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MEMOIRES DU MUSEUM NATIONAL D’HISTOIRE NATURELLE
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Source : MNHN. Pans
Ce volume des Resultats des Campagnes
MUSORSTOM est dedie a Rene GRANDPERR1N,
directeurde recherches de I'lRD, qui prend sa retraite en
I 'an 2000 apres 35 annees de recherches oceano-
graphiques presque entierement consacrees au Pacifique
Slid, en Nouvelle-Caledonie el a Vanuatu. Membre du
Comite de Pilotage du "programme devaluation des
ressources sous-marines de la zone economique de
Nouvelle-Caledonie" et responsable de la partie
halieutique de ce programme, directeur du programme
"monts sous-marins" consacre a I 'etude faunistique des
pentes recifales externes et des monts sous-marins et a
1'evaluation de leurs potentialites halieutiques. Rene
GRANDPERRIN a toujours soutenu le programme
MUSORSTOM et est intervenu chaque fois qu'il
convenait d'en faciliter /'execution.
Sa qualification pour les laches dont il etait
responsable. sa remarquable conscience professionnelle,
son devouement discret et ejficace envers ceux travail-
lant avec lui, sa courtoisie constante. font unanimenent
regretter son depart.
Resultats des Campagnes MUSORSTOM
Volumes deja parus :
Volume 1 : Mem. ORSTOM. 91 : 1-558, 225 fig., 39 pi. (1981). ISBN : 2-7099-0578-7.
Volume 2 : Mem. Mus. natn. Hist. nat.. (A), 133 : 1-525, 126 fig., 37 pi. (1986). ISBN : 2-85653-136-9.
Volume 3 : Mem. Mus. natn. Hist, nat., (A), 137 : 1-254, 82 fig., 9 pi. (1987). ISBN : 2-85653-141-5.
Volume 4 : Mem. Mus. natn. Hist, nat., (A), 143 : 1-260, 103 fig., 23 pi. (1989). ISBN : 2-85653-150-4.
Volume 5 : Mem. Mus. natn. Hist, nat., (A), 144 : 1-385, 128 fig., 35 pi. (1989). ISBN : 2-85653-164-4.
Volume 6 : Mem. Mus. natn. Hist, nat., (A), 145 : 1-388, 190 fig., 4 pi. couleur (1990). ISBN : 2-85653-171-7.
Volume 7 : Mem. Mus. natn. Hist, nat., (A), 150 : 1-264, 587 fig. (1991). ISBN : 2-85653-180-6.
Volume 8 : Mem. Mus. natn. Hist, nat., (A), 151 : 1-468, 198 fig. (1991). ISBN : 2-85653-186-5.
Volume 9 : Mem. Mus. natn. Hist, nat., (A), 152 : 1-520, 283 fig., 6 pi. couleur (1992). ISBN : 2-85653-191-1.
Volume 10 : Mem. Mus. natn. Hist, nat., 156 : 1-491, 163 fig., 2 pi. couleur (1993). ISBN : 2-85653-206-3.
Volume 11 : Mem. Mus. natn. Hist, nat., 158 : 1-426, 159 fig., (1993). ISBN : 2-85653-208-X.
Volume 12 : Mem. Mus. natn. Hist, nat., 161 : 1-569, 269 fig., 11 pi. couleur ( 1994). ISBN : 2-85653-212-8.
Volume 13 : Mem. Mus. natn. Hist, nat., 163 : 1-517, 132 fig., 4 pi. couleur (1995). ISBN : 2-85653-224-1.
Volume 14 : Mem. Mus. natn. Hist, nat., 167 : 1-647, 987 fig., 3 pi. couleur (1995). ISBN : 2-85653-217-9
Volume 15 : Mem. Mus. natn. Hist, nat., 168 : 1-539, 205 fig., 6 pi. couleur (1996). ISBN : 2-85653-501-1.
Volume 16 : Mem. Mus. natn. Hist, nat.. Ill : 1-667, 432 fig., 2 pi. couleur (1997). ISBN : 2-85653-506-2.
Volume 17 : Mem. Mus. natn. Hist, nat., 174 : 1-213, 93 fig., (1997). ISBN : 2-85653-500-3.
Volume 18 : Mem. Mus. natn. Hist, nat., 176 : 1-570, 458 fig., 7 pi. couleur (1997). ISBN : 2-85653-51 1-9.
Volume 19 : Mem. Mus. natn. Hist, nat., 178 : 1-255, 70 fig., 4 pi. couleur (1998). ISBN : 2-85653-517-8.
Volume 20 : Mem. Mus. natn. Hist, nat., 180 : 1-588, 192 fig., 2 pi. couleur ( 1999). ISBN : 2-85653-520-8.
Volume 21 : Mem. Mus. natn. Hist, nat., 184 : 1-813, 384 fig., 5 pi. couleur (2000). ISBN : 2-85653-526-7.
Source
resultats des campagnes
Volume 21
ISBN : 2-85653-526-7
ISSN : 1243-4442
© Publications Scientifiques du Museum, Paris, 2000
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Source : MNHN, Paris
MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE
TOME 184
ZOOLOGI E
Resultats des Campagnes MUSORSTOM
Volume 21
Coordonne par
Alain CROSNIER
Museum nalional d'Histoire naturelle
Laboratoire de Biologie des Invertebres marins et Malacologie
55 rue Buffon
F-75005 Paris
BIBL. OU
\MUSEUM/'
.PARIS,
PUBLICATIONS SCIENTIF1QUES
DU MUSEUM
PARIS
2000
-
Source : MNHN, Parts
SOMMAIRE
CONTENTS
Pages
1 . La campagne MUSORSTOM 10 dans I'archipel des iles Fidji.
Compte rendu et liste des stations . 9
Bertrand Richer DE FORGES, Peter Newell, Monika Schlacher-Hoenlinger.
Thomas SCHLACHER. Dako NATING, Frederic Cesa & Philippe BOUCHET
2. La campagne BORDAU 1 sur la ride de Lau (iles Fidji).
Compte rendu et liste des stations . 25
Bertrand Richer de Forges, Philippe Bouchet. Benoit Dayrat, Anders Waren
& Jean-Sebastien Philippe
3. Porifera Hexactinellida: On Euryplegma auriculare Schulze, 1886, and formation
of a new order . 39
Konstantin R. TABACHNICK & Henry M. REISWIG
4. Porifera Hexactinellida: Amphidiscophora off New Caledonia . 53
Konstantin R. Tabachnick & Claude Levi
5. Crustacea Cirripedia Thoracica: Chionelasmatoidea and Pachylasmatoidea
(Balanomorpha) of New Caledonia, Vanuatu and Wallis and Futuna Islands.
with a review of all currently assigned taxa . 141
Diana S. JONES
6. Crustacea Decapoda: A revision of the family Polychelidae . 285
Bella S. Galil
7. Crustacea Decapoda: Porcellanopagurus Filhol and Solitariopagurus Tiirkay
(Paguridae), from New Caledonian area, Vanuatu and the Marquesas: new records,
new species . 389
Patsy A. McLaughlin
8. Crustacea Decapoda: Species of the genera Crosnierila Macpherson, 1998,
Munida Leach, 1820, and Paramunida Baba, 1998 (Galatheidae) collected
during the MuSORSTOM 9 cruise to the Marquesas Islands . 416
Enrique MACPHERSON
9. Crustacea Decapoda: New species and new records of Ethusinae (Dorippidae)
from Vanuatu . 425
Chen Huilian
10. Crustacea Decapoda: A revision of the Indo-West Pacific species
of the family Palicidae (Brachyura) . 437
Peter CASTRO
11. Pycnogonida: Pycnogonids from the MUSORSTOM cruises to the South Pacific . 611
Roger N. Bamber
12. Echinodermata Crinoidea: Comatulid Crinoids of the Karubar Expedition
to Indonesia. The families Comasteridae, Asterometridae, Calometridae and
Thalassometridae . 627
Charles G. MESSING, Nadia AMEZIANE & Marc ELEAUME
13. Ascidiacea: Plurellidae collected in the Pacific Ocean by the cruises
MUSORSTOM, Karubar and the Coral Reef Research Foundation . 703
Fran^'oise MONNIOT & Claude MONNIOT
14. Pisces Gadiformes: Grenadier Fishes of the New Caledonian region, Southwest
Pacific Ocean. Taxonomy and distribution with ecological notes . 723
Nigel R. Merrett & Tomio Iwamoto
15. Pisces Pleuronectiformes: Flatfishes from New Caledonia and adjacent waters.
- Genus Arnoglossus . 783
Kunio AMAOKA & Eiji Mihara
Source : MNHN, Paris
5ULTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES C.
La campagne MUSORSTOM 10
dans l'archipel des lies Fidji.
Compte rendu et liste des stations
Bertrand RICHER DE FORGES * Peter NEWELL ** Monika SCHLACHER-
HOEN LINGER ** Thomas SCHLACHER ** Dako NATING **
Frederic CESA* & Philippe BOUCHET ***
*IRD
B. P. A5, 98848. Noumea Cedex
Nouvelle-Caledonie
**University of the South Pacific
School of Pure and Applied Sciences
Suva, Fiji
***Museum national d'Histoire naturelle
Laboratoire de Biologie des Invertebres marins et Malacologie
55, rue Buffon. 75005 Paris, France
RESUME
La campagne Musorstom 10. realisee a bord du N. O. "Alis", s'est deroulee dans les eaux des Fidji. du 5 au 19 aout
1998. Quatre-vingt-deux operations de dragages et de chalutages ont eu lieu dans les zones bathyale superieure et
circalittorale de file de Vitu Levu et dans la zone de Bligh Water. L'essentiel des stations realisdes dans Bligh Water
montre des fonds de vase, gdn^ralement parsemes de pierres ponces. Des fonds durs ont ete rencontres dans la zone de Beqa
Channel. Les peuplements presentent une diversity specifique plus faible qu'en Nouvelle-Caledonie, en particular dans
les groupes d'invertebres fix6s (spongiaires, stylasterides, crinoides).
ABSTRACT
The Musorstom 10 Cruise in the Fijian Archipelago. Report and list of stations.
The Musorstom 10 cruise, using the R.V. "Alis11, was carried out in the Fijian Archipelago during 5-19 August 1998.
A total of 82 samples were collected by dredging and trawling from the upper bathyal zone and in the eircaliitoral depths,
on the outer reef slopes of Vitu Levu and from Bligh Water. The bottom of Bligh Water is muddy and covered with pumice
stones. Hard bottoms were sampled in the Beqa Channel. The invertebrate biodiversity of the benthic communities
sampled is much lower than those sampled from the New Caledonian area, especially sessile epifaunal groups like
sponges, stylasterine hydrocorals (Cnidaria, Sty lasterina) and crinoids.
Richer de Forges, B., Newell, R. Schlacher-hoenlinger, M.. Schlacher. T., Nating. D.. Cesa, F. & Bouchet. P..
2000. — La campagne Musorstom 10 dans l'archipel des lies Fidji. Compte rendu et liste des stations. In : A. Crosnier
(ed.), Resultats des Campagnes Musorstom, Volume 21. Memoires du Museum national d'Histoire naturelle. 184 : 9-23.
Paris ISBN-2-85653-526-7.
10
B. RICHER DE FORGES ET AL.
INTRODUCTION
La campagne MUSORSTOM 10 s'est deroulee du 5 au 19 aout 1998 a bord du N.O. "Alls" dans 1'archipel des iles
Fidji. Cette campagne fait partie d'un programme destine a explorer la faune benthique de la zone bathyale
superieure dans l'lndo-Ouest Pacifique. Un recapitulate des huit premieres campagnes cst presente en introduction
du compte-rendu concernanl MUSORSTOM 9 aux iles Marquises (RICHER DE FORGES & Laboute. 1998; RICHER
de Forges et ai, 1999).
Les observations bathymetriques et geographiques ainsi que les resultats taxonomiques concernant I'ensemble de
ces campagnes sont stocks dans des bases de donnees informatisees, tenues a jour au Centre IRD (anciennemenl
ORSTOM) de Noumea. La plus grande partie des resultats zoologiques et biogeographiques ont ete publies dans la
serie "Resultats des Campagnes MUSORSTOM" qui, dans ses 20 premiers volumes traite de 4587 especes parmi
lesquelles 1324 etaient nouvelles pour la Science et ont necessite la creation de 126 genres ou sous-genres
nouveaux. Cette importante amelioration des connaissance sur les faunes de profondeur de l'lndo-Ouest Pacifique
tropical est le resultat d'une collaboration active d'un reseau international de taxonomistes (24 nations,
92 institutions de recherche et 181 chercheurs).
Bien que l'analyse de ce vaste ensemble de donnees originates soit encore en cours, certaines remarques peuvent
etre faites (RICHER DE FORGES, 1998) :
— les profondeurs bathyales (200-2000 m) sont tres discontinues et de nombreuses zones demeurent
inexplorees (Fig. 1);
— dans l'lndo-Ouest Pacifique, ces profondeurs presentent une richesse specifique elevee. Dans I’ensemble de la
zone etudiee, la faune etait en grande partie inconnue avec 28,9 % d'especes nouvelles; pour la Nouvelle-Caledonie,
60,7 % des 1619 especes d'invertebres actuellement etudices etaient nouvelles;
— les faunes benthiques de la "province Indo-Pacifique" sont differentes d'un archipel a 1'autre et la repartition
geographique de nombreuses espbces est plus restreinte que prevue dans les travaux de biogeographie (Ekman,
1953; Briggs, 1974). Ainsi, la faune recoltee dans la partie nord de 1'archipel de Vanuatu pendant la campagne
MUSORSTOM 8 se revele bien differente de celle de Nouvelle-Caledonie, distante de moins de 300 km;
— dans plusieurs groupes zoologiques les especes considerees comme archai'ques ou reliques sont
particulierement abondantes (Spongiaires, Brachiopodes, Echinodermes, Mollusques, Crustaces...) ;
— les sommets des monts sous-marins sont particulierement riches en especes et en biomasse et semblent
favoriser I'endemisme. Cette observation met en evidence la tres grande vulncrabilitc de certaines especes benthiques
qui ont des aires de repartitions limitees a quelques km2. Cette vulnerabilite est encore accrue du fait d'une
Fig. 1. — Cartographie des profondeurs bathyales de l'lndo-Ouest Pacifique. En noir, les fonds de 0 a 200 m; en hachures,
les zones de profondeurs bathyales (200-2000 m). Carte realisee par Richer de Forges (1998) a partir des donnees
d'altimetrie satellitaires publiees, sous forme d'une carte au 1/40 millionieme, par Sloss (1994).
A-I ; Arabo-Indian; EA : Easl Australia; EAf : East Africa; FP : French Polynesia; H : Hawaii; M : Marianas, Madagascar;
NC ; New Caledonia; O ; Ontong Java; RS : Red Sea; S : Seychelles; SJ : South Japan.
Source .
CAMPAGNE MUSORSTOM 10
1 I
croissance lcnte de ces organismes de profondcur (VACELET et al., 1992; KOSLOW & EXON, 1995; Zezina. 1997;
Koslow & Gowlett-Holmes, 1998; Richer de Forges et al., 1998).
GENERALITIES
L'archipel des Ties Fidji s'etend entrc 16 et 20°S et 177°E a 178°W. 11 est constitue d'environ 300 Ties et dots
dont les plus grands sont Vitu Levu et Vanua Levu qui ont des superficies respectives de 10400 km2 et 5538 km2
et qui culminent a 1323 m et 1 1 1 1 m. Ces ties, d’origine volcanique, sont entourees de formations coralliennes qui
delimitent par endroits des lagons vastes et profonds (Antheaume & Bonnemaison, 1988; Chandra &
Mason, 1998).
L'Tle de Vitu Levu comporle trois grands fleuves dont le plus important est, au sud-est, la Rewa River (Wai
Levu), qui draine un bassin versant d'environ 3000 km2. II s'agit du plus grand cours d'eau de tout le Pacifique
insulaire (Papouasie Nouvelle-Guinee exceptee). Le debit de la Rewa River a ete evalue pour la premiere fois par
Dana (1872) a 236 m'/s (Moseley, 1879). Des donnees plus recentes font etat de debits moyens mcnsuels en
1998 (mesures a Navolau) variant de 277 m’/s en janvier & 29 m'/s en aout. Ce debit aurait atteint 9000 m'/s
pendant le cyclone Kina en janvier 1991 (GOUYET, com. pers.). Son delta actuel s'est forme pendant I'Holocene et
le transport solide de ce fleuve est estime a 3200 tonnes/km2/an (Nunn, 1998).
D'apres les vestiges archeologiques et en particulier les poteries, il semble que l'archipel des ties Fidji ait ete
peuple par les hommes il y a plus de 3000 ans (Irwin, 1992; Gravelle, 1996; Kirch, 1997). Bien avant
I'arrivee des occidentaux, de puissantes chefferies occupaient toutes les lies et avaient deja profondement modifie
1'environncment (Kirch & Hunt, 1997). Les lies Fidji (Feedjee) furent revelees au rnonde occidental par le
navigateur hollandais Abel Tasman en 1643 (Dunmore, 1991). Au cours de ses periples dans le Pacifique, le
Capitaine Cook fit escale longuement (mai-juillet 1777) aux "lies de 1'Amitie" (Tonga) mais evita les Fidji alors
reputees tres dangereuses pour leurs feroces cannibales (COOK, 1980; FROST, 1998). L’un des premiers contacts
entre les civilisations est reste celebre. Il s'agit du passage du Capitaine William BLIGH (qui avail ete aux Tonga
avec COOK a bord du "Resolution") en 1789 dans la mer qui porte aujourd'hui son nom. Bligh Water, a la suite de
la mutinerie a bord de son bateau, la "Bounty".
Conditions hydroclimatiques. — Les lies Fidji sont soumises a un climat maritime tropical. Les vents
alizes de Sud-Est apportent des pluies qui sont done plus marquees sur le versant Est des Ties (JONES & PlNHEIRO,
1997; Chandra & Mason, 1998). La saison des pluies s'etend de novembre a avril et la pluviosite moyenne
annuelle est de 3100 mm a Suva. A Nandi, situe sur le versant sous le vent de Vitu Levu, la pluviosite moyenne
n'atteint pas 2000 mm/an. La temperature moyenne a Suva est de 25°C mais depasse frequement 30°C en etc
(decembre-mars) et descend occasionnellement en dessous de 20°C en hiver (juillet-aout).
Les caracteristiques hydrologiques de cette partie du Pacifique sud-ouest ont ete decrites par DELCROIX et Henin
(1989). Ces auteurs decrivent les variations saisonnieres et interannuelles, mettant en evidence I'evenement ENSO
(El Nino Southern Oscillation) tres fort de 1982-83. Les temperatures de surface de I'ocean sont comprises entre 25
et 30°C avec une moyenne annuelle en surface de 27,5°C. En temps ordinaire (hors El Nino), la temperature
minimale en surface s'observe en aout et la salinite minimale en mars, environ trois mois apres le maximum de
precipitations. Entre 300 et 400 m de profondcur, les temperatures fluctuenl, d’une annee a l'autre. entre 10 et
12 C. La salinite moyenne annuelle observee aux Fidji est de 35,1 %o. L'amplitude maximale des vents alizes de
SE est de 6-7 m.s'1.
Dans la zone de Bligh Water, les sondes XBT lancees donnerent les indications suivantes en aout 1997 : 24°C
en surface, thermocline a environ 100 m de profondeur et 8°C a 550 m de profondeur.
Pendant les grandes fluctuations climatiques du Quaternaire, le niveau marin a considerablement varie (Hopley,
1982; PAULAY, 1990, 1997; CABIOCH et al., 1996; Nunn, 1994, 1998). Les niveaux les plus bas furent situes
entre 100 et 150 m en dessous du niveau actuel. La quasi-disparition des lagons durant ces niveaux bas a du reduire
la repartition geographique des especes marines typiquement lagonaires. D'apres Paulay ( 1 990), la zone profonde
de Bligh Water aurait pu jouer le role de refuge pour cette faune lagonaire et particulierement pour les bivalves de
12
B. RICHER DE FORCES ET AL.
fonds meubles. 11 semble egalement quc dcs variations de temperature relativement rapides (1500 ans) dc plus dc
5°C aient eu lieu dans ces latitudes tropicales (BECK el al., 1997),
HISTOIRE GEOLOGIQUE DES ILES FIDJI
D'importants travaux de geophysique et dc geologie marine ont eu lieu dans le sud-ouest Pacifique, de mars a
juin 1982, a bord du R.V. "Kana Keoki", sous l’egide de la CCOP/SOPAC. Lcs rcsultats obtenus concernent la
bathymetrie, la nature des fonds et des datations des principals ties volcaniques. L'esscnticl dc ces resultats sont re-
groupes dans l'ouvrage de BROCHER (1985). Lcs grands traits de l'histoire geologique des ties Fidji ont etc relates
par Rodda (1994). Les plus anciennes roches volcaniques des Ties Fidji sont trouvees sur Vitu Levu et datent de
l'Eocene supericur soit environ 34 M.A. (COULSON, 1974; MYERS, 1985; Rodda, 1994). Depuis, de nombreux
episodes volcaniques se sont succedes jusqu'a la periode actuelle, les rochcs s'ctant souvent formees en milieu
marin.
Fig. 2. — Geomorphologie du Pacifique sud-ouest montrant les grands traits structuraux, plaques, fosses de subduction
(d'aprfcs AUZENDE el al., 1996).
Source :
CAMPAGNE MUSORSTOM 10
13
Les mouvements tectoniques complexes de la bordure orientale de la plaque australo-indienne out ete decrits a la
suite de nombreux travaux de geologie marine (Springer, 1982; Lewis, 1993; AUZENDE et al., 1995). Le groupe
principal des Ties Fidji, Vitu Levu et Vanua Levu, est situe juste a Tangle oriental de cette plaque ct faisait partie de
l'alignement Nouvelles-Hebrides - Fidji - Tonga lors de la periode active de la "fosse de Vitiaz", avant I'ouverture
du Bassin nord fidjicn, il y a 10 M.A. (Fig. 2).
CONNAISSANCES ANTERIEURES SUR LA FAUNE MARINE DES ILES FIDJI
WiENS (1962) attire l'attention sur le fait que la faune marine des Ties Fidji est encore typique du "Melanesian
Plateau" mais differe nettemenl de cclle des Ties Samoa, pourtant toutes proches, qui font partie de 1'ensemble
Polynesien.
L'expedition du H.M.S. "Challenger" realisa des prelevements benthiques dans I'archipel des Fidji entre
le 24 juillet et le II aout 1874 (MOSELEY, 1879; Spry, 1880; Lintaker, 1972). Les premiers dragages furent
realises pres de l'Tle Maluka, au SE de Vitu Levu, par 585 nr et un specimen de nautile (Nautilus pompilius ) fut
capture ainsi que de nombreux specimens de Polycheles. Les Ties de Kadavu et Ovalau furent ensuite explorees.
La faune bathyale n'etait done connue, avant MUSORSTOM 10, pratiquement que d'apres l'etude du materiel
recolte au cours des 3 stations profondes du "Challenger" pres de Matuka et Kandavu ( 19°09,35'S- I79°4I.50'E;
1 9°07,50'S- 1 78° 1 9,35'E; 1 9°05,50'S- 1 78° 1 6,20’E).
Harper et Smith (1989) donnent quelques indications sur les 32 operations de dragages realisees dans le Sud-
Est de Vitu Levu et sur la Ride de Lau, a bord du F.N.S. "Babale", dans le but de rechercher des "coraux precieux"
du genre Corallium. Dix-huit stations ramenerent des echantillons qui furent confies a l'Universite d'Hawaii. Les
quelques organismes (spongiaires, gorgones, echinodermes, scleractiniaires) signales dans ce rapport (GRIGG. 1996)
ne semblent pas avoir fait l'objet de publications.
De fait, au moment de la campagne MUSORSTOM 10, les faunes profondes de mollusques de I'archipel fidjien
restaient pratiquement terra incognita : les trois dragages realises par le "Challenger", en 1874, entre 385 et 1115 m
de profondeur avaient fait connaTtre en tout et pour lout six especes de gasteropodes, toutes nouvelles a l'epoque. et
aucun representant des autres classes de mollusques. Sur ces six especes, Bolma henica et Cerithium matukense se
sont averces avoir une vaste repartition Indo-Paciflque, et Bursa fijiensis n'a ete que tres recemment redecouverte en
Nouvelle-Caledonie (BEU, 1998); par contre, Fusus libratus, Gaza daedala et Clathurella contpsa restent. 125 ans
apres, connues uniquement par les recoltes du "Challenger" devant Kandavu et Matuka. Apres le "Challenger", il ne
se passe plus rien pendant plus de cent ans, hormis la decouverte, fortuite, d'un nouveau pleurotomaire sur la Ride
de Lau par le navire neo-zelandais "Tangaroa" en 1980 (BOUCHET & METIVIER, 1982). Cependant. I'evenement le
plus marquant des recentes decennics est la decouverte de communautes biologiques associees aux manifestations
hydrothermales dans les bassins de Lau et nord-fidjien (DESBRUYERES et al., 1994). Cette decouverte a entraine la
realisation de campagnes de prospection par submersibles (campagnes franco-japonaise Starmer 2 et frangaise
Biolau) et par benne telecommandee (campagne allemande du "Sonne"). La faune malacologique de ces sites,
relativement bien inventoriee; comprend une vingtaine d'especes de gasteropodes (BOUCHET & Waren, 1991;
Waren & BOUCHET, 1993; OKUTANl & Ohta, 1993; BECK, 1996) et plusieurs grands bivalves (METIVIER &
Cosel, 1993; COSEL et al., 1994), dont les affinites biogeographiques sont a rechercher dans le bassin de Manus et
le bassin des Mariannes.
S'y ajoutent quelques specimens benthiques recoltes lors de l'etude des sources hydrothermales du Bassin de Lau
par la campagne Biolau ou les reperages qui ont precede tels que la campagne du R.V. "Sonne" en 1987
(Yamaguchi & Newman, 1990; Baba & Turkay, 1992).
La faune benthique littorale et lagonaire des Ties Fidji n'est guere mieux etudiee. VERON (1995) dans son
ouvrage de biogeographie des scleractiniaires constructeurs, mentionne 60 genres ct 250 especes aux Fidji.
A 1'occasion d'un Symposium sur la biodiversite des zones littoralcs des Ties du Pacifique, un bilan des
connaissances avait ete presente par les specialistes de certains groupcs zoologiques (Maragos et al., 1995). Pour
14
B RICHER DE FORGES ET AL
la region des ties Fidji, les donnees proviennent soit du "Challenger", soit de quelques petiles recoltes
occasionnelles plus rccentes. En 1965, le R.V. "Te Vega" collecta un pcu de faune marine littorale de Viiu Levu,
ameliorant ainsi les rares donnees sur la faune marine de cette region. Une liste de 41 especes de spongiaires
actuellement signaldes des Fidji est donnee par KELLY-BORGES et Valentine (1995). Vingt-quatre especes de
Polychetes seraicnt signalees (BAILEY-BROCK, 1995). Pour les Crustaces, l'etude realisee par Myers (1985) sur les
amphipodes Gammaridea des ties Fidji a permis de recolter 77 especes donl 40 % etaient nouvelles. Les affinites
geographiques observees par cet auteur sont plutot avec Hawaii. 1'ouest du Pacifique (Indonesie) ou meme l'ocean
Indien qu'avec la Polynesie.
La faune malacologique marine des Fidji est sans doute l'une des micux connues de tous les archipels du
Pacifique. Cette situation est essentiellement le resultat du travail d'un seul homme, Waller O. CERNOHORSKY, qui
pendant 15 ans fut employe comme cadre par l'Emperor Gold Mining Co. a Vatukoula (Viti Levu), avant de
devenir en 1969 curator of molluscs a 1' Auckland Institute and Museum. CERNOHORSKY a public de nombreuses
monographies regionales sur les gasteropodes des Fidji (CERNOHORSKY, 1964a, 1964b. 1965, 1967b, 1967c.
1968, 1969, 1971), et ses revisions contiennent egalement une foule de donnees geographiques nouvelles
concernant I'archipel (CERNOHORSKY, 1976, 1984, 1991). Ses trois ouvrages d'identification des faunes du
Pacifique (CERNOHORSKY, 1967a, 1972, 1978) demeurent des classiques, et plus de la moitie des 600 especes
traitees dans le deuxieme volume (CERNOHORSKY, 1972) sont illustrees avec des specimens des Fidji. Dans tous
ses travaux, CERNOHORSKY a localise son interet et ses efforts sur les especes littorales des families qui inleressent
les collectionneurs : les micromollusques, les especes subtidales accessibles seulement par dragage et en plongee;
les bivalves restent peu ou pas inventories.
Fig. 3. — L'itineraire de la campagne Musorstom 10 dans I'archipel des ties Fidji.
Source : MNHN, Paris
CAMPAGNE MUSORSTOM 10
15
PARKINSON (1982) dans unc liste preliminaire des coquillages potcntiellement exploitables des Ties Fid j i.
signalc la capture de Cypraea valentia sur les cotes de ltle de Kadavu. Cette espece rare a ete retrouvee au cours de
MUSORSTOM 10 a l'embouchure de la Rewa River, sur l'tle de Vitu Levu. Un recent travail d'inventaire des
gasteropodes Conidae des Fidji, d'apres une compilation bibliographique, fait etat de 99 especes (Seeto. 1998).
L'etude biogeographique des echinodcrmes de l'lndo-Pacifique de Clark et Rowe (1971) signalc de nombreuses
especes des ties du Pacifique Sud mais ne renseigne pas specialement sur celles des ties Fidji.
Pour les poissons, KULBICKI et Rivaton (1997) recensenl. d'apres la litterature, 822 especes des Ties Fidji.
D’apres les compilations d'inventaires non publics, il y aurait actuellement 960 especes connues des Fidji
(KULBICKI, com. pers.).
Quelques travaux d'ecologie ont eu lieu dans le lagon du recif de 1' "Astrolabe" sur l'tle de Kadavu. Ils onl ete
effectues de fason quantitative avec une benne de 0,1 m2 ou en plongee sous-marine et apportent peu dedications
sur la composition specifiquc (Clavier et ai, 1996; Newell et ai, 1996; Newell et at ., 1997; Newell &
Clavier, 1997; Schlacher et ai, 1998).
DEROULEMENT DE LA CAMPAGNE MUSORSTOM 10
La campagne MUSORSTOM 10 avait pour objectif l'echantillonnage de la faune benthique de la zone bathyale
supericure des lies Fidji, de 200 a 1500 m (cf. liste des stations en annexe; Fig. 3-4).
Le navire oceanographiquc de l'ORSTOM, N.O. "Alis", base a Noumea, etait deja dans l'archipel des Fidji
depuis un mois et avait effectue deux autres campagnes concernant les effets anthropiques en zone littorale (Suva
1) et les paleoclimats (Paleofiji). La campagne MUSORSTOM 10 debuta le 4 aout au soir au depart de Suva et la
premiere operation de dragage (DW 1308) eut lieu au nord-ouest de 1'ile d'Ovalau dans le chenal sud de Bligh Water.
Les fonds de ce chenal, d'une profondeur de 700 a 900 m, sont composes de vases argileuses grises contenant de
nombreuses coquilles de pteropodcs. Des les premiers chalutages, nous remontions des pierres ponces et des bois
coulds, tres abondants dans les eaux fidjiennes a proximite des grandes lies hautes. Ces substrats particuliers
permettent la presence de faunes associees : des pagures symetriques de la famille des Pylochelidae (Pylocheles
incisus, Cheiroplatea pumicicola), foreurs dans les bois, les ponces et parfois les eponges siliceuses (FOREST.
1987) et, profitant des substrats organiques, des amphipodes, des chitons ( Leptochiton ) et des gasteropodes
cocculiniformes (Richer de FORGES et al., 1996). Les specimens de Pylochelidae recoltes appartiennent a l'espece
Cheiroplatea pumicicola Forest, 1987 qui n'etait jusqu'alors connue que par une trentaine de specimens des lies
Kermadec (29°S- 1 77°W).
II faut signaler egalement la decouverte, a plusieurs reprises de bees de cephalopodes portant des gasteropodes du
genre Bathysciadium.
Bligh Water. — La zone dite de Bligh Water est tres particuliere (Wingfield et ai, 1977). 11 s'agit d'une sorte
de "mer interieure" situee entre les deux lies principales, Vitu Levu et Vanua Levu. Elle est presque entierement
bordee par les barrieres coralliennes delimitant les lagons de ces deux lies et sa profondeur atteint pres de 1000 m en
son centre. Les chenaux d'acces a Bligh Water, situes au nord et au sud, sont toutefois tres profonds (>600 m) et ne
presentent done pas de seuils qui auraient pu y confiner une faune relique. Les etudes sismiques realisees dans cette
mer ont montre une epaisseur de sediments d'environ 2800 in (MAUNG & Eden, 1983).
Pratiquement la totalite des operations dans Bligh Water s'est deroulee sur des fonds vaseux entre 600 et 900 m
de profondeur. Les peuplements caracteristiques sont composes de gros oursins irreguliers. de mollusques
Pectinidae du genre Propeamusium , de crustaces des genres Platymaia et Glyphocrangon, de poissons de la famille
des Macrouridae. Des specimens du gasteropode Volutidae Calliotectum egregium ont ete trouves.
Sur les pentes recifales externes delimitant les contours de Bligh Water, entre 300 et 500 m de profondeur. les
fonds sont plus sablo-vaseux et presentent les organismes suivants : gasteropodes Xenophoridae. Crustaces
decapodes des genres Munida, Mttnidopsis, Hexaplax, Ethusina, des crustaces cirripedes pedoncules, poissons
Scorpaenidae, Triglidae, Bothidac.
16
B. RICHER DE FORGES ET AL.
Fig. 4. — Les participants it la campagne Musorstom 10 : de gauche a droite, Frederic Cesa, Dako Nating, Philippe
Bouchet, Bertrand Richer de Forges, Monika Schlacher-Hoenlinger, Peter Newell (Photo B. Richer de Forges).
Dans les fonds de moins de 300 m, on remarque I'abondance des crustaces decapodes brachyoures Goncplacidae
( Carcinoplax , Hexapus, Psopheticus) et Leucosiidae ( Iphiculus ) et dans certaines stations plus envasees de
brachyoures Retroplumidae, Palicidae et Raninidae ( Lyreidius , Raninoides). Les crevettes peneides du genre
Metapenaeopsis sont egalement frequentes.
Signalons la presence entre 300 et 400 m d'especes connues jusqu'alors uniquement de Nouvelle-Caledonie et du
sud de Vanuatu : e'est le cas du crabe Majidac Oxypleurodon orbiculatus.
Rewa Roads. — Quelques operations furent realisees en face de l'estuairc de la Rewa River, dans la zone
denommee Rewa Roads sur les cartes, entre 80 et 160 m de profondeur (dont trois operations de chalutages en
presence de 1 Ambassadeur de France et du Director of the National Trust of Fiji). Les fonds de vases noires y sont
tres riches en debris vegdtaux avec une faune associee tres diversifiee : mollusques Muricidae, une dizaine d'especes
de nudibranches, bivalves du genre Abra et Scaphopodes, crustaces Metapenaeopsis, Raninoides, Arcania, Myra,
Macropodia et Stomatopodes, poissons Scorpaenidae.
Beqa Channel. — La zone de Beqa Channel consiste en un etroit passage (< 5 milles) situe entre la cote sud de
I ile de Vitu Levu et 1 He haute de Beqa, entouree d'un vaste lagon delimite par une barriere corallienne. Le fond de
ce chenal se situe vers 240 m de prolondeur et presente des zones rocheuses et de sables grossiers, consequence des
forts courants qui y regnent. Les organismes les plus abondants dans cc type de fonds sont, les mollusques du genre
Liotia, des crustaces Munida et Chirostylus , des crabes Dynomenidae, Xanthidae, des Scyllariidac, de nombreux
pagures Pylochelidae forant des roches calcaires, des Brachiopodes. Signalons la decouverte de la volute Lyria
plamcostata qui etait jusqu'alors connue uniquement de Vanuatu et plus a l'ouest (Ladd, 1982).
Au cours de la campagne Musorstom 10, des gonades ont ete prelevees sur certaines especes de Crustaces afin
den decrire les spermatozoides et d'utiliser ces nouveaux caracteres pour preciser leur position phylogenetique
Source : MNHN, Paris
CAMPAGNE MUSORSTOM 10
17
(Jamieson et al 1995). Les prelevements, actuellement a l'etude, ont porte sur : Polychelidae ( Polycheles
typhlops), Pylochelidae ( Cheiroplatea pumicicola), Lithodidae (Lithodes sp. ), Galalhcidae (. Bathymunida sp.),
Lcucosiidae (Arcania sp., Iphiculus spongiosus , Myra sp.).
CONCLUSIONS
La connaissance de la faune de profondeur des lies Fidji s'accroTt considerablement avec les recoltes de la
campagne MUSORSTOM 10 et ces resultats vont combler une vaste lacune geographique. Les remarques
preliminaires qui peuvent etre faites avant l'etude detaillee par les specialistes de chaque groupe sont les suivantes :
— la faune bathyale des Fidji est nettement plus pauvrc en especes que celle de Nouvelle-Calcdonie, mais plus
riche que celle de Polynesie fran?aise et en particulier que celle de I'archipel des Ties Marquises (RICHER DE FORGES
etal ., 1999);
— certains groupes fixes tels que les stylasterides, les crinoides pedoncules et memo les spongiaires semblent
rares et peu diversifies. II faut toutefois moduler ces remarques car la majorite des zones echaniillonnees presentait
des fonds meubles peu favorables a ce type d'organismes.
REMERCIEMENTS
Nous remercions vivement les personnes qui ont contribue a la reussite de cette campagne : pour la preparation,
l'obtention des financements et les commandes, Patrice Cayre, Rene GRANDPERRIN, Alain CROSNIER et Philippe
Maestrati. L'Ambassade de France a joue un grand role dans la reussite de cette campagne en collaboration avec
les chercheurs de I'University of the South Pacific. Nous remercions tout particulierement Monsieur
l'Ambassadeur, Michel Jouvet et son Attache culturel et scientifique, Didier SAVIGNAT.
A bord du N.O. "Alis", le Commandant Raymond PRONER a fait le maximun pour realiser de bons
prelevements dans des zones souvent non hydrographiees; le MaTtre d'equipage. Loic LEGOFF, et le second
capitaine, Jean-Fran^ois Barazer ont inlassablement ramende les chaluts; les cartes sont dues a Yves Penvern et
Marika Tortelier, dessinateurs au Centre IRD de Noumea.
Nous remercions egalement la bibliothecaire du CCOP/SOPAC qui a pu nous procurer des copies de differents
rapports d'exploration geologique de la ZEE de Fidji et notre collegue hydrologue R. GOUYET pour les donnees sur
les debits de la Rewa River.
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ANNEXES
LISTE DES PARTICIPANTS A LA CAMPAGNE MUSORSTOM 10
Chef de mission : B. Richer de Forges.
A u ires participants : P. BOUCHET, P. Newell, M. Schlacher. T. SCHLACHER, D. Nating. F. Cesa.
LISTE DES STATIONS DE LA CAMPAGNE MUSORSTOM 10
Source : MNHN, Paris
B. RICHER DE FORGES ET AL.
Source :
CAMPAGNE MUSORSTOM 10
23
Source : MNHN. Paris
Source : MNHN. Paris
TATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULT ATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CA
La campagne BORDAU 1 sur la ride de Lau
(lies Fidji). Compte rendu et liste des stations
Bertrand RICHER DE FORGES *, Philippe BOUCHET ***
Benoit DAYRAT***, Anders WAREN** & Jean-Sehastien PHILIPPE*
*1RD
B.P. A5. 98848 Noumea Cedcx
Nouvelle-Caledonie
**Naturhistoriska Riksmuseet
Stokholm. Suede
***Museum national d'Histoire naturclle
Laboratoire de Biologie des Invertebres marins et Malacologie
55. rue Buffon, 75005. Paris. France
RESUME
La campagne Bordau 1, realis6e h bord du N. O. "Alis", s'est deroulee dans les eaux des Fidji. du 22 fevrier au 14 mars
1999. Cent dix-huit operations de dragages et de chalutages ont eu lieu dans les zones bathyale superieure et cirealittorale
des Ties et sur les monts sous-marins de la ride de Lau. La partie superieure des pentes presente des londs rocheux jusqu'a
600 m de profondeur. Plus profondement. on retrouve les fonds envases couverts de pierres ponces. Dans certaines Ties
particulibrement isolees (Vanua Balavu. Yacata, Aiwa et Yagasa), un echantillonnage de la malacofaune terrestre a 6te
realise.
ABSTRACT
The Bordau Cruise on the Lau Ridge (Fijian Archipelago). Report and list of stations.
The Bordau 1 cruise was carried out in the Fijian Archipelago from 22 February to 14 March 1999 on board ot
R.V. "Alis". A total of 1 18 samples were made by dredging and trawling in the upper bathyal zone and in the circalittoral
depths of the islands and on the seamounts in the Lau Ridge. The upper part of the slope to 600 m consists ot hard
bottoms and deeper muddy bottoms with pumice. In some islands particularly isolated (Vanua Balavu, Yacata. Aiwa and
Yagasa), the landsnails were sampled.
Richer de Forges, B.. Bouchet. P., Dayrat, B.. War£n, A. & Philippe. J.-S.. 2000. — La campagne Bordau 1 sur la
ride de Lau (Ties Fidji). Compte rendu el liste des stations. In : A. Crosnier (ed.), Resultats des Campagnes MUSORSTOM.
Volume 21. Memoires du Museum national d'Histoire naturelle, 184 : 25-38. Paris ISBN-2-85653-526-7.
26
B. RICHER DE FORGES ET AL.
Fig. 1. — Geomorphologie de la partie orientale de la plaque Ausiralo-lndienne montrant la ride de Lau-Colville el les
bassins environnants (d'apr&s Anonyme, 1982).
Source MNHN, Paris
CAMPAGNE BORDAU I
27
INTRODUCTION
La campagne BORDAU 1 est la premiere de deux campagnes destinees a echantillonner la faune bathyale de la
bordure orientale de la plaque Australo-Indicnne. Apres une serie de 10 campagnes d'exploration sous le nom de
MUSORSTOM, auxquelles se sont ajoutees d'autres campagnes en Nouvelle-Calcdonie et en Indonesie. les collections
accumulees permettent, une fois les especes decriles, de comprendre la repartition geographique et bathymetrique dc
ces especes. Alors que l'objectif des campagnes MUSORSTOM et assimilees etait de faire l'inventaire des faunes
bathyales et de demontrer l'existence, en zone bathyale. d'un gradient dccroissant de richesse speciftque d'Ouest en
Est. les campagnes Bordau 1 et BORDAU 2 ont d'autres objectifs. Elies doivent permettre d’observer la faune
bathyale situee en bordure des plaques.
Profitant de la disposition geographique et geomorphologiquc particuliere des rides de Lau-Colville et de Tonga-
Kennadec, l'echantillonnage permettra. de plus, de decrire la variation latitudinale de la faune benthique entre les lies
franchement tropicales par 16° S et les monts sous-marins situes hors de la zone inter-tropicale, a la limite des eaux
temperees.
La campagne MUSORSTOM 10 a permis d'obtenir les premieres donnees sur la faune de profondeur des lies Fidji
avec 82 stations reaiisees autour de file de Vitu Levu et, en grande partie, dans la zone de Bligh Water (RICHER DE
FORGES et al., 1999). La campagne Bordau 1 a ete consacree a la partie est de l’archipel en echantillonnant les
abords des lies de Lau (Fig. 1).
GENERALITIES SUR LES ILES DE LAU
La ride dc Lau correspond a fare d'tles forme par la subduction ancienne de la plaque Pacifique sous la plaque
Australo-Indienne. Cette ride, appelee aussi Lau-Colville Ridge, s'etend sur pres de 2400 km du nord-est des Fidji
jusqu'a file nord de la Nouvelle-Zelande (Fig. 1). Elle s'est formee au Miocene, de 22 it 5 M.A.. lorsque ce
volcanisme d'arc concernait l'alignement Nouvelles-Hebridcs, Fiji, Tonga (Woodhall, 1985).
Actuellement. la subduction est active plus a l'est, au niveau de la ride Tonga- Kermadec. La ride de Lau separe
des bassins de plus de 3000 m de profondeur, lc Bassin sud fidjien, a l'ouest et le Bassin de Lau. a l'est. Elle porte
dc nombreuses ties hautes d'origine voleanique (fin Oligocene). mais egalement des atolls coralliens et des recifs
soulcves. Le groupe des ties de Lau, parfois nomine "Eastern group" des Fidji. s'etend. du nord au sud. des
formations recifales situees par 16°S jusqu'au Minerva Reef par 24°S. Les ties emergees les plus au sud de cel
alignement sont Ono-i-Lau et Tuvana-i-Colo par 21°S (Fig. 3).
Cette region, d'une grande complexity tectonique, a fait l'objet dc nombreuses etudes de geologic et geophysique
(Pelletier et al., 1998; Pelletier. 1999). Le bassin de Lau est en cours d'ouverture tres active et presente des
sources hydrothermalcs (TIFFIN, 1991 ). Les vitesses d'ouverture du bassin dc Lau sont de l'ordre de 6.5 a 13 cm/an
(Pelletier et al ., 1998).
La faune des sources hydrothermales du bassin de Lau. recoltee au cours des campagnes en submersibles,
Biolau 1989 et Nautilau 1990 (Desbruyeres et al.. 1994), a donne lieu a plusieurs travaux de zoologie
(GuiNOT, 1990; GEISTDOERFER, 1991). Desbruyeres et Segonzac (1997). dans leur ouvrage presentant tin
inventaire illustre des faunes des sources hydrothermales, recensent 22 especes du Bassin de Lau.
DEROULEMENT DE LA CAMPAGNE BORDAU I
Les participants (Fig. 2) : Pour la realisation de cette premiere campagne BORDAU, I'equipe scientifique se
composait, de trois anciens des campagnes MUSORSTOM, P. BOUCHET. B. RICHER DE FORGES et B. DAYRAT. de
J.S. Philippe, un etudiant travaillant actuellement sur les bases dc donnees de profondeur au Centre IRD de
Noumea et du Professeur A. Waren (l'inventeur de la drague DW). Le Professeur P. Newell, de f University of
the South Pacific, qui devait embarquer comnte representant des Fidji. en fut cmpeche par sa charge d'enseignement.
28
B RICHER DE FORGES ET AL.
Fig. 2. — Les participants de la campagne Bordau 1 sur la plage arrifcre du N.O. "Alis", dans le port de Suva. Do gauche a
droite : Jean-Sdbastien Philippe; Benoit Dayrat; Bertrand Richer de Forges; Philippe Bouchet; Anders Waren.
L'itineraire : Apres un depart du N.O. "Alis" de Suva en fin de journee du 22 fevrier, les premieres
operations debuterenl dans Somo-somo Strait, chenal situe entre les deux grandes Ties hautes de Taveuni ct Vanua
Levu (Nunn, 1994, 1998). Cc passage profond de 600 a 800 m, correspondrait, comme Natewa Bay, a une faille.
II presente un etranglement etroit avec des fonds de 300 m dans lequel on observe, en periode de vives eaux, de ires
forts courants de marees (> 3 noeuds). Les debris vegetaux flottants y sont abondants en surface et, une fois coules,
s'accumulent sur les fonds, de part et d'autre du seuil (Fig. 3-4a). La premiere station realisee lc 23 fevrier. faisant
suite aux operations des campagnes Musorstom, porte le numero DW 1391. Deux journees furent consacrees a
l'exploration de ce chenal, lc 23 fevrier (DW 1391-DW 1399) et le 4 mars (CP 1447-DW 1455). La premiere
journee rencontra principalcment des fonds vaseux a debris vegetaux, mais aussi un fond de sable grossier coquillier
vers 400 m. Pour la deuxieme journee, les tentatives de dragages sur le seuil. de 300 a 400 m, s'avererent tres
difficiles et peu fructueuses.
Vanua Levu. — Une journee fut consacree a l'echantillonnage de la vaste baie Natewa qui entaille sur plus de
35 milles, au nord-est, Pile de Vanua Levu (CP 1400-CP 1407). Dans l'axe de la baie Natewa on trouve des
profondeurs de plus de 1000 m et des fonds vaseux. Les stations de 1'extremite de la baie sont par petits fonds, de
moins de 150 m.
II faut rappclcr que le mcridien 180° coupe Pile de Vanua Levu et passe exactement par Somo-somo Strait. En
consequence, certaines stations de Somo-somo Strait ont des coordonnees avec une longitude est (DW 1391 -CP
1395; CP 1447-DW 1455), ainsi que celles du fond de la baie Natewa (CP 1402-CP 1407), alors que toutes les
autres stations de la campagne ont des coordonnees en longitude ouest.
En quittant Vanua Levu, le cap fut mis sur un groupe de recifs situe par 16°S ou se dcroulerent les prelcvements
de la journee du 26 fevrier (DW 1408-CP1413). Les prelevemcnts sur les penles, vers 500-600 m, ramenerenl des
pierres ponces et leur faune associee : cirripedes, pagures Pylochelidae, crevettes Glyphocrangonidae, crabes
Cyclodorripidae et Ethusinae, mollusques Propeamussium , Xenophoridae; vers 400 m de profondcur, des eponges
Hexactinellides, des crabes Pleistacantha , Macropodia. Psopheticus , Mursia et des Brachiopodes. Signalons la
CAMPAGNE BORDAU I
29
decouverte a la station CP 1412, par 400 m de profondeur, de l'espece Lyria planicostata, qui devient ainsi la volute
la plus a I'est du Pacifique (BOUCHET & POPPE, 1995).
La journee du 27 fevrier se deroula en operations (DW 1414-DW 1418) sur un mont sous-marin qui culmine
vers 220 m de profondeur (16°31'S - 178°59'W). La partie sommitale, couverte dune epaisse couche de manganese
(> 15 cm), se revela ires difficile a draguer. Parmi la petite quantite d'organismes recoltes, furent trouves deux
specimens du crinoi'de pedoncule Gymnocrinus richeri. Ce "fossile vivant", appartenant a une famille supposee
eteinte au Jurassique (Hemicrinidae), decrit d'un mont sous-marin de la ride de Norfolk en 1985. avait jusqu’alors ete
retrouve seulement aux lies Loyaute et sur la pente externe de rile Wallis (RICHER DE FORGES, 1990: Richer de
Forges & Menou, 1993).
Lcs quatre jours suivant furent consacres a I'exploration des pentes de 1'Tle de Vanua Balavu (CP 1419-
CP 1446).
Fig. 3. — Itineraire de la campagne BORDAU 1 dans les lies de Lau. Chaque point represente une zone ou plusieurs
operations dc dragages et de chalutages ont eu lieu (cf. liste des stations en Annexe).
30
B RICHER DE FORGES ET AL.
Vanua Balavu. — L'Tle de Vanua Balavu, d'une superficie de 54,6 km2, culmine a 283 m. Elle est constitute de
plusieurs Hots entoures d’une longue barriere corallienne delimitant un vaste lagon (Fig. 4 a). La partie nord de cet
ensemble d’lles semblait propicc au chalutage d’apres les cartes bathymetriques etablics par le CCOP/SOPAC
(Mineral Resources Dept. Fiji; 1/250 000 bathymetric map series). En fait les pentes, relativement douces entre
500 et 700 m de profondeur, sont herissees de nombreux pinacles volcaniques rendant toule operation avec des
engins trainants hasardeuse. Les quelques prelevements reussis montrent des fends envases et recouverls dc pierres
ponces avec des mollusques Xenophora et Propeamussium.
Sur le versant oucst, dans lc chenal separant file Kanacea de Vanua Balavu, les fonds de 350 a 420 m sont
graveleux, avec des debris coquilliers. Les especes remarquables sont des crustaces : la langouste fouet ( Puerulus
angulatus), des crabes Raninidae ( Lyreidius , Cosmonotus ), Goneplacidae ( Hexaplax ), Cyclodorripidae, Palicidae,
Majidae et des Galatheidae (Munich); des poissons, Soleidac, Callionymidae, Scorpenidae.
Au sud-est de File, se trouvent des fonds a debris coquilliers et coquilles dc pteropodes, assez riches en
micromollusques et, par endroil. des pierres ponces avec dc tres nombreux pagures Pylochelidae foreurs. Le trait
CP 1433 rapporta une grande quantite de pierres ponces et un gros morceau de dalle de gres avec une faune riche en
crustaces : Paromola crosnieri (trouvee pour la premiere fois hors de Nouvelle-Caledonie). Platymaia, Randallia
granulosa , Munida spp., cirripedes pedoncules.
Le versant est de File presente une pente relativement douce ou plusieurs chalulages fructueux furent realises
entre 300 et 400 m : crevettes Peneidae ( Metapenaeopsis ), Crangonidae, Pandalidae; crabes Leucosidae ( Randallia ),
Majidae (. Pleistacantha , Achaeus ); nombreux echinodermes ophiurides. Par ailleurs. il existe une passe etroite et
profonde (American Passage), dune morphologie tout a fait exceptionnelle. En effet, une entaille penetre sur plus
de 4 milles nautiques a Finterieur du lagon, formant une sorte dc canyon de pres de 200 m de profondeur dans un
lagon ou les profondeurs environnantes sont de 50 a 60 m. Les stations DW 1435 a DW 1439 lurent consacrees a
Fechantillonnage de ce canyon ; les fonds y sont composes d’un sable vaseux fin et blanc. de debris d 'Halimeda et de
foraminiferes; la faune y est largement domince par une espece d’echinide (du genre ? Maretia), si abundant que les
chaluts a perche (CP 1437-1438) en
furent entierement colmates.
Aprcs etre retourne le 4 mars dans
Somo-somo Strait pour tenter d’y
retrouver la faune des fonds sableux,
Fechantillonnage des Ties de Lau repril
le 5 mars, au niveau de File Yacata
(Fig. 3, 4a). Cette Tie, calcaire, isolee,
culmine a 248 m, entre Taveuni et
Vanua Balavu; ses pentes sont tres
raides et les prelevements ont eu
lieu entre 600 el 1300 m sur des fonds
envases a pierres ponces (la sonde XBT
indiquait 5°C a 850 m). Les organismes
recoltes furent essentiellemcnt des
crustaces : crabes Cyclodorripidae,
Ethusinae; crevettes ( Glyphocrangon ,
Heterocarpus longirostris ); anomoures
( Lithodes , Bathy munida, Munida );
isopodes Serolidae et Antarcturidae.
Fig. 4 a. — Carte des Ties de Lau. 1 : Somo-somo Strait (DW 1391-1399;
CP 1447-DW 1455); 2 : Natewa Bay (CP 1400-1407); 3 : Real's par
16°S-179°29'W (DW 1408-CP 1413); 4: Haul fond par 16°27’S-
1 78°55’W (DW 1414-1418); 5 : Vanua Balavu (CP 1419-1446);
6 : Yacata (DW 1456-1459).
Lakeba. — Le groupe de Lakeba (ou
Lakemba) comporte plusieurs Ties dont
la plus grande a une superficie de
56 km2 et culmine a 220 m (Fig. 4b).
Deux journees furent consacrees a la
faune bathy ale, la journee du 6 mars
Source MNHN . Paris
CAMPAGNE BORDAU 1
31
Fig. 4 b. — Carte des ties de Lau. 7 : Lakeba
(CP 1460-1468; CP 1502-1507); 8 :
Vatoa (DW 1469-CP 1476); 9 : Tuvana-i-
Colo (DW 1477-1483); 10 : Yagasa
cluster (CP 1484-1501).
Figures 4 a el 4b d'aprbs la carte du
"Hydrographic Service of the Navy,
England". n°4632 .
(CP 1460-1468) puis celle du 13 mars (CP 1502-1507). La
journee du 7 mars, jour de repos, fut utilisee pour des recolies de
mollusques terrestres sur l'Tle corallicnne surelevee d’Aiwa (60 m
d’altitude), au sud-est de Lakeba. Les stations realisees dans le
chenal compris enlre l'Tle Lakeba et le grand atoll de Bukatanoa.
de 250 a 450 m. ramenerent du sable grossier et des blocs
calcaires. Sur ces fonds fut trouvee une faune riche en crustaces
avec des crabes Majidae (Oxypleurodon. Pugetia, Platymaia,
Pleistacantha ), des Calappidae (Mursia), des crcvettes
Slylodactylidae et des anomoures Pylochelidae et Galatheidae
( Munida , Munidopsis). Signalons, a la station CP 1504, vers
430 m de profondeur. la presence de crabes de la famille rare des
Retroplumidae.
Plus profondement. vers 650 a 800 m, on retrouve. sur les
fonds a pierres ponces, des crevettes (Glyphocrangon,
Heterocarpus spp.) et des crabes (Randal I ia, Platymaia).
Vatoa. — Apres une nuit de navigation, directement cn
quittant l'Tle d'Aiwa, la petite Tie de Vatoa (19°49,7'S -
178°13,4'W) fut atteinte le 8 mars. Plusieurs epaves sur les cotes
de cette He rappellent les reels dangers de cette region mal cario-
graphiee. Les pentes furent echantillonnees entre 250 et 400 m
(DW 1469-CP 1476). Les stations les moins profondes
ramenerent des blocs de calcaires et des sediments grossiers
detritiques (DW 1472). Parmi ces sediments figuraienl des debris
d'eponges calcifiees du groupe des Sphinctozoaires ainsi que des
crabes Raninidae et Palicidae.
Ono-i-Lau et Tuvana-i-Colo. — ProLiant toujours d'une
navigation de nuit pour descendre en latitude, le groupe d'ilots de
Ono-i-Lau el Tuvana-i-Colo fut atteint le 9 mars (DW 1477-
1483). II s'agit des reliefs emerges les plus au Sud et les plus
isoles de l'archipel des Fidji. Ono-i-Lau (20°38'S- 1 78°4 1 'W) est
forme d'un groupe de 6 Tlots entoures de recifs alors que Tuvana-
i-Colo (21°00'S-178°43'W) est une ile solitaire, egalement
entouree de recifs. Les fonds explores, de 350 a 500 m. permirent
la recolte de crustaces Leucosidae, Majidae. Parthenopidae,
Dromiidae, Raninidae, de mollusques Turridae. Pectinidae et
scaphopodes.
Yagasa cluster. — Le groupe d'iles du sud de la ride de Lau,
centre autour de Yagasa Cluster, fut etudie au retour de Tuvana-i-
Colo, du 10 au 12 mars (CP 1484-1501). Par lui-meme, l'atoll
de Yagasa Cluster est forme d'un red!' barricre d’environ 7 milles
de diametre, ouvert au Nord, qui enserre 4 Ties dont Yagasa Levu
qui culmine a 1 19 m (ANONYME. 1996). Comme dans le reste de
l'archipel, on observe des fonds envases a pierres ponces en
dessous de 500 m et dans les petites profondeurs (400-200 m) des
blocs calcaires et des sables grossiers. La station DW 1488 fut
particulierement riche en microgasteropodes et en Pylochelidae.
Signalons a la station CP 1490 la recolte du crabe Majidae
32
B. RICHER DE FORGES ET AL.
Cyrtomaia fund, decrit des lies Chesterfield (GuiNOT & Richer DE Forges. 1988). Un dragage realise dans la
passe de Yagasa. par 240 a 319 m de profondeur, sur un fond de sable fin vaseux, rapporta des crabes Raninidae
( Lyreidius . Raninoides), Leucosidae etdes crevetlcs Peneides (Metapenaeopsis, Solenocera), Crangonidae.
REMARQUES SUR LES RECOLTES
La faune marine. — Les peuplcments des funds vaseux vers 500 a 700 m soni frequemment domines par de
gros oursins irrcguliers, rouge-violet, portant de petites ophiures commensales et des cirripedes pedoncules. Sur ce
type de fond a pierre ponces, les Paguridae du genre Parapagunts, associes a une Actinie. sont frequenis.
Une fois encore, la faune associee aux vegetaux coules est tres interessante : mollusques cocculiniformes dans
les noix de coco. Leptochiton, crustaces Galatheides et Amphipodcs.
Parmi les crustaces recoltes, certaines espcces, decrites de Nouvelle-Caledonic, ont ete retrouvees pour la
premiere fois. C'est le cas d'Oxypleurodon orbiculatus. decrite du sud de la Grande Terre (GuiNOT & Richer de
FORGES, 1985) et de Paromola crosnieri. Par conlrc, dans le genre Pleistacantha, c'est l'espece indonesiennc,
P. terribilis , qui est presente sur la ride de Lau et non celle decrite de Nouvelle-Caledonie. P. exophthalmus.
Le crinoi'de pcdoncule Gymnocrinus richeri est relrouve de nouveau et son aire de repartition connue devient
maintenant : ride de Norfolk, ties Loyaute, tie Wallis et ride de Lau. Ces localites sont situees sur la plaque
Australo-Indienne, a l'exception de Wallis. Cependant, si Ton prend en compte revolution teclonique recente du
sud-ouest Pacifique (AUZENDE et al., 1995), la zone situee au sud de la fosse de Vitiaz, entre celle-ci et la fosse des
Nouvelles-Hebrides est de formation recente. La presence de cette espece archaique semble done bien liee a la plaque
Australo-Indienne.
La malacofaune terrestre. — Les ties du Pacifique sont celebres pour leurs faunes de mollusques terrestres,
tres diversifies et presentant de hauts niveaux d’endemisme. Au sein de I'archipel fidjien, le groupc de Lau est
original, tant par la composition gencralc des peuplements, que par l'endemisme local. Par rapport aux grandes ties,
les PlacOStylus manquent, les grandes Orpiella (Helicarionidae) et les Diplommatinidae sont pratiquement absents;
par contre, les Partula n'existent a Fidji qu’aux ties de Lau, et c’est dans cet archipel que les Assimineidae atteignent
leur plus grande diversification, avec en particulier le genre endemique Fijianella. Les Endodontidae, famille
polynesienne par excellence, n’existent a Fidji qu'aux ties de Lau, avec notamment les genres endemiques
Priceconcha et Zyzzyxdonta. Les sous-genres d’Helicarionidae Diastole (Laua) et Lamprocystis (Naiaua) et les
genres de Charopidae Maafu et Lauopa sont egalement endemiques de Lau. Par suite de la disparition des forets
naturelles et de l'introduction d’especes envahissantes, les malacofaunes terrestres des ties du Pacifique connaissent
des taux d'extinction parmi les plus eleves de la planete. Or nos connaissances sur les mollusques terrestres des ties
de Lau sont fragmentaires et anciennes : elles sont fondees sur les collectes d’E. Graeffe (1870), E.H. Bryan
(aout-septembre 1924), H. Ladd (juillet-aout 1934), Y. KONDO (aout 1938) cl enlin L. Price (1970). Les
difficultes de deplacement a l'interieur de I'archipel et les conditions meteorologiques au moment de leur visite font
que toutes les ties sont loin d'avoir ete convenablement echantillonnees. C'est pourquoi la campagne Bordau 1 a
ete mise a profit pour debarquer sur six ties calcaires : Evuevu et Cikobia-i-Lau (groupe de Vanua Balavu), Yacata,
Aiwa, et enfin Yagasa Levu et Navutu-i-Loma (groupe de Yagasa). Sur chacune des ties, les recoltes comprennent
des ramassages a vue et des prelevements de litiere, et representent un echantillonnage adequat de la malacofaune.
De plus, des Partula lirata de Cikobia-i-Lau ont etc conservees vivantes jusqu'au retour a Suva et expedites au Zoo
de Londres (Dr P. Pearce-Kelly) qui maintient en elevage ex situ plus de 30 cspeces de Partula des ties du
Pacifique.
La pollution. — On pourrait croire que ces ties, isolees au milieu du Pacifique et relativement peu peuplees,
echappent a l'un des Beaux de notre temps, la pollution (BENTON, 1991, 1995). II n'en est rien ! Une coutume
ancestrale oc^anienne semble considerer l'ocean comme une vaste poubelle ou tout disparatt. A plusieurs reprises
pendant les deux campagnes realisees aux Fidji, les chaluts, dans la zone bathyale, ont rapporte de nombreux
detritus, sacs en plastiques, vieux pneus... Devant la passe de Suva, par 400 m de profondeur, cela pouvait passer
Source MNHN, Paris
CAMPAGNE BORDAU I
33
pour une consequence logique de l'activite portuaire el de la proximite de la grosse decharge urbaine de la ville de
Suva. Dans la zone de Bligh Water, la presence de detritus en plastique etait deja plus surprenante. Mais le
summum de la surprise fut atteint entre Taveuni et Vanua Levu, dans la zone de Somo-somo Strait. En 30 minutes
de chalutage par 600 m de profondeur (CP 1 392) fut remontee une impressionnante quantile de boites de conserves,
de torches electriques ct de bouteilles en matieres plastiques (Fig. 5).
Bien sur, cette pollution n'atteint pas encore le niveau catastrophique des mers d'Europe mais compte tenu des
petites populations responsables, il y a de quoi etre inquiet pour I’avenir.
FIG. 5. — Detritus trouves dans un trait de chalut & perche d'une demi-heure dans Somo-somo Strait, par 600 m de
profondeur (CP 1392).
CONCLUSIONS
Au terme des deux campagnes MUSORSTOM 10 et BORDAU 1, e'est 200 operations de dragages et de chalutages
qui ont ete realisees, pour la plupart, dans la zone bathyale de l'archipel des Ties Fidji. Les collections zoologiques
ainsi faites ameliorent considerablement nos connaissances sur cette region du Pacifique sud-ouest. precisant la
repartition d'especes decrites de Nouvelle-Caledonie ou enrichissant d'especes nouvelles certains genres connus plus
a l'ouest.
Les observations preliminaires, faites pendant la campagne, semblent confirmer l'existence, en zone bathyale,
d’un fond faunistique du sud-ouest Pacifique s'appauvrissant d'ouest en est.
REMERCIEMENTS
Nous remercions vivement les personnes qui ont contribue a la reussite de cette campagne :
Au siege de l'IRD (Institut de Recherche pour le Developpement; ex-ORSTOM), Raymond Lae et Patrice
Cayre; a Noumea, Rene GRANDPERRIN. Au Museum national d'Histoire naturelle. Alain CROSNIER et Philippe
MaeSTRATI ont assure les expeditions de materiel. A I'Ambassade de France a Suva, nous avons rei,'u un accueil
chaleureux de Monsieur l'Ambassadeur, Michel Jouvet, ct une aide precieuse de son Attache culturel et
scientifique, Didier Savignat.
34
B. RICHER DE FORGES ET AL.
Nos collegucs geologues B. PELLETIER, J.-M. AUZENDE et S. Calmant nous onl aiguille parmi la litterature
geophysique, foisonnante, du Pacifique sud-ouest. M. Segonzac nous a communique les references
bibliographiques concernant les campagnes de biologie sur les sources hydrothermales.
A bord du N. O. "A! is", le Second Capitaine Jean-Fran^ois BaRAZER, qui a dirige seul toutes les manoeuvres, a
fait le maximun pour realiser de bons chalutages, sans trop de casse; le Maitre d'equipage. Loi'c LEGOFF , a
beaucoup ceuvre pour le maintien en etat des engins de peche .
Les cartes sont dues a Marika TORTELIER, dessinateur au Centre ORSTOM de Noumea.
Nous remercions specialemcnt le Professeur Peter NEWELL qui, bien que n'ayant pu embarquer sur le
N.O. "Alis" cette fois, nous a accueilli avec sa convivialite habituelle.
REFERENCES
Anonyme, 1982. — Contribution & l'etude geodynamique du sud-ouest Pacifique. Travaux et Documents de iORSTOM,
Paris, (147), 649 p.
Anonyme, 1996. — lies de I'Ocean Pacifique (partie centrale). Instructions Nautiques, volume K9. Service Hydrographique
et Oceanographique de la Marine, Paris, 584 p.
Auzende, J.-M., Pelletier, B. & Eissen, J.-P., 1995. — The North Fiji Basin. Geology, structure, and geodynamic
evolution. In : B. Taylor (ed.), Backarcs Basins : tectonics and maginatism. Plenum Press. New York : 139-175.
Benton, T., 1991. — Oceans of garbage. Nature, (352) : 1 13.
Benton, T.G., 1995. — From castaways to throwaways: marine litter in the Pitcairn Islands. In : T.G. Benton & T.
Spencer (eds). The Pitcairn Islands: biogeography, ecology and prehistory. Biological Journal of the Linnean
Society, 56 : 415-422.
Bouchet, P. & Poppe, G.T., 1995. — A review of the deep-water volute genus Callioiectum (Gastropoda : Volutidae). In :
P. BOUCHET (ed.), Resultats des Campagnes Musorstom. Volume 14. Memoires du Museum national d'Histoire
naturelle, (A), 167 : 499-525.
Desbruyeres, D., Alayse-Danet, A.M.. Ohta, S. & the scientific parties of Biolau and Starmer cruises, 1994. — Deep-
sea hydrothermal communities in the southwestern Pacific back-arc basins (the North Fiji and Lau basins):
composition, microdistribution and food web. In : J.M. Auzende & T. Urabe (eds). The STARMER French-Japanese
Joint Project 1987-1992. Marine Geology, (116) : 227-242.
Desbruyeres, D. & Segonzac, M., 1997. — Handbook of deep-sea hydrothermal vent fauna. Editions IFREMER. Brest,
279 p.
Geistdoerfer. P.. 1991. — Ichtyofaune associee a l'hydrothermalisme ocdanique et description de Thermobiotes
mytilogeiton, nouveau genre et nouvelle espece de Synaphobranchidae (Pisces, Anguilliformes) de I'Ocean Pacifique.
Comptes Rendus de I'Academie des Sciences, Paris, ser. III. 312 : 91-97.
GutNOT, D., 1990. — Austinograea alayseae sp. nov. Crabe hydrothermal decouvert dans le Bassin de Lau ("BIOLAU"
1989), Crustacea, Decapoda, Brachyura. Bulletin du Museum national d'Histoire naturelle, Paris. 4eme serie. 11 (4) :
879-903.
Guinot, D. & Richer DE Forges, B., 1985. — Crustaces decapodes : Majidae (genres Platymaia, Pleistacantlui,
Sphenocarcinus et Naxioides). In : Resultats des Campagnes Musorstom 1 et II, Philippines. Memoires du Museum
national d'Histoire naturelle, (A), 133 : 83-177.
Guinot. D. & Richer de Forges, B., 1988. — Description de trois especes de Cyrtomaia Miers. 1886, de Nouvelle
Caledonie et des lies Chesterfield (Crustacea, Decapoda, Brachyura). Bulletin du Museum national d'Histoire naturelle,
Paris, 4eme serie, 10 (4), sect. A (1) : 39-55.
Nunn, P.D., 1994. — Oceanic Islands. Blackwell, Oxford & Cambridge. 413 p.
Nunn, P.D., 1998. — Pacific Island Landscapes. Institute of Pacific Studies. The University of the South Pacific, 318 p.
PELLETIER, B., 1999. — Subduction de rides et ouvertures arriere-arc dans le Pacifique Sud-Ouest (arcs des Tonga-Kermadec
et du Vanuatu, bassins de Lau et Nord-Fidjien). Memoire d'Habilitation a diriger des recherches. University Paris 6.
102 p.
Source MNHN , Paris
CAMPAGNE BORDAU I
35
Pelletier, B.. Calmant, S. & Pillet, R., 1998. — Current tectonics of the Tonga-New Hebrides region. Earth and
Planetary Science Letters , (164) : 263-276.
Richer DE Forges., B., 1990. — Les campagnes d'exploration de la faune bathyale dans la zone economique de Nouvelle
Caledonie (1984 & 1987). In : A. Crosnier (ed.), Resultats des Campagnes MUSORSTOM, Volume 6. Memoires du
Museum national d'Histoire naturelle. A, 145 : 9-54.
Richer de Forges, B., Mf.nou, J.-L., 1993. — La campagne MUSORSTOM 7 dans la zone economique de Wallis et Futuna.
In : A. CROSNIER (ed.), Resultats des Campagnes MUSORSTOM, Volume 10. Memoires du Museum national d'Histoire
naturelle. (A), 145 : 9-25.
Richer de Forges. B., Newell, P„ Schlacher-Hoenlinger. M., Schlacher, T., Nating. D.. Cesa, F. & Bouchet. P..
1999. — La campagne Musorstom 10 dans I'archipel des Ties Fidji. Compie rendu et liste des stations. In :
A. CROSNIER (ed.). Resultats des Campagnes MUSORSTOM. Volume 21. Memoires du Museum national d'Histoire
naturelle. 184 : 1-23.
Tiffin, D.. 1991. — Bibliography of published and unpublished work in the Lau Basin from 1985 to early 1991. South
Pacific Applied Geoscience Commission. Technical Report, (124), 18 p.
Woodhall, D.. 1985. — Geology of the Lau Ridge. In : D.W. Scholl & T.L., Vallier teds), Geology and offshore
resources of Pacific Island arcs - Tonga Region. Circum-Pacific Council for Energy and Mineral Resources. Earth
Science Series. 2 : 351-404.
ANNEXES
LISTE DES PARTICIPANTS A LA CAMPAGNE BORDAU 1
Chef dc mission : B. Richer de Forges.
AuLres participants : P. BOUCHET, A. WAREN, B. DayRAT, J.- S. PHILIPPE.
LISTE DES STATIONS DE LA CAMPAGNE BORDAU 1
(DW : drague Waren; CP : chalut a perche)
Source . MNHN, Paris
36
B. RICHER DE FORGES ET AL.
Source MNHN, Paris
CAMPAGNE BORDAU I
37
Source MNHN, Paris
38
B. RICHER DE FORGES £T AL.
0'S DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 21 — RESULTATS DES CAf
Porifera Hexactinellida: On Euryplegma auriculare
Schulze, 1886, and formation of a new order
Konstantin R. TABACHNICK
Institute of Oceanology
Academy of Sciences of Russia
Moskow, Russia
&
Henry M. REISWIG
Redpath Museum and Department of Biology
McGill University, 859 Sherbrooke St.
Montreal, Quebec, Canada H3A 2K6
ABSTRACT
Euryplegma auriculare, previously known off the coast of New Zealand, has recently been collected from the vicinity
of New Caledonia, Fiji. Wallis and Futuna Islands. The description of spicules, structure and speculation on the origin of
this species as well as reinvestigation of the related genera, Tretopleura and Fieldingia, has led to revision of the family
Aulocalycidae and modification of its diagnosis. The scope of the family has been restricted to live genera: Euryplegma,
Aulocalyx, Rhabdodiciyon , Ijimadictyon , and Leioplegma. A new taxon Aulocalycoida, with the same rank as the orders
Hexactinosa, Lychniscosa. Lyssacinosa. has been proposed to receive the reorganized family. It is distinguished by an
aulocalycoid framework. The order Hexactinosa is thus restricted to include all of its former families except for the family
Aulocalycidae. Two genera previously included in the Aulocalycidae, Tretopleura and Fieldingia. are reassigned because ol
their lack of an aulocalycoid framework. Tretopleura is referred to the hexactinose family Euretidae without a clear closest
relative. Fieldingia (sensu stricto), still poorly known, is provisionally referred to the order Reticulosa, as incertae sedis.
and possibly represents the only extant member of that group. It has some similarity to the family Stromatidiidae, but
cannot be included in that taxon with the limited information now available. The position of Reticulosa in relation to the
more well-known recent Hexactinellida taxa remains unclear.
Tabachnick, K. R. & Reiswig, H. M., 2000. — Porifera Hexactinellida: On Euryplegma auriculare Schulze, 1886, and
formation of a new order. In: A. Crosnier (ed.), Resultats des Campagnes Musorstom, Volume 21. Memoires du Museum
national d'Histoire naturelle. 184: 39-52. Paris ISBN-2-85653-526-7.
40
K. R. TABACHNICK & H. M. REISWIG
RESUME
Porifera Hexactinellida : Sur Euryplegma auriculare Schulze, 1886. Creation d’un ordre
nouveau.
Euryplegma auriculare, connu jusqu'a present du large de la Nouvelle-Zelande, a recemment ete recolte au voisinage
de la Nouvelle-Caledonie et des lies Fidji. Wallis et Futuna. La description des spicules et de la structure de ccttc espece,
liee k des hypotheses sur son origine. ainsi que le reexamen des genres voisins Trelopleura et Fieldingia, ont conduit a une
revision de la famille Aulocalycidae et k la modification de sa diagnose. Le champ de la famille a etd reduit k cinq genres :
Euryplegma, Aulocalyx, Rhabdodictyon, Ijimadictyon et Leioplegma. Un nouveau taxon Aulocalycoida. ayant le
meme rang que les ordres Hexactinosa, Lychniscosa, Lyssacinosa, est propose pour recevoir la famille ainsi revisee.
11 se distingue par une organisation du squelette aulocalycoide. L'ordre Hexactinosa se trouve ainsi reduit a ses anciennes
families k l'exception de celle des Aulocalycidae. Deux genres inclus precedemment dans les Aulocalycidae, Trelopleura
et Fieldingia, sont rcclasses a cause de leur manque de structure aulocalycoide. Trelopleura est place, k litre provisoire.
dans la famille Euretidae appartenant aux Hexactinosa. bien qu'aucune autre forme de cette famille ne lui soit clairement
trks proche. Fieldingia (sensu stricto), encore mal connu. est provisoirement rattache k l'ordre Rcticulosa. cn
tant qu'incertae sedis et represente pcut-etre le seul membre de ce groupe encore existant. II possede quelques
similitudes avec la famille Stromatidiidae, mais ne peut etre inclus dans ce taxon avec le pen d'information dont nous
disposons actuellement. La position des Reticulosa par rapport aux Hexactinellida rdccnts, bien mieux connus, reste
obscure.
A large number of interesting Hexactinellida have been collected in recent years by French expeditions in the
southwestern part of the Pacific Ocean. Some of these species were previously known only by a few specimens
from restricted locations. One of them — Euryplegma auriculare — turns out to be a common species in this area.
The numerous specimens of E. auriculare made available by these expeditions provide opportunity to redescribe the
species and to discuss the syslematics of the Aulocalycidae.
Abbreviations: MNHN, Museum national d'Histoire naturelle (Paris); NHM, The Natural History Museum
(London); IORAS, Institute of Oceanology of Russian Academy of Sciences (Moscow).
1. — DESCRIPTION OF EURYPLEGMA AURICULARE Schulze
Genus EURYPLEGMA Schulze, 1886
Euryplegma Schulze, 1886: 80.
Type Species. — E. auriculare Schulze, 1886.
DIAGNOSIS (from SCHULZE 1887, emended here). — Fan-like, thin- walled sponge with aulocalycoid frame¬
work of choanosomal skeleton. Dermalia and atrialia are pentactins. Loose microhexactins are rarely found
in the choanosomal framework. Microscleres are discohexasters, hemidiscohexasters and sometimes
discohexactins.
Euryplegma auriculare Schulze, 1886
Figs 1-13; Table 1
MATERIAL EXAMINED. — New Zealand. "Challenger", sin 170 A, off Raoul or Sandy islands. 29°45'S.
1 78° 1 l'W, 1120 m: Lectolype (BMNH 1887.10.20.075), paraleciolype (BMHH 1887. 10.20.075A).
New Caledonia. Biocal: stn CP 17. 20°34.54,S, 167°24.68'E, 3680 m, 14.08.1985: 2 specs (MNHN-HCL 366-
368). — Stn CP 26. 22°39.66'S, 166027.41'E, 1618-1740 m, 28.08.1985: 2 specs (MNHN-HCL 369-370). —
Source : MNHN, Pans
THE HEXACTINELLID EURYPLEGMA
41
Figs 1-8. — Fragments of Euryplegma auriculare: 1-2, MNHN-HCL 369. upper and side view (scale 2 cm); 3-5. MNHN-
HCL 395, upper part side view and lower parts from two sides (scale 1 cm); 6-7. MNHN-HCL 392, lower parts, views
from 2 sides (scale 2 cm); 8. MNHN-HCL 392, basal part (scale 1 cm).
Source MNHN, Paris
42
K R TABACHNICK & H. M. REISWIG
Stn DW 33, 23°09.7rS, 167°10.27'E, 675-680 m. 29.08.1985: 2 specs (MNHN-HCL 372-373). — Stn DW 51.
23°05.27'S. 1 67°44.95'E. 700-680 m. 31.08.1985: 1 spec. (MNHN-HCL 374). — Sin CP 52, 23°05.79'S, 1 67°46.54'E.
600-540 m, 31.08.1985: 1 spec. (MNHN-HCL 371).
Musorstom 4: sin CP 217, 22°3.60'S, 167°27'E, 850 m, 29.09.1985: 1 spec. (MNHN-HCL 396).
Biogeocal: sin CP 238, 2r27.64’S, 166 23.41’E, 1300-1260 m, 13.04.1987: 10 specs (MNHN-HCL 375-382.
384-385). — Sin CP 265, 21°04.09'S. 167°00.40'E, 1760-1870 m, 18.04.1987: 6 specs (MNHN- HCL 386-391). —
Stn CP 290. 20°36.91'S, 167°03.34’E, 920-760 m, 27.04.1987: 1 spec. (MNHN-HCL 392). — Stn CP 297, 20°38.64’S.
167°10.77'E, 1230-1240 m, 28.04.1987: 2 specs (MNHN-HCL 393-394)
Calsub: Dive 12, 21°28'S. i66°21.50’E. 1265-700 m, 3.03.1989: 1 spec. (MNHN-HCL 395)
Loyally Islands. Musorstom 6: stn CP 427, 20°23.35'S, 166°20'E, 800 m, 17.02.1989: 1 spec. (MNHN-
HCL398). — Stn CP 438. 20°23'S. 166°20.10'E, 780 m, 18.02.1989: 1 spec. (MNHN-HCL 397).
Fiji. DK-88-4-NBF, 1988. 12-13, stn 29, 16°39.89'S. 173°49.32'E, 2580 m (MNHN-HCL 383)
Wallis and Futuna Islands. MUSORSTOM 7: stn DW 539. 12°27.30'S, 177°27.30'W, 700 m, 17.05.1992: 1 spec.
(ZMMU). — Stn CP 550, 12°14.80'S. 177 28’W, 800-810 m, 18.05.1992: 1 specs (MNHN-HCL 399). — Stn CP 564,
1 1°46. 10'S, 1 78°27.40'W, 1015-1020 m, 20.05.1992: 2 spec. (MNHN-HCL 400-401). — Stn 636. 13°39.40'S,
179°55.50'W, 650-700 m, 30.05.1992: 1 spec. (MNHN-HCL 402).
Type Material. — Lectotype (BMNH 1887.10.20.075). Paralectotype (BMNH 1887.10.20.075A). Both
collected by the HMS "Challenger", stn 170 A, 29°45'S, 178° 1 LW, NE New Zealand, depth 1120 m.
DESCRIPTION. — Body : The sponge is tongue-like or similar to an ear-shaped plate; flabellate. Sections of
most specimens, and especially the upper parts, are sicklc-like with one surface convex and the other concave.
Some fragments of the lower part of the body have a secondary funnel produced by ingrowth and union of the
opposite edges (Fig. 1). Each of these fragments has a narrow, longitudinally directed slit enclosed by the union of
the ingrown parts above. Oval and round orifices approximately 1-3 mm in diameter are located on both concave
and convex surfaces. Investigation of transverse and longitudinal sections of specimen MNHN-HCL 395 showed
that all internal interconnected canals (schizorhyses) are lined with pentactins identical with those of the dermal and
atrial linings. Some orifices are covered by a mesh-layer containing the same pentactins.
Figs 9-12. — Sections of Euryplegma auriculare: 9-11, transverse sections through basal and upper parts; 12,
longitudinal section.
Source MNHN, Paris
THE HEXACTINELLID EURYPLEGMA
43
Status of existing type specimens has not been previously addressed. The specimen 1 887. 10.20.075A stored in
The Natural History Museum (London) is marked with a type label. Since it is a fragment of macerated skeleton of
the lower part of the body (SCHULZE, 1887, pi. 102, fig. 2) it is here designated as a paralectotypc. The other
specimen 1887.10.20.075 is 80 mm in length. It consists of the lower part of a sponge (SCHULZE, 1887. pi. 102.
fig. 1) and contains the loose spicules. It is here designated as the lectotype.
The newly collected specimens are all fragments of the lower part of sponges. Some of them are represented by
macerated skeletons, while others contain loose spicules which correspond in size to the type material. The largest
complete fragment (MNHN-HCL 400) is over 140 mm in length. The thickness of the upper lamellar-like parts is
3-9 mm. The most complete specimen (MNHN-HCL 395) was broken during capture by the manipulator of the
submersible. The lower part is 50 mm in length while the rest of the body part is represented by numerous flexible
laminae 3-5 mm in thickness. The specimens contain none of the allochthonous loose spicules which have been
reported from the type materials by REID (1957).
Spiculation : The choanosomal skeleton consists of a framework of aulocalycoid skeleton with numerous
synapticular junctions. All central spicule nodes are hexaradiate. Since the walls between internal canals are thin
(1-3 mm), most rays arc bent to lie in a single plane, as occurs in framework dictyonalia of Aphrocallistes. The
diameter of the aulocalycoid beams is 0.03-0. 1 1 mm. They are smooth or covered with short spines. Free rays of
these hexactins have rough tips or they are entirely rough.
The loose hexactins are numerous in the lectotype but are less abundant in all of the new specimens. These
small hexactins are never fused to the framework as in most Farreidae and Euretidae. Choanosomal hexactins have
rays 0.023-0.084 mm — probably up to 0.266/0.002-0.005 mm — covered with short spines. These choanosomal
hexactins arc numerous in the lectotype, common in the specimens HCL 387 and HCL 388, rare in HCL 395,
HCL 389, HCL 392 and apparently absent in HCL 372, HCL 396. Only one large choanosomal hexactin was
found in the lectotype while there were several in HCL 395. These may belong to the dermal or atrial hexactin
population.
Spicules underlying the dermal, atrial and canalar surfaces are pentactins, rarely stauractins and sometimes the
hexactins mentioned above. Pentactins from all parts of the body are identical in range ot shape and size. The
tangential rays are 0.080-0.448 mm long; the proximal rays arc 0.068-0.669 mm long; their diameter near the base
is 0.007-0.034 mm. The rays are more or less spinous, but always have dense short spines ornamenting their ends.
Pentactins from concave and convex surfaces of specimens HCL 387 and HCL 389 were compared — they are
indistinguishable in most dimensions (Table 1).
Table 1. — Some measurements of the spicules of Euryplegma auriculare (in mm). L: length. — D: diameter.
44
K R. TABACHNICK & H. M. REISWIG
0.4
2-8, 14-17
14 15 16 17
Fig. 13. — Spicules of Euryplegma auriculare: 1, portion of choanosomal aulocalycoid skeleton (MNHN-HCL 395);
2. spherical discohexaster (NHM 1887.10.20.075); 3. stellate discohexaster (NHM 1887.10.20.075);
4,8, spherical discohexaster (MNHN-HCL 395); 5. discohexactin; 6-7. stellate discohexaster (MNHN-HCL 395);
9, pentactin (NMH 1887.10.20.075); 10-12, pcntactins (MNHN-HCL 395); 13, choanosomal hexactin (NHM
1887.10.20.075); 14-17, tips of pentactin tangential rays (MNHN-HCL 395).
Source MNHN, Paris
THE HEXACTINELLID EURYPLEGMA
45
Microscleres: Discohexastcrs usually prevail over the other types of microscleres with discoidal ends —
hemidiscohexastcrs and discohcxactins. Regular discohexasters 0.029-0.180 mm in diameter often have stellate
form, i.e. their secondary rays are arrayed in rather compact tufts, or they sometimes have secondary rays organized
in broad tufts, rendering their external form spherical. Discohexactins 0.029-0.088 mm in diameter are almost
absent in the lectotype, rare in some specimens e.g. HCL 392, HCL 396, HCL 395 and predominant in HCL 372.
The hemidiscohexasters are rare or entirely absent in most specimens but they were regularly observed in
HCL 372. The uncinates (spicules important for taxonomy) found in the type material by Reid (1957) were
absent in the sponge HCL 372 captured by submersible. It is now possible to confirm the suggestion of REISWIG
and TSURUMI (1996) that these spicules have an allochthonous origin.
2. — THE AULOCALYCOID STRUCTURE OF EURYPLEGMA
The taxonomic position of Euryplegma has been changed many times. SCHULZE (1886) described and first
referred his new genus to the family Tretodictyidac, but the next year he (SCHULZE, 1887) reassigned it to the
Rossellidae. Lima (1898) included Euryplegma in his Leucopsacinae. which, at that time, was a subfamily of
Rossellidae. Not long after, Schulze (1904) again changed the position of the genus, assigning it to the
Dactylocalycidae. Later Lima (1927) created a new family Aulocalycidae and included the genus Euryplegma in it.
The features of Euryplegma, described by SCHULZE (1887), i.e. the presence of epirhyses and aporhyses, are
reflected in Lima's differential key characters, but he did not provide a generic diagnosis. LauBENFELS (1955)
proposed a special family, Euryplegmatidae, for this genus but it had no diagnostic difference from the previous
Aulocalycidae and thus, no reasonable basis could be found to distinguish it. REID (1957) reinvestigated the type
materials and found uncinates, which he considered to be autochthonous, and schizorhyses instead of epirhyses and
aporhyses. These features led him to transfer the genus to the family Tretodictyidae. REISWIG & TSURUMI (1996)
dismissed the autochthonous nature of the uncinates and changed the diagnosis of Aulocalycidae by adding the
possibility of schizorhyses in order to include Euryplegma here. With the new specimens, we can now determine
the exact set of spicules and confirm the suggestion made by Reiswig & TSURUMI (1996) on the spicules of
Euryplegma. However, the mode of wall canalization casts doubt on the present taxonomic position of the genus
and requires special analysis. Unfortunately the juvenile features and young stages of E. auriculare are unknown,
hence speculation on the structure of this genus have to be based mainly on theoretical considerations. We present
3 schemes of interpretation of origin and nature of the internal canals of Euryplegma (Fig. 14).
Lima (1927) interpreted the wall structures of Euryplegma as epirhyses and aporhyses, apparently solely on the
basis of the description and figure provided by SCHULZE (1887). The structure of the reviewed material does not
agree with this interpretation. Reid's (1957) opinion, supported by REISWIG & TSURUMI (1996), on the develop¬
ment of true schizorhyses is more reliable (Scheme A, Fig. 14). The initially tubular archetype or unknown
ancestor became tongue-like due to asymmetrical growth of a tube's sector. Walls became thicker to provide greater
support for an extended body. Thickened walls required elaboration of an internal aquiferous system hence the
system of extensive canals were developed. The formation of the tongue-like body which is sickle-like in section,
is the result of the increase in longitudinal rigidity of whole sponge. In this case the convex side is dermal while
the opposite concave-atrial, similar to other bilateral-symmetrical forms of Hexactinellida. The secondary funnel
sometimes, but not always, present in the lower part of a sponge is produced by ingrowth and union of the
opposite edges of the a tongue-like growth. This enclosed, narrow, longitudinally-directed slit must be considered,
in this scheme, as a parietal osculum. Some objections can be raised against Scheme A (Fig. I4A). All of the
internal wall passages observed in sections of Euryplegma are lined with pentactins similar to those of the exposed
convex and concave surfaces. Are these specific canalaria equivalent to dermal and atrial spicules? Such canalaria are
unknown in other Hexactinellida, moreover true schizorhyses arc "subdermal canals always covered with dermal
layer" (Lima, 1927). Thus the interpretation of structure and derivation by Scheme A (Fig. 14), a pattern
consistent for Tretodictyidae. does not seem to be applicable to Euryplegma.
Scheme B of Fig. 14 suggests an interpretation of body form and canalization by development of schizorhyses
which arc lined with surface tissues originating from both dermal and atrial layers. This process began with
46
K. R. TABACHNICK & H. M. REISWIG
asymmetrical growth of only a sector of an ancestral tubular form as in the previous scheme (A). Walls of a
tongue-like sponge became thicker, its opposite surfaces developed small furrows and depressions which included
dermal and atrial layers. Some depressions became connected with those directly opposite and also with internal
cavities forming the structure typical of the genus Euryplegma. In this case dermal and atrial spicules are situated
on the concave and convex surfaces of the tongue-like sponge and, in addition, underlie the linings of the structures
similar to schizorhyses. The occurrence of the secondary funnel in the lower part of a body does not change the
location of layers of the initial structure. The canal-like structures in this case are not homologous to true
schizorhyses of Tretodictyidae and instead are specific aulocalycoid structures, interpretable as tortuous oscula —
orifices between dermal and atrial layers. This interpretation is improbable since intermediate forms of such
"pseudocanalization" are so far unknown. The primitive canalization of Tretopleura with superficial epirhyses and
aporhyses is considered unrelated to that of Euryplegma and offers no support here. If Scheme B is accepted, then
the family Aulocalycidae possesses unique "pseudocanalar" structures.
A
B
C
Fig. 14. — Schemes of morphogenesis of Euryplegma (thin line: dermalia; thick line: atrialia; dotted line: canalaria):
A, scheme of development through wall canalization of an asymmetrical form. B. scheme of development through
wall "pseudocanalization" of an asymmetrical form. C. Scheme of development through flattening of a tubular form
with lateral parietal oscula and marginal oscula.
1,6, hypothetical tubular form, side view and section. 5,7.9,13, hypothetical tubular form with lateral
parietal oscula, side view and section. 2, 17-19, hypothetical tubular form with lateral oscula, side view and section.
3. real tongue-like form, side view. 4. 10, 11, 14, 15, 20. 21, real tongue-like forms with secondary funnel in
lower part, side view and upper section. 12, 16, 22, sections through the lower secondary funnel of form 4.
Source : MNHN, Paris
THE HEXACTINELLID EORYPLEGMA
47
According to Scheme C (Fig. 14) the ancestral tubular form developed numerous lateral parietal oscula.
Bilateral compression produced a tongue-like sponge. To increase local rigidity the walls on opposite sides of the
atrial surfaces were connected by broad anastomoses. The sickle-like form in transverse section of the tongue-like
sponge developed to increase longitudinal rigidity of the entire organism. Also the secondary funnel in the base of
some specimens was produced by ingrowth and union of the opposite edges of a tongue-like body. According to
this interpretation, the convex and concave (or the inner and the outer) surfaces are lined with dermalia. The parietal
slit enclosed in the secondary funnel by union of the ingrown parts is not homologous to a parietal osculum. but
is an inter-cavaedium — an entrance into a dermally-lined cavity (term of Reid, 1964, for some Euretidae).
The atrial cavity in this form must be situated between convex and concave surfaces, subdivided into a system of
branching canals. The primary osculum of Euryplegma must also be subdivided into numerous apertures at the
growing edge of the tongue-like sponge. The functions of the atrial cavity and the oscula have presumably been
altered as a result of this major shape change. Water How might maintain its original pattern, moving into the
walls from all exposed surfaces, and transported to the distal edge within the atrial canal system. Alternately water
flow might have been grossly modified to pass from one dermal surface to the other across the now internalized
atrial cavity, as in the simple ear-like processes of "metameric" claviscopulid described as Chonelasma
(Tabachnick, 1988, 1991). These alternatives can only be decided when live specimens are directly examined or
specimens with very well-fixed tissues are analyzed histologically.
The three schemes presented should be viewed as hypotheses of possible derivation of the specialized
Euryplegma body form. At present there is no strong evidence, or known intermediate, to suggest that Scheme C.
calling for body flattening and atrial bridging, is more likely scenario than Scheme B. The canals of Euryplegma
are still termed schizorhyses, with the understanding that they arc unlikely to be homologous with the similar
canals of the Tretodictyidae.
3. — THE SCOPE OF THE FAMILY AULOCALYCIDAE
The family Aulocalycidae was created by Ijima (1927). It included 5 genera; Aulocalyx, with two species
A. irregularis Schulze, 1887, and A. serialis Dcndy, 1916; Rhabdodictyon, with two species R. delicatum
Schmidt, 1880, and R. kurense Ijima, 1927, and three monospecific genera, Tretopleura candelabrum Ijima, 1927.
Euryplegma auriculare Schulze, 1886, and Fieldingia lagettoides Kent, 1870. Only two taxa were added later:
Tretopleura styloformis Tabachnick, 1988, and Leioplegma polyphyUon Reiswig & Tsurumi, 1996. R. kurense
was formally transferred to a new genus Ijimadictyum Mehl, 1992. The family, according to Ijima (1927), was
determined by an irregular-meshed skeletal framework (later denoted as "aulocalycoid" by REID. 1963): the presence
of scopules and uncinates was permitted, as well as epirhyses and aporhyses in Tretopleura and Euiyplegma. The
diagnosis of the family given by BARTHEL and Tendal (1994) was simplified to the aulocalycoid skeleton
without any other features and without speculations on the scope of the family. Reiswig and TSURUMI (1996)
clarified the misunderstood distinction of the aulocalycoid skeleton and suggested an expanded diagnosis which also
allowed uncinates and scopules. Thus the family Aulocalycidae is presently contained within Scopularia and
includes forms with a great range of canalization modes where other Scopularia have one special mode, or perhaps
two modes, within a family. The mode of canalization is generally regarded as the most significant feature in the
family definition - e.g., families Euretidae (sensu IJIMA) or Euretidae and Coscinoporidae (sensu ZlTTEL. 1877;
Barthel & Tendal, 1994) and Tretodictyidae. Instances of body-form complication by development of inter-
cavaedia or atrial expansion in sponges such as Myliusia (Reid, 1964) are usually not reflected in the diagnosis of
families.
Except for the aulocalycoid type of skeleton, there is no other character authentically defining the family. The
obligatory presence of dermal and atrial pentactins are common in other Scopularia. From the contributions of
Ijima (1927), Mehl (1992), Reiswig & Tsurumi (1996), the genera recognized until the present as constituents
of the Aulocalycidae can be characterized by the following specific features (except choanosomal skeleton and
dermal and atrial pentactins):
1 . Euryplegma - see above.
48
K. R. TABACHNICK & H. M. REISWIG
2. Aulocalyx is a tubular stellate form with vertically directed lolds bearing lateral oscula; microscleres are
distinctive rhopalasters and various discohcxasters.
3. Rhabdodictyon is a cup-like or tubular sponge with lateral oscula (similar to e.g.. Regadrella in shape),
microscleres are discohexasters, spirodiscohexasters. hemidiscohexasters and discohexactins.
4. Fieldingia, as originally described by Kent (1870), is an incomplete sponge represented by a minute and
thin lamellar fragment composed of pentactins, rarely stauractins and hexactins; these spicules are fused to each
other by tangential rays and numerous synapliculae.
5. Tretopleura is a fan-like elongate sponge. Only this genus of Aulocalycidae (, sensu UlMA) has epirhyses and
aporhyses (Ijima, 1927), scopules. uncinates and often hexactins (rarely pentactins) among dermalia and atrialia
(TABACHNICK, 1988).
6. Ijimadictyum is composed of branching and anastomosing tubes provided with additional lateral oscula,
microscleres are discohexasters, spirodiscohexasters, hemidiscohexasters and discohexactins.
7. Leioplegma is fan-like with dichotomously branching lobate processes; microscleres are discohexasters and
hemidiscohexasters.
Loose choanosomal hexactins have been reported from some specimens of Euryplegma , from Aulocalyx
serialis and from Leioplegma polyphyllon. It is worth noting that they are always loose, never fused to the
aulocalycoid framework.
Of these seven genera, five of them, Euryplegma, Aulocalyx, Rhabdodictyon, Ijimadictyum and Leioplegma,
form a well distinguished group of taxa and it is suggested here that the family Aulocalycidae be limited to this
group. An emended family diagnosis, based upon those given by IJIMA (1927) and Reiswig & TSURUMI (1996), is
offered here.
Diagnosis of the Family Aulocalycidae (from Ijima, 1927 emended). — Funnel-like, or tubular
dichotomously branching-fusing, or fan-like thin-walled sponges with or without lateral oscula. The choanosomal
skeleton is of aulocalycoid type — irregular-meshed framework made of hexactins which are orientated irregularly
and have elongate rays, usually curved, intersecting at various angles and fusing at intersections and by
synapticular junctions. Dermalia and atrialia are pentactins. Canalization is usually lacking but may include unique
schizorhyses. Microscleres have spined club ends (rhopalaster) or discoidal ends (discohexasters, hemidiscohexasters
and discohexactins).
Remarks. — The following features seem to be absent in the suggested list of the representatives of the
family Aulocalycidae: no sceptrules (scopules or clavules), no uncinates. Except tor the schizorhyses canalization
of Euryplegma, true wall canalization appears to be absent in the other members of the family. Dermalia and
atrialia could include some stauractins or even hexactins. Rather small loose choanosomal hexactins could be
present within the meshes of the choanosomal framework. The wide spectrum of external body shape does not
seem to be an extraordinary feature of Aulocalycidae as it also occurs in the close families, Eurelidae and
Euplectellidae.
4. — PLACEMENT OF TRETOPLEURA AND FIELDINGIA
Two genera, Tretopleura Ijima and Fieldingia Kent, presently members of the family Aulocalycidae, lack the
required aulocalycoid choanosomal skeleton. They are to be transferred to other families.
We propose referring Tretopleura to the Euretidae (or Coscinoporidae if one chooses to accept division of the
former family). Both species of Tretopleura have small hexactins fused to the framework similar to representatives
of the Euretidae and Farreidae. The genus is poorly investigated and it remains uncertain whether the skeleton of
primary regular framework known in the Euretidae/Coscinoporidae representatives (Reid, 1958a, 1958b, 1961) can
be demonstrated in the type species, T. candelabrum. The skeleton in the central, thicker part of T. styloformis is
fairly regular and may be considered as euretoid. Since the latter is subjectively assigned to Tretopleura, its
characters cannot be used to set the characters of the genus. The spicule composition and overall body form of
Tretopleura, necessarily set by its type species, remains unknown; canalization includes extradictyonal epirhyses
Source MNHN, Pans
THE HEXACTINELLID EURYPLEGMA
49
and aporhyses. Spiculation, body form and canalization of T. styloformis are similar to those of Bathyxiphus ,
differing only in microscleres. We prefer to maintain the genus as a distinct taxon pending finding the type
specimen and comparing its characters to those of the other genera of Eurclidae.
The genus Fieldingia has been described twice. The holotype of F. lagettoides is a poor fragment described by
Kent (1870) from the Atlantic ocean, the remnants stored as NHM 1872.2.3.178 (Figs 15-17). It does not corre¬
spond to the other specimen from the Pacific ocean described under the same species name by SCHULZE (1887),
1887.10.20.127. The characters of the specimen described by Kent are summarized above (3. 4). It is impossible
to decide whether junctions between the tangential rays of the dermal pentactins begin as sympodial ray branching,
spines produced only in the tangential plane, or simply represent synapticular junctions; these alternatives may
only be extreme examples of a continuous spectrum of secondary silicification processes. We cannot even be
certain that small stauractins arc not involved as well. If this latter feature is true, the skeleton may resemble (in
plane view) some skeletons of Reticulosa composed of stauractins of several orders of size. "The largest are
arranged so as to enclose square meshes, and the smaller grades lie within these meshes so as to subdivide them
regularly" (Reid, 1958a) - into so-called "quadrules" (Hall & Clarke, 1898). In spite of the fact that Reticulosa
are considered to be lyssacine sponges, some members of this group may have had spicules cemented together
(Reid, 1958a).
Figs 15-16. — Fieldingia lagettoides: holotype, attached to coral (scale 5 mm).
Small spherical aggregations of fused hexactins are also sporadically found attached to the main skeleton of
lagettoides. These spherical bodies are similar to the siliceous aggregations which have been reported from
diactins of some Rossellidae (e.g., so called "small basidictyonal masses" in Staurocalyptus glaber Ijima 1904.
pi. 15, fig. 13), and are also likely to have an allochthonous origin.
dlie specimen described by SCHULZE as a second representative of F. lagettoides is a fragment of fused irregular
Iramework of large and small hexaradiate spicules with numerous thick spherical aggregations probably having
autochthonous origin. These spherical aggregations must be simple knots of the main spicule framework. The
loose spicules are scopules, uncinates, diactins; microscleres are discohexasters, hemihexasters and hexactins. It is
impossible to find similarities shared by these fragmentary specimens of KENT and SCHULZE, except for the
spherical siliceous bodies which indeed differ in detail in the two forms. Hence Schulze's specimen cannot even
50 K. R. TABACHNICK & H. M. REISWIG
be considered a representative of Fieldingia. According to its skeleton and spicules, some of which seem to have
allochtonous origin, it could not be referred to any existing genus or family and its allocation must await a more
detailed analysis of the specimen.
Fig. 17. — "Quadrules" of Fieldingia lagettoides (tangential surface). The large spicules are pentactins or hexactins, ihe
small ones are stauractines and synapticulae. This diagram, made from the NHM slide ol ihe holotype described by
Kent, resembles the figured skeletons of reticulosans (scale 1 mm).
The holotype of Fieldingia is somewhat similar to fossil Stromatidium Girty, 1908, which contains a single
species — S. typicale Girty. It is a cylindrical sponge with one osculum and walls built with several separate
layers of rough pentactins fused into a continuous net within each layer. The fusion is formed with dichotomously
branching pentactin tangential rays (up to three branchings) and probably with synapticular junctions. Analogous
dichotomous branching of tangential rays is described for the dermal pentactins of Aulochone clathroclada (family
Rossellidae) (Levi & Levi, 1982). On the photo of a non-type fragment of S. typicale (Finks, 1960, pi. 44,
fig. 7), the first inner spicular layer, which must be considered to be juvenile or neanic and which is situated close
to the atrial cavity, appears to be composed of fused stauractins. This fact is omitted in the description and should
be reviewed in both the photographed non-type fragment and GlRTY's type specimens. Dermal pentactin fusion
is known also for Lychniscosa (Mehl, personal communication). Finks (1960) suggested a new family
Stromatidiidae for this unique genus and the family itself is tentatively included into his superfamily
Brachiospongioidae. In our opinion both of Reid's systems (1958b), one developed chiefly from Lima's system
and the other from that of SCHRAMMEN, offer more suitable placement for Stromatidiidae. It may be included into
the order Reticulosa, characterized as having a skeleton of "coherent or connected megascleres of dermal origin or
partly dermal origin" (REID, 1958b). The similarity of Fieldingia to Stromatidium may be only superficial in that
there is an internal dictyonal framework of hexactins in Fieldingia but no such component in Stromatidium or
other Reticulosa. For the present, we consider Fieldingia doubtful member of the order Reticulosa, possibly related
to Stromatidiidae but not able to be placed within that family on the basis of limited knowledge of both groups.
The Reticulosa may be a hexasterophoran group with the same range as Lyssacinosa, Hexactinosa and Lychniscosa
or a separate stock with the range equal to Hexasterophora and Amphidiscophora.
Source MNHN, Paris
THE HEXACTINELLID EURYPLEGMA
51
5. — THE CLASSIFICATION OF HEXASTEROPHORA
Since the Aulocalycidae are distinguished from the other Hexactinosa by a distinct pattern of framework
construction, their removal from that order results in the formation of two coherent and consistent laxa, the
Aulocalycoida for the family Aulocalycidae, and the Hexactinosa for the remaining members. The Aulocalycoida
should have the same rank as Ijima's (1927) tribes or Reid's (1958b, 1961) orders: Hexactinosa, Lyssacinosa and
Lychniscosa. The new taxon, order Aulocalycoida, can be characterized by the diagnosis of Aulocalycidae without
specific description of loose spicules. Hence the Aulocalycoida is offered for sponges with specific aulocalycoid
skeleton in which uncinates and sceptrules are absent. The diagnosis of the Hexactinosa should be reformed to
delimit its scope. Hexactinosa were defined by Reid (1958a) as "dictyonina in which the connected hexactins are
all simple". Thus sponges with lychniscosan skeletons (all Lychniscosa) and sponges with basidictyonalia (some
Lyssacinosa) do not belong to Hexactinosa. The major diagnostic character separating the members of the
Hexactinosa from the removed group would be "with farreoid or eurctoid type of primary diclyonal skeleton". There
is some defense for adding the presence of sceptrules and/or uncinates to the Hexactinosa diagnosis but this would
only restrict forms like Daclylocalyx Gray, 1867, from this taxon. The modified higher classification of
Hexactinellida is presented in Fig. 18.
Class Subclass Order (Tribe)
Hexactinellida
Hexactinosa
Lyssacinosa
Lychniscosa
Aulocalycoida
Fig. 18. — Taxonomic scheme of highest taxa of recent Hexactinellida.
relationships; dashed lines indicate uncertain relationships.
Complete lines denote well supported
ACKNOWLEDGMENTS
We are grateful to C. VALENTINE and S. STONE (NHM), J. VACELET and N. BOURY-ESNAULT (Observatoire
oceanologique de Marseille). We also much appreciate the help of C. Levi (MNHN) who reviewed this
manuscript, exposed several problems and suggested improvements.
We thank M. Debrenne (MNHN) and J.-F. Dejouannet (1RD) for help with the illustrations, R. von
Cosel and C. Reynes (MNHN) for help with the photos.
This work was supported by grants of the CNRS, the Museum national d'Histoire naturelle (Paris) and the
Royal Society (London).
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Reid, R.E.H., 1958b. A monograph of the Upper Cretaceous Hexactinellida of Great Britain and Northern Ireland Part
II. P aleontographical Society of London, 112: xlvii-xlviii.
RE!D R E H., 1961. — A monograph of the Upper Cretaceous Hexactinellida of Great Britain and Northern Ireland. Part
III. P aleontographical Society of London, 115: 27-48.
Reid, R.E.H., 1963. — Notes on a classification of the Hexactinosa. Journal of Paleontology, 37: 218-231.
RE|D, R^E H., 1964. — A monograph of the Upper Cretaceous Hexactinellida of Great Britain and Northern Ireland Part
IV. P aleontographical Society of London, 117: xlix-cliv.
Reiswig, H.M & Tsurumi, M„ 1996. — A new genus and species of Aulocalydicae, Leioplegma polyphyllon (Porifera:
Hexactinellida) trom the Blake Ridge otf South Carolina, U.S.A. Bulletin of Marine Science, 58: 764-774.
Schmidt, O., 1880. — Die Spongien des Meerbusen von Mexico (und des Caraibischen Meeres). Zweites (Schluss-) Heft
G. Fischer, Jena: 33-90.
Schulze, F.E 1886. — Uber den Bau und das System der Hexactinelliden. Abhandlungen der Koniglichen Akademie der
Wissenschaften zu Berlin, Physikalisch-Mathematische Klasse, 1886: 1-97.
Schulze, F.E. 1887. — Report on the Hexactinellida collected by H.M.S. "Challenger" during the years 1873-1876
Report on the Scientific Results of the voyage of H.M.S. Challenger during tlie years 1873-76, Zoology. 21: 1-513. '
^""Valdivia'7 ySs-/799ie4aClti266ida' Wissenschafliche ErS^nisse der deutschen Tiefsee Expedition a, if dem Dampfer
rABACHNiCK K R 1988. — (Hexactinellid sponges from the mountains of West Pacific.). In: Shirshov P.P. (ed.)
(Structural and Functional Researches of the Marine Benthos.) Academy of Sciences of the USSR. Moscow : 49-64 (in
russian ).
TA Fossn'^nri 0™^' “ AdaPIalion of lhc hexactinellid sponges to deep-sea life. In: Refiner J. & Keufp H. (eds).
Fossil and Recent Sponges. Spnnger-Verlag. Berlin: 378-386.
Z'TTEL K.A 1877 — Studien uber fossile Spongien; I. Hexactinellidae. Abhandlungen der
Akademie der Wissenschaften, Mathematisch-Physikalische Klasse, 13: 1-63.
Eon igl ich Bayerischen
Source MNHN, Paris
rATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CA
4
Porifera Hexactinellida: Amphidiscophora
off New Caledonia
Konstantin R. TABACHNICK
Institute of Oceanology, Academy of Sciences of Russia
Moscow, Russia
&
Claude LEVI
Museum national d'Histoire naturellc
Laboratoire de Biologie des Invertebres marins et Malacologie
55 rue Buffon, 75231 Paris Cedex 05, France
ABSTRACT
EJur-mg the "MUSORSTOM" cruises in the southwestern Pacific, and particularly off New Caledonia. 19 species of
Hexactinellida Amphidiscophora have been found. Twelve species are considered as new: Hyalonema spaiha, H. uncinata,
H. microstauractina, Sericolophus calsubus, S. neocaledonicus, Semperella abyssalis, S. crosnieri, S. varioactinci.
roliopogon micropentactinus, P. claviculus, P. zonecus, and Pheronema pseudogiganteum.
■ , ,W° 01 r’ew sPec,es werc collected near Hawaii: Sericolophus hawaiicus and off eastern Australia: Sericolophus
uc ancus. The description of the holotype of Pheronema giganteum Schulze is completed.
RESUME
Porifera Hexactinellida: Amphidiscophora du large de la Nouvelle-Caledonie.
P *U‘°UrS.de lexamen des collections de spongiaires des campagnes "Musorstom" dans le sud-ouest de I'ocean
i-nnivi autour de la Nouvelle-Caledonie, 19 especes d'Hexactinellida Amphidiscophora vivant entre 200
li , 6 Prolondeur ont dte identifies. Douze especes sont nouvelles : Hyalonema spaiha. H. uncinata
Pnl,ZZ'l 'raCt'n^ Sencol°Phus calsubus. S. neocaledonicus, Semperella abyssalis. S. crosnieri. S. varioactina.
pi g n micropeniaciinus, P. claviculus, P. zonecus et Pheronema pseudogiganteum.
rAl especes nouvelles de Sericolophus provenant 1'une des Hawaii, S. hawaiicus. et 1 autre de la cote est de
complete. ,C“S' som 68alemen' Writes et la description de I'holotype de Pheronema giganteum Schulze est
A Crosnifr'^pH f LE^' Z 200°' ~ Porifera Hexactinellida: Amphidiscophora off New Caledonia. In:
naturelle. 184: ^ ^
54
K. R. TABACHN1CK & C. LEVI
INTRODUCTION
The Hexactinellida of the southwestern Pacific were almost unknown until the cruise of N/O "Vauban" in 1977
off southern New Caledonia (Levi & LEVI, 1982). However, we have two excellent documents on ihc
Hexactinellida of the central Pacific (VON Lendenfeld, 1915) and of Indonesia (Ijima, 1927). The species from
the Japanese coasts are known thanks to Ijima & Okada (1938).
The large collections of deep-water fauna ("Challenger", "Investigator", "Valdivia", "Galathea") do not include
sponges from this area, except two species collected by the "Challenger" near the Kermadec Islands.
The many French oceanographic expeditions around New Caledonia since 1985, obtained an interesting
collection of Hexactinellida Amphidiscophora rich in new species, which are the objective of this study.
The systematic study of Hexactinellida is still very difficult because of the small number of collected
specimens. Most of the described amphidiscophoran species are defined from a single specimen and sometimes
from fragments. If most of the genera are well defined, taxonomy at ihe species level is rather approximative, and
the degree of intraspecific variability is unknown. The works of VON LENDENFELD (1915), based on a very rich
photographic documentation, show the variability of spicular characteristics. Expecting results of the comparative
analysis of isoenzymes and nucleotide sequences, the taxonomist must underscore the observed morphological
variants. The description of atypical, abnormal shapes and the indications of presence vs absence or differences in
spicule densities lead to an artificial grouping of specimens into temporary "species".
Collection of Amphidiscophora from the "MUSORSTOM" expeditions mainly around New Caledonia and in
other areas of the southwestern Pacific is similar to the better known collection from the Indian Ocean. It is,
however, too early to extract pertinent biogeographical information and to determine its major and original
characteristics. This will be easier to achieve when all Hexactinellida genera have been studied.
The collection of Amphidiscophora contains 18 species, including 12 new species. Two other new species:
Sericolophus hawaiicus and Sericolophus cidaricus came respectively from Hawaii and the Eastern coast of
Australia. The type specimen of Pheronema giganteum Schulze (stored in the BNMH) has been redescribed.
Information on the cruises where the samples studied here were collected can be found in:
— Richer de Forges (1990) for Biocal. Biogeocal, Chalcal 2, Musorstom 4, 5 and 6. Smib 1 and 4;
— Richer de Forges & Menou (1993) for Musorstom 7;
— ROUX (1994) for Calsub;
— Richer de Forges & Chevillon (1996) for Bathus 4;
— Grandperrin et al. (1997) for Halipro 2.
No published reports are known concerning Cidaris 1 and Hurl 88 and 92.
When no place of deposit is specified, the material is in the collection of the Museum national d'Histoire
naturelle (MNHN).
The abbreviations for the institutions whose collections were used in this study are as follows:
BPBM - Bernice P. Bishop Museum, Honolulu.
IORAN - Institute of Oceanology, Russian Academy of Sciences, Moscow.
MTQ - Museum of Tropical Queensland, Townsville, Australia.
NHM - The Natural History Museum, London [formerly the Bristish Museum (Natural History)].
USNM - National Museum of Natural History. Smithsonian Institution, Washington.
ZMA - Zoologisk Museum, Amsterdam.
In the list of material examined, the capital letters preceding the station number refer to the gear used:
DC: Charcot dredge; DW: Waren dredge; CP: beam trawl; CC: shrimp otter trawl; BT: benthic fish trawl.
The research vessels' names are in italics, quoted in commas.
Source MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
55
LIST OF SPECIES
The new species are in bold.
AMPHIDISCOPHORA F.E. Schulze, 1886
I. Family HYALONEMATIDAE J.E. Gray, 1857
Genus HYALONEMA J.E. Gray, 1835
Hyalonema (Leptonema) spatha sp. nov.
Hyalonema (Onconema) uncinata sp. nov.
Hyalonema (Pteronema) topsenti Ijima, 1927
Hyalonema (Oonema?) microstauractina sp. nov.
Hyalonema (Cyliconema) pateriferum Wilson, 1904
II. Family MONORHAPHIDIDAE Ijima, 1927
Genus MONORHAPHIS F.E. Schulze, 1904
Monorhaphis chuni F.E. Schulze. 1904
III. Family PHERONEMATIDAE J.E. Gray, 1 872
Genus SERICOLOPHUS Ijima, 1901
Sericolophus calsubus sp. nov.
Sericolophus neocaledonicus sp. nov.
Sericolophus hawaiicus sp. nov.
Sericolophus cidaricus sp. nov.
Genus SEMPERELLA J.E. Gray, 1868
Semperella schultzei (Semper, 1868)
Semperella varioactina sp. nov.
Semperella abyssalis sp. nov.
Semperella crosnieri sp. nov.
Genus POLIOPOGON Wyville Thomson, 1873
Poliopogon micropentactinus sp. nov.
Poliopogon claviculus sp. nov.
Poliopogon zonecus sp. nov.
Genus PHERONEMA Leidy, 1868
Pheronema pilosum Levi, 1964
Pheronema semiglobosum Levi & Levi, 1982
Pheronema conicum Levi & Levi. 1982
Pheronema giganteum F.E. Schulze, 1887
Pheronema pseudogiganteum sp. nov.
Source : MNHN. Paris
56
K. R. TABACHNICK & C. LEVI
DEPTH DISTRIBUTION
The depth ranges where the species were found are given below.
The mean bottom temperatures and salinities observed were:
SYSTEMATIC ACCOUNT
Family HYALONEMATIDAE Gray, 1857
Genus HYALONEMA Gray, 1835
Subgenus LEPTONEMA von Lendenfeld, 1915
Diagnosis (Ijima, 1927). — Dermal pinular ray short-spiny and whip-like, up to 0.800 mm long or shorter
(up to 0.300 mm long) and either short spiny or moderately long-spiny and narrowly plumose in appearance;
rachis thickest at base. Choanodermal macrohexactin and intermedial microhexactin generally present. Without
ambuncinate. Largest macramphidisc relatively narrow, with umbel longer than broad. Sponge with or without
oscular sieve plate or a covering layer to sunken gastral surface.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
51
TYPE Species. — H. campanula von Lcndcnfeld. 1915.
Hyalonema (Leptonema) spatha sp. nov.
Fig, 1-6; Tab. 1
MATERIAL EXAMINED. — New Caledonia. Biocal: stn CP 31, 23°08.70'S. 166°51.55’E. 850-1005 m, 29.08.
1985: HCL 495. — Stn CP 54. 23°10.08'S, 167°43.54'E, 950-1000 m, 1.09.1985: HCL 420-421. — Stn CP 75
22°42'S, 167°23.41 'E, 825-860 m, 4.09.1985: HCL 416-419.
Halipro 2: stn BT 066, 24°44.86'S, 168°29.39'E, 885 m, 19.11.1996: HCL 422.
Loyalty Islands. Biogeocal: stn CP 290, 20°36.9LS. I67°03.34'E, 920-760 m. 27.04.1987: HCL 415.
Calsub: stn 7, W of Lifou Island and N of Santal Bay, 20°48'S. 167°05'E. 970-489 nt, 24.02.1989: HCL 413. 414.
Types. — Holotype : HCL 413 (Calsub, stn 7, W of Lifou Island and N of Santal Bay, 20°48'S, I67°05’E.
970-489 m).
Paratypes : all the other specimens cited above.
Description. — The holotype (fig. 1-2) is cone-like, 100 mm long, with a slightly convex atrial surface and
a short apical cone. The diameter of the lower part is 15 mm. the maximum diameter of 65 mm is between dermal
and atrial surfaces. Basalia arc twisted in a tuft over 60 mm long, about 7 mm in diameter. The body shape of the
paratypes varies: they are all bell-like whereas the atrial surface is flat (HCL 417) (fig. 3) or slightly concave
(HCL 414) (fig. 1-4). The canalar-like depressions on atrial surface are numerous and small in HCL 417, restricted
to 5 and deeply penetrating the body in HCL 414. The length of the body of paratypes is 50-80 mm. the diameter
15-90 mm.
Spicules : The choanosomal skeleton consists of smooth diactins (fig. 6.10) 0.6-1.6/0.006-0.008 mm with
a widening medially. Prostalia lateralia (fig. 6.1 1) are pinular diactins, with four rudimental tubercles in the
middle. Pinular ray 0.21-0.43/0.014 mm similar to dermal or atrial pentactins but the finely pointed ends do not
protrude far beyond the last spines. The ray directed inside the body 0.07-0.38/0.012-0.014 mm is smooth,
sometimes with rare short spines. These pinular diactins are often found among the dermal and atrial spicules. All
the anchorate basalia (if present) are broken and their ends are unknown. Dermalia and atrialia are pinular
pentactins, rarely hcxactins. The base of the pinular ray is thickest at the base. Pinular ray of dermalia (fig. 6.1)
0.085-0.380/ 0.004 mm, the tangential rays are 0.019-0.060/0.003 mm. The pinular ray of atrialia (fig. 6.2-3) is
0.074-0.270/0.004 mm, tangential rays 0.019-0.050/0.003 mm. The whip-like ends of the pinular rays freely
project lar beyond the last spine. The tangential rays are usually smooth, sometimes with spaced short spines.
Hypodermal and hypoatrial pentactins (fig. 6.12) have smooth rays 0.09-0.33/0.015-0.038 mm. the ray directed
inside the body is usually longer than tangential ones.
Microstauractins (fig. 6.4-9) are sword-like, with rays covered with dense spines, three rays are short 0.008-
0.026/0.003 mm and one is long 0.044-0.145/0.003 mm. Two rays of these stauractins are sometimes long,
rarely all lour rays are short and a spicule is similar to stauractin acanlhophore of some other representatives of
Hyalonema or very rarely these spicules have form of paratetractin.
The amphidiscs (fig. 6.13-14) fall into two groups, despite of some overlap of maximal length of
micramphidiscs and minimal length of macramphidiscs. They are sometimes hardly distinguished from each other
in shape and size. Nevertheless, most of them are well distinguished by size classes and the macramphidiscs often
have umbels more elongate than the micramphidisc (fig. 6.15) which umbels arc nearly equal in length
and diameter. The macramphidiscs are rare in the specimen HC1 415. Total length of macramphidiscs
0.025-0. 148 mm, umbel length 0.008-0.063 mm, umbel diameter 0.008-0.059 mm. Total length of
micramphidiscs 0.014-0.041 mm, umbel length 0.004-0.015 mm, umbel diameter 0.005-0.013 mm. Shafts
of macramphidiscs and micramphidiscs covered with spines. Several macramphidiscs, probably of foreign
origin, were found in one specimen. They have very long teeth which cross the equator and overlap with the
opposite teeth. One huge macramphidisc, probably foreign too, 0.315/0.1 1 1/0.104 mm. was found in specimen
58 K. R. TAB ACHNICK & C. LEVI
Etymology. — The specific name alludes to the shape of microholactine spicules.
Fig. 1-5. — Hyalonema ( Leptonema ) spaiha sp. nov.: 1-2 (HCL 413); 3 (HCL 417); 4-5 (HCL 414). Scales = 2 cm.
Source MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
59
Fig. 6. Spicules of Hyalonema ( Leptonema ) spailia sp. nov.: 1, dermal pentactin (HCL 413); 2-3. atrial pentactins
(HCL 413); 4-6. microstauractins (HCL 413); 7-9. microstauractins (HCL 418); 10. choanosomal diaclin (HCL 413);
ll.prostalia lateralia - pinular diactin (HCL 413); 12. hypodermal pentactin (HCL 413); 13. macramphidisc
(HCL 413); 14, macramphidisc (HCL 417); 15. micramphidisc (HCL 413).
60
K. R. TABACHNICK & C. LEVI
Remarks. — The species is referred to the subgenus Leptonema. The most outstanding feature of the
new species is the presence of sword-like stauractins. In several species of Amphidiscophora the derivatives of
hexactins to monactins are usually known as additional to the hexactins or pentaetins spicules (for example
Semperella schultzei) (MARSHALL & MEYER, 1879). Another species of the amphidiscophoran family
Hyalonematidae, Lophophysema australicum sp. now, has mainly whip-like monactins (TABACHNICK & Levi,
1999).
Subgenus ONCONEMA Ijima, 1927
DIAGNOSIS. — Choanosomal macrohexactins and microhexactins always present. Pinular pentaetins with
relatively short distal ray, terminating with an apical cone. Macramphidiscs with umbels slightly shorter than 1/3
the whole length of the spicules. Uncinate rhabdoxydiactins in the peripheral parts of the body.
Hyalonema (Onconema) uncinata sp. now
Fig. 7-9; Tab. 2-3
MATERIAL EXAMINED. — New Caledonia. BlOGEOCAL: stn CP 283, 2r22.25’S. ]66°31.07'E, 2375-2370 m,
26.04.1987: HCL 423-424.
TYPES. — Holotype: HCL 424 (BlOGEOCAL, stn CP 283, 21°22.25'S. 166°31.07'E. 2375-2370 m).
Paratype: HCL 423.
Description. — The holotype (fig. 7-8) is similar in shape to H. (Onconema) agassizi "form A"
(Lendenfeld, 1915, pi. 41.2) and to H. ( Onconema ) obtusum var. gracilis (Lendenfeld, 1915, pi. 33.16).
The body is oval or lemon-like, 17 mm long and 17x8 mm in maximal diameter. The apical cone is surrounded
by a circular wall which ends in a narrow frill, with a circular margin. The circular wall is separated from the
apical cone by a circular fissure which is the atrial cavity. The outer surface is smooth. The paratype is more
damaged than the holotype and measures approximately 18 mm long and about 10x4 mm in diameter.
Spicules : Choanosomal skeleton of smooth diactins (fig. 9.11-12), 0.5-1.7/0.004-0.019 mm with or with¬
out medial widening. Uncinates diactins (fig. 9.6-7) with medial widening and uncinates monactins (fig. 9.8.10)
without any widening are 0.3-0.7/0.004-0.090 mm. Some of these spicules could be considered as ambuncinates;
they have very short spines, which seem to be bent to the center of the spicule. The uncinates diactins are rare
in the holotype and abundant in the paratype, the opposite situation is observed with the uncinates monactins.
The special description of the types of uncinates is given in the "REMARKS" later. Choanosomal hexactins
are smooth with rays. 0.18-0.36/0.008-0.015 mm, which can be unequal in each spicule. Dermalia and atrialia
are pentaetins, very rarely hexactins. Dermal pentaetins can be separated into two types. The first type is a pinular
pentactin (fig. 9.1) which has the pinular ray 0.076-0.137/0.004 mm, thickest at base, with long or rarely
short spines and thin, whip-like end which protrudes far beyond the last spines. The tangential rays are covered
with dense short spines, they are 0.030-0.091/0.004 mm. These spicules are similar to the dermal, pinular
pentaetins from the lower part of the body of H. ( Onconema ) agassizi (Lendenfeld. 1915). The second type
which is also considered to be dermal pentaetins (fig. 9.2-3) are spicules with nearly equal rays, 0.084-
0. 122/0.004 mm, similar to the tangential rays of dermal pentaetins. Similar spicules were described in
H. ( Onconema ) agassizi as minute pentaetins. Furthermore, the rays of acanthophores in H. ( Onconema ) obtusion
arc similar in shape to the second type of dermal pentaetins of H. ( Onconema ) uncinata. The atrial pinular
pentaetins (fig. 9.4) differ strongly from the dermal ones. The pinular ray 0.129-0.167/0.014 mm is spindle-like,
often thickest in the middle, with relatively short spines, the end has an apical cone. The tangential rays
0.019-0.034/0.006 mm are short, usually with sparse small spines or entirely smooth. Their ends are rounded or
finely pointed.
Source MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
61
Fig. 7-8. — Hyalonema ( Onconema ) uncinaia sp. nov. (HCL 424). Scales = I cm.
The hypodermal and hypoatrial pentactins have smooth rays, 0.14-0.68/0.006-0.038 mm, the ray directed
inside the body is usually longer than the tangential ones. The tangential rays can be of different length and
sometimes with unusual spherical ends.
Microhexactins (fig. 9.17) have 0.023-0.065/0.002 mm rays. The rays are covered with dense minute spines
and often they have curved ends, or rarely straight.
Amphidiscs fall into two groups: Macramphidiscs (fig. 9.15) arc 0.236-0.334 mm long, umbel 0.084-
0.122 mm long and 0.061-0.099 mm in diameter. The shaft has a whorl of spines in the middle and some spines
in other places. Umbel's teeth have lancet-like ends. Total length of micramphidiscs (fig. 9.16), 0.014-0.032 mm,
umbel length 0.004-0.011 mm, umbel diameter 0.005-0.007 mm. Shafts of micramphidiscs smooth or covered
with spines.
Remarks. — This species belongs to the subgenus Onconema Ijima. A great similarity is found between
this and previously described species: Hyalonema (O.) agassizi and H. 10.) obtusum known from the central-
eastern Pacific at 4069-4504 m depth (Lendenfeld, 1915). The new species is closer to H. (Onconema) agassizi
than H. (O.) obtusion. The diagnostic feature of the new species is the presence of dermal pentactins with
pinularray thickest at the base and with whip-like outer end. Among other peculiarities is the narrower range
ol variation of both types of amphidiscs. Each type of macramphidisc and micramphidisc of LENDENFELD's
species is divided into two size classes, which notably overlap by the borders. Atrial pinular pentactins of
H. (Onconema) uncinata are dissimilar to other species due to the proportion of their rays and slighter because of
their measures. The measures of other spicules in all known species of the subgenus Onconema have similar
parameters.
Etymology. — The species name refers to the presence of the uncinate spicules.
62
K. R. TABACHNICK & C. LEVI
0.2 mm 0.4 mm _ 0.05 mm _
1-12, 15 13^14 16-17
Fig. 9. — Spicules of Hyalonema (Onconema) uncinata. sp. nov.: 1. dermal pinular pentactin (HCL 424); 2-3. dermal
pentactin (HCL 424); 4, alrial pinular pentactins (HCL 424); 5. atrial pinular pentactins (HCL 423); 6-7. uncinates
diactin (HCL 424); 8. 10. uncinates (HCL 424); 9. ambuncinate (HCL 424); 11-12. diactins (HCL 424);
13. hypodermal pentactin with different rays (HCL 424); 14. choanosomal hcxactin (HCL 424); 15, macramphidisc
(HCL 424); 16, micramphidisc (HCL 424); 17. microhexactin (HCL 424).
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
63
Subgenus PTERONEMA Ijima, 1927
Diagnosis. — Dermal pinular ray plumose or with moderately long lateral spines giving a conical or
spindle-like shape to their entire ray under 0.170 mm length. Rachis thickest at base. Ambuncinate present.
Microholactin hexactin sometimes pentactin and stauractin. Macramphidiscs with umbel broader than long.
Hyalonema (Pteronema) topsenti Ijima. 1927
Fig. 10-12; Tab. 4
Hyalonema < Pteronema ) topsenti Ijima. 1927: 61, pi. 1 fig. 5-6. pi. 2 fig. 11, pi. 3 fig. 1-10.
Hyalonema topsenti - LEVI & Lfivt. 1989: 36, fig. 5.
Fig. 10-12. — Hyalonema f Pteronema ) topsenti Ijima, 1927 (HCL 425). Scales = 2 cm.
nlItn™EX,^llNED' ~ ' New Caled«nia- Dragages "Vauban": stn 39, dragage 9. 22°29'S. 166°23'E. 375-
jju m, /.Uo. 1978; HCL 427.
Calsub: stn 9, 20°53’S, 167°03'E, 602 m, 27.02.1989: HCL 425
Bathus 4: stn CP 892, 21°01'S, 164°27'E. 580-602 m. 2.08.1994: HCL 426.
Sm n°w iiwld«.oMUSORSTOM 6: sln CP 490' 20°48.88’S, 167°06.13'E. 750 m. 24.02.1989: HCL 428. —
Stn DW 493. 20°48.35'S, 167°05.80'E. 700 m. 25.02.1989: HCL 429.
64
K. R. TABACHN1CK & C. LEVI
DESCRIPTION. - The complete specimens are HCL 425 and HCL 426. The first is oval, 55 mm long and
iS-nearIy quadrangular and the surfact see™ divided hTfour
is not an nare itT Tl n d p The lattice 's similar to Hyalonema globus Schulze. The atrial cavity
Basa ia are 10 ) im n 1 f ^ T ^ SUbatrial Canals pcnetrate vertical|y deep inside the bod/
Kim tS nonce HrT 5? t ° 7'? Str°ng 0r denticular rid8es llke in Cholaronema sibogae
S hulze I If l t CS TCra' CharaCterS With IWonema topsenti Ijima and Hyalonema globus
Schulze. Ijima (19.7) already suggested a close affinity between these species.
UMAHWUrZ TSl g,enera"y' 'h?SeTSp0ngeS aPe VCry Similar t0 sPcc'mcns of the same species described by
IJIMA (1927) from the Indonesia and by Levi & Levi ( 1 989) from the Philippine Islands
The external shape of this spec.es displays a high variability: presence of apical cone in one specimen and atrial
depression in another. This variation is large enough and may be rather observed amon» related ^ ^aecies thin
ctween intraspecific populations. All specimens are nearly identical in spicule content despite being collected far
Zn, 1 , , if rCimenS °ff NCW Caled°nia have m*croPentactins and microstauraSs iS of
microhexactms, which the previously known specimens exhibited.
Distribution. — Indonesia, Philippines, New Caledonia.
Hyalonema (Oonema?) microstauractina sp. nov
Fig. 13-16; Tab. 5
1.0949A8T5:RHCL vfSm TeVVIT? S’44'5'5' '^54.94'E. 1490-1620 m.
Stn CP 61, 24° 1 0.67'S, ,67°33.65'E, 1070 m 2 .09 1985 HCI^,’ ^ 2-°9'I985: HCL 432 61 433' ~
wfnE0CAa T CP 2l4- 22°43 09's’ 166°27-19'E, 1665-1590 m, 9.04 1987- HCL 435
HCLW4a34.S and FU‘Una IS,andS' MUS0RST0M 7' «" CP 564. N=46.,0'S. 178*27.4™, .0,5-, 020 m, 20.05.1992:
Types. Holotype: HCL 432 (Biocal, stn CP 60, 23°58.87'S, 167°08.43’E
raratypes: all the other specimens mentioned above.
, 1530-1480
m).
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
65
long and 40-70 mm in diameter. The paratype HCL 433, which contains the greatest number and variation of
macramphidiscs, is a thin lamellar fragment 45 x 75 mm. No notable canals were observed beneath the atrial
surface.
' ' Spicules ot Hyalonema ( Oonema ) microstauractina sp. nov,: 1-2, small dermal pentactins (HCL 433);
7- 'Small atrial pentactins (HCL 433); 5. large dermal pentactin (HCL 433); 6, dermal pinular diactins (HCL 433);
. rmal pinular diactins (HCL 432); 8-9, choanosomal diactins (HCL 432); 10. hypodermal pentactin (HCL 432);
II, choanosomal hexactin (HCL 432).
Spicules-. Choanosomal skeleton of smooth diactins (fig. 15.8-9), 0.76-1.8/0.006-0.015 mm. with a widening
in the central part. Choanosomal hexactins (fig. 15.10-11) smooth with rays 0.25-0.49/0.01 mm. Dermalia and
ria ia pinu ar pentactins (fig. 15.1-2) of one or two kinds and pinular diactins. Small dermal pentactins have
66
K. R. TABACHNICK & C. LEVI
pinular ray with apical cone and spines longest near the outer end or in the upper part. Pinular ray of small dermal
pentactin 0.076-0.258/0.008 mm at base, 0.010 mm near the outer end; tangential rays 0.023-0.046/0.006-
0.008 mm, covered with dense small spines. Large dermal pinular pentactins (fig. 15.5), with pinular ray spindle-
like 0.167-0.517/0.016-0.018 mm. The longest spines are medially situated and the ray has an apical cone or is
finely pointed. Tangential rays of large dermal pinular pentactins 0.030-0.122/0.008 mm, densely covered with
short spines. Dermal pinular diactins (fig. 15.6-7) have the pinular ray similar to large pinular pentactins in shape
and size (0.213-0.608/0.01-0.02 mm); the other ray (0.182-0.608/0.008-0.01 mm) is smooth or with rare spines
near the outer end. These diactins have four rudimentary median tubercles and are similar to the large pentactins.
Small atrial pentactins (fig. 15.3-4) have spindle-like pinular ray 0.084-0.228/0.01-0.015 mm and tangential rays
0.023-0.068/0.01 mm. Large atrial pentactins are similar to dermal ones, with pinular ray 0. 1 37-0.532/0.0 1 6-
0.018 mm and tangential rays 0.030-0.175/0.008 mm. Atrial pinular diactins with spindle-like pinular ray 0.205-
0.836/0.01-0.02 mm, the other rays 0.137-0.851/0.008-0.01 mm. All these dermal and atrial spicules are present
in specimens HCL 433 and in HCL 432. Only small pentactins were found in HCL 435, while pinular diactins
were present in this specimen as in broken fragments. In other specimens the small dermal and atrial pentactins
seem to be absent. Hypodermal and hypoatrial pentactins have smooth rays 0.3-0.5/0.01-0.03 mm.
Microstauractins (fig. 16.26-36) prevail to other spicules with similar rays from microhexactins to micro-
monactins. The stauractins often have a rudimentary fifth ray or a spine which lies in the central crest. In the
specimen HCL 433 microstauractins usually have one pair of opposite rays longer than the other. Other specimens
often have the stauractins with equal rays and more pentactins and hcxaclins. The 0.01 1-0. 1 12/0.005 mm rays are
densely covered with short spines.
The amphidiscs fall within three groups. Total length of macramphidiscs (fig. 16.1-19): 0.038-0.350 mm,
umbel length 0.015-0.1 14 mm, umbel diameter 0.021-0.129 mm. The shaft of largest amphidiscs has a whorl of
tubercles in the middle but sometimes these spicules have tubercles in other places. The smallest kinds of
amphidiscs were found in HCL 433. They usually have smooth shafts, often represented by amphidiscs derivatives
(fig. 16.12-18). paradises, hemidiscs, paradises without serrated or toothed discs, amphidiscs with additional rows
of teeth, sometimes with teeth reduced in number, sometimes with teeth of different length. The umbel of
macramphidiscs has 6, rarely 5, teeth which are oval in shape. Mesamphidiscs (fig. 16.20-23) have numerous teeth
and elongate umbel, the teeth of opposite umbels may meet at the equator. The shafts of mesamphidiscs are
densely covered with spines and have a whorl of spines and a widening in the middle. Total length of
mesamphidiscs 0.040-0.153 mm, umbel length 0.013-0.072 mm, umbel diameter 0.01 1-0.054 mm.
Micramphidiscs (fig. 16.24-25 ) have umbels nearly equal in length and diameter, shafts with spines in the middle
and some spines in other parts of the shaft. Total length of micramphidiscs 0.0 1 1-0.050 mm, umbel length 0 004-
0.016 mm, umbel diameter 0.004-0.020 mm. In HCL 433, HCL 432 and HCL 435 the largest mesamphidiscs can
be larger than the smallest macramphidiscs, the same phenomenon is observed in mesamphidiscs and
macramphidiscs in HCL 433, HCL 432 and HCL 431. In these cases amphidiscs are referred to one or another
kind, according to their umbel proportion. In other specimens the different types of amphidiscs have no overlap in
their length parameters and they are easily differentiated by their size.
Remarks. This new species can be easily distinguished from the many other species of the genus by the
presence of microstauractins with rays of different length which predominate over other microhexactins derivatives.
It is the only feature which unite all these very dissimilar specimens into one species. The greatest similaritv is
observed between HCL 432 and HCL 433 collected at the same station. The other group includes all other
specimens. Maybe, these two groups should be divided into different subspecies. The presence of stauractins is
reported for an another genus (or subgenus) of Hyalonematidae: Chalaronema (Ijima, 1927). H. microstauractina is
tentatively placed into the subgenus Oonema , but this new sponge possesses some features of the other subgenera,
Paradisconema and even Cyliconema. The main diagnostic feature of the doubtful subgenus Paradisconema is the
mandatory presence of rare paradises among micramphidiscs (Ijima, 1927), whereas in H. microstauractina
paradises belong to the kind of smallest macramphidiscs and were found only in two out of six specimens. The
outer ends of pinular pentactins in H. microstauractina are similar to those of both Paradisconema with the rachis
o the ray of dermal pinular pentactin "thickest at base, and tapering towards tip" [see H. ( Paradisconema ) vosmaeri
Source MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
67
Fig. 16. — Spicules of Hyalonema (Oonema) microstauractina sp. nov,
1 -19’ fnac^mpfiidises and derivatives: 1 (HCL 430); 2 (HCL 432); 3-19 (HCL 433).
2-18, paradises; 16, 18. paradises with unserrate and not-toothed umbels; 19. macramphidisc with unserrate and
not-toothed umbels.
20-23, mesamphidiscs: 20-21 (HCL 434); 22 (HCL 430); 23 (HCL 433).
24-25. micramphidiscs (HCL 433).
26-36. microstauractins and derivatives: 26, 30, 34, 36 (HCL 433) ; 32 (HCL 434); 33 (HCL 431),
68
K. R. TABACHNICK & C. LEVI
with some rachis slightly swollen in the middle and an apical cone at the end (IJIMA, 1927)] and to Oonema with
dermal pinular ray wearing the rachis "distinctly swollen in the middle parts or, toward the end, ending in an apical
cone" (Ijima, 1927). As in Oonema species, there are two kinds of small and large macramphidiscs, i.e.
H. sequoia, H. densum, H. crassipinulum, H. bianchoratum (all LENDENFELD, 1915). In these species some of
the small macramphidiscs are abnormal: with a tendency to develop paradises by having teeth of different length,
by having curved and additional teeth, some macramphidiscs may be found with one rounded outer part of the
umbel, and other being conically pointed. Such abnormal spicules are also known in H. umbraculum and
H. aequatoriale (LENDENFELD, 1915), which were initially described as subgenus Skianema but that later turned to
be junior synonyms of Oonema (Ijima, 1927). Two types of pinular pentactins are known in H. (Oonema)
bipinnulum (LEVI, 1964). Some specimens of H. microstauractina , which have the only type of dermal pinular
pentactins, have no small macramphidiscs and hence the paradises are similar to the subgenus Cyliconema. The
only notable difference is that the pentactin pinular rays of Cyliconema species have a finely pointed end which
usually protrudes far beyond the last spines. Thus the new species presents the features that draw together.
Paradisconema, Oonema and Cyliconema subgenera.
Etymology. — The species name refers to the cruciform microholactin spicules.
Subgenus CYLICONEMA Ijima, 1927
Diagnosis. — Dermal pillule ray short spiny and slender, in most cases under 0.250 mm length or longer and
whip-like. Rachis thickest at base. Without ambuncinate. Microholactin hexactine.
Sponge more or less cup-like, without an oscular sieve plate or a gastral covering layer, so that the excurrent
canals all open freely into the gastral cavity or on the gastral surface, which may be nearly externally exposed or
even entirely everted.
Hyalonema ( Cyliconema ) pateriferum Wilson, 1904
Fig. 17
Hyalonema pateriferum Wilson, 1904: 28, pi. 1 fig. 1-13.
Hyalonema (Phialonema) pateriferum - von LENDENFELD, 1915: 362, pi. 50 fig. 6-15, pi. 51 fig. 1-28. pi. 52 fig. 1-29.
Coscinonema pateriferum - Levi, 1964: 86, fig. 32.
Material EXAMINED. — Loyalty Islands. CALSUB, E. slope off Lifou Island, 20°35.4'S, 167°12E, 2697 m.
23.02.1989: HCL 436.
Description. — This fine bright white saucer-shaped specimen is 90 mm long, 70 mm wide and 10 mm
thick. The root-tuft reaches 330 mm long.
Its shape and structure are similar to the eastern Pacific sponges described by WILSON (1904) and VON
Lendenfeld (1915). They are flat but one surface is slightly convex. It is very difficult to observe the excurrent
orifices but the marginalia pinular diactins are found in the apical part of the sponge.
Spicules : The spiculation is quite similar to WlLSON’s specimens. Distal ray of gastral pentactins range 0.300-
0.410 mm long; average width is 0.088 mm. Characteristic macramphidics 0.130-6.220 mm long; umbels 0.020-
0.035 mm long and 0.060-0.070 mm broad. Teeth broad. 6 in number. Shaft with few central tubercules.
Abundant mesamphidics 0.025 -0.066 mm long. Scarcely found microxyhcxaclins with rays, straight or sligthly
bent. Acanthophora similar to the type present in the collar pad.
Remarks. Von Lendenfeld (1915) created the subgenus Phialonema for Hyalonema pateriferum Wilson
and Hyalonema brevancora Lendenfeld. But Luma (1927) maintains only H. brevancora in Phialonema and assigned
H. pateriferum to the subgenus Coscinoderma. used further by Levi (1964).
Source : MNHN , Pan’s
AMPHIDISCOPHORA OFF NEW CALEDONIA
69
In fact, Hyalonema pateriferum does not have ambuncinates and the distal rays of pinular pentactins are short
spiny and slender as in Cyliconema. If we do not keep H. pateriferum in the doubtful subgenus Phialonema defined
by only one character, this species is referable to Cyliconema.
Distribution. — Northern Peru 82-83°W - eastern tropical Pacific 104°- 1 17°W; 381 1-4063 m depth. Bottom
temperature: 1.5-3°C.
Discussion. — One of the most
outstanding features in the material
examined is the presence of uncinates
diactins, uncinates (or uncinates
monactins) and spicules similar to
ambuncinates in H.(Onconema)
uncinata. According to Ijima (1927)
the family Hyalonematidae is char¬
acterized, besides other features, by
the "absence of the uncinates" (p. 42)
or "as a rule without uncinate"
(p. 41). Instead of uncinates. which
are common for the close family
Pheronematidae, it is possible to find
ambuncinates in the Hyalonema
subgenera as in Pteronema. Cosci-
nonema and Euhy alone ma and
uncinate diactins ("uncinate rhabdo-
diactins") in the subgenus Onconema.
The poorly described H. pellucidum
(Ijima, 1927, see footnote of p. 6) has
some uncinates. called "ambuncinate
diactin with all the spines bent in the
same direction". Relatively rare
uncinates of H. agassizi were
especially discussed by Ijima (1927).
They have some vertically or even
retroverted inclined spines near one
end of a spicule. Such spicules are
numerous and even prevail over the
other types in H. (Onconema)
uncinata. Some of these uncinates,
the largest ones, have one smooth end
and they are similar to some types of
uncinates, which are present in the
family Pheronematidae. The smaller
type of uncinates are entirely covered
with spines, and they are similar to
those described for H. (Onconema)
agassizi and especially discussed by
JIMA. The smaller type of uncinates have a widening in their central part and only the smallest ones have
rudiments in shape of two tubercles. Does this morphological series of uncinates reflect their ontogenesis and
p \logencsis? If it is a result of ontogenesis, the observed morphological differences represent development stages
o t e adult. If it is a result of recapitulation, the smallest type of uncinate is likely to be derived from the "archaic
Fig. 17. Hyalonema ( Cyliconema ) pateriferum Wilson. 1904 (HCL
436). Scale = 1 cm.
70
K. R. TABACHNICK & C. LEVI
stauractin" (Ijima, 1927), and the largest type of uncinates which are described as uncinates monactins are
uncinates diactins. Are the uncinates of Pheronematidae homologous to these large uncinates diactins? If so, the
existence of uncinates monactins is questionable. Formally, according to morphology (asymmetry of spines) they
are monactins, being however real diactins according to their origin. The only argument to their monactin nature is
the absence of axial crest traces inside these spicules. Hence, we can conclude that it is very likely that the
morphological series of uncinates observed in the subgenus Onconema displays some features of its phylogenesis
and all the uncinates observed in other groups of Hexactinellida are likely to be uncinates diactins and only using
the formal morphological criteria, it is possible to call them uncinates monactins. In the ontogenesis of Farrea the
youngest uncinates show a widening in the middle (OKADA, 1928) and hence the uncinates of Hexasterophora may
be also considered to be uncinates diactins according to their origin. Moreover, uncinates diactins were found
together with usual uncinates (uncinates monactins) in mature Tretocalyx polae (Schulze, 1900). It is very likely
that uncinates diactins, uncinates monactins and ambuncinates are close types of spicules, homologous not only in
Hyalonematidae and Pheronematidae but even among Amphidiscophora and Hexasterophora. It is very difficult to
come to a conclusion in favour of separate origin of uncinate-like spicules and some other smooth choanosomal
spicules.
These facts point out the similarities of evolutionary trends even in quite separate branches of Hexactinellida,
when similar but independently evolved structures give parallel series of variability.
Family MONORHAPHIDIDAE Ijima, 1927
Genus MONORHAPHIS Schulze, 1904
Diagnosis (after Ijima, 1927). — Amphidiscophora with uncinate, choanosomal supporting spicules
predominantly tauactins, elongate in the complete axis and sometimes passing over into rhabdodiactin form by
suppression of the unpaired ray; basalia a single, excessively strongly developed needle. Sponge body somewhat
like a laterally compressed cylinder, with excurrent canals orifices, irregularly distributed on the surface, and
showing on one body edge a longitudinal series of niche-like depressions covered with a sieve-like lattice.
Monorhaphis is one of the most peculiar genera of Hexactinellida. Among the specimens collected by the
"Valdivia" in the Indian Ocean, Schulze (1904) identified two species, M. chuni and M. dives, based on the
presence or absence of mesamphidiscs and abnormal micramphidiscs.
Since then, other specimens collected in Indonesia (IJIMA, 1927) and the Indian Ocean (BURTON. 1959) have
been linked to these two species. We agree with SCHULZE (1886, 1893) and Ijima (1927) that the very small
fragment of "Hyalonema sp." collected off Luzon (Philippine Islands) renamed by SCHULZE as "Hyalonema
fruticosum" belongs to Monorhaphis.
More recently, Li JlNHE (1987) described M. intermedia, a new species from the China Sea. It has all types of
spicules which exist in the two species described by SCHULZE.
Finally, Monorhaphis was observed during dives of the submarine "Cyana" in the Loyalty Basin (ROUX et ai,
1991a, 1991b). It was also found on the Norfolk Rise (LEV! et al., 1989) and on the outer slope of the Great
Barrier Reef (HOOPER & WlEDENMAYER, 1994).
The Musorstom and Cidaris I cruises provided complete specimens and numerous raw basalia. The study of
these collections allowed us to determine that M. dives and M. intermedia are junior synonyms of M. chuni
Schulze, the type species of the genus (designation by de Laubenfels, 1936).
Monorhaphis chuni Schulze, 1904
Fig. 18; Tab. 6-9
Monorhaphis chuni Schulze, 1904: 112, fig. 3-4, pi. 40-42, 44-48. — Ijima, 1927: 37. fig. 3, pi. 6 fig. 11-12.
Source : MNHN, Paris
AMPH1DISC0PH0RA OFF NEW CALEDONIA
71
Monorhaphis dives Schulze, 1904: 121, pi. 43 fig. 1-20. — Burton, 1959: 176.
Monorhaphis intermedia Li Jinhe, 1987: 130, pi. 1-2.
MATERIAL EXAMINED. — Red Sea. "Anton Brum": stn 7-370 D, 28°00'N, 35°60'E, 880 m: kt476 (USNM).
Somalia. "Valdivia": stn 257, 1°48.2'N, 45°42.5'E, 1644 m: fragment of the holotypc of Monorhaphis dives
(NHM 1908.09.24,065). — Stn 264: 6°18.8'N, 49°32.5'E, 1079 m: fragment of Monorhaphis chuni (NHM 1908.09.
24.064).
Madagascar (NW coast). "Vitjaz II": stn 2601, 12°25.04'S, 48°08.00'E, 700 m: 5/2/1351, 5/2/1422, 5/2/1352
(10RAN). — Stn 2606. I2°26.70’S. 48°06.40'E, 700-720 m: 5/2/1408 (TORAN).
Australia (NE coast). Cidaris I: stn 1-4, 18<I08.69’S, 147°33.79'E, 962-966 m. 6.05.1986: p540 (MTQ). —
Stn 4-1, 1 8° 1 1.52'S. I47°52.12'E, 1012-998 m, 6.05.1986: fr767 (MTQ). — Stn 8-1, 18°07.82'S, 148°15.39'E. 1115-
1119 m, 7.05.1986: fr867.1 (MTQ).
Chesterfield Islands. Musorstom 5: stn CP 324, 21o15.01'S, 157°51.33'E, 970 m, 14.10.1986: HCL 408-409.
New Caledonia. Biocal: stn CP 61, 24°10.67'S, I67°33.65'E, 1070 m. 2.09.1985: HCL 445. — Stn CP 75.
22°20.42'S, 1 67°23.4 1 'E, 825-860 m, 4.09.1985: HCL 405-407.
Loyalty Islands. Calsub: stn 7, W of Lilou Island and N of Santal Bay, 20°48'S, 167°05'E, 970-489 m,
25.02.1989: HCL 41 1. — Stn 8, W of Lifou Island and N of Santal Bay. 20°48'S, 167°05'E, 880-516 m. 26.02.1989:
HCL 412. — Stn 13, 21°26'S, 166°22.7'E, 1807-1567 m, 4.03.1989: HCL 410.
Types. — Those of Monorhaphis chuni are perhaps deposited at the Hamburg Museum.
A fragment of the holotype of Monorhaphis dives (" Valdivia ", stn 257: 1°48.2’N, 45°42.5'E, 1644 m) is
deposited at The Natural History Museum, in London (NHM 1908.09.24.065).
Description. — Most of material examined consisted of fragments, but the typical shape of the genus may be
often deduced. Only two sponges are complete specimens similar in shape to those described as M. chuni
(Schulze, 1904) or observed and photographed from the submarine "Cyana" (ROUX et al., 1991a, 1991b).
Spicules: Choanosomal spicules are chiefly tauactins (triactines, typical for the genus and family) with the
opposite rays longer than the third one. These tauactins occur together with paratetractins and stauractins with two
opposite rays notably longer than the other ones. Uncinatcs arc also present in all specimens. Basalium, if present,
is a single giant monaxonal diactine spicule.
Hypodermalia and hypoatrialia are represented by pentactins (fig. 18.1), with the ray directed inside the body
longer than the tangential ones; the tangential rays are similar in length.
Dermalia and atrialia are pinular pentactins. They are very similar in shape and size to each other (see Tab. 6-
9). The pinular ray of these pentactins is slightly widened toward the end. often with an apical cone which
protrudes beyond the last spines. Canalaria are pinular pentactins observed in some, probably the biggest
specimens. These pentactins are shorter than the dermal and atrial ones; the pinular ray is thicker at the base with
an outer end finely pointed, the spines are notably shorter than the spines of dermal and atrial pentactins. All the
spicules mentioned above are similar in all the examined specimens and in all the known descriptions (for
references see synonymies). They can never be used for the species definition (see Tab. 9).
Microhexactins (fig. 18.3-5) (rarely micropentactins and microstauractins) are usually numerous in all the
examined specimens. They are rare in kt476. The microhexactins have rays covered with more or less dense short
spines.
The micramphidiscs (fig. 18.14-16) are more or less similar in all of these species. They are mostly similar
in shape and size but in some specimens they may have more or less spinous shafts. Since curious
micramphidiscs are rarely found among the normal ones it is possible to mention the following cases, but
they seem to have no taxonomic significance. Micramphidiscs in the form of paradise were observed in specimen
kt4 1 7, the ratio of its umbels diameters is 0.7. Micramphidiscs with one tooth at each umbel (sigmoidal in shape)
were lound in NHM 5/2/1408, whereas in HCL 41 1 only two teeth were found at one side while at the other
there was no umbel but a small spherical outer end instead. Micramphidisc in the form of tylodisc (fig. 18.18)
with one umbel reduced to spherical end while the other is "normal" were found in NHM 1908.09.24.065.
Micramphidisc (fig. 18.21) with one hemispherical umbel and the other conically pointed was found in
HCL 410. And finally hexadiscs (fig. 18.13) corresponding to micramphidiscs in shape and size were found
in fr767.
72
K R TABACHNICK & C. LEVI
Fig. 18. — Spicules of M. chuni Schulze, 1904: 1, hypodermal pentactin (5/2/1351); 2, uncinate (5/2/1422);
3-4, microhexactins (5/2/1351); 5. microhexactins (fr867. 1 .); 6, amphidisc intermediate between macramphidiscs
and mesamphidiscs (fr867. 1 .); 7, barrel-shaped mesamphidisc (HCL 409); 8-9. mesamphidiscs with fused teeth
(HCL 409) (9, optical section); 10-11. amphidiscs intermediate between mesamphidiscs and micramphidiscs
(10, HCL 411; 11, HCL 408); 12, hexadiscs (HCL 41 1); 13, hexadiscs (fr767); 14-15. micramphidiscs (5/2/135 1 );
16. abnormaly small micramphidisc (fr867.1); 17, hemidisc (1908.09.24.064); 18, tylodisc (1908.09.24.065);
19-20. paradises (1908.09.24.064); 21, micramphidisc with different umbels (HCL 410).
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
73
Remarks. — Species of Monorhaphis were distinguished earlier by the presence or absence of macramphidiscs
and mesamphidiscs. The macramphidiscs are usually notably larger (over 0.3 mm in length) than mesamphidiscs,
and have fewer teeth (about 8) in each umbel; they often have smooth shafts or are provided with sparse short
spines. Among macramphidiscs, forms with all or some of opposite teeth fused at equator (barrel-shaped. Ijima.
1927) are present. All mentioned exceptions were found in all the observed materials. Mesamphidiscs arc
distinguished by their length, shafts densely covered with spines and often more elongate umbels than in
micramphidiscs. Sometimes they can be found in the form of hexadiscs. Rarely mesamphidiscs (fig. 18.8-9) are
found with fused opposite teeth as macramphidiscs: HC1 409, 406, 405. In one specimen. HCL 409, all the teeth
of some mesamphidiscs were fused into a continuous silica layer. All the exceptions of mesamphidisc shapes are
very similar to those of macramphidiscs. Moreover, forms of amphidiscs which are transitional between
macramphidiscs and mesamphidiscs, and which could be hardly referred to as these kinds of spicules were widely
observed in HCL 409, 406, 407. Kinds of amphidiscs transitional between micramphidiscs and macramphidiscs
were observed in NHM 1908.09.24.065, HCL 408 and 410 as well.
The only known difference between three well described species of Monorhaphis — M. chuni, M. dives and
M. intermedia — is the presence or absence of macramphidiscs and mesamphidiscs. Using this criterion, the
existing material was arranged into six groups (Tab. 4). The first group consisted of M. chuni according to its
original description (SCHULZE, 1904). The second group is M. chuni according to the description of IJIMA ( 1927).
Unfortunately these sponges were not found in the investigated collections and the sponge NHM 1908.09.24.064
labeled as M. chuni turned out to be a suspected M. dives. The third group consisted of four specimens of
Monorhaphis which were closer to M. chuni from its typical location, in shape and in spiculation except
mesamphidiscs that are similar to M. dives (rare in 5/2/1408, and numerous in 5/2/1352). The fourth group
included the investigated fragment of the holotype of M. dives (NHM 1908.09.24.065) and two sponges that could
be doubtfully referred to this species by shape and by the fact that they were collected in nearby stations (NHM
1908.09.24.064 and kt476); and in NHM 1908.09.24.064 some macramphidiscs were found. The fifth group was
M. intermedia according to the description of Ll JlNHE (1987). These include sponges that contain both
macramphidiscs and mesamphidiscs together with their abnormal forms. The last and sixth group consisted of
numerous specimens which arc similar to M. intermedia collected off New Caledonia and Australia. Ii is possible
to distinguish specimens without macramphidiscs (HCL 405, 410, 41 1, p540), specimens that contained neither
macramphidiscs nor mesamphidiscs (HCL 412) and specimens that contained them both (all other specimens). It is
obvious from this list that the diagnostic features distinguishing the species docs not work and that the material
showed all the possible variations. Concerning the length of microhexactin rays, the third group of M. chuni had
rays about two times longer, but this type of spicule was poorly described in the holotype and such spicules of
M. chuni described by Ijima (1927) (second group) have the shortest rays of microhexactins comparable with
those of most other specimens. Hence, the investigated and described species must be considered as synonyms of
one polymorphic species, M. chuni.
Family PHERONEMATIDAE Gray, 1 872
Genus SERICOLOPHUS Ijima, 1901
Diagnosis (slightly emended after Reiswig & ZCRAGGEN, 1991). — Body shape like a thick tongue or
spoon borne upon a compact root tuft of approximately equal length. Frontal and abfrontal surfaces respectively as
gastral and dermal, with gastral surface strongly reflexed over the dermal surface around body margins. Gastral
surface with open excurrcnt canals. Hexactins and pentactins as spicules principalia. With genuine uncinates.
Pleuralia lateralia as monactine sceptres. Basalia include anchors and distinctive monactinc "crook" spicules.
Internally basalia radiate to body margins in several strands, without a columella or associated acanthophore.
Type Species. — Hyalonema reflexion Ijima, 1894.
74
K. R. TABACHNICK & C. LEVI
Sericolophus calsubus sp. nov.
Fig. 19-21; Tab. 10, 14
MATERIAL EXAMINED. — New Caledonia. BlOCAL: stn CP 75, 22°18.65'S. 167°23.30'E, 825-860 m, 4.09.
1985: HCL 446.
Calsub: stn 8, 20°48.3'S. 167°05'E, 880-516 m, 26.02.1989; HCL 155.
TYPES. — Holotype: HCL 155 (Calsub, stn 8, 20°48.3'S, 167°05’E, 880-516 m).
Paratype: HCL 446 (Biocal, stn CP 75, 22°18.65'S, 167°23.30'E. 825-860 m).
Description. — The shape and structure of the body are typically those of the genus Sericolophus. The
holotype is a well preserved sponge 100 mm in long, 70 mm wide and 12 mm thick; a long single tult of basalia
is about 100 mm long. The paratype is a poor fragment.
Fig. 19-20. — Sericolophus calsubus sp. nov. (HCL 155). Scales = 4 cm.
Source : MNHN, Paris
0.2 mm 0.1 mm 0.05 mm
1-6, 10-11, 15-19 14 7-9, 12-13
AMPHIDISCOPHORA OFF NEW CALEDONIA
75
Fig. 21. — Spicules of Sericolophus calsubus sp. nov. (HCL 155): 1-2, dermal pinular pentaclins; 3-4. airial pinular
pentactins; 5-6, microhexactins; 7-9, micramphidiscs; 10. choanosomal pentactin; 11, outer end of choanosomal
pentactin; 12-13, micruncinates; 14, macruncinate; 15-16, crooks; 17. shaft of crook; 18. anchorate basalia;
19. shaft anchorate basalia.
76
K. R. TABACHN1CK & C. LEVI
Spicules: The choanosomal skeleton consists of pentactins (fig. 21.10-1 1) with rays 0.7-1.3/0.02-0.03 mm.
Uncinates are of two types. Macruncinates (fig. 21.14) several mm long and about 0.005 mm in diameter, they are
covered with spines. Micruncinates (fig. 21.12-13) 0.032-0.108/0.004 mm, smooth or with some sparse spines.
Basalia are anchorate spicules, crooks and diactins.
Diactins are straight, smooth spicules not less than 27 mm long and about 0.015-0.038 mm in diameter.
Crooks (fig. 21.15-17) several mm long and 0.005-0.008 mm in diameter.
Anchorate basalia (fig. 21.18-19) rare in the microscopic slides. Only several one-toothed anchorate spicules
with the shaft covered with spines were found.
Dermalia canalaria and atrialia are pinular pentactins (fig. 21.1-4).
Pinular ray slightly widened toward the outer end. It is sharply pointed. Upper end free, projecting beyond the
last spines. Spines present at a short distance from the base. Some pinular rays of both dermalia and atrialia have
short spines. Tangential rays are smooth. Pinular ray of dermal pentactin 0.099-0.361/0.007-0.019 mm. of the
atrial one 0.144-0.608/0.007-0.022 mm. Tangential rays usually covered with spines. Tangential ray of dermalia
0.034-0. 1 14/0.004-0.006 mm, of the atrial one 0.046-0.087/0.004-0.01 1 mm.
Microhexactins (fig. 21.5-6) covered with spines, they have rays 0.068-0.175/0.007-0.01 1 mm. Amphidiscs
(fig. 21.7,9) are of only one type that corresponds to micramphidiscs of other Pheronematidac. Total amphidiscs
0.018-0.056 mm long, umbel length 0.007-0.018 mm, umbel diameter 0.006-0.014 mm, shafts covered with
spines and rare widenings. The amphidiscs have usual shape, except some cases of asymmetry of umbels similar
to paradises. One macramphidisc, probably of foreign origin, was found in the paratype.
ETYMOLOGY. — The species name refers to the name of the cruise, Calsub. during which it was collected.
Sericolophus neocaledoiticus sp. nov.
Fig. 22-25; Tab. 11,14
Material EXAMINED. — Chesterfield Islands. MUSORSTOM 5: stn CP 323, 2 1 ° 1 8.52’S, 1 57° 1 8 62'E 970 m
14.10.1986: HCL 173-175.
New Caledonia BlOCAL: stn CP 34, 23°12.44'S, 167°1I.87'E, 710-700 m, 29.08.1985: HCL 157-161. —
Stn CP 54, 23°10.08'S, 167°43.54'E, 1000-950 m. 1.09.1985: HCL 162, 447-451. — Stn CP 60. 23°58.87'S,
167°07.72'E, 1530-1480 m, 2.09.1985: HCL 452. — Stn CP 75. 22°40.42'S, I67°23.41'E. 825-860 m: HCL 163-164
Loyalty Islands. Biogeocal: stn CP 297, 20°38.64'S, 167°10.77'E. 1230-1240 m. 28.04.1987- HCL 156
165-172.
Types. — Holotype : HCL 425 (Biocal, stn CP 60, 23°58,87’S, 167°07,72'E, 1530-1480 m).
Parahpes: all the other specimens mentioned above.
Description. — The form of the body is typical for the genus Sericolophus.
Most specimens are lamellar fragments of different parts of the body. The largest fragment is the one of the
holotype HCL 452 (BlOCAL, stn CP 60) now broken in fragments. Alive, it was very soft and similar to
S. leflexus Ijima, and it is 190 mm long, 90 mm in diameter and 15 mm thick. The slightly convex and smooth
surface is gastral, with very fine meshes of pinular pentactins. On each side, the reflexed parts of the gastral surface
is 25-30 mm wide. The paratype HCL 156 is 80 mm long, 30 mm in diameter and 7 mm thick.
Spicules: Choanosomal spicules usually pentactins (fig. 24.8-11) with rays about 5/0.02-0.05 mm. The outer
ends are often smooth but sometimes rough. Three types of uncinates were found. Macruncinates (fig. 24.14)
several mm long and 0.015 mm in diameter. Mesuncinates (fig. 24.13) 0.135-0.426/0.004 mm. Both macrun-
cinates and mesuncinates are covered with spines. Spindle-like micruncinates (fig. 24.12, 25.8-10) 0.041-
0.137/0.004 mm smooth with a single widening. Some small mesuncinates (HCL 174) could have features of
both types, they have spines and widenings. Sceptres (fig. 24.15) rare, they are several mm long and 0.009 mm in
diameter. Prostalia basalia crooks (fig. 24.16-18) about 0.006 mm in diameter, long diactins 0.03-0.06 mm in
diameter and anchorate spicules (fig. 24,19-20). The anchorate spicules are rare. The only complete anchor has no
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
77
tooth, it has the shape of a clavule with serrated pileate end, the rachis 0.010-0.015 mm. Dermal and atrial
spicules pinular pentactins (fig. 24.1-2) similar in shape. Pinular ray slightly widened medially; the outer end is
sharply pointed and freely projects not far from the last spines. The spines appear on a short distance from the
base. Pinular ray of dermal pentactincs 0.065-0.342/0.007-0.015 mm, of atrial ones 0.076- 1 .079/0.0 1 5-
0.019 mm. Tangential rays covered with short spines. Tangential ray of dermal pentactins 0.030-
0.137/0.006 mm, of atrial ones 0.01 1-0.175/0.007 mm.
