129 mm <MNHN-B 22867). — St. CP 35, 06°08'S, 132°45'e!

390-502 m, 27.10.1991: 1 d 5.8 x 5.6 mm (MNHN-B 22872)

KT* U B A R : st- CP 38- 07°40'S. 132°27'E, 620-666 m, 28.10.1991: 1 d 5.8 x 5.8 mm

™ ‘sfrp s7 n8°?rQ53i avc' * °4 ‘ l026‘1053 m- 30. 10.1991: 1 ovig. 9 8.6 x 8.7 mm (MNHN-B

ifn nnS CL5Mu!,8D2 Ji 3 43 E- 836’869 m- 30.10.1991: 1 d 8.4 x 8.3 mm. 10 9 (2 ovig.) 10.5 x 10 8 -

12 0 x 12.0mm (MNHN-B 22862). — St. CC 56. 08°16'S, 131°59E. 552-549 m, 31 10 1991 2 d50x49-59x

?^Tn(McNnHN B 22864)' - St CC 57‘ 08°19'S x 13I°53'E- 603-620 m. 31.10.1991: 4 3 5 5 x 54 6 5 x 64 mm

d 8.0 x 8.0 mnJ (MNHN-B 22857). - St. CP 59. 08°20'S. 132°H'E. 399-405 m. 31.10.1991: 7 9 (3 ovig.) 74 x 7 7

109 x lYmmfpni i c ™ 66)' ~ St- CP 73' 08°29'S’ 131°33'E, 840-855 m, 02.11.1991: 1 ovig. 9

°79 x 1 T;0 mm (P0LIP|)- — st. CP 75, 08 46'S, 131°36'E, 400 m, 03.11.1991: 2 d 60 x 6 3 -60 x 65 mm 7 o in

9 7 0 x 7 7 Tpti t N ' B 2 2 8 3 8 ’ • - St' CP 76, 08°50'S. 131°33 E. 400-401 m, 03.1 1.1991: 1 d 6.3 x 6.3 mm 1

9 7.0 x 7.2 mm (POLIPI). — St. CP 87, 08°47’S, 130°49'E. 1017-1024 m 05 1 1 1991- 29 83x84 mm 8SvSS

mm (MNHN-B 22859) - St. CP 89. 08°39'S, 13U08 E. ,048-1084 m. 05 .1,991: 4 d 7.8 x 7 4 9 0 Ts 9 mm 6 9

7 40xV7^ 7 5Xx 7 3' ’jr8%ll8Oni8nMmHrB1n27860>' ~ S‘' CP 91' °8°44'S' 131°05'E- 884-890 m, 05.1 1.1991: 2 d

/.4 X I.Z /.3 X 7.3 mm, 8 9 8.0 x 8.0 - 10.3 x 10.3 mm (MNHN-B 22863).

Distribution. — This species is very common

and Sri Lanka eastward to the Philippines, China and

in the Indo-Pacific Region, ranging from the Maidive Islands

Japan, and southward to Indonesia and New Caledonia.

Source :

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

619

Elhusa indonesiensis sp. nov.

Fig. 4

MATERIAL EXAMINED. — Indonesia. Kai Islands. KaRUBar: st. DW 18. 05°18'S. 133°01'E. 205-212 m,

24.10.1991: 1 6 hololype 7.5 x 7.5 mm (MNHN-B 22888).

DESCRIPTION. — Carapace as long as broad, dorsal surface covered with very fine granules and short

pubescence. Grooves shallow and broad. Regions distinct: protogastric, metabranchial and cardiac regions strongly

convex, mesogastric, metagastric, urogastric and intestinal regions slightly convex. Posterior part of orbit and mid¬

line of front depressed. Front divided into 4 subequal teeth. Exorbital teeth short and broad at base, distal ends

sharp.

Male chelipeds very unequal: merus of larger cheliped 3 times longer than high, palm swollen, longer than

high. Fingers shorter than palm, cutting edges without teeth ; merus of smaller cheliped 4 times as long as high,

palm twice as long as high. Fingers longer than palm, cutting edges without teeth.

Third pereiopods longest, meri 7 times longer than high. Propodi slightly shorter than dactyli and 5.5 times

longer than high. Dactyli claw-shaped.

Male abdomen consisting of five segments (3rd-5th segments fused): first as long as second and sixth as long

as telson. Telson blunty triangular.

Male first pleopods stout, basal 4/5 stouter, distal part tapered and with some spines. Second pleopods longer

than first, distal half lamelliform.

Etymology. — This species is named after the country where it has been found, Indonesia.

REMARKS. — This species is similar to Elhusa paragymaea Chen, 1993 but differs from the latter in having

the carapace slightly longer, the male first pleopods with a foot-shaped tip, and the palm of the smaller cheliped as

long as the fingers.

Elhusa longidentata sp. nov.

Fig. 5

MATERIAL EXAMINED. — Indonesia. Kai Islands. KARUBAR: st. DW 28. 05°31'S, 132°54'E, 448-467 m.

26.10.1991: 1 6 holotype 6.4 x 6.0 mm (MNHN-B 22884); 1 9 paratype 7.0 x 7.0 mm (MNHN-B 22889).

DESCRIPTION. — Carapace longer than broad in male, as long as broad in temale, dorsal surface covered with

fine granules, borders of exorbital and frontal teeth with sharp granules. Grooves and regions distinct, posterior part

(behind frontal and orbital regions) concave. Protogastric and mesogastric regions slightly convex, metagastric and

urogastric regions depressed. Cardiac-intestinal and branchial regions convex. Front divided into 4 teeth by one

V-shaped and 2 U-shaped notches. Exorbital teeth long and acute, stouter in male than in female.

Merus, carpus and palm of male cheliped (right only) covered wilh fine granules. Palm longer than high and

slightly shorter than fingers. Fingers smooth, cutting edges without teeth.

Pereiopods covered with fine granules. Third pleopod longest, merus of medium length, 4 times (right), 5

times (left) as long as high. Carpus long. Propodus 3.8 times as long as high. Dactylus longer than propodus.

Merus of P2 4 times and palm 3 times longer than high. Last two legs slender and short.

Male abdomen with fine granules, consisting of five segments (3rd-5th fused): first segment stout, 3 times as

long as second. Fused segments trapezoid, base swollen. Sixth segment rectangular. Telson bluntly rounded.

Male first pleopods stout, slightly curved, twisted and gradually narrowed from base to distal end: distal part

with spines. Second pleopods slender, distal end thin and sharp.

ETYMOLOGY. — The name is formed by a combination of the Latin longus (long) and dentata (toothed), in

reference to the shape of the exorbital tooth.

Source : MNHN, Paris

620

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

Fig. 4. — Ethusa indonesiensis sp. nov. 6 holotype (MNHN-B 22888): a, carapace; b. larger cheliped; c. smaller

cheliped; d, ihird pereiopod; e, fifth pereiopod; f, male abdomen; g-i, first pleopod; j, second pleopod. Scales: a-g,

j = 1 mm; i = 0.1 mm.

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

621

FIG. 5. — Ethusa longidentata sp. nov. a-1: 6 holotype (MNHN-B 22884): a. carapace (damaged): b. cheliped; c. third

pereiopod; d. fifth pereiopod; e, abdomen; f-i, first pereiopod; j-1. second pereiopod. — m: 5 paratype (MNHN-B

22889), anterior portion of carapace. Scales: a-h, j, m = 1 mm; i, k-1 = 0.1 mm.

Source : MNHN Paris

622

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

REMARKS. — This new species is similar to Ethusa makasarica Chen. 1993 but they can be distinguished as

follows:

Ethusa sexdentala (Stimpson, 1858)

Dorripe sexdentata Stimpson, 1858: 163.

Ethusa sexdentata - Stimpson, 1907: 168, pi. 19, fig. 4. — Chen, 1993: 335, fig. 14. — Nagai. 1995: 59. pi. I,

fig. 4.

Material EXAMINED. — Indonesia. Kai Islands. Karubar: st. CP. 25, 05o30’S, 132°52'E, 336-346 m

26.10.1991: I 6 13.4 x 13.0 mm (POLIP1).

Tanimbar Islands. Karubar: st. CP. 79, 09°16'S, 131°22'E, 239-250 m, 03.11.1991: 2 9 8.0 x 8 I mm 95 x

9.5 mm. ovig. (MNHN-B 22880).

Distribution. — Japan, China, Philippines, Indonesia, New Caledonia, Andaman Sea and Nicobar

Islands.

Genus ETHUSINA Smith, 1884

Ethusina bicornuta sp. nov.

Fig. 6

Material EXAMINED. — Indonesia. Tanimbar Islands. Karubar: st. CP 87, 08°47'S. 130°49'E 1017-KP4 m

05.11.1991: 1 6 holotype 7.6 x 7.0 mm (MNHN-B 22885).

Description. — Carapace covered with fine sparse granules and pubescence, longer than broad. Grooves and

regions distinct: protogastric, mesogastric and cardiac-intestinal regions slightly convex, but lower than

metabranchial regions. Front swollen, divided into 4 teeth by 3 notches: median frontal teeth short, half as long as

lateral teeth. Exorbital teeth needle-like, only reaching to base of lateral frontal teeth. Lateral borders of carapace

arched. Posterior border slightly convex.

Chelipeds slightly unequal, right somewhat larger than left: right palm 1.43 times (left 1.54 times) as long as

high. Both palms as long as fingers. Right movable finger with blunt, obscure teeth and a large tooth near base ;

immovable finger with blunt teeth ; cutting edges of smaller cheliped with blunt teeth.

Second and third pereiopods smooth and hairless. Third pereiopod longest: merus 9 times as long as high and

propodus 7.5 times as long as high, dactylus very long. Last two pereiopods slender and short, with short, sparse

hairs.

Male abdomen consisting of five segments (3rd-5th fused): first two segments subequal. Fused segment

depressed at middle of base. Sixth segment rectangular. Telson semicircular.

Male first pleopods stout, basal 2/3 swollen, twice as broad as distal 1/3, distal end divided into two horns by a

V-shaped notch.

Etymology. — The name is formed by a combination of the latin bis (two) and commits (horned) in

reference to the distal end of the first pleopods.

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

623

Fig. 6. — Ethusina bicornuta sp. nov„ 8 holotype (MNHN-B 22885): a, carapace; b. anterior portion of carapace

(ventral); c, larger cheliped; d, smaller cheliped; e, fifth pereiopod; f. abdomen; g-h, lirst pleopod; i-j, second

pleopod. Scales: a-g, i = 1 mm; h, j = 1.1 mm.

Source : MNHN, Paris

624

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

Remarks. — This new species closely resembles Ethusina investigatoris Alcock, 1895. but differs from the

latter in having shorter exorbital and lateral frontal teeth, the first two segments of male abdomen subequal, and the

distal end of the first pleopod with two horns.

Ethusina brevidentata Chen, 1993

Ethusina brevidentata Chen. 1993: 337. fig. 16.

Material EXAMINED. — Indonesia. Kai Islands. Karubar: st. CC 21. 05°14'S, 133°00’E. 688-694 m

25.10.1991: I 9 8.5 x 8.7 (MNHN-13 22882).

DISTRIBUTION. — Known only from New Caledonia and Indonesia.

Ethusina desciscens Alcock, 1896

Ethusina desciscens Alcock. 1896: 286. — Alcock & McArdle, 1903: 62, fig. 2,2a. — Chen, 1993: 337.

Material EXAMINED. — Indonesia. Tanimbar Islands. Karubar: st. CP 53, 08°18'S, 131°41'E, 1026-1053 m.

30.10.1991: 1 ovig. 9 8.9 x 9.3 mm (MNHN-B 22876). — St. CP 87. 08°47'S, I30°49’E. 1017-1024 m, 05 1! 199L

1 9 8.4 x 8.5 mm (MNHN-B 22875). — St. CP 89. 08°39'S. I31°08'E, 1058-1084 m, 05.1 1.1991: 3 ovig. 9 9.0 x

9.6 mm. 9.1 x 9.7 mm, 9.9 x 10.1 mm (MNHN-B 22877). — St. CP 91. 08°44'S. 13r05'E. 884-891 m. 05 II 1991

2 9 9.0 x 9.2, 9.2 x 9.4 mm (ovig.) (POLIPI).

DISTRIBUTION. — China, the Philippines. Indonesia. Andaman Sea. Laccadive Sea and Madagascar.

Ethusina investigatoris Alcock, 1896

Ethusina investigatoris Alcock. 1896: 285. — Alcock & McArdle. 1903: fig. 3,3a. — Chen, 1986: 135. fig. 14.

Material EXAMINED, — Indonesia. Tanimbar Islands. Karubar: st. CP 87, 08°47'S, I30°49'E 1017-1024 m

05.11.1991: 2 6 7.4 x 7.2, 7.9 x 7.6 mm (MNHN-B 22883).

Distribution. — East China Sea. Bay of Bengal and Laccadive Sea, at depths of 1 1 15 to 2378 m.

Ethusina pubescens Chen, 1993

Ethusina pubscens Chen, 1993: 341, fig. 19.

„„„^ATERIAL EXAMINED- — Wallis and Futuna Islands. MuSORSTOM 7: st. DW 565. 1 1°47'04"S 178°25'03''W

900 m, 20.05.1992: 1 8 7.6 x 7.5 mm (MNHN-B 22879)

-28787 Ca'ed0nia BATHUS 3: St' CP 842’ 23°05'S' l66°48'E- 830 m. 01.12.1993: 1 <J 10.9 x 11.0 mm (MNHN-B

Distribution. — New Caledonia, Wallis and Futuna Islands.

ACKNOWLEDGEMENTS

l am very grateful to Alain Crosnier and Bertrand Richer DE FORGES (ORSTOM) for providing the material

or this study; to Prof. J. Y. Liu (Institute of Oceanology. Academia Sinica. Qingdao) for reading the manuscript ;

to Mrs Liang (Youp.ng Marine Product Museum, Qingdao) for drawing some of the figures ; and to

R.B. Manning (Natural H.story Museum, Washington) and Peter Ng (Singapore University) for reviewing the

Source :

ETHUSINAE MAINLY FROM THE KARUBAR EXPEDITION

625

REFERENCES

Alcock, a., 1894, — Natural history notes from H.M. Indian marine survey steamer "Investigator", Series 2, No. 1. On

the results of deep-sea dredging during the season of 1890-1891, Annals and Magazine of Natural History, ser. 6, 13:

225-245, 321-334. 400-411.

Alcock, A„ 1896. — Materials for a carcinological fauna of India. No. 2. The Brachyura Oxystomata. Journal of the

Asiatic Society of Bengal, 65 (2): 134-296, pis 6-8.

Alcock, A. & Anderson, A.S.R., 1895. — Crustacea Part III. In: Illustrations of the zoology of the Royal Indian marine

surveying steamer "Investigator", pis 9-15. Calcutta.

ALCOCK, A. & McARDLE, A.F., 1903. — Crustacea Part X. In: Illustrations of the zoology of the Royal Indian marine

surveying steamer "Investigator", pis 56-67. Calcutta.

CHEN Huilian, 1986. — Studies on the Dorripidae (Crustacea Brachyura) of Chinese waters. Transactions of the Chinese

Crustacean Society, (1): 118-139, figs 1-15. [In Chinese with English summary]

Chen Huilian & Xu Zhenxiong, 1991. — Studies on the crabs of the Nansha Islands, China. In: Contributions on marine

biological research of the Nansha Islands and the neighbouring waters, volume 3: 48-106, 36 figs.

Chen Huilian, 1993. — Crustacea Decapoda: Dorippidae of New Caledonia, Indonesia and the Philippines. In:

A. CROSNIER (ed.), R<§sultats des Campagnes MUSORSTOM, Volume 10. Memoires du Museum National d'Histoire

Naturelle. 156: 315-345, figs 1-20.

Nagai, S., 1995. — Some remarkable crabs of Wakayama prefecture IV. Nanki Seibutu, 37 (1): 58-64, 1 pi. [in Japanese].

Stimpson, W., 1858. — Prodromus descriptionis animalium evertebratorum, quae in expeditione ad oceanum Pacificum

septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers ducibus, observavit et

descripsit W. Stimpson. Pars VI. [Preprint from] Proceedings of the Academy of Natural Sciences of Philadelphia, 10:

159-163 [57-61],

Stimpson, W„ 1907. — Report on the Crustacea (Brachyura and Anomura) collected by the North Pacific Exploring

Expedition, 1853-1856. Smithsonian Miscellaneous Collections, 49: 1-240, pis 1-26.

Source : MNHN, Paris

;SULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 16 — RESULTATS

Echinodermata Crinoidea : Les Pentacrines recoltees lors

de la campagne KARUBAR en Indonesie

Nadia AMEZIANE

Laboratoire de Biologie des lnvertebres Marins el Malacologie.

Museum national d'Histoire naturelle

55 rue Buffon, 75005 Paris.

RESUME

Les crinoides pedoncules recueillis par la campagne Karubar (lies Kai et Taninibar : Indonesie) sont des Pentacrines

(Pentacrinidae) appartenant au genre Saracrinus. Trois espbces dont une nouvelle ( S . moosai) sont decrites. Celle taible

diversite detonne avec l'abondante richesse mise en evidence par les auteurs du debut du siecle dans les eaux indonesiennes.

Cette faune setage entre 210 et 430 m de profondeur. Le materiel important (165 specimens) permel, entre autres, de preciser

le champ de variation des principaux caracteres morphologiques de la couronne de bras et du p6doncule de ces trots especes et

de les comparer avec les Pentacrines du reste du sud-ouest Pacifique.

ABSTRACT

F.chinodermata Crinoidea: Pentacrinidae collected during the Karubar Cruise in Indonesia.

The stalked crinoids collected during the Karubar cruise (Kai and Taninibar Islands: Indonesia) belong to the genus

Saracrinus (Pentacrinidae). Three species in this genus are described, one of which, S. moosai, is new. The samples come trom

depths between 210 and 430 m. This pentacrinid fauna is abundant (165 specimens) but less diverse than the rich launa

collected elsewhere in Indonesia (Sulu Sea, Celebes Sea, Timor Sea, Banda Sea, Java Sea). Twelve species ol pentacnmd were

sampled by previous cruises: Saracrinus angulalus, S. suluensis, S. aculus , S. cingulatus, S. murrayu S. batheri, S.nobilis,

S. superbus, S. varians, Metacrinus serraius, Diplocrinus sibogae, Hypalocrinus naresianus. Data on morphological features,

biometry, arm branching and ossicle articulations are given for each species collected during the Karubar cruise. Stalk, arm

and pinnular joints were observed using scanning electron microscopy with regard to their taxonomic importance. As

numerous individuals of each species were collected, it was possible to study the variation of crown and stem characters. It

appears that the stems and arms of these species, and especially for S. angulatus, exhibit intraspecific polymorphism. One ot

the consequences of these studies is a reduction of the number of recognised species. Some of these forms are interpreted as a

consequence of ecophenotypic or geographical variations. Thus, for example. 5. suluensis, S. aculus, S. cingulatus and

5. batheri are synonyms of S. angulatus, while S. superbus and S. varians are synonyms of 5. nobilis.

AmDziane. N„ 1997 — Echinodermata Crinoidea : Les Pentacrines recoltees lors de la campagne Karubar en Indonesie.

In : A. CROSNIER & P. BOUCHET (eds), Resultats des Campagnes MUSORSTOM. Volume 16. Mem. Mus. natn. Hist, not., 172 :

627-667. Paris ISBN: 2-85653-506-2.

Source : MNHN, Paris

628

N. AMEZIANE

132°E

Source : MNHN Paris

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

629

SOMMAIRE

Introduction .

LlSTE DES STATIONS A PENTACRINES .

Etude systematique .

Genre Saracrinus A.H. Clark, 1923 .

Saracrinus angulatus (Carpenter, 1884)

Saracrinus nobilis (Carpenter, 1 884) .

Saracrinus moosai sp. nov .

CAS DE REGENERATION .

Conclusions .

References bibliographiques .

631

633

645

648

656

657

659

INTRODUCTION

Les differentes expeditions du siecle dernier [celles du "Challenger", 1873-1876 (P.H. Carpenter, 1884) et de

la "Siboga", 1899-1900 (L. DODERLEIN, 1907, A.H. Clark, 1908a)] et la campagne oceanographique danoise de

1922 (Th. MORTENSEN, 1923) ont permis de recolter, pour la premiere fois, un materiel assez abondant de

crinoi'des pedoncules dans les eaux indonesiennes. Parmi ce materiel, le genre Saracrinus , autrefois ddcrit sous le

nom de Metacrinus, dtait particulierement bien represente. L’extreme variabilite de ces organismes avait conduit

les auteurs k une multiplication du nombre des especes. Des travaux recents (Roux, 1981 ; Bourseau & Roux,

1989 ; AMEZIANE-COMINARDI. 1991) nous conduisent a regrouper le materiel recolte lors du siecle dernier en trois

especes (tabl. 1).

Tableau 1. — Inventaire et denomination actuelle des espfcces de Pentacrines recoltes dans les eaux indonesiennes lors des

campagnes du siecle dernier.

Une campagne rdeente (octobre 1991), KARUBAR, s'est deroulee au large des ties Kai, Tanimbar et Aru. Un

abondant materiel (165 specimens recoltes dont 158 mesures) de crinoi'des pedoncules a ete recueilli entre 210 et

430 m de profondeur (fig. 1). Cette faune comporte un seul genre de Pentacrine ( Saracrinus ), reparti en trois

espbees dont une est nouvelle pour la science. L’etude de cette faune est coni^ue comme directement

complementaire de celle des faunes du sud-ouest Pacifique et une comparaison sera dtablie entre les populations

de Saracrinus provenant des differentes regions du Pacifique occidental (fig. 2). Grace au grand nombre de

specimens recoltes, l’analyse quantitative des variations des principaux caracteres morphologiques du pedoncule

et de la couronne a pu etre realisee. Une telle demarche permet ainsi de cerner la variabilite intraspecifique qui

s’avere etre importante pour les Pentacrines.

Source : MNHN Paris

630

N. AMEZ1ANE

Le lecteur voudra bien se reporter aux ouvrages suivants pour ce qui concerne tout specialement la

morphologie des Pentacrines, leur systdmatique, leur ecologie (ROUX. 1981 ; BOURSEAU & Roux, 1989 ;

BouRSEAUera/., 1991 ; Ameziane-Cominardi, 1991).

Source

PENTACRINES DE LA CAMPAGNE KARUBAR EN IND0NES1E

631

LISTE DES STATIONS A PENTACRINES

Karubar. Stations situees au large destles Kai.

Station CP 05, 22.10.91, 05o49'S-132°18'E, 296-299 m : Saracrinus nobilis! 11 exemplaires numerates de sp. n°2

& sp. n°12), Saracrinus angulatus (26 exemplaires numerates de sp. n°l a sp. n°25 et sp. n°61), Saracrinus

moosai sp. nov. (24 exemplaires numerates de sp. n°l it sp. n°24).

Station CP 06, 22.10.91, 05°49'S-132°21'E, 298-287 m : Saracrinus nobilis (2 exemplaires numerates de sp. n°14

a sp. n°15), Saracrinus angulatus (1 exemplaire numerate sp. n°26).

Station DW 13, 24.10.91, 05°26'S-132°38'E, 417-425 m : Saracrinus angulatus (1 exemplaire non mesure).

Station CP 16, 24.10.91, 05° 17'S-132°50'E, 315-349 m : Saracrinus nobilis (1 exemplaire numerate sp. n°13),

Saracrinus angulatus (27 exemplaires numerates de sp. n°27 a sp. n°49 et sp. n°58 a sp. n°60),

Saracrinus moosai sp. nov. (38 exemplaires numerates de sp. n°25 a sp. n°62).

Station CP 25, 26.10.91, 05°30'S-132o52'E, 336-346 m : Saracrinus angulatus (6 exemplaires numerates de sp.

n°50 a sp. n°55).

KARUBAR. Stations situees entre les ties Aru et Tanimbar.

Station CP 46, 29.10.91, 08°0rS-132°5rE, 271-273 m : Saracrinus nobilis (9 exemplaires numerates de sp. n°16

a sp. n°22, deux specimens n’ont pas ete mesures).

Station CP 48, 29.10.91, 08°00'S-132°58'E, 223-218 m : Saracrinus nobilis (1 exemplaire numerate sp. 1).

KARUBAR. Stations situees au large des iles Tanimbar.

Station CP 82, 04.1 1.91, 09o32'S-131°02'E, 219-215 m : Saracrinus nobilis (10 exemplaires numerates de sp.

n°23 a sp. n°32), Saracrinus angulatus (2 exemplaires numerates de sp. n°56 a sp. n°57).

Station CP 83. 04.1 1.91, 09o23'S-131°00'E, 285-297 m : Saracrinus nobilis (1 exemplaire numerate sp. 33).

Station CP 86, 04. 1 1.91, 09°26'S- 131 °13'E, 225-223 m : Saracrinus nobilis (5 exemplaires numerates de sp. n°34

& sp. n°35, trois specimens n’ont pas ete mesures).

ETUDE SYSTEMATIQUE

Genre SARACRINUS A H. Clark, 1923

Rappelons rapidement les principals caracteristiques du genre Saracrinus, avant d'entreprendre une analyse

plus ddtaillee des differentes esp&ces : la premiere division des bras se situe, le plus frequemment, au niveau de la

quatriemc brachiale (I Br 4 ax), il s’ensuit que le nombre de primibrachiales (I Br) est toujours inferieur a 6 ; les

synostoses (articulations non fonctionnelles) des primibrachiales sont localises au niveau de la premiere et de

deuxieme primibrachiale (I Br 1+2).

Saracrinus angulatus (Carpenter, 1884)

Fig. 3-6, 11-12

Synonymie (limitie aux references importantes).

Metacrinus angulatus Carpenter, 1884 : 344. — A. H. Clark, 1908b : 671. — Roux, 1977 : 45.

Metacrinus tuberosus Carpenter, 1884 : 369.

Metacrinus cingulatus Carpenter, 1884 : 347.

Metacrinus acutus Doderlein, 1907 : 35.

Metacrinus suluensis Doderlein, 1907 47.

Metacrinus batheri A. H. Clark, 1909 : 85.

Saracrinus acutus - A. H. CLARK, 1923 : 9.

Saracrinus angulatus - A. H. CLARK, 1923 : 9. — Roux, 1981 : 484. — BOURSEAU & ROUX. 1989 : 1 17. — AmEziane-

Cominardi. 1991 : 73.

Source : MNHN, Paris

632

N. AMEZIANE

Tableau 2. — Principaux caracteres morphologiques du pedoncule et de la couronne chez Saracrinus angulatus (Ties Kai

et Tanimbar).

* : en exposant, nombre de cas observes ; ++ : pores interarticulaires presents sur toute la longueur du pedoncule ;

les caractferes de type continu sont exprimes en mm.

Epaisseur maximale

dc I'inicmodalc

Epaisseur maximale

de la nodalc

Dernier nodi lax is

avcc pores

Longueur ties cincs

9b d'articles/cirre

Longueur couronne

S° du specimen

Station

DiamEtre proximal

DiamEire distal

Nombre

d’iniemodalcs*

-ongueur nodi lax Ls

Nombre dc bras

IBr ax*

IVBrax*

VBrax*

Largeur IBr

Largeur IIBr

Largeur II IBr

-ongucurde PI

Nb d'anicles/Pl

Longueur de PBr

Nb d'arucles/PBr

Sp. 13

CP 05

5.4

9,2

1.2

40.8

113

>63

3*44

5 1 :62

7s ;8 1

92;ll3

123;136

14'

14*15*

17“:182

197;213

232;27'

5,6

3.9

2,7

Sp 14

CP 05

7,1

7,0

Sp. 15

CP 05

JLL.