Microhexactins (fig. 24.3-5, 25.1-7) are more numerous than other forms: pentactins, stauractins, monactins
and polyactins spicules with more than six rays. Their rays are usually covered with spines, rarely smooth with
sparse spines or entirely smooth. The microhexactins are usually represented by two types: with thick rays 0.046-
0.205/0.015-0.030 mm and with thin rays 0.046-0.213/0.004-0.007 mm. The proportion of the two types of
microhexactins varies in different specimens. The thick-rayed forms usually predominate but in HCL 175 and 160
the proportion is the opposite.
The amphidiscs (fig. 24.6-7) are represented by two types: mesamphidiscs and micramphidiscs. They both have
shafts covered with spines. Total length of mesamphidiscs 0.032-0.122 mm, umbel length 0.01 1-0.044 mm.
umbe! diameter 0.010-0.035 mm. Total length of micramphidiscs 0.015-0.050 mm, umbel length 0.005-
U.UZO mm, umbel diameter 0.004-0.018 mm.
etymology. — This specific name refers to the recorded locality.
0.2 mm 0.1 mm 0.05 mm
1-5, 8-11, 14-20 13 6-7, 12
78
K. R. TABACHNICK & C. LEVI
Fig. 24. — Spicules of Sericolophus neocaledonicus sp. nov. (HCL 156): 1. dermal pinular pentactin; 2. atrial pinular
pentactin; 3-5, microhcxactins; 6, micramphidisc; 7. mesamphidisc; 8. choanosomal pentactin; 9-11. outer ends of
choanosomal pentactin; 12, micruncinate; 13, mesuncinate; 14, macruncinate; 15, sceptre; 16-17, crooks;
18, shaft of crook; 19, anchorate basalia; 20, shaft of anchorate basalia.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
79
_ _ 0-2 mm _ 0.05 mm
1-7 8U0
Fig. 25. Spicules of Sericolophus neocaledonicus sp. nov,: 1-7, deformed microhexactins; 8-10. micruncinates
1 1 (HCL 175), 2(HCL 167). 3-4 (HCL 162), 5 (HCL 162), 6 (HCL 173), 7-9 (HCL 167), 10 (HCL 174)].
Sericolophus hawaiicus sp. nov.
Fig. 26; Tab. 12, 14
Mater IAL EXAMINED. — Hawaii. HURL-88: sin P5-057, 19°37'N. 156°02'W, 580 m: co47l (BPBM).
HURL-92, stn 5201, 20°53'N, 157°38'W, 380 m: co475 (BPBM).
Types. — Holotype: co475 (HURL-92, sta. 5201, 20°53’N, 157°38'W, 380 m).
Paratype: co471 (HURL-88: stn P5-057, 19°37'N, 156°02'W, 580 m).
Fig. 26. — Spicules of Sericolophus hawaiicus sp. nov.: 1, dermal pinular pentactin (co475); 2. atrial pinular pentactin
(co475); 3. pinular ray of atrial pinular pentactin (co475); 4, microhexactin (co475); 5, micropentactin (co475);
6, micromonactin (co475); 7-9, amphidiscs (co475); 10, choanosomal pentactin (co475); 11-13, outer ends of
choanosomal pentactin (co475); 14.16, micruncinates (co475); 15, macruncinate (co475); 17, crook (co471);
18-20, shafts of basal anchorate spicule (co471).
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
DESCRIPTION. — The holotype is a fragment 12 mm in diameter of the lower part of the sponge with a tuft of
basalia about 200 mm long and 17 mm in diameter. The paratype is a small fragment.
Spicules: The choanosomal spicules are pentactins (fig. 26.10-13) with rays, 0.24-1.37/0.008-0.016 mm.
Uncinates are represented by two types. Macruncinates (fig. 26.15) are several mm in length and about 0.008-
0.016 mm in diameter. The micruncinates (fig. 26.14-16) (according to the spicule size in other species) or
mesuncinates (according to their shape) are about 0.1-0.5/0.004 mm. All uncinates are covered with spines.
Basalia consists of crooks (fig. 26.17) and anchorate spicules. A single, one-toothed, anchorate spicule
(fig. 26.18-20) was found. It is not clear which side of the fragment is dermal and which is atrial, but spicules
(fig. 26.1-2) of both sides are similar in shape and size. The pinular rays (0.068-0.182/0.004-0.01 I mm) are
sharply pointed. The upper end of the pinular ray is usually long but sometimes relatively short. The outer ends of
pinular rays are always sharply pointed and project beyond the ends of last spines. The tangential rays (0.038-
0.072/0.004 mm) are covered with spines or smooth.
Microhexactins (fig. 26.4-6) covered with spines predominate over micropentactins and micromonactins. Their
rays arc 0.063-0. 1 1 7/0.007 mm.
The only type of amphidiscs (fig. 26.7-9) corresponds to micramphidiscs of other species. The shafts of
micramphidiscs are covered with spines. Total length of micramphidisc 0.020-0.065 mm, umbel length 0.005-
0.021 mm, umbel diameter 0.005-0.019 mm.
Etymology. — This specific name refers to the recorded locality.
Sericolophus cidaricus sp. nov.
Fig. 27; Tab. 13-14
Material EXAMINED. — Australia. Northeastern coast. ClDARIS 1: stn 4.1, 1 8°1 1 52'S, 147°52. 1 2'E, 1012-
998 m, 6.05.1986: HCL 180 (MTQ).
Types. — Holotype: HCL 180 (ClDARIS I, stn 4.1. 18° 11. 52'S, I47°52.12'E, 1012-998 m).
Description. — The holotype is a fragment of a nearly complete specimen, 200 x 1 10 x 20 mm, with a tuft
of basalia. The sponge is damaged but the body shape, typical for the genus Sericolophus. is recognizable.
Spicules: The choanosomal spicules arc usually pentactins (fig. 27.8-10), rarely their derivatives such as
tauactins. Their rays are 0.008-0.023 mm in diameter. The uncinates are represented by two types. The
macruncinates (fig. 27.12) are long, 0.003-0.016 mm in diameter. They are covered with spines. The
micruncinates (fig. 27.13-14), 0.061-0.120/0.011-0.015 mm, are smooth (smallest ones) or partly serrated (larger
ones), they both have widening. The spicules of basalia are represented by crooks (fig. 27.1 1) (0.004 mm in
diameter) only. The dermal and atrial spicules are pinular pentactins (fig. 27.1-3) similar in shape. The pinular ray
is constant in thickness, sharply pointed and projects freely beyond the last spines. The spines begin al a short
distance from the base. The pinular ray of dermal pentactins is 0.2 1 7-0.8 1 3/0.004-0.022 mm, of atrial ones 0. 1 86-
1.904/0.015-0.019 mm. The tangential rays of dermal pentactins are pointed and covered with short spines. The
tangential rays of atrialia are covered with spines longer and denser than those of the dermalia; they are pointed or
rounded. The tangential ray of dermal pentactins is 0.038-0.129/0.004-0.013 mm. of atrial ones 0.057-
0.148/0.013 mm.
Microhexactins (fig. 27.4) predominate over other types of analogues spicules (fig. 27.5-7): micropentactins,
microstauractins, micromonactins and polyactins. All these spicules are covered with spines. Their rays are 0.076-
0.107/0.007-0.009 mm. Two types of amphidiscs can be observed. Mesamphidiscs (fig. 27.16) have shafts with
spines or tubercles. Total length of mesamphidisc 0.058-0.104 mm, umbel length 0.013-0.032 mm, umbel
diameter 0.018-0.030 mm. Micramphidiscs have shafts with spines or widenings. Total length of micramphidiscs
0.016-0.040 mm, umbel length 0.005-0.013 mm, umbel diameter 0.005-0.014 mm. Some macramphidiscs
(fig. -7.I5) 0.312/0.099/0.122 mm with tuberculatcd shafts are not certain to belong to this species.
0-2 mm _ _ 0.1 mm _ 0.05 mm
1-11 12 13-16
82
K. R. TABACHN1CK & C. LEVI
Fig. 27. — Spicules of Sericolophus cidaricus sp. nov. (HCL 180): 1-2. dermal pinular pentactin; 3. airial pinular
pentactin; 4, microhexactin; 5, micropentactin; 6. microslauraclin; 7, micromonactin; 8, choanosomal
pentactin; 9. choanosomal tauactin; 10, outer end of choanosomal spicule; 11. crook; 12. macruncinate;
13-14, micruncinates; 15, macramphidisc; 16. mesamphidisc.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
83
Remarks. — All specimens of Sericolophus described earlier (for references see Reiswig & Zgraggen.
1991) were collected off Japan. All belong to S. reflexus. The best descriptions of the Japanese materials were
made by Ijima and Okada (1938). Reiswig & ZGRAGGEN (1991) completed the previous descriptions and
clarified the problem of synonymies. The specimens from Hawaii, New Caledonia and Australia obviously belong
to the same genus. Their shape is similar to that of S. reflexus and they have crooks that are never found in
representatives of other genera. Species of Sericolophus are distinguished by a combination of size classes of
amphidiscs, shapes and size variations of microhexactins, dermal and atrial pinular pentactins and uncinates
(Tab. 14). These features are distributed mosaically among these species. S. neocaledonicus and S. cidaricus are
similar in size of amphidiscs and variations of microhexactins (up to monactins). They differ in the proportions of
dermal and atrial pinular pentactins and the presence of thick-rayed microhexactins in .S', neocaledonicus.
S. calsubus has microamphidiscs smaller than those of S. neocaledonicus and S. cidaricus. Dermal and atrial
pinular pentactins of 5. calsubus differ in size. The maximal diameter of the microhexactins of S. calsubus is
intermediate between those of the two other species. S. hawaiicus differs from the others by more significant
features: small-sized dermal and atrial pentactins, and short and thin-rayed microhexactins and micruncinates
bearing spines similar to macruncinates. The anchorate spicules of basalia of 5. reflexus are best known. The
anchors in S. reflexus have one, sometimes two, teeth with serrations on the anchorate margin. Numerous
accessory spines, even similar to teeth, seem to be secondary elements as a result of "developed serration"
(REISWIG & ZGRAGGEN, 1991). A unique but predictable form was found in S. neocaledonicus with pileate head
and serrated margin, devoid of teeth. It may be considered as a derivate of the "normal" toothed anchors. This
spicule is analogous in shape to some clavules of Farrea (Ijima, 1927). These features of basalia have low
taxonomic value and after careful investigation of other materials, the difference among basalia may disappear.
Etymology. — The species is named after the Cidaris cruise along the N.E. Australian slope.
Genus SEMPERELLA Gray, 1868
Diagnosis (after Ijima, 1927). — Pheroncmatidae of a columnar or a club-like body shape, without gastral
concavity, consisting internally of alternating incurrenl and excurrent systems of anastomosing canals, which stand
in relation respectively with several dermal and oscular areas distinguishable in all parts of the external surface.
Dermal areas covered with uniformly fine-meshed lattice; oscular areas cither covered with a larger-meshed lattice
or opening through relatively large orifices, found isolated or in close groups. Basalia arising in separate tufts, join
in a matted mass.
Type Species. — Hyalonema schultzei Semper, 1868.
Semperella schultzei (Semper, 1868)
Fig. 28
Hyalonema sclmhzei. Semper. 1868: 272,
wTTe//a schulJzei - J E- Gray' 1868: 373. — W. Marshall, 1875: 212, fig. 67-83. pl. 12 fig. E. pi. 16-17. —
Ri ^Y^R’ l877: 276' fig- l8-'9- P'- 14, pl. 15. — Schulze, 1886: 67; 1887: 261. pl, 51-52. —
i c!,CjKB|URN' : 57’ p ’ L ~ 0kada, 1932: 16. — Ijima & Okada. 1938: 441. pi. 15 fig. 1-3, pl. 16 tie. 22-32. —
Lfivi & Levi. 1982: 289. fig. 3-4. pl. 4 fig. 2, pl. 5; 1989: 29, fig. 1. P g . P g
20 m im"* '^rNED' ~ Chesterfield Islands. Musorstom 5: stn DC 378. 19°53.74'S, 158°38.30'E, 355 m,
Me,Wc^aled0nilV Dragagcs " Vauban stn 39, 22°29'S, 166°23 E, 375-550 m, 5-7.06.1979: HCL 201.
18°58 0n°R Son'75, l8°59-30'S- 1 63° 1 7.50'E, 370 m. 17.09.1985: HCL 183-185. — Stn CC 202.
3 ,0 ,985S 580 m' 20 091985: HCL 182- — Sln CC 241 ■ 22°09.00'S, 167°12.20'E. 470-480 m.
84
K. R. TABACHNICK & C. LEVI
Biocal: stn CP 42, 22°46.09'S, 167°13.80'E, 380 m, 30.08.1985: HCL 181. — Stn DW 44, 22°47.35 S
167°14.50'E, 440-450 m, 30.08.1985: HCL 188.
Fig 28. — Spicules of Semperella schultzei (Semper. 1868): 1-4, curious amphidiscs (1-3, HCL 198; 4. HCL 182);
5-7, micromonactins (5-6, HCL 200; 7, HCL 182); 8, sceptre (HCL 186); 9. choanosomal stauractm;
10-12, choanosomal pentactins; 13-14, outer ends of choanosomal spicules; 15, choanosomal hypodermal or
hypoatrial pentactin.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
85
Loyalty Islands. MUSORSTOM 6: stn DW 391, 20°47.35'S, 167°05.70'E. 390 m, 13.02.1989: HCL 197. —
StnDVV 393, 20°48.29,S, 167°09.54'E, 420 m. 13.02.1989: HCL 198. — Stn DW 410, 20°38.05'S. I67°06.65'E,
490 m, 15.02.1989: HCL 190. — Stn DW 412, 20°40.60'S, 167°03.75'E. 15.02.1989: HCL 195. — Stn CP 415,
20°40.20'S, I67°03.95'E, 461 m, 15.02.1989: HCL 196. — Stn DW 428, 20°23.54,S, 166°12.57'E, 420 m, 17.02.
1989: HCL 189. — Stn DW 448. 20°55.66'S. 167°22.34'E, 410 m, 19.02.1989: HCL 191-194.
Wallis and Futuna Islands. MUSORSTOM 7: stn DW 510, 14°14.50'S, 178°1I.50'W, 280-370 m, 12.05.1992:
HCL 199. — Stn DW 605, 13°21.3'S, 176°08.3'W, 335-340 m. 26.05.1992: HCL 200.
DISTRIBUTION. — S. schultzei is widely distributed from New Caledonia and the southwestern Pacific to
Japan. Nearly all the specimens show similar morphological features except some from the Australian region
which may be considered as a new subspecies (unpublished data).
Fig. 29. — Semperella varioactina sp. nov. (HCL 202). Scale = 1 cm.
Fig. 30. — Semperella abyssalis sp. nov. (HCL 203). Scale = 3 cm.
Semperella varioactina sp. nov.
Fig. 29, 31; Tab. 15
Material
HCL 202.
EXAMINED. — New Caledonia. Biocal: stn CP 61. 24°10.67'S, 167°33.65'E, 1070 m. 2.09.1985:
86
K. R. TABACHN1CK & C. LEVI
Types. — Holotype: HCL 202 (BlOCAL: sin CP 61, 24°10.67'S, 167°33.65'E, 1070 m).
DESCRIPTION. — The general shape is elongated, columnar, sliglhly broader in ihe middle. The holotype is
140 mm long, 8-12 mm in diameter.
The body presents a rectangular cross-section and the general surface of the sponge is divided into four longitu¬
dinal areas without sieve plates. In all specimens or fragments, oscular and dermal areas cannot be recognized.
Openings of transverse canals (1-1.5 mm in diameter), are scattered.
Spicules: The choanosomal skeleton consists of pentactins (fig. 31.4-9) and rare stauractins with rays
0.3-4.6/0.008-0.180 mm. The rays are smooth, rarely with rough outer ends. Uncinates (lig. 31.10) are rare, and
only fragments about 0.004 mm in diameter were found. Basalia and other prostalia were not found. Dermalia and
atrialia are pinular pentactins (fig. 31.1-3). The atrial pinular pentactins were not found with certainty. The pinular
ray 0.099-0.198/0.007 mm is sharply pointed, the upper end projects freely beyond the ends of last spines. The
spines are relatively short, being bent upward; they begin at a short distance from the base. The tangential rays
(0.061-0.137/0.007 mm) are curved and acutely intersect in pairs, as in other Semperella species. Tangential rays
mainly smooth. Outer ends or even more than half of a ray covered with short spines directed upward.
Microhexactins (fig. 31.11-22) and their derivatives, including monactins and spheres (pearls), are rough or
spinous. Some spicules have enclosing unequal angles. Stauractins, monactins and pentactins are numerous. Ray
length of these spicules is 0.061-0.137/0.007 mm. The monactins have one spherical, the other finely pointed
outer ends. Only one type of amphidisc (fig. 31.23-25) is present, but their length varies greatly. The shafts of
these amphidiscs are covered with short spines. Total length of amphidiscs 0.032-0.108 mm, umbel length 0.009-
0.038 mm, umbel diameter 0.007-0.032 mm.
ETYMOLOGY. — The species is named after the diversity of microholactine spicules.
Semperella abyssalis sp. nov.
Fig. 30. 32; Tab. 16
MATERIAL examined. — Loyalty Islands. Biogeocal: stn CP 306, 20°35.I8’S, 167°13.99'E, 2960-3036 m.
1.05.1987: HCL 203-208.
Types. — Holotype: HCL 203 (Biogeocal, stn CP 306, 20°35.18’S. 167°13.99'E. 2960-3036 m).
Paratypes: HCL 205-208.
Description. — The sponge is subcylindrical. The holotype is over 210 mm long and 35 x 25 mm
in maximal section. The paratypes are over 100-170 mm long, and are Hatter than the holotype (15 x 10 -
25 x 5 mm). They were probably squeezed. No basalia were found in these specimens. The atrial surface is band¬
like, and situated along the dermal surface. In the holotype two such bands merge in the upper part oi the sponge.
Spicules: The choanosomal skeleton consists of pentactins (fig. 32.7-1 1) and rare stauractins with smooth rays
0.2-13/0.02-0.09 mm, equal or unequal in length. Uncinates (fig. 32.12) 0.002-0.004 mm in diameter are rarely
found. Basalia and prostalia marginalia are absent. Dermal and atrial spicules are very similar; most are pinular
pentactins (fig. 32.1-6) or hexactins. The dermal pentactins of the holotype have pinular ray 0.129-0.296/0.007-
0.030 mm (avg. 0.183 mm, std. 0.047 long), tangential ray 0.061-0.167/0.005-0.007 mm (avg. 0.092 mm,
std. 0.026 long). The atrial pentactins of the holotype have similar parameters; the pinular ray is 0.122-
0.303/0.007-0.030 mm (avg. 0.177 mm, std. 0.036 long), the tangential ray 0.053-0.21 1/0.005-0.007 mm (avg.
0.103 mm, std. 0.035 long). Two types of spicules with different pinular outer ends are present among dermal and
atrial pentactins. The first has the pinular ray widening toward the outer end with conical tip which do not project
freely beyond the ends of the last spines. The long spines bent upward and the longest ones are often situated at the
upper part of the ray. The second has pinular ray widening slightly toward the end and covered with short spines.
The conical upper end freely projects beyond the ends of last spines. The spines appear close to the base ot the
shaft. The tangential rays are curved and smooth, with outer ends covered with short spines directed upward.
Source : MNHN, Pans
AMPHIDISCOPHORA OFF NEW CALEDONIA
87
Fig
’ Semperella varioactina sp. nov. (HCL 202): 1, dermal pinular pentactin; 2-3. tangential rays of dermal
pinu at pentactin; 4, choanosomal pentactin; 5. choanosomal stauractin; 6-9, outer ends of choanosomal spicules;
, uncinate; 11, micropcntactin; 12, 16 microtauactins; 13, microstauractin; 14. microtauactin with rudimental
ray; 15, micropentactin with rudimental ray; 17-19. micromonactin; 20. microsphere; 21, microhexactin;
22, nucrodiactin; 23-24, macramphidiscs; 25, micramphidisc.
88
K. R. TABACHNICK & C. LEVI
Fig. 32. — Semperella abyssalis sp. nov,: 1, 6. atrial pinular pentactins (HCL 203); 2-3. pinular rays of atrial
pentactins (HCL 203); 4-5, tangential rays of atrial pentactins (HCL 203); 7-8, choanosomal pentactins (HCL 203);
9. choanosomal stauractin (HCL 203); 10-11, outer ends of choanosomal spicules (HCL 203); 12, uncinate
(HCL 203); 13, micropentactin (HCL 203); 14, microstauractin (HCL 203); 15, microhexactin (HCL 206);
16, microtauactin with rudimental ray (HCL 205); 17, micromonactin (HCL 203); 18, micromonactin (HCL 205);
19, micromonactin (HCL 204); 20. macramphidisc (HCL 203); 21, micramphidisc (HCL 203).
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
89
Micropentactins (fig. 32.13) have rays (0.046-0.167/0.009-0.011 mm) entirely covered with short spines.
Micropentactins coexist with rare other spicules (fig. 32.14-19): microhexactins, microstauractins. microtauactins
and micromonactins. A micromonactin has spherical, finely pointed outer end. The micromonactins are partly or
entirely covered with spines as well as other spicules mentioned above. The size of amphidiscs (fig. 32.20-21)
varies widely, but probably belongs to one type. The largest are about 5 times longer than the smallest. The shafts
of these amphidiscs are covered with short spines. In the smallest amphidiscs the shaft may be slightly curved and
the umbels slightly asymmetrical. Total length 0.016-0.1 19 mm, umbel length 0.004-0.036 mm, umbel diameter
0.005-0.025 mm.
Etymology. — This species name refers to its deep-sea habitat.
Semperella crosnieri sp. nov.
Fig. 33-37; Tab. 17-18
MATERIAL EXAMINED. — New Caledonia. BlOCAL: no other data: HCL 210.
Loyalty Islands. BiOGEOCAL: stn CP 297, 20°38.64'S, 167°10.77'E, 1230-1240 m. 28.04.1987: HCL 212.
Calsub: sin 6, W of Lifou Island and N of Santal Bay, 20°48'S, 167°02.4'E, 1 150-400 m. 24.02.1989: HCL 211.
Combes Bank ( near Wallis and Futuna Islands). MUSORSTOM 7: stn CP 550, 12°14.8'S. I77°28.0'W. 800-810 m.
18.05.1992: HCL 209.
Types. — Holotype: HCL 209 (Musorstom 7, stn CP 550, 12°14.8'S, 177°28.0'W, 800-810 m).
Paratypes : all the others specimens mentioned above.
Description. — The sponge is cylindrical, ovoid or flattened in section. The holotype is a tongue-like sponge
over 240 mm long and about 160 x 40 mm in section. It has three short rows of prostalia in the upper part. The
paratypes are cylindrical, over 280 and 330 mm in length and about 100-180 mm in diameter. The paratype HCL
211 is a complete specimen 330 mm long and 70 mm in diameter, with small rows of pleuralia on the upper end
(some scattered pleuralia are located on the lateral side). Surfaces occupied by the atrialia are irregularly located.
They probably correspond to the areas of the surface covered with large-meshed latticework, whereas dermalia -
areas ol the surface are covered with small-meshed latticework. The small meshed area (dermal area) is located on
both sides, on one side together with occasionally situated areas of large-meshed surfaces (atrial area).
Spicules: The choanosomal skeleton consists of pentactins (fig. 37.1 1-15) and rare stauractins with smooth
rays 0.1-4.6/0.008-0.26 mm. Their rays are often bent in an X-form. The uncinates (fig. 37.17-18) include two
types. Macruncinates arc long and about 0.002-0.01 mm in diameter. Micruncinates are shorter and thinner and
0.002 mm in diameter. The macruncinates were rarely found in preparations of the holotype's spicules, the
micruncinates are rare in the paratype HCL 211. Basalia (fig. 37.19) are present in the paratype HCL 21 1 only.
They are anchor-like two-toothed spicules covered with spines. Prostalia are sceptres (fig. 37. 16) as usual for other
Pheronematidae. Dermal and atrial spicules are pinular pentactins (fig. 37.1-4), similar in shape. The pinular ray is
sharply pointed and bears short spines appearing close to the base. The pinular ray of dermal pentactins is 0.072-
0.300/0.004-0.006 mm, the pinular ray of atrial pentactin is 0.076-0.342/0.004-0.006 mm. The tangential rays
are curved and smooth, with outer ends covered with short spines. The tangential ray of dermal pentactin is 0.038-
0.103/0.004-0.006 mm, the tangential ray of the atrial pentactin is 0.038-6.133/0.004-0.006 mm.
Micropentactins (fig. 37.5-10) and rare microhexactins have rays 0.031-0.171/0.01 1-0.022 mm usually thick
and entirely covered with numerous or rarely sparse spines. These spines are long, and are directed toward the base
ol the ray. The amphidiscs (fig. 37.20) are nearly uniform in shape and size. Total length 0.014-0.029 mm, the
u"1 'Cnclh 0.005-0.009 mm, the umbel diameter 0.005-0.01 1 mm. The shafts of these amphidiscs arc covered
with short spines.
Remarks. It is often difficult to decide if a specimen belongs to Semperella or to Poliopogon. Typical
specimens ol Semperella have columnar body shape whereas Poliopogon are plate-like, with dermal and atrial
sui aces on the opposite sides. The atrial surface of Semperella is not continuous, and it shows numerous areas
90
K. R. TABACHNICK & C. LEVI
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
91
E
E
in
o
o
Si
Fig. 37. Semperella crosnieri sp. nov.: 1-2, dermal pinular pentactins (HCL 209); 3-4, tangential rays of dermal
pentactins (HCL 209); 5-7,9-10. micropentactins (5-6. 9-10, HCL 209; 7, HCL 212); 8. microhexactin (HCL 212);
11-13, choanosomal pentactins (HCL 209); 14, choanosomal stauractin (HCL 209); 15. outer end of choanosomal
spicule (HCL 209); 16. sceptre (HCL 211); 17, micruncinate (HCL 210); 18, macruncinate (HCL 210); 19. anchorate
basalia (HCL 211); 20, amphidisc (HCL 209).
92
K. R. TABACHNICK & C. LEVI
isolated as single or in groups on all sides of the conical body (Ijima, 1927). If the specimen is damaged it may be
difficult to find dermal and atrial surfaces and hence difficult to relate the specimen to one of the two genera. The
atrial surfaces vary notably. 5. schultzei has atrial surfaces shaped as several thin bands (Ijima & Okada, 1938).
Atrial surfaces of S. stomata (Ijima & Okada, 1938) and probably of S. spicifera (Schulze, 1904) underline
numerous separate oscules. Atrial surfaces of 5. cucumis appear as numerous rounded spots of excavated atrial
cavities (SCHULZE, 1904). That of S. alba is represented by two opposite sides of squared section body
(Tabachnick, 1988). The external body shape of the latter is similar to the lower square part of P. micro-
pentactinus (described below). The new species of Semperella have the following external body shape: S. abyssalis
is similar to S. schulzei; S. crosnieri similar to S. cucumis but the spots of atrial surfaces of S. crosnieri are
larger; S. vdrioactina is probably close to S. spicifera. The shapes and combinations of spicules are more
important for the identification of the new species of Semperella.
S. varioactina is distinguished by the following features: micropentaclins, microstauractins and micro-
monactins prevail upon the analogous spicules; two types of amphidiscs are present; maeramphidiscs may have
spherical umbels (due to curved teeth). S. crosnieri has pinular pentactins equal to those of S. varioactina and
similar in shape to those of S. schultzei. A single type of amphidisc is known: micramphidisc. Micropentactins
have thick rays with numerous long spines, as in S. stomata but sometimes the spines are sparse. S. abyssalis
has pinular pentactins with widened upper part, two types of amphidiscs, and the maeramphidiscs with an
extremely elongated shaft.
Etymology. — This species name refers to Alain Crosnier, zoologist and oceanographer.
Genus POLIOPOGON Thomson, 1873
Diagnosis (after Schulze, 1886). — The body has the form of either a thick-walled goblet or an ear-shaped
involute plate. It has a broad basal tuft and an oscular fringe of marginalia, but no laterally projecting pleuralia.
The parenchyma contains small and extremely rough or spinose oxyhexacts and uncinates. In one species, even
small smooth oxydiacts vary in size and abundance. The two teeth of the basal anchors are arranged approximately
at the right angles of the long, almost smooth shaft. The marginalia end externally in club-shaped thickenings.
Type Species. — Poliopogon amadou Schulze, 1887.
Poliopogon micropentactinus sp. nov.
Fig. 38-44; Tab. 19-20
Material EXAMINED. —New Caledonia. Biocal: stn CP 60, 23°58.87'S, 167°07.72'E, 1530-1480 m, 2.09.
1985: HCL 214. — Stn CP 62, 24°19.35'S, 167°49.43'E, 1410 m, 2.09.1985: HCL 215. — Stn CP 63. 24°26.97'S.
168°08.17'E, 2160 in, 2.09.1985: HCL 454-457.
Loyalty Islands. BtOGEOCAL: stn CP 272, 21°00.04'S, 166°56.94'E, 1615-1710 m, 20.04.1987: HCL 213.
TYPES. — Holotype: HCL 213 (BlOGEOCAL, stn CP 272, 21°00.04'S, 166°56.94’E, 1615-1710 m).
Paratypes: all the other specimens mentioned above.
Description. — The holotype is a tongue-like sponge 480 mm in long and 80x30 mm in section. Basalia are
about 60 mm long. The paratype HCL 214 is over 270 mm long, 60 x 20 mm in section. The paratype HCL 215
is over 600 mm long. In section it shows a square with one opposite convex pair and the other concave as some
Semperella. Other paratypes are represented as fragments that are rather shaped like Poliopogon.
Spicules: The choanosomal skeleton consists of pentactins (fig. 44.4-8), with rays from 0.2 to over several
mm in length and 0.02-0.76 mm in diameter. The length of these rays differs notably. Uncinates (fig. 44.9) are
Source : MNHN, Paris
AMPHID1SCOPHORA OFF NEW CALEDONIA
93
Fig. 38-39. — Poliopogon micropentactinus sp. nov. (HCL 213). Scale = 10 cm.
Source . MNHN, Paris
94
K. R. TABACHN1CK & C. LEVI
Fig. 40-43. — Poliopogon micropentaciinus sp. nov.: 40-41. (HCL 215); 42-43, (HCL 214). Scales = 10 cm.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA 95
JJ’Ulj2 iym diameter. Basalia dig. 44.10-1 1) were found in the holotype only; they are anchor-like, two
6 SPICU es' 1 c shaft 0.01-0.04 mm in diameter covered with spines. Which surface between two opposite
pans is atrial and which is dermal, as it must be in Poliopogon , has not be determined so that spicules are
uiscussea as spicules from different sides. The careful examination of the pinular pentactins in the specimen
96
K. R. TABACHNICK & C. LEVI
HCL215 has shown the great similarity of statistical parameters of all four surfaces (two convex and two
concave), whereas the spicules of the ribs between two neighbours surfaces are notably larger. If they corresponded
to the atrial surface, this sponge should be transferred to the genus Semperella. The increased size of the pinular
pentactins from the ribs in HCL 215 can be a result of their marginal position. Pinular pentactins and
rare hexactins from both surfaces in other species of the genus are similar in size and shape. The pinular ray
(fig. 44.1-3) widens toward the end, ending with a relatively thick end with several short spines. The spines are
directed upward and they occur near the base. The tangential rays could make a regular crest or two, the opposite
rays are bent in X-form. The tangential rays are smooth with outer ends or even half of a ray covered with short
spines directed upward. The pinular ray of pentactins from one side is 0.122-0.334/0.01 1-0.026 mm, from the
other. 0.114-0.262/0.0011-0.026 mm. The tangential rays of pentactins from one side is 0.034-0.148/0.007-
0.01 1 mm, from the other, 0.034-0.122/0.007-0.01 1 mm.
Micropentactins and rare microstauractins (fig. 44.12-14) are covered with short spines. They have thick or thin
rays. The rays (0.052-0.198/0.006-0.017 mm) are covered with spines.
All the amphidiscs (fig. 44.15-17) belong to one type. The largest are about two times larger than the
smallest. The amphidisc shafts are covered with short spines. Total length of amphidiscs is 0.015-0.1 15 mm,
umbel length 0.005-0.041 mm, umbel diameter 0.005-0.026 mm.
Remarks. — This new species of Poliopogon differs notably from other species of the genus [ P . amadou
(Schulze, 1887). P. amadou pacifica (Tabachnick, 1988), P. maitai (Tabachnick, 1988)]. P. micropentactinus has
micropentactins instead of microhexactins and micropentactins, which are rarely found in other species. It has one
type of amphidisc, dermal and atrial pentactins may have the tangential rays curved.
Etymology. — The species name refers to the prevalence of micropentactins.
Poliopogon claviculus sp. nov.
Fig. 45, 48; Tab. 21
Material EXAMINED. — New Caledonia. Halipro 2: stn BT 25. 25°17.45'S. 170°23.93'E, 1100-1348 m,
11.1 1.1996: HCL 458.
TYPES. — Holotype: HCL 458 (HALIPRO 2, stn BT 25, 25°17.45’S, 170°23.93'E, 1 100-1348 m).
Description. — The holotype is a conical sponge 300 mm long and about 120 x 100 mm in section.
Atrialia is a flattened concave area about 200 x 50 mm. Basalia are very short, up to 10 mm long. The sponge is
fixed by its basal part to a dead cylindrical fragment of hexactinellid sponge with fused skeleton, probably
Tretopleura.
Spicules : The choanosomal skeleton consists of pentactins (fig. 48.18) and rare stauractins with rays
0.072-0.972/0.014-0.022 mm. Uncinates (fig. 48.1 1-12) are 0.191-0.828/0.002-0.007 mm. Basalia (fig. 48.7) are
anchor-like, two-toothed spicules and monaxons (fig. 48.6), with one outer rounded end. The shall of anchorate
basalia is about 0.006 mm in diameter, probably smooth. Basal monaxons with rounded end are thick, about
0.050 mm in diameter. Other spicules of prostalia are very specific to this species. They are monaxons with
clavate distal ends (fig. 48.8-10). which can have spines only on the top and beyond the clavate part. The prostal
parts of these spicules are common with other representatives of Pheronematidae. Clavate monaxons are numerous
in dermal area and particularly numerous among basalia, they are 0.220-0.396 mm long, 0.004-0.005 mm in
diameter of main shaft and 0.014-0.022 mm in diameter of their clavate parts. Dermal and atrial pentactins are
pinular pentactins (fig. 48.1-5) similar to each other in shape. Dermal pentactins can be about two times larger
than the atrial ones. The pinular ray of these spicules often has a thin spine or its outer end which protrudes far
beyond the last spines. The pinular ray of dermal pentactins is 0.078-0.392/0.01 1-0.025 mm, tangential ones are
0.044-0.096/0.007-0.016 mm. The pinular ray of atrial pentactins is 0.078-0.174/0.022-0.029 mm, tangential
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
97
Fig. 45. — Poliopogon claviculus sp. nov. (HCL 458). Scale = 10 cm.
Fig. 46-47. — Poliopogon zonecus sp. nov. (HCL 459). Scale = 10 cm.
ones are 0.030-0.078/0.01 1-0.014 mm. Micropentactins (fig. 48.13) are covered with short spines. They have rays
0.061-0.122/0.004 mm.
Amphidiscs are of three types. Macramphidiscs (fig. 48.14-15) are oval, almost round, with smooth shafts and
teeth that nearly meet at the equator. The umbel teeth are often broad but sometimes thin and long with some
short spines on their surface. Total length of macramphidiscs 0.131-0.287 mm, the umbel diameter 0.070-
0.183 mm. Mesamphidiscs and micramphidiscs (fig. 48.16-17) are similar in shape. They have spiny shafts and
are different in sizes. Total length of mesamphidiscs 0.061-0.200 mm. the umbel length 0.017-0.078 mm, the
umbel diameter 0.010-0.052 mm. Total length of micramphidiscs 0.032-0.058 mm, umbel length 0.012-
0.018 mm, umbel diameter 0.008-0.016 mm.
Remarks. Poliopogon claviculus can be easily distinguished from the other species. It has oval
macramphidiscs and prostalia lateralia, and monactins with clavate outer ends. A similar sponge with similar
c aracteristic spicules was found off Hawaii Islands. It is very likely that it belongs to Poliopogon claviculus
(REIS WIG, personal communication).
Etymology. The specific name claviculus refers to the shape of the prostalia spicules.
98
K. R. TABACHNICK & C LEVI
Fig. 48. — Poliopogon claviculus sp. nov. (HCL 458): 1-3, dermal pinular peniactins; 4-5. atrial pinular pentactins;
6, basal monaxon; 7, anchorate basalia; 8-10, prostalia lateralia (clavate); 11-12, uncinates; 13. micropentactin;
14, common macramphidisc; 15, thin-toothed macramphidisc; 16, mesamphidisc; 17, micramphidisc; 18, choano
somal pentactin.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA 99
0.2 mm
1-5, 11-16
0.2 mm
6-10, 18
0.05 mm
17
loHopogon zonecus sp. nov. (HCL 479): 1-2, dermal pinular pcntactins; 3-4. atrial pinular pentactins;
3-0. anchorate basalia; 7. choanosomal pentactin; 8, prostalia lateralia (cuspidate); 9. uncinate; 10. microhexactin;
. imcrotnactin; 12-13, micromonactins; 14, microorthodiactin; 15. microdiactin; 16-17. microspheres;
in. micramphidisc.
100
K. R. TABACHNICK & C. LEVI
Poliopogon zonecus sp. nov.
Fig. 46-47, 49; Tab. 22
Material EXAMINED. — New Caledonia. Halipro 2: stn BT 25. 25°I7.45'S, 170°23.93'E. 1100-1348 m,
11.11.1996: HCL 459. — Sin BT 32. 25°19.95'S. 168°54.75'E. 697-1340 m, 12.11.1996: HCL 460.
TYpES. _ Holotype : HCL 459 (Halipro 2, stn BT 25, 25°17.45'S. I70°23.93’E, 1 100-1348 m).
DESCRIPTION. — The holotype is a tongue-like, elongated, 170 mm long sponge. It is about 75 x 40 mm in
section. Tuft of basalia protrudes at about 20 mm. The specimen HCL 460 is a poor fragment.
Spicules: The choanosomal skeleton consists of pentactins (fig. 49.7) rarely of stauractins and tauactins, with
rays 0.216-1.080/0.011-0.025 mm. Uncinates (fig. 49.9) (0.367-0.576/0.004-0.005 mm) are covered with very
short spines, their surface seems to be rather rough than spinous. Basalia (fig. 49.5-6) are anchor-like, two-toothed
spicules with shaft about 0.007 mm in diameter. Prostalia lateralia (fig. 49.8) are monaxons with rounded (not
clavate) outer ends. Dermal and atrial pentactins (fig. 49.1-4) are pinular pentactins similar to each other in shape
and size. The outer end of pinular ray of these spicules is conical, equal to or not much longer than the last spines.
The pinular ray of dermal pentactins is 0.113-0.200/0.022-0.032 mm, tangential rays are 0.070-0. 1 3 1/0.01 3-
0.018 mm. The pinular ray of atrial pentactins is 0.104-0.191/0.018 mm. tangential ones are 0.06 1 -0. 1 22/0.0 1 4-
0.018 mm.
Microhexactins and various derivatives (fig. 49.10-17) (micropentactins, microstauractins, microdiactins,
micromonactins and spheres) are covered with short numerous spines. These spicules have rays 0.078-
0.191/0.014 mm in length. A single type of amphidisc (fig. 49.18) of a more constant total length than that of
Poliopogon micropentactimis was found in P. zonecus. These amphidiscs have shafts covered with numerous short
spines and often have a widened part in the middle. Total length of micramphidiscs 0.025-0.047 mm. umbel
length 0.006-0.010 mm, umbel diameter 0.007-0.012 mm.
Remarks. — The specimen HCL 460 is a poor fragment. Its spicule content is incomplete and it is
tentatively identified as P. zonecus because of similarities of pinular pentactins and micramphidiscs. Micro¬
hexactins prevail in this specimen. As this specimen is tentatively identified, it does not receive the paratype
status. Poliopogon zonecus differs from P. micropentactinus in having different microscleres: amphidiscs are
smaller; microhexactins and various derivatives are found in P. zonecus whereas there are mainly pentactins and
some stauractins in P. micropentactinus. Some analogues can be found between microhexactins and their
derivatives in P. zonecus and Semperella abyssalis but most other spicules differ strongly in these species.
Uncinates with short spines are also one of the characters of this new species.
ETYMOLOGY. — The species is named after the HALIPRO 2-ZONECO cruise.
Genus PHERONEMA Leidy. 1868
Pheronema pilosum Levi, 1964
Fig. 50-54; Tab. 23
Pheronema pilosum Levi, 1964: 99, fig. 54.
Pheronema giganteum - BURTON, 1959: 159 (in part).
MATERIAL EXAMINED. — Madagascar. Northwestern Coast. "Akademic Kurchatov", stn 3754, 12°29'S. 48°05'E,
670 m: 5/2/390.9 (10RAN).
Zanzibar area. John Murray Expedition: stn 122, 5°21.24'S, 39°23.00'E. 745 m. 22.01.1934: NHM
1936.03.04.080.
Australia. Eastern Coast. Cidaris I: stn 24-2, 17°19.58'S, I47°47.61E, 1187-1200 m. 11.05.1986: fr870
(provisional number) (MTQ).
Source : MNHN, Paris
AMPH1D1SC0PH0RA OFF NEW CALEDONIA
101
Fig. 50-53. — Pheronema pilosum Levi, 1964: 50-51 (HCL 216); 52 (HCL 256); 53 (HCL 252). Seales = 2 cm.
Source . MNHN, Paris
102
K. R. TABACHNICK & C. LEVI
Indonesia (Flores Sea). " Snellius sin 164, 7°27.0'S. I24°27.5'E, 200 m: por9402.
Chesterfield Islands. MUSORSTOM 5: sin CP 323. 21°I8.52'S, 157°57.62'E 970 _m H<r*r
— Stn CP 324, 21o15.01'S, 157°51.33’E, 970 m, 14.10.1985: HCL 242-244. — Stn CP 387. 20 53.41 S. 160 5-.14E.
650Ne6w C a 1 e^ 3 on i'a B I OCA L : ^st n" CT 29, 23°08.70'S. 166°39.70'E. 1100 m, 29.08.1985 HCL 222. - Sin CP 30.
23°09 76'S 1 66°40.85'E. 1140 m; 29.08.1985: HCL 225. — Sin CP 57. 23°44.51 S. 166°54.94E. 1490-1620 m.
1 09 198V HCL 462. — Sin CP 61. 24°11.67’S, 167°31.37'E. 1070 m. 2.09.1985: HCL 226-227. — Sin CP 62,
24°19 35'S 1 67°49.43'E, 1395-1410 m. 2.09.1985: HCL 223-224. — Sin CP 69, 23°52.21'S. 167°57.82'E, 1220-
1225 m 3.09.1985: HCL 228. — Sin CP 75, 22°20.42'S, 167°23.41'E, 825-860 m, 4.09.1985: HCL 216-221, 461.
Loyalty Islands. Musorstom 6: stn CP 427, 20°23.35'S, 166°20.00'E. 800 m. 17.02.1989: HCL 245-247.
SmCP 438. 20°23.00'S. 166°20.10'E, 780 m, 18.02.1987: HCL 248-249.
BlOGEOCAL: stn CP 297, 20°38.64'S. I67°10.77'E, 1230-1240 m, 28.04.1987: HCL 256.
Combe Bank (near Wallis and Futuna Islands). MUSORSTOM 7: stn CP 551, 12°15.3S, 1 77°28. 1 W, 791 m.
18.05.1992: HCL 250-252.
Types. — Holotype at the Zoologisk Museum in Copenhagen. A spicular slide (MNHN-HCL 28) from the
holotype at the Museum national d'Histoire naturelle in Paris.
DESCRIPTION. — Ovoid sponge from 23 to 120 mm in length, 12 to 65 mm in diameter, with relatively large
osculum (8-50 mm in diameter), deep atrial cavity (20-70 mm) and relatively thin walls about 6-25 mm thick.
The sponge usually has well developed prostalia latcralia which protrudes at 20-50 mm from the surface and
prostalia oscularia. Basalia arc different in length but usually not very long.
Spicules : The choanosomal skeleton consists of pentactins and rare hcxactins, stauractins and triactins. Basalia
and sceptres are similar to those of other species of Pheronema. The uncinates belong to three types (lig. 54. 1 3-
15). The macruncinates are several mm long and about 0.009 mm in diameter, with spines developed normally.
This kind of uncinate is absent in some specimens. The mesuncinates are about 0.7-1.2/0.008 mm. with short
spines, and arc present everywhere. Small, whip-like micruncinates 0.091-0.912/0.003 mm are tuberculated or
rough rather than spinous. This whip-like uncinate is absent in some specimens (HCL 242 and NHM 1936.03.
04.080) including the holotype.
Dermalia and atrialia are pinular pentactins (fig. 54.1-8) usually similar to each other in shape and size. The
pinular ray of dermal pentactins is 0.046-0.304/0.006-0.026 mm, ol atrial, 0.053-0.356/0.006-0.013 mm. The
pinular ray is sharply pointed. The whip-like outer end projects freely beyond the last spines. The spines begin at a
short distance from the base of the shaft. The pinular ray of atrial pentactins is often curved whereas the dermal one
is usually straight. In both specimens collected from the western Indian Ocean (NHM 1936.03.04.080 and IORAN
5/2/390.3) the pinular ray is straight and only the outer end, which projects freely beyond the last spines, may
curve slightly. The pinular ray of dermal pentactins of HCL 256 is straight and not very thick, whereas that ol the
atrial ones is curved or straight. Thickness of atrial pinular ray is similar to that of the dermal one. The atrial
pinular ray may be whip-like with rare spines or oval in shape with conical or rounded outer end. Some long
spines of pinular ray are dichotomously branched. The tangential rays are rough and sharply pointed. I he
tangential ray of dermal pentactins is 0.038-0.236/0.006-0.008 mm, of atrial. 0.038-0.205/0.006-0.007 mm.
Micropentactins (fig. 54.9) with rough rays are rare in most specimens and they were not described from the
holotype. These micropentactins are canalaria or non developed pinular pentactins as their rays are similar to the
tangential rays of pinular pentactins. The rays of these micropentactins are 0.026-0.152/0.006 mm. The
microhexactins and microstauractins in addition to micropentactins are present in HCL 256. The microhexactins
prevail in HCL 462.
The amphidiscs (fig. 54.15-18), with shafts covered with numerous spines, belong to one type. Total length
0.022-0.189 mm, umbel length 0.007-0.052 mm, umbel diameter 0.008-0.045 mm. Their length varies about
3-4 times between different specimens. However, according to figures of the holotype they also vary about
3 times. The largest amphidiscs were observed in HCL 220. 232, 248, the smallest ones in HCL 217, 221, 222.
226, 243. According to the length of amphidiscs of other species of Pheronema. the amphidiscs of P. pilosum
correspond both to micramphidiscs and mesamphidiscs. They were described as mesamphidiscs in the original
description. In the close species, P. amphorae (Reiswig, 1992), these spicules are divided into micramphidiscs and
mesamphidiscs. Some specimens have fragments of these micramphidiscs, probably of allochtonous origin.
Source : MNHN, Pans
AMPHIDISCOPHORA OFF NEW CALEDONIA
103
Pher°”e™a P'^sum Levi. 1964: 1-4, dermal pentactins (1-2. HCL 461; 3. HCL 242: 4. HCL 220); 5-8. atrial
Li). ,',nS (5'6' HCuL.fl; 7- 5/2/390.9; 8, HCL 461); 9. micropentactin (HCL 461); 10. anchor-like basalia (HCL
IS ma/n,n^eP,re in1 ,4,6,V: 12- Scep,re <5/2/390-9); 13. mesuncinate (HCL 461); 14. micruncinate (HCL 461):
• acruncinate (HCL 461); 16, amplndisc (HCL 461); 17. amphidisc (HCL 243); 18. amphidisc (HCL 220).
104
K R. TABACHNICK & C. LEVI
Remarks. — Pheronema pilosum is widely distributed in the Indo-west Pacific. The closest species are
P. echinatum (Ijima, 1927), P. amphorae (Reiswig, 1992), P. barbulosclera (Levi. 1964) and P. carpenter i
(Schulze, 1887). Unfortunately, the specimens collected from the coast of Zanzibar by the "Valdivia" and identified
as P. carpenteri (SCHULZE, 1904) were not correctly described. They could also belong to P. pilosum since they
were collected in the Indian Ocean. If so, P. carpenteri inhabits only the central and north Atlantic Ocean.
Distribution. — Indian Ocean, Indonesia, western Pacific.
Pheronema semiglobosum Levi & Levi, 1982
Fig. 55-57; Tab. 24
Pheronema semiglobosum Levi & Levi, 1982; 286, fig. 2, pi. 2-3.
Material EXAMINED. — New Caledonia. Biocal: stn DW 08, 20°35.09'S. 166°54.05'E, 435 m. 12.08.1985:
HCL 261-267. — Stn DW 37, 23°00.07'S. 167°16.34’E. 350 m. 30.08.1985; HCL 268-270. 496. — Stn DW 44.
22°47 35'S, 167°14.50'E, 440-450 m. 30.08.1985: HCL 271. — Stn CP 45. 22°47.87'S, 1 67° 1 5.94'E, 430-465 m,
30.08.1985: HCL 260. — Stn DW 46, 22°53.27’S, 167°I7.41'E. 570-610 m, 30.08.1985: HCL 272-275. — Stn DW 51.
23°05.43'S. 1 67°45.35'E. 680-700 m, 31.08.1985: HCL 276. — Stn DW 83. 20°35.25'S. 1 66°54.03'E. 460 m.
6.09.1985: HCL 278-279, 293-294.
Musorstom 4: stn CP 213. 22°5 1.30'S. 167°12.00'E, 405-430 m, 28.09.1985: HCL 284-286. — Stn CP 214.
22°53.80'S, 167°13.90'E. 425-440 m. 28.09.1985: HCL 287-288. — Stn CP 215. 22°55.70'S. 167°17.00'E. 485-
520 m. 28.09.1985: HCL 289. — Stn DW 229, 22°51.60'S, 167°13.50'E, 445-460 m, 30.09.1985: HCL 290. —
Stn DW 230, 22°52.50'S, 167°11.80'E. 390-420 m, 30.09.1985: HCL 291. — No station number: HCL 292.
Smib 1: sin DW 2, 22°51.9'S. 167°13’E. 415 m, 5.02.1986: HCL 306.
Smib 4: stn DW 54, 23o40.1'S. 168°00.2'E, 235 m. 9.03.1989: HCL 307.
Loyalty Islands. BiOGEOCAL: stn DW 307, 20°35.38'S, 166°55.25'E. 470-480 m. 1.05.1987: HCL 280-283.
Musorstom 6: stn DW 391, 20°47.35'S, 167°05.70'E, 390 m, 13.02.1989: HCL 295-298, 463-465. — Stn DW 406.
20°40.65’S, 167°06.80'E. 373 m, 15.02.1989: HCL 299-300. — Stn DW 448, 20°55.66'S, 167°22.34'E. 410 m,
19.02.1989: HCL 301. — Stn DW 458, 21°00.93'S, 167°29.96'E, 400 m. 20.02.1989: HCL 302. — St. DW 487.
21 °23.30'S. 167°46.40'E. 500 m. 23.02.1989: HCL 303-304. — No station number: HCL 305.
Fig. 55-56. — Pheronema semiglobosum L6vi & Levi, 1982: 55 (HCL 298); 56 (HCL 294). Scales 2 cm.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
1 05
Types. — Holotype : HC1 48. Paratypes: HC1 49 - 17 specimens (New Caledonia. 22°46'S. 167°I4'E. 400-
410 m).
Description. — The body is usually hemispherical, rarely oval (HCL 282) or spherical (HCL 304). The tufts
of pleuralia oscularia and pleuralia lateralia protrude at about 5-40 mm. In one spherical sponge (HCL 304),
pleuralia atrialia similar to those of dermalia are found over the atrial surface. Basalia are usually about 30 mm in
length. The sponge (one of the paratypes) about 13 mm long and 13x8 mm in diameter has no prostalia or
basalia. The other paratype is 7 mm long and 12x10 mm in diameter, it has no basalia and no prostalia oscularia.
The specimen HCL 304 mentioned above is about 15 mm in diameter. The oval sponge HCL 282 is rather large,
about 55 mm long and 50 x 30 mm in diameter. One of the small specimens (HCL 496) has a slightly depressed
atrial cavity. The largest specimen (HCL 278) is 1 20 mm long and 130x155 mm in diameter.
FlG,57'. . ^picules of Pheronema semiglobosum Levi & Ldvi. 1982 (types, except 4-5. HCL 278): 1-2. anchorate
oasaiia, s, macruncinate; 4-6, micruncinates; 7. sceptre.
106
K R TABACHNICK & C. LEVI
Additional remarks on the skeleton spicules :
Hexactins prevail over pentactins in the choanosomal skeleton in some specimens (e.g. HCL 304). The basalia
sometimes have some teeth located at a short part of the rachis.
Prostalia oscularia and lateralia are sceptres (fig. 57.7) as in other Pheronema. The micruncinates (fig. 57.4-6)
are rare in the smallest specimens (HCL 304, 287, 282) and in two smallest paratypes; they seem to be absent in
HCL 278. Only two mesuncinates have been found in the smallest paratype specimen but they belong to
macruncinates (fig. 57.3) rather than to micruncinates. The micruncinates are 0.084-0.646 mm long.
Dermalia and atrialia are pinular pentactins and, rarely, hexactins similar in shape. The pinular ray of dermal
pentactins is 0.053-0.236 mm, of atrial, 0.076-0.372 mm. The pinular ray is straight or curved. The tangential
rays are entirely or partly (the upper end) covered with numerous spines. The tangential rays are straight or two
opposite pairs bent in an X-form. From side view the tangential rays are always slightly deviated away from the
pinular ray. The tangential ray of dermal pentactins is 0.023-0.099 mm, of atrial 0.027-0.133 mm.
Total length of macramphidisc 0.068-0.239 mm, umbel length 0.015-0.068 mm, umbel diameter 0.027-
0.076 mm. Total length of micramphidisc 0.017-0.065 mm, umbel length 0.005-0.013 mm, umbel diameter
0.005-0.013 mm. Micramphidiscs are rare in the smallest specimen (HCL 496).
Distribution. — New Caledonia, Loyalty Islands.
Pheronema conicum Levi & Levi, 1982
Fig, 58-63; Tab. 25
Pheronema conicum Levi & L6vi, 1982: 284. pi. I fig. I.
Material EXAMINED. — New Caledonia. Biocal: stn DW 08, 20°35.09'S, I66°54.05'E. 435 m, 12.08.1985:
HCL 312-315. — Stn DW 38, 22°59.94'S, 167°15.42'E. 360 m, 30.08.1985: HCL 316. — Stn DW 44. 22°47.35'S.
167°14.50'E, 440-450 m, 30.08.1985: HCL 317-318. — Sin CP 45, 22°47.87'S. 167°I5.94'E. 430-465 m. 30.08.1995:
HCL 310. — Stn CP 47, 22°53.10'S, 167°16.82'E, 550 m, 30.08.1985: HCL 311. — Stn DW 83. 20°35.25'S.
166°54.03'E, 460 m. 6.09.1985: HCL 319-326.
Musorstom 4: stn DW 228. 22°47.00'S, 167°18.20’E, 420 m. 30.09.1985: HCL 333-335. — Stn DW 230.
22°52.50'S, 1 67°1 1 ,80'E, 390-420 m, 30.09.1985: HCL 337-338. 467. — No station number: HCL 472.
Chalcal 2: stn CP 25, 23°38.60'S. 167°43.12'E. 418 m, 30.10.1986: HCL 329-332.
Calsub: stn 19, S to SW of the Isle of Pines, 22°46’S, 167°20'E. 416-404 m, 10.03.1989: HCL 328, 336.
Smib 1: no station number: HCL 343.
Loyalty Islands. Biogeocal: stn DW 307, 20°35.38'S. 166°55.25’E, 470-480 m, 1.05.1987: HCL 327.
Musorstom 6: sin DW 396, 20°48.05'S, 1 67°00.59E. 1400 m, 13.02.1989: HCL 469. — Stn CP 427, 20°23.35'S,
166°20.00'E, 800 m. 17.02.1989: HCL 471. — Stn CP 438, 20°23.00'S, 166°20.10'E, 780 m. 18.02.1989: HCL 470
— Sin DW 487. 21°23.30S, 167°46.40’E, 500 m. 23.02.1989: HCL 341-342.
Chesterfield Islands. Musorstom 5: stn DW 272. 24°40.9I’S. 159°43E. 500-540 m. 9.10.1986: HCL 466. —
Stn DW 300, 22°48.27'S, 159°23.94’E, 450 m, 11.10.1986: HCL 468. — Stn DW 301, 22°06.90'S. 159°24.60'E 487-
610 m. 12.10.1986: HCL 339-340.
TYPES. — Holotype : HCL 46. Paratypes : HCL 47 - 5 specimens collected from two stations (New Caledonia.
22°49'S, 167°12'E, 390-395 m).
Description. — The body is conical or spherical, with a relatively small osculum and atrial cavity. The body
length is 15-150 mm, maximal diameter is 19-130 mm, diameter of the osculum is 1-23 mm, length of the atrial
cavity is 12-80 mm. The holotype is 45 mm long and 60 mm in diameter, the diameter of the osculum is 8 mm.
The sponge has few basalia and pleuralia lateralia variously developed in different specimens. Pleuralia lateralia
protrude at 10-40 mm from the surface. They are found everywhere on the lateral surface and are absent in the
vicinity of the osculum. Pleuralia are concentrated sometimes in several tufts. Pleuralia lateralia seem to be
entirely absent in some specimens.
Spicules: The choanosomal spicules are pentactins (fig. 62.20) or rarely hexactins and stauractins (fig. 62.18-
19), with smooth, rarely rough rays several mm long and 0.02-0.09 mm in diameter. Prostalia lateralia arc
Source :.MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
107
diactins, partly smooth and partly rough or entirely smooth, several mm long and about 0.15 mm in diameter.
Basalia (fig. 62.22-24) are usual for other Pheronematidae anchorate spicules, with rare spines on the rachis.
The macruncinate (fig. 62.1 1) is thick, 0.010-0.012 mm in diameter, and long, with normally developed spines.
The whip-like micruncinates (fig. 62.12) are relatively short, with short spines or rough surface. They are approx¬
imately 0.17/0.0015 mm. Dermal spicules differ from the atrial. They both are pinular pentactins (fig. 62.1-10) or
rare hexactins. In some specimens (as in one paratype) the dermal hexactins arc notably numerous and atrial ones
even predominate over the pinular pentactins. The dermal pinular ray is sharply pointed, the upper end
108
K. R. TABACHN1CK & C. LEVI
terminates at the same level with the upper spines or sometimes projects beyond the ends of the last spines. Most
of the spines are long, hook-like, and are directed upward except the closest one to the base, which may be directed
downward. The spines begin at the upper 2/3 of the ray or at a short distance from the base. The tangential rays are
not sharply pointed, and are smooth with outer ends rough or short-spiny. Some rarely found dermal and atrial
pentactins have few thick and long spines. The pinular ray of the atrial pentactins is long, conically pointed, and
the upper end freely projects beyond the ends of last spines. The spines are directed upward, and they cover all the
shaft, being relatively short. Some spines near the base may be directed downward; they may split into two spines.
The pinular ray of dermal pentactins is 0.049-0.251/0.004-0.007 mm, of atrial 0.053-0.468/0.004-0.01 1 mm. The
tangential rays of the atrial pentactins are covered with numerous short spines. The tangential ray of dermal
pentactins is 0.038-0.190/0.004-0.006 mm, of atrial 0.038-0.462/0.004-0.006 mm.
Microhexactins (fig. 62.13-17) covered with short or long spines prevail over the analogous micropentactins
and microstauractins. The ray of microhexactins is 0.12-0.22/0.007-0.013 mm. The microtriactins and micro-
diactins with thick ray covered with spines described in the primary description (Levi & Levi. 1982) seem to
belong to another sponge, probably Seniperella schulzei. They were observed only in the holotype and paratypes
of P- conicum. Nearly all the other specimens that were examined had no such spicules.
Amphidiscs located in the vicinity of dermal and atrial surface arc different. Two types are found. Macram-
phidiscs with great variation in size and micramphidiscs more uniform in length. The macramphidiscs (fig. 63.1-
20) of the dermal surface are usually larger then the atrial ones, their size variation within a specimen is less
distinct but the size variation among specimens is distinct. Total length of dermal macramphidiscs 0.046-
0.312 mm, umbel length 0.019-0.106 mm, umbel diameter 0.023-0.106 mm. Most of these amphidiscs are
"normal" with the umbels spherical or elongated, whereas some rare forms have the shaft projecting beyond the
end of the umbel. Some other dermal macramphidiscs have different sizes of umbels but this difference is not as
distinct as in atrial macramphidiscs. Such proportions were previously known for some other species of the genus.
The shalts ol dermal macramphidiscs are often covered with tubercles or rarely with a few curved spines. In some
specimens (e.g. HCL 321) the dermal macramphidiscs were not found. The micramphidiscs from the dermal
surface are entirely similar to atrial ones. Total length of dermal micramphidiscs 0.014-0.063 mm, umbel length
0.005-0.020 mm, umbel diameter 0.005-0.014 mm. The atrial macramphidiscs are usually numerous but in some
paratypes they are rare. Total length of atrial macramphidiscs 0.038-0.243 mm, the umbel length 0.018-
0 084 mm, the umbel diameter 0.014-0.084 mm. In nearly all specimens the atrial macramphidiscs show a very
similar variation. If the atrial macramphidiscs are normally developed, the teeth of the umbels may be flattened,
tongue-like or rarely spinous, hook-like. Some macramphidiscs have one normally developed umbel, whereas the
other has a deformed upper surface, with the shaft protruding over the surface and the lack of some teeth. The other
amphidiscs have one reduced umbel, they end with a prolongation of the shaft and have some teeth situated in an
irregular manner, often finger-like. Such spicules were described in Hyalonema thomsoni (Schulze, 1905). The
other deformed macramphidiscs are paradise-shaped. This type of spicules was previously known in the family
Hyalonematidae, only in Hyalonema ( Paradisconema ) (Lima, 1927). The next deformed form, hemidiscs, have
umbels ol different sizes (length and diameter). This type of amphidisc, or birotulates, according to Reid (1958a;
1958b; 1961), were previously known in the fossil order Hemidiscosa (SCHRAMMEN. 1924). The proportion
between the diameters of the umbels of Hemidiscella schrammeni is 3.3 (Reid, 1958b). whereas in the paradises
ol P. conicum it is not more than 1.2. Another type of deformed macramphidisc has umbels with teeth which
meet or even overlap in the equator. These macramphidiscs are usually smaller in size than others but nevertheless
they cannot be considered mesamphidiscs because of the absence of overlap in size with such other amphidiscs. As
most other deformed amphidiscs, they are usually smaller than normal ones and they may be considered as a type
ol macramphidiscs rather than mesamphidiscs. The long teeth of macramphidiscs show several deformations. They
can turn to be paradises. Others demonstrate some fusions. Teeth at each end may merge to form an entire
spherical surface. The ends of these teeth are accreted to the middle of a shaft. The whole spicule looks like a pair
ol joined balls with latitudinal depressions. Fusion of the umbels of the amphidiscs was previously known in
Monorhaphis chuni (Schulze, 1905) but it is the fusion of the opposite teeth. The shafts of atrial macramphidiscs
are usually tuberculated, rarely smooth with a whorl of tubercles in the middle. All the strongly deformed
macramphidiscs are smaller than the complete ones. It nevertheless seems to be impossible to distinguish separate
Source :.MNHN, Paris
(12, HCL 334V 5 7n ZZ7 "7"'"” Levi & Levl <>yPes °'hers specimens): 1-4, dermal pinular pentactins
12, micruncinate ^ 1^ 1 S f 3 P'nU af (5‘ HCL 315; 6‘8' HCL 3I4; 9' HCL 324'^ "• macruncinate;
17. micropen tacttn <HCL aSl-TT"'18 * ' HCL 34°; l5‘ HCL 439): 16' -"icrolauactin ,HCL 439);
(HCL 331) 20 choano.S 18’ fhoa“al furaC.m (HCL 331); 19, outer end of choanosomal staurac.in
(24, HCL 322), ' h°anosomal pentacI,n; 21, choanosomal hexactin (HCL 332); 22-24. anchorate basalia
1 10
K R. TABACHNICK & C. LEVI
CM
Fig. 63. — Spicules of Pheronema conicum Levi & L6vi, 1982 (types and others specimens): 1-2. dermal
macramphidiscs; 3-13. atrial macramphidiscs (3, HCL 330; 7. atrial macramphidisc with prolonged shaft: 8-9. atrial
macramphidiscs with prolonged shaft and deformed umbel; 9. HCL 340; 10. paradise. HCL 339; II. atrial
macramphidisc with prolonged shaft and deformed umbel; 12-13. hemidiscs, HCL 339); 14-20, atrial oval
macramphidiscs (14, HCL 322; 15. HCL 323; 17, atrial oval macramphidisc with fused teeth. HCL 322; 18. optical
section of atrial oval macramphidisc with fused teeth. HCL 322; 19. atrial oval paradise, HCL 313; 20. HCL 314);
21-29. micramphidiscs (21, HCL 322; 24, micramphidisc with prolonged shaft; 25. paradise with prolonged shaft;
26, HCL 322; 27. hemidisc, HCL 337; 28. HCL 337; 29. HCL 314).
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
I 1 I
types of these curious forms. Total length of atrial micramphidiscs (fig. 63.21-29) 0.016-0.043 mm. umbel
length 0.004-0.013 mm, umbel diameter 0.005-0.012 mm. The micramphidiscs are mostly located in the vicinity
of dermal and atrial surfaces. They are usually regular in shape and size but sometimes they display some particular
deformations. Sometimes the umbels are different: one is pileate or spherical, whereas the other has a conical tip.
Sometimes some of the teeth of an umbel are of different length, so that the spicule is nearly a paradise. The
shafts are usually covered with tubercles or, rarely, they have widening in the middle. Some tubercles of the shafts
are long up to the length of the teeth, and have the teeth-like shape of some atrial macramphidiscs.
REMARKS. — Despite the specific forms of newly found amphidiscs, P. conicum is close to P. gigantemn
Schulze (stricto sensu). They both have similar uncinates of two types, one of which is small and whip-like, with
a rough surface rather than spinous. The microhexactins in both species predominate over other forms.
P. giganteum, like P. conicum, has oval amphidiscs (mesamphidiscs) with teeth meeting at the equator and
macramphidiscs with both pileate and spherical umbels. At least some deformations of the macramphidiscs were
found in the reexamined holotype of P. giganteum.
The description of "polyactin" spicules in some Pheronematidae that resulted from microhexactins
that developed extra rays is not usual for the subclass Amphidiscophora. This character belongs to another
subclass, Hexasterophora, which has "mainly or all astral microtriaxones" (Reid, 1958a). In the Amphidisco¬
phora, "the parenchymal microtriaxones are always holactins, never astral" (REID, 1958a). Does this curious case
again bring into doubt the system ol subclasses of Hexactinellida? These asterous spicules are very rare, and
they have been found among their ancestral forms the "holactines", so it is better to consider them as abnormal
forms and to ignore them as a problem of macrotaxonomy. The finding of hemidiscs in Pheronema conicum
leads to the discussion of the taxonomic status of the order Hemidiscosa (Reid, 1958a). or Hemidiscaria
(Schrammen, 1924). Reid (1958a) defined this order as "Amphidiscophora with birotulates in the form
of hemidiscs; other characters unknown". The hypothesis on the convergent evolution of hemidiscs and
amphidiscs (Mostler & Mehl, 1990) is true if only the hemidisc of Pheronema conicum evolved convergently
with "true" hemidiscs of Hemidiscosa. The only reference on the difference between Amphidiscosa (Reid, 1958a),
or Amphidiscaria (Sehrammen, 1924), and Hemidiscosa is the note that hemidiscs occupy a "different position
within the sponge body than the amphidiscs of Recent Amphidiscophora. and thus the hemidiscs might have been
also functionally different" (Mostler & Mehl, 1991). The definition does not seem to be sufficient enough for
a taxon of order rank and requires further study of data on spicule content. To give the order rank for
Hemidiscosa it must distinctly differ from that of Amphidiscosa. It is very likely that Hemidiscosa is a lower
taxon of the order Amphidiscosa of Schrammen ( 1924) since Pheronema conicum is clearly a representative of
Amphidiscosa.
Distribution. — New Caledonia, Loyalty and Chesterfield Islands.
Pheronema giganteum Schulze, 1886
Fig. 64; Tab. 26
^4,7“ SChU‘Ze’ 1886: 66; '887: 250' Pl‘ 45'46; 1893: 563‘ ~ 0KADA' ,932: 6‘ - |JIMA & OKADA.
Noi Pheronema giganteum - Lima, 1927: 10.
Not Pheronema giganteum - Burton, 1959: 176 (= Pheronema pilosum Levi, in part).
NHM A1887lfoL.20X10l"NED' ~ Indonesia "Challenger", stn 192, Little Kai Island, 5°49'15"S. 132°14'15"E, 235 m:
235 m^S ~H°IOtyPe NHM l887 10-20101 C Challenger ", stn 192, Little Kai Island, 5°49'15"S- 132°14'15"E,
Lolo““2y0L~b3;iT^0n,y S"Pplem“B ,hat of SC"“LZE <1887> s'"“ besides .he
112
K. R. TABACHNICK & C. LEVI
Fig. 64. — Spicules of Pheronema giganteum Schulze, 1887 (holotype): 1-4. dermal pinular pentactins; 5-8, atrial
pinular pentactins; 9-10, microhexactins; 11-12. macramphidiscs; 13-14, oval mesamphidiscs; 15, micram-
phidisc; 16, macruncinate; 17, whip-like micruncinate; 18, mesuncinate; 19, anchorate basalia; 20, diactin with
smooth and rough parts.
Source : MNHNJ Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
I 13
Spicules: The uncinates belong to three types. Macruncinates (fig. 64.16) are several mm long and about
0.005-0.010 mm in diameter. Mesuncinates (fig. 64.18) are relatively short, about 0.35-0.72/0.005 mm. Both
macruncinate and micruncinate are covered with spines typical for these types of spicules as in other
Pheronematidae. Micruncinates (fig. 64.17) (0.13-0.30/0.001 mm) are whip-like. They are covered with very
shallow and obtuse spines. Dermal and atrial pinular pentactins (fig. 64.1-8) are similar. The pinular ray is oval or
whip-like in shape, rarely slightly curved. The outer end is usually sharply pointed or conical. It can project far
beyond the last spines or more often be equal in length to the last spines. The lower spines sometimes branch into
two spines. The pinular ray of dermal pentactin is 0.068-0.334/0.004-0.040 mm, of atrial one 0.076-0.266/0.004-
0.040 mm. The tangential rays are often smooth near the base and on the first half of the ray, whereas the second
half is rough or covered with spines. The tangential ray of dermal pentactin is 0.030-0.084/0.004. of atrial one
0.038-0.068/0.004 mm.
Microhexactins (fig. 64.9-10) and micropentactins have rays (0.061-0.152/0.005-0.012 mm) covered with
spines. Amphidiscs are divided into three classes. The macramphidiscs (fig. 64. 1 1-12) often have teeth of different
length, spherical umbels and tuberculated shafts. Total length of macramphidisc is 0.167-0.289 mm, the umbel
length 0.068-0.084 mm, the umbel diameter 0.053-0.084 mm. The mesamphidiscs (fig. 64.13-14) are oval with
teeth that nearly meet at the equator. All were found among the atrial spicules. Total length of mesamphidisc is
0.032-0.054 mm, the umbel length 0.016-0.027 mm, the umbel diameter 0.016-0.032 mm. Micramphidiscs (fig.
64.15) have spiny or tuberculated shafts. Total length of mesamphidisc is 0.023-0.041 mm, the umbel length
O. 007-0.018 mm, the umbel diameter 0.005-0.012 mm.
Remarks. In most of the features mentioned above P. giganteum is closer to P. conicum than to
P. giganteum described by Ijima (1927). Both P. giganteum and P. conicum have deformations of amphidiscs, the
lormcr has some deformations not so "deep" as the latter. These abnormal amphidiscs were not described by
SCHULZE (1887) despite that they prevail over "normal" ones. The uncinate composition, sizes of microhexactins
and micro-pentactins of P. giganteum of SCHULZE are similar to P. conicum rather than to P. giganteum of IJIMA.
These facts distinguish the latter as a separate species. The status of the other specimens of P. giganteum described
in the literature is not clear. The specimen described by Ijima and Okada (1938) is likely to be P. giganteum.
The specimen described by Okada (1932) is less clear because of its poor description. The reexamination of the
specimen described by Burton (1959) leads to transfer it to P. pilosum. As for the specimen described by Ijima
(1927), it seems better to refer it to a new species. Some specimens from the eastern and western Indian Ocean
lkcly belong to P. giganteum. but probably to a new subspecies. They have no mesamphidiscs and sometimes no
macramphidiscs (unpublished material).
Distribution. — Indonesia, Japan.
Pheronema pseudogiganteum sp. nov.
Fig. 65; Tab. 27
Pheronema giganteum Ijima, 1927: 10, pi. 5 fig. 1-7. (Non Schulze. 1886).
P- ma™n by I^mT 1927.' ~ "Siboga": s,n 25>- 5°28-4’S- 132°00.2'E, 204 m: PQR 5099 (ZMA). Identified as
1 ypes. — Holotype: PQR 5099 (" Siboga ", stn 251, 5°28.4'S, 132°00.2’E, 204 m) (ZMA).
mirnEMARKS JllC ,ollowing notes on spicules complete the excellent description of Ijima (1927). The
MicmhexTr-t S /f- arC Sh°rt’ USually with a widening in the middle. They are covered with minute spines.
rav„ 'nS I 'S’ , ‘ 4) prevai1 over analogous micropentactins and other derivatives. Microhexactins have
the snh • C^'n engt whictl carry spines of different length. Some rays in the microhexactins are reduced up to
end of I01""' ends\covered with nu™rous spines (nearly pinular). Such outer ends are characteristic of one
lcromonactins, whereas the other is sharply pointed. These micromonactins are rare. Two kinds of
K. R. TABACHNICK & C. LEVI
] 14
amphidiscs arc present. The macramphidiscs (fig. 65.10) have umbels that arc regular in shape, with teeth equal in
length. The shafts are tubcrculated. The micramphidiscs (fig. 65. 1 1 ) are similar in shape to the macramphidiscs.
Fig. 65. — Spicules of Pheronema pseudogiganteum nov. sp. (holotype): 1-4, microhexactins; 5, micromonactin;
6, mesuncinate; 7. macruncinate; 8-9, micruncinates; 10, macramphidisc; 11, micramphidisc.
The reasons for distinguishing a new species P. pseudogiganteum were given by the revision of the holotype.
Ijima did not revise the holotype, which was described insufficiently (Schulze, 1887). The features specific for P.
pseudogiganteum are regular macramphidiscs, absence of mesamphidiscs, size and shape of micruncinates and
microhexactins. Stronger similarities were found between P. giganteum and P. conic uni on one hand and
P . pseudogiganteum of Ijima and its forma from the New Caledonian area described below on the other hand.
Etymology. — This specific name alludes to the differences with Pheronema giganteum.
Pheronema pseudogiganteum forma nuda nov.
Fig. 66-67, 70; Tab. 28
Material EXAMINED. — New Caledonia. Biocal: stn DW 51, 23°05.43'S, 167°45.35'E. 700-680 m.
31.08.1985: HCL 360. — Stn CP 52, 23°06.18'S, 167°47.07'E, 600-540 m, 31.08.1985: HCL 358-359.
Description. — The sponge is ovoid, with a deep atrial cavity. The specimen HCL 358 is 70 mm in length,
45 mm in diameter. The osculum is 20 mm in maximal diameter. The walls are about 10 mm thick. Prostalia are
represented by basalia only. Basalia are about 30 mm long. The specimen HCL 359 is 100 mm in length. 30 mm
in diameter. The specimen HCL 360 is a fragment.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
Fig 66-67. — Pheronema pseudogiganteum f. nuda (HCL 358). Scales = 2 cm
FIG. 68-69. — Pheronema pseudogiganteum f. vaubana. (HCL 353). Scales = 2 c
in c\p!ml,eS choanosomal skeleton consists of pentactins and sparse stauractins. with rays 0.014-0.100 mm
ietcr. Ihree types of uncinates are known. Macruncinates (fig. 70.16) are several mm in length and
116 K. R. TABACHNICK & C. LEVI
Fig. 70. — Pheronema pseudogiganteum f. nuda (HCL 358): 1-2, dermal pinular pentactins; 3-7. alrial pinular
pentactins; 8, micropentactin; 9. microhexaetin; 10, macramphidiscs; 11-12, micramphidiscs; 13-14. micrun-
cinates; 15. mesuncinate; 16. macruncinale; 17, anchorate basalia.
0.008 mm in diameter. The mesuncinates (fig. 70.15) are about 0.45-0.52/0.004-0.009 mm. They are both
covered with spines. The micruncinates (fig. 70.13-14) are spindle-like, 0.041-0.1 17/0.004-0.007 mm. entirely or
partly serrated. The basalia are anchorate spicules with smooth rachis. Dermalia are pinular pentactins (fig. 70. 1-7)
with the pinular ray rather thin, 0.061-0.220/0.009-0.015 mm. The outer ends are finely pointed and project not
far beyond the last spines. The tangential rays of dermal pentactins 0.046-0.152/0.005-0.007 mm are usually
smooth at the base and covered with spines near the ends of the rays. Atrialia are pinular pentactins. The pinular
ray (0.084-0.144/0.007-0.037 mm) varies in shape; sometimes it is similar to that of dermal pentactins but
Source : MNHN. Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
117
usually ii is rather thick and has more or less distinct oval shape. The outer ends are longer or equal or smaller
than the last spines. The tangential rays (0.030-0.091/0.004-0.01 1 mm) are usually covered with spines. Pinular
hexactins are sparse dermal and atrial spicules.
The microhexactins and micropentactins (fig. 70.8-9) have rays (0.046-0.213/0.004-0.01 1 mm) covered with
spines. Amphidiscs are divided into two types. Umbels of macramphidiscs (fig. 70. 10) are regular in shape, shafts
smooth, usually straight or rarely with widenings. Total length of macramphidiscs 0.152-0.236 mm. the umbel
length 0.053-0.084 mm, the umbel diameter 0.061-0.099 mm. Shafts of micramphidiscs (fig. 70.1 1-12) are
covered with spines. Total length of micramphidiscs 0.027-0.049 mm, umbel length 0.008-0T)16 mm, umbel
diameter 0.007-0.013 mm.
Etymology. — The forma name refers to the absence of prostalia except basalia.
Pheronema pseudogiganteum forma vaubana nov.
Fig. 68-69, 71; Tab. 29
Material EXAMINED. —New Caledonia. Musorstom 4: stn CP 217, 22°03.60'S, 167°27.00'E, 850 m,
29.09,1985: HCL 363.
Description. — The sponge is ovoid, 55 mm long, 25 x 35 mm in diameter. The atrial cavity is deep,
the walls are about 5 mm in thickness. The sponge has two longitudinal rows on the dermal surface that extend
trom the base to the osculum. They are the result of growth rather than a character of the subspecies. Prostalia
oscularia are about 3 mm long, they edge the osculum regularly. Basalia are a broad tuft with same diameter as
the lower part of the sponge. They are about 15-20 mm long.
Spicules: I he choanosomal spicules are pcntactins with rays 0.05-0.11 mm in diameter. The uncinates
demonstrate three varieties. Macruncinates (fig. 71.10) are several mm long and 0.005-0.010 mm in diameter The
mesuncinates (fig. 71.11) are about 0.70-1.00/0.001-0.002 mm. They are both covered with spines. The
micruncinates (iig. 71.12-18) are 0.038-0.182/0.004-0.010 mm, spindle-like and partly serrated. Prostalia oscularia
are sceptres (fig. 718) 3-5/0.02 mm. Basalia are anchorate spicules (fig. 71.9) with smooth rachis. Dermalia are
pinular pcntactins (fig. 71.1-5) with pinular ray 0.068-0.1 14/0.007-0.010 mm slightly widening. Its outer end is
f. m/n nnso mn Pr°JeCtS "0t far fr°m lhc ends of ,he last sPines- The Pinular ray of lllt atria' pentactins 0.084-
. ' ' mm 1S similar to that of dermal ones, sometimes it is curved or oval in shape. If oval, the
u er end is equa in length with the last spines. Tangential rays both of dermalia and atrialia are covered with
0P007Smm8entia **** °f ** demia' penlactins are 0.046-0.084/0.005 mm, of the atrial ones 0.038-0.1 14/0.005-
tvn,MT,heXSnS (f'g' 7‘'6~7) 3nd microPentactins have rays (0.076-0.167/0.004 mm) covered with spines. Two
and hav e , T mSCh f" dlstmgulshed- Macramphidiscs (fig. 71.19-20) have umbels that are regular in shape
length of lb in'!'nrfmeS haVC S°me widenings’ Total len§th of macramphidiscs 0.167-0.251 mm,
do 'at 1 h f 5mm’ Umbe' diame'er 0068-°106 mni. The micramphidiscs (fig. 71.21-22) are
0 007*0 018 mm a ,L')V,ered Wlth sP,nes- ToIal lenSth of micramphidiscs 0.029-0.058 mm, length of umbel
U.UU/-0.018 mm, umbel diameter 0.007-0.013 mm.
and^h Jcond Sea N3med at'er lhe "Vauban"- which has been used in many research cruises in New Caledonia
Pheronema pseudogiganteum forma stellata nov.
Fig. 72; Tab. 30
2 7 . 0T 198^ ' H C L *^355^3 5^ ^7 T4 7 9 3 ' * ^ IS'andS Bl0GE0CAL; *tn CP 290, 20°36.91'S, 167°03.34'E, 920-760 m,
118
K. R. TABACHNICK & C. LEVI
0.4 mm 0.1 mm 0.05 mm
9 10-18 21-22
Fig. 71. — Spicules of Pheronema pseudogiganteum f. vaubana (HCL 353): 1, dermal pinular pentactin; 2-5. atrial
pinular pentactins; 6-7, microhexactins; 8. sceptre; 9, anchorate basalia; 10. macruncinate; 11, mesuncinate;
12-18, micruncinates; 19-20, macramphidiscs; 21-22, micramphidiscs.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
1 19
Description. — This sponge is represented by different fragments. Since they were collected from the same
station, these fragments must belong either to a single or numerous specimens. In the first case, the observed
variability of spicules results from the different location of fragments in the sponge, in the other, from fragments
from separate specimens. Most fragments are parts of the 5-10 mm thick walls. The fragment assigned as holotype
is 7-8 mm thick. The other fragments are parts of dermalia and subdermalia with tufts of basalia.
Spicules: The choanosomal skeleton consists of pentactins with rays 0.002-0.004 mm in diameter.
Macruncinates (fig. 72.5) are several mm long and 0.01 mm in diameter. The mcsuncinates (fig. 72.6) are about
0.3/0.002 mm. Both macruncinates and mesuncinates are covered with spines. The micruncinates (fig. 72.7-8)
0.041-0.1 18/0.004-0.008 mm appear smooth or slightly serrated. The basalia are anchoratc spicules (fig. 72.4)
with smooth rachis. Dermalia and atrialia are pinular pentactins (fig. 72.1-3) similar in shape. The pinular ray is
rather thin, finely pointed with the outer end that does not project far beyond the last spines. The pinular ray of
dermal pentactin is 0.042-0.198/0.007-0.020 mm, of the atrial one 0.087-0.555/0.010-0.026 mm. The tangential
rays are smooth, with outer ends covered with spines. The tangential rays of dermal pentactin are 0.034-
0.106/0.006 mm, of the atrial ones 0.038-0.144/0.006 mm.
Microhexactins (fig. 72.16-21) with thick rays 0.046-0.198/0.01 1-0.015 mm, prevail on the other types of
analogous spicules: micropentactins, microstauractins, and spicules with more than six rays. These microscleres
have rays usually covered with spines, seldom smooth. They are conically pointed or rarely have a rounded end.
The other spicules presented by microhexactins only have thin rays about 0.004 mm in diameter and covered with
spines. Ray length is equal to that of thick-rayed forms. The thin-rayed forms are sparse. Amphidiscs can be
divided into three types. Macramphidiscs (fig. 72.9-10) have umbels regular in shape, shafts are smooth. Total
length of macramphidiscs 0.152-0.228 mm, umbel length 0.042-0.068 mm, umbel diameter 0.068-0.1 18 mm.
Mesamphidiscs (fig. 72.1 1-13) and micramphidiscs (fig. 72.14-15) are similar in shape and size. Their shafts are
covered with numerous spines. Total length of mesamphidiscs 0.034-0.092 mm, umbel length 0.013-0.034 mm.
umbel diameter 0.012-0.030 mm. Total length of micramphidiscs 0.020-0.043 mm, umbel length 0.005-
0.018 mm, umbel diameter 0.005-0.014 mm.
Etymology. — The forma name refers to the shape of microhexactin spicules.
Pheronema pseudogiganteum forma variodisca nov.
Fig. 73; Tab. 31
Material EXAMINED. — New Caledonia. Biocal: stn CP 54, 23°10.30'S, I67°42'E, 1000-950 m, 1.09.1985:
Hv_L ^04.
description. — The sponge is ovoid. 55 mm long, 40 mm in diameter. The osculum is 20 mm in diameter,
the atrial cavity is about 20 mm long.
Spicules: The choanosomal spicules are pentactins with rays 0.04-0.08 mm in diameter. The uncinates are of
!/ee^f Macrun^'nates (fig. 73.10) are several mm long and 0.005 mm in diameter. The mesuncinates
f 7 , ;?nfe ab°Ut 070- 1.00/0.004-0.010 mm. They are both covered with spines. The micruncinates
.'g' 4) 0.030-0.084/0.004 mm are spindle-like, and appear smooth under light microscope. Dermalia are
pinular pentactins (fig. 73.1-6) and sparse hexactins with pinular ray (0.068-0.137/0.004-0.012 mm) sharply
Si^r^r1- 6nd lhat pr°jects free|y bey°n(J ihe last spines. Pinular ray in the atrial pentactins 0.091-
<h- ,7, 3 mm' P‘nular ray of alrial Pentactin ^ similar to that of the dermal ones or is rather ovoid in
H pm’i Wlt| llC term'nal'n£ at ’^e enc*s op 'ast spines. The tangential rays are covered with spines in both
t, ,d. an atna P,nular spicules. The tangential ray of the dermal pentactin is 0.042-0.080/0.002-0.007 mm. of
the atrial pentactin 0.049-0.1 14/0.006-0.010 mm.
snin^CxT‘Cr0helaCJlinS and micr°Pentactins (fig- 73.7-8), with rays 0.057-0.129/0.004 mm, are covered with
have iden?Cra|mP k 'i CS ^ j®' 73-15-20) have umbels and teeth of various shapes. The "normal" macramphidiscs
different mlmh UIT1f ? S a" QVal °r sonietimes lhin hook-like teeth. Umbels of some macramphidiscs have a
ei o teet i ( - ) in each umbel. Umbels of other amphidiscs differ in diameter because the teeth of
120
K. R. TABACHNICK & C. LEVI
476); 3. alrial pinular pentactin (HCL 355); 4, anchorate basalia (HCL 357); 5. macruncinate (HCL 355);
6, mesuncinate (HCL 355); 7-8, micruncinate (HCL 355); 9-10, macramphidiscs (9. HCL 478; 10. HCL 357);
11-13 mesamphidisc (11-12, HCL 475; 13, HCL 478); 14-15. micramphidiscs (HCL 478); 16-21, microhexactins
(16-19, HCL 478; 20. HCL 356; 21, HCL 478).
Source : MNHN, Paris
AMPHID1SCOPHORA OFF NEW CALEDONIA
121
CD
F'O-73. Spicules ol Pheronema pseudogiganteum f. variodisca (HCL 364): 1-2, dermal pinular pentactins;
-6, atrial pinular pentactins; 7, microhexactin; 8. micropentactin; 9, anchorate basalia; 10. macruncinate;
11-13, mesuncinates; 14, micruncinate; 15-20. amphidiscs.
the umbel are closer to the shaft, which makes the umbel smaller. The shafts are smooth but sometimes with
wtdenings or tubercles. Total length of macramphidiscs 0.137-0.228 mm, umbels length 0.046-0.076 mm.
umbels diameter 0.053-0.091 mm. Only two micramphidiscs were found in a microscopic preparation. They
probably belonged to another specimen.
Etymology. — The forma name refers to the abnormal umbels of some amphidiscs.
Remarks. All these specimens belong to Pheronema pseudogiganteum sp. nov. It is possible that the
\ ariation of shape and size ot dermal and atrial pinular pentactins, of microhexactins and its derivatives, together
^iti tie combination and shape of amphidiscs will be good features for a future taxonomic subspecific definition.
But these features are mosaically expressed in the different specimens.
122
K. R. TABACHNICK & C. LEVI
ACKNOWLEDGMENTS
We thank D. DOUMENC, A. CROSNIER, C. Valentine, R. Van SOEST, and M. Segonzac for the
possibility of studying the collections at the Museum national d'Histoire Naturclle (Paris), The Natural History
Museum (London) and the Zoologisch Museum (Amsterdam), J. Vacelet and M. Roux for the loan of colour
photographs of the CALSUB expedition, R. VON COSEL and P. LOZOUET for the photographs, T.A. SAVILOVA
for her thorough and numerous measurements of spicules, J.-F. DEJOUANNET for setting up photographs and
drawings and P. Castro for improving the language.
Special thanks are extended to H.M. REISWIG, Redpath Museum (Montreal) for critically reading draft of the
manuscript and suggesting several improvements.
This work was supported by grants from CNRS, Museum national d'Histoire naturclle (Paris), International
Science Foundation and Royal Society (London).
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LEvt, C. & LEvi. P., 1989. — Spongiaires (Musorstom I ct 2). In: J. Forest (ed.), Resultats des Campagnes Musorstom,
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Li Jinhe, 1987. — Monorhaphis intermedia, new species of Hexactinellida. Oceanologia and Limnologia sinica, 18 (2):
130-137, pi. 1-2.
Marshall, W. & Meyer, A.B., 1879. — Ueber einige neue und wenig bekannte Philippinische Hexactinelliden.
Mitteilungen aus dem K. Zoologischen Museum zu Dresden. 2: 263-279.
Mostler. H. & Mehl, D.. 1990. — On the origin of Hexasterophora and Amphidiscophora (Hcxactinellidae). A future
discussion of their phylogenetic significance. Fossil Cnidaria. International Newsletter, 19 (2): 13-15.
Okada, Y.. 1928. — On the development of a Hexactincllid sponge Farrea sollasi. Journal of the Faculty of Science,
Tokyo University, sect. 4 Zoology. 2(1): 1-27.
Okada, Y., 1932. — Report on the Hexactinellid sponges collected by the United States Fisheries steamer "Albatros" in
the Northwestern Pacific during the summer of 1906. Proceedings of the United Stales National Museum. 81 (12):
1-118. pi. 1-6.
Reid, R.E.H., 1958a. — A Monograph of the upper Cretaceous Hexactinellida of Great Britain and Northern Ireland.
Parti. Palaeontographical Society, London, 111: i-xlvi.
Reid, R.E.H., 1958b. — A Monograph of the upper Cretaceous Hexactinellida of Great Britain and Northern Ireland.
Part II. Palaeontographical Society, London, 112: xlvii-xlviii + 1-26, pi. 1-4.
Reid, R.E.H., 1961. — A Monograph of the upper Cretaceous Hexactinellida of Great Britain and Northern Ireland.
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Reiswig, H.M., & Zgraggen, M.N., 1991. — Sericolophus (Pheronematidac) and Hernandeziana (Corythophora) are
synonymes (Hexactinellida: Porifera). Zoological Journal of the Linnean Society. 103: 61-73.
Reiswig. H.M., 1992. — First Hexactinellida (Porifera) (Glass sponges) from the Great Australian bight. Record of the
South Australian Museum, 26 (1): 25-36.
Reiswig, H.M.. & Champagne, P., 1995. — The NE Atlantic glass sponges Pheronema carpenteri (Thomson) and P. gravi
Kent (Porifera: Hexactinellida) are synonymes. Zoological Journal of the Linnean Society of London. 115: 373-384.
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ouvelle-Caledonie. In: A. Crosnier (ed.), Resultats des Campagnes Musorstom, vol. 6. Memoires du Museum
national d Histoire naturelle, s6r. A, 145: 9-54.
RtOHER DE Forges, b. & Chevillon, C„ 1996. — Les campagnes dechantillonnage du benthos bathyal en Nouvelle-
caiedome, en 1993 et 1994 (Bathus I a 4, Smib 8 et Halipro 1). In: A. Crosnier (ed.), Resultats des Campagnes
MUSORSTOM, vol. 15. Memoires du Museum national d'Histoire naturelle, 168: 33-53.
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t-utuna. Compte rendu et liste des stations. In: A. Crosnier (ed.), Resultats des Campagnes Musorstom, vol. 10.
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R° FoRrpi rOUd,1EI’/oBOURSEAU’ J R- Gaillard- C„ Grandperrin, R.. Laurin. B„ Monniot. C.. Richer de
x \By R'°- M" Segonzac, M.. Vacelet, J„ Zibrowius, H„ 1991a. - L'&agement du benthos bathyal observe
a 1 aide de la soucoupe Cyana". Documents Travaux I.C.A.L., 15 151-165
DE
R° Forges Br°Ur?oTmR'cBOURSEAU; ,J P ' Gaillard- c- Grandperrin, R„ Laurin, B„ Monniot, C.. Richer
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de la SociM S ' S c'S P^'™res de la campagne Calsub et consequences paltSoecologiques. Bulletin
ae ta doctete geologique de France, 162 (4): 675-685.
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K. R. TABACHNICK & C. LEVI
Schrammen, A. 1924. — Die Kieselspongien der oberen Kreide von Nordwestdeutschland. III. und Letzler Toil mit
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Akademie der Wissenschaften zu Berlin, aus dem Jahre 1886: 1-97.
Schulze, F.E., 1887. — Report on the Hexactinellida collected by HMS "Challenger" during the years 187.1-1876.
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Schulze, F.E., 1893. — Revision des Systemes der Hyalonematiden. Sitzungsberichte der Koniglich-Preussischen
Akademie der Wissenschaften zu Berlin, 30: 541-589.
Schulze, F.E., 1894. — Hexaclinelliden des Indischen Oceans. 1. Theil. Die Hyalonematiden. Abhandlungen der
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Schulze, F.E., 1895. — Hexactinelliden des Indischen Oceans. II. Theil. Die Hexasterophora. Abhandlungen der
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SCHULZE, F.E., 1899. — Uber Hyalonema affine W. Marshall. Sitzungberichte der Gesellschaft Naturforschender Freunde
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Schulze, F.E., 1900. — Hexactinelliden des Rothen Meeres. Bericht der Commission fiir Ocectnographische
Forschungen. Zoologische Ergebnisse, 16: 31 1-324, 3 pi.
SCHULZE, F.E., 1902. — An Account of the Indian Triaxonida collected by the Royal Indian Marine Survey Ship
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Schulze, F.E., 1904. — Hexactinellida. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer
"Valdivia" 1898-1899, 4: 1-266, 52 pi.
Semper, C., 1868. — Einige neue Kieselschwamme der Philippinen. Verhandlungen der pliysikalisch und medicinischen
Gesellschaft zu Wurzburg, 1: 29-30.
Tabachnick, K.R., 1988. — Hexactinellid sponges from the mountains of West Pacific. In: Structural and functional
researches of the marine benthos. Academy of Science of the USSR, P.P. Shirshov Institute of Oceanology: 49-64.
Tabachnick, K.R., 1990. — Hexactinellid sponges from the Nasca and Sala-y-Gomez ridges. Trudy Institute i
Okeanologii. Akademiya Nauk USSR. Moscow. 124: 161-173.
Tabachnick. K.R. & Levi, C., 1999. — Revision of the genus Lophophysema with the description of two new species.
Invertebrate Taxonomy, 13: 495-509.
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THOMSON, C.W., 1877. — The voyage of the 'Challenger'. The Atlantic. A preliminary account of the general result of
the exploring voyage of H.M.S. 'Challenger' during the year 1873 and the early part of the year 1876, Macmillan and
Co., London, 1: i-xxx + 1-424, frontispiece, 14 pis.
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Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
125
ANNEXE
MENSURATIONS OF SPICULES
In all the tables : L = length; D = diameter. All measurements in mm.
Table 1. — Some measurements of the spicules of Hyalonema ( Leptonema ) spatha.
Table 2. — Some measurements of the spicules of Hyalonema (Onconema) uncinata.
L dermal pentactin pinular ray
L dermal penctatin tangential ray
L atrial pentactin pinular ray
L atrial pentactin tangential ray
L acanthophore pentactin ray
L microhexactin ray
L macramphidisc
L macramphidisc umbel
D macramphidisc umbel
L micramphidisc
L micramphidisc umbel
D micramphidisc umbel
126
K. R. TABACHNICK & C. LEVI
Table 3. — Some measurements of the spicules of the subgenus Onconema. From von Lendenfeld, 1915.
Table 4. — Some measurements of the spicules of Hyalonema (Ptefonema) topsenti.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
127
Table 5. Some measurements ol the spicules of Hyalonema ( Oonema ?) microstauractina.
L small dermal pentactin pinular ray
L small dermal pentactin tangential ray
L large dermal pentactin pinular ray
L large dermal pentactin tangential ray
L dermal diactin pinular ray
L dermal diactin proximal ray
L small atrial pentactin pinular ray
L small atrial pentactin tangential ray
L large atrial pentactin pinular ray
L large atrial pentactin tangential ray
L atrial diactin pinular ray
L atrial diactin proximal ray
L microstauractine ray
L macramphidisc
L macramphidisc umbel
D macramphidisc umbel
L mesamphidisc
I. mesamphidisc umbel
D mesamphidisc umbel
L micramphidisc
L micraniphidisc umbel
D micratnphidisc umbel
Lilian ucinidi pentactin pinular ray
small dermal pentactin tangential ray
L large dermal pentactin pinular ray
L large dermal pentactin tangential ray
L dermal diar.rin ninnlnr row
~ v.vnnai punamn tun
L dermal diactin ninular rav
I A . I 1 71 r - - r^~
_ muvuii t’liiuidi my
L dennal diactin proximal rav
L small atrial pentactin pinular rav
I . s; m nil ofriol _ .• i
anidii auidi pentactin pinular ray
L small atrial pentactin tangential ray
L large atrial pentactin pinular ray
L large atrial nentnnin tonrr»>nti»i r
iu|sc cunai pentactin pinular ray
_L large atrial pentactin tangential ray
L atrial diactin pinular rav
— . lav
L atrial diactin proximal ray
L microstauractin rav
L macramphidisc
1. macramphidisc umbel
D macramphidisc utnbeT
t -
L mesamphidisc
L mesamphidisc umbel
I'V _ _ I • - - - 7
— - .■■■.p.t.viljy, U 1 1 ILH71
D mesamphidisc umbel
L micramphidisc
L micramphidisc umbel
D mierainphidisc umbel
Source :
128
K. R. TABACHNICK & C. LEVI
Table 6. — Some measurements of the spicules of Monorhaphis which could be referred to M. chuni sensu stricto.
Table 7. —
Some measurements of the spicules of Monorluipliis which could be referred to previous M. dives.
Source : MNHN , Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
129
Table 8. — Some measurements of the spicules of Monorhaphis which could be referred to previous M. intermedia.
130
K. R. TABACHNICK & C. LEVI
Table 8. — Some measurements of the spicules of Monorhaphis which could be referred to previous M. intermedia (suite).
Table 9. — Comparative measurements of various spicule parameters of Monorhaphis collected from different locations
and of previously different species.
I - " M. chum " according to SCHULZE (1904)
II - " M. chuni " according to IJ1MA (1927)
III - specimens similar to " M. chum 5/2/1422; 5/2/1352; 5/2/1408; 5/2/135 1
IV - " M. dives 1908.09.24.065; 1908.09.24.064; kt476
V - " M. intermedia " according to Ll JINHE (1987)
VI - specimens off New Caledonia and Australia similar to " M. intermedia "
Table 10. — Some measurements of the spicules of Sericolophus calsubus.
Source : MNHN , Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
131
Table 11. — Some measurements of the spicules of Sericolophus neocaledonicus.
Table 12. — Some measurements of the spicules of Sericolophus hawaiicus.
Table 13. — Some measurements of the spicules of Sericolophus cidaricus.
Source : MNHN, Paris
132
K. R. TABACHNICK & C. LEVI
Table 14. — Ranges of spicule measurements of Sericoloplius species.
Table 15. — Some measurements of the spicules of Semperella varioactina.
Table 16. — Some measurements of the spicules of Semperella abyssal is.
Source : MNHN, Paris
AMPHIDISCOPHORA OFF NEW CALEDONIA
133
Table 17. — Some measurements of the pinular pentactins of Semperellci crosnieri (HCL 209) collected from different
places of the surface.
It - areas of the surface covered with small-meshed laticework - probably dermalia
* - areas of the surface covered with big-meshed laticework - probably atrialia
Table 18. — Some measurements of the spicules of Semperellci crosnieri.
Table 19, — Some measurements of the pinular pentactins of Poliopogon micropentaciinus collected from different
places of the surface.
Source : MNHN, Paris
134
K. R. TABACHNICK & C. LEVI
Table 20. — Some measurements of spicules of Poliopogon micropentactinus.
Table 21. — Some measurements of spicules of Poliopogon claviculus.
L dermal pentactin pinular ray
L dermal pentactin tangential ray
L atrial pentactin pinular ray
L atrial pentactin tangential ray
L microhexactin ray
L macramphidisc
D macramphidisc umbel
L mesamphidisc
L mesamphidisc umbel
D mesamphidisc umbel
L micramphidisc
L micramphidisc umbel
D micramphidisc umbel
HCL 458
Table 22. — Some measurements of spicules of Poliopogon zonecus ( HCL 459).
Source : MNHN, Paris
AMPHIDISCOPHORA OFT NEW CALEDONIA
135
Table 23. — Some measurements of the spicules of Pheronema pilosum.
Source : MNHN, Paris
136
K. R TABACHNICK & C. LEVI
Table 24. — Some measurements of the spicules of Pheronema semiglobosum.
HCL 48 HCL 220 HCL 49 (p4324)
Source : MNHN, Paris
AMPH1D1SCOPHORA OFF NEW CALEDONIA
137
Table 25. — Some measurements of the spicules of Pheronema conicum.
Source :
138
K. R. TABACHNICK & C. LEVI
Table 26. — Some measurements of the spicules of holotype of Pheronema giganieum.
Table 27. — Some measurements of the spicules of holoype of Pheronema pseudogiganteum
Table 28. — Some measurements of the spicules of Pheronema pseudogiganteum
L dermal ia pinular ray
L dermal ia tangential ray
L atrial ia pinular ray
L atrialia tangential ray
L microhexactin ray
L macramphidisc
L macramphidisc umbel
D macramphidisc umbel
L micramphidisc
L micramphidisc umbel
D micramphidisc umbel
L micruncinate
Source :
AMPHIDISCOPHORA OFF NEW CALEDONIA
139
Table 29. — Some measurements of the spicules of Pheronema pseudogiganteum forma vaubana.
TABLE 30. — Some measurements of the spicules of Pheronema pseudogiganteum forma stellata.
140
K. R. TABACHNICK & C. LEVI
Table 31.
Some measurements of the spicules of Pheronema pseudogiganteum forma variodisca.
Source : MNHN, Paris
1 TS DF.S CAMPAGNES MUSORSTOM. VOLUME 21 — RESULT ATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES
Crustacea Cirripedia Thoracica: Chionelasmatoidea and
Pachylasmatoidea (Balanomorpha) of New Caledonia,
Vanuatu and Wallis and Futuna Islands, with a review
of all currently assigned taxa
Diana S. JONES
Western Australian Museum
Francis St., Perth, Western Australia 6000, Australie
ABSTRACT
Balanomorph barnacles of the superfamilies Chionelasmatoidea and Pachylasmatoidea collected by various French
deep-sea expeditions in the waters of New Caledonia, Vanuatu, and the Wallis and Futuna Islands are discussed. One
sample from the Marianas Islands is also included. Of the 21 species reported herein. 18 are new to science, 2 are
recognised as relictual, and I represents a northward range extension within the waters of the southwestern Pacific
Ocean. In addition 4 new genera and 1 new subfamily are described. An exceptional diversity of species occurs in the
subfamilies Pachylasmatinae and Hexelasmatinae of the family Pachylasmatidae. The number of new pachylasmatines
described represents 46% of the known species and that of the new hexelasmatines 40%, indicating the richness of these
waters. Of the 17 new species described from the waters of New Caledonia, Vanuatu, and the Wallis and Futuna Islands.
14 are considered presently to be endemic to the Vanuatu/New Caledonian region and the remaining 3 occur in a broader
area which includes the Futuna and Wallis Islands region. The richest fauna occurs at the Loyalty Islands (15 species), the
Norfolk Ridge (11 species) and New Caledonia (11 species). The occurrence of 2 relictual species, the chionelasmatine
Chionelasmus darwini and the eolasmalincWaikalasma boucheli , in the waters of the New Caledonian region supports
the hypothesis that the southwestern Pacific is a relictual area.
RESUME
Crustacea Cirripedia Thoracica : Chionelasmatoidea et Pachylasmatoidea (Balanomorpha) de
Nouvelle-Caledonie, du Vanuatu et des Ties Wallis et Futuna, avec un examen de toutes les
especes reconnues actuellement.
Les cirripedes balanomorphes des supcrfamilles Chionelasmatoidea et Pachylasmatoidea, recoltes par diverses
expeditions franfaises en eau profonde, au large de la Nouvelle-Caledonie. du Vanuatu et des iles Wallis et Futuna, sont
Jones, D.. 2000. _ Crustacea Cirripedia Thoracica: Chionelasmatoidea and Pachylasmatoidea (Balanomorpha) of
New Caledonia. Vanuatu and Wallis and Futuna Islands, with a review of all currently assigned taxa. In: A. CROSNIER (ed ),
R6sultats des Campagnes Musorstom, Volume 21. Memoires du Museum national d'Histoire naturelle, 184: 141-283.
Paris ISBN 2-85653-526-7.
142
D. JONES
<5tudi<5s. Une recolte provenant des ties Mariannes est egalement examinee. Des 21 especes comprises dans ce travail,
18 sont nouvelles pour la Science, deux sont considerees comme des especes reliques et une voit son aire de repartition
accrue vers le Nord. En outre, quatre nouveaux genres et une nouvelle sous-fainille sont decrits. Une diversity d'especes
exceptionnelle est rencontrec dans les sous-familles Pachylasmatinae et Hexelasmatinae de la famille Pachylasmatidae.
Les nouvelles pachylasmatines ddcrites representent 46% des espbces connues et celles des hexelasmatines 40%, ceci
montrant bien la richesse des eaux prospectees. Sur les 17 especes decrites des eaux de la Nouvelle-Caledonie. du Vanuatu
et des lies Wallis et Futuna, 14 sont considdrees actuellement comme enddmiques de I'ensemble Nouvelle-Caledonie et
Vanuatu; les trois restantes ont ete trouvees dans une zone plus etendue incluant les ties Wallis et Futuna. Les faunes les
plus riches sc trouvent aux ties Loyauld (15 espdces), sur la ride de Norfolk (11 especes) et en Nouvelle-Caledonie
(11 especes dgalement). La prdsence, dans les eaux de la Nouvelle-Caledonie, de deux "fossiles vivants",
le chionelasmatine Chionelasmus darwini et I’eolasmatine Waikalasma bouclteti, renforce I'hypothese que le Pacifique
sud-ouest est une zone refuge.
INTRODUCTION
Since 1978 numerous specimens of balanomorph barnacles have been collected in the deep waters off Vanuatu
(Musorstom 8, 1994), New Caledonia, the Chesterfield and Loyalty Islands ("Vauban", 1978-79; Biocal, 1985;
MUSORSTOM 4, 1985; LAGON. 1984-89; MUSORSTOM 5, 1986; CHALCAL 2, 1986; Smib 2, 1986; SMIB 3, 1987;
Corail 2, 1988; Musorstom 6, 1989; SMIB 6, 1990; Beryx 2, 1992; Bathus 2, 1993; Smib 8, 1993;
Halipro 2, 1996), the Wallis and Futuna Islands, Combe, Field. Tuscarora and Waterwich Banks (Musorstom 1
1992), the Norfolk Ridge (SMIB 4. 1989; SMIB 5, 1989; BATHUS 3, 1993; Bathus 4, 1994) and the Matthew and
Hunter Islands (Volsmar. 1989).
Informations on the cruises where the samples studied here were collected can be found in Richer de Forges
ei al. (1988) for Corail 2; Richer de Forges (1990) for "Vauban" 1978-79, Biocal, Chalcal 2. Musorstom
4, 5 and 6, Smib 2, 3 and 4; Richer de Forges (1991) for Lagon; Richer de Forges (1993) for Volsmar,
Smib 5 and 6, Beryx 2; Richer de Forges & Menou (1993) for Musorstom 7; Richer de Forges, Faliex
& MENOU (1996) for Musorstom 8; Richer de Forge & Chevillon (1996) for Bathus 2, 3 and 4 Smib 8-
Grandperrin et al. (1997) for Halipro 2.
Examination of these collections has yielded an exceptional diversity of thoracican cirripeds. Of particular
interest are the numerous specimens that are herein assigned to the balanomorph superfamily Pachylasmatoidea.
especially within the pachylasmatine genus Pachylasma and the hexelasmatine genus Hexelasma, both of which
are included in the family Pachylasmatidae. These specimens are reported on herein, together with specimens from
the super! annly Chionelasmatoidea. A separate paper will report on the remaining balanomorph taxa collected by
the above expeditions.
The Indo- Pacific deep-sea benthos was investigated by major expeditions such as those of "Challenger" ( 1 873-
1876), "Investigator" ( 1 884- 1 887), "Valdivia" (1898-1899), "Siboga " ( 1 899- 1 900), "Albatross" (1907-1910) and
C«/ar/,ea" (1950-52). However, none or these expeditions collected Cirripedia in the waters of Vanuatu.
New Caledonia or the Wallis and Futuna Islands. The cirriped fauna of the region is known from the brief report of
FISCHER (1884), who described the shallow water barnacles of New Caledonia. More recently, Buckeridge (1994)
provided a comprehensive account of the deep-sea Verrucomorpha from collections made by several French cruises
in the New Caledonia area and the Wallis and Futuna Islands, Ross and Newman (1995) described a new Hoekiini
from the shallow waters of New Caledonia and Buckeridge ( 1998) described a new chionelasmatine from the
The present paper includes the description of 23 new taxa — 1 new subfamily, 4 new genera and 18 new
species Also included are diagnoses and discussions of the previously reported species included in the
lone asmatoidca and the Pachylasmatoidea. Thus, in toto, this account covers 1 family, 2 genera, 3 species and
2 subspecies of Chionelasmatoidea and, within the Pachylasmatoidea, I family, 5 subfamilies, 13 genera and
. secies. My evaluation of certain species suggests some are transferred into new genera, other are re-assigned
elsewhere. The diagnosis of Tetrapachylasma Foster, 1988 is emended to reflect the intra-specific variation now
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACH YLASMATOIDEA
143
recognised within this genus. Wall structure in the Chionelasmatoidea and the Pachylasmatoidea is discussed in
relation to balanomorph wall organisation. Endemism and species diversity, and the significance of antenniformy,
reproductive state and regression and/or loss of the penis are also discussed.
Informations on the types of the species mentioned in this paper are supplied where possible. When a chapter
"TYPES" is missing, no information was obtained for the species considered.
MATERIALS and Methods. — Specimens were examined with the aid of microscopy and dissection.
Illustrations of the shell and whole animal were made with aid of a camera lucida. Soft parts were cleared, stained
and mounted following the method of JONES (1993a) and mouthparts and appendages were drawn with a camera
lucida. Measurements given are all in millimeters. The terminology follows that of Newman (1987). Jones
(1990) and Yamaguchi & Newman (1990).
Abbreviations. — Institutions are abbreviated as follows:
AM: Australian Museum, Sydney, New South Wales, Australia;
AR: New Zealand Geological Survey Type Numbers, Auckland, New Zealand;
AU: Geology Department, University of Auckland, New Zealand;
BMNH: The Natural History Museum. London, U.K.;
GS: New Zealand Geological Survey, Auckland, New Zealand;
MNHN: Museum national d'Histoire naturelle, Paris, France;
NIWA: National Institute of Water and Atmospheric Research, Wellington. New Zealand;
NMNZ: Museum of New Zealand, Wellington. New Zealand;
NZOI: New Zealand Oceanographic Institute, Wellington, New Zealand;
SOSC: Smithsonian Oceanographic Sorting Center, Washington, U.S.A.;
UMUT: University Museum of the University of Tokyo, Japan;
UPIBM: Invertebrate Museum of the Institute of Biology, College of Science, University of the Philippines;
USNM: National Museum of Natural History, Washington, D.C., U.S.A;
WAM: Western Australian Museum. Perth, Western Australia;
YPM: Peabody Museum of Natural History, Yale University, New Haven, CT., U.S.A.;
ZMC: Zoologisk Museum, Copenhagen, Denmark;
ZMUA: Zoologisch Museum, Universileit van Amsterdam, Holland.
Within the text the following abbreviations are used: c.a. for caudal appendage; C for carina, CL for
carinolateral, CL1 for primary carinolaleral. CL2 for secondary carinolateral, Cl to CVI for cirri I to VI: LD for
lateral dimension: m for meter; mis for miles; R for rostrum. RC for rostro-carinal; RL for rostrolateral. rg for
range; S for scutum; T for tergum; x for average.
In the lists of material examined the capital letters preceding the station numbers refer to the gear used:
BT bottom trawl (large otter trawl for fishes); CC otter trawl (shrimps), CH otter trawl (fishes), CP beam
trawl, DE epibenthic sledge, DW Waren dredge.
The acronyms for the cruises that collected the material reported on herein are as follows:
ANZARE = Australian and New Zealand Antarctic Research Expedition. 1913:
BATHUS = the Greek Latinised name for deep;
BERYX = cruises on the seamounts around New Caledonia studying stocks of the fish Beryx,
BIOCAL = BlOlogic cn Nouvelle CALedonie (a trawling cruise);
CHALCAL = CHALutages en Nouvelle CALedonie (trawling cruises around New Caledonia);
CORAIL = cruises around the Chesterfield Islands which are in the CORAIL (Coral) Sea;
HALIPRO = HALIeutiquc PROfonde (deep fisheries);
LAGON = LAGON dc Nouvelle Caledonie (study of the benthos);
MUSORSTOM = a contraction of MUSeum (national d'Histoire naturelle) and ORSTOM (Office de la
Recherche Scientifique et Technique Outre-Mer);
NAGA = the name of a mythical sea dragon.
144
D. JONES
SMIB = Substances Marine d'Interet Biologique;
TUI - a New Zealand cruise of 1962 on board of the "Tuiw" a ship in the Royal New Zealand Navy. Tui is the
vulgar tongue for a large passerifonn bird, known also as the "Parson Bird".
"Vauban" = name of the ORSTOM research vessel working in New Caledonia from 1976 to 1988;
VOLSMAR = VOLcans Sous MARins (volcanoes under the sea).
Type Species. — All holotypes of the new species are held by the MNHN and. when numbers of specimens
available permit, paratypes by the WAM, the BMNH and the USNM.
Registration NUMBERS. — Due to computerisation of the WAM crustacean collection, starting from 1997,
WAM crustacean registration numbers are preceded by C [e.g. WAM C 23243 for specimens of Chionelasmus
darwini (Pilsbry, 1907) from SMIB 8: stn CP 180], Prior to this date a different numbering system existed (e.g.,
WAM 261-96 for C. darn'ini from Bathus 3: stn CP 812). MNHN registration numbers are prefixed by Ci and
ZMUA registration numbers by Cirr, which refer to the respective cirriped collections. NMNZ registration
numbers are prefixed by Cr and ZMC numbers by Cru, which refer to the respective crustacean collections. AM
numbers prefixed by E refer to dry specimens and prefixed by P to wet (alcohol) specimens.
The samples without any registration number are kept at the Museum national d'Histoire naturelle. Paris.
COMBINED LIST OF STATIONS AND OF SPECIES OBTAINED PER STATION
LAGON. New Caledonia.
Station 444. — 28.02.1985, 18°15’S, 162°59'E, 300-350 m: Pachylasma ovatum sp. nov.
Station 491. — 03.03.1985, 18°56'S, 163°20'E, 450 m: Hexelasma sandaracum sp. nov.
BlOCAL. Loyalty Islands.
Station DW 36. — 29.08.1985, 23°09'S, 167° 1 l'E, 650-680 m: Hexelasma sandaracum sp. nov.
Station DW 44. — 30.08.1985, 22°47.3'S, 167°14.3'E, 440 m: Eutomolasfna maclaughlinae sp. nov.
Station CP 45. — 30.08.1985, 22°47'S, 167°15'E, 430-465 m: Eutomolasma maclaughlinae sp. nov., Hexelasma
globosum sp. nov.
Station DW 66. — 03.09.1985, 24°55’S, 168°22'E, 505-515 m: Chionelasmus darwini (Pilsbry), Hexelasma
persicum sp. nov.
Station CP 67. — 03.09.1985, 24°55'S, 168°22’E, 500-510 m: Chionelasmus darwini (Pilsbry).
Station DW 83. — 06.09.1985, 20°35'S, 166°54'E, 460 m: Eutomolasma orbiculatum sp. nov.
Musorstom 4. New Caledonia.
Station DW 156. 15.09.1985, 18°54.0'S, 163°18.8’E, 525 m: Hexelasma aureolum sp. nov., H. globosum
sp. nov., H. sandaracum sp. nov.
Station CP 157. — 15.09.1985, 18°52.5'S, 163°16.9'E, 575 m: Hexelasma sandaracum sp. nov.
Station DW 159. 15.09.1985, 18°45.9'S, 1 63° 1 5.6'E, 585 m: Hexelasma persicum sp. nov., H. sandaracum
sp. nov.
Station CP 167. 16.09.1985, 18°35.8'S, 163°06.4'E, 575 m: Hexelasma globosum sp. nov., /-/. sandaracum
sp. nov.
Station CP 169. — 17.09.1985, 18°54.3'S, 163°11.2'E, 590 m: Hexelasma sandaracum sp. nov.
Station CP 170. 17.09.1985, 18°57.0'S, 1 63° 1 2.6'E, 460 m: Eutomolasma maclaughlinae sp. nov.,
Hexelasma sandaracum sp. nov.
Station CP 178. — 18.09.1985, 18°56.3'S, 163°12.9'E, 520 m: Hexelasma sandaracum sp. nov.
Station CP 179. 18.09.1985, 18°56.6'S, 163°13.7'E, 475 m: Eutomolasma maclaughlinae sp. nov.,
Hexelasma persicum sp. nov.
Station DW 183. - 18.09.1985, 19°01.8’S, 163°25.8'E, 280 m: Eutomolasma maclaughlinae sp. nov.
Source ; MNHN , Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
145
Station CP 190. — 19.09.1985, 19°06'S, 163°29.50'E, 215 m: Pachylasma laeviscutum sp. nov.
Station CP 193. — 19.09.1985, 18°56.3'S, 163°23.2'E, 415 m: Pachylasma ovatum sp. nov.
Station CP 194. — 19.09.1985, 18°52.8'S, 163°21.7'E, 545 m: Hexelasma sandaracum sp. nov.
Station CP 197. — 20.09.1985, 18°51.3'S, 163°2I.0'E, 550 m: Hexelasma globosum sp. nov., H. persicum
sp. nov., H. sandaracum sp. nov.
Station DW 215. — 28.09.1985, 22°55.7'S, 167°17.0'E, 485-520 m: Eutomolasma maclaughlinae sp. nov.
Station DW 221. — 29.09.1985, 22°58.60'S, 167°36.80'E, 535-560 m: Eutomolasma maclaughlinae sp. nov.
Station DW 222. — 30.09.1985, 22°57.6'S, 167°33.0'E, 410-440 m: Eutomolasma maclaughlinae sp. nov.,
Hexelasma persicum sp. nov.
Station DW 223. — 30.09.1985, 22°57.0'S, 167°30.0'E, 545-560 m: Eutomolasma maclaughlinae sp. nov..
Hexelasma persicum sp. nov.
Station DW 229. — 30.09.1985, 22°51.5'S, 167°13.5'E, 445-460 m: Eutomolasma maclaughlinae sp. nov.
Station DW 230. — 30.09.1985, 22°52.5'S, 167°11.8'E, 390-420 m: Eutomolasma maclaughlinae sp. nov..
Pachylasma laeviscutum sp. nov.
Station CP 238. — 02.10.1985, 22°13.0'S, 167°14.0'E, 500-510 m: Chionelasmus darwini (Pilsbry). Hexelasma
aureolum sp. nov.
SMIB 2. Norfolk Ridge.
Station DW 1. — 17.09.1986, 22°53'S, 167°13'E, 438-444 m: Eutomolasma maclaughlinae sp. nov.
Station DW 2. — 17.09.1986, 22°55'S, 167°14'E, 438-448 m: Eutomolasma maclaughlinae sp. nov.
Station DW 12. — 18.09.1986, 22°53'S, 167°14'E, 445-460 m: Eutomolasma maclaughlinae sp. nov.
MUSORSTOM 5. Chesterfield Islands.
Station DW 255. — 07.10.1986, 25° 15.40'S, 159°54.80’E, 280-295 m: Tetrapachylasma arcuatum sp. nov.
Station DW 301. — 12.10.1986, 22°06.90'S, 159°24.60'E, 487-610 m: Tetrapachylasma arcuatum sp. nov.
Station DW 304. — 12.10.1986, 22°10.34'S, 159°25.51'E, 385-420 m: Tetrapachylasma arcuatum sp. nov.
Station DW 338. — 15.10.1986, 19°51.6'S, 158°40.40'E. 540-580 m: Hexelasma aureolum sp. nov.,
H. persicum sp. nov.
Station DC 388. — 22.10.1986, 20°45.35'S, 160°53.69'E, 500-510 m: Eutomolasma maclaughlinae sp. nov.
Station CP 389. — 22.10.1986, 20°44.95'S, 160°53.67'E, 500 m: Hexelasma aureolum sp. nov.
Chalcal 2. Norfolk Ridge.
Station CC 1. — 28.10.1986, 24°54.96'S, 1 68°2 1 .9 1 'E, 500 m: Chionelasmus darwini (Pilsbry).
Station CC 2. — 28.10.1986, 24°55.48'S, 168°21.29'E, 500 m: Chionelasmus darwini (Pilsbry).
Station CH 4, — 27.10.1986, 24°44.31'S, 168°09.32'E, 253 m: Tetrapachylasma arcuatum sp. nov.
Station CH 9. — 31.10.1986, 23°15.64'S, 168°03.06'E, 300 m: Hexelasma persicum sp. nov.
Station CP 21. — 28.10.1986, 24°54.00'S, 1 68°2 1.61 'E, 500 m: Chionelasmus darwini (Pilsbry).
Station DW 72. — 28.10.1986, 24°54.5’S, 168°22.3'E, 527 m: Hexelasma aureolum sp. nov.
Station DW 74. — 29.10.1986, 24°40.36'S, I68°38.39'E, 650 m: Chionelasmus darwini (Pilsbry).
Station DW 75. — 29.10.1986, 24°39.31'S, 168°39.67'E, 600 m: Chionelasmus darwini (Pilsbry).
Station DW 76. — 30.10.1986, 23°40.50'S, 167°45.20'E, 470 m: Chionelasmus darwini (Pilsbry). Hexelasma
aureolum sp. nov.
Station DW 77. — 30.10.1986, 23°38.35'S, 167°42.68'E, 435 m: Chionelasmus darwini (Pilsbry).
Smib 3. Norfolk Ridge.
Station DW 5. — 21.05.1987, 24°55'S, 168°22'E, 502-512 m: Hexelasma aureolum sp. nov.
Station DW 21. — 24.05.1987, 22°59'S, 167°19'E, 525 m: Eutomolasma maclaughlinae sp. nov.
Station DW 22. — 24.05.1987, 23°03'S, 167°19'E, 503 m: Eutomolasma maclaughlinae sp. nov.
Station DW 24. — 24.05.1987, 22°59'S, 167°19'E, 525 m: Chionelasmus darwini (Pilsbry), Eutomolasma
maclaughlinae sp. nov.
146
D. JONES
Station DW 25. — 24.05.1987, 22°56’S. 167°16'E, 437 m: Eutomolasmci maclaughlinae sp. nov.
Station DW 27. — 24.05.1987, 22°55'S, 167°16'E, 457 m: Eutomolasma maclaughlinae sp. nov.
Corail 2. Chesterfield Islands.
Station DE 16. — 21.07.1988, 20°47.75'S, 160°55.87'E, 500 m: Hexelasma aureolum sp. nov.
Station CP 17. — 21.09.1988, 20°48.14’S, 160°57.14'E, 500 in: Hexelasma aureolum sp. nov.
MUSORSTOM 6. Loyalty Islands/Loyalty Ridge.
Station DW 400. — 14.02.1989, 20°42.18'S. 167°00.40'E, 270 m: Hexelasma foratum sp. nov.
Station CP 401. — 14.02.1989, 20*42. 15’S, 167°00.35'E, 270 m: Pachylasma bacum sp. nov.. Hexelasma
sandaracum sp. nov.
Station DW 407. — 15.02.1989, 20°40.70'S. 167°06.60'E, 360 m: Tetrapachylasma arcuatum sp. nov.
Station DW 421. — 16.02.1989, 20°26.27'S, 166°40.I7'E, 245 m: Microlasma fragile sp. nov.
Station DW 422. — 16.02.1989, 20°26.20'S, 166*40.3 l'E, 257 m: Hexelasma aureolum sp. nov.
Station DW 424. — 17.02.1989, 20°24.30'S, 166°24.70'E, 599 in: Hexelasma sandaracum sp. nov.
Station DW 451. — 20.02.1989, 20°59.00'S, 167°24.50'E, 330 m: Hexelasma foratum sp. nov.
Station DW 460. — 20.02.1989, 21°0I.70'S, 167°31.45'E, 420 m: Metalasma crassum sp. nov.. Hexelasma
aureolum sp. nov.
Station CP 465. — 21.02.1989, 21°03.55'S, 167°32.55’E, 480 m: Eutomolasma maclaughlinae sp. nov.
Station CP 466. — 21.02.1989, 21°05.25'S, 167°32.2'E, 540 m: Eutomolasma maclaughlinae sp. nov.
Station CP 467. — 21.02.1989, 21°05.13’S, 167*32.1 l'E, 575 m: Eutomolasma maclaughlinae sp. nov.
Station CC 470. — 21.02.1989, 21°04.40'S, 167°33.20'E, 560 m: Hexelasma sandaracum sp. nov.
Station CP 471. — 22.02.1989, 21°08.00'S, 167*54. 10'E, 460 m: Tetrapachylasma arcuatum sp. nov.,
Hexelasma sandaracum sp. nov.
Station DW 472. — 22.02.1989, 21°08.60'S, 167*54.70 E, 300 m: Tetrapachylasma arcuatum sp. nov.,
Hexelasma flavidum sp. nov.
Station DW 476. — 22.02.1989, 21*09. 36'S, 167*56. 40'E, 300 in: Metalasma crassum sp. nov., Hexelasma
aureolum sp. nov.
Station DW 478. — 22.02.1989, 21*08. 96'S, 167*54. 28'E, 400 in: Eurylasma angustum sp. nov., Tetrapachy¬
lasma arcuatum sp. nov., Metalasma crassum sp. nov., Hexelasma aureolum sp. nov., H. persicum sp. nov.
Station DW 479. — 22.02.1989, 21*09.13’S, 167°54.95'E, 310 in: Metalasma crassum sp. nov.
Station DW 483. — 23.02.1989, 2 1*1 9.80'S, 1 67*47. 80'E, 600 in: Hexelasma sandaracum sp. nov.
Station DW 486. — 23.02.1989, 20*21. 40'S, 167°47.65'E, 370 in: Hexelasma globosum sp. nov.
SMIB 4. Norfolk Ridge.
Station DW 44. — 08.03.1989, 24*46. 0'S, 168*08. 2'E, 300 m: Hexelasma sandaracum sp. nov.
Station DW 55. 09.03.1989, 23*21. 4'S, 168*04. 5'E, 260 m: Chionelasmus darwini (Pilsbry), Eutomolasma
orbiculatum sp. nov.
Station DW 58. — 09.03 1989, 22°59.8'S, 167*24.2'E, 560 m: Eutomolasma maclaughlinae sp. nov.
Station DW 60. — 10.03.1989, 22*00. l'S, 167*21. 6'E, 535 m: Hexelasma sandaracum sp. nov.
Station DW 61. 10.03.1989, 22*59. 9'S, 167*22. 8'E, 550 m: Eutomolasma maclaughlinae sp. nov.. Hexelasma
aureolum sp. nov.
Station DW 62. — 10.03.1989, 23*00.4'S, I67°21.8'E, 540 m: Hexelasma sandaracum sp. nov.
Station DW 63. — 10.03.1989, 22°58.7’S, 167*21. l'E, 520 m: Hexelasma sandaracum sp. nov.
Station DW 64. 10.03.1989, 22*55. 3'S, 167*16. 4'E, 460 m: Eutomolasma maclaughlinae sp. nov., Hexelasma
sandaracum sp. nov.
Station DW 65. 10.03.1989, 22*55. 3'S, 167*14. 5'E, 420 m: Eutomolasma maclaughlinae sp. nov.
Station DW 68. - 10.03.1989, 22*55.0’S, 167°16.0'E, 440 m: Eutomolasma maclaughlinae sp. nov.,
E. orbiculatum sp. nov.
Source : MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
147
VOLSMAR. New Hebrides Arc.
Station DW 16. — 03.06.1989, 22°25.1'S, 171°40.7'E, 500 m: Tetrapachylasma arcuatum sp. nov., Metalasma
crassum sp. nov., Hexelasma foratum sp. nov.
SMIB 5. Norfolk Ridge.
Station DW 71. — 07.09.1989, 23°41.3’S, 168°00.7'E, 265 m: Eurylasma pyramidale sp. nov.
Station DW 75. — 07.09.1989, 23°40.9'S, 168°00.8’E, 270 m: Eurylasma pyramidale sp. nov.
Station DW 85. — 13.09.1989, 22°20.0'S, 163°42.9’E, 260 m: Pachylasma laeviscutum sp. nov.
Station DW 87. — 13.09.1989, 22°18.7'S, 168°41.3'E, 370 m: Tetrapachylasma arcuatum sp. nov.. Hexelasma
flavidum sp. nov.
Station DW 88. — 13.09.1989, 22°18.6'S, 168°40.2’E, 350 m: Pachylasma laeviscutum sp. nov.
Station DW 93. — 13.09.1989, 22°20.0'S, 168°42.3'E, 255 m: Eurylasma pyramidale sp. nov., Tetrapachylasma
arcuatum sp. nov., Hexelasma flavidum sp. nov.
Station DW 94. — 13.09.1989, 22°19.6'S, 168°42.8'E, 275 m: Eurylasma pyramidale sp. nov., Hexelasma
persicum sp. nov.
Station DW 101. — 14.09.1989, 23°21.2'S, 168°04.9'E, 270 m: Eurylasma pyramidale sp. nov,
BERYX 2. New Caledonia.
Station DW 34. — 19.10.1992, 23°33'S, 1 67° 1 7'E, 560-570 m: Hexelasma sandaracum sp. nov.
Station DW 35. — 19.10.1992, 23°33'S, 1 67° 1 6'E, 550-570 m: Hexelasma sandaracum sp. nov.
Station DW 38. — 19.10.1992, 23°38'S, 167°39'E, 550-690 m: Hexelasma sandaracum sp. nov.
Station CP 46. — 20.10.1992, 23°42’S, 168°01'E, 300-350 m: Hexelasma persicum sp. nov.
MUSORSTOM 7. Wallis and Futuna Islands, Combe, Field, Tuscarora and Waterwich Banks.
Station CP 505. — 1 1.05.1992, 1 4° 1 9’S, 178°04’W, 245-400 m: Eutomolasma maclaughlinae sp. nov.
Station CP 508. — 1 1.05.1992, 14°20’S, 178°06'W, 245-440 m: Pachylasma laeviscutum sp. nov.
Station CP 515. — 12.05.1992, 14°13'S, 178°10'W, 224-252 m: Pachylasma laeviscutum sp. nov.
Station DW 522. — 13.05.1992, 1 3° 1 1 'S, 176°15'W, 650-765 m: Hexelasma sandaracum sp. nov.
Station DW 537. — 16.05.1992, 12°30’S, 176041'W, 325-400 m: Eutomolasma maclaughlinae sp. nov.
Station DW 539. — 17.05.1992, 12°27'S, 177°27’W, 700 m: Hexelasma sandaracum sp. nov.
Station DW 547. — 17.05.1992, 14°20'S, 178°06'W, 455 m: Eutomolasma maclaughlinae sp. nov.
Station DW 559. — 19.05.1992, i 1°48'S, 178°19'W, 547-552 m: Hexelasma sandaracum sp. nov.
Station DW 582. — 22.05.1992, 1 3° 1 0'S, 176°14'W, 360 m: Hexelasma sandaracum sp. nov.
Station DW 586. — 22.05. 1992, 13°1 l'S, 1 78° 1 l'W, 510-600 m: Hexelasma sandaracum sp. nov.
Station CP 622. — 28.05.1992, 12°34'S, 178°11'W, 1280-1300 m: Hexelasma persicum sp. nov.
SMIB 8. New Caledonia.
Station DW 150. — 27.01.1993, 24°54.3'S, 168°22.2'E, 519-530 m: Hexelasma aureolum sp. nov.
Station DW 167. — 29.01.1993, 23°38.1'S, 168°43.1'E, 430-452 m: Eutomolasma maclaughlinae sp. nov.
Station CP 180. — 30.01.1993, 23°47.7’S, 1 68° 18.1 'E, 425-460 m: Chionelasmus darwini (Pilsbry). Hexelasma
persicum sp. nov.
Station DW 185. — 31.01.1993, 23°16'S, I68°04.3'E, 305-355 m: Hexelasma persicum sp. nov.
Station DW 189. — 31.01.1993, 23°17.6'S, 168°05.5'E, 400-402 m: Hexelasma sandaracum sp. nov.
Station DW 193. — 01.02.1993, 22°58.7'S. 168°20.1'E, 500-508 m: Hexelasma persicum sp. nov.
Station DW 194. — 01.02.1993, 22°59.6'S, 168°22.5'E, 491 m: Hexelasma persicum sp. nov.
Station DW 197. — 01.02.1993, 22°51.3'S, 168°12.5'E, 414-436 m: Eutomolasma maclaughlinae sp. nov.
Station DW 198. — 01.02.1993, 22°51.6'S, 167°12.4'E, 414-430 m: Eutomolasma maclaughlinae sp. nov.
Station DW 199. — 01.02.1993, 22°51.6'S, 168°12.22'E, 408-410 m: Eutomolasma maclaughlinae sp. nov.
148
D. JONES
BATHUS 2. New Caledonia.
Station DW 718. 1 1.05.1993, 22°46.70'S, 167°14.45'E, 430-436 m: Eutomolasma maclaughlinae sp. nov.
Station DW 719. — 1 1.05.1993, 22°47.57'S, 167°14.58'E, 444-455 m: Eutomolasma maclaughlinae sp. nov.
Station DW 720. — 1 1.05.1993, 22°51.62'S, 167°16.40'E, 530-541 m: Eutomolasma maclaughlinae sp. nov.
Station DW 729. — 12.05.1993, 22°52.42'S, 167°11.90'E, 400 m: Eutomolasma maclaughlinae sp. nov
Hexelasma sandaracum sp. nov.
Station DW 738. — 13.05.1993, 23°02.09'S, 166°56.61'E, 588-647 m: Eutomolasma maclaughlinae sp. nov.
Bathus 3. Norfolk Ridge, Loyalty Ridge.
Station DW 778. — 24.1 1.1993, 24°43'S, 170°07'E, 750-760 m: Waikalasma boucheti Buckeridge.
Station CP 805. — 27.11.1993, 23°41'S, 168°OI'E, 278-310 m: Eurylasma pyramidale sp. nov.''
Station DW 809. — 27.1 1.1993, 23°39'S, I67°59'E, 650-730 m: Hexelasma aureolum sp. nov.
Station DW 811. 28.1 1.1993, 23°41'S, 1 68° 1 5'E, 383-408 m: Hexelasma sandaracum sp. nov.
Station CP 812. — 28.1 1.1993, 23°43.38’S, 168°15.98'E, 391-440 m: Chionelasmus darwini (Pilsbry)
Station CP 815. - 28.1 1.1993, 23°47'S, 168°16'E, 460-470 m: Chionelasmus darwini (Pilsbry), Eurylasma
ferulum sp. nov.
Station DW 818. — 28.1 1.1993, 23°44'S, 168°16'E, 394-401 m: Hexelasma persicum sp. nov.
Station DW 819. — 28.1 1.1993, 23°45'S, 168°16'E, 478-486 m: Hexelasma aureolum sp. nov.
Station CH 820. — 28.1 1.1993, 23°43’S, 168°16'E, 405-41 1 m: Hexelasma persicum sp. nov
Station DW' 829. - 29.1 1.1993, 23°21'S, 168°02'E, 386-390 m: Eutomolasma maclaughlinae sp. nov.
Stat.on DW 830. - 29.11.1993, 23°20'S, 168o01'E, 361-365 m: Hexelasma persicum sp. nov.. Eurylasma
pyramidale sp. nov.
Station CP 833. 30.1 1.1993, 23°03'S, 168°58'E, 441-444 m: Eutomolasma maclaughlinae sp. nov.
Bathus 4. New Caledonia, Norfolk Ridge.
Station CP 909. - 14.08.1994, 18°57.64'S, 163°10.30'E, 516-558 m: Eutomolasma maclaughlinae sp nov
Station CP 910. - 05.08.1994, 16°69.32'S, I63°08.47’E, 560-608 m: Hexelasma flavidum sp. nov
H. sandaracum sp. nov.
Station CP 91 1. - 05.08.1994, 18°58'S, 163°09'E, 558-566 m: Hexelasma sandaracum sp. nov.
Station DW 914.-05.08.1994, 18°48.79'S, 163°15.23'E, 600-616 m: Pachylasma ova, urn sp nov
Stat.on DW 916. - 05.08.1994. 18°53.30'S, 163°19.55’E, 600-616 m: Hexelasma sandaracum sp. nov.
Station DW 923. - 06.08.1994, 1 8°5 1 .5 1 'S, 163°24.17'E, 470-502 m: Hexelasma sandaracum sp nov
Stat.on DW 924. - 07.08.1994, 18°54.85'S. 163°24.34'E, 344-360 m: Pachylasma ova, urn sp nov
Station DW 927. - 07.08.1994, 18°55.48'S, 163°22.1 l’E, 452-444 m: Hexelasma sandaracum sp. nov.
Stat.on DW 928. - 07.08.1994, 18°55’S, I63°24’E, 420-452 m: Hexelasma sandaracum sp. nov
Station DW 929. - 07.08.1994, 18°51.55'S, 163°23.27'E, 502-516 m: Hexelasma persicum sp. nov.
H. sandaracum sp. nov.
Station DW 932. — 08.08. 1994, 19°07.91'S
Station CP 951. — 10.08.1994, 20°31.44'S,
, 163°29.38'E, 170-190 m: Hexelasma sandaracum sp. nov.
I64°54.97'E, 960 m: Hexelasma sandaracum sp. nov.
MUSORSTOM 8. Vanuatu.
Station CP 962. - 21.09.1994, 20°19.70'S, 169M9.02'E, 370-400 m: Pachylasma laeviscutum sp. nov
Station CP 963. - 21.09.1994, 20°20.10'S, 169°49.08’E, 400-440 m:' Pachylasma laeviscutum sp. nov
/ etrapachylasma arcuatum sp. nov.
si3!1011 S™7' ~ 2I 09-1994- 20°19.45'S, 169°52.87'E, 295-334 m: Eurylasma angustum sp. nov.
t '0n rl III ~ 22-091994’ 19°21-94'S, 169°28.07'E, 487-507 m: Hexelasma sandaracum sp. nov.
3 !°n „„ Q ' ()91994’ 19°21-30S, 169°27.03’E, 460-480 m: Hexelasma sandaracum sp. nov.
I ,0n Zt l™ '"4’ I9°21-5I'S’ 169°28-26'E- 492-520 m: Hexelasma sandaracum sp. nov.
Slat, on DW 978. - 22.09.1994. 19°22.62'S, 169°27.l l'E, 413-408 m: Hexelasma sandaracum sp. nov.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
149
Station DW 988. — 23.09.1994, 19°16.04'S, 169°24. 12'E, 372-466 m: Pachylasma laeviscutum sp. nov.
Station DW 989. — 23.09.1994, 19°13.92’S, 169°20.25'E, 650-669 m: Hexelasma sandaracunt sp. nov.
Station CP 1026. — 28.09.1994, 17°50.35'S, 168°39.33'E, 437-504 m: Hexelasma sandaracum sp. nov.
Station CP 1027. — 28.09.1994, 17°53.05'S, 168°39.35'E, 550-571 m: Hexelasma sandaracum sp. nov.
Station CP 1045. — 30.09.1994, 15°57.63'S. 166°38.43'E, 459-488 m: Hexelasma sandaracum sp. nov.
Station DW 1046. — 30.09.1994, I6°53.62'S, 168° 1 1.08'E, 461-480 m: Hexelasma sandaracum sp. nov.
Station DW 1060. — 02.10.1994, 1 6° 1 3.82'S, 167°20.80'E, 375-397 m: Eurylasma angustum sp. nov.
Station DW 1062. — 02.10.1994, 1 6° 1 4.82'S. 167°18.64'E, 619-658 m: Bathylasma alearum sp. nov.
Station DW 1067. — 02.10. 1994, 1 6° 1 6.78'S, 167°21 ,52'E, 344-366 m: Hexelasma sandaracum sp. nov.
Station CP 1080. — 05.09.1994, 15°57.30'S, 167°27.73'E, 799-850 m: Hexelasma sandaracum sp. nov.
Station CP 1083. — 05.10.1994, 15°5I.9FS, 167°19.42'E, 397-439 m: Hexelasma sandaracum sp. nov.
Station CP 1086. — 05.10.1994, I5°36.58’S, 167°16.32'E, 182-215 m: Pachylasma bacum sp. nov.
Station CP 1 1 14. — 08.10.1994. 14°52.39'S, 167°03.40’E, 647 m: Hexelasma sandaracum sp. nov.
Station CP 1125. — 10.10.1994, I5°57.63'S, 166°38.43'E, 1 160-1220 m: Hexelasma sandaracum sp. nov
Station CP 1 127. — 10.10.1994, 15°58.86'S, 166°37.82'E, 1052-1058 m: Hexelasma sandaracum sp. nov
Station CP 1 131. — 1 1.10.1994, 15°38.41'S, 167°03.52'E, 140-175 m: Pachylasma laeviscutum sp. nov.
Station CP 1 136. — 1 1.10.1994, I5°40.62'S, 167°01.60'E, 398-400 m: Hexelasma sandaracum sp. nov.
HAL1PRO 2. New Caledonia.
Station BT 49. — 16.1 1.1996, 25°23.66'S, 168°19.62’E, 605-628 m: Chionelasmus darwini (Pilsbry).
Station BT 70. — 20.1 1.1996, 24°42.96'S, I68°08.I7'E, 226-238 m: Tetrapachylasma arcuatum sp. nov.
Station BT 71. — 20.1 1.1996, 24°50.12'S, I68°09.97'E, 820-1220 m: Tetrapachylasma arcuatum sp. nov.
LIST OF SPECIES DESCRIBED OR DISCUSSED HEREIN
t = fossil species
CHIONELASMATOIDEA
Family CHIONELASMATIDAE
Genus EOCHIONELASMUS
Eochionelasmus ohtai Yamaguchi, 1990
Genus CHIONELASMUS
Chionelasmus crosnieri Buckeridge, 1998
Chionelasmus darwini (Pilsbry, 1907)
Chionelasmus dam-ini darwini (Pilsbry, 1907)
Chionelasmus darwini cantelli Yamaguchi, 1998
PACHYLASMATOIDEA
Family PACHYLASMATIDAE
Subfamily EOLASMATINAE
Genus EOLASMA
t Eolasma maxwell i Buckeridge, 1983
t Eolasma rugosa Buckeridge, 1985
Genus WAIKALASMA
Waikalasma boucheti Buckeridge, 1996
t Waikalasma juneae Buckeridge, 1983
Subfamily PACHYLASMATINAE
Genus EUTOMOLASMA gen. nov.
Eutomolasma cltinense (Pilsbry, 1912)
Eutomolasma japonicum (Hiro, 1933)
Eutomolasma maclaughlinae sp. nov.
Eutomolasma orbiculatum sp. nov.
Genus MICROLASMA gen. nov.
Microlasma arwetergum (Rosell, 1991)
Microlasma crinoidophilum (Pilsbry. 1911)
Microlasma fragile sp nov.
Microlasma ochriderma (Foster, 1981)
Genus PACHYLASMA
Pachylasma bacum sp. nov.
Pachylasma darwinianum (Pilsbry, 1912)
Pachylasma ecaiulatum Hiro, 1939
Pachylasma giganteum (Philippi. 1836)
Pachylasma integrirostrum Broch. 1931
Pachylasma laeviscutum sp. nov.
Pachylasma ovatum sp. nov.
Pachylasma scutistriata Broch. 1922
Source : MNHN, Paris
150
D. JONES
Genus EURYLASMA gen. nov.
Eurylasma angustum sp. nov.
Eurylasma ferulum sp. nov.
Eurylasma pyramidale sp. nov.
Genus TETRAPACHYLASMA
Tetrapachylasma arcuatum sp. nov.
Tetrapachylasma aurantiacum (Darwin. 1854)
Tetrapachylasma ferrugomaculosa (Jones, 1993)
Tetrapachylasma ornatum sp. nov.
Tetrapachylasma trigortum Foster, 1988
Subfamily METALASMATINAE subfam. nov.
Genus METALASMA gen. nov.
Metalasma crassum sp. nov.
Subfamily BATHYLASMATINAE
Genus BATHYLASMA
Bathylasma alearum (Foster, 1978)
t Bathylasma aucklandicum (Hector, 1888)
Bathylasma coroll forme (Hock, 1883)
Bathylasma hirsutum (Hoek, 1883)
Genus MESOLASMA
Mesolasma fosteri (Newman & Ross, 1971)
Genus TETRACHAELASMA
Tctracliaelasma southwardi Newman & Ross. 197 1
Genus TESSARELASMA
t Tessarelasma pilsbryi Withers, 1936
Subfamily HEXELASMATINAE
Genus HEXELASMA
Hexelasma americanum Pilsbry, 1916
Hexelasma arafurae Hoek. 1913
Hexelasma aureolum sp. nov.
Hexelasma brintoni (Newman & Ross, 1971 )
Hexelasma callistoderma Pilsbry. 1911
Hexelasma flavidum sp. nov.
Hexelasma foratum sp. nov.
Hexelasma globosum sp. nov.
Hexelasma gracilis Foster, 198 1
Hexelasma leptoderma Newman & Ross, 1971
Hexelasma nolearia (Foster, 1978)
Hexelasma persicum sp. nov.
Hexelasma sandaracum sp. nov.
Hexelasma triderma Newman & Ross, 1971
Hexelasma velutinum Hoek, 1913
WALL ORGANISATION IN THE BALANOMORPHA
In the systematics of balanomorph barnacles the number of wall plates and the manner in which they articulate
or coalesce to one another are of major importance. In early balanomorphs the wall was surrounded by one or more
whorls of smaller imbricating plates that protect the suture between the base of the primary wall and the
substratum, and the lower portion of the sutures between the wall plates (Buckeridge & Newman 1992-
Newman, 1993). In most Tertiary forms the imbricating plates are lost and significant concomitant changes
appear in the structure of the compartments themselves (e.g., the development of alae and radii), the number of
wall plates and in the way these articulate or coalesce with one another.
The phylogeny of the balanomorphs indicates that various plates have been added to the wall, with 8 being the
maximum complement in a few surviving early forms. This is repeated in their ontogeny to a remarkable extent.
However the general pattern is one of reduction, from 8-plated forms to 6-plated forms, of which some derived
forms independently went to 4 plates.
Although the 4 plates seen in derived forms have evolved through reduction by elimination or coalescence
current concepts on the evolution, ontogeny and homologies of the wall plates in balanomorphs indicate that
4 plates are the basic number (Ross & Newman, 1996). Since Darwin (1854) the primitive 8-plated wall in
balanomorphs such as Catophragmus s. I., Pachylasma, Octomeris and Chelonibia has been considered to consist
ot K-KL-L-CL-C with CL overlapping C and underlapped by L. The 6-plated balanomorph wall (R-L-CL-C) was
^ fr°m 8 P'ateS’ dUC 10 ,hC develoPmenl ^ a bipartite or compound rostrum
' , K f^ ^ar y 'n the ontogen>' of aM balanomorph barnacles, however, the wall consists of only 4 plates,
, , and C (Darwin, 1854; Glenner & H0eg, 1993). Yamaguchi & Newman (1990) consider this 4-plated
wa I to have a so been the case m balanomorph phylogeny and these and subsequent authors have provided evidence
. YeAt^ r-l"; tUS” (L>iS aCtUal,y ,hC CL’ as L Was incorP°rated into the operculum and subsequently-
lost (Yamaguchi & Newman, 1990; Buckeridge & Newman, 1992; Newman, 1993). Therefore, the basic
balanomorph wall should be designated R-CL-C (rather than R-L-C) with CL overlapping C and R.
Source : MNHN. Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
151
Catophragmus - Octomeris
Tetrapachylasma
Pachylasma
C = carina
CL = carinolatus (= CLi)
CL2 = carinolatus 2
R = rostrum
RL = rostrolatus
+ = fused plate
-- = suture line
urbalanomorph
Fig. l. — Schematic representation of wall organization in selected genera of lower Balanomorpha. viewed from right
side (basal whorls of imbricating plates, where present, not shown). These reflect grades of wall construction rather
than a strict phylogeny, but the polarity indicated by arrows reflects the general evolution of the balanomorph wall.
Proceeding from the bottom, the urbalanomorph wall of four plates in which CL overlaps R and C (imbricating plates
were present, radii absent). Chionelasmus, in which RL, derived from an imbricating plate, has been added to protect
the R/L suture, upper portion of RL not incorporated into sheath and lower portion not fully incorporated into wall
(imbricating plates present). Waikalasma, in which CL2 has been added by replication of CL1 rather than by transfer!
from the imbricating whorls; CL2 but not RL enter the sheath. Pachylasma, in which RL and CL2 are incorporated into
wall but RL is not yet incorporated into sheath (imbricating plates absent in Recent forms). Tetrapachylasma, in
which RL fuses with R to form compound rostrum (RL+R+RL), and CL2 fuses with CL1. Catophragmus-Octomeris
(imbricating plates lost in latter), in which RL is incorporated into sheath. [From Ross & Newman. 1996, fig. 2
(part), reproduced with the permission of Dr Arthur HUMES, Editor. Journal of Crustacean Biology],
152
D. JONES
Ontogenetic studies have also shown the development of a second pair of latera, which are known to develop
3-4 days after metamorphosis in balanoids (Darwin, 1 854; Costlow, 1965). These arc a replicate of the first pair
(CL1) and, therefore, are designated CL2 (Yamaguchi & Newman, 1990). It thus follows that the generalised
8-plated wall of balanomorphs is R-RL-CU-CL2-C and in 6-pIated forms with a compound rostrum the wall is R-
CLi-CL2-C (Ross & Newman, 1996).
SYSTEMATIC ACCOUNT
Superorder THORACICA Darwin, 1854
Order SESSILIA Lamarck, 1818
Suborder BALANOMORPH A Pilsbry, 1916
Superfamily CHIONELASMATOIDEA Buckeridge, 1983
diagnosis. — Symmetrical sessile barnacles having primary shell wall composed of 6 distinct compartmental
plates, including R, paired RL and CL, C (R-RL-CL-C) in contact with substratum, surrounded by single whorl
or several distinctly separate whorls, of basal imbricating plates; sheath formed by R. C and CL; basis thinlv
calcareous or membranous. J
Remarks. — See Remarks under Pachylasmatoidea.
Distribution. — Upper Eocene to Recent (Buckeridge, 1983).
Family CHIONELASMATIDAE Buckeridge, 1983
Diagnosis. — As for the superfamily.
Remarks. — The family comprises 2 genera, Chionelasmus and Eochionelasmus.
Distribution. — Upper Eocene to Recent.
Genus EOCHIONELASMUS Yamaguchi, 1990
Eochionelasmus Yamaguchi, 1990 (in Yamaguchi & Newman, 1990: 136).
Type Species. — Eochionelasmus ohtai Yamaguchi, 1990.
0»uSAapNpe°n<lageTpr™„7 S“Ir°“"dKl ^ S<!,i!raI diS,i"C*' WhOT'S imbricating plgj;
UMUT RA-18634 “ usnm 22i862;
Holotype depository: UMUT.
Paratypes depository: UMUT, USNM, BMNH, MNHN.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
153
REMARKS. — Until recently only 1 species, E. olitai, was attributed to this genus. In 1997 YAMAGUCHI and
Newman (1997a) described E. paquensis from the East Pacific Rise. In a subsequent paper, Yamaguchi and
Newman (1997b) proposed E. ohtai mqnusensis for the Manus Basin population, distinct from populations of
E. ohtai ohtai in the North Fiji and Lau Basins, based on small but distinct differences in the ontogenetic-
development of the basal imbricating plates.
Distribution. — Recent, from hydrothermal vents. South-west Pacific; North Fiji Basin; Lau Basin; Manus
Basin, 1990-2500 m; East Pacific Rise, N of Easter Island, 2578 m.
Genus CHIONELASMUS Pilsbry, 1911
Cat oph rag in us (Chi on el asm us ) Pilsbry. 1911: 82.
Chionelasmus - Nilsson-Cantf.LL. 1928: 445.
Type Species. — Chionelasmus darwini (Pilsbry, 1907).
Diagnosis. — Primary wall surrounded by basal, trimorphic, imbricating plates condensed into a single
whorl; caudal appendages present.
Remarks. — Until recently, only 1 species was attributed to this genus. However, during the final stages of
preparation of the present manuscript, 2 papers were published that described a new subspecies and a new species,
respectively. YAMAGUCHI (1998) proposed Chionelasmus darwini cantelli for the Indian Ocean population, distinct
from the Hawaiian population of Chionelasmus darwini darwini, based on the morphology of the opercular plates
and the ontogenetic development of the basal imbricating plates. In the same year, BUCKERIDGE ( 1998) described
Chionelasmus crosnieri, a facultative coral-dwelling species from a guyot on the northern part of the Norfolk
Ridge. In the light of these recent publications, re-examination of the material included below under Chionelasmus
darwini may be warranted.
Distribution. — Eocene, Tonga; Upper Eocene. New Zealand; Recent, western Indian Ocean to northern
Pacific; 260-650 m.
Chionelasmus darwini (Pilsbry, 1907)
Figs 2-3, 4 a
Catophragmus darwini Pilsbry, 1907: 188, fig. 4, pi. 5 figs 1-8; 1916: 335.
Catopliragmus ( Chionelasmus ) darwini - PILSBRY, 1911: 82.
Chionelasmus darwini - Nilsson-Cantell, 1928: 445, figs 1-2; 1938: 14. — Pope, 1965: 10. — Gordon, 1970: 9, 105,
— Newman & Ross, 1976: 40, frontispiece. — Stanley & Newman, 1980: 177. — Foster. 1981: 352, fig. 3 —
Buckeridge, 1983: 61, fig. 46. — Hui & Moyse, 1984: 91, fig. 1. — Newman. 1987: 23, figs 4, 5a. 6. —
Yamaguchi & Newman, 1990: 148, fig. 8.
Material EXAMINED. — New Caledonia. Musorstom 4: stn CP 238, 500-510 m: I specimen.
Smib 8: stn CP 180, 425-460 m: several specimens (WAM C 23243).
Halipro 2: stn BT 49, 605-628 m: several specimens attached to gorgonian.
Loyalty Islands. Biocal: stn DW 66, 505-515 m: 1 specimen. — Stn CP 67, 500-510 m: several specimens.
Norfolk Ridge. Chalcal 2: stn CC 1, 500 m: 1 specimen. — Stn CC 2, 500 m: 1 specimen. — Stn CP 21. 500 m:
1 specimen. — Stn DW 74, 650 m: 1 specimen. — Stn DW 75, 600 m: I specimen. — Stn DW 76. 470 m: 4 specimens.
— Stn DW 77, 435 m: 2 specimens.
Smib 3: stn DW 24, 535 m: 1 specimen, attached to sponge.
Smib 4: stn DW 55, 260 m: 1 specimen.
Bathus 3: stn CP 812, 391-440 m: many specimens, attached to coral rubble (WAM 261-96). — Sin CP 815, 460-
470 m: many specimens, attached to coral rubble (MNHN-Ci 2694). Drawn.
154
D. JONES
Types. — Syntypes : (from incomplete specimens) USNM 32407 [hard parts], USNM 3240K [soft parts);
"Albatross" station 3998. near Kauai, Hawaiian Is, 417-430 m.
Syntypes depository: USNM (material is missing).
Diagnosis. — Wall consisting of 6 white, porcellanous plates, without radii. R with alae. S without adductor
ridge; S and T with strongly developed articular ridges. Caudal appendages multi-articulate, 8-12 segments.
2.0 mm
a-d
Fig. 2. — Chionelasmus darwini (Pilsbry, 1907). Specimen from Bathus 3, stn CP 815 (MNHN-Ci 2694): a. S. external
view; b, S, internal view; c, T, external view; d. T. internal view.
Supplementary Description. — Measurements of 10 specimens examined, randomly selected from several
stations, as follows:
Cirrus I shortest, rami unequal. Cirrus II resembling cirrus III more than cirrus I. Cirrus III unmodified, similar
to cirri IV-VI. Chaetotaxy ctenopod. Cirral formula as follows:
Remarks. — This species was collected previously from the western Indian Ocean and from the southwestern
and northern Pacific Ocean. The present specimens from the waters of New Caledonia expand the distribution of
the species in the south-western Pacific Ocean and extend the depth range to between 260 and 650 m. The well
preserved specimens in the present collection agree in general with previous descriptions of C. darwini from
the Hawaiian Islands (Pilsbry, 1907) and Rodriguez Island (Nilsson-Cantell. 1928). However, some variation
exists in the form of the mandible. The material examined herein shows the pectinated inferior angle of
Source : MNHN, Paris
CIRRIPED1A THORACICA: CHIONELASMATOIDEA AND PACHYL ASM ATO I DE A
155
the mandible bifid on the right side, but roundly molariform on the left side. In addition, both mandibles lack the
2 small additional teeth between teeth 1 and 2 which were illustrated by Nilsson-Cantell (1928). These
differences may reflect intra-specific variation.
distribution. — Western Indian Ocean, south-western to northern Pacific Ocean: Rodriguez Island (9.5 mis
N, 4.0 mis W of Port Mathurin), western Indian Ocean, 527 m; Kermadec Islands, south-west Pacific Ocean
(30°34’S, 178°30'W), 501 m; Hawaiian Islands, north Pacific Ocean, 417-430 m. Now also known from New
Caledonia, 425-510 in, the Loyalty Islands, 500-515 m and Norfolk Ridge, 260-650 m.
40°N
20°
0°
20°
40°S
Fig. 3. — Chionelasmus darwini (Pilsbry, 1907). Distribution map (depth 260-650m).
Fig. 4. — a. Chionelasmus darwini (Pilsbry. 1907). Specimen from Bathus 3, stn CP 812 (WAM 261-96): lateral view.
Fig. 4. — b-c, Waikalasma boucheti Buckeridge, 1996. Specimen (incomplete) from Bathus 3, stn DW 778 (MNHN-Ci
2410): opercular plates articulated together; b, external view; c, internal view. Scales = 0.5 cm for a; 1 cm for b-c.
156
D JONES
Superfamily PACHYLASMATOIDEA Utinomi, 1968 (emend.)
Diagnosis. — Wall composed of 8 distinct compartmental plates (R, paired RL. CL1 and CL2, and C).
without (excepting Waikalasma) extra whorls of basal imbricating plates; rostrum compound, RL not entering
sheath; parietes solid; radii, when present, solid; basis commonly membranous, or if calcareous solid, and not
forming complex inter-digitations with wall.
Distribution. — Paleocene to Recent; cosmopolitan (Buckeridge. 1983).
Remarks. — Prior to the work of Newman and Ross (1971) the extant deep-sea balanomorph genera
Pachylasma, Hexelasma, Chionelasmus and Bathybalanus , and the fossil genus Tessarelasma, were included^ in the
Chthamalidae (Darwin, 1854; Pilsbry, 1916; Utinomi, 1968). Utinomi (1968) included 3 subfamilies within
the family Chthamalidae, placing Pachylasma, Hexelasma, Tessarelasma and, curiously, Bathybalanus in the
subfamily Pachylasmatinae and Chionelasmus in the Catophragminae. The third subfamily, the Chthamalinae,
was composed of shallow-water species.
The revisionary work of Newman and Ross (1971) correctly removed Bathybalanus from the Chthamalidae and
placed it in the Balanidae. These workers further proposed the family Bathylasmatidae to include the genera
Hexelasma and the extinct Tessarelasma as well as 3 new deep-sea genera ( Bathylasma , Tetrachaelasma and
Aaptolasma), whilst retaining Pachylasma in the Chthamalidae. In a later revision Newman and Ross (1976)
lurther separated Pachylasma and Chionelasmus from the Bathylasmatidae by placing the Bathylasmatidae in the
superfamily Coronuloidea, and retaining Pachylasma and Chionelasmus in the Chthalamoidea (in the Chthamalidae
and the Catophragmidae, respectively). The Bathylasmatidae was divided into the Bathylasmatinae, which contained
Bathylasma, Tessarelasma and Tetrachaelasma and the Hcxelasmatinae which contained Hexelasma and
Aaptolasma.
Foster (1978) maintained that Pachylasma and Hexelasma were closely related, as had been noted previously
by Pilsbry (1916). Furthermore, basing his conclusions upon shell construction. Foster (1978) proposed that
Chionelasmus should be grouped with Pachylasma, Hexelasma, Aaptolasma and Tetrachaelasma in the family
Pachylasmatidae, as distinct from the Chthamalidae which contained Chthamalus. Catpphragmus, Octomeris.
Tetrachthalamus and Chamaesipho. In a further revision, Foster (1981) synonymised Aaptolasma with Hexelasma
and transferred Bathylasma and Mesolasma to the Pachylasmatidae.
The superfamilial rank Pachylasmatoidea was proposed by Buckeridge (1983) for the families
Pachylasmatidae and retrachtidae, but the latter was later assigned to the Tetraclitoidea (Newman. 1993). Within
die Pachylasmatidae 5 subfamilies were recognised, i.e. Chionelasmatinae, Pachylasmatinae, Eolasmatinae,
Bathylasmatinae and Hexelasmalmae. BUCKERIDGE (1983) noted that both Chionelasmus and Pachylasma had
weakly articulated plates without radii and, in this respect, showed more in common with Hexelasma and
Bathylasma than the Chthamalidae. He also commented that the hexelasmatines and the living bathylasmatines
O.e. Hexelasma, Bathylasma, Tetrachaelasma) had a more specialised body morphology than Chionelasmus and
Pachylasma, as they did not possess caudal appendages. Buckeridge (1983) suggested that Pachylasma
represented an intermediate stage between an 8-plated and extra whorled balanomorph and the 6-plated, single
whorled Hexelasmalmae and, furthermore, that Chionelasmus was a plesiomorphic living balanomorph His
phylogenetic tree (Buckeridge, 1983) showed Pachydiadema as the most primitive balanomorph and, on p 21
he derived Chionelasmus '... perhaps [evolving] directly from Pachydiadema". BUCKERIDGE, however, based his
ideas on the now-considered false notion of Newman (pers. comm.) that Chionelasmus possibly changed from 8-
to 6-plated during ontogeny (as suggested by Newman 1987). Chionelasmus became .he “most primitive living
balanomorph m the works of Yamaguch. and Newman (1990) and Buckeridge and Newman (1992) In 1991)"
desrrL' Tf ^ ™St 'ivinS member °f ,he suboide'' Balanomorpha. Eochionelasmus, was
Serfemilii mnk cV "I FlJI Jasin (Yamaguch. & Newman, 1990). Newman (1987) proposed the
superfamilial rank Chionelasmatoidea to embrace the Chionelasmatidac and Chionelasmus and Eochionelasmus
Yamaguchi and NEWMAN (1990, and Newman and Yamaguch. (1995) proposed a new hypothesis
concerning the evolution of the 8-plated balanomorph wall, based on evidence concerning the organisation of the
Source : MNHN, Paris
CIRRIPED1A THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
157
shell of Eochionelasmus and Neobrachylepas, the ontogeny of the shell wall of Chionelasmus and the ontogeny or
Balanus s.l. This evidence suggested that the true (median) lateral plate is replaced by the CL plate in the
balanomorphs, and that an additional, secondary CL plate (CL2) is added by replication in advanced balanomorphs.
In the light of these findings. BUCKERIDGE and Newman (1992) reinterpreted the structure of the wall of the fossil
eolasmatine Waikalasma juneae, proposing that Waikalasma was amongst the most plesiomorphic of balanomorph
barnacles and not merely an evolutionary offshoot. Subsequently. BUCKERIDGE (1996) described an extant species
of Waikalasma , the first living representative of the eolasmatines.
In the lower balanomorphs (i.e. the Chionelasmatoidea) imbricating plates are present, as cither a basal whorl
( Chionelasmus ) or whorls ( Eochionelasmus ). The wall is 6-plated (R-RL-CL-C), with RL derived from an
imbricating plate which is added to protect the R/CL suture. The upper portion of RL is not incorporated into the
sheath and the lower portion is not fully incorporated into the wall.
In the higher balanomorphs (e.g., the Pachylasmatoidea), CL2 is added to the wall structure. Within the
Pachylasmatidae, two whorls of imbricating plates are present in the eolasmatine Waikalasma. The wall is 8-plated
(R-RL-CLI-CL2-C), with CL2 added by the replication of CL1, rather than by transfer from the imbricating
whorls. CL2 only slightly enters the sheath as a very narrow sliver. In the pachylasmatine Pachylasma,
imbricating plates are absent in Recent forms. Imbricating plates are also absent in the new pachylasmatine genera
described in the present paper ( Eutomolasma gen. nov., Eurylasma gen. nov. and Microlasma gen. nov.). In these
pachylasmatincs the wall is 6-plated (R-CL^CL—C), RL and CL2 are incorporated into the wall but RL is not
incorporated into the sheath. In Eurylasma CL2 is partially coalesced with CL1, but CL2 enters the sheath as a
very narrow zone. The pachylasmatine Tetrapachylasma has a wall that may be 6- or 4-plated, depending on the
degree of coalescence of the plates. In the 6-plated form (R-CL'-CL2-C). RL coalesces with R to form a compound
rostrum (RL+R+RL). In the 4-platcd wall (R-CL-C). RL and R form a compound rostrum (RL+R+RL) and CL2
is united with CL1 to form a compound CL plate. CL2 enters the sheath as a narrow zone but RL is not
incorporated into the sheath (except in T. ornatum sp. nov.).
Imbricating plates are absent in the new subfamily Metalasmatinae described herein, and in the Bathylasmatinae
and the Hexelasmatinae. In the Metalasmatinae, the Hexelasmatinae and the bathylasmatines Bathylasma.
Mesolasma and Tessarelasma, the wall is 6-plated (R-CL'-CL2-C), RL and CL2 are incorporated into the wall but
RL is not incorporated into the sheath. The bathylasmatinae Tetrachaelasma has a 4-plated wall (R-CL-C). due to
the absence of CL2. Thus in the Pachylasmatoidea the RL does not enter the sheath (with the exception of the
tetrapachylasmatid T. omatum, which shows the incipient inclusion of RL into the sheath).
Family PACHYLASMATIDAE Utinomi. 1968 (emend.)
Tables 1-2
Diagnosis. — Wall composed of 8 distinct, calcareous plates (R-RL-CL'-CL2-C); adults externally with 8. 6
or 4 plates differentiated; diametric growth of shell effected by alar growth of R (when distinct), CL1 and CL2, and
C; radii absent; sutural edges not complexly interlocked.
Remarks. — Within the Pachylasmatidae 5 subfamilies were recognised by Buckeridge (1983), i.e.
Chionelasmatinae, Pachylasmatinae, Eolasmatinae, Bathylasmatinae and Hexelasmatinae. Newman (1987) later
proposed the Chionelasmatoidea, containing the Chionelasmatidae. A new pachylasmatid subfamily, the
Metalasmatinae, is defined and described herein. The chionelasmatoids possess caudal appendages and, in the
pachylasmatoids, the metalasmatines and pachylasmatincs, but these structures are absent in the eolasmatines,
bathylasmatines and hexelasmatines.
Adult pachylasmatoids have a variety of shell plate patterns when viewed externally, i.e. 8, 6 or 4 plates.