-ILL

1,5

To-

45,0

148

7S;93

92;1 13

I33;142

15.2

26

-LL

7s:8l3;93

I0':112:I2

13,4

1.2

1.9

44,3

>56

5*63:74

iw

14':15'

13*.I54:16'

I73;18*192

20*.2I2;22‘

25> _

19'

6.4

4.4

3.4

18,9

16

Sp. 16

CP 05

6.1

';8'

7,8

1.2

1,5

30.6

>24

3' ;42;5 1

l';74

8*io'

7':8‘

12'

1 1 ' : 1 2'

13';I7'

16'

12.2

Sp. 17

CP 05

6,7

1.4

1.6

52;63;72

82:9 '

9 ' ; 1 T:12'

I37;14'

I5';163

IsTTT'

182:192

236:252

Sp. 18

CP 05

7I2:85

44.3

Sp. 19

CP 05

6-0

8'°;92

1.2

1,5

39.9

110

76;94

92ao':ii®

i2‘a35a5'

17'

n'a2'a36

is2a62a77

1912;21 1

233

5,0

LL

Sp. 20

CP 05

6.2

67

7";1

1.6

43..0

44:5 1

79-s'

9'.n7a2'

i3*as'

9'a32a4'

is'a62a74

19';202;21s

232:252;31 '

5.0

4,0

Sp. 21

CP 05

S3

7J;8'°:9'

74;82;94

n2a23

i3*a43

i53a64a7

182a97;21 '

25'

5.2

3.1

Sp. 22

CP 05

59

6';7

1.3

Sp. 23

CP 05

-Li-

16

14.o|0

714;8

1.2

2.1

47.0

>81

63;75;92

93ao2ai*

124,132a5'

n'a22a32a58

176a83;194:213

23 *25 *272;29'

31'

15';l7'a92

23' :25 '

5,2

3.8

46,4

64;72:94

92ai4a36

154:17'

i4'assa73

I8'.19s:202

212:232:27'

31'

2,1

TT/g«de iSSSf morphologiques du pddoncul. e, de la ™ro„„e ehez Sarecrinu, mSutam (suile). Mdme

Source

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

633

Lors d'un precedent travail (AMEZIANE-COMINARDI. 1991), je signalais que Saracrinus angulatus etait une

espece polymorphe. De nombreuses especes telles que Saracrinus aculus , S. cingulatus, S. batheri, S. suluensis ,

especes typologiques, exprimaient cette variabilite, raison pour laquelle J.-P. Bourseau & M. ROUX (1989)

integraient tous ces taxons au sein de S. angulatus. Le faible nombre de specimens recoltes alors, restreignait notre

champ d’ investigation, l'apport de quelques specimens nouveaux m'avait permis de mettre en evidence les points

suivants :

- 1’ espece S. tuberosus correspond egalemcnt a un variant de S. angulatus ;

- la grande variabilite de S. angulatus s’exprime aussi bien au niveau des caracteres morphologiques externes

qu’au niveau des articulations ;

- enfin, bien que les diverses populations de S. angulatus du Pacifique occidental soient tres proches, elles se

differencient les unes par rapport aux autres par au moins une caracteristique. II semblerait qu’il existe une

Evolution des caracteres morphologiques en fonction de la localite geographiquc.

Les soixante et un specimens de S. angulatus prelev£s au large des ties Kai et Tanimbar vont permettre de

fournir un complement appreciable a la comprehension de cette espece.

DESCRIPTION DE LA MORPHOLOGIE EXTERNE. — Les specimens indonesiens de Saracrinus angulatus tendent a

etre robustes (fig. 12 A) et leurs principaux caracteres morphologiques se trouvent dans les Tableaux 2 a 6.

N° du specimen

Station

Diamfetre proximal

Dtamfegt distal

Nombre

d'intemodales*

^ongueur noditaxis

Epaisseur maximalc

dc 1‘iniemodale

Epaisseur maximalc

dc la nodale

Dernier nodi lax is

avec pores

Longueur des cirrcs

Nb d'articles/cirrc

Nombre de bras

Longueur couronne

VBr ax*

Largeur IBr

l.argeur IlBr

UrgeurlllBr

Longueur de PI

Nb d'articles/Pl

Longueur dc PBr

Nb d'atticles/PBr

Sp.24

CP 05

6,9

6,0

62;716

87:10i

10.4

1.2

2.0

52

5*7'

13l:14l

15l

18*;21

Sp.25

CP 05

8.1

TTS9-

6";7

1.3

1.8

46.4

4':5‘

63:76

9'

I13;I24

135.144

15*:I6*

14*;15>;162

173;184;19*

204;223;232

252

5.9

33

12.5

6,6

Sp.26

CP 06

5.9

~&t*r

8.5

1.2

1.4

46.1

>63

3 '-A4

51 :6l:7‘

83 1*3 1 : 1 1 1

10- ;1 13:12>

133;14i;161

132;142:15*

163;175;18'

193;212;22*

232:26‘ ;272

3.8

173

3,8

Sp. 27

CP 16

5.5

7,0:8I0;9*

10.5

1.2

1.4

35.5

100

60

7* :8 1 :94

6';97

lll0;132

9*:122;137

14* ; 154

16* : 1 72

18*:19*

Sp. 28

CP 16

6.5

6~

1.2

1.4

50

74 ;8 *

95

1 1 *°;13

142;1 5 *

4,8

11.0

1 1 1 3 3

157:16*

I7";27*

5.5

Sp. 29

CP 16

4,3

2.8

9i3;in#

11'

11.2

1.2

1.4

>26

7 * :8 * :93

I0*:1 1*

1 1 ' :1 3*

152;16*

172

15*

3.2

21.6

14,6

2.7

2.0

13.1

Sp. 30

CP 16

5.6

10.9

1.3

1.7

150

74;83

93

Sp. 31

CP 16

6,9

“

43:52

11*:I35

157

9';ll4

13* 1 : 1 5

3

136;157

173;192

23*

3.9

5*;83

9S;7*

1 1 1 :l 3'

I56;162

174;192

23* :25

5,0

16,4

Sp. 32

CP 16

5.9

5.5

712:84

117

1.3

1.7

125

>58

44 ;5 1

72.98

U4;12*

136:15'

16*

9*:113

133:14*

157;16

I73:212

15'

4.9

3.5

-2JL

20.0

13.2

Sp. 33

CP 16

3.6

4.9

72;84

11.2

1.2

1,5

50,0

120

>44

7*:8*

93

ll3:136

Sp. 34

CP 16

5,7

713;86

94

10.0

1.4

17

19

44;6*

73;82

9'

154

1 1 2 : 1 35

1 54 ; 1 7 1

19*

2,9

1 1 2 : 1 2 '

132:I43

154:16*

15':17*

I8';I9*

23* ;252

5.0

4.0

3.0

16.2

Sp. 35

CP 16

6.2

6,5

39.7'6

>47

15“

72;82

9s

1 1 7 : 1 2 1

133:152

12* ; 1 3*

1SS.16*

174;192

21 ' :22*

Sp. 36

CP 16

5.9

63;714

83

1.2

1.5

>59

' :97

8*:S

10*

9';116;12*

135;14*

15*

13*;142:15*

162;173;193

20*;21273*

24*75*

6.3

3.2

5.4

2.9

14.0

Tableau 4. — Principaux caracteres morphologiques du pedoncuie et de la couronne chez Saracrinus angulatus (suite). Meme

16gende que celle du tableau 2.

Le pedoncuie.

La section transversale du pedoncuie est tres etoilee ; quelquefois elle tend a prendre une forme pentagonale.

Les aretes se marquent fortement (fig. 12 E). Les pedoncules sont, en general, de forte taille. En effet, en partie

proximale, leur diametre varie de 3,6 a 9 mm (la moyenne se situant a 6,1 mm). Leur ornementation s exprime

sous differentes formes :

- carene fine, reguliere, continue et non tuberculde ;

- carene fine, reguliere, discontinue s'estompant au niveau des aretes ou elle est remplacee par une

protuberance isol6e ;

Source : MNHN, Paris

634

N. AMEZIANE

Tableau 5. Principaux caracteres morphologiques du pedoncule el de la couronne chez Saracrinus angulatus (suite) Meme

legendc que celle du tableau 2.

N" du specimen

Station

DiamStre proximal

Jiamfetre distal

Nombrc

d'intcrnodales*

Longueur nodi tax is

Epaisseur maximalc

dc I'imemodalc

Epaisseur maximalc

dc la nodale

Dernier noditaxis

avee pores

Longueur dcs dries

Nb d'articlcs/cirre

Longueur couronne

Sombre dc bras

UBrax*

IVBrax*

VBrax*

Largcur IBr

Largcur HBr

^argeur IHBr

-ongueurde PI

Nb (Taniclcs/Pl

Longueur de PBt

Nb d'articles/FBr

Sp. 49

CP 16

6,3

mm

s'

9,5

1.4

1.8

47,0

68

>47

44;6'

6':74

8‘;9'

92;ll'

12';13S

142:152

16'

TsTie2

17s;182

192;20'

25 2

Sp. 50

CP 25

5.7

32:42

743:82

12,3

1.3

2.0

47.8

P

6';73

94

93;10'

115;122

132;I52

172

4.7

3.4

1.9

13,6

20

rim1

14':I54

173;I8'

22‘;23>

25';28'

23'

2,9

-ii.

Sp. 51

CP 25

72;85

9s

1.3

2.0

134

>60

6';76

8ii92

9':l Is

122;I36

14';152

1 1 ';12I

134;153

162: i 7s

212;22'

3.8

ZS_

11.6

20

Sp. 52

CP 25

6,0

6' ;722

87

10.0

1.4

Sp. 53

CP 25

AL

62;7'6

11.5

1.9

40,7

115

>57

7-':84

92;10'

lO' .l 1 7

122;I3S

14':153

17'

133;153

16':174

I82;192

212

5.0

14,0

20

1.8

42

66

43 ;5 ' :6 '

7">

10':ll2

12 1 ; 13s

14';162

172;182

136;I5'J

176;19s

23'

Sp. 54

CP 25

6,2

8‘

10.0

1.7

152

>62

8';97

I02

94;1 12

I2';137

154

5.2

3,5

15,6

1 1 3 : 1 3 4

14s ;154

174;193

Sp. 55

CP 25

69;72"

1.2

1.5

43,7

48

110

>65

4*1

74:8'

95

1 13;122

I3'°;14'

152

15';18‘

3.7

2,6

15.5

1 0 ': 1 1 4

I2':139

14';I54

17':I8'

19l:20'

Sp. 56

CP 82

7.2

64.

6';7*

1,7

49.3

4J:6'

7I0

1 12; 1 3 "

142

4.2

3.6

3,1

12.7

13';152

17s:19‘

2 1 4 ;23 '

242;25'

33'

19'

5.3

3.9

19.9

15

11.9

Sp. 57

CP 82

5.4

87:9'5

11,7

1.3

1.5

>29

45

77;9'

TTW

17'

15' : 17'

19';21 1

23'

Sp. 58

CP 16

6.9

6,0

5';6''

79

1.4

1.6

20

>51

45

62;72

93

7';9*

Il»;133

4,1

3.6

10,2

17

1 12:134

154;162

174;18'

19' ;21 '

29'

19*;20‘:21 1

5.2

3.9

3,2

9,8

Sp. 59

CP 16

7,0

■44

55:6

1.3

1.7

46.2

63;73

8 ' - 10'

94 ; 102 ; 1 17

12' :13 1

132:I43;15"

16';173;182

192;212;22'

23';242;252

26' ;29 '

6,0

4.0

3,3

21.6

16.5

20

Sp. 60

CP 16

6.5

6,5

6^

8.3

1.5

1.9

47.1

3':44

6' ;72

95

92‘.1 14

132

133;14'

1 5 1 ; 1 72

192;20'

Sp. 61

CP 05

642

1.6

2.1

73

3':43;5'

7S;92

92:117;122

137;14l

15'

13':14';155

162;172;1S4

193;213;24‘

21 ' :24 ' 23';252;33'

4.6

3.5

2J

18.5

I8':19':25'

6.2

“ 2reS d" “ * 1* — Che, anSu,a,u, (S„ite). Meme

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

635

- carene irreguliere, tuberculee ;

- enfin, apparition ponctuelle de quelques tubercules.

Le nombre d'internodales par noditaxis mature presente egalement une importante variability (tabl. 7). II est

compris entre 3 et 13 et se localise lc plus frequemment a 7. En dehors de la partie la plus proximate du

pedoncule, certains specimens presentent une forte heterometrie des internodales qui rdsulte souvent de

l'adjonction de nouvelles columnales. Les pores interarticulaires restent ouverts en partie distale sur une grande

portion du pedoncule. Ils pcuvent, occasionnellement, etre ouverts sur toute la longueur du pedoncule comme le

montrent deux specimens (n°15 et n°49). Le specimen n°59, quant a lui, se caracterise par des apparitions et

disparitions regulieres sur tout le pedoncule.

Tableau 7. — Variability de quelques caractfcres morphologiques chez Saracrinus angulatus (lies Kai et Tanimbar).

Le calice et la couronne de bras.

Les basales jointives montrent des formes differentes :

- basale aussi haute que large avec une apophyse distale pointue qui se marque fortement et qui recouvre les

premieres columnales ;

- basale piriforme, renflee et avec une apophyse distale peu marquee ;

- enfin, basale trapue possedant une apophyse peu marquee et emoussee, ainsi que deux apophyses externes

peu accentudes.

Le rapport de la largeur sur la hauteur des basales est en moyenne dgal a 1,2. Les radiales, quant a elles, sont

jointives, lisses et de forme rectangulaire. Le rapport de leur largeur sur leur hauteur est egal a 3. Les radiales du

specimen n° 52 presentent une apophyse inferieure.

De soixante it quatre-vingts bras constituent la couronne (fig. 12 B). Les brachiales sont le plus souvent lisses

mais parfois prennent un aspect rugueux. Le nombre de dichotomies atteint rarement cinq. Celui des brachiales

par serie augmente progressivement et regulierement des 1 Br aux V Br (tabl. 7). Les coefficients de variation sont

tres eleves a l'exception de celui des I Br. La largeur des brachiales decroTt depuis les 1 Br jusqu aux IV Br. Pour

les I Br, cette largeur est comprise entre 3,2 mm et 6,1 mm (moyenne = 5,1 mm) ; pour les 11 Br, elle se situe entre

2.7 mm et 5,1 mm (moyenne = 3,8 mm) et pour les III Br, elle est comprise entre 1,9 mm et 3,9 mm (moyenne =

2.8 mm). t

Les premieres pinnules sont composees en moyenne de 15 articles et ont une longueur d’environ 17.5 mm. Ces

pinnules sont massives et servent a proteger la masse viscerale. Les pinnules medianes (des II Br aux III Br)

possedent en moyenne 20 articles pour une longueur de 12,9 mm. Elies servent a vehiculer les parttcules

nutritives. La laille des pinnules est en relation avec celle des pedoncules ; plus l'individu est gros et plus les

pinnules tendent a etre importantes. C’est pourquoi, contrairement a certains auteurs (MacKnight, 1973 ; CHANG

& Liao, 1963), la taille des pinnules ne peut en aucun cas etre un critere discriminant differentes especes. Entin,

un S. angulatus (specimen n° 29) presente la particularity de posseder une pinnule bifide (fig. 12 C-D). En partie

distale, une des pieces de cette pinnule se comporte comme une axillaire et il y a dichotomic.

En introduction, je rappelle que les S. angulatus recoltes auparavant dans differentes regions du Pacifique

occidental se caracterisent par leur importante variabilite intraspecifique et leurs caracteres morphologiques qui

evoluent suivant le lieu geographique. Les S. angulatus preleves au large des ties Kai et Tanimbar prysentent

Source : MNHN, Paris

636

N. AMEZIANE

egalement une importante variabilite morphologique. II serait maintenant interessant d’effectuer une analyse

globaie en comparant, entre elles, ces populations de divers secteurs geographiques. Cette demarche permettrait de

verifier ainsi la validite des resultats obtenus lors de mes precedents travaux (Ameziane-Cominardi, 1991).

Analyse geographique de la diversite specifique de S. angulatus.

La localite geographique (pour des coordonnees precises, se referer a AMEZIANE-COMINARDI, 1991), le

nombre de specimens, l’appartenance au museum et l’auteur des mesures des S. angulatus utilises pour I’ analyse

globaie se trouvent dans le Tableau 8.

~t~ RdPanition geographique et nombre de specimens de tous les Saracrinus angulatus recoltds dans le Pacifique

* ; determination et mesures de A.H. Clark ; ** : determination et mesures de J.-P. Bourseau & M. Roux •

: determination et mesures personnelles.

Ces populations sont done regroupees en quatre grands secteurs. LTndonesie est decoupee en deux sous-

secteurs : la mer de Timor et le secteur oriental de la mer de Banda (lies Kai et Tanimbar). Bien que

1 echantillonnage soil tr£s restreint pour la population de la mer de Timor (deux individus), je traiterai ce secteur

geographique independamment de celui, proche, de la mer de Banda. En effet, les deux specimens de la mer de

comespondre"16111 ^ ^ Variabilit6 morPhologique et des caracteristiques particulieres qui peuvent

- son a des orgamsmes qui se trouvent a un pole morphologique extreme d'une population plus homogene

population comprenant entre autres les individus de la mer de Banda ;

' so'1 a des orgamsmes qui appartiennent k une population dont les caracteres morphologiques se distinguent

bien de ceux de la population de la mer de Banda.

La dermere hypothese se justifie par certaines observations faites sur les Saracrinus nobilis dans les eaux

indonesiennes (Amez.ane-Com.nardi, 1991). En effet, ces populations de 5. nobilis se caracterisent par des

J1°rph0l0g,qUeS bien distinctes suivant les localit« geographiques (ex. mer de Banda, mer de Timor,

Le pedoncule.

tr, laoneSffde Vm°I’ d’Australie et d'indonesie se caracterisent par les diametres pedonculaires les plus gros

us^ral enne 3 6mm T"' COmpn* eiUre : 6>1 mm et 6’8 mm (^nne = 6,4 mm) pour la faune

faune I t ’ 3’6 o 8’ 7 m°yenne = 6’5 mm) Pour la faune de la mer de Banda et egaux a 9 mm pour la

" diam6lreS P'd°nCUlaireS d£S f3UneS JaP°naise et Philippine restent dans

e m! (m IT Sltuent’ jespectivement, entre 4,1 mm et 5 mm (moyenne = 4,5 mm) et entre 2,8 mm

lennH ^ L* ^ de mer de Banda pr6senle Ie chamP de variation le plus large et dans

lequel s inseren. parfaitement les autres champs. La faune de la mer de Timor (Indonesie) se distingue des autres

par son tres large intervalle de valeurs, excentre par rapport au reste des autres faunes car les diametres

LceDtionreSm0"/^11' Ce *Ven,ail de Valeurs Se traduit Par un coefficient de variation

Oab 7) E f rTes vale H ** T ““ ^ ^ SCCteUrS ^aP^es sont beaucoup plus faibles

Phil n Enfm’ lesDVa'eurs des "odes sont so.t egales (mer de Timor. Australie), soil superieures (Japon

Philippines, mer de Banda) it celle de la population totale. P ' P°n’

En effetmdbans^’IeXiSte dU n°mbre d’inter"oda|cs nodi.axis mature en fonction de la latitude

En effet, dans le domaine intertropical, Australie et Indonesie (mer de Banda et de celle de Timor), les populations

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

637

de S. angulatus possedent un faible nombre d’intemodales (autour de 7). Quand nous nous eloignons de ces basses

latitudes pour monter vers le Nord, Philippines puis Japon, le nombre d’internodales croTt (8-9). II serait

interessant de voir revolution de ce caractere vers les hautes latitudes de l’hemisphere Sud, mais malheurcuscment

nous n’avons pas encore de populations de ces regions. Involution du nombre d’internodales par noditaxis

mature semble etre symetrique de part et d’autre de l’Equateur. En outre, cette evolution latitudinale globale peut

etre masquee partiellement par une variability locale plus ou moins importante.

Fig. 3. — Variation du nombre d'iniernodales par noditaxis mature en fonction du diam&tre maximal du pedoncule et de la

latitude chez Saracrinus angulatus.

In : mode du nombre d’internodales par noditaxis mature ; N : nombre d’individus ; • : mode pour le nombre

d’intemodales. la grosseur du point represente la taille de la population.

La partie proximale du pedoncule se caracterise chez les Pentacrines par la presence de pores interarticulaires.

Cette zone, qui caractdrise la zone de forte croissance des pedoncules, est toujours mieux developpee chez des

individus adultes que sur des formes juveniles. L'etude du parametre “zone d extension des pores interarticulaires

dans la partie proximale du pedoncule” (fig. 4) montre que :

- cette zone d’extension des pores interarticulaires peut etre tr£s developpee (faune de Banda) ;

- au contraire, elle peut etre restreinte (faune japonaise et Philippine) ;

- enfin, le reste de la faune se repartit entre ces deux extremites.

L’analysc de ce parametre indique que la faune du Japon et celle des Philippines presentent pour ce caractere

une tendance pedomorphique.

La couronne de bras.

L’organisation de la couronne de bras varie suivant les secteurs (fig. 5). Les taunes philippine et japonaise

presentent une organisation statistiquement plus reguliere que montre une variability globale plus restreinte. En

effet, le nombre d’ossicules par serie brachiale augmente regulierement et progressivement. Les faunes de Timor,

de la mer de Banda et d'Australie tendent a posseder moins d'ossicules par tronc brachial.

Source : MNHN, Paris

638

N. AMEZIANE

Tableau 9. — Variability de quelques caractdres morphologiques chez Saracrinus angulaius selon les secteurs geographiques.

N = nombre d'individus ; J. = Japon ; P. = Philippines ; T. = Timor ; A. = Australie ; E. = Ensemble des S. angulaius

recoltes dans le Pacifique occidental.

24

22 .

■5 20

u

! 18

e

^ 16-J

o

a

V)

•S 14 -

12 .

o

N

10 -

8 -

□ o □

o

o □

□

□ rm

□ □

□

□ □ □ □ □

□ □ □ CTD □

cm* c» a □

erf • cn □

cm □

□ □ □ □ □ □

oo

O

234 5 6 7 8 9

Diametre maximum (en mm)

P°'eS d„ diamdtre chdz Saracrinu,

Source : MNHN, Paris

Nombre de brachiales Nombre de brachiales

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

639

6cart-typc

mode

valeur maximale

el minimale

Fig. 5. — Organisation de la couronne de bras chez Saracrinus angulatus en fonction de la repartition geographique.

1 Br ax, II Br ax, III Br ax et IV Br ax : position la plus frequente des axillaires de la premise, deuxibme, troisifeme et

quatribme serie brachiale.

La repartition selon les secteurs geographiques de S. angulatus montre des differences importantes au niveau

des coefficients de variation. Ainsi, les faunes du Japon, des Philippines et de la mer de Banda, presentent la

memc progression de leurs coefficients de variation ; avec cependant, une grande stabilite pour la laune des

Source : MNHN, Paris

640

N. AMfiZIANE

Philippines. En revanche celle de Timor se caracterise par une importante variability pour toutes les sdries

brachiales alors que celle d'Australie a les plus forts coefficients de variation pour les deux premieres series

brachiales et les deux plus faibles pour les troisieme et quatrieme.

Bien qu'irreguliere, la position des dichotomies n'est pas le fait du hasard (Ameziane-Cominardi, 1991 ;

BOURSEAU et al. , 1991). Celie-ci pourrait etre l'expression d'une organisation plus complexe qui serait

significative de l'optimalisation de la fonction "filtration". Ainsi, lorsque la repartition des axillaires dcs deux

premiers troncs est reguliere, celle des derniers troncs est irreguliere. En revanche, lorsque la repartition des

premiers troncs brachiaux est irreguliere, celle des derniers troncs est reguliere. Ainsi, l'importante variability de la

repartition des I Br ax et II Br ax de la faune australienne se trouve correlee a une faible variability des III Br ax et

IV Br ax. Le decalage des dichotomies pennet une meilleure repartition des pinnules au sein du cone forme par la

couronne deployee.

Pour la faune des Philippines, alors que le diametre du pedoncule augmente en fonction de la profondeur, le

nombre de bras decrott (tabl. 10) ; le maximum de bras se situant vers 210 m. En revanche, a profondeur egale et a

diametre pedonculaire equivalent, le nombre de bras de la faune japonaise est plus faible. A profondeur similaire

et diametre du pedoncule egal, les faunes de la mcr de Banda et d'Australie possedent un nombre de bras plus

eleve que celui de la faune des Philippines. II semble que le creneau bathymetrique le plus propice au

developpement des S. angulatus change suivant le lieu geographique en fonction des parametres de

l’environnement, notamment des courants. Nous constatons que le nombre de bras de la couronne decroTt du sud

vers le nord.

Tableau 10. Nombre de bras chez Saracrinus angulatus en fonction de la profondeur du milieu et de la repartition

gyographique.

Caracteres des articulations. — Les zones petaloides des symplexies (fig. 12 H-I) du pedoncule se

caractynsent par des areolas lanceolees (en moyenne deux fois plus longues que larges) a trbs fortement lanceolees

(environ trois fois plus longues que larges). Certains specimens presentent, au sein du reseau de leur areola, de

grosses mailles. Le crenularium interne s'individualise bien. Le nombre de cryneaux varie de 12 a 18 (tabl. 1 1). De

plus, ce nombre de creneaux diminue avec la reduction du diametre du pedoncule. La forme et la largeur des

zones mterpetaloides varient egalement (tabl. 1 1). Le perilumen de tous les specimens se marque fortement. Le

canal axial est toujours circulaire.

Les zones petaloides des synostoses du pedoncule (fig. 12 F-G) se caracterisent par des areolas lanceolees

environ deux tois plus longues que larges) a ires fortement lanceolees (en moyenne trois fois plus longues que

larges). L ensemble du crenularium est bien individualise. Le nombre de creneaux fluctue entre 12 et 15. Le reseau

secondaire contenu dans le canal axial est regulier et de densite plus ou moins lache. La forme du lumen, quant a

elle, est pentalobee de fa?on plus ou moins reguliere (tabl. 11). Les synostoses de la partie distale s'ankylosent

souvent. J

Conclusion. - La grande variability de 5. angulatus s'exprime tres fortement au niveau des caracteres

morphologiques externes et plus faiblement au niveau des articulations. Bien que tres proches, les diverses

populations presentent toutes des differences les unes par rapport aux autres (fig. 6).

‘ aune d Australl®el celle de la mer de Timor presentent les plus forts pourcentages de variation cumulee

alors que la faune des Philippines possede le plus faible. II est etabl. que les specimens considers comme adultes

se differencient des formes plus juvyn.les par : de plus gros diametres, une zone d’extension des pores

641

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

interarticulaires plus importante, un nombre d'internodales par noditaxis mature plus pent, un nombre de bras plus

eleve et un nombre d'ossicules par serie brachiale moindre. Pour l’ensemble de ces caracteres. les faunes du Japon

et des Philippines ont des individus qui correspondent a des formes plus juvemles. Ces deux populations montrent

done une tendance pedomorphique pour la majority des caracteres morphologiques. Ces tendances evolut.ves a

grande variabilite intraspecif.que pourraient traduire une adaptation phenotyp.que aux cond.tions ecologies.

Tableau 11.- Principaux caracteres des articulations du pedoncule chez Saracnnu angula « (p ™v,n“ S^^que)-

* = rapport de la largeur de la zone petalo.de sur celle de la zone .nterpetalo.de ,0 - diamfct « •

Les dchantillons n° 34900 et 36134 = USNM Washington, le sp. 18 = mus.Se de Sydney, le sp. 21 - musde Leiden.

Source : MNHN Paris

642

N. AMEZIANE

Tableau 12. Pnncipaux caracibres morphologiques du pedoncule et de la couronne chez Saracrinus nobilis. Meme lcgendc

que celle du Tableau 2, 6

morphologiques du pedoucule e, de la couronue chee Saracrinus no tm (sui.e). Memo

Source :

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

643

Saracrinus nobilis (Carpenter, 1884)

Fig. 7,13

Synonymie (limitde aux references importantes).

Meiacrinus nobilis Carpenter, 1 884 : 35 1 . — DOderlein, 1912:31. — GlSLEN, 1 922 : 1 47.

Metacrinus murrayi Carpenter, 1884 : 349.

Saracrinus nobilis - A. H. Clark, 1923 : 9. — Roux. 1981 : 489. — Oti, 1986 : 355. — Bourseau & Roux, 1989 : 130. —

AmBziane-Cominardi, 1991 : 65, — Bourseau et al., 1991 : 254.