The 8-plated adult pattern is represented by R, paired RL. paired CL1 and CL2, and C (R-RL-CL1-CL2-C). This
pattern occurs in the eolasmatines. In the 6-plated adult pattern there is a compound rostral plate, paired CL1 and
CL2, and C (R-OJ-CL2-C). The compound rostral plate may have the rostral suture between R and RL visible
158
I). JONES
TABLE 1. — Pachylasmatidae: characters of the subfamilies.
ext. = external; int. = internal; + = present; - = absent or not applicable.
TABLE 2 , — Pachylasmatidae: characters of the subfamilies (continued).
artic. - articular; c.a. - caudal appendage; proj. = projecting; prom. = prominent.
(occasionally obscure) externally and internally; internally a total of 8 plates is differentiated since R and RL still
retain their integrity. Rostral sutures arc absent when the coalescence of R and the paired RL is complete;
internally a total of 6 plates is differentiated. In the 6-plated adult pattern, CL' and CL? may be distinct entities, or
partially coalesced, but in the latter condition CL2 still retains a discrete, reduced ala and the suture line is visible
internally and externally. The 6-plated pattern occurs in the subfamily Metalasmatinae described herein and in the
hexelasmatines, some pachylasmatines ( Eutomolasma gen. nov„ Microlasma gen. nov„ Eurylasma gen. now,
Source : MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
159
Pachylasma) and some bathylasmatines ( Bathylasma , Mesolasma). In the 4-plated adult pattern CL2 may never
develop this condition is seen in some bathylasmatines ( Tetrachaelasma , Tessarelasma).
However, whilst the shell wall pattern is regarded as "fixed" at either 8, 6 or 4 plates for the majority or
pachylasmatoid genera (e.g., 8 for Waikalasma , 6 for Pachylasma, 4 for Tetrachaelasma), one genus
( Tetrapachylasma ) exhibits a "variable" pattern, where adult populations contain individuals with 6 or 4 plates. In
the 6-plated individuals there is a compound rostral plate, and CL1 and CL2 can be separate or show degrees of
coalescence. In the 4-platcd individuals CL1 and CL2 are coalesced, there is a compound rostral plate, and a C.
Distribution. — Upper Eocene to Recent; cosmopolitan. All living species inhabit shallow to deep waters.
Key to the Subfamilies of the Family PACHYLASMATIDAE
Tables 1-2
1. Primary shell wall of adult externally 8-platcd; R and RL plates separate; with or without
extra whorls of basal imbricating plates . EOLASMATINAE
Primary shell wall of adult externally 6- or 4-platcd; R and RL forming a compound rostral
plate; without extra whorls of basal imbricating plates . 2
2. Alar welting absent; caudal appendages present, sometimes reduced or vestigial; mandible
tridentoid . ^
— Alar welting present; caudal appendages absent; mandible quadridentoid . 4
3. Chitinous rods or lamina absent in wall plates; horizontal growth ridges low, lacking
small setae; superior alar margins of C, CL2 and CL1 not thickened distally . .
• . PACHYLASMATINAE
— Chitinous rods or lamina present in wall plates; horizontal growth ridges prominent, with
small setae; superior alar margins of C, CL2 and CL1 thickened distally .
. METALASMATINAE subfam. nov.
4. Chitinous tubes or lamina absent in wall plates; articular ridge of S prominent
. BATHYLASMATINAE
— Chitinous tubes or lamina present in wall plates; articular ridge of S not prominent
. HEXELASMATINAE
Subfamily EOLASMATINAE Buckeridge, 1983
Tables 1-2
Diagnosis. — Primary wall with 8 solid calcareous compartmental plates (R-RL-CLi-CL2-C); with or
without whorls of imbricating plates; basis membranous.
Type Genus. — Eolasma Buckeridge, 1983.
Distribution. — Upper Paleocene to Lower Miocene: New Zealand, Australia. Recent: South-west Pacific
(off Vanuatu and off the Loyalty Ridge), 700-850 m.
Remarks. — Presently the Eolasmatinae contains 2 genera, Eolasma and Waikalasma. Eolasma, and hence the
type genus of the eolasmatines, is based on a handful of disarticulated plates, which may or may not be from the
same species. Thus, there is no guarantee that the genus is 8-plated. Intact specimens o (Eolasma, as yet unknown,
would indicate whether Waikalasma, which is 8-platcd and which has additional whorls of basal imbricating plates,
is related to Eolasma or whether Waikalasma belongs elsewhere, and not in this subfamily.
160
D. JONES
Genus WAIKALASMA Buckeridge, 1983
Waikalasma Buckeridge. 1983: 64.
Diagnosis. — Wall with 8 solid, weakly articulated, calcareous compartmental plates including narrow R,
paired RL, CL1 and CL2, and wide C; alae well developed, particularly on C, barely inset from paries; radii absent;
4 whorls of imbricating plates present; basis membranous; caudal appendages absent.
Type Species. — Waikalasma juneae Buckeridge, 1983.
FOSSIL Species. — Waikalasma juneae Buckeridge, 1983.
Recent Species. — Waikalasma boucheti Buckeridge, 1996.
TYPES of Waikalasma juneae. — Holotype : AU 5622 R13/f7078; A 131, articulated shell with all eight
compartments present but in varying stages of decortication, from the Waikawua Formation (Otaian), collected by
J. and J. Buckeridge.
Paratypes : from AU 5622, A 132 carina; from AU 6849, A 133 carina; from AU 5622, A 134 incomplete
rostrum.
Holotype and para type depository: AU.
Distribution. — Lower Miocene to Recent. Oceania.
REMARKS. — Until recently the genus contained a single fossil species, W. juneae. However, the first known
living representative of the Eolasmatinae, W. boucheti Buckeridge, has recently been described from the seas off
Vanuatu, in the south-western Pacific Ocean (BUCKERIDGE, 1996). In the present report an incomplete specimen
collected from the Loyalty Ridge is also attributed to W. boucheti.
Waikalasma boucheti Buckeridge, 1996
Figs 4 b-c. 5-7
Waikalasma boucheti Buckeridge, 1996: 449.
MATERIAL EXAMINED. — Loyalty Ridge. Bathus 3: stn DW 778, 750-760 in: 1 specimen (incomplete) (MNHN-
Ci 2410). Drawn.
Types. — Holotype: MNHN-Ci 2428; Vanuatu, Musorstom 8, stn CP 1080, I5°57.4'N, I67°27'E. 799-
850 m; 05.10.1994.
Paratype: MNHN-Ci 2506; Vanuatu, MUSORSTOM 8, stn DW 1113, 14°53'N, 167°06’E. 700-736 m; 08.10.
1994.
Holotype and paratype depository: MNHN.
Diagnosis. — Waikalasma with 2 or more whorls of large imbricating plates, inner whorl with total of
8 plates; S triangular, very elongate, with long, low articular ridge, adductor muscle scar weak, central; T narrow,
inverted 'V' shaped, basal margin deeply excavated, spur at basi-scutal angle.
Supplementary Description. — Incomplete specimen consisting of C. paired opercular plates and soft body;
R. CL1 and CL2 not recovered. C solid, large; external growth ridges regular, low, giving plate smooth appear¬
ance, faint indications of 2 (?3) weak longitudinal ribs; C height 13.7; plate sloping inward gently toward orifice,
apex slightly spout-like, retroverted; alae triangular, large, placed approximately at right angle to paries, basal out¬
line rectangular, basal widths of alae and paries equal, alar growth ridges sinuously paralleling inferior alar margin,
alar welting absent; internally sheath covering 2/3 height of valve, inner surface of C below sheath smooth; base
of C slightly inflected inward, with lamina of chitin close to inner surface. Basis unknown. S triangular; elongate,
height more than twice width, externally with many spaced, regular, growth ridges; articular ridge barely
Source : MNHN. Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
161
projecting; internally adductor muscle scar slightly above middle of valve; depressor muscle scars indistinct.
I narrow, height 6 times width; "boomerang" shape due to deep indention of basal margin; externally well spaced,
regular, growth ridges; articular ridge projecting; internally tergal muscle crests lacking. Internal articular junction
between opercular plates sinusoidal. Color of preserved material dirty cream, with some brown markings.
Fig. 5. — Waikalasma boucheti Buckeridge, 1996. Specimen (incomplete) from BaTHUS 3. stn DW 778 (MNHN-Ci 2410):
a. T. internal view; b, T, external view; c, S. external view; d. S, internal view.
Labrum with shallow medial depression, numerous small teeth (approximately 47) across margin. Mandibular
palp ovate; dense setae terminally. Mandible with 3 teeth, tooth I hugest, slightly separated from teeth 2 and 3;
upper margins of 2 and 3 smooth; inferior angle 1/3 length of margin, bluntly molariform, sharply dentate.
Maxillule with 1-2 pairs of long, stout setae at upper angle; slight notch indicated 1/3 length down cutting
margin, dense spines on either side of notch, 7-9 pairs of smaller spines above notch, 7-9 pairs of larger setae
below notch; inferior angle small, straight, not protuberant, with 5-6 pairs of small setae. Maxilla wide; lobes
indistinct; long serrulate setae terminally.
Cirrus I with rami subequal; anterior ramus slightly longer and wider than posterior; both rami thickly setose,
setae finely serrulate distally. Cirrus II similar to cirrus III. longer than cirrus I; rami subequal; both rami with
proximal segments setose, setae simple or serrulate, both rami with distal segments tending toward antenniformy,
segments becoming oblong. Cirrus III with rami subequal, segments becoming oblong distally; posterior ramus
with 2-3 pairs of long, finely serrulate setae on anterior faces of distal segments; anterior ramus with 2-3 pairs of
long, finely serrulate setae on anterior faces of distal segments, proximal segments with 4 pairs. Cirrus IV to VI
similar, longer than cirrus III; segments oblong, with 2-4 pairs of setae on anterior faces, distal 2 pairs of setae
longest, finely serrulate. Chaetotaxy ctenopod. Cirral formula as follows:
Caudal appendages absent. Penis less than height of basal segment of pedicel of cirrus VI; annulated; naked;
without tufts of long setae distally.
REMARKS. — Externally there are faint indications of 2, possibly 3, weakly developed longitudinal ribs on
the C. Although no basal whorls of imbricating plates were found associated with the incomplete specimen,
the "chitinous lamina" in the slightly inwardly inflected base of C may be all that remains of the infolded, basal
portion of the imbricating plates, as described in W. boucheti (Buckeridge, pers. comm.), and thus the presence
of imbricating plates may be inferred.
162
D. JONES
Fig 6 Waikalasma boucheti Buckeridge. 1996. Specimen (incomplete) from Bathus 3. stn DW 778 (MNHN-Ci 2410V
a. labrum; b, maxilla; c right mandible; d, left mandible; e, right maxillule; f. left maxillule; g. mandibular palp:
h. cirrus III; i, cirrus II; j. cirrus I; k, cirrus VI. 1 H
There are some variations in the soft part morphology of the present material compared to the specimen
described from off Vanuatu by Buckeridge (1996). The inferior angle of the mandible is large (1/3 the length of
the cutting margin), bluntly molanform and sharply dentate; the maxillule has a slight notch 1/3 the distance down
the cutting margin, with dense spines on either side; the labrum has an extremely shallow medial depression with
numerous conical small teeth along the crest; cirrus I is much shorter than cirrus II and the proximal segments of
both ram. of cirrus I are much broader and narrower than the corresponding segments in cirrus II. The totality of
these differences would suggest that we are dealing with 2 distinct species. However, the paucity of complete
specimens precludes further discussion.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
163
Distribution. — Vanuatu (700-850 m); Loyalty Ridge (750-760 m).
Genus EOLA SMA Buckeridge, 1983
Eolasma Buckeridge. 1983: 62.
DIAGNOSIS. — Shell with 8 solid calcareous compartmental plates (R-RL-CL'-CL—C); external alar growth
ridges parallel inferior alar margin: T with prominent articular ridge, S with articular ridge weakly elevated; basis
membranous.
Type Species. — Eolasma maxwelli Buckeridge, 1983.
FOSSIL Species. — Eolasma maxwelli Buckeridge, 1983; Eolasma rugosa Buckeridge, 1985.
TYPES OF Eolasma maxwelli. — Holotype : GS1 1463 L35/F7537. AR 852 scutum [= number 5 on slide];
Eyre River Sands (Waipawan to Mangaorapan), White's Creek, collected by P. A. Maxwell.
Paratypes : AR 853-864 twelve scuta; AR 865-871 seven terga; AR 872-874 three rostrolatera; AR 875-876
two carinae or rostra; AR 877-878 two carinolatera; AR 879-880 two latera; all same locality as holotype.
Holotype and paratype depository. New Zealand Geological Survey. Lower Hutt. New Zealand.
Distribution. — Late Paleocene to early Eocene: New Zealand; Early Miocene: Victoria, Australia.
Remarks. — BUCKERIDGE (1983) considered that the distinct alae and the lack of extra whorls of imbricating
plates surrounding the primary shell wall make Eolasma more derived than Waikalasma.
Subfamily PACHYLASMATINAE Utinomi. 1968 (emend.)
Tables 1-4
Diagnosis. — Wall not surrounded by whorl(s) of imbricating plates. Adult externally with 6 or 4 solid
compartmental plates. Parietes with chitin absent; horizontal growth ridges low, lacking small setae. External alar
growth ridges parallel to inferior alar margin, alar welting absent. Opercular plates with articular ridges varying
from projecting to not projecting. Basis membranous or calcareous. Mandible tridentoid. Caudal appendages
present, sometimes reduced or vestigial.
Distribution. — Paleocene to Recent; Western Pacific, Africa. Mediterranean.
164
D. JONES
TABLE 3. — Pachylasmatidae: characters of the genera.
artic - articular: b-s - basi-scutal; ext. = external: fus. = fused; int. = internal; long. = longitudinal; n.a. = not applicable; part. fus. = partlv
fused; proj. = projecting; sep. = separate.
Remarks. Buckeridge (1983) considered that this subfamily was represented by a single genus,
Pachylasma, containing 12 species, of which 3 were fossil. Prior to the present study 3 more extant species of
Pachylasma were described (Foster, 1981; Rosell, 1981; Jones, 1993b). A second pachylasmatine genus,
Tetrapachylasma, was described by Foster (1988). Herein I describe 3 new genera within the Pachylasmatinae
( Eutomolasma gen. nov., Eurylasma gen. nov. and Microlasma gen. nov.) together with 1 1 new species. The
diagnosis of Tetrapachylasma Foster, 1988 is emended to reflect the infra-specific variation now recognised within
this genus.
Source . MNHN. Paris
CIRR1PEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
165
TABLE 4. — Pachylasmatidae: characters of the genera (continued).
The material examined herein indicates that some species have a fixed pattern of 6 wall plates in the adult,
whilst others exhibit a variable pattern of either 6 or 4. The species which have a fixed pattern fall into 4 species
groups. Given the other consistent morphological similarities exhibited by these groups of taxa. 3 can be
considered justifiably as genera — i.e. Eutomolasma gen. nov., Eurylasma gen. nov. and Microlasma gen. nov.
However the fourth group. Pachylasma , must be considered to be a grouping whose resolution has yet to be
determined. The affinities of the various species included in this genus are still unclear and. until further specimens
are collected and information on ontogenetic development and intra-specific variation assembled, it will be difficult
to resolve the relationships of the various taxa.
Adults of Eury lasma gen. nov., Eutomolasma gen. nov., Microlasma gen. nov. and Pachylasma have a fixed
pattern of 6-plates (compound rostral plate, paired CL1 and CL2, and C). In Eutomolasma. Microlasma and
Pachylasma, CL1 and CL2 are separate entities. These plates are coalesced to a certain degree in Eurylasma, but the
ala of CL2 is still evident, albeit reduced - thus the adult is still regarded as 6-plated.
One genus, Tetrapachylasma , is herein regarded as having a variable wall pattern. Within this genus CL1 and
CL2 may be separate entities, or coalesced to a certain degree, but the reduced ala of CL2 is still evident. Both of
these conditions represent a 6-plated shell (compound rostral plate, paired CL1 and CL2, and C). However. CL1 and
CL- can be totally coalesced, in which case the shell is regarded as 4-plated — i.e. compound rostral plate, paired
CL plates (coalesced CL1 and CL2) and C. Both the 4- and the 6-plated condition occur within populations of the
same species. No instances have been recorded of a 5-plated condition (i.e. CL1 and CL2 totally coalesced on one
side only). Thus, the adult shell pattern of this genus is considered to be a variable pattern of either 4 or 6 parietal
plates. Ontogenetic evidence has revealed that Tetrapachylasma arcuatum sp. nov. and 2 species which were
included previously within Pachylasma (i.e. P. aurantiacum Darwin. 1854 and P. ferrugomaculosa Jones. 1993b)
166
D. JONES
may have either 4 or 6 plates in the adult. Similarities in the form of the wall plates, the opercular plates and body
morphology suggest these species, along with T. ornatum sp. nov., can be included with T. Irigomwi Foster,
1988 to form a generic grouping.
Key to the genera of PACHYLASMATINAE
Tables 3-4
1 . Adults with CL1 and CL2 partially coalesced (CL2 retaining small ala) or coalesced (ala of
CL2 absent); rostral plate compound, sutures obscure or visible, basal width of R equal to
or smaller (1/3 or 1/6 basal width) than basal width of RL . 2
— Adults with CL1 and CL2 separate; rostral plate compound, sutures obscure or visible,
basal width of R equal to or wider (twice basal width) than basal width of RL . 3
2. Rostral plate compound, sutures obscure or visible, basal width of R smaller (1/3 or 1/6
basal width) than basal width ot RL; S concave, sub-pyramidal or pyramidal, due to angled
basal margin; articular ridge of T not projecting . EURYLASMA gen. nov.
— Rostral plate compound, sutures obscure or visible, basal width of R equal to basal width
ol RL; S not concave, not sub-pyramidal or pyramidal, basal margin straight; articular
ridge of T projecting . TETRAPACHYLASMA
3. Rostral plate compound, sutures visible, basal width of R equal to or wider (twice basal
width) than basal width of RL; growth ridges of S even, smooth, not cut by longitudinal
striae; basal width of CL2 smaller (1/5-2/3 basal width) than basal width of CL1 . 4
Rostral plate compound, sutures obscure or visible, basal width of R equal to basal width
ol RL; growth ridges of S convoluted, cut by longitudinal striae; basal width of CL2 equal
to basal width of CL! . PACH YLA SMA
4. Basal width of R equal to or wider (twice basal width) than basal width of RL; S convex
externally, elongated and narrow; S articular ridge not projecting; caudal appendages long
(Irom 1/4 longer than pedicel of cirrus VI to more than twice height of pedicel of cirrus
VI), 11-17 segments; basal width of CL2 1/5- 1/2 basal width of CL' .
. . . EUTOMOLASMA gen. nov.
— Basal width of R equal to basal width of RL; S not convex externally, S as wide as high;
S articular ridge projecting; caudal appendages short (from less than 1/2 height of basal
segment of pedicel of cirrus VI to slightly longer than height of pedicel of cirrus VI).
6-9 segments; basal width of CL2 1/4-2/3 basal width of CL' .
. M ICROLASMA gen. nov.
Genus EUTOMOLASMA gen. nov.
Fig. 8; Tables 3-6
,R rr,A|GpI°2lr\';Symri,riCal ShC" f0mi- °rifice large‘ Adu" with fixed -shell-plate pattern of 6 parietal plates
rt i Jrn 7 R°Slra p ate Compound- sulurcs visible- basal width of R equal to or twice basal width of RL
LL and CL- separate. S convex externally; elongated, narrow; with smooth, even, transverse growth ridges not
cut by longitudinal striae. Caudal appendages longer than pedicel of cirrus VI by 1/4 to at lea's! twice height of
pedicel ol cirrus VI; 11-17 segments. c
Type Species. — Eutomolasma chinense (Pilsbry. 1912).
m SPfCIE(S- “ Eutomolasma chinense (Pilsbry. 1912); E. japonicun, (Hiro, 1933); E. maclaughlinae
sp. nov., t. orbiculatum sp. nov.
Source . MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
167
Tables 5 and 6. — Eutomolasma gen. nov.: characters of the species
b-s = basi-scutal; ext. = external; int. = internal; long. = longitudinal; proj. = projecting.
artic. - articular; b-s = basi-scutal; ext. = external; long. = longitudinal; max. = maximum; segs = segments
168
D. JONES
Remarks. — The symmetrical shell form and the large orifice, the externally convex, elongated and narrow S
with smooth S growth ridges and the long caudal appendages (11-17 segments) distinguish this genus. Other
distinguishing characters are listed in Tables 3-6. The species included herein are moderate (maximum: RC length
10.5, C height 8.2) to large (maximum: RC length 23.0, LD 29.9. C height 9.6, R height 9.0) in size and occur
in association with gastropods, echinodcrms, brachiopods, balanomorph and scalpcllid barnacles, coral and pumice.
ETYMOLOGY. — From the Greek eutomos, "regular", in reference to the symmetrical form of the shells of
the species in this genus.
Distribution. — Western to central Pacific, 55-647 m: Loyalty Islands, Norfolk Ridge, 260-460 m
(E. orbiculatum)', Lord Howe Ridge, New Caledonia, Loyalty Islands, Norfolk Ridge, Futuna Island, Combe
Bank, Walerwich Bank, 245-647 m (£. maclauglilinae) ; South China Sea, 380 m (E. chinense ); SW Japan, 55-
364 m (E. japonicum).
FIG. 8. — Eutomolasma gen. nov. Distribution map: ■ , E. chinense (380 m); a , E. japonicum (55-364 m);
• , E. orbiculatum (260-460 m); O, E. maclauglilinae (245-647 m).
Eutomolasma chinense (Pilsbry, 1912)
Fig. 8; Tables 5-6
Pachylasma chinense Piisbry, 1912: 293; 1916: 329. — Newman & Ross, 1976: 40.
TYPES. — Holotype: USNM Cat. No. 43471; "Albatross", stn 5301. China Sea, near Hong Kong. 20°37‘N,
1 15°43’E, 380 m, on scutum of a living ScalpeUum stearnsii Pilsbry; mouthparts and caudal appendages on slide,
shell and opercular plates separate.
Holotype depository: USNM (shell and opercular plates have not been located).
DIAGNOSIS. — Orifice piriform, toothed. Parietal plates thin, covered with thin, yellowish epicuticle. Basal
width of R equal to basal width of RL, R separated from RL by sutures, inconspicuous externally, conspicuous
internally. Basal width of CL2 1/2 basal width of CL1. Superior alar margins arched, finely serrate. S convex
externally. T externally with longitudinal depression absent; articular ridge of T not projecting beyond articular
margin; T spur short, wide.
Remarks. — Eutomolasma chinense is a moderate sized species (maximum: RC length 16.5, C height 13.6),
characterised by the equal basal widths of R and RL, the basal width of CL2 being 1/2 the^basal width of CL', and
the S being externally convex.
Source : MNHN, Paris
CIRR1PEDIA TH0RAC1CA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
169
The calcareous rather than membranous basis, the inconspicuous rather than conspicuous external sutures of
the rostral plate, the basal width of CL2 being 1/2 the basal width ot CL1, rather than less than 1/2 the basal
width of CL'; the alae being arched rather than parallel to the basis, the short and wide, rather than indistinct,
tergal spur; the absence rather than presence of a longitudinal ridge externally on the T, close to the S margin;
and the thin parietal plates, white with pink apices and covered with a yellowish epicuticle. rather than glossy
pink plates, distinguish E. chinense from E. japonicum. The differences between E. maclaughlinae sp. nov. and
E. chinense are listed under E. maclaughlinae , and between E. orbiculatum sp. nov. and E. chinense are listed
under E. orbiculatum.
Distribution. — South China Sea, 380 m.
Eutomolasma japonicum (Hiro, 1933)
Fig. 8; Tables 5-6
Pachylasma japonicum Hiro, 1933: 65; 1937: 430. — Newman & Ross, 1976: 40. — ROSELL. 1991: 37.
Types. — Syntypes : SMBL Type; station 303, offToi-saki. Hiuga-nada, 31°16'00"N, 131 33 15 E. 364 m.
12.06.1928, bottom sand; 7 specimens attached to broken shells of Clypeaster japonicus Doderlein, together with
Scalpellum laccadivicum Annandalc.
Syntypes depository: SMBL (specimens have not been located).
DIAGNOSIS. — Form rounded oval from above. Parietal plates with numerous, smooth, regular growth ridges.
Basal width of R equal to basal width of RL; rostral sutures visible externally. Basal width of CL2 less than 1/2
basal width of CL1. Alae broad, triangular, distinctly striped verically, summits almost parallel to basal margins.
S elongate, triangular, externally feebly convex; articular ridge not projecting. T triangular, externally longitudinal
ridge set close to S margin; articular ridge slightly projecting; 6-7 muscle crests projecting beyond basal margin;
T spur indistinct.
Remarks. — The slightly projecting tergal articular ridge, the 6-7 tergal muscle crests projecting heyond the
tergal basal margin and the externally feebly convex S characterise E. japonicum. Differences between
E. japonicum and E. chinense are listed under E. chinense. between E. japonicum and E. maclaughlinae sp. nov.
under E. maclaughlinae. and between E. japonicum and £. orbiculatum sp. nov. under £. orbiculatum.
Eutomolasma japonicum is a large species (maximum: RC length 21.0, R height 1 1.0).
Distribution. — SW Japan, 55-364 m.
Eutomolasma maclaughlinae sp. nov.
Figs 8-11; Tables 5-6
Material EXAMINED. — Chesterfield Islands. MUSORSTOM 5: stn DC 388, 500-510 m: 2 specimens, attached
to coral.
New Caledonia. "Vauban": 12.04.1978: 8 specimens, some attached to shell of gastropod.
MUSORSTOM 4: stn CP 170, 460 m: 1 specimen. — Stn CP 179, 475 m: 1 specimen. — Stn CP 179. 475 m:
5 specimens, on pumice and shells of gastropod. — Stn DW 183. 280 m: 1 specimen, on Chirona ( Striatobalanus )
amar\llis (Darwin. 1854), latter attached to a sponge. — Stn CP 215. 485-520 m: 2 specimens. — Stn DW 221.
535-560 m: 1 specimen. — Stn DW 222, 410-440 m: 5 specimens. — Stn DW 223. 545-560 m: many specimens. —
Stn DW 229, 445-460 m: many specimens, some attached to gastropod shell (MNHN-Ci 2695). Drawn; 1 specimen
(MNHN-Ci 2391); 9 specimens, attached to gastropod shell (WAM C 23244); many specimens, attached to gastropod
shells (WAM 251-96). — Stn DW 230, 390-420 m: 3 specimens.
Smib 8: stn DW 167, 430-452 m: 1 specimen (WAM C 23245). — Stn DW 197. 414-436 m: 4 specimens.
Stn DW 198, 414-430 m: several specimens. — Stn DW 199, 408-410 m: 1 specimen, attached to gastropod.
Bathus'2: stn DW 718. 430-436 m: several specimens, attached to brachiopod shells and echinoderm tests tW'AM
C 23246). _ Stn DW 719. 444-455 m: many specimens (1 ovigerous). attached to gastropod shells and echinoderm
170
D. JONES
tests (WAM C 23247). — Stn DW 720. 530-541 m: 2 specimens (WAM C 23248); many specimens. — Stn DW 729.
400 m: many specimens. — Stn DW 738, 588-647 m: 1 specimen, attached to gastropod; 1 specimen.
Bathus 4: stn CP 909, 516-558 m: 2 specimens, attached to shells of gastropods.
Loyalty Islands. Biocal: stn DW 44. 440 m: many specimens, attached to shells of gastropods and test of
echinoderm. — Stn CP 45. 430-465 m: 17 specimens (6 juveniles), attached to fragment of test of echinoderm and
shells of gastropods.
MUSORSTOM 6: stn CP 465. 480 m: 1 specimen, attached to fragment of echinoderm test. — Stn CP 466. 540 m:
2 specimens, attached to gastropods (USNM); 2 specimens. — Stn CP 467, 575 m: many specimens, some attached to
shells of gastropods.
Norfolk Ridge. Smib 2: stn DW 1, 438-444 m: several specimens on gastropod shell. — Stn DW 2. 438-448 m:
I specimen (BMNH). — Stn DW 12. 445-460 m: 1 specimen, attached to shell of gastropod.
Smib 3; stn DW 21, 525 m: 2 specimens. — Stn DW 22, 503 m: 12 specimens. — Stn DW 24, 525 m: 1 specimen;
1 specimen (broken). — Stn DW 25. 437 m: 3 specimens. — Stn DW 27, 457 m; 1 specimen.
Smib 4: stn DW 58. 560 m: 11 specimens. — Stn DW 61, 550 m: 1 specimen. — Sin DW 64, 460 m: several
specimens on gastropod shell. — Sin DW 65, 420 m: many specimens attached to gastropod shells. — Stn DW 68,
440 m: 3 specimens, attached to gastropod shells.
Bathus 3: sin DW 829. 386-390 m: 1 specimen, attached to shell of gastropod (MNHN-Ci 2392). — Sin CP 833,
441-444 m: 11 specimens, attached to fragments of echinoderm test and shell of gastropod.
Futuna Island. Musorstom 7: sin CP 505, 245-400 m: 1 specimen, attached to fragment of shell of gastropod.
Combe Bank. MUSORSTOM 7: stn DW 547. 455 m: 1 specimen.
Waterwich Bank. MUSORSTOM 7: stn DW 537, 325-400 m: 1 specimen, attached to shell of gastropod.
Types. — Holotype: MNHN-Ci 2391 (MUSORSTOM 4, stn DW 229).
Paratypes : BMNH (Smib 2, stn DW 2). — MNHN-Ci 2392 (Bathus 3, sin DW 829). — MNHN-Ci 2695
(Musorstom 4. stn DW 229). Drawn. — USNM (Musorstom 6, stn CP 466). — WAM 251-96
(Musorstom 4, stn DW 229).
Fig. 9. — Eutomolasma maclauglilinae sp. nov. Paratype from Musorstom 4, stn DW 229 (MNHN-Ci 2695):
a. S, external view; b. S, internal view; c, T, external view; d. T, internal view; e. whole animal, lateral view;
f, whole animal, from above.
Source . MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
171
DIAGNOSIS. — Form rounded oval from above; orifice smooth, large, diamond-shape. Basal width ol R twice
basal width of RL; rostral sutures visible externally. Basal width of CL- 1/2 basal width ol CL1. Superior alar
margin subparallel to basal margin, finely serrate. S elongate triangle, feebly convex externally. Articular ridge ol
T scarcely projecting; externally with 1 parallel, longitudinal ridge on each side of spur; 7-9 muscle crests
extending beyond basal margin; T spur short.
Description. — Shell moderate sized
(maximum: RC length 10.5. C height 8.2). Parietal
plates with growth ridges barely visible, irregular,
most developed on rostral plate. Rostral plate
convex; basal width twice basal width of CL1,
height subequal to height of C; sutures between R
and RL visible externally and internally. CL1 and
CL2 separate; basal width of CL1 subequal to basal
width of R. twice width of CL2. Basal width of CL2
1/2 basal width or CL'; CL2 higher than CL1 by
1/3. Height of C equal to or slightly higher than R.
basal width equal to that of CL1; apex sloping
inward to orifice, not retroverted and spout-like. Alae
of C, CL1 and CL2 subequal, broad, triangular; alar
growth ridges regular, parallel to inferior alar
margin. Basis membranous; flat. Opercular plates
orientated from sub-parallel to basis to angle ol 40°
to basis. S elongate triangle, basal margin 7/15
length of occludent margin; apex acute; externally feebly convex, growth ridges regular, smooth, not cut by
longitudinal striae; internally adductor muscle and lateral depressor muscle pits absent; articular furrow narrow;
articular ridge long, extending 3/4 length of valve, not projecting beyond articular margin; adductor ridge indistinct.
T similar size to S, sub-triangular; externally with fine growth ridges, fine longitudinal striations in carinal area,
2 parallel, longitudinal ridges, 1 either side of spur; internally with 7-9 distinct muscle attachment crests extending
beyond basal margin; articular margin convex; articular ridge long, barely projecting beyond articular margin;
articular furrow wide; spur short, rounded, 1/9 width of basal margin, set at its own width Irom basi-scutal angle.
Color of parietal plates peachy orange, C and CL2 deeper orange; shell sometimes pale ochre-orange or dirty cream
color; opercular plates lemon yellow; eggs orange. Measurements of 10 specimens examined, randomly selected
Fig. 10. — Eutomolasma maclaughlinae sp. nov. Paratypes
from MUSORSTOM 4, stn DW 229 (WAM 251-96):
oblique lateral view. Scale = 1cm.
from several stations, as follows:
Labrum with shallow medial depression; small teeth present at edges of depression, teeth absent from
central area. Mandibular palp club-shaped; dense, serrate setae terminally. Mandible with 3 teeth, tooth I largest,
separated from teeth 2 and 3; upper margins of 2 and 3 with subsidiary cusps; interior angle molaritorm,
dentate. Maxillule with 1 pair of long, stout setae at upper angle; notch below upper angle 1/4 length of cutting
margin, with 5-6 pairs of smaller setae; straight cutting margin below with 5-7 pairs of longer, stout setae;
inferior angle slightly sinuous, with 5-6 pairs of smaller setae. Maxilla as wide as high, with long serrulate setae
terminally.
Cirrus I with rami unequal; anterior ramus longer than posterior; proximal segments ol anterior ramus barely
protuberant anteriorly; both rami with segments moderately setose. Cirrus II longer than cirrus I; rami subequal,
anterior ramus shorter than posterior; proximal segments of anterior ramus slightly protuberant anteriorly; distal
segments of posterior ramus antenniform; all segments with dense, simple setae, some finely serrulate setae
172
D JONES
distally. Cirrus III longer than cirrus II, rami subequai, segments becoming oblong distally; posterior ramus with
2-4 pairs of long, finely serrulate setae on anterior faces of distal segments; anterior ramus with 2-4 pairs of long,
finely serrulate setae on anterior faces of more distal segments, proximal segments with 8 pairs. Cirrus IV to VI
similar, longer than cirrus III; segments oblong, with 4 pairs of setae on anterior faces, distal 2 pairs longest,
finely serrulate. Chaetotaxy ctenopod. Cirral formula as follows:
Caudal appendages at least twice length of pedicel of cirrus VI; 11-17 segments; sparse circlet of long, fine
setae apically and at distal borders of segments, setal length equal to segmental length. Penis subequal to cirrus VI;
annulated. sparsely setose, with sparse circlet of long setae distally. Eggs large, 0.5 x 0.4 mm.
Remarks. — Eutomolasma maclaughlinae sp. nov. differs from all other members of the genus by the large
size of the R (basal width twice the basal width of RL); by CL2 being higher, not shorter, than CL1; and by T
possessing 2 external longitudinal ridges, 1 each side of the spur. It is further separated from E. cliinense by the
membranous rather than calcareous basis, by the rostral plate sutures being visible externally rather than being
inconspicuous, by the absence of an epicuticle, rather than a thin, yellowish epicuticle covering the parietal plates;
and by the color of the parietal plates (peachy orange or pale ochre/orange to dirty cream rather than white tinted
with pink apically).
The new species is similar to E. orbiculatum sp. nov. as both species have a convex tergal articular margin
(more pronounced in £. orbiculatum), but E. maclaughlinae may be separated from £. orbiculatum as follows: by
its smaller size (maximum: RC length 10.5, RC length 23.0, respectively), by the basal width of CL2 being 1/2
that of CL1 rather than 1/4- 1/5; by the T articular ridge scarcely projecting, rather than extending well beyond
the articular margin; by the length of the caudal appendages (at least twice the height of the pedicel of cirrus VI
rather than longer than the pedicel of cirrus VI by 1/4); and by the dentition of the labrum (small teeth present at
the edges of the median depression, rather than absent).
Eutomolasma japonicum differs from £. maclaughlinae sp. nov. in C being the shortest of all the plates (rather
than equal to or higher than R). by the basal width of R being equal to. rather than twice, the basal width of RL;
by the basal width of CL2 being less than 1/2 that of CL', rather than 1/2; by the height of CL2 being 8/9 the
height ol CL i rather than higher than CL' by 1/3; by the T spur being indistinct rather than short and rounded; and
by the color (glossy pink parietal plates, fading toward a whitish R, rather than peachy orange or pale
ochre/orange).
Etymology. — Named in honour of my friend and colleague Dr Patsy A. McLaughlin, in recognition of her
contributions to cirripedology. and who first noted this species on mollusc shells inhabited by hermit crabs.
Distribution. — Chesterfield Islands, 500-510 nr, New Caledonia, 280-647 m: Loyalty Islands. 430-575 m;
Norfolk Ridge, 386-560 m; Futuna Island, 245-400 m: Combe Bank. 455 m; Waterwich Bank. 325-400 m.
Eutomolasma orbiculatum sp. nov.
Figs 8. 12-14; Tables 5-6
Islands. Biocal: stn DW 83, 460 m: 3 specimens
Material examined. — Loyalty
Ci 2696). Drawn.
Norfolk Ridge. Smib 4: stn DW 55. 260 m: 13 specimens (ontogenetic series), attachec
caledomcus Bouchet & Metivier (WAM 252-96). - Stn DW 68. 440 m: 1 specimen (MNHN-Ci 2394)
(MNHN-Ci 2393,
to Petrotrochiis
Source : MNHN: Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
173
0.5 mm
- a-c
0.5 mm
d
0.1 mm
- e-k
Fig. 11. — Eutomolasma madaughlinae sp. nov. Paratype from Musorstom 4, sin DW 229 (MNHN-Ci 2695): a. cirrus I;
b, cirrus II; c, cirrus III; d, cirrus VI. penis and caudal appendage; e. labrum; f. right mandible; g, left mandible;
h. right maxillule; i, left maxillule; j, maxilla; k, mandibular palp.
Types. — Holotype : MNHN-Ci 2393 (Biocal, stn DW 83).
Paratypes: MNHN-Ci 2394 (Smib 4. stn DW 68). — MNHN-Ci 2696 (BlOCAL, stn DW 83). Drawn. —
WAM 252-96 (Smib 4, stn DW 55).
174
D. JONES
Diagnosis. — Shell large (maximum: RC length 23.0, C height 9.6); basal circumference subcircular. Orifice
smooth, rounded pentagonal. Parietal plates smooth, growth ridges faint. Basal width of R equal to basal width of
RL; rostral sutures visible externally and internally. Basal width of CL2 1/4-I/5 basal width of CL1. Alae
triangular, superior alar margins slightly serrate. Basis membranous; Hat. S convex externally. T articular ridge
strongly projecting; spur indistinct; externally with longitudinal ridge close to S margin; 15-18 small muscle
attachment ridges extending just beyond basal margin.
5.0 mm
e.f
Fig. 12. — Eutomolasma orbiculatum sp. nov. Paratype from Biocal, stn DW 83 (MNHN-Ci 2696): a. S. internal view;
b. S. external view; c, T, external view; d, T, internal view; e, whole animal, lateral view: f. whole animal, from
above (right S and T removed).
Description. — Parietal plates smooth, growth ridges faint. Rostral plate convex, largest plate; sutures
between R and RL visible internally and externally, dividing plate into 3, basal width of R equal to basal width of
RL. Basal width of CL' subequal to that of rostral plate; CL2 smallest plate; basal width of CL2 1/4 to 1/5 basal
width of CL1; height of CL2 4/5 height of CL1. C convex; narrower than CL1 and rostral plate, wider than CL2;
shorter than or subequal to height of R. Alae triangular, those of CL1 widest, alae of CL2 2/3 width of alae of
CL1, alae of C 1/3 width of alae of CL'; superior alar margins slightly serrate; alar growth ridges parallel to
inlerior alar margin, faintly marked. Basis membranous; flat. Orifice large, roundly pentagonal; not toothed.
Opercular plates lodged sub-parallel to basis. S convex externally, elongate triangle, occludent margin twice length
o! basal margin; apex acute; externally with regular, smooth, transverse growth ridges not cut by longitudinal
striae; internally adductor muscle pit faint, at slightly less than 1/2 height of valve; lateral depressor muscle pit
absent, articular furrow narrow, shallow; articular ridge extending 3/4 length of valve, not projecting beyond
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
175
articular margin; adductor ridge absent. T 2/3 size of S; externally growth ridges faint, except in area of articular
ridge (visible when whole animal viewed from above), with longitudinal ridge set at distance of 1/5 width of basal
margin from S margin; internally with 15-18 weak muscle crests projecting slighty beyond basal margin; articular
margin distinctly convex; articular ridge prominent, projecting beyond articular margin; articular furrow well-
developed; spur indistinct, width 1/4 width of basal margin, set at distance of 1/5 width basal margin from basi-
scutal angle. Color of parietal plates pale ycllow/cream with some ochre/orange on C and CL2; opercular plates
cream. Measurements of 10 specimens examined, randomly selected from several stations, as follows:
Labrum with shallow, small teeth absent; dense
patch of stiff, small, downwardly pointing setae on
either side of medial depression. Mandibular palp
broadly ovate; long serrate setae terminally.
Mandible with 3 teeth, equally sized, equally
separated; upper margin of teeth 2 and 3 with few
weak subsidiary cusps; inferior angle large, bluntly
molariform. strongly dentate. Maxillule setose;
several long, stout setae at upper angle; notch below
upper angle ill-defined, with 4-6 pairs of smaller
setae; cutting margin below irregular or slightly
stepped, with 7-8 pairs of longer, stout setae;
inferior angle sinuous, with 9-10 pairs of small
setae. In illustrated specimen left mandible showing
following variations — notch barely defined; stout,
serrulate setae interspersed between stout setae on
cutting margin; inferior angle small, with 4 pairs of
small setae. Maxilla wide, coxal endite well-developed; setae long, dense, simple or serrulate.
Cirrus I with rami unequal; anterior ramus longer than posterior; proximal segments of anterior slightly
protuberant anteriorly; both rami with segments setose; postero-distal angles of segments of both rami with small
tufts of simple and finely serrulate setae, surface of segments with combs of small, simple setae. Cirrus 11 longer
than cirrus I; rami unequal, anterior ramus shorter than posterior; proximal segments of anterior broader than those
of posterior and slightly protuberant anteriorly; distal segments of posterior ramus antenniform; all segments with
simple and finely serrulate setae, serrulate setae distally. Cirrus III longer than cirrus II; rami subequal, segments
becoming oblong distally; posterior ramus with 2-4 pairs of long, finely serrulate setae on anterior faces of distal
segments; anterior ramus with 2-4 pairs of long, finely serrulate setae on anterior faces of distal segments,
proximal segments with 10 pairs. Cirrus IV to VI similar, longer than cirrus III; segments oblong, with 4 pairs of
setae on anterior faces, distal 2 pairs of setae longest, finely serrulate. Chaetotaxy ctenopod. Cirral formula as
follows:
Fig. 13. — Eutomolasina orbiculatum sp. nov. Holotype,
Biocal, stn DW 83 (MNHN-Ci 2393): lateral view.
Scale = I cm.
Caudal appendages longer than pedicel of cirrus VI by 1/4 length of pedicel of cirrus VI; 14-15 segments;
sparse, long, fine setae apically and around distal margins of segments. Penis 1/4 length of cirrus VI; finely
annulated; sparsely setose, with circlet of dense long setae distally.
176
D. JONES
Fig. 14. — Eutomolasma orbiculatum sp. nov. Paratype from Biocal, stn DW 83 (MNHN-Ci 2696): a, cirrus VI, posterior
ramus, median segment; b, labrum; c, penis; d, mandibular palp; e. right maxillule; f. left maxillule; g. right
mandible; h, left mandible; i, maxilla; j, cirrus III, posterior ramus, median segment; k, cirrus III; 1. cirrus VI and
caudal appendage; m, cirrus II; n, cirrus I, posterior ramus, median segments; o, cirrus 1.
Remarks. — The strongly projecting tergal articular ridge, the 15-18 weak muscle crests projecting slightly
beyond the tergal basal margin and the externally convex S characterise Eutomolasma orbiculatum sp. nov.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
177
The species is similar to E. japonicum , but may be distinguished from that species by the narrow secondary
CL2, whose basal width is 1/4- 1/5, rather than less than 1/2, the basal width of CL1; by the basis being totally
membranous rather than calcareous at the edges and membranous centrally; by features of the S, which in
E. orbiculatum is convex externally, with the basal margin 1/2 the length of the occludent margin, and feebly
convex, with the basal margin less than 1/2 the length of the occludent margin in E. japonicum ; by the T having
a convex articular margin and a prominent articular ridge extending well beyond the articular margin
(£. orbiculatum) rather than a straight articular margin and a scarcely projecting articular ridge (£. japonicum ); by
the relative lengths of the caudal appendages, being longer than the pedicel of cirrus VI by 1/4 the height of the
pedicel of cirrus VI (14-15 segments) rather than twice the height of the pedicel of cirrus VI (13-14 segments); and
by the color - parietal plates pale yellow-cream with some ochre-orange (E. orbiculatum), rather than glossy pink
(E. japonicum). The new species differs from E. maclaughlinae sp. nov. in size, in the overall shell form and in
the soft part morphology (see remarks under E. maclaughlinae).
Eutomolasma orbiculatum sp. nov. may be distinguished from E. chinense by the rostral sutures being visible
externally and internally, rather than inconspicuous; by the basal width of CL2 being 1/4- 1/5 the basal width of
CL1 rather than 1/2 the basal width of CL1; by the membranous rather than calcareous basis; by the tergal articular
ridge strongly projecting, rather than not projecting; and by the color of the parietal plates — pale yellow/cream
with some ochre/orange, rather than white with pink apices and covered with a yellowish epicuticle.
ETYMOLOGY. — From the Latin orbiculatum, "circular", in reference to the circular circumference of the shell.
Distribution. — Loyalty Islands, 460 m; Norfolk Ridge, 260-440 m.
Genus MICROLASMA gen. nov.
Fig. 15; Tables 3-4, 7-8
Diagnosis. — Adult with fixed shell pattern of 6 weakly articulated, thin parietal plates (R-CL1-CL2-C).
Rostral plate compound, sutures visible, basal widths of R and RL subequal. CL1 and CL2 separate. S externally
flat, as wide as high, with smooth, even transverse growth ridges, not cut by longitudinal striae. Caudal appendage
length ranging from less than height of basal segment of pedicel of cirrus VI to slightly longer than pedicel ol
cirrus VI, 6-9 segments.
Type Species. — Microlasma fragile sp. nov.
Recent Species. — Microlasma arwetergum (Rosell, 1 99 1 ); M. crinoidophilum (Pilsbry, 1911); M. fragile
sp. nov.; M. ochriderma (Foster, 1981).
Remarks. — Microlasma gen. nov. is similar to Eutomolasma gen. nov. as both have an adult fixed wall
pattern of 6 plates, a compound rostral plate with visible sutures and CL1 and CL2 as separate entities. However,
the genera may be distinguished by characters of the S and the caudal appendages. The S ol Microlasma is as wide
as high and is flattened, that of Eutomolasma is elongated, narrow and convex externally. The caudal appendages of
Microlasma are 6-9 segmented and small, ranging from less than the height of the proximal segment of the pedicel
of cirrus VI to just longer than the pedicel of cirrus VI, whereas those of Eutomolasma arc 11-17 segmented and
long (longer than the pedicel of cirrus VI by 1/4 the height of the pedicel of cirrus VI, to at least twice the height
of the pedicel of cirrus VI). Other distinguishing characters are listed in Tables 3, 4, 7 and 8.
The species included within Microlasma gen. nov. are relatively small (maximum: RC length 9.0. LD 6.2,
C height 7.0. R height 2.6). Ovigerous specimens of M. arwetergum have an RC length of 4.2 and those of
M. fragile sp. nov. 5.3. The species occur in association with animal substrates — e.g., M. arwetergum and
M. crinoidophilum with crinoids, M. ochriderma with gorgonians, and M. fragile with gorgonians and sponges,
but further records may show these associations to be purely facultative.
178
D. JONES
Tables 7 and 8: Microlasma gen. nov; characters of the species
artic. = articular; b-s = basi-scutal; ext. = external; int. = internal; proj. = projecting.
artic. - articular; c.a. - caudal appendage; long. = longitudinal; max. = maximum; proj. = projecting.
Etymology. — From the Greek mikros, "small",
constitute this genus.
in reference to the small size of the
species which
Source : MNHN, Paris
CIRR1PEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
179
Distribution. — Western Pacific Ocean. East China Sea, Japan, 274-400 m ( M . crinoidophilum);
Philippines, 120-123 m (M. arwetergum)\ Norfolk I,, 392-423 m (M. ochriderma ): Loyalty Islands, 245 in
(M. fragile).
FlG. 15. — Microlasma gen. nov. Distribution map: ▲. M. crinoidophilum (274-400 m); • . M. arwetergum (120-123 m);
O, M. fragile (245 m); ■ , M. ochriderma (392-423 m).
Microlasma arwetergum (Rosell, 1991)
Fig. 15; Tables 7-8
Pachylasma arwetergum Rosell, 1991: 35, figs 6 j-k. 8 f-h, 9 a-f.
Material EXAMINED. — Philippines. Musorstom 3: stn 124, I2°02.6’N, 1 2 1 °35.3'E, 120-123 m: holotype
(MNHN-Ci 2103).
Types. — Holotype: MNHN Ci 2103; Philippines, Musorstom 3, stn 124, 12°02.6'N. 121°35.3'E. 120-
123 m; attached to cirri of crinoids, with Heteralepas cornuta (Darwin, 1851).
Paratypes : MNHN Ci 2104; Philippines, MUSORSTOM 3: stn 124, 12°02.6'N. 1 2 1 °35.3'E, 120-123 m. —
UPIBM Crust. Coll. No. 372; Philippines. Musorstom 3: sin 124, 12°02.6'N, 121°35.3'E, 120-123 m.
Holotype depository >: MNHN.
Paratypes depository. MNHN, UPIBM.
Diagnosis. — Shell conical, more or less elongated along rostrocarinal axis. CL2 sharply tapering, apex so
narrow that apex of CL1 almost touches C; basal width of CL2 1/4 basal width of CL1; CL2 shorter Ilian CL1 by
1/5. C deeply concave, almost V-shaped. S transversely elongated towards basioccludent angle; S basal margin
5/7 length of occludent margin, S articular ridge projecting. T smaller than S, arrow-shaped; long spur 1/2 height
of valve, width 2/9 width of basal margin, set at distance of 1/9 length of basal margin from basi-scutal angle:
3-5 muscle crests projecting beyond basal margin. Mandible with 3 teeth, upper margin of tooth 3 and/or 2 with
small subsidiary cusps. Caudal appendages slightly longer than pedicel of cirrus VI. 7-9 segments.
Remarks. — This species (maximum RC basal diameter 5.0; ovigerous specimens RC length 4.2)
is distinguished from all others in the genus by the unique, arrow-shape of the T, produced by the long T spur
and a deep incision on the tergal basal margin, and by the pinkish-brown color of the parietal plates. Microlasma
arwetergum resembles M. crinoidophilum , as both species have small, sharply tapering CL2 (1/4 the basal
width and 1/2 the basal width of CL1, respectively), the apices of which are shorter than the apices of CL1.
The caudal appendages of M. arwetergum are 7-9 segmented and are longer than the pedicel of cirrus VI,
180
D. JONES
whilst those of M. crinoidophilum are 8-segmented and not as long as the basal segment of the pedicel of cirrus
VI. Microlasma arwetergum is a smaller species than M. crinoidophilum (maximum RC length 5.0, 9.0,
respectively).
Microlasma arwetergum differs from M. fragile sp. nov. and M. ochriderma as both the latter have CL2 (basal
width of CL2 1/2 to 1/3 basal width CL1, basal width of CL2 2/3 basal width CL1, respectively) which do not
taper narrowly and whose height equals the height of CL1. Differences between M. arwetergum and M. fragile
sp. nov. are listed under M. fragile. The caudal appendages of M. ochriderma are shorter (equal to the height of
the basal segment of the pedicel of cirrus VI, 7-segmented) than those of M. arwetergum (longer than the pedicel
of cirrus VI, 7-9 segmented).
Distribution. — Philippines, 120-123 m.
Microlasma crinoidophilum (Pilsbry, 1911)
Fig. 15; Tables 7-8
Pachylasma crinoidophilum Pilsbry, 1911: 81, fig. 11, pi. 17 figs 1-11; 1916: 329. — KROGER, 1911: 460. —
Nilsson-Cantell, 1932; 14, fig. 5, pi. 1 figs 1-2. — Utinomi, 1958: 307; 1968: 24, fie. 3. — Newman & Ross.
1976: 40. — Rosell, 1991: 37.
MATERIAL EXAMINED. — Japan. " Albatross stn 4934: off Sala Misaki, Kagoshima Gulf. 274-400 m,
16.08.1906: hololype (USNM 3867).
TYPES. — Holotype: USNM 38675; Japan, "Albatross": sin 4934, off Sala Misaki, Kagoshima Gulf, 274-
400 m, 16.08.1906; dry (1 lot) and wet (1 lot).
Holotype depository: USNM.
Diagnosis. — Shell basal contour oblong. C and CL1 largest plates; CL2 shorter than CL1 by 1/5, apices of
former tapering abruptly, so that alae of CL1 almost touch alae of C; basal width of CL2 1/2 basal width of CL1.
C vertical, other parietal plates sloping inward. S width equal to 1/2 S length, S basal margin 2/3 width of
occludent margin, articular ridge projecting. T externally with longitudinal depression running along S border; spur
short, 2/5 width of basal margin, set at distance of 1/10 length of basal margin from basi-scutal angle; 5-6 muscle
crests projecting from basal margin. Mandible with 3 teeth, upper margins of teeth 3 and 2 often with small
subsidiary cusps, inferior angle dentate. Caudal appendages less than 1/2 height of basal segment of pedicel of
cirrus VI, 8 segments.
Remarks. — The species is small (maximum: RC length 9.0, C height 7.0). Differences between
M. crinoidophilum and M. arwetergum, and between M. crinoidophilum and M. fragilis sp. nov., are listed under
M. arwetergum and M. fragilis sp. nov., respectively.
Microlasma crinoidophilum differs from M. ochriderma in the form of CL2. In M. crinoidophilum CL2
is sharply tapering and shorter than CL1 by 1/5, whereas in M. ochriderma CL2 and CL1 are equal in height,
and the apex of CL2 does not taper narrowly. The basal width of CL2 is 1/2 the basal width of CL1^ in
M. crinoidophilum whereas it is 2/3 the basal width of CL1 in M. ochriderma. The T of M. crinoidophilum has
a longitudinal depression running along the S border and the tergal spur is 2/5 the width of the basal margin,
whilst the T or M. ochriderma has no longitudinal depression and the tergal spur is 1/2 the width of the basal
margin. The caudal appendages of M. crinoidophilum are 8-segmented and are less than 1/2 the height of the basal
segment of the pedicel of cirrus VI. whilst those of M. ochriderma are 7 segmented and equal to the height of the
basal segment of the pedicel of cirrus VI. The mandibular teeth of M. ochriderma are curved and smooth, whereas
those of M. crinoidophilum are not curved and the upper margins of teeth 2 and 3 may have small subsidiary
cusps.
distribution. — East China Sea, Japan, 274-400 m.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
181
Microlasma fragile sp. nov.
Figs 15-18; Tables 7-8
MATERIAL EXAMINED. — Loyalty Islands. Musorstom 6: stn DW 421, 245 m: 3 specimens, ovigerous (large
orange eggs), attached to sponge (MNHN-Ci 2395, Ci 2396). Drawn.
TYPES. — Holotype : MNHN-Ci 2395 (MUSORSTOM 6, stn DW 421).
Paratypes : MNHN-Ci 2396 (Musorstom 6, stn DW 421). Drawn.
Diagnosis. — Shell form upright; fragile, parietal plates thin, weakly articulated. Basal width of CL2 1/2 to
1/3 basal width of CL1; height of CL2 equal to height of CL1. C almost V-shaped. S as wide as high; articular
ridge extending 2/3 length of tergal margin, projecting. Tergal spur 1/2 height of valve, 1/5 width of basal margin,
set at 1/8 length of basal margin from basi-scutal angle; 6-8 muscle crests projecting from basal margin. Mandible
with 3 teeth, margins of teeth smooth. Caudal appendages slightly longer than basal segment of pedicel of CVI.
6-7 segmented.
Fig. 16. — Microlasma fragile sp. nov. Paratype (ovigerous) from Musorstom 6. stn DW 421 (MNHN-Ci 2396):
a. S, internal view; b. S, external view; c, T, external view; d. T. internal view; e, whole animal, from above;
f. whole animal, lateral view.
Description. — Size small (maximum: RC length 5.3, C height 3.0). Shell form upright, rostral plate and
CL1 sloping in toward orifice, apex of CL2 pointing toward C. Parietal plates thin, fragile, weakly articulated;
growth ridges fine, irregular, marked by minute setae. Rostral plate broad, convex; sutures between R and RL
visible internally and externally, dividing plate into 3 subequal parts. CL1 and CL2 separate; CL1 as broad as R.
C deeply concave, apex spout-like, retroverted; higher than and subcqual to width of R. Alae wide, those of C
widest, those of CL1 and CL2 subequal, 1/2 width of carinal ala; superior alar margins finely serrate; alar growth
ridges parallel to inferior alar margin. Basis membranous; flat. Orifice large, pentagonal, toothed. Opercular plates
placed approximately 30° to basis. S flat, thin; triangular, as wide as high, basal margin 7/10 length of occludent
margin; externally with well-spaced, even growth ridges, not convoluted or sinuous, not cut by longitudinal striae;
internally adductor muscle and lateral depressor muscle pits absent; articular furrow narrow; articular ridge
182
D. JONES
extending 2/3 length of valve, projecting well beyond articular margin; adductor ridge absent. T thin; sub-
triangular. smaller than S by 1/4; externally growth ridges fine, some longitudinal strialions in carinal area;
internally with 6-8 fine muscle crests projecting beyond basal margin; articular ridge prominent, extending well
beyond articular margin; articular furrow well-developed; spur roundly truncate, length 1/2 length of basal margin,
set at distance of 1/8 length of basal margin from basi-scutal angle, width 1/5 length of basal margin. Color of
parietal and opercular plates orange-ochre; eggs orange. Measurements of 3 specimens measured as follows:
Labrum with shallow medial depression, with
regular row of small, spaced teeth, some bidentate.
Mandibular palp oblong, narrow; long, serrate setae
terminally. Mandible with 3 teeth, tooth I largest,
well separated from teeth 2 and 3; margins of teeth
smooth; inferior angle moderately large, bluntly
molariform, strongly dentate. Maxillule setose;
2 long, stout setae at upper angle; notch below
upper angle wide, 1/4 length of cutting margin, with
3-4 pairs of smaller setae; stepped cutting margin
below with 3-4 pairs of longer, stout setae; stepped
inferior angle with 4-6 pairs of small setae. Maxilla
moderately wide, coxal endite barely developed; setae
simple and serrulate, long, dense.
Cirrus I with rami unequal; anterior ramus longer
than posterior; proximal segments of anterior
slightly protuberant anteriorly; both rami with
segments thickly setose, setae simple, stout;
postero-distal angles of segments of posterior ramus with small tufts of stout setae, surface with combs of small,
simple setae. Cirrus II longer than cirrus I, similar form to posterior cirri; rami subequal; proximal segments of
anterior ramus broader than those of posterior ramus, slightly protuberant anteriorly, all segments setose,
especially distally, setae simple or serrulate; posterior ramus antenniform, segments becoming elongated and
narrow distally, distal segments with 4-5 pairs of long, finely serrulate setae on anterior faces. Cirrus III similar to
cirri IV-VI; rami subequal, segments becoming oblong distally; posterior ramus with 2-5 pairs of long, finely
serrulate setae on anterior faces of distal segments; anterior ramus with 2-3 pairs of long, finely serrulate setae on
anterior faces distal segments, proximal segments with 5 pairs. Cirrus IV to VI similar, longer than cirrus III;
segments oblong, with 5 pairs of setae on anterior faces, distal 2 pairs longest. Cirral formula as follows:
Fig. 17. — Microlasma fragile sp. nov. Holotype
(ovigerous) from Musorstom 6, stn DW 421 (MNHN-
Ci 2395): lateral view. Scale = 1 cm.
Caudal appendages slightly longer than basal segment of pedicel of CVI, 6-7 segmented; sparse circlet of long
setae terminally; long setae around distal segmental margins, setal length equal to or longer than segmental length.
Penis subequal to length of cirrus VI; annulated; sparsely setose distally; circlet of long setae terminally. Eggs
large, rounded, 0.40 x 0.42 mm.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
183
0.5 mm
Fig. 18. — Microlasma fragile sp. nov. Paratype (ovigerous) from Musorstom 6. stn DW 421 (MNHN-Ci 2396): a. right
mandible; b, left mandible; c, right maxillule; d, left maxillule; e. maxilla; f, labrum; g. cirrus III. posterior ramus,
median segments; h. cirrus 11. posterior ramus, median segments; i, cirrus I. posterior ramus, median segments;
j, cirrus II; k, cirrus VI, caudal appendage and penis; 1. cirrus III; m, cirrus 1.
Remarks. — Microlasma fragile sp. nov. may be distinguished from other members of the genus by its
fragile, thin, weakly articulated parietal plates. The size of M. fragile is similar to that of M. arwetergum
184
D. JONES
(maximum RC length 5.3, 5.0 respectively), but the overall upright shell form and circular basis, rather than a
conical form with the basis elongated along the RC axis, distinguish the 2 species. The orifice of M. arwetergum
is rhomboidal and not toothed rather than pentagonal and toothed as in M. fragile. Furthermore, in M. arwetergum
CL2 is smaller (CL2 basal width 1/4 the basal width of CL1 rather than 1/2 to 1/3 basal width of CL1, and CL2
shorter than CL> by 1/5 rather than equal to height of CL1) and sharply tapering, with the apex so narrow that the
apex of CL1 almost touches C, rather than the height of CL2 being equal to the height of CL1 ( M . fragile). The
unique arrow-shaped T of M. arwetergum further separates the species. There are also differences in the soft parts
between these two species — the maxillule of M. arwetergum is slightly notched, rather than broadly as in
M. fragile, the upper margins of mandibular teeth 2 and 3 have small subsidiary cusps, rather than being smooth;
and the caudal appendages (7-9 segmented) of M. arwetergum are slightly longer than the pedicel of cirrus VI.
rather than slightly longer than the basal segment of the pedicel of cirrus VI (M. fragile).
Microlasma fragile is similar to M. crinoidophilum and M. ochriderma but differs from those species in size
(maximum RC lengths 5.3, 9.0, 7.8, respectively) and in the form and size of CL2 and T. In M. fragile the basal
width of CL2 is 1/2- 1/3 the basal width of CL1 and the height of CL2 is equal to the height of CL1. In
M. crinoidophilum the basal width of CL2 is 1/2 that of CLL and the apices of CL2 taper abruptly, so that the
alae of CL' almost touch the carinal alae; CL2 is shorter than CL1 by 1/5. In M. ochriderma the basal width of
CL2 is 2/3 that of CL1, and CL1 and CL2 are equal in height. The T spur of M. fragile is 1/2 the height of the
valve, its width is 1/5 that of the basal margin and it is set at 1/8 the length of the basal margin from the basi-
sculal angle. The tergal spur of M. ochriderma is short, 1/2 the length of the basal margin and set at a distance of
1/10 the width of the basal margin from the basi-scutal angle. In M. crinoidophilum the short tergal spur is
1/3 the length or the basal margin and is set at a distance of 1/12 the length of basal margin from the basi-scutal
angle. In none ol these species is the tergal basal margin excised to produce an arrow-shape.
ETYMOLOGY. — From the Latin fragilis, "fragile", in reference to the fragile nature of the parietal plates and
the opercular plates.
Distribution. — Loyalty Islands, 245 m.
Microlasma ochriderma (Foster, 1981)
Fig. 15; Tables 7-8
Pachylasma ochriderma Foster, 1981: 354, fig. 5 A-J. — RoSELL, 1991: 37.
Material EXAMINED. — Norfolk Island. NZOI: stn P35, 28°57.9'S. 167°45.5'W. 392-423 m. 28 01 1 977-
holotype (NZOIH-356).
Types. — Holotype: NZOI H-356; Norfolk Island, NZOI stn P 35, 28°57.9'S, 167°45.5'W 392-423 m
28.01.1977.
Holotype depository. NIWA.
Diagnosis. — Shell pale orange, externally smooth, internally with faint, irregular pegs along thickened basal
edge of plates. Basal width of CL2 2/3 basal width of CL1; height of CL2 equal to height of CL1. Alae wide. S as
wide as high, basal margin 2/3 length of occludenl margin; articular ridge extending 1/2 length of tergal margin,
projecting; growth ridges cut by few faint longitudinal striae. T spur short, width 1/2 length of basal margin, set at
distance of 1/10 length of basal margin from basi-scutal angle; 5-7 muscle crests projecting from basal margin.
Mandible with 3 sharp, curved teeth, margins smooth. Caudal appendages as long as basal segment of pedicel of
cirrus VI, 7 segments.
Remarks. The species (RC length 7.8) is distinguished by the dimensions and the manner of articulation of
the opercular plates, and by the orange color of the shell plates. Differences between M. ochriderma
and M. arwetergum are listed under M. arwetergum, between M. ochriderma and M. crinoidophilum under
M. crinoidophilum, and between M. ochriderma and M. fragile under M. fragile.
Source : MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
185
Distribution. — Norfolk Island, 392-423 m.
Genus PACHYLASMA Darwin, 1854 (emend.)
Fig. 19; Tables 3-4. 9-11
Pachylasma Darwin. 1854; 475.
Diagnosis. — Adult with fixed shell pattern of 6 parietal plates (R-CU-CL2-C). Rostral plate compound,
sutures obscure or visible (externally and internally or visible internally, inconspicuous externally), basal width of
R equal to basal width of RL. CL1 and CL2 separate. Length of S basal margin 1/2 or more length of occludenl
margin; externally growth lines smooth or convoluted, often cut by numerous longitudinal striae. Caudal
appendages varying from vestigial or reduced to 1/3 greater than length of pedicel of cirrus VI, from uniarticular to
19 segments.
TYPE Species. — Pachylasma giganteum (Philippi, 1836).
RECENT Species. — Pachylasma bacum sp. nov.; P. darwinianum Pilsbry, 1912; P. ecaudatum Hiro. 1939;
P. giganteum (Philippi, 1836); P. integrirostrum Broch, 1931; P. laeviscutum sp. nov.; P. ovatum sp. nov,;
P. scutistriata Broch, 1922.
Distribution. — Mediterranean Sea; Indian Ocean to central Pacific Ocean: Mediterranean Sea, "deep water"
(P. giganteum)-, Indian Ocean to New Zealand, Japan, South China Sea to Malaysia, 104-2050 m (P. scutistriata)-,
Kci Is, Indonesia, 140 m ( P . integrirostrum Broch. 1931); Japan, 200 m (P. ecaudatum)-, Philippines, 146 m
(P. darwinianum)-, Vanuatu, Loyalty Islands, 182-270 m ( P . bacum)-. New Caledonia. 300-616 m (P. ovatum)-,
Vanuatu, New Caledonia, Norfolk Ridge, Futuna I., 140-466 m (P. laeviscutum).
Fig. 19. — Pachylasma Darwin, 1854. Distribution map: ★, P giganteum ("deep water"); □. P. scutistriata (104-2050 m);
O, P. integrirostrum (140 m); • . P. ecaudatum (200 m); A. P. darwinianum (146 m); O. P. bacum ( 182-270 m);
<0>, P. ovatum (300-616 m); ■ , P. laeviscutum (140-466 m).
186
D. JONES
Remarks. — In the present paper Pachylasma is considered a grouping whose resolution has yet to be
determined. The affinities of the various species included under this genus are as yet unclear, but may be resolved
as more material comes to hand. Differences between Pachylasma , Eutomolasma, Microlasma, Eurylasma and
Tetrapachylasma are listed in tables 3 and 4. The species included herein are small (maximum: RC length 7.0, C
height 2.5), moderate (maximum: RC length 10.5, C height 7.2), and large (maximum: RC length 30.0,
C height 30.5) in size, and occur attached to gorgonians, sponges, bivalves, balanomorph barnacles, crinoids,
antipatharians and coral rubble.
Table 9. — Pachylasma: characters of the species
anic. - articular; b-s = basi-scutal; ext. = external; int. = internal; max. = maximum; n.a. = not applicable; proj. = projecting.
Source MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
187
Table 10. — Pachylasma: characters of the species (continued)
artic. = articular; long. = longitudinal; max. = maximum; proj. = projecting; segs = segments.
Table 11. — Pachylasma : maximum size, geographical and bathymetrical distribution of the species
188
D. JONES
Pachylasma bacum sp. nov.
Figs 19-22; Tables 9-11
MATERIAL EXAMINED. — Loyalty Islands. Musorstom 6: stn CP 401, 270 m: 1 specimen, attached to sponge
(MNHN-Ci 2503). Drawn.
Vanuatu. Musorstom 8; stn CP 1086, 182-215 m: 1 specimen (MNHN-Ci 2397).
Types. — Holotype : MNHN-Ci 2397 (Musorstom 8, stn CP 1086).
Paratype : MNHN-Ci 2503 (MUSORSTOM 6, stn CP 401). Drawn.
DIAGNOSIS. — Rostral plate broad, sutures obscure. Basal width of CL1 equal to basal width of CL2. S with
externally convoluted growth ridges cut by 10-14 radial striations; articular ridge slightly projecting. T spur short,
almost indistinguishable from basal margin; 11-15 muscle attachment ridges extending well beyond basal margin;
articular ridge barely projecting. Caudal appendages reduced, leaf-like.
FlG. 20. — Pachylasma bacum sp. nov. Paratype from MUSORSTOM 6, stn CP 401 (MNHN-Ci 2503): a, T, external view;
b, T, internal view; c, S, external view; d, S, internal view; e. whole animal, from above; f. whole animal, lateral
view.
DESCRIPTION. — Shell large (maximum: RC length 14.0, C height 7.0); form circular. Parietal plates with
irregular growth ridges not well-defined, except on rostral plate. Rostral plate slightly broader than C, sutures
obscure. Basal width of CL1 equal to basal width of CL2, 1/2 to 1/3 basal w'idth of rostral plate. C w ider than CL1
and CL2, narrower than rostral plate, height equal to or greater than rostral plate; plate sloping tow'ard orifice; apex
acute. Alae wide, triangular; growth ridges w'ell-developed, parallel to inferior alar margin; superior alar margins
oblique, not serrate; alae of C widest, alae of CL1 and CL2 equal in size, 1/2 width of carinal alae. Basis
calcareous. Orifice small, pentagonal, slightly toothed; apex of rostral plate curving in toward orifice, apices of
Sour ce : MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
189
CL1 and CL- sloping toward C. Opercular plates set at angle of approximately 20° to basis. S triangular, apex
acute, basal margin convex, 7/12 length of occludent margin; externally growth ridges extremely convoluted, cut
by 10-14 radial striations, giving 'beaded' appearance; internally with adductor muscle pit approximately medial;
lateral depressor muscle pit indistinct; articular furrow narrow; articular ridge extending 1/2 length of valve,
projecting slightly beyond articular margin; adductor ridge indistinct. T larger than S. triangular; as wide as high;
externally with prominent, regular growth ridges, few radial striations in carinal area; internally with 11-15 strong
muscle attachment ridges extending beyond basal margin; articular ridge barely projecting beyond articular margin;
articular furrow broad; spur short, almost indistinguishable from basal margin, 1/2 width of basal margin, set at
distance of 1/10 length of basal margin from basi-scutal angle. Color of parietal plates orange-yellow with rusty
orange markings; alae creamy yellow, those of C, CL> and CL2 with rusty orange markings. Measurements of
2 specimens as follows:
Labrum with shallow medial depression, small
teeth absent. Mandibular palp broadly ovate; dense,
serrate setae terminally. Mandible with 3 teeth, tooth
1 largest, separated from teeth 2 and 3; upper
margins of 3 or upper margins of 2 and 3 often with
small subsidiary cusps, 2 sometimes with
subsidiary, smaller tooth distally; inferior angle
large, bluntly molariform, dentate. Maxillule with
pair of long, stout setae at upper angle; notch below
upper angle wide (right maxillule of illustrated
specimen with notch barely defined), with 3-5 pairs
of smaller setae; straight cutting margin below with
6-8 pairs of larger setae; inferior angle protuberant,
with 5-6 pairs of smaller setae. Maxilla wide, coxal
endite well developed; long serrulate setae terminally.
Cirrus 1 with rami subequal; anterior ramus
slightly longer than posterior; proximal segments of
anterior ramus anteriorly protuberant; both rami with
segments thickly setose. Cirrus II longer than cirrus I; rami subequal, anterior ramus shorter than posterior;
proximal segments of anterior ramus anteriorly protuberant; posterior ramus distally antenniform, small spines
along distal margins of segments; all segments setose, setae simple or serrulate, especially distally. Cirrus III
longer than cirrus II, with rami subequal, segments becoming oblong distally; posterior ramus with 3-5 pairs of
long, finely serrulate setae on anterior faces of distal segments; anterior ramus with 3-5 pairs of long, finely
serrulate setae on anterior faces of more distal segments, proximal segments with 8 pairs. Cirrus IV to VI similar,
longer than cirrus III; segments oblong, with 4-5 pairs of setae on anterior faces, distal 2 pairs longest, finely
serrulate. Chaetotaxy ctcnopod. Cirral formula as follows:
^ Caudal appendages reduced, leaf-like, with small, stout setae apically. Penis equal to length of basal segment of
pedicel of cirrus VI; sparsely setose, with circlet of short setae distally.
FtG. 21. — Pachylasma bacum sp. nov. Paratype from
Musorstom 6, stn CP 401 (MNHN-Ci 2503): lateral
view. Scale = 5 mm.
190
D. JONES
Fig. 22. —Pachylasma bacum sp. nov. Paralype from MUSORSTOM 6, stn CP 401 (MNHN-Ci 2503): a. right mandible;
b. left mandible; c. right maxillulc: d. left maxillule; e, labrum; f, left maxilla; g. mandibular palp; h. cirrus VI and
penis; i. cirrus I, posterior ramus, median segments; j. cirrus II, posterior ramus, median segments; k. cirrus I.
I, cirrus Ill; m. cirrus II.
Remarks. — The circular form of the specimen from the Loyalty Islands is modified to an elongated oval, due
to its attachment position on the host sponge. Pachylasma bacum sp. nov. is similar to P. ovatum sp. nov., but
the 2 may be separated by the presence or absence of rostral plate sutures, visible in P . ovatum , obscure in
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
191
P. bacum. There are also differences in the relative sizes of the parietal plates - in P. bacum the basal width of
CL1 is equal to the basal width of CL2 and 1/2 to 1/3 the width of the rostral plate; in P. ovation the basal width
of CL2 is 7/8 the basal width of CL' and is equal to the basal width of the rostral plate. The basis is calcareous in
P. bacum but is membranous with calcareous edges in P. ovatum. In P. bacum the S articular ridge is 1/2
the length of the valve and slightly projects beyond the articular margin; in P. ovatum it is 3/4 the length of
the valve and projects beyond the articular margin. The tergal articular ridge of P. bacum barely projects beyond
the articular margin; in P. ovatum it is short, well-developed and prominent, projecting beyond the articular
margin. The caudal appendages are reduced and leaf-like in P. bacum, and absent in P. ovatum. The color of
P. bacum is orange-yellow with rusty orange markings; P. ovatum is a dirty creamy white color.
The new species is similar to P. ecaudatum Hiro, 1939. but there arc the following differences: the caudal
appendages are reduced and leaf-like, with small, stout setae apically in P. bacum and absent in P. ecaudatum ;
the basis is entirely calcareous in P. bacum rather than calcareous, thin and centrally membranous (P. ecaudatum );
cirral counts for cirri I1I-V1 are greater in P. bacum (CII1 21/19, CIV 22/23, CV 24/24, CVI 24/25) than in
P. ecaudatum (CIII 16/16, CIV 17/17, CV 18/18, CVI 20/20) and the colour of P. bacum is orange-yellow with
rusty orange markings, whereas in P. ecaudatum the parietal plates are whitish, mottled with pink reticulations and
the alae are uniformly rosy. Pachylasma bacum is larger in size (RC diameter 14.0, C height 7.0) and found
attached to sponges, whilst Pachylasma ecaudatum (RC diameter 10.0, C height 6.0) has been found attached to
the bivalve Malleus regulus (Forsk&l, 1775).
ETYMOLOGY. — From the Latin bacum, "bead", in reference to the beaded appearance of the scuta.
Distribution. — Loyalty Islands, 270 m; Vanuatu, 182-215 m.
Pachylasma darwinianum Pilsbry. 1912
Fig. 19; Tables 9-11
Pachylasma darwinianum Pilsbry. 1912: 293; 1916: 329. — Newman & Ross. 1976: 40. — Rosell, 1991: 37.
MATERIAL EXAMINED. — Philippines. " Albatross stn 5168. Sulu Arch., Tawi Tawi Group, 4°56'30"N.
1 19°45'40"E, 146 m, 25.02.1908: holotype (USNM 43465).
Types. — Holotype : USNM 43465; Philippines, "Albatross", stn 5168, Tawi Tawi Group, Sulu Arch..
146 m; dry (I lot) and slides (3).
Holotype depository. USNM.
Diagnosis. — Rostral plate with sutures visible internally but not externally. Basal width of CL2 more than
1/2 basal width of CL1. S growth ridges strongly convoluted, not cut by longitudinal striae; articular ridge barely
projecting. T spur absent, 8-10 muscle crests projecting beyond basal margin; articular ridge slightly projecting.
Mandible with upper margins of teeth 1 and/or 2 often with small subsidiary cusps. Caudal appendages
uniarticulate, represented by swollen mounds with few. minute bristles.
Remarks. — Pachylasma darwinianum is a large species (maximum RC length 20.5). It is most similar to
P. bacum sp. nov. and P. ecaudatum, as all have extremely reduced caudal appendages which are uniarticulate.
In P. darwinianum and P. ecaudatum the caudal appendages are represented by swollen mounds with a few, minute
bristles apically; in P. bacum they are leaf-like, with small, stout setae apically. In P. darwinianum the basal
width of CL> is greater than 1/2 the basal width of CL2, in P. bacum it is equal to the basal width of CL2,
in P. ecaudatum it is equal to or slightly less than the basal width of CL2. The basis of P. darwinianum is
membranous, that of P. bacum is calcareous and that of P. ecaudatum is membranous centrally, with calcareous
edges. The T spur is absent in P. darwinianum and there are 8-10 muscle attachment ridges projecting well beyond
the basal margin; in P. bacum the spur is short. 1/2 the width of the basal margin and set at 1/10 the length of
t e basal margin from the basi-scutal angle, and there are 11-15 muscle attachment ridges projecting well beyond
192
D. JONES
the basal margin; in P. ecaudatum it is indistinct, 3/8 the width of the basal margin and set at 1/8 the length of
the basal margin from the basi-scutal angle, and there are 8-11 muscle attachment ridges projecting well beyond
the basal margin. The dull red to whitish color of the parietal plates further distinguish P. darwinianum from all
others in the genus.
Distribution. — Tawi Tawi Group, Sulu Archipelago, Philippines, 146 m.
Pachylasma ecaudatum Hiro, 1939
Fig. 19; Tables 9-11
Pachylasma ecaudatum Hiro, 1939: 52, figs 3-4. — Newman & Ross, 1976: 40.
Hexelasma ecaudatum - UTINOMI, 1968: 31, fig. 6.
TYPES. — Holotype: SMBL Type 30; Japan, Ogasawara I., 200 m, associated with bivalves ( Malleus regains
Forskal, 1775).
Holotype depository. SMLB (specimen has not been located).
DIAGNOSIS. — Rostral plate lowest of all plates, sutures obscure. Basal width of CL2 equal to or narrower than
basal width of CL1. S externally with convoluted growth ridges cut by 8 longitudinal striae; articular ridge not
projecting. T articular ridge not projecting; externally with growth ridges prominent, spur indistinct, 3/8 width of
basal margin; 8-1 1 muscle attachment ridges extending well beyond basal margin. Caudal appendages uniarticulate,
represented by few minute bristles on swollen mounds.
REMARKS. — Pachylasma ecaudatum is a moderate sized species (maximum: RC length 10.0. C height 6.0).
The thin calcareous basis which is centrally membranous, the form of the T, the reduction of the caudal appendages
and the whitish parietal plates, mottled with pink reticulations, determine this species. Pachylasma ecaudatum is
most similar to P. baettm sp. nov.; differences between these 2 species are listed under P. bacum.
Distribution. — Ogasawara Is, Japan, 200 m.
Pachylasma giganteum (Philippi. 1836)
Fig. 19; Tables 9-11
Chthamalus giganteus Philippi, 1836: 250.
Pachylasma giganteum - Darwin. 1854: 477. pi. 19 figs 5 a-d. — Weltner, 1897: 273. — Gruvel, 1905: 198. fig. 218.
— Pilsbry, 1916: 329. — Kolosvary. 1942: 143; 1943: 77; 1952: 412. — Withers. 1953: 60. — Relink 1969:
169. — Stubbings, 1967: 263. — Newman & Ross, 1976: 40.
Types. — Holotype: BMNH Cat. No. 12060 Seguenza, in Jeffreys collection; Strait of Messina.
Holotype depository : BMNH. [specimen has not been located in the collections of (he BMNH (Ms A.
Morgan, pers. Comm. 05.02.1996)].
DIAGNOSIS. — Rostral plate sutures obscure. CL1 and CL2 equal sized. S with prominent, sinuous growth
ridges cut by 4-6, longitudinal striae; articular ridge not projecting. T with growth ridges prominent; spur short,
1/2 width of basal margin; 8-10 muscle attachment ridges projecting well beyond basal. Caudal appendages
1/3 longer than pedicel of cirrus VI. 19 segments.
REMARKS. — This species (maximum: RC length 30.0, C height 30.5) is distinguished from all others in
the genus by its large size, by the dirty white color of the parietal plates and the form of the opercular plates.
Pachylasma giganteum is similar to P. bacum as both species have a calcareous basis; the basis of P. giganteum
is of variable thickness. Differences between P. giganteum and P. bacum are listed under P. bacum.
Distribution. — Sicily, Mediterranean Sea; “deep water" (Darwin, 1854).
Source : MNHN. Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
193
Pachylasma integriroslrum Broch, 1931
Fig. 19; Tables 9-11
Pachylasma integriroslrum Broch, 1931: 50, fig. 18. — Newman & Ross, 1976: 40.
MATERIAL EXAMINED. — Indonesia. Danish Exped TO Kei Is 1922: Mollucas, Amboina Bay. 140 m,
22.02.1922: syntypes (ZMC Cru 1977).
TYPES. — Syntypes: ZMC Cru 1977; Indonesia, Mortensen s PACIFIC EXPED. 1914-16. Kei Islands,
Amboina Bay, 140 in, 22.02.1922; 4 specimens.
Syntypes Location: ZMC.
Diagnosis. — Parietal plates covered by persistent yellowish/brownish epidermis. Rostral plate sutures
obscure. Basal width of CL2 almost 1/2 basal width of CL1. S growth ridges feeble, radial striations absent;
articular ridge slightly projecting. T with growth ridges feeble, smooth, not cut by longitudinal striae; spur
indistinct, 1/2 width of basal margin; 4-6 weak muscle crests not extending beyond basal margin; S side of spur
marked by shallow groove; articular ridge projecting. Caudal appendages equal to height of basal segment ol
pedicel of cirrus VI; 6 segments.
REMARKS. — In his description of Pachylasma integrirostrum from Ambon. BROCH (1931: 50) referred to
the development of the alae of CL2 (determined as CL by Broch) as "....subject to variations: in three specimens
- among them also in the type specimen (fig. 18a) they are externally visible only in the uppermost part ot
the plate, but in the fourth they arc almost as conspicuous as those of the lateral plates". After examining
the 4 specimens of P. integrirostrum described by BROCH, the alae of CL2 of the 3 intact specimens are
conspicuous, especially when the small size of the specimens is taken into account. The alae of CL2 are
approximately 1/2 the width of the alae of CL1, and extend most of the height of the shell. The fourth specimen,
labelled 'type', is disarticulated, but was illustrated whole by BROCH prior to disarticulation. BROCH illustrated a
small CL2 ala (BROCH, 1931, fig. 18a). By inference from BROCH's written description, similar small-sized CL2
alae should also be found in 2 of the 3 remaining whole specimens. However, in these 3 intact specimens I regard
the CL2 alae as conspicuous.
This small species (maximum: RC length 7.0, C height 2.5) differs from all others presently included in
Pachylasma by 2 features; the basal width of CL2 is almost 1/2 the basal width of CL1, rather than equal or
subequal, as in the other species within Pachylasma ; and the S external growth lines are feeble, not sinuous or
convoluted, and not cut by longitudinal striae. In the remaining species within Pachylasma the S external growth
lines are sinuous or convoluted, and cut by longitudinal striae, which may be faint in some species (e.g.
P. laeviscutum sp. nov.).
Pachylasma integrirostrum is similar to P. bacum, P. ecaudatum, and P. giganteum as they all have a discrete,
rather than tripartite, rostral plate. However, the basis of P. bacum and of P. giganteum is calcareous, rather than
calcareous at the edges and membranous centrally, as in P. integrirostrum and P. ecaudatum. The S external growth
lines of P. ecaudatum are convoluted and cut by 8 longitudinal striae; those of P. integrirostrum are feeble, not
sinuous or convoluted, and not cut by longitudinal striae. These latter 2 species can be further separated by
the caudal appendages - uniarticulate and extremely reduced being represented by swollen mounds with a few
minute bristles in P. ecaudatum, 6-segmented and the height of the basal segment of the pedicel of cirrus VI in
P. integrirostrum.
Distribution. — Ambon, Mollucas, Indonesia; 140 m.
Pachylasma laeviscutum sp. nov.
Figs 19, 23-25; Tables 9-11
Material examined. — Norfolk Ridge. Smib 5: stn DW 85. 260 m: 1 specimen (broken and lacking R). —
Stn DW 88. 350 m: 1 specimen (dry), attached to gorgonian.
194
D. JONES
New Caledonia. MUSORSTOM 4: stn CP 190, 215 m: 1 specimen (shell only), attached to stem of dead gorgonian.
— Stn DW 230. 390-420 m: 3 specimens, attached to stem of dead gorgonian (MNHN-Ci 2399).
Vanuatu. MUSORSTOM 8: stn DW 962, 370-400 m: several specimens, attached to gorgonian (WAM C 23249). —
Stn CP 963, 400-440 m: 1 specimen. — Stn DW 988, 372-466 m: several specimens, attached to gorgonian. —
Stn DW 1131, 140-175 m: several specimens, attached to gorgonian.
Futuna Island. MUSORSTOM 7: stn CP 508. 245-440 m: 3 specimens, attached to stem of dead gorgonian (MNHN-Ci
2697). — Stn CP 515, 224-252 m: 2 specimens, attached to stem of gorgonian (MNHN-Ci 2398, WAM 253-96). Drawn.
Types. — Holotype: MNHN-Ci 2398 (MUSORSTOM 7, stn CP 515).
Paratypes : MNHN-Ci 2399 (MUSORSTOM 4, stn DW 230). — MNHN-Ci 2697 (MUSORSTOM 7, stn CP 508).
— WAM 253-96 (MUSORSTOM 7, stn CP 515). Drawn.
Diagnosis. — Rostral plate with basal width R equal to basal width of RL. Basal width of CL1 subequal to
basal width of CL2. S externally with growth ridges slightly sinuous, cut by traces of 2-3 slight radial striations;
S articular ridge slightly projecting. T spur indistinct, more than 1/3 length of basal margin; 13-18 muscle
attachment ridges extending slightly beyond basal margin; articular ridge projecting. Caudal appendages equal to
height of basal segment of pedicel of cirrus VI, 8-9 segments.
Fig. 23. — Pachylasma laeviscuium sp. nov. Paratype from Musorstom 7. stn CP 508 (MNHN-Ci 2697): a. S. external
view; b. S. internal view; c, T. internal view; d. T, external view; e. whole animal, lateral view; f. whole animal,
from above.
Description. — Size moderate (maximum: RC length 10.5, C height 7.2). Shell form upright, parietal plates
gently sloping toward orifice. Parietal plates smooth, growth ridges regular, faint, not marked by setae, in larger
specimens paries more rugged, growth ridges less regular. Tripartite rostral plate convex, basal width twice basal
width of C; rostral sutures visible externally and internally, basal width of R equal to basal width of RL. CL' and
CL- separate, basal width of CL1 subequal to basal width of CL2. C higher than rostral plate, basal width 1/2
basal width of rostral plate. Alae wide, those of C widest, alae of CL' 1/2 width of C alae, alac of CL2 1/4 width
Source : MNHN, Paris
CIRRIPED1A THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
195
of C alae; growth ridges parallel to inferior alar margins, well-marked, summits oblique. Orifice large, pentagonal,
not toothed. Basis thin, calcareous at edges, membranous centrally; flat. Opercular plates sloping at angle of
approximately 45° to basis. S subequal to T; elongated triangle, basal margin 1/2 length occludent margin;
externally growth ridges slightly sinuous, not fluted, cut by traces of 2-3 slight radial striations; internally adductor
muscle pit situated at 1/2 length of valve; lateral depressor muscle pit indistinct; articular furrow narrow; articular
ridge extending 2/3 length of valve, projecting slightly beyond articular margin; adductor ridge indistinct.
T subtriangular; width equal to height; externally growth ridges prominent, angularly upturned close along sculal
margin, longitudinal striations in carinal area over depressor muscle region; internally with 13-18 small muscle
crests projecting beyond basal margin; articular ridge long, slightly extending heyond articular margin: articular
furrow broad, shallow; spur short, truncate, width more than 1/3 length of basal margin, set at distance of 1/9
length of basal margin from basi-scutal angle. Color of parietal plates orange with some rusty-red markings.
Measurements of 10 specimens examined as follows:
Labrum with shallow medial depression; teeth
absent but irregular groups of minute denticles
present. Mandibular palp ovate, elongated; long
serrate setae terminally. Mandible with 3 primary
teeth, tooth 1 largest, well separated from 2 and 3;
upper margins of 2 and 3 with small subsidiary
cusps; inferior angle large, bluntly molariform,
dentate. Maxillule setose; 3 long setae at upper
angle; indistinct notch below with 2-3 pairs of
smaller setae; cutting margin below notch slightly
stepped, with 7-8 pairs of longer setae; inferior angle
slightly stepped, with 4-7 pairs of smaller setae.
Maxilla wide, with dense, long setae; coxal endite
more developed than basal endite.
Cirrus I with rami unequal; anterior ramus longer
than posterior; proximal segments of anterior ramus
protuberant anteriorly; both rami with segments
thickly setose. Cirrus II longer than cirrus I. similar
to cirrus III; rami subequal; proximal segments of anterior ramus broad, slightly protuberant anteriorly, all
segments thickly setose; posterior ramus antenniform, segments elongated, narrow, distal segments with 2-3 long
setae antero-distally. Cirrus III more similar to cirri IV-VI than to cirri I and II; rami subequal. segments becoming
oblong distal ly, more distal segments of both rami with 4 pairs of long setae on anterior faces. Cirrus IV to VI
similar, longer than cirrus III; segments oblong, with 4 pairs of setae on anterior faces, distal 2 pairs of setae
longest. Chaetotaxy ctenopod. Cirral formula as follows:
Fig. 24. — Pachylasma laeviscutum sp. nov. Holotype
from Musorstom 7. stn CP 515 (MNHN-Ci 2397):
lateral view. Scale = 1 cm.
Caudal appendages length of proximal segment of pedicel of cirrus VI; 8-9 segmented; long setae terminally
and around distal segmental margins. Penis more than 1/2 length of cirrus VI; finely annulated; sparsely setose,
with dense rosette ol moderately long, fine setae distally.
196
D. JONES
Fig. 25. — Pachylasma laeviscutum sp. nov. Paratype from Musorstom 7, stn CP 508 (MNHN-Ci 2697): a, right
mandible; b. left mandible; c, right maxillule; d. left maxillule; e, maxilla; f. cirrus I; g, cirrus II; h, cirrus III;
i, pedicel of cirrus VI, caudal appendage and penis; j, cirrus VI. posterior ramus, median segments: k. mandibular
palp; 1, cirrus III, posterior ramus, median segments; m. labrum.
Remarks. Pachylasma laeviscutum sp. nov., P. ovatum sp. nov. and P. scutistriata are similar because all
have the rostral sutures visible externally. However, there are differences in the form and length of the caudal
appendages, which are 8-9 segmented and the length of the basal segment of the pedicel of cirrus VI in
Source : MNHN, Paris
C1RRIPED1A THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
197
P. laeviscutum. ; absent in P. ovatum; and 15-18 segmented and a little longer than the pedicel of cirrus VI in
P. scutistriata. There are also differences in the ornamentation of the S - the scutal growth ridges are slightly
sinuous and cut by 2-3 slight radial striations in P. laeviscutum: moderately convoluted and cut by 12-15 radial
striations in P. ovatum: and convoluted and distinctly divided by many radial striations in P. scutistriata. Features
of the T also separate these species. In P. laeviscutum the indistinct tergal spur is more than 1/3 the width of
the basal margin, set at distance of 1/9 the length of the basal margin from the basi-scutal angle. 13-18 muscle
attachment crests slightly extend from the basal margin, and the articular ridge slightly projects beyond
the articular margin. In P. ovatum the short spur is more than 2/5 the width of the basal margin, set at distance of
less than 1/5 the length of the basal margin from the basi-scutal angle, 12-14 muscle attachment crests extend well
beyond the basal margin, and the articular ridge projects beyond the articular margin. In P. scutistriata the spur is
indistinct, not separated from the basi-scutal angle, the muscle attachment crests do not extend well beyond
the basal margin, and the articular ridge projects beyond the articular margin.
Pachylasma laeviscutum sp. nov. differs from P. darwinianum as the rostral sutures are visible internally
but not externally in the latter species. Additionally, P. darwinianum has a membranous basis, and the caudal
appendages are extremely reduced, being represented by swollen mounds, whereas P. laeviscutum has
8-9 segmented caudal appendages which are the height of the basal segment of the pedicel of cirrus VI. The S of
P. darwinianum externally has strongly convoluted growth lines which are not cut by longitudinal striae, whilst
those of P. laeviscutum are slightly sinuous and cut by traces of 2-3 faint longitudinal striations. The T spur of
P. laeviscutum is more than 1/3 the length of the basal margin, whereas the spur is absent in P. darwinianum.
Pachylasma laeviscutum differs from P. ecaudatum, P. bacum sp. nov. and P. giganteum because in these latter
species the rostral sutures are obscure. The form and length of the caudal appendages (equal to the length of
the basal segment of the pedicel of cirrus VI in P. laeviscutum, extremely reduced in P. ecaudatum and P. bacum,
and 1/3 longer than the pedicel of cirrus VI in P. giganteum ) further separate these species. There arc also
differences regarding the form of S and T between these species. The scutal growth ridges arc slightly sinuous, not
fluted, and cut by traces of 2-3 slight radial striations in P. laeviscutum: the S of P. ecaudatum has convoluted
growth ridges cut by 8 radial striations; the S of P. bacum sp. nov. has extremely convoluted growth ridges, cut
by 10-14 radial striations, giving it a 'beaded' appearance; and the S of P. giganteum has prominent, sinuous
growth ridges cut by 4-6 slight longitudinal furrows. The T of P. laeviscutum has prominent growth ridges which
are angularly upturned close along the scutal margin, longitudinal striations in the carinal area over the depressor
muscle region, and a short spur more than 1/3 the length of the basal margin and set at 1/9 the length of the basal
margin from the basi-scutal angle. The T of P. ecaudatum has prominent growth ridges and a slight ridge parallel
to the S margin where the growth ridges are angularly upturned, and an indistinct spur which is 3/8 the length of
the basal margin and set at 1/8 the length of the basal margin from the basi-scutal angle. The T of P. bacum
sp. nov. has prominent, regular growth ridges and a short, broad spur which is 1/2 the width of. and almost
indistinguishable from, the basal margin, set at distance of 1/10 the length of the basal margin from the basi-scutal
angle. The T ol P. giganteum has prominent growth ridges, which are angularly upturned close along the scutal
margin, and a short, broad spur which is 1/2 the width of the basal margin and set at a distance of 1/7 the length of
the basal margin from the basi-scutal angle.
Pachylasma laeviscutum sp. nov. differs from P. integrirostrum as in the latter the basal width of CL1 is
almost twice the basal width of CL2, rather than equal to the basal width of CL2. The S growth ridges of
P. integrirostrum are feeble with no radial striations, whilst those of P. laeviscutum are slightly sinuous and cut
by 2 to 3 faint longitudinal striae. The T of P. integrirostrum has feeble growth ridges, a broad spur which is 1/2
the width of the basal margin and 4-6 muscle attachment ridges which do not extend beyond the basal margin.
The T ol P. laeviscutum has well marked growth ridges, a broad spur which extends more than 1/3 the width of
the basal margin and 13-18 muscle attachment ridges which extend beyond the basal margin.
ET"i MOLOCY. From scutum, and the Latin laevis, ‘smooth', in reference to the smooth, slightly sinuous
scutal growth ridges.
Distribution. — Vanuatu, 140-466 m; New Caledonia, 215-420 m; Norfolk Ridee, 260-350 m; Futuna
Island, 224-440 m.
198
D. JONES
Pachylasma ovatum sp. nov.
Figs 19, 26-28; Tables 9-11
MATERIAL EXAMINED. — New Caledonia. Lagon: sin 444, 300-350 m: 1 specimen (MNHN-Ci 2401). Drawn.
Musorstom 4: stn CP 193, 415 m: 1 specimen, attached to a sponge (MNHN-Ci 2400); 1 specimen, attached to coral
rubble.
Bathus 4: stn DW 914, 600-616 m: 2 specimens, attached to sponges. — Stn DW 924, 344-360 m: 1 specimen,
attached to coral rubble (WAM 254 -96).
Types. — Holotype: MNHN-Ci 2400 (MUSORSTOM 4, stn CP 193).
Paratypes: MNHN-Ci 2401 (LAGON, stn 444). Drawn. — WAM 254-96 (Bathus 4, stn DW 924).
DIAGNOSIS. — Rostral plate sutures visible, basal width of R equal to basal width of RL. Basal width of CL2
7/8 basal width of CL1. S growth ridges convoluted, cut by 12-15 radial striations; articular ridge projecting.
T with regular growth ridges; spur almost indistinguishable from basal margin, width 2/5 length of basal margin;
12-14 muscle attachment ridges extending well beyond basal margin; articular ridge projecting. Caudal appendages
absent.
Fig. 26. — Pachylasma ovatum sp. nov. Paratype from Lagon, stn 444 (MNHN-Ci 2401): a. S. internal view;
b, S, external view; c, T. internal view; d. T, external view; e, whole animal, from above (right S and T removed);
f. whole animal, lateral view.
DESCRIPTION. — Size large (maximum: RC length 14.6, C height 10.7). Shell form oval from above. Parietal
plates thick, growth ridges not well-defined, irregular: apices of rostral and CL1 plates curved in toward orifice.
Rostral plate convex; sutures visible internally and externally (sometimes obscure externally), dividing plate into
3 subequal parts; apex curved in toward orifice. CL1 and CL2 separate; basal width of CL1 equal to basal width of
rostral plate; basal width of CL2 7/8 basal width of CL1; apices of CL1 and CL2 curved in toward orifice.
C convex; apex curving out from orifice; basal width of C equal to basal width of CL1 and of rostral plate; shorter
than, or approaching height of. R. Alae wide, triangular, those of C widest, alae of CL1 and CL2 equal in size,
Source . MNHN, Paris
CIRR1PEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
199
width 1/2 width of carinal ala; superior alar margins finely serrate, oblique; alar growth ridges parallel to inferior
alar margin. Basis thin, calcareous, centrally membranous. Orifice large, wide, roundly pentagonal; not toothed.
Basis calcareous at edges, membranous centrally. Opercular plates lodged subparallel to basis. S triangular,
elongated, basal margin greater than 1/2 length of occludent margin; apex bluntly pointed; externally with
convoluted growth ridges, cut by 12-15 radial striations; internally adductor muscle pit in middle of valve; few
short crests for lateral depressor muscle in shallow pit; articular furrow narrow, shallow; articular ridge running 3/4
length of valve, extending beyond articular margin; adductor ridge indistinct. T similar size to S, wide, triangular;
externally growth ridges regular; internally with 12-14 strong muscle crests projecting beyond basal margin;
articular ridge short, well-developed, prominent, extending beyond articular margin; articular furrow well-developed;
spur short, almost indistinguishable from basal margin, width 2/5 width of basal margin, set at distance of 1/5
width of basal margin from basi-scutal angle. Color of parietal and opercular plates dirty creamy white.
Measurements of 4 specimens examined as follows:
Labrum with shallow medial depression, small
teeth present, bi- or tricuspid. Mandibular palp wide,
ovate; dense, serrate setae terminally. Mandible with
3 teeth, tooth 1 separated from teeth 2 and 3; upper
margins of 2 and 3 with small subsidiary cusps;
inferior angle molariform, strongly dentate. Maxillule
with 1 pair of long, stout setae at upper angle; notch
below upper angle broad, with 4-5 pairs of smaller
setae; cutting margin below almost straight, with 5-7
pairs of longer, stout setae; inferior angle slightly
protuberant, with 7-9 pairs of small setae. Maxilla
wide; lobes subequal; long serrulate setae distally.
Cirrus I with rami subequal; anterior ramus
slightly longer than posterior; proximal segments of
anterior ramus anteriorly protuberant; both rami with
segments moderately setose. Cirrus II similar to but
longer than cirrus I; rami subequal; proximal
segments of anterior ramus anteriorly slightly
protuberant, all segments setose, setae simple or serrulate, serrulate setae distally; antenniform distal segments of
posterior ramus with tuft of setae and spines at posterior distal margins. Cirrus III longer than cirrus II; rami
subequal, segments becoming oblong distally; posterior ramus with 3-5 pairs of long, finely serrulate setae on
anterior faces of distal segments; anterior ramus with 3-5 pairs of long, finely serrulate setae on anterior faces of
distal segments, proximal segments with 10 pairs. Cirrus IV to VI similar, longer than cirrus III; segments
oblong, with 5 pairs of setae on anterior faces, distal 2 pairs longest, finely serrulate; distal segments of posterior
ramus with spines at distal margin. Chaetotaxy ctenopod. Cirral formula as follows:
Fig. 27. — Pachylasma ovaium sp. nov. Holotype from
Musorstom 4, stn CP 193 (MNHN-Ci 2400): lateral
view. Scale = 5 mm.
Penis 1/2 length ol basal segment of pedicel of cirrus VI; sparsely setose; tufts of long setae distally. Caudal
appendages absent.
200
D. JONES
Fig. 28. — Pachylasma ovation sp. nov. Paratype from Lagon, sin 444 (MNHN-Ci 2401): a. right mandible; b. left
mandible; c, right maxillule; d. left maxillule; e, cirrus 1; f. cirrus II; g. pedicel of cirrus VI and penis; h. cirrus VI.
posterior ramus, median segments; i, labrum; j, cirrus III; k. cirrus VI; 1, maxilla; m. mandibular palp.
Remarks. — Pachylasma ovation sp. nov. is similar to P. scutistriata but they may be separated by a number
of characters. In P. ovation the basal width of CL2 is 7/8 the basal width of CL', rather than equal to the basal
width (P. scutistriata). The caudal appendages are absent in P. ovation rather than 15-18 segmented and a little
longer than the pedicel of cirrus VI (P. scutistriata). In P. ovation the S growth ridges are moderately convoluted
Source : MNHN. Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
201
and divided by 12-15 radial striations; in P. scutistriata they are distinctly convoluted and divided by a few
radial striations. There are also differences in the form of the T - the muscle attachment crests do not project
beyond the basal margin in P. scutistriata, but there are 12-14 well-developed muscle crests projecting well
beyond the basal margin in P. ovatum', the tergal spur is 2/5 the width of the basal margin and set at a distance of
less than 1/5 the width of the basal margin from the basi-scutal angle in P. ovatum, but the spur is indistinct.
1/3 the width of the basal margin and not separated from the basi-scutal angle in P. scutistriata-, and the tergal
articular ridge extends much further beyond the articular margin in P. scutistriata than in P. ovatum. In addition,
the parietal plates of P. scutistriata are pale pink colored with white alae, whilst those of P. ovatum are dirty
creamy white.
Etymology. — From the Latin ovatum, 'oval', in reference to the ovate form of the basis.
Distribution. — New Caledonia. 300-616 m.
Pachylasma scutistriata Broch, 1922
Fig. 19; Tables 9-1 1
Pachylasma scutistriata Broch. 1922: 301. figs 48-50. — Nilsson-Cantell, 1927: 781. — Pope. 1965: 10. — UTINOMI,
1968: 26. figs 4-5. — Newman & Ross, 1976: 40. — Foster, 1978: 77. fig. 47. pi. 9 E-F.
Material EXAMINED. — Australia. " Endeavour 38°12'S. 149°40'E. 256-274 m. 16.09.1914: holotype (ZMC
Cru 1978). — E of East Sister Island. Bass Strait, 104-1 10 m: I dry specimen (AM E 6629). — SE Australia, few
specimens, on alcyonarian stock covered by anemones (AM P 23757).
type S. — Holotype'. ZMC Cru 1978; Australia, "Endeavour", 38°12'S, 149°40'E, 256-274 m. 16.09.1914;
2 slides plus material in alcohol.
Paratypes : AM E 6565; Australia, "Endeavour", 38°12’S, 149°40'E, 256-274 m. 16.09.1914; ZMC Cru 1978.
Holotype depository >: ZMC.
Paratypes depository: ZMC, AM.
Diagnosis. — Rostral plate shortest plate; sutures visible, basal width R equal to basal width RL. Basal width
ol CL1 equal to basal width of CL2. S externally with growth ridges convoluted, cut with many radial striations;
articular ridge slightly projecting. T with faint growth ridges, spur almost indistinguishable from basal margin,
width 1/3 length of basal margin, set at basi-scutal angle; prominent articular ridge projecting beyond scutal
margin; muscle attachment ridges not extending beyond basal margin. Caudal appendages slightly longer than
pedicel of cirrus VI, 1 5- 1 8 segments.
Remarks. — Size large (maximum: RC length 20.0, C height 15.0). Pachylasma scutistriata is similar to
P. ovatum sp. nov. but they may be separated by a number of characters, which are listed in the Remarks section
under P. ovatum sp. nov. The vivid pink color of the parietal plates, and the form of the caudal appendages
(15-18 segmented and a little longer than the pedicel of cirrus VI) separate P. scutistriata from (he remaining
species presently included within Pachylasma.
Distribution. — Indian Ocean to New Zealand, Japan, South China Sea to Malaysia, 104-2050 m.
Genus EURYLASMA gen. nov.
Fig. 29; Tables 3-4, 12-14
Diagnosis. — Adult externally with fixed shell pattern of 6 parietal plates (R-CL'-CL2-C). Rostral plate with
sutures obscure, or visible externally and internally with R reduced to narrow sliver, RL correspondingly well
eveloped, basal width of RL 3 times or 6 times basal width of R. CL1 and CL2 partially coalesced, coalescence
202
D. JONES
incomplete as CL2 retains minute ala. S concave externally; pyramidal or sub-pyramidal in form. Length of caudal
appendages ranging from 1/2 height of distal segment of pedicel of cirrus VI to longer than pedicel of cirrus VI
by 1/4; 5-13 segments.
Type Species. — Eurylasma angustum sp. nov.
Recent Species. — Eurylasma angustum sp. nov.; E. ferulum sp. nov.; E. pyramidale sp. nov.
Tables 12 and 13. — Eurylasma: characters of ihe species
b-s - basi-sculal; exl. = exiemal; ini. = internal: n. a. = not applicable: proj. = projecting.
artic. - articular; c.a. - caudal appendage; long. = longitudinal; max. = maximum: proj. = projecting; segs = segments.
Tables 14. — Eurylasma : maximum size, geographical and bathymetrical distribution of the species
Remarks. — The well developed carinal alae, and the partial coalescence of CL> and CL2, with the
concomitant reduction of the ala of CL2, are characteristic of the genus. In addition, the angle of lodgement of the
Source . MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
203
opercular plates in the orifice, and the curved form of the S, due to its external concavity, plus the almost right-
angle formed by the basal and tergal margins (resulting in the sub-pyramidal to pyramidal form), are unique
characters. Furthermore, in species where the rostral sutures are visible, the reduction of R to a narrow sliver,
associated with the great development of RL, are characters not found in any other pachylasmatine genera. Species
included herein are moderate (maximum: RC length 9.7, C height 7.5) to large (maximum: RC length 15.0,
C height 10.6) in size and have been found attached to gorgonians and Corallium species.
In Eutomolasma sp. nov., Microlasnia sp. nov. and Pachylasma , CL1 and CL2 are discrete entities, resulting in
the adult fixed wall pattern of 6 plates. CL' and CL2 are coalesced to a certain degree in Eurylasma sp. nov.
although the ala of CL2 is still evident, albeit reduced. This arrangement also results in an adult fixed wall pattern
of 6 parietal plates. In Tetrapachylasma, CL> and CL2 may be coalesced, partially or totally, resulting in
a variable shell pattern in the adult of 6 (partial coalescence of CL1 and CL2) or 4 (total coalescence of CL1 and
CL2) parietal plates.
ETYMOLOGY. — From the Greek eurys, 'wide’, in reference to the characteristically broad carinal alae of
species included in this genus.
Distribution. — Western Pacific Ocean: Norfolk Ridge, 255-365 m (£. pyramided e)\ Norfolk Ridge, 460-
470 m (E. ferulum)-, Vanuatu, Loyalty Islands. 295-400 m (E. angustum).
Fig. 29. — Eurylasma gen. nov. Distribution map: O, K pyramidale sp. nov. (255-365 in); ▲. E. ferulum sp. nov. (460-
470 m); ■. E. angustum sp. nov. (295-400 m).
Eurylasma angustum sp. nov.
Figs 29-32; Tables 12-14
Material EXAMINED. — Loyalty Islands. MUSORSTOM 6: sin DW 478. 400 m: several specimens, attached to
gorgonian (MNHN-Ci 2402, Ci 2504). Drawn.
Si nwi*o<AU °RST0M 8: stn DW 967' 295-334 m: 2 specimens, attached to gorgonian (WAM C 23250). —
n DWIU60, 375-397 m: 1 specimen, attached to gorgonian (MNHN-Ci 2698).
Types. Holotype : MNHN-Ci 2402 (MUSORSTOM 6. stn DW 478).
Paratypes : MNHN-Ci 2504 (MUSORSTOM 6, stn DW 478). Drawn. — WAM C 23250 (MUSORSTOM 8.
stn DW 967). — MNHN-Ci 2698 (MUSORSTOM 8, stn DW 1060).
Diagnosis. — Rostral plate with sutures visible. R reduced to sliver; basal width of RL 3 times basal width
otK; apical width RL equal to apical width R. CL> and CL2 partially united; CL2 3/4 height, 1/2 basal width
204
D. JONES
of CL'. S elongated, triangular tending to sub-pyramidal, growth ridges convoluted, longitudinal striations absent.
T with 7-9 muscle crests extending well beyond basal margin. Caudal appendages extending 1/2 height of distal
segment of pedicel of cirrus VI, 5-6 segments.
Fig. 30. — Eurylasma angustum sp. nov. Paratype from MUSORSTOM 6, stn DW 478, attached to axis of gorgonian
(MNHN-Ci 2504): a, S. external view: b, S. internal view; c. T, external view; d. T. internal view; e, whole animal,
lateral view; f, whole animal, from above (right S and T removed).
DESCRIPTION. — Size moderate (maximum: RC length 9.7, C height 7.5). Shell form rounded oval, depressed
at carinal end. Parietal plates rough textured basally, smoother apically, growth ridges not deeply marked; all
parietes except C curving in toward orifice; orifice pentagonal. Rostral plate convex, almost bowed, basal width
equal to basal width of CL1, wider and higher than C; apex well curved in toward orifice; rostral sutures visible
externally and internally, R reduced to narrow sliver; RL tapering, basal width 3 times basal width of R. apical
width equal to apical width of R. CL1 and CL2 partially united but CL2 retaining small ala. suture line visible
externally; basal width of CL1 subequal to basal width of rostral plate, apex of CL1 curving toward orifice;
CL2 3/4 height, 1/2 basal width of CL1; apex of CL2 slightly curving toward orifice. C bowed, basal width equal
to basal width of CL1; height 7/10 rostral height; apex erect, not spout-like. Alae of C wide, growth ridges
sinuously parallel to inferior alar margins, superior alar margins subparallel to basis; alae of CL2 1/4 width of
carinal alae, alae of CL1 1/2 width of carinal alae; superior alar margins minutely serrate. Basis calcareous, thin
centrally; flat, central groove axis of gorgonian host. Orifice pentagonal from above, not toothed. Opercular plates
placed parallel to basis; slightly sunken down in orifice in rostral area, then gently curving upward at angle of 30°,
due to concave form of scuta. S elongated, triangular tending to subpyramidal, basal margin 3/5 length of
occludent margin; basal margin without angled bend, but curved toward articular margin at point 2/3 length of
basal margin from occludent margin; externally valve concave in upper 1/3, lower 1/3 retroverted away from
concavity (thus curved in side-view); externally with regular, convoluted growth ridges, longitudinal striations
absent; apex acute; internally muscle scar visible at 2/3 height of valve, small lateral depressor muscle crests
developed; articular furrow narrow; articular ridge not projecting beyond articular margin; adductor ridge indistinct.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
205
T larger than S, triangular, slightly higher than wide; externally with well-spaced growth ridges; internally with 7-
9 strong muscle crests extending well beyond basal margin; articular ridge not extending beyond articular margin;
articular furrow shallow; spur indistinct, short, set at basi-scutal angle, occupying 5/9 width of basal margin.
Color of preserved material creamy lemon-yellow with darker pinkish-orange markings on opercular plates, C and
carinal alae; color after bleaching pearly white with hint of shell pink, C rose-pink with color suffusing onto
carinal alae; eggs orange-yellow. Measurements of 8 specimens examined, randomly selected from several stations,
as follows:
Labium with shallow median depression, small
teeth and setae present. Mandibular palp ovate, with
moderately dense, finely serrulate setae terminally.
Mandible with 3 teeth, tooth 1 largest, separated
from teeth 2 and 3; upper margins of 2 and 3 with or
without small subsidiary cusps; inferior angle
acutely molariform, dentate. Maxillule setose, with
pair of long setae at upper angle; wide notch below
upper angle with 4-6 pairs shorter setae, cutting edge
below almost straight, with 5-7 pairs longer setae,
inferior angle slightly protuberant, with 4-6 short
setae. Maxilla wide, coxal endite weakly defined;
long serrulate setae apical ly.
Cirrus I with rami subequal; anterior ramus
slightly longer than posterior; proximal segments of
anterior and posterior rami moderately protuberant
anteriorly; both rami moderately setose, setae finely
serrulate distally. Cirrus II longer than cirrus I; rami
subcqual; proximal segments of anterior ramus
moderately protuberant anteriorly; rami with segments moderately clothed with setae, some serrulate, especially
distally; posterior ramus with fine spines at antero-distal angles of segments. Cirrus III longer than and similar to
cirrus II, with rami subequal, segments becoming oblong distally; posterior ramus with 2-4 pairs of long, finely
serrulate setae on anterior faces of distal segments; anterior ramus with 2-4 pairs of long, finely serrulate setae on
anterior faces of more distal segments, proximal segments with 6 pairs. Cirrus IV to VI similar, longer than cirrus
III; segments oblong, with 3-4 pairs of setae on anterior faces, distal 2 pairs longest, finely serrulate. Chaetotaxy
ctenopod. Cirral formula as follows:
Fig. 31. — Eurylasma angustum sp. nov. Holotype from
Musorstom 6, stn DW 478. attached to axis
ofgorgonian (MNHN-Ci 2402): lateral view. Scale =
5 mm.
auda appendages 1/2 height of distal segment of pedicel of cirrus VI; 5-6 segmented; long setae apically. with
urc ets of sparse, fine setae around distal margins of segments, length of setae 2/3 length of segment.' Penis
ength of cirrus VI; finely annulated; sparsely setose with circlet of long setae distally. Eggs large, 0.37 x
Remarks. — Eurylasma angustum may be distinguished from E. pyramidale sp. nov. by the form of the
stral sutures (visible m the former, obscure in the latter); by differences in the relative proportions of the
206
D. JONES
CL plates (the basal width of CL2 is 1/2 the basal width of CL' in E. angustum, and 9/10 the basal width of CL'
in E. pyramidal e)\ by the form of the S (tending toward sub-pyramidal in E. angustum, pyramidal in
Fig. 32. — Eurylasma angustum sp. nov. Paratype from MUSORSTOM 6, stn DW 478, attached to axis of gorgonian
(MNHN-Ci 2504): a, right mandible; b, left mandible; c. right maxillule; d, left maxillule; e. maxilla; f. labrum;
g. cirrus I; h, cirrus II; i, cirrus III; j. cirrus VI; k. pedicel of cirrus VI. penis and caudal appendage; 1. cirrus I.
posterior ramus, median segments; m, cirrus VI, posterior ramus, median segment.
Source . MNHN, Pans
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
207
E. pyramidale ); by the form of the T (spur of E. angustum is 5/9 the width of the basal margin and there are
7-9 strong muscle crests extending well beyond the basal margin; the spur off. pyramidale is 2/3 the width of the
basal margin and there are 10-14 muscle crests barely extending beyond the basal margin); by the length of the
caudal appendages (5-6 segments and 1/2 the height of the distal segment of the pedicel of cirrus VI, rather than
13 segments and longer than the pedicel of cirrus VI by 1/4 ); by the dentition of the mandible (strong subsdiary
cusps on the upper margins of teeth 2 and 3 in E. pyramidale, small cusps often on upper margins of teeth 2 and 3
in E. angustum)-, by the size (maximum RC length 9.7, 14.1 respectively); and by the color (creamy lemon-
yellow with darker pinkish-orange markings).
Eurylasma angustum is similar to E. ferulum sp. nov. but R is more reduced in the latter (basal width RL
6 times basal width of R, rather than 3 times), and the basal width of CL2 is 1/2 the basal width of CL' in
£. angustum, and 3/4 the basal width of CL1 in E. ferulum. The rostral plate is higher than the C in E. angustum
(C 7/10 the height of the rostral plate), but the rostral plate is shorter (5/6 the height of the C) in E. ferulum.
The T spur of £. angustum is 5/9 the width of the basal margin and there are 7-9 strong muscle crests extending
well beyond the basal margin; the spur of £. ferulum is 1/2 the width of the basal margin and there are
16-20 strong muscle crests which do not extend beyond the basal margin. The form and length of the caudal
appendages (5-6 segments and 1/2 the height of the distal segment of the pedicel of cirrus VI in £. angustum.
11-12 segments and equal to the height of the pedicel of cirrus VI in E. ferulum) further separate these two species.
Etymology. — From the Latin an gust us. ‘very narrow', in reference to the very narrow form of the rostral
plate.
Distribution. — Vanuatu, Loyalty Islands, 295-400 m.
Eurylasma ferulum sp. nov.
Figs 29, 33-35; Tables 12-14
Dra!^nATERIAL EXAMINED' ~ Norfolk Ridge. Bathus 3: stn CP 815, 460-470 m: 1 specimen (MNHN-Ci 2405).
Types. — Holotype : MNHN-Ci 2405 (Bathus 3, stn CP 815). Drawn.
Scale = 5 mm.
visible externally and internally; R reduced
width of R, apical width of RL twice
Diagnosis. — Rostral sutures visible. R reduced
to narrow sliver; basal width of RL 6 times basal
width of R. CL1 and CL2 partially united; CL2 9/10
height of CL1, basal width of CL2 3/4 basal width
of CL1. S sub-pyramidal; externally with many
extremely convoluted growth ridges, longitudinal
striations absent. T with 16-20 muscle attachment
crests not extending beyond basal margin. Caudal
appendages height of pedicel of cirrus VI. 11-12
segments.
Description. — Size large (maximum: RC
length 15.0, C height 10.6). Shell form elongate
oval from above; all parietes curving in toward
orifice. Parietal plates with smooth, regular growth
ridges. Rostral plate convex, bowed, basal width
subequal to basal width of C. height 5/6 height of
C; apex curved in toward orifice; rostral sutures
to narrow sliver; RL tapering apically, basal width of RL 6 times basal
apical width of R. Basal width of CL1 subequal to basal width of RL. apex
208
D. JONES
curved toward orifice. CL' and CL2 partially united but CL2 retaining minute ala; suture lines visible externally;
height of CL2 9/10 height of CL', basal width of CL2 3/4 basal width of CL'; apex slightly curving in toward
orifice. C bowed, basal width subequal to basal width of rostral plate, C higher than rostral plate by 1/6; apex
slightly curved toward orifice. Alae of C widest, growth ridges sinuously parallel to inferior alar margin, superior
alar margins not parallel to basis, minutely serrate; alae of CL2 1/10 width of carinal alae, alae of CL' 2/5 width
of carinal alae. Basis calcareous, thick; slightly cup-shaped, with central groove. Orifice subtriangular from above;
not toothed. Opercular plates not parallel to basis, gentley curving upward from rostral end at angle of 40°, due to
concave form of S. Basal margin of S 2/5 length of occludent margin; sub-pyramidal shape produced by almost 90°
bend on basal margin toward articular margin, bend at 2/3 length of basal margin from occludent margin;
externally middle area of valve concave, areas above and below retroverted away from concavity (thus curved in
side-view); externally with many extremely convoluted growth ridges, producing frilled appearance; apex pointed;
internally shallow adductor muscle scar at 2/3 height of valve; few small lateral depressor muscle crests developed;
articular furrow narrow; articular ridge not projecting beyond articular margin; adductor ridge indistinct. T larger
than S, triangular; externally with well-marked growth ridges; internally with 16-20 muscle crests, not extending
beyond basal margin; articular ridge short, not projecting beyond articular margin; articular furrow wide; short spur
set at basi-scutal angle, rounded. 1/2 width of basal margin. Color of preserved material mottled creamy/lemon
yellow with C and CL alae more orange-ochre; opercular plates mottled orange-ochre and cream; basis creamy-
white. Measurements of holotype as follows:
Labrum with shallow medial depression; small teeth in irregular row, with some long fine setae; dense spicules
on both sides of depression below teeth. Mandibular palp oblong; dense, serrate setae terminally. Mandible with
Fig. 34. — Eurylasma ferulum sp. nov. Holotype from Bathus 3, stn CP 815 (MNHN-Ci 2405): a, S, external view;
b, S, internal view; c, T, external view; d. T, internal view; e. whole animal, lateral view; f. whole animal, from
above.
Source . MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
209
3 teeth, tooth 1 well-separated from 2 and 3; upper margin of 3 with small subsidiary cusps; inferior angle bluntly
molariform, dentate. Maxilla wide, coxal endite barely defined; long serrulate setae terminally. Maxillule with
1-2 pairs of long, stout setae at upper angle; notch below upper angle moderately large, indistinct, with 4-5 pairs
of smaller setae; cutting margin below almost straight, with 7-8 pairs of larger setae; inferior angle with 4-5 pairs
of smaller setae and 1 longer seta.
2.0 mm
a. d, g
0.2 mm
b, c
0.5 mm
e. f
h.k
0.5 mm
Fig. 35. — Eurylasma ferulum sp. nov, Holotype from BaTHUS 3, stn CP 815 (MNHN-Ci 2405): a, right mandible; b. left
mandible; c, right maxillule; d, left maxillule; e. labrum; f. mandibular palp; g. cirrus I, anterior ramus, median
segments; h, cirrus I; i, cirrus VI; j, cirrus III; k, cirrus II.
210
D. JONES
Cirrus I with rami subequal; posterior ramus slightly longer than anterior; proximal segments of anterior ramus
protuberant anteriorly; both rami thickly setose, setae finely serrulate distally. Cirrus II similar to cirrus III and
longer than cirrus I; rami subequal; proximal segments of anterior ramus protuberant anteriorly; rami with seg¬
ments thickly setose, setae finely serrulate, especially distally. Cirrus III with rami subequal, segments becoming
oblong distally; posterior ramus with 2-4 pairs of long, finely serrulate setae on anterior faces of distal segments;
anterior ramus with 2-3 pairs of long, finely serrulate setae on anterior faces of distal segments, proximal segments
with 7 pairs. Cirrus IV to VI similar, longer than cirrus III; segments oblong, with 4-5 pairs of setae on anterior
faces, distal 2 pairs longest, finely serrulate. Chaetotaxy ctenopod. Cirral formula of holotype as follows:
Caudal appendages length of pedicel of cirrus VI; 11-12 segmented; apically with sparse setae, circles o! sparse
setae around distal margins of segments, setal length 1/3 segmental length. Penis broken, basal remnant attached
to cirrus VI.
Remarks. — Only 1 specimen of this species has been collected, but the very narrow, sliver-like R. and
characters of the shell and of the soft part morphology are sufficiently distinct to warrant specific differentiation.
Differences between E.ferulum sp. nov. and E. angustum sp. nov. are listed under E. angustum.
Eurylasma ferulum sp. nov. is distinguished from E. pyramidale sp. nov. by the rostral sutures, obscure in the
latter, visible in the former. The rostral plate and C are of equal height in E. pyramidale , whereas the C is higher
than the rostral plate by 1/6 in E.ferulum. The form of the S is sub-pyramidal in E.ferulum. and the convoluted
S growth lines are not cut by longitudinal striae. In E. pyramidale the S is pyramidal and the convoluted growth
lines are cut by faint longitudinal striae, giving a slight beaded effect. In E. ferulum the T spur is 1/2 the width ol
the basal margin and has 16-20 muscle crests which do not extend beyond the basal margin; the spur of
E. pyramidale is 2/3 the width of the basal margin and has 10-14 muscle crests barely extending beyond the basal
margin. Cirrus I of E. pyramidale has the anterior ramus longer than the posterior, with the proximal segments of
both rami protuberant anteriorly, whereas that of E. ferulum has the posterior ramus longer than the anterioi with
only the proximal segments of the anterior ramus protuberant. The caudal appendages of E. ferulum are 11-12
segmented and are equal to the height of the pedicel of cirrus VI, whilst those of E. pyramidale are 13 segmented
and longer than the pedicel of cirrus VI by 1/4. The upper margins of mandibular teeth 2 and 3 have many small
subsidiary cusps in E. pyramidale ; only the upper margin of tooth 3 has small subsidiary cusps in E. ferulum.
Etymology. — From the Latin ferula, ‘rod’, referring to the rod-like nature of the R.
Distribution. — Norfolk Ridge, 460-470 m.
Eurylasma pyramidale sp. nov.
Figs 29. 36-38; Tables 12-14
MATERIAL EXAMINED. — Norfolk Ridge. Smib 5: stn DW 71. 265 in: I specimen, attached lo live coral
( Corallium sp.) (MNHN-Ci 2404). — Stn DW 75. 270 m: 1 specimen, attached to live coral ( Corallium sp.). —
Stn DW 93. 255 m: 1 specimen, attached to live coral ( Corallium sp.) (MNHN-Ci 2403); many specimens, attached to
live coral (.Corallium sp.) (BMNH); many specimens, attached to live coral ( Corallium sp.) (USNM). — Stn DW 94.
275 m: several specimens, attached to live coral ( Corallium sp.). — Stn DW lOi, 270 m: 2 specimens, attached to live
coral ( Corallium sp.).
Bathus 3: stn CP 805, 278-310 m: 3 specimens, attached to coral (WAM 255-96). — Stn DW 830. 361-365 m:
4 specimens, attached to coral. (MNHN-Ci 2699). Drawn.
TYPES. — Holotype: MNHN-Ci 2403 (Smib 5. stn DW 93).
Source . MNHN , Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
21 1
Paratypes: BMNH (SMIB 5, stn DW 93). — MNHN-Ci 2404 (Smib 5, stn DW 71). — MNHN-Ci 2699
(Bathus 3, stn DW 830). Drawn. — USNM (Smib 5, stn DW 93). — WAM 255-96 (Bathus 3. stn DE 830).
DIAGNOSIS. — Rostral plate sutures obscure. CL1 and CL2 partially united; basal width of CL2 9/10 basal
width of CL1; height of CL2 8/9 height of CL1. S elongated, pyramidal shape; externally convoluted growth ridges
cut with many faint longitudinal striations. T with 10-14 muscle attachment ridges barely extending beyond basal
margin. Caudal appendages longer than pedicel of cirrus VI by 1/4. 13 segments.
Fig. 36. — Eurylasma pyramidale sp. nov. Paralype from Bathus 3. stn DW 830 (MNHN-Ci 2699): a. S. external view:
b, S. internal view; c, T. external view; d. T. internal view; e, whole animal, from above (S and T removed); f. whole
animal, lateral view.
Description, — Size large (maximum: RC length 14.1. C height 6.7). Form rounded, all parietes except C
curving in toward orifice. Parietal plates rough textured; growth ridges regular, not deeply marked. Rostral plate
with sutures obscure; basal width less than basal width of C, slightly greater than basal width of CL1; apex curved
in toward orifice; height subequal to height of C. CL1 and CL2 united, CL2 still retaining small ala. suture lines
visible externally. Basal width of CL1 slightly less than basal width of rostral plate; apex curving toward orifice.
Height ol CL2 8/9 height of CL1, basal width of CL2 9/10 basal width of CL1 ; apex curving toward C. C widest
of all plates, slightly higher than R, bowed, apex slightly curved inward, not spout-like. Basis calcareous, cup-
shaped. Orifice triangular from above; not toothed. Opercular plates set parallel to basis, slightly sunken down in
oiifice, making gently concave depression due to concave form of scuta. S elongate, basal margin 5/12 length of
occludent margin; pyramidal shape, produced by almost 90° bend toward articular margin on basal margin, bend at
3/4 length ol basal margin from occludent margin; valve concave in upper 1/2 with upper 1/4 retroverted away
ftom concavity; externally with convoluted growth ridges, cut by many longitudinal striations, giving almost
beaded appearance; apex pointed; internally muscle scar visible at 2/3 height of valve; small lateral depressor
muscle crests present; articular furrow wide; articular ridge barely projecting beyond articular margin; adductor ridge
indistinct. T larger than S, triangular; externally with well-marked growth ridges; internally with 10-14 muscle
crests barely extending beyond basal margin; articular ridge short, not projecting beyond articular margin; articular
urrow wide, short spur set at basi-scutal angle, rounded, width 2/3 basal margin. Parietes of bleached specimens
212
D JONES
pearly white with hint of shell pink, covered by pearly white, moderately thick membrane; specimens from live
Corallium sp. with parietes, opercular plates and covering membrane yellowish-cream, sometimes C orange-
yellow. Measurements of largest specimen examined as follows:
Fig. 37. — Eurylasma pyramidale sp. nov. Paratype from BATHUS 3, stn DW 830 (MNHN-Ci 2699): a, right mandible;
b, left mandible; c, right maxillule; d. left maxillule; e, mandibular palp; f, labrum; g. maxilla; h, cirrus I;
i, cirrus II; j, cirrus VI, penis and caudal appendage; k, cirrus III; 1, cirrus I. posterior ramus, median segments.
Source : MNHN. Paris
CIRRIPED1A THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
213
Labrum with shallow medial depression; few scattered, small teeth present. Mandibular palp oblong; dense,
serrate setae terminally. Mandible with 3 teeth, tooth 1 largest and well-separated from 2 and 3; upper margin of 2
and 3 strongly dentate; inferior angle moderately large, bluntly molariform, dentate. Maxillule with 1-2 pairs of
long, stout setae at upper angle; notch below upper angle moderately wide, with 3-4 pairs of smaller setae; long
cutting margin below with 7-10 pairs of larger setae; inferior angle small, with 4-6 pairs of smaller setae. Maxilla
wide, coxal endite barely discernible; long serrulate setae terminally.
Cirrus 1 with rami subequal; anterior ramus
slightly longer than posterior; proximal segments of
anterior and posterior rami protuberant anteriorly;
both rami with segments moderately setose, setae
finely serrulate distally; distal segments of posterior
ramus with dense, stout setae at antero-distal angles
and anterior surface with combs of small setae. Cirrus
II longer than cirrus I; rami subequal; proximal
segments of anterior ramus protuberant anteriorly;
anterior and posterior rami antenniform distally; all
segments setose, setae simple or serrulate. Cirrus III
longer than but similar to cirrus II. with rami
subequal, segments becoming oblong distally;
posterior ramus with 2-5 pairs of long, finely
serrulate setae on anterior faces of distal segments;
anterior ramus with 2-5 pairs of long, finely serrulate
setae on anterior faces of distal segments, proximal
segments with 9 pairs. Cirrus IV to VI similar,
longer than cirrus III; segments oblong, with 4-5 pairs of setae on anterior faces, distal 2 pairs of setae longest,
finely serrulate. Chaetotaxy ctenopod. Cirral formula as follows:
Fig. 38. — Eurylasma pyramidale sp. nov. Paralypes from
Bathus 3, sin DW 830 (WAM 255-96): lateral view.
Scale = 5 mm.
CV1 c. a.
23. 24 1 1
22, 26 12
Caudal appendages slightly longer than pedicel of cirrus VI; 13 segmented; long setae apically, circlets of short
setae around distal margins of segments. Penis almost as long as cirrus VI; annulated; sparsely setose with circlet
of long setae distally.
Remarks. Differences between E. pyramidale and £ angustum sp. nov. are listed under E. angustum and
between £. pyramidale and E.ferulum sp. nov. are listed under E.ferulum.
Etymology. — From the Greek pyramidalis , ‘of a pyramidal shape’, relating to the pyramidal form of the
scuta.
Distribution. — Norfolk Ridge, 255-365 m
Genus TETRAPACHYLASMA Foster, 1988 (emend.)
Fig. 39; Tables 3-4, 15-17
Tetrapachylasma Foster, 1988: 226.
Diagnosis. — Adult with variable wall pattern of 6-plates (R-CL1-CL2-C), or 4-plates (R-CL-C); in 6-plated
wall, RL coalesces with R to form compound rostrum (RL+R+RL); in 4-plated wall RL and R form compound
214
D JONES
rostrum and CL2 coalesces with CL1 to form compound CL plate. Rostral plate sutures obscure; basal width of R
equal to basal width of RL. CL' and CL2 separate, partially coalesced, or totally concrescent. S with growth lines
cut by longitudinal striae. Caudal appendages varying from short (1/8 height of basal segment of pedicel of cirrus
VI) to long (approximately twice height of pedicel of cirrus VI).
Type Species. — Tetrapachylasma trigonum Foster, 1988.
Recent Species. — Tetrapachylasma arcuatum sp. nov.; T. aurantiacum (Darwin, 1854); T. ferrugomaculosa
(Jones, 1993b); T. ornatum sp. nov.; T. trigonum Foster, 1988.
REMARKS. — Foster's (1988) diagnosis of Tetrapachylasma defined the genus as having "...Four (adult) solid
calcareous compartmental plates including a carina, a rostrum and paired laterals ...". However, new material and
re-examination of P. aurantiacum Darwin, 1854 and P. ferrugomaculosa Jones, 1993 (all attributed to
Tetrapachylasma herein) confirm that the number of wall plates in the adults of these species is variable. The adult
may have a 4-plated pattern, consisting of a compound rostral plate, a pair of plates formed by the coalescence of
CL1 and CL2, and a C. However, within the same sample, 6-plated individuals of the same species also occur,
where CL' and CL2 are either separate or partially coalesced (but in the latter condition CL2 still retains a minute
ala). In these individuals the rostral sutures may be obscure or visible. I consider that Foster's (1988) diagnosis of
Tetrapachylasma applies to the 4-plated adult condition found in the above samples and I have broadened the
diagnosis of Tetrapachylasma accordingly, to embrace the variable plate numbers also found in the above samples.
The variable 4- or 6-plated pattern, however, makes specific identification and attribution difficult when dealing
with a single or few specimens. Tetrapachylasma trigonum may represent a condition where the 4-plated pattern
has become fixed - the small number of specimens (3) in Foster’s sample of T. trigonum is not significant to
determine intra-specific variation within this species.
In Tetrapachylasma, RL coalesces with R to form a compound rostrum (RL+R+RL) in both the 6- and 4-plated
wall forms. RL is not incorporated into the sheath, with the exception of T. ornatum sp. nov., where RL is clearly
incorporated into the sheath (contrary to the definition of the Pachylasmatoidea). Thus in Tetrapachylasma, as re¬
defined. the adult wall condition (6- or 4-plated) is achieved through a reduction in the number of plates by
coalescence, rather than by exclusion as was postulated by Foster (1988). The pattern of coalescence of the plates
has not become fixed, at least within T. arcuatum sp. nov., T. aurantiacum, T. ferrugomaculosa and T. ornatum
sp. nov., and the plates are disposed symmetrically rather than asymmetrically, since both the 4- and the 6-plated
condition occur within the same species and no instances have been found of a 5-platcd condition (i.c. CL1 and
CL2 totally coalesced on one side only). Thus the adult condition for this genus is considered to be a variable wall
pattern of either 4 or 6 parietal plates.
Such intra-specific variation is also seen in the Balanomorpha. Within the Chihamaloidea Jehlius gilmorei
Ross, 1971. an unusual species from San Ambrosia and San Felix Islands, Chile, exhibits an array of wallpatterns
in the adult condition (Ross, 1971). In Jehlius, development of the adults is in a transitional stage between 6 and
4 plates, with reduction in the number of plates by coalescence rather than by exclusion. In the 4-plated stage the
wall plates are not secondarily coalesced, the plates are disposed symmetrically or asymmetrically and the
coalescence pattern is variable throughout the population. It appears, from the few specimens known of this
species, that the pattern of coalescence of the plates has not become fixed. In the 4-plated Tetrachthamalus,
however, a different pattern is seen. Here RL are coalesced with R and, during the ontogeny of individuals, the
4 plates coalesce.
Darwin's (1854) diagnosis of Pachylasma defines the adult condition as 6- or 4-platcd viz. " ....when mature,
either six, or in appearance only four owing to the close union of the lateral compartments...". There is an implicit
assumption in Darwin's diagnosis that it applies to the fixed plate number in the adult condition (i.e. it refers to
inter-specific variation). Darwin's diagnosis was based on 2 species - P. giganteum (adult condition recognised as
fixed 6-plated in the present paper) and P. aurantiacum (adult condition recognised as variable 4- or
6-plated in the present paper). The fixed 6-plated adult condition is herein recognised for Pachylasma (type species
P. giganteum ) and the variable 4- or 6-platcd adult condition is herein recognised for Tetrapachylasma (type species
T. trigonum).
Source : MNHN, Paris
CIRR1PEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
215
Tables 15 and 16. — Tetrapachylasma; characters of the species
artic. - articular; b-s = basi-scutal; ext. = external; int. = internal; n.a. = not applicable; proj. = projecting.
artic. articular, c.a. - caudal appendage; long. = longitudinal; max. = maximum; proj. = projecting; segs = segments.
216
D. JONES
Distribution. — Southwestern and eastern Australia, Western and Central Pacific Ocean: Western Australia,
3-30 m (T. fe rrttg omaculosa); eastern Australia, Kermadec Islands, 122-490 m (T. aurantiacum)-, Vanuatu.
Chesterfield Islands, Loyalty Islands, Loyalty Ridge, Norfolk Ridge, 226-1220 m (T. arcuatum)-, Marianas Islands!
6 in ( T. ; ornatum sp. nov.); Cook Islands, 15 m (T. trigonum).
Table 17. — Tetrapachylasma : maximum size, geographical and bathymetrical distribution of the species
Fig. 39. — Tetrapachylasma. Distribution map: A, T. ornatum (6 m); ■ T. arcuatum (226-1220 m); •. T. aurantiacum
(122-490 m); ▲. T. ferrugomaculosa (3-30 m); O, T. trigonum (15 m).
Tetrapachylasma arcuatum sp. nov.
Figs 39-42; Tables 15-17
„. MATERiAL EXAMINED. — Chesterfield Islands. MUSORSTOM 5: stn DW 255, 280-295 m: 2 specimens ( MNHN-
n i a"f hed 10 COraL - Stn DW 301’ 487-610 ™ 5 specimens (2 juveniles), attached to sponge. -
Mn DW 304, 385-420 m; 3 specimens.
n Jr0yalg ISn.d7s: MUSORSTOM 6: stn DW 407, 360 m: 1 specimen. - Stn CP 471, 460 m: I specimen, attached to
sponge (MNHN-C^iil^^severa^sp^cimeris^battache^U) ^fpong^l [""spe'cimen. ^auached To dea^styDsterlc^'txfra'l0
d leacf coral"] M N FI N - C i* *2 700)° *D rawr a f se va ral 2il" ‘ °Vi8er°US *** <br°ken)' ^ °range 3,taChed "
MuS0RST0^ 6: s,n Dw 478' 4°0 m: 1 specimen, attached to 1 specimen of Hexelasma persicum
sp. nov. (MNHN); 1 specimen (WAM 256-96); 1 specimen (WAM C 23251). 7
Vanuatu. Musorstom 8: stn CP 963. 400-440 m: several specimens, attached to sponge
New Hebrides Arc. Volsmar: stn DW 16, 500m: 1 specimen.
Source : MNHN , Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
217
Norfolk Ridge. Chalcal 2: stn CH 4, 253 m: 1 specimen, attached to sponge.
Smib 5: stn DW 87. 370 m: I specimen, attached to coral rubble (USNM). — Stn DW 93. 255 m: many specimens.
Halipro 2: stn I3T 70, 226-238 m: 1 specimen, attached to gorgonian. — Sin BT 71, 820-1220 m: 2 specimens,
attached to gorgonian.
Types. — Holotype: MNHN-Ci 2406 (Musorstom 6, stn DW 472).
Paratypes : BMNH (MUSORSTOM 6, stn CP 471). — MNHN-Ci 2407 (MUSORSTOM 5, stn DW 255). —
MNHN-Ci 2700 (MUSORSTOM 6, stn DW 478). Drawn. — USNM (Smib 5, stn DW 87). — WAM 256-96
(Musorstom 6, stn DW 478).
Diagnosis. — Rostral plate sutures visible; basal width of R equal to basal width of RL. CU and CL?
separate, partially or totally coalesced; basal width of CL2 equal to basal width of CL'. Articular margin of S
curved toward occludent margin in upper 1/3; externally sinuous growth lines cut by 7-8 slight longitudinal striae;
articular ridge projecting. T articular ridge slightly projecting; 12-16 muscle attachment ridges projecting beyond
basal margin. Caudal appendages twice length of pedicel of cirrus VI; 16-18 segments.
Fig 40. Tetrapachyl asm a arcuatum sp. nov. Paratype from Musorstom 6, stn DW 478 (MNHN-Ci 2700): a. S. internal
view; b. S, external view; c, T, external view; d. T, internal view; e. whole animal, lateral view; f. whole animal,
rostral view.
Description. — Size moderate (maximum: RC length 1 1.4, C height 9.5). Shell form upright, all parietal
plates except C gently sloping in toward orifice; C sloping toward orifice at acute angle. Parietal plates solid,
growth ridges irregular, not marked by small setae. Rostral plate broad, slightly convex, basal width twice basal
width ol C; sutures between R and RL visible internally, partially visible or obscure externally; basal width of R
equal to basal width of RL. CL1 and CL2 separate in juveniles, partially or totally coalesced in mature specimens;
asal width of CL1 equal to basal width of CL2, height of CL2 subequal to height of CL1; external suture
sometimes indistinct, giving superficial impression of 1 plate, coalescence of plates incomplete with CL2
retaining minute ala. latter obscure in adults if plates apically eroded; ala of CL' 1/4 width of carinal ala. ala of
CL- minute, 1/6 width of alae of CL1. C alae widest; C highest of all plates, almost twice height of R. basal
wi 1 1 1/2 basal width ol R. Basis thin, flat; calcareous, centrally thin, sometimes membranous. Orifice large, sub-
triangu ar, not toothed. Basis thin, flat; calcareous, centrally thin, sometimes membranous. Opercular plates lodged
upright in orifice; C seemingly forming hood over opercular plates. S smaller than T; triangular, elongated,
218
D. JONES
convex basal margin 2/3 length of occludent margin; articular margin distinctly curved toward occludent margin in
upper 1/3; externally growth ridges sinuous, cut with 7-8 radial striations; apex blunt; internally adductor muscle
pit deep, situated 2/3 up length of valve; lateral depressor muscle pit indistinct; articular furrow narrow; articular
ridge 1/2 length of valve, projecting beyond articular margin; adductor ridge indistinct. T triangular, wider than S;
externally growth ridges well marked, articular margin with growth ridges raised to form elevated ridge; internally
with 12-16 strong muscle attachment crests projecting well beyond basal margin; articular ridge short, prominent,
projecting slightly beyond articular margin; articular furrow broad; spur set at 1/12 length of basal margin front
basi-scutal angle, truncate, 3/7 width of basal margin, barely defined from basal margin. Color of parietal plates
dirty cream with orange-brown markings on C and carinal alac. Eggs orange. Measurements of 6 specimens
examined, randomly selected from several stations, as follows:
Labrum with shallow medial depression, with regular row of small bi- or tri-lobed teeth. Mandibular palp
ovoid; long serrate setae terminally. Mandible with 3 teeth, tooth 1 largest, well separated from teeth 2 and 3:
upper margins ol 2 and 3 teeth with small subsidiary cusps; inferior angle bluntly molariform. dentate. Maxiilule
setose; 2-3 long, stout setae at upper angle; indistinct notch below upper angle with 23 pairs of smaller setae;
cutting margin below stepped, with 5-6 pairs of longer, stout setae; inferior angle stepped, with 5-6 pairs of
smaller setae. Maxilla wide; coxal endite not well developed; dense, long setae terminally.
Fig. 41. — Tetrapachylasma arcuatum sp. nov. Holotype
from Musorstom 6, stn DW 472 (MNHN-Ci 2406):
lateral view. Scale = 5 mm.
segments, proximal segments with 6 pairs. Cirrus IV to
4 pairs of setae on anterior faces, distal 2 pairs longest.
Cirrus I with rami unequal; anterior ramus longer
than posterior; proximal segments of anterior ramus
protuberant anteriorly; both rami with segments
thickly setose, setae strong, serrulate, stout; poslero-
distal angles of segments of posterior ramus with
tufts of stout, serrate setae, surface with combs of
small setae. Cirrus II similar to but slightly longer
than cirrus I; rami subequal; proximal segments of
anterior ramus broad, slightly protuberant anteriorly,
all segments thickly setose, setae simple, stout;
posterior ramus more antenniform than anterior,
segments becoming more elongated and narrower
distally, distal segments with 3-4 pairs of long,
finely serrulate setae on anterior faces. Cirrus III
similar to cirri IV-VI; rami subequal. segments
becoming oblong distally; posterior ramus with 2-4
pairs of long, finely serrulate setae on anterior faces
of distal segments; anterior ramus with 2-3 pairs of
long, finely serrulate setae on anterior faces of distal
VI similar, longer than cirrus III; segments oblong, with
Chaetotaxy ctenopod. Cirral formula as follows:
seImemajlZrdT "T ''f h °f l*dicel °U 16-18 segmented; long setae apieally and around distal
r ,cngft of dms vi; ”"uiaMd; ^ ^ —
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
219
G' 42' . Te,raPachylasma arcualum sp. nov. Paratype from Musorstom 6. stn DW 478 (MNHN-Ci 2700): a. cirrus VI,
posterior ramus, median segments; b. right mandible; c. left mandible; d, right maxillule; e. left maxillule; f. labrum;
g. maxilla, h. cirrus I, posterior ramus, median segments; i, mandibular palp; j. cirrus II; k. cirrus 111; 1, cirrus 1:
m, cirrus and caudal appendage; n, cirrus III, posterior ramus, median segments; o, penis.
Remarks. T. arcuatum sp. nov. can be separated from T. aurantiacum as follows: the S articular margin is
convex, the S basal margin is 2/3 the length of the occludent margin, the articular ridge is projecting and the scutal
220
D. JONES
growth ridges are sinuous and cut by 7-8 longitudinal striae in T. arcuatum. In T. aurantiacum the S articular
margin is not convex, the S basal margin is 1/2 the length of the occludent margin, the articular ridge is not
projecting and the scutal growth ridges are convoluted and cut by 5-6 slight longitudinal striae. The T articular
ridge of T. arcuatum is slightly projecting, and the spur is indistinct, 3/7 the width of the basal margin and set at
the basi-scutal angle, with 12-16 muscle crests projecting from the basal margin. In T. aurantiacum the T articular
ridge is not projecting, and the spur is 1/2 the width of the basal margin and set at a distance of 1/10 the width of
the basal margin from the basi-scutal angle, with 6-7 muscle crests projecting from the basal margin. The caudal
appendages of T. arcuatum are 16-18 segmented and twice the height of the pedicel of cirrus VI; those
of T. aurantiacum are 5 segmented and 1/5 the height of the pedicel of cirrus VI. Tetrapachylasma arcuatum
is small in size (RC length 1 1.4. C height 9.5), whereas T. aurantiacum is significantly larger (RC length 24.0,
C height 7.0).
T. arcuatum differs from T. ornatum sp. nov. as the S articular margin is curved, the S basal margin is 2/3 the
length of the occludent margin, the articular ridge is projecting and the scutal growth ridges are sinuous, and cut by
7- 8 longitudinal striae. In T. ornatum the S articular margin is not curved, the S basal margin is 1/2 the length of
the occludent margin, the articular ridge is not projecting and the growth ridges are complexly sculptured, cut by
4-6 deep, longitudinal striae. The T articular ridge of T. arcuatum is slightly projecting, and the spur is indistinct.
3/7 the width of the basal margin and set at the basi-scutal angle, with 12-16 muscle crests projecting from the
basal margin. In T. ornatum the T articular ridge is projecting, and the spur is short, more than 1/3 the width of
the basal margin and set at a distance of 1/12 the width of the basal margin from the basi-scutal angle, with
8- 12 muscle crests projecting from the basal margin. Soft parts of T. ornatum are unknown.
The long caudal appendages (approximately twice the length of the pedicel of cirrus VI) and the form of the
paries (without narrow ribs) prevent confusion of T. arcuatum with T. trigonum (uniarticulate caudal appendages,
1/8 the height of the basal segment of the pedicel of cirrus VI; outer surface of paries with narrow ribs
peripherally). The S articular margin is curved, the S basal margin is 2/3 the length of the occludent margin, the
articular ridge is projecting and the scutal growth ridges are sinuous, and cut by 7-8 longitudinal striae in
T. arcuatum. In T. trigonum the S basal margin is 5/8 the length of the occludent margin, the articular ridge is
not projecting and the scutal growth ridges are ruffled and overlapping, and not cut by longitudinal striae.
In T. arcuatum the T articular ridge is slightly projecting, and the spur is indistinct, 3/7 the width of the basal
margin and set at the basi-scutal angle, with 12-16 muscle crests projecting from the basal margin. In T. trigonum
the T articular ridge is projecting, and the spur is 1/3 the width of the basal margin and set at a distance of 1/7
the width of the basal margin irom the basi-scutal angle, with 6 muscle crests slightly projecting from the basal
margin, and the basal corner of the articular ridge is hollowed out for the reception of dwarf males.
The new species diflers from T. ferrugomaculosa as the caudal appendages of the latter are 7-9 segmented
and short (equal to the height of the basal segment of the pedicel of cirrus VI) rather than 16-18 segmented
and long (approximately twice the height of the pedicel of cirrus VI). In T. ferrugomaculosa the S articular margin
is not curved, the S basal margin is 7/10 the length of the occludent margin, the articular ridge is barely projecting
and the scutal growth ridges are very convoluted, and cut by 6-8 longitudinal striae. In T. arcuatum the S articular
margin is curved, the S basal margin is 2/3 the length of the occludent margin, the articular ridge is projecting
and the scutal growth ridges are sinuous, and cut by 7-8 longitudinal striae. The T articular ridge of
T. ferrugomaculosa is not projecting, the T spur is indistinct, 1/2 the width of the basal margin and set at a
distance of 1/12 the width of the basal margin from the basi-scutal angle, with 7-9 muscle crests projecting well
beyond the basal margin. In T. arcuatum the T articular ridge is slightly projecting, the T spur is indistinct,
3/7 the width of the basal margin and set at the basi-scutal angle, with 12-16 muscle crests projecting from the
basal margin.
Etymology. — From the Latin arcuatus, ‘curved’, in reference to the curved form of the articular marein of
the S.
Distribution. — Chesterfield Islands, 280-610 m; Loyalty Islands, 300-460 m; Loyalty Ridge, 400 m;
Vanuatu. 400-440 m; New Hebrides Arc, 500 m; Norfolk Ridge, 226-1220 m.
Source MNHN, Paris
CIRR1PEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
221
Tetrapachylasma aurantiacum (Darwin, 1854), comb. nov.
Fig. 39; Tables 15-17
Pachylasma aurantiacum Darwin, 1854: 480, pi. 20 figs 1 a-d. — Weltner, 1897: 273. — Gruvel. 1905: 199, fia. 219.
— Newman & Ross. 1976: 40. — Foster, 1981: 354, figs 2 E, 4 A-L.
TYPES. — Holotype: BMNH (no registration number) New South Wales, Australia, apparently from deep
water, attached to sandstone; dry specimen.
Holotype depositor y: BMNH.
Diagnosis. — R and RL closely united by linear sutures, latter often obliterated externally but visible
internally; basal width R equal to basal width of RL. CL' and CL2 closely united by linear sutures; basal width
CL2 equal to basal width CL1. S articular ridge not projecting; growth lines extremely convoluted, cut
by 5-6 slight longitudinal striae. T articular ridge not projecting; 6-7 muscle crests projecting from basal margin.
Caudal appendages 5 segmented, 1/5 the height of pedicel of cirrus VI.
Remarks. — The variable nature of the adult plate pattern has been noted previously for Pachylasma
aurantiacum but the significance of the variability has not been recognised. Darwin (1854) commented that '...the
compartments in appearance are only four; but on close examination, the lateral compartments are seen to be
divided by a very fine fissure into 2 equal compartments...' and FOSTER (1981 ), examining new collections of this
species, noted '... carinolatus/latus or lalus/rostral plate variously coalesced externally so that no intermediate alar
areas show (although always with distinct sutures internally).' The variable plate pattern of adults of
P. aurantiacum is recognised herein. Comparing shell plate, opercular plate and soft body morphology, this
species is now included in the genus Tetrapachylasma.
Tetrapachylasma aurantiacum is a large species (maximum: RC length 24.0, C height [inferred] 7.0).
Dillerences between 7. aurantiacum and T. arcuatum sp. nov. are listed under T. arcuatum. In T. ferrugomaculosa
the caudal appendages are 7-9 segmented and equal to the height of the basal segment of the pedicel of cirrus VI.
rather than 5 segmented and 1/5 the height of the pedicel of cirrus VI (T. aurantiacum). In T. ferrugomaculosa
the S basal margin is 7/10 the length of the occludent margin, the articular ridge is barely projecting and the scutal
growth ridges are convoluted, and cut by 6-8 longitudinal striae. In T. aurantiacum the S basal margin is 1/2 the
length ol the occludent margin, the articular ridge is not projecting and the scutal growth ridges are convoluted and
cut by 5-6 slight longitudinal striae. The T articular ridge of T. ferrugomaculosa is not projecting, the T spur is
indistinct, 1/2 the width of the basal margin and set at a distance of 1/12 the width of the basal margin from the
basi-scutal angle, with 7-9 muscle crests projecting well beyond the basal margin. In T. aurantiacum the
articular ridge is not projecting, and the spur is 1/2 the width of the basal margin and set at a distance of 1/10
t ie width ol the basal margin from the basi-scutal angle, with 6-7 muscle crests projecting from the basal margin.
Characters ol the T separate T. aurantiacum from. T. ornatum sp. nov. The T articular ridge of T. aurantiacum
is not projecting, and the spur is 1/2 the width of the basal margin and set at a distance of 1/10 the width of the
basal margin from the basi-scutal angle, with 6-7 muscle crests projecting from the basal margin. In T. ornatum
el articular ridge is projecting, and the spur is indistinct, more than 1/3 the width of the basal margin and set at
a 'stance ol 1/12 the width of the basal margin from the basi-scutal angle, with 8-12 muscle crests projecting
rom me basal margin. In T. aurantiacum the S basal margin is 1/2 the length of the occludent margin, the
ar icu ar ri ge is not projecting and the scutal growth ridges are extremely convoluted and cut by 5-6 slight,
g.tu ina striae. In 7. ornatum the S basal margin is 1/2 the length of the occludent margin, the articular ridge
mrK fr°jeC1'ng and the growlh ridSes are complexly sculptured, cut by 4-6 deep, longitudinal striae. Soft
. ° „ orna,um are unknown. Tetrapachylasma ornatum is small in size (RC length 6.0. C height 2.5).
wnereas T. aurantiacum is large (RC length 24.0. C height 7.0).
(5 can bc separated from T. trigonum by the form of the caudal appendages
the ned' ' I f ^ ^ hC‘ght °f lhe pcdlcel of c,rrus VI- and uniarticulate. 1/8 the height of the basal segment of
P o cirrus VI, respectively). In T. aurantiacum the paries are without narrow ribs; in T. trigonum
222
D. JONES
the surface of the paries have narrow ribs peripherally. In T. aurantiacum the S basal margin is 1/2 the length of
the occludent margin, the articular ridge is not projecting and the scutal growth ridges are extremely convoluted and
cut by 5-6 slight, longitudinal striae. In T. irigonum the S basal margin is 5/8 the length of the occludent margin,
the articular ridge is not projecting and the scutal growth ridges are ruffled and overlapping, and not cut by
longitudinal striae. The T articular ridge of T. aurantiacum is not projecting, and the spur is 1/2 the width of the
basal margin and set at a distance of 1/10 the width of the basal margin from the basi-sculal angle, with
6-7 muscle crests projecting from the basal margin. In T. trigonum the T articular ridge is projecting, and the spur
is 1/3 the width of the basal margin and set at a distance of 1/7 the width of the basal margin from the basi-scutal
angle, with 6 muscle crests slightly projecting from the basal margin, and the basal corner of the articular ridge
hollowed out for the reception of dwarf males.
DISTRIBUTION. — New South Wales, Australia; Kermadec Is. New Zealand; 122-490 m.
Tetrapachylasma ferrugomaculosa (Jones, 1993), comb. nov.
Fig. 39; Tables 15-17
Pacliylasma ferrugomaculosa Jones, 1993b: 116, fig. 1 a-o.
MATERIAL EXAMINED. — South-western Australia (between Rottnest I. and Albany). Rottncst
Island, 30 m: hololype (WAM 220-91). — Roe Reef. Rottnest Island, 30 m: paratype (WAM 223-91).
Ollier material : Duffield Ridge, Rottnest Island, 30 m (WAM 224-91). — Roe Reef. Rottnest Island, 30 m (WAM 225-
91). — Salmon Bay, Rottnest Island (WAM 427-95). — Kwinana. 3-5 m (WAM 180-75). — Kwinana. 3-5 m (WAM 1 86-
75). — Off Castle Rock, Dunsborough, 20 m (WAM 221-91). — From wreck off Dunsborough (WAM 464-92). — King
George Sound, Albany (WAM 222-91).
TYPES. — Holotype : WAM 220-91; Western Australia; Duffield Ridge, Rottnest Island, 30 m.
Paratype: WAM 223-91; Western Australia; Roe Reef, Rottnest Island, 30 m.
Holotype and paratype depository: WAM.
Diagnosis. — R and RL closely united by linear sutures, latter often obliterated externally but visible
internally; basal width R twice basal width of RL. Basal width CL2 1/2 basal width CL2; plates separate, or
partially or totally coalesced. S articular ridge barely projecting; growth lines convoluted, cut by 6-8 longitudinal
striae. T articular ridge not projecting; 7-9 muscle crests projecting from basal margin. Caudal appendages
7-9 segmented, equal to height of basal segment of pedicel of cirrus VI.
Remarks. — Re-examination of the type material as well as of new collections of specimens of Pacliylasma
ferrugomaculosa have revealed that the number of shell plates in the adult condition is variable within this species.
The adult form may be 6-plated (rostral plate, CL1 and CL2 distinct entities or partially coalesced, and C) or
4-plated (rostral plate, CL1 and CL2 totally coalesced, and C). Comparing wall, opercular plate and soft body
morphology, T. arcuatum sp. nov., T. aurantiacum (Darwin, 1854), T. ornatum sp. nov. and T. trigonum Foster,
1988 are closely related to P. ferrugomaculosa ( sensu JONES, 1993b) and the latter is now included within
Tetrapachylasma. JONES (1993b) noted the similarity between P. ferrugomaculosa and P. aurantiacum.
Tetrapachylasma ferrugomaculosa is a moderate sized species (maximum: RC length 1 1.4, C height 10.2).
Differences between T. ferrugomaculosa, T. arcuatum sp. nov. and T. aurantiacum (Darwin, 1854) are listed under
T. arcuatum and T. aurantiacum , respectively. Soft parts of T. ornatum are unknown; thus only hard parts can be
compared with T. ferrugomaculosa. In T. ornatum the S basal margin is 1/2 the length of the occludent margin,
the articular ridge is not projecting and the growth ridges are complexly sculptured, cut by 4 deep, longitudinal
striae. In T. ferrugomaculosa the S basal margin is 7/10 the length of the occludent margin, the articular ridge is
barely projecting and the convoluted scutal growth ridges are cut by 6-8 longitudinal striae. The T articular ridge of
T. ferrugomaculosa is not projecting, the T spur is indistinct. 1/2 the width of the basal margin and set at
a distance of 1/12 the width of the basal margin from the basi-scutal angle, with 7-9 muscle crests projecting well
beyond the basal margin. In T. ornatum the T articular ridge is projecting, and the spur is short, more than 1/3
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOI DEA AND PACHYLASMATOIDEA
223
the width of the basal margin and set at a distance of 1/12 the width of the basal margin from the basi-scutal
angle, with 8-12 muscle crests projecting from the basal margin. Tetrapachylasma ornatum is small in size (RC
length 6.0, C height 2.5), whereas T. aurantiacum is larger (RC length 1 1.4, C height 10.2).
Tetrapachylasma ferrugomaculosa can be separated from T. trigonum by the form of the caudal appendages
(7-9 segmented, equal to the height of the basal segment of the pedicel of cirrus VI, and uniarticulate. 1/8 the
height of the basal segment of the pedicel of cirrus VI, respectively). The paries of T. ferrugomaculosa are without
narrow ribs; in T. trigonum the surface of the paries have narrow ribs peripherally. In T. ferrugomaculosa the S
basal margin is 7/10 the length of the occludent margin, the articular ridge is barely projecting and the scuta!
growth ridges are very convoluted, and cut by 6-8 longitudinal striae. The S basal margin of T. trigonum is 5/8 the
length of the occludent margin, the articular ridge is not projecting and the scutal growth ridges are ruffled and
overlapping, and not cut by longitudinal striae. The T articular ridge of T. ferrugomaculosa is not projecting,
the T spur is indistinct, 1/2 the width of the basal margin and set at a distance of 1/12 the width of the basal
margin from the basi-scutal angle, with 7-9 muscle crests projecting well beyond the basal margin. In T. trigonum
the T articular ridge is projecting, and the spur is 1/3 the width of the basal margin and set at a distance of 1/7
the width of the basal margin from the basi-scutal angle, with 6 muscle crests slightly projecting from the basal
margin, and the basal corner of the articular ridge is hollowed out for the reception of dwarf males.
Distribution. — South-western Australia, 3-30 m.
Tetrapachylasma ornatum sp. nov.
Figs 39, 43-44; Tables 15-17
Material EXAMINED. — Marianas Islands. The Grotto. Puntan, Madog, Saipan, 6 m. 27.07.1969:
2 specimens attached to the sclerosponge Astrosclera sp. (YPM 9303 A. Bl; I specimen attached to the sclerosponee
Astrosclera sp. (YPM 9305).
Types. — Holotype: YPM 9303 A (The Grotto, Puntan, Madog, Saipan, 6 m, 27.07.1969).
Paratypes: YPM 9303 B (The Grotto, Puntan, Madog. Saipan. 6 m, 27.07.1969). — YPM 9305 (The Grotto.
Puntan, Madog, Saipan, 6 m. 27.07.1969).
Diagnosis. — R and RL closely united by inconspicuous linear sutures; basal width of R equal to hasal width
of RL; RL clearly incorporated in the sheath. Basal width of CL2 slightly less than basal width of CL1; CL' and
CL- partially or totally coalesced. S basal margin sinuous, externally growth ridges complexly sculptured, cut by
4-6 deep radial striations; articular ridge not projecting beyond articular margin. T articular ridge projecting;
8-12 strong muscle attachment ridges extending well beyond basal margin.
Description. — Size small (maximum: RC length 6.0. C height 2.5). Shell form low conical, higher at
cartnal end; almost circular when viewed from above. Parietal plates externally differentiated in adult as 6 plates -
rostral plate, paired CL1 and CL2 (sometimes partialy coalesced) and C - or 4 plates (rostral plate. CL 'and CL2
coa csced. C), plates solid, thicker basally, internally calcareous pegs developed along basal margins of paries:
externally growth ridges ill-defined, irregularly scalloped, more regular and defined on rostral plate; outer surface of
paries rough. Alae of C and CL1 wide, triangular, well-developed; alae of CL2 minute or absent; alar growth ridges
wi e y spaced, parallel to inferior alar margin; superior alar margins serrate; alar welling absent. Rostral plate
road, convex, smaller than C; apex pointing inward; sutures between R and RL not or slightly visible externally
an | interna y. In adults CL' and CL2 coalesced (4-plated condition) or incompletely coalesced (6-plated condition);
• 'ar8er l*ian CL2; apices sloping inward, toward C. CL2 triangular; apex sloping inward,
oward C. C largest, highest plate, occupying almost 1/2 circumference of shell; apex pointed. V-shaped from
Ve’ P al° s opm£ ‘nward toward orifice; alae wide. Basis unknown, presence of calcareous pegs alone basal
argms of par.etes indicative of interdigitation with calcareous basis. Orifice large, triangular, not toothed.
( ™ , , gf,C tr‘angUlar; basal mar8in sinuous, 3/7 length of occludent margin; externally growth ridges
p ex y sculptured, cut by 4 deep radial striations; apex pointed; internally smooth, adductor muscle pit absent.
224
D. JONES
C
Fig. 43. — Tetrapachylasma omatum sp. nov. Hololype from Puntan, Madog, Saipan (YPM 9303A): a. T. external view;
b, T, internal view; c. S. external view; d. S, internal view.
depressor muscle pits indistinct; articular furrow broad, deep; articular ridge high, not projecting beyond articular
margin; adductor ridge indistinct. T wider than S, triangular, basal margin twice height of valve; externally with
regular growth ridges; internally with 8-12 strong muscle attachment ridges extending well beyond basal margin;
articular ridge projecting beyond articular margin; articular margin with growth ridges raised to form elevated ridge;
articular furrow broad, deep; spur furrow shallow, spur 1/3 width of basal margin, truncate, pointing toward and set
a distance of 1/12 length of basal margin from basi-scutal angle. Color of parietal plates, alae and opercular plates
buff with dull ochre/orange. Measurements (approximate) of holotype:
Soft parts not known presently.
Remarks. — During the course of the present study 3 dry specimens from the Marianas Islands were sent to
me, for comparative purposes, by Professor W. Newman. The cirripeds had been collected by T. Goreau and
J. Lang who had sent them, and the associated sclerosponge, to Willard Hartman of YPM for identification.
The cirriped material is herein described as Tetrapachylasma omatum sp. nov. On examination it is clear that the
RL does enter the sheath, a unique attribute not found in any other tetrapachylasmatid or pachylasmatoid species
presently known. Apart from this unique character, the complex S sculpturing separates T. oruatum from all other
species in the genus and warrants specific status. Differences between T. omatum sp. nov. and T. arcuatum
sp. nov., T aurantiacum and T. ferrugomaculosa are listed under and T. arcuatum sp. nov., T aurantiacum and
T. ferrugomaculosa, respectively. Details of the soft part anatomy of T. omatum are presently unknown.
Tetrapachylasma omatum differs from T. trigonum in the form of the parietal plates (narrow peripheral ribs
present in T. trigonum, absent in T. omatum). The S basal margin of T. trigonum is 5/8 the length of the
occludent margin, the articular ridge is not projecting and the scutal growth ridges are ruffled and overlapping, and
not cut by longitudinal striae. In T. omatum the S basal margin is 1/2 the length of the occludent margin, the
articular ridge is not projecting and the complexly sculptured scutal growth ridges are cut by 4-6 deep, longitudinal
striae. The T articular ridge of T. omatum is projecting; the spur is indistinct, more than 1/3 the width of the basal
margin and set at a distance of 1/12 the width of the basal margin from the basi-scutal angle; and there are
8-12 muscle crests projecting from the basal margin. In T. trigonum the T articular ridge is projecting; the spur is
Source MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
225
1/3 the width of the basal margin and set at a distance of 1/7 the width of the basal margin from the basi-scutal
angle; and there are 6 muscle crests slightly projecting from the basal margin. Additionally, the basal corner of
the articular ridge is hollowed out for the reception of dwarf males in T. trigonum. Tetrapachylasma ornatum is
small in size (RC length 6.0, C height 2.5), T. aurantiacum is larger (RC length 10.0. C height 8.0).
FlG. 44. — Tetrapachylasma ornatum sp. nov.
a‘d- — Paratype from Puntan, Madog, Saipan (YPM 9305). Wall plates: a. C from above; b, CL-, internal view; c, CL1
internal view; d, R+RL+ missing RL, internal view.
e-i. — Holotype from Puntan, Madog, Saipan (YPM 9303A). e-h, Opercular plates: e. S. internal view; f. S. external
view; g, T, internal view; h. T. external view. — i, Compound rostrum, sutures absent, internal view.