Meiacrinus superbus Carpenter, 1884:435. — DOderlein, 1907 : 48. — A. H. Clark, 1908:674.

Saracrinus superbus - A.H. Clark, 1923 : 10. — Roux, 1981 : 486. — Bourseau & Roux, 1989 : 125.

Meiacrinus varians Carpenter, 1884 : 352.

Saracrinus varians - A.H. CLARK, 1923 : 9. — ROUX. 1981 : 489. — BOURSEAU & Roux, 1989 : 131.

Cette espece, largement etudiee (Am£ziane-Cominardi. 1991 ; Bourseau et al., 1991) possede une large aire

de repartition comprise entre le Japon et la Nouvelle-Zelande. S. nobilis presente de nombreuses affinites

morphologiques avec S. superbus et les S. varians recoltes dans les milieux profonds. Lors de precedents travaux

(AMEZIANE-COMINARDI. 1991), je signalais que ces trois taxons constituaient un gradient morphologique qui se

singularisait par sa vaste repartition geographique. Leur distribution met en evidence un double relais. Le premier

est un relais ecophenotypique geographique et il affecte seulement les variants de S. nobilis. En effet,.le nombre

d'internodales par noditaxis mature de ces derniers augmente suivant la ligne Sumatra-Timor-Mer de Banda-

Nouvelle-Caledonie. Le second relais ecophenotypique est bathymetrique. Ainsi, S. superbus domine vers 200 m,

S. nobilis vers 300-500 m et S. varians semble infeode aux milieux profonds (AMEZIANE-COMINARDI. 1991).

DESCRIPTION de la morphologie EXTERNE. — La faune de crinoi'des pedoncules recoltes lors de la mission

Karubar livre une quarantaine de Saracrinus nobilis dont la localisation geographique est precisee dans la liste

des stations. L'ensemble des caracteres morphologiques des individus est donne dans les tableaux 12 & 15.

Tableau 14. — Principaux caracteres morphologiques du pedoncule et de la couronne chez Saracrinus nobilis (suite). Meme

legende que celle du tableau 2.

Source : MNHN, Paris

644

N. AMEZIANE

Tableau 1 5. — Principaux caractfcres morphologiques du pedoncule et de la couronne chez Saracrinus nobilis (suile) Meme

legcnde que celle du tableau 2.

Les indi vidus sont de grande taille (fig. 13 A). En effet Ieur diametre varie, en partie proximale, entre 4,8 et

8 2 mm (moyenne - 6,4 mm). La section transversale du pedoncule est toujours etoilee dans sa partie proximale

alors quelle devient circulate ou subpentagonale en partie distale. Le pedoncule est tres souvent lisse (fig. 13 C)

mais quelquefois presente des tubercules plus ou moins marques. Le nombre d'internodales par noditaxis mature

fiuctue entre 8 et 15 et se trouve le plus souvent a 13 (tabl. 16). La longueur de la zone d'extension des pores

interarticulaires est importante (tabl. 16). Les premiers articles des cirres sont arrondis et peu epais puis

deviennent trfes rapidement subrectangulaires et legerement arrondis sous la face inferieure de 1'article formant

atnst une petite apophyse.

Nombre d'internodales

Dernier noditaxis

avec pores _

IBrax

IlBrax

MlBrax

IVBrax

VBrax

VIBrax

Valeur

minimale

8

13

3

5

7

10

13

20

Valeur

maximale

15

27

13

19

49

27

Mode

13

17

11

17

19

23

Coefficient de

variation (%)

8,6

19.4

7,6

18.8

18,4

22,9

18,3

10,7

Nombre

d'observations

550

33

164

304

561

809

190

TAB£?e, ^bVr(?nbdon6?e)qUdqUeS m0rPholo^-s chez les Saracrinus nobilis Voltes au large des Ties Aru,

inHivilbcaSaleSrnt dC f0rme l0oSangique el P°sscdent une legere apophyse. Elies sont jointives chez la plupart des

mats les specimens n 2, 25, 27 et 34 possedent des basales non jointives. Le rapport de la largeu^ sur la

hauteur est egal a 1,53. Les basales tendent tres frequemment ii etre plus hautes que les radiales (fig 7) §

Source :

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONFSIE

645

Les radiales de forme rectangulaire et jointives, sont en moyenne trois fois plus larges que hautes (rapport de la

largeur sur la hauteur = 3,1). La largeur des radiales tend a augmenter avec l'accroissement du diametre pedoncu-

laire. De meme, quoique moins marquee, cette tendance s'observe pour la hauteur et la largeur des basales. En

revanche, pour l'bventail de valeurs actuellement disponibles, la hauteur des radiales n’est pas fortement liee a la

taille du pedoncule. Le specimen n°7 se caracterise par des radiales de petite taille, notamment au niveau de leur

largeur. Cette particularite peut resulter d'un probleme de regeneration. Le specimen n°l presente, quant a lui, des

radiales exceptionnellement bien developpees.

E

e

0>

;§

1

■§ 3

V)

B

©

7

6

5

4

2

1

0

■ H radialc o H basale

□ 1 radiale • 1 basale

o

B □

.*

0

□D g0B nD

□□ °Bb dDb

• • •

0 0*

• ••■b: o

i o o

• • I

•8°'

- -OOj

« ■

°Bea09,

,o 1

o

— i —

7,5

□

4,5

5,5 6 6,5 7

Diametre du pcdoncule (en mm)

Fig. 7. — Dimensions du calice (hauteur et largeur des basales et des radiales) en fonction du diametre maximum du pedoncule

pour Saracrinus nobilis (iles Am. Kai et Tanimbar).

H : hauteur ; 1 : largeur.

Le nombre maximum de bras est compris entre 60 et 120, atteignant rarement six dichotomies (fig. 13 B). Des

verrues ornent les brachiales de quelques individus leur donnant ainsi un aspect rugueux. Le nombre d ossicules

contenus dans chaque serie brachiale augmente regulierement et progressivement depuis les I Br jusqu'au VI Br.

Les pinnules proximales se composent en moyenne de 16 articles pour une longueur de 18,6 mm. Leurs articles

proximaux sont massifs et trapezoidaux, alors que les articles plus distaux sont fins et rectangulaires. Le specimen

n° 24 se particularise par des pinnules proximales anormalement longues, qui peuvent atteindre voire depasser 31

mm de longueur. Les pinnules medianes, situees sur les II Br et III Br, ont en moyenne 21 articles pour une

longueur de 15,4 mm.

Les formes les plus robustes, dont le diametre pedonculaire est superieur a 6,5 mm, possedent les caracteris-

tiques suivantes : leur nombre d'internodales par noditaxis mature se situant tres frequemment a 13, une section

transversale du pedoncule arrondie, un nombre de bras important (souvent superieur a 80). Ces principaux criteres

ont permis a Carpenter (1884) de decrire une nouvelle espece Saracrinus superbus. Nous ne pouvons, pour la

faune indonesienne, discriminer les 5. nobilis des S. superbus. ce qui corrobore les hypotheses faites auparavant

(DODERLEIN, 1907 ; Bourseau & Roux, 1989 ; BOURSEAU et al.. 1991). S. superbus doit done etre mis en syno-

nymie avec S. nobilis. S. superbus representerait un morphotype stable, atteignant un stade “senile", de S. nobilis.

Ce morphotype stable s’epanouirait dans les milieux peu profonds et abrites, alors que S. superbus plus poly-

morphe (pole opportuniste) relaie le premier dans un creneau bathymetrique plus profond (300-500 m).

CARACTERES DES ARTICULATIONS. — Les zones petaloides des symplexies (fig. 13 F-G) du pedoncule peuvent

etre de forme lanceolee a bout triangulaire ou piriforme. De grosses mailles sont presentes au sein du reseau des

areolas de certains specimens. Les aires interpetalo'fdes sont en moyenne 1,2 fois moins larges que les zones

Source : MNHN, Paris

646

N. AMEZIANE

petaloi'des et s'evasent vers l'exterieur. Le crenularium interne est peu developpe. Le nombre de creneaux est

compris entre 12 et 15 (mode a 14). Le perilumen se marque fortement autour du canal axial subcirculaire.

Les synostoses possedent des areolas lanceolees et sont en moyenne 1,6 fois plus larges que les zones

interpdtaloides (fig. 13 D-E). Le nombre de creneaux est compris entre 9 et 10 (mode a 10). Le reseau du canal

axial est dense et irrdgulier. Le lumen, quant a lui, presente une ebauche de lobes.

Rapports et differences. — Les S. nobilis de Karubar entrent dans le champ de variation des 5. nobilis du

reste de la faune du Pacifique sud-ouest (Ameziane-Cominardi, 1991) et presentent les particularites suivantes :

- de faibles coefficients de variability pour les caracteres morphologiques externes tels que le nombre d'inter-

nodales par noditaxis mature et la position de 1'axillaire de la premiere serie brachiale (I Br ax) ;

- une zone d'extension des pores interarticulaires plus etendue ;

- un nombre de brachiales plus eleve pour les quatrieme et cinquieme series brachiales. Ces deux series, pour

le reste de la faune du Sud-Ouest Pacifique, sont constituees d'une quinzaine de pieces ;

- a diametre pedonculaire egal et a profondeur equivalente, un nombre de bras beaucoup plus important.

La faune indonesienne (lies Aru, Kai et Tanimbar) presente d’une part une plus grande stability morphologique

et possede d'autre part des caracteres beaucoup plus matures que le reste de la faune du Pacifique sud-ouest. Cette

laune indonesienne par rapport au reste de la faune se compose de formes que nous pouvons qualifier de "super

adultes" (Ameziane-Cominardi, 1991). Ces caracteristiques pourraient resulterde conditions environnementales,

telles que l'hydrodynamisme, l'aspect du substrat et les apports trophiques, qui seraient plus stables pour les sites’

indonesiens.

Saracrinus moosai sp. nov.

Fig. 8-10, 14-15

Synonymie.

Meiacrinus varians Doderlein, 1907 : 41 (non P. H. Carpenter, 1884).

Materiel examine. —

forid. Une autre station CP 05,

La serie-lype se constitue de 38 specimens provenant de la station CP 16, par 315-349 m de

par 296-299 m de fond, a foumi 24 spycimens.

LOCALITE-TYPE. — Au large de file Kai, Indonesie. Station CP 16 : 24.10.91, 05° 17'S-132o50'E, 315-349 m.

Etymologie. — L'espece est dediee au Dr K. Moosa, chef de mission indonesien de la campagne Karubar.

Diagnose de L'ESPECE. — Saracrinus de taille tres moyenne et de morphologic variable. Pedoncule de section

pentagonale et, le plus souvent, depourvu d’ornementation. La longueur du pedoncule est petite par rapport a celle

de la couronne de bras. Le nombre d'internodales par noditaxis mature est generalement de 5 ou 6. Les cirres sont

pctits, freles et trois a quatre fois plus longs que les internodes. La surface externe des bras est lisse. Les brachiales

preYinent souvent un aspect globuleux. L'organisation de la couronne est la suivante : 3 a 8 I Br (mode a 4), 4 a 13

II Br (mode a 9), 7 a 31 III Br (mode a 13). Les articulations du pedoncule et des bras sont differentes de celles de

5. angulatus et de S. varians (les plus profonds), especes avec lesquelles S. moosai presente le plus d'affinites.

description dela morphologie externe. - L’holotype (specimen n°45) est un specimen de taille plutot

moyenne (fig. 14 A). La longueur du pedoncule est de 1 1 cm pour 26 noditaxis ; son diametre en partie proximale

est de 5 mm et sa section transversale est pentagonale (fig. 14 E). Le pedoncule est de petite taille par rapport a la

couronne de bras. Le pedoncule tend a etre lisse et presente localement une tres faible omementation qui se traduit

par un leger renflement du centre de l'ossicule. Les columnales se caracterisent par une heterometrie. L'epaisseur

des nodales les plus grosses atteint 1,2 mm et celle des internodales 1,0 mm. La hauteur des noditaxis ne depasse

p 7,2 mm. Les pores interarticulaires s'observent jusqu'au douzieme internode. Le nombre total d'internodales

IZrZ T matUr" CSt ^ 5 (16/3S) Cl dC 6 <4 CaS)' 11 6St atleint d6s le douzi^e internode. La longueur des

cirres est d environ 26 mm pour 46 articles.

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

647

Tableau 17. — Principaux caracteres morphologiques du pedoncule et de la couronne chez Saracrinus moosai sp. nov. Meme

legende que celle du Tableau 2.

Tableau 18. — Principaux caract&res morphologiques du pedoncule et de la couronne chez Saracrinus moosai sp. nov,

(suite). Meme legende que celle du Tableau 2.

Source : MNHN. Paris

648

N. AMEZIANE

N° du specimen

Station

Diam&tre proximal

Di,im6tre distal

Sombre

d'intcniodales*

longueur noditaxis

Epaisseur maximale

dc I'intemodale

Epaisseur maximale

de ia nodale

Sp. 33

CP 16

4.7

5,2

3 1 ;4 1

5IS:65

0,8

1.0

Sp. 34

CP 16

4.6

4,0

4i7;53

5.1

1.0

1.1

Sp. 35

CP 16

4.4

4,0

w

5,5

0.8

1.0

Sp. 36

CP 16

4,5

4.0

53;6'2

6,0

0.9

1.0

Sp. 37

CP 16

5.0

4.7

53;6'3

7,9

1.2

1,5

Sp. 38

CP 16

4.9

4';53

Sp. 39

CP 16

5,2

4.6

42;5‘s

5.8

1.1

1.2

Sp. 40

CP 16

4,7

4.4

7?