Paratype from Puntan, Madog, Saipan (YPM 9303B). Compound rostrum, sutures present, internal view.
Etymology. — From the Latin ornatus , ‘adorned’, in reference to the complex sculpting of the growth ridges
on the S.
Distribution. — Marianas Islands, 6 m.
226
D. JONHS
Tetrapachylasma trigonuin Foster, 1988
Fig. 39; Tables 15-17
Tetrapachylasma trigonum Foster, 1988: 226. figs 1-3.
Material EXAMINED. — Cook Islands. Off Avarua. Rarotonga, 15 m, 09.1984: 2 paratypes, under coral rubble
(NMNZ Cr 3641).
Types. — Holotype : NMNZ CR 3640; Cook Islands, off Avarua, Rarotonga, 15 m, 09.1984; 2. dissected
appendages on microslide with dry disarticulated plates kept separately.
Paratypes: NMNZ Cr 3641; Cook Islands, off Avarua. Rarotonga, 15 m, 09.1984; 2 whole specimens,
attached to piece of coral rubble, in alcohol.
Holotype and paratype depository: NMNZ.
Diagnosis. — Adult with 4 parietal plates (rostral plate, CL' and CL2 coalesced, and C); plates solid, thin,
externally finely stellate basaily, outer surface of paries with narrow ribs peripherally. S articular ridge not
projecting; external growth ridges ruffled, overlapping, not cut by longitudinal striae. T articular ridge wide,
projecting wing-like from articular margin, basal corner hollowed for reception of dwarf males; 6 muscle
attachment ridges extending just beyond basal margin. Caudal appendages uni-articulate. 1/8 height of basal
segment of pedicel of cirrus VI.
Remarks. — The 4 plates composing the adult shell, the basal corner of the T articular margin being
hollowed for the reception of dwarf males, and the vestigial caudal appendages, distinguish this species.
Tetrapachylasma trigonum is a moderate sized species (maximum: RC length 10.0, C height 6.4). Differences
between 7. trigonum and T. arcuatum sp. nov., T. aurantiacum, T. ferrugomaculosa and T. ornatum sp. nov. are
listed under T. arcuatum, T. aurantiacum, T. ferrugomaculosa and T. ornatum, respectively.
Distribution. — Cook Islands, 15 m.
Subfamily METALASMATINAE nov.
Tables 1-2
Diagnosis. — Primary shell wall not surrounded by whorl(s) of imbricating plates; adult with 6 solid,
calcareous, loosely articulated plates (R-CU-CL2-C); thin lamina of chitin dividing paries into inner and outer
laminae; externally paries with prominent, irregular growth ridges lined with small setae; external alar growth
ridges sinuously parallel to inferior alar margin; superior margins of C alae thickened in distal halves and. to lesser
extent, superior margins of alae of CL1 and CL2 thickened in distal 1/3, giving dentate appearance; basis
membranous; articular ridges of S and T prominent, articular furrow of T well developed; caudal appendages
present; mandible tridentoid.
Type Genus. — Metalasma gen. nov.
Remarks. — Only 1 genus, Metalasma, is assigned to this subfamily. The metalasmatines have lost any trace
of the suture between the R and RL but retain the more plesiomorphic pachylasmatoid basic 6-plated wall. The
body morphology of the Metalasmatinae is intermediate between the Pachylasmatinae (retention of the caudal
appendages, a tridentoid mandible, form of T) and the Hexelasmatinae (chitin in the parietal wall, form of the cirri -
cirrus II resembling I more than III). The presence of caudal appendages, the presence of chitin in the parietal wall,
and the tridentoid nature ot the mandible, separate the Metalasmatinae from the Bathylasmatinae, which is also
phylogenetically between the Pachylasmatinae and the Hexelasmatinae.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
227
ETYMOLOGY. — From the Greek meta, 'between', in reference to the intermediate position of this subfamily
within the Pachylasmatidae.
DISTRIBUTION. — Recent; Western Pacific: Loyalty Islands; New Hebrides Arc; 300-500 m; attached to stones
and Stylasteridae.
Genus METALASMA gen. nov.
Fig. 45; Tables 18-19
Diagnosis. — As for the subfamily, which is monogeneric.
Type Species. — Metalasma crassum sp. nov.
Remarks. — One species is presently assigned to this genus.
Tables 18 and 19. — Metalasma gen. nov.: characters of M. crassum sp. nov.
artic. = articular; b-s = basi-scutal; distrib. = distribution; ext. = external; int. = internal; max. = maximum; n. a. = not applicable; prop =
projecting; segs = segments.
Distribution. — Western Pacific: Loyalty Islands, New Hebrides Arc; 300-500 m; attached to stones and
Stylasteridae.
Fig. 45. — Metalasma gen. nov. Distribution map: •, M. crassum (300-500 m).
228
D. JONES
Metalasma crassum sp. nov.
Figs 45-48; Tables 18-19
Material EXAMINED. — New Hebrides Arc. Volsmar: sin DW 16. 500 m: 8 specimens (MNHN-Ci 2708).
Loyalty Islands. Musorstom 6; stn DW 460, 420 m: many specimens, attached to rocks (MNHN-Ci 2701). —
Stn DW 476, 300 m: 2 specimens (ovigerous), attached to stone (MNHN-Ci 2408, Ci 2409). Drawn. —
Stn DW 478, 400 m: 1 specimen (broken), attached to Stylasteridae (MNHN-Ci 2505).— Stn DW 479, 310 m: many
specimens, attached to rocks (WAM C 23252).
Types. — Holotype : MNHN-Ci 2408 (MUSORSTOM 6, stn DW 476).
Paratypes : MNHN-Ci 2409 (MUSORSTOM 6, stn DW 476). Drawn. — MNHN-Ci 2505 (MUSORSTOM 6,
stn DW 478). — MNHN-Ci 2701 (MUSORSTOM 6, stn DW 460). — MNHN-Ci 2708 (VOLSMAR, sin DW 16).
— WAM C 232-52 (MUSORSTOM 6, stn DW 479).
Diagnosis. — Shell form low, rounded from above. Parietal plates with thin chitinous laminae dividing paries
into inner lamina (1/4 thickness) and outer lamina (3/4 thickness). Basal width of CL? 1/2 basal width of CL'.
Narrow remnants of radii on CL1, CL2 and rostral plate. Superior alar margins oblique; carinal superior alar
margins thickened in distal 1/2 giving dentate appearance, to lesser extent superior alar margins of CL1 and CL2
thickened in distal 1/3. Basis membranous. S externally with many regular growth ridges, not cut by longitudinal
striae; articular ridge projecting. Tergal spur indistinct; 13-16 muscle attachment crests, some extending below
basal margin; articular ridge projecting. Caudal appendages twice length of pedicel of cirrus VI; 10-12 segmented.
Fig. 46 .—Metalasma crassum sp. nov. Paratype from MUSORSTOM 6, stn DW 476 (MHNH Ci 2409): a. S, internal view;
b. S, external view; c, T, external view; d, T, internal view; e. whole animal, lateral view; f, whole animal, from
above.
description. — Size small (maximum: RC length 6.5, LD 6.9, C height 2.2, R height 2.6). Parietal plates
thick basally, externally roughened, irregular growth ridges widely spaced, lined with small setae; chitinous
laminae dividing paries into inner and outer laminae, inner lamina 1/4 thickness of outer lamina. Rostral plate
Battened, angled inward, apex not curved in toward orifice, basal width twice basal width of C. CL' and CL2
Source : MNHNJ Paris
CIRR1PEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
229
separate; basal width of CL1 just greater than 1/2 basal width of R, apex not curved in toward orifice; basal width
of CL2 1/2 basal width of CL1, height less than CL1 by 1/10; apex not curved in toward orifice. C highest plate,
basal width subequal to basal width of CL1, apex bowed, spout-like, curved away from orifice. Superior alar
margins oblique, growth ridges sinuously parallel to inferior alar margins; alae of C widest, alae of CL1 1/3 width
of carinal alae. alae of CL2 1/2 width of carinal alae; carinal superior alar margins thickened in distal 1/2 giving
dentate appearance, to lesser extent superior alar margins of CL1 and CL2 thickened in distal 1/3. Narrow remnants
of radii on CL1, CL2 and rostral plate. Basis membranous, flat. Orifice pentagonal, slightly toothed. Opercular
plates lodged subparallel to basis. S triangular; basal margin 3/4 length of occludent margin; externally with many
spaced, regular growth ridges, not cut by longitudinal striae; apex pointed; internally adductor muscle scar at 1/2
height of valve; lateral depressor muscle pit well developed; articular furrow moderately deep; articular ridge long,
projecting well beyond articular margin; adductor ridge indistinct. T subequal to S, sub-triangular; externally with
fine growth ridges; internally with 13-16 depressor muscle crests, some extending just beyond basal margin;
prominent tergal articular ridge projecting well beyond and standing clear of convex articular margin; articular
furrow broad, deep; indistinct spur, roundly truncate, 1/3 width of basal margin, set at distance of 1/6 width of
basal margin from basi-scutal angle. Color of preserved material creamy yellow, opercular plates more yellow;
eggs yellow. Measurements of 2 specimens examined as follows:
Labrum with shallow medial depression and
numerous small, pointed teeth. Mandibular palp
oblong; dense setae terminally. Mandible with
3 teeth, tooth 1 largest, separated from tooth 2 and 3;
upper margins of 2 and 3 with small subsidiary
cusps; inferior angle large, bluntly molariform,
dentate. Maxillule setose; 2 pairs of long, stout setae
at upper angle; notch below upper angle large, with
3-4 pairs of smaller setae; straight, stepped cutting
margin below with 5-6 pairs of larger setae; inferior
angle small, with 4-5 pairs of smaller setae. Maxilla
wide; endites indistinct; long, serrulate setae
terminally.
Cirrus I with rami subequal; anterior ramus
slightly longer than posterior; proximal segments of
anterior ramus protuberant anteriorly; both rami
moderately setose, setae finely serrulate distally.
Cirrus II similar to but slightly longer than cirrus I; rami subequal; proximal segments of anterior ramus slightly
piotuberant anteriorly; posterior ramus with distal segments antenniform; rami with segments moderately setose,
some distal setae serrulate. Cirrus III longer than cirrus II, with rami subequal, segments becoming oblong
distally; posterior ramus with 2-4 pairs of long, finely serrulate setae on anterior faces of distal segments, posterior
laces with short, spine-like setae; anterior ramus with 2-3 pairs of long, finely serrulate setae on anterior faces of
distal segments, proximal segments with 5 pairs. Cirrus IV to VI similar, longer than cirrus III; segments oblong,
with 4-5 pairs of setae on anterior faces, distal 2 pairs of setae longest, finely serrulate. Chaetotaxy ctenopod.
Cirral lormula as follows:
Fig. 47. — Meialasma crassum sp. nov. Holotype from
Musorstom 6, stn DW 476 (MNHN-Ci 2408): lateral
view. Scale = 5 mm.
230
D. JONES
Caudal appendages twice length of pedicel of cirrus VI; 10-12 segmented; apically with sparse, long setae,
distal margins of segments with sparse setae, setal length more than 1/2 segmental length. Penis length of
proximal segment of pedicel of cirrus VI; finely annulated; sparsely setose with irregular circlet of sparse, short
setae apically. Eggs large, ovoid, 0.36 x 0.24 mm.
Fig. 48. — Meiatasma crassum sp. nov. Paratype from Musorstom 6, stn DW 476 (MHNH Ci 2409): a, maxilla;
b.labrum; c, right mandible; d. left mandible; e, right maxillule; f. left maxillule; g. mandibular palp; h. cirrus I;
i. cirrus II, j, cirrus III, posterior ramus, median segments; k, cirrus VI, posterior ramus, median segments;
1, cirrus VI and caudal appendage; m, cirrus III; n, cirrus I. posterior ramus, median segments; o, penis.
Source : MNHN Paris
CIRRIPEDIA THORACICA; CHIONELASMATOIDEA AND PACHYLASMATOIDEA
231
REMARKS. — Specimens of M. crassum sp. nov. have been collected, by VOLSMAR and MUSORSTOM 6. from
five separate stations which are spatially close. Metalasma crassum is remarkable in having a prominent tergal
articular ridge extending well beyond and standing clear of the convex articular margin, and an extremely well
developed, broad and deep articular furrow. The manner of articulation of the S and T ensure that the T fits closely
into the trough formed by the broad carinal alae and the apical portion of the C, and is almost unexposed when the
opercular plates are closed.
Etymology. — From the Latin crassa, ‘thick’, in reference to the thickened areas on the upper edges of the
alae of the C and CL2 of this species.
Distribution. — New Hebrides Arc, Loyalty Islands, 300-500 m.
Subfamily BATHYLASMATINAE Newman & Ross, 1976
Tables 1-2
Diagnosis. — Primary shell wall not surrounded by whorl(s) of imbricating plates. Adult shell wall with
6 (R-CL'-CL—C), or4(R-CL-C), solid compartmental plates; paries with prominent horizontal growth ridges
lined with small setae; plates thin ( Bathylasma ) or thickened (. Mesolasma ); chitinous laminae in parietal plates
absent. External alar growth ridges diverging from inferior alar margin. Basis membranous or calcareous. S with
prominent articular ridge; “balanoid" in form. T articular ridge varies from not prominent to prominent. Mandible
quadridentoid. Caudal appendages absent.
Distribution. — Paleocene to Recent; Cosmopolitan.
Remarks. — Newman and Ross (1971) proposed the Bathylasmatidae, in which they included Bathylasma,
Hexelasma, Aaptolasma, Tetrachaelasma and Tessarelasma. FOSTER (1978) indicated that there was only scant
evidence for generic differentiation between Hexelasma and Bathylasma, these genera differing principally in the
shape of the spur of the T and the number of paired setae on the anterior edge of the segments of cirrus VI. FOSTER
(1978) also showed that there was considerable confusion concerning some of the holotypes of Hoek and
suggested that more than 1 taxon may have been involved in the initial description of H. velutinum Hoek, 1913.
FOSTER (1981) subsequently re-examined the type material, concluding that li. velutinum was an aaptolasman and
placing Aaptolasma in Hexelasma as a junior synonym. FOSTER (1981) further proposed the Mesolasma for a
genus intermediate between Bathylasma and Hexelasma. Mesolasma differed from Bathylasma in having opercular
plates which lacked prominent articular ridges and from Hexelasma in the less specialised setation of the segments
of the posterior cirri.
BUCKERIDGE (1983) remarked that the bathylasmatines retained the more primitive pachylasmatoid basic shell
construction but had lost any trace of the suture between the R and the RL and that the body morphology was
intermediate between the Pachylasmatinae and the Hexelasmatinae. BUCKERIDGE (1983) suggested that Bathylasma
be retained for those forms with pachylasmatid-like lerga, solid calcareous compartments and normal chaetotaxy of
the posterior cirri.
Hie following genera are included within the Bathylasmatinae herein: Batlivlasma, Mesolasma, Tetrachaelasma
and Tessarelasma.
Genus BATHYLASMA Newman & Ross, 1971
Fig. 49; Tables 20-21
Bathylasma Newman & Ross, 1971: 143.
Diagnosis. — Adult with fixed shell pattern of 6 thin, calcareous compartmental plates (R-CL'-CL2-C);
parietes with prominent horizontal growth ridges lined with small setae; chitinous laminae absent. External alar
232
D. JONES
ridges diverging from inferior alar margin; superior alar margin with welting. Basis membranous. Articular ridges
of T and S prominent. Mandible quadridentoid. Caudal appendages absent.
Type Species. — Balanus corolliformis Hoek, 1883.
RECENT Species. — Bathylasma alearum (Foster, 1978) Eastern Australia, New Zealand, Vanuatu; 414-
1750 m; B. corolliforme (Hoek, 1883) (includes Hexelasma antarcticum Borradaile, 1916) circum Antarctic;
to 1464 m; Pleistocene, to 70 m above sea level.; B. hirsutum (Hoek, 1883) North-east Atlantic, from Faeroe Is
south to Azores; 944-1829 m.
Fossil Species. — Bathylasma aucklandicum (Hector, 1888) Oligocene to Pliocene (Duntroonian or
Waitakian to Waitotaran); New Zealand. Bathylasma rangatira Buckeridge, 1983 Lower Palaeocene to Lower
Eocene; Chatham Islands.
Remarks. — Members of this genus have solid shells, pachylasmatid-like T and normal chaetotaxy of the
posterior cirri (FOSTER, 1981).
Tables 20 and 21. — Bathylasma : characters of the species
arI'lr: - articular; b-s = basi-scutal; c.a. = caudal appendage; ext. = external; int. = internal; long. = longitudinal; max. = maximum; n a = not
applicable; prop = projecting; prom. = prominent; segs = segments.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
233
DISTRIBUTION. — Palaeocene to Recent. Lower Palaeocene to Lower Eocene: Chatham Islands; Oligocene:
New Zealand, Victoria; Miocene to Pliocene: New Zealand; Lower Miocene: Burma; Pleistocene to Recent in
Antarctica.
Recent: North Atlantic and oceans bordering Antarctica (deep seas): Antarctica, to 1464 m (B. coroll forme)-.
New Zealand, eastern Australia, Vanuatu, 414-1750 m ( B . alearum)-, N E Atlantic, 944-1829 m (B. hirsutum).
Fig. 49. — Baihylasma Newman & Ross, 1971. Distribution map: O, B. corolliforme (to 1464 m); ▲. B. alearum (414-
1750 m); •, B. hirsutum (944-1829 m).
Bathylasma alearum (Foster, 1978)
Fig. 49: Tables 20-21
Hexelasma alearum Foster, 1978: 80, fig. 49 A-L, pi. 10 A-B.
Bathylasma alearum - FOSTER, 1981: 356.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn DW 1062, 619-658 m: 2 specimens (broken).
New Zealand. NZOI: sin F 132, 49°59'S, 177°32'E, 1335 m, 29.01.1965: holotype, on sponge Epipolasis
novaezelandiae (NZOI 205).
Australia. Vessel unknown: stn BSS 641 Q, SSE Cape Conran, Victoria, 38°46'S. 148°34.8'E. 1750 m, 17.11.
1981: several specimens (AM P 40930).
Types. — Holotype: NZOI 205; New Zealand, NZOI stn F 132, 49°59'S, 177°32'E, 1335 m, 29.01.1965: on
sponge Epipolasis novaezelandiae-, sandy substrate.
Holotype depository’-. NIWA.
Diagnosis. — Parietal plates not flared out from orifice; prominent rows of setae on prominent external
growth ridges. Basal width of CL2 1/4 basal width of CL1. Conspicuous narrow alar welting along superior alar
margin; alar growth lines not parallel with inferior alar margin, leaving smooth, triangular extension of internal
alar surface on outside towards alar angle beneath growth lines. S externally with regular, well-spaced growth lines,
not cut by longitudinal folds; basal margin 7/10 length of occludent margin; articular ridge projecting. T articular
ridge projecting well beyond articular margin; spur narrowly rounded, 1/7 width of basal margin, set at 1/14 width
234
D. JONES
of basal margin from basi-scutal angle; 8-9 strong muscle crests projecting just below basal margin. Mandible
with teeth 2-4 with small side teeth, small subsidiary cusps on upper and lower margins of tooth 4.
Remarks. — During the final preparation of this paper, A. CROSNIER showed me several samples recently
collected by the Musorstom 8 expedition in the waters of Vanuatu. Of this material, 2 specimens are identified as
Bathylasma alearum (Foster, 1978). This record extends the range of B. alearum in the waters of the south-western
Pacific Ocean, from eastern Australia and New Zealand northwards to Vanuatu.
Bathylasma alearum differs from B. corolliforme in a number of characters. In B. alearum the parietal plates are
not flared out from the orifice, but they are flared in B. corolliforme. The basal width of CL2 is 1/4 the basal
width of CL1 in B. alearum , and 1/10 the basal width of CL1 in B. corolliforme. In B. alearum the alar growth
lines are not parallel with the inferior alar margin; in B. corolliforme they are parallel with the inferior alar margin.
The S scutal growth lines of B. alearum are wide and regular, rather than regular and well-spaced in the upper half
of the valve and narrowly spaced in the lower half (B. corolliforme). In B. alearum the T is externally fiat, rather
than concave, as in B. corolliforme. The T spur of B. alearum is distinct, 1/7 the width of the basal margin and set
at a distance of 1/14 the length of the basal margin from the basi-scutal corner, with 8-9 muscle attachment crests
projecting below the tergal basal margin; in B. corolliforme it is indistinct, set at the basi-scutal corner, and
the 8-9 muscle crests do not project below the basal margin. In B. alearum mandibular teeth 2-4 have small side
teeth, and there are small subsidiary cusps on the upper and lower margins of tooth 4; the mandibular teeth of
B. corolliforme have no side teeth.
Bathylasma alearum is similar to B. hirsutum as in both species the parietal plates are not flared out from the
orifice and the alar growth lines are not parallel with the inferior alar margin, but there are the following differences
between these species. In B. alearum the basal width of CL2 is 1/4 the basal width of CL1, in B. hirsutum 1/7 the
basal width of CL1. The alar welting in B. alearum is conspicuous and narrow; in B. hirsutum it is wide, Hat and
1/2 the alar width. The T spur of B. alearum is distinct, 1/7 the width of the basal margin and set at a distance of
1/14 the length of the basal margin from the basi-scutal corner, with the muscle crests projecting below the basal
margin; in B. hirsutum it is 1/6 the width of the basal margin and set at a distance of 1/12 the length of the basal
margin from the basi-scutal corner, with the muscle crests not projecting below the basal margin. The mandible of
B. alearum has mandibular teeth 2-4 with small side teeth, and small subsidiary cusps on the upper and lower
margins of tooth 4; in B. hirsutum the lower margin of mandibular tooth 4 has small subsidiary cusps.
Foster (1978) noted similarities between B. alearum and the fossil B. aucklandicum , distinguishing the
2 species on size and the shape of the T articular ridge. Buckeridge (1983) similarly considered B. aucklandicum
to closely resemble B. alearum.
Distribution. — New Zealand, eastern Australia, Vanuatu; 414-1750 m.
t Bathylasma aucklandicum (Hector, 1888)
Scalpellum aucklandicum Hector, 1888: 440.
Pollicipes (?) aucklandicus - Benham, 1903: 1 1 1. — Park. 1910: 115, pi. 7 figs 3-7, 10-11.
PolUcipes aucklandicus - Clarke. 1905: 419.
Hexelasma aucklandicum - WITHERS, 1913: 841, pi. 86 fig. 1; 1924: 19, figs 7-8, pi. 4 figs 1-4, pi. 5 fig. I. -
Buckeridge, 1975: 121. figs 4 c, 5.1, 5.2. — Foster. 1978: 82, pi. 10 C-E.
Bathylasma aucklandicum - Newman & Ross, 1971: 151, pi. 25; 1976: 45; 1981: 356.
Types .—Lectotype: GS (locality no. 695); rostrum, figured on PI. LXXXV fig. 1 .
Syntypes : GS (locality no. 695); 6 rostral compartments, 9 carinal, 13 laterals (8 right and 5 left). 5 carino-
laterals (3 right and 2 left), 1 tergum (PI, LXXXV figs 2-13).
Lectotype and syntypes depository. GS.
diagnosis. — Shell tall, cylindrical; aperture not flared. Parietal plates solid, calcareous, thin. External alar
growth lines line, diverging from inferior alar margin before turning abruptly downwards towards alar angle then
Source : MNHN, Paris
CIRRIPED1A THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
235
deflecting apically to form narrow welting along superior alar margin. Sheath insignificant or wanting. S flat with
low articular ridge not projecting. T with elevated articular ridge projecting, short spur set close to basi-scutal
angle.
Remarks. — See remarks under B. alearum for similarities between B. aucklandicum and B. alearum.
Distribution. — Oligocene to Pliocene (Duntroonian or Waitakian to Waitotaran); New Zealand.
Bathylasma corolliforme (Hoek, 1883)
Fig. 49. Tables 20-21
Balanus corolliformis Hoek, 1883: 155, pi. 6 figs 21-22, pi. 13 figs 1-7.
Hexelasma corolliforme - Hoek, 1913: 245. — PlLSBRY, 1916: 330. — NlLSSON-CANTELL, 1930: 252, figs 11-1. —
Zevina, 1968: 94, fig. 6 a-i.
Bathylasma corolliforme - Newman & Ross, 1971: 143, figs 71-72, pis 15-22, 43 A; 1976: 46. — FOSTER, 1978: 80,
fig. 48 A-L; 1981: 356. — BUCKERIDGE, 1989: 333.
Hexelasma antarcticum Borradaile, 1916: 132, fig. 7. — Withers, 1924: 22. — Bage, 1938: 8, pis 5-8. — Speden, 1962:
746. — Weisbord, 1965: 1015 (part.). — Utinomi, 1965: 4, figs 1-4, pis 1-2. — Weisbord. 1967: 51 (part.), figs 1-
6, pis 1-2. — Zevina, 1968, fig. 1.
TYPES. — Holotype : BMNH 85.17; near Kerguelen I., "Challenger'’: stn 150, 52°4'S, 71°22’E, 274 m,
02.02.1874.
Paratype: BMNH 1917.2.9.59; "Terra Nova" Expedition.
Holotype and paratype depository. BMNH. (paratype specimen has not been located in the collection of
BMNH) [Ms A. Morgan, pers. comm., 05.02.1996).
Material EXAMINED. — Near Kerguelen Island. "Challenger": stn 150, 52°04'S, 7I°22’E, 274 m, 02.02.
1874: holotype (BMNH 85.17).
Antarctic. Anzare: stn III, 66°32'S, I41°39'E, 297 m, 31.12.1913: 1 specimen (AM P 7206); 1 specimen (AM P
7207); 1 specimen (AM P 7209); 1 incomplete specimen (AM P 7210); 1 specimen (AM P 7211). All identified
Hexelasma antarcticum Borradaile. 1916.
DIAGNOSIS. — Parietal plates flared out from orifice; rows of short setae on persistent membrane above coarse,
external growth ridges. Basal width of CL2 1/10 basal width of CL1. Alae oblique; conspicuous, narrow alar
welting along superior alar margin; alar growth lines parallel with inferior alar margin. S externally concave, with
regular, well-spaced growth lines in upper half of valve, growth lines narrowly spaced in lower half, growth lines
not cut by longitudinal folds; S basal margin 3/4 length of occludent margin; articular ridge slightly projecting.
T externally concave, articular ridge projecting well beyond articular margin; spur indistinct, set almost at basi-
scutal angle; 8-9 muscle crests not projecting below basal margin. Mandible with 4 teeth, small subsidiary cusps
on upper and lower margins of teeth absent.
Remarks. The flaring orifice, the coarse growth ridges on the parietes and the alar growth ridges running
along and closely paralleling the inferior alar margins distinguish this species. Differences between B. corolliforme
and B. alearum are listed under B. alearum.
Bathylasma corolliforme differs from B. hirsutum as the parietal plates are flared out from the orifice in the
former, rather than not flared in the latter; the growth lines of the parietal plates are coarse, rather than fine; and the
alar growth lines are parallel, rather than not parallel, with the inferior alar margin. In B. corolliforme the basal
width of CL2 is 1/10 the basal width of CL>, in B. hirsutum 1/7 the basal width of CL'. The alar welting in
B- corolliforme is conspicuous and narrow; in B. hirsutum it is wide, flat and 1/2 the alar width. The S sculal
crowt i incs are regular and well-spaced in the upper half of the valve and narrowly spaced in the lower half in
. corolliforme ; in B. hirsutum they are uniformly wide and regular. The T spur of B. corolliforme is indistinct,
SCt. 3 ™?stat t*le basi-scutal corner, with the muscle crests not projecting below the basal margin; in B. hirsutum
i is 1/6 the width of the basal margin and set at a distance of 1/12 the length of the basal margin from the
basi-scutal corner, with the muscle crests not projecting below the basal margin. The mandible of B. corolliforme
236
D. JONES
has no subsidiary cusps on the upper and lower margins of tooth 4; in B. hirsutum the lower margin of mandibular
tooth 4 has small subsidiary cusps.
Distribution. — Circum Antarctic, to 1464 m; Pleistocene, to 70 m above sea level.
Bathylasma hirsutum (Hoek, 1883)
Fig. 49; Tables 20-21
B alarms sp. Jeffreys, 1878: 414 ( fide Southward & Southward, 1958).
Bathylasma hirsutus Hoek, 1883: 158, pi. 13 figs 8-15. — Gruvel, 1920: 55. pi. 1 figs 13-14.
Hexelasma hirsutum - HOEK, 1913: 245. — PlLSBRY, 1916: 330. — SOUTHWARD & SOUTHWARD, 1958: 635. — UTINOMI,
1965: 1 1. — Foster, 1978: 79.
Batlivlasma hirsutum - Newman & Ross, 1971: 149, fig. 73. pis 23-24; 1976: 46. — Foster, 1981: 356. — Young,
1998: 66.
Types. — Holotype : BMNH 85.17; Faeroe Channel, "Triton": stn 10, 59°40’N, 7°21’W. 944 m, 24.10.1882.
Holotype depository’: BMNH.
Material EXAMINED. — Faeroe Channel. "Triton": stn 10, 59°40'N, 7°2r\V. 944 m, 24.10.1882: holotype
(BMNH 85.17).
DIAGNOSIS. — Parietal plates thin, not flared out from orifice; rows of setae on yellow membrane above fine
external growth ridges. CL1 and CL2 separate; basal width of CL2 1/7 basal width of CL1. Wide, flat alar welting
along superior alar margin, 1/2 alar width; alar growth lines not parallel with inferior alar margin. S externally
with regular, well-spaced growth lines, not cut by longitudinal folds; basal margin 5/8 length of occludent margin;
articular ridge projecting. T articular ridge projecting well beyond articular margin; spur truncate, 1/6 width of
basal margin, set at 1/12 width of basal margin from basi-scutal angle; muscle crests not projecting below basal
margin. Mandible with lower margin of tooth 4 with small subsidiary cusps.
REMARKS. — This species can be distinguished by the non-flared aperture and by the fine, external growth
ridges of the thin parietal plates. In addtion the alar growth ridges are removed from and do not closely parallel the
inferior alar margin, and the upturned alar margins of CL' and CL2 are wide, 1/2 the width of the entire ala,
and flat. Differences between B. hirsutum and B. alearum and B. corolliforme are listed under B. alearum and
B. corolliforme, respectively
Distribution. — North-east Atlantic, from the Faeroe Islands south to the Azores; 944-1829 m.
Genus MESOLASMA Foster, 1981
Fig. 50; Tables 22-23
Mesolasma Foster. 1981: 362.
DIAGNOSIS. — Adult with fixed shell pattern of 6 thickened, solid, calcareous compartmcntal plates including
discrete rostral plate, paired CL1 and CL2, and C (R-CL'-CL2-C); externally with prominent horizontal growth
ridges lined with small setae and small beads; paries thickened and inflected inward basally; chitinous laminae
absent. External alar growth ridges diverging from inferior alar margin; superior alar margin with narrow welting.
Basis thin, calcareous. Articular ridge of S not prominent. Articular ridge of T not prominent. Mandible tridentoid
or quadridentoid. Segments of posterior cirri longer than wide, with graded series of 3 or more setae on anterior
edge. Caudal appendages absent.
Type Species. — Mesolasma fosteri (Newman & Ross, 1971).
Source : MNHN, Paris
CIRRIPED1A THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
237
Tables 22 and 23. — Mesolasma: characters of Mesolasma fosteri.
artic. = articular; b-s = basi-scutal; c.a. = caudal appendage; ext. = external; int. = internal; n.a. = not applicable; proj. = projecting.
Remarks. — One species only is presently assigned to this genus. The genus occupies an intermediate
position between Bathylasma and Hexelasma (Foster, 1981). The opercular plates of Mesolasma are more
balanoid-likc, lacking the prominent articular ridges found in Bathylasma , whereas the setation of the segments of
the posterior cirri is less specialized than that found in Hexelasma.
DISTRIBUTION. — Oligocene to Recent. Oligocene: Victoria, Australia. Recent: Western Pacific Ocean: New
Zealand, Norfolk and Kermadec Islands; 490-1556 m.
Fig. 50. — Mesolasma Foster, 1981 and Tetrachaelasma Newman & Ross, 1971. Distribution map: •. M. fosteri (490-
556 m); A. T. southwardi (I 190-2328 m).
Mesolasma fosteri (Newman & Ross, 1971)
Fig. 50; Tables 22-23
Hexelasma fosteri Newman & Ross, 1971: 155, figs 75-76; 1976: 46
Aaptolasma fosteri - Foster, 1978: 83 (part). Not pi. 10F. tig 50 (= Hexelasma gracilis Foster. 1981).
Mesolasma fosteri - Foster, 1981: 362. figs 8 C. 9.
238
D. JONES
Material EXAMINED. — New Zealand. Tui Exped.: sin 011-14, north of Auckland, 30°45'S, 173°51'E, 538-
676 m: holotype (USNM Type 125310).
Types. — Holotype: USNM Type 125310; New Zealand, Tui Exped., stn 011-14, north of Auckland.
30°45'S, 173°51'E, 538-676 m, with Megalasma striatum Hoek and Trilasmis kaempferi (Darwin), the former on
the spine of echinoid, substratum of others unknown; slides (3).
Holoppe depository: USNM.
DIAGNOSIS. — As for the genus.
Distribution. — New Zealand, Norfolk and Kermadec Islands; 490-1556 m.
Genus TETRACHAELASMA Newman & Ross, 1971
Fig. 50; Tables 24-25
Tetraehaehisma Newman & Ross, 1971: 152.
DIAGNOSIS. — Adult with fixed shell pattern of 4 thick, solid, calcareous compartmental plates including
compound rostral plate, paired CL, and C (R-CL-C); parietes with fine horizontal growth ridges lined with small
setae; chitin absent. External alar ridges diverging from inferior alar margin; superior alar margin with narrow,
coarse welting. Basis membranous. S with prominent articular ridge. T similar to that of Bathylasma but articular
margin sinusoidal rather than straight or concave; lateral depressor muscle crests few, restricted to small area below
carinal angle. Rami of cirri II and III antenniform; intermediate articles of cirrus VI bearing 3 or 4 pairs of major
setae. Mandible quadridentoid. Caudal appendages absent.
Type Species. — Tetrachaelasma southwardi Newman & Ross, 1971.
Remarks. — Only one species is presently assigned to this genus.
Tables 24-25. — Tetrachaelasma: characters of the species
artic. - articular; b-s - basi-scutal; c.a. = caudal appendage; ext. = external; int. = internal; long. = longitudinal; max. = maximum: n.a. = not
applicable; proj. = projecting; segs = segments.
Distribution. — Southern Pacific Ocean, Antarctic Basin and off South America; 1 190-2328 m.
Tetrachaelasma southwardi Newman & Ross, 1971
Fig. 50; Tables 24-25
Hexelasma antarcticum Borradaile, 1916: 132 (part). — Weisbord. 1965: 1015 (part); 1967: 51. pi. 2 figs 7-8 (part).
Tetrachaelasma southwardi Newman & Ross, 1971: 152, fig. 74. pis 26-31.
Material EXAMINED. — South Pacific Ocean. "Eltanin ": SOSC sin 6. 52°10'S, 142°10'W. 1260-1273 m
20.03.1965: holotype (USNM 125305).
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
239
Types. — Holotype : USNM 125305; Soulh Pacific Ocean, "Eltanin", SOSC stn 6, 52°10'S. 142°10’W, 1 260-
1273 m, 20.03.1965; dry (1 lot) and slide (1 lot).
Paratypes: USNM 125306, South Pacific Ocean. " Eltanin ", SOSC stn 6, 52°10'S, 142°I0'W, 1260-1273 m,
20.03.1965. — USNM 125307, off Falkland Is, "Eltanin", SOSC stn 376. 54°03'S, 56°03'W. 1720-1739 m,
plates only. — USNM 125308, off southern Chile, "Eltanin", SOSC sin 216, 52°53'S, 75°36'W, 1 190-1263 m,
plates only. — USNM 125309, off Cape Horn, "Eltanin", SOSC stn 255, 59°45’S, 68°50'W. 1207-1591 m,
plates only, in alcohol.
Holotype and paratype depository: USNM.
Diagnosis. — As for the genus.
Remarks. — The 4-platedness is due to the compound R and the absence of CL2 (Newman & Ross. 197 1 ).
The morphology of this species is puzzling, not only because of the massive 4-plated wall and absence of radii,
but also because of the flaring orifice and exceptionally broad carinal alae that internally contribute to about 1/2
of the sheath’s circumference. Apparently, it reflects an early radiation into the deep sea, and has a combination of
plesiomorphie and apomorphic characters that have likely evolved from the chionelasmatid level of organisation
and, therefore, its relationship remains to be resolved.
Distribution. — Southern Pacific Ocean, Antarctic Basin and off South America; 1 190-2328 m.
Genus TESSARELASMA Withers, 1936
Tables 26-27
Tessarelasma Withers, 1936: 591.
DIAGNOSIS. — Adult with fixed shell pattern of 4 solid, calcareous compartmental plates including compound
rostral plate, paired CL, and C (R-CL-C). External alar ridges diverging from inferior alar margin; superior alar
margin with narrow welting. Basis calcareous, marginally thickened, appearing as inflection of wall. S lacking
prominent articular ridge, with area above shallow adductor muscle pit roughened as in Hesperibalanus.
Type Species. — t Tessarelasma pilsbryi Withers, 1936.
Remarks. — The T and the soft parts of this species are unknown. Newman and Ross (1971: 155)
commented that the S is more balanoid than chthamaloid in appearance, whereas the structure of the basis is
chthamaloid or bathylasmatid, suggesting an intermediate position between the chthamaloids and the balanoids.
For these reasons they tentatively suggested bathylasmatid affinities for Tessarelasma. The genus is only known
from the fossil record and is monospecific.
Distribution. — Lower Miocene; Burma.
Tables 26 and 27. — Tessarelasma : characters of the species
articular, c.a. caudal appendage; ext. - external; int. - internal; long. = longitudinal; n.a. = not applicable; proj. = projecting
240
D. JONES
t Tessarelasma pilsbryi Withers, 1936
Tables 26-27
Tessarelasma pilsbryi Withers, 1936: 591. — Newman & ROSS. 1971: 155; 1976: 46.
Types. — Holotype: No registration number. Lower Miocene. Eastern Bengal in grey shale and greenish-grey
sandstone, containing pebbles of shale, and forming part of the Surma series, about 2000 feet below the base of
the Tipam Sandstone.
Holotype depository’: Geological Department, BMNH.
Diagnosis. — As for the genus.
Remarks. — As for the genus.
Distribution. — Lower Miocene; Burma.
Subfamily HEXELASMATINAE Newman & Ross, 1976
Tables 1-2
Diagnosis. — Primary shell wall not surrounded by whorl(s) of imbricating plates. Adult shell wall with
6 plates (R-CL>-CL2-C) divided into inner and outer laminae by longitudinal chitinous rods; externally with
prominent horizontal growth ridges lined with small setae. External alar growth ridges parallel to or diverging from
inferior alar margin; superior alar margin with narrow to moderate welting. Basis calcareous, thin centrally;
sometimes membranous. Opercular plates with articular ridges not prominent. Mandible quadridentoid. Caudal
appendages absent.
Remarks. — Only one genus, Hexelasma, is contained in this sub-family. Heretofore 9 species were included
in Hexelasma. Six new species are described in the present report bringing to 15 the number of hexelasmatines
presently known.
DISTRIBUTION. — Eocene to Recent. Eocene: Tonga. Lower Miocene: Victoria. Recent: Western Atlantic,
Indo-west Pacific, Australasia.
Genus HEXELASMA Hoek, 1913
Figs 51, 64; Tables 28-30
Hexelasma Hoek, 1913: 244 (in part.). — Pilsbry, 1916: 329. — KRUGER, 1940: 30. — Withers, 1953: 99. — UTINOMI,
1965: 13; 1968: 30. — Zevina, 1968: 64. — Newman & Ross, 1971: 155.
Aaptolasma Newman & Ross, 1971: 158. — FOSTER, 1981: 356.
DIAGNOSIS. — As for the subfamily, which is monogencric.
TYPE Species. — Hexelasma velutinum Hoek. 1913.
RECENT Species. — Hexelasma americanum Pilsbry, 1916; H. arafurae Hoek, 1913; H. aureolum sp. nov.;
H. brintoni (Newman & Ross, 1971); H. callistoderma Pilsbry, 191 1; H. flavidum sp. nov.; H. foratum sp. nov.;
H. globosum sp. nov.; H. gracilis Foster, 1981; H. leptoderma (Newman & Ross, 1971); H. nolearia (Foster,
1978); H. persicum sp. nov.; H. sandaracum sp. nov.; H. triderma (Newman & Ross, 1971); H. velutinum Hoek,
1913.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
241
Table 28. — Hexelasma: characters of the species
242
D. JONES
Table 29. — Hexelasma : characters of the species (continued)
Source : MNHNJ Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATQ1DEA
243
artic. = articular; b-s = basi-scutal; c.a. = caudal appendage: ext. = external; int. = internal; n.a. = not applicable; long. = longitudinal; proj. =
projecting.
Table. 30. — Hexelasma: characters of the species (continued)
244
D. JONES
artic. = articular; b.-s. = basi-scutal angle; c.a. = caudal appendage; ext. = external; int. = internal; long. = longitudinal; n.a. = not applicable;
proj. = projecting; segs = segments.
Remarks. — Difficulties regarding the definition and familial placement of the deep-sea genera Aaptolasma ,
Bathylasma and Hexelasma were discussed by FOSTER (1978, 1981). Re-examination of the type material of
H. velutinum led FOSTER (1981) to conclude that Hexelasma was a valid genus, but that Aaptolasma was a junior
synonym of Hexelasma. FOSTER (1981) also described the new genus Mesolasma to accommodate H. foster!
(Newman & Ross, 1971). BUCKERIDGE (1983) included Mesolasma in the Bathylasmatinae.
HOEK (1913) established the genus Hexelasma but did not designate a type species. Hexelasma velutinum was
designated as the type species of the genus by Withers (1913) and by subsequent selection by Utinomi (1963)
who, probably not realising that Withers (1913) had already designated a type species for Hexelasma, similarly
designated Hexelasma velutinum as the type. Foster (1981) apparently also overlooked Withers’ (1913)
designation. Buckeridge (1983) incorrectly listed Balanus callistoderma Pilsbry, 191 1 as the type species of
Hexelasma. Balanus callistodenna was the type species of Aaptolasma originally designated by Newman and ROSS
(1971).
Much confusion has stemmed from the fact that Hoek (1913) confounded more than 1 species in his
description of Hexelasma velutinum — viz. H. velutinum s. str., H. leptodenna (Newman & Ross, 1971), and
H. arafurae (Hoek, 1913). Of the 4 specimens originally included by HOEK (1913) in H. velutinum, I specimen
had operculae which differed from those of the 2 specimens of H. velutinum (" Siboga ", stn 105) and the single
specimen from "Siboga", stn 251. This ’odd’ specimen and its operculae were illustrated by HOEK (1913: 247,
pi. 26 figs 3-5) and was later confirmed by Foster (1981) to be H. leptodenna Newman & Ross, 1971. FOSTER
(1981) also determined that the parietal plates of H. leptodenna did not contain chitinous laminae, but that strips of
orange chitin were present on the inner surface. The specimen from "Siboga", stn 251 was attributed to H. arafurae
by Foster (1981). The types of H. velutinum show no chitinous lamellae in the parietal shell, but do have a
lamina applied basally to the inner faces of the shell plates. These specimens are considered immature since, in the
further development of these laminae in larger specimens, calcite layers are found on either side of the laminae
(Foster, 1981).
Distribution. — Eocene to Recent. Eocene: Tonga; Lower Miocene : Victoria; Recent : Western Atlantic.
Indo-west Pacific, 70-1300 m.
Olf Florida, 734-770 m (H. americanum); Arafura Sea, 560 m (H. arafurae)-, Kei Is, 290 m (H. leptodenna)-,
Vanuatu, New Caledonia, Loyalty Is, Norfolk Ridge, Wallis Is, Combe Bank, Field Bank, Tuscarora Bank, 1 70-
1300 m ( Hexelasma sandaracum)- Chesterfield Is, Lord Howe Ridge, New Caledonia, Loyalty Ridge, Norfolk
Ridge, 257-730 m (H. aureolum)-. New Caledonia, Loyalty Is, Norfolk Ridge, 255-608 m (H. flavidum)-
New Caledonia, Loyalty Is, Norfolk Ridge, 275-585 m {H. persicum); New Caledonia, Loyalty Is, 370-575 m
(//. globosum )\ Vanuatu, Loyalty Is, 270-500 m (H. foratum)-, New Zealand, 741 m (H. nolearia)' New Zealand,
1024-1029 m (H. gracilis)-. Japan, 1 15-141 m (H. callistoderma); Japan, 549 m (H. triderma)-, Japan, Philippines
to South China Sea, 204-390 m (H. velutinum)-, Vietnam, 1 10-198 m (H. brintoni).
Source : MNHN, Paris
CIRRIPED1A THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
245
Table 31. — Hexelasma: maximum size, geographical and baihymetrical distribution of the species
Hexelasma americanum Pilsbry, 1916
Fig. 51; Tables 28-31
Hexelasma americanum Pilsbry, 1916: 330, fig. 98, pi. 69 figs 1-3 a. — Utinomi, 1965: 12. — Newman & ROSS, 1976:
46. — Foster, 1981: 356. — Young, 1998: 66, figs 1, 20-21.
Aaptolasma americanum - Newman & Ross, 1971: 161, pis 32 A-B, 36. 37. — Foster, 1978: 79.
Material EXAMINED. — United States. "Albatross"-, stn 2663, off E coast of Florida, 29°39‘N, 79°49’W.
770 m, 04.04.1886: holotype (USNM Type 14559).
Types. — Holotype: USNM 14559; United States, off S Carolina, "Albatross" stn 2663, 29°39'N, 79°49'W.
770 m, 04.04.1886, bottom temperature 42.7°F, seated on a branch of coral; complete specimen, dry.
246
D. JONES
Paratype : USNM 48093, from "Albatross", sins 2662-3-9, 2671-2; remains of 3 individuals.
Holotype and paratype depository. USNM.
Diagnosis. — Paries with longitudinal chitin-filled tubes, sub-arcuate in cross-section, regularly spaced,
separated from one another by pronounced calcareous bridges. Basal width of CL2 1/3 basal width of CL1. Inferior
alar margin not curved, with slight hook at free end; narrow welling along superior alar margin. Basis
membranous, with narrowly inflected calcareous borders. S internally smooth; few crests for lateral depressor
muscles present; externally smooth, regular growth lines not cut by longitudinal striae. T spur obliquely truncate,
1/3 width of basal margin, slightly removed from basi-scutal angle. Mandibular teeth acute, margins smooth.
Remarks. — Hexelasma americanum is a large sized species (maximum: RC length 17.0, C height 21.0),
with pale orange opercular plates.
Distribution. — Off Florida, 734-770 m.
FIG. 51. — Hexelasma Hoek, 1913. Distribulion map: A . H. brintoni (110-198 m); O.H. americanum (734-770 m);
A . H. callistoderma (115-141 in); •. H. arafurae (560 m); □. H. aureolum (257-730 m); ■. H. flavidum (255-608 m);
O, H. foratum (270-500 m).
Hexelasma arafurae Hoek, 1913
Fig. 51; Tables 28-31
Hexelasma arafurae Hoek, 1913: 251, pi. 25 figs 12-16. — Utinomi, 1965: 11. — Newman & Ross, 1971: 155: 1976:
46. — Foster, 1981: figs 6 F-H.
Aaptolasma arqfura - Foster, 1978: 79.
Material EXAMINED. — Indonesia. "Siboga": stn 262. 5°53.8'S, 132°48.8'E, 560 m. 18.12.1899: holotype
(ZMUA Cirr. 100.318).
Types. — Holotype-. ZMUA Cirr. 100.318; Indonesia, "Siboga", stn 262, 5°53.8'S, 132°48.8'E. 560 m,
18.12.1899, in alcohol.
Holotype depository. ZMUA.
Diagnosis. — Continuous chitinous lamina bisecting thickness of parietal plates. Basal width of CL2 more
than 1/2 basal width of CL1. Alae externally with strongly marked growth ridges; welting along superior alar
margin inconspicuous. Basis calcareous, thin. S internal surface rough; crests for lateral depressor muscles present;
externally smooth, even growth lines cut by a broad, longitudinal stria. T spur truncate, more than 1/3 width of
basal margin, set almost at basi-scutal angle; inner surface smooth. Mandible with upper margins of teeth 3 and 4
with small subsidiary cusps.
Source : MNHN. Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
247
Remarks. — Hexelasma arafurae is a “large" species, with orange to brick-red orange opercular plates. The tall
conical shell and the toothed orifice of H. arafurae are common to H. aureolum sp. nov., H. callistoderma,
H. persicum sp. nov. and H. velutinum. Differences between H. arafurae, H. aureolum sp. nov. and H. persicum
sp. nov. may be found under H. aureolum sp. nov. and H. persicum sp. nov., respectively.
Hexelasma callistoderma differs from H. arafurae as the former has parietal plates with longitudinal chitin-filled
tubes, sub-arcuate in cross-section, separated from one another by regularly spaced, broad calcareous bridges; and
the basal width of CL2 is 1/4-1/5 the basal width of CL1. In juvenile H. callistoderma the basis is membranous,
adults having a thick calcareous ledge. The opercular plates of H. callistoderma are white. The S internal surface is
smooth and externally the smooth, even growth lines are not cut by longitudinal striae. The T spur is roundly
truncated, 1/4 the width of the basal margin, and set at 1/2 its own width from the basi-scutal angle; the inner
surface is distally roughened in the carinal area above the muscle crests with linear beading. The mandible with no
small subsidiary cusps.
H. velutinum is a large species (maximum: RC length 10.0, C height 13.0), with orange chitin applied to the
inner surfaces of the parietal plates, which differs from H. arafurae as follows: the basal width of CL2 is 2/3 the
basal width of CL1; the welting along the superior alar margin is narrow and finely crcnated; the opercular plates
are white; the S is convex externally, with smooth, widely-spaced, growth lines which arc not cut by longitudinal
striae; internally the adductor ridge is weak, crests for the lateral depressor muscles are absent; and the occludent and
articular margins are rounded and folded; the T spur is set slightly apart from the basi-scutal angle; and the inner
surface is roughened.
Distribution. — Arafura Sea, 560 m.
Hexelasma aureolum sp. nov.
Figs 51-54; Tables 28-31
MATERIAL EXAMINED. — Chesterfied Islands. Corail 2: stn DE 16. 500 m: 2 specimens (WAM C 23253). —
Stn CP 17, 500 m: I specimen.
Lord Howe Ridge. Musorstom 5: stn DW 338, 540-580 m: 1 specimen (MNHN-Ci 2702). Drawn. — Stn CP 389.
500 m: 2 specimens (WAM C 23254, WAM C 23255).
New Caledonia. Musorstom 4: sin DW 156, 525 m: 2 specimens. — Stn CP 238, 500-510 m: 1 specimen.
Smib 8: stn DW 150, 519-530 m: 1 specimen.
Loyalty Ridge. Musorstom 6: stn DW 422, 257 m: 1 specimen, attached to sponge. — Stn DW 460, 420 m:
1 specimen (incomplete) (WAM C 23256). — Stn DW 476, 300 m: 2 specimens, attached to stones (WAM C 23257). —
Stn DW 478. 400 m: several specimens.
Norfolk Ridge. Chalcal 2: stn DW 72, 527 m: 1 specimen, attached to sponge (MNHN-Ci 241 1). — Stn DW 72.
527 m: 2 specimens, 1 attached to coral rubble, 1 to sponge (BMNH). — Stn DW 76, 470 m: 2 specimens.
Smib 3: stn DW 5, 502-512 m: 1 specimen (MNHN-Ci 2412).
Smib 4: stn DW 61, 550 m: 1 specimen, attached to coral.
Bathus 3: stn DW 809, 650-730 m: I specimen. — Stn DW 819. 478-486 m: I specimen, attached to sponge
(USNM).
Types. — Holotype: MNHN-Ci 2411 (Chalcal 2, stn DW 72).
Paratypes : BMNH (Chalcal 2, stn DW 72). — MNHN-Ci 2412 (Smib 3, stn DW 5). — MNHN-Ci 2702
(Musorstom 5, stn DW 338). Drawn. — USNM (Bathus 3, stn CP 819). — WAM C 23253 (Corail 2,
stn DE 16).
Diagnosis. — Thin parietal plates with chitin either as lamina on inner walls of parietes, or as rods within
paries separating paries into inner (1/4 thickness) and outer (3/4 thickness) lamina. Basal width of CL2 1/4- 1/3
basal width of CL1; bases of plates thickened, forming calcareous ring. Inferior alar margin curved with slightly
basally directed hook at free end; externally with fine, narrow growth ridges; narrow welting (sometimes
inconspicuous) along superior alar margin. Basis calcareous, thin. Opercular plates whitish, apical areas of S and T
pinkish orange-brown. S internal surface distally roughened with beading; crests for lateral depressor muscles
248
D. JONES
present; externally transverse, sinuous growth lines cut by 2 narrow, longitudinal striae. T spur truncate, 1/4 width
of basal margin, set at its own width from basi-scutal angle; inner surface distally roughened with 6-9 low ridges
and associated beading. Mandible with 1 small subsidiary tooth sometimes between teeth 3 and 4, lower margin of
tooth 4 with 1 small subsidiary cusp.
Fig. 52. — Hexelasma aureolum sp. nov. Paratype from Musorstom 5, stn DW 338 (MNHN-Ci 2702): a, S, external view;
b, S, internal view; c, T, external view; d. T, internal view.
Description. — Size large (maximum: RC length 19.0, C height 18.0). Shell conical. mature specimens tall.
Parietal plates, thin (especially in juveniles), all plates sloping inward toward orifice except outwardly curving C;
thick, golden yellow, persistent epicuticle covering paries; growth ridges definite, regular, narrow, marked by small
golden setae; bases of paries thickened, forming calcareous rim; internally walls ribbed basally. Paries with yellow
chitin, either as lamina on inner walls, or as rods within paries separating paries into inner (1/4 thickness) and
outer (1/4 thickness) laminae. Rostral plate flattened, basal width greater than basal width of CL'. CL1 wider than
CL2; CL2 narrow, elongated triangular shape, basal width 1/4 to 1/3 basal width of CLL C highest plate, not as
wide as rostral plate; apex recurved outward. Alae with summits oblique; fine, narrow growth ridges directed
upward parallel to inferior alar margin, then turning sharply downward toward alar angle, then upward again at
sharp angle to form narrow welting, latter inconspicuous in some specimens; inferior alar margin curved, slightly
basally directed hook at free end; alae visible externally for 3/4 (juvenile) to 2/3 (adult) length of parietes. Basis
thin, calcareous. Orifice toothed, diameter smaller than that of basis; basis as wide as or wider than total height of
shell. Opercular plates below level of orifice. S triangular, basal margin 9/14 length of occludent margin; apex
acute; externally slight concavity in middle area; occludent margin longest margin, basal margin sinuous, shorter
than articular margin, distinctly curved upward toward articular margin; sinuous growth ridges well defined, cut by
2 narrow, longitudinal ridges varying from slight to well marked, ridges not well separated, extending from apex
to basal margin, ridge nearest occludent margin most well-marked; inner S surface with large, deep adductor muscle
pit; adductor ridge weakly developed; lateral depressor muscle pit deep, with irregular muscle attachment crests;
roughened, irregular ridges lor attachment of rostral depressor muscle in deep pit; articular furrow deep; articular
ridge extending almost 2/3 length of margin, slightly projecting; articular margin rounded, folded; inner surface of
valve, including articular ridge, roughened with irregular beading distally. T smaller and narrower than S; externally
sinuous growth ridges prominent; articular margin slightly concave; carinal margin convex; basal margin slightly
hollowed, broad spur roundly truncated, 1/4 width of basal margin, set at its own width from basi-scutal angle;
spur furrow open, basally shallow, broad; inner surface with articular ridge prominent in upper 1/3, not projecting;
4 well developed muscle attachment crests for depressor muscles; inner surface of valve roughened with
Source : MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
249
6-9 irregular, low ridges and associated beading in distal half. Parietal plates pale ochre/dirty lemon color, basally
fading to whitish; alae whitish with lower areas peachy orange in young specimens. Opercular plates whitish; in
some specimens apical areas of S and T pinkish orange-brown internally and externally. Measurements of holotype
as follows:
Labrum with shallow medial depression with row
of small teeth and dense, short setae. Mandibular palp
oblong-ovoid; long, finely serrate setae terminally.
Mandible with 4 teeth, tooth I largest, well separated
from teeth 2-4; with or without 1 small subsidiary
cusp in angle between tooth 3 and 4; lower margin of
tooth 4 with 1 small subsidiary cusp; inferior angle
moderately large, molariform, with 2 protrusions,
upper tooth-like, lower peg-like. Maxillulc setose; 3
long, stout setae at upper angle; notch below upper
angle wide, with 3-5 pairs of smaller setae; cutting
margin below stepped, almost vertical, with 8-10
pairs of longer, stout setae; inferior angle not
protuberant or stepped, barely demarked. with 5-7
pairs of smaller setae. Maxilla wide, coxal endite
barely defined; setae long, finely serrulate.
CiiTus I with rami unequal; anterior ramus longer than posterior; proximal segments of anterior ramus not
protuberant anteriorly, segments wider than posterior ramus; both rami with segments thickly setose, setae finely
serrate. Cirrus II longer than cirrus I; rami unequal, anterior ramus shorter than posterior ramus; all segment,
clothed with setae. Cirrus III similar to cirri IV-VI; rami subequal; segments becoming oblong distally; posterior
ramus with 2-3 pairs of long, finely serrulate setae on anterior faces of distal segments, posterior segments densely
setose; anterior ramus with 2 pairs of long, finely serrulate setae on anterior faces of distal segments, proximal
segments with 3 pairs. Cirrus IV to VI similar, longer than cirrus III; rami subequal; segments oblong, with
3 pairs of long setae on anterior faces, distal 2 pairs of longest. Cirral formula as follows:
Fig. 53. — Hexelasma aureolum sp. nov. Holotype from
Chalcal 2, stn DW 72 (MNHN-Ci 2411): lateral view.
Scale = 4 cm.
Penis 1/2 length of cirrus VI; annulated; with few sparse setae; irregular circlet of longer setae distally.
Remarks. — The tall conical shell and the toothed orifice are common to H. aureolum sp. nov., H. arafurae,
H. callistoderma and H. velutinum. Hexelasma aureolum can be separated from H. arafurae as follows: in
H. aureolum the basal width ol CL2 is 1/3 to 1/4 the basal width of CL1, rather than being more than 1/2, and the
alae are narrower and exposed for 2/3-3/4 the height of the wall, rather than wide and exposed for the total height of
the wall, as in H. arafurae. In H. aureolum the chitin is either as a lamina applied on the inner surface of the
paries, or as rods which separate the paries into inner (1/4 thickness) and outer (3/4 thickness) laminae; in
H. a i af lira there is a continuous lamina of chitin bisecting the thickness of the parietal plates. The sinuous S
growth lines of H. aureolum are cut by 2 narrow striae, those of H. arafurae are smooth and cut by 1 broad stria;
the S basal margin of H. aureolum is distinctly curved upward toward the articular margin rather than being
S l§hlly rounded; and the S articular ridge is slightly, rather than not, projecting. In H. aureolum the T spur is 1/4
tie width of the basal margin and set at its own width from the basi-scutal angle, rather than more than 1/3 of the
250
D. JONES
width of the basal margin and set almost at the basi-scutal angle; the inner surfaces of S and T are roughened with
irregular beading distally, rather than being smooth, and are pale orange rather than brick-red. The cirral counts for
H. aureolum arc significantly higher than those for H. arafurae (Cl 14. 13; CII 17, 22; CIII 32. 37; CIV 43, 49;
CV 49, 49; CVI 49, 51 and Cl 14, 10; CII 12, 15; CIII 19. 43; CIV 30, 37; CV 38, 40; CVI 39, 41,
respectively).
Fig. 54. — Hexelasma aureolum sp. nov. Paratype from Musorstom 5, stn DW 338 (MNHN-Ci 2702): a. right mandible;
b. left mandible; c, right maxillule, d, left maxillule; e, mandibular palp; f, labrum; g. maxilla; h, cirrus I;
i, cirrus VI and penis; j, cirrus III; k. cirrus II.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
251
Hexelasma aureolum is superficially similar 10 H. callistoderma but differs as follows: the basal width of CL2
is 1/3 to 1/4 the basal width of CL1, rather than 1/4 to 1/5 the width; the chitin is either as a lamina applied on
the inner surface of the paries, or as rods which separate the paries into inner (1/4 thickness) and outer (3/4
thickness) laminae, rather than as long rods, sub-arcuate in cross section, separated by regularly spaced, broad
calcareous bridges. The basis of H. aureolum is calcareous and thin; that of H. callistoderma is membranous in
young specimens, with adults developing a thick calcareous ledge. In H. aureolum the T spur is set at a distance of
its own width from the basi-scutal angle rather than 1/2 its width; and the inner surfaces of S and T are orange-
brown in H. aureolum, white in //. callistoderma.
Hexelasma aureolum is superficially similar to H. velutinum but there are the following differences: in
H. aureolum the basal width of CL2 is 1/4 to 1/3 the basal width of CL1, rather than 2/3 the width; the S adductor
muscle pit is distinct rather than faint; the S articular ridge is slightly projecting, rather than not; the S depressor
muscle pit is distinct rather than being a shallow excavation, the T basal margin is slightly hollowed out rather
than straight; the T spur of H. aureolum is 1/4 the width of the basal margin and set at its own width from the
basi-scutal angle, rather than 1/3 the width of the basal margin and set at 1/2 its own width from the basi-scutal
angle; and the inner surface is roughened with 6-9 irregular, low ridges and associated beading in the distal half,
rather than smooth. Cirral counts for H. aureolum are also higher than those for H. velutinum (Cl 14. 13; CII 17,
22; CIII 32, 37; CIV 43, 49; CV 49, 49; CVI 49, 51 and Cl 9, 13; CII 11, 14; CIII 22. 28; CIV 37, 39; CV 40.
43; CVI 42, 43, respectively).
Etymology. — From the Latin aureolum, ‘golden’, referring to the golden color of the persistent epicuticle
present on the compartmental plates.
Distribution. — Chesterfield Islands, 500-580 m; New Caledonia, 500-530 m; Loyalty Ridge, 257-420 m;
Norfolk Ridge, 470-730 m.
Hexelasma brintoni (Newman & Ross, 1971)
Fig. 51; Tables 28-31
Aaptolasma brintoni Newman & Ross, 1971: 162, fig. 78, pis 33 E-F, 38, 39 A-E; 1976: 46. — FOSTER. 1978: 79.
Hexelasma brintoni - FOSTER, 1981: 356.
Material EXAMINED. — South China Sea. Naga Exped. 1959-61: stn 97, 15°40.0'N, 109°22.9'E. 110-198 m.
27.02.1960: holotype (USNM Type No. 125311).
Types . — Holotype: USNM 125311; South China Sea, Naga Exped. 1959-61, stn 97, I5°40.0'N.
109°22.9'E, 1 10-198 m, 27.02.1960; dry (1 lot) and slides (3).
Holotype depository ■: USNM.
Diagnosis. — Parietal plates with longitudinal chitin-filled tubes sub-arcuate in cross-section, regularly
spaced, separated from one another by broad calcareous bridges; tubes bisecting thickness of parietal plates. Basal
width of CL2 3/5 basal width of CL>. Inferior alar margin curved, without basally directed hook at free end;
externally with weakly marked growth ridges; ridged, narrow welling along superior alar margin. Basis calcareous,
thick. Opercular plates white. S internal surface smooth; several crests for lateral depressor muscles present;
externally smooth, even growth lines not cut by longitudinal striae. T spur truncate, 1/3 width of basal margin,
terming abrupt angle of 130° with basal margin, set at 1/2 its width from basi-scutal angle; inner surface smooth.
Mandibular teeth 2 and 3 with side teeth, upper and lower margins of tooth 4 with small subsidiary cusps.
Remarks. — Hexelasma brintoni is superficially similar to H. flavidum sp. nov., H.foratum sp. nov. and
H. sandaracum sp. nov. Differences between H. brintoni and these species may be found under H. flavidum
sp. nov., H.foratum sp. nov. and H. sandaracum sp. nov., respectively.
Distribution. — Off Da Nang, Vietnam, 1 10-198 m.
252
D. JONES
Hexelasma callistoderma Pilsbry, 1911
Fig. 51; Tables 28-31
Balanus callistoderma Pilsbry, 1911: 78, fig. 10, pi. 12 fig. 5, pi. 15 figs 3-7.
Balanus corolliformis - KROGER, 1911: 55, figs 112-114, pi. 1 fig. 1, pi. 4 fig. 38.
Hexelasma callistoderma - HoEK, 1913: 245. — Pilsbry, 1916: 332, fig. 99. — Utinomi, 1958: 307; 1965: 12 —
Foster, 1981: 356.
Aaptolasma callistoderma - Newman & ROSS. 1971: 159, fig. 77. pis 32 C-D, 35, 43 C; 1976: 46. — FOSTER. 1978: 79.
Material EXAMINED. — Japan. " Albatross stn 5068, off Ose Saki, 35°02'25"N, I38°46'55"E, 141 in,
15.10.1906: holotype (USNM Type 38690).
TYPES. — Holotype: USNM 38690; ‘■Albatross" stn 5068, off Ose-Saki, Japan, 35°02'25"N, 138°46'55"E,
141 m, 15.10.1906.
Paratype: USNM 38690; " Albatross " stn 3741, Japan, about the same place than above, 115-124 m,
17.05.1900; in alcohol (1 lot), and slide (1).
Holotype and paratype depository n USNM.
Diagnosis. — Parietal plates with longitudinal chitin-filled tubes, sub-arcuate in cross-section, separated from
one another by regularly spaced, broad calcareous bridges. Basal width of CL2 1/4- 1/5 basal width of CL'. External
alar growth ridges line; thin, rough welting along superior alar margin. Basis membranous (juveniles), adults with
thick calcareous ledge. Opercular plates white. S internal surface smooth; crests for lateral depressor muscles
present; externally smooth, even growth lines not cut by longitudinal striae. T spur roundly truncated, 1/4 width of
basal margin, set at 1/2 its own width from basi-scutal angle; inner surface distally roughened in carinal area above
muscle crests with linear beading. Mandibular teeth lacking small subsidiary cusps.
REMARKS. — Hexelasma arafurae and H. aureolum sp. nov are superficially similar to H. callistoderma.
Dilferences between these species may be found under H. arafurae and H aureolum sp nov., respectively.
Distribution. — Japan, 115-141 m.
Hexelasma flavidum sp. nov.
Figs 51, 55-57; Tables 28-31
Material EXAMINED. — New Caledonia. Bathus 4: stn CP 910, 560-608 m: 1 specimen (MNHN-Ci 2413).
Loyalty Islands. Musorstom 6: stn DW 472. 300 m: several specimens, attached to sponges (MNHN-Ci 2703).
Drawn. — Stn DW 472, 300 m: 2 specimens. 1 attached to sponge, 1 to coral rubble (WAM 257-96).
Norfolk Ridge. Smib 5: stn DW 87, 370 m: 6 specimens, attached to coral rubble (MNHN-Ci 2414). — Stn DW 93
255 m: 1 specimen, attached to live coral ( Corallium sp.) (MNHN-Ci 2704).
Types. — Holotype : MNHN-Ci 2413 (Bathus 4, stn CP 910).
Paratypes : MNHN-Ci 2414 (SMIB 5, stn DW 87). — MNHN-Ci 2703 (MUSORSTOM 6, stn DW 472). Drawn.
— MNHN-Ci 2704 (Smib 5, stn DW 93). — WAM 257-96 (Musorstom 6, stn DW 472).
Diagnosis. — Thin parietal plates with chitin present as lamina on inner walls of parietes between
longitudinal ribs, or within paries as longitudinal tubes of chitin, separating paries into inner (1/4 thickness) and
outer (3/4 thickness) lamina. Basal width of CL2 1/3 or less than basal width of CLf Inferior alar margin curved,
slight hook at Iree end; externally growth ridges well spaced; serrated, narrow, ridged welting along superior alar
margin. Basis with calcareous edges, membranous centrally; thin. Opercular plates orange, apices cream. S internal
surface smooth; crests for lateral depressor muscles slight; externally well-spaced, sinuous growth lines cut by
slight longitudinal striae. T spur truncate, more than 1/3 width of basal margin, forming abrupt angle of 130° with
basal margin, set at 1/4 its own width from basi-scutal angle; inner surface distally roughened. Mandible with
upper and lower margins of tooth 4 with irregular, small subsidiary cusps.
Source : MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
253
Fig. 55. — Hexelasma flavidum sp. nov. Paratype from Musorstom 6, stn DW 472 (MNHN-Ci 2703): a. S. external;
view; b, S, internal view; c. T, external view; d, T, internal view.
Description. — Size small (maximum: RC length 10.0, C height 6.0). Shell rounded, squat and flattened.
Parietal plates thin, fragile; growth ridges faint, regular, finely beaded, lined with small, fine, golden setae, giving
hirsute appearance; persistent epidermis absent; internally with longitudinal, low, spaced ribs extending 1/2 length
ol plates. Yellowish chitin applied either as lamina on inner surface of parietes, between longitudinal ribs, or as
longitudinal tubes of chitin within plates, dividing thicker outer (3/4 width) and thinner inner (1/4 width) laminae.
Rostral plate largest plate, convex; broad at base, height equal to height of C. Basal width of CL2 1/3 or less than
1/3 basal width of CL1. C larger than CL1; distinctly shaped, central longitudinal ridge dividing plate into halves,
ridge more distinct in juveniles, slight in adults. Superior alar margins sub-parallel to basis; growth ridges well
spaced, directed upward parallel to inferior alar margin, then curving outward toward alar angle and upward again at
sharp angle to form ridged welting, latter with serrated superior margin formed from wide spacing of growth ridges;
inferior alar margin curved, slight hook at free end. Basis membranous centrally, calcareous at edges, thin. Orifice
smaller in diameter than basis; opercular plates subparallcl to basis. S thin, triangular, S basal margin 9/14 length
of occludent margin, shorter than articular margin; occludent margin longest margin; externally sinuous growth
ridges prominent, well spaced, cut by slightly indicated external longitudinal striae; internal surface smooth;
adductor muscle and lateral depressor muscle pits shallow; adductor ridge weak; articular furrow shallow; articular
ridge extending for 3/4 length of margin, not projecting; articular margin rounded, folded; S when closed filling
aperture. F smaller than S; valve thin; externally growth ridges sinuous, less defined than S growth ridges;
articular and carinal margins almost straight; basal margin hollowed out, meeting spur at sharp angle; spur basally
truncated, more than 1/3 width of basal margin, forming abrupt angle of 130° with basal margin, set at 1/4 its own
width from basi-scutal angle; spur furrow open, shallow, flat, broadest basally; internally articular ridge not
projecting, traceable through 1/3 length of valve; 9 distinct, prominent muscle attachment crests for depressor
muscles, slightly extending below basal margin; inner surface distally roughened. Parietal plates yellowish-orange
with ochre yellow longitudinal tubes showing through thin parietal wall: alae yellowish-orange, welting whitish;
opercular plates externally golden-orange with cream apices, orange/cream internally. Measurements of holotype as
follows:
Labrum with shallow medial depression with row of 30-35 hi- or tri-cusped teeth. Mandibular palp ovate; long,
finely serrate setae terminally, plumose setae medially. Mandible with 4 teeth, tooth 1 largest, well separated from
254
D. JONES
2-4; margins of 1-3 smooth, upper and lower margins of 4 with irregular subsidiary cusps; inferior small,
molariform, slightly dentate. Maxillule setose; 2 long, stout setae at upper angle; notch below upper angle wide,
with 4 pairs of smaller setae; cutting margin below stepped, with 5 pairs of longer, stout setae; inferior angle not
protuberant or stepped, with 4 pairs of small setae. Maxilla wide, coxal endite well developed; setae long, finely
serrulate.
Cirrus I with rami subequal; anterior ramus slightly longer than posterior; proximal segments of anterior ramus
not protuberant anteriorly, segments wider than those of posterior ramus; both rami with segments moderately
setose. Cirrus II slightly longer than cirrus I; rami subequal; segments moderately setose, setae finely serrulate.
Cirrus III longer than cirri I and II, and intermediate in form between cirrus II and cirrus IV; rami subequal;
segments becoming oblong distally; anterior and posterior rami with 1-3 pairs of long, finely serrulate setae on
anterior faces of distal segments. Cirrus IV to VI similar, longer than cirrus III; rami subequal; segments oblong,
with 2 pairs of long, finely serrulate setae on anterior faces. Cirral formula as follows:
Penis 1/2 length of cirrus VI; annulated; with few sparse setae; circlet of longer setae distally.
Remarks. — This species, with its wide, high
alae and its low profile, is more similar to H.
brintoni, H.foratum sp. nov. and H. triderma than to
H. callistoderma, H. americanum or H. leptoderma.
Hexelasma flavidum sp. nov. can be separated
from H. brintoni as follows: the shell is rounded,
squat and flattened, rather than conical; the basal
width of CL- is 1/3 or less than 1/3 the basal width
of CL1, rather than 3/5 the basal width; at the free
end of the curved inferior alar margin a slight, basally
directed hook is present, rather than absent;
longitudinal tubes of chitin divide the paries into a
thicker outer (3/4) and thinner inner (1/4) lamina,
rather than into laminae of equal width; the basis is
thin, calcareous at the periphery and membranous
centrally, rather than thick and calcareous; the T spur
is more than 1/3 the width of the tergal basal margin
and set at 1/4 its own width from the basi-scutal
angle, rather than 1/3 the width of the basal margin and set at 1/2 its width from the basi-scutal anale; the inner
surface of T is roughened rather than smooth; and the color of S and T are orange-cream and not white. In addition
the cirral counts arc lower for H. flavidum than those for H. brintoni.
Hexelasma flavidum is distinguished from H. triderma by the following characters: the parietal plates are thin
and relatively fragile, are without a persistent epidermis, and have faint, regular growth ridges lined with golden
setae, in contrast to the robust, thicker plates of H. triderma, which are covered with a brownish cuticle marked by
heavy, transverse growth ridges supporting numerous chitinous setae; the basal width of CL2 is 1/3 or less than
1/3 the basal width of CL1 in H. flavidum. in H. triderma 1/2-2/3. In H. flavidum longitudinal, chilin-filled tubes
divide the paries into an outer (3/4 width) and thinner inner lamina (1/4 width), rather than an almost continuous
sheet ol chain dividing the paries into an outer (2/3 width) and an inner lamina (1/3 width), as in H. triderma.
he curved inferior alar margin of H. flavidum has a slight hook at its free end, rather than an abrupt, basallv
directed hook; and the basis of H. flavidum is thin, calcareous at the periphery and membranous centrally rather
t lan thin, calcareous and marginally thickened ( H . triderma). Opercular and soft parts are unknown for H. triderma.
Fig. 56. — Hexelasma flavidum sp. nov. Holotype from
Bathus 4. sin CP 910 (MNHN-Ci 2413): lateral view.
Scale = 5 mm.
Source : MNHN Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
255
Differences between H.flavidum and H. foreman sp. nov. are listed under H.foratum sp. nov.
Etymology. — From the Latin flavidum , 'yellow', in reference to the distinctive yellow scuta of this species.
Distribution. — New Caledonia, 560-608 m; Loyalty Islands, 300 m; Norfolk Ridge, 255-370 m.
u ' mm _ 0.1 mm K 0.1 mm ... 0.1 mm 0.1 mm
- - <= - a. b - d-g, j, k - h - i
Fig 57. — Hexelasma flavidum sp. nov. Paratype from Musorstom 6, stn DW 472 (MNHN-Ci 2703): a, right mandible;
b, left mandible; c, right maxillule; d. labrum; e. mandibular palp; f, maxilla; g. cirrus III; h. penis; i. cirrus VI;
j. cirrus II; k, cirrus I.
256
D. JONES
Hexelasma foratum sp. nov.
Figs 51, 58-60; Tables 28-31
MATERIAL examined. — Loyalty Islands. Musorstom 6: stn DW 400, 270 m; 3 specimens (2 incomplete)
(MNHN-Ci 2706. Ci 2707). — Stn DW 451. 330 m: 2 specimens (1 ovigerous, both incomplete) (MNHN-Ci 2415, Ct
2416). Drawn.
New Hebrides Arc. Volsmar: stn DW 16, 500 m; 1 specimen (WAM C 23258).
Types. — Holotype-. MNHN-Ci 2707 (Musorstom 6, stn DW 400).
Paratypes : MNHN-Ci 2415 (MUSORSTOM 6, stn DW 451). — MNHN-Ci 2416 (MUSORSTOM 6. stn DW 451).
Drawn. — MNHN-Ci 2706 (MUSORSTOM 6, stn DW 400). — WAM C 23258 (VOLSMAR, sin DW 16).
Diagnosis. — Thick parietal plates with chitin present as longitudinal, spaced rods, separating partes into
equal sized laminae. Basal width of CL2 1/4- 1/3 basal width of CL'. Inferior alar margin curved, without distinct
hook at free end; narrow, ridged welting along superior alar margin, increasing in width toward free end. Basts
calcareous, thick; minute, pinprick holes perforating junction of basis and inner surfaces of parietal plates.
Opercular plates whitish. S internal surface roughened distally; crests for lateral depressor muscles absent;
externally widely-spaced, sinuous growth lines not cut by longitudinal striae. T spur truncate, 3/10 width of basal
margin, set at less than 1/2 its own width from basi-scutal angle; inner surface roughened over 2/3 of surface.
Mandible with teeth 2 and 3 each with 1 small subsidiary cusp, upper and lower margins of tooth 4 with irregular,
small subsidiary cusps.
d
Fig. 58. — Hexelasma foratum sp. nov. Paratype (ovigerous) from MUSORSTOM 6, stn DW 451 (MNHN-Ci 2416):
a, S, external view; b. S, internal view; c, T, internal view; d, T, external view.
DESCRIPTION. — Size small (maximum: RC length 10.9, C height 4.4). Shell truncated cone shaped. Parietal
plates thick, rugged; externally with remains of yellow epidermis; fine growth ridges marked by long, golden,
chitinous setae; basal margins of plates sometimes deeply fluted; longitudinal, spaced tubes of chitin in middle ol
parietal plates; inner surfaces of paries with faint indications of longitudinal ribs; minute, regulatly spaced,
pinprick holes perforating junction of basis and inner surfaces of parietal walls, holes corresponding to edges of
longitudinal ribs. Rostral plate widest plate, flattened. Basal width ol CL1 less than basal width of rostral plate.
Basal width of CL2 1/4- 1/3 basal width of primary CL1. Carinal proportions similar to, but smaller than, those of
rostral plate. Alae small; superior alar margins sub-parallel to basis; growth ridges directed upward, parallel to
inferior alar margin, then curving outward toward alar angle and again upward at sharp angle to form narrow, ridged
welting; inferior alar margin curved, without distinct hook at free end. Orifice small compared to basis, slightly
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
257
toothed. Basis sub-circular; calcareous, thick. S triangular, apex acute; sinuous basal margin 4/7 length of
occludent margin; basal margin just shorter than straight articular margin; occludent margin longest margin;
externally sinuous growth ridges prominent, wide, not cut by external longitudinal ridges; internally adductor
muscle pit well defined, roughened area dislally above adductor muscle pit; lateral depressor muscle pit deep, crests
for lateral depressor muscles absent; adductor ridge feebly developed; articular ridge slightly projecting, extending
2/3 to 3/4 length of articular margin; articular margin rounded, slightly folded. T distinctly smaller than S; apex
bluntly beaked; externally sinuous growth ridges less defined than those of S; articular margin slightly curved;
carinal margin convex; basal margin slightly hollowed out; spur roundly truncated, rounded. 3/10 width of basal
margin, set at less than 1/2 its own width from basi-scutal angle; spur furrow open, deep, flat; articular ridge not
projecting, barely traceable; 6 prominent muscle attachment crests for depressor muscles not extending below basal
margin; inner surface of valve roughened over 2/3 of surface. Color of parietal plates dirty white, S and T whitish
externally and internally, S with golden yellow epidermis and golden, chitinous setae. Eggs orange. Approximate
measurements of holotype as follows;
Labrum with shallow medial depression, with fine
small setae centrally and few small teeth on each side.
Mandibular palp ovate; long, finely serrate setae
terminally. Mandible with 4 teeth, tooth 1 largest, well
separated from teeth 2-4; teeth 2 and 3 each with
subsidiary small tooth; margins of teeth 1-3 smooth,
upper and lower margins of 4 with irregular subsidiary
cusps; inferior angle small, molariform, slightly dentate.
Maxillule setose; 2 long, stout setae at upper angle;
notch below upper angle wide, with 3-4 pairs of smaller
setae; cutting margin below stepped, with 5-7 pairs of
longer, stout setae; inferior angle not protuberant or
stepped, with 4-5 pairs of small setae. Maxilla wide,
coxal endite well developed; setae long, dense, finely
serrulate.
Cirrus I with rami unequal; anterior ramus longer than
posterior ramus; proximal segments of anterior ramus not
protuberant anteriorly, segments wider than those of
posterior ramus; both rami with segments moderately setose. Cirrus 11 longer than cirrus I; rami subequal; all
segments moderately setose, setae finely serrulate. Cirrus III longer than cirri I and II and similar to cirri IV- VI;
rami unequal, anterior ramus shorter than posterior ramus; segments of both rami becoming antenniform distally;
posterior ramus with 2-3 pairs of long, finely serrulate setae on anterior faces of distal segments; anterior ramus
with 2-3 pairs of long, finely serrulate setae on anterior faces of distal segments. Cirrus IV to VI similar, longer
than cirrus III; rami subequal; segments oblong, with 2 pairs of long, serrulate setae on anterior faces. Chaetotaxy
ctenopod. Cirral formula is as follows:
Fig. 59. — Hexelasma foratum sp. nov. Holotype
from Musorstom 6, stn DW 400 (MNHN-Ci
2707): oblique view. Scale = 1 cm.
Penis 1/3 length of cirrus VI; finely annulated; with few sparse setae; circlet of longer setae distally. Eggs
large, rounded, 0.45 x 0.51 mm.
258
D. JONES
Fig. 60. — Hexelasma foratum sp. nov. Paratype (ovigerous) from Musorstom 6, sin DW 451 (MNHN-Ci 2416): a. right
mandible; b. left mandible; c, right maxillule; d. left maxillule; e, maxilla; f. mandibular palp; g. labrum; h, cirrus 1;
i. cirrus II; j, cirrus III; k. cirrus VI and penis.
Remarks. — The minute pinprick holes, which correspond to the edges of the longitudinal ribs on the inner
surface of the parietes, and which perforate the junction of the basis and the inner surfaces of the parietal plates, are
unique structures not found in other hexelasmatines. Hexelasma foratum sp. nov., with its low profile and high
alae nearly parallel to the basis, stands closer to H. brintoni , H. triderma and H. flavidum sp. nov. in form than to
the other known members of the genus, where the alae are oblique.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
259
Hexelasma foratum may be separated from H. triderma by the form of the chitin in the paries; in the former
long, spaced rods of chitin bisect the thickness of the paries, in the latter the chitinous layer is developed as
an almost continuous, thin sheet between the laminae which are about equal in thickness. In H. foratum the basal
width of CL2 is 1/4- 1/3 the basal width of CL1; in H. triderma the basal width of CL2 is 1/2-2/3 the basal width
of CL1. The parietal plates of H. foratum are thick and rugged, externally fine growth ridges marked by long,
golden, chitinous setae and with the remains of yellow epidermis; the robust, thicker plates of H. triderma are
covered with a brownish cuticle marked by heavy, transverse growth ridges supporting numerous chitinous setae.
In H. foratum the basis is calcareous and thick; in H. triderma it is thin, calcareous and marginally thickened. No
opercular or soft parts are known for H. triderma.
The shape of Hexelasma foratum sp. nov. is a truncated cone; that of H. flavidum sp. nov. is rounded, squat
and flattened. Longitudinal rods of chitin divide the paries into a thicker outer (3/4) and thinner inner (1/4) laminae
in H. flavidum. and into laminae of equal width in H. foratum. In H. foratum the basal width of CL2 is 1/4- 1/3 the
basal width of CL1, and there is a yellow, persistent epidermis covering the parietal plates; in H. flavidum the
basal width of CL2 is 1/3 or less the basal width of CL1 and a persistent epidermis is absent. The basis is thick
and calcareous, with pinprick holes perforating the parietal/basal junction in H. foratum-. that of H. flavidum is
thin, calcareous at the periphery and membranous centrally, and perforations are absent. In H. foratum the S basal
margin is 4/7 the length of the occludent margin, crests for the lateral depressor muscles are absent, the articular
ridge is slightly projecting, and the inner surface is roughened distally. In H. flavidum the S basal margin is 9/14
the length of the occludent margin, slight crests for the lateral depressor muscles are present, the articular ridge is
not projecting, and the inner surface is smooth. The T spur of H. foratum is 3/10 the width of the basal margin
and set at 1/2 its own width from the basi-scutal angle; in H. flavidum it is 1/3 the width of the basal margin, set
at 1/4 its width from the basi-scutal angle, and forms an abrupt angle of 130° with the basal margin. The color of
S and T arc whitish in H. foratum, orange-cream in H. flavidum.
Hexelasma foratum sp. nov. can be distinguished from H. brintoni as the basal width of CL2 of the former is
1/4- 1/3 the basal width of CL1, rather than 3/5 the width. The basis is thick and calcareous, with pinprick holes
perforating the parietal/basal junction in H. foratum-, the basis is thick and calcareous, with no perforations in
H. brintoni. The new species can be further separated by its low profile and truncated cone shape, compared to the
conical form of H. brintoni. The T spur of H. foratum curves into the basal margin, rather than forming an abrupt
angle of 130° with the basal margin, as in H. brintoni, and the T is internally roughened over 2/3 of the surface,
rather than smooth. In H. foratum the S basal margin is 4/7 the length of the occludent margin, the internal
surface is roughened distally, crests for the lateral depressor muscles are absent and the articular ridge is slightly
projecting. In H. brintoni the S basal margin is 2/3 the length of the occludent margin, the internal surface is
smooth, there are several crests for the lateral depressor muscles, and the articular ridge is not projecting.
ETYMOLOGY. — From the Latin foratum, ‘to perforate’, in reference to the small pinprick holes, correspond¬
ing to edges of parietal tubes, which perforate the junction between the basis and the inner surface of the parietal
walls.
Distribution. — New Hebrides Arc, Loyalty Islands, 270-500 m.
Hexelasma globosum sp. nov.
Figs 61-64; Tables 28-31
Material EXAMINED. — New Caledonia. Musorstom 4; stn DW 156, 525 m: 3 specimens. — Stn CP 167.
575 m: 6 specimens, attached to pumice and coral rubble (MNHN-Ci 2418). Drawn. — Stn CP 197. 550 m: 5 specimens
(MNHN-Ci 2417, WAM 258-96).
Loyalty Islands. BlOCAl.: stn CP 45. 430-465 m: 1 specimen.
Musorstom 6: stn DW 486. 370 m: 2 specimens, attached to surface of sponge.
Types. — Holotype : MNHN-Ci 2417 (Musorstom 4, stn CP 197).
260
D. JONES
Paratypes : MNHN-Ci 2418 (MUSORSTOM 4: sin CP 167). Drawn. — WAM 258-96 (MUSORSTOM 4.
stn CP 197).
Diagnosis. —Thin parietal plates with thin lamina of chitin developed in longitudinal strips on inner surface
of paries; bases of plates slightly thickened to form rim. Basal width of CL2 1/2 basal width of CL1. Basis
calcareous, thin. Inferior margin of alae curved with slightly basally directed hook at free end; externally growth
ridges well spaced, marked by fine tubercles; ridged welting along superior alar margin, increasing in width toward
free end. Opercular plates whitish. S internal surface smooth; crests for lateral depressor muscles absent; externally
well-spaced, sinuous growth lines cut by 3 slight longitudinal striae, middle stria most marked. T spur truncate,
1/4 width of basal margin, set at distance of less than 1/2 its own width from basi-sculal angle; inner surface
roughened over 2/3 of surface. Mandible with upper and lower margins of tooth 4 with irregular, small subsidiary
cusps.
Fig. 61. — Hexelasma globosum sp. nov. Paratype from Musorstom 4, stn CP 167 (MNHN-Ci 2418): a, S, external
view; b, S, internal view; c, T, internal view; d, T. external view.
Description. — Size moderate (maximum: RC length 12.5, C height 14.0). Shell form globose or sub
globose; rostral plate, CL1 and CL2 sloping inward toward orifice, C upright or sloping outward from orifice, apex
slightly retroverted; opercular plates erect in orifice. Thin parietal plates; externally irregular, faint growth ridges
defined by well spaced, small tubercles, and fine setae; bases of plates thickened to form rim. Thin lamina of chitin
applied in longitudinal strips to inner surface of paries; bases of plates slightly thickened to form rim. Rostral
plate wide, slightly convex. Basal width of CL' less than 1/2 basal width of rostral plate; basal width of CL2 1/2
basal width of CL1, narrow distally. C as wide as, but higher than, rostral plate; transversely bent, V-shaped from
above. Alae of C and CL2 well developed. Summits of alae oblique, especially those of C; growth ridges well
spaced, marked by fine tubercles, directed upward parallel to inferior alar margin then curving outward and slightly
downward to alar angle before upward again at sharp angle to form ridged welting, latter narrowing toward summit,
widening toward alar angle; superior margin of welting serrate due to well spaced growth ridges; inferior margin of
alae curved with hook slightly basally directed. Basis calcareous, thin. Orifice triangular; toothed; wider than basis.
S triangular, apically acute; basal margin 2/3 length of occludent margin; occludent margin longest margin, basal
margin longer than articular margin; externally sinuous growth ridges well spaced, cut by 3 slight longitudinal
ridges in some specimens, shallow middle ridge most obvious; internal surface smooth; adductor and depressor
muscle pits absent, crests for lateral depressor muscles absent; adductor ridge feeble, often visible only on 1 valve;
articular lurrow shallow; articular ridge extending 1/2 length valve, slightly projecting; occludent margin folded,
articular margin deeply folded. T thin, smaller than S, apex bluntly beaked; externally sinuous growth ridges
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
261
strong; articular margin almost straight, carinal margin convex; basal margin hollowed out; spur narrow, truncated,
1/4 width of basal margin, set at distance of less than 1/2 its own width from basi-scutal angle; spur furrow open,
shallow wide; internally articular ridge prominent in upper 1/4 of valve; 7 well developed depressor muscle
attachment crests, slightly extending below basal margin; inner surface of valve distinctly roughened over
2/3 surface; articular ridge not projecting. Parietal plates whitish with yellow tinge; alae pale yellow; opercular
plates whitish. Measurements of holotype as follows:
Fig. 62. — Hexelasma globosum sp. nov.
Holotype from MUSORSTOM 4, stn
CP 197 (MNHN-Ci 2417): lateral view.
Scale = 5 mm.
Medial depression of labrum with regular row of small teeth.
Mandibular palp ovate; long, finely serrate setae terminally.
Mandible with 4 teeth, tooth I largest, well separated from teeth
2-4; margins of teeth 1-3 smooth, upper and lower margins of 4
with irregular subsidiary cusps; inferior angle bluntly molariform,
slightly dentate. Maxillule setose; 2 long, stout setae at upper
angle; notch below upper angle wide, with 4-5 pairs of smaller
setae; cutting margin below stepped, with 5-7 pairs of longer,
stout setae; inferior angle slightly stepped, with 5-7 pairs of small
setae. Maxilla wide, coxal endite well developed; setae long, dense,
finely serrulate.
Cirrus I with rami unequal; anterior ramus slightly longer than
posterior; proximal segments of anterior ramus not protuberant
anteriorly, segments wider than posterior ramus; both rami with
segments thickly setose. Cirrus II longer than cirrus I: rami
subequal; all segments thickly setose, setae finely serrulate. Cirrus
III longer than cirri I and II, similar to cirri IV-VI; rami subequal,
segments slightly antenniform distally; anterior and posterior
ramus with 1-2 pairs of long, finely serrulate setae on anterior
faces of distal segments. Cirrus IV to VI similar, longer than
cirrus III; segments oblong, with 2 pairs of long, serrulate setae
on anterior faces. Cirral formula as follows:
Penis equal to length of proximal segment of pedicel of cirrus VI; finely annulated; with few sparse setae;
irregular circlet of longer setae distally.
Remarks. — Hexelasma globosum sp. nov. is similar to H. leptoderma but the form of the latter is conical
and high with the orifice much smaller than the basis, rather than sub-globose with a large, triangular, toothed
orifice which is wider than the basis. Hexelasma leptoderma has yellowish plates externally due to a persistent
yellow epicuticle; there is no epicuticle associated with H. globosum and the plates are whitish with a yellow
tinge. The alar growth ridges of H. globosum turn slightly downward before reaching the superior alar margin to
form a slight hook, rather than turning abruptly downward, forming a definite hook. A thin lamina of chitin is
developed in longitudinal strips on the inner surface of the parietal plates of H. globosum, rather than contiguous
chitinous rods forming a continuous layer between the inner and the outer chitinous laminae. The S and T of
H. globosum are not covered with a persistent yellow epidermis; the S and T of H. leptoderma are covered by
262
D JONES
a persistent yellow epidermis. The T spur of H. globosum is truncated; that of H. leptoderma pointed.
In H. globosum the scutal adductor muscle pit is not defined, rather than being distinct, the depressor muscle pits
are absent rather than present, the adductor ridge is feebly developed, often on one valve only, rather than being
developed on both plates, and the occludent and articular margins are deeply folded, rather than only the articular
margin being slightly folded. The labrum of H. globosum has a medial depression with a regular row of
approximately 70 small teeth, that of H. leptoderma has 42 teeth.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
263
ETYMOLOGY. — From the Latin globosus, 'globose’, in reference to the globular form of the shell.
Distribution. — New Caledonia, 525-575 nr, Loyalty Islands, 370-465 m.
Fig. 64, — Hexelasma Hoek, 1913 (continued). Distribution map: □, H. triderma (549 m); ★, H. velutinum (204-390 m);
A , H. leptoderma (290 m); O. H. globosum (370-575 m); •. H. persicum (275-585 m); ■ H. sandaracum (170-
1300 m); ▲. H. nolearia (741 m); O, H. gracilis (1024-1029 m).
Hexelasma gracilis Foster, 1981
Fig. 64; Tables 28-31
Hexelasma fasten Foster, 1978: 83 (part).
Aaptolasma fasten - FOSTER, 1978: (part), fig. 50. pi. 10 F.
Hexelasma gracilis Foster, 1981: 361.
Material EXAMINED. — New Zealand. NZOl: stn E 852, 33°38.00'S, 170°55.00'E, 1024-1029 m. 17.03.1968:
holotype (NZOl Type H-357).
Types. — Holotype: NZOl H-357; New Zealand, NZOl stn E 852, 33°38.00'S. 170°55.00'E, 1024-1029 m.
17.03.1968.
Holotype depository: NIWA.
Diagnosis. — Parietal plates with longitudinal chitin-fi lied tubes arcuate in cross-section, separated by
regularly spaced, broad, calcareous bridges, dividing plate into laminae of equal width. Basal width of CL2 sub¬
equal to basal width of CL1. Alae externally with growth ridges well marked, narrow ridged welting along superior
alar margin. Basis with wide calcareous ledge. Opercular plates internally dull orange. S internal surface smooth;
crests for lateral depressor muscles faint; externally widely-spaced, sinuous growth lines not cut by longitudinal
striae. T slender; spur rounded, 1/4 width of basal margin, set at 1/2 its own width from basi-scutal angle; inner
surface smooth. Mandibular tooth 2 with side tooth.
Distribution. — New Zealand. 1024-1029 m.
264
D. JONES
Hexelasma leptoderma (Newman & Ross, 1971)
Fig. 64; Tables 28-31
Aaptolasma leptoderma Newman & Ross, 1971: 165. figs 79-80, pis 33 A-B. 40; 1976: 46. — FOSTER, 1978: 79.
Hexelasma velutinum - Broch, 1931: 53 (? part). ? non Hexelasma velutinum Hoek, 1913.
Hexelasma leptoderma - FOSTER, 1981: 356.
Material examined. — Indonesia. Danish Exped. 1922: stn 58, Kci islands, 5°29’S, 132°37'E. 290 m,
12.05.1922: holotype (ZMUC Cru 1979).
Types. — Holotype: ZMUC Cru 1979; Indonesia, Danish Exped. 1922, stn. 58, Kei Islands, 5°29'S,
132°37'E, 290 m, 12.05.1922: 3 slides attached to brachiopod.
Holotype depository: ZMUC.
DIAGNOSIS. — Parietal plates with longitudinal chitin-filled tubes contiguous, forming almost continuous
layer between inner and outer calcareous laminae, inner lamina and chitinous layer thin compared to the outer
lamina. Basal width of CL2 1/2 basal width of CL1. Inferior margin of ala turning abruptly downward to form
hook at free end; externally with fine growth ridges, thin welting along superior alar margin. Basis calcareous,
thin. S internal surface roughened distally; crests for lateral depressor muscles present; externally widely-spaced,
smooth growth lines not cut by longitudinal striae. T spur pointed, 1/4 width of basal margin, set at distance of
1/2 its own width from basi-scutal angle; inner surface roughened distally. Mandible with upper and lower margins
of tooth 4 with small subsidiary cusps.
Remarks. — Hexelasma globosum sp. nov. is similar to //. leptoderma. The differences between the 2 species
are listed under Hexelasma globosum sp. nov.
Distribution. — Kei Islands, 290 m.
Hexelasma nolearia (Foster, 1978)
Fig. 64; Tables 28-31
Aaptolasma nolearia Foster, 1978: 85, fig. 51, pi. 10 G-H.
Hexelasma nolearia - Foster, 1981: 356, 361.
Material EXAMINED. — New Zealand. NZOI: stn DI59. 49°01.00'S, 164°30.00'E, 741 m, 17.01.1964:
holotype (NZOI Type H-206).
Types. — Holotype: NZOI H-206; New Zealand, NZOI stn D 159, 49°01.00'S, 164°30.00'E, 741 m,
17.01.1964.
Holotype depository: NIWA.
Diagnosis. — Parietal plates with pairs of chitinous ribbons, situated inward from grooves between
prominent ribs of inner surface, dividing plates into thicker outer and thinner inner laminae. Basal width of CL2
1/2 basal width of CLL Inferior margin of ala not curved, with no basally directed hook at free end; narrow welling
along superior alar margin. Basis calcareous. S occludent and articular margins folded; internal surface smooth;
faint crests for lateral depressor muscles; externally, well-spaced, smooth growth lines not cut by longitudinal
striae. T articular margin convex; spur indistinct, 1/3 width of basal margin, set at basi-scutal angle; inner surface
roughened distally. Mandible with upper and lower margins of teeth smooth.
Distribution. — New Zealand, 741 m.
Source : MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
265
Hexelasma persicum sp. nov.
Figs 64-67; Tables 28-31
MATERIAL EXAMINED. — Chesterfield Islands. Musorstom 5: sin DW 338, 540-580 m: I specimen (WAM
259-96).
New Caledonia. Musorstom 4: stn DW 159, 585 m: 1 specimen, attached to stone. — Stn CP 179, 475 m:
1 specimen (BMNH). — Stn DW 197, 550 m: 1 specimen, attached to shell of gastropod (USNM). — Stn DW 222. 410-
440 m: 1 specimen. — Stn DW 223, 545-560 m: 1 specimen.
Beryx 2: stn CP 46, 300-350 m: 2 specimens.
Smib 8: stn CP 180, 425-460 m: 1 specimen. — Stn DW 185, 305-355 m: 1 specimen, 1 incomplete specimen. —
Stn DW 193, 500-508 m: several specimens. — Stn DW 194, 491 m: 3 specimens.
Bathus 4: stn DW 929, 502-516 m: several specimens.
Loyalty Islands. Biocal: stn DW 66, 505-515 m: I specimen, attached to coral.
Musorstom 6: stn DW 478, 400 m: several specimens, some attached to sponge (MNHN-Ci 2419, Ci 2420). Drawn.
— Stn DW 478, 400 m: 1 specimen, attached to rocks.
Norfolk Ridge. CHALCAL 2: stn CH 9, 300 m: 2 specimens, attached to sponge.
Smib 4: stn DW 60, 535 m: 3 specimens.
Smib 5: stn DW 94, 275 m: 2 specimens, attached to Rhizotrochus sp.
Bathus 3: stn 818, 405-41 1 m: 2 specimens. — Stn CH 820, 394-401 m: 1 specimen, attached to gastropod shell. —
Stn DW 830, 361-365 m: 2 specimens.
Types. — Holotype : MNHN-Ci 2419 (Musorstom 6, stn DW 478).
Paratypes : BMNH (Musorstom 4, stn CP 179). — MNHN-Ci 2420 (Musorstom 6, stn DW 478). Drawn.
— USNM (Musorstom 4, stn DW 197). — WAM 259-96 (Musorstom 5, stn DW 338).
DIAGNOSIS. — Parietal plates with chitinous rods separating parietes into inner (1/3 thickness) and outer (2/3
thickness) laminae. Basal width of CL2 1/2 basal width of CL1. Inferior margin of ala curved, with blunt, basally
directed hook at free end; ridged welting along superior alar margin, increasing in width toward free end. Basis
calcareous; thin. Opercular plates peachy-orange. S internal surface smooth; crests for lateral depressor muscles
feeble or absent; externally sinuous, narrowly-spaced, growth lines cut by 3 faint longitudinal striae, 2 near
occludent margin. 1 near articular margin. T spur truncate, 1/4 width of basal margin, set at distance of 1/3 its own
width from basi-scutal angle; inner surface roughened distally. Mandibular tooth 2 with 1 or a few small subsidiary
cusps, upper and lower margins of tooth 4 with small, irregular subsidiary cusps.
Fig. 65. — Hexelasma persicum sp. nov. Paratype from Musorstom 6, stn DW 478 (MNHN-Ci 2420): a. S, external view;
b, S, internal view; c, T, internal view; d, T. external view.
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D. JONES
DESCRIPTION. — Size large (maximum: RC length 19.4, C height 16.9 ). Shell tall, nearly vertical. Parietal
plates thick, covered by thin, pale cream, persistent cpicuticle; growth ridges irregular, marked by small setae and
minute calcareous knobs; basally inflected, forming thin rim; internally ribbed parietal walls with ribs prominent
basally; chitinous rods separating parietes into inner (1/3 thickness) and outer (2/3 thickness) laminae. Rostial
plate convex, wide basally; apex curving in toward orifice. Basal width of CL1 1/2 basal width of rostral plate.
Basal width of CL? 1/2 basal width of CL'; triangular. C highest plate; apex curving toward orifice. Alae wide,
broadly triangular, extending full height of parietes; summits oblique; growth ridges directed upward parallel to
inferior alar margin before curving downward and outward toward alar angle, then sharply upward to form ridged
welting, increasing in width toward free end; inferior alar margin curved, with blunt, basally directed hook at Irec
end. Orifice triangular; deeply toothed. Basis thin, calcareous. S triangular, elongated, apically sharply acute;
occludent margin longer than other margins, basal margin 9/16 length of articular margin; externally with
persistent ochre-yellow epidermis; sinuous growth ridges narrowly spaced, cut by 3 narrow, longitudinal ridges
(some specimens with 1), 2 near to occludent margin, extending from apex to basal margin, I close to articular
margin, traceable through 3/4 of valve, well-marked in lower half; basal margin sinuous, sometimes notched at
bases of ridges; internally with indistinct, oval adductor muscle pit; adductor ridge feeble; lateral depressor muscle
pit faint, feeble muscle attachment ridges on S; articular furrow shallow; articular ridge extending 2/3 margin,
slightly projecting; articular and occludent margins rounded, folded, inner edges sinuous. T slightly smaller than S;
apex blunt; externally sinuous growth ridges less defined than on S; articular margin straight/slightly convex;
carinal margin straight; basal margin hollowed out; spur basally truncate, pointed toward and set close to basi-
scutal angle; spur furrow open, slightly sunken, broad basally; internally with articular ridge not projecting;
7 distinct and 3 smaller muscle attachment crests for depressor muscles, not extending below basal margin; inner
surface of valve lightly roughened with irregular beading in distal half. Color ol parietal plates dirty white; cleaned
specimens with some peachy-orange on parietes and alae. Apical and carinal areas ol 1 often peachy-orange color
internally and externally; remainder of valve whitish. Externally apical tip ol S. and internally and externally
central lower parts of valve tinged peachy-orange in some specimens, valve whitish elsewhere. Measurements ol
holotype as follows;
Fig. 66. — Hexelasma persicum sp. nov. Holotype from
Musorstom 6. stn DW 478 (MNHN-Ci 2419): lateral
view. Scale = 1 cm.
protuberant anteriorly, segments wider than posterior
Labrum with shallow medial depression provided
with row of bifid teeth and dense, short setae.
Mandibular palp ovoid; long, finely serrate setae
terminally. Mandible with 4 teeth, tooth 1 largest, well
separated from teeth 2-4; tooth 2 with 1 to several
small subsidiary teeth, tooth 3 with 2 subsidiary teeth,
tooth 4 with upper and lower margins irregularly
dentate; inferior angle large, molariform, irregularly
dentate. Maxillule setose; 2 long, stout setae at upper
angle; notch below upper angle with 3-5 pairs of
smaller setae; cutting margin below almost vertical,
stepped, with 8-10 pairs of longer, stout setae; inferior
angle barely delineated, with 4-6 pairs of smaller setae.
Maxilla wide, coxal endite barely defined; area between
basal and coxal endite with small spines; setae long,
finely serrulate.
Cirrus I with rami unequal; anterior ramus longer than
posterior; proximal segments of anterior ramus not
ramus, becoming antenniform distally; both rami with
Source : MNHN, Paris
CIRR1PED1A THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
267
1.0 mm
0.5 mm
0.5 mm
1.0 mm
k
Fig. 67. — Hexelasma persicum sp. nov. Paratype from Musorstom 6, stn DW 478 (MNHN-Ci 2420): a. right mandible;
b, left mandible; c, right maxillule; d, left maxillule; e, labrum; f. maxilla; g. mandibular palp; h. cirrus II; i. cirrus
VI and penis; j. cirrus III; k, cirrus I.
segments setose; setae finely serrulate distally. Cirrus II longer than cirrus I; rami unequal, posterior ramus longer
than anterior ramus; all segments setose, especially proximally. Cirrus III much longer than cirri I and II and
similar to cirri IV- VI; rami subequal, segments becoming oblong distally; anterior and posterior ramus with
2 pairs of long, finely serrulate setae on anterior faces of distal segments, proximal segments densely setose.
Cirrus IV to VI similar, longer than cirrus III; rami subequal; segments oblong, with 2 pairs of long, finely
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D. JONES
serrulate setae on anterior faces; proximal segments wider than long, becoming oblong distally. Cirral formula as
follows:
Penis 1/2 length of cirrus VI; annulated; with few sparse setae; irregular circlet of longer setae distally.
Remarks. — Hexelasma persicum sp. nov. is similar to //. arafurae but differs by several characteristics of the
hard parts. The shell of H. persicum is taller, more upright and with a more deeply toothed orifice than that of
H. arafurae. In H. persicum chitinous rods separate the plates into inner (1/3 thickness) and outer (2/3 thickness)
laminae, rather than a continuous chitinous lamina bisecting the thickness of the parietal plates as in H. arafurae
(FOSTER, 1981). A persistent epicuticle covers the shell plates of H. persicum but is absent in H. arafurae. The
basal width of CL2 is 1/2 the basal width of CL1 in H. persicum and more than 1/2 the basal width of CL1 in
H. arafurae. The welting of the alae of H. persicum is ridged, the welting increasing in width towards the free end;
in H. arafurae the alae have strongly marked growth ridges, but the welting along the superior alar margin is
inconspicuous. Externally the S of H. persicum has sinuous, narrowly spaced growth lines cut by 3 faint
longitudinal striae, whilst that of H. arafurae has smooth, even lines cut by I broad, distinct longitudinal stria; and
the S inner surface of H. persicum has an indistinct adductor muscle pit, rather than deep and distinct as in
H. arafurae. The T spur of H. persicum is 1/4 the width of the basal margin, basally truncate and pointed toward
the basi-scutal angle, rather than greater than 1/3 the width of the basal margin and basally rounded. In H. persicum
the inner surface of T is lightly roughened with irregular beading, rather than smooth, and both S and T are colored
peachy orange to white, rather than brick-red as in H. arafurae. There are also differences in the mouthparts and
cirri. The labrum of H. persicum bears small teeth and setae centrally and small setae laterally, rather than small
setae in the central region and small teeth laterally, and the cutting edge of the maxillule is distinctly, rather than
barely, notched. Cirrus III of H. persicum has subequal rami, rather than the posterior ramus being antenniform and
very much longer than the anterior ramus. The cirral counts for H. persicum are generally higher than those for
H. arafurae, especially for cirri IV-VI.
FOSTER (1981) concluded that HOEK's (1913) specimen from "Siboga", station 251 was H. arafurae. based on a
continuous chitinous lamina, an inflected basal ledge and a thin calcareous basis. The elongated S was as in
H. arafurae and cirral counts agreed better with H. arafurae than with the H. velutinum type specimens. However,
the tergal spur differed from that figured by Hoek for H. arafurae, so ".... a definite conclusion about the
synonymy must await further material and appreciation about the extent of variation." (FOSTER, 1981). The subtle
yet consistent inter-specific variation found within the genus Hexelasma, indicated by the present study and alluded
to by Newman and ROSS (1971), further supports Foster’s opinion.
ETYMOLOGY. — Named from the Latin persicus, 'peach-tree', in reference to the peachy-orange color on areas
of the shell and opercular plates.
Distribution. — Chesterfield Islands, 540-580 m; New Caledonia, 300-585 m; Loyalty Islands, 400-515 m;
Norfolk Ridge, 275-535 m.
Hexelasma sandaracum sp. nov.
Figs 64. 68-70; Tables 28-31
Material EXAMINED. — Vanuatu. Musorstom 8: stn CP 972, 487-507 m: 4 specimens. — Stn CP 973, 460-
480 m: 7 specimens. — Stn CP 974, 492-520 m: many specimens (WAM 260-96). — Stn DW 978, 413-408 m:
7 specimens. — Stn DW 989, 650-669 m: 2 specimens. — Stn CP 1026, 437-504 m: 1 specimen. — Stn CP 1027. 550-
571 m: 4 specimens. — Stn CP 1045, 459-488 m: many specimens. — Stn DW 1046, 461-480 m: 1 specimen. —
Stn DW 1067, 344-366 m: 2 specimens. — Stn CP 1080, 799-850 m: 1 specimen. — Stn CP 1083. 397-439 m:
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYl.ASMATOIDEA
269
1 specimen. — Stn CP 1114, 647 m: 1 specimen. — Stn CP 1125, 1160-1220 m: many specimens. — Stn CP 1127.
1052-1058 m: 1 specimen. — Stn CP 1136, 398-400 m: 2 specimens.
New Caledonia. — Lagon: stn 491, 450 m: 1 specimen, attached to solitary coral.
Musorstom 4: stn DW 156, 530 m: 5 specimens (3 adults, 2 juveniles). — Stn DW 156, 525 m: 4 specimens. —
Stn DW 156, 525 m: 3 specimens. — Stn CP 157, 575 m: 3 specimens, attached to pumice and gastropod shells. —
Stn DW 159, 585 m: 1 specimen, attached to rock. — Stn CP 167, 575 m: 2 specimens, attached to siliceous sponge. —
Stn CP 169, 590 m: 2 specimens. — Stn CP 170, 480 m: 2 specimens (1 attached to gastropod shell). — Stn CP 178,
520 m: 1 specimen, attached to fragment of echinoderm test. — Stn CP 194, 545 m: 3 specimens (MNHN-Ci 2421, Ci
2705). Drawn; many specimens (MNHN-Ci 2422, BMNH. USNM). — Stn CP 197, 550 m: 5 specimens (1 free. 2
attached to dead coral, 2 attached to gastropod shells); 3 specimens (2 attached to dead coral, I to shell of gastropod).
Beryx 2: stn DW 34, 550-570 m: 1 specimen. — Stn DW 35. 550-570 m: 1 specimen. — Stn DW 38. 550-690 m:
2 specimens (dry), from shell.
Smib 8: stn DW 189, 400-402 m: 2 specimens.
Bathus 2: stn DW 729. 400 m: 1 specimen (dry), from shell.
Bathus 4: stn CP 910, 560-608 m: 3 whole specimens; 2 incomplete specimens; 1 specimen (disarticulated). —
Stn CP 911, 566-558 m: 2 specimens (dry), from shell; many specimens, attached to pumice and gastropod shells. —
Stn DW 916, 600-616 m; 4 specimens, attached to dead coral rubble. — Stn DW 923, 470-502 m: many specimens,
attached to dead coral rubble. — Stn DW 927, 452-444 m: 4 specimens, attached to dead coral rubble. — Stn DW 928.
420-452 m: 3 specimens (dry), from shell. — Stn CP 929, 502-516 m: 4 specimens; 1 specimen, attached to gastropod
shell; 1 specimen (dry), broken, from shell. — Stn DW 932, 170-190 m: 1 specimen (juvenile). — Stn CP 951. 960 m: 1
specimen (disarticulated).
Loyally Islands. BiOCAL: stn DW 36. 650-680 m: 2 specimens, attached to rock (WAM C 23259).
Musorstom 6: stn CP 401, 270 m: 1 specimen. — Stn DW 424, 599 m: I specimen. — Stn CC 470, 560 m:
1 specimen, attached to stone. — Stn DW 471, 460 m: 2 specimens, attached to corals. — Stn DW 483. 600 m:
2 specimens, attached to rock.
Norfolk Ridge. Smib 4: stn DW 44, 300 m: 1 specimen, attached to sponge. — Stn DW 62. 540 m: 2 specimens.
— Stn DW 63, 520 m: 1 specimen, attached to dead coral. — Stn DW 64, 460 m: 4 specimens (2 adults, 2 juveniles).
Bathus 3: stn CP 811, 383-408 m: 1 specimen (dry), broken, from shell.
Combe Bank. Musorstom 7: stn DW 539, 700 m: 1 specimen. — Stn CP 622, 1280-1300 m; 1 specimen, attached
to rock.
Wallis Islands. MUSORSTOM 7; stn DW 522, 650-765 m: 1 specimen, attached to rock. — Stn DW 522. 650-
765 m: 1 specimen, attached to rock. — Stn DW 582, 360 m: 3 specimens, attached to rock.
Tuscarora Bank. Musorstom 7: stn CP 559, 547-552 m: 4 specimens, 2 attached to rock.
Field Bank. MUSORSTOM 7: stn DW 586, 510-600 m: I specimen, attached to rock (WAM C 23260).
Types. — Holotype : MNHN-Ci 2421 (MUSORSTOM 4, stn CP 194).
Paratypes : BMNH (MUSORSTOM 4, stn CP 194). — MNHN-Ci 2422 (MUSORSTOM 4, stn CP 194). —
MNHN-Ci 2705 (MUSORSTOM 4, stn CP 194). Drawn. — USNM (MUSORSTOM 4, stn CP 194). — WAM 260-
96 (Musorstom 8, stn CP 974).
DIAGNOSIS. — Thin parietal plates with inner surfaces covered with lamina of chitin. Basal width of CL2 1/3-
1/2 basal width of CL1. Inferior margin of ala hollowed out, blunt tooth at free end; external surface with parallel
growth ridges widely spaced; narrow, ridged welting, with serrated superior margin, serrations corresponding to
spacing of growth ridges. Basis calcareous; thin. Opercular plates orange (fresh material) or white (preserved
material). S internal surface smooth; low crests for lateral depressor muscles; externally sinuous, widely-spaced,
growth lines cut by 3 longitudinal striae, 2 near occludent margin, 1 near articular margin; occludent and articular
margins rounded, folded. T spur truncated, 1/4 width of basal margin, set at distance of 1/2 its own width from
basi-scutal angle; inner surface slightly roughened in upper 1/2. Mandible with upper margin of tooth 3 with 1 or
more small, subsidiary cusps, upper and lower margins of tooth 4 with small, irregular subsidiary cusps.
Description. — Size large (maximum: RC length 33.0, C height 27.0 ). Shell with paries erect, not curved
inward toward C. Parietal plates thin; covered with colorless, thin, persistent epidermis; growth ridges well spaced;
marked by fine, ochre setae and small tubercles; basal edges of plates irregularly toothed; inner surface of paries
smooth, covered with lamina of chitin. Rostral plate convex, triangular; height subequal to C height, basal width
greater than that of C. CL1 with apex erect. Basal width of CL2 1/2- 1/3 basal width of CL1; apex suberect.
C strongly bent transversely, more pronounced toward apex with lateral halves meeting at precise angle. Alae
wide, triangular, summits oblique; parallel growth ridges widely set, directed upward parallel to inferior alar margin
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D. JONES
Fig. 68. — Hexelasma sandaracum sp. nov. Paratype from Musorstom 4. stn CP 194 (MNHN-Ci 2705): a, S. external
view; b. S. internal view; c. T. external view; d. T, internal view.
before curving outward and slightly downward toward alar angle and then up again at sharp angle to form narrow,
ridged welting, with serrated upper margin, serrations corresponding to spacing of growth ridges; inferior alar
margin hollowed out, blunt tooth at free end. Orifice pentagonal; toothed. Basis calcareous, thin, uneven;
circumference circular. Opercular plates flush with orifice, inclined at angle to basis. S triangular, sharply pointed
apically, externally gently concave in middle area; basal margin 9/15 length of occludent margin; occludent margin
longest margin, basal margin slightly shorter than articular margin; sinuous growth ridges prominent, widely
spaced, cut by 3 slight longitudinal striae (some specimens with 1), 2 near occludent margin, extending from apex
to basal margin (stria nearest middle of valve shallower, longer than narrower stria nearest occludent margin),
1 near articular margin shallow, extending from apex for 2/3 length of valve; internally adductor muscle scar
shallow, low lateral depressor muscle ridges in slight pit; adductor ridge feeble, often distinct on one S only;
articular furrow shallow; articular ridge extending for 1/2 to 2/3 margin, slightly projecting; articular and occludent
margins rounded, folded. T thin, distinctly smaller than S; apex beaked; externally sinuous growth ridges less
defined than on S; articular margin slightly curved; carinal margin convex; basal margin weakly hollowed out;
broad spur truncate, 1/4 width of basal margin, pointed toward and set at distance of 1/2 its width from basi-scutal
angle; spur furrow open, shallow, flat, broad basally; articular ridge not projecting but traceable through 2/3 length
of valve; 5-7 distinct depressor muscle crests, some slightly extending below basal margin; inner surface of valve
slightly roughened in distal 1/2. Freshly preserved specimens with T vivid orange externally and internally in
middle regions of valve; S similarly colored orange apically, externally and internally; C internally with sheath
vivid orange, externally color suffusing through upper portions of C, including ala. through partly transparent
paries; elsewhere paries externally pale lemon yellow; internally with yellow chitinous layer applied to inner faces.
Long-preserved material with parietal plates externally creamy white-yellowish; summits of alae of C and CL2
orange-tinged; internally parietes yellow; external and internal surfaces of S and T white. Eggs orange.
Measurements of holotype and of largest specimen, respectively, as follows:
Shallow medial depression of labrum with row of well-developed teeth. Mandibular palp oblong; long, finely
serrate setae terminally. Mandible with 4 teeth, tooth 1 largest, well separated from teeth 2-4; tooth 3 with 1 or
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
271
more small subsidiary cusps on upper margin, upper and lower margins of tooth 4 with irregular, small subsidiary
cusps; inferior angle small, molariform, dentate. Maxillule setose; 3 long, stout setae at upper angle; notch below
upper angle wide, with 4 pairs of smaller setae; cutting margin below stepped, sloping, with 7-10 pairs of longer,
stout setae; inferior angle barely demarcated, with 4-6 pairs of smaller setae. Maxilla wide, coxal endite barely
defined; setae long, finely serrulate.
Fig. 69. — Hexelasma sandaracum sp. nov. Holotype from
Musorstom 4, stn CP 194 (MNHN-Ci 2421): lateral
view. Scale = 1 cm.
Cirrus I with rami subequal; anterior ramus longer
than posterior; proximal segments of anterior ramus
not protuberant anteriorly, segments wider than
posterior ramus; both rami with segments setose,
setae finely serrulate distally. Cirrus 11 longer than
cirrus I; rami subequal, all segments setose, setae
finely serrulate. Cirrus III longer than cirri I and II and
similar to cirri IV-VI; rami subequal, segments
slightly antenniform distally; posterior ramus with 1-
2 pairs of long, finely serrulate setae on anterior faces
of more distal segments; anterior ramus with
2-3 pairs of long, finely serrulate setae on anterior
faces of distal segments, proximal segments with
4 pairs of long setae. Cirrus IV to VI similar, longer
than cirrus III; rami subequal; segments becoming
oblong distally, with 2 pairs of finely serrulate, long
setae on anterior faces. Cirral formula as follows:
Penis 1/4 length of cirrus VI; annulated; with few sparse setae; irregular circlet of longer setae distally. Large
ellipsoidal eggs, few in number, 0.35 x 0.25 mm.
Remarks. — Hexelasma sandaracum sp. nov. is similar to H. brintoni but the two can be separated on
characters of the hard and soft parts. The basal width of CL2 of H. sandaracum is 1/2- 1/3 the basal width of CL1
rather than 3/5 the basal width. In H. sandaracum the inferior alar margin is hollowed out, with a blunt,
downwardly directed tooth at the free end, as opposed to curved and without a basally directed hook at the free end,
as in H. brintoni. Chitin is present as a lamina on the inner surface of the paries in H. sandaracum. In II. brintoni
the parietal plates have longitudinal, chitin-filled rods, sub-arcuate in cross-section, regularly spaced and separated
from one another by broad calcareous bridges, which bisect the thickness of the parietal plates. The calcareous
basis is thin in H. sandaracum and thick in H. brintoni. In H. sandaracum the tergal spur is 1/4 rather than 1/3 the
width of the basal margin; and the tergal basal margin rounds into the spur, rather than meeting it at an abrupt
angle as in H. brintoni. The S of H. sandaracum externally has sinuous growth lines cut by 3 longitudinal striae;
that of H. brintoni has smooth, even growth lines not cut by longitudinal striae. In addition the cirral counts are
higher for H. sandaracum than those for H. brintoni.
Hexelasma sandaracum sp. nov. is also similar to H. triderma but there are differences in the overall shell form.
The opercular plates and soft parts of//, triderma are undescribed. In //. sandaracum the shell plates are covered
with a colorless, thin, persistent epidermis, and the growth ridges are well spaced and marked by fine, ochre setae
and small tubercles, rather than covered with brownish cuticle and marked by heavy transverse growth ridges with
numerous setae. In H. sandaracum the alar welting is narrow rather than broad, and the inferior alar margin is
hollowed out, with a blunt, downwardly directed tooth at the free end, rather than curved, with an abrupt, basally
directed hook at the alar angle. The basal width of CL2 of //. sandaracum is 1/2- 1/3 the basal width of CL1 rather
than 1/2-2/3. Chitin is present as a yellow lamina on the inner surface of the paries in H. sandaracum. rather than
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D. JONES
as a thin, chitinous layer formed by laterally elongate, longitudinal, chitin-filled tubes, nearly straight in cross-
section, which separate the wall into equal sized inner and outer calcareous laminae, connected infrequently by
small, inconspicuous calcareous bridges. Opercular and soft parts of H. tridema are unknown.
0.2 mm
a, b
0.5 mm
e-g i,j 9 5 mm h, k
0.5 mm
c, d
Fig. 70. — Hexelasma sandaracum sp. nov. Paratype from Musorstom 4, stn CP 194 (MNHN Ci 2705)’ a right
mandible; b, left mandible; c right maxillule; d, left maxillule; e, labrum (torn); f. mandibular palp; g.' maxilla;
h, cirrus III; 1, cirrus I; j, cirrus II; k, cirrus VI and penis.
Etymology. — From the Greek sandaraca, ’orange-colored’, in reference to the vivid orange color of the
opercular plates and of the large eggs.
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
273
Distribution. — Vanuatu, 344-1220 m; New Caledonia, 170-960 m; Loyalty Islands, 270-680 m; Norfolk
Ridge, 300-540 m; Combe Bank, 700-1300 m; Wallis Islands, 360-765 m; Tuscarora Bank, 547-552 m; Field
Bank, 510-600 m.
Hexelasma triderma (Newman & Ross, 1971)
Fig. 64; Tables 28-31
Aaptolasma triderma Newman & Ross, 1971: 164, pi. 33 C-D, 39 F-G. — Foster, 1978: 83.
Hexelasma triderma - Newman & Ross, 1976: 46. — Foster, 1981: 356.
MATERIAL EXAMINED. — Japan. Jyn (Japanyon) III Exp.: stn 51, E of Honshu Island, 34°54.3'N, 139°43.5'E, 549
m, 17.07.1961: holotype (USNM Type 38690).
Types. — Holotype : USNM 125312; Japan, E of Honshu Island, off Tateyama, Jyn III Exped., stn 51,
34°54.3'N, 139°43.5'E, 549 m, 17.07.1961.
Holotype depository : USNM.
DIAGNOSIS. — Thin parietal plates with thin chitinous layer formed by laterally elongate, thin, longitudinal,
chitin-filled tubes, nearly straight in cross-section, separating wall into inner and outer calcareous laminae
connected infrequently by small, inconspicuous calcareous bridges; inner and outer laminae equal in thickness.
Basal width of CL2 1/2-2/3 basal width of CL1. Inferior margin of alae with abrupt, basally directed hook at alar
angle, exterior surface strongly marked by growth lines; broad welting along superior alar margin. Basis
calcareous; thin centrally, marginally thickened. Opercular and soft parts unknown.
Remarks. — Hexelasma triderma is a large sized species (maximum: RC length 19.5, C height 12.0),
which is superficially similar to H.flavidum sp. nov., H.foratum sp. nov. and H. sandaracum sp. nov.
Differences between H. triderma and these species may be found under H. flavidum, H.foratum and H.
sandaracum , respectively.
Distribution. — Japan, 549 m.
Hexelasma velutinum Hoek, 1913
Fig. 64; Tables 28-31
Hexelasma velutinum Hoek, 1913: 246 (part). — Withers, 1913: 847. — Broch, 1931: 53 (? part). — Hiro, 1933: 70,
pi. 3 fig. 2. — Utinomi, 1968: 30. — Newman & Ross, 1971: 155; 1976: 4. — Foster, 1978: 79; 1981: 356, figs 6
A-E, 7. — Rosell, 1991: 37.
Material EXAMINED. — Indonesia. " Siboga ": stn 105, 6°08'S, 1 21 0 19’E, 275 m, 04.07.1899: 2 specimens
syntypes (ZMUA Cirr. 100.322 a), 2 dry specimens (ZMUA Cirr. 100.322).
Types. — Syntypes : Indonesia, "Siboga" stn 105, 6°08’S, 121°19E, 275 m, 04.07.1899: ZMUA Cirr.
100.322a : 2 specimens, in alcohol; ZMUA Cirr. 100.322: 2 specimens, dry.
Syntypes depository : ZMUA.
DIAGNOSIS. — Parietal plates with lamina of orange chitin applied to inner surfaces. Basal width of CL- 2/3
basal width of CL1. External alar surface with parallel growth ridges; narrow, finely crenated welting along
superior alar margin. Basis calcareous; thin. Opercular plates white. S convex externally; internal surface distal ly
roughened; crests for lateral depressor muscles absent; externally smooth, widely-spaced, growth lines not cut by
longitudinal striae; occludent and articular margins rounded, folded. T spur truncated, more than 1/3 width of basal
margin, set slightly apart from basi-scutal angle; inner surface roughened. Mandible with upper margin of tooth 3
and lower margin of tooth 4 with side teeth, lower angle produced into single oi bifid spine.
274
D. JONES
REMARKS. — Hexelasma velutinum is a large sized species (maximum: RC length 10.0. C height 13.0).
Hexelasma arafurae and H. aureolum sp. nov. arc superficially similar to H. velutinum. Differences between these
species may be found under H. arafurae and H. aureolum, respectively.
Distribution. — Japan, Philippines to South China Sea; 204-390 m.
DISCUSSION
Wall structure in Chionelasmatoidea and Pachylasmatoidea. — A basal whorl, or whorls, ol
imbricating plates is present in the chionelasmatoids Chionelasmus and Eochionelasmus . The wall is 6-plated (R-
RL-CL-C), with the RL derived from an imbricating plate which is added to protect the R/CL suture. I lie upper
portion of RL is not incorporated into the sheath and the lower portion is not fully incorporated into the wall.
In the higher balanomorphs, such as the Pachylasmatoidea. CL2 is added to the wall structure. Within the
Pachylasmatidae, four whorls of imbricating plates are present in the eolasmatine Waikalasma. The wall is 8-plated
(R-RL-CL'-CL2-C), with CL2 added by the replication of CL1, rather than by transfer irom the imbricating
whorls. CL2 only slightly enters the sheath as a very narrow zone. In the pachylasmatine Pachvlasma, imbricating
plates are absent in Recent forms. Imbricating plates are also absent in the new pachylasmatine genera described in
the present paper ( Eutomolasma gen. nov., Eurylasma gen. nov. and Microlasma gen. nov.). In (hese
pachylasmatines the wall is 6-plated (R-CL'-CL2-C), RL and CL2 are incorporated into the wall but RL is not
incorporated into the sheath. In Eurylasma CL2 is partially coalesced with CL1, but CL- enters the sheath as a
very narrow zone. The pachylasmatine Tetrapachylasma, has a wall that may be 6- or 4-plalcd, depending on the
degree of coalescence of the plates. In the 6-plated form (R-CLI-CL2-C). RL coalesces with R to form a compound
rostrum (RL+R+RL). In the 4-plated wall (R-CL-C), RL and R form a compound rostrum (RL+R+RL) and CL2
coalesces with CL1 to form a compound CL plate. CL2 enters the sheath as a narrow zone but RL is not
incorporated into the sheath, with the exception of T. ornatum sp. nov., which shows the incipient inclusion ol
RL into the sheath.
Imbricating plates are absent in the Metalasmatinae, the Bathylasmalinae and the Hexelasmatinae. In the
Metalasmatinae, the Hexelasmatinae and the bathylasmatines Bathylasma, Mesolasma and Tessarelasma , the wall
is 6-plated (R-CL>-CL2-C), RL and CL2 are incorporated into the wall but RL is not incorporated into the sheath.
The bathylasmatinae Tetrachaelasma has a 4 -plated wall (R-CL-C) due to the absence ot CL-.
Thus the RL does not enter the sheath in the Pachylasmatoidea, with the exception of T. ornatum sp. nov. The
early balanids ( Palaeobalanus ) and coronulids ( Chelonibia ) reflect the 8-patterncd form seen in some
pachylasmatoids, including a "tripartite” or compound rostrum (RL-R-RL). In Chelonibia DARWIN (1854) noted
vestigial sutures on the inner surface as evidence for the coalescence of RL with R. 1 he tripartite rostrum ol these
higher forms, however, is distinguished from that of pachylasmatoids by the fact that RL enters the sheath. For
this situation to occur RL must have been incorporated into the sheath before coalescence with R and, therefore,
the two types of tripartite rostra are convergent (BUCKERIDGE & Newman, 1992).
Morphological Characters. — The subtlety of the specific differences within the Pachylasmatoidea can
only be determined by examination of a large number of specimens and ontogenetic series. Intra- and inter-specific
variation can thus be recognised, definition relying on the combination of consistent diflercnces in a number ol
characters (e.g. the form of the S and T, the pattern of the wall plates, the length ot the caudal appendages). BROCH
(1931) first noted the subtle variation found within the Hexelasmatinae in a specimen which he considered to be
H. velutinum. although he commented that the external features of the S differed from HOEK's description of
H. velutinum. On re-examining BROCH’s material, Newman & Ross (1971) noted other subtle character
differences and re-described the specimen as a new species, Aaptolasma leptoderma (Newman & Ross, 1971). These
authors also suggested that HOEK (1913) had confounded 2 species in his description of H. velutinum and it was
Source : MNHN. Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
275
not improbable that the specimens from "Siboga" stations 59 and 251 represented A. leptoderma rather than
H. velutinum. A further re-examination of Hoek’s material by Foster (1981) led him to conclude that in fact
3 species had been confounded in the description - H. velutinum from "Siboga" station 105. A. leptoderma from
HOEK's specimen from "Siboga" station 59 and H. arafurae from "Siboga" station 251. However, FOSTER
commented that the identification of H. arafurae was provisional, as the tergal spur differed from that figured by
HOEK for H. arafurae , so ".... a definite conclusion about the synonymy must await further material and
appreciation about the extent of variation". The studies of Newman and Ross (1971) and Foster (1981) have
described a number of new species within Hexelasma. The present contribution significantly increases the number
of species within Hexelasma from 9 to 15 and further extends the known distribution of the genus in the western
Pacific Ocean.
Species diversity within Pachylasma has also been unrecognised heretofore, primarily due to a paucity of
collections and a lack of comparative material. Many of the original species descriptions have been based on an
individual or few specimens, with some species having never been re-collected. When a species has been re¬
collected at a later date, in some instances it has been from an area distant from the type locality and although
differences (such as in the form of the S, the T, the pattern of the shell plates, the length of caudal appendages)
between the more recently collected specimen(s) and the type specimen have been documented or illustrated, they
have received little comment and. presumably, have been attributed to intra-specific variation. However, re¬
examination of species such as P. scutistriata of Nilsson-Cantell, 1927 (p. 781, fig. 17. pi. 1 fig. 14);
P. scutistriatum orUTlNOMl, 1968 (p. 26, figs 4-5); and P. aurantiacum of FOSTER, 1981 (p. 354. figs 2 E. 4)
may be warranted.
Significance of antenniformy, reproductive state and regression and/or loss of the
penis. — In the Balanomorpha the regression and/or loss of the penis after the reproductive period has been
documented in some species (see review of Barnes. 1992). The amount of regression appears to be related to the
number of broods produced per year. In species producing I brood a year, or with a synchronised first brood of the
year, the penis may be completely lost during the long non-breeding period. However, in other species the variable
reproductive state of the population may mask this loss. A similar connection between the loss or regression of
the penis after reproduction, and which also seems to be related to the number of broods produced in a year, has
been found in the Littorinidae. Based on the above evidence. BARNES (1992) argued that the length of the penis of
an individual is irrelevant in a species description unless the reproductive state of the individual is also given.
A connection exists between antenniformy, the reproductive state of the cirriped and the regression and/or loss
of the penis (Ross, 1969). Ross (1969) discussed the functional significance of antenniformy and Barnes (1992)
reviewed the reproductive periods and condition of the penis. In cirripeds which have antenniform cirri the penis is
usually absent or much reduced. Antenniformy is not found in the Balanidae and is uncommon in the
Chthamalidae. In Chthamalus antennatus (Pope, 1965) and Tetrachthamalus oblitteratus (Newman. 1967) there is
clearly a connection between antenniformy and reproduction. Antenniformy is more common in the Tetraclitidae
e.g. Tetraclita and Newmanella hentscheli (Ross & Perreault, 1999); only Tetraclitella does not exhibit
antennilormy and it is rare in Tesseropora, although ROSS (1969) recorded antenniformy, a reduced penis and eggs
occurring together in the mantle cavity of T. pacifica (Pilsbry, 1928). Antenniform rami occur immediately before,
during, and directly after the reproductive period. ROSS (1969) suggested that individuals brooding eggs or nauplii.
having antenniform rami, and lacking or having reduced penes, assume the morphology of and act as females. The
penis remains non-functional until the beginning of the next reproductive season, whilst the antenniform ramus is
replaced by a functional feeding appendage after release of the nauplii.
In the present paper a connection between antenniformy, the reproductive state of the cirriped and the regression
of the penis is indicated for the Pachylasmatidae (Table 32). Antenniformy of cirrus II and/or cirrus III was recorded
in 1 1 species - in the pachylasmatines Eutomolasma maclaughlinae sp. nov., E. orbiculatum sp. nov.,
achylasma bacum sp. nov., P . laeviscutum sp. nov., P. ovatum sp. nov., Eurylasma pyramidale sp. nov. and
etrapachylasma arcuatum sp. nov.; in the metalasmatine Metalasma crassum sp. nov.; and in the hexelasmatines
exelasma foratum sp. nov., H. globosum sp. nov. and H. sandaracum sp. nov. Antenniformy was associated
276
D. JONES
Table 32. — Relationship between antenniformy, regression of penes and eggs.
with small numbers of large eggs in 5 species - i.e. £. maclaughlinae, T. arcuatum, M. crassum, H.foratum and
H. sandaracum. Regression of the penis was shown by M. crassum (penis equal to the length of the hasal segment
ol the pedicel of cirrus VI). Penile lengths for the remaining 4 species were as follows: subequal to length of
cirrus VI (£. maclaughlinae, M. fragile), slightly shorter than length of cirrus VI (T. arcuatum), 1/3 the length of
cirrus VI ( H . foratum) and 1/4 the length of cirrus VI (H. sandaracum). Of the remaining 6 species which showed
antenniformy, but which did not have eggs in the mantle cavities, 3 species (P. bacum, P. ovatum, H. globosum)
showed regression of the penis; penile lengths were 1/2 the length of the basal segment of the pedicel of cirrus VI
(P. ovatum), and equal to the length of the basal segment of the pedicel of cirrus VI (P. bacum, H. globosum).
The remaining 3 species (£. orbiculatum, E. pyramidale, P. laeviscutum) had larger penes; penile lengths were 1/4
the length of the pedicel of cirrus VI (£. orbiculatum), 1/2 the length of cirrus VI (P. laeviscutum) and equal to the
length of cirrus VI (£. pyramidale). One species with large eggs, Microlasma fragile sp. nov., showed neither
antenniformy nor regression of the penis (length of the penis subequal to cirrus VI). Southward & SOUTHWARD
(1958) recorded antenniformy of cirrus III in Bathylasma liirsutum (as Hexelasma hirsutum, then regarded as a
chthamalid but now as a bathylasmatine), and Newman & Ross (1971) recorded antenniformy of the rami of
cirri II and III ol the bathylasmatine Tetrachaelasma southward'!.
Source : MNHN, Paris
C1RRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
277
Although many cirriped nauplii are pelagic and planktotrophic, some can be lecithotrophic, and develop to
the cyprid stage within the mantle cavity of the parent. Animals producing lecithotrophic nauplii usually produce
small broods ol large size eggs, e.g. deep-water Scalpellum and Arcoscalpellum species (see Barnes. 1989
for review ol egg production in cirripeds). In the present study large eggs, few in number, were recorded in
7 species - the pachylasmatines Eutomolasma maclaughlinae, Microlasma fragile , Eurylasma angustum,
Tetrapachylasma arcuatum, the metalasmatine M. crassum and the hexelasmatines Hexelasma foratum and
H. sandaracum , suggesting that these species are likely to have lecithotrophic larvae. The large size of the eggs of
these species is similar to the size of those of hydrothermal vent barnacles (Table 33). Eggs which produce
planktotrophic larvae are ovoid, those which produce lecithotrophic larvae are more globular, due to an increased
amount of yolk. Lecithotrophy is advantageous for species, such as the deep-water pachylasmatids and
scalpellomorphs, and vent barnacles, where restricted dispersal in habitats with few suitable substrata for settlement
is advantageous. Abbreviated naupliar development occurs within the mantle cavity and, when released, the cyprid
remains in the vicinity of the parent group. No nauplii or cyprids were found in any of the material examined in
the present study.
TABLE 33. — Egg characters of hydrothermal vent barnacles.
Endemism and Species Diversity. — The material examined herein is both rich in numbers of species
and numbers of individuals, with an exceptional diversity of species occurring within the subfamilies
Pachylasmatinae and Hexelasmatinae. Of the new species described, 1 pachylasmatine (: Tetrapachylasma ornatum
sp. nov.) is endemic to the Marianas Islands, 8 pachylasmatines (Eutomolasma orbiculatum, Microlasma fragile
sp. nov., Pachylasma bacurn sp. nov., P. ovatum sp. nov., Eurylasma angustum sp. nov., £. ferulum sp. nov.,
E pyramidale sp. nov., Tetrapachylasma arcuatum sp. nov.), 1 metalasmatine (Metalasma crassum sp. nov.) and
5 hexelasmatines ( Hexelasma aureolum sp. nov., H.flavidum sp. nov., H. foratum sp. nov., H. globosum
sp. nov., H. persicum sp. nov.) are considered endemic to the region of Vanuatu/New Caledonia. Two
pachylasmatines (Eutomolasma maclaughlinae sp. nov., Pachylasma laeviscutum sp. nov.) and I hexelasmatine
( Hexelasma sandaracum sp. nov.) occur in a broader area, extending from the Indo-Australian Plate to the Futuna
and Wallis Islands on the Pacific Plate. The number of new pachylasmatines described in the present study
represents 46% of the known species and that of the new hexelasmatines 40%.
The most diverse chionelasmatoid and pachylasmatoid fauna occurs at the Loyalty Islands where 15 species
have been collected (1 chionelasmatoid, 1 eolasmatine, 6 pachylasmatines, 1 metalasmatine, 6 hexelasmatines).
At the Norfolk Ridge, 11 species have been collected (1 chionelasmatoid, 6 pachylasmatines and
hexelasmatines), and nine species are known from both Vanuatu and New Caledonia (1 eolasmatine.
_ pachylasmatines, 1 metalasmatine, 1 bathylasmatine, 2 hexalasmatines and 1 chionelasmatid,
- pachylasmatines, 5 hexelasmatines respectively). Three species are known from the Chesterfield Islands
U pachylasmatines, 1 hexelasmatine). On the Pacific Plate, 2 species were collected at Combe Bank
pac y asmatine, 1 hexelasmatine). A single pachylasmatine species was collected at both Futuna Island and
aterwich Bank and a single hexelasmatine at the Wallis Islands, at Field Bank and at Tuscarora Bank.
278
D. JONES
The present study has confirmed that 2 relictual species, the chionelasmalid Chionelasmus darwini and the
eolasmatine Waikalasma boucheti occur in the waters of New Caledonia and ol Vanuatu, respectively. However,
although C. darwini is represented by many specimens, collected at various localities throughout the New
Caledonian region, IV. boucheti is known only by an incomplete specimen from the Loyalty Islands (present
report) as well as by 2 specimens from the waters of Vanuatu (Buckeridge, 1996). Similarly, only
I bathylasmatine - B. alearum , represented by 2 specimens from the waters of Vanuatu - has been collected.
Sampling by the French deep-sea expeditions has been from comparable depths in the waters of Vanuatu,
New Caledonia and the Wallis and Futuna Islands. Whether sampling has been from comparable habitats,
however, is a more difficult question to answer, so the possibility exists that species which occur in specialised
habitats either have not been collected, or have been collected infrequently.
The cruises BiOCAL and MUSORSTOM 6 gathered a variety of faunas from the Loyalty Islands, of which many
arc relictual species, e.g. a group of crinoids that were assumed to have died out in the Jurassic - the pedunculate
crinoid Gymnocrinus and a small black crinoid Holopus reported previously from the Caribbean but not from the
Pacific Ocean (AMEZIANE-COMINARDI et al ., 1987). The presence of the relictual species Chionelasmus darwini
and Waikalasma boucheti at the Loyalty Islands further supports the hypothesis that this bathyal fauna is a vestige
of marine populations of the ancient Tethys Sea, and that the south-west Pacific is a relictual area (Newman,
1991).
An abundance and diversity of benthic material has also been collected at the Norfolk Ridge (Chalcal 2,
Smib 2-5, Bathus 3, 4). Several groups, such as the calcified sponges of the Sphinctozoa group, the pedunculate
crinoid Gymnocrinus, molluscs of the family Pleurotomariidae, as well as hydroids and echinoderms, show a clear
relationship with the Mesozoic fauna of the ancient Tethys (VaCELET. 1977, 1978; BOURSEAU et al., 1987;
Richer DE Forges, 1990). These living fossils are located on the eastern rim of the ancient continent of
Gondwanaland. Abundant zoological collections have also been made from New Caledonia (MUSORSTOM 4,
Beryx 2, Bathus 2), especially sponges, molluscs, stylasters and echinoderms. Many fossilised shark teeth,
especially those of the giant species Procarcharodon megalodon Agassiz. 1843. have also been collected (Richer
de Forges, 1990).
In contrast to the diverse chionelasmatoid and pachylasmatoid fauna occurring in the New Caledonian area, a
pachylasmatoid fauna low in diversity and in numbers of individuals and no chionelasmatoids have been found at
the Wallis and Futuna Islands on the Pacific plate. The pachylasmatoid species are not new, but are known also
from New Caledonian waters. In general, the deep sea faunas from these waters are diverse but quite poor compared
to those of New Caledonia, with species described from New Caledonian waters being rediscovered on the Pacific
plate (e.g. Sphinctozoa, Gymnocrinus, Amalda) (Richer deFORGES & Menou. 1993).
ACKNOWLEDGMENTS
The author is deeply indebted to A. CROSNIER, IRD (ex ORSTOM) and Museum national d'Histoire naturelle,
Paris, and B. Richer de FORGES, IRD. Noumea, for the opportunity to study the cirripcds from Vanuatu,
New Caledonia and the Wallis and Futuna Islands. The study was carried out partly at the Museum national
d'Histoire naturelle, Paris, with the support of grants from ORSTOM (August-October, 1994) and the Museum
national d'Histoire naturelle (September-October, 1997), and partly at the Western Australian Museum, Perth.
I also thank D. Defaye, Laboratoire de Zoologie (Arthropodes) Museum national d'Histoire naturelle, Paris, for
her assistance with this project. I am indebted to my colleagues W. NEWMAN and A. ROSS, Scripps Institution of
Oceanography, La Jolla, California, J. BUCKERIDGE, Auckland Institute of Technology, Auckland, and
P. McLaughlin, for critically reading drafts of the manuscript and for valuable discussions.
Thanks are also extended to Clay BRYCE, Western Australian Museum, and P. LOZOUET, Museum national
d'Histoire naturelle, Paris, for photography of specimens, to Simon JONES, Lisa KING, Jane MacRae and
Nadine GUTHRIE who inked some of my drawings, to Anne Nevin who typed the tables. At the Museum national
d'Histoire naturelle, J.-F. DEJOUANNET prepared ihe plates and at the IRD Center of Noumea. M. TORTELIER
prepared the distribution maps.
Source : MNHN, Paris
CIRRIPEDIA THORACICA: CHIONELASMATOIDEA AND PACHYLASMATOIDEA
279
I also wish to thank colleagues in the following institutions who kindly lent type specimens for comparisons
with the new material described in this study: P. Berents, The Australian Museum. Sydney; G. Boxshall and
A. Morgan, The Natural History Museum, London; N. BRUCE and P. JEPPESEN, Zoologisk Museum. University
of Copenhagen; S. O'Shea, National Institute of Water and Atmospheric Research, Wellington, New Zealand;
D. PLATVOET. Zoologisch Museum, Amsterdam; K.J. REED, Smithsonian Institution, Washington;
R. Webber, Museum of New Zealand, Wellington.
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rs DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CA
Crustacea Decapoda: Review of the genera and species
of the family Polychelidae Wood-Mason, 1874
Bella S. GAUL
Israel Oceanographic & Limnological Research
Nalional Institute of Oceanography
P.O.B. 8030, Haifa 31080, Israel
ABSTRACT
The polychelids are large, uncommon, primitive decapods that inhabit the depths of the world oceans down to
5000 m, between latitudes 50°N and 55°S. A study of major deep-sea collections led to a revision of the family. All
genera and species are redescribed and extended synonymies given. Two new genera are established: Cardus, for
Polycheles crucifer (Thomson, 1873) and Homeryon, for Polycheles asper Rathbun, 1906 and a new species. H.
armarium. The genus Pentacheles Bate, 1878, is revived to include polychelids in which the epipod on third maxilliped is
longer than the ischium: P. gibbus Alcock, 1894, P. laevis Bate. 1878, P. obscurus Bate, 1878, P. synderi (Rathbun.
1906) and P . validus A. Milne Edwards, 1880. Stereomastis Bate. 1888 is considered a synonym of Polycheles Heller.
1862. Willemoesia Grote, 1873 is retained with but four species: W. forceps A. Milne Edwards, 1880. VP. inornate Faxon.
1893. W. leptodactyla (Willemoes-Suhm, 1875). and W. pacifica Sund, 1920. In all, thirty-two species are recognized,
including six new species. The bathymetric and geographic ranges are amended and discussed. A key to the genera and
species of the family is provided.
RESUME
Crustacea Decapoda
1 874.
Revue des genres et des especes de la famille Polychelidae Wood-Mason,
Les Polychelidae sont des crustaces decapodes primitifs de grande tai lie et peu communs, qui vivent dans les
prolondeurs des oceans jusqu'it 5000 m et entre les latitudes 50°N et 55°S. Une etude des principales collections faites en
eau profonde a conduit h une revision de la famille. Tous les genres et especes sont redecrits et leurs synonymies
completes sont indiquees. Deux nouveaux genres sont exits: Cardus pour Polycheles crucifer (Thompson. 1873) et
Homeryon pour Polycheles asper Rathbun, 1906; une espece nouvelle, Homerion armarium sp. now, est decrite. Le genre
entacheles Bate. 1878, est ressuscit6 pour accueillir les especes dont 1'epipode des troisiemes maxillipedes est plus long
187a A/L1L" B" 2°00' — Crustacea Decapoda: Review of the genera and species of the family Polychelidae Wood-Mason.
I>w4. In: A. Crosnier (ed.), Resultats des Campagnes MUSORSTOM, Volume 21. Memoires du Museum national d'Histoire
naturelle. 184: 285-387. Paris ISBN-2-85653-526-7.
286
B. GALIL
que I'ischion : P. gibbus Alcock. 1894, P. laevis Bale, 1878, P. obscurus Bate, 1878. P. synderi (Rathbun, 1906) el
P. validus A. Milne Edwards, 1880. Stereomastis Bale, 1888, est considere comme etant synonyme de Polycheles Heller.
1862 Willemoesia Grote, 1873, est conserve mais avec quatre especes seulement: W. forceps A. Milne Edwards. 1880,
W inornata Faxon, 1893, W. leptodactyla (Willemoes-Suhm. 1875). cl IV. pacifica Sund, 1920. Au total 32 especes
sont reconnues, dont six sont nouvellcs. Les repartitions bathymetriques el geographiques sonl revues el discutees. Une
cle des genres et especes est proposee.
INTRODUCTION
The depths of the world ocean are home to the eyeless, claw-footed polychelids, "living fossils", kin to the
long vanished eryonids, tracing their ancestry back as far as the Mesozoic.
Bate (1888: 100) described his excitement following the finding of polychelids by the " Challenger "
Expedition: "The great depth from which it was dredged, a depth that was previously believed to be barren, if not of
all life, certainly of animals so high in the scale of existence, the apparent absence of the power of vision, and the
relationship of the animal to forms of Crustacea that were supposed to have been extinct since the period of the
Upper White Jura of Bavaria, gave a considerable degree of interest to the discovery".
Subsequent deep-sea expeditions added new species and extended the range ol known ones. However, many
species were known from few, damaged, or juvenile specimens, confusing workers searching lor valid taxonomic
characters. BATE (1888: 141) recognized the problem: "It is highly probable that many of the animals that we
determine as specifically distinct, because they arc found in widely separated localities, and exhibit some greater or
less deviation from each other, would cease to be considered such if they lived side by side, and there can. I think,
be little doubt that many of our museum specimens are not really species". A century later, it has remained
unsolved: "The taxonomy of the Polychelidae, especially of the generic level, is still very unsettled" (HOLTHUIS,
1991: 88).
The revision comprises adult forms only as "The larval forms of the Polychelidae are incompletely known"
(Firth & Pequegnat, 1971: 14). A polychelid larva was collected by Willemoes-Suhm early on the
"Challenger" Expedition, and labelled by him "Zoea of Brachyura", but the specimen was recorded by Bate (1882)
as a new genus, Eryoneicus. Recognized by Wood-Mason & Alcock (1891: 199) as "an immature form of
some species of Pentacheles", "the question of whether the peculiar forms known as Eryonicus are larvae of the
crayfish-like deep-sea crustaceans Polycheles" (SUND, 1915: 372) was long subject of debate - DE SAINT LAURENT
(1979) still argued they are adults secondarily adapted to the bathypelagic existence. Though Sund (1915) tried to
match larval forms with adult species using carapacial spine formula, only a few of the nearly 40 larval forms
known (GORDON, 1960), are firmly correlated with the adult.
Much of the material examined in this study was collected by the following cruises: MUSORSTOM 1. Philippine
Islands, April 1976; MUSORSTOM 2, Philippine Islands, November-December 1980: MUSORSTOM 3, Philippine
Islands, May-June 1985; MUSORSTOM 4, New Caledonia, September-October 1985; MUSORSTOM 5. Chesterfiled
Islands, October 1986; MUSORSTOM 6, Loyalty Islands, February 1989; MUSORSTOM 7, Wallis and Futuna
Islands, May 1992; MUSORSTOM 8, Vanuatu Archipelago, September-October 1994; Biocal, New Caledonia and
Loyalty Islands, August-September 1985; Biogeocal, New Caledonia and Loyalty Islands, April-May 1987;
Volsmar, Matthew and Hunter Islands, May-June 1989; Karubar, Indonesia, October-November 1991;
Beryx 2, Norfolk Ridge, Loyalty Islands, October 1991. Bathus 1-4, New Caledonia. March 1993-1994;
Halipro 1, New Caledonia. March 1994; (Forest, 1981, 1985, 1989; Richer DEFORCES, 1990, 1993; Richer
de Forges & Menou, 1993; Richer de Forges & Chevillon, 1996; Richer de Forges et al., 1996;
CROSNIER et al., 1997).
A study of the extensive collections of the Museum national d'Histoire naturelle. Paris (MNHN), National
Natuurhistorische Museum, Leiden (formerly Rijksmuseum van Natuurlijke History) (RMNH), National Museum
of Natural History, Smithsonian Institution, Washington (USNM), The Natural History Museum, London
(NHM), together with material made available by the Australian Museum, Sydney (AM), American Museum of
Natural History, New York (AMNH), Indian Museum, Calcutta (IM), Institut royal des Sciences naturelles de
Belgique, Bruxelles (IRScNB), Kitakyushu Museum & Institute of Natural History, Kitakyushu (KMNH), Natural
Source : MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
287
internal angle
of
orbital sinus
anterolateral angle
of
carapace
orbital sinus
gastro-orbital
region
cervical incision
postcervical
incision
branchial groove -7
first
pereiopod (PI)
rostrum
posterior margin
of
carapace
basal antennular segment
antennular peduncle
jxvl antennal scaphocerite
uropodal exopod
external angle
of
orbital sinus
median postrostral
carina
postorbital
carina
anterior branch
of
cervical carina
cervical groove
branchial carina
median tergal carina
median postcervical
carina
pleuron
submedian tergal groove
basal tubercle
uropodal endopod
telson
Fig. I. — Diagrammatic polychelid.
Source
288
B. GAL1L
History Museum of Los Angeles (LAM), Museum of Comparative Zoology, Harvard University, Cambridge
(MCZ)' Museo Zoologico, Universita degli studi di Firenze (MF). Naturhistorisches Museum, Wien (NHMW),
Nagasaki University Museum, Nagasaki (NUM), National Institute of Water & Atmospheric Research. New
Zealand (NIWA). National Fisheries University, Shimonoseki (NFU), South African Museum, Cape Town
(SAM). Scnckenberg Museum, Frankfurt (SMF), National Collections, Tel Aviv University (TAU), Tasmanian
Museum Hobart (TM). Western Australian Museum, Perth (WAM), Zoologisk Museum, Copenhagen (ZMC),
and Zoological Museum of Moscow University, Moscow (ZMMU) has enabled the reexamination of nearly all
type specimens and much of the published material.
The present study divides the Polychelidae into five genera, two of which are new, and describes six new
species. Descriptive and distributional information is presented as well as detailed references to literaiure. All laxa.
but Pentacheles gibbus, have been photographed, and a key is presented for their identification.
In the lists of Material examined, the names of the cruises are in capital letters. The names ol the vessels are in
italic letters and quoted.
In the chapter Distribution, when the depth ranges are given without attribution, they are taken trom the
material examined in the present work, all other records are cited by author.
TERMINOLOGY
The terminology used to describe the species of the family Polychelidae is illustrated on the figures 1-3.
Measurements refer to carapace length from the tip of the rostrum to the middle of the posterior border of the
carapace and are given in millimeters.
FIG. 2. — a-c, frontal margin of carapace: a. rostrum bifid; b, rostral spines separate; c. frontal subterminal tooth
prominent, longer than separate rostral spines. — d, ventral view showing the renal process ol the antenna (1) and
the eyestalk (2); e. dorsal view showing the aperture of the renal process.
Source : MNHN , Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
289
Source : MNHN , Paris
290
B. GALIL
Family POLY CHELID AE Wood-Mason, 1 874
Polychelidae Wood-Mason, 1874: 180: 1875: 132. — Carus, 1885: 485. — Balss, 1957: 1564. — Zariquiey Alvarez,
1968: 208. — Glaessner. 1969: 470. — FIRTH & PEQUEGNAT, 1971: 5. — Wenner. 1979: 435. — Abele &
Felgenhauer. 1982: 306. — Riedl. 1983: 481. — Squires, 1991: 354.
Eryonidae - Boas, 1880: 157 (pan). — Bate, 1888: 100 (part). — Ortmann, 1896: 428 (pari), — Young, 1900: 439. —
ALCOCK. 1901b: 164 (pan). — Stebbing, 1902: 35 (pan); 1910: 377 (part); 1917: 27 (part). — SELBIE, 1914: 8. —
De Man, 1916: 1 (part). — Bouvier, 1917: 26 (part). — PESTA, 1918: 161 (part). — Stephensen. 1923: 65 (part). —
Balss. 1925: 5 (part). — Boone, 1927: 90 (part). — Rodriguez, 1980: 194 (part).
Eryontidae Smith, 1880a: 345 (part). — Stebbing. 1893: 199 (part). — Schmitt, 1921: 105 (part),
Polychelida Scholtz & Richter, 1995: 296.
Diagnosis. — Carapace dorso-ventrally flattened, subrectangular or ovate. Anterolateral angle of carapace
produced, spiniform. Lateral margins well defined, spinose; cervical and postcervical incisions dividing margin into
three parts. One or two rostral spines. Orbital sinus absent, reduced, or well defined. Eyestalk present, lacking
cornea.
Cervical groove marked, arcuate, laterally bifurcate. Median postrostral and postcervical carinae well defined,
unarmed, granulate, or spinose. Postorbital and branchial carinae obsolescent or well defined, unarmed, granulate or
spinose.
Abdomen narrow or broad, laterally or dorsoventrally depressed. First abdominal tcrgite narrow, pleura
fused, abbreviated; latching into posterolateral margins of carapace. Abdominal tergites smooth or sculptured,
medially carinate or binodal. Abdominal pleurae diminishing in size posteriorly, smooth or sculptured.
Abdominal sternites with or without median knob. Telson lanceolate, bearing two submedian carinae.
Uropods lacking transversal suture; uropodal exopod lamellar, endopod bearing denticle or lappet basally on
interior margin.
Basal antennular segment flattened, mesial margin granulate or spinose; anterolateral angle with 1-2 spines or
without. Outer antennular flagellum short, inner flagellum as long as antennal flagellum. Renal process well
developed, tubular, fitting into hollow on underside of basal antennular segment. Antennal scaphocerite short,
lingulate or lanceolate; antennal peduncle five-jointed, first segment not fused to epistome.
Buccal opening subquadrate. Mandible concave, cutting edge with triangular teeth of uneven size; mandibular
palp biarticulate. Maxillula with two incurved lobes. Maxilla with two slender lobes; large scaphognathitc. First
maxilliped endopod slender; exopodal lobe membraneous, reniform, extending further back than scaphognathitc.
exopod anteriorly divided into two lobes enclosing efferent passage. Second and third maxillipeds slender, pediform;
epipod on basal segment of third maxilliped rudimentary or long.
Pereiopods 1-4 chelate. P5 either simple or chelate in both sexes, or chelate in female, simple or subchelate in
male. First pereiopod larger, more robust than P2-5; pereiopods successively shorter posteriorly. Fingers of first
chelipeds crossing, their inner margin furnished with row of overlapping cornute platelets, forming serrate edge.
First male pleopod spatuliform, distal margin folded internally. Pleopods 2-6 biramose, with rod-like appendix
interna, second pair in male with appendix masculina.
Remarks. — The Polychelidae is the sole extant family of the superfamily Eryonoidea, a group represented in
fossil records from the mid-Triassic period (GLAESSNER, 1969).
H. Milne Edwards (1837) described "Tribu des Eryons", and de Haan (1841) erected the family Eryonidae
for the fossil genus E/yon. Heller (1863) placed Polycheles typhlops in the family Astacidae though he noted
its affinity with the fossil Eryon. Only after additional material was collected, did WOOD-MASON (1875: 132)
write that those specimens “ cannot be placed in no existing family of crustaceans, recent or fossil, except
perhaps the Eryonidae. the structural characters of which are too incompletely known at present to admit of
their being included in it '. and established the family Polychelidae. However, most authors continued placing the
extant genera within the Eryonidae. The Eryonidae are now considered to be exclusivelv fossil (Balss, 1957;
Glaessner, 1969).
Source : MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
29 1
Key to the genera and species of the family POLYCHELIDAE
1. Frontal margin of carapace without orbital sinus . Willemoesia 4
— Frontal margin of carapace with orbital sinus . 2
2. Carapace ovate; postorbital and postcervical carinae swollen; fifth pereiopodal dactyl
simple in both sexes . CARDUS gen. nov. (including one species: C. crucifer)
— Carapace ovate or subrectangular; postorbital and postcervical carinae well-defined or
obsolescent, never swollen; dactyl of P5 simple or subchclate in male, chelate in female...
. .
3. Epipod ol third maxilliped longer than ischium; basal antennular segment proximally
quadrate, lamellar . PENTACHELES 7
— Epipod of third maxilliped half as long as ischium, basal antennular segment squat
. . . Homeryon gen. nov. 29
— Epipod of third maxilliped rudimentary; basal antennular segment proximally rounded .
. POLYCHELES 1 1
4. Sixth abdominal tergite sculptured . c
— Sixth abdominal tergite nearly smooth . g
5. Lateral margins of carapace posterior to postcervical incision with less than 10 spines;
upper margin of chela with 2 files of spines . W. inornata
— Lateral margins of carapace posterior to postcervical incision with more than 20 spines;
upper margin of chela with several files of spines . W. leptodactyla
6. Lateral margins of carapace anterior to postcervical incision with 15-19 spines; abdominal
tergites 2-5 with deep submedian grooves; tip of telson rounded . W. forceps
Lateral margins of carapace anterior to postcervical incision with 6-10 spines; abdominal
tergites smooth, lacking deep submedian grooves; tip of telson acuminate .. W. pacifica
7. Internal angle of orbital notch unarmed . g
Internal angle of orbital notch spinose . 9
8. Carapace depressed; abdominal terga, pleura nearly smooth . P. obscurus
Carapace convex; abdominal terga, pleura set with conic tubercles . P. gib bus
9. Lateral margins of carapace posterior to postcervical incision with 14-15 spines
. . . . . . P. laevis
- Lateral margins of carapace posterior to postcervical incision with more than 25 spines ...
. 10
10. Carapace subrectangular, densely spinulose; external orbital margin pectinate
. . P. snyderi
Larapace ovate, sparsely spinose; external orbital margin with 0-3 spines .... P. validus
11. One rostral spine . 12
— Two rostral spines . . j 3
12. Anterior pleuron ovate; inferior surface of uropodal exopod bicarinate . P. ty phi ops
Anterior pleuron rounded; inferior surface of uropodal exopod tricarinate . P. perarmatus
13. Antero-external margin of basal antennular segment unispinose . l 4
— Antero-external margin of basal antennular segment bispinose . ZZ l 5
Source : MNHN, Paris
292
B. GALLL
14. Carina of fifth abdominal segment with antrorse spine; carapace between branchial and
median poslcervical carinae spinose . P. phosphorus
— Carina of fifth abdominal segment unarmed; carapace between branchial and median post-
cervical carinae unarmed . P. politus sp. nov.
15. Frontal submarginal tooth prominent, longer than separate rostral spines . 16
— Frontal submarginal tooth shorter than rostrum, or rostrum bifid . 17
16. Gastro-orbital region bispinose; median postrostral and postcervical carinae irregularly
granulate; second pleuron ovate; upper margin of first chela prominently spinulose .
. P. baccatus
— Gastro-orbital region quadrispinose; median postrostral and postcervical carinae set with
antrorse tubercles; second pleuron anteriorly triangulate; upper margin of first chela
granulose . P. coccifer sp. nov.
17. Internal angle of orbital sinus unarmed . 18
— Internal angle of orbital sinus spinose . 19
18. Antrorse spine on fifth abdominal tergite large, overhanging anterior margin of fourth
tergite . P. ceratus
— Antrorse spine on fifth abdominal tergite not as above . P. helleri
19. Median carina on abdominal tergites 2-5 bicuspid, anteriorly spinose; gastro-orbital region
quadrispinose anteriorly, cluster of 5-6 spines posteriorly . P. suit mi
— Median carina on abdominal tergites 2-5 not cuspidate; spinulation of gastro-orbital region
otherwise . 2 0
20. Frontal margin with several spinules on either side of rostral spines . P. enthrix
— Frontal margin with single spine on internal angle of orbital sinus . 2 1
2 1 . Cheliped carpus one third as long as merus . P. tanneri
— Cheliped carpus half as long as merus . 2 2
22. Branchial groove with a distal spine . 2 3
— Branchial groove unarmed . 2 5
23. Carapace between branchial and median postcervical carinae unarmed posteriorly; median
carina on fifth abdominal tergite anteriorly bulbous . P. pacificus
— Carapace between branchial and median postcervical carinae posteriorly spinose; median
carina on third abdominal tergite bearing long antrorse spine; fourth and fifth abdominal
tergites unarmed . p. trispinosus
— Carapace between branchial and median postcervical carinae posteriorly spinose; median
carina on fifth abdominal tergite bearing antrorse spine . 24
24. Antrorse spine on third abdominal tergite largest; lyre-shaped carina on sixth abdominal
tergite prominently denticulate; basal tubercle on telson salient . P. nanus
— Antrorse spine on fifth abdominal tergite largest; lyre-shaped carina on sixth abdominal
tergite rounded; basal tubercle on telson blunt . P. evexus sp. nov.
25. Median carina of fifth abdominal tergite with antrorse spine . 2 6
— Median carina of fifth abdominal tergite without antrorse spine . 2 7
26. Sixth abdominal tergite bearing lyre-shaped, denticulate carina mesially; lateral margins of
carapace posterior to postcervical incision with 8-10 spines . P. talismani
Sixth abdominal tergite bearing parallel rounded carinae, confluent anteriorly and
posteriorly; lateral margins of carapace posterior to postcervical incision with 6-7 spines . .
. . . P. sculptus
Source : MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
293
27. Posterior margin of cervical groove with single antrorse spine midway between median
postcervical and branchial carinae; frontal submarginal tooth prominent, visible in dorsal
view; second pleuron anteriorly triangulate . P. aculeatus sp. nov.
— Posterior margin of cervical groove with 3 or 4 antrorse spines midway between median
postcervical and branchial carinae; frontal submarginal tooth small; second pleuron ovate..
. 28
28. Lateral margins of carapace posterior to postcervical incision with 7 spines; submedian
grooves on abdominal tergites marked; lyre-shaped carina on sixth tergite prominent .
. P. auriculatus
— Lateral margins of carapace posterior to postcervical incision with 10-14 spines;
submedian grooves on abdominal terga obsolescent; lyre-shaped carina on sixth tergite
obsolescent . P. surdus sp. nov.
29. Lateral margins of carapace posterior to postcervical incision cristate, serrulate; abdominal
carina blunt; second pleuron cordiform; uropods smooth . H. asper
— Lateral margins of carapace posterior to postcervical incision rounded, bearing rows of
antrorse spinules; abdominal carina bicuspid; second pleuron reniform; uropods granulate .
. H. armarium sp. nov.
Genus CARDUS nov.
Type Species. — Deidamia crucifer Thomson, 1873. Gender masculine.
DIAGNOSIS. — Carapace dorsoventrally flattened, ovate. Anterolateral angle of carapace produced, spiniform.
Lateral margins well defined, spinose; cervical and postcervical incisions markedly dividing margin into three parts.
Single rostral spine. Orbital sinus reduced, vermicular process of eyestalk filling orbit.
Cervical groove marked, arcuate, laterally bifurcate. Median postrostral and postcervical carinae well defined,
swollen, granulate. Postorbital and branchial carinae swollen, well defined, granulate.
Abdomen narrow, laterally depressed. First abdominal tergite narrow, pleuron abbreviated. Abdominal tergites
2-5 sculptured, medially binodal; sixth tergite medially bitubcrculatc. Abdominal pleura bearing granulate basal
protuberance; second and third pleura lingulate, fourth pleuron triangulate, fifth acuminate. Abdominal slernites
with median knob. Telson lanceolate, with a basal median tubercle and two submedian carinae. Uropodal exopod
lamellar, oar-shaped; cndopod lanceolate with a basal denticle on interior margin.
Mesial margin of basal antennular segment spinose, upcurved, anteriorly produced, not reaching base of
peduncle; anterolateral angle spinose. Antero-internal angle of second, third antennular segments prominently
spiniform. Outer antennulary flagellum short, inner flagellum as long as antennal flagellum. Renal process well
developed, tubular, fitting into hollow on underside of basal antennular segment. Antennal scaphocerite short,
lanceolate, extending beyond second peduncular segment.
Buccal opening subquadrate. Mandible concave, cutting edge with triangular teeth of uneven size; mandibular
palp biarliculate. Maxillula with two incurved lobes, spiniform setae distally on anterior lobe. Maxilla with two
slender lobes, internal lobe half as long as lateral; large scaphognathite. First maxilliped endopod slender, exopodal
lobe membraneous, reniform, extending further back than scaphognathite; exopod anteriorly divided into two lobes
enclosing efferent passage. Second and third maxillipeds, slender, pediform; basal segment of third maxilliped
bearing long epipod; dactyl as long as both preceding segments.
Pereiopods 1-4 chelate, P5 simple in both sexes. PI larger, more robust than P2-5; pereiopods successively
shorter posteriorly. Fingers of first, second chelipeds crossing, their inner margin furnished with row of
overlapping cornute platlets, forming serrate edge. Fingers of third, fourth pereiopodal chelae slightly upcurved,
their inner margins simple. First male pleopod spatuliform, spinulose swelling distally on inner margin, spinules
294
B. GALIL
distally on mesial margin, distal margin folded internally. Pleopods 2-6 biramose, with rod-like appendix interna,
second pair in male with appendix maseulina.
ETYMOLOGY. — From the Latin, carduus (= thistle), referring to the spinose, thistle-like carapace.
Remarks. — Cardus is distinguished from the other polychelid genera in having the fifth pereiopodal dactyl
simple in both sexes. It is further distinguished from Polyclieles by the antero-internal process ol the basal
antennular segment not reaching the base of the peduncle; the basal segment of the third maxilliped bearing a long
epipod; the fingers of the second and third pereiopods slightly upeurved, and their internal margin simple.
WlLLEMOES-SUHM (1875c: xxxiii) correctly distinguished crucifer: "hat nur Scheeren an vieren und wird
vielleicht mil dei Zeit ein eigenes Genus bilden mussen".
Cardus crucifer (Thomson, 1873)
Fig. 4
Deidamia crucifer Thomson 1873; 247, fig. 1. — Wood-Mason, 1874: 180.
Willemoesia crucifer - GROTE, 1873: 485. — HUMBERT, 1874: 130. - WlLLEMOES-SUHM. 1875b: 577: 1875c: xxxiii. —
Thomson. 1877: 256. fig. 59. — Giglioli, 1912: 160.
Willemoesia crucifera - Willemoes-Suhm, 1875a: 52. pi. 12 fig. 10. pi. 33 figs 10-11. — Norman. 1879: 174. —
Pagenstecher. 1879: 37.
Polxcheles crucifer - Wood-Mason, 1875: 132. — Bate, 1878a: 277. pi. 13 ligs 6-8; 1878b: 484; 1878c: 563. — Boas.
1882: 352, fig. 12. — Marshall, 1888: 267. — Faxon, 1896: 155. — Bouvier. 1905a: 480; 1905b: 644; 1905c: 3;
1905d: 2; 1917: 36: 1925: 428. figs 5-8, pi. 4 fig. 4. pi. 11 figs 7-14. — De Man. 1916: 5 (list). — Schmitt, 1935:
171, fig. 35. — Bernard. 1953: 86. — Chace, 1960: 30. fig. 9b. — Firth & Pequegnat. 1971: 42. — Pequegnat ei
ai, 1971: 4. — Beaubrun, 1979: 33. fig. 19. — Gonzalez, 1995: 124. — Dawson, 1997: 8.
Pentacheles crucifer - FlLHOL, 1884: 231.
Polxcheles crucifera - BATE, 1888: 127, fig. 31. pi. 13. — YOUNG. 1900: 440. — RICHARD, 1905: 3. — Turkay. 1976:
27, fig. 5.
Polychelles crucifer - RODRIGUEZ, 1980: 194 [erroneous spelling].
Material EXAMINED. — Ibero-Morrocan Gulf. Balgim: stn CP 10, 36°45’N. 9°32'W, 1582-1602 m.
29.05.1984: 1 d 15.4 mm (MNHN-Pa 1675). — Stn CP 69, 35°H'N. 7°50'W. 1978-2077 m, 5.06.1984: 2 juvs (MNHN-
Pa 1676). — Stn CP 99, 34°28’N, 7°43'W, 1848-1892 m, 9.06.1984: 3 juvs (MNHN-Pa 1677).
Canary Is. Cancar 2: stn 2.131, 27°40'N, I8°10'W, 1200-1800 m, 8.09.1977: 3 juvs (RMNH).
Cancap 4: stn 4.056, 28°47’N. 13°22'W, 1306-1345 m, 18.05.1980: 1 3 17.0 mm (RMNH). — Stn 4.057, 28°27'N.
13°24'W, 1300-1335 m, 19.05.1980: 1 ov. 9 22.9 mm (RMNH).
United States (East coast). "Gerda" (id. L.B. Holthuis): stn 223, Florida Sir.. 24°18'N. 80°29'W. 897-915 m,
23.01.1964: 1 juv. (USNM 136603). — Stn 296, Florida Str., 25°36'N. 79°23’W, 715 m. 5.04.1964: 1 9 21.3 mm
(USNM 136604). — Stn 266, Florida Str., 24°12'N, 81°I7'W, 679-709 m, 15.09.1964: 1 9 22.6 mm (RMNH 22396). -
Stn 374, Florida Str., 23°50'N, 81°37'W, 1208-1241 m, 17.09.1964: 2 9 20.5, 21.4 mm; 1 juv. (USNM 136605).
Bahama Is. "Albatross": stn 2629, 23°48'07"N, 75°10'07"W. 2104 m, 8.03.1886. id. M.J. RaTHBUN: 2 juvs
(USNM 11422). — Stn 2654, 27°57'05"N, 77°27'05"W, 1188 m. 2.05.1888, id. M.J. RaTHBUN: 2 juvs (USNM 11460).
"Iselin": stn 40, 23°46'N, 76°58'W, 1317 m, 22.02.1972, id. F.M. Bayer: Id 17.6 mm; 1 juv. (USNM 154313). —
Stn 51, 23°37'N, 76°49'W. 1290 m, 25.02.1972: 1 9 18.8 mm: 2 juvs (RMNH). — Stn 156, 23°43'N. 76°50’N, 1334 m,
5.02.1974: 1 3 14.8 mm; 2 9 18.3. 25.6 mm (RMNH). — Stn 163, 23°32.5'N, 77°09.9'W, 1342 m, 6.02.1974: 1 9
16.1 mm (RMNH). — Stn 301, 23°51.95'N, 76°52'W, 1304 m. 3.04.1975: I d 16.9 mm; 2 9 21.5. 25.7 mm; 1 juv.
(RMNH). — Stn 302, 24°H'N, 77°23.2'W. 1452 m, 3.04.1975: 1 juv. (RMNH). — Stn 316, 24°24'N. 76°26.75'W.
1567 m. 7.04.1975: 2 9 15.8, 19.5 mm (RMNH). — Stn 321, 23°39.25'N. 76°46.24'W. 1328 m, 8.04.1975. id.
F.M. Bayer: 2 9 19.3, 26.6 mm; 1 juv. (USNM 154320). — Stn 322, 23°40.5'N. 77°03.5'W. 1362 m. 8.04.1975: 1 9
19.3 mm (RMNH).
"Pillsbury": stn 634, 23°33’N, 82°47'W. 1638-1757 m. 25.03.1968: 5 9 13.3-24.8 mm (RMNH).
West Indies. "Challenger": stn 23, Leeward Is, Sombrero 1.. 18°24'N. 63°28'W, 810 m, 15.03.1873: 1 d 19.0 mm
(NHM 1888.22).
Caribbean Sea. "Albatross": stn 2117, Aves Ridee, 15°24'40"N, 63°3r30"W. 1229 m. 27.01.1884: I d
15.0 mm; 1 9 20.3 mm. 2 juvs (USNM).
"Oregon": stn 1907, 12°25'N, 82°23'W, 720-765 m, 11.09.1957: 1 9 27.5 mm (USNM 251942). — Sin 3561,
16°35'N. 80°04'W, 720 m, 18.05.1962: 1 d 16.6 mm; 2 9 21.8, 24.5 mm (USNM 244083). — Stn 4566, 23°05'N.
Source MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
295
86°09'W, 1260 m, 6.12.1963: 1 2 22.1 mm; 2 juvs (USNM 273140). — Stn 5753. 29°29'N. 79°53'W. 675 m.
19.11.1965: 1 2 26.9 mm (USNM 273140). — Sin 5930. I5°38'N. 61°07'W. 796 m. 5.03.1966: I 2 22.7 mm (USNM
273140). — Sin 6697. 17°47'N, 63°09'W. 792-828 m, 18.05.1967: 2 2 27.0 28.0 mm (USNM 273140). — Sin 6703.
1 6°53'N, 6I°53'W, 738-828 m. 21.05.1967: 1 2 29.6 mm (USNM 273140). — Stn 6705. I7°I4,N, 63°01'W. 693-
765 m. 22.05.1967: 2 ov. 2 28.5. 29.7 mm (USNM 273140).
"Oregon II": sin 10825, Martinique. I5°42'N. 61°08'W. 630 m. 1.12.1969: 1 2 25.7 mm (USNM 283646). —
Sin 10831, Saba I., 17°38'N, 63°48'W. 641 m, 3.12.1969: 1 2 24.6 mm (USNM 283650). — Sin 10835. Si. Martin I..
18°28'N, 63°23'W. 671 m. 6.12.1969: 1 2 27.4 mm (USNM 283649).
"Gilliss": stn 31. 16°58'N. 79°28.1'N, 1116-1088 m, 28.07.1972: 1 <3 17.5 mm; 3 2 19.8-21.2 mm; 6 juvs
(RMNH).
Gulf of Mexico. "Oregon": stn 3516. 29°33'N, 88°38'W, 720 m. 18.03.1962: 3 3 18.1-18.4 mm: 52 16.1-
26.9 mm; 4 juvs (USNM). — Stn 4570. 23°H'N, 86°28'W, 900 m. 7.12.1963: 1 2 20.8 mm (USNM 260368). —
Stn 4815. 25°05'N, 96°27'W, 720 m, 12.04.1964: 1 2 21.7 mm (USNM 260368).
Types. — The male (19.0 mm cl) (NHM 1882.22) collected by the "Challenger" stn 23 off Sombrero i.
(18°24'N. 63°28'W, 810 m), is the holotype.
Fig. 4. — Cardus crucifer (Thomson, 1873), 2 27.0 mm, Caribbean Sea. "Oregon" stn 6697. 17°47'N. 6°39'W. 792-
828 m (USNM 273140): a. dorsal view of the cephalothorax: b. lateral view of the abdomen.
Description. — Carapace ovate, granular, setose. Dorsal surface of carapace bearing swollen earinae. Two
pairs of granulate submedian earinae anterior to cervical groove, lateral pair short; granulated tubercles posterior to
cervical groove, some along branchial carina, some scattered. Posterior margin of cervical groove raised. Anterior
margin of carapace nearly straight; single rostral spine prominent, upeurved. Internal angle of orbital sinus
prominently bispinose, external angle unarmed. Eyestalk curved distad under anterolateral angle of carapace;
vermicular appendage projects dorsally into orbital sinus, with basal spine directed forward. Basal antennular
segment squat, produced anteriorly to a sharp point; mesial margin spinose, two distalmost spines largest; single
spine on anlero-external angle. Lateral margins of carapace armed with long, slender, upeurved spines; spine
Source
296
B. GALIL
formula 6,7:5:17; anterior spine in each section most prominent. Posterior margin of carapace with three pairs of
antrorse spines, lateral pair largest.
Abdominal tergites granulate, medially carinate. Anterior tergite bearing antrorse spine. Tergites 2-5 with
bispinose median carina, anterior spine antrorse, submedian shallow oblique grooves. Carina on sixth tergum
blunt. Pleura bearing granulate tubercle mesially, scattered granules posterioly. Telson distally with two obtuse
submedian carinae.
Merus of PI with margins smooth, lacking subterminal spine. Carpus triangular, very short, small
subterminal spines on upper and lower margins. Chela swollen, upper margin irregularly spinose, distalmost spine
largest.
Color. — Described by WiLLEMOES-SUHM (1875a: 53) as "very beautiful", "a fine red, while the hairs
bordering it were yellowish".
Bouvier's (1925: 432) description relies on a watercolor sketch taken aboard the "Talisman"-, "un ton jaunatre
clair un peu lave de rose; cette derniere devient plus forte en dehors de la ligne mediane sur la region gastrique; elle
est egalement assez nette sur la ligne mediane de ] 'abdomen. Les pattes anterieures sont d'un gris noiratre clair
egalement un peu lave de rose".
DISTRIBUTION. — Eastern Atlantic: Portugal, Morocco, Canary Is, Azores; Western Atlantic: Florida Str.,
Bahama Is, Gulf of Mexico, West Indies, Caribbean Sea; 549-2195 m.
Genus HOMERYON nov.
Type Species. — Homeryon armarium sp. nov. Gender masculine.
DIAGNOSIS. — Carapace dorsoventrally flattened, ovate. Anterolateral angle of carapace produced, spiniform.
Lateral margin divided into three parts by cervical and postcervical incisions. Two rostral spines. Orbital sinus slit¬
like. Eyestalk filling orbit, vermicular tip distad, beneath anterolateral angle of carapace.
Cervical groove marked, arcuate, laterally bifurcate. Postrostral and postcervical median carinae well defined,
granulate, spinulate. Branchial carina well defined.
Abdomen broad, depressed. First abdominal tergite narrow, pleuron abbreviated. Abdominal tergites 2-5
sculptured, medially carinate; sixth tergite medially tuberculate. Abdominal pleura 2-5 rounded, successively
diminishing in size posteriorly; sixth pleuron subquadrate. Abdominal sternites with median knob. Telson
lanceolate, with short median basal ridge, two submedian carinae. Uropodal exopod lamellar, rounded; endopod oar¬
shaped, with basal lappet on interior margin.
Mesial margin of basal antennular segment anteriorly produced, upeurved, spinose, shorter than peduncle;
segment proximally squat, anterolateral angle spinose. Outer antennular flagellum short, inner flagellum as long as
antennal flagellum. Renal process well developed, tubular, fitting into hollow on underside of basal antennular
segment. Antennal scaphocerite lingulate, reaching to base of flagellum.
Buccal opening subquadrate, lateral margins prominent, keel-like. Mandible concave, cutting edge with
triangular teeth of uneven size; mandibular palp biarticulate. Maxillula with two incurved lobes, distally setose.
Maxilla with two slender lobes, internal lobe half as long as lateral; large scaphognathite. First maxilliped endopod
slender, exopodal lobe membraneous, reniform, extending further back than scaphognathite, exopod anteriorly
divided into two lobes enclosing efferent passage. Second and third maxillipeds slender, pediform; cpipod on basal
segment of third maxilliped half as long as ischium; dactyl slightly longer than preceding segment.
Pereiopods 1-4 chelate; P5 chelate in female, subchelate in male. PI robust; pereiopods successively shorter
posteriorly. Fingers of PI -2 crossing, their inner margin furnished with row of overlapping cornute platelets,
forming serrate edge. Fingers of chelae of P3-5 markedly upeurved.
Etymology. — Named for Homer, the blind poet, and eiyon - the first name ever used for the fossil kin of
the polychelids.
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
297
Remarks. — Homeryon gen. nov. differs from the other polychelid genera in having epipod on basal segment
of third maxilliped half as long as ischium; basal antennular segment squat; lateral margins of buccal opening
prominent, keel-like; fingers of pereiopodal chelae 3-5 upcurved.
Homeryon armarium sp. nov.
Fig. 5
Polycheles sp. - Baba, 1986: 157, 283. pi. 108.
MATERIAL EXAMINED. — Kvushu-Palau Ridge, 700 m. 18.12.1979, coll. M. Toriyama: I 2 58.1 mm (NFU
530-2-0740); 1 2 56.0 mm (NFU 530-2-1489). — 28°04'N, 134°20'E, 520 m, 20.02.1978, coll. O. Tabata: 2 2 61.4.
61.1 mm (NFU 530-2-0975).
Japan (?). Coll. T. Sakai: 1 6 37.7 mm (SMF 24649).
TYPES. — The female (58.1 mm cl) (NFU 530-2-0740) collected on the Kyushu-Palau Ridge, is the hololype;
the other specimens (NFU 530-2-0975, NFU 530-2-1-489) are paratypes.
Fig. 5. — Homeryon armarium sp. nov., 2 58.1 mm, hololype. Kyushu-Palau Ridge. 700 m (NFU 530-2-0740): a. dorsal
view of the cephalothorax; b, lateral view of the abdomen.
DESCRIPTION. — Dorsal surface of carapace setose, covered with robust, antrorse spinules. Frontal margin con¬
cave. Internal angle of orbital sinus triangulate, dense with antrorse spinules. Outer orbital margin pectinate.
Eyestalk bearing vermicular process, pointing distad. Antennular segment bearing spinose tubercle on antero-cxter-
nal angle. Lateral margins of carapace anterior to postcervical incision cristate, bearing 1 1-14 spines. Posterolateral
angle of cervical incision set with robust, antrorse spinules. Lateral margins of carapace posterior to postcervical
incision rounded, with several irregular rows of robust, antrorse spinules. Median postrostral and postcervical cari-
298
B. GALIL
nae swollen, thickly set with antrorse spinules; spinules blunter, more numerous behind cervical groove.
Postorbital carina swollen, thickly set with antrorse spinules. Cervical carina set with row of spinules. terminating
in spinose tubercle. Posterior margin of cervical groove minutely spinulose. Branchial carina sinuous, set with
clumps of antrorse spinules. Posterior margin of branchial groove spinose. Posterior margin of carapace smooth.
Abdominal tcrgites 1-5 sculptured, prominently tuberculate, mesially carinate. Tergites 2-5 with deep
transversely oblique grooves laterally. Carinae 2-5 bicuspid, anterior cusp larger, prominently granulose. Sixth
tergite bearing obsolescent granules mesially and laterally. Second pleuron reniform. granulate, with short groove
mesially; pleura 3-4 posteriorly granulate, with short rib mesially. Basal tubercle on telson granulate, submedian
carinae prominently tuberculate, lateral margins thickly set with acuminate tubercles. Uropodal exopod and
endopod thickly set with spinules laterally.
Merus of PI serrulate on lower margin and with a subterminal spine on upper margin; carpus of P I 3/4 as long
as merus, spinose on upper margin; chela of PI spinose on upper and lower margins. Merus ol P2 with several
rows of spines on lower margin and subterminal spines on upper margin; carpus and chela of P2 spinose on upper
margin.
Color. — Uniformly orange-red. Color plate in Baba (1986. fig. 108).
ETYMOLOGY. — From Latin, armarium (= chest, box) refers to the stocky, robust carapace ol the species.
Remarks. — "These specimens are not referred to any of the species in the literature now available to me. The
granulate body, short basal segment of antennule and dactyli of 3rd and 4th pcrcopods curving outward seems to be
unique to this species" (Baba, 1986: 283). Indeed, the slit-like orbital sinus, squat basal antennular segment and
the curved dactyli of 3-4 pereiopods are among the features that distinguish Homeryofl gen. nov. lrom the other
polychelid genera.
Distribution. — Kyushu-Palau Ridge, Japan; 520-700 m.
Homeryon asper (Rathbun. 1906)
Fig. 6
Polycheles asper Rathbun. 1906: 899; pi. 24 fig. 11. — De Man, 1916: 5 (list). — Bouvier. 1917: 35. — Firth &
Pequegnat, 1971: 41. — Dawson, 1997: 7.
MATERIAL EXAMINED. — Hawaiian Is. "Albatross", stn 4174, Niihau I.. 1323-1557 m, 1902: 1 9 32.0 mm
(USNM 30323).
Type. — The female (32.0 mm cl) (USNM 30323) collected by the "Albatross", stn 4174, off Niihau I., 1323-
1557 m, is the holotype of Polycheles asper Rathbun, 1906.
DESCRIPTION. — Dorsal surface of carapace densely covered with slender antrorse spinules. Frontal margin
slightly concave. Internal angle of orbital sinus bearing slender spine. Outer orbital margin pectinate. Eyestalk
bearing vermicular process, pointing distad. Antennular segment bearing three closely set spines on antero-external
angle. Spine formula of lateral margins of carapace 14-15:8-10:28-29. Median postrostral and postcervical carinae
bearing irregular double row of antrorse spinules, blunter behind cervical groove. Gastro-orbital region without
carina. Branchial carina sinuous, minutely spinulose. Diagonal groove anteriorly between median postcervical and
branchial carinae. Posterior margin of cervical groove minutely spinulose. Posterior margin of carapace smooth.
Abdominal tergites sculptured, granulate. Tergites 2-5 with deep, transversely oblique grooves on each side of
blunt median carina. Sixth tergite bearing obsolescent, beaded, lyre-shaped carina mesially, low granules laterally.
Second pleuron cordiform, with short groove mesially; pleura 3-4 with short rib mesially. Telson margins
serrulate, uropods smooth.
Remarks. — Rathbun (1906: 899) had not recognized that the single, juvenile asper differs much from other
polychelid species: "This species has the form of P. granulatus Faxon, but the dorsal surface is much rougher, the
Source MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
299
carapace more convex, its side margins more finely and obscurely cut. abdominal carinae, except the seventh,
nonprojecting, antennal scale rounded at tip instead of pointed".
H. asper differs from its congener in having blunt abdominal carinae rather than bicuspid, the second pleuron
cordiform rather than reniform, smooth uropods rather than granulate, and the lateral margins of carapace posterior
to postcervical incision cristate and serrulate rather than rounded, bearing several irregular rows of robust, antrorse
spinules.
DISTRIBUTION. — Known only from the type specimen, collected in Hawaii, between 1323-1557 m.
Fig. 6. — Homeryon asper (Rathbun, 1906), 9 32.0 mm. holotype. Hawaii. "Albatross" stn 4174. 1323-1557 m (USNM
30323): a, dorsal view of the cephalothorax; b. lateral view of the abdomen.
Genus PENTACHELES Bate, 1878
Pentacheles Bate, 1878a: 276.
Type Species — Pentacheles laevis Bate, 1878. designated by Fowler. 1912 {Annual Report of the New
Jersey State Museum for year 1911: 569). Gender masculine.
Diagnosis. — Carapace dorsoventrally flattened, subrectangular or ovale. Anterolateral angle of carapace
produced, spiniform. Lateral margins well defined, spinose; cervical and postcervical incisions markedly dividing
margin into three parts. Front concave, bearing two rostral spines. Orbital sinus well defined; eyestalk filling
orbit, pointing distad.
Cervical groove marked, arcuate, laterally bifurcate. Postrostral and postcervical carinae well defined, granulate
or spinose. Postorbital carina obsolescent, branchial carina obsolescent or spinulose.
300
B. GALIL
Abdomen narrow, laterally depressed. First abdominal lergite narrow, pleura fused, abbreviated. Abdominal
tergites 2-5 obliquely grooved, medially carinate; carinae blunt or anteriorly spinose; sixth lergite smooth. Second
abdominal pleuron larger, clypeiform. smooth; pleura 3-4 cordiform, successively diminishing in size posteriorly;
fifth pleuron triangulate. Abdominal sternites with median knob. Telson lanceolate, with median basal tubercle,
two submedian carinae. Uropodal exopod lamellar, rounded, grooved; endopod rounded, grooved, basal lappet on
interior margin.
Basal antennular segment proximally quadrate, lamellar; mesial margin granulate or spinose, upeurved.
anteriorly produced, reaching base of peduncle. Outer antennular flagellum short, inner flagellum as long as
antennal flagellum. Renal process well developed, tubular, fitting into hollow on underside of basal antennular
segment. Antennal scaphocerite lanceolate, not extending beyond pedunculc.
Buccal opening subquadrate. Mandible concave, cutting edge with triangular teeth of uneven size; mandibular
palp biarticulate. Maxillula with two incurved lobes, spiniform setae distally on anterior lobe. Maxilla with two
slender lobes, internal lobe half as long as lateral; large scaphognathite. First maxilliped endopod slender, exopodal
lobe membraneous, reniform, extending further back than scaphognathite. exopod anteriorly divided into two lobes
enclosing efferent passage. Second and third maxillipeds, slender, peditorm; third maxilliped epipod longer than
ischium; dactyl as long as propodus.
Pereiopods 1-4 chelate, P5 chelate in female, subchelate in male. PI larger, more robust than P2-5; pereiopods
successively shorter posteriorly. Fingers of first and second chelipeds crossing, their inner margin furnished with
row of overlapping cornute platelets, forming serrate edge. Fingers of chelae of P3-4 slightly upeurved. First male
pleopod spatuliform, spinulose, swelling distally on inner margin, spinules distally on mesial margin, distal
margin folded internally. P2-6 biramose, with rod-like appendix interna; second pair in male with appendix
masculina.
ETYMOLOGY. — From Greek, pente (= five) and chele (= claw) referring to the chelae of all five pereiopods.
REMARKS. — Bate established Pentacheles for species where "All the pereiopoda are more or less perfectly
chelate in both sexes" (Bate, 1888: 143). However, all species ( agassizii , euthrix, gracilis, laevis, obscura,
sculptus, validus) put into Pentacheles by Bate (1888) have males with fifth pereiopod simple or subchelate.
ALCOCK (1901b: 165) characterized Pentacheles as having "epipodite of the external maxillipeds . of fair size".
This does not correspond with Pentacheles as described by Bate (1878a), and further, ALCOCK. mistakenly
included hextii , a junior synonym of Polycheles typhlops , in his Pentacheles. Kemp & Sewell (1912). Selbie
(1914), De Man (1916) rejected Pentacheles and it fell into disuse early in this century.
Pentacheles is distinguished from the closely related Polycheles in having third maxilliped epipod longer than
ischium, proximally quadrate basal antennular segment, whereas Polycheles has rudimentary epipod and rounded
basal antennular segment.
Pentacheles gibbus Alcock, 1894
Pentacheles sp. - WOOD-MASON, 1891: 19.
Pentacheles gibba Alcock. 1894: 234. — WOOD-MASON & ALCOCK. 1894. pi. 8 fig. 4. — ANDERSON. 1896: 98.
Polycheles gibba - ALCOCK, 1899: 33.
Pentacheles gibbus - ALCOCK. 1901b: 173.
Polycheles gibbus - Df. Man, 1916: 5. — BERNARD, 1953: 86. — HOLTHUIS, 1984: 4. — Firth & PEQUEGNAT, 1971: 46.
— Dawson, 1997: 10.
Material EXAMINED. — Andaman Sea. " Investigator ": stn 114, 13°21'N, 93°27'E. 1687 m. 13.11.1890: 1 juv.
(IM 6799/10); 1 juv. (IM 6800/10).
Arabian Sea. "Investigator": stn 192, 15°H'N, 72°28'45"E. 1668-1703 m, 14.01.1895: I <3 62.0 mm (IM 46/10);
1 juv. 26.0 mm (IM 792/10).
Type. — The juvenile (IM 6799/10) collected by the "Investigator", stn 1 14, in the Andaman Sea is the
holotype, the other juvenile in the sample is a paratype.
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
301
Description. — Carapace convex, gibbous; lateral margins convergent anteriorly and posteriorly. Dorsal
surface of carapace thickly tomentose, minutely granulose. Anterior margin of carapace sinuous, bearing small,
bifid rostral spine. Internal and external angles of orbital sinus unarmed. Eyestalk with small spine directed upward.
Basal antcnnular segment foliaceous; mesial margin raised above rostrum, irregularly granulate; single spine on
antero-cxtcrnal angle. Spine formula of lateral margins of carapace 5-6:3:16-19; spines diminish considerably in
size posteriorly, posteriormost no more than granules. Median postrostral Carina irregularly granulate, median
postcervical carina with two rows of antrorse spinules. Gastro-orbital region unarmed. Branchial carina sinuous,
obsolescent. Posterior margin of carapace smooth.
Abdominal tergites set with prominent conic granules, medially carinate. Carinae on tergites 2-5 bicuspid.
Sixth tergite set with prominent conic granules on each side of short crest. Tergites 2-5 bearing well-defined,
transversely oblique submedian grooves. Pleura set with prominent conic granules. Second pleuron anteriorly
produced, ovate. Telson with granulate tubercle medially, two smooth submedian carinae posteriorly, margins
minutely spinulate.
First pereiopod remarkably short, smooth except for subterminal spines on upper margin of merus, chela and
lower margin of carpus. Merus proximally dilated. Carpus 3/4 as long as merus. P5 subchelate in male. Female
unknown.
Color. — "bright pink" (ALCOCK, 1894: 235).
Remarks. — P. gibbus is distinguished from the closely related P. obscurus in its humped carapace and
prominent conic granulation on abdominal terga and pleura.
No photograph is available as the author was prohibited from photographing in the Indian Museum.
Distribution. — Andaman Sea. Arabian Sea; 1668-1703 m.
Pentacheles laevis Bate, 1878
Fig. 7
Pentacheles laevis Bate, 1878a: 278; 1878b: 484; 1878c: 563; 1888: 144. pi. 15 figs 4c. 5. — Faxon. 1895: 118. —
Murray, 1897: 388. — De Man. 1916: 5 (list). — Sund. 1920: 224. — Estampador, 1937: 497: 1959: 43.
Pentacheles gracilis Bate, 1878a: 279; 1878b: 484; 1878c: 563; 1888: 146. pi. 16 figs 1-2. — Faxon, 1895: 118. —
Sund. 1920: 224. — Bernard, 1953: 86.
Polycheles granulatus Faxon, 1893: 197; 1895: 123, pi. 32 fig. I, pi. 33 figs 2. 2a. — Rathbun, 1906: 899. fig. 54. —
Stebbing, 1910: 378. — Selbie, 1914: 23. pi. 3 figs 1-2. — De Man, 1916: 5 (list). — Bouvier. 1917: 45, pi. 2
figs 7-14. — Stephensen, 1923: 67. — Balss, 1925: 200. — Calman, 1925: 18. — Miranda & Rivera. 1933: 17.
— Le Danois. 1948: 150 (list). — Barnard, 1950: 569. — Bernard, 1953: 86. — Squires, 1965: 89. fig. 38;
1966: 2. — Allen. 1967: 55. — Zariquiey. 1968: 210. — Zarenkov, 1969: 81. — Firth & Pequegnat, 1971: 47. —
Thiriot, 1974: 344. — TOrkay, 1976: 27. — Wenner, 1979: 443. — Wenner & BOESCH. 1979: 1 1 1. — Beaubrun.
1979: 39, figs 17-18. — KENSLEY, 1981b: 29. — POHLE, 1985: 22. — SQUIRES. 1991: 355. figs 187-188. —
Lemaitre & Alvarez-Leon, 1992: 33. — Falciai & Minervini, 1992: 134. — Gonzalez, 1995: 126. — Griffin &
Stoddart, 1995: 240, figs 4-5. — Hendrickx. 1995: 156. — Dawson, 1997: 10.
Pentacheles beaumontii Alcock, 1894: 236; 1901b: 175. — Wood-Mason & Alcock, 1894. pi. 8 fig. 3.
Polycheles beaumontii - Faxon, 1895: 125. — ALCOCK. 1899: 33. — Bouvier, 1905b: 644: 1905d: 2. — Holthuis,
1984: 4.
Polycheles dubitts Bouvier. 1905a: 480; 1905c: 4. — Richard. 1905: 3; 1907: 322.
Polycheles eryoniformis Bouvier, 1905b: 644; 1905d: 2; 1907: 62.
Polycheles gracilis - De Man, 1916: 5 (list). — Bouvier. 1917: 35. — Firth & Pequegnat, 1971: 46.
Polycheles laevis - FlRTH & PEQUEGNAT, 1971: 49. — Dawson, 1997: 12.
Polycheles aff. gracilis - MACPHERSON, 1984: 74.
not Polycheles beaumontii - Stubbing, 1908: 25; 1910: 377 [= Polycheles validus (A. Milne Edwards. 1880)].
Material EXAMINED. — Ibero-Morrocan Gulf. Balgim: stn CP 98. 34°29'N, 7°42’W. 1721-1773 m.
9.06.1984: 2 juvs (MNHN-Pa 1358).
Azores. "Travailleur": 44°04'N, 9°81'W. 1600 m, 12.07.1882: 1 juv. damaged (MNHN-Pa 1 138).
"Hirondelle": 39°27'N, 33°29'W, 1557 m, 30.07.1888: 1 2 36.4 mm; 2 juvs (MNHN-Pa 34).
302
B. GALIL
CANCAP 5: stn 5.005. 37°55'N, 24°46'W. 1650-2050 m, 24.05.1981: I juv. (RMNH). — Sin 5.090. 38°09'N.
28°31'W. 1320-1350 m. 2.06.1981: I <5 35.3 mm; 1 2 37.8 mm; 1 juv. (RMNH).
Canary Is. Cancap 4: stn 4.057. 28°47'N, 13°24'W, 1300-1335 m. 19.05.1980: 1 <3 48.0 mm (RMNH). -
Stn 4 102. 29°08’N. 13°19"W. 1290 m, 23.05.1980: 1 2 34.6 mm (RMNH).
Cape Verde Is. Cancap 6: stn 6.049. 14°52'N. 24°32'W. 1100-1300 m. 10.06.1982: 2 juvs. (RMNH). —
Stn 6.050. 14°53'N. 24°32'W. 1100-1200 m. 10.06.1982: I 6 54.1 mm (RMNH). — Stn 6.055. 15°46’N. 22°33'W.
950-1040 m, 12-13.06.1982: 2 3 45.8, 37.7 mm (RMNH). — Sin 6.065. 15°58'N, 22°33'W. 950-1040 m.
12-13.06.1982: 2 3 45.8, 37.7 mm (RMNH).
SE Atlantic. Rio Grande Ridge. "Prof. Mesyalsev": 12.07.1974. coll. B.N. KOTENEV: 2 <J 36.5. 39.8 mm;
1 ov. 2 56.0 mm; 1 juv. (ZMMU).
SE Atlantic. Discovery Seamount. "Prof. Mesyalsev": stn 154. 760 m, 10.12.1979. coll. M.V. BONDARENKO:
I 6 33.7 mm; I juv. (ZMMU).
United States. "Gill is": stn 83. 36°4I’N. 74°29’W. 1500-1350 m. 9.11.1974: 1 juv. (USNM 314136).
Bahama Is. "Cerda": stn 182. 27°55'N, 78°40’W. 860-897 m. 2.07.1963: 1 2 55.4 mm (RMNH 25834).
Panama. "Albatross": stn 3380. 4°03'N. 81°3rW. 1645 m, 5.03.1891: I 2 44.2 mm (MCZ 4578).
Galapagos Is. "Albatross": stn 4641, Hood I., 0I°35'S. 89°30'W, 1139 m. 7.11.1904: 1 juv. (USNM).
Nazca Ridge. "Prof Mesvatsev": 25°51'S. 84°34'W, 1050 m. 5.09.1983: 1 6 55.2 mm (ZMMU). — Stn 44.
20°44,9'S. 80°52.3'W. 18.09.1983: 1 2 39.3 mm (ZMMU)
Hawaiian Is. "Albatross" (id. M.J. RaTHBUN as P. granulatus): stn 3887. Molokai I., 21°16'30"N. I56°29'55 'W.
997-1456 m, 17.04.1902: I 2 23.7 mm (USNM 30321). — Stn 3989. Kauai I.. 22°05'55"N. 159°16'50"W. 900-693 m.
II 06.1902: 1 juv. (USNM 30315). — Stn 4007, btw. Honolulu and Kauai I.. 21°50'20"N, 159°31'40"W. 914-1003 m.
17.06.1902: 1 2 23.0 mm (USNM 30319). — Sin 4019. Kauai I.. 22°05’15"N. 159°17'W. 990-736 m. 21.06.1902:
I juv. (USNM 30315). — Stn 4111, Kaiwi Channel, 21°24'40"N. 157°23’55"W, 841-860 m. 24.07.1902: 1 6 50.1
mm; 1 juv. (USNM 30320). — Stn 4138. Oahu. 22°00'N. 1 59° 1 9'20" W. 857-788 m. 1.8.1902: 2 juvs (USNM 30318).
Marquesas Is. Eiao I.. MusoRSTOM 9: sin CP 1278, 75°2-S. 140°39’W. 1000 m. 5.09.1997: 2 2 24.6, 37.5 mm
(MNHN).
Wallis & Futuna Is. Musorstom 7: stn CP 592, 12°32'S. I74°22'W, 730-775 m. 24.05.1992: 1 juv. (MNHN-Pa
1655). — Stn CP 623, 12°34’S. 178°15'W, 1280-1300 m. 28.05.1992: 1 juv. (MNHN-Pa 1656).
Fiji Is. " Challenger ": stn 174c. Kadavu I.. 19°07'50"S, 1 78° 19'35"E. 1098 m. 1 2 24.7 mm (NHM 1888.22).
South Fiji Basin. NZOI: stn Z8880, 30°00.1'S, 177°20.1'E. 29.07.1997. coll. J. Wills: I 6 30.3 mm (N1WA).
Vanuatu. Musorstom 8: stn CP 956. 20°33.41'S. I69°35.95'W. 1175-1210 m, 20.09.1994: 1 2 44.2 mm; 1 juv.
(MNHN-Pa 1647). — Stn CP 990. 18°51.63'S. 168°50.98'W, 980-990 m. 24.09.1994: 1 2 29.3 mm (MNHN-Pa 1648).
— Stn CP 996, 18°52.41'S. I68°55.73'W. 764-786 m. 24.09.1994: I 2 49.8 mm (MNHN-Pa 1649). — Stn CP 1037.
18°03.70'S. 168°54.40'W. 1058-1086 m. 29.09.1994: 1 6 32.8 mm (MNHN-Pa 1650). — Stn CP 1112. 14°52.57'S.
1 67° 1 1.89'W. 961-950 m. 8.10.1994: 1 juv. (MNHN-Pa 1651).
New Caledonia. Biocal: stn CP 30, 23°09'S. 166°41'E. 1140 m. 29.08.1985: I 2 37.6 mm; 3 juvs (MNHN-Pa
1657). — Stn CP 31, 23°08'S, 166C51'E, 850 m. 29.08.1985: 1 2 56.2 mm; 2 juvs (MNHN-Pa 1658). — Stn CP 32,
23°07'S. 166°51'E. 825 m, 29.08.1985: 1 6 47.2 mm. 1 juv. (MNHN-Pa 1659). — Stn CP 54. 23°10'S. I67°43'E. 950-
1000 m. 1.09.1985: 1 6 37.8 mm (MNHN-Pa 1660).
BtOGEOCAL: stn CP 290. 20°36.91'S. 167°03.34'E, 920-760 m. 04.1987: 1 6 46.1 mm (MNHN-Pa 1750).
VOLSMAR: stn CP 26, 22°22.8'S. I71°21.4'E, 980 m. 4.06.1989: 1 d 48.4 mm (MNHN-Pa 1662).
Bathus 1: stn CP 651, 21°41.8'S. 166°40.rE. 1080-1180 m, 11.03.1993: 1 3 22.7 mm (MNHN-Pa 1663). —
Stn CP 661, 21°05'S. I65°50.05'E. 960-1100 m. 13.03.1993: 1 2 44.9 mm (MNHN-Pa 1664).
Bathus 2: stn CP 767, 22°10.47’S. 165°59.10'E, 1060-1450 m, 17.05.1993: 4 3 22.1-33.7 mm; 1 2 22.7 mm
(MNHN-Pa 1665).
Bathus 3: stn CP 844. 23°60.38'S, 16°45.60'E, 908 m, 1.12.1993: 1 2 58.3 mm (MNHN-Pa 1666).
BATHUS 4: stn CP 951. 20°31.44'S. 164°54.57'E. 960 m. 10.08.1994: 1 juv. 16.6 mm (MNHN-Pa 1667).
Halipro 1: stn 867. 21°26.15'S, 166°18.17'E, 720-950 m. 22.03.1994: 1 3 juv. 24.9 mm (MNHN-Pa 1668). —
Stn 874. 23°05'S, 166°48'E. 708-830 m, 30.03.1994: 1 2 57.0 mm (MNHN-Pa 1669). — Stn 876. 23C10.41'S,
166°49. 16'E, 870-1000 m, 31.03.1994: 2 2 52.9. 63.4 mm (MNHN-Pa 1670); 2 3 juvs 22.7. 26.4 mm.
Halipro 2: stn BT 41. 25°51'S, 167°12'E. 1317-1383 m. 14.11.1996: 2 2 36.5. 60.4 mm (MNHN-Pa 1689). —
Stn BT 69. 24°40'S, 168°09'E. 943-1080 m, 19.11.1996: 1 2 38.3 mm (MNHN-Pa 1691). — Stn BT 73. 24°5 1 'S.
167°40'E, 1043-1102 m. 20.11.1996: 1 2 59.6 mm (MNHN-Pa 1692). — Stn BT 85. 23°40'S. 168°05'E. 935-1100 m.
23.11.1996: 1 ov. 2 49.3 mm (MNHN-Pa 1693). — Stn BT 97, 24°00'S. 161°49'E. 964-1031 m. 25.11.1996: 4 2
30.7-50.1 mm (MNHN-Pa 1694). — Stn BT 101, 24°19'S, 161°43'E. 970-1063 m, 26.11.1996: I ov. 2 59.1 mm
(MNHN-Pa 1695). — Stn 102, 24°31'S. 161°52'E. 1060-1130 m. 26.11.1996: 1 2 43.2 mm; 1 juv. (MNHN-Pa 1696). —
Stn BT 103. 24°54'S. 162°09'E. 1235-1256 m, 26.11.1996: 1 2 24.8 mm. 2 juvs (MNHN-Pa 1697). — Stn BT 104.
25°26’S, 162°27'E, 1118-1124 m, 27.11.1996: 1 ov. 2 44.3. mm; 2 juvs (MNHN-Pa 1698).
Chesterfield Is. Musorstom 5: stn 323. 21°I8.52'S. 157°57.62'E, 970 m. 14.10.1986: 1 3 44.7 mm; 2 2 48.4.
56.2 mm; II juvs (MNHN-Pa 1652). — Stn 324. 21°15.01S. 157°51.33'E, 970 m. 14.10.1986: 2 3 22.8. 42.3 mm:
Source MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
303
1 2 53.7 mm; 6 juvs (MNHN-Pa 1653). — Stn 326. 21°07.40'S, I57°47'E. 980 m, 14.10.1986: 1 3 33.5 mm (MNHN-
Pa 1654).
Indonesia. "Challenger": stn 214, Molucca Sea. 4°33'N, 127°06'E. 914 m. 10.02.1875: 1 2 17.9 mm (NHM
1888.22).
"Albatross"-, stn 5601, Sulawesi, 1°13. 10'N. 125°17.05'E. 1377 m. 13.11.1909: I 9 23.1 mm; 2 juvs (USNM).
Philippines. "Albatross": stn 5127, Panay. 10°22'45"N, 121°48'15"E, 1724 m. 4.02.1908: 1 3 23 4 mm
(USNM). — Stn 5428. Palawan, 9°13’N, I18051'15"E, 1989 m, 3.04.1909: 1 3 24.6 mm; 2 2 25.1. 39.6 mm; 7 juvs
(USNM). — Stn 5468, Lagonoy Gulf, San Bernardino Str., 13°35'39"N, 123°40'28"E, 1024 m. 18.06 1909- I 9
35.9 mm (USNM).
Musorstom 2: stn CP 56. 13°54’S, 1 19°57'E, 970 m, 28.1 1.1980: I 2 44.1 mm (MNHN-Pa 1741).
Australia. New South Wales. " Kapala stn K79-20-06. 33°36’S, 152°06'E, 914 m, 4.12.1979. id. H.E. Stoddart
as Polycheles granulatus: 1 2 52.7 mm (AM P44898). — 33°30'S. 152°I0'E, 880 m, Nov. 1992, id. H.E. Stoddart as
Polycheles granulatus: 1 3 43.7 mm; 1 2 52.4 mm (AM P44908).
Tasmania. Off St. Patrick’s Head. 920 m, 24.07.1983, coll. R.M. Green: 1 6 40.2 mm (TM G3278). — 4 1 0 1 4'S,
148°45'E, 950 m. 17.06.1991, coll. R. Brewster: 1 ov. 2 42.3 mm (TM G3489).
Tasman Sea. Lord Howe Rise. NZOI: stn U 197, 34°09.8'S, 163°36.7'E. 1 186 m. 25.9.1982: 1 juv. (NIWA).
New Zealand. NZOI: stn V 370, 42°41.70'S, 179°03.42'W, 1024 m, 12.09.1989: 1 2 24.1 mm (NIWA). — Stn X
508, 42°5I.86'S, 174°56.10’W, 1163-1173 m, 9.07.1994: 1 3 30.7 mm (NIWA). — Stn Z8445. 44°0I'S. 160°37'W
1000 m. 23.08.1995: I 2 54.7 mm (NIWA).
Madagascar. "Vauban": stn CH 131, 13°46'S. 47°33'E, 1490-1600 m. 20.01.1975: 3 juvs (MNHN-Pa 1742).
Types. — The juvenile female (17.9 mm cl) (NHM 1888.22) collected by the "Challenger" sin 214, off the
Moluccas (4°33'N, 127°6'E, 914 m), is the lectotype of Pentacheles laevis Bate. 1878, designated by Sund (1920:
225).
The female (24.7 mm cl) (NHM 1888.22) collected by the "Challenger" stn 1 74c, off Fiji (19°07.50'S.
178°19.35'E, 1098 m), is the holotype of Pentacheles gracilis Bate, 1878.
The female (44.2 mm cl) (MCZ 4578) collected by the "Albatross" stn 3380. off Panama (43'N. 8I°31'W.
1645 m), is the holotype of Polycheles granulatus Faxon. 1893.
The male (44.0 mm cl) (IM 520/7) collected by the "Investigator" off Colombo. Sri Lanka, at a depth of
1234 m, is the holotype of Pentacheles beaumonti Alcock, 1894.
The female (36.4 mm cl) (MNHN-Pa 34) collected by the "Hirondelle", off the Azores, is designated here as the
lectotype of Polycheles dttbius Bouvier. 1905; the two juvenile specimens of this sample and a juvenile collected
by the "Travaillettr" (MNHM-Pal 138) are paratypes.
Description. — Carapace ovate. Dorsal surface of carapace granulose, sparsely setose. Anterior margin of
carapace slightly concave, two long rostral spines adjoining at base. Internal and external angles of orbital sinus
prominently produced, spinose. Eyeslalk bearing spine, curved distad. Basal antennular segment lamellar, produced
anteriorly to an acute point; mesial margin prominently spinose; single spine on antero-exlernal angle. Lateral
margins of carapace lined with upeurved spines, anteriormost spine largest, diminishing gradually in size
posteriorly; spine formula 7-9:3-4:14-15. Median postrostral carina bearing irregular number of spines followed by
close-set granules; median postcervical carina with two rows of antrorse, mammilate tubercles. Gastro-orbital
region unarmed. Posterior margin of cervical groove unarmed. Branchial carina sinuous, spinulate; spinules
increasing in size posteriorly. Posterior margin of carapace smooth.
Abdominal tergites medially carinate; carinae on tergites 1-3 bearing antrorse beak; tergites 4-5 with blunt
carina; sixth tergitc noncarinate; tergites 2-5 with well defined, oblique submedian grooves. Surface of pleura
smooth, minutely punctate; anterior pleuron with mesial groove proximally; following pleura bearing short,
obsolescent mesial rib; margins smooth. Telson with rounded tubercle anteriorly, two smooth, confluent carinae
posteriorly.
Merus of cheliped finely granulate on upper margin with a subterminal claw; lower margin spinulate, spines
increasing in size proximally. Carpus nearly as long as merus, claw-like spine on interior and exterior margins.
Upper margin of chela with two (or more) serrulate rows, interior row with largest spines, claw-like subterminal
spine; lower margin with single serrulate row. P5 subchelate in male, chelate in female.
Color. — "Entire carapace, abdominal segments and telson light rose pink, spines tipped with white.
Antennular and antennal flagella and peduncles, pereopods, pleopods, endopods and exopods of uropods a darker
304
B. GALIL
pink to red." (Griffin & Stoddart, 1995: 242). Bouviers ( 1917: 150, fig. 7) colored drawing was done "d'apres
des aquarelles executees au moment de la capture".
Fig. 7 . — Peniacheles laevis Bate, 1878, <3 42.3 mm, New Caledonia. MUSORSTOM 5. sin 324. 21^1 5.01 S, 157 51. 33 E,
970 m (MNHN-Pa 1653): a. dorsal view of the cephalothorax; b. lateral view of the abdomen.
REMARKS. — P. laevis is easily distinguished from P. validus in having fewer postcervical lateral spinules,
bearing prominent spines anteriorly on the median carina, lacking the gastro-orbital spines, and lacking spines on
the posterior margin of the cervical groove.
The single female specimen of Pentacheles laevis was "not a very perfect one" (Bate. 1888: 146) and was
inaccurately described and inadequately drawn. Though a juvenile, as evinced from its subchelate fifth pereiopod.
the ovate carapace, the prominently produced internal and external angles ol the orbital sinus, the spinosc mesial
margin of the basal antennular segment, the single spine on its antero-external angle, and the form of the median
abdominal carinae leave no doubt as to its identity. Bate's (1888) description of P. gracilis was based on a slightly
larger specimen, but the accompanying drawing (pi. 16) has erroneously depicted the frontal margin much produced
and was criticized by SUND (1920). However, it was Faxon's (1893) description of P. granulatus that was
favoured by most subsequent authors. FAXON himself (1895: 124) observed: 'This species [P. granulatus ]
somewhat resembles P. laevis (Bate), P. validus (A.M. Edw.), and P. debilis (Smith)... an East Indian species,
Polycheles beaumontii , bears the closest likeness to P. granulatus. Indeed the resemblance between these two
forms is so great that I am inclined to regard P. beaumontii as, at most, but a geographical race of P. granulatus
scoring three out of four synonymies. SELBIE (1914: 25), following RaTHBUN (1906), wrote no doubt that
P. beaumontii , Alcock, is merely a synonym of P. granulatus. Faxon". BOUVIER (1905a: 481) described P.
dubius "elle se distingue par l'atrophie complete des carenes exogastriques, par ses carenes exocardiaques faiblement
indiquees, par lc bord posterieur incrme de la carapace, par la forte saillie aigue qui limite en dedans et en dehors
chaque sinus orbitaire, par les carenes obtuses ct peu saillantes de ses tergites abdominaux 4 et 5, par I'absencc de
Source MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
305
tout dessin sur le tergitc 6" - a perfect description of laevis. SELBIE (1914: 25) recognized that "P. dubius (Bouvier)
is also clearly a synonym of P. granulatus." Bouvier (1917: 51) conceded that :"unc etude plus scricusc me donne
la certitude que toutes ces formes [P. dubius, P. eryoniformis] doivent etre identifies avec le P. granulatus." -
indeed with laevis.
DISTRIBUTION. — Worldwide; SW Ireland, Ibero-Morrocan Gulf, Madeira, Canary Is, Azores, Cape Verde Is,
Mid-Atlantic bight, southwestern Atlantic, Rio Grande Ridge, Newfoundland. Nova Scotia, United States. Bahama
Is, Gulf of Panama, Colombia, Galapagos Is, Nazca Ridge. Marquesas Is, Hawaii, Fiji, Wallis & Futuna Is.
Vanuatu, New Caledonia, New Zealand, Tasmania, Australia, Indonesia, Molluca Sea, Philippines, Sri Lanka,
Madagascar; 347 m (DAWSON, 1997) - 2505 m (ALCOCK, 1899).
Pentacheles obscurus Bate, 1878
Fig. 8
Pentacheles obscura Bate, 1878a: 279; 1878b: 484; 1878c: 563; 1888: 143, pi. 15 fig. 2. — Pagenstecher, 1879: 63.
— Faxon, 1895: 12.
Pentacheles carpenteri Alcock, 1894: 235; 1901b: 174. — ALCOCK & Anderson, 1895. pi. 10 fig. I.
Polycheles carpenteri - ALCOCK, 1899: 33. — DE Man, 1916: 23, pi. 1 fig. 3. — BOUVIER. 1917: 35. — BERNARD. 1953:
86. — Zarenkov, 1969: 81. — Firth & Pequegnat. 1971: 42. — Dawson, 1997: 7.
Polycheles obscurus - De Man, 1916: 5 (list). — Bernard. 1953: 86. — Firth & Pequegnat. 1971: 49. — Dawson,
1997: 12.
Material EXAMINED. — Wallis and Futuna Is area. Musorstom 7. 51V Combe Bank: sin CP 622, 12°34'S,
178°H'W, 1280-1300 m, 28.05.1992: 1 2 55.6 mm (MNHN-Pa 1671).
New Caledonia. BiOGEOCAL: stn CP 260, 21°00'S, 166°58.34'E. 1820-1980 m, 17.04.1987: I 2 43.2 mm
(MNHN-Pa 1743).
BATHUS 1: stn CP 704, 20°55.8'S. 165°38.6'E, 1100-1200 m, 18.03.1993: I 3 39.0 mm (MNHN-Pa 1672).
New Guinea. " Challenger ". stn 218. 2°33'S. 144°4'E, 1857 m, 1.03.1875: 1 juv. (damaged) (NHM 1888.22).
Indonesia. " Siboga ": stn 126, Molucca Sea, 3°27.1’N. 125°18.7’E, 2053 m, 20.07.1899. id. de Man as Polycheles
carpenteri : 1 3 39.8 mm (ZMA).
"Galathea": stn 471, off the south coast of Java, 10°26'S, 1 14°15 E, 3080 m. 10.09.1951: I 2 20.8 mm (ZMC).
Gulf of Aden. "Meteor 5": stn Me5-257. 13°06.6'N. 47°0'E. 2227-2250 m. 11.03.1987: 1 2 29.4 mm (SMF
24659). — Stn Me5-271, 12°56.7'N, 47°47.0'E, 2276-2282 m. 14.03.1987: 2 2 43.9, 32.3 mm; 1 juv. (SMF 24660).
Madagascar. "Vauban": CH 138, 13°48.8'S, 47°29'E. 1800-2000 m, 27.02.1975, coll. A. Crosnier: I <J 50.2 mm
(MNHN-Pa 1673).
TYPES. — The damaged juvenile (NHM 1888.22) collected by the "Challenger" stn 218. off New Guinea
(2°33'S, 144°4'E, 1857 m), is the holotype of Pentacheles obscurus Bate, 1878.
The female (38.5 mm cl) (IM 76/7) collected by the "Investigator" off Carpenter's Ridge, Bay of Bengal, at
depths of 2505-2616 m, is the holotype of Pentacheles carpenteri Alcock, 1894.
Description. — Carapace ovate, lateral margins convergent anteriorly and posteriorly. Dorsal surface of
carapace thickly tomentose, minutely granulose laterally. Anterior margin of carapace sinuous, bearing small, bifid
rostral spine. Internal and external angles of orbital sinus unarmed. Eyestalk with small spine directed upward.
Basal antennular segment foliaccous, mesial margin raised above rostrum, irregularly granulate; single spine on
antero-external angle. Spine formula of lateral edges of carapace 5:3:27-28; spines diminish considerably in size
posteriorly, posteriormost no more than granules. Median postrostral carina irregularly granulate anteriorly, median
postcervical carina bearing two rows of anlrorse granules. Gastro-orbital region unarmed. Branchial carina sinuous,
obsolescent. Posterior margin of carapace smooth.
Abdominal tergites medially carinate; carinae on tergites 2-5 bicuspid; sixth tergitc with short crest posteriorly.
Tergites 2-5 bearing well-defined, transversely oblique, submedian grooves. Pleura nearly smooth; second pleuron
anteriorly produced, ovate. Telson with granulate tubercle medially, two smooth submedian carinae posteriorly.
First pereiopod remarkably short, smooth except for subterminal spines on upper margin of merus. palm and
lower margin of carpus; merus proximally thickened; carpus 3/4 as long as merus. P5 subchelate in male, chelate
in female.
306
B. GALIL
Fig. 8. — Pentacheles obscurus Bate, 1878, 3 50.2 mm, Madagascar. 13°48.8'S, 47°29'E. 1800-2000 m (MNHN-Pa
1673): a, dorsal view of the cephalothorax; b. lateral view of the abdomen.
Remarks. — The female holotype of Pentacheles obscurus was described by Bate (1878a: 279) as being "in a
very imperfect condition". Though a juvenile, as evident from its subchelate fifth pereiopod, the densely tomentose
carapace, the single spine on the antero-external angle of the basal antennular segment, the shallow orbital sinus,
the bicuspid carinae on the abdominal tergites, and the remarkably short first pereiopod with proximally thickened
merus and elongate carpus, leave no doubt that P. carpenter i Alcock (1899) is synonymous with P. obscurus.
Distribution. — Melanesia, New Caledonia, New Guinea. Molluca Sea. Madagascar. Gulf of Aden; 1 1 00-
3080 m.
Pentacheles snyderi (Rathbun, 1906)
Fig. 9
Polyeheles snyderi Rathbun, 1906: 898, pi. 24 fig. 9. — De Man, 1916: 6 (list). — Bernard. 1953: 86. — Firth &
Pequegnat, 1971: 50. — Dawson. 1997: 12.
Material EXAMINED. — Bay of Biscay: "Thalassa": stn 448. 48°45.0'N, 11°19.8'W. 1830-1870 m. 1973: 1 3
26.0 mm; 1 juv. (MNHN-Pa 358). — Stn 453. 48°34.0'N, 10°51.6'W. 1975-2070 m. 1973: I 3 21.1 mm (MNHN-
Pa 359).
Biogas III. CV 23: stn 1, 47°32.70'N, 8o34.20'W. 2034 m, coll. 1FREMER: 1 3 21.8 mm; 1 2 26.8 mm (MNHN-
Pa 540). — CV 32: stn 6. 44°07.60'N, 4°15.80'W. 2245 m. coll. IFREMER: 1 juv. (MNHN-Pa 548).
Biogas XI, CP 37: stn 1, 47°33.80'N, 47°34.27'W. 11.10.1981, 2175 m, coll. IFREMER: 1 damaged (MNHN-
Pa 1138).
Hawaiian Is. "Albatross": stn 4151, vicinity of Modu Manu. 573-1463 m, 7.08.1902: 1 3 34.5 mm (USNM
30333).
Source MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
307
Ohau. 21°I5.30’N, 158°15.20’W. 0-1100 m, 9.09.1977, coll. F. Clarke: 1 <5 36.0 mm (RMNH 32623).
Indian Ocean Ridge. "Prof. Mesyatsev": 11°31.0'S, 88°55.4'E. 1600-1700 m. 18.02.1979. coll. M.G. Kar-
pinski: 2 6 36.6, 42.7 mm; 1 2 50.7 mm (ZMMU).
Type. — The male (34.5 mm cl) (USNM 30333) collected by the "Albatross" stn 4151. off Hawaii, is the
holotype of Polycheles snyderi Rathbun, 1906.
Fig. 9. — Pentacheles snyderi (Rathbun, 1906). <3 42.7 mm, Indian Ocean Ridge. "Prof. Mesyatsev" 11°3 1.0'S.
88°55.4'E, 1600-1700 m (ZMMU): a. dorsal view of the cephalothorax: b. lateral view of the abdomen.
DESCRIPTION. — Carapace narrow, subrectangular, slightly convergent anteriorly. Dorsal surface of carapace
setose, wholly covered with slender spinules. Frontal margin concave, rostral spines adjoining at base. Internal
angle of orbital sinus produced, triangular, unispinose, external orbital angle with 3-4 upeurved spines. Eyestalk
bearing upeurved spine distad. Basal antennular segment lamellar, produced anteriorly to a sharp point, mesial
margin distally spinose, anterolateral angle unispinose. Spines on lateral margins of carapace upeurved,
anteriormost largest. Lateral spine formula 9-10:5-8:30-35, spines successively smaller posteriorly. Median
postrostral carina defined, irregularly spinulose, bearing half way to cervical groove pair of stouter spines.
Posterior margin of cervical groove spinulose. Postorbital carina obsolescent. Branchial carina obsolescent,
sinuous, minutely spinulose. Posterior margin of carapace smooth.
Abdominal tergites 1-5 medially carinate, carinae blunt, lacking antrorse spine, sixth tergite noncarinate,
smooth. Tergites 2-5 bearing transversely oblique submedian grooves. Pleura smooth. Second pleuron ovate.
Tclson with blunt median crest anteriorly, two smooth carinae posteriorly.
Merus of PI basally dilated, with granulate margins and subterminal claw-like spine on upper margin: carpus
nearly 2/3 as long as merus with subterminal spines on upper and lower margins; upper margin of propodus
minutely serrulate, with subterminal spine. P5 subchelate in male, chelate in female.
308
B. GAUL
Remarks. — P. snyderi was summarily described by RATHBUN (1906) from a single specimen from Hawaii.
P. snyderi is distinguished from P. validus by its subrectangular. spinose carapace and pectinate external orbital
margin.
DISTRIBUTION. — Bay of Biscay, Indian Ocean Ridge, Hawaii. Known reliably between I 100-2245 m,
collected too in a trawl between 573-1463 m.
Pentacheles validus A. Milne Edwards. 1880
Fig. 10
Pentacheles validus A. Milne Edwards, 1880: 65. — YOUNG, 1900: 443.
Polycheles validus - Mocquard, 1883: 153. — Faxon, 1895: 124. — BouvtER, 1905a: 480; 1917: 45; 1925: 434.
fig. 10, pi. 5. figs 1-2. — De Man. 1916: 6 (list). — Bernard. 1953: 86. — Sivertsen & Holthuis, 1956: 41. —
Dollfus, 1956: 135. — Firth & Pequegnat, 1971: 61. — Pequegnat et at., 1971: 4. — Beaubrun. 1979: 38.—
Wenner, 1979: 443. — Wenner & Boesch, 1979: 111. — Rodriguez, 1980: 194 (list). — Pohle, 1985: 21. —
De Saint Laurent, 1985: 475. — Dawson, 1997: 16.
Pentacheles debilis Smith, 1884: 360; 1886a: 188 (list); 1886b: 607, pi. 7. fig. 2. — Verrill. 1885: 554 (list).
Pentacheles laevis Bate, 1888: 144, pi. 15, fig. 5 (p.p.). — MURRAY, 1986: 388.
Polycheles debilis - Bouvier, 1905a: 480; 1905c: 3. — De Man, 1916: 5 (list). — Faxon, 1895: 124.
Polycheles debilis var. armatus Bouvier, 1905c: 4. — De Man, 1916: 5 (list).
Polycheles beaumontii (?) - Stebbing, 1908: 25; 1910: 377.
Pentacheles debilis - Fowler, 1912: 570 [erroneous spelling],
Polycheles - MURRAY & Hjort, 1912: 420 fide SlVERTSEN & Holthuis, 1956].
Polycheles laevis - De Man, 1916 (list) (p.p.).
Polycheles demani Stebbing. 1917: 28, pi. xcii. — Calman, 1925: 17. — Barnard, 1950: 570. figs I05a-c. —
Kensley, 1968: 292; 1974: 69; 1981b: 29. — Firth & Pequegnat. 1971: 45.
Polycheles chilensis Sund, 1920: 226. — Holthuis, 1952a: 78. — Bernard. 1953: 86. — Bahamonde. 1963: 4. —
Zarenkov, 1969: 81. — Firth & Pequegnat, 1971: 42. — Retamal. 1981: 15. — Wicksten, 1989: 304. —
Dawson, 1997: 8.
Material EXAMINED. — Bay of Biscay. Biogas IV, CP 01: 47°34.6'N. 8°36.8'W. 25.02.1974. 2245 m, coll.
IFREMER: 1 2 44.4 mm; I <3 damaged (MNHN-Pa 563).
Canary Is. "Talisman": stn 39, 30°08N, 14°02'W. 2300 m, 23.06.1883: I juv. (MNHN-Pa 15). — Sin 41. 30°01'N.
14°06'W, 2115 m, 24.06.1883: 2 juvs (MNHN-Pa 16). — Stn 43. 29°52'N. 14°04’W. 2075 m, 25.06.1883: 2 juvs
(MNHN-Pa 14).
Cancap 2: stn 26, 27°50'N, 14°29'W. 1980-2200 m. 25.08.1977: 8 2 20.0-49.7 mm (R.MNH).
Azores. Cancap 5: stn 5.004, 38°06'N, 25°49'W. 2400-3100 m. 24.05.1981: 6 juvs iRMNH). — Stn 5.052.
36°42'N. 25°09'W, 2500-3000 m. 30.05.1981: 1 8 42.6 mm (RMNH). — Stn 5.171, 39°20'N. 30°52'W. 1874-1887 m,
11.06.1981: 1 juv. (RMNH).
West Africa. Ivory Coast. "Pillsbury": stn 314, 4°58'N. 3°48’E. 2268-2332 in. 27.05.1965: 1 8 69.7 mm; 1 $
69.3 mm (R.MNH).
South Africa. Cape Point Lighthouse: NE 40 mi.. 1024-1280 m, coll. Dr. Gilchrist: I 2 66.9 mm (NHM
1928.12.1.337). — NE 43 mi., 1620 m, I 3 61.1 mm (SAM A4343).
South Atlantic. South Georgia 1., 53°26.7’S, 36°32.6'W, 1967-2186 m, 17.05.1975: 1 juv. (USNM).
Bahama Is. (F.M. Bayer). N of Andros I., 1737 m, 5.03.1973: 1 juv. (USNM 154324). — N of Andros !..
24°24.48'N, 76°11.24'W. 1757 nr, 13.02.1974: 3 juvs (USNM 154326). — Exuma Sound. 24°39'N, 76°26.54'W.
1624 m, 12.04.1975: 2 8 49.3.50.6 mm; 1 2 22.9 mm (USNM 154399). — 24°3I.54’N, 76°17.12’W. 1673 m.
12.04.1975: 1 8 53.7 mm; I 2 23.5 mm (USNM 154398). — 24°38.54'N, 76°26.15'W, 1628 m, 15.04 1975- I 2
28.5 mm (USNM 154328). — 23°55'N, 75°27.12'W, 2178 m, 16.04.1975: 1 2 33.8 mm (USNM 154329).
"Iselin": stn 73, 24°46'N. 75°41'W. 1792 m. 4.03.1972: 1 2 (RMNH 31659). — Stn 346. 22°55'N. 75°17.06'W.
2426 m. 17.04.1975: 12 37.1 mm (USNM 154331). — Stn 8007, 22°46'N, 75°34'W. 7.09.1980: 1 8 57.6 mm
(USNM 344508).
United States. East Coast. "Albatross": stn 2074. 41°43'N, 65°2r50"W, 2393 m, 3.09.1883: 1 8 20 0 mm
(USNM 7145).
Caribbean Sea. Jamaica. "Albatross": stn 2140, 17°36'10"N, 76°46'05"W. 1768 m, 11.03.1884, id
M.J. Rathbun as P. debilis: I juv. (USNM 23751).
"Oregon": stn 2571. 26°34'N, 90°31'W, 2340 m, 27.07.1959: 1 6 50.7 mm (USNM 232276). — Stn 2574, 26°34’N,
89'-53W, 2610 m, 28.07.1959: 1 8 32.2 mm; 1 2 77.0 mm (USNM 232276). — Stn 2575. 27°06’N, 89°13'W. 1980-
Source : MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
309
2160 m. 29.07.1959: 1 3 31.4 mm (USNM 232276). — Sin 2814, 28°53'N, 87°47'W. 1710-1890 m. 13.07.1960: 1 3
51.7 mm (USNM 232276).
"Pillsbury": stn 634, 23°33'N. 82°47'W, 1638-1757 m, 25.03.1968: 1 <J 35.8 (RMNH). — Sin 1178, 19'14'N,
73°I4'W, 1750 m, 30.06.1970: 1 <3 62.2 mm (RMNH).
Gulf of Mexico. "Albatross": stn 2379, 28°00'15"N. 87°42'W. 2684 m, 2.03.1885: 1 2 27.3 mm (USNM
23645). — Sin 2383. 28°32’N, 88°05'W. 2161 m, 3.03.1885, id. M.J. RATHBUN as P. debilis: 1 juv. (USNM 23752).
"Oregon": stn 2577, 27°48'N, 88°45'W, 1530-1980 m, 30.07.1959: I 3 48.1 mm (USNM 232276). — Sin 2819,
28°35'N, 88°16.5'W, 1620-1800 m, 15.07.1960: 1 ov. 2 56.8 mm (USNM 232276). — Sin 2820, 28°23'N. 88°-2l.5’W,
1800 m, 15.07.1960: 1 3 46.0 mm (USNM 232276).
"Oregon II": sin 10878, 28°54'N, 87°29'W, 1830 m. 16.01.1970, id. B. Andryzak as P. demani: 1 3 55.2 mm
(USNM 283665). — Sin 10881, 24°5I'N, 96°00'W. 1669 m, 24.01.1970, id. 13. Andryzak as P. demani : 4 3 39.9-
49.6 mm; I 2 63.2 mm (USNM 283667). — Stn 10897, 281,45'N, 88°23’W, 1647 m, 28.01.1970: I 3 42.2 mm
(USNM 283669).
Chile. "Challenger": sin 300, Juan Fernandez Is. 33°42'S. 78°18'W, 2516 m, 17.12.1875: 1 2 20.0 mm (NHM
1888.22).
"Albatross": sin 2788, 45°35'S. 75°55'W, 1921 m, 11.02.1888: 3 3 19.3-41.1 mm; 2 ov. 2 59.6. 66.9 mm
(USNM).
Wallis & Futuna Is area. MUSORSTOM 7. SW of Combe Bank: sin 621, 12°35.0'S. 178°11.5'W, 1300-1280 m,
28.05.1992: I 2 26.8 mm (MNHM-Pa 1734), — Sin 623, 12°34'S. 178°15'W. 1280-1300 m. 28.05.1992: 1 juv.
(MNHM-Pa 1735).
Vanuatu. Musorstom: 8: stn CP 1111, I4°5I.09'S. 167°14'E, 1210-1250 m. 8.10.1994: 1 3 31.6 mm (MNHN-Pa
1674).
New Caledonia. BlOCAL: stn CP 60. 24°01'S, 167°08'E. 1480-1530 m. 2.09.1985: 1 juv. (MNHN-Pa 1736).
Halipro 2: stn BT 22, 24°09'S, 167°47'E, 1349-1350 m, 21.11.1996: 1 ov. 2 58.4 mm (MNHN). — Sin BT 95,
24°00'S. 162°08'E. 1224-1233 m. 25.11.1996: 1 damaged (MNHN-Pa 1705). — Sin BT 103. 24°54'S. 162°09'E. 1235-
1256 m, 26.11.1996: 1 3 53.2 mm (MNHN-Pa 1706). — Stn BT 105. 24°45'S. 162°50’E. 1200-1218 m. 27.11.1996:
I 3 56.7 mm; 1 2 61.9 mm (MNHN-Pa 1707).
New Zealand. NZOI: stn P971, 41 °1 1 9S, 177°19.6'E. 2200-2328 m, 18.06.1980: 1 3 39.5 mm (NIWA). —
Stn P941 , 4 1 0 1 5.2’S. 167°07.2'E, 1463-1457 m, 23.04.1980: 1 2 37.4 mm; 1 juv. (NIWA). — Sin X188. 36°46.30'S.
177°15.06'E, 1500 m, 4.12.1989: 1 2 damaged (NIWA). — Stn U198, 35°59.3'S. 1 62° 1 1.21E. 1573 m, 26.09.1982:
1 juv. (NIWA). — Stn Z8391 , 44°21.72'S, 177°52.90'E, 1143-1145 m, 10.10.1995: I 2 48.9 mm (NIWA).
Australia. Tasmania. 34°28’S, 132°03'E, 1175 m, 11.01.1984: 1 ov. 2 75.1 mm (TM G3279). — 4I°53.6'S.
144°24.6'E. 1384-1416 m, 18.05.1986: I 2 45.8 mm (TM G3482). — 42°22.4'S. 144°37.9'E. 1376-1404 m.
8.01.1988: 1 d 39.9 mm; 1 2 39.2 mm (TM G3978).
Australian Bight. "Galathea": stn 554, 37°28'S, I38°55'E. 1350 m. 12.05.1951: 1 3 38.5 mm (ZMC).
Indian Ocean. East Indian Ridge. "Prof. Mesyatsev": 31°55'S, 87°54'E, 1645-1600 m, 30.03.1979. coll.
M. Karpinskl 1 2 49.1 mm (ZMMU).
Types. — The large female (82 mm cl) (MCZ 4041) collected by the "Blake" stn 236. off Bequia (Antilles.
Windward Islands), 2909 m (BOUVIER, 1925: 437). is the holotype of Pentaclteles validus A. Milne Edwards,
1880.
The male (20.0 mm cl) (USNM 7145) collected by the "Albatross" stn 2074, off New England (4I°43'N,
65°21'50"W, 2393 m), is designated here as the lectotype of Pentaclteles debilis Smith, 1884: the male (19.5 mm
cl) (USNM 7146) collected at stn 2084 (40°16.50'N, 67°05.15'W, 2359m) is the paralectotype.
The female (20 mm cl) (NHM 1888.22) collected by the "Challenger" stn 300. off Juan Fernandez I.(33°42'S.
78°18'W, 2516 m), is the holotype of Polycheles chilensis Sund, 1920.
The female (66.9 mm cl) (NHM 1928.12.1.337) collected off Cape Point Lighthouse, 1024-1280 m. South
Africa, is the holotype of Polycheles demani Stcbbing, 1917.
The juvenile collected by the "Talisman" stn 39, (MNHN-Pa 15) off the Canaries (30°08'N. I4°02’W. 2300 m).
is designated here as the lectotype of Polycheles armatus Bouvier. 1905; the specimens from stns 41 (MNHN-
Pa 16) (30°0I'N, 14°06'W, 21 15 m) and 43 (MNHN-Pa 14)(29°52’N. 14°04'W, 2075 m) are paralectotypes.
Description. — Carapace ovate. Dorsal surface of carapace setose, minutely granulose. Anterior margin
concave, rostrum bifid, antrorse. Internal and external angles of orbital sinus irregularly spinose (0-2), sometime
differing between left and right sides. Eyestalk bearing upeurved spine. Basal antennular segment lamellar, produced
anteriorly to a sharp point, prominently spinose along its mesial edge, single spine on antero-external angle. Spine
formula of lateral margins of carapace 7-9:4:20-32, spines posterior to postcervical incision successively decreasing
310
B. GALIL
in size. Median postrostral carina with antrorse spinules, pair of larger spines 2/3 way to cervical groove; median
postcervical carina with two files of antrorse tubercles, pair of larger spines one third distance to posterior margin.
Gastro-orbital region bearing two (sometimes three) prominent spines anteriorly, some six spinules posteriorly.
Posterior margin of cervical groove spinose. Branchial carina obsolescent. Carapace between median postcervical
and branchial carinae with conical tubercles, between branchial carina and lateral margin minutely granulate.
Posterior margin of carapace smooth.
Abdominal tergiles 1-5 bearing blunt median carina, 1-3 armed anteriorly with obtuse tooth, sixth tergite
noncarinate. Tergites 2-5 with deep, transversely oblique, submedian grooves. Pleura smooth. Telson with
triangulate basal tubercule, smooth submedian carinae convergent posteriorly.
Subterminal claw-like spine on upper margin of merus and propodus of PI . Carpus, 3/4 as long as merus, with
small subterminal spine on lower margin. P5 subchelate in male, chelate in female.
Fig. 10. — Peniacheles validus A. Milne Edwards, 1880, <3 49.3 mm, Bahama Is, 24°39'N. 76°26.54'W. 1624 m (USNM
154399): a. dorsal view of the cephalothorax; b, lateral view of the abdomen.
Remarks. A. Milne Edwards (1880: 65) described P. validus as having "Lcs cinq premiers articles dc
1 abdomen portent en dessus une carenc obtuse terminee en avant par une pointe mousse; de cette pointc part de
chaque cote un sillon profond dirige en arriere et en dehors". Similarly, Smith (1884: 361) described his two
immature males ot P. debilis: "The first live somites of the pleon are perceptibly, though very inconspicuously,
carinated, and on the second, third, and fourth somites there is a narrow sulcus each side, extending from near the
carina outward and backward", and readily admitted "it is probably most nearly allied to P. validus A. M. Edwards".
Bouvier (1905c: 4), whose type series consisted of juvenile specimens only, distinguished the east Atlantic
var. armatus from P. debilis by its "armature spiniforme sur le bord externe du sinus orbitaire", an unstable
character. By 1917, BOUVIER himself synonymized both debilis and var. armatus with P. validus , and in 1925 (: 4)
Source : MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
31 I
wrote: "Les specimens recucillis par le Talisman et la Princesse Alice m'ont permis d'etablir. avec une entiere
certitude, que les excmplaires captures par l’Albatross et decrits sous le nom de P. debilis Smith ne soni rien autre
chose quc des P. validus de mediocre ou de petite taille". Examination of the single juvenile specimen from
"Challenger" stn 300, assigned to laevis by Bate (1888: 145) and then to P. chilensis by SUND (1920: 226).
revealed it to be P. validus.
Distribution. — Worldwide: Bay of Biscay, Canary Is. Azores, West Africa, South Africa.. S. Georgia I.,
Mid Atlantic Bight, United States, Bahama Is, Caribbean Sea, Gulf of Mexico, Chile. Vanuatu. Wallis & Futuna
Is, New Caledonia, New Zealand. Tasmania, Australian Bight. East Indian Ridge; 914 m (present work) - 3365 m
(FlRTH & PEQUEGNAT. 1971).
POLYCHELES Heller, 1862
Polycheles Heller, 1862: 389; 1863: 209. Gender masculine.
Type species, by monotypy. Polycheles typhlops Heller. 1862: 392.
Name placed on the official list of Generic Names in Zoology (p. 153) in Opinion 519 of the International
Commission on Zoological Nomenclature.
Eryoneicus Bate, 1882: 456. Gender masculine.
Type species, by monotypy, Eryoneicus caecus Bate. 1882: 457.
Name suppressed by the International Commission on Zoological Nomenclature in Opinion 702 (1964. Bulletin of
Zoological Nomenclature, 21(2): 111, 112).
Stereomastis Bate. 1888: 154. Gender feminine.
Type species, designated by Holthuis. 1962 (Bull. Zool. Notnencl., (19): 183): Pentacheles suhmi Bate, 1878: 278.
Name validated under the plenary powers in Opinion 702 of the ICZN and placed on the Official List of Generic Names
in Zoology (p. 1 12).
DIAGNOSIS. — Carapace dorsoventrally flattened, subrectangular. Anterolateral angle of carapace produced,
spiniform. Lateral margins well defined, spinose; cervical and postcervical incisions dividing margin into three
parts. One or two rostral spines. Orbital sinus simple or compound, eyestalk filling orbit, continuing distad,
beneath anterolateral angle of carapace.
Cervical groove marked, arcuate, laterally bifurcate. Postrostral and postcervical carinae well defined, granulate,
spinulate. Gastro-orbital, posterior branchial carinae well defined. Sublateral carina starting near posterolateral
margin of carapace, running to base of antenna and First cheliped.
Abdomen laterally depressed. First abdominal tergite narrow; pleura fused, abbreviated. Abdominal tergites 2-5
sculptured, medially carinate; sixth tergite medially bicarinate. Second abdominal pleuron clypeiform; pleura 3-5
Ungulate, successively diminishing in size posteriorly. Abdominal sternites with median knob. Tclson lanceolate,
with median basal tubercle, two submedian carinae. Uropodal exopod lamellar, rounded; endopod oar-shaped, basal
lappet on interior margin.
Mesial margin of basal antcnnular segment anteriorly produced, upeurved, reaching beyond peduncle; basal
antennular segment proximally rounded, its anterolateral angle uni- or bispinose. Outer antennular flagellum short,
inner flagellum as long as antennal flagellum. Renal process well developed, tubular, fitting into hollow on
underside of basal antennular segment. Antennal scaphoccritc lanceolate, extending to base of flagellum.
Buccal opening subquadrate. Mandible concave, cutting edge with triangular teeth of uneven size; mandibular
palp biarticulate. Maxillula with two incurved lobes, distally setose. Maxilla with two slender lobes, internal lobe
half as long as lateral; large scaphognathite. First maxilliped endopod slender; exopodal lobe membraneous,
reniform, extending further back than scaphognathite; exopod anteriorly divided into two lobes enclosing efferent
passage. Second and third maxillipeds, slender, pediform; epipod on basal segment of third maxilliped rudimentary;
dactyl as long as preceding segment.
Pereiopods 1-4 chelate, P5 simple or subchelate in male, chelate in female. PI longer, more robust than P2-5;
pereiopods successively shorter posteriorly. Fingers of first and second chelipeds crossing, their inner margin
furnished with row of overlapping cornute platelets, forming serrate edge. Fingers of chelae of P3-4 slightly
312
B. GALIL
upcurved, their inner margins simple. First male pleopod spatuliform, spinulose. swelling distally on inner
margin; spinules distally on mesial margin; distal margin folded internally. Pleopods 2-6 biramose. with rod-like
appendix interna; second pair in male with appendix masculina.
Remarks. — "There seems to be some little doubt as to the right nomenclature of the various species of this
group" - an understated comment by Alcock (1901b; 165) that fails to convey the nomenclatural confusion in the
genus.
Heller (1862), described the fifth pereiopodal dactyl of the single male specimen of Polycheles typhlops as
simple. With the rush of excitement engendered by the findings of the "Challenger", an error was introduced when
Bate (1878a: 276) described Pentacheles as differing from Polycheles "in having the last pair of pereiopods always
more or less perfectly chelate". Norman (1878; 382) intuited "Has not my friend mistaken sexual for generic
characters?" but was brusquely rebuffed, though BATE (1878b: 484) admitted "I was hesitating where several
species of Pentacheles should be placed, as there is a regular gradation from the imperfect to the perfect chelate
character of the fifth pereiopod; but as I found Polycheles, both male and female, with simple non-chelate foot, at
present it appears to me that there is no arrangement so constant as that which 1 propose." This last statement was
patently false - Bate examined both male and female specimens of Polycheles baccatus, which fifth pereiopod is
simple in male, chelate in female and P. helleri, which fifth pereiopod is subchelate in male, chelate in female.
Faxon (1895: 1 18) recognized that chelation of the fifth pereiopod in this genus was both sex and age-dependent
and was gravely critical of Bate: "That author was most arbitrary in the disposal of his own species".
A decade later. Bate (1888: 154) erected Stereomastis: "This genus differs in nothing externally from
Pentacheles, but is established to receive those species in which the mastigobranchial lash does not exist".
ALCOCK (190 lb: 165) acknowledged the value of the third maxilliped epipod. recognizing Polycheles (rather
than Stereomastis) as having "epipodite of the external maxillipeds .. a mere papilla". Adding to the confusion,
ALCOCK kept shifting species from one genus to the other: phosphorus was placed first in Pentacheles (ALCOCK &
Anderson, 1894: 165), then in Polycheles (1901b: 168); hextii, placed in Pentacheles (1901b: 171). was found
to be synonymous with Polycheles typhlops.
No wonder then that BALSS (1925: 16) wrote in consternation: "Alle Unterschiede generischer Art. die man
zwischen einzelnen Formen hat frnden wollen, sind. wie sich herausgestelt hat, so geringfugiger Natur und zudem
noch durch Ubergange verbunden, das man am besten alle zu den obigen drei Gattungen gerechneten Arten unler
dem altesen Namen Polycheles zusammenfast".
Eryoneicus Bate 1882, early presumed "a young and immature form of some species allied to Polycheles"
(Bate, 1888: 125). Yet, although "In all essential characteristics" resembling Polycheles (ALCOCK, 1901b: 176).
it was considered a separate genus. Faxon (1895: 108), on examining the eight specimens collected by the
•' Albatross ” wrote "The existence of well developed external sexual organs... at once disposes of the theory that
Eryonicus is an immature stage in the development of some other genus". SUND (1915: 372) believed the close
correspondence in carapace spine lormula "evidence of the Eryonicus species being in fact larval Polycheles" .
Gordon (1960) recognized 37 "species" of Eiyoneicus, many unrelated to adult polychelids. Following the ruling
ot the ICZN to suppress Eiyoneicus (ICZN 702), it is used to denote polychelid larval stages, though DE Saint
Laurent ( 1 979) maintains that they are adults secondarily adapted to life in the bathypelagial.
Polycheles differs irom the other polychelid genera in having only a rudimentary epipod on the basal segment
of 'fie 'f’lrcf maxilliped. It is distinguished from Willemoesia and Cardus in having the fifth pereiopodal dactyl
simple or subchelate in the male, chelate in the female, and from Pentacheles and Homeryon in having the basal
antennular segment proximally rounded.
Polycheles aculeatus sp. nov.
Fig. 11
Stereomastis phosphorus - GRIFFIN & Stoddart, 1995: 246. figs 9-1 1 (p.p.).
Material examined. — Vanuatu. Musorstom 8: stn CC 992. I8°52.34'S.
1994: 1 juv. (MNHN-Pa 1425). — Stn CC 994, 18°47.72'S, 168°17'W. 649-641 m.
168°55.I6'W. 775-748 m. 24.09.
24.09.1994: 4 3 20.6-44.6 mm
Source : MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
313
(MNHN-Pa 1426). — Stn CC 996, 18°52.41'S, I68°55.73’W, 764-786 m, 24.09.1994: 2 <5 33.2. 34.4 mm; 1 2
34.7 mm (MNHN-Pa 1427). — Stn CC 1033, 17°54.75'S, 168°40.66'W, 650-691 m, 29.09.1994: 3 2 23 0-47 7 mm
(MNHN-Pa 1428). — Stn CC 1056, 16°33.H'S, 167°55.64'W. 602-620 m, 1. 10.1994: 2 <3 46.6, 53.9 mm; 2 2 52 5
57.7 mm (MNHN-Pa 1429). — Sin CC 1073, 15°45.70'S, 167°22.24'W. 630-650 m, 4.10.1994: 2 <3 33.1, 42.5 mm;
1 2 48.9 mm; 1 juv. (MNHN-Pa 1430). — Stn CC 1124, 15°0I.72'S, 166°56.51'W, 532-599 m, 9 10 1994' 1 2’
49.2 mm; 2 juvs (MNHN-Pa 1431).
Loyalty Islands. MUSORSTOM 6: stn DW 426, 20°24.60'S, 166°22.90'E. 610 m. 17.02.1989: 1 3 22 5 mm
(MNHN-Pa 1434).
New Caledonia. Bathus 1: stn CP 663, 20°58.66'S, 165°38.27'E, 730-780 m. 13.03.1993: 2 2 28.3, 5 1 5 mm
(MNHN-Pa 1435). — Stn CP 702, 20°55.97'S, 165°34.67'E, 591-660 m, 18.03.1993: 1 2 69.8 mm (MNHN-Pa 1436)
Bathus 2: stn CP 741, 22°35.53'S, 166°26.56'E, 700-950 m. 14.05.1993: 2 <3 48.3, 49.1 mm; 2 2 45.5. 57.0 mm.
2 juvs (MNHN-Pa 1437). — Stn CP 743, 22°35.6'S, 166°26.2'E. 713-950 m, 14.05.1993: 3 <3 23.7-28.6 mm: 1 2
40.6 mm, 1 juv. (MNHN-Pa 1438). — Stn CP 751, 22°24’S. 166°12’E. 1300-1500 m, 15.05.1993: 1 3 42.4 mm; I 2
46.3 mm, (MNHN-Pa 1604). — Stn DW 753, 22°35.6'S. 166°26.2'E. 144-155 m, 15.05.1993: 1 3 45.6 mm (MNHN-Pa
1423). — Stn CP 765, 22°09.6’S. 166°02.8’E. 600-630 m, 17.05.1993: I <3 43.0 mm; 2 2 28.4, 61.8 mm (MNHN-Pa
1439). — Stn CP 766, 22°10'S. 166°01.7'E, 650-724 m, 17.05.1993: 3 <3 25.9-32.6 mm; 3 2 24.7-33.8 mm, I juv
(MNHN-Pa 1440). — Stn CP 771, 22°09.52'S. 166°0I.75'E. 610-800 m, 18.05.1993: 2 <3 24.7. 48.3 mm; 1 2
60.9 mm; 1 juv. (MNHN-Pa 1441).
Bathus 4: stn CP 899, 20°16.68'S, 163°50.26'E, 500-600 m. 3.08.1994: 1 2 19.4 mm (MNHN-Pa 1442) —
Stn CP 910, 1 8°59.32'S, 163°08.47'E, 560-608 m, 5.08.1994: 1 2 20.3 mm (MNHN-Pa 1443). — Stn CP 912.
1 8°55.61'S. 163°07.68'E, 702-690 m, 5.08.1994: 1 2 19.9 mm (MNHN-Pa 1444). — Stn CP 948. 20°33.13'S.
1 64°57.03’E, 533-610 m, 10.08.1994: 1 2 52.2 mm (MNHN-Pa 1445). — Stn CP 949. 20°32.01'S. 164°56.85'E. 616-
690 m, 10.08.1994: 1 <3 42.9 mm (MNHN-Pa 1446). — Stn CP 950. 20°3I.93'S. I64°56.1 1'E. 705-750 m, 10.08.1994:
3 3 29.7-46.7 mm; 1 2 54.9 mm (MNHN-Pa 1468).
Halipro 1: stn CP 854. 21°45.77'S, I66°38.34,E. 650-780 m, 19.03.1994: 1 juv. (MNHN).
Chesterfield Islands. MUSORSTOM 5: stn 324, 21°1 5.01S, I57°51.33'E, 970 m, 14.10.1986: 3 juvs (MNHN-Pa
1432). — Stn CC 384, I9°42.40'S. 158°50.80'E, 21.10.1986: 1 juv. (MNHN-Pa 1433).
Indonesia. "Siboga": stn 297. Timor Sea. 10°39'S. 123°40'E, 520 m. 27.01.1900: 1 2 27.7 m (ZMA).
Off south coast of Bali, 08°34'S, 1 15°36'E, 27.06.1981: 1 2 62.0 mm (USNM 356448).
Karubar. Tanimbar I. : stn CC 39, 7°47'S, 132°26'E, 477-466 m, 28.10.1991: 2 <3 39.7, 45.8 mm; 5 2 43.3-
58.7 mm (MNHN-Pa 1447). — Stn CC 40, 7°46'S, 132°31'E, 443-468 m, 28.10.1991: 2 <3 38.7. 50.5 mm: 4 2 34.9-
63.0 mm (MNHN-Pa 1448). — Stn CC 41. 7°45'S, 132°42'E. 401-393 m, 28.10.1991: 2 <3 39.7. 45.8 mm; 5 2 43.3-
58.7 mm (MNHN-Pa 1449). — Stn CC 42, 7°53'S. I32°42’E. 354-350 m. 28.10.1991: 1 2 56.5 mm (MNHN-Pa 1450).
— Stn CC 56, 8°16'S, 131°59'E, 552-549 m, 31.10.1991: 2 2 43.5, 44.0 mm (MNHN-Pa 1451). — Sin CP 69. 8°42'S.
1 3 1°53'E, 358-368 m, 2.11.1991: 3 <3 25.1-35.1 mm; 2 2 44.8-64.0 mm (MNHN-Pa 1452). — Stn CP 72. 8°36'S.
131°33'E, 699-676 m. 2.11.1991: 1 2 58.3 mm (MNHN-Pa 1453). — Stn CP 76. 8°50'S. 131°33'E. 401-400 m,
3.11.1991: 2 <3 31.3, 36.4 mm; 4 juvs (MNHN-Pa 1455). — Stn CP 78. 9°06'S. 132°4'E. 295-284 m, 3.11.1991: 5 3
18.0-28.0 mm; 4 2 19.4-26.6 mm (MNHN-Pa 1456).
Kai /.: stn CP 9. 5°23'S, 132°29,E, 368-389 m, 23.10.1991: 4 <3 26.6-46.8 mm (MNHN-Pa 1457). — Stn CC 10.
5°21'S, 1 32°30'E, 329-389 m. 23.10.1991: 5 <3 36.6-48.6 mm; 4 2 27.6-62.2 mm (MNHN-Pa 1458). — Sin CP 12.
5°23'S, 1 32°37'E, 436-413 m. 23.10.1991: 1 <3 42.2 mm; 1 2 47.5 mm (MNHN-Pa 1459). — Stn CP 35. 6°08'S.
132°45'E. 390-502 m. 27.10.1991: 4 <5 27.9- 43.4 mm; 1 2 37.0 mm (MNHN-Pa 1460).
East China Sea. "Ryoan Man,": 29°44'N, 127°48'E. 495-510 m. 19.03.1978, id. Y. Miya as Stereomastis
andatnanensis: 1 2 57.3 mm (NUM Cr 10239).
Australia. North West Coast. N.N.W. of Cape Leveque, "Solea": 13°07.2'S, 123°15.7'E. 400 m, 15.02.1984:
1 ov. 2 60.3 mm (WAM C23588). — N.W. Collier Bay. "Solea": 13°44'S, I22°13.3'E, 496-494 m. 13.02.1984: 1 ov. 2
51.2 mm (WAM C23515). —N.W. York Sound, "Solea": I2°48.1'S. 122°56.7'E. 496-504 m, 15.02.1984: 1 8 48.1 mm
(WAM C23514). —N.W. of Beagle Bay, "Solea": 15°12.8'S. 121°05.9'E. 410-404 m. 11.02.1984: 2 3 38.9, 38.7 mm;
1 ov. 2 54.9 mm (WAM C23587). — W. of Lacepede Arch., "Solea": 16°55.2'S, 1I9°52.8'E. 432-430 m, 20.02.1984:
2 3 52.4, 52.2 mm; 1 2 56.2 mm (WAM C23526). — 16°56.8'S. !I9°51.0'E. 480-432 m, 20.02.1984: 2 <3 48.8.
45.7mm (WAM C23525). — 14°50.2'S, 12I°31.4'E, 356 m. 12.02.1984: I ov. 2 64.4 mm (WAM C235I7).
— 16°56'S. 1 19°53.0'E, 436 m, 22.02.1984: I 2 50.7 mm (WAM C23528). — Off Port Hedland. "Solea": 16°46.1'S.
1 19°44'E. 495-502 m. 19.08.1983: I <3 58.5 mm; 1 3 juv. 31.7 mm (WAM C23596). — 17°55'S. I18°22.0'E. 437-
442 m. 21.08.1983: 1 2 51.9 mm (WAM C23521). —"Solea": 18°29.I'S. 117°42.5'E. 352-356 m. 18.08.1983: 1 3
51.6 mm (TM G3777), — 18°34.1'S, 1I7°27.5'E. 400-402 m, 16.08.1983: I 3 48.8 mm (TM G3776). — Off Port
Hedland, "Solea": 17°42'S. II8°38'E, 402-404 m. 18.08.1983: I 2 53.7 mm (WAM C23519). — I7°42'S. 1 1 8°38 E.
402-404 m. 18.08.1983: 1 3 44.1 mm (WAM C23520).
Types. — The holotypc is the male (45.6 mm cl) (MNHN-Pa 1423) collected in New Caledonia. Bathus 2
stn DW 753 (22°35.6’S, 166°26.2'E, 144-155 m). Two males and one female (MNHN-Pa 1427) collected in
Source
314
B. GALIL
Vanuatu. MUSORSTOM 8 stn CC 996 (18°52.41’S, 168°55.73'W. 764-786 m), an ovigerous female (WAM
C23588) collected in Australia (NW coast), and a female (USNM 356448) collected in Indonesia off Bali, arc
paratypes.
Fig. 11. — Polycheles aculeatus sp. nov., <3 42.8 mm, Vanuatu. MUSORSTOM 8, stn 994. I8°47.72'S. 168°17'W, 649-
641 m (MNHN-Pa 1426): a. dorsal view of the cephalothorax; b, lateral view of the abdomen.
Description. — Lateral margins of carapace subparallel, slightly convergent anteriorly. Dorsal surface of
carapace tomentose. Anterior margin of carapace straight. Median submarginal tooth prominent, upeurved. fronting
small, bifid rostrum. Internal angle of orbital sinus bearing small spine, external angle unarmed. Eyestalk with
mammilate tubercle, curved distad. Basal antennular segment produced anteriorly to a sharp point, mesial margin
smooth; antero-external margin angled, bispinose, anterior spine longer than posterior. Spines on lateral margins
of carapace decreasing successively in size posteriorly; spine formula 6-7:3:8-10. Spine formula of median
postrostral carina 1:1:2: 1; postcervical carina with two pairs of spines. Posterior margin of cervical groove with
antrorse spine midway between median and branchial carinae. Gastro-orbital region bearing five spines in curved
row, anterior and posterior spines largest. Branchial carina sinuous, bearing 6 or 7 spinules, posteriormost largest.
Posterior margin of carapace bearing two prominent, antrorse submedian spines. Anterior margin of first tergile
with two pairs of lateral spines.
Abdominal tergites medially carinate; carinae 1-4 with antrorse spines; third and fourth prominent, salient; fifth
carina bicuspid; sixth bearing denticulate double carina, confluent posteriorly in somewhat larger tooth. Tergites 2-
5 with oblique submedian grooves obsolescent. Pleura 2-5 with curved rib mesially, margins denticulate. Second
pleuron anteriorly produced, triangulate, prominently spinose. Telson with short, denticulate crest anteriorly; two
minutely serrulate, submedian carinae posteriorly.
Merus of PI with mostly two (1-3) spines proximally, subterminal claw-like spine on upper margin,
occasional spinule on lower margin. Carpus more than half as long as merus, with prominent subterminal spines
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
315
on upper and lower margins. Upper margin of propodus granulate, spinose, subtcrminal spine long; lower marsin
with two minutely granulate rows. P5 subchelate in male, chelate in female.
Etymology. — The Latin aculeatus (- sharp, pointed) refers to the prominently spinose anterior margin of
the second abdominal pleuron.
Remarks. — P. aculeatus is distinguished from the closely allied auriculatus and surdus in having the second
pleuron anteriorly triangulate, and a single antrorse spine midway between the median postcervical and the
branchial carinae on the posterior margin of the cervical groove.
According to GRIFFIN & STODDART (1995: 247) "The east Australian specimens differ from Alcock's original
descriptions of 5. phosphorus in having two spines on the antero-external angle of the basal antennular segment
instead of one, and nine to thirteen spines on the lateral edges of the carapace (behind the cervical groove) instead of
six or seven.... also differ in having a conical tooth, quite large in some specimens, projecting from the frontal
wall of the carapace but concealed by dense setae". The accompanying photograph (fig. 11) clearly shows the
anteriorly triangulate second pleuron typical of P. aculeatus.
Distribution. — Vanuatu, New Caledonia, Loyalty Is, Chesterfield Is, Indonesia, Australia (NW Coast). East
China Sea; 144 m (present work) - 1053 m (Griffin & Stoddart. 1995).
Polycheles auriculatus (Bate, 1878)
Fig. 12
Pentacheles auriculatus Bate. 1878a: 280; 1878b: 484: 1878c: 563.
Stereomastis auriculata - Bate, 1888: 159. — De Man. 1916: 4 (list). — Sund, 1920: 224. — Bernard. 1953- 87 —
Firth & Pequegnat, 1971: 64. — Dawson. 1997: 17.
Pentocheles auriculata - Bate, 1888, pi. 16 figs 3-4 [erroneous spelling],
Polycheles auriculatus - BouviER, 1917: 35 (list).
Stereomastis phosphorus - GRIFFIN & STODDART, 1995: 246, figs 9-11 (p.p.).
Material EXAMINED. — Marquesas Islands. Eiao. MUSORSTOM 9: stn CP 1278. 7°52'S 140°39'W 1000 m
5.09.1997: 3 juvs (MNHN).
Fiji Islands. "Challenger": stn 174c. Kandavu 1., 19°07,50'S, 178°19.35'E. 1116 m. 3.08.1874: 1 <? 21 7 mm
(NHM 1888.22).
Musorstom 10: stn CP 1309, 17°32,05'S, 178°53.24'E, 830 m. 5.08.1998: 2 <3 25.0. 39.8 mm; 3 juvs (MNHN)
Vanuatu. MUSORSTOM 8: stn CC 991, 1 8°5 1 .26S, 168°52,19'W. 936-910 m, 24.09.1994- 1 $ 53 7 mm (MNHN-
Pa 1424). — Stn CC 1124, 15°01.72'S, 166056.5I'W. 532-599 m, 9.10.1994: 1 juv. (MNHM-Pa 1680). - Stn 1125.
1 5°57.63'S, 166°38.43'W, 1160-1220 m. 10.10.1994: 1 juv. (MNHN-Pa 1461). — Stn CP 1126 1 juv (MNHN-Pa
1462).
New Caledonia. BATHUS 1: stn CP 651, 21°41.8'S, 166°40. 1 E. 1080-1180 m. 11. 03.1993: I 5 20 6 mm
(MNHN-Pa 1464). — Stn CP 661, 21°05'S, 1 65°50.05'E. 960-1100 m. 13.03.1993: 2 3 20.5. 20 9 mm- 1 $ 17 3 mm
(MNHN-Pa 1465).
BATHUS 2: stn CP 751, 22°24.35'S. 166°12.83'E, 1300-1500 m. 15.05.1993: 1 2 24.8 mm (MNHN-Pa 1466). —
Stn CP 767, 22°10'S, 1 65°59'E. 1060-1450 m, 17.05.1993: 2 3 24.3, 31.2 mm; 3 2 19.7-22.7 mm (MNHN-Pa 1467).
Philippines. "Albatross": stn 5467, Lagonoy Gulf. San Bernardino Str., 13°35'27''N, 123°37'18"E. 864 m
18.06.1909: I 2 23.4 mm (USNM). - Stn 5468, 13°35'30"N, !23°40'28"E. 1024 m, 18.06.1909: 2 3 31.5. 40.2 mm;
1 2 29.8 mm (USNM).
Australia. Torres Sir. "Franklin". 10°37.!7'S. 144°21.99'E, 990-1053 m, 21.08.1988. id. H E. STODDART as
Stereomastis phosphorus: 13 31.6 mm (AM P44920).
North West coast. " Solea ": N.VV. of Port Hedland, 18°42'S. 116°21'E. 694-704 m. 5.04.1982: 1 3 45 3 mm (WAM
C23583). - 18°40'S, 116°27'E, 720-724 m. 5.04.1982: 1 <3 44.2 mm; 1 2 31.7 mm (WAM C13435). - I8°45'S.
1 1 6°26.5'E, 590-592 m, 4.04.1982: 2 2 52.6, 54.1 mm (WAM C13434). — 18°41'S 116°29 5'E 696-704 m
6.05.1982: 1 <3 47.7 mm (WAM Cl 3433).
"Southern Surveyor": Exmoulh Plateau, 20°55.4’S, !12°51.5'E, 1139-1128 m ?3 ] 1991 1 2 ^8 4 mm
(TMG3779). — 20°16.5'S, 1 13°13.5 E. 913 m, 23.1.1991: 2 ov. 2 47.9. 51.2 mm (TM G3778).
West coast. "Southern Surveyor": W. of Cape Farquhar, 23°44.6'S. 112°35.5'E. 612-620 m. 26.1.1991: 1 2 55.0 mm
(TM G3780). — W, of Freycinet Estuary, 26°40.4'S. 1 12°32.7'E, 478-456 m. 30.1.1991: I 2 45.1 mm (TM G3843).
316
B GALIL
Western Australian Basin. "Lira": 22°08'S, I13°02'E. 880 m. 7.05.1973. coll. O. Petrov: 2 2 51.4, 53.2 mm
(ZMMU). — 20°10.3'S, 1 12°58. 1'E, 880-862 m. 7.05.1973. coll. O Pf.TROV: 2 2 56.4. 38.3 mm (ZMMU).
Type. — The male (21.7 mm cl) (NHM 1888.22) collected by the "Challenger" stn 174c. off Kandavu I., Fiji.
(19°07.50'S. 178°19.35'E, 1 1 16 m) is the holotype.
Fig. 12. — Polycheles auriculatus (Bate. 1878), 2 51.4 mm. West Australian Basin. "Lira". 22°08'S. 1 13°02'E. 880 m
(ZMMU): a, dorsal view of the cephalothorax; b. lateral view of the abdomen.
DESCRIPTION. — Carapace oblong, lateral margins subparallel, slightly convergent anteriorly. Dorsal surface of
carapace tomentosc. Anterior margin of carapace straight. Median submarginal tooth upeurved, in front of small,
bifid rostrum. Internal angle of orbital sinus bearing small spine, external angle unarmed. Eyestalk with
mammilate spine, curved distad. Basal antennular segment produced anteriorly to a sharp point, mesial margin
smooth; antero-external margin bispinose, anterior spine twice as long as posterior. Lateral spines decreasing
successively in size posteriorly; spine formula 6-7:3:7. Spine formula of median postrostral carina 1: 1:2:1;
postcervical carina with two pairs of spines. Posterior margin of cervical groove with 3 or 4 antrorse spines
midway between median postcervical and branchial carinae. Gastro-orbital region bearing five spines in curved row,
posteriormost spine largest. Branchial carina sinuous, bearing 10 spinules, posteriormost largest. Carapace
between branchial and median postcervical carinae minutely granulate. Posterior margin of carapace bearing two
prominent, antrorse. submedian spines.
Anterior margin of first abdominal tergile bearing two pairs of lateral spines. Abdominal tergites medially
carinate; carinae ol tergites 1-4 bearing antrorse spine; spine of third tergite elongate, salient, longer than the one
of fourth; carina of fifth tergite simple; sixth tergite bearing prominent, denticulate, lyre-shaped carina. Tergites 2-
5 with marked oblique submedian grooves. Pleura 2-5 with curved rib mesially. Anterior pleuron anteriorly
produced, ovate. Tclson with short, denticulate, crest anteriorly, two minutely serrulate submedian carinae
posteriorly.
Source MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
317
Merus of PI bispinose proximally, with subterminal claw-like spine on upper margin. Carpus, more than half
as long as merus, with prominent subterminal spines on upper and lower margins. Upper margin of propodus
smooth, except for a subterminal spine; lower margin with two minutely granulate rows. P5 subchelate in male,
chelate in female.
Remarks. — P. auriculatus differs from the closely allied P. surdus in having the lateral margins of carapace
posterior to postcervical incision bearing 7 spines rather than 10-14; the branchial carinac with 10 spines rather
than 5 or 6; the submedian grooves on the abdominal tergites well marked rather than obsolescent; and the sixth
abdominal tergite bearing prominent rather than obsolescent carinae.
The single specimen collected by the "Challenger" was a juvenile male. SUND (1920: 224) was mistaken in
describing the holotype as female. Comparing the type specimen with Bate's drawing, a discrepancy is apparent in
the form of the second pleuron and the pleural mid rib. Bate (1878a: 161) dcscribed^the "Coxal plates ridged with
markings like small cars," or (1888: 161) as "an elevated line or ridge that somewhat resembles in form the outline
of the human ear" however, the rib appears in the illustration bow-like (pi. 16. fig. 4).
DISTRIBUTION. — Marquesas Is, Fiji Is, Vanuatu, New Caledonia, Philippines, Australia. Western Australian
Basin. 532-1500 m.
Polycheles haccatus Bate, 1878
Fig. 13
Polycheles haccatus Bate, 1878a: 278; 1878b: 484; 1878c: 563. — De Man, 1916: 26. pi. 1 figs 4. 4a. — Bouvier.
1917: 35 (list). — Gunther & Deckert, 1950: 6. — Bernard. 1953: 86. — Firth & Pi quegnat 1971- 41
Griffin & Stoddart, 1995: 238.
Polycheles haccata - Bate, 1888: 131, fig. 32. pi. 14 fig. 1. — Stebbing, 1893: 200. — Hickson. 1894- 136 fig jg
— Sund, 1920: 226. '
Polycheles buccatus - Zarf.nkov. 1983: 110, fig. 46 [erroneous spelling).
nol Polycheles haccatus - Chan & Yu. 1989: 168. pi. 1 figs c-d; 1993: 109 [= P. coccifer sp. nov.].
Material EXAMINED. — Wallis and Futuna Islands area. MUSORSTOM 7. Wallis Is: sin DW 606 13°?1'S
176°08'W. 420-430 m. 22,05.1992: 1 2 31.8 mm (MNHN-Pa 1469).
Banc Tuscarosa : stn DW 555. 11°47'S. 178°19'W. 540-542 m. 19.05.1992: 1 d 26.6 mm (MNHN-Pa 1470) —
StnCP 559. 1 1 °48'S. 178°19'W, 547-552 m. 19.05.1992: 5 d 16.6-30.0 mm; 1 $ 36.2 mm (MNHN-Pa 1471)
Banc Bayonnaise: stn DW 625, 11°52'S, 179°34'W. 425-430 m. 29.05.1992: 1 3 31.5 mm (MNHN-Pa 1473) _
Stn DW 627, 11°54'S, 179°31'W, 597-600 m, 29.05.1992: 5 m 17.3-21.4 mm. 6 $ 19.0-25.0 mm; 9 juvs (MNHN-Pa
1473). - Stn CP 629. 1 U54'S. 179°32'W, 400-420 m, 29.05.1992: 2 d 17.6, 21.5 mm; 2 juvs (MNHN-Pa 1474) -
Stn CP 630, 1 1 °54'S, 179°32'W, 500 m, 29.05.1992: 1 d 23.7 mm (MNHN-Pa 1475). — Stn CP 631. 11°54'S
179°32'W, 600 m. 29.05.1992: 2 3 16.8, 19.0 mm; 1 $ 21.3 mm (MNHN-Pa 1476). — Stn CP 632, II°54'S.
179°31'W. 595-600 m. 29.05.1992: 1 3 19.4 mm; 4 9 21.1-31.9 mm (MNHN-Pa 1477).
Fiji Islands. Maluku I. "Challenger": stn 173. 19°09.35’S, 179°41.50'E, 567-576 m, 24.07 1874- 3 3 17 5-
27.8 mm; 3 2 16.9-29.3 mm (NHM 1888.22).
Vanuatu. Musorstom 8: sin DW 1014, 17°54.53’S. 168°19.08'E. 495-498 m. 27.09.1994: 1 d 28 8 mm (MNHN-
Pa 1478). — Stn CP 1089, 15°08.82'S. 167°17.23'E, 494-516 m, 6.10.1994: 1 juv. (MNHN-Pa 1479) — Stn CP 1111
14°5 1 ,09'S, I67°14.00'E. 1210-1250 m, 8.10.1994: 1 3 28.6 mm (MNHN-Pa 1480).
New Caledonia. Musorstom 4: stn CP 169. I8054.03'S, I63°l 1.20'E. 575-600 m. 17.09.1985: 3 3 15.9-
24.6 mm; 1 2 24.2 mm, 3 juvs (MNHN-Pa 1481). — Stn 200. 18°53.8’S, 163°14.1'E. 535 m 20 09 1985- 1 2
25.5 mm (MNHN-Pa 1482). — Stn 202, 18°58'S. 163°59.3'E, 560 m, 20.09.1985: 4 3 246-3? 7 mnr 3 2 32 9-
38.8 mm (MNHN-Pa 1483). — Stn 236. 22°11.35’S, 167°15'E. 495-550 m. 2.10.1985: 1 2 36.4 mm (MNHN-Pa 1484)
Halipro 1: stn CP 872. 23°02'S, 166°52'E. 620-702 m, 30.03.1994: 1 2 29.0 mm (MNHN-Pa 1488)
Bathus 4: stn 909, 18°57.64'S, 163°10.30'E, 516-558 m, 4.08.1994: 2 2 33.2, 37.0 mm (MNHN-Pa 1489) —
Stn 911. 1 8°57.80'S. 163°08.47'E, 566-558 m, 5.08.1994: 1 d 33.3 mm (MNHN-Pa 1490) — Stn 921 I8°46 7?'S
163°17.81'E, 613-610 m. 6.08.1994: 1 2 29.4 mm (MNHN-Pa 1491).
Chesterfield Islands. Musorstom 5: stn 383, 19°40.85'S. 158°46.10'E. 615-600 m 21 10 1986' 3 3 iuvs
17.5-20.7 mm (MNHN-Pa 1486).
Australia. New South Wales. "Kapala": off Newcastle. 360-540 m, 27-29.04.1971: 1 ov, 2 35.9 mm (WAM
C 1 3430). — Stn K77-13-12, 29°52'S. 153°43'E. 495 m. 23.08.1977, id. D.E. Brown: 1 3 34.8 mm (AM P26549). —
Stn K78-17-21, 28°37'S, 153°50'E. 502 m. 19.08.1978. id. H.E. Stoddart: 1 2 33,5 mm (AM P44749).
318
B. GALIL
Philippines. Musorstom 1: stn CP 50. 13°49.2’N, I20°01.8'E, 415-510 m. 25.03.1976: 1 3 26.4 mm (MNHN-Pa
1492).
MUSORSTOM 2: stn CP 75, 13°50.5'N, 120°30.3'E. 300-330 m. 1.12.1980: 1 9 19.4 mm (MNHN-Pa 1493).
Musorstom 3: stn CP 118. 11°58'N. !2!°06'E. 448-466 m. 3.06.1985: 1 juv. (MNHN-Pa 1494).
Indonesia. " Siboga stn 38, Bali Sea. 7°35.4'S, 117°28.6'E, 730-915 m. 1.04.1899: 1 3 26.3 mm: 1 9 26.9 mm
(ZMA).
Karubar. Kai Is: stn CP 17. 5°15'S. 133°01’E. 459-439 m. 24.10.1991: 5 3 19.8-23.8 mm; 7 9 20.7-27,1 mm;
12 juvs (MNHN-Pa 1496). — Stn CP 25. 5°30'S, 132°52 E. 336-346 m. 26.10.1991: 1 9 35.5 mm (MNHN-Pa 1497). —
Stn CP 27, 5°03'S, 132°51'E, 304-314 m. 26.10.1991: 1 9 29.8 mm (MNHN-Pa 1498).
Tanimbar Is: stn CC 42, 7°52'S, 132°42’E, 354-350 m. 28.10.1991: 2 3 30.1. 33.0 mm; 3 9 29.8-32.4 mm (MNHN-
Pa 1564).
Types. — The male (27.8 mm cl ) (NHM 1888.22) collected by the "Challenger" stn 173. off Maluku I.. Fiji
(19°09.35'S. 179°41.50'E, 567-576 m), is the lectotype; all other specimens from that sample arc paralectolypes.
Fig. 13. — Polycheles baccatus Bate, 1878. 3 32.7 mm. New Caledonia. Musorstom 4. stn 202. 18°58'S. !63°59.3'E.
560 m (MNHN-Pa 1483): a. dorsal view of the ccphalothorax: b, lateral view of the abdomen.
Description. — Lateral margins of carapace parallel, slightly convergent anteriorly. Dorsal surface of carapace
densely granulose, setose. Frontal margin concave. Median submarginal tooth upcurved, prominent. Hanked by
two slender rostral spines. Internal angle of orbital sinus produced, triangulate; external angle pectinate, 6-spinose.
Eyestalk bearing vermicular tubercle. Basal antennular segment produced anteriorly to a sharp point, mesial margin
spinose, antero-external margin with one or more minute spinules. Spines on lateral margins of carapace
decreasing successively in size posteriorly; spine formula 10-12:4-6:25-26. Median postrostral carina irregularly
granulate; granules on postcervical carina larger, paired. Poslorbital carina obsolescent, granulate, bispinose.
Posterior margin of cervical groove between branchial carinae granulate. Branchial carina finely granulate. Posterior
margin of carapace bearing conic granules, submedian pair largest.
Source : MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
319
Abdominal tergites granulate. Tergites 2-5 bearing oblique submedian grooves. Anterior margin of first tergite
beaded. Tergites 1-4 bearing antrorse beaks; fifth carinate; sixth with submedially two beaded rows, confluent
posteriorly. Proximal half of pleura 2-5 granulate, margins smooth. Second pleuron medially scooped; margin
anteriorly produced, ovate. Pleura 3-5 with short, curved, granulate rib medially. Telson with granulate tubercle
anteriorly; two minutely serrulate, confluent carinae posteriorly.
Merus of PI minutely serrulate on upper and lower margins, with subterminal claw-like spine on upper
margin. Carpus hall as long as merus, with two serrulate rows on upper margin; spines on external row more
prominent, distal spines largest; spinule subterminally on lower margin. Upper margin of propodus with several
rows of antrorse conic granules; lower margin with two distal serrulate rows. P5 simple in male, chelate in female.
Color. — "Dorsal surface of the carapace and abdomen a rich burnt orange, area below lateral edges of carapace
and lower half of the abdominal pleura white. Interspaces between abdominal segments as well as grooves in
abdominal terga also white. Second to fifth pereopods white, chelae of second and third pereopods pinkish red.
Ischium of chcliped pinkish, colour continuing to proximal portion of merus; distal portion of merus, carpus and
propodus burnt orange; joints between merus and carpus, carpus and propodus and tips of finaers whitish"
(Griffin & Stoddart, 1995).
Remarks. — Bate's (1878a: 278) preliminary description, though brief and inaccurate, presents the essential
characters of P. baccatus: "armed with teeth on the inner side of the antero-lateral angle, central ridge projected into
a rostriform tooth supported by two small teeth;. ..Pleon carinated on each of the four anterior somites and
projected into an anteriorly pointed tooth". De Man's (1916) description is detailed and meticulous, pointing out
many of Bate's inaccuracies.
Chan & Yu (1989: 1 10) clearly describe another species IP. coccifer) which has the "anterolateral angles armed
with 2-3 strong spines... Abdominal pleuron II with anterior margin sharply pointed".
Distribution. — Wallis Is, Melanesia, Fiji Is. Vanuatu. New Caledonia. Australia (east coast). Philippines,
Indonesia. 300-1250 m.
Polycheles ceratus (Alcock. 1894)
Fig. 14
Pentacheles sp. Wood-Mason, 1891: 19.
Pentaclieles ceraia Alcock, 1894: 241. — Wood-Mason & ALCOCK. 1894. pi. 8 fig. 1.
Polycheles ceraia - ALCOCK, 1899: 33.
Polycheles ceratus - ALCOCK 1901b: 170. — BouviER, 1917: 35.
Stereomastis ceraia - De Man, 1916: 4 (list). — Bernard. 1953: 87. — Firth & Pequegnat. 1971: 64. — Dawson.
1 997: 1 8.
Pentacheles certa - Dawson, 1997: 74 [erroneous spelling].
Material EXAMINED. — Indonesia. Sumatra. "Te Vega": stn 96, 3°25'N. 95°40'E, 1150 m. 21.11.1963: 1 $
33.5 mm (R.V1NH).
Andaman Sea. " Investigator ": stn 114. 13°21'N, 93°27'E. 1686 m: I 9 32 mm (IM 6880/9) — Sin 310
13°29'30"N. 95°29'E, 1757 m. 5.11.1902: I 2 33 mm (IM).
Type. — The female (32 mm cl) (IM 6880/9) collected by the "Investigator" stn 114 in the Andaman Sea
(13°21'N, 93°27'E, 1686 m) is the holotype.
Description. — Lateral margins subparallel, slightly convergent anteriorly. Dorsal surface of carapace
smooth, nearly glabrous. Frontal margin straight. Rostrum short, bifid. Internal and external angles of orbital
sinus unarmed. Eyestalk without vermicular tubercle. Basal antennular segment with mesial margin smooth,
antero-external margin bispinose. Spine formula of lateral margin of carapace 5-6:2:3; lateral margins of carapace
posterior to postcervical incision smooth. Median postrostral carina set with antrorse spines; spine formula
1 : 1 :2: 1 . Pair of antrorse spines medially on posterior margin of cervical groove. No postorbital carina and spines.
320
B. GAUL
Small spine on posterior margin of cervical groove at junction of anterior and posterior branches. Branchial carina
distinct, unarmed. Posterior margin of carapace with pair of antrorse submedian spines.
Abdominal tergites smooth. Oblique submedian grooves on tergites 2-5 obsolescent. Anterior margin of first
tergite smooth. Tergites 2-5 medially crested; tergites 2-4 with small antrorse spine; fifth tergite with large, ram¬
like. antrorse spine reaching forward beyond fourth tergite. Sixth tergite bearing mesially parallel pair of rounded
carinae, confluent posteriorly. Pleura nearly smooth; second pleuron anteriorly produced, ovate. Telson with
obsolete tubercle anteriorly, two smooth confluent carinae posteriorly.
Merus of PI smooth. Carpus half as long as mcrus with subterminal spine on upper and lower margins. Upper
margin of propodus with two subterminal spines. P5 chelate in female.
Fig. 14. — Polycheles ceratus (Alcock, 1894), 2 33.5 mm. Sumatra, "Te Vega", stn 96, 3°25'N, 95°40'E. 1150 m
(RMNH): a, dorsal view of the cephalothorax; b. lateral view of the abdomen.
Remarks. — P. ceratus is easily distinguished from its congeners by the large antrorse spine on fifth
abdominal tergite, overhanging the anterior margin of the fourth tergite.
Distribution. — Known from Sumatra (Indonesia) and the Andaman Sea, 1 150-1757 m.
Polycheles coccifer sp. nov.
Fig. 15
Polycheles baccatus - Tung et ai. 1988: 47. pi. 46. — Chan & Yu, 1989: 168, pi. 1 figs c-d; 1993: 109 [non Bate,
1878].
Material EXAMINED. — Vanuatu. Musorstom 8: stn CP 1123, 15°07.19'S. 1 66°55.20'E, 262-352 m,
9.10.1994: 1 juv. (MNHN-Pa 1500).
Source : MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
321
New Caledonia. MUSORSTOM 4: stn 247, 22°09'S. 167°13.3'E, 435-460 m, 4.10.1985: 1 3 40.6 mm (MNHN-
Pa 1499).
H alipro I: stn CP 868, 21°14.53'S, 165°55.84'E, 430-450 m. 21.03.1994: I 3 26.1 mm (MNHN-Pa 1501).
Bathus 1: stn CP 698, 20°34.2'S, I64°57.3'E, 491-533 m. 17.03.1993: 2 3 34.5. 38.3 mm; 3 2 27.6-40 5 mm
(MNHN-Pa 1502).
Bathus 4: stn 947, 20°33.72'S. 164°57.72'E. 470-490 m, 10.08.1994: 3 2 30.2-33.5 mm; 4 juvs (MNHN-Pa
1503) . — Stn 948, 20°33.13'S, 164°57.03-E. 533-610 m. 10.08.1994: 1 3 39.7 mm; 1 2 40.1 mm; 1 juv. (MNHN-Pa
1504) .
Indonesia. Karubar. Kai Is: stn CP 16, 5°17'S, 132°50'E, 315-349 m. 24.10,1991: 1 2 44.6 mm (MNHN-
Pa 1733).
Tanimbar /.: stn CC 42, 7°53’S, 132°42'E, 354-350 m, 28.10.1991: 1 2 46.7 mm (MNHN-Pa 1511). — Sin CP 62,
9°0I'S, I32°42'E, 246-253 m, 1.11.1991: 3 3 39.8-44.6 mm; 2 2 37.1, 47.9 mm (MNHN-Pa 1512). — Sin CP 78.
9°06'S. I31°24'E, 295-284 m, 3.11.1991; 2 2 43.7, 45.8 mm (MNHN-Pa 1513). — Stn CP 79. 9°I6'S. ni°22'E 250-
239 m, 3.11.1991; 1 3 33.4 mm; 1 2 53.8 mm (MNHN-Pa 1514).
Philippines. " Albatross stn 5112, Luzon. 13°48'22,,N, 120°47'25"E. 319 m, 17.01.1908: 2 3 25.5. 33.5 mm:
2 2 34.6, 37.2 mm (USNM). — Stn 5118, Balayan Bay, 13°48'45"N, 12Q°41'5r'E, 284 m. 21.01.1908: 1 2 38.3 mm
(USNM). — Sin 5221, btw Marinduque and Luzon, 13°38'15"N. 121°48'15"E, 351 m. 24.04.1908: 3 3 28.2-40.1 mm;
2 2 30.1, 57.2 mm (USNM). — Stn 5222, btw Marinduque and Luzon, 13°38'30”N, 12i°42'45''E. 351 m. 24.04.1908:
1 2 21.5 mm (USNM). — Stn 5256, Illana Bay. Bongo I., 7°2r45"N, 124°07'15"E. 284 m. 22.05.1908: 1 3 22.7 mm;
2 2 25.2, 46.7 mm (USNM). — Stn 5289, Luzon, 13°41'50"N, 120°58'30"E. 310 m. 21.07.1908: I 2 30.1 mm
(USNM). — Stn 5297. Luzon. 13°41'20"N. 120°58'E. 356 m, 24.07.1908: 1 dry (USNM). — Stn 5364. Luzon. Balayan
Bay, 13°48'30"N, 120°43'45"E, 288 m. 20.02.1909: 10 3 19.8-34.5 mm; I 2 18.1-28.0 mm (USNM). — Stn 5365.
Luzon. Balayan Bay, 13°44'24"N, 1 20°45'30'’E. 385 m, 22.02.1909: I 2 56.1 mm (USNM). — Sin 5374. Marinduque I.,
13°45'45''N, 121°35'08"E, 342 m. 2.03.1909: 3 3 17.5-37.0 mm; 4 2 17.7-39.3 mm (USNM). — Sin 5421. btw Panay
and Guimaras, 10°33'30"N, 122°26'E, 247 m, 30.03.1909: 1 3 25.1 mm; 1 2 45.9 mm (USNM). — Luzon. Balayan Bay,
Calaca Town, 13°44'N, 120°45'E, 99-177 m, 24.06.1966: 1 3 41.9 mm; 1 2 30.9 mm (USNM 322873). —
Nouongeasto, 13°44'N, 120°45'E. 144-155 m. 25.06.1966: 2 3 26.8. 33.5 mm (USNM 322873). — S. of Sapating.
1 3°44'N, 120°45'E, 266-301 m. 29.07.1966: 2 3 27.3, 37.7 mm; I juv. (USNM 322873).
MUSORSTOM I: stn 42, 13°55.1'N. 120°28.6'E. 379-407 m, 24.03.1976: 1 2 damaged (MNHN-Pa 1505).
Musorstom 2: stn CP 26. 13°49.6’N, 120°51.0'E, 299-320 m, 23.11.1980: 8 3 23.6-33.4 mm; 7 2 25.1-48.2 mm
(MNHN-Pa 1506). — Stn CP 74. 13°54'N, I20°27'E, 300-370 m: 1 2 47.9 mm (MNHN-Pa 1507).
Musorstom 3: stn CP 105. !3°52'N, 120°30’E. 398-417 m. 1.06.1985: 2 3 20.0, 25.0 mm; I 2 50.4 mm; 1 juv.
(MNHN-Pa 1508). — Stn CP 119, 1 1°59'N, 121°13'E, 320-337 m, 3.06.1985: 1 3 36.3 mm (MNHN-Pa 1509). —
Stn CP 125. 1 1°57'N. 121°28'E. 388-404 m, 4.06.1985: 19 3 22.7-38.1 mm; 16 2 24.7-51.6 mm; 14 juvs (MNHN-
Pa 1510).
Taiwan. Tai chi. I-lan County, 300 m, 10.06.1993, coll. & id. T.Y. Chan as P. baccatus: 1 <5 34.3 mm (NTOU).
Type. — The male (36.3 mm cl) (MNHN-Pa 1509) collected by MUSORSTOM 3 at stn 119 ( 1 1 °59'N.
1 2 1 0 1 3'E, 320-337m) in the Philippines is the holotype.
DESCRIPTION. — Lateral margins of carapace parallel, slightly convergent anteriorly. Dorsal surface of carapace
densely granulose, tomentose. Frontal margin concave. Median submarginal tooth upeurved. prominent, reaching
considerably beyond pair of non contiguous rostral spines. Internal angle of orbital sinus produced, triangulate;
externa] angle bispinose, occasionally with auxiliary spinules. Eyestalk bearing vermicular tubercle. Basal
antennular segment produced anteriorly to a sharp point, mesial margin minutely serrulate, antero-external margin
tuberculate. Spine formula of lateral margins of carapace 6-7:4-6:20-23, spines decreasing in size posteriorly.
Median postrostral carina set with antrorse mammilate tubercles; postccrvical carina set with larger granules,
paired, anteriormost largest. Postorbital carina obsolescent, quadrispinose, anteriormost spine largest. Posterior
margin of cervical groove between branchial carinae granulate. Branchial carina serrulate. Posterior margin of
carapace granulate, with pair of antrorse submedian spines.
Abdominal tergites granulate; tergites 2-5 with transversely oblique submedian grooves. Anterior margin of
First tergite irregularly beaded. Tergites 1-4 bearing antrorse spines; fifth crested; sixth bearing submedially two
beaded rows, confluent posteriorly. Proximal 3/4 of pleura 2-5 granulate, margins granulate. Second pleuron
medially scooped, margin anteriorly triangulate. Pleura 3-5 with short, curved, granulate rib medially. Telson with
granulate tubercle anteriorly; two minutely serrulate, confluent carinae posteriorly.
Merus of PI spinulate on upper margin with longer proximal spines and subterminal claw-like spine, minutely
serrulate on lower margin. Carpus half as long as merus, with two minutely serrulate rows on upper margin and
Source :
322
B. GAL1L
sublerminal spinule on lower margin. Upper margin of propodus minutely granulate, tiny spinules on lower
margin. P5 simple in male, chelate in female.
Color. — "Body generally ivory and covered with large patches of orange-red on dorsal surface. Non-articulate
surface of dorsal abdomen, posterior parts of abdominal pleura, telson margins and bases of uropods also orange-
red. Setae light brown." (Chan & Yu, 1989: 109, color photo).
Fig. 15. — Polycheles coccifer sp. nov„ $ 53.8 mm, Indonesia, Tanimbar I., Karubar sin 79. 9°16'S. 131°22'E, 250-
239 m (MNHN-Pa 1514): a. dorsal view of the cephalothorax; b. lateral view of the abdomen.
ETYMOLOGY. — The Latin coccifer (- berried) refers to the densely granulose carapace of this species.
REMARKS. — P. coccifer is distinguished from its closely related congener, P. baccatus , in having the external
angle of orbital sinus bispinose, the lateral margins of carapace anterior to cervical incision bearing 6-7 spines, the
carpus and propodus of the first cheliped indistinctly granulose, the gastro-orbilal carina quadrispinose, and the
anterior margin of the second pleuron triangulate.
Distribution. — Vanuatu. New Caledonia, Taiwan, Indonesia, Philippines; known reliably between 155-
533 m, collected too in trawlings between 99-610 and 533-610 m.
Polycheles enthrix (Bate, 1878)
Fig. 16
Willemoesia eulhrix Willemoes-Suhm. 1875c: xxxiii. (see remarks).
Pentacheles enthrix Bate, 1878a: 280. pi. 13 figs 1-3; 1878b: 484; 1878c: 563. — Smith, 1880b: 272.
Pentacheles anthrax - SMITH, 1880a: 353 [erroneous spelling].
Source MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
323
Pemacheles euihrix - Bate, 1888: 149. figs 14-27. 33-36. — Stebbing, 1893: 200. — Faxon. 1895' 1 19 — Hutton
1904: 253. — Sund. 1920: 226.
Polycheles helleri Bate, 1878a: 277 (p.p.); 1878b: 484 (p.p.); 1878c: 563 (p.p.); 1888: 138 (p.p.).
Pentocheles enihrix - BATE. 1888, pi. 17 [erroneous spelling].
Pemacheles helleri - Bate, 1888, pi. 15 fig. 1 [non Bate. 1 878J.
Polycheles euihrix - De Man. 1916: 5 (list). — Bouvier, 1917: 35 (list). — Bernard. 1953: 86. — Firth & Pequegnat.
1971: 45. — Griffin & Stoddart, 1995: 239, figs 2-3. — Dawson. 1997: 9.
Polycheles enllirix - Chan & Yu, 1993: 107.
Stereomasiis kermadecensis Sund. 1920: 224. — Bernard, 1953: 87.
Polycheles amemiyai Yokoya. 1933: 44. fig. 23. — Miyake, 1972: 65; 1982: 190. — Takeda. 1982: 38 — Dawson
1997: 7.
Polycheles anaemayai - Firth & Pequegnat, 1971: 41. — Dawson. 1997: 78 [erroneous spelling],
Polycheles kermadecensis - Firth & Pequegnat, 1971: 49. — Dawson, 1997: 1 1.
Stereomasiis nana - Baba el ai, 1986 pi. 109 [non Smith, 1880].
MATERIAL EXAMINED. — Wallis and Futuna Is area. MUSORSTOM 7 . Banc Combe: stn DW 539, 1 2°27'S.
1 77°27'W, 700 m. 17.05.1992: 1 9 59.1 mm (MNHN-Pa 1515). — Stn CC 551, 12°15'S. 177°28'W. 791-795 m,
18.05.1992: 1 <3 20.1 mm (MNHN-Pa 1516). — Stn CC 552, 12°16'S. 177°28'W. 786-800 m, 18.05.1992: 1 3
21.7 mm (MNHN-Pa 1517). — Stn CC 553. 12°17'S. 177°28'W. 780-794 m. 18.05.1992: 1 9 31.3 mm (MNHN-Pa
1518). — Stn CC 554. 12°14'S, 177°28'W, 795-820 m, 18.05.1992: 1 9 25.7 mm (MNHN-Pa 1519).
Banc Bayonnaise: stn CP 631, 11°54'S. 179°32'W. 600 m. 29.05.1992: 1 9 54.1 mm (MNHN-Pa 1520). — Sin CP
632, 11°54'S. 179°31'W, 595-600 m, 29.05.1992: I <3 39.0 mm (MNHN-Pa 1521).
Fiji Islands. "Challenger": stn 173. off Matuku I.. 19°09.35'S. 179°41.50'E. 576 m, 24.07.1874: 1 9 27.4 mm
(NHM 1888.22).
New Caledonia. Biocal: stn CP 32, 23°07'S. 166°51'E. 825 m. 28.08.1985: 4 9 24.9-56.3 mm (MNHN-
Pa 1536).
Beryx 2: stn CP 58. 23°19.20'S, I67°59.35 E. 850-920 m. 22.10.1992: I 9 23.7 mm (MNHN-Pa 1537). —
Stn CP 59, 23°19.'S, I68°00'E. 750-800 m, 22.10.1992: 1 <3 26.2 mm; I 9 47.6 mm (MNHN-Pa 1538).
Bathus 1: stn CP 663, 20°58.66'S. 165°38.27'E, 730-780 m. 13.03.1993: I 9 54.5 mm (MNHN-Pa 1539) —
Stn CP 709, 21°41.78'S. 166°37.88'E, 650-800 m, 19.03.1993: 2 9 56.7, 59.5 mm (MNHN-Pa 1540).
Bathus 2: stn CP 741, 22°35.53'S, 166°26.56'E, 700-950 m, 14.05.1993: 1 9 42.8 mm (MNHN-Pa 1541). —
Sin CP 743, 22°35.6'S, 166°26.2'E, 713-950 m, 14.05.1993: 1 <5 20.0 mm; 2 9 57.2, 48.0 mm (MNHN-Pa 1542). —
Stn CP 765, 22°09.6'S. 166°02.8'E, 600-630 m. 17.05.1993: 1 9 46.6 mm (MNHN-Pa 1543). — Stn CP 766. 22°I0'S.
166°01.7'E, 650-724 m. 17.05.1993: 1 9 52.7 mm (MNHN-Pa 1679). — Stn CP 771. 22°09.52'S. 166°01 ,75'E. 610-
800 m, 18.05.1993: 1 9 47.3 mm (MNHN-Pa 1544).
Bathus 3: stn DW 790, 23°48.94'S, 169°47.60'E, 685-715 m. 25.11.1993: 1 9 29.7 mm (MNHN-Pa 1545). —
Stn DW 793. 23°47.50'S, 169°48.75'E. 731-751 m. 26.11.1993: 2 3 26.0-34.5 mm (MNHN-Pa 1546). — Stn DW 794.
23°48.35'S. 169°49.10'E. 751-755 m. 26.11.1993: 1 <3 34,1 mm (MNHN-Pa 1547). — Stn CP 831. 23°04'S. 166°56'E.
650-658 m. 30.11.1993: 6 <3 23.2-43.1 mm; 2 9 31.2, 45.0 mm (MNHN-Pa 1548). — Stn CP 832, 23°03'S. I66°54'E.
650-659 m, 30.11.1993: 2 9 50.7, 51.7 mm (MNHN-Pa 1549). — Stn CC 841, 23°03'S. 166°53'E, 640-680 m.
30.11.1993: 3 <3 26.4-47.7 mm; 7 9 22.0-52.5 mm (MNHN-Pa 1550). — Sin CP 845, 23°03.30'S. 166°59.29’E. 592-
622 m. 1.12.1993: 1 3 29.1 mm; 1 9 41.2 mm (MNHN-Pa 1551). — Stn CC 848. 23°02.50'S. 166°52.80'E. 680-
700 m, 1.12.1993: 5 3 28.6-46.9 mm; 10 9 40.0-58.5 mm (MNHN-Pa 1552).
Halipro 1: stn CP 854, 21°40'S. 166°38'E, 650-780 m, 19.03.1994: 2 9 59.1. 57.8 mm (MNHN-Pa 1554). —
Stn CH 872, 23°02'S, 166°52'E. 620-702 m, 30.03.1994: 9 3 28.9-42.7 mm; 10 9 36.3-57.8 mm (MNHN-Pa 1555).
— Stn CH 873, 23°01'S. 166°53'E, 640-680 m. 30.03.1994: 1 3 29.7 mm; 2 9 50.8, 57.2 mm (MNHN-Pa 1556). —
Stn CH 874, 23°05'S, 166°48'E, 708-830 m, 30.03.1994: 1 <3 45.7 mm; 1 9 52.1 mm (MNHN-Pa 1557). —
Stn CH 876. 23°10'S. 166°49'E. 870-1000 m. 31.03.1994: 1 3 37.5 mm (MNHN-Pa 1558).
Bathus 4: stn CP 913, 18°56.23 S, I63°04.86'E. 777-820 m, 5.08.1994: 1 <3 32.6 mm; I 9 43.2 mm (MNHN-
Pa 1553).
Loyalty Islands. MUSORSTOM 6: stn DW 483, 2I°19.80'S. 167°47.80'E, 600 m, 23.02.1989: 2 3 39.3. 44.5 mm
(MNHN-Pa 1535).
Chesterfield Islands. MUSORSTOM 5: stn DC 321, 21°20.40'S. 158°02.20'E, 1000 m. 14.10.1986: 1 juv.
12.2 mm (MNHN-Pa 1522). — Stn CP 323, 21°18.52'S. 157°57.62'E, 970 m, 14.10.1986: 2 3 31.4. 32.2 mm; 2 9
41.6, 48.4 mm; 35 juvs (MNHN-Pa 1523). — Stn CP 359, 19°39.00'S, 158°49.00'E. 700-720 m. 18.10.1986: 1 3
31.8 mm (MNHN-Pa 1524). — Stn CP 360. 19°36.40'S. 158°49.60'E. 770-810 m. 18.10.1986: 1 9 damaged: 2 juvs
(MNHN-Pa 1525). — Stn CP 363, 19°47.90'S, 158°44.30'E, 700-685 m, 19.10.1986: 1 9 47.3 mm (MNHN-Pa 1526).
— Stn CC 365. 19°42.82'S, 158°48.00'E, 710 m. 19.10.1986: 1 <3 37.2 mm; 3 9 53.2-65.1 mm (MNHN-Pa 1527). —
Stn CC 366, 19°45.40'S. 158°45.62’E, 650 m. 19.10.1986: 2 <3 35.2, 22.9 mm; 1 9 37.0 mm (MNHN-Pa 1528). —
Stn CC 367, 19°36.80'S. 158°53.20'E. 855-830 m, 19.10.1986: 1 <3 42.9 mm; 1 9 31.8 mm (MNHN-Pa 1529) —
324
B. GALIL
SinCC 383. I9°40.85'S. 158°46.10'E, 615-600 m, 21.10.1986: 2 2 46.7. 52.5 mm (MNHN-Pa 1530). — Sm CC 384.
19°42.40'S, 158°50.80'E, 772-756 m. 21.10.1986: 3 3 13.3-45.2 mm; 6 2 15.8-59.3 mm (MNHN-Pa 1531). —
Sin DC 385, 20°53.60'S. 160°49.40'E. 740-750 m, 22.10.1986: 1 3 23.2 mm (MNHN-Pa 1532). — Stn CP 386.
20°56.21'S, 1 60°5 1 . 1 2'E. 770-755 m. 22.10.1986: 2 juvs 14.6. 15.4 mm (MNHN-Pa 1533). — Sin CP 387.
20°53.41'S, 160°52.4I'E, 650-660 m. 22.10.1986: I juv. 14.4 mm (MNHN-Pa 1534).
Kermadec Is. " Challenger ": sin 170. 29°55’S. 178°14'E, 951 m. 14.07.1874: 1 2 15.3 mm (NHM 1888.22). —
Stn 170a, 29°45'S, 178°H'W. 1152 m. 14.07.1874: 1 2 22.0 mm. badly broken specimen (NHM 1888.22).
New Zealand. NZOI: sin K5804. 29°14.8'S, 177°49.6’W. 590-490 m. 22.07.1974: 2 juvs (NIWA). — Stn Z8989.
37°32.55'S. 1 76°48.6 1 'E. 550-577 m, 18.01.1998: 1 ov. 2 42.6 mm (NIWA). — Sm Z9263, 34°09.2'S. 162°51.9'E.
791 m, 18.09.1998: 1 3 36.2 mm (NIWA).
Australia. New South Wales. "Kapala": stn K78-09-05. 28°02'S. 153°59'E. 549 m. 2.06.1978. id. D.E. Brown as
P. euthrix : 1 2 73.2 mm (AM P44750). — Sin K78-23-08. 28°03'S. 154°04'E. 732 m. 6.1 1.1978. id. H.E. Stoddart as
P. euthrix : 2 3 61.4, 69.5 mm: I 2 51.2 mm (AM P44754).
East China Sea. "Yuryo-maru" (del. Y. Miya as Stereomastis nana): 28°27.20'N. 126°56.36'E, 320-340 m.
7.02.1978: I 2 62.4 mm (NUM cr. 10243). — 25°48'N, 124°25.5'E. 420-400 m. 14.09.1979: 1 2 57.5 mm (NUM
cr. 10406). — 28°31'N. 126°57'E. 310 m, 12.10.1979: 1 ov. 2 60.9 mm (NUM cr.10407).
Taiwan. Su-Aon, I-Lan County, 500 m, 10.04.1991, coll. T.Y. Chan: 1 2 25.4 mm (NTOU).
Japan.Tosa Bay. 16.01-14.02.1961, coll. K. Sakai: 1 3 45.7 mm; 1 2 49.3 mm (SMF 24647).
Types. — The female (15.3 mm cl) (NHM 1888.22) collected by the "Challenger" stn 170 off Kermadec Is
(29°55'S, 178°14'E, 951 m) is designated here as the lectotype of Pentacheles enthrix Bate. 1878. The female
(22.0 mm cl) (NHM 1888.22) collected by the "Challenger" off Kermadec Is (sin 170a) is a paralectotype.
About the types of S. kermadecensis, see remarks.
Description. — Margins of carapace subparallel, slightly convergent anteriorly. Dorsal surface of carapace
setose, minutely granulose, granules more prominent anteriorly. Frontal margin of carapace slightly convex;
median submarginal tooth half as long as rostral spines; several spines and spinulcs on either side of rostral spines.
Internal angle of orbital sinus spinose, external angle unarmed. Eyestalk bearing small mammilate spine, curved
distad. Basal antennular segment produced anteriorly to a sharp point, mesial margin serrulate, two spines on
antero-external angle, lateral spine smaller. Anterolateral angle of carapace prominently produced. Spine formula of
lateral margins of carapace 8-9:4-5:12-14. Median postrostral carina ill-defined anteriorly, spine formula 1 : 1 :2: 1 ;
postcervical carina with one pair of spines followed by two rows of mammilate tubercules. Gastro-orbital region
bearing single spine. Branchial carina sinuous, obsolescent, granulose. Posterior margin of carapace bearing three
pairs of antrorse spines, submedian pair largest.
Abdominal tergites without submedian grooves, medially carinate. Carina on tergites 1-4 with antrorse spine;
carina on filth tergitc with small tooth; sixth tergite crested distally. Anterior tergite with pair of lateral spines.
Pleura with crescentic mid-rib nearly obsolete, margins setose. Second pleuron anteriorly produced, triangulate,
lelson with mammilate tubercle anteriorly followed by deep furrow; two smooth, conlluent carinae posteriorly.
Merus of cheliped bispinose proximally on upper margin, clawed subterminal spine; lower margin serrulate,
proximal spinules longer. Carpus 3/4 as long as merus, upper margin distally granulate, small subterminal spine
on lower margin. Upper margin of chela with two serrulate rows, distalmost spine on inner row prominent, lower
margin with single serrulate row. P5 subchelate in male, pollex very short; chelate in female.
Color. — Carapace and abdomen red, first chela pale distally (S. O'Shea, pers. comm.). "Body generally
orange-red. Lateral surface and grooves on carapace, antennal and antennular flagella, pereiopods, ventral surface,
articulated surface and lateral margins of abdominal tergites slightly orange-red to somewhat whitish" (Chan &
Yu. 1993: 107, color photo). Color photo also in Baba, 1986, pi. 109.
Remarks. — During the "Challenger" expedition Willemoes-Suhm collected two juvenile female specimens
otl the Kermadec Is on July 14th. 1874. He described them in his notes (cited in Bate, 1888: 153) and referred to
them by name, Willemoesia euthrix, in his letter to VON SlEBOLD, sent September 1874 and published in 1875.
However, as he failed to include even the briefest description, the name could not be accepted. Bate (1878a).
working trom Willemoes-Suhm's notes, described the species under the name Pentacheles enthrix. Including, in
addition to the two Kermadec specimens, a male (in fact a female) specimen collected off Matuku I„ Fiji. A farther
complication arose as it seems that Bate (1878a), in his preliminary account of the Challenger Polychelids, pul
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
325
down the same female specimen from the Kermadec Islands as syntype of both Polycheles helleri and Pentdcheles
enthrix - actually depicting it (Bate, 1888, pi. 15 fig. 1). SUND (1920: 224) recognized that "the other specimens
(from stn 170, near Kermadec Islands, N. of New Zealand) is certainly of a species distinct from the type" [from
the type of Polycheles helleri ]. However the species SUND erected, Stereomastis kermadecensis , is a junior
synonym of P. entlirix. Bate's misplacement serves to explain Sund's (1920) confusion as to the number of
enthrix specimens available to Bate, of which, he reported, only two survived. However, all three syntypes of
P. enthrix are extant. The three are very young specimens, as attested by their small size and the fact that "The
posterior pair of pereiopoda is chelate, the dactylos and pollex being subequal in both sexes" (Bate. 1 888: 1 50) - a
condition existing in juveniles.
Yokoya's (1933) description and illustration leave no doubt that P. amemiyai is but a synonym of enthrix.
Unfortunately, a search in the Kitakyushu Museum for the type was unsuccessful (K. Baba, pers. com).
Distribution. — Melanesia, Fiji Is, Kermadec Is, New Caledonia, New Zealand. Australia, East China Sea.
Taiwan, Japan; 229 m (Griffin & Stoddart, 1995) - 1 152 m.
Fig. 16. — Polycheles entlirix (Bate, 1878), 9 46.7 mm. New Caledonia. MUSORSTOM 5: stn 383. 19°40.85'S.
158°46.10'E, 615-600 m (MNHN-Pa 1530): a. dorsal view of the cephalothorax; b. lateral view of the abdomen.
Polycheles evexus sp. nov.
Fig. 17
Polycheles nanus - Faxon, 1895: 121, pi. 33, figs 1, la-b [non Smith, 1884],
Stereomastis nana - WlCKSTEN, 1989: 304. — HENDR1CKX, 1995: 156 [non Smith, 1884],
Material EXAMINED. — Peru. "Anton Bruun stn 766. 04°10'S. 81°27'W. 1860-1815 m. 9.09.1966: 3 9 22.8-
23.9 mm (USNM).
326
B. GALIL
Chile. " Albatross stn 2788, Clionos Arch., 45°35'S. 75°55'W, 1890 m, 1 1.02.1883. id. M.J. Rathbun: 3 6 19.7-
21.9 mm (USNM 28350). — Stn 2789. Chiloe I.. 45°36'S, 75°28 W, 2416 m. 12.02.1883. id. W.L. Schmitt: I spec,
damaged (USNM 58644).
Types. — The male (21.9 mm cl) (USNM 28350) collected by the "Albatross", stn 2788 off Chile (45°35’S,
75°55'W, 1890 m), is the holotype; the other males from that sample are paratypes.
Fig. 17. — Polycheles evexus sp. nov. 9 22.8 m, Peru. "Anton Bruun", stn 766. 4°10S. 81°27'W. 1860-1815 m
(USNM): a. dorsal view of the cephalothorax: b. lateral view of the abdomen.
Description. — Dorsal surface of carapace setose. Anterior margin straight, rostrum bifid. Internal angle of
orbital sinus with slender spine, external angle unarmed. Eyestalk bearing small spine, curved distad. Basal
antennular segment produced anteriorly to a sharp point, mesial margin smooth, anterolateral margin bispinose,
posterior spine half as long as anterior. Spines on lateral margins of carapace upeurved, anteriormost largest.
Lateral spine formula 5-6:3:6-7, spines further apart posteriorly. Spine formula of median postrostral carina
1: 1:2:1; postcervical carina with two pairs of spines. Gastro-orbital region with crescentic row of five prominent
antrorse spines. Posterior margin of cervical groove with antrorse spine between median and branchial carinae.
Branchial carina prominent, subparallel to lateral margin, with five spines. Posterior margin of branchial groove
spinose; several antrorse spines posterior to groove. Posterior margin of carapace spinose, submedian spines
largest.
Abdominal tergites medially carinate. Carinae on tergites 1-5 bearing antrorse spine; spine of tergites 3-5
prominent; sixth tergite bearing mesially pair of obsoletely granulate carinae, joined posteriorly in a low tubercle.
First tergite bearing two pairs of spinules distally on anterior margin. Tergites 2-5 with transversely oblique
submedian grooves. Pleura 2-4 with curved rib mesially, margins minutely granulate. Second pleuron ovate.
Telson with obsolescent median crest anteriorly, two convergent carinae posteriorly.
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
327
Merus ol PI proximally unispinose, subterminal claw-like spine on upper margin, finely granulose on lower
margin, carpus hall as long as merus with subterminal spines on upper and lower margins; upper margin of
propodus with subterminal spine. P5 subchelate in male, chelate in female.
Etymology. — The Latin evexus (= rounded at the top) refers to the blunter, less prominent lubercles on the
sixth abdominal tergite and telson.
Remarks. — Faxon (1895: 121) correctly observed that his Pacific specimens "differ slightly, but
constantly, from the type specimens from the Atlantic. The edges of the sulcated carina on the sixth abdominal
somite are less prominently denticulated and the tubercle at the posterior end of this carina is lower. Further, the
spine near the base ol the dorsal surface of the telson of the type specimens is reduced to a blunt tubercle in the
Pacific examples." However, he failed to recognize them as a distinct species.
Distribution. — Eastern Pacific. Mexico, Peru, Chile; 1544 m (Wicksten. 1989) - 2418 m (Faxon
1895).
Polycheles helleri Bate, 1878
Fig. 18
Polycheles helleri Bate, 1878a: 277 (p.p.); 1878b: 484 (p.p.); 1878c: 563 (p.p.); 1888: 138, pi. 14 fig 2 (pp) —
Pagenstecher. 1879: 63. — Sund. 1920: 224. — Hutton. 1904: 253.
S tereomastis helleri - De Man. 1916: 4 (list). — Bernard. 1953: 87. — Firth & Pequegnat. 197L 65 — Griffin &
Stoddart. 1995: 245. — Dawson, 1997: 18.
Siereomaslis nana - Baba, 1986: 157 [not Smith. 1884J.
Material EXAMINED. — Japan. "Albatross": stn 4957. Kagoshima Gulf. 32°36'N. 132°23'E, 787 in.
23.08.1905: 1 d 23.2 mm (USNM). — Stn 4958. Kagoshima Gulf, 32°32'N. 132°25'E. 1296 m. 23.08.1905: 1 d
20.2 mm (USNM). — Sin 4969, Wakayama, 33°23'40"N. 135°33'E. 1057 m, 29.08.1905: 3 d 19.1-24 3 mm- I 5 ">4 9
mm (USNM). — Stn 4971. Wakayama, 33°23'30"N, 135°34'E. 1168 m. 30.08.1905: 1 d 23.5 mm; 1 2 ->8 2 mnv 4
juvs (USNM). — Sm 4972, Wakayama, 33°25'45"N, 135°33'E. 792 m, 30.08.1905: 1 d 21.5 mm: 1 2 35 9 mm
(USNM). — Stn 4973, Wakayama, 33°24'15"N, I35°30’30"E, 1080 m, 30.08.1905: 2 d 28.2, 31.2 mm: 2 2 ?9 5 4^ 5
mm (USNM). — Stn 4975, Wakayama. 33°21‘30"N. 135°38'50"E. 1282-981 m, 31.08.1905: 1 d 22.5 mm (USNM). —
Stn 4976, Wakayama, 33°22'50"N, 135°38'30"E, 981 m. 31.08.1905: I d 22.6 mm (USNM). — Sm 4977, Wakayama,
33°23'N, 135°37'40"E. 979 m. 31,08.1905: I 2 18.5 mm (USNM). — Stn 5082. 34°05'N, 137°59'E, 1192 m.
20.10.1905: 1 d 28.3 mm; 1 2 34.8 mm: 1 ov. 2 45.6 mm (USNM).
NE Honshu I.. Iwate Pref., 39°10.3'N, 142°25'E. 980-1000 m. 19.09.1979. det. K. Baba as Siereomaslis nana: 1 2
41.3 mm (NFU 530-2-1498).
Locality uncertain. 1000 m, 26.01.1975, coll. Toriyama, del. K. Baba as Stereomaslis nana: 1 2 40.5 mm ((NFU
530-2-1497).
Indonesia. "Albatross": stn 5601. Sulawesi, lo13'10"N, 1 25° 1 7-05"E, 1399 m, 13.11.1909- 1 ov 2 35 4 mm
(USNM).
New Guinea. "Challenger": stn 218, 2°33'S, 144°04'E, 1957 m, 1.03.1875: 1 d 20.8 mm damaged (NHM
1888.22).
New Caledonia. Biogeocal: sin CP 260, 21°00.00'S, 166°58.34'E. 1820-1980 m. 17.04.1987: 2 2 24.0 mm:
1 damaged (MNHN-Pa 1749).
Australia. Coral Sea. "Franklin": ll°33.02'S. 145°19.34'E. 1611-1584 m. 22.08.1988, id. H.E. Stoddart as
Siereomaslis helleri: Id 18.7 mm (AM P40360).
Madagascar. "Vauban": stn CH 126, 17°50'S. 43°07'E. 1475-1530 m, 16.01.1975: I d 27 9 mm (MNHN-Pa
1559). — Stn CH 127. 17°50'S, 43°07'E. 1715-1750 m. 16.01.1975: 1 2 36.7 mm (MNHN-Pa 1560). — Stn CH 131.
1 3°46'S, 47°33'E, 1490-1600 m. 20.01.1975: 1 d, 22.8 mm; I juv. (MNHN-Pa 1561). — Stn CH 132 I3°43 8'S
47°29’E, 1950-2150 m, 20.01.1975: 1 d 25.7 mm (MNHN-Pa 1562).
I YPE. — I he male (20.8 mm cl) (NHM 1888.22) collected by the "Challenger" north of New Guinea is the
holotype.
Description. — Lateral margins of carapace subparallel, convergent anteriorly. Dorsal surface of carapace
sparsely setose, minutely granulate. Anterior margin of carapace straight, bearing bifid rostrum. Internal, external
328
B. GAUL
angles of orbital sinus unarmed. Eyestalk with mammilate spine curved distad. Basal antennular segment with
smooth mesial margin, bispinose antero-cxternal angle. Spine formula of lateral margins of carapace 6:3:8-9,
spines diminish in size posteriorly to form indistinct serrulation. Spine formula of median postrostral carina
1: 1:2:1; postcervical carina with two paired spines. Posterior margin of cervical groove with spine midway
between median and branchial carinae. Gastro-orbital region bearing five spines: anterior and posterior spines
largest. Branchial carina sinuous, well defined, prominently spinose: posteriormost spine largest. Posterior margin
of carapace bearing submedian spines.
Abdominal tergites without oblique submedian grooves, medially carinate; carinae on tergites 2-5 bearing
antrorse spine, increasing in size posteriorly; fifth carina noticably larger; margins of double carina on sixth tergite
smooth, confluent posteriorly. Anterior margin of first tergite with pair of lateral spinules. Pleura 3-5 with
obsolescent rib mesially; second pleuron anteriorly ovate, medially faintly depressed, sparsely denticulate. Telson
with two smooth, confluent carinae posteriorly.
Merus of PI bispinose proximally on upper margin, lower margin smooth; carpus, half as long as merus, dis-
tally with claw-like spine on upper and lower margins; upper margin of propodus smooth except for sublerminal
spine, lower margin with single row of minute granules distally. P5 subchelate in male, chelate in female.
Fig. 18. Polyclieles helleri Bate, 1878, 2 36.7 mm. Madagascar, stn 127. 17°50'S, 43°07'E. 1715-1750 m (MNHN-Pa
1560): a. dorsal view of the cephalothorax; b. lateral view of the abdomen.
Remarks. Bate (1888: 141) wrote "I can detect no distinction of sufficient importance to separate
Pentacheles sculptus from Polyclieles helleri, nor would it have taken much consideration to decide their identity
had it not been lor the generic character of the fifth pair of pereiopoda, and that the specimens were procured from
localities so widely apart". He was mistaken in all three points: P. helleri is easily distinguished from sculptus in
lacking a spine on the internal orbital margin; both helleri and sculptus have the fifth the pereiopod subchelate in
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHEL1DAE
329
the male, chelate in the female - Bate was mistaken because he examined immature specimens; and lastly, both
sculptus and helleri are present in the Indo-Pacific Ocean.
GRIFFIN & Stoddart (1995: 246) thought helleri "very similar to S. nana (Smith. 1884)... especially in the
absence of a spine on the inner orbital angle". However, P. nanus bears a spine on the inner orbilal angle, though
it was ommited by Smith (1884), and is further distinguished by possesing a prominent spine anteriorly on
branchial groove, and having the carapace spinulose between the median postcervical and branchial carinae.
Distribution. — Japan, New Caledonia. Australia, New Guinea. Indonesia. Madagascar; 787-2150 m.
Polycheles nanus (Smith, 1884)
Fig. 19
Pentacheles nanus Smith. 1884: 359; 1886a: 188; 1886b: 607. 612, 619, 651. pi. 7 figs I. la. — Verrill, 1885: 554.
— Fowler, 1912: 569.
Pentacheles andamanensis Alcock. 1894: 239. — AlCOCK & ANDERSON. 1895 pi. 10 fig. 3.
Polycheles nanus - Caullery, 1896: 386. pi. 16 fig. 12. — Koehler. 1896: 723. — Bouvier, 1905a: 480; 1905c: 5:
1915: 2; 1917: 35. — Hansen, 1908: 42. — Stebbing. 1908: 27; 1910: 377. — Murray & Hjort. 1912: 539. —
Stephensen. 1912: 578; 1913: II. — Selbie. 1914: 21. — Sund. 1915: 372. — Williamson. 1915: 457. — Jensen.
1928: 24. — Miranda & Rivera, 1933: 17. — Heegaard, 1941: 20, fig. 6. — Le Danois. 1948: 255. — Kramp.
1963: 57. — Allen, 1967: 55. — Thiriot, 1974: 344 (list).
Polycheles andamanensis - ALCOCK, 1899: 33; 1901b: 169. — Bouvier. 1905a: 481; 1905c: 5; 1917: 35. — Ramadan.
1938: 124. — Sewell, 1955: 205. — Dong et al.. 1986: 203.
Polycheles grimaldii Bouvier, 1905a: 481; 1905c: 4; 1917: 52, pi. 3 figs 2-4. — Boas, 1939: 4. pis 2,4.5.6.9. —
Danois. 1948: 255.
Polycheles nanus var. grimaldii - Selbie, 1914: 21. pi. I figs 14-15. — Richard. 1907: 322.
Stereomastis nana - De Man. 1916: 4 (list). — Calman, 1925: 19. — Barnard, 1950: 573, fig. 105e. — Bernard.
1953: 87. — Dahl, 1953: 46. — Kensley. 1968: 293; 1981b: 29. — Firth & Pequecnat, 1971: 65. — Baez &
Andrade. 1979: 226. — Wenner, 1979: 436. — Wenner & Boesch, 1979: 131. — George, 1983: 16. — De Saint
Laurent, 1985: 474. — Squires, 1991: 354 (key). — Takeda & Hanamura. 1994: 32. — Dawson. 1997: 19.
Stereomastis grimaldii - De Man, 1916: 4 (list). — STEPHENSEN, 1923: 67. — Zariquiey. 1968: 211. — Beaubrun.
1979: 42. figs 22-23. — Macpherson. 1991: 405. fig. 7.
Stereomastis nanus - STEBBING, 1917: 30. — ABELE & Felgenhauer, 1982: 31 1 [non Smith. 1884].
Stereomastis grimaldii - THIRIOT, 1974: 344 (list).
Stereomastis andamanesis - Ohta, 1983: 230 (list) [erroneus spelling] [non Alcock. 1895],
Stereomastis andamanensis - De Man, 1916: 16. pi. 1 figs 2, 2b (p.p.). — Dahl. 1953: 46. — HOLTHUIS. 1984: 4. —
Griffin & Stoddart. 1995: 244 (p.p.). — Dawson, 1997: 17 (p.p.) [non Alcock, 1895].
not Polycheles nanus - Faxon, 1895: 121. pi. 33 figs 1, la-b [= P. evexus sp. nov.].
not Polycheles grimaldi - Wolff, 1967: 236 (photo) [= P. talismani (Bouvier. 1917)].
not Stereomastis nana - Miyake, 1982: 78. pi. 26 fig. 6 [= P. typhlops Heller. 1862].
not Stereomastis nana - Baba et ah. 1986: 284 [= P. helleri Bate. 1878].
not Stereomastis nana - Baba et al.. 1986, pi. 109 [= P. enthrix (Bate. 1878)].
not Stereomastis nana - WlCKSTEN. 1989: 304 [= P. evexus sp. nov.].
not Stereomastis nana - HENDRICKX. 1995: 156 [= P. evexus sp. nov.].
Material EXAMINED. — Bay of Biscay. "Thalassa": stn 333, 44°10.5'N, 4°32.'W. 1900-1950 m, 14.10.1971:
1 juv. (MNHN-Pa 346). — Stn 453, 48°34'N, 10°51.6'W. 1975-2070 m: 1 6 20.8 mm; 2 2 21.9. 19.4 mm (MNHN-Pa
352).
Biogas IV: stn CV 33, 44°04.6'N, 4°18.1'W, 1913 m. Feb. 1974: 3 $ 25.7-26.3 mm (MNHN-Pa 644).
Biogas XL CP 37: stn 1, 47°33.80'N. 8°40.5'W. 2175 m. 11.10.1981. coll. IFREMER: 1 2 damaged (MNHN-Pa
1138).
Western Sahara. -Meteor": stn M36-I00 AT 152, 21°27.rN. 18°I6.rW. 2049-2110 m. 26.02.1975: 2 spec.
(SMF 24646).
West Africa. Senegal. "Talisman'1: stn 98. 17°16'N. 20°17'W. 2324 m. 17.07.1883: 1 3 damaged (MNHN-Pa 22).
— Stn 100. 17°16'N, 19°19'W, 1550 m. 17.07.1883: 1 2 35.1 mm (MNHN-Pa 21 ).
Ivory Coast. "Pills bury stn 233. 5°19'N. 4°I4'E, 2013-1464 m. 12.05.1965: 1 2 49.4 mm (RMNH).
Azores. "Talisman": stn 698. 39°I1'N, 33°04'W. 1846 m. 18.07.1883: 1 <5 18.6 mm (MNHN-Pa 23). — Stn 863.
39°19'N. 29°15'W. 1940 m, 1.08.1897: 1 2 21.7 mm (MNHN-Pa 24).
330
B. GALIL
Cancap 5: stn 5.016. 37°21'N. 25°29'W. 2000-2100 m, 26.05.1981: 5 <5 19.5-20.7 mm; 4 9 19.8-20.0 mm
(RMNH). — Stn 5.123, 38°56'N. 27°44'W, 2300-2400 m. 4.06.1981: 1 6 29.8 mm (RMNH). — Stn 5.162. 39°21'N,
30°51’W. 900 m, 11.06.1981: 1 juv. (RMNH).
"Atlantis": cr. 152, stn 20, 37°50.5'N, 26°00'W, 2560 m. 18.08.1948. coll. L.W. Hutchins, id. F.A. Chace as
P. grimaldii: 1 3 22.0 mm (USNM 88693).
SE Atlantic. Off Cape Province. "Africana": 32°26'S. 16°H'E. 1302 m. 27.01.1993: 2 ov. $ 26.1. 27.1 mm
(SAM). — 32°22'S. 1 6°09'E. 1150 m, 14.01.1995: 3 $ 23.2-27.7 mm (SAM).
Canada. "Albatross": stn 2706, Nova Scotia, 41°28’05"N, 65°35'05''W. 2138 m. 27.08.1886: 2 3 17.7. 23.1 mm;
3 2 19.1-22.7 mm (USNM 11887).
United States. "Albatross": sin 2102, 38°44'N. 72°38'W. 2212 m. 5.11.1883: I <? 22.8 mm (USNM 5714). -
Stn 2035. 39°26'12"N, 70°02'37"W, 2490 m, 17.07.1883: 1 2 damaged (USNM 7142). — Stn 2077, 41°09'40"N,
66°02'W, 2296 m, 4.09.1883: 3 <5 19.0-22.3 mm (USNM 7143). — Stn 2102. 38°44'N, 72°38'W. 2212 m, 5.1 1.1883:
1 3 22.6 mm; 1 2 22.5 mm (USNM 5714). — Stn 2103. 38°47'20"N. 72°37’00"W, 1964 m. 5.1 1.1883: 2 juvs (AMNH
15329). — Stn 2105. 37°50'N. 73°03'50"W, 2551 m, 6.11.1883: 1 2 18.7 mm (USNM 5711). — Stn 2106.
37o41'20"N. 73°03'20"W. 2838 m, 6.11.1883: 1 2 19.0 mm (USNM 5710). — Stn 2115. 35°49'30"N. 74CI34'45''W.
1541 m. 11.11.1883: 1 3 20.4 mm; 12 19.5 mm (USNM 5709). — Sin 2116, 35°45'23"N. 74°3i'25"W. 1624 m.
11.11.1883: 3 9 19.0-35.6 mm (USNM 5708). — Stn 2205. 39°35'00"N, 71°18'45"W, 1931 m. 20.08.1884: 2 ov. 2
27.8. 27.6 mm (AMNH 15239); 2 2 23.5, 26.7 mm (USNM 8237). — Stn 2206. off New Jersey. 39o35'00”N.
71°24'30"W, 1877 m. 20.08.1884: I ov. 2 29.5 mm; 1 juv. (USNM 8238). — Stn 2230. 38°27'00"N, 73°02'00"W.
2102 m, 12.09.1884: 1 2 23.2 mm (AMNH 15589). — Stn 2533. 40°16'30"N. 67°26'15"W. 1490 m, 15.07.1885: 1 ov.
2 31.3 mm (AMNH 15176). — Stn 2728, 36°30’N, 74°33'W, 1546 m. 25.10.1886: 1 ov. 2 31.8 mm (AMNH 15104).
— Stn 2729, 36°36'N, 74°32’W. 1222 m, 25.10.1886: 1 2 22.1 mm (AMNH 15104). — Stn 2731. 36°45'N, 74°28'W.
1406 m, 25.10.1886: 1 ov. 2 32.3 mm (USNM 12270). — Stn 2732. 37°27'N. 73°33'W, 2074 m, 26.10.1886: I ov.
2 27.3 mm (USNM 12274). — Stn 2734. 37°23'N, 74°33'W. 1514 m. 26.10.1886: I ov. 2 29.7 mm (AMNH 15106).
"Gilliss": stn 71. 36°37.36'N, 74°28'W, 1695-1734 m. 17.11.1974: 1 3 29.6; 1 2 damaged (USNM 314136). —
Stn 72. 36°36. 18’N. 74°24.48'W. 1730-1790 m. 17.11.1974: 1 3 27.9 mm (USNM 314136). — Stn 84, 36°36'N.
74°24.24'W, 1740-1775 m, 19.11.1974: 1 3 18.1 mm (USNM 314136). — Stn 95, 37°05'N. 74°12.30'W. 2250 m.
22.11.1974: 1 3 20.7 mm (USNM 314136). — Stn 97. 37°00.18'N, 74°15'W. 1400-1460 m. 23.11.1974: 2 3 19.0,
20.4 mm; 3 2 19.1-26.9 mm; 2 ov. 2 26.1, 30.6 mm (USNM 314136).
Wallis & Futuna Is area. — Musorstom 7. SW of Combe Bank: stn CP 621, 12°34.4'S. 178°1 1.5'W, 1280 m,
28.05.1992: 1 2 27.0 mm (MNHN-Pa 1712).
Vanuatu. Musorstom 8: stn CP 1124. 15°01.72'S. I66°56.51'E. 532-599 m, 9.10.1994: I 2 43.7 mm (MNHN-Pa
1565). — Stn CP 1125. 15°57.63'S, 166°38.43'E, 1160-1220 m. 10.10.1994: I 3 22.8 mm; 1 juv. (MNHN-Pa 1566).