6.3

1.0

1.1

Sp. 41

CP 16

4,7

4,5

5 1 ;6 1 s:7 1

8.3

1.1

1.4

Sp. 42

CP 16

5.1

5':62

Dernier noditaxis

avec pores

14

12

13

Longueur ties cirres

35.0

Mb d'ariieles/cirre

27.5

Longueur couronne

40

Mombre dc bras

135

35

112

29,0

22.0

42

37

IBrax*

HBrax*

>17

43

103

>39

3‘;43;5'

40

104

39

34.0

39

114

140

40

43;52

45

>34

45

95

>24

44

12

10

25.7

39

103

4]

115

>34

126

40

MlBrax*

92; 1 1 7

137

IVBrax'

136;143;153

202;23 * ;243

26'

VBrax*

Largeur IBr

Largeur HBr

4.6

4.1

Largeur IIIBr

3.6

2.8

Longueur dc PI

2.7

Nb d'aniclcs/Pl

16.8

11.5

Longueur de PBr

Mb d'articles/PBr

15

11,0

9,9

17

20

5 1 :6*

77;9 1

8' ;98:10'

1 1 1 ;I310

142;154

162;18‘

9>;1 ls;133

143;15':162

184;202

72;9 1

II7

92;ll5

1 37: 1 53

4.7

4.3

3.2

3,5

2-3

2,5

15,2

16

9.0

16

Tableau 20. —

174

5.0

3.7

2.9

13.0

13.5

15

7,;83:94

9'-.10‘

ll9:13s

5.0

3.7

2.9

10.0

16

61 ;7J ;9*

1 O’ : 1 2 1

12,;132;15*

162;172:18'

19'

4.5

3.9

3.2

13.1

14

9,6

17

7l;8l

98

92;135

155;17>

193;204

6*;72

95

1 14;I36

I4L.161

17'

11'

4.5

4.8

3.5

3.4

2.6

2.7

16.4

17

10,0

9.8

16

96: 1 1 3

I3>

92; 1 1 4

12';137

1 54; 1 72

4.6

3.4

2.9

11.9

19

sp. nov.

Source : MNHN Pahs

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

649

Tableau 21 . — Principaux caracteres morphologiques du pedoncule et de la couronne chez Saracrinus moosai sp. nov. (suite).

Meme legende que celle du Tableau 2.

Tableau 22. — Principaux caracteres morphologiques du pedoncule et de la couronne chez Saracrinus moosai sp. nov, (suite).

Meme legende que celle du Tableau 2.

Source : MNHN, Paris

650

N. AMEZIANE

Les basales, d aspect trapezoidal, sont jointives. Les radialcs, joinlives ct de forme rectangulaire, sont deux fois

plus larges que hautes. Le noinbre de bras est de quarante (fig. 14 B). La hauteur de la couronne est de 121 mm.

Son organisation est la suivante : I Br 1+2 (5 cas), 4+5 (1 cas), 4 ax (4 cas), 5 ax (1 cas) ; II Br 3+4 (9 cas), 1+2

(1 cas). 5+6 (1 cas), 7 ax (2 cas), 8 ax (1 cas), 9 ax (7 cas) ; III Br 3+4 (19 cas), 4+5 (1 cas), 1 1 ax (6 cas), 13 ax

(6 cas), 14 ax (1 cas), 15 ax (3 cas), 16 ax (4 cas). Deux troncs brachiaux presentent une regeneration ; l'une, au

stade final, affecte la septieme secondibrachiale et l'autre se localise au niveau de ia quatri&me seric brachiale. Les

premieres pinnules (PI) presentent une surface externe qui tend parfois a etre rugueuse et une section

subrectangulaire. Leur longueur atteint 17,1 mm pour environ 15 articles. A partir des II Br, les pinnules

deviennent plus petiles (12,2 mm pour 16 articles) et leur surface externe perd I'aspect rugueux. L'extremite des

bras est depourvue de pinnule fonctionnelle sur environ 26 mm. Les brachiales les plus distales portent

effectivement des pinnules rudimentaires, constitutes de deux ou trois ossicules.

8

2 7

S

c

U

a

<S1

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■s

c

u

V

6 ■

5 ■

4 ■

-- 3 i

<u

u

JS

E

Z

2 ■

'll11

iruru

'2,3 '3,6 ’3,7 '3,9 '4,0 '4,1 '4,2 '4.3 '4,4 '4,5 '4,6 ' 4,7 ' 4,8 ' 4,9 ' 5,0 ' 5,1 '5,2 '5,4 '

Diametre maximum (en mm)

Rc. S. — Variations du nombre d’intemodales par noditaxis mature en fonction du diametre maximal du pedoncule chez

Saracrinus moosai sp. nov. (lies Kai, Indonesie). 1

Le rond noir correspond au mode.

Les Tableaux 17 a 22 regroupent les caracteres morphologiques externes de tous les specimens, y compris

ceux de la sene-type. Pour lensemble de ces individus, la section transversale du pedoncule est pentagonale

parfois une tendance arrondie se dtveloppe chez quelques individus. La section transversale des nodales peut

tendre vers une fomie dtoilee. La plupart des specimens a un pedoncule plutot gracile par rapport a leur couronne

de bras. Les diametres atteignent cxceptionnellcment 6,0 mm et varient en partie proximale entre 2,3 mm et 6 0

mm. Le diametre moyen est de 4,7 mm et 32 individus on. un diametre compris entre 4,0 et 5,0 mm Cetie

?e°xPcfntinnn ITT** ™*f^/™*«** ™tures. En effet. les jeunes specimens sont inexis.ants, a

2T ' i " md‘re V3rie Cnlre 23 mm (en P3™*5 Proximale) et 1,9 mm (en partie

do nC : L > C0lumnales SOnt geoeralement depourvues d'ornementation. Cependant, certains specimens possedent

des ossicules qui peuvent _ porter un tubercule. Ce dernier peut, sur quelques rares ossicules, evoluer vers une fine

carene discontinue. L epa.sseur des columnales varie plus ou moins fortement d'une piece a l’autre. Cette variation

(2 foTnlis neSeCWlainS P3f ra,ternance d'une columnale tres epaisse et d’une tres mince

tabl 23 fio 8? t n°m COlUT eSt comPris entre 3 e> 7 1 le premier mode e.ant a 6 e. le second a 5

40 arM's)- L“'s ~ »

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

651

Les basales sont soit jointives (79 % des cas), soil disjointes (21 % des cas). Elies peuvent parfois posseder une

apophyse distale. Le rapport de leur largeur sur leur hauteur est egal a 1,46. Les radiales, jointives, sont en

moyenne deux fois et demie plus larges que hautes, et ont ainsi un aspect rectangulaire. Les infrabasales sont

egalement presentes (fig. 14 C-D) et se trouvent a l'interieur du cercle de basales. La presence des infrabasales,

decrite pour d’autres espkces (Ameziane-Cominardi et ai, 1991 ; Clark, 1908b ; Doderlein, 1907), semble

etre une caracteristique des Metacrininae.

Tableau 23. — Variability de quelques caractferes morphologiques chez Saracrinus moosai sp. nov.(tles Kai, Itidonesie).

Fig 9. — Morphologie du systeme entoneural (systeme nerveux aboral) dans le calice chez Saracrinus moosai sp. nov. (ties

Kai, Indonesie).

A : projection horizontal ; B : vue de profil.

L'analyse detaillee des pieces de la coupe dorsale permet de transcrire le parcours, dans le calice, du systeme

nerveux entoneural (fig. 9). De l’organe cloisonnd, partent cinq troncs nerveux qui se divisent chacun au niveau

des infrabasales. Deux branches nerveuses pen£trent done dans chaque basale, dies ont pour origine deux

infrabasales differentes. Ces branches sont reliees entre elles, vers le tiers posterieur (cote pedoncule), par un

troisieme cordon nerveux. Ce dernier ne relie entre elles que les branches nerveuses appartenant a la nieme basale,

e'est pourquoi ce cordon nerveux est discontinu. Dans chacune des cinq radiales se trouvent deux ramifications

qui proviennent de basales differentes. Au sommet de la partie anterieure (cote couronne de bras), les deux

branches se soudent dans la radiale et un cordon nerveux circulaire apparait egalement a ce niveau, ce qui permet

une meilleure cohesion du cercle de radiales. Par consequent, une seule branche nerveuse sort de la partie

anterieure de la radiale et entre dans la premiere brachiale. La position des cordons nerveux se revele semblable a

celle de Neocrinus decorus et Metacrinus rotundus (A.H. Clark, 1908c).

Source : MNHN, Paris

652

N. AMEZ1ANE

La couronne brachiale est frequemment constitute d'une quarantaine de bras. Les dichotomies, au niveau de la

quatrieme strie brachiale, sont rares (tab!. 23) et sont exceptionnelles pour la cinquieme serie brachiale (I cas). Le

nombre d ossicules par serie brachiale augmente regulierement et constamment depuis les I Br jusqu'au IV Br (fig.

10). Pour les II Br, un second mode s’exprime au niveau de la septieme brachiale (fig. 10). De meme, pour les III

Br, il existe un second mode au niveau de la onzieme el de la treizieme brachiale et pour IV Br, ce second mode se

trouve aux treizieme, quinzieme et dix-ncuvieme brachiales. La repartition du nombre d’ossicules par serie

brachiale evolue depuis les I Br ou il existe un seul mode (82 % des cas), jusqu’au IV Br ou s’exprime un mode

majcur (24 % des cas) et plusieurs modes secondaires. Ces modes se localisent, a I’exception de la premiere serie

brachiale, toujours au niveau d’un nombre impair de pieces. Les brachiales ont souvent un aspect globuleux. La

largeur des brachiales varie entre 2,9 mm et 5,6 mm pour les I Br (moyenne = 4,7 mm), entre 1,9 mm et 4,1 mm

pour les II Br (moyenne = 3,6 mm) et entre 1,4 mm et 3,7 mm pour les III Br (moyenne = 2,7 mm). La largeur

des ossicules, pour chaque serie, a ete mesuree generalement sur la deuxieme brachiale. La largeur des brachiales

decro.t done regulierement depuis les I Br jusqu’au IV Br et elle est en relation avec la taille du diametre

pedonculaire, exception faite des interferences dues a la regeneration des ossicules. Les premieres pinnules,

massives, sont composees en moyenne de 14 articles et ont une longueur d'environ 14,5 mm. Les pinnules

medianes (des II Br aux III Br) possedent en moyenne 17 articles pour une longueur de 10,7 mm.

Nombre de brachiales par serie

Pn^.j,1 °; 0r§anisallon de la couronne de bras chez Saracrinus moosai sp. nov. (He Kai, Indonesie)

En haut et a droite : mode et dean-type pour chaque strie brachiale. Meme legende que celle de la figure 5.

5. moosa, sp. nov. possedc le plus faible coefficient de variation. nod.tax.s avec pores ,

Caracteres des articulations. - Les symplexies du pedoncule (fig. 14 G-H • fig 15 A) oresentent des

“ ToTb 'T' ento“r&s d'™ ££££

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

653

s'6vasant vers l'exterieur de la piece (en moyenne 1,5 fois moins larges que la zone petaloi'de). Le perilumen

s'exprime tres fortement. Le canal axial est de forme circulaire.

Les zones petaloi'des des synostoses du pedoncule sont egalement lanceolees (fig. 15 B-C). Ces zones sont

egalement soit ouvertes vers l'exterieur, soit fermees. Le nombre de creneaux varie entre 8 et 10 (mode a 10). Les

zones interpetalo'ides sont bien differences (en moyenne 1,75 fois moins large que la zone petaloTde). Ces zones

interpetaloi'des sont beaucoup plus larges au niveau des synostoses distales. Le reseau secondaire qui remplit le

canal axial est de forme pentagonale, lache et regulier. Le lumen secondaire est irregulier et pentalobe.

Les synarthries des bras ont une Crete fulcrale qui est nettement oblique et bien en relief (fig. 15 E, G). Le

sillon interne, qui separe les aires musculaires, est etroit. La zone ligamentaire externe est non seulement bien

developpee mais est egalement fortement evidee. Les aires ligamentaires internes presentent une limite franche

avec les aires musculaires. Ces dernieres sont bien developpees et leurs zones de croissance sont tres apparentes.

L'insertion pinnulaire provoque une dissymetrie des aires musculaires. Une evolution de la microstructure des

synarthries existe depuis les brachiales proximales jusqu'aux brachiales distales (fig. 15 E, G, H, J). En effet, le

perilumen qui entoure le canal axial est bien individualist et a une forme trapezoidale pour les ossicules

proximaux, alors qu’il est pratiquement inexistant pour les ossicules distaux. Les zones musculaires deviennent de

plus en plus restreintes et sont pratiquement inexistantes en partie distale. L'evidement de l'aire ligamentaire

externe est de plus en plus faible, de meme que la crete fulcrale se difference de moins en moins, au fur et a

mesure que les brachiales apparaissent en position distale.

Les synostoses des bras sont generalement planes (fig. 15 D, I). Le sillon median est bien marque. Les

synostoses proximales (fig. 15 D) presentent, des la partie mediane de l'articulation, des epaississements

concentriques de reseau syzygial. Ces epaississements sont absents des synostoses distales (tig. 15 I).

Pour les deux types d'articulation, le canal axial dvolue depuis des formes rectangulaires pour des brachiales

proximales jusqu'a des formes circulaires en partie distale.

Les premiers ossicules des pinnules proximales ou mddianes possfedent des synarthries qui sont beaucoup plus

evoluees que celles des brachiales les plus distales (fig. 15 K, M). Leurs cretes fulcrales, bien que peu

differenciees au niveau microstructure, sont nettement individualists. De meme, les aires musculaires sont

visibles. En revanche, les ossicules plus distaux des pinnules proximales ou medianes sont tres peu differences

(fig. 15 L).

RAPPORTS et DIFFERENCES. — Saracrinus moosai est associe avec S. nobilis et S. angulatus. S. moosai

sp. nov. est tres different de S. nobilis , alors que certains specimens ont un aspect general qui presente parfois

quelques analogies avec S. angulatus. Cependant l'etude detaillee des caracteres morphologiques permet de

separer les deux especes. En effet, meme si quelques noditaxis matures du pedoncule de certains S. moosai

sp. nov. se composent parfois de 7 columnales, la majorite d'entre eux possedent 5 S 6 ossicules. De plus, la zone

d’extension des pores interarticulaires est beaucoup plus restreinte chez S. moosai sp. nov. En revanche,

difference S. moosai sp. nov. de 5. angulatus seulement avec la couronne de bras se revele difficile. Toutefois, il

semble que S. moosai sp. nov. possfede plus d'ossicules au niveau de la deuxieme serie brachiale. Au niveau des

articulations du pedoncule, S. moosai sp. nov. presente par rapport a S. angulatus les differences suivantes : des

zones petaloi'des des symplexies et des synostoses beaucoup moins lanceolees ; un nombre de creneaux plus

faible ; un crenularium interne peu developpe et un perilumen moins large. Les principales differences au niveau

de l'articulation des bras se situent essentiellement au niveau de l'obliquite de la crete fulcrale (plus forte chez 5.

moosai) et la presence, chez S. angulatus, d'apophyses calcitiques sur les bords internes de l'aire ligamentaire.