— Stn CP 1126, 15°58.35'S, 166°39.98'E. 1210-1260 m. 10.10.1994: 1 2 30.2 mm (MNHN-Pa 1567). — Stn CP 1127.
15°58.86’S. 166°37.82'E. 1052-1058 m, 10.10.1994: 1 2 32.3 mm (MNHN-Pa 1568).
New Caledonia. Biocal: stn CP 30. 23°09'S, 166°41'E, 1140 m, 29.08.1985: 1 2 36.2 mm (MNHN-Pa 1569).
Biogeocal: stn CP 205, 22°40.61'S. 166°28.01'E, 1350-1380 m, 8.04.1987: I juv. (MNHN-Pa 1713).
Halipro 1: stn CP 858, 21°42'S. 166°41'E, 1000-1120 m. 20.03.1994: 1 2 (MNHN-Pa 1570).
Bathus 1: stn CP 651, 2I°41.8'S. 166°40.1'E. 1080-1 180 m. 11.03.1993: 1 3 25.1 mm (MNHN-Pa 1678).
Halipro 2: stn BT 95, 24°00’S. 162°08'E, 1224-1233 m. 25.11.1996: 1 3 22.4 mm; 1 2 29.7 mm (MNHN-Pa 1699).
— Stn BT 103. 24°54'S. 162°09'E, 1235-1256 m, 26.11.1996: I 2 41.3 mm (MNHN-Pa 1700). — Sin BT 105. 25°45'S.
162°50'E, 1200-1218 m, 27.11.1996: 1 2 32.4 mm (MNHN-Pa 1701). — Stn BT 106. 25°27'S. 163° 1 6 E. 1315-1357
m, 27.11.1996: 2 2 25.2. 23.8 mm (MNHN-Pa 1702).
Chesterfield Islands. MUSORSTOM 5: sin 323, 21°I8.52’S. 157°57.62'E. 970 m, 14.10.1986: 1 2 21.9 mm
(MNHN-Pa 1571). — Stn 324, 21°15.0TS, 157°51.33'E. 970 m, 14.10.1986: 1 2 22.7 mm (MNHN-Pa 1572).
New Zealand. NZOI. stn U 198. 34°59.3'S, 162°11.2rE. 1573 m. 26.09.1982: 1 juv. (N1WA).
Australia. Lord Howe Rise. "Franklin ": 27°59.3'S, 162°48.6'E. 1250 m. 6.05.1989. id. H E. Stoddart as
Stereomastis andamanensis: 1 2 42.6 mm (AM P40370).
Japan. "Albatross": stn 4909. Kagoshima Gulf. Koshiki Is, 31°38'30"N, 129°27'30"E. 540 m, 11.08.1906: 1 2
24.0 mm (USNM).
East China Sea. "Ryoan Maru": 29°1I'N, 1270H'E, 300-310 m, 17.03.1978. det. Y. MlYA as Stereomastis
andamanensis: 2 2 20.6, 34.9 mm (NUM cr 1 0238).
Philippines. "Albatross": stn 5468, Lagonoy Gulf, San Bernardino Str., 13°35'30"N. 1 23°40'28"E. 1024 m,
18.06.1909: 1 2 32.7 mm (USNM).
Indonesia. "Siboga": stn 48, 8°47'S. 188°44.3'E, 2000 m. 13.04.1899: 1 juv. (ZMA).
Corindon: stn CH 241. 0°58'S, 119°15'E, 1525-1550 m: 1 juv. (MNHN). — Stn CH 286, 2°03'S. 118°45'E. 1710-
1730 m: 3 juvs (MNHN-Pa 1573).
Arabian Sea. "Investigator": stn 108, off Cape Comorin, 7°04’N. 76°34'15"E. 1877 m; 24.10.1890: I 2
20.5 mm (1M 532/7). — Stn 250, SW Ceylon. 6°54'30''N, 79°34'30"E. 878 m. 20.10.1898. det. Ai.COCK as P.
andamanensis: 1 2 31.0 mm; 3 juvs (1M 3421/10).
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
331
Gulf of Aden. "Meteor 5": stn Me5-257. !3°06.6'N, 47°54.0'E, 2227-2250 m. 11.03 1987- 2 8 19 6 4 8 mm
(SMF 24653). — Sin 262, 13°19.7'N, 47°29.2'E. 1830-1837 m. 12.03.1987: 1 <J 22.0 mm (SMF 24652).
Types. The male (22.8 m cl) (USNM 5714) collected by the "Albatross" at stn 2102 (38°44'N. 72°38'W.
2212 m) is designated here as the lectotype of P. nanus Smith. 1884.
The female (20.5 mm cl) (IM 532/7) collected by the "Investigator" stn 108, off Cape Comorin (7°04'N.
76 34 15 E, 1908 m) is the holotypc of Pentacheles andamanensis Alcock. 1894.
The female (35.1 mm cl) (MNHN PA21) collected by the "Talisman" off Senegal is the holotype of
P. grimaldii Bouvier, 1905.
Description. Dorsal surface of carapace setose. Anterior margin straight, rostrum bifid. Internal angle of
oibital sinus with slender spine, external angle unarmed. Eyestalk bearing small spine, curved distad. Basal
antennular segment produced anteriorly to a sharp point, mesial margin smooth, anterolateral margin bispinose.
posterior spine half as long as anterior. Spines on lateral margins of carapace upeurved, anteriormost largest.
Lateral spine formula 5-6:3:6-7, spines further apart posteriorly. Spine formula of median postrostral carina
1 : 1 2: 1 anterior to cervical groove, postcervical carina with two pairs of spines. Gastro-orbilal region with
crescentic row of five prominent antrorse spines. Posterior margin of cervical groove with antrorsc spine between
median and branchial carinae. Branchial carina prominent, subparallel to lateral margin, with five spines. Carapace
between branchial and median carinae unarmed anteriorly, obliquely grooved medially, spinulose posteriorly. Large
spine distally on posterior margin of oblique groove. Posterior margin of carapace spinose. submedian spines
largest. First abdominal tergitc bearing two pairs of spines distally on anterior margin.
Fig. 19. — Polycheles nanus (Smith, 1884), 2 26.1 mm, Off Cape Province. "Africana", 32°26'S, 1 6° 1 l'E, 1302 m
(SAM): a, dorsal view of the cephalothorax; b. lateral view of the abdomen.
332
B. GALIL
Abdominal icrgiies medially carinate. Carinae on tergites 1-5 bearing antrorsc spine; spine of tergites 3-5
longer, salient; sixth tergite bearing mcsially pair of denticulate carinae, joined posteriorly in a prominent tooth.
Tergites 2-5 with transversely oblique submedian grooves. Pleura 2-4 with curved rib mesially, margins minutely
granulate. Second pleuron ovate. Telson with denticulate median crest anteriorly, two minutely serrulate,
convergent carinae posteriorly.
Merus of PI proximally bispinose with subterminal claw-like spine on upper margin, finely granulose on
lower margin; carpus half as long as merus, subterminal spines on upper and lower margins; upper margin of
propodus with subterminal spine, lower margin with two minutely granulate rows distally. P5 subchelate in male,
chelate in female.
Color. — BouviER (1917: 54) wrote "D'apres deux aquarelles . l'animal est rouge avee une leinte d'un blanc
jaunatre sur lc cephalothorax". But he adds "il est probable que cette tcinle (blanc jaunatre) est due a la vase".
Remarks. — P. nanus is distinguished from P. sculptus in having the region between the branchial and
median carinae spinulose posteriorly, and having a prominent spine anteriorly on the branchial groove.
Smith (1884: 359) described P. nanus : "the surface of the branchial region on both sides of the sublateral carina
is armed with many small spines or spinules, and on the anterior part of the oblique ridge between the dorsal and
sublateral carinae there is one spine as large as the spines of the sublateral carina itself", and added "the distinctive
characters are well marked and very constant in all the large number of specimens seen". Yet he hesitated: "This
species is very closely allied to P. sculptus and will possibly prove to be only a dwarf deep-water variety of it".
Faxon (1895: 121) reiterated: "P. nanus may be only a dwarf deep-water variety of P. sculptus". HANSEN (1908:
42), who examined the large series of specimens collected by the "Ingolf ' expedition, disagreed : "P. nanus must
remain an independent species well-marked off from P. sculptus". Yet, KENSLEY (1968: 294) persisted: "The
possibility exists that nana is a juvenile form of sculpta and that some of the spines are lost with development".
However, the distinguishing characters separating P. sculptus and P. nanus were found to be constant in the many
specimens studied; this, together with the comparable vertical distributions of both species, and the ovigerous
nanus specimens found, convince us that sculptus and nanus are indeed distinct species.
Smith's (1884) omission of the spine on the interior margin of the orbital sinus (though the type specimens
possess them) moved Bouvier (1905a) to describe P. grimaldii. Bouvier (1905c: 4) gallantly complimented the
type specimen on being "une tres jolie femclle" and pronounced "j'ai pu constater que le Polycheles grimaldii est
une forme representative... dune espece americaine decrite et figuree par Smith, le P. nanus". Indeed. SELBIE
(1914: 2) regarded grimaldii as "merely... a variety" of P. nanus, and Calman (1925: 19) surmised it "may even
be identical with the typical form of that species".
De Man (1916: 20) recognized the similarity between andamanensis and nanus: "This species [andamanensis]
now bears such a close resemblance to Stereom. nana (S.I. Smith), that I wonder why Col. Alcock has made no
mention of it". Yet the specimens De Man described as andamanensis are in fact P. phosphorus.
DISTRIBUTION. — Worldwide: Iceland. Irish Sea, Bay of Biscay, Portugal. West Africa, Azores. South Africa,
Mid Atlantic Bight, Greenland, Canada. United Stales, Wallis & Futuna Is area, Vanuatu, New Caledonia, New
Zealand, Australia. Japan, East China Sea, Philippines, Indonesia, India, Arabian Sea, Gulf of Aden; 300 m
(present work) - 4000 m (Stephensen, 1923).
Polycheles pacificus (Faxon. 1893)
Fig. 20
Polycheles sculptus pacificus Faxon. 1893: 196; 1895: 122, pi. 50 figs 1, la. — Caullery. 1896: 386. — Hansen,
1908: 42. Murray & Hjort, 1912: 587. fig. 427. — Bouvier, 1915: 2. — Sund. 1915: 372 — Luke 1977- to
— Pretzmann, 1983: 316.
Stereomastis sculpta pacifica - De Man, 1916: 5 (list). — Firth & Pequegnat. 1971: 71. — Wicksten. 1980: 914: 1989:
293. — Wicksten & Mendez, 1982: 110. — Hendricks, 1995: 156. — Dawson, 1997- 23
Polycheles sculptus - Del Solar. 1972: 11 [non Smith, 1880],
Stereomastis pacifica - Retamal, 1981: 16. — Quintana & Retamal, 1984: 133.
Source :
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
333
M WiJSSfa^ fXAMINED- - United States. California. SW Pi Conception. 900-1080 m. 10.05.1979 id
pbw (?6 W ! 080 ^ mm ,LACM ex AHF '«*■*>• - Tanner Bank, 3-52.56^
■ oVV , ■ '°80 ' 979, d‘ M- WlCKSTEN as Stereomastis sculpius pacificus: 1 9 39.7 mm (LACM ex AHF)
TaCM ex AHn0Sa ■ °80*1260 "• 18 101979- id M- W— N « S^LL sculp, us packet ™ 32.2 ™
(da “SuaSh Sr*' ~A'ba"'OSS"- “ 5685' 25”42'45"N’ 1 13°38,30"W, 1161 m, 22.04.1911, 1 4,1*
U3 JWItfSMKSSrr '■ ,125',53° m- 210U97°- id- M- W,CKSIE“
ILA^SlIjSr 1110 1,,197S' iA ” W'«STE»a sS,e,eomas,i, sculp, us pacificus: 1 ov, 2 46.9 -
Gulf of Panama. " Galathea sin 724, 5°44'N, 79°20'W, 2712-3330 m 12 05 1952 2 9 18 9 71 8 ,7un
- Sin 739. 7°22'N. 79°32'W. 938 m. 15.05.1952: 25 <3 31.0-34.7 mm- 12 ov 9 4 9 53 3 J J 2
<ZMCr>:~ ?ln 743: 7°15'N- 79°25'W, 938 m. 16.05.1952: I c3 33.0 mm;’ 1 "ov. 9 46.3 mnn 2 9 ^6 2 40 6 ZIzMC)"1
: s‘n 22‘ 7 28-5'N, 79°12'W, 741-823 m. 18.01.1972: 1 <3 33.9 mm; 3 9 42.5-43 7 mm (RMNhT
21076) : ^ 7°21'N’ 79°35'W- 920 m' 1003 l891: >3 <? 31.5-34.4 mm; 19 9 37.1-48.1 mm (USNM
34.2Emma1°9 3TTL1%SNM)Stn ^ GU‘f °f Guayaquil’ 3°i5'S' 80°55'W- 945-960 m. 10.09.1966: 3 <3 33.9-
450-500 m, 2.05.1967. coll. G. Ricauter: 1 ov. 9 42.4 mm (RMNH 73445)
509 l9“: 3 * 3I J-34J mm: 3 8 39-2-40-3 ■» (USNM), -
3S4 m\,S'i 8 3m°i5 W' m' 23 01 1974- ld- M- WlCKSTEN as Siereonuislis sculpius pacificus: 2 6 33 3
35.4 mm, 1 9 40.4 mm (LACM ex AHF 104-01). - 7°46'S. 80°31'W, 800 m. 23.01.1974. id. M. Wicksten as
Stereomastis sculpius pacificus : 2 c3 35.6. 35.4 mm (LACM ex AHF 103-05)
Chile. "Anton Bruun": sm 714. 25°00'S, 70°40'W, 950 m. 16.08.1966: 3 <J 32.7-35 5 mm (USNM)
Fig. 20. — Polycheles pacificus (Faxon, 1893). <3 32.0 mm. Panama. "Albatross" stn 3394. 7°2I'N. 79°35'W. 920 m
(USNM 21076): a. dorsal view of the cephalothorax: b. lateral view of the abdomen.
334
B. GALIL
TYPES. — The male (34.4 mm cl) (USNM 21076) collected by the "Albatross" stn 3394. off Panama (7°2I'N,
79°35'W, 920 m) is the holotype. The other specimens of the sample are paratypes
DESCRIPTION. — Dorsal surface of carapace densely setose. Anterior margin of carapace nearly straight, bearing
two rostral spines. Internal angle of orbital sinus spinose, external angle convex, unarmed. Eyestalk bearing distad
mammilate tubercle. Basal antennular segment produced anteriorly to a sharp point, mesial margin smooth,
anterolateral angle bispinose. Spine formula of lateral margin of carapace 6:3:7. Spine formula of median
postrostral carina 1:2:1; postcervical carina bearing two paired spines. Gastro-orbital region with five spines.
Branchial carina well-defined, bearing 5 prominent spines. Single spine midway between second spine of branchial
carina and median postcervical carina. Posterior margin of carapace bearing pair of prominent submedian spines.
First abdominal tergites bearing two pairs of spines distally on anterior margin, lateral pair minute,
mammilate. Median carina on tergites 1-4 bearing antrorse spine, increasing in size posteriorly. Fifth carina
without antrorse spine, bulbous. Sixth tergite bearing lyre-shaped carina. posteriorly with raised knob. Shallow,
transversely oblique grooves on either side of median carina 2-5. Pleura 2-5 with cresentic carina mesially; margins
setose, occasionally denticulate. Second pleuron not produced anteriorly. Telson with median blunt crest anteriorly;
rounded, convergent carinae posteriorly.
Merus of PI with two spines proximally and a single subterminal claw-like spine on upper margin; minutely
serrulate lower margin; carpus with subterminal claw-like spine on upper and lower margins; lower margin of
propodus with two rows of spines distally, upper margin with subterminal antrorse spine. P5 subchelate in male,
chelate in female.
Color. — "Pale rose purple on the dorsal side of the carapace and abdomen, deepening to a brighter orange red
on the thoracic appendages, the branches of the posterior abdominal appendages, and the flagella of both pairs of
antennae, and fading to a delicate bluish white on the hepatic and anterior brnachial regions. The raised ridges of the
carapace and abdomen are whitish, the setae on the margins of the basal parts of antennae, swimmerets. etc., are
yellow". (Faxon, 1895: 123, pi. C).
Remarks. — Faxon (1893: 196) separated P. scitlptus pacificits from the typical form citing the "small spine
on each branchial region inside of and on a level with the second spine of the submarginal carina ... the pleurae of
the second abdominal somite have a different shape... These differences, although slight, are constant". Other
distinguishing characters are the mammilate lateral spine on the first abdominal tergite. the bulbous median carina
on the fifth abdominal tergite and the prominent knob posteriorly on the median carina of the sixth abdominal
tergite.
Distribution. — Eastern Pacific, from California to Chile; 600 m (present work) - 3380 m (WiCKSTEN,
1981).
Polycheles perarmatus Holthuis, 1952
Fig. 21
Polycheles ryphlops perarmata Holthuis. 1952b: 7. figs 1-2. — Burukovsky, 1983: 134, fig. 178.
Polycheles perarmatus FOREST, 1963: 627. — FraNSEN et al.. 1997: 61.
Polycheles typlilops - SPRINGER & BULLIS. 1956: 14 (p.p.).
Polycheles typlilops perarmatus - Firth & Pequegnat. 1971: 56. fie. 8. — Griffin & Stoddart. 1995: 243. — Dawson.
1997: 15.
Material EXAMINED. — West Africa. Ivory Coast and Nigeria. "Pillsburv": stn 44, 5°05'N, 4°00'W, 586-
403 m, 30.05.1964: 3 juvs (RMNH). — Stn 255, 3°49’N, 7°38'E. 269-264 m, 14.05.1965: 1 <5 40.2 mm: 2 2 41.4.
49.8 mm (RMNH). — Stn 256. 3°49'N. 7°38'E, 267-264 m, 14.05.1965: 8 3 48.0-54.4 mm; 2 2 54.4. 51.1 mm
(RMNH).
Ghana. Guinean Trawling Survey I: stn 28\8. 4°16'N, 2°09.30’W. 380-400 m. 4.10.1963: I 3 24 9 mm ( MNIIN-
Pa 559).
Equatorial Guinea. " Atlantide stn 120. 2°09'N, 9°27'E. 260-650 m, 1.03.1946: 1 3 19.0 mm (ZMC).
Gabon. 1°64'S. 8°44'E, 396 in. 3.09.1963, id. R.B. Manning: 1 3 37.6 mm (USNM 125986). — "Mbizi"-
stn AS 154, 0°I5'S. 8°47'E. 239 m. 15.03.1949: 2 2 50.7. 45.8 mm (RMNH 8622).
Source . MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
335
- S4'8' ilSSt.34”50. » 707 1967- *• 1 * ^ - imwn* 1122).
1 ' .7 W’ 30?:^° m- 30.10.1969. coll. A. Crosnier: 1 <3 50.8 mm; 1 2 46.2 mm (MNHN-Pa 1575)
Gulf of Mexico Albatross stn 2376, 29°03'03"N, 88°16'W. 583 m, 11.02.1885, id. M.J. Rathbun as
1 1 of Txrs” ‘ I vi |2p“ mm' 2d31-‘ mm; 1 -iuv' (USNM 23470)- — Sln 2377, 29°07'05"N. 88°08'W. 378 m
86°26'W8409 m l4 mYsTVL , T™Z": 1 5 33 7 mm; 2 9 44-3- 47.6 mm (USNM 9654). _ Stn 2398. 28»45’n!
86 -6W, 409 m, 14.03.1885, id. M.J. Rathbun as P. agassizii: 1 2 60.1 mm (USNM 9743)
•Oregon'': stn 126, 29°02'N, 88°34.5'W. 351 m. 23.09.1950: 3 3 33.2-41.2 mm: 2 2 3^6 41 4 mm (USNM
9 44 • " ?n *2’ 27°41'N' 94°59’W- 450-540 m- i6 04-1952' id. F.A. CHACE as />. ypVops ii 38 5 mm USNM
936r61)- 7,S '4°q7: 07'N- 89°59'W’ 464 20.09.1955. id. F.A. ChaCE as P. typhlops: 1 <J 35.2 n m ov 2
40.8 mm (USNM 99454). - Stn 3076. 28°30'N. 89°23'W, 41 m. 25.10.1960: 2 <3 34.7. 41.3 mm: 5 2 37 7-54 4 mnr
I ov. 2 45.6 mm (USNM 273140). — Stn 3651, 29°12'N, 88°03'W 450-540 m 25 07 I96">- ? o sc r
(USNM 244083). - Stn 4005. 29*07.5'N, 88’09’W. 540 m, 23. 10 f96i: 1 6 34 8 mm (USNM 273 140) Stn 470^
27°55'N, 90°28'W, 540 m. 22.02.1964: 1 <3 28.3 mm; 5 2 22.0-33.7 m, (USNM 273140 - s ^729 SS’
16 if 19644°>m^24nn19^ ' 3 2mtZ7' ' 2 3°'6 mm (USNM 260368)- - Stn 5107. 29°I5'N, 80°05'W?378 m.'
i L40^ ~ $,n 524°’ 29°39'N- 80°"-5'W- 342 - '0-02,965: 1 3
"Silver Bay'': sin 2070, 29°13'N, 79°59'W. 369 m. 2.05.1960: 1 2 52.5 mm; 1 ov. 2 43.1 mm (USNM 932276)
Stn 5464. 29°25'N, 80°07'W. 310 m. 1 1.02.1964: 4 <3 38.8-42.2 mm; 6 2 40.1-45.6 mm (USNM 260368)
, * ;?S : imcu!”21 3r'N' 88°2 1 48 w- 379-430 m. 24.02.1989. id. as Stereo, nasus nana by M.K. WiCKSTEW
88"25 ,2'w- 4,,-4,g m' “• as s—“ — b7
P a n a raa. Go Ifo de Escudo de Veraguas. "Oregon": stn 3583. 9°16'N, 81°37'W. 512 m. 25.05.1962: 3 <3 38 6-
4L0 mm ^ ^ 8,°22'W' 36°’396 *• 3I05'**- id F-M- Bayer as P. agassizir^
,,KCw!;b,b“*ea' "°Je8°"l Sln 2774- n°32'N- 62°40'W- 351-382 m- * 904. 1 960, 1 <3 41.4 mm: I ov. 2 48.2 mm
232276) 232276 ' S‘n 27?9, 35 N- 62°59'W- 468-432 m. 20.04.1960: I 3 45.4 mm: I 2 40.1 mm (USNM
"Pillsbury stn 776, 12°13.3'N, 72°50'W, 408-576 m. 29.07.1968: 1 <5 35.5 mm; 1 2 35.4 mm (RMNH)
56°9tURr,wan^' 7°2L6'N’ 55°22-2'W' 400-420 m. 27.04.1966: 2 juvs. (RMNH). - Stn F36. 7°26’N.
56 -1.8 W, 365-410 m. 5.05.1966: 2 juvs. (RMNH).
-TYPESo' 7 The male 1131 mm tolal lenglh> (irScNB 16808) collected by the "Mbizi" stn AS 35. off West
Africa (71 6'S 1 2C01'E, 440 tn) is the hololype. The females (50.7. 45.8 mm cl) (RMNH 8622) collected at
stn AS 154 (0°15'S. 8°47'E, 239 m) are paratypes.
Description. — Dorsal surface of carapace setose, prominently spinulate. Frontal margin concave. Median
submarginal tooth prominent. Single rostral spine antrorse. Internal angle of orbital sinus'" triangular, spinose;
external margin pectinate. Internal margin of orbit mesially pectinate, dividing orbit into two. Eyestalk bearing
upcurved spine. Basal antennular segment produced anteriorly to a shaip point, mesial margin spinose. anterolateral
margin rounded, bispinose. Spines on lateral margins of carapace upcurved. Spine formula of lateral margin of
carapace 7-9:4-5:17-20; spines successively decrease in size posteriorly. Median postrostral carina granulate,
irregularly spinulate, postcervical carina with paired spinules. Postorbital carina obsolescent, comprising five
antrorse spines, anteriormost largest. Posterior margin of cervical groove spinose between branchial carinae.
Branchial carina prominent, parallel with lateral margin, bearing 12-15 spines; region between branchial and
median postcervical carinae prominently spinulose. Posterior margin of carapace bearing seven or more pairs of
antrorse spines, submedian spines largest.
Abdominal tergites densely granulate, medially carinate. Carinae on tergites 2-5 bearing antrorse spines, fourth
and fifth more prominent, salient. Sixth tergite prominently granulate, bearing mesially twin parallel rows of
granules, confluent posteriorly. Anterior margin of first tergite bearing nearly 30 denticles. Anterior and posterior
margins of tergites 2-5 prominently denticulate. Pleura prominently granulate, margins denticulate. Second pleuron
rounded; pleura 3-5 with curved, granulate rib mesially. Telson with granulate median crest anteriorly, two
minutely serrulate, convergent carinae posteriorly. Inferior surface of uropodal expopod tricarinatc.
Upper margin of merus of PI spinulate with subterminal claw-like spine, lower margin finely granulose;
carpus half as long as merus. two serrulate rows distally on upper margin, subterminal spines on upper and lower
margins; upper margin of propodus with several rows of antrorse spinules, lower margin with two minutelv
spinulate rows. P5 simple in male, chelate in female.
336
B. GALIL
Fig. 21. — Polycheles perannatus Holthuis, 1952. 3 50.8 mm. Congo. 4°49'S. 11°17'W, 300-600 m (MNHN-Pa 1575):
a. dorsal view of the cephalothorax; b, lateral view of the abdomen.
Remarks. — P. perannatus is distinguished from P. typhlops by the rounded second pleuron. the three carinae
venlrally on the uropodal expopod. the prominently granulate sixth tergite, the seven pairs of spines on the
posterior margin of the carapace and the evenly denticulate anterior margin of the first abdominal tergite.
Distribution. — Eastern Atlantic: Gulf of Guinea to Angola, Azores; Western Atlantic: Gulf of Mexico,
Caribbean, Panama; 41-650 m.
Polycheles phosphorus (Alcock, 1894)
Fig. 22
Pentacheles phosphorus Alcock, 1894: 240. — Wood-Mason & ALCOCK, 1894, pi. 8 fig. 2. — ALCOCK & ANDERSON,
1894: 165. — Anderson. 1896: 98. — Thomson. 1901 : 18. — Alcock. 1901a: 74; 1902: 134. — Lloyd. 1907: 6.
— Hansen, 1925: 142. — Boas. 1939. fig. 1.
Polycheles phosphorus - Alcock. 1899: 33; 1901b: 168. — Calman, 1910: 59. fig. 38: 1911. fig. 46. — KEMP &
Sewell, 1912: 24. — Balss, 1924: 177, fig. 2. — Duncan, 1948: 69.
Stereomastis andamanensis - De Man, 1916: 16. pi. 1 figs 2-2b (p.p.). — Takeda & Hanamura, 1994: 31. — Griffin &
Stoddart, 1995: 244 (p.p.) [non Alcock, 1894],
Stereomastis phosphorus - Calman, 1927: 55, fig. 37. — Dahl, 1953: 46. — Firth & Pequegnat, 1971: 68. — Monod.
1973: 126, figs 40-44 (p.p.). — Holthuis, 1984: 4.
not Polycheles phosphorus - Rathbun, 1906: 898 [= P. surdus sp. nov.].
not Stereomastis phosphorus - De Man. 1916: 15 [= P. aculeatus sp. nov.].
not Stereomastis phosphorus - Griffin & Stoddart, 1995: 246 [= P. aculeatus sp. nov.. P. auriculatus Bate. 1878.
P. surdus sp. nov.].
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
337
Material EXAMINED. — Central Eastern Pacific. "Te Vega": stn 58. 1°31’N, 128°13’W. 525-570 m
27.09.1963: I $ 47.0 mm (USNM).
Vanuatu. MUSORSTOM 8: stn CP 1074. 15°48.42'S. 167°24.27'E. 775-798 m. 4.10.1994: I juv. (MNHN-Pa 1738).
— Stn CP 1076. 15°53’S. 167°30'E, 1100-1191 m, 4.10.1994: 1 juv. (MNHN-Pa 1739). — Stn CP 1089, 1 5°57.30'S
167°27.73'E. 799-850 m, 5.10.1994: 2 juvs (MNHN-Pa 1740).
New Caledonia. 22°02'S, 165°57'E. 800 m. 1971. det. Th. Monod Stereomastis phosphorus: 1 3 27.3 mm
(MNHN-Pa 1576).
Bathus 2: stn CP 743. 22°35.6'S. I66°26.2'E. 713-950 m. 14.05.1993: 2 3 32.4. 37.1 mm (MNHN-Pa 1578). —
Stn CP 751. 22°24.35'S, 166°12.83’E, 1300-1500 m, 15.05.1993: 1 2 22.5 mm (MNHN-Pa 1579).
Chesterfield Islands. Musorstom 5: stn 324. 21°15.0rS, 157°51.33'E. 970 m. 14.10.1986: 1 $ 20.5 mm
(MNHN-Pa 1577).
Australia. Coral Sea. "Franklin": 10°34.28'S. 144°21.99'E. 990-1053 m, 21.08.1988, id. H.E. Stoddart as
Stereomastis andamanensis: 1 3 33.3 mm (AM P40367).
N.W. Coast. "Solea" N.W. Port Headland. 18°48'S, 1 16°21'E, 694-704 m. 5.04.1982: 1 3 44.8 mm (WAM C 1 3436).
Japan. "Albatross": stn 4906, Koshiki Is, 31°39'N, 129°20’E, 664-731 m, 11.08.1906: 2 2 26.3. 44.5 mm
(USNM). — Stn 4911, Koshiki Is, 31°38'30"N, 129°19'E. 794 m, 12.08.1906: 1 3 37.9 mm; I 2 45.3 mm. 2 juvs
(USNM). — Stn 4912. Koshiki Is, 31°39'40"N. 129°20'E. 704 m. 12.08.1906: 2 3 32.0. 38.5 mm; 1 2 46.3 mm;
3 juvs (USNM). — Stn 4915. Koshiki Is, 31°31'N, 129°25'30"E, 769 m. 12.08.1906: 2 3 33.6, 41.2 mm (USNM).
Suruga Bay. off Fuji River, 200 m, 9.12.1975: 1 3 broken (NFU 530-2-1495). — S.W. Kyushu. 30°32'N. 128°25'E.
700-750 m. 15.06.1978. colls O. Tabeta & K. Hayashi: 3 3 39.9-40.0 mm (NFU 530-2-1490); 1 2 47.8 mm: 2 ov. 2
49.7. 51.3 mm (NFU 530-2-1527).
East China Sea. "Yuryo Maru": 29°38.92'N. 127°55.95'E, 750 m. 4.02.1978. det. Y. Miya as Stereomastis
andamanensis: 6 3 ; 1 ov. 2 ; 3 2 (NUM crl0229). — 29°23.01'N. I27°30.92'E, 650-642 m. 5.02.1978. det. Y. Miya as
Stereomastis andamanensis: 1 3 37.8 mm; I 2 36.8 mm (NUM cr 1 023 1 ).
"Ryoan Mam" (det. Y. Miya as Stereomastis andamanensis ): 28°53'N, 127°18'E, 820-830 m. 16.03.1978: 6 3 : 1 2 ;
2 ov. 2 (NUM crl0230). — 28°43'N. 127°14’E. 750-755 m, 11.03.1978: 6 3 31.3-36.5 mm; 3 ov. 2 47.6-53.0 mm;
1 2 47.4 mm (NUM crl0232). — 28°44'N, 127°01'E. 610-640 m, 11.03.1978: 1 3 41.6 mm; 3 ov. 2 47.4-51.3 mm;
3 2 50.5-51.6 mm (NUM crl0233). — 28°42'N. 127°09'E. 500-535 m. 11.03.1978: 2 3 26.0. 26.5 mm (NUM
cr 1 0234). — 28°35'N. I27°10'E. 600 m. 16.03.1978: 3 3 ; 2 ov. 2 ; 4 2 (NUM cr!0235). — 28°45'N, 127°07'E. 542-
530 m, 16.03.1978: 2 3 34.1, 39.5 mm; 3 ov. 2 47.1-59.2 mm; 2 2 41.0. 51.0 mm (NUM crl0236). — 28°50’N.
1 27°14'E, 700-740 m. 16.03.1978: 9 3 26.7-40.1 mm; 3 ov. 2 41.4-50.8 mm; 6 2 27.7-58.7 mm (NUM cr!0237). —
29°46'N, 1 27°59'E. 710-713 m. 20.03.1978: 3 ov, 2 26.7-54.8 mm (NUM crl0240). — 29°44'N. 128°03'E. 815 m.
20.03.1978: 1 2 35.5 mm (NUM crl0241). — 29°49'N. I28°06'E. 915-932 m, 20.03.1978: 2 2 damaaed (NUM
crl0242).
South China Sea. W. Borneo. 6°05'N. 110°07'E. 6.11.1964, coll. A.J. BRUCE: 1 3 35.8 mm; 1 2 50.9 mm
(RMNH 23484).
Indonesia. "Siboga": (del. De Man as Stereomastis andamanensis): stn 18. 7°28.2'S. 1 1 5°24.6'E. 1018 m.
18.03.1899: 1 juv. (ZMA). — Stn 85. 0°36.5'S, II9°29.5'E, 724 m. 17.06.1899: 2 3 43.9. 44.4 mm (ZMA). —
Stn 137, 0°23'S, 127°29’E, 472 m. 3.08.1899: I 3 20.3 mm (ZMA). — Stn 314, 7°36’S. lH^O.S’E. 694 m.
17.02.1900: 1 3 41.6 mm; 1 2 38.3 mm (ZMA).
"Albatross": stn 5613, Sulawesi, Gulf of Tomini. 0°42'S. 121°44'E. 1376 m. 2.11.1909: I juv. (USNM). — Stn
5646, 5°31’30"S. 1 22°22'40"E, 834 m, 16.12.1909: 1 juv. (USNM). — Stn 5658. Gulf of Boni. 3°32'40"S.
120°31'30"E. 918 m. 19.12.1909: 1 3 44.7 mm (USNM).
Corindon: stn CH 209, 0°07.3’S, I17°53.8'E. 490 m, 31.10.1980: 10 3 20.8-36.2 mm; 10 2 20.1-51.7 mm
(MNHN-Pa 1588). — Stn CH 214, 0°3I.4'N, 117o50.1'E, 595 m, 1.11.1980: 1 3 42.9 mm (MNHN-Pa 1589). —
Stn CH 217. 0°38.2'N, 117°59.6'E, 470-447 m, 1.11.1980: 1 3 39.2 mm; 1 2 19.8 mm (MNHN-Pa 1590). —
Stn CH 240, 0°37'N. 1I9°33'E, 675 m. 5.11.1980: 1 3 43.2 mm: 2 juvs (MNHN-Pa 1591). — Stn CH 276. 1°54.6'S.
1 1 9°1 3.8'E, 456-395 m, 8.11.1980: 1 3 29.7 mm; 3 2 24.7-48.4 mm (MNHN-Pa 1592). — Stn CH 280. 1°59.0'S.
1 19°09.9'E, 715-800 m. 8.11.1980: 1 juv. (MNHN-Pa 1593). — Stn CH 281, 1°57.5'S. 1 19°02.0'E. 1120-1150 m.
8.11.1980: 1 juv. (MNHN-Pa 1594).
Snellius 2: stn 4.267, 8°17.6'S, 118°21.3'E, 650 m, 29.10.1984: 4 3 24.7-35.4 mm: 4 2 19.2-37.3 mm (RMNH).
— Stn 4.276, 8°12'S, 118°12'E, 750 m, 31.10. 1984: 1 3 40.2 mm; 2 ov. 2 47.8.43.3 mm (RMNH).
Karubar: Tanimbar /.: stn CP 38. 7°40'S, 132°27'E. 620-666 m. 28.10.1991: 2 juvs (MNHN-Pa 1580). —
Stn CP 54. 8°21'S, 131°43'E, 836-869 m, 30.10.1991: 2 3 20.3, 34.7 mm; 3 2 21.7-64.5 mm (MNHN-Pa 1582). —
Sin CP 73, 8°29'S. 131°33,E, 855-840 m, 2.11.1991: 1 3 33.0 mm; 1 2 49.2 mm (MNHN-Pa 1583). — Stn CP 81.
9°35'S. 131°02'E, 200-207 m, 4.11.1991: 2 3 31.2. 50.7 mm; 2 juvs (MNHN-Pa 1584).
Philippines. "Albatross": stn 5114, Balayan Bay, 13°36'11"N, 120°45'26"E, 622 m. 20.01.1908: 2 2 27.1.
32.7 mm (USNM). — Stn 5286. Luzon. 13°38'I5"N, 120°34'20"E. 823 m. 20.07.1908: 1 3 44.8 mm (USNM). —
Stn 5349, Palawan Passage. 10°54'N, 118°26'20"E. 1335 m, 27.12.1908: 2 3 28.8, 42.8 mm; 2 juvs (USNM).
338
B. GAUL
MUSORSTOM 1: stn 47. 13°40.7'N, I20°30'E, 757-685 m. 25.03.1976: 1 <3 44.2 mm; 3 2 34.9-44.8 mm (MNHN-Pa
1587).
Musorstom 2: stn CP 50. 13°37'N, 120°33'E. 810-820 m. 27.11.1980: I 6 28.9 mm (MNHN-Pa 1585).
S'i CP 55, 13°54'N. 119°58'E. 865 m. 27.11.1980: 1 6 22.8 mm (MNHN-Pa 1586).
Bay of Bengal. "Investigator": stn 112, 13°47'30''N. 92°36'E, 1026 m. 7.11.1890: 1 2 54.0 mm (IM 6873/9).
Ceylon. "Investigator": stn 377. 5°51'N. 80°26'E. 1107 m. 25.04.1907: 2 2 26.0, 28.0 mm (IM 5814/10).
Laccadive Sea. "Investigator": stn 177. 13°47'49"N, 73°07'E, 1166 m, 5.05.1894: 1 2 38.7 mm (USNM 19012;
ex. IM).
Type. — The female (54.0 mm cl) (IM 6873/9) collected by the "Investigator" at stn 1 12, Bay of Bengal
(13°47'30"N. 92°36'E. 1026 m). is the holotype of Pentacheles phosphorus (Alcock, 1894).
Fig. 22. — Polycheles phosphorus (Alcock, 1894), 2 43.6 mm. East China Sea, "Ryoan Marti", 28°50'N. 127°14'E. 700-
740 m (NUM 10237): a. dorsal view of the cephalothorax; b. lateral view of the abdomen.
Description. — Lateral margins of carapace anteriorly convergent. Dorsal surface of carapace setose. Rostrum
bifid. Small spine on internal angle of orbital sinus, external angle unarmed. Eye stalk bearing mammilate
tubercle, curved distad. Basal antennular segment produced anteriorly to a sharp point, mesial margin smooth,
anterolateral margin unispinose. Spines on lateral margins of carapace upcurved. Spine formula of lateral margin of
carapace 6:3:6-7, posteriormost spine further apart. Spine formula of median postcervical carina (excluding rostral
spines) 1: 1:2:1; median postcervical carina bearing two pairs of spines. Gastro-orbital region with five prominent
antrorse spines in arcuate row. Posterior margin of cervical groove with antrorse spine midway between median
postcervical and lateral carinae. Branchial carina prominent, bearing 6 or 7 spines. Anteriorly between branchial and
median carinae shallow diagonal groove, 5 or 6 spines posterior to groove. Posterior margin of carapace bearing
antrorse submedian spines.
Abdominal tergites medially carinate. Carinae on tergites 1-5 bearing antrorse spines, 3 and 4 largest. Sixth
tergite boldly sculptured, raised lateral margins, mesially bearing prominently denticulate double carinae, confluent
at posterior margin. Anterior margin of first tergite with two pairs of lateral spines. Tergites 2-5 bearing deep,
Source MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
339
transversely oblique, submedian grooves with raised margins. Pleura 2-5 with curved rib mesially. bolder
posteriorly, margins minutely denticulate. Second pleuron ovate. Telson with short crest anteriorly, two minutely
serrulate, confluent carinae posteriorly.
Merus of PI proximally bispinose, claw-like spine subterminally on upper margin, finely granulose on lower
margin; carpus 2/3 as long as merus, subterminal spines on upper, lower margins; upper margin of propodus with
subterminal spine, lower margin with two mintely spinulate rows. P5 subchelate in male, chelate in female.
Color. — "bright pink" (ALCOCK, 1894: 241).
REMARKS. — ALCOCK (1894: 240) clearly distinguished phosphorus as having on its "basal joint of the
antennules . a single spine at the antero-external angle", and further (1901b): "some spines flanking the cardio-
branchial grooves". Misunderstanding the significance of these characters, authors mistook various polychelid
species for phosphorus. De Man (1916) and GRIFFIN & STODDART (1995) mingled both nanus and phosphorus
and misidentified them as andamanensis , whereas their phosphorus are different species - in fact. GRIFFIN &
STODDART (1995: 246) had three different species identified as Stereomastis phosphorus.
DISTRIBUTION. — Vanuatu, New Caledonia, Australia, Japan, China Sea, Indonesia. Philippines, Bay of
Bengal, Ceylon, Laccadive Sea; 200 m (present work) - 1354 m (Takeda & Hanamura, 1994).
Polycheles politus sp. nov.
Fig. 23
MATERIAL EXAMINED. — Philippines. "Albatross": stn 5219, btw Marinduque & Luzon, 1 3°2 1 ’N. 122°I8'45"E.
969 m, 23.04.1908: 7 2 25.6-49.6 mm (USNM). — Stn 5423. Cagayan Is, 9°38'30"N, 121°H'E, 914 m. 31.03.1909:
1 3 19.1 mm; 2 juvs (USNM). — Stn 5424. Cagayan Is. 9°37'05"N, 121°12'37"E, 622 m. 31.03.1909: 1 3 19.1 mm;
1 2 24.9 mm (USNM). — Stn 5488, Panaon l„ Leyte. 10°00'N, 125°06'45"E, 1412 m. 31.07.1909: 1 3 18.4 mm;
3 juvs (USNM). — Stn 5492, Mindanao, 9°12'45"N, 125°20'E, 1344 m. 1.08.1909: I 2 34.3 mm (USNM). —
Stn 5494, Mindanao, 9°06'30"N, 125°18'40"E, 1220 m, 2.08.1909: 2 3 17.9, 22.4 mm (USNM). — Sin 5495,
Mindanao, 9°06’30"N, 125°00'20"E, 1785 m. 2.08.1909: 1 2 23.1 mm (USNM). — Stn 5511. Mindanao. 8°15'20"N,
123°57'E, 750 m. 7.08.1909: I 2 23.1 mm (USNM). — Stn 5514, Mindanao, 8°32'42"N, 123°58'36"E. 1275 m,
8.08.1909: 1 2 damaged (USNM).
Musorstom 2: stn CP 38, 12°53.5'N, 122°26.6'E. 1650-1660 m. 25.11.1980: 2 3 21.5. 25.1 mm: 5 2 20.7-
37.0 mm (MNHN-Pa 1595). — Stn CP 39, 10°02.8'N, 122°37.1'E, 930-1190 m. 25.11.1980: 19 juvs (MNHN-Pa 1596).
— Stn CP 40, 13°07.7'N, 122°39.1'E, 280-440 m, 25.11.1980: 1 2 21.8 mm (MNHN-Pa 1597). — Stn CP 42,
13°04.2’N, 122°25.0'E, 1580-1610 m, 25.11.1980: 1 3 21.7 mm; 2 2 22.3,31.7 mm (MNHN-Pa 1 598). — Stn CP 44.
13°23.2'N, 1 22°20.7'E, 760-820 m, 26.11.1980: 1 3 27.5 mm: 5 juvs (MNHN-Pa 1599).
Gulf of Thailand. " Galathea stn 324, Sir. of Malacca. 6°06'N. 96°00'E, 1130 m, 9.05.1951: 1 2 25.3 mm
(ZMC). — Stn 443. Mindanao Sea, 8°48'N, 124°09'E, 1570 m, 16.08.1951: I c? 22.6 mm: 3 2 26.4-28.3 mm; 3 juvs
(ZMC).
Indonesia. "Albatross": stn 5629, Moluccas, S. Doworra I., 0°50'S, I28°12'E, 375 m. 2.12.1909: 1 3 17.3 mm
(USNM).
Types. — The male (25.1 mm cl) (MNHN) collected by Musorstom 2, stn CP 38 (12°53.5'N. 122°26.6'E.
1650-1660 m) off the Philippines, is the holotype. The other specimens in the sample are paratypes.
DESCRIPTION. — Margins of carapace anteriorly convergent. Dorsal surface of carapace setose. Rostrum bifid.
Internal angle of orbital sinus with minute spine, external angle unarmed. Eyestalk bearing minute mammilate
tubercle, curved distad. Basal antennular segment elongate, produced anteriorly to a sharp point, mesial margin
smooth, anterolateral margin unispinose. Spines on lateral margins of carapace upeurved. anteriormost largest.
Spine formula of lateral margin of carapace 6:3:6-7, posteriormost spine further apart. Spine formula of median
postrostral carina (excluding rostral spines) 1: 1:2: 1, median postcervical carina bearing two pairs of spines. Gastro-
orbital region quadrispinose, anterior spine larger by far. Posterior margin of cervical groove with antrorse spine
between median postcervical carina and lateral margin of carapace. Branchial carina obsolescent, prominent spine
(sometime preceded by spinules) near posterior margin of carapace. Posterior margin of carapace bearing antrorse
submedian spines.
340
B. GALIL
Abdominal tergites medially carinate. Anterior margin of first tergite with two pairs of prominent lateral
spines. Carinae on tergites 1-4 bearing antrorse spine, spine of third tergite longest; carina of fifth tergite unarmed.
Sixth tergite smooth, mesially bearing lyre-shaped denticulate carina. Tergites 2-5 bearing shallow, transversely
oblique, submedian grooves. Pleura 2-5 with obsolescent curved rib mesially, margins minutely denticulate.
Second pleuron ovate, bearing spine on anterior margin. Telson with short crest anteriorly, two minutely serrulate,
confluent carinae posteriorly.
Merus of PI proximally bispinose with subterminal claw-like spine on upper margin, finely granulose on
lower margin; carpus 2/3 as long as merus, subterminal spines on upper and lower margins; upper margin of
propodus with subterminal spine, lower margin granulate. P5 subchelate in male, chelate in female.
Etymology. — The Latin politus (- polished) refers to the smooth, unarmed carapace of the species.
REMARKS. — Both P. phosphorus and politus have but a single spine on the anterolateral margin of the basal
antennular segment, however, politus is clearly distinguished from phosphorus by its unarmed fifth abdominal
carina, the obsolescent branchial carinae, and the postcervical part of the carapace, between the branchial and
median carinae, lacking spines.
Distribution. — Philippines, Gulf of Thailand, Indonesia; 280-1785 m.
Polycheles sculptus Smith. 1880
Fig. 24
Polycheles typhlops - Norman, 1878: 382 (p.p.); 1879: 175 (p.p.).
Polycheles sculptus Smith. 1880a: 346. pi. 7 figs 1-6; 1880b: 269. — Faxon, 1893: 196; 1895: 121. pi. C fig. 2; 1896:
155. — Caullery, 1896: 385. — Koehler. 1896: 721 (list). — Alcock, 1901b: 170. — Stebbing, 1902: 36. —
Senna. 1902: 338. — Joubin. 1905: 75, fig. 58. — Bouvier, 1905a: 480: 1905c: 3; I905d: 4; 1915: 2: 1917: 51.
pi. 3. fig.l; 1925: 438. fig 11. pi. 7 fig. 1. — Hansen, 1908: 41. — Stebbing. 1910: 377. — Andrews. 1911: 428.
figs 11-15. — Murray & Hjort, 1912: 538. — Selbie, 1914: 18. pi. 2. figs 1-9. — Sund. 1915: 372. —
Williamson. 1915: 457. — Balss, 1925: 201. — Boone. 1930: 86. pi. 23 fig. b. — Miranda & Rivera, 1933: 17.
— Delphy & Magne. 1938: 83 (list). — Boas, 1939: 4, figs 3. 7-8. — Zariquiey, 1946: 99. — Le Danois, 1948: 150
(list). — Springer & Bullis, 1956: 14. — Holthuis, 1962: 183. — China, 1964: 1 1 1. — Barnard, 1964: 12 (list).
— P£r£s & Picard, 1964: 105. — Allen. 1967: 56. — Peres, 1967: 519. — Rodriguez, 1980: 194.
Pentacheles spinosus A. Milne Edwards. 1880: 66. — FlLHOL, 1885: 139, fig. 44. — W. Marshall, 1888: 267, fig. 88.
— YOUNG, 1900: 442. — Seeliger. 1901: 43. — Bouvier, 1925: 438, figs 1 1-12, pi. 7 fig.l.
Willemoesia leptodactyla - Giglioli, 1881: 358; 1882: 5; 1912: 186 [non Willemoes-Suhm. 1875],
Pentacheles sculptus - Smith, 1882: 23, pis 3-4; 1884: 358 (list); 1886a: 188; 1886b: 607 (list). — Verrill, 1882: 364;
1884: 653; 1885: 554; pi. 34 fig. 152. — Agassiz. 1888: 42. fig. 239. — Alcock & Anderson. 1899: 289.
Stereomastis sculpta - De Man, 1916: 8. — Stephensen, 1923: 66. — Calman, 1925: 18; 1927: 55. — Boone. 1927: 90.
— Barnard, 1950: 572. fig. 105d. — Holthuis, 1952b: 11, fig. 3; 1962: 182; 1984: 4; 1987: 297, fig. 2; 1991,
fig. 169. — Bernard, 1953: 87. — Siversten & Holthuis, 1956: 41. — Dollfus, 1956: 135 (list). — Squires,
1965: 88, figs 37-38; 1966: 2. — Kensley. 1968: 29; 1981a: 60 (list); 1981b: 29 (list). — Zariquiey. 1968: 210. —
Carpine, 1970a: 11 (list); 1970b: 135 (list). — Firth & Pequegnat, 1971: 69. — Relini-Orsi & Relink 1972: 59.
fig. 5. — Relini-Orsi, 1973: 29. — Lagardere, 1973: 93; 1977a: 399. — Thiriot. 1974: 344 (list). — Williams &
Wigley, 1977: 8. — Baf.z & Andrade, 1979: 225. pi. 1 fig. 6. — Pequegnat & Jeffrey, 1979: 72. — Wenner,
1979: 441. — Beaubrun, 1979: 40. fig. 21. — Andrade & Baez, 1980: 262 (list). — Burukovsky et ah, 1982:
523. — Burukovsky. 1983: 134, fig. 179. — Takeda & Okutani, 1983: 75. — Pohle, 1985: 21. — Andrade,
1986: 45. — Duris. 1987: 9. — Macpherson. 1988: 58. fig. 5b. — Abello & Valladares. 1988: 98. — Pequegnat
et ah, 1990: 63. — Squires, 1991: 360. figs 190-191. — Abello & Cartes, 1992: 109. — Cartes & Abello, 1992:
139. — Cartes & Sarda, 1992: 1315. — Falciai & Minervini, 1992: 134. — Cartes, 1993: 32 (list). — Cartes et
ah, 1993: 210. — Emmerson, 1993: 181. — Quackenbush. 1994: 85. — Gonzalez, 1995: 126. — Griffin &
Stoddart. 1995: 248. — Tiefenbacher. 1995: 8. fig. 3.
Stereomastis sculptus - STEBBING, 1917: 30.
Stereomastis sculpta - Firth & Pequegnat, 1971: 69. — Wenner & BoESCH, 1979: 131. — Dawson. 1997: 21.
Stereomastis andamanensis - TUNG el ah, 1988: 45, fig. 45a-f.
not Polycheles sculptus - Bruun, 1950: 24, fig. 16 [= P. talismani Bouvier, 1917],
Source MNHN, Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
341
not Siereomaslis sculpta - HOLTHUIS, 1952b: 11. — LONGHURST, 1958: 32. — FOREST, 1963: 627. — Macpherson, 1983:
48. — RUCABADO & Bas, 1984: 19 1= P. lalismani Bouvier, 1917],
not Stereomastus sculpta - Gauld, 1960: 63), [erroneous spelling] [= P. lalismani Bouvier. 1917],
not Polvcheles sculptus - Del Solar, 1972: 11 [= P. pacificus Faxon, 1893].
Fig. 23. — Polycheles politus sp. nov., 3 25.1 mm. holotype, Philippines. Musorstom 2 stn CP 38. 12°53.5'N.
122°26.6'E, 1650-1660 m (MNHN-Pa 1595): a, dorsal view of the cephalothorax; b. lateral view of the abdomen.
MATERIAL EXAMINED. — Mediterranean Sea. "Washington": 41°15'09"N, 8°10'41"E. 2150 m. 4.08.1881.
coll. & id. E.H. GiGLlOLl as Willemoesia lepiodaciyla : 1 juv. damaged (MF). — 41°14’38"N. 8°18'05' E, 2145 m.
8.08.1881, coll. & id. E.H. GiGLlOLl as W. leptodaciyla : 1 3 21.2 mm (MF). — 41°24'42"N, 7°43’28"E. 2836-2809 m.
9.08.1881, coll. & id. E.H. Giglioli as W. leptodactyla : 1 $ 25.1 mm (MF).
Ibero-Moroccan Gulf. BALGIM: stn CP 69. 35°11'N, 7°50’W. 1978-2077 m. 5.06.1984: 2 juvs (MNHN-Pa 1600).
— Stn CP 92, 34°24'N, 7°30'W. 1182 m. 8.06.1984: 1 juv. (MNHN-Pa 1601). — Stn CP 98. 34°29'N. 7°42'W. 1721-
1773 m, 9.06.1984: 2 juvs (MNHN-Pa 1602). — Stn CP 108. 36°H'N. 8°06'W. 1521-1534 m. 10.06.1984: 1 2
31.6 mm (MNHN-Pa 1603).
"Talisman": Cap Bogador, 11.07.1883, 1400 m: 1 damaged (MNHN).
Canary Is. Cancap 2: stn 27. 27°56'N, 14°28'W. 1440-1540 m, 25.08.1977: 2 juvs (RMNH).
Cancap 4: stn 4.056, 28°47'N, 13°22'W, 1306-1345 m, 18.05.1980: 2 3 28.5. 41.5 mm; 3 2 26.2-46.0 mm
(RMNH). — Stn 4.057, 28°47'N. 13°24'W, 1300-1335 m, 19.05.1980: 3 3 26.8-35.2 mm: 2 2 22.8. 28.0 mm
(RMNH).
Western Sahara. "Meteor": stn M36-98 AT149. 25°3I.5’N. 16°02.2'W. 658-888 m, 24,02.1975: 1 juv. (SMF
24645).
Mauritania. Tyro: stn 104, 19°43'N, 17°30'W. 1500 m. 17.06.1988: 1 3 42.9 mm; 1 2 53.2 mm (RMNH).
West Africa. "Pillsbury": stn 4, 5°37'N, 3°19'E, 1876-1263 m. 24.05.1964: 1 2 29.1 mm (RMNH). — Stn 44.
5°05'N. 4°00'W, 586-403 m. 30.05.1964: 1 juv. (RMNH). — Stn 52, 4°54'N. 4°58’W. 952-915 m. 31.05.1964: I 2
63.1 mm (RMNH). — Stn 53, 4°50'N. 4°55W, 1579-1519 m, 31.05.1964: 1 3 24.8 mm; 1 2 36.1 mm (RMNH). —
Stn 233, 5°19'N. 4°I4'E. 2013-1464 m, 12.05.1965: 1 3 29.3 mm; 6 2 28.9-33.1 mm (RMNH). — Stn 309. 4°15'N,
4°27’E, 678-720 m, 26.05.1965: 19 3 22.2-54.0 mm; 26 2 22.1-71.4 mm (RMNH).
342
B. GALIL
SE Atlantic. Off Cape province. 34°54.45'S. 18°12.06'E. 1650 m. 14.03.1988: 2 3 33.6, 43.1 mm (USNM
235172).
"Africana": 29038.8'S. 14°25.4'E, 861 m, 23.01.1990: 5 <3 32.7-41.5 mm; 5 2 25.2-50.5 mm (SAM). — 32°26'S.
I6°H'E, 1302 m. 27.01.1993: 1 5 30.7 mm (SAM). — 32°22'S, 16°09'E, 1150 m. 14.01.1995: 3 3 31.7-37 6 mnv
5 9 20.1-32.1 mm (SAM).
Labrador Sea. "Dana"-, stn 16439. off Hudson str., 62°49'N, 60°20'W. 980-1000m, 26.07.1974: 1 3 35 7 mm
(ZM CRU3275).
Canada. Nova Scoiia. "Epes Tarr": SE Sable I., 43°10'N, 61°20'W, 450 m. coll. Capt. Th. Olsen: I 3 38 2 mm
(USNM 39957).
United States. "Albatross" (id. S.I. Smith as Pentacheles sculptus ): stn 1123, off Massachusetts, 1417 m
26.08.1882: 1 <3 31.6 mm (AMNH 15234). — Stn 2115, 35°45'30"N. 74°34'45''W. 1517 m, 11.11.1883- 1 juv
20.6 mm (AMNH 15535). — Stn 2233, 38°36'30"N. 73°06'00"W. 1134 m, 12.09.1884: 1 <3 22.3 mm (AMNH 15745)
— Sin 2550, 39°44'30"N, 70°30'45"W, 1811 m, 9.08.1883: 1 $ 26.5 mm (AMNH 15445). — Stn 2628 950 m 1 885-
1 2 35.5 mm (AMNH 5074).
Off Florida. 448-450 m. 3.08.1932, coll. W.L. Schmitt: 1 3 32.6 mm (USNM 78272).
Gulf of Mexico. "Albatross": stn 2396. 28°34'N, 86°48'W. 603 m, 13.03.1885. id. M J Rathbun- 1 3 29 2 mm
(AMNH 3983).
"Cape Hatteras": 28°23.3'N. 86°43.8'W, 750-818 m, 23.04.1987, coll. H.G. Kuck & J.F. Quinn- 2 <3 41 7
43.1 mm; 1 9 38.0 mm (LACM 87-2691).
"Oregon": stn 2814. 28°53'N, 87°47'W. 1710-1890 m. 13.07.1960: 2 2 38.0. 39.0 mm; 3 juvs (USNM 232276). —
Stn 2816. 28°38'N. 88°18'W. 1710 m. 14.07,1960: 3 juvs (USNM 232276). — Stn 2820. 28°23’N. 88°21.5'W. 1800 m
15.07.1960: 5 2 26.5-39.0 mm (USNM 232276). — Stn 2824, 29°07'N, 88°04'W. 657-711 m, 17.07 I960- 2 <? ?? 4
29.2 mm; 2 2 42.1. 43.5 mm (USNM 232276). — Stn 4005, 29°07.5'N, 88°09'W. 540 m, 23.10.1962: 1 2 51 4 mnv
2 °v $ 43.8, 44.2 mm (USNM 273140). — Stn 4147, 24°12'N. 83°32'W. 900 m. 15.12.1962: I c3 28.5 mm;
3 2 27.1- 56.4 mm; 1 ov. 2 53.2 mm. 2 juvs (USNM). — Stn 4531, 24°32'N, 83°36'W. 360 m, 24 I 1 1963- ^ 5 469
51.2 mm; 2 juvs (USNM 279502). — Stn 4561, 24°03'N, 83°15'W, 900 m. 3.12.1963: 4 3 22 0-27 0- ’ 2 22 6'
31.9 mm; 1 juv. (USNM 260368). — Stn 4570. 23°11'N. 86°28'W, 900 m. 7.12.1963: 2 2 34.8. 41.3 mnv 1 ov 2
62.5 mm; 1 juv. (USNM 260368). — Sin 5790. 24°22'N. 83°34'W. 648 m, 26.11.1965: 3 3 31.5-44.0 mm (USNM
273140).
Caribbean Sea. "Oregon": stn 1907, 12°25'N, 82°23'W. 720-765 m, 11.09.1957: 2 2 33.7. 44.2 mm (USNM
251942). — Sin 2777. 11°36'N, 62°46'W, 522 m, 19.04.1960: 1 2 47.9 mm (USNM 232276). — Stn 3561. 16°35'N,
80°04'W, 720 m, 18.05.1962: 1 2 44.0 mm (USNM 244083). — Stn 3601, 9°07'N, 81°00’W. 720 m 31 05 1962- 4 2'
32.7-56.5 mm (USNM 244083). — Stn 4293, 7°14'N, 52°55'W, 720 m, 21.03.1963: 1 3 43.0 mm; 3 2 49 ’-60 8 mnv
I ov. 2 48.5 mm (USNM 273140). — Stn 4300, 7°44'N, 54°19'W, 540 m, 23.03.1963: 1 2 53 9 mnv 2 ov 2 53 0
55.6 mm (USNM 273140). — Stn 4412, 11°49'N. 69°24'W. 540 m, 3.10.1963: II 5 30.0- 39 6 mnv 12’ 2 28 7-
44.5 mm (USNM 273140). — Stn 4413, 11°53'N, 69°25'W. 630 m, 3.10.1963: 2 5 28.0-32.1 mm: 2 2 48.7.~62.8
,2„ juvs (USNM 260368); 4 3 28.7-37.8 mm; 2 2 43.5 mm, damaged; 3 ov. 2 43.4-50.1 mm; 4 juvs (USNM
f a™ 7,oS‘" ' ^lLn°54'N- 69°23'W' 720 3-10.1963: 2 3 30.0 34.6 mm; 1 2 damaged (USNM 260368). -
■? 57 W' 720 m' 6I01963; 3 5 22.1-30.4 mm; 3 2 27.7-50.5 mm (USNM 260368). — Stn 4441.
II 03 N, 67 53 W. 900 m, 9.10.1963: 1 2 70.3 mm; 1 ov. 2 65.6 mm (USNM 273140)
n>*ZZ[SbUryc: S“45£, °J? Guajira’ Colombia- l3°01'N- 2! °55'N. 27.07.1966: 1 <J 42.0 mm; I 2 42.4 mm; I juv.
(RMNH). — Stn 478, W. Grenadine Is, 11°34.4'N. 62°I0.7’W. 598-597 m. 2.08.1966: 6 3 23 5-46 1 mnv 5 2 ’3 3-
47.4 mm; 1 juv. (RMNH). - Stn 846, 11°37.48'N, 60°37.24'W, Tobago I., 659-1126 m. 2.07.1969 2 <5 25 4
Ilf mn74-, ?n29-2'5,0-5 mm; 2 Juvs (LACM ex AHF 1969-31). — Stn 850, 1I°45.30'N. 6I°29.30'W. Tobago I . 896-
923 m, 3.07.1969: 2 6 24.4. 29.1 mm; 3 2 30.7-40.5 mm (LACM ex AHF 1969-31).
Guyana. "Luymes": stn 48, 7°45'N. 57cOFW, 500 m. 30.08.1970: 1 2 41.6 mm (RMNH)
Ch*1f- "A",on Bc.n,un": stn 687- 34°07’s- 72°19’W, 750-730 m. 5.08.1966: 3 <3 25.4-30.9 mm; 1 2 26 2 mnv
3juvs (USNM). - Sin 703. 32°09'S, 71°43'W. 960 m, 12.08.1966: 3 5 25.4-26.4 mm; 4 2 31.7-32.4 mm (USNM) -
?Z02T966: f 2 ' * 26J ' *"•»* (USNM)‘ “ 0ff praise. 400 m.
163°36.7'E, 1186 m, 25.09.1982: 1 3 26.2 mm (NIWA) ’ '8S'
Tasman Sea. "Kamenskoe": 40°32'S, 167°23.5'E, 1140 m. 9.11.1977: 1 <3 30.3 mm; 1 juv. (ZMMU)
Vanuatu. Musorstom 8: stn 1075, 15°53.26 S, 167027.21'E. 956-944 m, 4.10.1994- 4 2 ’3 3-36 5 mm (MNHN
Pa ,605). - Stn 1076, 15“53.81'S, 167“30.42'E, 1100-119. m. 4.10.1991: 7 juvs (MNHN-Pa 17.4)
Australia. New South Wales. "Kapala": stn K77-23-13, 33°27'S, 152°09'E. 882-914 m, 8.12.1977. id. D Brown
Ai& iQMH??uuCUJpta: 2 48;8 mm; 2 JUVS' (AM P40365F — Sln K88-08-04. 32°06'S. 153°08'E. 1025-1080 m.
4.U6.iy88, id. H.E. Stoddar r as Stereomastis sculpta: 1 2 41.9 mm (AM 40363).
Japan. "Albatross": stn 3696, off Honshu I., 902-1346 m. 5.05.1900: 1 2 35.2 mm (USNM).
Source MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
343
East China Sea. "Yuryo Maru": 29°38.92,N, 127°55.95'E, 750 m, 4.02.1978: 2 ov. $ 51.6. 56.0 mm (NUM
crl 0229).
Philippines. "Albatross": sin 5468, Lagonoy Gulf, 13°35'39"N. 123°40'28'E, 1040 m. 18.06.1909: I 2 66.8 mm
(USNM). — Sin 5470, Lagonoy Gulf, 13°37'30"N, 123o41'09"E. 1024 m, 18.06.1909: 1 2 51.3 mm (USNM).
Indonesia. "Albatross": stn 5607, Sulawesi, 00°04'S, 121°36'00"E, 1392 m, 18.1 1.1 909: 1 juv. (USNM). —
Sin 5619, Molucca Passage, 00°35'00"N, 1 27°14'40"E, 795 m, 27.11.1909: 2 juvs (USNM). — Sin 5638. Moluccas.
3°47'15"S. 126°23'40"E, 946 m, 10.12.1909: 1 2 24.3 mm (USNM). — Sin 5648. Buion sir., 5°35'00"S, 122°20’00"E.
1022 m. 16.12.1909: I juv. (USNM). — Sin 5654, Gulf of Boni, 3°42'00"S, I20°45'50"E, 1472 m. 18.12.1909: 1 juv.
(USNM). — Stn 5657, Gulf of Boni, 3°19'40''S, 120°36'30"E, 898 m. 19.12.1909: 1 juv. (USNM). — Stn 5658. Gulf of
Boni, 3°32'40"S, 120°31'30"E, 933 m, 19.12.1909: 1 ov. 2 56.3 mm (USNM).
"Siboga": stn 18, 7°28'02"S, 115o24'06"E. 1018 m. 18.03.1899: 1 3 40.6 mm (ZMA). — Sin 211, 5°40'07"S.
120°45'05"E, 1158 m, 25.09.1899: 1 juv. (ZMA).
KarubaR: Tanimbar Is: sin CP 52, 8°03'S, 131°48'E, 1244-1266 m. 30.10.1991: 2 juvs (MNHN-Pa 1715). —
Stn CP 54, 8°21'S, 131°43'E, 836-869 m, 30.10.1991: 1 3 37.4 mm (MNHN-Pa 1716). — Stn CP 73, 8°29'S. 131°33'E,
855-840 m. 2.11.1991: 1 3 32.8 mm (MNHN-Pa 1606). — Sin CP 81. 9°35'S. 131°02’E. 200-207 m, 4.11.1991: 1 3
40.4 mm (MNHN-Pa 1607).
Bali Sea. "Galathea": stn 489, 7°38'S, 116°08'E. 1195-1165 m, 13.09.1951: 1 <5 40.2 mm (ZMC).
Ceylon. "Investigator": stn 321, 5°04'08"N. 80°22'E. 1208 m, 15.11.1903: 1 ov. 2 59.5 mm (IM 4003/10).
Gulf of Aden. "Meteor 5": stn Me5-268, 13°1I.2'N, 47°10.9'E. 1654-1686 m. 13.03.1987: 1 3 27.2 mm; 1 2
42.2 mm; 1 juv. (SMF 24651). — Stn Me5-277. 12°37.5'N, 46°09.4'E, 1552-1554 m, 14.03.1987: 1 <3 32.9 mm; I 2
46.7 mm; 2 juvs (SMF 24650).
Comoro Is. Benthedi: stn CA 31, 12°37.4’S, 45°25.2'E. 1800 m. 25.03.1977: 2 3 20.4. 24.4 mm (MNHN-Pa
1709).
Madagascar. "Vaitban" (A. Crosnier coll.): stn CH 126, 17°50'S. 43°07'E. 1475-1530 m. 16.1.1975: 1 3
29.1 mm; 1 2 31.7 mm (MNHN-Pa 1717). — Stn CH 133, 13°02'S. 48°02'E. 1000-1525 m. 21.01.1975: I 3 42.4 mm
(MNHN-Pa 1608). — Stn CH 142, 13°45.6'S, 47°34.2'E, 1250-1300 m, 29.02.1975: 1 juv. (MNHN).
Fig. 24. — Polycheles sculptus Smith, 1880. 3 42.4 mm. Madagascar, 13°02’S. 48°02'E. 1000-1525 m (MNHN-Pa
1608): a. dorsal view of the cephalothorax; b, lateral view of the abdomen.
344
B GALIL
Type. — The male (38.2 mm cl) (USNM 39957) collected by the "Epes Tan " off Nova Scotia. Canada
(43°10'N. 61°20'W. 450 m) is the holotype of Polycheles sculptus Smith. 1880.
Four syntypes of Pentacheles spinosus A. Milne Edwards, 1880 are at the Museum of Comparative Zoology
(Harvard) under the registration numbers MCZ 2794 {"Blake" stn 29, 2 specimens), MCZ 2795 ("Blake" stn 175,
1 specimen), MCZ 3076 ("Blake" stn 162. 1 specimen).
DESCRIPTION. — Dorsal surface of carapace setose, minutely granulate. Frontal margin straight, rostrum bifid.
Internal angle of orbital sinus minutely spinose, external angle unarmed. Eyestalk bearing papilla, curved distad.
Basal antennular segment produced anteriorly to a sharp point, mesial margin smooth, anterolateral margin
bispinose. Spines on lateral margins of carapace upcurved, anteriormost largest. Spine formula of lateral margins
of carapace 5-7:3:6-7; spines successively further apart posteriorly. Spine formula of median postrostral carina
(excluding rostral spines) 1:2:1; median postcervical carina bearing two pairs of spines. Gastro-orbital region with
crescentic row of five antrorse spines, anteriormost largest. Anterior margin of cervical groove with antrorse spine
between median postcervical and branchial carinae. Branchial carina prominent, parallel to lateral margin, with
5-7 spines, postcriormost largest; region between branchial and median postcervical carinae unarmed, obliquely
grooved anteriorly. Posterior margin of carapace bearing pair of antrorse submedian spines.
First abdominal tergite bearing two pairs of prominent spines laterally on anterior margin. Abdominal lergites
medially carinate. Carinae on lergites 1-5 bearing antrorse spines, spines 1-4 increasing in size posteriorly; antrorse
spine on fifth tergite smaller than fourth. Sixth tergite bearing mesially parallel pair of rounded carinae, confluent
posteriorly. Tergitcs 2-5 with transversely oblique submedian grooves. Pleura 2-5 with curved rib mesially,
margins minutely granulate. Anterior margin of second pleuron produced, triangulate. Telson with median crest
anteriorly, two minutely serrulate, convergent carinae posteriorly.
Merus of PI proximally bispinose, claw-like spine subterminally on upper margin, finely spinulose on lower
margin; carpus half as long as merus with subterminal spines on upper and lower margins; upper margin of
propodus with subterminal spine, lower margin with two minutely spinulate rows. P5 subchelate in male, chelate
in female.
Color. — "rosy pink" (ALCOCK, 1901b: 171). In BouviER's color plate of P. sculptus (1917. pi. 3 fig. I) the
carapace and abdomen of P. sculptus are ivory colored with spines, tail fan. antennae and legs light orange red.
Bouvier (1925: 439) described it as "rouge rose presque uniforme".
Remarks. — Smith's description of P. sculptus (1880a, 1882) is both detailed and meticulous. A. Milne
Edwards' (1880) description of Pentacheles spinosus , though brief, allowed ALCOCK (1901b) to synonymize it
with sculptus.
The "magnificent specimen of that strange blind crustacean" collected in the Mediterranean and described by
Giglioli (1881: 358) as differing "only in one or two minor details, which may be sexual differences" from
Willemoesia leptodactyla, is in fact P. sculptus.
Distribution. — Worldwide: Mediterranean, Ibero-Moroccan Gulf, Canary Is, Mauritania. Cape Verde Is,
West Africa, South Africa, Labrador Sea, Iceland. Canada (Nova Scotia), United States. Gulf of Mexico. Caribbean
Sea, Guyana, Chile, New Zealand, Tasman Sea.Vanuatu, Australia. Japan, China Sea, Philippines, Indonesia,
Malay Archipelago, Ceylon, Arabian Sea, Gulf of Aden, Comoro Is, Madagascar. East Africa; 200 m (present
work) - 4000 m (Stephensen, 1923).
Polycheles suhmi (Bate, 1878)
Fig. 25
Pentacheles sulmii Bate, 1878a: 278; 1878b: 484; 1878c: 563. — Norman, 1879: 180. — Holthuis, 1962- 183 —
China. 1964: 111.
Stereomastis suhmi - Bate, 1888: 154. figs 13. 37-38, pi. 15 figs 3-4. — Stebbing. 1893: 200. — De Man, 1916:
5 (list). — Sund, 1920: 223. — Calman, 1925: 19. — Barnard. 1950: 574. fig. 105f. — Holthuis. 1952a: 78. —
Source MNHN. Parts
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
345
Bernard, 1953: 87. — Bahamonde, 1963: 4. — Zarenkov, 1964: 8; 1969: 81. — Firth & Pequegnat, 1971: 72. —
Retamal, 1974: 6; 1981: 16. — Baez & Andrade, 1979: 225, pi. I fig. 5. — Andrade & Baez, 1980: 262. —
Kensley, 1981b: 29. — Macpherson. 1983: 48 fig. 27; 1984: 74; 1991: 405. — Andrade. 1986: 45. 50. —
Takeda, 1990: 358. — Tiefenbacher. 1994: 17. fig. 1. — Griffin & Stoddart, 1995: 249, figs 12-15. — Dawson
1997: 23.
Polycheles suhmi - Bouvier, 1917: 35.
Sieromaslis suhmi - Wicksten, 1989: 304 [erroneous spelling].
Material EXAMINED. — SE Atlantic. Off Cape province. "Africana": 32°50'S. 16°45'E. 495 m. 25.08.1988:
1 ov. 2 28.5 mm (SAM). — 29°50.0'S, 1 5C03. 1 E. 345 m, 12.08.1989: 1 3 18.7 mm; 1 2 23.1 mm (SAM). —
30°05'S, I4°54.1'E, 465 m. 13.08.1989: 1 2 27.0 mm (SAM). — 32°20.4'S. I6°31.7'E. 367 m, 14.01.1990' I 3 18 2
mm (SAM). — 34°14'S. 17°39'E. 419 m, 12.02.1992: 3 ov. 2 27.7-30.5 mm (SAM). — 34°34'S. 17°57'E. 438 m.
12.02.1992: 1 ov. 2 31.4 mm (SAM). — 31°20'S, 16°09'E, 462 m, 20.02.1992: 2 3 20.5, 23.1 mm; 1 2 31.2 mm;
1 ov. 2 26.5 mm (SAM). — 29°56'S, 14°51'E, 446 m. 22.02.1992: 1 3 23.6 mm; 1 ov. 2 27.0 mm (SAM). — 34°12'S.
1 7°37'E, 416 m, 11.01.1995: 1 3 25.1 mm; 2 ov. 2 28.6, 22.6 mm (SAM). — 31°45'S. 15°42'E. 468 m, 25.01.1995:
1 2 27.3 mm; 2 ov. 2 26.7, 24.2 mm (SAM). — 35°14'S, 18°42’E. 424 m, 5.02.1996: 4 3 21.2-26.0 mm; 4 2 22 9-
29.5 mm (SAM).
33°20'S, 1 7°32.48'E, 451 m, 10.03.1988, coll. & id. B. Kensley: 2 3 18.7, 18.9 mm; 2 2 20.3, 28.8 mm; 10 ov 2
25.5-29.4 mm (USNM 235173).
Discovery Seamount. "Prof. Mesyatsev stn 154, 760 m, 10.12.1979, coll. M.V. Bondarenko: 1 3 18.3 mm; 2 2
20.9. 20.1 mm (ZMMU).
Chile. " Challenger stn 305b. Gulf of Penas. 47°48'S. 74°46'W, 292 m. 1.01.1876: 1 3 17.1 mm (NHM 1888.22).
— Stn 311, 52°45'30"S. 73°46'W, 441 m, 11.01.1876: 5 3 15.2-18.6 mm; 4 2 17.3-22.3 mm (NHM 1888.22).
"Albatross": stn 2780. 53°01'S, 73°42'30"W, 664 m. 2.02.1888, id. W.L. Schmitt; 4 3 19.9-22.6 5 mm; 5 2 17.2-
32.4 mm (USNM 58641). — Stn 2781, 51°52’S, 73°41’W. 626 m, 4.02.1888. id. W.L. Schmitt: 1 damaged (USNM
58642). — Stn 2782, 51°12'S. 74°13’05"W, 464 m. 6.02.1888, id. M.J. RaTHBUN: 3 3 21.1-23.6 mm; 4 2 22.7-
37.7 mm; i juv. (USNM 28348). — Stn 2784, 48°4I'S, 74°24'W. 349 m, 8.02.1888. id. M.J. Rathbun: 2 3 19.5,
26.8 mm; 3 2 28.8-29.3 mm; 1 ov. 2 26.6 mm (USNM 28349). — Stn 2785. 48°09'S, 74°36'W. 808 m. 8.02.1888,
id. M.J. Rathbun: 1 2 19.3 mm (USNM 58643).
Valparaiso. 10.02.1956: 2 ov. 2 24.5, 20.6 mm (USNM 99776); 1 3 21.3 mm (RMNH).
"Eltanin stn 1605, 52°53'S, 74°05'W, 522-544 m, 1.04.1966: 1 <J 22.1 mm; I 2 27.1 mm; 1 ov. 2 27.7 mm
(USNM).
"Anton Bn, un": stn 687. 34°07'S. 72°19'W, 750-730 m, 5.08.1966: 1 2 21.3 mm (USNM).
New Zealand. NZOI: stn E747, 40°43.2'S, 176°48.4'E, 548-563 m. 29.03.1967: 1 2 34.6 mm (N1WA). —
Stn PI 20. 35°45.7'S, 165°04.1'E, 950 m, 3.06.1977: 1 juv. (NIWA). — Stn 1699. 48°16.0'S, 1 79°E, 532 m,
19.03.1979: 1 2 20.1 mm; 2 juvs (NIWA). — Stn P927, 40°50.1'S, 168°14.8'E. 1009-1005 m. 18.04.1980: 1 3
25.9 mm (NIWA) — Stn T33, 48°30.2,S, 179°43.5’W, 721-725 m, 13.03.1981: 1 juv. (NIWA). — Stn V227, 39°33.9'S.
1 69°04.7'E, 604 m, 18.10.1982: 1 3 15.4 mm (NIWA). — Stn V391. 42°49.97'S, 1 76°59.9 1 E. 476 m. 17.09.1989:
1 3 19.3 mm (NIWA). — Stn X5I0, 42°56.29'S. 175°41.39'W. 856-860 m. 10.07.1994: 1 3 19.2 mm (NIWA). —
Stn Z8974, 37°01.29'S. 176°19.58'E, 566-578 m. 11.04.1996: 1 3 22.9 mm (NIWA). — Sin Z8566. 43°04'S.
1 75°39’E, 460 m, 1.10.1996: 5 2 35.2-39.0 mm (NIWA). — Stn Z8990, 37°3 1.70'S, 176°46.96'E. 523-527 m.
17.01.1998: 1 c? 25.1 mm (NIWA). — Stn Z9014, 36°54.78'S. 176°20.43'E. 588-599 m, 23.01.1998: I 2 19.8 mm
(NIWA).
Tasman Sea. "Galathea": stn 626. 42°10'S, 170°I0’E, 610 m. 20.01.1952: 2 3 16.8, 26.5 mm; 6 2 17.9-31 I mm
(ZMC).
Australia. SE Port Hunter. "Kapala": stn K77-23-10, 33°11'S, 152°24'E. 732 m. 7.12.1977, id. D.E. Brown: I 3
26.8 mm; 1 ov. 2 30.8 mm (AM P26756). — Stn K87-23-03, 34°42'S. 1 5 1 ° 1 6E. 760-855 m, 3.12.1987.
id. H.E. Stoddart: 1 2 37.4 mm (AM P38726).
Type. — The male (17.1 mm cl) (NHM 1888.22) collected by the "Challenger" at stn 305b in the Gulf of
Penas (47°48'S. 74°46'W. 292 m), is the holotypc.
DESCRIPTION. — Dorsal surface of carapace with short setae, minutely granulate. Anterior margin of carapace
straight, two obliquely set rostral spines. Internal angle of orbital notch bearing spine, external angle unarmed. Eye
stalk with short spine, curved distad. Mesial margin of basal antennular segment with 2-3 spines, antero-external
angle bearing two long spines. Spines on lateral margins of carapace upcurved. diminishing in size posteriorly,
spine formula 5-6:2-3:7-9. Spine formula of mid-dorsal carina (excluding rostral spines) 1: 1:2:2: 1 before cervical
groove, 2,2,2 behind cervical groove. Posterior margin of cervical groove with three spines midway between
346
B. GAUL
median, branchial carinae. Gastro-orbital region with quadrispinose row. anteriormost largest, followed by cluster
of 5-6 spines. Superior branchial carina prominently spinose, 7 spines between median and branchial carinae.
Posterior margin of carapace with pair of submedian spines.
Anterior margin of first abdominal tergite laterally with two pairs of prominent spines, antrorse median spine.
Median carina on abdominal tergites 2-5 bicuspidale, anteriorly spinose; sixth tergite with prominently denticulate
double carina joined posteriorly in larger denticle. Terga 2-5 bearing shallow oblique grooves on each side oi
median carina. Pleura with cresentic rib mesially, margins sparsely denticulate, ventrally acute. Anterior margin of
second pleuron produced, bearing prominent spine. Telson with median denticulate crest anteriorly, serrulate carinae
posteriorly.
Merus of PI with single proximal spine and subterminal claw-like spine on upper margin, lower margin
smooth. Carpus half as long as merus, subterminal spine on upper and lower margins. Margins of chela smooth.
P5 subchelate in male, chelate in female.
Color. — "Carapace, abdominal segments and telson a uniform pale whitish pink; all spines of carapace and
abdominal terga a darker rose pink. Endopods and inner three-quarters of exopods of uropods whitish (almost
transparent), entire outer edge of exopods trimmed in dark rose pink. Ischium, anterior portion of merus and palm
of propodus of cheliped pale, almost white; distal end of merus and carpus pale pink; fingers of cheliped also pale
pink, fading to w'hite at tips." (GRIFFIN & STODDART, 1995: 250).
Remarks. — P. suhrni is easily distinguished from its congeners in having a cluster of 5 or 6 spinules
posteriorly on the gastric region, bicuspid median carinae on the abdominal tergites 2-5, the posterior cusp more
prominent on fourth and fifth tergites.
Distribution. — Southern Ocean: South Africa. Chile. New Zealand. Australia; 292 (present work) - 2195 m
(Calman, 1925).
Fig. 25. — Polycheles suhmi IBate, 1878): a, 9 31.2 mm, off Cape Province, "Africana", 31°20’S, 16°09'E, 462 m
(SAM), dorsal view of the cephalothorax; b. 6 26.5 mm. Tasman Sea. "Galathea" stn 626. 42°10’S, 170°10'E, 610 m
(ZMC), lateral view of the abdomen.
Source MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLY C HELID AE
347
Polycheles surdus sp. nov.
Fig. 26
Polycheles phosphorus RATHBUN, 1906: 898 [non Alcock, 1894J.
Stereomastis phosphorus - Griffin & Stoddart. 1995: 246. figs 9-1 1 (p.p.).
Material EXAMINED. — Southeast Pacific. Nazca Ridge. "Prof. Mesvatsev": 24°58'S, 88°28'W 565 m
1.09.1983: 2 2 45.8, 55.7 mm (ZMMU).
"Prof. Stockman": stn 1964. 580-564 m. 30.04.1987: 3 8 38.2-45.7 mm; 1 ov. 2 47.8 mm (ZMMU). — Sin 1965.
565-570 m. 30.04.1987: 1 $ 43.1 mm; 2 2 43.3, 27.6 mm (ZMMU). — Sin 1971. 570-580 m. 1.05.1987: 2 6 28.5.
39.4 mm; 1 2 49.4 mm; 7 juvs (ZMMU). — Sin 1976. 569-590 m, 1.05.1987: 1 6 38.2 mm; I 2 33.3 mm; 1 ov. 2
45.9 mm. 5 juvs (ZMMU). — Stn 1977, 545-600 m, 1-2.05.1987: 1 <3 28.8 mm; 4 juvs (ZMMU). — Sm 1996. 750-
800 m, 5.05.1987: 2 6 31.9, 39.9 mm; 1 ov. 2 52.0 mm; 1 juvs (ZMMU).
Sala-v-Gomez Ridge. "Prof. Stockman": sm 2018, 760 m, 7.05.1978,: 1 6 44.8 mm; 1 2 49.2 mm; 9 juvs (ZMMU).
— Stn 2019. 25°05.6'S, 99°28.9'W, 750 m, 7.05.1978. colls Mironov & Zarenkov: 1 2 45.4 mm (ZMMU).
Marquesas Is. MUSORSTOM 9: stn CP 1191, 8°46.3'S. I40°07.2'W. 350 m 26.08.1997: 1 ov. 2 35.1 mm (MNHN-
Pa 1688). — Stn CP 1271, 7°53.6'S, 140°42.2'W 600 m. 4.09.1997: 4 juvs (MNHN-Pa 1684). — Stn CP 1272. 7°55.5'S.
140°43.6'W 660-680 m. 4.09.1997: 1 <3 31.7 mm; 1 ov. 2 39.3 mm (MNHN-Pa 1683). — Stn CP 1276. 7°51.9'S.
1 40°38.2'W, 800-805 m, 5.09.1997: 3 juvs (MNHN-Pa 1685). — Stn CP 1278. 7°52.)'S. 140°38.6'W, 1000 m.
6.09.1997: 1 juv. (MNHN-Pa 1687). — Stn CP 1307, 8°57.9'S. I40°15.8'W, 708-738 m. 10.09.1997: I juv. (MNHN-Pa
1686).
Hawaii. "Albatross"! all the specimens identified as P. phosphorus by M.J. Rathbun): stn 3467. Kauai Channel,
21°I3'N, 157°43'37"W, 558 m, 3.12.1891: 1 2 39.4 mm (USNM 30273). — Stn 3476, Kauai Channel, 21°09'N.
1 57°53'W. 536 m. 6.12.1891: 1 2 43.6 mm (USNM 30274). — Stn 3816. Oahu l„ surface, 29.03.1902: 1 6 39.1 mm
(USNM 30275). — Stn 3824, Molokai I„ 400-880 m, 1.04.1902: 1 ov. 2 52.8 mm (USNM 30276). — Stn 3836.
Molokai I.. 428-459 m, 3.04.1902: I <3 38.5 mm (USNM 30277). — Stn 3839. Molokai I.. 466-479 m. 4.04.1902: 1 <3
33.8 mm (USNM 30278). — Stn 3867, Pailolo Channel. 511-522 m. 10.04.1902: 1 <3 33.6 mm: 3 2 31.1-58.0 mm
(USNM 30282). — Stn 3883, Pailolo Channel. 499-511 m. 16.04.1902: 1 6 38.0 mm (USNM 30283). — Stn 3884,
Pailolo Channel. 511-522 m. 16.04.1902: 1 <3 47.6 mm (USNM 30284). — Stn 3907. Oahu I., Diamond Head. 547-
567 m, 5.05.1902: 1 <3 39.4 mm (USNM 30286). — Stn 3910. Oahu 1.. Diamond Head. 560-607 m. 5.05.1902: 2 8
37.0, 49.7 mm; 1 2 39.9 mm (USNM 30287). — Stn 3911, Oahu 1.. Diamond Head. 601-607 m. 5.05.1902: 1 <3
43.1 mm (USNM 30288). — Stn 3917, Oahu I.. Diamond Head. 529-594 m, 6.05.1902: 1 2 26.8 mm (USNM 30289).
— Stn 3920, Oahu I.. Diamond Head, 477-504 m. 6.05.1902: 1 2 39.4 mm (USNM 30290). — Stn 3988. Kauai I.. 297-
844 m, 11.06.1902: I 8 31.5 mm; 1 2 48.6 mm (USNM 30296). — Stn 4028. Kauai I.. 799-862 m. 24.06.1902: 1 <3
21.5 mm; 1 2 46.0 mm (USNM 30300). — Stn 4097, Pailolo Channel. Mokuhooniki Islet. 515 m. 22.07.1902: I <3
43.7 mm (USNM 30285). — Stn 4187. Kauai 1„ 914-1265 m. 13.08.1902: 1 2 41.2 mm (USNM 30314).
Banc Tuscarora. Musorstom 7: sin CP 562, 11°48'S, 178°22'W, 775-777 m, 19.05.1992: 2 8 49.0. 49.5 mm;
1 2 45.1 mm (MNHN-Pa 1610). — Stn CC 550, 12°15'S, 177°28'W. 800-810 m. 18.05.1992: 1 2 27.8 mm (MNHN-Pa
1718). — Stn CC 554. 1 2°14'S. 177°28'W, 795-820 m. 18.05.1992: I juv. (MNHN-Pa 1719). — Stn CP 567. 1 1 °47 S,
1 78°27'W. 1010-1020 m, 20.05.1992: 1 juv. (MNHN-Pa 1720).
Banc Combe. Musorstom 7: stn CC 553. 12°17'S. 177°28'W. 780-794 m. 18.05.1992: 2 8 46.6. 34.7 mm: 1 2
45.0 mm (MNHN-Pa 1609).
Banc Field. Musorstom 7: stn CP 592, 12°32’S, 174°22'W. 730-775 m. 24.05.1992: 3 juvs (MNHN-Pa 1721).
Vanuatu. Musorstom 8: stn CP 991, 1 8°5 1 .26'S, 168°52.I9'E. 936-910 m, 24.09.1994: I <3 29.4 m: 1 juv.
(MNHN-Pa 1724). — Stn CP 996, 18°52'S, I68°56'E. 764-786 m, 24.09.1994: 1 juv. (MNHN-Pa 1725). — Stn CP 1008.
1 8°53.29’S, 1 68°52.65'E, 919-1000 m. 25.09.1994: 4 juvs (MNHN-Pa 1726). — Stn CC 1035. I7°58.02’S.
168°44.06'E, 765-780 m, 29.09.1994: 1 2 44.7 mm (MNHN-Pa 1727).
New Caledonia. B iogeocal: stn CP 290, 20°36.91 'S. I67°03.34'E. 920-760 m, 27.04.1987: 1 2 28.4 mm
(MNHN-Pa 1746).
Bathus 2: stn CP 751. 22°24.35’S, 166°12.83E, 1 300- 1500m. 15.05.1993: 1 2 26.8 mm (MNHN-Pa 1747). —
Stn CP 765. 22°09'S, 166°02'E, 600-630 m, 17.05.1993: 1 8 29.7 mm (MNHN-Pa 1722).
Bathus 4: stn CP 950, 20°3I.93'S. 164°56.H'E. 705-750 m. 10.08.1994: I 8 37.7 mm; I 2 54.6 mm (MNHN-
Pa 1728).
New Zealand. NZOI: stn P927, 40°50.I'S. 168°14.8'E. 1009-1005 m. 18.04.1980: 1 juv. (NIWA). — Stn Z8256.
36°09.1'S, 176°15.5'E. 470 m. January 1994: 1 8 25.6 mm (NIWA). — Stn Z8267. 36°49.7'S. 1 76° 1 8.3 E. 510 m.
December 1994: 1 2 50.6 mm (NIWA). — Stn Z8268. 36°52.9'S. 1 76°1 9. 1 E. 520 m. December 1994: 2 2 43 1,
43.8 mm (NIWA). — Stn Z28996, 37°23.23'S, 176°32.85'E, 525-552 m, 19.01.1998: 1 2 52.6 mm (NIWA). —
Stn Z9007. 37°08.61'S, I76°19.64’E, 472-475 m. 21.01.1998: 2 8 31.1. 30.3 mm; 1 2 49.0 mm (NIWA). —
Stn Z9008, 37°09.87'S, 176°21.74'E. 518-536 m, 22.01.1998: 1 <5 34.3 mm; 1 2 29.2 mm (NIWA). — Stn Z9009.
348
B. GAL1L
37°13.40'S, 176°14.37'E, 224-264 m, 22.01.1998: 1 8 damaged (NIWA) — Stn Z9014. 36°54.78'S. I76°20.43'E, 588-
599 m, 23.01.1998: I 8 32.4 mm (NIWA). — Stn Z9017, 36°46.53'S. 176°16.56'E. 465-474 m. 23.01.1998: I 2
41.8 mm (NIWA)
Australia. New South Wales. "Kapala": stn K78-23-08. 28°03'S, 154°04’E, 732 m. 6.11.1978. id. H.E. Stoddart
as Stereomastis phosphorus: 1 8 42.4 mm; 1 2 49.4 mm (AM P44915).
Indonesia Makassar Strait. Corindon 2: stn 209. 00°07'S. 117°53'E. 490 m. 31.10.1980: 1 2 45.9 mm (MNHN).
Indian Ocean . Mascarene Ridge."Vitiaz": stn 2817. 9°26'S, 60°05'E. 350-430 m. 10.01.1989: 1 ov. 2 45.0 mm
(USNM). — Stn 2819, 8°48'S, 59°47'E. 525-530 m. 10.01.1989: 2 2 27.3. 32.0 mm (USNM).
Reunion "Marion Dufresne stn CP 113. 20°47.8'S, 55°08.5'E. 712-790 m. 31.08.1982: 2 8 28.5. 40.0 mm
(MNHN). — Stn CP 120. 20°54'S, 55°14.4'E, 940-1 180 m, 31.08.1982: I 8 28.1 mm; 2 2 damaged (MNHN).
Madagascar. "Vauban" (coll. A. Crosnier): stn 48. 1 5° 1 8'S. 46°1 2. 1’E. 480-510 m, 8.11.1972: 2 8 32.6.
42.4 mm; 5 2 46.6-56.7 mm (MNHN-Pa 1613). - Stn 59. 23°36'S, 43°29.6'E. 600-610 m. 27.02.1973: 1 <3 37.0 mm.
1 2 50.9 mm (MNHN-Pa 1614). — No stn number, 21°17'S. 43°23'E. 490-530 m. 6.06.1973: 2 <J 32.0. 45.8 mm; 1 2
53.5 mm (MNHN-Pa 1615). — Stn CH 133. 13°02'S, 48°02'E. 1000-1525 m. 21.01.1975: 3 juvs (MNHN-Pa 1612). —
Stn CH 139, 13°50'S. 47°37'E. 850-1125 m. 27.02.1975: 4 juvs (MNHN-Pa 1616). — Stn CH 142. 13°45.6'S.
47°34.2'E. 1250-1300 m. 28.02.1975: 10 juvs (MNHN-Pa 1617).
Mascareignes ///"(coll. R. Cleva): stn 5, 22°25'S. 43°04.8'E. 500-520 m. 21.12.1985: 3 <3 49.1-58.0 mm (MNHN-
Pa 1145). — Stn 7. 22°26.rS. 43°06.3'E. 400-425 m. 21-22.12.1985: 1 2 50.4 mm (MNHN-Pa 1147). — Stn 9.
22°25.3'S, 43°04.9'E. 525-550 m. 22.12.1985: 3 2 42.4-56.3 mm (MNHN-Pa 1148). — Stn 12, 22°23.5'S. 43°04.9'E.
400-425 m, 23.12.1985: 1 juv. (MNHN-Pa 1142). — Stn 25. 22°23.5'S. 43°05'E. 500-520 m, 15.01.1986: 3 8 48.4-
57.7 mm; 2 2 31.1. 57.8 mm (MNHN-Pa 1144). — Stn 26. 22°18.7'S. 43°04.1'E. 450-600 m. 15.01.1986: 2 8 43.0.
49.3 mm (MNHN-Pa 1143). — Stn 32. 22°25.8’S. 43°04,3'E. 450-475 m. 19.01.1986: 1 8 52.7 mm (MNHN-Pa 1 149).
"Mascareignes ///"(coll. R. VON Cosel): stn CH 55. 22°17.2'S. 43°03.4'E. 430 m. 16.10.1986: 1 8 50.7 mm
(MNHN Pa 1618). — Stn CH 60. 22°25.6'S, 43°06.2'E, 475 m. 18.10.1986: 1 8 50.9 mm (MNHN-Pa 1619). —
Stn CH 63. 22°26.8'S. 43°05.4'E. 530 m. 20.10.1986: 1 <3 44.8 mm; 1 damaged (MNHN-Pa 1620). — Stn CH 113.
22°1 1.35'S. 43°02.3'E. 650 m. 27.11.1986: 5 juvs (MNHN-Pa 1621).
Mozambique Channel. "Anton Bruttn": sin 398, 22°25'S. 35°54'E, 740 m. 1.10.1964: 1 2 26.9 mm (USNM).
Mozambique. "Algoa": 18°14'S. 37°31'E, 472 m, 17.06.1994: 2 8 39.5. 38.1 mm (SAM A41702); I 2 44.7 mm
(SAM A41700). — 19°27'S. 36°51'E, 486 m. 14.06.1994: I 2 39.6 mm (SAM A4I703). — 16°20'S. 40°08'E. 500 m.
16.06.1994: 1 ov. 2 51.8 mm (SAM A41701).
Types. — The male (39.5 mm cl) (SAM A41 702) collected off Mozambique ( 1 8° 1 4'S. 37°31’E. 472 m) is the
holotype; the smaller male from that sample is a paratype.
Description. — Carapace oblong, lateral margins subparallel, slightly convergent anteriorly. Dorsal surface of
carapace tomentose, minutely granulose. Anterior margin of carapace straight. Median submarginal tooth upeurved,
in front of bifid rostrum. Internal angle of orbital sinus unispinose. external angle unarmed. Eyestalk bearing
mammilate spine, curved distad. Basal antennular segment produced anteriorly to a sharp point, mesial margin
smooth; antero-external margin curved, bispinose, anterior spine longer than posterior. Spines formula ot lateral
margins of carapace 6-7:3-4:10-14; spines decreasing successively in size posteriorly. Spine formula of median
postrostral carina (excluding rostral spines) 1 : 1 :2: 1 ; median postcervical carina bearing two pairs of spines.
Posterior margin of cervical groove with 3 or 4 antrorse spines midway between median postcervical and branchial
carinae. Gastro-orbital region bearing five spines in curved row. Branchial carina prominent, sinuous, bearing five
spinules, posteriormost largest. Carapace between branchial and median carinae minutely granulate. Posterior
margin of carapace bearing two prominent, antrorse submedian spines.
Anterior margin of first abdominal tergite bearing two pairs of lateral spines. Abdominal tergites medially
carinate. Carinae of tergite 1-2 bearing small antrorse spine; carina of tergite 3 with a prominent, salient spine
longer than the spine of the fourth; carina of tergite 5 simple. Sixth tergite bearing low, granulate, double carinae,
confluent posteriorly in rounded knob. Tergites 2-5 with submedian grooves obsolescent. Pleura 2-4 with curved
rib mesially. Margins of pleura denticulate; second pleuron anteriorly produced, ovate. Tclson with short,
denticulate crest anteriorly, two minutely serrulate submedian carinae posteriorly.
Merus of PI bispinose proximally, claw-like spine subterminally on upper margin, occasionally spinulose on
lower margin; carpus more than half as long as merus, with prominent spines subterminally on upper and lower
margins. Upper margin of propodus nearly smooth. P5 subchelate in male, chelate in female.
Color. — "Dorsum pale opaque rose madder, darkest on abdomen; ridges of carapace opaque white;
swimmerets, thoracic legs and mouth parts deeper madder yet still pink." (RATHBUN, 1906: 898).
Source MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
349
Fig. 26. — Polycheles surdus sp. nov„ 3 38.4 mm, off Mozambique. "Algoa". I8°I4.0'S. 37°3I.0'E, 472 m (SAM
A41702): a. dorsal view of the cephalothorax; b. lateral view of the abdomen.
Etymology. — From the Latin surdus (- deaf), refers to its relation to auriculatus (Bate, 1878).
Remarks. — Rathbun (1906: 898) observed that in her Hawaiian specimens "The lateral marginal spines of
the carapace... are frequently seven in front of the first sinus, sometimes five between the sinus and the cervical
groove, and behind the groove ther may be as many as ten". Indeed, the increased number of spines on the
postcervical lateral margin distinguishes surdus from auriculatus (see above).
DISTRIBUTION. — Nazca Ridge in the southeast Pacific, Marquesas Is, Hawaii, New Caledonia. New Zealand.
Australia, Indian Ocean, Reunion, Madagascar, Mozambique; 350-1525 m.
Polycheles talismani (Bouvier, 1917)
Fig. 27
Polycheles sculptus var. talismani Bouvier, 1917: 53.
Polycheles sculptus - Bruun, 1950: 24, fig. 16 [non Smith, 1880],
Stereomastis sculpta - Hoi.THUIS, 1952b: 1 1. — LoNGHURST, 1958: 32. — Forest, 1963: 627. — Macpherson. 1983: 48.
— Rucabado & BaS, 1984: 19 [not Smith. 1880].
Stereomastus sculpta - Gauld 1960: 63 [erroneous spelling] [non Smith. 1880].
Polycheles grimaldi - Wolff, 1967: 236 (photo) [non Bouvier, 1905],
Stereomastis talismani - MACPHERSON, 1988: 57. fig. 5a; 1991: 405, fig.7. — Dawson. 1997: 24.
Stereomastis sculpta talismani - Holthuis, 1991: 88.
Material EXAMINED. — West Africa. Cancap 3: 24°17'N. 16°52'W, 1100-1150 m, 26.10.1978' 1 3 26 3 mm
(RMNH).
350
B. GALIL
"Talisman": stn 78. 23°57’N, 19°35'W, 1400-1435 m. 11.07.1883: I 3 28.1 mm; 1 2 28.8 mm (MNHN-Pa 29). —
Stn 93, 20°24'N. 20°28'W, 1435 m. 14.07.1883; 1 3 damaged (MNHN-Pa 28).
Tyro: stn 104. 19°43'N, 17°30'W. 1500 m. 17.06.1988: 1 2 32.9 mm; 7 juvs (RMNH).
"Meteor": sin 192, 17011.8’N, 16°50.6’W. 620 m, 11.02.1977: 1 2 (SMF 24644).— Sin 265. 17°17.1’N,
16°27.7'W, 95 m. 13.02.1982: 1 <3, IF (SMF 24643).
"Prof. Bogucki": sin 31. 15°02'N. 17°33'W, 807 m, 30.05.1979. coll. P. CaYRE: 1 <3 28.7 mm; I 2 35.1 mm
(MNHN-Pa 655).
"Pillsbury": stn 41. 4°47'N. 3°33'W, 641-842 m, 30.05.1964: II 6 23.5-29.5 mm; 12 2 26.2-47.8 mm (RMNH). —
Stn 44. 5°05’N. 4°00'W. 586-403 m. 30.05.1964: 10 3 24.5-30.1 mm; 9 2 29.8-49.9 mm (RMNH). — Stn 51. 4°56'N.
4°01'W, 494-329m. 31.05.1964: 19 <3 23.5-30.1 mm; 21 2 27.9-40.2 mm (RMNH). — Stn 52. 4°54'N. 4°58'W. 952-
915 m. 31.05.1964: 1 (3 27.9 mm (RMNH). — Stn 74. 4°20'N, 9°26'W. 733-641 m. 4.06.1994: 33 (3 24.6-28.0 mm;
94 2 23.5-48.5 mm (RMNH). — Stn 256. 14.05.1965: 20 <3 24.1-26.1 mm; 26 2 26.2-36.8 mm (RMNH). — Sin 309.
4°1 5'N. 4°27'E. 678-720 m. 26.05.1965: 1 <3 26.1 mm; 2 2 27.1. 29.5 mm (RMNH).
"Atlantide": stn 120. 2°09’N, 9°27'E, 260-650 m, 1.03.1946: 21 3 24.6-29.4 mm; 16 2 26.8-50.1 mm (ZMC). —
Stn 135. 7°55'S, 12°38’E. 235-460 m, 17.03.1946: 16 <3 23.7-28.3 mm; 26 2 26.5-51.5 mm; 1 ov. 2 42.3 mm (ZMC).
"Galathea": stn 32, 4°05'N, 2°13'W. 2100 m. 20.11.1950: 3 <3 22.1-23.8 mm (ZMC). — Stn 101. 8°50'S. 12°32’E.
1000 m. 12.12.1950: 1 2 47.7 mm (ZMC). — Stn 110. 12°05'S, 13°08'E. 1000 m. 19.12.1950: 2 <3 26.9. 27.0 mm;
4 2 28.0-54.3 mm (ZMC).
Guinean Trawling Survey I (GTS I): "Thierry": stn 18/6. 4°28'N, 8°28.5'W, 100 m. 31.10.1963: 1 3 29.8 mm; 2 2
28.9, 49.1 mm (USNM). — Stn 29/8. 4°13'N, 1°23'W. 380-535 m, 4.09.1963: 2 2 43.5, 48.5 mm (USNM). — Stn 34/8.
5°57'N. 1 °33'W. 400 m. 23.02.1964: 3 2 33.5-51.2 mm (USNM).
Stn 31/8. 5°06'N. 0°17'W. 400 m, 8.09.1963: 1 <5 28.1 mm; 3 2 29.4-47.1 mm (MNHN-Pa 1622). — Stn 34/8.
5°57’N. 1°34'E. 370 m. 2.10.1963: 1 2 29.0 mm (MNHN-Pa 1623). — Stn 36/8. 6°08'N. 2°56'E. 350-600 m.
6.10.1963: 1 2 39.2 mm (MNHN-Pa 1624). — Stn 46/8. 3°45'N. 8°22'E, 425 m. 2.1 1.1963: 1 <3 28.0 mm (MNHN-Pa
1625). — Stn 59/8. 3°52'S. 10°14’E. 400 m. 2.12.1963: 3 2 39.8-47.8 mm (MNHN-Pa 1116). — Stn 60/8. 4°25’S,
1°04'E. 400 m. 3.12.1968: 1 2 41.2 mm (MNHN 1627).
Guinean Trawling Survey 11 (GTS II): stn 28/8. 4°2I'N. 2°08'W. 400 m. 18.03.1963: 3 2 26.6-42.9 mm (MNHN-Pa
562). — Stn 31/8, 5°05'N. 0°18'W. 400 m. 5.03.1964: 2 2 33.0. 44.0 mm (MNHN-Pa 1628).
"Andre Nizery": stn 28. 1°51S. 8°38 E, 950 m, 16.04.80: 4 2 48.5- 58.7 mm (MNHN-Pa 530). — Stn 29. 1°55'S.
8°33.5'E, 1200-1430 m. 16.04.80: 2 2 53.3, 57.8 mm (MNHN-Pa 533). — Stn 32, 2°4rS. 8°5 1 E. 900-930 m.
17.04.80: 2 2 43.1. 43.2 mm (MNHN-Pa 528).
"Geronimo": stn 2-240. 4°08'S, 10°08'E, 1134 m. 8.09.1963: 1 2 42.2 mm (MNHN-Pa 1108). — Stn 2-247,
4°38.4'S. 11°01.2'E, 402 m. 9.09.1963: 1 2 43.9 mm (MNHN-Pa 1108).
TYPES. — The male (28.1 mm cl) (MNHN-Pa 29) collected by the "Talisman" stn 78 off West Africa
(23°57'N, 19°35'W. 1400-1435 m). is the lectotype of Polycheles talismani Bouvier. 1917: (he female in the same
sample is the paralectotype.
DESCRIPTION. — Dorsal surface of carapace setose, minutely granulate. Frontal margin straight, rostrum bifid.
Internal angle of orbital sinus minutely spinose, external angle unarmed. Eyestalk bearing papilla, curved distad.
Basal antennular segment produced anteriorly to a sharp point, mesial margin smooth, anterolateral margin
bispinose. Spines on lateral margins of carapace upeurved, anteriormost largest. Spines formula of lateral margins
of carapace 5-6:3:8-10; spines successively further apart posteriorly. Spine formula of median postrostral carina
(excluding rostral spines) 1:2:1, median postcervical carina bearing two pairs of spines. Gastro-orbital region with
crescentic row of five antrorse spines, anteriormost largest. Posterior margin of cervical groove with antrorse spine
between median postcervical and branchial carinae. Branchial carina prominent, parallel to lateral margin, with 5-7
spines, posteriormost largest; region between branchial and median carinae unarmed, obliquely grooved anteriorly.
Posterior margin of carapace bearing pair of antrorse submedian spines.
First abdominal tergite bearing two pairs of prominent spines distally on anterior margin. Abdominal tergites
medially carinate. Carinae on tergites 1-5 bearing antrorse spines, spines 1-4 increasing in size posteriorly; antrorse
spine on fifth tergite smaller than on the fourth. Sixth tergite bearing lyre-shaped denticulate carina mesially.
Tergites 2-5 with transversely oblique submedian grooves. Pleura 2-5 with curved rib mesially. margins minutely
granulate. Anterior margin of second pleuron produced, triangulate. Telson with procumbent crest anteriorly, two
minutely serrulate, convergent carinae posteriorly.
Merus of PI proximally bispinose, with subterminal claw-like spine on upper margin, finely spinulose on
lower margin; carpus half as long as merus, subterminal spines on upper and lower margins. Upper margin of
Source MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE 35 1
propodus with subterminal spine, lower margin with two minutely spinulatc rows. P5 subchelate in male, chelate
in female.
Color Carapace pale pink anterior to cervical groove, spines red, abdomen pale pink carinae, uropods dark
red, legs red, antennae and antennulae red (Holthuis, pers. obs.).
Fig. 27. — Polycheles talisman! (Bouvier, 1917), 2 35.1 mm. Senegal, "Prof. Bogucki". stn 31, 15°02'N, 17°33'W.
807 m (MNHN-Pa 655): a. dorsal view of the cephalothorax; b. lateral view of the abdomen.
Remarks. — P. talismani is distinguished from P. sculptus in having 8-10 spines on the lateral margins of
carapace posterior to postcervical incision rather than 6 or 7; the sixth tergal carina denticulate and lyre-shaped
rather than parallel and rounded; and the median crest on telson prominent, keel-like, rather than rounded.
Bouvier (1917: 53) offered but a cryptic description of Polycheles sculptus var. talismani "qui
habite...d'Allantique oriental". Holthuis (1952: 11) noticed that his West African specimens differed in the
number of spines on the lateral margin of the carapace from the North Atlantic sculptus. MaCPHERSON (1988),
examined BouviER's specimens and finding that they differed from sculptus in the number of lateral spines of the
carapace, and the median dorsal keels on the sixth abdominal segment, assigned them specific status.
Distribution. — West Africa: Mauritania to Angola; 100-2100 m.
Polycheles tanneri Faxon. 1 893
Fig. 28
Polycheles tanneri Faxon. 1893: 196; 1895: 119. pi. 31. — Seeliger. 1901: 46. — Bouvier. 1915: 2. — Sund. 1915:
372. — De Man, 1916: 6 (list). — Bernard, 1953: 86. — Firth & Pequegnat. 1971: 50. — Del Solar, 1972: II.
— Wicksten. 1989: 303. — HendriCKX. 1995: 156. — Dawson, 1997: 12.
352
B. GALIL
Material EXAMINED. — Panama. "Albatross": sin 3354, 7°09'07"N, 80°50'W, 580 m. 23.02.1891, id.
W. Faxon: I juv. (USNM 21068).
Galapagos Is. "Albatross": sin 3402, San Cristobal 1., 0°57'03"S. 89°03'30"W, 758 m. 28.03.1891. ,d.
W Faxon: 4 2 19.6-25.9 mm (USNM 21069). — Sin 3409, 0°18'40"N, 90°34'W. 589 m, 3.04.1891: 1 2 41.8 mm
(USNM 21070). — Sin 4642. Hood 1., I°30'30"S. 89°35'W. 540 m, 7.11.1904: 1 juv. (USNM).
Ecuador ."Albatross"-, sin 2792. 0°37'N, 81°00'W. 722 m. 2.03.1888. id. W.L. SCHMITT: 1 <J damaged; 7 juvs
(USNM 58646).
Peru. S. of Mancora Bank, 700-830 m, 1 1.01.1971, coll. E.M. DEL Solar: 1 6 31.7 mm: 1 2 24.5 mm (RMNH
27349).
Type. — The female (41.8 mm cl) (USNM 21070) collected by the "Albatross" sin 3409 off the Galapagos Is
(0°18'40"N, 90°34'W, 589 m) is the holotype of Polycheles tanner i Faxon, 1893.
Fig. 28. — Polycheles tanneri Faxon, 1893, 2 41.8 mm, holotype. Galapagos Is, "Albatross" stn 3409. 0°I8'40"N.
90°34'W, 589 m (USNM 21070): a, dorsal view of the cephalothorax; b. lateral view of the abdomen.
DESCRIPTION. — Dorsal surface of carapace tomentose, laterally granulose. Median submarginal tooth
upcurved, reaching beyond twin rostral spines. Internal angle of orbital sinus pronounced, triangular; external angle
convex, spinose, pectinate. Eyestalk distally granulose, bearing mammilate tubercle. Basal antennular segment
produced anteriorly to a sharp point, mesial margin prominently spinulose; antero-external angle bispinose. Lateral
spines diminishing in size posteriorly. Spine formula of lateral margins of carapace 5:3:13-14. Spine formula ol
median postrostral carina (excluding rostral spines) 1:1:1 :(0- 1 ):2: 1 ; median postcervical carina bearing two pairs of
spines. Gastro-orbital carina quadrispinose, anteriormost spine largest. Anterior branch of cervical carina spinulose.
Posterior margin of cervical groove between median postcervical and branchial carinae spinose. Branchial carina
prominent, bearing 13 or 14 antrorse spines. Carapace between gastro-orbital, branchial and median carinae with
antrorse spinules. Posterior margin of carapace bearing antrorse spines, median pair largest.
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
353
First abdominal tergitc with mammilate tubercle laterally. Abdominal tergites without oblique, submedian
grooves, medially carinate. Tergites 1-5 bearing prominent, keel-like antrorsc spine, spine of fourth tergite most
prominent. Sixth tergite with conical tubercles, prominently granulate lateral margins, medially with two parallel
rows of large conical granules meeting posterior margin in prominent denticle. Second pleuron furnished anteriorly
with prominent spine. Pleura 3-5 with granulate rib mesially. Telson anteriorly with median denticulate crest,
posteriorly with convergent, minutely serrulate carinae.
Merus of PI spinulose on upper and lower margins, two distal claw-like spines on upper margin. Carpus one
third as long as merus, distally spinose on upper margin, distal claw-like spine on upper and lower margins. Upper
and lower margins of propodus prominently spinose. P5 simple in male, chelate in female.
Remarks. — P. tanneri was described (1893), and not-quite-accurately illustrated (1895) by FaXON. P. tanned
is distinguished by its short cheliped carpus, the spinulose anterior branch of cervical carina and the prominent
keel-like carinae on abdominal terga.
Distribution. — Eastern Pacific: Panama, Ecuador, Peru and Galapagos Is; 540-830 m.
Polycheles trispinosus (De Man, 1905)
Fig. 29
Pentaclieles trispinosus De Man, 1905: 587.
Stereomastis trispinosa - De Man, 1916: 10. pi. I figs 1, la-b. — Bernard. 1953: 87. — Firth & Pequegnat, 1971: 74.
— Macpherson, 1991: 405. — Takeda & Hanamura, 1994: 32.
Material EXAMINED. — Indonesia. "Siboga": stn 38, 7°35.4'S, 1 1 7°28.6E, 732-915 m, 1.04.1899: 1 3
34.8 mm (ZMA); 1 $ 23.6 mm and 4 juvs (ZMA). — Stn 314, 7°36'S, 1 17°30.8'E. 694 m. 17.02.1900: I 9 34.0 mm;
7 juvs (ZMA).
Snellius 2: stn 4267. 8°17.6'S. 1 18°2 1 .3 E, 650 m, 29.10.1984: 6 3 21.2-34.6 mm; 6 $ 21.4-35.2 mm; I juv.
(RMNH).
Types. — The male (34.8 mm cl) (ZMA) collected by the "Siboga" stn 38 off Indonesia (7°35.4’S.
117°28.6'E. 732-915 m) is designated here as the lectotype; the female (23.6 mm cl) in the same sample is a
paratype.
DESCRIPTION. — Carapace oblong, lateral margins subparallel, slightly convergent anteriorly. Dorsal surface of
carapace tomentose. Frontal margin straight. Median submarginal tooth upeurved, in front of bifid rostrum.
Internal angle of orbital sinus spinose, external angle unarmed. Eyestalk bearing mammilate tubercle, pointing
distad. Basal antennular segment produced anteriorly to sharp point, mesial margin smooth; antero-external angle
bispinose, anterior spine nearly twice as long as lateral. Spines of lateral margins of carapace decreasing
successively in size posteriorly. Spine formula of lateral margins of carapace 5:3:9. Spine formula of median
postrostral carina (excluding rostral spines) 1: 1:2:1; median postcervical carina bearing two pairs of spines. Gastro-
orbital region with row of five antrorse spines curved distad. Posterior margin of cervical groove with single
prominent antrorse spine midway between median postrostral carina and lateral margin. Branchial carina slightly
sinuous, bearing 1 1 spines. Carapace bearing 5 spines parallel with branchial groove; additional spine posteriorly,
near branchial carina. Posterior margin of carapace bearing antrorse submedian spines.
Anterior margin of first abdominal tergite bearing two pairs of lateral spines. Carinae on tergites 1-3 bearing
antrorse spine; spine of tergite 3 considerably longer, overhanging second tergite. Carinae on tergites 4-5 unarmed.
Tergite 6 granulate, medially with denticulate double carina, confluent posteriorly. Tergites 2-5 with deep,
transversely oblique, submedian grooves. Pleura 2-5 granulate, with curved, granulate rib medially, margins
denticulate. Second pleuron rounded, bearing spine anteriorly. Telson bituberculate anteriorly, two minutely
serrulate, confluent carinae posteriorly.
Merus of PI proximally bispinose, subterminal claw-like spine on upper margin, finely spinulose on lower
margin. Carpus half as long as merus, subterminal spines on upper and lower margins. Upper margin of propodus
with 6-7 spinules, lower margin finely spinulose distally. P5 subchelate in male, chelate in female.
354
B. GAUL
Fig. 29. — Polycheles trispinosus (dc Man, 1905), 2 34.7 mm, Indonesia, SNELLIUS 2, stn 4267. 8°17.6'S. 1 18°21.3'E,
650 m (RMNH): a. dorsal view of the cephalothorax; b, lateral view of the abdomen.
Remarks. — Polycheles trispinosus is distinguished from its congeners: "The carinae of only the three first
abdominal terga are culminating in antrorse spine" (De Man. 1916: 10).
Distribution. — Known only from Indonesia; 650-915 m (de Man, 1905).
Polycheles typhlops Heller, 1862
Fig. 30
Polycheles typhlops Heller. 1862: 392, pi. 1 figs 1-6; 1863: 211, pi. 7 figs 1-2. — Carpenter & Jefpreys. 1870: 154.
— Wood-Mason, 1874: 180; 1875: 134. — Bate, 1878a: 273; 1878c: 563; 1888: 100. — Norman. 1878: 382
(P-P-): 1879: 175 (p.p.); 1886: 7. — Carus, 1885: 486. — Steindachner. 1891: 438. — Stebbing. 1893: 200. —
Adensamer. 1898: 621. — Lo Bianco, 1903: 250. — Senna, 1903: 332. fig. 7, pi. 18 figs 1-11. — Magri, 1904: 7.
— Bouvier, 1905a: 480; 1905c: 3; 1915: 2; 1917: 35. pi. 2 figs 1-6; 1925: 433, fig. 9; 1940: 75. pi. 2. fig. 5. —
Kemp, 1906: 7; 1910: 413. — Richard, 1907: 322. — Pesta. 1912: 103; 1918: 162, fig. 53. — Kemp & Sewell,
1912: 24. — Selbie, 1914: 12, pi. 1 figs 1-13. — Sund. 1915: 372. — H.C. Williamson, 1915: 437. — De Man.
1916: 6. — Miranda & Rivera. 1921: 184; 1933: 18. — Stephensen. 1923: 67. — Balss. 1924: 179. figs 5. 8;
1925: 201, pi. 19. — Calman. 1925: 18. — Dieuzeide, 1929: 103; 1950: 46; 1960: 77. — Bolivar. 1930: 21, fig.
2. — Seurat. 1930: 37; 1940: 145. — Santucci. 1931: I; 1932: 1; 1933: 1. — Schmitt, 1935: 172. — Zariquiey
Cenarro, 1935: 6. — Zariquiey Alvarez, 1946: 99, figs 123-124; 1955: 407; 1962: 30. 33; 1968: 209, fig. 86b.
— Vilela, 1936: 224. — Ramadan. 1938: 129. — Parenzan, 1940: 139 (list). — Le Danois. 1948: 139. pi. 3. —
Barnard, 1950: 568. — Tortonese, 1951: 220; 1958: 195; 1959: 448; 1962: 112; 1965: 85. — Bernard, 1953:
86. — Marshall, 1954: 120. — PLr£s & Picard, 1955: 51. — Sewell, 1955: 203. — Holthuis. 1956: 1 14; 1987:
297. fig. 2. — Springer & Bullis, 1956: 14 (p.p.). — Dieuzeide & Roland, 1957: 19; 1958: 23. — Hemming. 1958:
137. — Holthuis & Gottlieb. 1958: 1 14. — Longhurst. 1958: 32. — Rossi, 1958: 7. — Forest & Gantes, 1960:
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHEL1DAE
355
348. — Russell, 1962: 6. — Maurin, 1962: 180; 1968a: 45; 1968b: 479. — Forest, 1963: 627: 1965: 346. —
Lewinsohn & Holthuis, 1964: 54. — P£r£s, 1964: 22. — Torchio, 1964: 120; 1967: 122. — Buu.is & Thompson.
1965: 8. — Dawson, 1965: 1. — Grindley & Penrith, 1965: 286. — Kotthaus, 1966: 348. figs 1-3. — Allen.
1967: 55. — Massuti. 1968: 306. — Berry, 1969: 46. — Stevcic, 1969: 128; 1990: 210. — Carbine, 1970b: 135.
— Rubio Lois, 1971: 10. — Firth & Pequegnat. 1971: 50. figs 9-10. — Pequegnat ct al., 1971: 4. — Lagardere.
1972: 672, fig. 5; 1977a: 399. — Relini-Orsi & Relink 1972, fig. 5. — Koukoura. 1973: 753. — Relink 1973: 29.
— Neves, 1974: 7. — Thiriot, 1974: 344. — Pastore, 1976: 107. — Turkay, 1976: 26; 1993: 284. — Voss &
Jeuniaux. 1978: 127. — Beaubrun, 1979: 35, figs 15-16. — Rodriguez, 1980: 194. — Soto. 1980: 87. —
Burkenroad. 1981: 263. — Moncharmont, 1981: 66. — KENSLEY. 1981b: 29. — Burukovsky. 1983: 134. fig.
178. —George, 1983: 16. — Ohta, 1983: 230. — Riedl. 1983: 481.pl. 176. — Takeda. 1983: 74. — Lemaitre.
1984: 427. — O'Riordan, 1984: 322. — Omori, 1985: 63. — Baba, 1986: 157, pi. 107. — Relini et al., 1986: 156,
tab. 4. fig. 4. — Manning & Froglia. 1982: 321. — Duris, 1987: 9. — Garcia & Massudi. 1987: 75. — Gonzalez-
Gurriaran & Olaso. 1987: 328 (list). — Abello & Valladares, 1988: 98. — Abello e/ al., 1988: 41. — Morgan
& Jones. 1988: 15. — Gonzales et al., 1988: 28. — Chan & Yu. 1989: 166, pi. 1; 1993: 105. — Intes & Bach.
1989: 21. — Cartes & Abello, 1990: 35; 1992: 139 . — Falciai & Minervini. 1992: 134, pi. 9. fig. 3. — Cartes &
Sarda, 1992: 1315. — Cartes, 1993: 32. — Cartes et al., 1993: 210. — Emmerson. 1993: 181. — Kocatas &
Katagan, 1993: 35. — Rodriguez, 1993: 197 (list). — Vaso & Gjiknuri. 1993: 397. — Pipitone & Tambiolo. 1993:
361 (list). — Galil & Goren, 1994: 45, fig. 2b. — Quackenbush, 1994: 85. — Gonzalez. 1995: 126. fig. 65.
Pentacheles agassizii A. Milne Edwards. 1880: 65. — Faxon. 1896: 155. — Young. 1900: 441.
Polycheles doderleini Riggio, 1885: 103, pi. 3 figs 1-5.
Pentacheles hextii Alcock, 1894: 237; 1899: 35; 1901b: 172; 1902: 264. — Alcock & ANDERSON. 1894: 165; 1895.
pi. 10 fig. 2.
Palyclieles agassizii - Faxon, 1895: 120.
Polycheles agassizi - Bouvier. 1905a: 480; 1905c: 3.
Polycheles hexti - Bouvier, 1905a: 481.
Policheles typhlops - Maori, 191 1: 32. — Bombace. 1968: 113. — Relini-Orsi & Relink 1972: 59 [erroneous spelling],
Polycheles intermedins Balss, 1914: 599.
Polychelles lypltlopis - Nobre, 1931: 246, fig. 137; 1936: 153. pi. 51 fig. 130. [erroneous spelling].
Polycheles typhlops typhlops - Firth & Pequegnat, 1971: 51, fig. 7. — Wadley & Evans. 1991: 27. pi. n.n. — Griffin
& Stoddart. 1995: 242, figs 6-8. — Tiefenbacher. 1995: 5. — Dawson, 1997: 13.
Polycheles typhlos - Arena & Greci. 1973: 164 [erroneous spelling].
Polycheles thyphlos - Arena & Greci, 1973: 173. [erroneous spelling],
Polycheles tiphlops - Froglia, 1976: 78 (list) [erroneous spelling].
Polycheles sp. - Gamo. 1980, pi. 7b. — Utinomi, 1983: 105.
Stereomastis nana - Miyake, 1982: 78. pi. 26 fig. 6 [non Smith, 1884].
Polycheles hextii - HOLTHUIS. 1984: 4.
Polycheles thyphlops - Gonzalez-Gurriaran & Olaso. 1987: 329. — Jacques, 1989: 4. [erroneous spelling].
Polycheles typholps - Miyake, 1982, pi. 26 fig. 6. [erroneous spelling],
nol Polycheles typhrops - Takeda, 1982, pi. 38.
Material examined. — Mediterranean Sea.
Israel. 32°31'N, 34°12'E. 1347-1500 m, 4-7.11.1998. coll. B. Galil: 13 <3 18.3-37.6 mm; 16 2 17.0-43.2 mm
(TAU).
Spain. Costa Brava. 09.1960, id. L.B. Holthuis: 1 ov. 2 33.2 mm (USNM 106118).
France. Banyuls stir nter, 1978: 1 2 (SMF 24637).
Atlantic Ocean.
North Atlantic. 49°24.52'N. H°29.14'W, 581 m, 20.07.1996: I 3 27.7 mm; 1 2 25.9 m; 10 juvs (RMNH).
Balgim: stn CP 124. 35°46'N, 3°52.'W, 1405 m, 14.06.1984: 2 2 22.6. 37.1 mm (MNHN-Pa 1630).
West Africa. "Meteor": stn M36-98 AT-149. 25°31.5'N. 16°02.2'W. 658-888 m. 24.02.1975: 8 spec. (SMF
24640). — Stn 180. 33°37.5'N, 9°02.2'W, 953-1038 m. 12.03.1975: 1 juv. (SMF 24641). — Stn M36-I27 AT-261.
1 7°20.6'N, 16°27.TW, 105 m. 10.02.1982: 10 spec. (SMF 24642).
"Talisman": Cape Verde, 16°52'N, 27°30’W. 400-580 m. 27.07.1883, det. E.L. Bouvier: 1 <3 40.4 mm; I 2 48.4 mm
(MNHN-Pa 1169).
"Pillsbury": stn 73, 4°40'N. 9°20'W, 366-311 m, 4.06.1964: I 3 32.0 mm (RMNH). — Stn 255. 3°49'N. 7°38'E.
269-264 m. 14.05.1965: 1 3 35.5 mm (RMNH).
SE Atlantic. "Prof. Mesyatsev": Rio Grande Ridge, 12.07.1974, coll. B.N. Kotenev: 1 2 38.8 mm (ZMMU).
Bermuda. E. St. Davids, 540 m. July-Aug. 1960: 1 2 damaged (USNM 106099).
United States. SE Cape Fear, 34°18'N, 75°51'W. 396 m. 14.06.1957, id. F.A. Chace as Stereomastis nr grimaldii:
1 3 29.8 mm (USNM 101642).
"Oregon": stn 5750, 29°33'N. 79°58’W. 556 m, 19.11.1965: I 2 26.7 mm (USNM 273140).
356
B. GALIL
" Gerda ": sin 67, 25°31'N, 79°57'W. 351 m. 26.09.1962: 1 $ 30.8 mm (RMNH).
West Indies. "Blake": sin 246. 277 m, 1878-79: I 9 26.2 mm; 1 juv. (MNHN-Pa 32).
"Silver Bay": stn 2458, 23°40'N, 79°18'W, 522 m. 5.11.1960: 2 3 23.5, 46.1 mm (USNM 273140). — Sin 2483.
26°25'N, 79°0’l'W. 540 m, 9.1 1.1960: 5 3 29.7-33.3 mm; 2 9 32.6, 51.3 mm (USNM 251943). — Sin 3472. 27°20'N.
78°19'W, 450-612 m. 25.10.1961: 1 ov. 9 34.9 mm (USNM 273140).
Gulf of Mexico. Dry Toriugas. 522 m, 3.07.1931. coll. & id. W.L. Schmidt: 4 3 22.1-30.2 mm; 2 9 23.6.
37.9 mm; 1 juv. (USNM 78283). — Ibidem, 448 m. 3.08.1932, coll. & id. W.L. Schmidt: 1 9 37.0 mm (USNM 78260).
"Oregon": sin 489. 27°44'N, 85°09’W. 457 m. 29.09.1951: 1 9 damaged (USNM 92625). — Stn 1324. 24°23'N.
82°22'W. 360 m. 07.1955: 1 9 48.5 mm (USNM 98652). — Sin 1407, 28°07'N. 89°59'W. 464 m. 20.09.1955: 1 3
34.1 mm (USNM 99454). — Sin 3565, 14°10'N, 81°55'W, 432-450 m. 21.05.1962: 2 9 38.6. 46.8 mm; 1 ov. 9
49.3 mm (USNM 244083). — Stn 3628. 16°57'N. 8P19’W, 450 m. 7.06.1962: 1 9 43.4 mm (USNM 244083). —
Stn 4337. 24°26'N. 83°23'W. 342 m. 26.07.1963: 1 9 46.2 mm (USNM 273140). — Sin 4343, 24°33'N, 83°38'W.
378 m. 27.07.1963: 2 9 28.4, 43.9 mm (USNM 273140). — Sin 4703. 27°55'N. 90°28'W. 540 m, 22.02.1964: 2 9
32.3, 39.2 mm (USNM 273140) together with P. perarmatus. — Stn 5168, 29°23'N. 80°22'W. 43 m. 12.01.1965: 1 3
40.9 mm (USNM 260368). — Stn 5233, 29°54.5'N. 80°10'W, 342 m. 9.02.1965: I 9 50.7 mm (USNM 273140).
Caribbean Sea ."Albatross": stn 2134, 19°56'05"N, 75°47'32',W. 457 m, 27.02.1884: 1 3 28.1 mm (USNM).
Puerto Rico. 18°32'N. 66°21.I5'W, 468-648 m. 4.02.1933. id. W.L. Schmitt: 2 3 26.4. 27.2 mm; 4 juvs (USNM
82183).
"Silver Bav": stn 5142, I9°52'N, 71°58.5’W. 630 m. 12.10.1963: I 3 41.6 mm; I 9 29.9 mm (USNM 260368). —
Stn 5166. 19°48.5’N, 70°30.5'W. 396-540 m. 15.10.1963: 2 <3 30.7, 38.1 mm; 1 ov. 9 36.1 mm (USNM 260368).
"Oregon": stn 2774. 1 1 °32'N, 62°40’W. 351-382 m, 19.04.1960: 1 ov. 9 42.2 mm (USNM 232276) together with
P. perarmatus. — Stn 2776. 1I°36'N, 62°42'W. 423 m, 19.04.1960: 1 3 40.7 mm; 1 9 43.1 mm; I ov. 9 62.2 mm
(USNM 232276). — Stn 2780. 11°36'N. 62°52’W. 387-414 m, 20.04.1960: 1 9 41.5 mm (USNM 232276). —
Sin 3552, I7°40'N. 77°55'W. 522 m, 16.05.1962. id. F.M. Bayer as P. agassizii: 2 9 30.6. 40.5 mm (USNM 154322).
— Stn 3570. 14°08'N, 81°55'W, 360-432 m, 05.1962. id. F.M. Bayer as P. agassizii: 1 c? 37.0 mm (USNM 154323).
— Stn 3627. 16°50'N, 81°21'W. 360 m, 7.06.1962, id. F.M. Bayer as P. agassizii: 3 3 28.9-40.2 mm; 1 ov. 9
43.0 mm (USNM 154397). — Stn 3635, I6°58'N. 87°53'W, 450-720 m. 10.06H962, id. F.M. Bayer as P. agassizii:
1 9 55.2 mm (USNM 154396). — Stn 4407. 11°59'N, 69°30'W. 414 m, 27.09.1963: 1 3 32.7 mm; 2 9 45.3. 50.8 mm
(USNM 273140). — Stn 4411. 11°55'N, 69°27'W. 468 m. 3.10.1963: 2 9 39.3. 44.0 mm (USNM 273140). — Stn
4417, 1 1°46'N, 69°15'W, 432 m. 4.10.1963: 1 3 35.0 mm; 1 juv. (USNM 273140). — Stn 4419. 11°43'N. 69°13'W,
450 m. 4.10.1963: 1 3 24.1 mm; 2 9 40.7, 43.8 mm (USNM 273140). — Sin 4421. I1°49'N. 69°24'W. 360 m.
4.10.1963: 2 3 32.3. 33.7 mm; 2 9 27.1, 30.2 mm (USNM). — Stn 4424. 1 1 °50'N. 69°23'W. 369 m, 5.10.1963: I 9
48.7 mm (USNM 273140). — Stn 4425. I1°46'N, 69°17'W. 432 m. 5.10.1963: 2 9 49.7. 58.5 mm (USNM 273140).
"Pillsbury": stn 374. 9°57'N, 76°10.6’W. 439-377 m. 14.07.1966: 4 9 46.1-52.3 mm; 2 juvs (RMNH). — Stn 394.
9°28.6'N, 76°26.3'W. 421-641 m. 16.07.1966: 1 9 38.5 mm; 1 juv. (RMNH). — Stn 776. I2°13.3'N. 72°50’W. 408-
576 m. 29.07.1968: 1 9 40.4 mm; 9 juvs (RMNH). — Stn 929. 15°29.5'N. 61°11.5'W. 503 m. 15.07.1969: 1 3
38.3 mm (RMNH).
Indo-west Pacific.
Fiji Is. Musorstom 10: stn CP 1316, 17°14.84'S. 178°21.99'E. 478-490 m, 6.08.1998: 2 3 38.5, 39.5 mm;
2 ov. 9 48.3, 47.2 mm (MNHN). — Stn CP 1331, 17°02.45'S, 178°01.84'E, 694-703 m. 8.08.1988: 14 3 20.0-
44.4 mm; 14 9 20.7-37.7 mm; 7 juvs (MNHN). — Stn CP 1332, 16°56.17’S, 178°07.86'E. 640-687 m. 8.08.1998:
3 3 24.9-6.8 mm; 4 9 23.8-40.6 mm; 2 juvs (MNHN). — Stn CP 1368, I8°10.92'S. 178°23.47'E. 380-469 m,
15.08.1998: 1 9 32.4 mm (MNHN). — Stn CP 1369, 1 8°1 1 . 1 3'S. 178°23.44'E. 392-433 m, 16.08.1998: I 3
39.7 mm; 2 ov. 9 4.32. 4.30 mm (MNHN).
New Caledonia. Bathus I: stn CP 671, 20o51.l8’S. 1 65°28. 1 7E. 450-470 m. 14.03.1993: I 9 32.4 mm (MNHN-
Pa 1730).
Bathus 2: stn CP 760, 22°18.9'S. 166°10.5'E, 455 m. 16.05.1993: I 9 48.0 mm (MNHN-Pa 1631). — Stn CP 764.
22°09.41'S. 1 66°02.93'E. 560-570 m, 17.05.1993: I 3 43.1 mm (MNHN-Pa 1632). — Stn CP 765, 22°09.6'S.
1 66°02.8'E. 600-630 m. 17.05.1993: 1 3 44.8 mm (MNHN-Pa 1633). — Stn CP 771. 22°09.52'S. I66°01 75’E 610-
800 m, 18.05.1993: 1 <3 50.5 mm (MNHN-Pa 1634).
Japan. Kyushu L, Miyazaki Pref., 300 m, 20.12.1953. det. S. Miyake as P. amemiyai Yokoya: 1 9 (K.MNH 419). —
Shikoku I.. Tosa Bay. nr Mimase. 16.01-14.02.1961. coll. K. Sakai: 2 9 (SMF 24638).
East China Sea. 20.10.1989, coll. M. Yamada: 1 3 54.0 mm (NFU).
Taiwan. I-Lan County, 400 m. 21.05.1992, coll. & det. T.Y. Chan: 2 9 34.6. 53.3 mm (NTOU). — Ta-Chi, 400 m.
13.06.1995. coll. & det. T.Y. Chan: 4 9 40.3-52.5 mm (MNHN-Pa 1682).
Philippines. "Albatross": stn 5111, Luzon. 13°45'I5"N. 120°46'30"E. 432 m. 16.01.1908: 1 3 22.0 mm; 4 9
16.0-37.5 mm (USNM). — Stn 5122, Mindoro, 13o2L30"N. 1 20°30'33''E. 402 m. 2.02.1908: 8 3 21 1-26 6 mnr 7 9
18.1-45.2 mm (USNM). — Stn 5123. Mindoro, 13°12'45"N, I21°38'45"E. 518 m, 2.02.1908: I <J 30.0 mm; 4 9 34.6-
43.8 mm (USNM). — Stn 5124. Mindoro. 12°52'N. 121°48'30"E. 514 m, 2.02.1908: 3 3 27.5-29.7 mm; 4 9 18.0-
33.7 mm (USNM). — Stn 5221, btw Marinduque and Luzon, 13°38'15"N, 12P48'15"E. 357 m. 24.04.1908: 3 9 41.8-
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
357
49.5 mm (USNM). — Sin 5222. btw Marinduque and Luzon, 13°38'30"N, I2I°42'45"E. 357 m. 24.04.1908: 2 <3 28.6,
29.5 mm; 2 2 20.9, 47.3 mm (USNM). — Sin 5364, Luzon. Balayan Bay, 13°48'30"N, 120°43'45"E. 293 m,
20.02.1909: 4 3 32.0-34.1 mm; 5 2 27.6-43.5 mm (USNM). — Stn 5374, Marinduque I„ 13°46'45"N. 121°35'08"E,
347 m. 2.03.1909: I 3 31.2 mm; 2 2 32.9, 36.0 mm (USNM). — Stn 5404, Lcyle, Dupon Bay. 10°50'N. 124°26'I8"E,
347 m, 17.03.1909: 1 2 39.9 mm (USNM). — Sin 5409. blw Cebu and Lcyle, 10°38'N. 124°13'08"E. 346 m,
18.03.1909: 2 2 31.3, 31.9 mm (USNM).
Musorstom 1: sin 50. 13°49.2'N. 120°01.8'E, 415-510 m, 25.03.1979: I 3 43.4 mm; 1 2 56.5 mm (MNHN-
Pa 1631).
MUSORSTOM 3: sm CP 105. 13°52'N, 120°30'E. 398-417 m, 1.06.1985: 1 <3 25.9 mm: 3 2 26.1-38.0 mm (MNHN-
Pa 1636). — Stn CP 118, 11°58'N, 121°06'E. 448-466 m. 3.06.1985: 1 <3 25.6 mm; 2 2 44.2. 44.4 mm (MNHN-
Pa 1637). — Stn CP 127, ll°48’N, 121°30'E. 464-475 m, 4.06.1985: 1 <5 41.3 mm; 3 2 28.6-50.1 mm (MNHN-
Pa 1638).
Indonesia. "Albatross": stn 5622. blw Gillolo and Makyan Is. 0°19'20"N, 1 27°28'30,,E. 503 m. 29.11.1909: 1 2
50.2 mm; 2 ov. 2 49.0, 49.8 mm (USNM). — Stn 5624. btw Gillolo and Makyan Is. 0°12T5"N, 127°29'30"E. 518 m.
29.1 1.1909: 2 3 37.2. 41.8 mm; 5 2 24.0-63.1 mm; 1 ov. 2 49.8 mm (USNM). — Stn 5625, btw Gillolo and Kayoa Is.
0°07'00"N. 1 27°28'00"E. 421 m. 29.11.1909: 1 3 25.8 mm; 1 2 65.8 mm; 2 juvs (USNM). — Stn 5626. btw Gillolo
and Kayoa Is, 0°07’30"N, I27°29'00,,E, 485 m, 29.11.1909: 4 3 18.5-29.1 mm; 5 2 18.5-34.1 mm (USNM).
MORTENSEN Expedition 1929: 8°30'S. 1 I4°38'E, 450 m. 7.04.1929: 1 3 31.7 mm (ZMC). — 7°29'S. 114°49'E.
240 m, 10.04.1929: 1 3 24.8 mm (ZMC).
Java, 8°34'S. 1 14°36'E, 27.06.1981: 1 ov. 2 65.4 mm (USNM 356448).
Karubar. lies Kai: stn CP 12. 5°23'S, 132°37'E. 436-413 m, 23.10.1991: 1 3 44.8 mm (MNHN-Pa 1639). —
Stn CP 35, 6°08'S, 132°45'E, 390-502 m. 27.10.1991: 1 2 47.0 mm (MNHN-Pa 1640). — Tanimbar Is: stn CC 40.
7°46'S, 132°31'E. 443-468 m, 28.10.1991: 1 2 56.1 mm (MNHN-Pa 1641).
Australia. South East Coast. "Kapala": sin K78-17-21, 28°37'S. 153°50 E. 502 m, 19.08.1978. id. H.E. Stoddart.
as Polycheles typhlops typhlops: 1 3 43.8 mm (AM 44911). — Stn K84-15-02. 33°34'S. 1 5 1 °57 E. 549-568 m.
10.09.1984, id. H.E. Stoddart as Polycheles typhlops typhlops: 1 2 37.6 mm (AM P39741).
North West Coast. "Solea": N.W.E. Cape Lambert, 18°32'S. 1 1 7°2 I E, 496-504 m, 8.04.1982: 1 2 46.3 mm (WAM
Cl 3427). — N.W. of Beagle Bay, 15°13.5'S, 121°08.9'E. 352 m, 11.02.1984: I 3 45.7 mm (WAM C169I2). —
16°55.0'S. 1 19°54.0'E, 435-434 m, 18.02.1984: 1 3 47.0 mm (WAM C23523). — 16°55. 1 S. 119°55.1'E. 432-434 m,
19.02. 1984: 1 2 57.6 mm (WAM C23524). — 16°54.1'S. 119°55.6'E. 434 m. 21.02.1984: 1 2 49.9 mm (WAM
C23527). — W. of Lacepede Arch.. 16°55.1'S. 119°54.8'E. 432-430 m, 22.02.1984: 1 3 41.6 mm (WAM C23589). —
W. Roebuck Bay, 18°04’S, 1 18°14E, 400-396 m. 24.02.1984: 1 3 (WAM C235I6). — 1 6°5 1 S, 1 19°48'E. 466-464 m.
28.02.1984: 1 2 35.2 mm (WAM C23529).
"Courageous": N.W. of Port Hedland. 17°54'S, 1I8°28'E. 426-427 m, 21.08.1983: 1 ov. 2 57.2 mm (WAM
C23597). — 17°28'S. 118°52'E. 428-433 m, 18.08.1983: 1 2 60.0 mm (WAM C23599). — 18°54'S, II7°02'E, 300-
306 m. 13.04.1984: 2 2 49.1. 60.1 mm (WAM C13428). — 18°44'S. 116°59'E. 404-406 m, 23.08.1983: 1 3 45.8 mm;
1 2 60.6 mm (WAM C23522). — 18°05'S, I18°10’E. 401-400 m, 17.08.1983: 1 3 38.9 mm; I 2 45.0 mm (WAM
C23518).
Indian Ocean. 29°49.82'S, 41°39'E, 486 m. 1.06.1994, coll. S.l. Luswazi: 1 2 56.1 mm (SAM).
Comoro Is. Benthedi: stn 61F. Mayotte, W de la grande passe de l'Ouest, 510-474 m. 29.03.1977: 2 3 21.5.
27.5 mm; 2 2 30.3, 38.5 mm (MNHN-Pa 1708). — Stn 114 F. 12°22.3'S. 46°28.2'E. 300-600 m. 11.04.1977: 1 <3
32.6 mm (MNHN-Pa 1710).
Madagascar. "Vauban": stn CH 29, 12°43 I'S, 48°ll.rE. 540 m, 13.09.1972, coll. & det. A. Crosnier: I 3
46.0 mm (MNHN-Pa 1642). — Stn 58, CH 23°36.2'S. 43°30.5'E. 510 m, 27.02.1973. coll. & det. A. Crosnier: 1 3
48.3 mm (MNHN-Pa 1643).
"Mascareignes III": stn 63, 22°26.8'S. 43°05.4’E. 530 m. 20.10.1986. coll. R. von Cosel: 1 2 72.4 mm (MNHN-Pa
1644). — Stn 68, 22°24.7'S. 43°04.4'E, 525 m, 21.10.1986, coll. R. von Cosel: 1 3 57.9 mm (MNHN-Pa 1645).
Mozambique. "Algoa": 22°07'S. 35°45'E, 500 m. 9.06.1994: 1 <3 31.0 mm (SAM A41697). — 21°22'S, 35°40.E.
481 m. 13.06.1994: 3 (3 27.6-35.0 mm; 1 2 25.3 mm (SAM A41698). — 17°33'S, 38°27'E, 433 m. 14.06.1994:
1 ov. 2 54.7 mm. — I9°27'S. 36°51'E. 486 m, 14.06.1994: 1 5 30.6 mm; I 2 damaged (SAM A4I703). — 17°33.0'S.
38°27.0'E. 433 m. 15.06.1994: 1 ov. 2 54.3 mm (SAM A41704). — 16°20.0'S. 40°08.0'E. 500 m, 16.06.1994: 1 ov.
2 63.7 mm (SAM A41656). — 14°00'S, 37°31'E. 472 m. 17.06.1994: 1 ov. 2 51.6 mm; 1 2 54.4 mm (SAM A41700).
— 24°02'S, 35°45'E. 517 m, 20.06.1994: 1 ov. 2 47.4 mm: 1 2 37,8 mm (SAM A4I695). — Off Bazaruto I., 400-
500 m. 19.06.1994: 2 3 25.0. 35.2 mm (SAM A41699).
"Galathea ": stn 202, 25°20’S. 35°17’E, 535-580 m. 21.02.1951: 1 2 34.1 mm; 8 juvs (ZMC). — Stn 203, 25°20'S.
35°1 7'E, 680-730 m. 21.02.1951: 1 2 26.6 mm (ZMC).
Kenva. Mombassa. 4°31.5'S, 39°42.5'E. 21.03.1981: 1 2 40.3 mm (RMNH 34706).
Gulf of Aden. "Meteor 5": stn 249. I2°29.5'N. 45°38.7’E, 1299-1314 m, 8.03.1987: I 3 24.9 mm; 1 9 20.2 mm;
7 juvs (SMF 24654). — Stn 250. 12°40.4'N. 45°22.6'E. 907-917 m, 8.03.1987: 2 juvs (SMF 24655). — Stn 279.
12°52.5'N, 45°53.3'E. 1185-1186 m, 15.03.1987- 6 juvs (SMF 24656). — Stn 281. 12°38.5'N. 45°28.5'E. 1063-
358
B. GALIL
1068 m, 15.03.1987: 1 juv. (SMF 24657). — Stn 286, !2°20.9'N. 44°44.7’E. 725-751 m. 16.03.1987: 1 ov. 2
31.1 mm; 1 juv. (SMF 24658).
Types. — The male (21.1 mm cl) (NHMW 9712) collected off Sicily by von Grohmann, is the holotype of
Polycheles typhlops Heller. 1 862.
The female (26.2 mm cl) (MNHN-Pa 32) collected by the "Blake" stn 246 off Grenada, is the holotype of
Pentacheles agassizii A. Milne Edwards, 1880; the juvenile of the sample is the paratype. Other paratypes are at
the Museum of Comparative Zoology under the registration numbers MCZ 2789 ("Blake" sin 216, I specimen),
MCZ 2790 ("Blake" stn 47, 1 specimen). MCZ 2792, ("Blake" stn 274, 2 specimens, one broken), MCZ 3075
("Blake" stn 285, 1 specimen), MCZ 4042 ("Blake" stn 151, 1 male).
The largest male (80.0 mm total length) (IM 6767-6770/9 collected in the Andaman Sea by the "Investigator"
stn 115(1 lo3T40"N, 92°46'40"S, 344-402 m) is the lectotype of Pentacheles hextii Alcock, 1994; the other males
and the single female of the sample are paralectotypes.
The female (40 mm cl) found at Palermo's fish market, and deposited in the Zoological Museum of Palermo
(registration number C 44) is the holotype o( Polycheles doderleini Riggio. 1895 (M. Sara, pers. comm.).
Description. — Dorsal surface of carapace setose, minutely spinulate, granulate. Frontal margin concave.
Median submarginal tooth prominent. Single rostral spine antrorse. Internal angle of orbital sinus triangular,
spinose; external angle of orbital sinus pectinate. Internal margin of orbit mesially pectinate, dividing orbit into
two. Eyestalk bearing upcurvcd spine. Basal antennular segment produced anteriorly to a sharp point, mesial
margin spinose, anterolateral margin rounded, bispinose. Spines on lateral margins of carapace upcurved: lateral
spine formula 7-9:4-5:17-20; spines successively decrease in size posteriorly. Median postrostral carina granulate,
irregularly spinulate, median postcervical carina bearing paired spinules. Gastro-orbital carina obsolescent,
comprising five antrorse spines, anteriormost largest. Posterior margin of cervical groove spinose between
branchial carinae. Branchial carina prominent, parallel with lateral margin, bearing 12-15 spines. Region between
branchial and median postcervical carinae with shallow, oblique groove anteriorly, minutely spinulose posteriorly.
Posterior margin of carapace bearing three or four pairs of antrorse spines, submedian spines largest.
Abdominal tergites medially carinate. Carinae on tergites 2-5 bearing antrorse spine, increasing in size
posteriorly. Tergite 6 bearing mesially twin parallel rows of granules, confluent posteriorly. Anterior margin of
first tergite smooth or granulate, never more than irregularly disposed 18 granules. Anterior and posterior margins
of tergites 2-5 irregularly granulate. Second pleuron ovate, margins denticulate. Pleura 3-5 with curved granulate
rib mesially, margins denticulate. Telson with granulate median crest anteriorly, two minutely serrulate,
convergent carinae posteriorly. Inferior surface of uropodal expopod bicarinate.
Upper margin of merus of PI spinulate with subterminal claw-like spine, lower margin finely granulose.
Carpus half as long as merus, two serrulate rows distally on upper margin, subterminal spines on upper and lower
margins. Upper margin of propodus with several rows of antrorse spinules, sometimes reduced to granules; lower
margin with two minutely spinulate rows. P5 simple in male, chelate in female.
Color. — "Medial gastric and frontal region, scaphocerite and antennular peduncles, anterolateral edges and
posterior border of carapace, cervical groove, mid-dorsal carina of carapace, carinae and granulate edges of abdominal
terga bright orange. Remaining portions of carapace, abdominal terga and pleura white. Cheliped with proximal
portion of merus white, distal third of merus and carpus orange. Upper surface of propodus and fingers orange,
shading to pinkish white on underside and distal portions. All other pereopods white." (GRIFFIN & STODDART,
1995: 243). "Body ivory. Setae light brown. Ridges, margins, tips of spinules and granules on dorsal surface
orange-red. Anteromedian carapace, tail-fan and pleopods marked with orange-red patches. Eggs whitish." (CHAN &
Yu, 1993: 106). Color photos in Baba (1986); CHAN & Yu (1993); JONES & MORGAN (1994).
Remarks. Heller (1862) described Polycheles typhlops from a single specimen collected off Sicily. Not
only his description is remarkably accurate, but he realized its significancc:"Die meiste Uebereinstimmung zeigt
die Gattung noch mil dem von Desmarest beschrieben fossilen Krebse aus dem Solenhofer Kalkschiefer, eryon
Cuvieri. indem auch bei diesem ein abgeflachter Cephalothorax, ahnlich gestalte Fuhler und Fusse sich vorfinden"
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
359
(Heller, 1863: 21 1). Wood-Mason (1874: 180) thought that:"a species differing in no particular of generic
value from Deidamia leptodactyla et crucifer... had, years before, been described by Prof. C. Heller under the name
of Polycheles typhlops". Adensamer (1898) went further and put Willemoesia leptodactyla in synonymy with
P. typhlops.
A. Milne Edwards (1880) described Pentacheles agassizii from the West Indies. Bouvier (1905a: 480,
footnote) compared "les types des P. agassizi, recueillis par le "Blalce” avec un exemplaire de P. typhlops donne au
Museum par le Musee de Vienne" and considered them one species.
RIGGIO (1885: 101) described Polycheles doderleini from a specimen found in Palermo fish market, though
aware of Heller's opinion that "era certamente la femina del Polycheles typhlops da lui descriito".
ALCOCK (1894: 239) described Polycheles hextii quite accurately and noted "the peculiar formation of the
ophthalmic notches, which superficially are divided into two portions by the meeting of the edges across the
ophthalmic peduncle". BOUVIER (1905a: 481) recognized that "le P. Hexti Ale., presente les affinites les plus
etroites avec notre P. typhlops ", but Kemp and Sewell (1912: 24) realized that they "correspond so precisely that
the specific identity of the two forms cannot be doubted".
BALSS (1914: 599) described Polycheles intermedins and noted: "der frontal ist konkav; in seiner Mine stcht ein
Rostraldorn". This species is without doubt a junior synonym of P. typhlops.
DISTRIBUTION. — Worldwide: Mediterranean, Irish Sea, Alboran Sea, West Africa, Cape Verde Is, Rio Grande
Ridge, Bermuda, United States, West Indies, Gulf of Mexico, Caribbean Sea, New Caledonia. Australia, Japan,
East China Sea, Taiwan, Philippines, Indonesia, Indian Ocean. Maidive Is, Comoro Is. Madagascar, South Africa,
East Africa , Gulf of Aden; 77 m (present work) - 2055 m (Adensamer, 1898).
Fig. 30. _ Polycheles nplilops Heller, 1862. 2 38.8 mm, Rio Grande Ridge, "Prof. Mesyatsev" (ZMMU): a. dorsal view
of the cephalothorax; b. lateral view of the abdomen.
360
B. GALIL
Genus WILLEMOESIA Grote, 1873
Willemoesia Grote, 1873: 485. Type species by monotypy through the replaced name Deidamia Thomson. 1873:
Deidamia leptodactyla Thomson, 1873: 51. Placed on the Official List of Generic names in Zoology in Opinion 519
of the ICZN. Gender feminine.
Deidamia Thomson, 1873: 51. Type species by monotypy, Deidamia leptodactyla Thomson. 1878: 51. A junior
homonym of Deidamia Clemens, 1859. Placed on the Official Index of rejected generic names in Zoology in Opinion
579 of the ICZN. Gender feminine.
DIAGNOSIS. — Carapace dorsoventrally flattened, subrectangular or ovate. Antero-lateral angle of carapace
produced, spiniform. Lateral margins well defined, spinose; cervical and postcervical incisions dividing margin into
three parts. Front bearing single rostral spine, lacking well defined orbits. Eyestalks globose, fixed beneath and
parallel with anterior margin of carapace.
Cervical groove marked, arcuate, laterally bifurcate. Median postrostral and postcervical carinae well defined,
unarmed, granulate or spinose. Gastro-orbital and branchial carinae present, obsolescent or well defined, unarmed,
granulate or spinose. Two sublateral carinae starting near posterolateral margin of carapace, running to base of
antenna and base of PI; latter more distinct.
Abdomen somewhat laterally depressed. First abdominal tergite narrow, pleura fused, abbreviated. Abdominal
tergites 2-5 smooth or sculptured, medially carinate, carinae blunt or anteriorly spinose; sixth tergite smooth or
sculptured. Second abdominal pleuron larger, clypeiform, smooth; pleura 3-4 cordiform, successively diminishing
in size posteriorly; fifth pleuron triangulate. Abdominal sternites lacking median knob. Telson lanceolate, basally
swollen; two submedian carinae. Uropodal exopod lamellar, posteriorly rounded, bicarinate; endopod unicarinate,
lappet basally on interior margin.
Interior process of basal antennular segment lamellate, rounded, shorter than peduncle; mesial margin spinose,
perpendicular to segment; anterolateral angle not spinose. Outer antennular flagellum short, inner flagellum as
long as antennal flagellum. Renal process well developed, tubular, fitting into hollow on underside of basal
antennular segment. Antennal scaphocerite not extending to end of peduncle.
Buccal opening subquadrate. Mandible concave, cutting edge with triangular teeth of uneven size, lateral,
median teeth largest; mandibular palp biarticulate. Maxillula with two incurved lobes; anterior lobe more robust,
terminating in cornute claw. Maxilla with two slender protognathal lobes of equal length: scaphognathite large,
reniform, membraneous. First maxillipcd endopod slender; exopodal lobe membraneous, extending further back
than scaphognathite; exopod anteriorly divided into two lamellar lobes enclosing efferent passage. Second and third
maxillipcds slender, pediform; interior margins of basal segments denticulate; dactyls ending in cornute claw; third
maxilliped bearing long epipodite basally, cornute subterminal spines on interior margin of dactyl.
All pereiopods chelate in both sexes; fingers of fifth pereiopodal chela one sixth as long as propodus, whereas
in preceding pereiopods fingers about half as long as propodus. PI elongate, slender; P 2-5 successively shorter
posteriorly. Fingers of first cheliped crossing, pollex bearing on inner margin subtcrminal spine at right angle to
finger; inner margins of both pollex, dactyl furnished with row of overlapping cornute platelets, forming serrate
edge. Fingers of second and third pereiopodal chelae crossing distally, their inner margins set with row of
overlapping platelets; platelets not overlapping in fingers of fourth chela, missing entirely in fingers of fifth
pereiopodal chela. First male pleopod oar-shaped, spinulose swelling distally on inner margin, spinules distally on
mesial margin, distal margin folded internally. Pleopods 2-6 slender with rod-like appendix interna, second pair in
male with appendix masculina.
REMARKS. — Willemoesia is distinguished from other polychelid genera in having the five pairs of pereiopods
chelate in both sexes; lacking well defined orbits, its globose eyestalks fixed beneath and parallel with the anterior
margin of the carapace; and the pollex of the first cheliped bearing on its inner margin a subterminal spine at right
angle to the finger.
Nearly a century after it was established, it was felt that "a careful revision of the entire genus is badly needed"
(Wolff, 1961: 148). Kensley (1968: 297) considered the "Specific delimitation in this genus ... very difficult,
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
361
particularly as the number of specimens available is very low". Gore (1984: 1 ) declared that "Because the range of
morphological variation within the genus Willemoesia is not yet known, owing to paucity of specimens in the
8 known species, the taxonomic validity of several species within the genus remains uncertain". Of the eight
species described, only four are recognized as valid in this work.
Willemoesia forceps A. Milne Edwards, 1880
Fig. 31
Willemoesia forceps A. Milne Edwards, 1880: 64. — Faxon, 1895: 126. — Bouvier. 1905a: 480 (p.p.): 1905c: 3 ( p.p. );
1907: 62; 1917: 32, figs 1-4, pi. 10 figs 1-2, pi. 11 figs 1-6; 1925: 424. — De Man. 1916: 6 (list). — Siversten &
Holthuis, 1956: 41, figs 30-31. — Firth & Pequegnat, 1971: 75. figs 3, 13. — Pequegnat ei ai, 1971: 4. —
Rodriguez. 1980: 194. — Gore, 1984: 2. — Dawson. 1997: 25.
Willemoesia sp. - Gamo, 1980: 94. fig. 7a.
not Willemoesia forceps - ALCOCK, 1894: 230; 1899: 33; 1902: 155 [= W. leptodactyla (Willemoes-Suhm. 1875)].
not Willemoesia forceps - Bernard, 1953: 88. fig. 36 [= IV. pacifica Sund. 1920].
MATERIAL EXAMINED. — West Africa. "Talisman": 29°01’N. 19°15'E. 2713 m, 10.07.1883: I <3 37.0 mm
(MNHN).
"Pillsbury": stn 18. 5°01'N, 0°12'E. 3047-3129 m. 26.05.1964: 1 9 77.8 mm (RMNH). — Stn 266. I°12.5'N.
7°46'E, 2507-2562 m. 17,05.1965: 1 2 71.8 mm (RMNH).
"Waldo": 0°20.7'S, 5°48.4'E, 3244 m. 30.07.1971: 1 2 74.7 mm (MNHN-Pa 537).
West Indies. "Pillsbury": stn 1178. 19°14'N. 73°14'W. 1760 m. 30.06.1970: 1 <3 40.1 mm (RMNH).
"Iselin": stn 8, 21°34'N, 74°32’W, 2745 m, 18.11.1981: 1 <3 73.1 mm (USNM 347932). — Sin 5, 22°03'N.
74°48'W, 2728 m. 16.11.1981: 1 <3 60.0 mm (USNM 347932); 2 <3 46.6 59.8 mm (USNM 347932). — Stn 9. 21°27'N,
74°21'W. 2714 m. 18.11.1981: 1 2 40.7 mm (USNM 347932).
Caribbean Sea. "Pillsbury": stn 748. 1 1 °24.8'N, 67°10.rW, 1878-1784 m, 25.07.1968: 1 juv. (RMNH).
Venezuela Basin, "Bartlett" (id. R.H. Gore): stn 39. 15°08.93'N. 69°13.33'W, 3977-4047 m, 27-28.10.1981: I juv.
33.2 mm (NHM 1984:54). — Stn 93, 13°32.18'N, 64°40.54'W. 3406-3453 m, 28.11.1981: 1 2 52.7 mm (USNM
210820). — Stn 94, 13°32.2'N. 64°42'W. 3427-3475 m, 28-29.11.1981: 2 2 67.7. 80.8 mm (RMNH 35840).
Type. — The female (72 mm total length) (MCZ 4045) collected by the "Blake" stn 31 off Santa Cruz, Cuba
(24°33'N, 84°23'W, 3512 m) is the holotype.
Description. — Dorsal surface of carapace nearly glabrous, densely set with antrorse spinules. Rostral spine
upeurved. Frontal margin of carapace concave, external angles produced, spinose. Eyestalk globose, bearing spine
curved distad. Mesial margin of basal segment of antennule prominently spinose, pectinate. Lateral margin of
carapace spinose; spine formula 14-19:14-15:29-40. Median postrostral carina granulose, bearing pair of larger
spinules 2/3 way to cervical groove, followed by single spine. Median postcervical carina granulate, bearing larger
antrorse spines posteriorly. Gastro-orbilal carinae sinuous, minutely spinulate. Branchial carina prominent,
spinose. Posterior margin of carapace smooth.
First abdominal tergite tuberculate medially. Median dorsal carina on tergites 2-4 bearing antrorse spine. Fifth
carina blunt. Sixth tergite smooth, medially on posterior margin slight hump. Tergites 2-5 bearing well-defined,
submedian transverse groove. Second pleuron subcircular, smooth. Pleura 3-5 with obsolescent rib mesially,
margins unarmed. Telson with triangular hump anteriorly, smooth convergent carinae posteriorly. Margin of
uropodal endopod triangular.
First chelipeds thin, elongate; upper margin of merus minutely spinose, two claw-like spines subterminally.
lower margin prominently spinose; carpus 4/5 as long as merus, wider distally, spinose on upper margin. Upper
margin of chela bearing several irregular files of spines; lower margin bearing two files, spines of interior file
longer; single file along lower margin of finger.
Color. — "deep brick red overall, more brownish-red on cephalothorax. brighter red on abdominal somites.
Telson and uropods almost cherry red. Antennae and antcnnulac deep scarlet, including flagella. Perciopods brick
red, tending toward orange-red distally. Long bunched setae and hairs ornamenting carapace ranging from pale to
golden yellow." (GORE, 1984: 3).
362
B. GALIL
REMARKS. — W. forceps differs from the other three species of the genus in having the most spines along the
lateral margins of carapace, and a smooth sixth abdominal tergite except for a blunt median carina posteriorly.
Distribution. — West Africa, Azores, Saragasso Sea, West Indies, Caribbean Sea; 1760 m (present work) -
4064 m (GORE, 1984).
Fig. 31. — Willemoesia forceps A. Milne Edwards, 1880, 2 71.8 mm "Pillsbury" stn 266. I°12.5'N. 7°46'E. 2507-
2562 m (RMNH): a, dorsal view of the cephalothorax; b, lateral view of the abdomen.
Willemoesia inornata Faxon, 1893
Fig. 32
Willemoesia leptodactyla Bate, 1878a: 280 (p.p.); 1878c: 563 (p.p.); 1888: 163, pis 18P. 19c" (p.p.). — De Man,
1916: 6 (list) [non Willemoes-Suhm, 1875].
Willemoesia inornata Faxon, 1893: 195; 1895: 125, pi. 32 fig. 2. pi. 33 fig. 3. — Seeliger, 1901: 46. — Bouvier,
1917: 32 (key). — De Man, 1916: 6 (list). — Zarenkov, 1969: 81. — Bayer et a!.. 1970: 98. — Firth &
Pequegnat, 1971: 77. — Gore, 1984: 6. — Dawson. 1997: 26.
Willemoesia challenged Sund. 1920: 223. — Holthuis, 1952a: 79. — Bahamonde. 1963: 4. — Firth & Pequegnat.
1971: 75. — Zarenkov, 1969: 81. — Retamal. 1981: 16. — Gore. 1984: 6. — Wicksten, 1989: 304. — Dawson,
1997: 25.
Willemoesia - Wolpf, I960: 179; 1961: 148. fig. 17.
Material examined. — Eastern Pacific
Costa-Rica. "Galathea": stn 716. 9°23'N. 89°32'W, 3680 m, 6.05.1952: 3 3 41.6-46.1 mm; 2 2 47.8. 56.) mm;
1 ov. 2 47.5 mm (ZMC).
Gulf of Panama. "Albatross": stn 3382, 6°21'N, 80°4FW, 3227 m. 7.03.1891: 2 3 34.4. 37.9 mm; 3 2 42.7-
48.9 mm (USNM 21078).
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
363
"Pillsbury": sin 526. 6°53'N. 79°27'W, 3193-3200 m. 5.05.1967: 2 3 39.9. 37.7 mm; 6 2 35.9-49.5 mm (RMNH
23527).
"Gilliss": sin 1, 6°52'N, 79°28'W, 3200-3229 m. 13.01.1972: 2 <3 37.5, 39.4 mm; 5 2 43.3-54.6 mm (RMNH). —
Stn 3. 6°47'N. 79°13'W, 3045-3158 m, 14.01.1972: 3 3 38.3-44.4 mm; 5 2 41.6-51.0 mm (RMNH). — Sin 4. 6°42'N,
78°56'W, 3173-3208 m. 14.01.1972: 5 3 32.5-40.6 mm; 4 2 40.4-45.5 mm (RMNH). — Stn 6. 6°12'N. 78°33'W.
3475-3517 m, 15.01.1972: 2 3 31.5. 38.6 mm: 2 2 39.8, 51.5 mm (RMNH). — Stn 18. 6°31'N, 78°20'W. 3940-
3986 m. 17.01.1972: 1 3 34.9 mm; 5 2 43.5-58.0 mm (RMNH). — Stn 20. 6°45.2'N, 78°31.7'W. 3977-4005 m.
18.01.1972: 2 3 34.0. 38.3 mm; 5 2 40.0-50.6 mm (RMNH).
Ecuador. " Albatross ": stn 3399, W Cabo de San Francisco. 1°07'N, 81°04'W. 3132 m. 24.03.1891: 2 3 34.8.
38.6 mm (USNM 21079).
Colombia. "Galathea": stn 724, 5°44'N. 79°20’W. 2712-3330 m, 12.05.1952: 1 ov. 2 55.8 mm (ZMC). —
Stn 726. 5°49'N, 78°52'W. 3800 m, 13.05.1952: 5 3 39.6-42.9 mm: 4 2 43.3-57.8 mm (ZMC).
Galapagos Is. "Albatross": stn 3400, 0°36'S, 86°46'W, 2380 m. 27.03.1891: 1 2 34.9 mm (USNM 21080).
Arcturus Expedition: stn 53. 1°51'S, 89°50'W. 31 19 m, 25.04.1925. id. L. Boone Willemoesia intermedia, type
(unpublished): 1 3 42.3 mm (AMNH 12402). — Stn 40, 1 °5 1 S. 89°50'W, 2966 m. id. L. BOONE Willemoesia intermedia.
paratypes (unpublished): 2 3 35.1, 34.5 mm (AMNH 12403).
Chile. "Challenger": stn 298, 34°07'S. 73°56'W, 4005 m. 17.11.1875: 1 ov. 2 53.5 mm; I 2 45.6 mm (NHM
1888.22).
Juan Fernandez Is. "Challenger": stn 300. 33°42’S. 78°18’W, 2475 m. 17.12.1875: 1 3 33.7 mm (NHM
1888.22).
TYPES. — The male (37.9 mm cl) (USNM 21078) collected by the "Albatross" stn 3382 off Panama (6°21'N.
80°41'W, 3227 m), is designated here as the lectotype of Willemoesia inornata Faxon. 1893; the other specimens
in the samples are paralectotypes, as well as the specimens (USNM 21079) collected by the "Albatross" stn 3399
off Ecuador and those (USNM 21080) from stn 3400 off Galapagos Is.
The ovigerous female (53.5 mm cl) (NHM 1888.22) collected by the "Challenger" stn 298. off Chile (34°07'S,
73°56'W. 4005 m), is designated here as the lectotype of W. challengeri Sund. 1920, the female of the sample is a
paralectotype, as well as the male (33.7 mm cl) (NHM 1888.22) collected at stn 300 off Juan Fernandez.
Description. — Carapace oblong, margins subparallel, anteriorly convergent; surface densely covered with
antrorse spinules. Frontal margin transverse. Rostral spine upeurved. Internal orbital angle bearing small granulate
spine, lacking in some larger specimens. Anterolateral angle of carapace formed by large spine. Mesial margin of
basal antennular segment prominently spinose, pectinate. Spine formula of lateral margins of carapace 5-8:2-4:0-
10, lateral margins of carapace posterior to postcervical incision carinate. Median postrostral carina irregularly
spinulate, granulate anteriorly; median postcervical carina unarmed. Gastro-orbital carinae obsolescent, unarmed.
Branchial carina prominent, sinuous, unarmed, polished. Posterior margin of carapace unarmed.
First abdominal tergite smooth, bearing tubercle medially on anterior margin. Tergites 2-5 sculptured, bearing
deep submedian grooves, medially carinate, carinae culminating in antrorse spine anteriorly, spines increasing in
size posteriorly. Sixth tergite with margins, submedian crescents and lyre-shaped median carina swollen. Second
pleuron heart-shaped, pleura 3-5 with blunt rib mcsially, indented posteriorly. Telson with T-shaped hump
anteriorly, smooth convergent carinae posteriorly, unarmed margins, tip rounded. Posterior margin of uropodal
endopod rounded.
First chelipeds thin, elongate. Merus swollen proximally; upper margin granulose, subterminally trispinose;
lower margin spinose, spines larger proximally. Carpus 4/5 as long as merus, expanded distally; upper margin
spinose, distalmost spine largest. Upper margin of chela bearing two files of spines, spines of interior lile longer;
single file on lower margin of finger.
Color. — Carapace anteriorly pale orange-red, posterior to cervical groove almost colorless, with ridges, spines
darker, interior organs purple-red. Abdomen pale to bright reddish-orange; uropods, margin of pleura, ridges and
spins darker; legs, pleopods, antennae and antennulae bright reddish-orange (HOLTHUIS pers. obs.. specimens
collected by the "Gilliss" stn 3).
REMARKS. — Holthuis (1952a: 79) found "Willemoesia inornata Faxon, (1895) ... closely related to, if not
identical with W. challengeri." Gore (1984: 6) tentatively placed the "Galathea" specimens described by Wolff
(1961), in W. inornata. Having examined both the "Galathea" and type specimens, I confirm Gore's identification.
364
B. GAUL
W. inornata and IV. leptodactyla both have the sixth abdominal tergite with margins, submedian crescents and
lyre-shaped median carina swollen. W. inornata differ from IV. leptodactyla in having the lateral margins of
carapace posterior to postcervical incision bearing less than 10 spines, the upper margin of the first chela with two
files of spines, the lower margin with a single file; whereas in leptodactyla , the lateral margins of carapace
posterior to postcervical incision bear more than 20 spines, the upper margin of the first chela bears several files of
spines, the lower margin bearing two files.
Distribution. — Eastern Pacific: Gulf of Panama, Colombia, Ecuador, Chile, Juan Fernandez and Galapagos;
2380-4005 m.
FIG. 32. — Willemoesia inornata Faxon. 1893. 3 37.9 mm. Eastern Pacific, Panama. "Albatross" stn 3382. 6°21'N,
80°41W, 3227 m (USNM 21078): a, dorsal view of the cephalothorax; b. lateral view of the abdomen.
Willemoesia leptodactyla (Willemoes-Suhm. 1875)
Fig. 33
Deidamia leptodacnla Willemoes-Suhm. in Thomson, 1873: 51, 247, fig. 2. — Wood-Mason. 1874: 180. — Holthuis
1956: 114. — Hemming, 1958: 138. — Wolff, 1960: 180.
Willemoesia leptodactyla - GROTE, 1873: 485. — HUMBERT, 1874: 130. — WiLLEMOES-SUHM, 1875a: 50. pi 13 figs 1-9'
1875b: 577; 1875c: xxxiii. — THOMSON. 1877: 187, fig. 42. — Bate, 1878a: 280. pi. 13 figs 4-5 (p.p); 1878b: 484
(p.p); 1878c: 563 (p.p); 1888: 163, pis 18-20 (p.p.). — Norman, 1879: 175. — Pagenstecher. 1879: 37. —
Carus, 1885: 486. — Perrier, 1886: 293. — W. Marshall, 1888: 266. — Stebbing, 1893: 201 — Faxon 1 895-
118. — Murray, 1897: 388. — Giglioli, 1912: 158. — De Man, 1916: 5 (list). — Bouvif.r. 1917: 33, pi. I figs 6-
15. Sund, 1920: 221. — Stephensen, 1923: 64. — Sivertsen & Holthuis, 1956: 42. — Zarfnkov 1969- 81 _
Firth & Pequegnat, 1971: 79. — Gore, 1984. tab. 2. — Dawson. 1997: 27.
Polycheles leptodactyla - Wood-Mason, 1875: 132 (p.p.).
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
365
Willemoesia forceps - Wood-Mason & Alcock, 1891: 199. — Alcock, 1894: 230; 1899: 33; 1902: 155. — Bouvier.
1905a: 480 (p.p); 1905c: 5 (p.p) [non A. Milne Edwards, 1880],
Willemoesia indica Alcock, 1901b: 178, pi. 1 figs 1-la. — Alcock & McArdle, 1903, pi. 57 fig. 5. — Bouvier. 1917:
32 (key). — De Man, 1916: 6 (list). — Firth & Pequegnat, 1971: 77. — Gore, 1984: 5. — Dawson, 1997: 26.
Willemoesia secunda Sund, 1920: 223. — Firth & Pequegnat. 1971: 80.
not Willemoesia leptodactyla - Giglioli, 1881: 358; 1882: 5 [= P. sculplus Smith, 1880].
not Willemoesia ( Polycheles ) leptodactyla - GIGLIOLI, 1912: 186 [= P. sculptus Smith. 1880].
not Willemoesia leptodactyla - Bage, 1938: 9 [= W. pacifica Sund, 1920].
Material examined. — Atlantic Ocean. NE Atlantic. "Talisman". 42°19'N, 23°36'E. 4060 m,
24.08.1883: 1 3 35.0 mm; 2 $ damaged (MNHN-Pa 3). — 1 2 52.2 mm (MNHN-Pa 1172).
Dora 2: 46°59.5'N, 14°58.3'W, 4776 m. 29.03.1984: 1 juv. (RMNH).
Incal 2. 48°20.4'N, 15°14.6'W, 4823 m. 1.08.1976: 1 2 42.0 mm (MNHN-Pa 535).
BBC-96. 49°00.37'N. 13°41.42'W. 4488 m. 29.07.1966: 1 juv. (RMNH).
Canary Is. Cancap 3: stn 3.105, 29°50'N. 15°46'W, 3400-3550 m, 24.10.1978: 1 2 49.8 mm (RMNH).
Cancap 4: stn 4.135, 28°27’N. I8°27'W, 4000 m, 1.06.1980: 1 ov. 2 71.1 mm (RMNH). — Stn 4.166. 29°48’N,
1 5°56'W, 3427-3570 m. 6.06.1980: I 2 37.8 mm (RMNH). — Sin 4.180, 32°48'N, 15°18'W, 3315-3499 m,
9/10.06.1980: 1 <3 43.7 mm; 1 2 60.7 mm (RMNH).
Cape verde Is. Cancap 6: stn 6.002, I4°39'N, 23°21'W, 3650-3840 m, 4-5.05.1982: 1 2 53.8 mm (RMNH). —
Sin 6.023, 14°40'N, 24°51'W. 3900-3875 m, 6.06.1982: 3 juvs. (RMNH). — Stn 6.036, 15°04'N. 25°01'W. 4030-
4130 m, 8.06.1982: 1 3 49.0 mm; 1 2 41.5 mm (RMNH)
West Africa. "Gilliss": stn 93, 6°30.5'N, 22°06'W. 3446 m, 19.08.1973: 3 juvs. (RMNH 31671). — Stn 94,
5°03.8'N, 20°48.2'W. 2784 m. 20.08.1973: 1 juv. (RMNH). — Stn 99. 0°37.8'N, 18°20.8'W, 4620 m, 22.08.1973: 1
juv. (RMNH 31672).
"Pillsbury": stn 66, 3°31'N, 7°34'W, 4045-4008 m, 2/3.06.1964: 1 2 65.4 mm (RMNH).
Walda: 18°52.1'S, 7°23.1'E. 5124 m, 06.06.1971: 1 3 54.4 mm; 3 2 49.6-64.4 mm; 2 juvs (MNHN-Pa 653). —
18°28.5'S, 10°31.5'E, 3530 m. 28.06.1971: 1 2 57.9 mm (MNHN-Pa 536). — 17°30'S. 9°27.5'E. 4335 m. 30.06.1971:
1 2 47.4 mm (MNHN-Pa 650). — 12°21.4'S, 1 1°02.7'E, 3431 m. 11.07.1971: I 3 40.7 mm; 2 2 52.3, 54.3 mm
(MNHN-Pa 652). — 9°26'S. 10°33'E, 4080 m, 15.07.1971: 1 3 45.9 mm (MNHN-Pa 649). — 3°59'N, 3°42.9'E. 3225 m.
14.08.1971: 1 2 51.5 mm; 1 juv. (MNHN-Pa 651).
S. Atlantic. "Challenger": stn 133, 35°41'S, 20°55'W. 3420 m, 11.10.1873: 1 2 38.1 mm (NHM 1888.22).
NW Atlantic. "Challenger": stn 13, 21°38'N. 44°39'W, 3420 m. 4.03.1873: I 2 broken (NHM 1888.22).
Jamaica. "Pillsbury": stn 1180, 18°55'N, 73°53'W. 3109-3493 m. 1.07.1870: 1 3 46.0 mm (RMNH).
Venezuela Basin. "Bartlett": stn 39. 15°08.93'N, 69°13.38'W. 3977-4047 m, 27-28. 10.1981, id. R.H. Gore as
VV indica: 1 3 51.5 mm; 1 2 58.8 mm (NHM 1984.52); 2 3 35.5, 48.4 mm; 2 2 37.5, 38.5 mm; 4 juvs (NHM
1984.55). — Sin 65, 13°51.8'N. 67°52.4'W, 5000 m. 10.11.1981. id. R.H. Gore as VV. indica: 1 3 68.2 mm; 1 2
69.3 mm (LAM ex AHF 2491-02); 1 3 38.0 mm (LAM ex AHF 2491-03). — Stn 87, W Aves ridge. 13°36.36'N,
64°6. 1 2'W, 3640 m. 25.11.1981: 1 3 33.8 mm; I 2 66.5 mm; 2 juvs (USNM 210821); 1 ov. 2 82.2 mm (USNM
210822); 3 2 55.6-78.5 mm; 2 juvs (USNM 210823).
Indian Ocean. Tasman Sea." Galathea": stn 607. 44°18'S. 166°46'E. 3930 m, 7.01.1952: 1 2 35.6 mm (ZMC).
Java Deep. "Galathea": stn 474, 9°49'S, 114°13'E, 3920 m, 11.09.1951: 1 2 58.2 mm (ZMC).
Bay of Bengal. "Investigator": stn 117, 11°58'N, 88°52'17"E, 3197 m. 13.12.1890: 1 3 53 mm (1M 6871/9).
"Galathea": stn 299, 17°10'N. 84°34'E. 2935 m, 24.04.1951: 7 3 37.2-51.9 mm; 1 2 80.9 mm; 1 ov. 2 80.1 mm
(ZMC). — Stn 314, 15°54’N, 90°17'E. 2610 m, 3.05.1951: 5 3 35.3-42.0 mm; 7 2 37.2-45.3 mm (ZMC).
Mid Indian Basin. Safari 2: stn 2-CP 03. 5°48'N, 78°43’E, 3450 m, July-Scpt. 1981: 1 2 58.7 mm (MNHN). —
Stn 35-CP 28. 12°52.9'S, 79°32.1'E, 4950 m, July-Sept. 1981: 1 2 (MNHN-Pa 1751).
Madagascar. "Galathea": stn 235, 4°47'S, 46°19'E, 4940 m. 11.03.1951: 1 juv. (ZMC).
South Africa. "Galathea": stn 192, 32°00'S, 32°41'E, 3615 m. 5.02.1951: 3 3 25.4-50.1 mm (ZMC).
Types. — The female (broken) (NHM 1888.22) collected by the "Challenger" stn 13 in the NW Atlantic
(21°38'N, 44°39'W. 3420 m), is the holotype of Willemoesia leptodactyla (Willemoes-Suhm, 1875).
The male (53 mm cl) (IM 6871/9) collected by the "Investigator" stn 117 in the Bay of Bengal (1 1°58'N.
88°52T7"E, 3197 m), is the holotype of Willemoesia indica Alcock, 1901.
The female (38.1 mm cl) (NHM 1888.22) collected by the "Challenger" stn 133 in the South Atlantic
(35°41'S, 20°55'W, 3420 m), is the holotype of Willemoesia secunda Sund. 1920.
Description. — Carapace oblong, lateral margins subparallel, anteriorly convergent. Dorsal surface of
carapace densely covered with antrorse spinules. Rostral spine upcurved. Frontal margin of carapace transverse.
Internal angle of orbital sinus spinose. Eyestalk bulbous, bearing short curved spine. Mesial margin of basal
366
B. GALIL
antennular segment spinose, pectinate. Lateral margins of carapace spinose. spines increasing in size anteriorly,
anteriormost largest; spine formula 8-10:5-7:15-25. Median postrostral carina prominent, granulate; spine formula
variable; spines on median postcervical carina larger posteriorly. Gastro-orbital carina sinuous, sparsely spinulose.
Branchial carina sinuous, prominent, spinose. Posterior margin of carapace smooth.
First abdominal tergite smooth, bearing tubercle medially on anterior margin. Tergites 2-5 sculptured, with
deep submedian grooves, medially carinate, carinae culminating in antrorse spine anteriorly, spines increasing in
size posteriorly. Sixth tergite with swollen margins, submedian crescents and lyre-shaped median carina; median
carina more prominent posteriorly. Second pleuron heart-shaped; pleura 3-5 with blunt rib mesially, indented
posteriorly. Telson with T-shaped hump anteriorly, smooth convergent carinae posteriorly, unarmed margins, tip
rounded. Posterior margin of uropodal endopod rounded.
First chelipeds thin, elongate. Merus swollen proximally; upper margin granulose, distal ly bispinose,
distalmost spine larger; lower margin spinose, spines larger proximally. Carpus 3/4 as long as merus expanded
distally; upper margin spinose, distalmost spine largest. Upper margin of chela bearing several irregular files of
spines; lower margin bearing two files, spines of interior file longer; single file along lower margin of finger.
Color. — "very beautiful... it was of a fine red, while the hairs bordering it were yellowish" (Willemoes-
SUHM, 1875a: 53). "..brick orange to light pinkish orange overall... juveniles white, or translucent pink with
reddish highlights on cephalothorax. Sensory appendages and pereopods usually a solid salmon red or pink in larger
individuals, translucent white in smaller specimens." (GORE, 1984: 7).
Fig. 33. — WiUemoesia leptodactyla (Willemoes-Suhm, 1875), $ 66.5 mm, Venezuela Basin, "Bartlett" stn 87,
13°36.36'N, 64°46.12'W, 3640 m (USNM 210821): a. cephalothorax, dorsal view; b. abdomen, lateral view.
Remarks. — "W. leptodactyla does not have the embossment, as is clearly seen from Willemoes-Suhm's
(1875) illustration" (GORE, 1984: 8) - an omission of the characteristic lyre-shaped median carina and submedian
crescents from the early illustrations of IV. leptodactyla. coupled with Bate's (1878a: 280) erroneous count of the
lateral spines, contributed to the creation of several synonymies.
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
367
DISTRIBUTION. — Worldwide: east Atlantic and west Atlantic (Jamaica to Venezuela), Indo-West Pacific
(Tasman Sea. Philippines, Bay of Bengal, Madagascar. South Africa): 2396 m (ALCOCK, 1906) - 5124 m (present
work).
Willemoesia pacifica Sund, 1920
Figs. 34
Willemoesia leptodactyla Bate. 1878a: 280 (p.p); 1878c: 563 (p.p.); 1888: 163, pi. 19 fig. c (p.p). — De Man. 1916: 6.
— Bage, 1938: 9 [non Willemoes-Suhm, 1875],
Willemoesia pacifica Sund. 1920: 223. — HOLTHUIS. 1952a: 79. — BAHAMONDE, 1963: 4. — Zarenkov. 1969: 81. —
Firth & Pequegnat. 1971: 80. — Retamal. 1981: 16. — Wicksten, 1989: 304.
Willemoesia forceps - Bernard. 1953: 88. fig. 36 [non A. Milne Edwards. 1880].
Willemoesia bonaspei Kensley, 1968: 294. figs 4-5; 1974: 69; 1981b: 29. — Gore, 1984, tab. 2. — Griffin &
Stoddart. 1995: 251, figs 16-17. — Dawson, 1997: 25.
MATERIAL EXAMINED. — South Africa. Cape Point. 34°36'S. 17°00'E, 2745 m, 10.12.1959: 1 ov. 2 48.4 mm
(SAM A10543).
Juan Fernandez Is. " Challenger stn 300. 33°42'S. 78°18'W. 2475 m. 17.12.1875: 1 <J 46.3 mm (NHM
1888.22).
Kermadec Trench. "Dana": stn 3627, 30°08'S, 176°50'W. 5000 m, 14.12.1928: 1 juv. (ZMC 3274).
New Zealand. NZOI: stn U195. 34°31.5'S, 166°21.0'E. 2930-2928 m, 23.09.1982: 1 3 32.5 mm (NIWA). —
Stn VI 96, 33°03'S. 165°22.4'E. 3118-3120 m, 24.09.1982: 1 6 38.4 mm (NIWA).
TYPE. — The male (46.3 mm cl) (NHM 1888.22) collected by the "Challenger" stn 300. off Juan Fernandez Is
(33°42'S, 78°18'W, 2475 m), is the holotype of Willemoesia pacifica Sund. 1920.
The female (48.4 mm cl) (SAM A 10543) collected off Cape Point, South Africa, is the holotype of
Willemoesia bonaspei Kensley. 1968.
Description. — Carapace oblong, lateral margins subparallel, convergent anteriorly. Dorsal surface of
carapace close-set with antrorse spinules. Rostral spine long, upeurved. Frontal margin of carapace trigonally
excavate, internal angle of orbital sinus prominently spinose. Eyestalk bulbous, bearing curved spine. Mesial
margin of basal antennular segment prominently spinose. Lateral margins of carapace spinose, spines increasing in
size anteriorly, spine formula 6-10:5-8:18-30. Median postrostral carina well defined, spine formula variable;
postcervical median carina irregularly spinose, spines larger posteriorly. Gastro-orbital carina obsolescent, sinuous,
spinose. Branchial carina sinuous, prominent, spinose. Posterior margin of carapace smooth.
Abdominal tergites lacking obliquely transverse grooves, medially carinate; carinae 1-3 anteriorly dentate,
denticle increasing in size posteriorly. Tergites 4-5 carinate, lacking antrorse denticle. Sixth tergite smooth.
Abdominal pleura smooth, margins unarmed; second pleuron subcircular. Telson with triangular hump anteriorly,
polished submedian carinae posteriorly, margins spinulate, tip acuminate. Margin of uropodal endopod triangulate.
First cheliped long, thin. Upper margin of merus granulate, distally with claw-like spine; lower margin
spinose, spines longer proximally. Carpus 3/4 as long as merus; upper margin spinose, sublerminal claw-like
spines on upper and lower margins. Upper margin of chela with two spinose rows, spines on interior row longer;
lower margin with single row of spinules, continuing to Finger.
Remarks. — Bate (1888: 169) described one of the Juan Fernandez specimens as having "the telson longer
and more pointed and the outer foliaceous plates of the rhipidura somewhat pointed at the extremity" but thought it
"a decided variety of Willemoesia leptodactyla" . Kensley (1968: 297) found W. bonaspei "closely related to several
described species, such as leptodactyla (Willemoes-Suhm), pacifica Sund, or challengeri Sund", but mostly it
"resembles Bate’s 1888 plate 19 C (a specimen captured off the coast of Chile and named pacificus by Sund in
1920)". Griffin & Stoddart (1995: 251) thought W. bonaspei "is very close to W. pacificus and. as suggested
by Kensley, may be found to be synonymous with it when more material becomes available". Examination of
both types proved them right.
IV. pacifica differs from its congeners in lacking the deep submedian grooves on its abdominal tergites, in its
entirely smooth sixth abdominal tergite, and its acuminate telson.
368
B. GALIL
DISTRIBUTION. — The southern ocean: South Africa, Juan Fernandez Is, Kermadec Trench, New Zealand;
2475-5000 m.
Fig. 34. — Willemoesia pacifica Sund, 1920, <3 46.3 mm. holotype. Juan Fernandez I.. "Challenger" stn 300, 33°42'S,
78°18W. 2475 m (NHM 1 888.22): a. dorsal view of the ccphalothorax: b. lateral view of the abdomen.
CONCLUSIONS
The polychelids are prominent members of the deep sea biota, yet, choosing the bathyal as their abode, they arc
considered rare. Though there are a few reports of abundant hauls, like that of the "Pillsbury" off West Africa,
where surplus specimens were shoveled overboard (HOLTHUIS, 1991), most are known from a small number of
specimens scattered among distant museums. The polychelid collections assembled through die Musorstom deep-
sea cruises allowed for examination ol large series of specimens and analysis of the validity of characters used to
separate species and genera. The resulting taxonomic revision replaced the long established "three living genera in
the Polychelidae : Polycheles Heller, 1862; Stereomastis Bate, 1888; and Willemoesia Grote 1873" (Firth &
PEQUEGNAT, 1971:5), with five genera ol which only Willemoesia retains its primary meaning. The study of
extensive collections and most type specimens revealed many synonymies, and helped reduce the number of
polychelid species known from single specimens from twelve (Firth & PEQUEGNAT. 1971: 32) to one -
Homeryon asper (Rathbun, 1906). This taxonomic rearrangement resulted in reshuffling of geographic and
bathymetric ranges. Accordingly, five polychelid species are now known to occur only in the Atlantic, nineteen
occur solely in the Indo-Pacific, whereas eight are found in both oceans - contrary to the notion "Most polychelid
species appear to be endemic to particular ocean basins" (Firth & PEQUEGNAT. 1971: 16). Of the five species
listed by Firth & PEQUEGNAT (1971: 16) as endemic to the Atlantic, one, validus , occurs from the eastern
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHELIDAE
369
Pacific to the Indian Ocean, another, leptodactyla, from the Tasman Sea to South Africa, and a third, secunda, is
but a junior synonym of the latter. Two taxa, Polycheles perarmata and Willemoesia forceps, occur on both sides
of tropical Atlantic, whereas Willemoesia inornata was found on both sides of the isthmus of Panama.
Willemoesia pacifica and Polycheles suhmi inhabit the southern oceans, both Atlantic and Indo-Pacific. Some
species such as the above mentioned Homeryon asper, Pentacheles gihba, Polycheles cerata and Polycheles
irispinosus, all described nearly a century ago, are presumably rare and of limited range.
Table 1. — Geographical distribution of the Polychelidae.
1. Mediterranean Sea; 2. Northeast Atlantic; 3. West Africa; 4. South Africa; 5. Northwest Atlantic; 6. Tropical West
Atlantic; 7. East Pacific; 8. Northeast Pacific; 9. Central Pacific; 10. Southwest Pacific (New Guinea, Australia.
New Caledonia, New Zealand); 11. Southeast Asia (South China Sea, Philippines. Indonesia); 12. Indian Ocean;
13. East Africa
370
B. GALIL
All but four polychelid species occur only beyond the continental shelf, most are found in the mid-bathyal
depths of 500-1500 m. However, the four species of Willenioesia inhabit the abyssal, occurring exclusively at
depths greater than 2000 m - a specimen of leptodactyla even hauled from 5060 m.
On the 125th anniversary of the first deep-sea cruise, we note that the challenge of the "Challenger" has not
been met - there is still much to discover on the bottom of the sea.
Table 2. — Vertical distribution of the Polychelidae
ACKNOWLEDGMENTS
This work is dedicated to Prof. L.B. HOLTHUIS, who encouraged me to study the polychelids and provided me
with his own bibliographical notes.
Source : MNHN. Paris
REVIEW OF THE GENERA AND SPECIES OF THE FAMILY POLYCHEUDAE
371
I would like to thank Bertrand RICHER DE FORGES (IRD, Noumea) and Philippe BOUCHET (BIMM, Museum
national d'Histoire naturcllc, Paris) , who collected many of the specimens used in this study.
I am grateful to K. Baba, P. Berents, P. Clark, D. Eibye-Jacobsen, C.H.J.M. Fransen, D. Guinot,
K.-I. Hayashi, G. Innocenti, A. Johnston, R. Manning, J. Martin, M. van der Merwe, Y. Miya.
S. O'Shea, D. Platvoet, T. Roy, A. Sampey, V. Spiridonov, M. Turkay, E. Turner and K. Ueda for
entrusting me with valuable material from their collections. I thank M. Sara and P. Dworschak for their help
in locating type material. A. SHOOB took the photographs. J.-F. DEJOUANNET prepared the illustrations. I am
indebted to Alain CROSN1ER for his generous hospitality and careful editorship.
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Jl-TATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CAMPAGNES MIJSORSTOM, VOLUME 21 — RESULTATS DES C
Crustacea Decapoda: Porcellanopagurus Filhol
and Solitariopagurus Tiirkay (Paguridae),
from the New Caledonian area, Vanuatu
and the Marquesas: new records, new species
Patsy A. MCLAUGHLIN
Shannon Point Marine Center
Western Washington University
1900 Shannon Point Road
Anacortes, Washington, 98221 -90S IB. U.S.A.
ABSTRACT
The very interesting and rather specialized hermit crab genera Porcellanopagurus and Solitariopagurus are represented
in collections from the Musorstom cruises to New Caledonia and the Marquesas by four species of the former and three of
the latter. Among the species of Porcellanopagurus, three species, P. tridentatus Whitelegge, P. filholi de Saint Laurent &
McLaughlin, and P. chiltoni de Saint Laurent & McLaughlin have heretofore been reported only from Australia and
New Zealand; P. haptodactylus sp. nov. is a distinctive species, new to science. Solitariopagurus triprobolus Poupin &
McLaughlin is reported for the first time beyond the islands of French Polynesia, and the range of S. tuerkayi
McLaughlin is extended from the Kai and Tanimbar Island of Indonesia to New Caledonia. Vanuatu and Okinawa. A new
species, 5. trullirostris sp. nov., is described from New Caledonia and the Marquesas. The similarities and differences of
the two genera are elucidated, and an apparently rare attribute, a terminal anus, common to some species of both is
discussed.
The new species are fully described and illustrated, while diagnoses and illustrations of principal diagnostic characters
are provided for the previously described species. Keys to the Indo- and western Pacific species of Porcellanopagurus and
to the genus Solitariopagurus are included.
RESUME
Crustacea Decapoda : Porcellanopagurus Filhol et Solitariopagurus Tiirkay (Paguridae) de la
region neo-caledonienne, de Vanuatu, et des ties Marquises : nouvelles observations et nouvelles
espeees.
Les genres tres interessants et plutot specialises de bernard-l'ermite Porcellanopagurus et Solitariopagurus sont
representes. dans les collections rassemblees lors des campagnes Musorstom en Nouvelle-Caledonie et aux ties
Marquises, par quatre espfeces appartenant au premier genre et trois au second. Parmi les espeees de Porcellanopagurus.
McLaughlin, P.A., 2000. — Crustacea Decapoda: Porcellanopagurus Filhol and Solitariopagurus Tiirkay (Paguridae).
from the New Caledonian area, Vanuatu and the Marquesas: new records, new species. In: A. Crosnier (ed.), Resultats
des Campagnes Musorstom, Volume 21. Menioires du Museum national d'Histoire naturelle, 184: 389-414. Paris ISBN-
2-85653-526-7.
390
P.A. MCLAUGHLIN
irois especes, P. tridentatus Whitelegge, P. fillioli de Saint Laurent & McLaughlin, cl P. chiltoni de Saint Laurent &
McLaughlin n'avaient ete recoltees, jusqu'a present, qu'en Australie et en Nouvelle-Zelande. La quatrieme espece,
P. haplodactylus sp. nov., est une espdce distincte, nouvelle pour la Science. Solitariopagurus triprobolus Poupin &
McLaughlin est signale pour la premiere fois en dehors de la Polynesie franyaise, et la repartition de .S', tuerkayi
McLaughlin est etendue des lies Kai et Tanimbar, en Indonesie, a la Nouvelle-Caledonic. au Vanuatu et a Okinawa.
Une espece nouvelle, S. trullirostris sp. nov., est decrite de la Nouvelle-Caledonie et des Ties Marquises. Les simi larites
et differences des deux genres sont mises en lumiere et un caractbre apparemment rare, un anus terminal, commun a
quelques especes dcs deux genres, est discute.
Les especes nouvelles sont d6crites et illustrees en detail, tandis que des diagnoses et des illustrations des principaux
caracteres diagnostiques des especes dej& decrites sont publiees. Des cles d'identi fication pour les especes indo-ouest
pacifiques de Porcellanopagurus et pour les especes du genre Solitariopagurus sont proposees.
INTRODUCTION
The genera Porcellanopagurus Filhol, 1885a. and Solitariopagurus Tiirkay, 1986, have been distinguished from
other pagurid genera principally by their broad, vaulted carapaces, each provided with prominent lateral carapace
projections. In contrast to most other pagurid genera, it is the cephalothorax that provides the most reliable
diagnostic characters, e.g., size, shape and development of the rostrums, lateral projections and lateral carapace
lobes. Phylogenetically, Porcellanopagurus and Solitariopagurus have been considered classic examples of
carcinization in the Anomura (e.g., Borradaile, 1916b; WOLFF, 1961; Turkay. 1986), although
the carcinization hypothesis has recently been disputed by McLaughlin & Lemaitre (1997. in press).
Porcellanopagurus also has aroused behavioral and ecological interest because females have been reported as
carrying their eggs dorsally, and species of this genus utilize limpet or halves of bivalve shells as protective
coverings. Solitariopagurus is a more recently recognized genus, with fewer representatives; however, females
similarly appear to carry their eggs dorsally, and for at least two species the protective covering of choice is
a bivalve shell.
A seemingly rare morphological attribute shared by species of Porcellanopagurus and Solitariopagurus. but
overlooked until now, sets these genera apart, not only from most other hermit crabs, but from the vast majority
ol decapods as well. In their description of Porcellanopagurus adelocercus McLaughlin & Hogarth, 1998. these
authors commented that the telson was separated from the tergite of the sixth abdominal somite by membranous
tissue, and that the amount of separation increased with increased animal size. Additionally, although they did not
comment, they illustrated (MCLAUGHLIN & HOGARTH, 1998, fig. 29) the very unusual condition of a terminal
anus. In the new species of both Porcellanopagurus and Solitariopagurus described herein, the anal opening is also
terminal. In these two taxa, the telson similarly would seem to be separated by some distance from the posterior
margin of the tergite of the sixth abdominal somite, and carried ventrally under the abdomen. Thus it might appear
that the terminal position of the anus could be a function of this correspondingly distinctive telsonal development.
However, in the course of examining the three other species of Porcellanopagurus occurring in New Caledonia and
the Marquesas, as well as both Solitariopagurus tuerkayi McLaughlin, 1997, and 5. triprobolus Poupin &
McLaughlin, 1996, the terminal, or nearly terminal, anal position appears to be characteristic of both genera and
not simply correlated with the positioning of the telson.
Although the two genera are, in other morphological aspects, superficially also very similar, several aspects of
their morphology immediately demarcate them. The most obvious are the lateral carapace projections. In
Porcellanopagurus (Fig. la) two pairs of lobes develop from the lateral margins of the shield, anterior to
the cervical groove; a pair of prominent posterior carapace lobes develop directly behind the cervical groove,
connected usually by a narrow median element. In contrast, three pairs of lobes develop anterior to the cervical
groove in Solitariopagurus (Fig. lb) while the posterior lobes are represented by a moderately broad median and
two small lateral elements. McLaughlin (1997) referred to the tinea transversalis of S. tuerkayi as being
represented by a well calcified transverse rod. In reality, the tinea transversalis , a chitinous hinge (cf. PILGRIM.
1973) separating the posterior margin of the shield from the median portion of the posterior carapace, is rarely
visible externally in Solitariopagurus species. McLaughlin’s (1997) tinea transversalis is actually the median
clement of the posterior carapace lobes of that species.
Source . MNHN, Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOL1TARIOPAGURUS
391
Fig. 1. — Diagrammatic cephalothorax: a, Porcellanopagurus edwardsi Filhol, 1885: b. Solitariopagurus triprobolus
Poupin & McLaughlin, 1996.
ac 1: anterior "carapace lobe 1; ac 2: anterior carapace lobe 2; ac 3: anterior carapace lobe 3; cs: cardiac sulcus;
cbs: sulcus cardiobranchialis ; eg: cervical groove; Ip: lateral projection; It: location ot linea transversalis ;
pci: posterior carapace lateral lobe or element; peme: posterior carapace median element; pip: posterolateral plate;
pmp: posteromedian plate; r: rostrum.
A less obvious, but equally significant difference is the gill number. Eleven pairs of biserial phyllobranchiate
gills are present in species of Porcellanopagurus but only ten pairs in Solitariopagurus species. Pleopods are
entirely absent in males of both genera; however, different levels of secondary sexual modification occur. In some
Porcellanopagurus species, one or both coxae of the fifth pereopods may be somewhat drawn out or elongated, and
rarely a very slight extrusion of the vas deferens produces a short right sexual tube (cf. McLaughlin &
HOGARTH, 1998). In contrast, species of Solitariopagurus all have very well developed male sexual tubes. These
tubes appear also to be true coxal extensions but with development comparable to that seen in certain species of
the coenobitid genus Coenobita Latreille, 1829. At least in S. triprobolus, the tubes exhibit a slight amount of
calcification, whereas in 5. trullirostris sp. nov. they are membranous. Females of both genera have only three
unpaired pleopods, pseudo-dorsally positioned (cf. McLaughlin & Hogarth, 1998). However, females ol
Porcellanopagurus have paired gonopores, while females of Solitariopagurus have a single left gonopore. Although
the use of halves of bivalves shells as carcinoecia, is common to species of both genera, the structure of the fourth
pereopods in Porcellanopagurus species is usually semichelate. and that ot the fifth, chelate. In contrast both the
fourth and fifth pereopods of species of Solitariopagurus are subchelate.
Currently 12 species are assigned to Porcellanopagurus. These include four from the north and central western
Pacific: P.japonicus Balss, 1913 and P. truncatifrons Takeda. 1981, from Japan, P. nihonkaiensis Takeda, 1985.
from the Ogasawara Islands, and P. belauensis Suzuki & Takeda, 1987, Irom the Palau Islands. The description of
one additional new species from Guam is in preparation (Kropp & Eldredge, personal communication). Two
species have been described from the eastern Pacific; P. platei Lenz, 1902, from Juan Fernandez Island (incorrectly
reported as Eastern Island by dc SAINT LAURENT and McLaughlin, 2000). and P.foresti Zarenkov. 1990 from
the Sala-y-Gomez Ridge. Four species have been reported from Australian-New Zealand waters: P. edwardsi Filhol.
1885, P. tridentatus Whitelegge, 1900, Porcellanopagurus filholi de Saint Laurent & McLaughlin, 2000. and
Porcellanopagurus chiltoni de Saint Laurent & McLaughlin, 2000. Porcellanopagurus jacquesi McLaughlin, 1997,
was recently described from the Kai Islands of Indonesia, and in the Indian Ocean, P. adelocercus, a dwarf species.
392
P.A. MCLAUGHLIN
was described from the Seychelles. One species, P. haptodactylus sp. nov„ is described herein, and range
extensions are reported for P. tridentatus, P. chiltoni and P.filholi. A key to the Indo- and Western-Pacific species
of the genus is provided.
To date, only three species have been assigned to Solitariopagurus. The type species, S. profundus Tiirkay,
1986, was described from the Red Sea. More recently S. triprobolus was described from French Polynesia and
S. tuerkayi from the Kai and Tanimbar Islands of Indonesia. Range extensions are reported for S. triprobolus and
S. tuerkayi , one new species is described, and a key to species of the genus is provided.
MATERIALS AND METHODS
Information on the cruises where the samples studied herein were collected can be found in Richer de FORGES.
1990 for BiOCAL, Chalcal 2. MUSORSTOM 4, 5 and 6, Smib 2; LEHODEY et al, 1992 for Beryx 1 1 : RICHER DE
Forges, 1993 for Volsmar and Smib 5; Bouchet, 1994 for the Montrouzier Expedition; Richer de
Forges & Chevillon, 1996 for Bathus 1; Richer de Forges et al., 1999 for Musorstom 9.
The specimens are deposited in the Museum national d'Histoire naturelle, Paris (MNHN). Comparative
materials of the Australian, New Zealand, and South African species of Porcellanopagurus have come from the
collections of the Australian Museum, Sydney (AM), the New Zealand Oceanographic Institute, now part of the
National Institute of Water and Atmospheric Research, Wellington (NZOI), the Museum of New Zealand Te Papa
Tongarewa (formerly the National Museum of New Zealand), Wellington (NMNZ), and the South African
Museum, Cape Town (SAM). One additional specimen of Solitariopagurus tuerkayi from Okinawa is deposited in
the collections of the National Museum of Natural History, Smithsonian Institution. Washington. D.C. (USNM).
In the lists of material examined the capital letters preceding the station numbers refer to the gear used:
CC otter trawl (shrimps), CP beam trawl, DR rock dredge. DC Charcot dredge; DW Warcn dredge, while the
abbreviation ov. indicates ovigerous female.
General terminology follows that of McLaughlin (1974); the terms semi- and subchelate are used according
to the definition of McLaughlin (1997), and gill type as defined by MCLAUGHLIN & DE SAINT LAURENT
(1998). Terminology for the carapace grooves, linca, and sulci follows that given by PILGRIM (1973) and
Morgan & Forest (1991). One measurement, shield length, indicative of animal size, is given in parentheses
following the number and sex of the specimens.
SYSTEMATIC ACCOUNT
PORCELLANOPAGURUS Filhol, 1885
Porcellanopagurus Filhol, 1885a: 47; 1885b: 23; 1885c: 410. — Borradaile, 1916b: 111. — BENNETT, 1932: 520. —
FOREST. 1951a: 82; 1951b: 182. — Wolff, 1961: 28. — Miyake, 1978: 117. — Turkay, 1986: 140. —
McLaughlin, 1997: 464. — De Saint Laurent & McLaughlin, 2000: 105.
Type Species. — Porcellanopagurus edwardsi Filhol. 1885, by monotypy. Genus masculine.
DIAGNOSIS. — Eleven pairs of biserial phyllobranchiate gills. Anterior carapace vaulted and well calcified;
lateral margins of shield developed into two pairs of blunt or spinose lobes. Rostrum and lateral projections widely
separated. Anterolateral plates of posterior carapace calcified anteriorly and usually drawn out into projecting lobes;
remainder of carapace membranous or with areas of some calcification. Ocular acicles reduced, simple; obscured
from dorsal view. Maxillule with external lobe of endopod slightly produced, not recurved. Third maxilliped with
well developed crista dentata and 1 accessory tooth.
Chelipeds unequal; propodal-carpal articulation of right with considerable clockwise rotation. Ambulatory legs
generally similar. Fourth pereopods usually semichelate; with single row of scales in propodal rasp. Fifth
pereopods chelate.
Source : MNHN, Paris
CRUSTACEA DECAPODA: PORCELIANOPA G UR US AND SOLITARIOPAGURUS
393
Males usually without extrusion of vas deferens forming membranous sexual tube on coxa of one or both fifth
pereopods, but often with coxae slightly drawn out ventromesially; without paired or unpaired pleopods. Females
with paired gonopores; no paired pleopods, unpaired left pleopods on somites 2 to 4. Abdomen usually globular,
membranous, but with tergites at least faintly delineated. Uropods symmetrical or slightly asymmetrical. Telson
partially calcified or entirely membranous, telsonal tergite contiguous or not with tergite of sixth abdominal
somite; anus terminal or nearly so.
Remarks. — McLaughlin (1997) and McLaughlin & HOGARTH (1998) referred inaccurately to the weakly
calcified anterolateral plates of the posterior carapace as the "cardiac sulci". As clearly indicated by MORGAN &
FOREST (1991), the cardiac sulci laterally delineate the posterior median plate, which in Tisea grandis Morgan &
Forest, 1991 is represented by a calcified median triangular region. In Porcellanopagurus species, a pair of
triangular lateral plates are frequently weakly calcified (Fig. la). These appear to represent the posterolateral plates
ol PILGRIM (1973) that are bounded laterally by the sulci cardiobranchialis, and mcsially. by the cardiac sulci.
which also delineate the lateral boundaries of the weakly calcified posteromedian plate.
Reference to the telson was intentionally omitted in the generic diagnosis of Porcellanopagurus given by
McLaughlin (1997). As pointed out by McLaughlin & Hogarth (1998), although in most species of the
genus, a typical pagurid-like telson is developed, in two Japanese species, P. truncatifrons and P. nihonkaiensis.
the telson was described or implied as being absent. Of 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. McLaughlin &
HOGARTH (1998). in their description of P. adelocercus , noted the fact that while the telson would also appear to
be lacking if a specimen was viewed only dorsally, a membranous telson, apparently not contiguous with the sixth
abdominal somite could be detected ventrally. A similar condition occurs in P. haptodelctylus sp. nov.. and such
well may prove to be the case for the two Japanese species.
Key to the Indo- and Western Pacific species of Porcellanopagurus
1 . Rostrum truncate . 2
— Rostrum triangular or sublriangular . 4
2. Second lateral carapace lobe fringed with long setae . 3
— Second lateral carapace not fringed with long setae . P. adelocercus
3. First lateral carapace lobe with distal and median spinule; rostrum usually trilobed. Dactyls
of ambulatory legs each with 10 or 11 corneous spines on ventral margin .
. P. haptodactylus sp. nov.
— First lateral carapace lobe with only distal spinule; rostrum not trilobed. Dactyls of
ambulatory legs each with 7 corneous spines on ventral margin . P. truncatifrons
4. Posterolateral projection of carapace acute, spinulose or spinose . 5
— Posterolateral projection of carapace bluntly rounded . 8
5. Rostrum obtusely triangular; shield length (including rostrum) slightly less to slightly
more than shield width (including lateral projections) . 6
— Rostrum acutely triangular; shield length (including rostrum) considerably less than shield
width (including lateral projections) . p. filhoti
6. Midpoint of rostrum distinctly produced, often giving quasi trilobed appearance . 7
— Midpoint of rostrum not distinctly produced, not giving quasi trilobed appearance .
. P . jacquesi
1. Ventromesial surfaces of palms of both chelae thickly set with tufts of long setae. Propodi
of ambulatory legs longer than dactyls . />. japonicus
394
P.A. MCLAUGHLIN
— Ventromesial surfaces of palms of both chelae not thickly set with lulls of long setae.
Propodi of ambulatory legs approximately equal to length of dactyls . P . tridentatus
8. Tergite of sixth abdominal somite divided into 4 subequal lobes . 9
— Tergite of sixth abdominal somite divided into 2 subequal lobes . P. nihonkaiensis
9. Second lateral carapace lobes well developed, produced distinctly beyond lateral level of
first . 10
— Second lateral carapace lobes not well developed, not produced distinctly beyond lateral
level of first . P- chiltoni
10. Ambulatory legs stout; propodi and carpi each with irregular rows of prominent often
spinulose protuberances . P- edwardsi
— Ambulatory legs slender; propodi and carpi without irregular rows of prominent often
spinulose protuberances . P. belauensis
Porcellanopagurus haptodactylus sp. nov.
Fig. 2
MATERIAL EXAMINED. — New Caledonia. Montrouzier Expedition: stn 1250. Touho, 20°46.7'S. 1 65° 1 3.7'E.
3-6 m, vase sableuse. 1.09.1993: 1 2 (1.4 mm) (MNHN-Pg 5861). — Stn 1318, Kournac, 20°41.4'S, 164°14.8’E, 20-
30 m, reef outer slope, 24.10.1993: 1 6 (2.3 mm) (MNHN-Pg 5862).
Types. — The male from Stn 1318 is the holotypc; the female from Touho is the allotype.
DESCRIPTION. — Anterior carapace (Fig. 2a) with shield length slightly shorter than maximum breadth;
anterior margin between rostrum and lateral projections slightly concave; lateral carapace margins each with short
acute spine in distal 0.35 of first lobe; roundly triangular, unarmed lobe at mid-length; moderately short, blunt,
subacute or denticulate lobe posterior to cervical groove, median element very narrow; dorsal surface of shield
strongly calcified, posterolateral regions weakly delineated; posterior margin truncate. Posterolateral plates weakly
calcified, cardiac sulci and sulci cardiobranchialis extending nearly to posterior carapace margin; posteromedian
plate not noticeably delineated. Rostrum well developed, broad, truncate or prominently trilobed (Fig. 2a-b),
reaching considerably beyond bases of ocular peduncles. Lateral projections triangular; well developed but produced
only to mid-rostral length.
Ocular peduncles short, stout, less than half length of shield, with submedian constriction; corneas slightly
dilated, diameter approximately 0.5 length of peduncle. Ocular acicles very small, acutely triangular.
Antennular peduncles when fully extended, overreaching ocular peduncles by approximately 0.5 length of
penultimate segment. Ultimate segment with few setae dorsally. Basal segment unarmed.
Antennal peduncles overreaching ocular peduncles by approximately 0.5 length of ultimate segment. Fifth and
fourth segments with few scattered, short setae. Third segment unarmed. Second segment with dorsolateral distal
angle produced, terminating acutely; dorsomesial distal angle rounded. First segment produced, unarmed or with
tiny terminal spinule. Antennal acicle reaching to or beyond distal margin of fourth peduncular segment,
terminating acutely and with few moderately long setae. Antennal flagella missing.
Thoracic sternites 3-8 (Fig. 2c) unarmed. Sternite of third maxillipeds (third thoracic) bluntly triangular on
either side of median concavity. Sternite of chelipeds with left side enlarged, partially fused with sternite of second
pereopods. Sternite of second pereopods broad, plate-like, with incomplete median longitudinal groove. Sternite of
third pereopods with subrectangular anterior lobe. Sternite of fourth pereopods concealed from direct ventral view
by sternite of third. Sternite of fifth pereopods narrow, well separated from preceding sternites.
Right cheliped (Fig. 2d) stout; longer and considerably stronger than left. Dactyl slightly shorter than palm; ar¬
ticulating obliquely; cutting edge with 3 or 4 calcareous teeth; terminating in small calcareous claw, slightly over¬
lapped by fixed finger; dorsal surface convex, unarmed or minutely granular, dorsomesial margin with very low,
minutely spinulose or denticulate ridge; ventral surface with few tufts of setae. Palm longer than carpus; somewhat
Source : MNHN, Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOLITARIOPAGURUS
395
swollen dorsoventrally; dorsal surface convex, dorsomesial and dorsolateral margins slightly elevated, unarmed or
minutely spinulose or scalloped; fixed finger with few tufts of short setae dorsally and ventrally; cutting edge with
2-4 distinct calcareous teeth. Carpus slightly longer than merus; dorsomesial and dorsolateral margins not
distinctly delimited; lateral and mesial surfaces with low, weakly rugose or microscopically spinulose, transverse
ridges extending onto dorsal surface. Merus broadly subtriangular; dorsal margin with several transverse ridges and
Fig. 2. — Porcellanopagurus haptodactylus sp. nov. a, c-i. holotype S (2.3 mm) from Montrouzier Expedition Stn 1318;
b, allotype (1.4 mm) from Montrouzier Expedition Stn 1250: a. cephalothorax and cephalic appendages (aesthctascs
omitted); b, rostrum and anterior margin of shield; c, thorax (ventral view); d. carpus and chela of right cheliped
(dorsolateral view); e, carpus and chela of left cheliped (dorsal view); f. right second pereopod (lateral view); g. left
third pereopod (lateral view); h. propodus and dactyl of left fourth pereopod (lateral view); i, tergile of sixth
abdominal somite, uropodal protopods, and telson. Scales equal 1 mm (a-g, i) and 0.5 mm (h).
396
P.A. MCLAUGHLIN
few setae; ventromesial margin with I prominent tubercle or subacute spine at proximal angle; ventrolateral
margin with row of few to several acute spines. Ischium with row of small spinules on ventromesial margin.
Left cheliped (Fig. 2e) with 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 or minutely
granular, dorsomesial margin not delimited or microscopically serrate; mesial and ventral surfaces with numerous
long setae. Palm approximately 0.5 length of carpus; dorsal surface smooth or microscopically granular,
dorsolateral margin with row of very tiny spinules, dorsomesial margin unarmed or microscopically denticulate;
ventral surface with scattered long setae; cutting edge of fixed finger with row of very small calcareous teeth,
terminating in corneous claw. Carpus slightly longer than merus; dorsolateral surface with low transverse ridges,
dorsal surface mcdianly with few low, short transverse ridges, in female forming longitudinal row of short rugose
ridges; dorsomesial margin with very low granular or very weakly tuberculate ridge, mesial face with few short
transverse low ridges and moderately long setae. Merus subtriangular; dorsal surface with few short transverse
ridges and setae; ventromesial margin with 1 small spine adjacent to proximal angle; ventrolateral margin with row
of widely-spaced stronger spines. Ischium with row of very small tubercles on ventromesial margin; row of small
spines on ventrolateral margin.
Ambulatory legs (Figs 2f-g ) moderately short; generally similar. Dactyls 0.60-0.75 length of propodi; in dor¬
sal and lateral views, nearly straight; dorsal margins with moderately sparse setae; mesial and lateral faces with few
short setae; ventral margins each with row of 9-1 1 corneous spines. Propodi equal to or slightly longer than carpi;
dorsal surfaces with row of low protuberances and sparse setae; ventral margins each with row of 5-7 spinose setae.
Carpi shorter than nreri; dorsal margins with few low protuberances, no distinct spine at distal angle; lateral faces
each with longitudinal ridge dorsally; ventrodistal margins each with spiniform seta. Meri with low protuberances
on dorsal margins, ventral surfaces slightly oblique (mesial view), ventromesial margins unarmed; ventrolateral
margin unarmed, but formed as distinct ridge. Fourth pereopods with dactyl articulating perpendicularly with lateral
face of propodus (Fig. 2h). Propodus of fifth pereopods with small rasp of corneous scales.
Males with coxae of fifth pereopods (Fig. 2c) slightly asymmetrical, somewhat produced posteromedially,
gonopores each masked by long setae, no sexual tubes.
Abdomen short, globular. Tcrgite of first abdominal somite subrectangular, membranous and weakly delineated.
Tergites of somites 2 to 5 also membranous, moderately broad, weakly indicated. Tergite of sixth somite (Fig. 2i)
partially calcified, divided into subrectangular anterior and somewhat broader subquadrate posterior lobes by
transverse furrow, distal lobe with incomplete median longitudinal furrow. Uropods symmetrical; protopods each
with posteriorly directed subacute protuberance. Tergite of telson (Fig. 2i) carried ventrally and not contiguous
with posterior tergal margin of sixth somite; entirely membranous, with complete transverse suture separating
poorly defined anterior portion from posterior portion; posterior lobes separated by small median cleft, terminal
margins rounded, unarmed; anus terminal.
Variation. — As P. haptodactylus is known from only two specimens, it is not clear whether certain of the
variations noted are size or sex related or both. Most noteworthy is the difference in the rostrum of the male
holotypc and female allotype. As may be seen in Figure 2a, the rostrum of the male is clearly trilobed. whereas
that of the female (Fig. 2b) is only weakly produced medianly. The right chela of this much smaller female is
considerably narrower, but the armature of both chelipeds is stronger. The tergal portion of the telson of the female
is nearly contiguous with the tergite of the sixth abdominal sternite, while in the male it is widely separated by a
membranous area. This type of separation was reported by McLaughlin & Hogarth (1998) to be size-related in
the similar species, P. adelocercus.
COLOR. — Unknown.
Habitat. — One specimen was accompanied by part of a bivalve mollusk shell.
Distribution. — Known only from New Caledonia (East coast: Touho; West coast: Koumac); 3-30 m.
Etymology. — The specific name is taken from the Greek liapto meaning grasp, and daktylos , meaning
finger and denotes the very characteristic articulations of the dactyls of the fourth pereopods, clearly adapted for
grasping the crab’s carcinoecium.
Source : MNHN, Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOUTARIOPAGURUS
397
REMARKS. — Porcellanopagurus haptodactylus is most closely allied to P. truncatifrons and P. adelocercus,
sharing with both a truncated rostrum. While the rostrum of the holotype male more closely resembles that
of P. adelocercus in being distinctly trilobed, the rostrum of the allotype female more closely approaches that of
P. truncatifrons. It is probable that these rostral differences are size related, as McLaughlin & Hogarth (1998)
reported similar size-related rostral differences in the rostrums ot P. adelocercus. From the limited data available,
it would appear that the truncate rostrum of P. truncatifrons does not represent a subadull developmental condition.
Takeda’s (1981) specimen has a shield length greater than either specimen of P. haptodactylus and only 0.1 mm
smaller than the largest specimen ol P. adelocercus. Porcellanopagurus haptodactylus is also distinguished from
P. truncatifrons by the smoother, less setose shield surface of (he former species and the greater number of spines
on the ventral margins of the ambulatory dactyls. Similarly, P. haptodactylus has a greater number of dactylar
spines than is seen in P. adelocercus, and the carpus of the left cheliped is more strongly armed in the former
species. Additionally, the length of the antennular peduncles is considerably greater in P. haptodactylus.
McLaughlin (1997) remarked that Takeda’s [1981, Fig. 3(3)] illustration of the fourth percopod of
P. truncatifrons appeared to have been inverted, thus giving the appearance of a dorsally directed dactyl. Although
the dorsally directed propodal rasp in Takeda’s ligure does suggest inversion, it is very possible that the author
was trying to illustrate a dactyl articulated away from the plane of the propodus. A similar condition is seen in the
dactyls of the fourth pereopods of P. haptodactylus , although the dactylar articulation is 90° from (perpendicular to)
the plane of the propodus, not 180° as suggested by Takeda’s description.
Porcellanopagurus filholi de Saint Laurent & McLaughlin, 2000
Fig. 3
Porcellanopagurus edwardsi - Borradaile 1916a: 97 [not Porcellanopagurus edwardsi Filhol. 1885b],
Porcellanopagurus sp., "probably P. edwardsi" - BORRADAILE, 1916b: 111, figs 1-13.
Porcellanopagurus edwardsi - BALSS, 1930: 196 (in part). — BENNETT, 1932: 470 (in part). — GORDAN, 1956: 339 (in
part) (lit.). — Zarenkov, 1990: 239 (in part).
Porcellanopagurus sp. - KENSLEY, 1977: 167, fig. 2.
Porcellanopagurus filholi de Saint Laurent & McLaughlin. 2000: 1 14, fig. 36. pi. 3, fig. 3.
Material EXAMINED. — New Caledonia. Musorstom 4: stn CC 201. 18°55.8'S, 163°13.8'E 490 m
20.09.1985: 1 damaged (8.2 mm) (MNHN-Pg 5863).
South Africa. "Mering Naude" : stn 43, (north of Richards Bay, KwaZulu-Natal), 28°45 5'S 32°24 5'E 4'>0-
360 m, 29.05.1975: 1 $ (3.1 mm) (SAM 15302).
diagnosis. — Cephalothorax (Fig. 3a) strongly calcified; lateral margins each with 3 distinct lobular or wing-
like processes: first lobe moderately broad, anteriorly flat or rounded with prominent spike-like or spiniform
projection; acute or subacute triangular or subtriangular lobe at mid-length; and considerably larger, elongate, acute
or subacute lobe posterior to cervical groove, median element not always clearly delineated; posteromedian and
posterolateral plates weakly calcified, sulci cardiobranchialis and cardiac sulci not reaching to posterior carapace
margin. Dorsal surface of shield and lateral processes marked by numerous short transverse low ridges fringed with
setae (not illustrated). Rostrum triangular with short dorsal keel distally, separated from subacute, moderately
short, lateral projections by straight or slightly concave anterior margin. Ocular peduncles short, stout. Antennular
peduncles overreaching distal margins of corneas by 0.2-0.5 length of penultimate segment. Antennal peduncles
overreaching distal margins of corneas by 0.25-0.65 length of ultimate segment.
Ischia of third maxillipeds each with prominent ovate patch of dense setae dorsolaterally. Thoracic sternites
(Fig. 3b) with scattered very small protuberances or granules Sternite of third maxillipeds with u-shaped median
cleft, flanked on both sides by pair of widely-separated small spines. Sternite of chelipeds with left side markedly
larger, partially fused with subrectangular sternite of second pereopods. Sternite of third pereopods incompletely
divided into anterior and posterior lobes. Sternite of fourth pereopods hidden from view beneath sternite of third.
Sternite of fifth pereopods slender, rod-like, widely separated from fourth.
398
P.A. MCLAUGHLIN
Chelipeds very unequal, right much stronger, although not appreciably longer. Dactyl of right cheliped with
row of small granules or denticles in proximal 0.65 of dorsomesial margin; dorsal surface rounded, granular or
minutely tuberculate and with sparse tufts of moderate to long setae, with short subacute ridge proximally. Palm
with convex dorsal surface minutely granular, particularly in mesial half and with numerous sparse tufts of
moderately long setae, dorsomesial margin with low ridge of tiny tubercles or simply crenulations, dorsolateral
margin with distinct low ridge of crenulations or granules, becoming somewhat more prominent on fixed finger.
Carpus somewhat roundly elevated in dorsal midlinc with several short transverse piliferous ridges, dorsomesial and
dorsolateral margins and mesial and lateral faces each with transverse piliferous crenulated low ridges. Merus with
row of moderately small spines on ventromesial and ventrolateral margins, ventral surface with very dense patch
setae in mesial half.
Left cheliped with surfaces
of dactyl unarmed but with
sparse tufts of setae adjacent to
cutting edge and ventrally.
Palm with minutely granular
dorsal surface; dorsolateral
margin only weakly delimited
by very low granular ridge, not
extending onto fixed finger;
dorsomesial margin with
granular ridge. Carpus with
several transverse, weakly
raised piliferous ridges;
dorsomesial and dorsolateral
margins each with low,
transverse, somewhat
crenulated ridges and sparse
tufts of setae. Merus with row
of small spines in proximal
0.65 of ventromesial margin;
ventrolateral margin with row
of smaller, more widely-spaced
spines.
Ambulatory legs generally
similar. Dorsal margins of
dactyls each with sparse setae
(second) or transverse rows of
moderately long setae (third);
ventral margins each with row
of 8-13 corneous spines.
Propodi with short somewhat
denticulate ridges forming
irregular longitudinal rows on
dorsal surface and lateral and mesial faces dorsally, ventral margin with single or double row of long spiniform
setae. Carpi each with dorsodistal spine, rows of denticulate protuberances on dorsal faces; lateral faces each with
protuberances forming distinct ridge near mid-width and with few lower denticulate protuberances ventrally. Meri
each with row of very small spines on ventrolateral margins; ventromesial margins minutely granular.
Males with coxae of fifth pereopods roundly subquadrate, gonopores each encircled by tuft of long setae. Tergite
of sixth abdominal somite (Fig. 3c) divided into anterior and posterior halves by prominent transverse furrow, with
small area of calcification laterally. Anterior and posterior portions each divided by longitudinal median furrow.
Fig. 3. — Porcellanopagurus filhoti de Saint Laurent & McLaughlin, 2000. a, c.
damaged <3 (8.2 mm) from MUSORSTOM 4, Stn 201; b, 2 (3.1 mm) from "Mering
Nattde" Stn 43: a, cephalothorax and cephalic appendages (aesthetascs omitted);
b, thorax (ventral view); c, tergite of sixth abdominal somite and telson.
Scales = 1 mm.
Source : MNHN, Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOUTARIOPAGURUS
399
Felson (Fig. 3c ) with small lateral indentations; posterior lobes separated by very minute median cleft, terminal
margins rounded; anus opening very slightly anterior to terminal telson margins.
COLOR. — Ambulatory legs with broad red bands on cream background; carapace, rostral apex and ocular
peduncles with red patches (after Kensley, 1977).
Habitat. — Commonly found utilizing halves of bivalve shells, and occasionally limpets. Animals have
been found with the telson and endopods of the uropods securely lodged in the umbos of the bivalves.
Distribution. — Eastern South Africa, eastern Australia, New Zealand from north of Three Kings Islands
south to off the Canterbury Bight, and east to the Chatham Islands, New Caledonia; 64-1392 m, possibly as deep
as 1430 m.
Remarks. — As pointed out by De Saint Laurent & McLaughlin (2000), Porcellanopagurus filholi
has, for many years, been confounded with P. edwardsi. Porcellanopagurus fillioli is most easily disitnguished
from P . edwardsi by the bilobed first lateral carapace process, and shorter, stouter ambulatory legs of the latter
species.
Porcellanopagurus sp. of KENSLEY (1977), a female with regenerating chelipeds has been reexamined and can
be assigned to P. filholi. Although KENSLEY cited the specimen as a male in his material examined, in his station
data it is listed as a female, and his discussion and illustration correctly indicate female. This specimen, from north
of Richards Bay, KwaZulu-Natal, South Africa, extends the range of this species.
Porcellanopagurus tridentatus Whitelegge, 1900
Fig. 4
P o reel lano-pag urns tridentatus Whitelegge, 1900: 181, figs 13- 13b.
Porcellanopagurus tridentatus -Takeda, 1985: 141. — Turkay, 1986: 140. — Suzuki & Takeda. 1987: 17. —
Zarenkov, 1990: 239. — De Saint Laurent & McLaughlin, 2000: 106, fig. 33.
Not Porcellanopagurus tridentatus - Chilton, 1911: 352 (= Porcellanopagurus chihoni de Saint Laurent & McLaughlin
2000).
Material EXAMINED. — New Caledonia. BlOCAL: stn DW 64, 24°48'S. I68°09'E, 250 m, 3.09.1985: 2 8 (2.7,
3.7 mm). 1 ov, $ (3.2 mm) (MNHN-Pg 5864).
Musorstom 4: stn DW 230, 22°52.5'S, 1 67° 1 1 ,8'E, 390-420 m, 30.09.1985: 1 2 (3.4 mm), I ov. 2 (5.3 mm)
(MNHN-Pg 5865).
Smib 2: stn DW 3, 22°56.0'S. 167°14.8'E, 428 m, 17.09.1986: 1 <3 (5.8 mm) (NHM 1999.2169).
Chalcal 2: stn CP 18, 24°47.0'S, 168°09.4'E. 274 m, 20.10.1986: 1 8 (4.7 mm), 1 ov. 2 (4.3 mm) (USNM
276173). — Stn DW 69, 24°43.7'S. 168°07.9'E, 260 m, 27.10.1986: 1 2 (3.8 mm) (MNHN-Pg 5866). — Stn DW 71.
22°42.3'S, 1 68°09.5'E, 230 m, 27.10.1986: 1 o (4.2 mm) (MNHN-Pg 5867).
Bathus 2: stn DW 729, 22°52'S, 167°H'E. 400 m. 12.05.1993: 1 2 (5.3 mm) (MNHN-Pg 5868). — Stn DW 730,
23°02'S, 166°58'E, 397-400 m, 12.05.1993: I 2 (3.0 mm) (MNHN-Pg 5869).
Chesterfield Islands. Musorstom 5: stn DC 361, 19°52.5'Sr 158°38. I E, 400 m. 19.10.1986: 1 8 (6.8 mm)
(MNHN-Pg 5870). — Stn DC 379, 19°53.2'S, 158°39.5'E. 370-400 m, 20.10.1986: 1 ov. 2 (6.7 mm) (MNHN-Pg 5871).
Diagnosis. — Anterior carapace (Fig. 4a) broad, smooth, with few scattered tufts of short setae: lateral
margins each with slender acute spine on triangular or rounded first lateral lobe, slightly smaller spine distally and
often 3 or 4 spinules laterally on short subtriangular prominence of second lateral lobe, and moderately strong
subtriangular, spinulose projection posterior to cervical groove, median element very narrow; posteromedian plate
partially calcified, posterolateral plates distinct, weakly calcified, sulci cardiobranchialis and cardiac sulci extending
nearly to posterior carapace margin. Rostrum very broadly triangular and usually weakly irilobed, reaching nearly
to mid-length of ocular peduncles, broad basally, narrowing abruptly in distal 0.24-0.35, terminating subacutely.
Lateral projections prominent, acute, forwardly or slightly anterolaterally directed. Ocular peduncles moderately
short and stout; corneas slightly dilated. Antennular peduncles when fully extended overreaching distal margins of
corneas by approximately 0.25 length of penultimate segment. Antennal peduncles overreaching distal margins of
corneas by 0.5-0.65 length of fifth segment.
400
P.A. MCLAUGHLIN
Ischium of third maxilliped
without distinctive tuft of
setae dorsolaterally. Thoracic
sterniles (Fig. 4b) with nu¬
merous scattered tubercles.
Sternite of third maxillipeds
with acute spine on either side
of broad, u-shapcd median
cleft. Sternite of chelipeds
largest on left side; incom¬
pletely fused to subquadrate
sternite of second pereopods.
Sternite of third pereopods
with elongate subrectangular
anterior lobe and shorter
subrectangular posterior lobe.
Sternite of fourth pereopods
very slender, often obscured
from view beneath third.
Sternite of fifth pereopods
slender, rod-shaped, well sepa¬
rated from fourth.
Right cheliped stout,
considerably stronger, but not
appreciably longer than left.
Dactyl articulating somewhat
obliquely with palm; dorsal
and mesial surfaces rounded.
Fig. 4. — Porcellanopagurus tridentatus Whitelegge, 1900. a-c, 6 (3.7 mm) from granular or minutely tubercu-
Biocal Sin DW 64; a, cephalothorax and cephalic appendages (aesthetascs ]ate Dorsomesial margin of
omitted); b. thorax (ventral view); c, tergite of sixth abdominal somite and pa|m wjth low tuberculate or
telson. Scales = 1 mm. granular ridge, dorsal surface
granular or minutely tuberculate and with few scattered setae, dorsolateral margin with very faint granular ridge.
Carpus trapezoidal, dorsomesial and dorsolateral margins and mesial and lateral faces dorsally with series ol low.
transverse protuberant ridges, strongest mesially. Merus without distinctive tuft of setae on ventromesial margin;
ventrolateral and ventromesial margins each with row of small spines, strongest laterally.
Left cheliped with slight granular ridge on proximal half of dorsomesial margin ol dactyl, dorsal surlace with
few very small tubercles in midline proximally. Palm with granular or minutely spinulose dorsal suitace.
dorsolateral margin with faint ridge. Carpus with mesial and lateral faces marked by low transverse ridges, lorming
longitudinal row on both dorsomesial and dorsolateral margins, strongest dorsomesially; dorsal surface with low,
short transverse, sometimes granular or spinulose ridges. Merus with spinose ventromesial margin.
Ambulatory legs generally similar. Dactyls slightly shorter to slightly longer than propodi; each with
9-12 corneous spines on ventral margin. Propodi with low, sometimes spinulose protuberances on dorsal surlaces,
lateral faces with short or very short transverse protuberances or ridges; ventral margins each with row of 5 or
6 spinose setae, longest distally. Carpi each with 1 or 2 irregular rows ol blunt or spinulose protuberances on
dorsal surfaces, dorsodistal spine sometimes slightly better developed; lateral faces each with prominent denticulate
ridge in dorsal half and smaller ridges or low protuberances ventral. Meri each with row of spinules on
ventromesial and ventrolateral margins, strongest laterally.
Coxae of fifth pereopods in males drawn out posteromedially and fringed with setae, but no membranous sexual
tubes developed. Sixth abdominal somite (Fig. 4c) with prominent transverse suture at mid-length; anterior and
Source : MNHN, Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOLITARIOPAGURUS
401
posterior lobes each incompletely divided by longitudinal grooves; marginal areas of anterior lobes and median
portions of posterior lobes membranous; terminal margin entire. Telson (Fig. 4c) membranous; with slight lateral
indentations; posterior lobes indicated by very slight median cleft; terminal margins unarmed; anus opening just
anterior to terminal margins.
Color. — Unknown.
Habitat. — Although most New Caledonian specimens were without any protective covering, three
specimens carried limpet shells, while another utilized a bivalve shell.
DISTRIBUTION. — Eastern Australia, Norfolk and Kermadec Islands; New Caledonia; 99-250 m in Australian
and New Zealand waters, 250-428 m in New Caledonia.
Remarks. — De Saint Laurent & McLaughlin (2000) discussed at length the inaccuracies in
WniTELEGGE's (1900) original description as well as Chilton’s (1911) misidentification of material as
P. tridentatus. De Saint Laurent & McLaughlin (2000) described Chilton’s specimens as the new species,
P. chiltoni de Saint Laurent & McLaughlin. The New Caledonia specimens differ from the lectotype of
P. tridentatus , and from the other specimens examined by De Saint Laurent & McLaughlin in having longer
and slightly thinner ocular peduncles, and slightly longer antennular peduncles. One specimen showed only a hint
of the somewhat trilobed rostrum characteristic of this species.
Porcellanopagurus chiltoni de Saint Laurent & McLaughlin. 2000
Fig. 5
Porcellanopagurus tridentatus - CHILTON, 191 1: 352 (not Porcellanopagurus tridentatus Whitclegge. 1900).
Porcellanopagurus chiltoni de Saint Laurent & McLaughlin, 2000: 107, fig. 34.
Material EXAMINED. — New Caledonia. Volsmar: stn DW 48, 21°00.1'S. 170°03'E, 200 m, 4.07.1989: 1 <?
(2.9 mm) (MNHN-Pg 5872).
Smib 5: stn DW 99, 23°24.7'S. 168°05.4'E. 58 m, 14.09.1989: 1 <3 (2.1 mm) (MNHN-Pg 5873).
DIAGNOSIS. — Cephalothorax (Fig. 5a) usually with denticulate scale-like protuberances, particularly on
rostrum and laterally and anteriorly on shield. First lateral carapace lobe weakly produced, with small acute,
subacute or blunt spine; second lateral carapace lobe not produced beyond lateral level of first, subtriangular,
marginally denticulate; slightly stronger subtriangular, subacute projection posterior to cervical groove, median
element very narrow; posteromedian plate not delineated, posterolateral plates weakly calcified, sulci
cardiobranchialis and cardiac sulci reaching nearly to posterior carapace margin. Rostrum very broadly triangular,
weakly, subacutely or acutely trilobed. reaching beyond mid-length of ocular peduncles. Lateral projections
moderately short, subtriangular, anterolaterally directed, terminally acute or blunt. Ocular peduncles short, stout;
with weak constriction adjacent to corneas. Antennular peduncles when fully extended overreach distal margins of
corneas by slightly less to slightly more than full length of ultimate segment. Antennal peduncles overreaching
distal margins of corneas by 0.25-0.75 length of ultimate segment.
Ischia of third maxillipeds usually without distinctive tuft of setae dorsolaterally. Thoracic sternites (Fig. 5b)
slightly granular. Sternite of third maxillipeds bilobed, with anterior margins straight; small spinule on either side
of v-shaped median cleft. Sternite of chelipeds with left side larger, incompletely fused to roundly subquadrate
sternite of second pereopods. Sternite of third pereopods incompletely divided into anterior and posterior lobes;
anterior lobe more elongate, posterior lobe with median indentation on posterior margin. Sternite of fourth
pereopods very narrow. Sternite of Fifth pereopods rod-shaped; considerably distant from fourth.
Right cheliped stout, considerably stronger than left. Dactyl with dorsal and mesial surfaces rounded, granular
or minutely tuberculate, dorsomesial margin sometimes defined by faint ridge. Palm with dorsomesial margin
marked by low tubercles or granules forming slightly elevated ridge, dorsal surface granular or tuberculate,
sometimes with few scattered setae, dorsolateral margin with crenulate, granular, or minutely tuberculate ridge.
402
P.A. MCLAUGHLIN
Dorsomesial and dorsolateral
margins of carpus and mesial
and lateral faces dorsally with
series of low transverse scale¬
like, granular or spinulose
protuberant ridges, strongest
mesially. Merus with few tiny
spinulcs or tubercles on
dorsodistal margin; without
distinctive tuft of setae on
ventromesial margin; ventro¬
lateral and ventromesial margins
each with row of subacute
spines.
Left cheliped with row of
very small tubercles on
dorsomesial margin of dactyl;
dorsal midline with short lon¬
gitudinal row of tiny tubercles.
Palm with covering of low
tubercles or granules on dorsal
surface not extending to distal
half of fixed finger; dorsomesial
and dorsolateral margins each
with low spinulose or crenulate
ridge. Carpus with transverse
ridges forming longitudinal row
on dorsomesial margin, dorsal
surface w'ith numerous scale-like
tuberculate or spinulose, pro¬
tuberant short ridges; mesial and
lateral faces with low transverse
ridges. Merus with row of subacute tubercles on ventromesial margin; ventrolateral margin with irregular double
row of blunt or subacute spines or tubercles.
Ambulatory legs generally similar. Dactyls 0.75-0.85 length of propodi; dorsal surfaces with low protuberances
and sparse setae; ventral margins each with 8-11 corneous spines. Propodi with numerous low, sometimes
spinulose, protuberances on dorsal surfaces; ventral margins each with row of 5 or 6 long spiniform setae, and
usually additional pair at ventrodistal margin. Carpi each with 1 or 2 irregular rows of spinulose protuberances on
dorsal surface, no distinct dorsodistal spine; lateral faces each with prominent tuberculate or spinulose ridge in
dorsal half and row of smaller ridges or low protuberances ventrally. Meri with ventromesial margins unarmed or
granular; ventrolateral margins with 1 or 2 rows of spines or spinules.
Coxae of fifth pereopods in males slightly drawn out posteromedially and fringed with setae, but no sexual
tubes developed. Sixth abdominal somite (Fig. 5c) with prominent transverse suture at or near mid-length; anterior
and posterior lobes each incompletely divided by longitudinal groove; posterior lobes with subtriangular or ovate
terminal margins and adjacent, sometimes well-calcified accessory plates. Telson (Fig. 5c) longer than broad, often
membranous, with slight transverse indentation; rounded terminal margins with very slight median indentation;
anus opening ventral to, but very near, terminal margin.
COLOR. — Unknown.
FlG. 5. — Porcellanopagurus chiltoni de Saint Laurent & McLaughlin, 2000. a-
c, 6 (2.1 mm) from Smib 5 Stn DW 99: a, cephalothorax and cephalic
appendages (aesthetascs omitted); b. thorax (ventral view); c. tergite of sixth
abdominal somite and telson. Scale = 1 mm (a-b) and 0.5 mm (c).
Source : MNHN, Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOL1TARIOPAGURUS
403
Habitat. — Reportedly living under stones between the tidemarks in Kermadec Islands, and carrying half of a
bivalve shell "or a vacant Siphonaria or limpet shell" (Chilton, 1911).
Distribution. — Kermadec Islands, New Zealand, intertidal; New Caledonia, 58-200 m.
Remarks. — Porcellanopagurus chiltoni was described by De Saint Laurent & McLaughlin (2000) from
three specimens collected intertidally from the Kermadec Islands, New Zealand. As noted by these authors, this
species bears considerable resemblance to P. tridentatus, but is distinguished by its longer antennular peduncles,
shorter and stouter ocular peduncles, and most particularly by the distinct ovate appearance of the posterior lobes
of the sixth abdominal tergite. Both species are reported for the Kermadec region, but with a distinct bathymetric
separation. Although in New Caledonia Porcellanopagurus chiltoni occurs at considerably greater depths, i.e. 58-
200 m. P. tridentatus similarly occupies a greater depth zone (250-428 m). Thus the bathymetric separation of the
species appears to be maintained.
SOLITARIOPAGURUS Tiirkay, 1986
Solitariopagurus Tiirkay, 1986: 139. — Poupin & McLaughlin, 1996: 212. — McLaughlin, 1997: 461.
TYPE Species. — By original designation: Solitariopagurus profundus Tiirkay, 1986.
DIAGNOSIS. — Ten pairs of biserial phyllobranchiate gills. Anterior carapace vaulted and strongly calcified;
lateral margins of shield developed into 3 blunt or spiniform lobes, posterior carapace lobe consisting of elongate
median and small lateral elements; posterolateral and posteromedian plates usually well calcified; sulci
cardiohranchialis and cardiac sulci usually not readily distinguishable. Rostrum and lateral projections widely
separated. Ocular acicles reduced, simple; hidden from dorsal view. Maxillule with external lobe of endopod
obsolete or absent. Third maxilliped with well developed crista dentata and 1 accessory tooth.
Right cheliped much stronger, but not appreciably longer than left. Ambulatory legs similar. Fourth and fifth
pereopods subchelate.
Males with stout, moderately long, unequal sexual tubes developed as extensions of coxae of both fifth
pereopods, right longest; each with long setae subterminally and terminally; no paired or unpaired pleopods.
Females with single gonopore posteriorly on coxa of left third pereopod; no paired pleopods; unpaired left
uniramous pleopods on abdominal somites 2-4.
Abdomen reduced; tergal plate of first somite chitinous or very faintly calcified; tergal plate of second weakly
delineated; tergal plates of somites 3-5 clearly defined, chitinous or very weakly calcified; tergite of sixth somite
weakly calcified, subdivided into narrow anterior transverse rod, and posterior pair of broad plates incompletely
divided by median groove. Uropods symmetrical; protopods each with very prominent, posteriorly directed spine;
exopods each with large, circular rasp of corneous scales; endopods each with ovate rasp of corneous scales. Telson
with transverse suture or at least with distinct to weakly indicated mid-lateral indentation; terminal margin entire.
Anus terminal or nearly so.
Key to the species of Solitariopagurus
1 . Dorsal surface of carapace with transverse row of 4 distinct tubercles or spines proximal to
anterior margin . 2
— Dorsal surface of carapace without transverse row of 4 distinct tubercles or spines proximal
to anterior margin . S. profundus
2. Rostrum triangular, terminating acutely or subacutely . 3
— Rostrum broad, "trowel-shaped" . S. trullirostris sp. nov.
404
P.A. MCLAUGHLIN
3. Antennular peduncles overreaching ocular peduncles by 0.35-0.75 length of penultimate
segment. Carpi of chelipeds, with slightly elevated and weakly armed longitudinal median
crests on dorsal surfaces . S. triprobolus
— Antennular peduncles overreaching ocular peduncles by length of ultimate segment or only
slightly more. Carpi of chelipeds with row of spines or tubercles on prominent
longitudinal crests on dorsal surfaces . S. tuerkayi
Solitariopagurus trullirostris sp. nov.
Fig. 6
MATERIAL EXAMINED. — New Caledonia. Mbere Reef, 22°19.9’S, !66°13.2'E. 30 m. 7.01.1993: 1 3
(2.1 mm), 1 ov. $ (3.3 mm) (MNHN-Pg 5874).
Marquesas Islands. Musorstom 9: stn DR 1259, 09°25.6'S, 140°08.3'W. 90-180 m, 3.09.1997: 1 immature 2
(1.6 mm) (MNHN-Pg 5875).
Types. — The ovigerous female from Mbere Reef is the holotype; the male from the same locality is the
allotype.
DESCRIPTION. — Shield (Fig. 6a) length shorter than breadth; anterior margin between rostrum and lateral
projections straight or only very slightly concave; dorsal surface well calcified, with transverse row of 4 broad
tubercles proximal to anterior margin; lateral margins each with relatively short, but prominent spiniform first
lobe adjacent to anterolateral angle, broad, sometimes weakly bilobed second lobe at mid-length, and moderately
slender subacute, somewhat hooked lobe adjacent to cervical groove; posterior margin roundly truncate. Posterior
carapace with median element of posterior lobe moderately narrow, lateral elements subrectangular, unarmed; small
posteromedian and posterolateral plates well calcified. Rostrum broad, roundly subacute, usually reaching to
proximal 0.25 of ocular peduncles; slightly upturned, with elevated lateral margins. Lateral projections narrowly
triangular; not as long as rostrum, directed anterolaterally.
Ocular peduncles laterally directed, slightly less than half shield length, with 3 prominent protuberances (not
developed in immature specimen); corneas slightly dilated, diameter approximately 0.25 length of peduncle. Ocular
acicles small, acutely triangular.
Antennular peduncles when fully extended exceeding length of ocular peduncles by almost entire length of
ultimate segment. Ultimate segment with 2 or 3 long plumose setae on dorsodistal margin. Penultimate segment
with few scattered short setae. Basal segment with tiny spinule on distolateral margin. Epistomial plate well
calcified, broad, with pair of large subacute projections.
Antennal peduncles slightly shorter than ocular peduncles, and 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 in small spine or spinule; dorsomesial distal angle rounded. First
segment with row of tubercles on ventrolateral margin. Antennal acicle moderately short, reaching nearly to or
slightly beyond distal margin of fourth peduncular segment, terminating in tiny spinule, with 1 or 2 minute
spinules on lateral margin. Antennal flagellum long, overreaching outstretched chelipeds, 1 or 2 short setae every 2
to 6 articles.
Thoracic sternites (Fig. 6b) smooth. Sternite of third maxillipeds produced into acute spinose process on either
side of midline. Sternite of chelipeds moderately broad, subtriangular, surface concave, with blunt projection on
either side of deep median groove. Sternite of second pereopods broad, plate-like, with faint median longitudinal
groove. Sternite of third pereopods with narrowly subrectangular anterior lobe.
Right cheliped (Fig. 6c) elongate; considerably stronger but not appreciably longer than left; propodal-carpal
articulation perpendicular. Dactyl slightly shorter than palm; 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 or tuberculate; ventral surface smooth or
Source : MNHN. Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOLITARIOPAGURUS
405
Fig. 6. — Solitariopagurus irullirostris sp. nov. a-i. holotype 6 (2.1 mm) from Mbere Reef: a, eephalothorax and
cephalic appendages (aesthetascs omitted); b. thorax (ventral view); c, carpus and chela of right cheliped (dorsolateral
view); d. carpus and chela of left cheliped (dorsal view); e, right second pereopod (lateral view); f. right third
percopod (lateral view); g. propodus and dactyl of left fourth pereopod (lateral view); h. coxae and sternite of fifth
pereopods; i, tergite of sixth abdominal somite and telson. Scales equal 1 mm (a-h). and 0.5 mm (i).
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 with 1 large and
several smaller calcareous teeth; ventral surface of fixed finger with obliquely longitudinal spinulose or tuberculate
ridge extending onto palm as row of distinct small spines; mesial face minutely spinulose. Carpus equal to or
slightly longer than merus; trapezoidal (in dorsal view), with spinulose or tuberculate dorsomesial and dorsolateral
margins slightly elevated; dorsal surface with irregular double median row of spinulose tubercles or blunt spines.
406
p.a. McLaughlin
also with faint longitudinal ridge or crest, most distinct distally; mesial and lateral faces minutely granular or
spinulose; ventral surface with numerous small spines, ventromesial and ventrolateral margins each with row of
very small tubercles or blunted spines. Merus subtriangular; dorsal margin spinose or spinulose; lateral and mesial
faces spinulose or granular; ventrolateral margin with row of small tooth-like spines, strongest proximally:
ventromesial margin with row of small subacute spines, 1 prominent spine on ventral surface near mesioproximal
margin. Ischium with row of small tubercles or spinulcs on ventromesial margin.
Left cheliped (Fig. 6d) with chela somewhat dorsoventrally compressed; dactyl and fixed finger curved ventrally.
Dactyl 1.25 to 1.75 length of palm; cutting edge with row of very small calcareous teeth; terminating in corneous
claw and very slightly overlapped by fixed finger; dorsal surface unarmed; dorsomesial margin serrate; mesial face
and ventral surface each with few scattered moderately long setae. Palm approximately 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 longitudinal row of small tubercles, ventral surface of fixed
finger with longitudinal oblique row of minute spinulose tubercles; cutting edge of fixed finger with row of small
calcareous teeth; terminating in corneous claw. Carpus approximately equal to length of merus; dorsal surface
trapezoidal (dorsal view); dorsomesial and dorsolateral margins slightly elevated and minutely serrate, midline with
single or irregularly double row of subacute tubercles; mesial and lateral surfaces smooth or minutely granular,
ventromesial, ventrolateral and ventrodistal margins each with row of very small spines or spinules. Merus
subtriangular; dorsal surface spinose or spinulose, mesial and lateral faces spinulose or tuberculate particularly near
ventral margins; ventromesial margin with row of small tubercles; ventrolateral margin with row of spines, ventral
surface with prominent spine at mesioproximal angle. Ischium with row of small tubercles or spines on
ventromesial margin.
Ambulatory legs (Figs 6e-f) moderately long and slender; generally similar. Dactyls approximately equal to
length of propodi; laterally compressed; dorsal margins with few scattered short setae; ventral margins each with
row of 10 or 11 corneous spines. Propodi 1 .25 to 1 .5 length of carpi; dorsal margins each with few widely-spaced
low protuberances; mesial and lateral faces unarmed; ventral margins each with 2 or 3 widely-spaced spiniform
setae and 1 or 2 spiniform setae at distal angle. Carpi less than half length of meri; dorsal margins minutely
granular, no distinct spine at distal angle. Meri with few low protuberances on dorsal margins, strongest on third
pereopods; ventral surfaces oblique, ventromesial and ventrolateral margins microscopically tuberculate (not
apparent in figures). Fourth pereopods strongly subchelate; propodal rasp (Fig. 6g) consisting of single row of
spiniform corneous scales. Fifth pereopods weakly subchelate; dactyl and propodus each with small rasp of
corneous scales dorsally.
Males with markedly unequal sexual tubes on coxae of fifth pereopods (Fig. 6h); each with terminal long setae.
Females with uniramous, unpaired left pleopods on somites 2 to 4.
Abdomen with segmentation clearly delineated dorsally. Tergitcs 1-5 chitinous. 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 few small spinules; exopods
subcircular, each with large circular rasp of corneous scales; endopods appreciably smaller, roundly subrectangular,
each with small rasp. Telson (Fig. 6i) with anterior lobes appearing to becoming more elongate with increasing
size; complete, but very faint transverse suture separating membranous anterior and posterior lobes: terminal
margin entire, with rounded external angles; anus terminal.
COLOR. — Unknown.
Habitat. — Utilizing half of bivalve shell.
DlSIRlBUTlON. — Presently known from Mbere Reef in New Caledonia, and one locality in the Marquesas;
30-90, possibly as deep as 180 m.
ETYMOLOGY. The specific name is taken from the Latin India, meaning trowel, and rostrum, and denotes
the broad, trowel-like rostrum of this species.
Remarks. — Solitariopagurus trullirostris sp. nov. shares with S. triprobolus and S. tuerkayi the distinctive
character of four prominent tubercles on the anterior portion of the shield, an attribute that sets all three species
Source : MNHN, Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOLITARIOPAGURUS
407
apart from S. profundus. However, 5. trullirostris is immediately distinguished from all three other species by its
broad, trowel-like rostrum.
Solitariopagurus triprobolus Poupin & McLaughlin. 1996
Fig. 7
Solitariopagurus triprobolus Poupin & McLaughlin, 1996: 212, figs 1-2. — McLaughlin. 1997: 464.
Material EXAMINED. — New Caledonia. BERYX 11: stn CP 23. 24°43.4'S, 168°07.8'E, 270-290 m. 17,10.
1992: 1 ov. 2 (3.8 mm) (MNHN-Pg 5876).
Chesterfield Islands. Musorstom 5: stn CP 287, 24°05.4'S, 159°36.5'E, 270 m, 10.10.1986: 1 3 (5.2 mm)
(MNHN-Pg 5877).
Vanuatu. Musorstom 6: stn CP 455, 21°01'S, 167°26. 1 E. 260 m, 20.02.1989: 1 2 (5.7 mm) (MNHN-Pg 5878).
DIAGNOSIS. — Shield (Fig. 7a) shorter than broad, well calcified; dorsal surface with transverse row of 4
prominent, blunt or subacute spines proximal to anterior margin, sometimes 1 smaller spine anteriorly in midline
near base of rostrum,
and frequently additional
small tubercles ante¬
riorly and/or laterally;
lateral margins each
with prominent spinose
first lobe at anterolateral
angle, slightly shorter,
simple or faintly bi-
lobed, weakly serrate, or
denticulate spinose sec¬
ond at mid-length and
strong, often elongate
and slender, usually
anteriorly directed,
spinose third lobe ad¬
jacent to cervical
groove. Posterior cara¬
pace with median
element of posterior
lobe moderately broad,
elongate, lateral ele¬
ments, each armed with
spine and/or projecting
laterally as short,
spinose process; pos¬
terolateral and postero¬
median plates small,
weakly to moderately
well calcified. Rostrum
triangular, elongate,
usually reaching to distal half of ocular peduncles. Lateral projections extremely well developed, often reaching to
or beyond tip of rostrum, directed anteriorly. Ocular peduncles moderately short. Antennular peduncles overreaching
distal margins of corneas by at least 0.35 length of penultimate segment. Antennal peduncles overreaching distal
margins of corneas by at least 0.25 length of ultimate segment. Sternite of chelipeds subtriangular, with terminal
notch or indentation medially.
Fig. 7. — Solitariopagurus triprobolus Poupin & McLaughlin, 1996. a-c, 3 (5.2 mm) from
Musorstom 5 Stn CP 287: a, cephalothorax and cephalic appendages (aesthetascs
omitted); b, coxae and sternite of fifth pereopods; c, tergite of sixth abdominal somite
and telson. Scales = 1 mm.
Source : MNHN. Paris
408
P.A. MCLAUGHLIN
Right cheliped stronger, but not appreciably longer than left. Dorsal surfaces of palm, fixed finger and dactyl,
smooth or minutely granular. Carpus trapezoidal (in dorsal view), dorsomesial and dorsolateral distal angles
frequently produced as wing-like projections; dorsal surface often with scattered small tubercles or spinules, also
with median or submedian longitudinal row of tubercles, strongest in proximal half.
Left cheliped with dorsal surfaces of palm, fixed finger and dactyl smooth or minutely granular. Carpus
frequently with scattered tubercles on dorsal surface; dorsomesial and dorsolateral distal angles often produced as
wing-like projections, with raised, serrate dorsomesial and dorsolateral margins, midline with longitudinal row of
simple or multidenticulate, spinulose tubercles or small spines, at least proximally.
Ambulatory legs long and slender; dactyls usually distinctly shorter than propodi, each with row of 9-17
corneous spines on ventral margin. Propodi with minutely granular or spinulose dorsal and ventral surfaces; ventral
margins with 3 widely-spaced spiniform setae. Carpi with minutely granular or spinulose dorsal surfaces; lateral
faces each with median longitudinal ridge and often longitudinal band of minute spinules. Meri with minutely
granular of spinulose dorsal and ventral margins. Fourth pereopods each with row of 6-8 club-shaped corneous
scales on ventral margin of propodus.
Male sexual tubes (Fig. 7b) weakly calcified. Telson (Fig. 7c) with anterior portion moderate to elongate;
transverse suture weakly delineated; terminal margin entire, rounded; anus terminal or nearly so.
COLOR. — Overall generally orange-brown. Ocular peduncles with mesial and lateral longitudinal red stripe.
Antennular peduncles with longitudinal red stripe dorsally. Ambulatory legs with three continuous or interrupted
red stripes on lateral surfaces of meri; carpi often with one median longitudinal red stripe on lateral face and one on
dorsal surface (Poupin & McLaughlin, 1996).
Habitat. — Usually utilizing halves of bivalve shells, occasionally univalve shells.
Distribution. — French Polynesian islands; New Caledonia; 200-380 m.
Remarks. — The small, ovigerous female from Beryx 1 1 station CP 23 resembles the male paratype figured
by POUPIN & McLaughlin (1996, fig. 2A) in lacking accessory tubercles on the carapace and in having shorter
lateral projections and lateral carapace spines. Additionally, the ocular peduncles are shorter and stouter in this
specimen than seen in most representatives of the species; however, this development of the ocular peduncles is
not an uncommon characteristic of smaller specimens of many species. In these respects, this specimen
approximates S. luerkayi ; but is distinguished from that species by its longer antennular peduncles, and more
anteriorly directed lateral projections. Although the right cheliped is missing, the armature of the carpus of the left
cheliped agrees with that of the other specimens of S. triprobolus found in New Caledonia.
Solitariopagurus tuerkayi McLaughlin, 1997
Fig. 8
Porcellanopagurus sp. - TUDGE, 1995: 256, fig. 1G.
Solitariopagurus tuerkayi McLaughlin, 1997: 461. fig. 8.
Material EXAMINED. — Okinawa. 26°30'N, 1 27°50.9'E, 61-65 m, collector R.F. Bolland: 1 ov 9 (2 9 mm)
(USNM 276172).
Pg 58e7W9)Caledonia- Bathus 1: stn 684. 20°34'S, 165°07'E, 110-130 m. 16.03.1993, 1 «J (2.1 mm) (MNHN-
Diagnosis. — Shield (Fig. 8a) length shorter than breadth; dorsal surface with transverse row of 4 moderately
low, narrow to broad tubercles proximal to anterior margin; posterolateral region weakly delineated and usually
somewhat globular; lateral margins each with relatively short, but prominent spinose first lobe adjacent to
anterolateral angle, slender or broad, simple or weakly bilobed, acute or subacute spinose second lobe at mid¬
length, and short, broad or moderately slender spinose third lobe adjacent to cervical groove; posterior margin
broadly rounded. Posterior carapace (damaged in New Caledonia specimen) with posterior lobe consisting of
Source : MNHN. Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOUTARIOPAGURUS
409
moderately slender median element and pair of unarmed lateral elements, sometimes projecting as short blunt or
spinose processes; posterolateral and posteromedian plates not delineated or sometimes very weakly calcified.
Rostrum elongate, usually reaching to proximal half of ocular peduncles; broad, usually terminating bluntly.
Lateral projections triangular; moderately elongate, slightly shorter to slightly longer than rostrum, directed
anterolaterally. Ocular peduncles short or moderately short, corneas somewhat dilated. Antennular peduncles when
fully extended, elongate, overreaching ocular peduncles by full length of ultimate segment to 0.25 of penultimate
segment. Antennal peduncles overreaching ocular peduncles by 0.1 to 0.5 length of ultimate segment.
Right cheliped elongate;
considerably stronger than
left. Dactyl with dorsal
surface convex, smooth or
minutely granular, dorso-
mesial margin minutely
serrate; ventral surface smooth
or microscopically granular.
Palm with 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; ventral surface of
fixed finger with obliquely
longitudinal and micro¬
scopically tuberculate ridge
extending onto palm in distal
half. Carpus with spinulose
dorsomesial and dorsolateral
margins slightly elevated;
dorsal surface with scattered
small tubercles or spinules,
also with longitudinal median
ridge or crest, most distinct
distal ly and armed with
spinules or tuberculate spines,
one strongest in proximal
half; mesial and lateral faces
minutely granular or spin¬
ulose; ventromesial and
ventrolateral margins each
with row of very small
tubercles or blunted spines.
Merus with row of small spines on ventrolateral margin strongest proximally; ventromesial margin with row of
small spines distally, 3 or 4 very prominent spines proximally.
Left cheliped with dorsal and ventral surfaces of dactyl unarmed; dorsomesial margin serrate, at least in
proximal half. Palm with 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. Carpus with few scattered very small tubercles or spinules:
dorsomesial and dorsolateral margins raised and serrate, midline with somewhat elevated crest armed with row of
Fig. 8. — Solitariopagurus tuerkayi McLaughlin, 1997. a. 3 (2.1 mm) from
Bathus I Stn 684; b. ov. 2 (2.9 mm) from Okinawa: a. cephalothorax and
cephalic appendages (aeslhetascs omitted): b. tergite of sixth abdominal somite
and telson. Scales = 1 mm (a) and 0.5 mm (b).
Source : MNHN. Paris
410
P.A. MCLAUGHLIN
simple or multidenticulate spinulose tubercles or small spines, strongest proximally, and with I very prominent
tuberculate spine at mid-length; mesial, lateral, and ventral surfaces spinulose or tuberculale. ventrodistal margin
with row of tubercles. Merus with row of small spinules, becoming strong spines proximally on ventromesial
margin; ventrolateral margin minutely spinulose distally becoming row of stronger spines in proximal half.
Ambulatory legs moderately long and slender; generally similar. Dactyls approximately equal to length of
propodi; dorsal margins with few scattered short setae; ventral margins each with row of 9 to 12 corneous spines.
Propodi with serrate dorsal margins; mesial and lateral faces minutely spinulose, particularly ventrally; ventral
margins each with I or 2 corneous spiniform setae at distal angle and 2 additional widely-spaced on ventral margin.
Carpi with 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. Mcri with serrate or spinulose dorsal margins; ventromesial and ventrolateral margins spinulose.
Males with subequal or markedly unequal sexual tubes on coxae of fifth pereopods (missing in New Caledonia
specimen); each with subterminal and terminal long setae. Abdomen (missing in New Caledonia specimen)
reduced, segmentation clearly delineated dorsally. Tergite of sixth somite with moderately well calcified anterior
rod-like and posterior paired rectangular plates. Telson (Fig. 8b) (missing in New Caledonia specimen) elongate;
transverse suture clearly delineated; terminal margin entire, with rounded external angle; anus opening directly
beneath terminal margin.
COLOR (in preservative). — After four years in alcohol only a faint orange tint remains on chelae and dactyls of
ambulatory legs; propodi of ambulatory legs with one or two very faint bands (McLaughlin, 1997).
Habitat. — Unknown.
DISTRIBUTION. — Known from the Kai and Tanimbar Islands, Indonesia, New Caledonia and Okinawa;
61-212 m.
Remarks. — In his study of the ultrastructure of spermatozoa in'lhe Thalassinidea and Anomura, TUIX3E
(1995) utilized a specimen borrowed form the Museum national d'Histoire naturelle, Paris, and incorrectly
identified for him as Porcellanopagunts sp. Now, after personally examining the remaining cephalothorax,
chelipeds and ambulatory legs, it has been possible to assign his specimen to Solitariopagurus tuerkayi. Tudge's
findings should now be referred to Solitariopagurus rather than Porcellanopagunts.
Among the collections of the former Smithsonian Oceanographic Center (SOSC) is an ovigerous female
specimen collected in Okinawa that is also identifiable as S. tuerkayi. This specimen represents the first record of
Solitariopagurus in the northern hemisphere.
DISCUSSION
The matter ot the separation of the telson from the tergite of the sixth abdominal somite and the location of the
anal opening require turther commentary. In the course of crustacean development, somite addition occurs between
the dignathan head and the telson (e.g. Calman,1909; ANDERSON, 1979). The telson is, by that criterion, not
considered a true somite (McLaughlin, 1980). However, Sharov (1966) and Bowman (1971) contended that
two types ol non-homologous telsons occurred in crustaceans, one in which no appendages were present and the
anus was subterminal, and a second in which the anus was terminal and caudal rami were present. These authors
considered that the latter type of telson was actually a terminal (anal) body somite. Although SCHMINKE (1976),
argued rather convincingly that Bowman's (1971) interpretation of the telson was incorrect. Schram (1986)
continued to make the distinction. According to Bowman (1971), while the anus may open distally on the ventral
surface of the telson in decapods, no decapod has a terminal anus. To the contrary, in some species of
Porcellanopagunts and Solitariopagurus the anus is clearly terminal, while nearly so in others. Consequently by
Sharov (1966), Bowman (1971) and Schram's (1986) definition, the telson in at least certain species in these
two genera would be the seventh, and anal, body somite, albeit lacking appendages. While SCHMINKE (1976) did
Source : MNHN. Paris
CRUSTACEA DECAPODA: PORCELLANOPAGURUS AND SOLITARIOPAGURUS
41 1
not address the condition in decapods, he did suggest that the position of the anus was not an important feature for
the definition of the telson. As he noted, a change in the position of the anus is not an uncommon phenomenon
and might even change within the same animal during the course of its development. Roberts (1972) clearly
indicated in his description of the larval stages of Porcellanopagurus edwardsi that the anus was located anteriorly
on the ventral surface of the telson. MCLAUGHLIN (in press) reported that the telson was still in an anteroventral
position in the megalopa of this species, but shifted to a slightly more posterior position in the first crab stage.
Thus it is obvious that a positional change does occur in Porcellanopagurus, and presumably does as well in
Solitariopagurus.
A change in the position of the anus does not, in itself, fully explain its unusual migration to a terminal
location, nor does it offer any exegesis for the apparent separation of the telson from the sixth somite. To submit
an unequivocal explanation would require the examination of a complete series of juvenile crab stages, which
unfortunately are not available. However, from the data now at hand, a possible scenario can be put forth. It would
appear that, contrary to development in most other pagurids [e.g., Pagurus kennerlyi (Stimpson, 1864),
cf. McLaughlin et a!., 1989] there is a marked shortening of the abdomen that accompanies the molt from
megalopa to first crab stage in Porcellanopagurus (cf. McLaughlin, in press) and presumably also
Solitariopagurus species. As a result, the existing abdominal tissues expand laterally, which accounts for the semi-
globular shape of the abdomen. With increased animal size, abdominal growth appears to be somewhat volumetric.
Such growth would explain the ostensible dorsal migration of the female pleopods and perhaps even the posterior
shift of the anus. Although the telson is most commonly described simply in terms of its tergal structure, it is a
body unit just as any thoracic or abdominal somite, but differing from a true somite in the absence of paired
ganglia and appendages (ANDERSON, 1979). In P. adelocercus (cf. MCLAUGHLIN & Hogarth. 1998. fig. 29).
P. haptodactylus sp. nov. (Fig. 2i), and S. trullirostris sp. nov. (Fig. 6i), the tergite of the telson is no longer the
posterior-most dorsal structure. It appears to almost "float" on the upper surface of an extended telsonal unit, with
the anal opening becoming terminal.
An explanation of the separation of the telson and sixth abdominal somite it less straightforward, and may not
be comparable in the two genera. In P. adelocercus and P. haptodactylus sp. nov. there is no clear indication that
the tergite of the sixth somite is the most posterior part of that somite, whereas there is a clear delineation of the
telsonal unit. Thus it appears as if there is additional posterior growth of the sixth somite beyond the calcified
tergal plate in some Porcellanopagurus species. In contrast, it seems as though the additional growth occurs in the
anterior portion of the telsonal unit, including the anterior lobes of the tergite in Solitariopagurus trullirostris
sp. nov. That the telson is carried ventrally is probably associated with these phenomena and the actions of the
uropods in holding the carcinoecia.
ACKNOWLEDGMENTS
I am deeply indebted to Alain CROSNIER and Bertrand RICHER DE FORGES for making the MUSORSTOM
material available for study. Thanks are due Rafael LEMAITRE. National Museum of Natural History. Smithsonian
Institution, for the loan of the SOSC specimen, and Michelle van DER Merwe and Elizabeth Louvv, South
African Museum, for the loan of their Porcellanopagurus specimen. This is a scientific contribution from the
Shannon Point Marine Center, Western Washington University.
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ILTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 - RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 - RESULTATS DES C
Crustacea Decapoda: Species of the genera
Crosnierita Macpherson, 1998, Munida Leach, 1820,
and Paramunida Baba, 1988 (Galatheidae)
collected during the MUSORSTOM 9 cruise
to the Marquesas Islands
Enrique MACPHERSON
Centro de Estudios Avanzados de Blanes (CSIC)
Camf de Santa Barbara s/n
17300 Blanes (Girona), Spain.
ABSTRACT
During the Musorstom 9 eruise (August-September 1997) to the Marquesas Islands, galatheids belonging to the
genera Crosnierita Macpherson, 1998 (one species). Munida Leach. 1820 (4 species, one of which is new) and
Paramunida Baba, 1988 (one new species) were collected. The two new species, Munida glabella and Paramunida echinata,
are described and illustrated. M. glabella is close to M. pusiola Macpherson, 1993 from the Philippines. It belongs to the
group of small size species that have five spines on the lateral borders of the carapace posterior to the cervical groove, no
spines on the second abdominal somite and the distolateral spine of the basal antennal segment shorter than the
distomesial spine. P. echinata is dose to P. labis Macpherson, 1996 from the Wallis and Futuna Islands It is charac¬
terised by the absence of ridges on the thoracic sternites and the presence of a spinous process on the distomesial spine of
the second antennal segment.
RESUME
Crustacea Decapoda : Especes des genres Crosnierita Macpherson, 1998, Munida I.each, 1820,
et Paramunida Baba, 1988 (Galatheidae) recoltees durant la campagne Musorstom 9 aux ties
Marquises.
Lors de la campagne Musorstom 9 (aout-septembre 1997) aux lies Marquises, des galatheides appartenant aux genres
Crosnierita Macpherson, 1998 (1 espece), Munida Leach. 1820 (4 especes dont 1 nouvelle) et Paramunida Baba!^ 1988
(1 espece nouvelle) ont 6t6 recoltees. Les deux especes nouvelles. Munida glabella et Paramunida echinata , sont deerites
Macpherson. E.. 1999. — Crustacea Decapoda: Species of the genera Crosnierita Macpherson. 1998. Munida Leach.
1820. et Paramunida Baba, 1988 (Galatheidae) collected during the Musorstom 9 cruise to the Marquesas Islands.
In. A. Crosnif.r (ed. ). Resultats des Campagnes Musorstom. Volume 21. Memoires du Museum national d'Histoire
naturelle. 184: 415-423. Paris ISBN 2-85653-526-7.
416
E. MACPHERSON
et illustrees. M. glabella est proche de M. pusiola Macpherson, 1993. des Philippines el appartient au groupe d'especes
dc petite taillc ayant cinq Spines sur les bords latSraux de la carapace en arrive du sillon cervical, le deuxieme segment
abdominal sans epines, et 1‘epine distolaterale du segment antennulaire basal plus courte que la distomesiale. P. echinata
est proche de P. labis Macpherson. 1996. des lies Wallis et Futuna; elle se caracterise par les sternites thoraciques sans
stries et P Spine distomesiale du second segment antennaire avec un processus Spineux.
INTRODUCTION
The species of the genus Munida Leach, 1820 and related genera from the Central Pacific are poorly
known, although a few publications have been published in the last decades (Baba, 1981; Zarenkov &
Khodkina, 1981; Titgen, 1987; Macpherson & de Saint Laurent, 1991; Macpherson, 1996; Baba &
de Saint Laurent, 1996; Poupin, 1996). During the MUSORSTOM 9 cruise to the Marquesas Islands (RICHER
DE FORGES etal., 1999) in August-September 1997, some galatheids of the genera Crosnierita Macpherson, 1998,
Munida Leach, 1820 and Paramunida Baba, 1988 were collected. The study of these specimens revealed the
presence of one species of Crosnierita, four species belonging to the genus Munida, including a new species, and
one new species of the genus Paramunida.
The types of the new species and other material have been deposited in the collections of the Museum
national d'Histoire naturelle, Paris (MNHN) and National Museum of Natural History, Washington (USNM). The
samples without any registration are kept at the Museum national d'Histoire naturelle, Paris.
Measurements given are of carapace length, excluding rostrum, and the terminology used mainly follows
that used in previous publications (Zariquiey Alvarez, 1952; Macpherson & de Saint Laurent,
1991; Macpherson, 1994; Baba & de Saint Laurent. 1996). Colour notes are taken from slides by
P. Laboute.
COMBINED LIST OF STATIONS AND OF SPECIES OBTAINED PER STATION
Stn CP 1 146. — 22.08.1997, 09°18.8'S, 140°06.2'W, 200 m: C. yante, P. ecliinata.
Stn CP 1 148. — 22.08.1997, 09°18.9'S, 140°06.3'W, 300 m: M. plexaura, P. echinata.
Stn CP 1 154. — 23.08.1997, 07°58.5'S, 140°43.7'W, 102 m; P. echinata.
Stn CP 1 172. — 25.08.1997, 08°44.8'S, 140°15.3'W, 300-302 m: P. echinata.
Stn CP 1 176. — 25.08.1997, 08°45.8'S, 140°14.5'W, 260 m: P. echinata.
Stn CP 1177. —25.08.1997, 08°45. US, 140°14.1'W, 108-112 m; M. glabella.
Stn CP 1 190. — 26.08.1997, 08°46.3'S, 140°07.2'W, 350 m: P. echinata.
Stn CP 1201. — 28.08.1997, 09°50.6’S, 139°09.2'W. 275-300 m: P. echinata.
Stn CP 1208. — 28.08.1997, 09°48.9'S, 139°09.5'W, 1 17 m; M. glabella.
Stn CP 1237. — 31.08.1997, 09°41.9'S, 139°03.6’W, 95-305 m: C. yante.
Stn CP 1238. — 31.08.1997, 09°41.4'S, 139°03.8'W, 280-370 m: C. yante, P. echinata.
Stn CP 1262. — 03.09.1997, 09°19.8’S, 140°08.3'W, 850-905 m: M. ty’phle.
Stn CP 1268. — 04.09.1997, 07°55.8'S, 140°42.6'W, 285-320 m: M. tuberculata.
Stn CP 1269. — 04.09.1997, 07°56.3'S, 140°43.3'W, 420-430 m: M. tuberculata, P. echinata.
Stn CP 1278. — 05.09.1997, 07o52.1'S, 140°38.6’W, 1000 m: M. typhle.
Sin CP 1282. — 07.09.1997, 07°51.7'S, 140°30.6'W, 416-460 m: C. yante. M. plexaura. M. tuberculata.
Stn CP 1287. — 07.09.1997, 07°54.5'S. 140°40.2'W, 163-245 m: M. plexaura.
Source : MNHN, Pans
CROSNIERITA, MUNIDA AND PARAMUNIDA FROM MARQUESAS ISLANDS
417
SYSTEMATIC ACCOUNT
Genus CROSNIERITA Macpherson, 1998
Crosnierita yante (Macpherson, 1994)
Munida yante Macpherson, 1994: 555, figs 62, 97.
Crosnierita yante - MACPHERSON. 1998: 353.
MATERIAL EXAMINED. — Marquesas Islands. MUSORSTOM 9: stn 1146, 200 m: 1 ov. 9 8.2 mm (MNHN-Ga
4352). — Stn 1237, 95-305 m: 7 6 5.8 to 10.3 mm; 3 ov. 2 8.2 to 8.4 mm; 8 2 5.0 to 6.4 mm (MNHN-Ga 4353). —
Stn 1238, 280-370 m: 1 3 10.3 mm; 3 ov. 2 7.5 to 9.3 mm; 1 2 6.7 mm (USNM). — Stn 1282, 416-460 m: 2 3 7.5
and 9.7 mm; 1 ov. 2 7.5 mm; 1 2 7.4 mm (MNHN-Ga 4354).
REMARKS. — At first, the species was originally known from the female holotype collected in New Caledonia
from 400 m (Macpherson, 1994). A second specimen was collected later during the SMIB 8 cruise (unpublished
data) in 310-460 m depth. The specimens collected in the Marquesas Islands agree well with these two specimens.
DISTRIBUTION. — New Caledonia from 310-460 m. The specimens from Marquesas Islands were collected
in 95-460 m.
Genus MUNIDA Leach, 1 820
Munida glabella sp. nov.
Fig. 1
MATERIAL EXAMINED. — Marquesas Islands. Musorstom 9: stn 1177, 108-112 m: 1 ov. 2 3.6 mm (MNHN-Ga
4355). — Stn 1208, 117 m: 1 3 3.3. mm (MNHN-Ga 4373); 1 ov. 9 3.4 mm (MNHN-Ga 4356).
Types. — The ovigerous female of3.4 mm (MNHN-Ga 4356) from stn 1208 (09°48.9'S, 139°09.5'W. 117m)
has been selected as the holotype, the other specimens are paratypcs.
ETYMOLOGY. — From the Latin diminutive of glaber , smooth, in reference to the smooth surface of the
carapace and the small size of the species.
DESCRIPTION. — Carapace slightly longer than wide. Transverse ridges mostly interrupted, with dense, very
short, non-iridescent setae and few long, iridescent setae. Main transverse striae on posterior part of carapace
interrupted in cardiac region. Few scales and secondary striae between main striae. Scales on intestinal region
absent. Gastric region with 5 pairs of epigastric spines. One parahepatic and one postcervical spine on each side.
Frontal margins transverse. Lateral margins slightly convex. Anterolateral spine short, at anterolateral angle, not
reaching level of sinus between rostrum and supraocular spines. Second marginal spine before cervical groove two
times smaller than preceding one. Branchial margins with five spines. Rostrum spiniform, half as long as
remaining carapace, slightly sinuous and horizontal. Supraocular spines reaching midlength of rostrum and not
overreaching end of corneas, parallel, directed upwards.
Thoracic sternites smooth, without striae, granules or carinae. Anterior part of fourth sternite slightly narrower
than third. Transverse ridges between fifth, sixth and seventh sternites obtuse, feebly granulated.
Second abdominal somite unarmed. Second and third somites each with one transverse stria. Fourth abdominal
somite without stria.
418
E. MACPHERSON
Eyes moderately large, maximum corneal diameter about one-third distance between bases of anterolateral
spines.
Basal segment of antennule (distal spines excluded), about one quarter carapace length, elongate, slightly
overreaching corneae, with two distal spines, mesial spine slightly longer than lateral spine; two spines on lateral
margin, proximal one short, located at midlength of segment, distal one long, overreaching distolateral spine.
D
1 mm
A, B, E, F
0.5 mm
C, D.G
Fig. 1. — Munida glabella sp. nov., ov. 2 3.4 mm, holotype from stn 1208: A. carapace, dorsal view; B, sternal
plastron; C, ventral view of cephalic region, showing antennula and antennal peduncles; D, right third maxilliped.
lateral view; E, left cheliped, dorsal view; F. right first walking leg, lateral view; G, dactylus of right first walking
leg, lateral view.
Source : MNHN, Pans
CROSN1ERITA, MUNIDA AND PARAMUNIDA FROM MARQUESAS ISLANDS
419
First segment of antennal peduncle with one distal spine on mesial margin nearly reaching end of second
segment; second segment with two distal spines, mesial spine slightly longer than lateral spine, overreaching
penultimate segment, although not overreaching antennal peduncle; penultimate segment unarmed.
Ischium of third maxilliped about 1.5 times length of merus measured along dorsal margin, distoventrally
bearing spine. Merus of third maxilliped with two well developed spines on flexor margin, distal smaller; extensor
margin with small distal spine.
Chelipeds subequal, squamous, with numerous short uniramous setae denser on mesial borders of articles. Palm
slightly shorter than fingers. Merus armed with some spines, strongest spine on distal border short, not
overreaching proximal quarter of carpus. Carpus with several spines on dorsal side and several spines scattered on
mesial and ventral sides. Palm with several spines scattered on mesial and dorsal sides and one row of lateral
spines, continuing onto along fixed finger. Fingers distally curving and crossing, ending in a sharp point; movable
with some spines along mesial border; cutting edges with small teeth of different sizes.
Second pereiopod about twice carapace length; merus shorter than carapace, about 6 times as long as high,
about 3 times carpus length and 1.5 times as long as propodus; propodus 5.5 times as long as high, about
1.5 times dactylus length. Merus with small spines on dorsal border, increasing in size distally, ventral margin
with one long distal spine. Carpus with few dorsal spines and one distoventral spine. Propodus with 8-10 movable
ventral spines. Dactylus with dorsal margin slightly convex on proximal half, slightly curving distally, with
6-7 movable spinules along entire ventral margin. Third pereiopod similar to second; fourth pereiopod shorter
than second and third. Merus of fourth pereiopod 2/3 length of second pereiopod. Epipods absent from all
pereiopods.
Remarks. — Munida glabella belongs to the group of small species having five spines on the lateral margins
of the carapace behind the cervical groove, moderately large eyes, an unarmed second abdominal somite, lateral
parts of the posterior thoracic sternites without granules or carinae, spiniform rostrum, the distolateral spine of the
basal antennular segment shorter than the distomesial, and the dactylus of the walking legs bearing movable
spinules along the entire ventral border. The new species is closely related to M. pusiola Macpherson, 1993, from
(he Philippines (MACPHERSON, 1993).
However, the two species are distinguished by several characters:
— The frontal margins are clearly more oblique in M. pusiola than in the new species.
— The distal spines of the basal segment of the antennular peduncle are subequal in M. pusiola, whereas the
distomesial spine is longer than the distolateral spine in the new species.
— The dactylus of the walking legs are as long or slightly shorter than the propodus in M. pusiola. but are
about 2/3 propodus length in the new species.
Munida plexaura Macpherson & de Saint Laurent, 1991
Munida plexaura Macpherson & de Saint Laurent 1991: 396, fig. 7. p). IE. — Poupin, 1996: 24, pi. I la.
MATERIAL EXAMINED. — Marquesas Islands. MUSORSTOM 9: stn 1148. 300 m: 1 3 6.1 mm; I juv. 5.0 mm
(MNHN-Ga 4357). — Stn 1282, 416-460 m: 5 d 8.8 to 13.2 mm; 1 juv. 6.0 mm (MNHN-Ga 4358). — Stn 1287, 163-
245 m: 1 <3 12.6 mm (USNM).
Distribution. — Previously known from French Polynesia: Tuamotu (Fakarava and Mururoa) and Tubuai
(Raevavae) Islands in 350-398 m. The present material was collected from 163-460 m.
Munida tuberculata Henderson, 1885
Munida tuberculata - MACPHERSON, 1996: 408 (references); 1999: 424.
Material EXAMINED. — Marquesas Islands. Musorstom 9: stn 1268. 285-320 m: 1 ov. 2 3.8 mm (MNHN-Ga
4359). — Sin 1269, 420-430 m: 1 ov. 2 6.8 mm (MNHN-Ga 4360). — Stn 1282, 416-460 m: 2 3 4.3 and 5.0 mm
(MNHN-Ga 4361).
420
E. MACPHERSON
DISTRIBUTION. — The species has previously been reported from Fiji, Vanuatu, New Caledonia, Matthew and
Hunter Islands, Wallis and Futuna area, between 420 and 650 m. The material studied here was collected from
285-460 m.
Munida typhle Macpherson, 1994
Munida typhle Macpherson 1994: 549, fig. 60; 1999: 425.
Material EXAMINED. — Marquesas Islands. MUSORSTOM 9: stn 1262, 850-905 m: 1 2 8.3 mm (MNHN-Ga
4362). — Stn 1278, 1000 m: 4 3 4.5 to 5.7 mm; 2 ov, 2 5.4 and 6.7 mm; 3 2 5.0 to 5.5 mm (MNHN-Ga 4363).
Remarks. — The specimens from Marquesas Islands agree with the types and the material reported earlier from
New Caledonia and Vanuatu. However, the small spines situated along the anterior ridge of the second abdominal
somite may be absent in the specimens from Marquesas Islands (present in the type material).
Distribution. — Previously known from New Caledonia and Vanuatu, between 1210 and 1470 m. The
specimens from Marquesas Islands were collected from 850-1000 m.
Genus PARAMUNIDA Baba, 1988
Paramunida echinata sp. nov.
Fig. 2
Material EXAMINED. — New Caledonia. MUSORSTOM 9: stn 1146, 200 m: 17 3 5.6 to 11.7 mm; 5 ov. 2 9.1
to 12.6 mm; 2 2 7.8 and 8.5 mm (MNHN-Ga 4364). — Stn 1148. 300 m: 4 <3 9.3 to 10.6 mm; 3 2 7.4 to 1 1.6 mm
(USNM). — Stn 1 154, 102 m: 1 6 13.3 mm (MNHN-Ga 4365). — Stn 1172, 300-302 m: 3 3 1 1.4 to 12.2 mm (MNHN-
Ga 4366). — Stn 1176, 260 m: 17 3 10.8 to 13.4 mm; 13 ov. 2 10.8 to 12.7 mm; 1 2 9.1 mm (MNHN-Ga 4367). —
Stn 1190, 350 m: 1 3 15.5 mm (MNHN-Ga 4368). — Stn 1201, 275-300 m: I 3 13.4 mm (MNHN-Ga 4369). —
Stn 1238, 280-370 m: 1 3 6.8 mm (MNHN-Ga 4370). — Stn 1268, 285-320 m: 33 specimens (MNHN-Ga 4371). —
Stn 1269, 420-430 m: 1 2 6.9 mm (MNHN-Ga 4372).
Types. — The male 13.3 mm (MNHN-Ga 4365) from the stn 1154 has been selected as the holotype. the
other specimens are paratypes.
ETYMOLOGY. — From the Latin echinatus, spiny, in reference to the presence of numerous spinules on the
carapace and abdomen.
Description. — Carapace, excluding rostrum, as long as broad. Dorsal surface covered with numerous
spinules. Gastric region with 3 well developed spines in midline. Cervical groove distinct. Cardiac region slightly
circumscribed, with a median row of 3 well-developed spines, first slightly thicker than the others. Frontal margin
transversal. Lateral margins convex, with some spines and iridescent setae on anterior half. Anterolateral spine well
developed. Rostrum triangular; supraocular spines half as long and more slender than rostrum.
Fourth thoracic stemite with few arcuate striae; fifth to seventh smooth.
Abdomen with two median well-developed spines and one smaller lateral spine on each side on anterior and
posterior ridges on second and third somites. Fourth abdominal somite similar to preceding, but posterior ridge
with distinct single median spine. Some small spines and spiniform tubercles between median and lateral spines.
Eye large, maximum corneal diameter about 1/3 distance between bases of external orbital spines.
Basal segment of antennule (distal spines excluded) exceeding corneae, with distomesial spine shorter than
distolateral. Anterior prolongation of first segment of antennal peduncle clearly overreaching antennular peduncle
by about 1/4 of its length; second segment (spines excluded) about 2 times length of third segment and about
2 times longer than wide, distomesial spine slightly exceeding third segment, distally mucronated, distolateral
spine not reaching end of third segment; third segment slightly longer than wide and unarmed.
Source : MNHN, Paris
CROSN1ERITA. MUNIDA AND PARAMUNIDA FROM MARQUESAS ISLANDS
421
Chelipeds and walking legs long and slender. First walking leg with propodus about 10 times as long as wide
and more than 1.5 times dactylus length.
Fig. 2. — Paramunida echinata sp. nov., <? 13.3 mm. holotype from stn 1 154: A, carapace, dorsal view; B. upper margin
of carapace and rostrum, lateral view; C, sternal plastron; D, ventral view of cephalic region, showing antennula and
antennal peduncles; E, right first walking leg, lateral view.
Colour. — Carapace and abdominal segments reddish; transverse rows of spinules red, with several white
spots on cervical groove. Pereiopods whitish with numerous small red and white spots; chelipeds with reddish band
on articulation of merus-carpus, carpus-propodus and hand-fingers. One specimen with a large transverse whitish
band on gastric and anterior branchial regions.
422
E. MACPHERSON
Remarks. — Paramunida echinata is closely related to P. labis Macpherson, 1996 from Wallis and Futuna
Islands. Both species are characterized by having a rostral spine that is larger than the supraocular spines, the
median gastric region bearing a row of 3-4 spines, the thoracic sternites smooth and the distomesial spine on the
second antennal segment not evenly tapering, but distally mucronated. The two species differ in several characters:
— The rostrum is triangular in the new species, whereas it is spiniform in P. labis.
— The propodus of the walking legs is more than 1.5 times the dactylus length in the new species, whereas
the propodi are slightly longer only than the dactyli in P. labis.
— The carapace and abdominal segments are reddish in the new species but with numerous small, white and
yellow spots in P. labis (see MACPHERSON, 1996).
The new species also resembles P. tricarinata (Alcock, 1894) from the Indian Ocean, but they differ in the
following aspects:
— The thoracic sternites have numerous arcuate striae in P. tricarinata-. in the new species the sternites are
smooth.
— The distomesial spine of the second antennal segment is distally mucronated and not overreaches the
antennal peduncle in the new species; in P. tricarinata this spine is evenly tapering to a sharp tip. clearly
exceeding antennal peduncle.
Size. — Males examined ranged from 5.6 to 14.3 mm; females from 6.8 to 12.8 mm. Females ovigerous
from 9.1 mm.
ACKNOWLEDGEMENTS
I am grateful to A. CROSNIER and B. Richer de Forges (IRD) for their support and help and for making this
interesting material available to me. Thanks are also due to K. Baba (Kumamoto University) for improvements to
the manuscript. The species have been illustrated by J. MACPHERSON and P. LABOUTE (IRD) took the colour
photographs.
REFERENCES
Baba, K., 1981. — A new galatheid crustacean (Decapoda, Anomura) from the Hawaiian Islands. Journal of Crustacean
Biology, 1: 288-292.
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.). Resultats des Campagnes Musorstom.
Vol. 15. Memoires du Museum national d'Histoire naturelle, 168: 433-502.
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 genera Munida Leach, 1820 and Paramunida Baba, 1988
(Galatheidae) from the seas around the Wallis and Futuna Islands. In: A. Crosnier (ed.), Resultats des Campagnes
Musorstom, Vol. 15. Memoires du Museum national d'Histoire naturelle. 168: 387-421.
Macpherson, E., 1998. — A new genus of Galatheidae (Crustacea, Anomura) from the Western Pacific Ocean.
Zoosystema . 20: 351-355.
Macpherson, E., 1999. — Crustacea Decapoda: Species of the genera Agononida Baba & de Saint Laurent. 1996 and
Munida Leach, 1820 (Galatheidae) collected during the Musorstom 8 cruise in Vanuatu. In: A. Crosnier (ed.),
Resultats des Campagnes MUSORSTOM, Vol. 20. Memoires du Museum national d'Histoire naturelle, 180: 407-426.
Source : MNHN, Pans
CROSNIERITA. MU N ID A AND PARAMUNIDA FROM MARQUESAS ISLANDS
423
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 , Paris, (4), 13. sect. A, (3-4): 373-422.
POUPIN, I, 1996. — Atlas des Crustaces marins profonds de Polynesia Frangaise. Recoltes du navire Marara 1 1986/1996).
Service mixte de Surveillance radiologique et biologique (SMSRB). Montlhery. 59 p.
Richer de Forges, B., Poupin, J. & Laboute, P.. 1999. — La campagne Musorstom 9 dans l’archipel des lies Marquises
(Polyndsie franfaise). Compte rendu et liste des stations. In: A. Crosnier (ed.). R6sultats des Campagnes Musorstom.
Vol. 20. Memoires du Museum national d'Histoire naturelle, 180: 9-29.
Titgen, R.H., 1987. — New decapod records from the Hawaian Islands (Crustacea. Decapoda). Pacific Science 41
141-147.
Zarenkov, N.A. & Khodkina. I.V., 1981. — Decapoda. In: A.P. Kuznetsov & A.N. Mironov (eds). Benthos of the
submarine mountains Marcus-Necker and adjacent Pacific Regions. Academija Nauk SSSR, Moscow: 83-93.
Zariquiey Alvarez, R., 1952. — Estudio de las especies europeas del gen. Munida Leach 1818. Eos, 28: 143-231.
Source : MNHN, Paris
JLTATS DES CAMPAGNES MUSORSTOM, VOLUME 21 - RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 21 - RESULTATS DES
Crustacea Decapoda : New species and new records of
Ethusinae (Dorippidae) from Vanuatu
CHEN Huilian (H. L. CHEN)
Institute of Oceanology,
Chinese Academy of Sciences.
7 Nan-Hai Road, Qingdao 266071, China
ABSTRACT
During the MUSORSTOM 8 cruise in Vanuatu, in September and October 1994, 12 species belonging in the genera
Ethusa and Ethusina were collected. Two of them, Ethusina microspina and E. vanuatuensis , are new; all the others are
recorded for the first time in Vanuatu.
RESUME
Crustacea Decapoda : Nouvelles especes et nouvelles recoltes d'Ethusinae (Dorippidae) en
Vanuatu.
Lors de la campagne MUSORSTOM 8 faite en Vanuatu en septembre et octobre 1994, 12 especes appartenant aux genres
Ethusa et Ethusina (Dorripidae) ont 6t6 recoltees. Deux de ces especes, Ethusina microspina et E. vanuatuensis. sont
nouvelles. Les dix autres sont mentionnees pour la premiere fois en Vanuatu.
This report is based on the samples collected in Vanuatu during the MUSORSTOM 8 cruise in September and
October 1994. A total of 12 species belonging to 2 genera (Ethusa and Ethusina) of the subfamily Ethusinae have
been identified, of which two species, Ethusina microspina and Ethusina vanuatuensis, are described as new. The
other species are reported for the first time from Vanuatu.
Species of genus Ethusa were taken from depths of 180-956 m and those of the genus Ethusina from 437-
1620 m.
Chen Huilian (H. L. Chen), 2000. — Crustacea Decapoda : New species and new records of Ethusinae (Dorippidae)
from Vanuatu. In: A. Crosnier (ed.), Resultats des Campagnes MUSORSTOM, Volume 21. Memoires du Museum national
d'Histoire naturelle , 184: 425-435. Paris ISBN 2-85653-526-7.
Contribution no. 3843, Institute of Oceanology, Chinese Academy of Sciences, Qingdao, China.
426
H.L. CHEN
LIST OF SPECIES
Genus ETHUSA Roux, 1830
Ethusa furca Chen, 1993
Ethusa granulosa Ihle, 1916
Ethusa indica Alcock, 1 896
Ethusa izuensis Sakai, 1937
Ethusa major Chen, 1993
Ethusa makasaricaC hen, 1993
Ethusa minuta Sakai, 1937
Genus ETHUSINA Smith, 1884
Ethusina desciscens Alcock, 1 896
Ethusina longipes Chen, 1987
Ethusina microspina sp. nov.
Ethusina robusta Miers, 1886
Ethusina vanuatuensis sp. nov.
SYSTEMATIC ACCOUNT
Subfamily ETHUSINAE Guinot, 1977
Genus ETHUSA Roux, 1830
Ethusa furca Chen, 1993
Ethusa furca Chen, 1993: 321, fig. 4.
Material EXAMINED. — Vanuatu. Musorstom 8: st. CP 980. 19°21.02'S, 169°25.22'E. 433-450 m.
22.09.1994: 1 3 5.8 x 5.4 mm (MNHN-B 27493).
Distribution. — New Caledonia and Vanuatu.
Ethusa granulosa Ihle, 1916
Ethusa granulosa Ihle, 1916: 143, fig. 76. — SERENE, 1968: 40. — Chen. 1993: 323, fig. 5.
Material EXAMINED. — Vanuatu. Musorstom 8: St. DW 1046, 16°53.62’S, 168°11.08’E, 461-480 m,
30.09.1994: 1 $ 8.0 x 7.5 mm (MNHN-B 27494). — St. DW 972, 19°21.94’S. 169°28.07'E, 487-507 m, 20.09.1994:
1 9 8.9 x 9.0 mm (MNHN-B 27495). — St. CP 975, 19°23.60'S, 1 69°28.93'E, 536-566 m, 22.09.1994: 1 o' 6.8 x
6.1 mm (MNHN-B 27496). — St. DW 1045, 16°54.50'S, 168°20.37'E, 459-488 m, 30.09.1994: 1 3 8.9 x 8.1 mm
(MNHN-B 27497).
Distribution. — Indonesia, New Caledonia and Vanuatu.
Ethusa indica Alcock, 1896
Ethusa indica Alcock, 1894: 405; 1896: 283. — Alcock & Anderson, 1895, pi. 14 fig. 2. — Ihle, 1916: 136. — Sakai.
1965: 24, pi. 11 fig. 4; 1976: 64. fig. 27. — SerEne, 1968: 40. — Chen, 1985: 189, figs 8-9. pi. 1 fig. 1, pi. 2
fig. 5; 1986: 128. fig. 10(45-49); 1993: 324; 1997: 618. — Dai & Yang, 1991: 59, figs 27(3-4), pi. 6(4).'
Ethusa gracilipes - SerEne & Lohavanijaya, 1973: 35, figs 56-59, pi. 14 figs c-d. — SerEne & Vadon. 1981: 119. 121
[Not Ethusa I Ethusina ) gracilipes Miers, 1886],
Ethusa serenei Sakai, 1983: 4.
Material EXAMINED. — Vanuatu. Musorstom 8: st. CP 991, 18°51.26'S, 168°52.19'E. 910-936 m.
24.09.1994: 1 2 12.0 x 12.5 mm (MNHN-B 27498). — St. CP 992, 18°52.34’S, 168°55.16'E. 748-775 m, 24.09.1994:
1 3 10.8 x 11.5 mm; 1 $ 9.5 x 9.6 mm; 1 ovig. $ 11.6 x 12.5 mm (MNHN-B 27499). — St. CP 1007, 18°51.97'S,
1 68°55.92'E. 720-830 m, 25.09.1994: 1 ovig. 2 9.9 x 11.0 mm (MNHN-B 27500). — St. CP 1075, 15°53.26'S,
I67°27.21'E, 944-956 m, 04.10.1994: 1 3 (broken); 3 2 9.9 x 11.0 - 10.0 x 11.0 mm (MNHN-B 27501). —
St. CP 1080. 15°57.30'S, 167°27.73'E, 799-850 m. 05.10.1994: 2 3 8.3 x 8.0, 8.5 x 8.4 mm (MNHN-B 27502).
Source : MNHN, Paris
ETHUSINAE FROM VANUATU
427
Distribution. — East China Sea, South China Sea, Japon, the Philippines, Indonesia, New Caledonia and
Vanuatu.
Ethusa izuensis Sakai, 1937
Eiliusa izuensis Sakai, 1937: 80. fig. 4; 1965: 23, pi. 12 figs 1-2; 1976: 66, figs 26d, 29. — Serene, 1968: 40. —
Takeda & Miyake, 1972: 67. — SerEne & Vadon, 1981: 119. — Chen. 1985: 193, figs ll-12.pl. 1 fig 2- 1986
131, fig. 12(59-61); 1993: 324. — Dai & Yang, 1991: 58, figs 27(5-6). pi. 6(5).
Material EXAMINED. — Vanuatu. Musorstom 8: st. DW 1 060, 16°13.82'S. 167°20.80’E 375-397 m O’’ 10
1994: 1 juv. 2 4.6 x 4.8 mm (MNHN-B 27503).
DISTRIBUTION. — China, Japan, the Philippines, New Caledonia and Vanuatu.
Ethusa major Chen, 1993
Ethusa major Chen, 1993: 326, fig. 8.
Material EXAMINED. — Vanuatu. Musorstom 8: st. CP 1124, 15°0I.72'S, 166“56.51'E, 512-599 m, 09 10
1994: 1 $ 15.5 x 15.0 mm (MNHN-B 27504).
Distribution. — Coral Sea, Indonesia, Chesterfield Islands, New Caledonia and Vanuatu
Ethusa makasarica Chen, 1993
Ethusa makasarica Chen, 1993: 327, fig. 9.
Material examined. — Vanuatu. Musorstom 8: st. CP 1124, 15°0I.72'S, 166056.51'E. 532-599 m.
09.10.1994: 4 <3 14.0 x 15.0 - 15.6 x 16.3 mm; 1 2 17.5 x 19.0 mm (MNHN-B 27505). — St. 991. 18°5I.26'S.
168°52. 19'E, 910-936 m, 24.09.1994: 2 3 14.0 x 15.0 mm; 16.0 x 17.5 mm (MNHN-B 27506).
Distribution. — Indonesia and Vanuatu.
Ethusa minuta Sakai, 1937
Ethusa minuta Sakai, 1937: 81, pi. II fig. 2; 1965: 23. pi. 11 fig. 4. — Chen, 1985: 193, figs 11-12, pi. 1 fig. 2; 1 986:
131, figs 12(55-58); 1993: 329, fig. 10. — Takeda & Miyake, 1972: 68. — Dai & Yang, 1991: 58, figs 27(1-2)
pl. 6(3).
Material EXAMINED. — Vanuatu. Musorstom 8: st. 1071, 15°36.63'S, 167°16.34'E, 180-191 m, 04.10.1994:
\ 3 4.0 x 4.0 mm (MNHN-B 27507). — St CP 963, 20°20.10'S, 169°49.08'E, 400-440 m, 21.09.1994: 2 3 9 0 x 8 7
7.7 x 7.3 mm (MNHN-B 27508).
Distribution. — East China Sea, Japan, the Philippines, Indonesia, Coral Sea, Chesterfield Islands,
New Caledonia and Vanuatu.
Genus ETHUSINA Smith, 1884
Ethusina desciscens Alcock, 1 896
Ethusina desciscens Alcock, 1896: 286. — Alcock & McArdle, 1903: pl. 2 figs 2-2a. — Chen. 1985: 197: 1986' 136.
figs 15(71-73); 1987: 689. fig. 7, pl. 2 F.
Source
428
H L. CHEN
Material EXAMINED. — Vanuatu. Musorstom 8: st. CP 1109, 14°52.20'S, 1 67° 1 8.00'E, 1550-1620 m.
08.10.1994: 1 2 10.7 x 10.3 mm (MNHN-B 27509).
DISTRIBUTION. — Madagascar, Laccadive Sea, Andaman Sea, Indonesia, the Philippines, China. Vanuatu.
Ethusina longipes Chen, 1987
Ethusina longipes Chen, 1987: 687, fig. 6. pi. 2 D-E.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: st. CP 1110, I4°49.16'S. 167° 1 5. 1 8'E. 1360 m, 08.10.1994:
1 <3 7.8 x 7.1 mm (MNHN-B 27510). — St. CP 956, 20°33.41'S. 169°35.95'E. 1175-1210 m. 20.09.1994: I <5 7.3 x
7.1 mm (MNHN-B 27511).
Distribution. — Madagascar (N.W. coast) and now Vanuatu.
Ethusina tnicrospiita sp. nov.
Figs 1-2, 5
Material EXAMINED. — Vanuatu. Musorstom 8: st. CP 992, 18°52.34'S. 168°55.16'E. 775-748 m.
24.09.1994: <3 holotype 6.0 x 6.2 mm (MNHN-B 27512). — St. CC 996, 18°52.41'S, 168°55.73'E, 764-786 m.
24.09.1994: 2 2 paratypes 9.0 x 9.2 mm, 8.9 x 9.0 mm ovig. (MNHN-B 27513). — St. CP 1074. 15°48.42'S.
167°24.27'E. 775-798 m. 04.10.1994: 1 <3 paratype 6.1 x 6.2 mm (MNHN-B 27514).
Carapace slightly broader than long, anterior narrow, posterior broad, female carapace more swollen than male.
Dorsal surface covered with short pubescence, mesogastric and branchial regions slightly convex. Middle line of
cardiac region depressed, its lateral sides convex. Cervical, branchial and branchiocardiac grooves broad and shallow
in male, thin in female. Front divided into 4 teeth by one V-shaped and two U-shaped notches: median teeth and
lateral teeth short spine-shaped; outer border of lateral teeth to exorbital spine obliquely straight. Exorbital tooth
minutely spined, part of eyestalk visible from dorsal view, more distinct in female than in male. Anterolateral
border slightly sinuous, with a small incision located between anterolateral and posterolateral borders. Anterior half
of posterolateral borders outwardly swollen, posterior half inwardly convergent. Median part of posterior border
concave.
Male right cheliped slightly larger than left. Merus triprismatic. Larger palm swollen. 1.32 times as long as
high ; smaller palm 1.46 times as long as high. Fingers longer than palm, cutting edges with blunt teeth. Female
chelipeds symmetrical.
Second pair of pereiopods longest, merus 7 limes as long as high. Carpus shorter than propodus. Dactylus
longer than propodus. Last two legs slender and short, with sparsely distributed short setae. Merus of last leg
6 times as long as high.
Male abdomen consists of five segments (3rd-5th fused): first segment 3.5 times as broad as long; second
segment 8 times as broad as long; distal half of lateral border of fused section (segments 3rd-5th) inwardly
convergent, its basal half swollen; sixth segment square, median line of its anterior border concave, anterior outer
border of each side bluntly rounded; telson semi-rounded. Female abdomen divided into seven segments: length of
segments 1-5 about equal; segments 1-3 gradually broadening; segments 4-6 to telson gradually convergent; sixth
segment trapezoid, and telson broadly triangular.
Male first pleopods stout, proximal part stouter than distal part, with 2 blunt processes near distal part. Second
male pleopods slender, not thin, distal half narrower than proximal half.
REMARKS. — This new species is similar to Ethusina brevidentata Chen, 1993 but it may be distinguished
from the latter by its carapace being broader than long, by having short pubescence, lacking sharp granules; only
part of eyestalk visible from dorsal view; minutely spined exorbital tooth; asymmetrical male chelipeds; semi-
rounded telson of male abdomen; male first pleopods with two processes near distal part and slightly stouter second
pleopods.
Source : MNHN, Paris
ETHUSINAE FROM VANUATU
429
Fig
. 1. — Ethusina microspina sp. nov.: holotype 6 6.0 x 6.2 mm (MNHN-B 27512): a. carapace; b, chelipeds (ventral
view); c, anterior sternal shield; d. abdomen; e-f, first pleopod; g. second pleopod. Scales = 1 mm.
Source : MNHN, Paris
430
H.L. CHEN
Fig. 2. — Ethusina microspina sp. nov.: paratype 2 9.0 x 9.2 mm (MNHN-B 27513): a. carapace; b. abdomen.
Scale = 1 mm.
ETYMOLOGY. — From the Greek mikros, small, and the Latin spina, spine, in reference to the shape of the
exorbital tooth.
Ethusina robusta Micrs, 1 886
Fig. 3
Ethusina (Ethusina) var. robusta Miers, 1886: 333, pi. 29 figs 2a-c.
Ethusina gracilipes robusta - SERfeNE, 1968: 40.
Ethusina robusta - Chen, 1986: 133, fig. 13(62-66); 1993: 342.
Material examined. — Vanuatu. Musorstom 8: st. DW 1128, 16°02.14'S, 166°38.39'E, 778-811 m.
10.10.1994: 1 2 8.0 x 8.0 mm (MNHN-B 27515). — St. CP 1037, 18°03.70'S, 168°54.40’E, 1058-1086 m,
29.09.1994: 1 3 11.5 x 11.0 mm (MNHN-B 27516). — St. 1125, 15°57.63'S, 166°38.43'E, 1160-1120 m, 10.10.
1994: 1 2 8.1 x 8.2 mm (MNHN-B 27517). — St. CP 1111, 14°51.09'S. 167°14.00'E. 1210-1250 m. 08.10.1994: 1 2
12.3 x 12.5 mm (MNHN-B 27518). — St. CP 1026, 17°50.35'S, 168 39.33'E. 437-504 m, 28.09.1994: 1 2 7.9 x
7.9 mm (MNHN-B 27519).
Distribution. — China, the Philippines, Indonesia, New Caledonia and Vanuatu.
Ethusina vanuatuensis sp. nov.
Fig. 4
Material examined. — Vanuatu. Musorstom 8: st. CP 1026, I7°50.35'S. 168°39.33'E, 437-504 m,
28.09.1994: 3 holotype 6.0 x 5.6 mm (MNHN-B 27520). — St. CP 1129, 16°00.73'S, I66°39.94'E, 1014-1050 m.
10.10.1994: 1 3 paratype 5.9 x 5.7 mm (MNHN-B 27521).
Carapace slightly longer than broad, covered with short pubescence. Cervical groove narrow, branchial groove
broad and shallow. Protogastric, mesogastric, metagastric and branchial regions slightly convex. Cardiac region
Source : MNHN, Paris
ETHUSINAE FROM VANUATU
431
Fig. : l. — Ethusina robusta Miers, 1886. S 11.5 x 11.0 mm (MNHN-B 27516): a, carapace; b. right chelipcd (ventral
view); c. left chela (ventral view); d. right second leg; e. right last leg; f. abdomen; g. first' pleopod: h. second
pleopod. Scales = 1 mm.
Source MNHN, Paris
432
H.L. CHEN
I _ 1
0.5 mm
h
Fig. 4. — Ethusina vanuatuensis sp. nov.: holotype <J 6.0 x 5.6 mm (MNHN-B 27520): a. carapace and left cheliped;
b. righi cheliped (ventral view); c, left chela (ventral view); d, right fifth leg; e, anterior portion of carapace (ventral
view); f, anterior sternal shield; g, abdomen; h, first pleopod; i, second pleopod. Scales = 1 mm.
Source MNHN, Paris
ETHUSINAE FROM VANUATU
433
cordiform, convex. Front with 4 teeth: median teeth short and sharp, lateral teeth long, blunt. Exorbital teeth long,
sharp, and curved inwards. Posterior half of anterolateral borders and anterior half of posterolateral border gradually
convex, posterior half of posterolateral border gradually convergent. Posterior border concave.
Male chelipeds equal in size and shape. Merus triprismatic, 4 times as long as high. Palm swollen. Finger
longer than palm, movable finger with blunt teeth and a larger tooth and with a small gap near the base;
immovable finger with blunt teeth.
Anterior two pairs of ambulatory legs smooth, second legs longest. Merus 8.5 times as long as high. Carpus
as long as propodus. Dactylus longer than propodus; last two pairs of legs covered with short and dense
pubescence. Merus of last leg 6 times as long as high. Dactylus very short.
Male abdomen consists of five segments (3rd-5th fused): first segment 3 times as long as second; distal half of
lateral border of fused section (segments 3rd-5th) inwardly convergent, its basal half swollen; 6th segment
2.2 times as broad as long; telson bluntly triangular.
Male first pleopods stout, distal third narrower than basal two-thirds, with sparsely distributed short setae, distal
end with 2 lobes. Second pleopods relatively stouter, distal part hooked.
Remarks. — This new species is similar to Ethusina dilobotus Chen, 1993, but may be easily distinguished
from it by the 4 front teeth being slightly longer and the exorbital teeth being inwardly curved, by the anterior two
pairs of pereiopods being much longer and by the different shape of the male first and second pleopods.
Etymology. — This species is named after the locality, Vanuatu, where it has been found.
ACKNOWLEDGMENTS
This study has been supported by a visitors' grant from IRD (ex ORSTOM). The author is very grateful to
A. CROSNIER, IRD and Museum national d'Histoire naturelle, Paris, for providing the material for this study and
reviewing the manuscript; to B. RICHER DE FORGES, IRD Noumea, New Caledonia, for collecting most of the
specimens studied here; to J.F. DEJOUANNET for the drawings of Ethusa microspina , to R. VON COSEL for the
photos.
REFERENCES
Alcock, A., 1894. — Natural History Notes from H.M. Indian Marine Survey Steamer Investigator. Ser. II, No I. On the
results of deep-sea dredging during the season of 1890-1891. Annals and Magazine of Natural History, (6) 13: 225-
245, 321-334. 400-41 1.
ALCOCK. A,, 1896. — Materials for 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. — Illustrations of the Zoology of Royal Indian Marine Surveying Steamer
Investigator under the command of Commander A. Carpenter, R.N., D.S.O., of the Late Commander R.F. Hoskyn,
R.N., and of Commander C.F. Oldham, R.N., Crustacea. Part III. pis 9-15. Calcutta.
ALCOCK, A. & McARDLE, A.F., 1903. — Illustrations of Zoology of Royal Indian Marine Survey Ship Investigator, under
the command of Commander T.H. Hemming, R.N., Crustacea. Part X, pis 56- 67. Calcutta.
Chen, H.L., 1985. — Decapod Crustacea: Dorippidae. In: Rdsultats des Campagnes Musorstom I et II. — Philippines
(1976, 1980), vol. 2. Memoires du Museum national d'Histoire naturelle, (A), 133 : 178-203, figs 1-6. pis 1-2.
CHEN, H.L., 1986. — Studies on the Dorippidae (Crustacea Decapoda Brachyura) of Chinese waters. Transactions of the
Chinese Crustacean Society, (1): 118-139, figs 1-15 (in Chinese with English summary).
Chen, H.L., 1987. — Dorippidae (Crustacea Decapoda Brachyura) collected in Madagascar waters. Bulletin du Museum
national d'Histoire naturelle, Paris, 4e ser., 9, section A. (3) : 677-693, figs 1-7. pis 1-2.
434
H.L. CHEN
Chen, H.L., 1993. — Crustacea Decapoda: Dorippidae of New Caledonia, Indonesia and the Philippines. In: A. Crosnier
(ed.), Resultats des Campagnes MUSORSTOM, vol. 10. Memoires clu Museum national d'Histoire naturelle. 156: 315-
345, figs 1-20.
Chen. H.L., 1997. — Crustacea Decapoda: Ethusinae (Dorippidae), mainly from the Karubar Cruise. In: A. Crosnier &
P. Bouchet (eds), Resultats des Campagne MUSORSTOM, vol. 16. Memoires du Museum national d'Histoire naturelle,
172: 613-625, figs 1-6.
Dai, A.Y. & Yang, S.L., 1991. — Crabs of the Chinese Seas. China Ocean Press. Beijing and Springer-Verlag. Berlin,
i-xxi + 1-682 pp., figs 1-295, pis 1-74.
iHLE, J.E.W., 1916. — Die Decapoda Brachyura der Siboga-Expedition. II. Oxystomata, Dorippidae. Sijoga Expeditie
Monograph, 39bl. livr. 78: 97-158, figs 39-77.
MiERS, E.J., 1886. — Report on the Brachyura collected by H.M.S. Challenger during the years 1873- 1876. Report of
the scientific Results of the Voyage of H.M.S. ‘’Challenger". Zoology, 17, London. Edinburgh and Dublin: 1-L +
1-362, 29 pis.
Sakai, T., 1937. — Studies on the Crabs of Japan II. Oxystomata. Scientific Reports Tokyo Bunrika Daigaku, sect. B.
suppl. 3 (2): 67-192, 45 figs, 19 pis.
Sakai, T., 1965. — The Crabs of Sagami Bay collected by His majesty the Emperor of Japan. Tokyo, Maruzen Co.
Pp. i-xvi + 1-206, figs 1-27 (in English), pi. 1-100, pp. 1-92 (in Japanese), pp. 1-26 (bibliography and index in
English), pp. 27-32 (index in Japanese), 1 map.
Sakai, T., 1976. — Crabs of Japan and the Adjacent Seas. Tokyo, Kodansha Ldt, 3 vol.: i-xxix + 1-773 pp., figs 1-379
(English text), 1-461 pp. (Japanese text), 1-16 pp., 1-251 pis.
Sakai, T., 1983. — Description of new genera and species of Japanes crabs, together with systematically and
biogeographically interesting species. Researches on Crustacea, (12): 1-44. pis 1-8.
Serene, R., 1968. — The Brachyura of the Indo-West Pacific Region. In: Prodromus for a check list of the (non-
p lane tonic) marine fauna of the South East Asia. UNESCO, Singapore, Special publication n°l. Fauna HlCc3: 34-112.
Sf.rCne, R. & Lohavanijaya, P., 1973. — The Brachyura (Crustacea: Decapoda) collected by the Naga Expedition,
including a review of the Homolidae. Naga Report. 4 (4): 1-187, 186 figs, 21 pis.
SerLne, R. & Vadon, C., 1981. — Crustaces Decapodes: Brachyures. Liste prdliminaire, description de formes nouvelles
et remarques taxonomiques. Resultats des Campagnes MUSORSTOM 1- Philippines (18-28 Mars 1976). Memoires
ORSTOM, (91): 117-140, fig. 1-3, pi. 1-4.
Takeda, M. & Miyake, S.. 1972. — Crabs from the East China Sea, V. A remaining collection. OHMU I Occasional
Papers of Zoological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan), 3 (8): 63-91, fig 1
pi. 3.
Source : MNHN , Paris
ETHUSINAE FROM VANUATU
435
Fig. 5. — Ethusina microspina sp. nov.: a, hololype <J (MNHN-B 27512); b. paralype $ (MNHN-B 27513).
Source : MNHN, Paris
Source : MNHN, Pahs
-TATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES
Crustacea Decapoda: A revision
of the Indo-west Pacific species of palicid crabs
(Brachyura Palicidae)
Peter CASTRO
Biological Sciences Departmeni
California State Polytechnic University
Pomona, California 91768-4032, USA
ABSTRACT
The taxonomy of the crabs belonging to the family Palicidae Bouvier. 1898 from the Indo-west Pacific region is
revised. On the basis of extensive material collected by French expeditions in the Coral Sea and other regions of the
Pacific and Indian oceans, as well as material from numerous museums, including most of the types, the present study
recognizes two subfamilies, 10 genera, and 43 species. Of these taxa. four are new genera: Exopalicus. Miropalicus.
Paliculus, and Rectopalicus. Manella is synonymized with Crossotonotus A. Milne Edwards. 1873. Parapleurophricoides
Nobili, 1906, sometimes believed to be a palicid. is a xanthoid and it is removed from the Palicidae. Nine nominal
species described by previous authors are synonymized and an additional 17 species are described.
RESUME
Crustacea Decapoda : Revision des especes indo-ouest pacifiques de crabes palicides
(Brachyura Palicidae).
La taxonomic des crabes de la famille Palicidae Bouvier, 1898. provenant de 1’Indo-ouest Pacifique est revisee. Basee
sur l'abondant matdriel recolte par les expeditions frangaises en mer de Corail et dans d'autres regions des oceans Pacifique
et Indien. ainsi que sur du materiel provenant de nombreux museums renfermant la plupart des types, la presente etude
reconnait deux sous-familles, dix genres et 43 especes. Parmi ces taxa, quatre sont des genres nouveaux : Exopalicus.
Miropalicus. Paliculus. et Rectopalicus. Le genre Manella est mis en synonymie avec Crossotonotus A. Milne Edwards,
1873. Parapleurophricoides Nobili, 1906, parfois considcre comme etant un palicide, est un xanthide et est retire de la
famille Palicidae. Neuf especes decriles anterieurement sont mises en synonymie, tandis que 17 autres sont decrites
comme nouvelles.
Castro. P., 2000. — Crustacea Decapoda : A revision of the Indo-west Pacific species of palicid crabs (Brachyura
Palicidae). In: A. CROSNIER (ed.). Resultats des campagnes MUSORSTOM, Volume 21. Memoires du Museum national
d'Histoire naturelle, 184: 437-610. Paris ISBN: 2-85653-526-7.
438
P CASTRO
TABLE OF CONTENTS
Introduction . 439
Presentation and Terminology . 440
Systematic Account . 443
Family PALICIDAE Bouvier, 1898 . 443
Key to the subfamilies of the family Palicidae . 445
Subfamily PALICINAE Bouvier, 1898 . 445
Key to the Indo-west Pacific genera of the subfamily Palicinae . 448
Genus PSEUDOPALICUS Moosa & Serene, 1981 . 448
Key to the species of Pseudopalicus . 451
Pseudopalicus acanthodactylus sp. nov . 452
Pseudopalicus amadaibai ( Sakai, 1963) . 455
Pseudopalicus declivis sp. nov . 458
Pseudopalicus glaber sp. nov . 462
Pseudopalicus oahuensis ( Rathbun, 1906) . 465
Pseudopalicus sexlobatus (Kensley, 1969) . 467
Pseudopalicus investigatoris (Alcock, 1900) . 469
Pseudopalicus serripes (Alcock & Anderson, 1895) . 473
Pseudopalicus sp . 474
Pseudopalicus pictus sp. nov . 476
Pseudopalicus macromeles sp. nov . 480
Pseudopalicus undulatus sp. nov . 483
Genus PARAPALICUS Moosa & Serene, 1981 . 486
Key to the species of Parapalicus . 488
Parapalicus ambonensis Moosa & Serene, 1981 . 489
Parapalicus armatus sp. nov . 492
Parapalicus clinodentatus sp. nov . 495
Parapalicus denticulatus sp. nov . 499
Parapalicus inermis sp. nov . 502
Parapalicus elaniticus (Holthuis, 1977) . 505
Parapalicus inanis sp. nov . 506
Parapalicus microphthalmus sp. nov . 509
Parapalicus nanshaensis Dai and Xu, 1991 . 510
Parapalicus piruensis Moosa & Serene, 1981 . 512
Parapalicus trispiralis sp. nov . 5 1 3
Parapalicus trituberculatus (Chen, 1981) . 517
Parapalicus unidentatus (Zarenkov, 1968) . 519
Genus MIROPALICUS new genus . 520
Miropalicus vietnamensis (Zarenkov, 1968) . 522
Genus PALICULUS new genus . 525
Key to the species of Paliculus . 527
Paliculus kyusyuensis (Yokoya, 1933) . 527
Paliculus foliatus sp. nov . 530
Paliculus sp . .
Genus RECTOPALICUS new genus . 533
Key to the species of Rectopalicus . 535
Rectopalicus woodmasoni ( Alcock, 1900) . 535
Rectopalicus amphiceros sp. nov . 533
Rectopalicus ampullatus sp. nov . 54]
Source : MNHN, Paris
1ND0-WEST PACIFIC SPECIES OF PALICID CRAfSS
439
Genus EXOPALICUS new genus . 543
Exopalicus maculatus (Edmondson, 1930) . 546
Genus NEOPALICUS Moosa & Serene, 1981 . 548
Key to the species of Neopalicus . 550
Neopalicus contractus (Rathbun, 1902) . 550
Neopalicus jukesii (While, 1847) . 554
Genus PALICOIDES Moosa & Serene, 1981 . 558
Key to the species of Palicoides . 561
Palicoides longimanus (Miyake, 1936) . 561
Palicoides whitei (Miers, 1884) . 565
Subfamily CROSSOTONOTINAE Moosa & Serene, 1981 . 568
Key to the genera of the subfamily Crossotonotinae . 569
Genus CROSSOTONOTUS A. Milne Edwards, 1873 . 569
Key to the species of Crossotonotus . 57 1
Crossotonotus compressipes A. Milne Edwards, 1873 . 571
Crossotonotus spinipes (De Man, 1888) . 574
Crossotonotus ceramensis (Moosa & Serene, 1981) . 578
Crossotonotus lophocheir sp. nov . 579
Genus PLEUROPHRICUS A. Milne Edwards, 1873 . 582
Key to the species of Pleurophricus . 583
Pleurophricus cristatipes A. Milne Edwards, 1873 . 583
Pleurophricus longirostris (Moosa & Serene, 1981) . 585
Ecology and other Aspects of the Biology of the Indo-West Pacific Palicids . 585
Habitat . 585
Biogeography . 586
Diet . 592
Sexual dimorphism and mating behavior . 592
Role of the fifth pair of pereopods . 592
Acknowledgments . 593
References . 594
Index . 606
INTRODUCTION
Brachyuran crabs belonging the family Palicidae Bouvier, 1898 are a group rather neglected by taxonomists and
ecologists alike, notwithstanding their sometimes frequent occurrence in several marine environments. The feeding
habits, locomotion, reproduction, and larval development of palicids remain practically unknown. The evolution
and biological significance of the reduced last pair of pereopods of most species remain vexing questions. The
taxonomy of the family and its position within the Brachyura have also remained much neglected subjects.
The family Palicidae consisted of four ill-defined genera until the revision of the Indo-west Pacific species by
Moosa & Serene (1981): Palicus Philippi, 1838, Crossotonotus A. Milne Edwards, 1873, Pleurophricus
A. Milne Edwards, 1873, and Manella Rathbun, 1906. A fifth genus, Parapleurophrycoides Nobili, 1906 was
questionably included in the family by Serene (1968, as Pleurophrycoides [sic]). Most species were grouped into
one genus, Palicus, with Rathbun (1918) recognizing 21 western Atlantic and eastern Pacific species in six
"groups”. Three more species from this area (all in Palicus) were subsequently described. An additional species,
P. caronii (Roux, 1830), is Mediterranean and eastern Atlantic in distribution. In the Indo-west Pacific, 24 species
(16 in Palicus) had been described until 1981 (SERENE, 1968; Zarenkov, 1968). While describing two new
western Pacific species, Zarenkov (1968: 765) briefly commented on Rathbun's groups of Palicus species
without revising her classification.
440
P. CASTRO
The first and until now the only major revision of Indo-west Pacific palicids was that of MOOSA & SERENE
(1981). They separated the family into two subfamilies, Palicinae s. str. and Crossotonotinac Moosa & Serene.
1981. For the Palicinae, MOOSA & SERENE (1981) restricted Palicus for the Atlantic, Mediterranean, and eastern
Pacific species, and established four new genera ( Neopalicus , Palicoides, Parapalicus , and Pseudopalicus) for the
26 species they recognized from the Indo-west Pacific. Six new species were described from these genera. Within
the subfamily Crossotonotinae, MOOSA & SERENE (1981) initially recognized four genera. Crossotonotus ,
Parapleurophrycoides , Pleurophricus, and Manella , with a total of six species. Parapleurophrycoides and its only
species, however, were "discarded" since the description of the species was based on a juvenile "which cannot be
used for the definition of the genus" (MOOSA & SERENE, 1981 : 52).
Unfortunately, the revision of MOOSA & SERENE (1981) was based on limited material. They examined
specimens belonging to only nine of the 24 Indo-west Pacific species known at the time and the type material of
only one of the nine species. In addition, the identities of several species (such as Parapleurophrycoides roseus
Nobili, 1906) were left unresolved.
Three Indo-west Pacific species of palicids were described after MOOSA & Serene's revision: Palicus
trituberculatus Chen, 1981 (its description was apparently published before the revision). Palicus bidentatus Sakai,
1983. and Parapalicus nanshaensis Dai & Xu, 1991, bringing the total number of nominal species known from the
region to 33. About 58% of all the known species of palicids (34 out of 58) are Indo-west Pacific in distribution.
The study of a large collection of palicids from numerous French expeditions (mostly from the Coral Sea but
also from several other areas of the Indo-west Pacific region) plus type material and other specimens from
museums, institutions, and private collections from around the world provided the opportunity to undertake a more
comprehensive revision of the Indo-west Pacific Palicidae.
PRESENTATION AND TERMINOLOGY
A list of complete references is given for each of the species treated, while references for the family are
restricted to those where characters or the status of the family are discussed. All geographic names in English
follow, whenever possible, the orthography given in the ninth revised edition (1993) of the Times Atlas (Times
Books, London).
The measurements given in the text, unless specified, refer to carapace length (CL) and carapace width (CW).
The length of the carapace was measured across the middle of the carapace from the tip of the longest lobe of the
frontal border to the posterior border, including any tubercles along the posterior border. The width of the carapace
was measured across the widest breadth, including the longest anterolateral tooth on each side.
The total length of the walking legs (P2-4) was measured along each segment from the proximal edge of the
merus to the tip of the dactylus. The fifth pair of pereopods were measured from the proximal edge of the basis-
ischium to the tip of the dactylus. The abdomen was treated as consisting of six segments and a telson.
The terminology used to refer to parts of the dorsal surface of the carapace and pereopods is summarized in
Fig. 1 , while that used to refer to the anterior and ventral surfaces of the carapace, male and female abdomens, and
male first pleopods is summarized in Fig. 2.
The depth range reported for each species gives the minimum and maximum depths at which each species was
collected with accuracy. Extreme depth values are also given for some species since dredgings depths in some of
the stations varied widely and it was impossible to know the exact depths of collection.
The following abbreviations are used to refer to the deposit of the examined material.
AM. — Australian Museum. Sydney, Australia.
ASIO. — Institute of Oceanology, Academia Sinica, Qingdao, China.
ASIZB. — Academia Sinica, Beijing, China.
ASIZT. — Institute of Zoology, Academia Sinica, Taipei, Taiwan.
BMNH. — The Natural History Museum [formerly British Museum (Natural History)], London, U.K.
BPBM. — Bishop Museum. Honolulu. Hawaii, U.S.A.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
441
postorbital
angle
PI (cheliped)
P2 (first walking leg)
retina
peduncle
(second walking leg)
inner outer
lobe lobe
supraorbital
border
P4 (third walking leg)
episternal process
Fig. 1. — Terminology used to describe ihe carapace and the pereopods.
CBM. — Natural History Museum and Institute, Chiba, Japan.
HSM. — Hayama Shiosai Museum, Hayama, Japan.
KMNH. — Kitakyushu Museum of Natural History, Kitakyushu. Japan.
KPM. — Kanagawa Prefectural Museum of Natural History, Odawara, Japan.
LACM. — Natural History Museum of Los Angeles County, Los Angeles, California. U.S.A.
MCZ. — Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A.
MNHN. — Museum national d'Histoire naturelle, Paris, France.
MZUF. — Museo di Zoologia "La Specola", Universita di Firenze, Italy.
NTOU. — National Taiwan Ocean University, Keelung, Taiwan.
PMBC. — Phuket Marine Biological Center. Phuket, Thailand.
QM. — Queensland Museum. Brisbane, Australia.
RDC. — Research & Development Center for Oceanography, Jakarta, Indonesia.
RMNH. — Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van Natuurlijke Historic). Leiden,
The Netherlands.
SAM. — South African Museum, Cape Town, South Africa.
SMF. — Forschungsinstitut Senckenberg, Frankfurt am Main, Germany.
UMZ. — University Museum of Zoology, University of Cambridge, Cambridge. United Kingdom.
USNM. — United States National Museum, Smithsonian Institution, Washington, D.C., U.S.A.
Source : MNHN , Paris
442
P. CASTRO
pterygostomial lobe
sperm channel
E: MALE FIRST PLEOPOD
B: SUBORBITAL BORDER
complete transverse
Fig. 2. — Terminology used to describe: A, anterior end of the carapace; B, suborbital border of the carapace in
Pseudopalicus and most other genera of palicids; C. male abdomen; D. female abdomen; E. male first pleopod.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
443
WAM. — Western Australian Museum, Perth, Australia.
ZLKU. — Zoological Laboratory, Kyushu University, Fukuoka, Japan (now at Kitakyushu Museum of
Natural History, Kitakyushu, Japan).
ZMA. — Zoologisch Museum, Universiteit van Amsterdam, The Netherlands.
ZMB. — Museum fur Naturkunde (Zoologisches Museum), Universitat Humboldt, Berlin, Germany.
ZMMU. — Zoological Museum, Moskow State University.
ZMUC. — Zoologisk Museum, Kpbenhavns Universitet, Copenhagen, Denmark.
ZMUH. — Zoologisches Institut und Zoologisches Museum, Universitat Hamburg, Germany.
ZRC. — Zoological Reference Collection, Raffles Museum, Department of Biological Sciences, National
University of Singapore, Singapore.
The expedition stations from which material was examined were described in the following publications:
Bathus l to Bathus 4: Richer de Forges & Chevillon, 1996.
Beryx 1 1: Lehodey et al., 1992.
Biocal: Richer deForges, 1990.
Bordau i : Richer de Forges et al., 2000b
Chalcal 1: Richer deforces, 1991.
Chalcal 2: Richer deForges, 1990.
Corail 2: Richer de Forges, 1991.
Corindon: Moosa, 1984.
Expedition Montrouzier: Bouchet, 1994.
Halical 1: Grandperrin etal, 1995.
Halipro l: Richer de Forges & Chevillon, 1996.
Karubar: Crosnier etal., 1997.
Lagon: Richer deForges, 1991.
Musorstom l: Forest, 1981.
Musorstom 2: Forest, 1985.
Musorstom 3: Forest, 1989.
Musorstom 4 to Musorstom 6: Richer deForges, 1990.
Musorstom 5: Richer deforces etal., 1986.
Musorstom 7: Richer de Forges & Menou, 1993.
Musorstom 8: Richer de Forges etal., 1996.
Musorstom 9: Richer de Forges etal., 1996.
Musorstom 10: Richer de Forges et al., 2000a
Reves 2: Marchal etal., 1981.
Smib 3, 4: Richer de Forges, 1990.
Smib 5, 6: Richer deforces, 1991; 1993.
Smib 8: Richer de Forges & Chevillon, 1996.
"Vauban" dredgings: Richer de Forges, 1990.
VoLSMAR: LABOUTEera/., 1989.
There is no available report on the LlTHIST cruise carried out by R.V. "Alis" on the Norfolk Ridge
Seamounts (4-8 August 1999).
SYSTEMATIC ACCOUNT
Family PALICIDAE Bouvier, 1898
Cymopoliidae Faxon, 1895: 38. — Rathbun, 1918: 182. — Sakai, 1939: 607. — Bouvilr, 1940: 303. — MONOD,
1956: 387. — Balss, 1957: 1661.
444
P CASTRO
Pallets Bouvier, 1897: 785 [vernacularl.
Palici Bouvier, 1898a: 105.
Palicidae - Rathbun, 1898: 600. — ALCOCK, 1900: 285, 450. — Zarenkov, 1968: 762, 765. — Takeda & ShimazaKI,
1974: 75. — Guinot, 1978: 249; 1979: 1 1 1. — Moosa & Serene, 1981 : 22. — Williams, 1984:481.
Palicae Bouvier, 1898b: 56, 58.