S. moosai sp. nov. tend egalement a presenter de tres fortes affinites avec trois des cinq S. various decrits par

L. DODERLEIN (1907). Ces individus ont ete recoltes a la station 253 de l'expedition de la "Siboga". Cette station

se trouve a proximite de la station CP 05 de Karubar et a une profondeur similaire. Les specimens decrits par L.

DODERLEIN presentent non seulement la meme allure generate mais egalement des caracteristiques

morphologiques identiques. Ce sont done bien des 5. moosai. Le nombre limite de S. various et la faible

connaissance sur l'importante variabilite inherente a chaque espece a masque cette nouvelle espece. Les deux

autres S. various, preleves entre 500 et 600 m. sont des morphotypes profonds de S. nobilis (AMEZIANE-

COMINARDI, 1991).

Source : MNHN, Paris

654

N. AMEZIANE

CAS DE REGENERATIONS

Les crinoi'des pedoncules ont une grande capacity k se regenerer (Bourseau et al., 1991 ; OJI. 1986) et peu-

vent le faire plusieurs fois au cours de leur existence. Une taille plus petite et une couleur differente, souvent plus

claire, indiquent que 1 individu est en cours de regeneration. La regeneration traduit un instant t qui ici est le

moment du prelevement. En effet, un specimen peut s'etre deja regenere plusieurs fois et ne porter aucune trace

significative de cette regeneration lors de sa recolte, car il se trouve dans une phase de “non-regeneration”. La fre¬

quence de regeneration sur I'ensemble d’une population recoltee dans une meme station refletera les perturbations

subies par les mdividus. Theoriquement, toutes les parties de l’organisme presentent la possibility de se regenerer

Ma.s la presence des basales (Ameziane-Cominardi, 1991 ; Amemiya & Oji. 1992) et de la partie proximale du

pedoncule se revele indispensable pour que la regeneration s’effectue. En effet, le centre nerveux aboral contenu

dans ces elements est necessaire pour que le processus se declenche. Les results d'experimentation (Amemiya &

OJI. 1992) montrent egalement qu’une resorption de la partie proximale du pedoncule se manifeste lorsque

I'ensemble de la couronne de bras est en cours de regeneration. Cette particularity a ete observee chez un

Metacrinus levii (Bourseau et al., 1991), un M. wyvillei et un M. serratus (BOURSEAU & ROUX, 1989).

Le pedoncule. L’observation du pedoncule en cours de regeneration est rare. Aucun des individus recoltes

lors de la mission Karubar ne presente de regeneration visible du pedoncule.

Lecalice. — Un seul specimen ( Saracrinus moosai, specimen n°l) montre une anomalie au niveau du calice

En effet cet indiv.du possede six radiales ; l'une d'entre-elles etant plus petite et surdlevee par rapport aux autres.

Chacune de ces six radiales porte un tronc brachial. Cette anomalie peut resulter soil d'un probleme lors d’une

regeneration, soit d'un defaut de naissance.

La couronne de bras. — La regeneration des bras, quant il elle, est frequente et les cassures s'effectuent gdnd-

ralement au niveau des synostoses. Une aberration particuliere, due certainement a un probleme de regeneration

est observe pour la premiere fois chez un 5. angulatus (specimen n°2). Au niveau de la quatrieme primibrachiale

(1 Br 4) ou normalement se situe la division brachiale, se trouve une axillaire avortee (fig. 11). En effet, cette

piece a failure generale dune axillaire par sa forme losangique, accentuee par un bee central. A fendroit’ou se

localise habituellement la pinnule, apparaissent deux brachiales singulieres : la premiere (1 Br 5), tres petite se

niche dans le creux de la I Br 4 lui conferant ainsi une vocation d'axillaire ; la seconde (I Br 6) recouvre non

seulement la I Br 5 ma.s egalement une partie de la I Br 4. De cette situation, il resulte une dissymetrie de la I Br 6

qu. est bien dSveloppee du cote ou se trouve I Br 5, mais qui est tres fine sur le cote oppose. La fonc.ion de cette

piece serait de comger 1 anomalie induite par I Br 4. Pour ce tronc brachial, faxillaire se localise, en fait, au niveau

de la huitieme brachiale (I Br 8 ax).

' septa de S«raZr^toS2SSrP'im4e “ P°“ge- " ‘“8 d“ l«*f~ trn les tsois

pourcentage total ; - : pas de regeneration observee. de 2 ou 3 ossicules) par rapport au

PENTACR1NES DE LA CAMPAGNE KARUBAR EN INDONESIE

655

Les frequences de regenerations s’obticnnent a partir du nombre total de regenerations obscrvees sur une

meme serie brachiale compare au nombre total de troncs brachiaux de tous les individus (regeneres ou non) pour

cette meme serie brachiale. Sur l’ensemble des bras, S. nobilis presente les plus forts pourcentages de

regenerations et S. angulatus les plus faibles (tabl. 24). Pour les trois especes, ce sont au niveau des II Br et III Br

que s’observent le plus grand nombre de regenerations, alors qu'elles sont les plus rares au niveau des I Br et V

Br. Toutes les frequences de tres jeunes regenerations augmentent depuis les II Br jusqu’au V Br, k l’exceplion

des III Br de S. moosai sp. nov. II semble que les parlies distales, plus fragiles, se r6g6nerent plus souvent. Si nous

comparons ces donndes avec celles obtenues dans le SW Pacifique (AMEZIANE-COMINARDI, 1991), nous

constatons que les S. angulatus indonesiens (lies Kai et Tanimbar) se caracterisent par un taux de regeneration

plus faible que celui observe chez ceux recoltes dans le reste du SW Pacifique. Les S. nobilis d'Indonesie (lies

Aru, Kai et Tanimbar), quant a eux, montrent un taux de regeneration plus important pour les trois premieres

series brachiales que celui obtenu chez les individus recoltes dans le reste du SW Pacifique. Les S. nobilis

indondsiens se caracterisent egalement par une plus forte variabilite de leurs caracteres morphologiques.

A B

Br 8 a x

Bre

Br 5

Br4

Bri

Fig. 11.— Schema representant l’anomalie de la premiere serie brachiale (1 Br) du specimen Saracrinus angulatus n°2 (Ties

Kai, Indondsie).

A : vue du cote lateral ; B: vue du cote dorsal.

I Br 1, I Br 4, I Br 5 et I Br 6 = premiere, quatrieme, cinquieme et sixieme primibrachiale ; 1 Br 8 ax = huitieme

primibrachiale axillaire ; P = pinnule.

La relation qui existe entre les fortes frequences de regeneration et les forts coefficients de variation chez de

nombreuses especes (BOURSEAU et al., 1991) se verifie de nouveau. Les mysostomes ont ete invoquds, de meme

que la predation et faction des courants (A.H. CLARK, 1921 ; CONAN et al., 1981 ; OJI, 1986), pour expliquer la

cassure des bras. Aucun mysostome n'a ete trouve chez S. moosai. En revanche, chez certains specimens de

5. angulatus et S. nobilis, un certain nombre de mysostomes sont presents. Ces organismes affectent aussi bien les

brachiales que les pinnules. Un individu peut etre infeste par de tres nombreux mysostomes ; par exemple,

quatorze individus parasitent le S. nobilis n°19 et douze parasitent le n°21.

CONCLUSIONS

L’etude du materiel recueilli par la campagne Karubar confirme qu'il existe une grande variabilite

intraspecifique chez les Metacrininae et que de nombreuses especes sont, en fait, des ecophenotypes qui sont

susceptibles de traduire les variations du milieu environnant. Sur les huit especes de Metacrininae recoltees aux

abords des lies Kai lors de precedentes campagnes ("Challenger", "Siboga", expedition danoise), trois especes

Source : MNHN, Paris

656

N. AMEZIANE

seulement sont indiscutablement presentes dans cette region. Ainsi, Saracrinus acutus, S. cingulatus, S. tuberosus

deviennent synonymes de S. angulatus, de meme que S. superbus , S. murrayi, S. varians (certains specimens) sont

synonymes de S. nobilis. Les S. varians, dccrits par L. Doderlein, ont ete mal identifies et correspondent a des

5. moosai sp. nov.

Lors des diverses campagnes, passees ou recentes, les specimens preleves dans une meme station appar-

tiennent souvent a des especes et a des genres differcnts. Quelquefois, S. nobilis et 5. angulatus sont associes dans

un meme site, mais la encore la faune de Karubar se singularise par la presence simultanee de trois especes

appartenant au meme genre.

La faune recoltee lors de la campagne Karubar se caracterise par des populations homogenes depourvues de

juveniles. En effet, les Saracrinus nobilis et angulatus, par rapport au reste de la faune SW Pacifique, ont de fortes

taiHes. De meme, S. moosai, espece plus gracile que les deux prccedentes, semble se situer a un pole maximal de

croissance. Plusieurs hypotheses peuvent etre envisagees pour tenter d'expliquer cette situation :

- il peut exister un probleme au niveau de l'echantillonnage. Les techniques de prelevements utilisees sont les

memes que celles employees lors des campagnes effectuees au large de la Nouvelle-Caledonie, region ou de

nombreux juveniles ont etc recoltes alors que le nombre total d'individus preleves est moindre que celui de la

campagne Karubar. Cette premiere hypothese semble done peu vraisemblablc ;

les larves des Saracrinus des Ties Kai se deplacent, de fa^on active ou passive (action des courants), et

s'installent ailleurs ;

- enfin, ces populations ont des vitesses de renouvellemeni faunistique lentes car leur environnement est stable.

Lors des campagnes effectuees auparavant au large des Ties Kai, Democrinus weberi et un Rhizocrinus

(stations 56 et 58 de l’expedition danoise, MORTENSEN, 1923) ont ete preleves en meme temps que des

Saracrinus. GlSLEN (1925) decrit un Democrinus globularis et un Porphyrocrinus verrucosus (station 56 de la

"Siboga "). Des prelevements ont ete faits aux memes endroits et a dcs profondeurs similaires, mais

malheureusement aucune de ces especes n'a ete recoltee.

De nombreuses autres especes sont potentiellement presentes dans ces eaux. Au sein des Pentacrinidae, les

diverses recoltes passees ont permis d' identifier Metacrinus serratus (mer de Sulu), M . wyvillei (Celebes),

M. costatus (Celebes), Diplocrinus sibogae (mer de Timor), Hypalocrinus naresianus (Celebes). L’absence de

Metacrinus est surprenante car ce genre se trouve tres frequemmcnl et plus regulierement que Saracrinus dans

tout le SW Pacifique. Enfin, parmi les Bathycrinus, B. minimus (Celebes), B. nodipes (Celebes et Banda) et

B-Poculum (Banda) ont ete preleves lors du passage de la "Siboga". Parmi la quinzaine de stations de Karubar

effectuees a des profondeurs ou se developpent generalement de telles especes, aucune n’a fourni de crinoides

pedonculcs. Deux hypotheses peuvent expliquer I ’absence de ces organismes : soit ces formes sont effectivement

presentes aux abords des lies Kai et n’ont pas ete recoltes pour des raisons diverses telles que : zones de

developpement tres restreintes, deplacement des organismes, probleme d’echantillonnage ; soit alors ces especes

ne vi vent pas dans ce secteur. Nous constatons que la faune des diverses mers qui separent les lies (Timor,

Celebes, Banda..) differe suivant les secteurs (especes rencontrees qu’a un seul endroit ; cf. Diplocrinus sibogae)

et de plus elle presente des variations morphologiques importantes pour des especes deja connues (cf. Saracrinus

nobilis et S. angulatus). Ainsi, les crinoides pedoncules pourraient etre caracteristiques de chacune de ces regions,

mais en 1 etat actuel de nos connaissances ceci reste une simple supposition.

REMERC1EMENTS

Je remercie les membres de la mission Karubar, et plus particuiierement les personnes appartenant au

Museum national d H.sto.re naturelle de Pans et a l’ORSTOM, pour toutes les precautions prLs lors du

S5S—*8 SPCC,menS- permeUam 3inSi d'°btemr d6S mdividuS P^tement preserves. Je remercie

I R, rapporteurs, plus specialement Jean-Paul Bourseau, d’avoir lu avec attention ce manuscrit

Leurs critiques constructives ont permis d’en ameliorer sa qualite. Je tiens egalement & remercier L. Martini

PENTACRINES DE LA CAMPAGNE KARUBAR EN IND0NES1E

657

de son aide pour les photographies, Mesdames D. GuiLLEMlN et F. Devienne pour leur gentillesse et leur

competence au microscope electronique h balayage et A. FOUBERT pour ses precieux conseils informatiques.

REFERENCES

Amemiya, S. & Oji.T., 1992. — Regeneration in sea lilies. Nature, 357 (6379) : 546-547.

AmEziane-Cominardi, N., 1991. — Distribution bathymetrique des Pentacrines du Pacifique Occidental. Essai

de modelisation et d'application au Lias. Documents du Laboratoire de Geologie, Universite de Lyon, 116 : 1-253, 5 pis.

BOURSEAU, J.-P. & Roux, M„ 1989. — Echinodermes : Crinoi'des Pentacrinidae (Musorstom 2 & Corindon 2). In : Resultats

des Campagnes MUSORSTOM, Volume 4. Memoires du Museum National d'Histoire Naturelle, (A), 143 : 1 13-201, 1 1 pis.

BOURSEAU, J.-P., AmEziane-Cominardi. N„ Avocat, R. & ROUX, M.. 1991. - Echinodermata : Les cnnoides pddoncules

de Nouvelle-Caledonie. In : A. Crosnier (ed.). Resultats des Campagnes Musorstom, Volume 8. Memoires du Museum

National d’Histoire Naturelle, (A), 151 : 229-333, 16 pis.

Carpenter, P. H., 1884. — Report upon the Crinoidea collected during the voyage of H.M.S. Challenger during the years

1873-1876. Part I - General morphology with descriptions of the stalked crinoids. Challenger Reports, Zoology , 11 (32) .

1-442,62 pis.

Chang, F. Y. & Liao, Y. L., 1963. — On the Recent stalked crinoids of China. Acta Zoologica Sinica, 15 (2) : 282-288, 2 pis.

Clark. A. H., 1908a. — The stalked crinoids of the Siboga Expedition. American Naturalist, 42 : 203-206.

Clark, A. H., 1908b. — Infrabasals in Recent genera of the crinoid family Pentacrinidae. Proceedings of the United States

National Museum, 33 : 671-676.

Clark, A. H., 1908c. — The axial canals of the Recent Pentacrinidae. Proceedings of the United States National Museum. 35 :

87-91.

Clark, A. H„ 1909a. — New Recent crinoid from the Indian ocean. Proceedings of the Biological Society of Washington. 22 :

73-86.

Clark, A. H.. 1909b. — On a collection of Recent crinoids from the Philippines Islands. Proceedings of the United States

National Museum, 36 : 391-410.

Clark, A. H„ 1909c. — Description of seventeen new species of Recent crinoids. Proceedings of the United States National

Museum, 36 : 633-651.

CLARK, A. H„ 1909d. — New Recent Indian crinoids. Proceedings of the Biological Society of Washington, 22 : 143-156.

Clark, A. H., 1921. — Sea lilies and feather-stars. Smithsonian Miscellaneous Collections, 72 (7) : 1-43. 16 pis.

Clark, A. H„ 1923. — A revision of the Recent representatives of the crinoid family Pentacrinidae with the diagnoses of two

new genera. Journal of the Washington Academy of Sciences, 13 (1) : 8- 1 2.

Conan, G.. Roux, M. & Sibuet, M.. 1981. — A photographic survey of the stalked crinoid Vipjocri™3 {Annacrmus)

wyvillethomsoni (Echinodermata) from the bathyal slope of the Bay of Biscay. Deep Sea Research, 28A (5): 441 -45J.

DOderlein, L., 1907. — Die gestielten Crinoiden der Siboga-Expedition. Siboga-Kxpeditie, 42a : 1-54, 23 pis.

Gislen, T„ 1922. — The crinoids from Dr S. Bock's expedition to Japan - 1914. Nova Acta Regiae Societatis Scientiarum

Uppsaliensis, sdr. 4. 5 (6) : 183, 2 pis.

GlSLEN, T., 1924. — Echinoderm studies. Zoologiska Bidrag fran Uppsala, 9 : 1-330. 349 text-fig.

GISLEN, T., 1925. — Two new stalked crinoids from the Kei Islands. Videnskabelige Meddelelser fra Dansk Naturhistorisk

Forening, 79 : 85-95, text figs 1-22.

MacKnight. D. G., 1973. — Stalked crinoids from the New Zealand region. New Zealand Oceanographic Institute, Records,

1 (14) : 199-210.

MORTENSEN, Th„ 1923. — The Danish expedition to the Kei Islands 1922. Videnskabelige Meddelelser fra Dansk

Naturhistorisk Forening, 76 : 55-100.

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Oji, T„ 1986. — Skeletal variation related to arm regeneration in Metacrinus and Saracrinus, Recent stalked crinoids. Lethaia

19 (4) : 335-360.

Oji. T„ 1989. — Of the stalked crinoids from Japanese and nearby waters. In : H. Obha, I. Hayami, & K. Mochizuki (eds).

Current aspects of biogeography in West Pacific and East Asian regions, Volume 1 : 27-43. University Museum

University of Tokyo.

ROUX, M., 1977. — The stalk joint of Recent Isocrinidae (Crinoidea). Bulletin of the British Museum ( Natural History)

Zoology, 32 (3) : 45-64.

Roux, M.. 1980. — Les articulations du pddoncule des Hyocrinidae (Echinodermes, Crinoides pddoncules) : interet

systi5matique eI consequences. Bulletin du Museum National d'Histoire Naturelle, ser. 4, Zoologie, 2 : 31-57, pis 1-5.

ROUX, M„ 1978. — Ontogenese et evolution des crinoides pedoncules depuis le Trias. Implications oceanographiques. These

University Paris-Sud. 167 pp., 1 annexe. Inedit.

Roux. M., 1981. Echinodermes : Crinoides Isocrinidae. In : Resultats des Campagnes Musorstom. I - Philippines (18-28

mars 1976). Memoires ORSTOM, 91 : 477-543, 15 pis.

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

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PHOTOGRAPHIES

Source : MNHN, Paris

660

N. AMEZIANE

FIGURE 12

Saracrinus angulatus (Carpenter, 1884)

A. — Specimen n°9, station CP 05. Vue generale (x 0,6).

B. — Specimen n°9, station CP 05. Vue de detail montrant le calice et l'organisation de la base de la couronne de bras.

Les premieres pieces axillaires sont les quatriemes primibrachiales (x 1,4).

C. — Specimen n°29, station CP 16. Vue gEnErale de la pinnule bifide (x 4).

D. Specimen n°29, station CP 16. Vue de detail de la pinnule bifide. La piEce ou se localise la division se comporte comme

une axillaire (x 7).

E. Specimen n°9, station CP 05. Vue de detail montrant l'organisation du pedoncule et l’ornementation des columnales.

Le nombre d'intemodales par noditaxis mature est de 7 sur les noditaxis complets (x 1,4).

F. — Specimen n°34900 (collection de l’USNM a Washington), station 5623, "Albatross". Vue gEnErale au microscope

Electronique a balayage d’une synostose de nodale distale (x 7,5).

G. — Specimen n°34900 (collection de l’USNM h Washington), station 5623, "Albatross". Vue de detail de la figure

precedente. Vue du canal axial de la synostose (x 60).

H. — Specimen n°56, station CP 82. Vue gEnErale au microscope Electronique it balayage d’une symplexie d’une nodale

distale (x 5,6).

1 ~ sP^clmen n°56- s'a'ion CP 82. Vue gEnErale au microscope Electronique h balayage d’une symplexie d’intemodale distale

Les vues au microscope electronique ont EtE effectuEes au C.I.M.E., dEpartement M.E.B., Paris VI.

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

661

Source : MNHN, Paris

662

N. AMEZIANE

FIGURE 13

Saracrinus nobilis (Carpenter, 1884)

A. — Specimen n°27, station CP 82. Vue gdndrale (x 0.6).

B. — Specimen n°27, station CP 82. Vue de detail montrant le calice et I'organisation de la base de la couronne de bras. Les

premieres pibces axillaires sont les quatribmes primibrachiales (x 1,4).

C. Specimen n°27, station CP 82. Vue de detail montrant I'organisation du ptSdoncule et l’omementation des columnales.

Le nombre d’intemodales par noditaxis mature est de 12 sur les noditaxis complets (x 1.4).

D. — Specimen n°!6. station CP 46. Vue generale au microscope Electronique h balayage d’une synostose de nodale distale

(x 7).

E. Specimen n°16, station CP 46. Vue de detail de la figure prEcEdente. Vue du canal axial de la synostose (x 115).

F. — Specimen n°l, station CP 48. Vue generale au microscope Electronique a balayage d’une symplexie d’une nodale distale

(x 6,5).

T ftPff'men h0*1 staIlon cp 48- Vue generale au microscope electronique b balayage d'une symplexie d’intemodale distale

(X 0,j).

Les vues au microscope electronique ont EtE effectuees au C.I.M.E., departement M.E.B., Paris VI.

.iui y’ii

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONES1E

663

Source MNHN Paris

664

N. AMEZIANE

FIGURE 14

Saracrinus moosai sp. nov.

A. — Holotype (specimen n°45), station CP 1 6. Vue generale (x 0,8).

Holotype (specimen n°45), station CP 16. Vue de detail montrant le calice et (’organisation de la base de la couronne de

bras. Les premieres pieces axillaires sont les quatridmes primibrachiales (x 1,4).

C. — Specimen n°15, station CP 05. Vue gendrale au microscope electronique k balayage du calice (x 7). R = radiales ;

B = basales ; I = infrabasales ; C = premidres columnales.

D. — Specimen n°6, station CP 05. Vue de ddtail au microscope electronique k balayage des infrabasales (x 35).

E- Holotype (specimen n°45), station CP 16. Vue de ddtail montrant I'organisation du pedoncule et l'omementation des

columnales. Le nombre d'intemodales par noditaxis mature est de 5 (7 cas), 6(1 cas) sur les noditaxis complets (x 1.4).

F. Specimen n°15, station CP 05. Vue generale au microscope dlectronique k balayage d'une ieune columnale proximale

(x 30).

G. Specimen n°l, station CP 05. Vue gdndrale au microscope dlectronique k balayage de la symplexie d’une intemodale

distale (x 8,5).

H. Spdcimen n°15, station CP 05. Vue de detail de la figure precedente. Vue du canal axial et du perilumen (x 45).

Les vues au microscope dlectronique ont dte effectuees au C.I.M.E., departement M.E.B., Paris VI.

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

665

Source : MNHN} Paris

666

N. AMEZIANE

FIGURE 15

Saracrinus moosai sp. nov.

A. — Specimen n°15. station CP 05. Symplexie d'une intemodale distale (x 8.5).

B. — Specimen n°15, station CP 05. Synostose d'une infranodale distale (x 8.5).

C. — Specimen n°15, station CP 05. Vue de detail de la figure prdcddente. Vue du canal axial de la synostose (x 1 10).

D. — Specimen n°15, station CP 05. Vue geniirale de la synostose de la deuxiemc primibrachiale, I Br 2, (x 1 1).

E. — Specimen n°15, station CP 05. Vue generate de la synarthrie de la troisteme primibrachiale, I Br 3, (x 10).

F- — Specimen n°15, station CP 05. Synarthrie de la quatrteme primibrachiale axillaire, I Br 4, (x 9).

G. — Specimen n°15, station CP 05. Synarthrie d'une brachiale distale appartenant h la troisteme s6rie brachiale. III Br, (x 20).

H. — Specimen n°15, station CP 05. Synarthrie d'une brachiale distale (x 35).

I. — Specimen n°15, station CP 05. Synostose d'une brachiale distale (x 35).

J. Specimen n°15, station CP 05. Synarthrie d'une brachiale terminale qui porte une pinnule non fonctionnelle (x 40).

K. — Sp6cimen n°15, station CP 05. Vue ventrale d’un <§tement proximal appartenant & une pinnule proximale (x 20).

6- Specimen n°15, station CP 05. Articulation d’un 6tement distal d'une pinnule proximale (x 45).

M. Specimen n°15, station CP 05. Articulation d'un element proximal d'une pinnule distale (x 45).

N. Specimen n°15, station CP 05. Vue ventrale d’un ossicule d'une pinnule distale non fonctionnelle (x 100).

Les vues au microscope 61ectronique ont 6te effectudes au C.I.M.E., d<5partement M.E.B., Paris VI.

Source : MNHN, Parts

PENTACRINES DE LA CAMPAGNE KARUBAR EN INDONESIE

667

□ IE.

MU

PAH.:

Source : MNHN. Pahs

Source : MNHN, Paris

Remerciements aux rapporteurs / acknowledgements to referees

La Redaction tieni it remercier les experts extdrieurs au Museum national d'Histoire naturelle dont les noms suivent, d'avoir bien voulu

contribuer, avec les rapporteurs de rEtablissement, A revaluation des manuscrits (1988-1997) :

The Editorial Board acknowledges with thanks the following referees who, with Museum referees, have reviewed papers submitted to the

(vtemoires du Museum (1988-1997):

Source :

\ Paris

SIMON Joseph L.

SPIRIDONOV V.

STEFANESCU Mihai O.

STORK N. E.

TAKEDA M.

TAN C. G. S.

TAYLOR P D.

THIBAUD Jean-Marc

THURSTON M. H.

TOULMOND Andre1

TRICART J

TUDGE Chrisiopher

VACELETJ.

VAN AMERON H. W. J.

VAN SOEST R. W. M.

VICKERY Vernon R.

VOICES E.

VOVELLEJean

VUL M. A.

WAGELE J. W.

WARfiN A.

WARREN Lynda

WATSON J.

WATSON Nikki

WESTHEIDE Wilfried

WILLIAMS A.

WILSON Mike

WILSON Robin

WITTMANN K.

ZEVINA G. B.

ZIBROWIUS Helmut

ZIEGLER Peter A.

New Orleans

Paris

Lvov

Bielefeld

Stockholm

Cardiff

Essendon

Armidale

Osnabriick

Washington

Cardiff

Victoria

Vienne

Moscou

Marseille

Bale

U. S. A.

France

Ukraine

Allemagne

Subde

Grande-Bretagne

Australie

Allemagne

U. S. A.

Grande-Bretagne

Australie

Autriche

Russie

France

Suisse

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La Cliapelle Monlligeon - Depot legal 1" trimestre 1997 - N" Imp. 18467 - Distribue le 28 fevrier 1997

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Ever since the explorations of the Danish Expedition to the Kei Islands, led by the famous zoologist

Th. MORTENSEN in 1922, the eastern seas of Indonesia have been renowned as a hotspot of marine biodiversity.

In 1991, the French-Indonesian KARUBAR cruise took place on board R.V. Banina Jaya 1 in the Banda and

Arafura Seas, off the Kai and Tanimbar Islands (Eastern Indonesia). The expedition, which had a dual zoological

and fisheries goal, collected a rich material of deep-sea benthos between 200 and 1200 m. Scattered faunistic

results have already been published elsewhere, but the present volume is the first consolidated report, consisting

of a narrative and 13 contributed articles on the systematics and biogeography of Scleractinia (1 paper), Mollusca

(5 papers), Crustacea Decapoda (6 papers) and Crinoidea (1 paper). Among the more remarkable results is

a review of the highly diverse azooxanthellate coral fauna from the Philippines and Indonesia, and a description of

a major new hermit crab assemblage.

Altogether the contributions report new data on over 400 species of invertebrates, of which 97 species and

1 1 genera are described as new. The volume is richly illustrated with numerous photographic plates, line

drawings and 2 colour plates, representing living animals. Because of the key biogeographical position of

Indonesia, the results presented in this volume bear relevance to a vast region in the tropical Indian and Pacific

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