Type Genus. — Palicus Philippi, 1838.
DIAGNOSIS. — Dorsal surface of carapace typically depressed, granular, often with high bosses of varying
sizes; confluence of branchial and mesogastric regions depressed. Frontal border of carapace divided into 2-4 lobes.
Anterolateral borders each with a varying number (1-4) of triangular, rounded, or truncate teeth. Supraorbital and
suborbital borders long, usually with lobes (may be dentiform); orbits deep, wide. Antennae with basal segment
rectangular or variously expanded; flagellum well developed. Retina of eyes dorsoventrally flattened or spherical;
peduncles typically long, with soft or granular tubercles. Chelipeds small to moderate in size, equal or unequal
(particularly in males). Antennules long, transversely folded beneath front; interantennular septum narrow.
Epistome ventrally inclined (sunken) or dorsoventrally expanded. Buccal cavity square, mostly but not completely
covered by third maxillipeds. Mcri of third maxillipeds small, with palp on each inner border. Fifth pair of
pereopods (P5) characteristically reduced in size (less than CL in practically all species), conspicuously more
slender or of same morphology as anterior walking legs (P2-4). Male opening sternal; sperm ducts under sternal
plates, penis soft, curved, free on inner side of coxae. Male first pleopods with helicoidal or sinuous basal parts,
distal parts uniramous or biramous. Male and female abdomens with six segments plus telson, some (segments 1-
2) can be dorsoventrally compressed, others fused. Male abdomen narrower than sternum, with both sides parallel
to each other or triangular; locking mechanism with very low to medium-size tubercle at edge of thoracic sternite 5
and shallow concavity on underside (ventral surface) of abdominal segment 6. Abdomen of mature females rounded,
broad, thoracic sternite 8 visible laterally; triangular in immature females. Vulva large, visible on thoracic sternite
5 but actually in sternite 6.
Subfamilies included. — Palicinae Bouvier, 1898; Crossotonotinae Moosa & Serene. 1981.
Discussion. — The use of the name Palicidae (with Palicus Philippi. 1838 as the type genus) for the family
instead of Cymopoliidae (with Cymopolia Roux, 1830 as the type genus) is by now accepted. The name
Cymopolia is preoccupied by an alga, Cymopolia Lamouroux, 1816 (see HOLTHU1S & Gottlieb, 1958: 104).
The use of Cymopolia and Cymopolidae has been rejected and the name "Palicidae Rathbun" placed in the official
list of valid names ( Bulletin of Zoological Nomenclature , 1964, 21: 336-351; 338, 341, 342, 344).
The correct authorship of the name Palicidae is a complicated question that fortunately has been clarified by
L.B. Holthuis (in litt.). Rathbun (1898: 600) is generally credited as first using the family name Palicidae. It is
the author of the family name Palicidae that is placed in the official list of valid names in zoology (see above).
BOUVIER (1897: 785) for the first lime used the term "Palices" for a group based on the name of the genus Palicus
but the name is not valid since it is a vernacular name (International Code of Nomenclature, fourth edition. 1999,
article 1 1.3). An English translation (Bouvier, 1898a) of Bouvier's 1897 paper uses "palicids" as a translation of
"Palices" and as a vernacular name it is also invalid. A footnote on the same paper (Bouvier. 1898a: 105) that is
absent in the original paper in French states that the subfamily Dorippinae was to be subdivided into two tribes:
"Palici ( Palicus ) and Dorippi ( Ethusa , Ethusina , Dorripe)". The name "Palici" is an available name for the family
since it was used for a suprageneric taxon and it was written in Latin. The name "Palicae" was also used by
Bouvier (1898b: 56, 58), which was published in the same year. Since its exact publication date could not
be determined, 31 December 1898 is adopted as the publication date (International Code of Nomenclature, fourth
edition, 1999, article 21.3.2). Palicidae BOUVIER. 1898 (a correction of "Palici"), which was published in January
1898 (Bouvier, 1898a), thus has priority over Palicidae Rathbun, 1898, which was published in June 1898.
The systematic position of the family Palicidae within the Brachyura remains uncertain. The placement of
the family in relation to other families has varied from author to author (see Moosa & Serene, 1981: 22:
Williams, 1984: 481). Before the establishment of the family, some of the species now included in the
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
445
subfamily Crossotonotinae were initially placed in various groups of the Brachyura (see discussion of ihe sub¬
family below).
Most evidence places the family close to the family Dorippidae (superfamily Dorippoidca of Heterotremala
following GuiNOT. 1977). There are similarities in the functional morphology of the male and female reproductive
systems (Guinot, 1979: 1 12, 1 13; 1978: 250) and in the circulation of respiratory water (Moosa & Serene.
1981: 23) as well as in what little is known about the morphology of the larvae (Gurney, 1942: 275). The
investigation of larval development and DNA analyses should provide some fresh evidence to support this or other
alternate hypotheses.
The family was divided into two subfamilies by Moosa & SERENE (1981: 23). Although radically different in
the shape of their walking legs, Crossotonotus and the allied Pleurophricus , which are grouped togelher as the
subfamily Crossotonotinae, share many characters with the remaining palicids so their inclusion in the family
Palicidae is not in doubt.
Key to the subfamilies of the family Palicidae
1. Fifth pair of pereopods (P5) conspicuously reduced, shorter, more slender than anterior
walking legs (P2-4), with articulation to carapace dorsal to those of P2-4. Segments 1-2 of
male and female abdomens dorsoventrally compressed and very narrow in comparison to
remaining four segments of abdomen . PALICINAE Bouvier. 1898
— Fifth pair of pereopods (P5) shorter but similar in shape to anterior walking legs (P2-4),
with articulation to carapace at same level as those of P2-4. Segments 1-2 of male and
female abdomens wide, not dorsoventrally compressed .
. CROSSOTONOTINAE Moosa & Serene, 1981
Subfamily PALICINAE Bouvier, 1898
Palicinae Moosa & Serene, 1981: 23.
TYPE Genus. — Palicus Philippi, 1838
DIAGNOSIS. — Fifth pair of pereopods (P5) reduced in size to less than length of carapace, different in shape to
anterior walking legs (P2-4); articulation to carapace dorsal to walking legs. Carapace with very conspicuous teeth
or pointed tubercles only along anterolateral border. Dorsal surface of carapace typically convex. Episternal process
of thoracic sternite 7, sometimes reduced, overhanging each posterolateral border of carapace posterior to fifth pair
of pereopods. Epistome dorsoventrally broadened. Abdomen of males and females with segments 1-2 dorsoventrally
compressed, some segments may be fused, complete or incomplete, transverse ridges (carinae) may be present.
Genera included. — Nine genera are included, by alphabetical order: Exopalicus new genus, Miropalicus
new genus, Neopalicus Moosa & Serene, 1981, Palicoides Moosa & Serene, 1981. Paliculus new genus. Palicus
Philippi, 1838, Parapalicus Moosa & Serene, 1981, Pseudopalicus Moosa & Serene, 1981. Rectopalicus new
genus.
Discussion. — The subfamily was recognized by Moosa & Serene, 1981 as a group for all palicids with
a reduced and slender fifth pair of pereopods. The subfamily includes all Atlantic, Mediterranean, and eastern
Pacific species (all of which make up Palicus) as well as eight genera that are restricted to the Indo-west Pacific
region. Palicus [with Palicus granulatus Philippi, 1838, a junior subjective synonym of Cymopolia caronii Roux.
1830, as its type species by monotypy] needs to be revised. As it now stands, it is undoubtedly heterogeneous,
a suggestion indirectly made by RATHBUN (1918: 184) with her grouping of 21 American species into six
different groups.
446
P. CASTRO
Table 1. — Morphological differences between Palicus Philippi, 1838 (as exemplified by P. caroni Roux, 1830).
Parapalicus Moosa'' & Serbne, 1981, Pseudopalicus Moosa & Serene, 1981, Parapalicus Moosa & Serene, 1981.
Miropalicus new genus, Paliculus new genus, Rectopalicus new genus, Rectopalicus new genus. Neopalicus Moosa &
Serene, 1981, and Palicoides Moosa & Serene, 1981.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
447
Source : MNHN, Paris
448
P. CASTRO
Key to the Indo-west Pacific genera of the subfamily Palicinae
1. Walking legs (P2-4) not slender, with flattened carpi and propodi; meri with unequal
tubercles or teeth (rounded tubercles if equal or nearly equal) along upper and lower borders;
dactyli may be armed with teeth or spines along complete length of posterior borders . . 2
— Walking legs (P2-4) typically very slender, with elongate meri, filiform (long, very
narrow) propodi and dactyli; meri with equal or nearly equal, small (or Hat, rounded if not
small) tubercles along upper and lower borders; dactyli without teeth or spines along
complete length of posterior borders . 6
2. Carapace transversely hexagonal (clearly wider than long), with conspicuously curved
anterolateral borders that are armed with triangular and pointed or rounded teeth . 3
— Carapace subquadrate (almost as wide as long), with straight or only slightly curved
anterolateral borders that are armed with truncate teeth . 4
3. Dorsal border of carapace slightly convex. Eye peduncles long. Walking legs long to very
long, P3 much longer than carapace length (1.5 CL or more) .
. . . . P .S' E l! DO PA LI C U S Moosa & Serene. 1981
— Dorsal border of carapace conspicuously convex, almost hemispherical in outline. Eye
peduncles very short (Fig. 38a). Walking legs conspicuously short. P3 only slightly
longer than carapace length (1.2 CL or less) . EXOPALICUS new genus
4. Supraorbital borders v/ith rounded or rectangular lobes. Posterior borders of propodi and
dactyli of walking legs (P2-4) with spines . RECTOPALIC'US new genus
— Supraorbital borders with triangular, dentiform lobes. Posterior borders of propodi and
dactyli of walking legs (P2-4) entire, without spines . 5
5. Eye peduncles with rounded, granular tubercles. Anterior borders of propodi of P3-4 with
wide, convex enlargement . NEOPALICUS Moosa & Serene, 1981
— Eye penduncles with thin, calcified, crescent-shaped process on anterior border. Anterior
borders of propodi of P3-4 without conspicuous enlargement .
. PALICOIDES Moosa & Serene. 1981
6. One triangular or rounded tooth on anterolateral side of carapace. Abdomen of males
triangular. Basal antennal segment slightly expanded distally, forming scale-like structure
. PARAPAL1CUS Moosa & Serene, 1981
— Two to four narrow, pointed teeth or 2-5 rounded tubercles on anterolateral side of carapace.
Abdomen of males elongate (sides almost parallel to each other). Basal antennal segment
rectangular or, if expanded distally, not forming scale-like structure . 7
7. Supraorbital borders with two very short lobes. Two to five rounded tubercles on
anterolateral side of carapace. Abdomen of mature females with all segments free .
. M1ROPALICUS new genus
— Supraorbital borders with 1-2 rounded or pointed lobes. Two to four narrow, pointed teeth
on anterolateral side of carapace. Abdomen of mature females with segments 3-6 or 4-6
fused . PALICULUS new genus
Genus PSEUDOPALICUS Moosa & Serene, 1981
Pseudopalicus Moosa & Serene, 1981: 35.
Type Species. — Palicus serripes Alcock & Anderson. 1895 by original designation (MOOSA & SERENE,
1981). Gender: masculine.
INDO-WEST PACIFIC SPECIES OF PALIC1D CRABS
449
Diagnosis. — Frontal border of carapace divided into four elongate or rounded lobes, inner pair more advanced
anteriorly than outer pair. Anterolateral borders each with 3-4 triangular or rounded teeth. Dorsal surface of carapace
typically with high, granular bosses. Eyes dorsoventrally flattened or spherical; peduncle with small, typically
granular tubercles. Supraorbital borders each with two triangular or rounded lobes. Suborbital borders each
typically with two rounded or square lobes. Each basal antennal segment rectangular, typically slender, without
distal expansion. Epistome narrow, vertically inclined; border with at least two triangular or rounded median teeth.
Chelipeds increasingly unequal with increasing body size (particularly in males), smallest typically more slender in
females. First three pairs of walking legs (P2-4) with flattened (not filiform) carpi, propodi, and (with one
exception) dactyli. First pair (P2) shorter than second and third pairs (P3-4); third pair slightly shorter or about as
long as second pair. Borders of meri of P2-4 with thick tubercles or triangular teeth; posterior border of dactyli
smooth or armed with teeth or spines. Fifth pair of pereopods (P5) reduced (0.6-0.9 CL), slender. Abdomen of
males elongate (sides almost parallel to each other), with segments free, one transverse ridge each along some or
all segments. Male first pleopods long, slender; basal parts straight or sinuous but not helicoidal; each distal part
biramous, rarely uniramous. Abdomen of mature females with all segments free, one transverse ridge along each
segment 1-4.
REDESCRIPTION. — Carapace transversely hexagonal, broader than long, with conspicuous, granular bosses
that vary in sizes and arrangements. Anterolateral borders each with 3-4 triangular or rounded teeth, below which
carapace slightly rounded. Confluence of branchial and mesogastric regions depressed, smooth; no conspicuous
sulcus between hepatic and branchial regions. Thoracic stemite 7 typically with reduced process (episternal process)
at each outer edge and posterior to insertion of fifth pair of pereopods (P5), visible dorsally as very short process
below posterolateral border of carapace.
Frontal border divided into four elongate or rounded lobes, inner pair more advanced anteriorly than outer pair.
Supraorbital borders each with two triangular or rounded lobes. Postorbital angles conspicuous, typically with
long, pointed tips. Cornea of eyes dorsoventrally flattened, wider than base of eye peduncle, or spherical and as
wide as base of peduncle; peduncle with granular tubercles at least on distal border. Orbits deep.
Suborbital borders each with two broadly rounded or rectangular lobes. Pterygostomial lobe at each anterolateral
angle of buccal frame often projects ventrally.
Each basal antennal segment rectangular, usually slender, without distal expansion. Epistome narrow, not
expanded dorsoventrally, vertically inclined. Border of epistome with two median teeth (plus two triangular outer
processes in most species) connecting distally with each pterygostomial lobe. Meri of third maxillipeds smaller
and narrower than ischia.
Chelipeds increasingly unequal with increasing size (particularly in males), smallest typically more slender in
females. Fingers of larger cheliped with cutting edge, sometimes with broadly rounded teeth; those of smaller
cheliped with short, triangular teeth, pollex becoming flatter with increasing size. First pair of walking legs (P2)
shorter, more slender than second and third pairs (P3-4); third pair slightly shorter or as long as second. First three
pairs of walking legs (P2-4) with flattened carpi, propodi, and (with one exception) dactyli; anterior (dorsal) and
posterior (ventral) borders of meri with thick tubercles or triangular teeth, often rows of tubercles on dorsal and
ventral surfaces; posterior border of dactyli smooth or armed with teeth or spines. Last two pairs of walking legs
(P3-4) each with broad coxae, having laminar, thin-edged anterior and posterior borders. Fifth pair of pereopods
(P5) reduced (0.6-0. 9 CL), slender; basis-ischia and meri often short, tuberculate; one row of small spines along
posterior border of dactyli.
Abdomen of mature males elongate, with both sides almost parallel to each other, with all segments free,
segments 1-2 dorsoventrally compressed, one transverse ridge each along some or all segments. Male first pleopods
long, slender; basal parts straight or sinuous but not helicoidal; each distal part biramous (with two typically
flattened processes) or uniramous (with simple tip). Second male pleopods short, thin, slightly curved; distal
segment with blunt tip.
Abdomen of mature females broad, rounded, with all segments free, segments 1-2 dorsoventrally compressed,
one transverse ridge each along segments 1-4. Abdomen of immature females triangular (but broader than
in males), segments 1-2 dorsoventrally compressed, 3-5 or 3-6 fused, transverse ridge along each segment 1-3
or 1-4.
450
P. CASTRO
SPECIES included. — There are 1 1 known species of Pseudopalicus , by alphabetical order: P. acanthodactylus
sp. nov., P. amadaibai (Sakai, 1963), P. declivis sp. nov., P. glaber sp. nov., P. investigatoris (Alcock, 1900),
P. macromeles sp. nov., P. oahuensis (Rathbun, 1906), P. pictus sp. nov., P. serripes (Alcock & Anderson,
1895), P. sexlobatus (Kensley, 1969), P. undulatus sp. nov. A possible new species, being referred to as
Pseudopalicus sp., is known from only one incomplete specimen.
Sexual Dimorphism. — There is a considerable difference in size between males and females, with females
being larger. The chelipeds are unequal in size in both sexes but the difference is more apparent in males. The
smaller cheliped is usually more slender in females. There is no evidence of differences between the sexes in the
length or shape of the walking legs or fifth pair of pereopods.
Discussion. — Pseudopalicus was established by Moosa & Serene (1981) for eight Indo-west Pacific-
species formerly included in Palicus Philippi, 1838. However, they actually examined specimens of only two
species, P. amadaibai (Sakai, 1963) and P. serripes (Alcock & Anderson, 1895). Two of the original eight species,
P.fisheri (Rathbun, 1906) and P. cyrenae (Ward, 1942), are being placed in synonymy with P. investigatoris
(Alcock, 1900). The present definition of the genus has therefore been amended to include characters present in the
12 recognized species, including six new ones, that were examined in this study.
The shape of the carapace and walking legs of Pseudopalicus agree with the same characters of group 1 (or "typ¬
ical" group) as defined by Rathbun (1918: 184) for six western Atlantic and eastern Pacific species o (Palicus.
Four of these species have four anterior lobes as in Pseudopalicus : Palicus affinis A. Milne Edwards & Bouvier,
1899. P. alternants Rathbun, 1897, P. faxoni Rathbun, 1897, and P. lucasii Rathbun, 1898. They all have two
anterolateral teeth on each side of the carapace, however, unlike the four typical of Pseudopalicus. None of these
species seem to have the convex carapace with conspicuous bosses typical of Pseudopalicus (see Rathbun, 1918;
Williams, 1984; Melo, 1996). P. velerae (Garth, 1939) from the eastern Pacific has a carapace with four anterior
lobes, five anterolateral teeth on each side, and short walking legs similar to Pseudopalicus , but prominent epister-
nal processes as in Parapalicus. It perhaps belongs to group 4 of Rathbun on account of the presence of conspic¬
uous episternal processes. There are also similarities with the three western Atlantic and eastern Pacific species in
group 5 of Rathbun (1918: 184): P. dentatus (A. Milne Edwards, 1880), P. obesus (A. Milne Edwards. 1880).
and P. tuberculatus (Faxon, 1893). These species have four anterior lobes, convex carapace, conspicuous bosses,
and short walking legs. They differ from Pseudopalicus in having two or three anterolateral teeth (see RATHBUN,
1918; MELO, 1996). There is unfortunately incomplete information on the morphology of the abdomen and no
published information on the male first pleopods of these western Atlantic and eastern Pacific species.
An unidentified palicid (USNM) from off the coast of northern Peru in the eastern Pacific had a carapace very
similar to that of Pseudopalicus, with four anterior lobes, well-developed bosses, four anterolateral teeth on each
side, and two supraorbital and two suborbital lobes on each side. The male first pleopods, however, had long,
helicoidal basal parts and each distal part consisting of a rounded, Hat, proximal process and a very long, slightly-
curved, pointed process, quite unlike the male pleopods of Pseudopalicus species. Furthermore, the epistome lacked
teeth, each basal antennal segment had a flat, scale-like, distal extension, the fifth pereopods were thicker than
those in Pseudopalicus, and the fingers of the chelipeds were much longer than in any species of Pseudopalicus.
There is very little information on the male pleopods of the Atlantic and eastern Pacific species of palicids. The
male first pleopods of Palicus caronii (Roux, 1830) of the Mediterranean and eastern Atlantic were illustrated by
Monod (1956, figs 549-551). The pleopods of a specimen from an unknown location (SMF 4712) had sinuous
basal parts and hatchet-like, uniramous distal parts. The male first pleopods of Palicus alternants Rathbun, 1897
were described by Williams (1984: 482) as "stout and twisted, tip bilobed, inner lobe thinner and longer than
outer" and "weaker, not twisted, and tip less spreading".
The elongate male abdomen with all segments free suggests a less specialized, and perhaps more plesiomorphic
condition. The morphologies of the male first pleopods differ strikingly, although they share a sinuous, non-
helicoidal basal part. The distal part can be uniramous or of several biramous patterns. Species that share a general
body type and many other characters have very dissimilar male pleopods. The significance of this remains
unknown. It may be suggested that the group is speciating rapidly. Another possibility is that Pseudopalicus may
not be a natural group after all. The uniramous male pleopods of P. investigatoris, P. serripes, and P. pictus
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
451
sp. nov., for example, sets these species apart from their congeners. The three species, plus an undescribed species
close to P. serripes, also share a squarish carapace and conspicuous teeth on the pterygostomial ridge.
Key to the species of PSEU DO PALI CU S
1. Anterolateral border of carapace with four typically pointed teeth (last tooth may be very
small) . 2
— Anterolateral border of carapace with three rounded teeth (last tooth may be very small) 1 0
2. Dactyli of walking legs (P2-4) with teeth or spines along posterior border . 3
— Dactyli of walking legs (P2-4) entire, without teeth or spines along posterior border . 8
3. Suborbital border with one triangular or pointed tooth on inner edge. One or two sharp
teeth on proximal portion of ridge that connects pterygostomial lobe with pterygostomial
region of carapace (Figs 9a-b, 11b) . 4
— Suborbital border with one rectangular or rounded lobe on inner edge. No sharp teeth on
proximal portion of ridge that connects pterygostomial lobe with pterygostomial region of
carapace . 6
4. Dorsal surface of carapace with few, low, granular bosses. Male first pleopods with thick
distal parts that are sharply bent distally (Figs 1 ld-e) . P. pictus sp. nov.
— Dorsal surface of carapace with conspicuous bosses. 10 of which cross branchial and
metagastric regions. Male first pleopods with simple, straight distal parts (Figs 4b-d) ... 5
5. Dense cluster of setae on inner surface of chcliped propodi of males. Supraorbital border
with lobes having rounded tips (Fig. 12c). Dactyli of walking legs broad (P4 0.2-0.3 CL)
. P. serripes (Alcock & Anderson, 1895)
— Inner surface of cheliped propodi of males smooth. Supraorbital border with lobes having
narrow, pointed tips (Fig. 12b). Dactyli of walking legs slender (P4 0. 5-0.6 CL) .
. P. investigatoris (Alcock. 1900)
6. Posterior third of carapace smooth, without conspicuous, high bosses. Posterior border of
carapace entire, without tubercles (Fig. 5d) . P. glaber sp. nov.
— Posterior third of carapace with conspicuous, high bosses. Posterior border of carapace
with six low tubercles . 7
7. Two very short supraorbital lobes (Fig. 3a) . P. acanthodactylus sp. nov.
— Two conspicuous supraorbital lobes with rounded tips (Fig. 5b) .
. P. amadaibai (Sakai, 1963)
8. Second, pointed anterolateral tooth on side of carapace larger than first, which has rounded
or lobate borders (Fig. 12a). Upper and lower borders of meri of walking legs (P2-4) with
thin, pointed, nearly equal tubercles . P. oahuensis (Rathbun. 1906)
— Anterolateral teeth gradually decrease in size posteriorly. Upper and lower borders of meri
of walking legs (P2-4) with tubercles of different sizes and shapes . 9
9. Male first pleopods with distal part consisting of two processes, one broad and one thin,
armed with minute teeth (Fig. 6d-e) . P. declivis sp. nov.
— Male first pleopods with distal part uniramous and teeth along inner and outer borders
(KENSLEY, 1969, fig. 2d) . P. sexlobalus Kensley, 1969
10. Posterior border of carapace with 4-6 rounded, granular tubercles. Supraorbital border with
two triangular lobes (Fig. 13a) . P. macromeles sp. nov.
— Posterior border of carapace with six low, elongate tubercles (Figs 12f. 14a). Supraorbital
border with two short lobes, inner rectangular, outer rounded . P. undulatus sp. nov.
Source : MNHN, Paris
452
P. CASTRO
Pseudopalicus acanthodactylus sp. nov.
Figs 3, 5a, 55. 60a
MATERIAL EXAMINED, — New Caledonia. CHALCAL 2: stn CC 2. 24°55.48'S, 168°21.29'E. 500 m, 28.10,1986:
1 <3,2 2 (MNHN-B 26861); 1 2 (MNHN-B 26862). — Stn CP 21, 24°54.00'S. 168°21 ,61'E, 500 m, 28.10.1986: 1 2
(MNHN-B 26863). — Stn DW 72, 24°54.5'S, !68°22.3,E, 527 m, 28.10.1986: 3 2 (MNHN-B 26864). — Stn DW 74,
24°40.36'S, 168°38.38'E. 650 m, 29.10.1986: 1 2 (MNHN-B 26676). — Stn DW 75, 24°39.31'S, 168°39.67'E. 600 m.
29.10.1986: 2 <3 paratypes 6.9 x 9.0 mm, 7.3 x 9.7 mm (MNHN-B 26675), 1 2 (MNHN-B 26865). — Sin DW 76.
23°40.5'S, 167°45.2’E. 470 m, 30.10.1986: 2 3 , 5 2 (MNHN-B 26866). — Stn DW 77, 23°38.35'S. I67°42.68'E.
435 m. 30.10.1986: 3 2 (MNHN-B 26867).
Smib 3: stn DW 6, 24°56'S, 168°21'E, 505 m. 21.05.1987: 1 <3 . 4 2 (MNHN-B 26868). — Stn DW 9, 24°42'S.
168°08'E, 265 m. 21.05.1987: 1 3 (MNHN-B 26869). — Stn DW 12, 23°37.7'S, I67°41.5'E. 470 m. 22.05.1987: I <3,
2 2 (MNHN-B 26870). — Stn DW 13, 23°37.5'S, 167°4I.6'E, 448 m, 22.05.1987: \ 3,2 9 (MNHN-B 26871).
Smib 4: stn DW 58, 22°59.8'S, 167°24.2'E, 560 m. 9.03.1989: I 2 paratype 10.6 x 13.0 mm (MNHN-B 26674).
BERYX 11: stn CP 7, 24°55'S, 168°21'E. 510-550 m, 15.10.1992: 1 2 (MNHN-B 26872). — Stn CP 8. 24°54'S.
1 68°21’E. 540-570 m, 15.10.1992: 1 3 (MNHN-B 26873). — Stn CP 10, 24°53'S, 168°21'E. 565-600 m, 15.10.1992:
2 2 (MNHN-B 26874). — Stn DW 27, 23°37'S. 167041'E, 460-470 m, 18.10.1992: 1 2 paratype 12.3 x 14.9 mm
(MNHN-B 26673), 2 3,2 2 (MNHN-B 26737). — Stn CP 32, 23°38'S. 167°43'E. 420-460 m. 18. 10. 1992: 1 2 (MNHN-
B 26875).
Smib 8: stn DW 146, 24°55.2'S, 168°21.7'E, 514-522 m, 27.01.1993: 1 holotype 3 8.9 x 10.5 mm (MNHN-B
26672). — Stn DW 147, 24°54.9'S, 168°21.8’E, 508-532 m, 27.01.1993: 1 2 9.6 x 13.5 mm (MNHN-B 26876). —
Stn DW 150. 24°54.3'S, 168°22.2'E, 519-530 m, 27.01.1993: 1 3 , 1 2 (MNHN-B 26877). — Stn DW 152. 24°54.3'S.
168°22.2'E, 514-530 m, 27.01.1993: 1 2 (MNHN-B 26878). — Stn DW 166. 23°37.8'S, 167°42.7'E. 433-450 m.
29.01.1993: 3 2 (MNHN-B 26879). — Stn DW 167, 23°38.rS, 168043.1'E. 430-452 m, 29.01.1993: I <3,32
(MNHN-B 26880). 1 2 13.8 x 16.7 mm (MNHN-B 26738). — Stn DW 168, 23°37.7'S, 168°42.5'E. 433-450 m.
29.01.1993: 1 3 , 5 2 (MNHN-B 26881). — Stn DW 169, 23°37.7'S, I67°42.5'E. 447-450 m. 29.01.1993: 4 2
(MNHN-B 26882).
Smib 10: stn DW 202. 24°55'S, 168°22'E, 525-513 m, 10.01.1995: 1 2 (MNHN-B 26883).
Lithist: stn CP 2, 23°37.07'S, 167°41.14'E, 442 m, 10.08.1999: 3 2,12 (MNHN-B 26969). — Stn CP 3,
23°37.01'S, 1 67°49’E. 447 m, 10.08.1999: 1 3 , 3 2 (MNHN-B 26967). — Stn DW 5. 23°38.25’S, 167°42.94'E, 433-
500 m, 11.08.1999: 8 2 (MNHN-B 26968), 1 <3 paratype 8.4 x 10.3 mm, 1 2 paratype 12.0 x 14.8 mm (USNM). 1 3
paratype 8.4 x 10.4 mm, 1 2 paratype 11.8 x 14.4 mm (ZRC 1999.1425-1426). — Stn CP 8. 24°54.24'S. 168°2I.35'E.
540 m, 11.08.1999: 1 2 (MNHN-B 26970). — Stn CP 11, 24°46.69'S, 168°08.33'E, 254-283 m, 11.08.1999: 1 <3 , 1 2
(MNHN-B 26971).
Types. — Holotype : 1 6 8.9 x 10.5 mm, Smib 8, stn DW 146 (MNHN-B 26672).
Paratypes'. IS 6.9 x 9.0 mm, 7.3 x 9.7 mm, CHALCAL 2, stn DW 75 (MNHN-B 26675); 1 2 10.6 x
13.0 mm, Smib 4, stn DW 58 (MNHN-B 26674); I 2 12.3 x 14.9 mm, BERYX 11, sin DW 27 (MNHN-B
26673); 1 S 8.4 x 10.3 mm, 1 2 12.0 x 14.8 mm. LlTHlST, stn DW 5 (USNM); 1 <3 8.4 x 10.4 mm, 1 2
1 1.8 x 14.4 mm, Lithist, stn DW 5 (ZRC 1999.1425-1426).
Type Locality. — Norfolk Ridge, south of New Caledonia, 24°55.2'S, 168°21.7'E, 514-522 m.
Diagnosis. — Carapace (Figs 3a, 5a, 60a) with four anterolateral teeth on each side, first tooth usually
rounded, blunt, second and third larger, pointed, and fourth very small. Dorsal surface of carapace with medium-size
bosses, 12 of which form an almost straight row across branchial and metagastric regions in larger specimens;
posterior border with six low, elongate tubercles. Supraorbital borders each with two very short lobes. Suborbital
borders (Fig. 3b) each with one broad, square outer lobe; one higher, narrower, rounder inner lobe. Outer surfaces
of cheliped propodi each with 2-3 rows of blunt tubercles. Dactyli of walking legs (P2-4) with short, thick spines
along posterior border (2-4 spines in P4; Fig. 3c). Abdomen of mature males (Fig. 3d) with all segments free, one
complete transverse ridge along each segment 1-3, incomplete ridge along median portion of segment 4. Male first
pleopods (Figs 3e-f) with sinuous basal parts; each distal portion with broad ventral process having small teeth,
two dorsal, wing-like extensions. Abdomen of mature females with all segments free (Fig. 3g), one complete
transverse ridge along each segment 1 -4.
Source : MNHN, Pans
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
453
Color: The carapace of the live male holotype (Fig. 60a) had a pattern of irregular, very fine yellow blotches on
a red-brown background. The same pattern was repeated on the chelipeds and walking legs. The distalmost tubercle
on each eye peduncles was red, color still visible in specimens that had been preserved for one month.
Fig. 3. — Pseudopalicus acanihodactylus sp. nov.: a-f, 3 holotype 8.9 x 10.5 mm. Norfolk Ridge, south of
New Caledonia. Smib 8, stn DW 146, 514-522 m (MNHN-B 26672): a. carapace, dorsal surface; b. suborbital border;
c. dactylus, right fourth pereopod, dorsal view; d, abdomen; e. left male first pleopod, ventral view; f. left male first
pleopod. distal part, lateral (inner side) view. — g. $ paratype 10.6 x 13.0 mm. Norfolk Ridge, south of
New Caledonia, Smib 4, stn DW 58, 560 m (MNHN-B 26674): abdomen.
454
P. CASTRO
Description. — Carapace (Figs 3a, 5a, 60a) transversely hexagonal, wider than long (CW/CL = 1.2- 1.3);
dorsal surface covered with coarse granules and rows of conspicuous granular bosses. Confluence of branchial and
mesogastric regions depressed, granular, usually darker in color. Anterolateral borders of carapace each with four
anterolateral teeth, first usually rounded, blunt, second and third larger, pointed, and fourth very small. Dorsal
surface of carapace with medium-size bosses, 12 of which form an almost straight row across branchial and
metagastric regions in larger specimens (up to 14, not in straight row, in smaller specimens). Posterior border
with six low, elongate tubercles (almost continuous in larger specimens) and no plumose setae.
Frontal border of carapace divided into four narrow, rounded lobes, inner lobes longer and typically more
pointed, particularly in larger specimens. Borders between frontal lobes and supraorbital borders anteriorly rounded
(slightly pointed in some specimens), folded upward slightly, and ending in sharp angle, forming very shallow, U-
or L- shaped fissure before supraorbital border. Supraorbital borders each with two very short, nearly straight, or
slightly rounded lobes. Postorbital angles short (not extending beyond dorsal border of retracted eye), slightly
pointed inward. Cornea of eyes dorsoventrally flattened, wider than base of short eye peduncle; each peduncle with
two flat, granular tubercles on distal border.
Suborbital borders (Fig. 3b) each with one broad, square outer lobe; one higher, narrower, rounder inner lobe.
Ptcrygostomial lobes project ventrally, forming flat, semicircular structure posterior to each inner suborbital lobe.
Each basal antennal segment slender, rectangular; flagellum long, with lew, simple setae. Epistome vertically
inclined, with two broad (rounded or pointed), flat median teeth, each flanked by narrow, pointed outer process and
thin, rounded margin before pterygostomial lobe. Inner border of ischia of third maxillipeds straight; surface
coarsely granular, upper borders rounded. Meri much narrower than ischia; straight-edged, triangular upper borders.
Dorsal and outer borders of cheliped propodi with rows of rounded, sometimes elongate tubercles. Fingers of
largest cheliped with cutting edges (rounded teeth in small specimens), 4-8 rounded to triangular teeth in smallest
cheliped. Carpi short, outer borders with low,' rounded tubercles; meri slender, outer borders with low, rounded
tubercles.
First three pairs of walking legs (P2-4; Fig. 5a, 60a) flattened. Upper and lower borders of meri with tubercles
of different sizes and shapes (some pointed, some rounded; smaller but on two anterior rows in P2); distalmost
tubercle on each anterior border much wider at base, much higher, slender, pointed, directed distally (only in P2-3
in some small specimens). Anterior borders of carpi tuberculate; borders of propodi entire. Each P2 with one
dorsal, one ventral carina on carpus; two dorsal, two ventral carinae on propodus; one dorsal, two ventral carinac,
0-3 spines along posterior border of dactylus. Each P3 with one dorsal, one ventral row of tubercles on merus; two
dorsal, two ventral carinae on carpus; one dorsal, two ventral carinae on propodus; one dorsal, two ventral carinae,
0-4 spines along posterior border of dactylus. Each P4 with two dorsal, one ventral rows of tubercles on merus
(absent in some); two dorsal, two ventral carinae on each carpus and propodus; one dorsal, two ventral carinae, 2-4
short, thick spines along posterior border of dactylus (Fig. 3c). Meri of all walking legs with scattered plumose
setae; dorsal surfaces of propodi and dactyli of P3-4 each with one row of conspicuous plumose setae along anterior
border.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair slightly
shorter than second. Fifth pair of pereopods (P5) short (0.7-0.8 CL); each merus wide in larger specimens (slender
in smaller specimens), tuberculate surface, 4-5 triangular tubercles (proximal largest) along posterior border, and
scattered plumose setae; each propodus with low tubercles (2-4 short spines in largest specimens) along posterior
border; each dactylus with row of 1 1-13 short, thick spines along posterior border, 2-3 distally on anterior border,
one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature females (Fig. 3g) with all segments free. One transversal ridge along each segment 1-4.
Source : MNHN, Pahs
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
455
Abdomen of mature males (Fig. 3d) with all segments free. One complete transverse ridge along each segment
1-3, incomplete ridge along median portion of segment 4. Male first pleopods (Fig. 3e-f) with sinuous basal parts;
each distal part with broad ventral process with small teeth and two dorsal, wing-like extensions.
DISCUSSION. — P. acanthodactylus can be easily distinguished from congeners by its short supraorbital lobes,
which make the eyes seem prominent, and the short, thick spines on the posterior border of the dactyli of the
walking legs (P2-4; Fig. 3c).
SIZE. — Maximum size among specimens examined: 13.8 x 16.7 mm (female, MNHN-B 26738); 9.4 x
1 1 .5 mm (male, MNHN-B 26737).
ETYMOLOGY. — From akantha, Greek for spine, and daktylos. Greek for finger, in reference to the diagnostic
spiny dactyli of the walking legs (P2-4).
DISTRIBUTION. — Known only from the Norfolk Ridge south of New Caledonia (Fig. 55). Depth: 265-650 m.
Pseudopalicus amadaibai (Sakai, 1963)
Figs 4a, 5b, 53
P aliens oahuensis - Balss, 1922: 120, fig. 6. — Yokoya, 1933: 206, 212 [non Palicus oahuensis Rathbun, 1906],
Cymopolia oahuensis - Sakai, 1939: 609, fig. 90a; 1956: 52. — Nakazawa & Sakai. 1947: 665. unnumb. fig. [non
Palicus oahuensis Rathbun, 1906).
Palicus amadaibai Sakai, 1963: 227, fig. 7a. — Sakai, 1965: 183, pi. 89, fig. 1; 1976: 593, pi. 205. fig. 1. — Serene,
1968: 96. — Takeda & Miyake. 1969: 464. — Muraoka, 1998: 49.
Pseudopalicus amadaibai - Moosa & SerEne, 1981: 39, fig. 6. pi. 2. fig. B. — Takeda, 1982a: 205, fig. 608.
MATERIAL EXAMINED. — Japan. Tosa Bay. Mimase, 1959, dry specimen with red tag, possible pari of lype
material: 1 2 14.2 x 17.0 mm (SMF 24698). — Kochi, Mimase, K. Sakai coll.: 1 3 12.6 x 14.5 mm (KPM-NH 107132).
— K. Sakai coll., 15.04.1963: 1 9 15.5 x 18.6 mm (SMF 24699); 20.03.1964: 1 9 (SMF 24700); 10.1961-04.1963:
1 3 (SMF 24701).
Sagami Bay. Okinoyama Bank. 34°59'N, 139°39'E. 105-113 m, coarse sand and shell fragments, T. KOMAI coll..
21.04.1995: 1 9 (CBM-ZC 2004).
Kii Peninsula. Off Shionomisaki, 33°26.I3'N, 135°40.64'E, 160 m, S. Nagai coll., 16.01.1997: 1 9 (CBM-ZC
3602); 33°26.72'N, 135°29.20'E, 180 m: 1 juv. 3 (CBM-ZC 4953). — Off Kirimezaki, 150-180 m. H. Ikeda coll..
09.1981: 1 3 (HSM). — Kii Strait, trawl, M. Ozaki coll, from Sakaihama fish market: I 3 , 1 9 (SMF 24702).
Goto Islands. "Sovo Man,": stn 440, 152 m. 19.07.1929: 1 3 (KMNH).
No location. 1 $ (SMF 24703). — 1 2 (SMF 24704). — 1 9 (SMF 24705). — 1 2 (SMF 24706).
Philippine Islands. Verde Island Passage. MUSORSTOM 2: stn DG 32, 13°40'N, 120°54'E. 192-220 m.
24.11.1980: 1 3 12.1 x 14.7 mm (MNHN-B 26804).
Indonesia. Tanimbar Islands. Karubar: stn DW 49. 08°00'S, I32°59’E, 206-209 m, 29.10.1991: 1 3 (MNHN-B
26739). — Stn CP 86. 09°26'S. 131°13’E, 223-225 m. 4.11.1991: 1 2 12.3 x 14.5 mm (MNHN-B 26828).
New Caledonia. Bathus 1: stn DW 690, 20°32.94'S. 165°00.83'E, 352 m, 16.03.1993: 1 juv. 3 (MNHN-B
26815).
Fiji. Bordau 1: stn DW 1450. 16°44'S, 179°58’E, 327-420 m, 4.03.1999: 1 3 , 1 2 (MNHN-B 27133).
Types. — Holotype of Palicus amadaibai Sakai, 1963: I 3 12x15 mm (Sakai, 1963), Emperor of Japan
coll. (Showa Memorial Institute, National Science Museum, Tsukuba. Japan; NSMT-Cr R: 2506).
Allotype: 1 2 , same locality as holotype. Deposit unknown.
Paratypes: [6,29, west of Jyogashima light, Sagami Bay, Japan, 65-85 m, Emperor of Japan coll. —
2 6 , 1 9 , Tosa Bay, Japan, K. Sakai coll. Deposit unknown.
Possible paratype: 1 9 14.2 x 17.0 mm, Tosa Bay, Japan (SMF 24698).
TYPE Locality. — Amadaiba fishing grounds off Hayama (Kannonzuka-dashi. Aoyama-dashi), Sagami Bay,
Japan, 65-85 m.
456
P. CASTRO
DIAGNOSIS. — Carapace (Fig. 5b) with lour broad, slightly pointed anterolateral teeth on each side; teeth
decrease in size posteriorly. Dorsal surface of carapace with large granular bosses, 12 of which form an almost
straight row across branchial and mctagastric regions in large specimens; posterior border with six low, elongate
tubercles (almost continuous in larger specimens). Supraorbital borders each with two rounded lobes. Suborbital
borders each with two broad, slightly rounded to straight-edged lobes; inner border slightly higher than outer.
Dorsal borders of cheliped propodi each with row of thin, elongate, rounded tubercles; outer surface with scattered
blunt tubercles. Dactyli of walking legs (P2-4) with short, thick spines along posterior border (2-3 in P4).
Abdomen of mature males with all segments free, one complete transverse ridge along each segment 1-3,
incomplete ridge along median portion of segment 4. Male first pleopods (Fig. 4a; MOOSA & SERENE, 1981.
fig. 6) with sinuous basal parts; each distal portion with four distal tips (three thick, mammillate; one slender
with spinules). Abdomen of mature females with all segments free, one complete transverse ridge along each
segment 1-4 (very low ridge may be present along segment 5).
Fig. 4. — a , Pseudopalicus amadaibai (Sakai, 1963), 3 12.1 x 14.7 mm, Verde Island Passage, Philippine Islands.
Musorstom 2, stn DG 32, 192-220 m (MNHN-B 26804): left male first pleopod, ventral view. — b-e. Pseudopalicus
invesiigatoris (Alcock. 1900), 3 10.3 x 12.0 mm. South China Sea, Philippine Islands, Musorstom 1, stn 57.
107-96 m (MNHN-B 26805): b, left male first pleopod. ventral view; c, left male first pleopod, distal part, dorsal
view. — d. Pseudopalicus serripes (Alcock & Anderson, 1895). 3 10.4 x 11.0 mm, Seychelles, Reves 2, stn 21,
55-60 m (MNHN-B 26750): left male first pleopod, lateral (inner side) view. — e-f. Pseudopalicus oahuensis
tRathbun, 1906), 3 7.8 x 10.4 mm. Norfolk Ridge, south of New Caledonia. Smib 4. DW 56. 260 m (MNHN-B
26806): e. left male first pleopod, lateral (outer side) view; f. left male first pleopod, distal part, ventral view.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
457
Fig. 5. — a. Pseudopalicus acanthodactylus sp. nov., 2 9.6 x 13.5 mm, Norfolk Ridge, souih of New Caledonia.
CHALCAL 2, stn DW 74, 650 m (MNHN-B 26676): dorsal view. — b, Pseudopalicus amadaibai (Sakai. 1963):
2 12.3 x 14.5 mm, Tanimbar Islands, Indonesia. Karubar, sin CP 86. 223-225 m (MNHN-B 26828): dorsal view. —
c, Pseudopalicus declivis sp. nov., 2 paratype 15.5 x 18.1 mm. Grand Passage, north of New Caledonia. MUSORSTOM
4, stn DW 196, 450 m (MNHN-B 26680): dorsal view. — A, Pseudopalicus glaber sp. nov., 2 paratype 9.8 x
12.3 mm, Norfolk Ridge, north of New Caledonia, Smib 5, stn DW 103. 315 m (MNHN-B 26687): dorsal view.
Discussion. — Comparison of material collected throughout the western Pacific with material from the type
locality, central Japan (including material examined by T. Sakai and most probably some of his type material),
confirms previous characterizations of the species. Sakai (1963) named his new species after the Amadaiba fishing
grounds in Sagami Bay, Japan, but erroneously used "amadaibai" instead of the correct "amadaibaiensis" to name
the species.
MOOSA & SERENE (1981) noted that no mention was made in Sakai's description of the presence of spines
along the posterior border of the walking legs, even though they were shown in Sakai (1963, fig. 7a). It is
assumed that Moosa & Serene were referring specifically to the dactyli of the walking legs, which Sakai (1963:
228) described as having "two or three denticles".
Other characteristics of P. amadaibai include a carapace with a frontal border that is divided into four narrow and
rounded lobes (inner lobes are more pointed in large specimens), followed by a broad, rounded, and folded-upward
lobe before the supraorbital border. The postorbital angles are short (not extending beyond the dorsal border of the
retracted eye) and only slightly pointed inward. Each basal antennal segment is slender and rectangular (but slightly
expanded along the outer border in large specimens) and slightly tuberculatc. Each eye peduncle has two low.
458
P. CASTRO
granular tubercles on the distal border and, in alcohol-preserved specimens, large red-orange spots ventrally. The
pterygostomial lobes project ventrally, forming a flat, semicircular structure (triangular when seen ventrally)
posterior to each inner suborbital lobe. The epistome is vertically inclined, with two short, rounded median teeth
flanked on each side by a triangular outer process and a thin, rounded margin before joining pterygostomial lobe.
The third pair of walking legs (P4) have slender meri (0.4-0. 5 CL; Fig. 5b) each armed with one row of teeth
along upper border and one conspicuously pointed distal tooth; the dactyli are slender (0.3 CL), each armed with 2-
3 microscopic teeth along the posterior border. The fifth pair of pcreopods (P5) are short (0.8 CL), each merus is
smooth or with several low tubercles along the posterior border, and dactylus with a row of 4-5 thick spines along
the posterior border, 1 -3 distally on the anterior border.
P. amadaibai was initially confused with P. oaliuensis (Rathbun, 1906) (Balss, 1922; Yokoya, 1933;
Sakai, 1939; 1956; Nakazawa & Sakai. 1947) but their resemblance is only superficial. In addition to the
clear differences between the male first pleopods, the dactyli of the walking legs (P2-4) are unarmed in P.
oaliuensis but armed in P. amadaibai.
Moosa & Serene (1981) pointed out rather superficial similarities between P. amadaibai and
P. investigatoris, which unfortunately they did not examine. P. investigators can be easily distinguished by its
very pointed anterolateral teeth, presence of a pointed tooth on the inner border of the suborbital border, and very
simple male first pleopods (see discussion for P. investigatoris below).
The figure of the male first pleopods given by Moosa & Serene (1981. fig. 6) shows three thick and pointed
processes, two being of the same size, with the third smaller. Most of the specimens examined in this study only
had two equal processes (Fig. 4a).
An unusual abnormality, the presence of one inner lobe along the anterior border of the carapace instead of the
normal two (for a total of three lobes instead of four), was observed in a male from the Tanimbar Islands, Indonesia
(MNHN-B 26739). It was probably due to damage and subsequent growth.
Size. — Maximum size among specimens examined: 15.5 x 18.6 mm (female, SMF 24699); 12.6 x 14.5 mm
(male, KPM-NH 107132).
Distribution. — Japan (Balss, 1922; Yokoya, 1933; Sakai, 1939. 1956, 1963, 1965, 1976; Nakazawa
& Sakai, 1947; Takeda & Miyake, 1969; Takeda. 1982a; Muraoka, 1998), and the Philippine Islands
(Moosa & SerLne, 1981). It is here recorded for the first time from the Arafura Sea (Indonesia). New Caledonia
and Fiji (Fig. 53). Sakai (1976) recorded it from soft bottoms. Depth: known reliably between 65 m (Sakai.
1963) and 352 m, also collected in a trawl between 327-420 m.
Pseudopalicus declivis sp. nov.
Figs 5c. 6. 55
Material EXAMINED. — Philippine Islands. South China Sea. Musorstom 2: sin CP 19. 14°00'N, 120°16'E,
189-192 m, 22.11.1980: \ 6.2 9 (MNHN-B 26740).
Indonesia. Kai Islands. Karubar: stn DW 18, 05°18'S, 133°OI'E, 205-212 m, 24.10.1991: 1 juv. S (MNHN-B
26884).
Chesterfield Islands. Musorstom 5: stn 379. 19°53.20'S. 158°39.50'E, 370-400 m, 20.10.1986: 1 6 paratype
10.2 x 12.0 mm (MNHN-B 26678).
Vanuatu. Musorstom 8: stn CP 1024, 17°48.21'S, 168°38.77'E, 335-370 m, 28.09.1994: 1 2 (MNHN-B 26885).
New Caledonia. "Vauban" Dredgings 1978: stn D3, 22°17'S, 167°12'E, 390 m, 23.05.1978: 1 2 (MNHN-B
26886). — Stn D4, 22°17'S, 167°13'E, 400 m. 23.05.1978: 1 6 (MNHN-B 26887). — Stn DI6. 22°17'S. 167°!2'E,
390 m. 23.05.1978: 1 6 (MNHN-B 26888).
Biocal: stn CP 105, 21°31'S. 166°22’E, 330-335 m, 8.09.1985: 1 6 (MNHN-B 26889). I 6 paratype 10.5 x
12.9 mm, 1 2 paratype 14.5 x 17.8 mm (ZRC 1999.1427-1428).
Musorstom 4: stn CP 180. 18°56'S, 163°17.7'E. 440 m. 18.09.1985: I 6 (MNHN-B 26890). — Stn DW 181.
1 8°57.2’S. 1 63°22.4'E, 350 m, 18.09.1985: 3 2 (MNHN-B 26891). — Stn CP 193, 18°56.3'S. 163°23.2'E, 415 m,
19.09.1985: 2 6 . 3 2 (MNHN-B 26892). — Stn DW 196. 18°55.0'S. 163°23.7'E, 450 m, 20.09.1985: 1 2 paratype
15.5 x 18.1 mm (MNHN-B 26680), 1 3 , 1 2 (MNHN-B 26893). — Stn DW 222, 22°57.6'S, 167°33.0'E. 410-440 m.
30.09.1985: 1 2 paratype 14.2 x 16.3 mm (MNHN-B 26681).
Source :
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
459
Lagon: stn DW 1152, 18°58.2'S, 163°23.9’E, 335 m, shark teeth and stylasterids. 29.10.1989- I 3 (MNHN-B
26894).
Smib 6: stn DW 120, 18°58.5'S, 163°25.6'E, 310-325 m, sponges and stylasterids, 3.03.1990: 1 2 paratype 11 5 x
12.6 mm (MNHN-B 26682). F yv
Smib 8: stn DW 189, 23°17.6'S. 168°05.5'E, 400-402 m, 31.01.1993: 1 $ (MNHN-B 26895)
Bathus 1: stn CP 701, 20°57.54'S. 165°35.86'E, 302-335 m, 18.03.1993: 1 <5,32 (MNHN-B 26896). I <5
paratype 10.4 x 12.6 mm, 1 2 paratype 11.4 x 14.1 mm (USNM). — Stn CP 710, 21°43.16'S, 166°36.35'E 320-386 m
19.03.1993: 1 2 (MNHN-B 26897).
Bathus 2: stn DW 718, 22°46.70'S, 167°14.45'E. 430-436 m. 11.05.1993: 1 <5 (MNHN-B 26898). — Stn DW 729.
22C52.42'S, 167°1 1.90'E, 400 m, 12.05.1993: 1 <5 (MNHN-B 26899). — Stn DW 730, 23°02.56'S, 166°58.30'E. 397-
400 m, 12.05.1993: 2 3 , 1 2 (MNHN-B 26900). — Stn 737. 23°03.42'S. 166°59.97’E, 350-400 m, 13.05 1993- I <5
12.4 x 14.7 mm (MNHN-B 26741).
Bathus 3: stn DW 827, 23°22'S, 167°01'E, 381-469 m, sand and gravel. 29.1 1.1993: 1 <J holotype 11.1 x 13.4 mm
(MNHN-B 26677), — Stn CP 847, 23°03'S, 166°58'E, 405-411 m. 1.12.1993: 3 2 (MNHN-B 26901)
Bathus 4: stn DW 894, 20°15.77'S, 163°52.03'E, 245-268 m, 3.08.1994: 1 <5 (MNHN-B 26902). — Stn DW 925.
1 8°54.55'S, 1 63°23.75'E, 370-405 m, 7.08.1994: 1 <5 , 1 2 (MNHN-B 26903). — Stn DW 926, I8°56.80'S.
1 63°25.36'E. 325-330 m. 7.08.1994: 2 3 , 1 2 (MNHN-B 26904). — Stn DW 927. I8°55 48'S 1 63°'>'> I I'E 452-
444 m, 7.08.1994: 1 3 (MNHN-B 26905).
Halical I: stn DW I. 18°54.96'S, J63°24'E, 380-400 m. 23.1 1.1994: 1 <5 (MNHN-B 26906). — Stn DW 4. 18°55'S.
163°23.29'E, 350-365 m, 29.11.1994: 1 3 paratype 10.4 x 11.0 mm (MNHN-B 26679).
Loyalty Islands. Musorstom 6: stn DW 406, 20°40.65'S. 167°06.80’E, 373 m. 15.02.1989: I 2 (MNHN-B
26907). — Stn DW 460, 21°01.72'S, 167°31.45'E, 420 m. 20.02.1989: 1 <3 (MNHN-B 26908). — Stn CP 464
21 °02.30'S, 167°31.60'E, 430 m. 21.02.1989: 1 2 (MNHN-B 26909).
types. — Holotype-. 1 <5 1 1.1 X 13.4 mm, Bathus 3, stn DW 827 (MNHN-B 26677).
Paratypes: 1 3 10.2 x 12.0 mm, Musorstom 5, stn 379 (MNHN-B 26678); 1 3 10.4 x 11.0 mm.
Halical 1, stn DE 4 (MNHN-B 26679); 1 2 15.5 x 18.1 mm, Musorstom 4, stn DW 196 (MNHN-B 26680);
19 14.2 x 16.3 mm, MUSORSTOM 4, stn DW 222 (MNHN-B 26681); 1 2, New Caledonia 11.5 x 12.6 mm,
Smib 6, stn DW 120 (MNHN-B 26682); 1 3 10.4 x 12.6 mm, 12 11.4 x 14.1 mm, Bathus 1, stn CP 701
(USNM); Biocal, stn CP 105, I 3 paratype 10.5 x 12.9 mm, 1 2 paratype 14.5 x 17.8 mm (ZRC 1999.
1427-1428).
Type Locality. — Norfolk Ridge, south of New Caledonia, 23°22’S, 167°01'E. 381-469 m, sand and gravel.
DIAGNOSIS. — Carapace (Figs 5c, 6a) with four pointed anterolateral teeth on each side; teeth gradually
decrease in size posteriorly. Dorsal surface of carapace with small granular bosses, 10 of which form an almost
straight row across branchial and metagaslric regions; posterior border with 6-8 low, elongate tubercles.
Supraorbital borders each with two slightly rounded to triangular lobes. Suborbitai borders (Fig. 6b) each with one
square or slightly rounded outer lobe; one slightly higher, rounded, or obliquely-directed inner lobe. Outer surfaces
of cheliped propodi with scattered blunt tubercles of various sizes. Dactyli of walking legs (P2-4) very slender,
with posterior borders entire (Fig. 6c). Abdomen of mature males with all segments free, one complete transverse
ridge along each segment 1-4 (low ridge may be present along median portion of 5-6). Male first pleopods (Figs
6d-e) with sinuous basal parts; each distal part with two processes, inner broad, longer, outer narrow, shorter,
armed with minute teeth. Abdomen of mature females with all segments free, one complete transverse ridge along
each segment 1-4 (low ridge sometimes along segment 5).
Description. — Carapace (Figs 5c, 6a) transversely hexagonal, wider than long (CW/CL = 1. 1-1.2).
Anterolateral borders of carapace each with four pointed anterolateral teeth gradually decreasing in size posteriorly
(last tooth may be missing or greatly reduced in smaller specimens). Dorsal surface of carapace with medium-size,
rounded, granular bosses, 10 of which form an almost straight row across branchial and metagaslric regions.
Posterior border with 6-8 low, elongate tubercles and elongate plumose setae.
Frontal border of carapace divided into four narrow, rounded lobes, inner lobes longer and typically more
pointed (outer lobes always shorter but may be pointed in some specimens). Borders between frontal lobes and
supraorbital borders straight, folded upward, slightly pointed anteriorly in some specimens, and ending in sharp
angle, forming U-shaped fissure before supraorbital border. Supraorbital borders each with iwo large, slightly
rounded to triangular lobes. Postorbital angles short (not extending beyond dorsal border of retracted eye), only
460
P. CASTRO
slightly pointed inward. Cornea of eyes dorsoventrally flattened, wider than base of short eye peduncle; each
peduncle with three granular tubercles on distal border (most ventral tubercle conspicuous, protuberant).
Suborbital borders (Fig. 6b) each with one slightly rounded outer lobe; one slightly higher, narrower,
obliquely-directed inner lobe; wide gap or notch at confluence of outer and inner lobes. Pterygostomial lobes
project vcntrally, forming flat, semicircular (sometimes notched), or pointed structure posterior to each inner
suborbital lobe.
Each basal antennal segment thick, tuberculate, rectangular, slightly expanded distally; flagellum long, with
few, simple setae. Epistome vertically inclined, with two broadly rounded (narrower, triangular in small
specimens), short median teeth flanked on each side by very broad, curved margin connecting directly with
pterygostomial lobe. Inner border of ischia of third maxillipeds straight; surface coarsely granular, upper borders
obliquely cut. Meri much narrower than ischia; upper lobes straight with rounded outer borders.
Outer border of each cheliped propodus with one row of 1-5 thin, rounded tubercles. Fingers of largest cheliped
with cutting edges (rounded teeth in small specimens), 3-5 rounded to triangular teeth in smallest cheliped (only
sinuous edge in largest specimens). Carpi short, outer borders with low', rounded tubercles; meri slender, distal
portions of outer borders with rounded or pointed tubercles.
First three pairs of walking legs (P2-4: Fig. 5c) flattened. Upper and lower borders of meri with tubercles of
different sizes and shapes (some pointed, some rounded; smaller but on two anterior rows in P2); distalmost
tubercle on each anterior border much wider at base, much higher, slender, pointed, directed distally (only rounded
to triangular in P4). Anterior border of carpi of walking legs (P2-4) tuberculate; borders of propodi and dactyli
(Fig. 6c) entire. Each P2 with one row of tubercles of dorsal, two rows on ventral surface of merus; two dorsal,
two ventral carinae each on carpus and propodus; one dorsal, one ventral carinae on dactylus. Each P3 with
one dorsal row of tubercles on merus; two dorsal, tw'o ventral carinae each on carpus and propodus; one dorsal,
2-3 ventral carinae on dactylus. Each P4 with two dorsal rows of tubercles on merus; two dorsal, two ventral
carinae each on carpus and propodus; one dorsal, 2-3 ventral carinae on dactylus. Meri of all walking legs with
scattered to numerous plumose setae; dorsal surfaces of propodi and dactyli of P3-4 with one row of conspicuous
plumose setae along anterior border.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair slightly
shorter than second. Fifth pair of pereopods (P5) short (0. 7-0.8 CL); each merus slender, surface with microscopic
tubercles, many microscopic tubercles and 1-6 short spines along posterior border; each propodus with 4-7 spines
along posterior border; each dactylus with 2-6 short, thick spines along posterior border (up to 7 spines along
anterior border in some specimens), one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males with all segments free. One complete transverse ridge along each segment 1-4. and
incomplete, low ridge along median portions of segments 5-6. Male first pleopods (Figs 6d-e) with sinuous basal
parts; each distal part with two bilobed processes, inner one broad, longer and outer one (with gonopoda! duct)
narrow, shorter, and provided with minute teeth.
Abdomen of mature females with all segments free. One transversal ridge along each segment 1-4 (low ridge
along segment 5 in most specimens).
DISCUSSION. — P. declivis is characterized by anterolateral teeth that decrease in size from first to fourth, and
walking legs (P2-4) with entire, unarmed dactyli. The dactyli are also unarmed in P. oahuensis but they arc not as
long and slender as in P. declivis. P. declivis shares with P. sexlobatus (Kensley, 1969) anterolateral teeth that
decrease in size and unarmed dactyli. All three species can be easily distinguished by examining their characteristic
male first pleopods.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
461
Fig. 6. — Pseudopalicus declivis sp. nov„ 6 holotype 11.1 x 13.4 mm, Norfolk Ridge, south of New Caledonia.
Bathus 3, stn DW 827, 381-469 m (MNHN-B 26677): a, carapace, dorsal surface: b, suborbital border; c. dactylus,
right fourth pereopod, dorsal view; d, left male first pleopod, dorsal view; e. left male first pleopod. apex, ventral
view.
SIZE. — Maximum size among specimens examined: 18.5 x 22.6 mm (female. MNHN-B 26740); 12.4 x
14.7 mm (male, MNHN-B 26741).
ETYMOLOGY. — From declivis, Latin for downhill, in reference to the decrease in size in a posterior, or
downward, direction of the anterolateral teeth.
DISTRIBUTION. — Known from the South China Sea (off the Philippine Islands), Banda Sea (Indonesia), and
the Coral Sea (Vanuatu, Chesterfield Islands, New Caledonia, and the Loyalty Islands) (Fig. 55). Depth: 189-
469 m.
462
P. CASTRO
Pseudopalicus glaber sp. nov.
Figs 5d, 7, 55, 60b
MATERIAL EXAMINED. — New Caledonia. Musorstom 4: sin DW 164, 18°33.2'S. 1 63° 1 3.0'E. 255 m.
16.09.1985: 1 3 (MNHN-B 26991). — Sin DW 183, 19°01.8'S. I63°25.8'E, 280 m. 18.09.1985: I 3 holotype 9.1 x
10.9 mm (MNHN-B 26683).
CHALCAL 2: stn DW 78, 23°41.3'S, 167°59.6'E, 233 m, 30.10.1986: 1 2 (MNHN-B 26985). — Stn DW 83,
23°20.3’S. 168°05.5'E. 200 m, 31.10.1986: 1 3 , 1 9 (MNHN-B 26986).
Smib 3: stn DW 14, 23°40'S, 168°00’E. 246 m, 22.05.1987: 1 3 , 3 2 (MNHN-B 27028). — Stn DW 18. 23°41.5'S,
167°59.4'E, 338 m, 23.05.1987: 1 2 (MNHN-B 27016). — Stn DW 20, 23°40'S, 168°00'E, 280 m, 23.05.1987: 1 <3,
1 2 (MNHN-B 27029).
Smib 4: stn DW 57. 23°21.5'S, 168°04.6'E, 260 m, 9.03.1989: 1 2 8.4 x 10.6 mm (MNHN-B 26812).
Smib 5: stn DW 70, 23°40.6'S, 168°01 .I E, 270 m, 7.09.1989: 1 2 (MNHN-B 26980). — Stn DW 71. 23°41.3'S,
168°00.7'E, 265 m, 7.09.1989: 2 3,2 9 (MNHN-B 27026). — Stn DW 72, 23°42.0'S. 168°00.8'E. 400 m. 7.09.1989:
1 2 (MNHN-B 27019). — Stn DW 73, 23°41.4'S. I68°00.6'E. 240 m, 7.09.1989: 1 9 paratype 11.7 x 15.0 mm
(MNHN-B 26686). — Stn DW 74, 23°40.2'S. 168°00.9'E, 245 m, 7.09.1989: I 2 (MNHN-B 26981). — Stn DW 75.
22°40.9'S. 1 68°00.8'E, 270 m, 7.09.1989: 6 2 (MNHN-B 27030). — Stn DW 76, 23°41.2'S, 168°00.5’E. 280 m.
7.09.1989: 1 3 paratype 11.0 x 13.5 mm (MNHN-B 26684), 2 2 (MNHN-B 26983). — Sin DW 77. 23°40.0,S.
168°01.1'E, 270 m, 7.09.1989: 2 2 (MNHN-B 27020). — Stn DW 78, 23°40.8'S, 168°00.2'E. 245 m, 7.09.1989: 1 2
(MNHN-B 26988). — Rocky bottom, no station number, 250 m, 13.09.1989: 1 3 paratype 7.7 x 9.0 mm (MNHN-B
26685), 2 2 (MNHN-B 26977). — Stn DW 85, 22°20.0'S, 168°42.9'E, 260 m. 13.09.1989: 1 2 (MNHN-B 26987). —
Sin DW 88, 22°1 8.6'S, 168°40.2'E. 350 m. 13.09.1989: 1 2 (MNHN-B 27018). — Sin DW 91, 22°18.4'S. 168°41.1'E,
340 m, 13.09.1989: 1 2 (MNHN-B 26973). — Stn DW 101, 23°21.2'S, 168°04.9'E, 270 m, 14.09.1989: 4 2 (MNHN-B
27027), 1 3 paratype 7.1 x 8.4 mm. 1 2 paratype 8.5 x 10.8 mm (ZRC 1999.1429-1430). — Stn DW 102, 23°19.6'S,
168°04.7'E, 305 m, 14.09.1989: 1 3 . 1 2 (MNHN-B 27007). — Stn DW 103, 23°17.4'S, I68°04.8’E. 315 m,
14.09.1989: 1 2 9.8 x 12.3 mm (MNHN-B 26687). — Stn DW 105, 23°14.3'S, I68°04.5’E. 310 m. 14.09.1989: 1 2
(MNHN-B 27024).
BERYX 11: stn DW 40, 23°41'S. 168°01'E, 240-300 m. 20.10.1992: 2 2 (MNHN-B 27023).
Smib 8: stn DW 155, 24°45.7'S, 168°08.2’E, 257-262 m, 28.01.1993: 1 2 (MNHN-B 26978). — Stn DW 163,
24°49.1'S, 168°08.9'E. 310-460 m. 28.01.1993: 2 3 (MNHN-B 26984). — Stn DW 170, 23°41.2'S. 168°00.5’E. 241-
244 m. 29.01.1993: 4 2 (MNHN-B 27011), 1 3 paratype 11.4 x 14.0 mm, I 2 paratype 10.6 x 13.4 mm (USNM). —
Stn DW 181, 23°1 7.7'S, 168°04.8'E. 31 1-330 m, 31.01.1993: 2 2 (MNHN-B 26975). — Stn DW 182, 23°19.3'S,
168°04.8'E. 314-340 m. 31.01.1993: 2 <5 (MNHN-B 27015). — Stn DW 183, 23°18.3’S. 168°04.9'E. 330-367 m,
31.01.1993: 1 2 (MNHN-B 27021). — Stn DW 184. 23°18.3'S, 168°04.8'E. 305-320 m, 31.01.1993: 1 3 (MNHN-B
26974). — Stn 185, 23°16'S. 168°04.3'E, 305-355 m, 31.01.1993: 1 3 , 1 2 (MNHN-B 26989). — Stn DW 187,
23°17.7’S, I68°05.6'E, 390-540 m, 31.01.1993: 1 2 (MNHN-B 27014). — Stn 190. 23°18.5'S, 168°04.9'E. 305-
310 m, 31.01.1993: 2 3 (MNHN-B 27022).
Bathus 2: stn DW 727. 22°48.03'S, 167°29.03’E, 299-302 m, 12.05.1993: 1 2 (MNHN-B 27013).
bathos 3: stn CP 804, 23°41'S, 168°00'E. 244-278 m, 27.11.1993: 1 2 (MNHN-B 26979). — Stn CP 805. 23°41'S.
168°01'E, 278-310 m. 27.11.1993: 6 2 (MNHN-B 27009). — Stn DW 829, 23°21'S, 168°02'E, 386-390 m, detritic
rubble, 29.11.1993: 3 8,22 (MNHN-B 26742). — Stn DW 830. 23°20'S, 168°01'E. 361-365 m. 29.11.1993: 1 3 . 1 3
feminized by sacculinid, 6 2 (MNHN-B 27008).
Bathus 4: stn DW 924, 18°54.85'S, 163°24.34'E, 344-360 m, 7.08.1994: 2 3 (MNHN-B 26982). — Stn DW 925.
18°54.55'S, 163°23.75'E. 370-405 m, 7.08.1994: 4 2 (MNHN-B 27025). — Stn DW 926, I8°56'S, 163°25'E. 325-
330 m. 7.08.1994: 1 3 , 1 2 (MNHN-B 27010). — Sin DW 931, 18°55.38'S, 163°24.36'E, 360-377 m. 7.08.1994: 2 2
(MNHN-B 26972). — Stn CP 936, 19°03.67'S, 163°28.05'E, 258-252 m, 8.08.1994: 1 2 paratype 16.5 x 20.4 mm
(MNHN-B 26688).
Halical 1: stn DW 1, 18°54.95'S, 163°24'E. 380-400 m. 23.11.1994: 1 2 (MNHN-B 26990). — Stn DW' 2.
18°55.86'S, 163°24.18'E, 252-397 m, 23.11.1994: I 2 (MNHN-B 27012). — Stn DW 4. 18°55'S. 163°23.29'E. 350-
365 m. 29.11.1994: 1 3 , 1 2 (MNHN-B 26976).
Gemini Seamounts. Volsmar: stn DW 51, 20°58.5'S, 170°03.4’E, 450 m, 4.07.1989: 1 2 (MNHN-B 27017).
Types. — Holotype: 1 3 9.1 x 10.9 mm, MUSORSTOM 4, stn DW 183 (MNHN-B 26683).
Paratypes : 1 3 1 1.0 x 13.5 mm, Smib 5, stn 76 (MNHN-B 26684); 1 6 7.7 x 9.0 mm, Smib 5, no station
number (MNHN-B 26685); 1 2 11.7 x 15.0 mm SMIB 5. stn 73 (MNHN-B 26686); 1 9 9.8 x 12.3 mm.
SMIB 5, stn DW 103 (MNHN-B 26687); 1 9 16.5 x 20.4 mm, BATHUS 4, stn CP 936 (MNHN-B 26688); 1 S
11.4 x 14.0 mm, 1 2 10.6 x 13.4 mm, Smib 8, stn DW 170 (USNM); 1 3 7.1 x 8.4 mm, I 2 8.5 x 10.8 mm,
Smib 5, stn DW 101 (ZRC 1999.1429-1430).
Source : MNHN, Paris
IN' DO- WEST PACIFIC SPECIES OF PALICID CRABS
463
Type Locality. — Grand Passage, north of New Caledonia, 19°0I.8’S, 163°25.8'E, 280 m.
Diagnosis. — Carapace (Figs 5d, 7a, 60b) with four pointed anterolateral teeth on each side; first three teeth
almost equal in size, larger than very small last tooth. Dorsal surface of carapace with few, mostly low, granular
bosses, leaving posterior third of carapace relatively smooth. Supraorbital borders each with two rounded lobes.
Suborbital borders (Fig, 7b) each with two broad, slightly rounded lobes, Dorsal borders of cheliped propodi each
with one row of thin, rounded tubercles. Dactyli of walking legs (P2-4) with short spines along posterior borders
(3-5 spines in P4; Fig. 7c). Abdomen of mature males with all segments free, one complete transverse ridge along
each segment 1-3, incomplete ridge along each median portions of segments 4-6. Male first plcopods (Figs 7d-f)
with sinuous basal parts; each distal part with pointed, spinous dorsal tip, ventral process with two superimposed
foliaceous portions. Abdomen of mature females with segments free, one complete transverse ridge along each
segment 1-4 (low ridge may be present along segment 5).
Colon A female from New Caledonia photographed live (Fig. 60b) had a white vertical band through the center
of the carapace. A brown spot was located on each area anterior and posterior of the white band. The rest of the
carapace was light green and light brown, with small blue spots throughout the branchial regions. The walking
legs were light green with light brown bands.
DESCRIPTION. — Carapace (Figs 5d, 7a, 60b) transversely hexagonal, wider than long (CW/CL = 1.2- 1.3);
dorsal surface covered with small granules and few, mostly low, granular bosses (most conspicuous bosses 3-4 on
branchial region, particularly very large specimens), leaving posterior third of carapace relatively smooth.
Anterolateral borders of carapace each with four pointed teeth, first three of which almost equal in size (first
slightly larger, rounder in many specimens) and larger than very small last tooth (differences more apparent in
larger specimens). Posterior border with an evenly raised edge, no separate tubercles (except in very small
specimens), scattered plumose setae.
Frontal border of carapace divided into four narrow, rounded lobes (inner lobes narrower, longer, and very close
to each other; outer lobes may be broad in some specimens). Borders between frontal lobes and supraorbital borders
straight, folded upward, slightly pointed to conspicuously pointed anteriorly (giving anterior border six-lobcd
appearance in largest specimens), ending in sharp angle, forming V-shaped fissure before supraorbital border.
Supraorbital borders each with two rounded lobes. Postorbital angles moderately long (extend almost to dorsal
border of retracted eye), slender, slightly pointed inward. Cornea of eyes dorsoventrally flattened, wider than base of
short eye peduncle; each peduncle with one conspicuous, granular tubercle on distal border.
Suborbital borders (Fig. 7b) each with broad, rectangular outer lobe, similar in shape (but slightly rounded in
larger specimens); slightly higher, inner lobe. Pterygostomial lobes project ventrally. forming flat, semicircular
structure posterior to each inner suborbital lobe.
Each basal antennal segment thick, rectangular, rounded distal margin; flagellum long, with few, simple setae.
Epistome vertically inclined, with two triangular, pointed (short, rounded in small specimens) median teeth, each
flanked by triangular outer process and thin, rounded margin before pterygostomial lobe. Inner border of ischia of
third maxiliipeds straight; surface coarsely granular, upper borders rounded. Meri much narrower than ischia;
anterior upper lobes much reduced, with rounded outer borders.
Outer surfaces of cheliped propodi each with one row of 1-4 thin, rounded tubercles. Fingers of largest cheliped
with cutting edges (broadly rounded teeth in some specimens), 2-7 rounded to triangular teeth in smallest cheliped
(only sinuous edge in largest specimens). Carpi short, outer borders with one thin, broadly rounded or triangular
tubercle and several low, rounded tubercles; merus slender, distal portions of outer borders with low. rounded
tubercles.
First three pairs of walking legs (P2-4; Figs 5d, 60b) flattened. Upper borders of meri with mostly large,
triangular tubercles; lower borders with few, low tubercles; distalmost tubercle on each anterior border much wider
at base, much higher, slender, pointed, directed distally (only rounded to triangular in P4). Anterior border of carpi
of walking legs (P2-4) each with two large, triangular tubercles; borders of propodi entire. Each P2 with one
dorsal, 1-2 ventral rows of tubercles on merus; one dorsal, one ventral carinae on carpus; one dorsal carina on
propodus; two dorsal, 1-2 ventral carinae, 4-5 short, thick spines on dactylus. Each P3 with one dorsal, two ventral
rows of tubercles on merus (very low in some specimens); two dorsal, one ventral carinae on carpus; two dorsal.
464
P. CASTRO
two ventral carinae on propodus; two dorsal, 2-3 ventral carinae, 3-4 short, thick spines on dactylus. Each P4 with
1-2 dorsal rows of tubercles on merus (very low in some specimens); two dorsal, one ventral carinae on carpus;
two dorsal, two ventral carinae on propodus; one dorsal, 2-3 ventral carinae, 3-5 short, thick spines on dactylus
(Fig. 7c). Meri of all walking legs with scattered plumose setae; dorsal surfaces of propodi and dactyli of P3-4 each
with one row of conspicuous plumose setae along anterior border.
Fig. 7. — Pseudopalicus glaber sp. nov., <S paratype 7.7 x 9.0 mm, Norfolk Ridge, south of New Caledonia, Smib 5,
unknown station, 250 m (MNHN-B 26685): a. carapace, dorsal surface; b, suborbital border; c, dactylus. right fourth
pereopod. dorsal view; d, left male first pleopod, dorsal view; e, left male first pleopod, apex, lateral (outer side)
view; f, left male first pleopod, apex, ventral view.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair slightly
shorter than second. Fifth pair of pereopods (P5) short (0.8 CL); each merus slender, surface with microscopic
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
465
tubercles, many microscopic tubercles, plumose setae along posterior border; each propodus with 4-6 spines along
posterior border; each dactylus with 3-8 short, thick spines along posterior border (mostly along distal portion),
5-10 short spines along anterior border, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males with all segments free. One complete transverse ridge along each segment 1-3, and
incomplete ridge along each median portions of segments 4-5 (low, incomplete ridge in segment 6 in some
specimens). Male first pleopods (Figs 7d-f) with sinuous basal parts; each distal part with pointed, spinous dorsal
tip and ventral process with two superimposed foliaceous portions.
Abdomen of mature females with all segments free. One transversal ridge along each segment 1-4 (low ridge
may be present along segment 5).
DISCUSSION. — P. glaber is easily distinguished from other species of Pseudopalicus by the smooth appearance
of the posterior third of the carapace. Only a few small specimens showed low tubercles along the posterior border
of the carapace.
SIZE. — Maximum size among specimens examined: 16.5 x 20.4 mm (female paratype. MNHN-B 26688);
1 1.0 x 12.8 mm (male, MNHN-B 26742).
Etymology. — From glaber, Latin for smooth, in reference to the relatively smooth dorsal surface,
particularly the posterior portion, of the carapace.
DISTRIBUTION. — Known from New Caledonia (including the Norfolk Ridge south of New Caledonia) and the
Gemini Seamounts east of New Caledonia (Fig. 55). Depth: known reliably between 200 and 450 m; also
collected in a trawl between 390 and 540 m.
Pseudopalicus oahuensis (Rathbun, 1906)
Figs 4e-f, 12a, 54, 60c
Palicus oahuensis Rathbun, 1906: 836, fig. 2. pi. 7, fig. 4. — EDMONDSON, 1933: 268; 1946: 309. — McNeill, 1968:
81, pi. 2, figs 5-6.
Cymopolia oahuensis - EDMONDSON, 1962: 9.
Pseudopalicus oahuensis - Moosa & SERENE, 1981: 37 (in key).
non Palicus oahuensis - BALSS, 1922: 120, fig. 6. — YOKOYA, 1933: 206. 212 [= Pseudopalicus amadaibai (Sakai.
1963)].
non Cymopolia oahuensis - Sakai. 1939: 609. fig. 90a; 1956: 52. — Nakazawa & Sakai, 1947: 665. unnumb. fig.
[= Pseudopalicus amadaibai (Sakai, 1963)].
Material EXAMINED. — Taiwan. Tai-Shi. M.-S. JENG coll., 29.04.1998: 1 9 (AS1ZT 72007).
Indonesia. Kai Islands. KARUBAR: stn CP 17, 05°15'S, 133°01'E, 459-439 m. 24.10.1991: 1 6 (MNHN-B 26925).
Australia. Great Barrier Reef Expedition: stn 15. Cook's Passage, east of Lookout Point, 384 m, 8.03.1929: 1 9
(BMNH 1950.12.1.23).
Chesterfield Islands. MUSORSTOM 5: stn DW 306. 22°07.66'S, 1 59°2 1 ,40'E, 375-415 m, 12.10.1986: 2 $
(MNHN-B 26931).
New Caledonia. Biocal: stn CP 67, 24°55'S. 168°22'E, 500-510 m, 3.09.1985: 1 9 (MNHN-B 26928).
Musorstom 4: stn CP 194. 18°52.8'S, 163°21.7’E, 545 m, 19.09.1985: 1 9 (MNHN-B 26929). — Stn CP 195.
1 8°54,8'S, 163°22.2'E, 465 m. 19.09.1985: 2 6,5 9 (MNHN-B 26923). 1 6 10.6 x 16.9 mm (MNHN-B 26744). —
Stn CP 238, 22°13.0'S, 167°14.0'E. 500-510 m. 2.10.1985: 1 9 11.7 x 15.8 mm (MNHN-B 26930).
466
P. CASTRO
Chalcal 2: sin DW 73, 24°39.9'S, 168°38. l'E, 573 m. 29.10.1986: I juv. 3 . I 3 , 2 2 (MNHN-B 26932). —
Stn DW 74, 24°40.36'S, 168°38.38'E, 650 m, 29.10.1986: 1 2 (MNHN-B 26933). — Sm DW 75, 24°39.31'S.
168°39'67'E. 600 m, 29.10.1986: 1 2 8.2 x 10.7 mm (MNHN-B 26813).
Smib 4: stn DW 56. 23°20.6’S. 168°05.2’E, 260 m, 9.03.1989: 1 3 7.8 x 10.4 mm (MNHN-B 26806).
Beryx 11: stn CP 7, 24°55'S. 168021'E. 510-550 m. 15.10.1992: I 2 (MNHN-B 26934). — Sm DW 9, 24°52'S,
168°22'E. 635-680 m, 15.10.1992: 2 2 (MNHN-B 26935).
Smib 8: stn DW 146, 24°55.2'S, I68°21.7'E, 514-522 m, 27.01.1993: 1 2 (MNHN-B 26936). — Stn DW 149,
24°54.9'S. 168°21.8'E, 508-510 m, 27.01.1993: 1 3 (MNHN-B 26937). — Stn DW 150, 24°54.3'S. 168°22.2'E. 519-
530 m, 27.01.1993: 1 2 (MNHN-B 26938), 2 3 (MNHN-B 26743).
Bathus 4: stn DW 927, 18°55.48'S, 163°22.11'E, 452-444 m, 7.08.1994: 1 2 (MNHN-B 26924).
Loyalty Islands. "Vauban" Dredgings 1977: 20°30'S, 166°48'E, 375 m. 22.02.1977: 1 3 (MNHN-B 26992).
Loyalty Ridge. Bathus 3: stn DW 798, 23°35'S, 169°37'E. 657-660 m, rocks, 26.11.1993: 1 2 (MNHN-B 26993).
— Stn DW 800, 23°35’S. 169°37'E, 655 m. rocks and gravel, 26.11.1993: 1 2 (MNHN-B 26994).
Vanuatu. Musorstom 8: stn CP 983, 19°21 .61 S. 169°27.76’E. 480-475 m, 23.09.1994: 3 3 (MNHN-B 26926). —
Stn CP 1026. 1 7°30.35'S. 168°39.33'E, 437-504 m. 28.09.1994: 1 3 (MNHN-B 26927).
French Polynesia. SMCB (J. Poupin coll.). Socien Islands. Maupiti Island: stn D 92. 16°28.40'S, 151°14.00'E,
397 m, 12.05.1991: 1 2 (MNHN-B 26273).
Tubal Islands: sin 350. 23°20.7'S. 147°32.4'W, 350-200 m. 5.12.1990: I 2 (MNHN-B 26274).
Hawaiian Islands. "Albatross": stn 3919. Oahu, off south coast, 470-402 m, : 1 2 holotype of Palicus oahuensis
Rathbun, 1906 7.2 x 10.0 mm (USNM 29374).
TYPES. — Holotype of Palicus oahuensis Rathbun, 1906: 1 9 7.2 x 10.0 mm, "Albatross", stn 3919 (USNM
29374).
Type Locality. — Off south coast, Oahu, Hawaiian Islands, 470-402 m.
Diagnosis. — Carapace (Figs 12a, 60c) with four anterolateral teeth on each side; first tooth rounded, blunt,
followed by two larger, pointed teeth (second usually slightly larger that third) and one very small last tooth.
Dorsal surface of carapace with large, granular bosses. 10 of which form an almost straight row across branchial
and metagastric regions in large specimens; posterior border with four salient, elongate tubercles, 4-6 smaller,
anterior tubercles in large specimens. Supraorbital borders each with two short, straight-edged lobes. Suborbital
borders each with two broad, rounded lobes; inner slightly higher. Anterior borders of cheliped propodi with
rounded tubercles of various sizes. Anterior and posterior borders of meri of walking legs (P2-4) with thin, nearly
equal, dentiform tubercles (Figs 12a, 60c). Dactyli of walking legs (P4) with posterior border entire. Abdomen of
mature males with all segments free, sutures between segments 4-5 and 3-4 with median U-shaped constriction,
one complete transverse ridge along each segment 1-3, incomplete ridge on segment 4. Male first pleopods (Figs
4e-f) with sinuous basal parts; each distal part broad, flattened, pointed ventrally, two small ventral processes at
base. Abdomen ol mature females with all segments free, one complete transverse ridge along each segment 1-4
(low ridge may be present along segment 5).
Color: Both the carapace and walking legs of a female from New Caledonia photographed live (Fig. 60c) were
pink and light blue. Most tubercles on the dorsal surface of the carapace were light orange.
Discussion. — In addition to the male first pleopods, diagnostic for P. oahuensis is a first anterolateral
tooth that is short and rounded in contrast to the more prominent and pointed second and third teeth. The meri
of the walking legs (P2-4) arc prominent and have nearly equal, dentiform tubercles along both the anterior
and posterior borders that give the legs a frilly, almost lace-like appearance (Figs 12a, 60c; McNeill, 1968,
pi. 2, fig. 5).
Other characters of P. oahuensis include a carapace with a frontal border that is divided into a pair of narrow and
pointed inner lobes, and broad and rounded outer lobes (but narrow and pointed in some small specimens). The
Irontal lobes are followed by a folded-upward and rounded border before the supraorbital border. The postorbital
angles are short (not extending beyond the dorsal border of the retracted eye) and blunt. Each basal antennal
segment is slender, rectangular, and tuberculate on its ventral surface. Each eye peduncle has two granular tubercles
on the distal border. The pterygostomial lobes project ventrally, forming a flat, semicircular structure posterior to
each inner suborbital lobe. The epistome is vertically inclined, with two broadly rounded median teeth flanked on
INDO-WEST PACIFIC SPECIES OF PALIC1D CRABS
467
each side by a triangular outer process and a thin, rounded margin before joining the anterior process of the buccal
frame. The third pair of walking legs (P4) has relatively slender mcri (0.5-0.6 CL), pointed teeth all along both
borders, and one blunt distal tooth on each anterior border. The fifth pair of pereopods (P5) is short (0.7-0.8 CL),
each merus and propodus has numerous pointed tubercles along the posterior border, and each daclylus with 5-12
short spines along the posterior border and two thick distal spines in addition to the terminal tooth.
Size. — Maximum size among specimens examined: 12.7 x 17.3 mm (female, MNHN-B 26743); 10.6 x
16.9 mm (male, MNHN-B 26744).
DISTRIBUTION. — This species has a relatively wide distribution throughout the Pacific basin (Fig. 54).
It has been recorded from the Hawaiian Islands (RATHBUN, 1906; EDMONDSON, 1933, 1946, 1962) and the Great
Barrier Reef, Queensland, Australia (McNeill, 1968). It is now also known from Taiwan, Banda Sea (Indonesia),
Coral Sea (Vanuatu, New Caledonia, and the Loyalty Islands), and French Polynesia. A record of a specimen from
a shallow-water reef in the Hawaiian Islands (EDMONDSON, 1930: 17; 1962: 9) is most probably a
misidentification. Depth: known reliably between 260 and 660 m; also collected in trawls between 200-350
and 635-680 m.
Pseudopalicus sexlobatus (Kensley, 1969)
Fig. 52
Palicus sexlobatus Kensley. 1969: 156, fig. 2a-e; 1981: 50.
Pseudopalicus sexlobatus - Moosa & Serene, 1981: 37 (in key).
MATERIAL EXAMINED. — Mozambique Channel. "Anton Bruun": stn 371F. 24°46'S. 35°18'E, rock dredge.
1 10 m. 18.08.1964: 1 8 holotype of Palicus sexlobatus Kensley, 1969 8.2 x 9.7 mm (SAM A 12642).
Mauritius. Tombeau Bay, stn 43, 238 m, T. MORTENSEN coll., 1 1.10.1929: 1 9 12.5 x 15.8 mm (ZMUC).
TYPES. — Holotype of Palicus sexlobatus Kensley, 1969: 1 $ 8.2 x 9.7 mm (SAM A 12642).
Type Locality. — Mozambique Channel, 24°46'S, 35°18'E, 1 10 m.
Diagnosis. — Carapace (Kensley, 1969, fig. 2a) with four pointed anterolateral teeth on each side; teeth
decrease in size posteriorly. Dorsal surface of carapace with relatively low, granular bosses; posterior border with
six low, elongate tubercles. Supraorbital borders each with two triangular, pointed lobes. Suborbital borders
(KENSLEY, 1969, fig. 2b) each with one broad outer lobe and one rectangular inner lobe. Dorsal borders of cheliped
propodi (KENSLEY, 1969, fig. 2c) with pointed tubercles. Dactyli of walking legs (P2-4) with posterior border
entire (Kensley, 1969, fig. 2e). Abdomen of mature males and females unknown. Male first pleopods (KENSLEY.
1969, fig. 2d) with sinuous basal parts; each distal part uniramous, with triangular teeth along inner and outer
borders and square tip. Abdomen of mature females with all segments free, one complete transverse ridge along
each segment 1-4.
DISCUSSION. — The holotype and an adult female are the only specimens examined. Other characters include a
frontal border that is divided into two pointed inner and two rounded outer lobes, followed by a border that is
slightly pointed near the middle before the supraorbital border. The postorbital angles are relatively long (they
extend to the dorsal border of the retracted eye) and pointed. The walking legs (P2-4) have slender meri, each armed
with one row of teeth along the anterior upper border and one large, pointed distal tubercle.
Size. — Size of the only two specimens examined: 12.5 x 15.8 mm (female, ZMUC); 8.2 x 9.7 mm
(holotype male, SAM A 12642).
Distribution. — Known only from the Mozambique Channel (KENSLEY, 1969) and Mauritius (Fig. 52).
Depth: 1 10-238 m.
468
P. CASTRO
Fig. 8. — a, Pseudopalicus serripes (Alcock & Anderson. 1895). — b, Pseudopalicus investigatoris (Alcock, 1900). —
c. Rectopalicus woodmasoni (Alcock, 1900). Plate 67 from Alcock & McArdle, 1903.
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
469
Pseudopalicus investigatoris (Alcock, 1900)
Figs 4b-c, 8b, 9a, 12b, 52, 60d
Palicus investigatoris Alcock, 1900: 455. — ALCOCK & McArdle. 1903: pi. 67, fig. 2.
Cymopolia fisheri Rathbun, 1906: 835, fig. 1, pi. 7, fig. 5.
Palicus fislieri - EDMONDSON, 1933: 268; 1946: 310.
Cymopolia cyrenae Ward, 1942a: 46. pi. 4, figs 1-2; 1942b: 53. — Michel, 1964: 33; 1974: 134. — Guinot, 1967:
280.
Pseudopalicus fisheri - MOOSA & SERENE, 1981: 36 (in key).
Pseudopalicus cyrenae - MOOSA & Serene, 1981 : 36 (in key).
Pseudopalicus investigatoris - MOOSA & SerEne, 1981: 36 (in key).
non Palicus investigatoris - MACGlLCHRIST, 1905: 265 [= Pseudopalicus serripes (Alcock & Anderson, 1895)].
Material EXAMINED. — Madagascar. Off Nosy Be. " Vauban stn 52, trawl, 15°21.0'S, 46°12.5'E, 150 m,
A. Crosnier coll., 8.11.1972: 1 juv. <3 (MNHN-B 26910).
Mauritius. Tombeau Bay. stn 34, 366 m, T. MoRTENSEN coll.. 26.09.1929: 1 2 (ZMUC).
Western Australia. "Diamantina": stn 33, west of Garden Island. 32°15.7'S, 115°06.7'E, 176-182 m.
18.03.1972: 1 2 (WAM C.23578). — Stn 56, west of Dongara. 29°18'S. 1 14°04'E, 170-174 m. 20.03.1992: 1 2 (WAM
C. 23571). — Stn 66, northwest of Green Head. 29°59'S, 1 14°25'E, 146 m, 22.03.1972: 1 2 (WAM C.23569).
Japan ?. No location: 1 <5 18.7 x 24.4 mm (SMF 24707).
Philippine Islands. Musorstom 1: stn 57. 13053.1'N. 120°13.2'E, 107-96 m, 26.03.1976: 1 (3 10.3 x 12.0 mm
(MNHN-B 26905).
Landsdowne-Fairwav Banks. Chalcal 1: stn D 3, 120-150 m, 21°14.00'S, 162°16.40'E, 18.05.1984: I 2
(MNHN-B 26915).
New Caledonia. BiOCAL: stn CP 84, 20°43'S. 167°0I’E, 150-210 m, 6.09.1985: 1 2 22.3 x 26.3 mm (MNHN-B
26745).
LAGON: stn 1 16, 43 m, sand and gravel, 22°25.2'S, 166°43.7'E, 08.1984: I 2 (MNHN-B 26916). — Stn 390, 155 m.
stylasterids and boulders, 22°42.6’S, 167°01.6'E, 22.01.1985: 1 2 (MNHN-B 26917).
Musorstom 4: stn DW 183, 280 m, 19°01.8'S, 163°25.8'E, 18.09.1985: 1 <3 (MNHN-B 26918).
Smib 5: stn DW 95, 200 m, 22°59.7'S. I69°19.8'E, 14.09.1989: 1 2 (MNHN-B 26919).
Bathus 4: stn CP 938, 280-288 m. 19°00.I6’S, 163°26.45'E, 8.08.1994: 1 2 (MNHN-B 26920). — Stn DW 942.
270-264 m, 19°04.26'S. 163°27.36’E, 8.08.1994: 3 2 (MNHN-B 26921).
Vanuatu. Musorstom 8: stn CP 1077, 180-210 m, 16°04.00'S, 167°06.09'E. 5.10.1994: 1 <3 (MNHN-B 26911),
12 11.7 x 14.5 mm (MNHN-B 26746), 1 2 (MNHN-B 26912). — Stn CP 1078, 194-230 m, 16°03.65'S, 167°26.18'E.
5.10.1994: 1 2 (MNHN-B 26913). — Stn CP 1086, 182-215 m, 15°36.58’S. 167°16.32'E, 5.10.1994: I 2 8.8 x
10.1 mm (MNHN-B 26747), I 2 (MNHN-B 26914).
Fiji. Musorstom 10: stn 1323, 17°16.10’S. 177°45.75'E. 143-173 m. 7.08.1998: 1 juv. <3 (MNHN-B 26922). —
Stn. 1324, 1 7° 1 7.37'S, 177°47.05'E, 102-104 m, 7.08.1998: I 2 8.2 x 9.5 mm (MNHN-B 26748).
Hawaiian Islands. Kauai. ", Albatross stn 3987. 73-426 m: 1 <3 holotype of Cymopolia fisheri Rathbun. 1906.
11.9 x 14.0 mm (USNM 29368).
Oahu. 30-300 m, M. King coll., 1959: 1 2 (LACM).
TYPES. — Holotype of Palicus investigatoris Alcock. 1900: 1 <3 7 x 8 mm, Andaman Islands, India (ALCOCK.
1900). Deposit unknown: Zoological Survey of India, Calcutta ?
Holotype of Cymopolia fisheri Rathbun, 1906: 1 S 1 1.9 x 14.0 mm, off Kauai. Hawaiian Islands. 73-426 m
(USNM 29368).
Holotype of Cymopolia cyrenae Ward, 1942: 1 2 CW 15 mm (WARD, 1942), Mauritius. Deposit unknown:
Mauritius Institute, Port Louis, Mauritius ?
Type Locality. — Andaman Islands, India, unknown depth.
Diagnosis. — Carapace (Figs 8b, 12b, 60d; Alcock & McArdle, 1903, pi. 67, fig. 2) with four pointed
anterolateral teeth on each side; first three teeth usually of similar size but always much larger than fourth (second
and third teeth may be fused into wider tooth in smaller specimens). Dorsal surface of carapace with large granular
bosses, 10 of which form an almost straight row across branchial and metagastric regions in large specimens;
posterior border with 8-10 pointed or elongate tubercles. Supraorbital borders each with two triangular, pointed
lobes. Suborbital borders (Fig. 9a) each with one broad, rounded outer lobe; long, pointed tooth on inner portion.
470
P. CASTRO
Typically two (sometimes one), sharply pointed teeth on pterygostomial ridge connecting pterygostomial lobe
with pterygostomial region of carapace (Fig. 9a). Dorsal borders of cheliped propodi each with several rounded or
pointed tubercles, at least one sharp tooth on proximal portion; inner surface of propodi of males smooth, without
dense cluster of plumose setae; sharp teeth on dorsal surfaces of carpi and meri. Propodi and dactyli of walking legs
(P3-4) with teeth along anterior borders. Dactyli of walking legs slender (P4 0. 5-0.6 CL), with large teeth along
posterior borders (2-3 teeth in P4; Figs 8b, 12b). Abdomen of mature males with all segments free, one complete
but low ridge along segment 3. Male first pleopods (Figs 4b-c) with sinuous basal parts; simple distal part with
small teeth along inner border. Abdomen of mature females with all segments free, one complete transverse ridge
along each segment 1-4 (low ridge may be present along segment 5).
Color. A large female (11.7 x 14.5 mm; MNHN-B 26746) from Vanuatu photographed live (Fig. 60d) had
a multicolored but mostly red-brown carapace and dark green depressed areas at the confluence of the branchial and
mesogastric regions. A small immature female (8.8 x 10.1 mm; MNHN-B 26747) from the same region was light
brownish yellow with bright orange depressed areas. An almost identical color pattern, light brownish yellow with
bright orange depressed areas, was recorded in a female from Fiji that had been preserved for only nine months
[8.2 x 9.5 mm (MNHN-B 26748), Fig. 12b],
Fig. 9. — a. Pseudopalicus investigatoris (Alcock, 1900), $ 8.2 x 9.5 mm, Fiji, MUSORSTOM 10, stn CP 1324, 1 02-
104 m (MNHN-B 26748): suborbital region. — b. Pseudopalicus serripes (Alcock & Anderson. 1895). <3 10.4 x
11.0 mm, Seychelles, Reves 2, stn 21, 55-60 m (MNHN-B 26750): suborbital region.
Discussion. — Additional morphological characters of P. investigatoris are a spherical or slightly flattened
cornea of the eyes, which are wider than the base of the eye peduncle. Each peduncle is long and it has two
conspicuous granular tubercles on the distal border. The inner edge of each suborbital border is marked by one very
conspicuous, pointed tooth slightly higher than the postorbitai angle. The tooth is often but not always separated
dislally from the outer portion of the suborbital border by a narrow notch. The anterior tip of each pterygostomial
lobe projects ventrally, thus forming a flat, semicircular structure (often with 1-3 pointed tips when seen ventrally;
Fig. 9a) posterior to the inner suborbital tooth. The epistome is vertically inclined, with two narrow, triangular
median teeth with very pointed tips Hanked on each side by a similar outer process and a rounded margin before
joining the pterygostomial lobe. The fingers of the larger cheliped have a cutting edge and no teeth: the fingers of
the smaller cheliped are flattened, have triangular teeth, and thick tips. The third pair of walking legs (P4) has
relatively short meri (0.5 CL), each with one row of teeth along upper border and one large and conspicuously
pointed distal tooth; dactyli are slender (0.3 CL), each with 2-3 large teeth along the posterior border. The fifth pair
of pereopods (P5) are short (0.7-0.8 CL), meri each with 5-6 microscopic teeth and tubercles along the posterior
border, propodi each with 6-8 slender spines along the posterior border, and dactyli each with 3-5 short spines
(thicker distal ly ) along the posterior border, 5-12 along anterior border, and one terminal pointed tooth.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
471
P. investigatoris was described from only one specimen, a "non-adult" male (7x8 mm) from the Andaman
Islands. Its detailed description (ALCOCK, 1900) and excellent figure (ALCOCK & McARDLE, 1903. pi. 67, fig. 2:
reproduced here as Fig. 8b) has confirmed the status of this species as a distinct one, notwithstanding its close
proximity to P. serripes (Alcock & Anderson, 1895) and the unavailability of the hololype.
P. investigatoris shares with P. serripes very similar characters: 10 large granular bosses that form an almost
straight row across the branchial and metagastric regions of the carapace of large specimens; four pointed
anterolateral teeth, first three usually of similar size but always much larger than the fourth; a squarish carapace;
two triangular and pointed supraorbital lobes; suborbital borders each with one broad and rounded lobe continuous
with narrow and straight or slightly rounded inner portion; long and pointed postorbital angles; teeth on
the posterior border of the dactyli of the walking legs; epistome with two triangular, pointed median teeth
(although long, thin tips are characteristic of P. investigatoris and only in some small specimens of P. serripes).
They also share very similar male first pleopods: simple distal parts armed with small teeth along the inner border.
Several characters of P. investigatoris, however, separate it from P. serripes, some of which are apparent in the
figures of ALCOCK & McARDLE (1903, pi. 67, figs 1-2; reproduced as Figs 8a-b). Differences are summarized
in Table 2.
TABLE 2. - Morphological differences between Pseudopalicus investigatoris (Alcock, 1900) and serripes (Alcock &
Anderson, 1895).
472
P. CASTRO
The male first pleopods are only slightly different from those of P. serripes in contrast to the sharp differences
between other species of palicids. The male first pleopods of P. investigaioris arc slender, each distal part
dorsovenlrally flattened, slightly curved, and armed with one dorsal row of small teeth (Figs 4b-c), whereas they are
of similar shape (each distal part typically sharply curved and S-shaped) but armed with two dorsal rows of smaller
teeth in P. serripes (Fig. 4d; MOOSA & SERENE. 1981, fig. 5).
Some of the characters given by ALCOCK (1900) as diagnostic for P. investigatoris were found to be absent in
the largest or smallest specimens. The bosses of the carapace, although described by ALCOCK (1900: 455) as
"aureolae... capped by sharp little tubercles" in contrast to "granules" in P. serripes, were found to be mostly
covered with granules in the larger specimens of P. investigatoris. The suborbital border, described by ALCOCK
(1900: 455) as having "no fissure" along its inner end in contrast to an inner "fissure" and an outer notch in
P . serripes, is a variable character in both species. A notch may be present in some individuals. Similarly, the
anterior borders of the carpi of the second and third pairs of walking legs (P3-4) may be serrated in P. investigatoris
as in P. serripes, and they are not always smooth as indicated by ALCOCK (1900: 455).
The examination of the holotype of Cymopolia fisheri Rathbun, 1906 shows that it is a subjective junior
synonym of P . investigatoris. RATHBUN (1906: 836) differentiated her new species from P. investigatoris by
following ALCOCK's premise that the carapace of P. investigatoris is covered by tubercles, not granules as in P.
serripes. She also stated that the frontal lobes of her species were less acute than in P. investigatoris. Although
the holotype (USNM 29368) has granules, the specimen clearly shows the characters diagnostic for
P. investigatoris that are illustrated in ALCOCK's figure of the species (ALCOCK & McArdle, 1903, pi. 67, fig.
1 : reproduced as Fig. 8b), including the more pointed frontal lobes and the slender dactyli of the walking legs.
Although the holotype of Cymopolia cyrenae Ward, 1942 (supposedly still at the Mauritius Institute, Port
Louis, Mauritius) was not available for examination, Ward's description and poor photograph (Ward, 1942a: 46.
pi. 4, figs 1-2) nevertheless provide enough evidence to conclude that his only female specimen is actually
conspecific with P. investigatoris. Ward referred to C. cyrenae as "allied to C. investigatoris Alcock" and
differentiated the two species by three characters: the presence in C. cyrenae of "two petaloid spines" on the carpi of
the walking legs, dactyli each with three spines, and by "all the spines" being longer in the new species. Only one
distal, pointed tubercle is visible on the merus of each of the second walking legs, the only legs shown on Ward’s
photograph, not two as he indicated. Although the figure of P. investigatoris given by ALCOCK & McArdle
(1903, pi. 67, fig. 2; reproduced as Fig. 8b) and the only that could have been available to Ward shows only two
teeth on each second leg, this is a variable character.
The most important diagnostic characters of P. investigatoris are apparent in Ward's photograph:
conspicuously pointed supraorbital lobes and postorbital angles, and slender dactyli of the walking legs. Other
characters are typical of P. investigatoris. The tubercles along the posterior border of the carapace were described by
Ward ( 1942a: 46) as "ten or eleven dentiform spines" (but shown as pointed tubercles on the photograph), each
supraorbital border as "cut into three broad spines, the median of which is broadly truncate, the other two sharp,
and there is a large spine at the external orbital angle" and shown in the photograph as two pointed suborbital
lobes and an elongate, pointed postorbital angle. Each anterolateral border, however, was described as having "three
broad acuminate spines". Although only three can be seen in the photograph, it is very possible that the fourth,
always the smallest, was much reduced (and therefore not visible in the poor photograph) and not defined as an
acuminate spine" by Ward. The eye peduncles were described as having "spines" which is assumed to refer to
pointed tubercles. The character was nevertheless used as a diagnostic feature in the key to the species of
Pseudopahcus that was given by MOOSA & SERENE (1981: 36).
Size. — Maximum size among specimens examined: 22.3 x 26.3 mm (female, MNHN-B 26745)- 18 7 x
24.4 mm (male, SMF 24707).
Distribution. — Mauritius (Ward, 1942a, 1942b; as Cymopolia cyrenae), Andaman Islands (ALCOCK
1900) and the Hawaiian Islands (Rathbun, 1906; EDMONDSON, 1933, 1946; all as Cymopolia fisheri).
Its distribution is now extended to Madagascar, Western Australia, Japan ?, South China Sea (off the
Philippine Islands), Coral Sea (Vanuatu, New Caledonia, and the Landsdowne-Fairway Banks), and Fiji (Fig. 52).
Depth: known reliably between 43 and 366 m; also collected in trawls between 30-300 and 73-426 m.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
473
Pseudopalicus serripes (Alcock & Anderson, 1895)
Figs 4d, 8a, 9b, 12c, 52
Cymopolia serripes Alcock & Anderson, 1895: 208; 1896: pi. 24, fig. 7. — Sakai, 1935: 86. fig. 17: 1936: 208
fig. Ill, pi. 58, fig. 1; 1939: 608, 726, pi. 71, fig. 1; 1956: 52.
Palicus serripes - Bouvier, 1897: 65. — Alcock. 1900: 454. — Calman, 1900: 32. pi. 2, figs 20-22. — De Man. 1902:
546. — ALCOCK & McArdle, 1903: pi. 67, fig. 1. — Laurie, 1906: 431. — Sakai, 1976: 594, pi. 205. fis. 2.
Palicus invesligatoris - MACGlLCHRIST, 1905: 265 [non Pseudopalicus investigators (Alcock, 1900)].
Pseudopalicus serripes - MOOSA & SerEne, 1981: 37, fig. 5, pi. 2, fig. A. — Takeda, 1982a: 205. fig. 607.
'? Palicus aff. serripes - Takeda & KURATA, 1976: 132, fig. 6e.
Material EXAMINED, — Somalia. "Anton Bruun ", cruise 9: sin 447. 10°00'N, 5I°I5'E. 59-61 nr. 16.12 1 964
2 2 (USNM).
Seychelles. Reves 2: sin 21, 05°22.9'S, 56°10.4'E, 55-60 m, 6.09.1980: 1 juv. 6,16 10.4 x 1 1.0 mm iMNHN-B
26750).
Sri Lanka. Gulf of Manaar, W.A. Herdman coll.: 1 2 (BMNH 1934.1.16.165).
Japan. Boso Peninsula. Tateyama Island. 30-60 m, sand. M. Osawa coll., 22.05.1990: 1 6 (CBM-ZC 307).
"Misago": Izu Peninsula (between Ito and Hatsushima), 06.1934: 1 6 (SMF 24708).
Indonesia. Moluccas. Ternate. W. K0KENTHAL coll.: I 2 (SMF 4714),
Australia. Torres Strait. A.C. Haddon coll.: 1 2 (BMNH 1900.11.26.516).
Chesterfield Islands. Corail 2: sin DW 9, 20°53.00'S, 161°35.32,E. 62 m. Halimeda sand, 20.07.1988: 1 2
(MNHN-B 26829). — Stn DW 34. 19°21.62'S, 158°55.77'E. 47 m. foraminiferans and boulders, 23.07.1988: I 2
(MNHN-B 26830). — Stn DW 61, 19°14.96'S, 158°53.60'E. 54 nr. Halimeda sand, 24.08.1988: 1 6 (MNHN-B 26831).
Chesterfield-Bellona Plateau. Chalcal 1: stn D 55. 21°23.90'S. 158°59.60'E, 55 m, 25.07.1984- 1 6 1 2
(MNHN-B 26832).
New Caledonia. MUSORSTOM 4: stn DW 231, 22°33.7’S, 1 67°1 0.5'E. 75 m. 1.10.1985: 1 2 (MNHN-B 26833).
Smib 5: stn DW 100, 23°22.9'S. 168°05.2'E, 120 m. 14.09.1989: 1 2 10.4 x 11.3 mm (MNHN-B 26749).
Futuna Island. MUSORSTOM 7: stn DW 498, 14°19'S, I78°03'W. 105-160 m. 10.05.1992: 1 6 (MNHN-B 26834).
TYPES. — Holotype of Palicus serripes Alcock & Anderson, 1895: 1 9 9.5 x 1 1 mm, India (ALCOCK &
Anderson, 1895). Deposit unknown: Zoological Survey of India. Calcutta ?
Type Locality. — Off Madras coast, southeastern India, shallow water.
Diagnosis. — Carapace (Figs 8a, 12c; Alcock & McArdle, 1903, pi. 67, fig. I) with four pointed
anterolateral teeth on each side; first three teeth usually of similar same size but always much larger than fourth.
Dorsal surface of carapace with relatively low, conspicuously granular bosses; posterior border with 10-12 low.
elongate tubercles. Supraorbital borders each with two narrow, rounded lobes. Suborbital borders (Fig. 9b) each
with one broad, rounded outer lobe; long, pointed tooth on inner portion. One rounded tooth on each
pterygostomial ridge connecting pterygostomial lobe with pterygostomial region of carapace (Fig. 9b). Dorsal
borders of cheliped propodi with several rounded tubercles; inner surface of propodi of males with dense cluster of
plumose setae (see Sakai, 1936, fig. 111c); rounded tubercles on dorsal surfaces of carpi and meri. Propodi and
dactyli of walking legs (P3-4) with teeth along anterior borders. Dactyli of walking legs short, broad (P4 0.2-0.3
CL), with teeth along posterior borders (3-5 teeth in P4; Fig. 8a, 12c). Abdomen of mature males with all
segments free; one almost complete transverse ridge along segment 3. Male first pleopods (Fig. 4d) with sinuous
basal parts; each distal part, with small teeth along inner border. Abdomen of mature females with all segments
free, one complete transverse ridge along each segment 1-4 (low ridge may be present along segment 5).
DISCUSSION. — Besides the characters outlined in the diagnosis and those discussed in its differentiation from
P. invesligatoris (see discussion for P. invesligatoris above), other characters for P. serripes include a frontal border
of the carapace that is divided into four rounded lobes (inner pair narrower and pointed in some specimens),
followed by a border that is folded upward and very slightly pointed anteriorly before the supraorbital border. The
postorbital angles are relatively long (they extend to the dorsal border of the retracted eye) and pointed. Each basal
antennal segment is slender and rectangular (but expanded along its outer border in larger specimens), and with a
row of rounded tubercles along the ventral surface. Each eye peduncle is long and has two granular tubercles on the
474
P. CASTRO
distal border and several along the dorsal border. The inner edge of each suborbital border is marked by one blunt to
pointed tooth that is slightly higher than the suborbital border; the tooth is sometimes separated distally from the
outer portion of the suborbital border by a narrow notch. Each pterygostomial lobe projects ventrally, forming a
flat, semicircular structure (with 2-3 pointed tips when seen ventrally; Fig. 9b) posterior to each inner suborbilal
tooth. The epistome is vertically inclined, with two narrow, triangular median teeth with rounded lips (thin and
pointed in some small specimens) flanked on each side by a narrow, triangular outer process. It continues ventrally
as two rounded lobes around the buccal region. The third pair of walking legs (P4) has short meri (0.4 CL), one
row of teeth along the upper border, and one small and rounded distal tubercle. The fifth pair of pereopods (P5) is
short (0. 6-0.7 CL). Each mcrus has 5-7 microscopic teeth (some of which with one spine) arranged as two rows
along the posterior border; propodus with 6-9 thick spines along the posterior border; each dactylus with 3-4
slightly more slender spines (longer distally) along the posterior border, 2-3 along the anterior border and one
terminal pointed tooth.
P. serripes is very close to P. investigatoris . Differences between the two species are outlined in the discussion
of P. investigatoris (see above) and summarized in Table 2. Reliable characters for the identification of P. serripes
are the presence of a conspicuous cluster of plumose setae on the inner surfaces of the chelipeds of males (Sakai,
1936. fig. 1 1 lc) and the short and broad dactyli of the walking legs (P2-4; Figs 8a, 12c). Juvenile specimens,
however, are difficult to identify.
The male first pleopods are very close in both species. Their shape is very similar, with dorsoventrally Fattened
distal parts, although it was found to be sharply more sinuous (S-shaped) in most of the specimens of P. serripes
that were examined in this study. It is armed with two distal rows of smaller teeth in P. serripes (Fig. 4d) but only
one row of slightly coarser teeth in P. investigatoris (Figs 4b-c). The figure given by Moosa & SERENE (1981.
fig. 5), however, does not clearly show the two rows of teeth and the distal part is not as sinuous as in
most specimens of P. serripes. The three specimens examined by them certainly belong to P. serripes, not
P. investigatoris (which they did not examine), because the chelipeds were described as having "inner surface of the
palm setose" (Moosa & SERENE, 1981: 39).
A male specimen from the Persian Gulf identified as an undescribed variety of Palicus investigatoris by
M.ACGlLCHRIST (1905: 265) seems from its description to belong instead to P. serripes. The location of the
specimen is unknown and could not be examined, but the presence of tubercles along the posterior border that are
"broad and blunt" (indeed described as to "resemble more those of P. serripes") and outer frontal lobes that are
"broad, blunt, and rounded at the end" strongly suggests P. serripes. Two other characters that were given, the
presence ot a fissure on each suborbital border and "densely and finely granular" surface between "sharp little
tubercles on the aureolae of the carapace", are present in specimens of both species.
Size. — Maximum size among specimens examined: 10.4 x 11.3 mm (female, MNHN-B 26749); 10.4 x
1 1.0 mm (male, MNHN-B 26750).
Distribution. — Persian Gulf (MacGilchrist, 1905; as Palicus investigatoris ); southern India and Sri
Lanka (ALCOCK, 1900; Laurie, 1906); Ternate (DE Man, 1902) and Banda Sea (Ceram, Aru Islands, and Kai
Islands), Indonesia (Moosa & Serene, 1981); Japan (Sakai, 1935, 1936, 1939, 1956, 1976; Takeda. 1982a),
and Torres Strait, Queensland, Australia (Calman, 1900). It is now recorded for the first time from Somalia,
Seychelles, Coral Sea (Chesterfield Islands, Chesterfield-Bellona Plateau, and New Caledonia), and Futuna Island,
southwestern Pacific Ocean (Fig. 52). Sakai (1976: 594) records it from "muddy sand or broken shells" in Japan.
Depth: known reliably between 30 m (Sakai, 1976) and 120 m; also collected in a trawl between 105-160 m.
Pseudopalicus sp.
Fig. 10
MATERIAL EXAMINED. — Japan. Ryukyu Islands. Okinawa, off Horseshoe Cliffs, west-northwest Onna Village,
stnRFB 966, 26°30.0'N, 127°50.9'E, 79 m, R.F. Bolland coll., 14.11.1981: I $ 6.3 x 7.5 mm (USNM).
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
475
DIAGNOSIS. — Carapace (Fig. 10a) with four pointed anterolateral teeth on each side; first three teeth of similar
same size but much larger than fourth. Dorsal surface of carapace with relatively low, conspicuously granular
bosses; posterior border with eight low, elongate tubercles. Supraorbital borders each with one rounded inner lobe;
one very short, rounded outer lobe. Suborbital borders (Fig. 10b) each with one pointed inner lobe; outer lobe
divided into two pointed, close-together lobes. Dorsal borders of Cheliped propodi with several rounded tubercles.
Propodi and dactyli of walking legs (P3-4) with conspicuous teeth along anterior borders. Dactyli of walking legs
(Fig. 10c) slender (P4 0.2 CL), with triangular teeth along posterior borders. P5 thick, with thick spines along
posterior borders of propodi and anterior and posterior borders of dactyli.
b
Fig. 10. — Pseudopalicus sp., 2 6.3 x 7.5 mm. Okinawa, Ryukyu Islands. Japan, stn RFB 966. 79.2 m (USNM):
a, carapace, dorsal surface; b. suborbital border; c. dactylus, right fourth pereopod, dorsal view.
DISCUSSION. — An incomplete female specimen (missing the abdomen and one each of the chelipeds, P3. P4.
and P5) from the Ryukyu Islands was found to share some superficial characters with P. serripes, but the presence
of several features unique to the genus suggests that it probably represents a new species. As there is only one
incomplete female specimen available, however, the species is not described as new here.
Similar to P. serripes are the morphology of the dorsal surface of the carapace, chelipeds, and walking legs.
Significant differences are the broad and rounded frontal lobes (narrower in P. serripes), eight elongate tubercles
along the posterior border of the carapace (10-12 in P. serripes), supraorbital borders, each with one rounded inner
lobe and one very broad and short outer lobe (both lobes narrow, rounded, and nearly equal in P. serripes),
suborbital borders each with one pointed inner lobe and an outer lobe that is uniquely divided into two pointed,
close-together lobes (pointed tooth on inner border and one broad, rounded outer lobe in P. serripes), and the
pterygostomial lobes are straight and pointed (ventrally inclined and with 2-3 pointed tips in P. serripes-. Figs 9b.
10b). The posterior borders of the propodi and dactyli of P2-4 are armed with spines and teeth. Another unique
feature is the fifth pair of pereopods (P5). They are thick, short (0.7 CL), the surface of each merus is covered with
small tubercles and long, plumose setae. The mcri and propodi are also unusually spiny, with several rows of thick
spines of various sizes (a total of 17 along posterior border of each propodus) and similar spines along both
borders of the thick and short (0.01 CL) dactyli (three thick spines along each posterior side and one along anterior
side, plus several smaller spines). There are also spines on the basis-ischia and carpi.
Size. — Only specimen known: 6.3 x 7.5 mm (female, USNM).
Distribution. — Known only from Okinawa, Ryukyu Islands, Japan. Depth: 79 m.
476
P CASTRO
Pseudopalicus pictus sp. nov.
Figs 11, 1 2d, 54, 60c
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn DW 965. 20°20.40'S. 169°51.36'E, 361-377 m.
21.09.1994: 1 2 paralype 9.7 x 11.2 mm (MNHN-B 26693). — Stn DW 1042, 16°52.61'S, 168°27.65'E, 200-265 m.
30.09.1994: 1 3 holotypc 8.6 x 10.4 mm (MNHN-B 26690).
French Polynesia. Marquesas Islands. Eiao Island. MUSORSTOM 9: sin CP 1159, 07°58.3'S, 140°43.7'W. 145 m,
23.08.1997: 1 2 (MNHN-B 26818). 1 3 paralype 7.4 x 8.9 mm (MNHN-B 26692). 1 2 paralype 9.0 x 9.9 mm (USNM).
Hiva Oa Island. MUSORSTOM 9: sin DW 1218, 09°44.5'S, 138°50.9'W. 125-135 m, 30.08.1997: I juv. 2 paralype
8.8 x 10.8 mm (MNHN-B 26694), 1 2 (MNHN-B 26816). — Stn DW 1236, 09°41'S. 139°04'W, 250-400 m.
31.08.1997: 1 3 (MNHN-B 26817). — Sin CP 1237, 09°41.9'S, 139°03.6'W, 95-305 m. 31.08.1997: I 3 paralype
6.8 x 8.2 mm (MNHN-B 26691).
Fatu Hiva Island. MUSORSTOM 9: sin DW 1242, 10°28.1'S, 138°41.1'W, 119-122 m, 1.09.1997: 1 juv. 3 . 2 juv. 2
(MNHN-B 26819).
Types. — Holotype: 1 S 8.6 x 10.4 mm, Musorstom 8, stn DW 1042 (MNHN-B 26690).
Paratypes: 1 3 6.8 x 8.2 mm, MUSORSTOM 9, stn CP 1237 (MNHN-B 26691); 1 6 7.4 x 8.9 mm,
Musorstom 9, stn CP 1159 (MNHN-B 26692); 1 2 9.7 X 1 1.2 mm, MUSORSTOM 8, stn DW 965 (MNHN-B
26693); I juv. 2 8.8 x 10.8 mm, Musorstom 9. stn DW 1218 (MNHN-B 26694); 1 2 9.0 x 9.9 mm,
Musorstom 9. stn CP 1 159 (USNM).
Type Locality. — Off southeast coast of Epi Island. Vanuatu, 16°52.61'S, 168°27.65'E, 200-265 m.
Diagnosis. — Carapace (Figs 1 la, 1 2d, 60e) with four pointed anterolateral teeth on each side; second tooth
slightly wider than the rest. Dorsal surface of carapace covered by large granules and few small, low granular
bosses, mostly on branchial and metagastric regions; posterior border with 9-10 elongate tubercles. Supraorbital
borders each with two triangular, pointed lobes. Suborbital borders (Fig. 1 lb) each with one slightly rounded lobe;
long, pointed tooth on inner portion. Dorsal borders of cheliped propodi with several rounded or pointed tubercles.
Propodi and dactyli of walking legs (P3-4) with teeth along anterior borders (1-4 large teeth on each dactylus of P4;
Fig. 1 lc). Abdomen of mature males with all segments free, each with complete transverse ridge, one complete
but low ridge along segments 1-3. Male first pleopods (Figs I Id-e) with sinuous basal parts; each distal part thick,
with pointed, toothed tip sharply bent distally. Abdomen of mature females with all segments free, one complete
transverse ridge along each segment 1-3 (short ridge at each edge of segment 4).
Color : Live specimens (Fig. 60e) with irregular, dark-brown pattern on the carapace and dark-brown legs
distally banded with white, which remains in preserved specimens as a variegated red-brown pattern.
Description. — Carapace (Figs 11a, 1 2d, 60e) transversely hexagonal, slightly squarish, wider than long
(CW/CL = 1. 1-1.2); dorsal surface covered with large granules and few, low, granular bosses (most conspicuous
bosses on branchial and metagastric regions). Anterolateral borders of carapace each with four pointed teeth, second
slightly wider, fourth much smaller than first three. Row of low, rounded tubercles along each posterolateral
border. Posterior border with 9-10 elongate or rounded tubercles, iridescent plumose setae.
Frontal border of carapace divided into four pointed lobes (inner lobes narrower). Borders between frontal lobes
and supraorbital borders slightly pointed anteriorly, folded upward, concave, ending in pointed tip, forming U- or
V-shaped fissure before supraorbital border. Supraorbital borders each with two pointed lobes with slightly rounded
tips. Postorbital angles moderately long (extend just to dorsal border of retracted eye), slender, pointed. Cornea of
eyes dorsoventrally flattened, wider than base of moderately long eye peduncle; each peduncle with four
conspicuous, granular tubercles: two median (most ventral conspicuously elongate, pointed), two on distal border.
Suborbital borders (Fig. lib) each with one broadly rounded outer lobe (limited on outer and inner edges by
notch or wide fissure); conspicuous, pointed tooth on inner limit of suborbital border. Pterygostomial lobes
project ventrally, forming flat, semicircular structure (sometimes notched on inner side) as long as or slightly
shorter than tooth on inner limit of each suborbital border. Sharp tooth on each pterygostomial ridge connecting
pterygostomial lobe with pterygostomial region of carapace.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
All
Each basal antennal segment slender, rectangular (slightly pointed distal border in ventral view); flagellum
long, with few, simple setae. Epistome vertically inclined, with two triangular median teeth with thin, pointed*
tips, each flanked by triangular outer process and thin, rounded margin before pterygostomial lobe. Inner border of
ischia of third maxillipeds straight; surface coarsely granular, upper borders rounded. Meri much narrower than
ischia; upper lobes straight with rounded outer borders.
Dorsal and outer borders of chcliped propodi each with one row of 2-9 thin, rounded (some pointed) tubercles, at
least one of which expanded as high, thin, semicircular process. Fingers of largest cheliped with cutting edges
(broadly rounded teeth in some specimens); smallest chcliped only with sinuous edge except some males (5-7
rounded teeth). Carpi short, outer borders with one thin, broadly rounded tubercle, several pointed tubercles; meri
slender, with few pointed tubercles.
First three pairs ol walking legs (P2-4; Figs 1 2d, 60e) flattened. Upper and lower borders of meri with tubercles
of different sizes and shapes (mostly pointed, some rounded; smaller but on two anterior rows in P2-3): distalmost
tubercle on each anterior border much wider at base, much higher, slender, pointed, directed distally. Anterior
borders of carpi of walking legs (P2-4) with pointed tubercles; anterior borders of propodi and carpi with teeth.
Each P2 with one ventral row of tubercles on merus; 1-2 dorsal, 1-2 ventral row of tubercles on carpus; two dorsal,
2 ventral carinae, 4-5 teeth along posterior border of broad propodus; 1-2 dorsal, two ventral carinae, two teeth on
dactylus. Each P3 with one ventral row of tubercles on merus; two dorsal, one ventral row of tubercles on carpus;
1-2 dorsal, two ventral carinae, 2-6 teeth along posterior border of broad propodus; 1-2 dorsal, two ventral carinae.
1-4 teeth on dactylus. Each P4 with two dorsal, one ventral row of tubercles on carpus; one dorsal, two ventral
carinae, 4-5 teeth along the posterior border of broad propodus; 1-2 dorsal. 1-2 ventral carinae, 1-4 teeth on dactylus
(Fig. 1 lc). Meri of all walking legs (and carpi, propodi of P3-4) with numerous iridescent plumose setae; distal
border of propodi and dorsal surface of anterior borders of propodi and dactyli of P3-4 each with one row of
conspicuous plumose setae.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair slightly
shorter than second. Fifth pair of pereopods (P5) short (0.8-0.9 CL); each merus slender, surface with microscopic
tubercles, many microscopic tubercles and plumose setae along posterior border; each propodus with 4-6 spines
along posterior border; each dactylus with 3-9 short, thick spines along posterior border, 3-8 short spines along
anterior border, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males with all segments free. One complete transverse ridge along each segment 1-3. Male
first pleopods (Figs 1 Id-e) with sinuous basal parts; each distal part thick, with pointed, toothed tip sharply bent
distally.
Abdomen of mature females with all segments free. One transversal ridge along each segment 1-3; short ridge
at each outer edge of segment 4.
Discussion. — P. pictus shares with P. investigatoris and P. serripes a similar shape of the carapace (squarish,
with pointed anterolateral teeth), teeth on the propodi and dactyli of the walking legs (P2-4), a similar arrangement
of the suborbital borders (broad outer lobe and large tooth on the inner limit of the border), and a tooth on the
proximal portion of the ridge that connects the pterygostomial lobe with the pterygostomial region (Fig. 1 lb).
It can be clearly differentiated from both species, however, by its unique male first pleopods, which have thick,
sharply bent distal parts (Figs lld-e), and the smoother appearance of the surface of its carapace. As in
P. investigatoris, the inner surfaces of the male chelipeds lack clusters of setae, in contrast to the cluster of
plumose setae in the male of P. serripes.
478
P. CASTRO
Fig. 11. — Pseudopalicus pictus sp. nov„ 6 hololype 8.6 x 10.4 mm. southeast of Epi Island, Vanuatu, Musorstom 8.
stn DW 1042, 200-265 m (MNHN-B 26690): a, carapace, dorsal surface; b, suborbital border; c. dactylus, right fourth
pereopod, dorsal view; d, left male first pleopod. ventral view; e, left male first pleopod. distal part, dorsal view.
Size. — Maximum size among specimens examined: 9.7 x I 1.2 mm (female, MNHN-B 26693); 8.6 x
10.4 mm (male, MNHN-B 26690).
Etymology. — From the Latin pictus , for painted, in reference to the variegated color pattern of the carapace
and the banded legs, which remain evident in preserved specimens, and the color effect that results from the
numerous iridescent plumose setae.
Distribution. — Coral Sea (Vanuatu) and French Polynesia (Fig. 54). Depth: known reliably between 120
and 370 m; also collected in trawls between 95-305 and 250-400 m.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
479
Fig. 12. — a, Pseudopalicus oahuensis (Rathbun, 1906): $ 11.7 x 15.8 mm. off soulhwest coast of New Caledonia.
MUSORSTOM 4. stn CP 238, 500-510 m (MNHN-B 26749): dorsal view. — b, Pseudopalicus investigaioris (Alcock.
1900), 2 8.2 x 9.5 mm, Fiji. MUSORSTOM 10, stn CP 1324, 102-104 m (MNHN-B 26748): dorsal view. —
c, Pseudopalicus serripes (Alcock & Anderson, 1895), 2 10.4 x 11.3 mm, Norfolk Ridge, south of New Caledonia,
Smib 5, stn DW 100, 120 m (MNHN-B 26749): dorsal view. — d. Pseudopalicus picius sp. nov., juv. 2 paratype 8.8 x
10.8 mm, Marquesas Islands, French Polynesia. Musorstom 9, stn DW 1218, 125-135 m (MNHN-B 26694): dorsal
view. — e, Pseudopalicus macromeles sp. nov., 2 9.7 X 1 1.7 mm, "Diamantina" , west of Rottnest Island. Western
Australia, stn 78. 146-150 m (WAM C. 23580): dorsal view. Unidentified, tubiculous polychaete worm on right eye.
— f, Pseudopalicus undulatus sp. nov., 3 paratype 7.3 x 9.0 mm, Vanuatu. MUSORSTOM 8, stn DW 977. 410-505 m
(MNHN-B 26696): dorsal view.
480
P CASTRO
Pseudopalicus macromeles sp. nov.
Figs 12e, 13, 52
Material EXAMINED, — Western Australia. "Diamantina", cruise DM 6/63: stn 225. wesi of Rottnest Island.
32°00'S, 1 15°16'E. 137-143 m, 12.10.1963: I 3 paratype 7.5 x 8.6 mm (WAM C.24386), 1 9 paratype 10.2 x 12.1 mm
(WAM C. 24387), 1 3 8.0 x 9.4 mm, 21 9 (WAM C. 23575); I 3 paratype 6.8 x 7.5 mm. I 9 paratype 8.5 x 10.3 mm
(MNHN-B 26689).
"Diamantina", cruise DM 1/72: stn 8. northwest of Bunbury, 32°37.5'S. 114°48'E. 139-122 m, 15.03.1972: 2 9
(WAM C. 23582). — Stn 34. west of Garden Island, 32°19'S, 115°07'E, 148-154 m, 18.03.1972: I 9 (WAM C. 23574).
— Stn 55. west of Dongara. 29°15’S, 1 14°01'E. 146 nt. 20.03.1972: 5 9 (WAM C.23577). — Stn 61, west southwest of
Dongara, 29°31'S, 1 14°1 1 'E, 227-221 m, 21.03.1972: 1 9 (WAM C.23572). — Stn 64. northwest of Green Head.
29°58'S. 1 14°27'E. 197-219 m, 22.03.1972: 1 9 (WAM C.23570). — Stn 66. northwest of Green Head. 29°59'S.
I I4°25'E, 146 m. 22.03.1972: 1 9 10.4 x 12.6 mm (WAM C.23569). — Stn 68(1). northwest of Green Island. 30°34'S,
1 14°44'E. 146-139 m. 22.03.1972: 1 9 (WAM C.23567). — Stn 68(3), northwest of Green Island. 30°34'S. 1 14°44'E.
128 m. 22.03.1972: 3 9 (WAM C.23568). — Stn 73(1). west southwest of Lancelin, 31°04'S. 113°50'E, 256 m,
23.03.1972: 1 3,2 9 (WAM C.23576). — Stn 78. west of Rottnest Island. 32°00'S, 115°15'E, 146-150 m.
23.03.1972: 1 <3 holotype 7.4 x 8.6 mm (WAM C.24385), 1 9 paratype 8.1 x 10.0 mm (WAM C.24388). 1 3 , 9 9
(WAM C. 23580). — Stn 78(2). west of Rottnest Island. 32°00'S, 1 15°15 E. 137-146 m. 23.03.1972: 2 9 (WAM
C. 23581). — Stn 79. west of Rottnest Island. 31°59'S, 115°14'E. 182 m, 23.03.1973: 2 3 (WAM C.23579). -
03.1972: 18,49 (WAM C.23573).
"Sprightly": stn 22, 75 km west of Cliff Head. 29°31'S. I14°15'E, 145 m, 18.02.1976: 1 9 (WAM C.14325).
TYPES. — Holotype : 1 $ 7.4 x 8.6 mm, "Diamantina", stn 78 (WAM C.24385).
Paratypes : 1 6 7.5 x 8.6 mm, "Diamantina", stn 225 (WAM C.24386); 1 2 10.2 x 12.1 mm, "Diamantina".
stn 225 (WAM C.24387); 1 2 8.1 x 10.0 mm, "Diamantina", stn 78 (WAM C.24388); 1 <3 6.8 x 7.5 mm. 1 2
8.5 x 10.3 mm, "Diamantina", stn 225 (MNHN-B 26689).
Type Locality. — West of Rottnest Island. Western Australia, 32°00'S, 1 15°15'E. 146-150 m.
DIAGNOSIS. — Carapace (Figs 12e, 13a) with three granular anterolateral teeth on each side; teeth decrease
posteriorly in size. Dorsal surface of carapace covered by large granules and several granular bosses, some rounded,
high, particularly 6-7 on each branchial region; posterior border with 4-6 rounded, granular tubercles. Supraorbital
borders each with two triangular lobes. Suborbital borders (Fig. 13b) each with two rounded lobes (high, narrow
inner lobe, broad outer lobe). Dorsal borders of cheliped propodi with several rounded or pointed tubercles. Dactyli
of walking legs (P2-4) very slender, with posterior borders entire (Fig. 13c). Abdomen of mature males w ith all
segments free, one complete transverse ridge along each segment 1-3, incomplete ridge along segment 4. Male first
pleopods (Figs 13d-e) with sinuous basal parts; each distal part with two broad lateral processes, inner one broader,
bordered by thicker portion, outer one narrower, armed with teeth. Abdomen of mature females with all segments
free, one complete transverse ridge along each segment 1-4.
DESCRIPTION. — Carapace (Figs 12e, 13a) transversely hexagonal, wider than long (CW/CL = 1.1 -1.2); dorsal
surface covered with large conspicuous granules and granular bosses (most conspicuous 6-7 on each branchial
region, continuing as a row of smaller bosses to two salient bosses on mctagastric region). Sulcus between hepatic
and branchial regions apparent. Anterolateral borders of carapace each with three granular, anteriorly pointed teeth
decreasing posteriorly in size (last one can be very small, as in holotype; Fig. 13a). Epistcrnal processes granular,
reduced (short, rounded) in males and immature females, more pronounced (elongate) in females. Posterior border
with 4-6 rounded, granular tubercles, scattered plumose setae.
Frontal border of carapace divided into four rounded to pointed lobes (inner lobes longer). Borders between
frontal lobes and supraorbital borders folded upward, nearly straight (slightly rounded in some specimens),
and ending in sharp angle, forming U- or V-shaped fissure before supraorbital border. Supraorbital borders each
with two triangular, pointed (lips slightly rounded in some specimens) lobes. Postorbital angles long (extend
beyond dorsal border of retracted eye), pointed inward. Cornea of eyes dorsoventrally flattened, wdder than base
of short eye peduncle; each peduncle with three granular tubercles on distal border (two dorsal tubercles most
conspicuous).
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
481
Suborbital borders (Fig. 13b) each with high, narrow, rounded inner lobe; broadly rounded outer lobe.
Pterygostomial lobes project ventrally, forming semicircular structure posterior to each inner suborbital lobe.
Each basal antennal segment slender, rectangular (slightly expanded distally, with pointed distal border in
ventral view), and luberculate; flagellum long, with few, simple setae. Epistome vertically inclined, with two
short, triangular median teeth and wide, rounded margin before pterygostomial lobe. Inner border of ischia of third
maxillipeds straight; surface coarsely granular, upper borders obliquely-directed. Meri much narrower than ischia;
upper lobes much reduced, with straight outer borders.
Chclipeds equal or slightly unequal in females, conspicuously unequal in males. Dorsal borders of propodi of
small chelipeds of both sexes each with one row of pointed, thin, elongate tubercles, outer border with 1-2 rows of
low, rounded tubercles. Large cheliped of males with high propodus armed with 3-4 rows of conspicuous, rounded
tubercles decreasing in size ventrally. Fingers of small chelipeds of both sexes each with 3-4 broadly rounded or
triangular (particularly in males) teeth or cutting edges. Carpi of small and large chelipeds of both sexes short,
outer borders with conspicuous pointed, rounded tubercles; meri slender, with pointed, rounded tubercles.
First three pairs of walking legs (P2-4; Fig. 12e) very slender, without filiform propodi. Upper and lower
borders of meri with tubercles of different sizes and shapes (mostly rounded, some pointed; smaller in P2);
distalmost tubercle on each anterior border of meri of P2-3 much wider at base, much higher, slender, pointed or
rounded, directed distally (short, truncate in P4). Carpi and dactyli of walking legs (P2-4) elongate; anterior borders
of carpi tuberculate; borders of propodi and dactyli (Fig. 13c) entire. Each P2 with one ventral row of tubercles on
merus; one dorsal, one ventral carinae on carpus; one dorsal, 1-2 ventral carinae on propodus; one dorsal, two
ventral carinae on dactylus. Each P3 with one dorsal, one ventral row of tubercles on merus (very low in some
specimens); two dorsal, two ventral carinae each on carpus, propodus, and dactylus. Each P4 with two dorsal, one
ventral row of tubercles on merus; one dorsal, one ventral carinae on carpus; two dorsal, two ventral carinae on
propodus; one dorsal, two ventral carinae on dactylus. Meri of all walking legs with scattered plumose setae, few
on carpi; dorsal surfaces of propodi and dactyli of P3-4 each with one row of conspicuous plumose setae along
anterior border.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair slightly
shorter than second. Fifth pair of pereopods (P5) short (0.9 CL); each merus slender, surface with microscopic
tubercles, 2-4 spines along posterior border, 0-2 thick spines along anterior border; each propodus with 6-9 long
spines along anterior border; each dactylus with 0-3 spines along posterior border, 0-2 thick spines along anterior
border, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males with all segments free. One complete transverse ridge along each segment 1-3, low,
incomplete ridge along most of segments 4-6 (very low, if present, in segments 5-6). Male first pleopods (Figs
13d-e) with sinuous basal parts; each distal part with two broad lateral processes, inner one broader and bordered by
thicker portion, outer one (with sperm channel) narrower and armed with teeth.
Abdomen of mature females with all segments free. One transversal ridge along each segment 1-4 (very low in
segment 4 and, if present, in segment 5).
Discussion. — An unusual feature of P. macromeles is its very long walking legs (P2-4; Fig. 12e),
a characteristic of species of Parapalicus and Miropalicus. However, in contrast to Miropalicus. and as in
Pseudopalicus and Parapalicus , the last pair of walking legs (P4) is about the same length as the second pair (P3)
and much longer than the first pair (P2). The total length of the walking legs, however, is a function of the shape
of the dactyli, which varies among the species of Pseudopalicus but are always slender and long in Parapalicus.
Thus, total length of the P3 varied between 2. 3-2. 5 CL in males and females of Pseudopalicus macromeles ,
482
P. CASTRO
Fig. 13. — Pseudopalicus macromeles sp. nov., 8 holotype 7.4 x 8.6 mm, west of Rottnest Island. Western Australia,
"Diamantina", stn 78, 146-150 m (WAM C. 24585): a, carapace, dorsal surface; b. suborbital border; c. dactylus,
right fourth pereopod. dorsal view; d. left male first pleopod. ventral view; e. left male first pleopod, distal part,
dorsal view.
2.0 CL in Pseudopalicus declivis (a species with slender dactyli), 1 .7 CL in Pseudopalicus serripes (a species with
short dactyli), and 2.7 CL in both Parapalicus ambonensis (Moosa & Serene, 1981) and P. unidentatus (Zarenkov.
1968). Furthermore, the walking legs of P. macromeles are morphologically identical to the legs of the other
species of Pseudopalicus. They all have relatively short and broad meri with unequal tubercles arranged as rows
along anterior and posterior sides and as a single row of unequal (some much broader and triangular) tubercles along
both the upper (dorsal) and lower (ventral) borders; the carpi are short and their upper borders have irregular,
triangular tubercles, particularly on the proximal and distal portions; and the coxae of each P3-4 are broad, laminar,
with thin anterior and posterior borders. In contrast, the walking legs among the species of Parapalicus have
longer, narrower meri with more homogenous tubercles throughout their surface and much longer, more slender
carpi with only microscopic tubercles (an exception being P. armatus, with equal, pointed tubercles on the carpi of
P3-4; see Fig. 1 7a); and the coxae are not broad and laminar, with the anterior and posterior borders rounded.
Another character P . macromeles shares with Parapalicus is the presence in females of conspicuous episternal
processes that protrude from each posterolateral margin of the carapace. It is reduced, as in other species of
Pseudopalicus, in the male and immature females. The functional significance of this character is unknown. It may
act as stabilizers in larger, heavier individuals, like mature females, for sliding over soft sediments as live
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
483
individuals of Parapalicus species have been observed doing (B. Richer de FORGES, personal communication).
Miropalicus vietnamensis (Zarenkov, 1968), however, has very long, slender walking legs but the episternal
process is substantially reduced. It may be suggested that P. macromeles has departed from the basic
Pseudopalicus- type walking legs and has secondarily evolved by convergence longer Parapalicus- type legs, as an
adaptation for sliding. No other species of Parapalicus have ever been collected along ihe Indian Ocean coast of
Western Australia, where P. macromeles is only known to occur.
Otherwise, the structure of the carapace (including anterior, anterolateral, supraorbital, and suborbital borders),
antennae, epistome, and abdomen of P. macromeles are like those characteristic of Pseudopalicus and different from
Parapalicus (see Table 1). The chelipeds of the males are like those in the other species of Pseudopalicus, where
the fingers of the larger cheliped have cutting edges or rounded teeth and the smaller cheliped has triangular teeth
and becomes flatter with increasing size. In females, however, both chelipeds are about the same size, with slender
fingers armed with triangular teeth.
P. macromeles is close to P. undulatus sp. nov. They are the only two species of Pseudopalicus with only
three conspicuous anterolateral teeth. They also share male first pleopods where the sperm channel follows
a strongly sinuous (S-shaped), although not helicoidal, path along each basal part of the pleopods. This is
a departure from the slightly sinuous basal part that is characteristic of congeners. The remaining characters of
the pleopods, however, are very different in both species. In P. undulatus (Figs 14d-e). there is a unique median
tubercle (present also in Rectopalicus amphiceros sp. nov.; Fig. 35d) and the distal part is dorsoventrally-flattened
and bordered with teeth. In P. macromeles, in contrast, there are two broad processes similar to those in
P. acanthodactylus (Figs 3e-f), P. declivis (Figs 6d-e), and P. oahuensis (Figs 4e-f).
Size, — Maximum size among specimens examined: 10.4 x 12.6 mm (female. WAM C. 23569); 8.0 x
9.4 mm (male, WAM C.23575).
Etymology. — From makros , Greek for long, and melos, Greek for limb, to refer to the diagnostic, very
long walking legs (P2-4).
Distribution. — Known only from the eastern Indian Ocean off the coast of Western Australia (Fig. 52).
Depth: 128-256 m.
Pseudopalicus undulatus sp. nov.
Figs 12f, 14, 53
MATERIAL EXAMINED. — Japan. Izu Islands. Hyotan-se Bank, 34°20.75'N. 139°20.00'E, 275-350 m. T. Komai
coll., 15.10.1997: I <3 8.2 x 10.0 mm paralype (CBM-ZC 4625).
Philippine Islands. Musorstom 2: stn CP 15, 13°55.1'N, 120°28.4’E. 326-330 m, 2.11.1980: I 2 9.9 x
11.9 mm (MNHN-B 26697). — Stn CP 83, 13°55.2'N, 120°30.5E, 320 m. 2.12.1980: I 2 (MNHN-B 26820).
Indonesia. Kai Islands. Karubar; stn DW 14, 05°18'S, 132°38'E. 245-246 m, 24.10.1991: 1 3 feminized by
sacculinid (MNHN-B 26821). — Stn DW 32, 05o47'S, 132°51'E. 170-206 m. 26.10.1991: 1 2 (MNHN-B 26822).
Tanimbar Islands. Karubar: stn CP 46, 08°01'S. 132°51'E. 271-273 m, 29.10.1991: 1 3 holotype 8.4 x 10.8 mm
(MNHN-B 26695).
Vanuatu. Musorstom 8: stn DW 977, 19°24.89'S. 169°28.6rE. 410-505 m, 22.09.1994: 1 3 paratype 7.3 x
9.0 mm (MNHN-B 26696).
Fiji. Bordau 1: stn Dw 1448, 16°45'S, 179°59'E, 410-500 m, 4.03.1999: 18.42 (MNHN-B 27132). —
Sin DW 1465. 18°09'S. 178°39'W, 290-300 m. 6.03.1999: 1 3 7.8 x 9.4 mm (MNHN-B 27134).
TYPES. — Holotype-. 1 S 8.4 x 10.8 mm, Karubar, sin CP 46 (MNHN-B 26695).
Paratypes : 18 7.3 x 9.0 mm. MUSORSTOM 8, stn DW 977 (MNHN-B 26696); 18 8.2 x 1 0.0 mm,
Hyotan-se Bank, Japan (CBM-ZC 4625); 1 $ paratype 9.9 x 1 1.9 mm, Musorstom 2, stn CP 15 (MNHN-
B 26697).
Type Locality. — Tanimbar Islands, Indonesia. 08°0I'S, 132°51'E. 271-273 m.
DIAGNOSIS. — Carapace (Figs 12f, 14a) transversely subovate with three anterolateral teeth on each side; teeth
(rounded or slightly pointed upward in shape) decrease posteriorly in size. Dorsal surface covered by large granules
484
P. CASTRO
and several granular bosses, some rounded, high, particularly two on branchial region close to border of carapace;
branchial region slightly inflated; posterior border with 4-6 elongate, curved tubercles, giving border sinuous
appearance. Supraorbital borders each with one short, rectangular inner lobe; short, rounded outer lobe. Suborbital
borders (Fig. 14b) each with high, narrow inner lobe; high, rounded outer lobe. Dorsal borders of cheliped propodi
with rounded or pointed tubercles. Dactyli of walking legs (P2-4) very slender (P4 0.3-0. 4 CL), with posterior
borders entire (Fig. 14c). Abdomen of mature males with all segments free, one complete transverse ridge along
segments 1-3, incomplete ridge along segment 4. Male first pleopods (Figs I4d-e) with sinuous basal parts;
each distal part dorsoventrally flattened, with tubercle on ventral side, long row of teeth along outer edge (shorter
row on inner edge). Abdomen of mature females with all segments free, one complete transverse ridge along each
segment 1-4.
DESCRIPTION. — Carapace (Figs 12f, 14a) transversely subovate, wider than long (CW/CL = 1.2-1. 3); dorsal
surface covered with small granules and granular bosses (conspicuous and high are two rounded bosses on each
branchial region along anterolateral borders, 2-3 similar bosses on center and upper border of each branchial region,
two elongate bosses on metagastric region flanked by one rounded boss on each side, and one elongate boss on
intestinal region immediately above the posterior border of carapace). Anterolateral borders of carapace each with
three rounded or slightly pointed teeth decreasing posteriorly in size. Branchial region slightly inflated. Posterior
border with 4-6 elongate, curved tubercles (giving border sinuous appearance), scattered plumose setae.
Frontal border of carapace divided into four narrow, rounded lobes (inner lobes narrower, much longer). Two
conspicuous, broadly rounded bosses behind outer lobes. Borders between frontal lobes and supraorbital borders
folded upward, rounded or slightly pointed medially, ending in sharp angle, forming V-shaped fissure before
supraorbital border. Supraorbital borders each with one short, rectangular inner lobe; short, rounded outer lobe.
Postorbital angles short (extending two-thirds of dorsal border of retracted eye), slightly narrower proximally,
pointed. Cornea of eyes dorsoventrally flattened, wider than base of short eye peduncle; each peduncle with two
granular tubercles on distal border.
Suborbital borders (Fig. 14b) each with high, narrow, inner lobe; high, rounded, outer lobe only slightly lower
than postorbital angle. Pterygostomial lobes project ventrally, forming Hat, semicircular structure posterior to each
inner suborbital lobe.
Each basal antennal segment slender, rectangular (slightly pointed distal border in ventral view); flagellum
long, with few, simple setae. Epistome vertically inclined, with two triangular median teeth with slightly rounded
tips, each flanked by triangular, narrow, pointed outer process and thin, rounded margin before pterygostomial
lobe. Inner border of ischia of third maxillipeds straight; surface coarsely granular, upper borders rounded. Meri
much narrower than ischia; upper lobes broadly rounded.
Outer borders of cheliped propodi each with 3-4 rows of rounded tubercles (more pronounced in small cheliped).
Fingers of largest cheliped with 3-4 broadly rounded teeth, 4-8 rounded to triangular teeth (or none) in smallest
cheliped. Carpi short, outer borders with several low, rounded tubercles; meri slender, each with one row of
conspicuous rounded tubercles (distalmost one more salient) and other rounded tubercles throughout.
First three pairs of walking legs (P2-4; Fig. 12f) flattened. Upper and lower borders of meri with tubercles of
different sizes and shapes (mostly rounded, some pointed; smaller in P2); distalmost tubercle on each anterior
border much wider at base, much higher, slender, pointed or rounded, directed distally. Carpi of walking legs (P2-4)
elongate; borders of propodi and dactyli (Fig. 14c) entire. Each P2 with one dorsal, 1-2 ventral rows of tubercles on
merus; two dorsal, 1-2 ventral carinae on carpus; one dorsal, two ventral carinae on propodus; one dorsal, one
ventral carinae on dactylus. Each P3 with 1-2 dorsal rows of tubercles on merus (very low in some specimens);
two dorsal, one ventral carinae on carpus; two dorsal, two ventral carinae on propodus; one dorsal, two ventral
carinae on dactylus. Each P4 with two dorsal, one ventral (very low) rows of tubercles on merus; two dorsal, two
ventral carinae on carpus; one dorsal, one ventral carinae on propodus; one dorsal, two ventral carinae on dactylus.
Meri, carpi, and propodi of all walking legs with scattered plumose setae; dorsal surfaces of propodi and dactyli of
P3-4 each with one row of conspicuous plumose setae along anterior borders.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair shorter than
second. Fifth pair of pereopods (P5) short (0.8-0. 9 CL); each merus very slender, surface with microscopic
tubercles, many microscopic teeth, plumose setae along posterior border; each propodus with many microscopic
Source : MNHN, Paris
1ND0-WEST PACIFIC SPECIES OF PALICID CRABS
485
teeth, 0-3 short spines, plumose setae along posterior border; each dactylus with 2-9 thick spines (thicker distally)
along posterior border, 1-5 small spines along anterior border, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males with all segments free. One complete transverse ridge along each segment 1-3; low,
incomplete ridge along most of segment 4. Male first pleopods (Figs 14d-e) with sinuous basal parts; each distal
part dorsoventrally flattened, with tubercle on ventral side, long row of teeth along outer edge, shorter row on inner
edge.
Abdomen of mature females with all segments free. One transversal ridge along each segment 1-4.
Fig. 14. _ Pseudopalicus undulatus sp. nov., <J holotype 8.4 x 10.8 mm, Tanimbar Islands. Arafura Sea. Indonesia.
Karubar, stn CP 46, 271-273 m (MNHN-B 26695): a, carapace, dorsal surface: b. suborbital border; c. dactylus.
right fourth pereopod, dorsal view; d, left male first pleopod, lateral (outer side) view; e, left male first pleopod, apex,
lateral (inner side) view.
Source
486
P. CASTRO
DISCUSSION. — Diagnostic for P. undulatus is the presence of only three anterolateral teeth in contrast to four
in most other known species of Pseudopalicus, although in some of these species the last tooth may be very small
in some specimens. P. undulatus and P. macronieles are exceptional in the genus for having only three
conspicuous anterolateral teeth. Also diagnostic for P. undulatus is the presence of conspicuously unequal
supraorbital lobes, the inner lobe being short and the outer higher and rounded. The branchial region is slightly
higher (in part accentuated by the three high bosses on its surface) than in other species of Pseudopalicus so that
the carapace has a transversely subovatc appearance (Fig. 12f) rather than the more hexagonal appearance
characteristic of the genus.
More significant than these characters are the unusual male first plcopods. The basal parts are strongly sinuous
(although not helicoidal as in Parapalicus) and a tubercle is present on each median, ventral surface. P. macronieles
has male first pleopods with strongly sinuous basal parts (Fig. 13d); those of Rectopalicus amphiceros sp. nov.
also have a median tubercle (Fig. 35d). The distal ends were thin, straight, and obliquely oriented in an inner
direction in a small, perhaps juvenile male from Fiji (7.8 x 9.4 mm; MNHN-B 27134).
Size. — Maximum size among specimens examined: 9.9 x 11.9 mm (female, MNHN-B 26697); 8.4 x
10.8 mm (male, MNHN-B 26695).
ETYMOLOGY. — From unda, Latin for wave, in reference to the wavy appearance of the posterior border of
the carapace due to the presence of elongate and curved tubercles along the margin.
DISTRIBUTION. — Japan, South China Sea (off the Philippine Islands). Banda and Arafura seas (Indonesia),
Coral Sea (Vanuatu) and Fiji (Fig. 53). Depth: known reliably between 206 and 410 m; also collected in trawls
between 170-206 and 410-505 m.
Genus PARAPALICUS Moosa & Serene, 1981
Parapalicus Moosa & Serene, 1981: 25.
Type Species: Palicus trituberculatus Chen, 1981. Gender: masculine
DIAGNOSIS. — Frontal border of carapace divided into two small lobes (sometimes secondarily notched).
Anterolateral borders each with one triangular or rounded tooth. Dorsal surface of carapace typically with small
granules and high granular bosses. Eyes dorsoventrally flattened; peduncles with small, typically soft tubercles.
Supraorbital borders each with one or two notches but no conspicuous lobes. Suborbital borders each typically
with two dentiform lobes; outer lobe, if present, smaller than inner lobe. Each basal antennal segment slightly
expanded distally (scale-like). Epistome expanded dorsoventrally; thin carina-like process along median portion
without protruding median teeth. Chelipeds small, propodi and fingers slender; equal or nearly equal in males and
females. First three pairs of walking legs (P2-4) very slender, with elongate meri and filiform (long, very narrow)
propodi and dactyli. First pair (P2) much shorter than second and third pairs (P3-4); third pair slightly shorter or
about as long as second pair. Borders of meri of P2-4 with equal or nearly equal tubercles; carpi entire, without
teeth (tubercles rare); propodi and dactyli entire. Fifth pair of pereopods (P5) reduced (0.8-0.9 CL) and very slender.
Abdomen of mature males triangular, with segments 3-5 fused into segment much wider than segment 6 and
telson; typically two tubercles on segment 3, one on 4. Male first pleopods short; basal parts helicoidal; each
distal part with one blade-like process or helicoidal pointed tips. Abdomen of mature females with segments 3-5
fused; typically one complete or reduced transverse ridge along each segment 1-4.
REDESCRIPTION. — Carapace transversely subovate, broader than long, conspicuously covered with granules of
varying size, often grouped in clusters forming bosses. Anterolateral borders of carapace each with one typically
conspicuous, triangular or rounded, tooth, below which carapace slightly rounded. Confluence of branchial and
esogastric regions depressed, smooth; sulcus between hepatic and branchial regions. Thoracic sternite 7 with
conspicuous, protuberant process (episternal process) at each outer edge posterior to insertion of fifth pair of
pereopods (P5), visible dorsally as rounded or oblong process below posterolateral border of carapace.
Source : MNHN, Paris
IN DO- WEST PACIFIC SPECIES OF PALICID CRABS
487
Frontal border divided into two small lobes (sometimes secondarily notched). Supraorbital borders each with
one or two notches but no conspicuous lobes. Postorbital angles short, slightly pointed or rounded tips. Cornea of
eyes dorsoventrally flattened, much wider than base of eye peduncle (reniform): peduncles with small, soft or
granular tubercles. Orbits deep.
Suborbital borders each typically with two pointed or rounded, dentiform lobes; outer lobe, if present, smaller
than inner lobe. Pterygostomial lobe at each anterolateral angle of buccal frame projects ventrally, rounded to
pointed in shape.
Each basal antennal segment slightly expanded
distally (scale-like); flagellum long with conspic¬
uous setae. Epistome (Fig. 15) expanded dorsoven¬
trally, forming broad, nearly flat surface between
antennular fossae and anterior border of endostome;
no median teeth, only low, thin, carina-like process
along median portion flanked on each side by an
outer process connecting with pterygostomial lobe.
Meri of third maxillipeds smaller and narrower than
ischia.
Chelipeds small, equal or nearly equal in males
and females. Cheliped propodi slender; dorsal
borders with small teeth or tubercles, or smooth;
fingers with pointed tips, typically with triangular
or rounded distal teeth. First three pairs of walking
legs (P2-4) very slender, with elongate meri and
filiform (long and very narrow) propodi and dactyli.
Upper and lower borders of meri with equal or
nearly equal tubercles; carpi entire, without teeth
(tubercles rare); borders of propodi and dactyli
entire. First pair of walking legs (P2) much shorter and slender than second and third pairs (P3-4); third pair
slightly shorter or about as long as second pair. Last two pairs of walking legs (P3-4) each with rounded coxae,
having rounded (not expanded, thin-edged) anterior and posterior borders. Fifth pairs of pereopods (P5) reduced
(0. 8-0.9 CL) and very slender; basis-ischia slender, smooth; few spines along posterior border of dactyli.
sometimes on propodi.
Abdomen of mature males with segments 1-2 dorsoventrally compressed, 3-5 fused (but traces of sutures
usually visible) into segment much wider than segment 6 and telson, giving abdomen triangular appearance.
Typically two tubercles on segment 3, one on 4. Telson with fine granules. Male first plcopods short; basal parts
helicoidal, each distal part with one blade-like process or hclicoidal pointed tips. Second male pleopods short, thin,
slightly curved; distal segment with blunt tip.
Abdomen of mature females broad, rounded, segments 1-2 dorsoventrally compressed. 3-5 fused (but traces of
suture usually visible), typically one complete or reduced transverse ridge along each segment 1-4. Abdomen ol
immature females triangular (but broader than in males), segments 1-2 dorsoventrally compressed. 3-5 or 3-6 fused.
Fig. 15. — Parapalicus clinodentatus sp. nov., $ 8.6 x
12.2 mm, off southeast coast of New Caledonia. Bathus
2, stn CP 742, 340-470 m (MNHN-B 26753): epistome.
transverse ridge along each segment 1-4.
SPECIES INCLUDED. — There are 13 known species of Parapalicus, by alphabetical order: P. ambonensis
Moosa & Serene, 1981, P. annatus sp. nov.. P. clinodentatus sp. nov., P. denticulatus sp. nov., P. elaniticus
(Holthuis, 1977), P. inanis sp. nov., P. inermis sp. nov., P. microphthalmus sp. nov., P. nanshaensis Dai & Xu.
1991, P. piruensis Moosa & Serene, 1981, P. trispiralis sp. nov., P. trituberculatus (Chen, 1981), P. unidentatus
(Zarcnkov, 1968).
SEXUAL Dimorphism. — There is only a marked difference in size, females being much larger. The chelipeds
arc equal or almost equal in size in both males and females. There is no evidence of differences in the length or
shape of the walking legs or fifth pair of pereopods.
488
P. CASTRO
DISCUSSION. — Parapalicus is a relatively homogenous genus described by Moosa and SERENE (1981) to
include seven species, two of which are now placed in other genera. The description of 10 additional species (eight
here) and the description of new genera, however, necessitates the addition of characters to the brief original
definition.
In defining the genus and separating it from other genera of palicids, MOOSA and SERENE (1981: 24, 25)
described the first pair of walking legs (P2) of Parapalicus as "much slenderer" than the remaining two pairs in
contrast to the other three genera of the subfamily Palicinae they described, where the first pair is only "a little
smaller" than the remaining pairs. This is misleading since, although in Parapalicus the second pair is slightly
more slender (that is, narrower) than the other legs, there is actually no difference in the total length between the
genera.
The long, slender legs and general carapace shape of Parapalicus are closer to Rathbun’s group 2 of the west¬
ern Atlantic species of Palicus Philippi, 1838 (Rathbun, 1918: 184): P. acutifrons (A. Milne Edwards, 1880), P.
cursor (A. Milne Edwards, 1880), P.floridanus (Rathbun, 1918), P. gracilis (Smith, 1883), and P. gracilipes (A.
Milne Edwards, 1880). None of these five species fully agree with the characters used in the definition of
Parapalicus and the other Indo-west Pacific genera of palicids. Nevertheless, Zarenkov (1968) placed his species,
Parapalicus unidentatus (Zarenkov, 1968), under group 1 of RATHBUN. He suggested that several species had inde¬
pendently evolved the long legs of group 2 as a specialization for greater depths (Zarenkov, 1968: 766).
The three western Atlantic species placed by Rathbun (1918: 184) in her group 4 [P. angustus Rathbun,
1897, P. depressus Rathbun, 1897, and P. sica (A. Milne Edwards, 1880)1 are characterized by a structure ("broad,
projecting plates of abdomen and sternum which are conspicuous in dorsal view"; "laminate crest projecting behind
third and fourth walking legs" in WILLIAMS. 1984: 484) that is analogous, and perhaps homologous, to the
episternal process of Parapalicus. None of these Atlantic species, however, have the very slender walking legs with
filiform propodi and dactyli, the single anterolateral tooth on each side of the carapace, and the dentiform suborbital
lobes diagnostic of Parapalicus.
All known species of Parapalicus appear to be restricted to muddy sediments in the Indo-west Pacific region.
They have been collected at depths of 55-778 m. Recently dredged, live specimens have been observed to slide over
sediment by using their long and slender walking legs (B. RICHER DE FORGES, personal communication).
MOOSA and SERENE (1981) designated P. marielae Moosa & Serene, 1981 as the type species of their new
genus Parapalicus. It has been found to be a junior subjective synonym of P. trituberculatus (Chen, 1981)
(see below). The type species is now fixed (under Article 70.3 of the International Code of Nomenclature, fourth
edition. 1999) as Parapalicus trituberculatus (Chen, 1981) misidentified as Parapalicus marielae Moosa & Serene,
1981 in the original designation of Moosa & SERENE (1981).
Key to the species of PARAPALICUS
1. Dactyli of chelipeds with blunt teeth closed together at tip of fingers (Dai & Xu, 1991.
fig. 28-4) . P. nanshaensis Dai & Xu, 1991
— Dactyli of chelipeds with triangular teeth extending along two-thirds to one-half (rarely
one-third) of fingers . 2
2. Dorsal border of cheliped propodi with conspicuous, pointed tubercles (Fig. 17c). Outer
border of carpi of walking legs (P2-4) with conspicuous pointed tubercles (Fig. 17a) .
. P. armatus sp. nov.
— Dorsal border of cheliped propodi smooth, with minute teeth, or two rounded, proximal
tubercles. Outer border of carpi of walking legs (P2-4) smooth or with microscopic
tubercles . 3
3. Only one suborbital lobe, leaving straight border between tooth-like inner lobe and
postorbital angle (Fig. 23b) . P. inanis sp. nov.
— Two dentiform suborbital lobes before postorbital angle . 4
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
489
4. Gap between inner and outer suborbital lobe long, narrow, V-shaped . 5
— Gap between inner and outer suborbital lobes wide, U-shaped, or short when outer lobe
much smaller than outer lobe . 6
5. Anterolateral tooth on side of carapace rounded, bent at slight angle in relation to carapace
(Figs 18a, 19c) . p. clinodentatus sp. nov.
— Anterolateral tooth on side of carapace triangular (Zarenkov, 1968. fig. 3 A) .
. P. unidentatus (Zarenkov. 1968)
6. Dorsal border of cheliped propodus with two proximal tubercles . 7
— Dorsal border of cheliped propodus smooth or with microscopic teeth . 8
7. Rounded tubercle on distal edge of supraorbital border and proximal to postorbital angle,
giving angle bilobed appearance (HOLTHU1S. 1977, fig. 3a) .
. P. elaniticus (Holthuis, 1977)
— Supraorbital angle without conspicuous, distal tubercle proximal to postorbital angle .
. P. ambonensis sp. nov.
8. Outer suborbital lobe very small, less than one-third as long as inner suborbital lobe ... 9
— Outer suborbital lobe at least one-half as long as inner suborbital lobe . 10
9. Supraorbital border sinuous, with two notches but no lobes. Dorsal borders of cheliped
propodi smooth . P. piruensis Moosa & Serene. 1981
— Supraobital border with two very small lobes (Fig. 27a). Dorsal borders of cheliped
propodi with microscopic teeth (Fig. 27c) . P. trispiralis sp. nov.
10. Inner suborbital lobes narrow, pointed, bordered by pointed tubercles (Fig. 24b). Male
first pleopods with helicoidal distal ends (Figs 24d-e). Eyes narrow, small (Fig. 24a) .
. P. microphthalmus sp. nov.
— Inner suborbital lobes not narrow, with rounded tips. Male first pleopods with simple,
blade-like distal ends. Eyes not narrow . 11
11. Eye peduncles covered with small, granular tubercles. Gap between inner and outer
suborbital lobes very broad, nearly circular (Chen, 1981, pi. 2, fig. 1) .
. P. trituberculatus (Chen, 1981)
— Eye peduncles covered with few, soft tubercles. Gap between inner and outer suborbital
lobes broad but V- or L-shaped (Figs 20b, 22b) . 12
12. Dorsal borders of cheliped propodi with rows of microscopic teeth (Fig. 20c). Trigger-like
process at proximal portion of distal part of male first pleopods (Fig. 20e) .
. P. denticulatus sp. nov.
— Dorsal borders of cheliped propodi with barely visible microscopic teeth (Fig. 22c).
Proximal portion of distal parts of male first pleopods smooth, without process (Fig. 22e)
. P. inermis sp. nov.
Parapalicus ambonensis Moosa & Serene, 1 98 1
Figs 16, 19a, 58
Parapalicus ambonensis Moosa & Serene, 1981: 29, figs 2a, 3a, pi. 1, fig, D.
MATERIAL EXAMINED. — Thailand I Andaman Sea coast). Off Phuket Island. Stn 8. 80 m. triangular dredse.
18.02.1998: 1 9 (PMBC).
Indonesia. Haruku Island. Mariel King Memorial Expedition: stn AH 1/4. north of Cape Batang Kapal, 03°36'S.
I28°24'E, shell sand and rubble. 1 10-115 m, 31.05.1970: 1 9 holotype 5.4 x 7.6 mm IRDC CB2733).
490
P. CASTRO
Kai Islands. Karubar: stn DW 29. 05°36'S, 132°56'E. 181-184 m. 26.10.1991: 1 3 (MNHN-B 26963).
Vanuatu. MUSORSTOM 8: stn CP 963. 20°20.IO'S, 169°49.08'E, 400-440 m. 21.09.1994: 2 2 (MNHN-B 27078). —
Stn CP 971. 20°18.87’S. 169°53.12'E. 250-315 m. 21.09.1994: 1 3 (MNHN-B 27079). — Stn CP 1017, 17°52.80'S.
168°26.20'E. 294-295 m, 27.09.1994: 1 3.2 2 (MNHN-B 27080). — Stn CP 1018. 17°52.88'S. 168°25.08'E. 300-
301 m. 27.09.1994: 1 $ (MNHN-B 27081). — Stn CP 1086, 15°36.58'S, I67°16.32'E. 182-215 m. 5.10.1994: 1 2
6.3 x 8.7 mm (MNHN-B 26707). 1 3 5.1 x 7.2 mm (MNHN-B 26708). 1 3 . 1 2 (MNHN-B 27082). — Stn DW 1093.
15°10 91 ’S. 1 67° 11.1 9'E. 275 m. 6.10.1994: 1 3 5.2 x 7.4 mm (MNHN-B 26709). — Stn DW 1097. 1 5°05. 1 3'S.
167°10.76'E. 281-288 m, 7.10.1994: 2 3 (MNHN-B 27083). — Stn CP 1102, I5°03.82'S, 167°08.68'E. 210-208 m.
7.10.1994: 1 2 (MNHN-B 27084). — Stn CP 1134. 15°39.94'S. 167°02.79'E, 230-287 m, 11.10.1994: 1 2 (MNHN-B
27085). — Stn CP 1135, 15°40.50'S. 167°02.43'E. 282-375 m. 11.10.1994: 3 2 (MNHN-B 27086).
Chesterfield-Bellona Plateau. Chalcal 1: stn CP 11. 20°04.40’S, I58047.41'E. 300 m. 07.1984: I <J (MNHN-
B 26964).
MUSORSTOM 5: stn 345. I9°39.70'S, 158°32.40'E, 305-310 m. 16.10.1986: 1 d , 1 2 (MNHN-B 27087).
New Caledonia. MUSORSTOM 4: Grand Passage, stn 173. 19°02.50’S. 163°18.80’E. 250-290 m. 17.09.1985: I 3
5.2 x 7.3 mm (MNHN-B 26706).
"Kandjar" . dredging between 22°40'-22°50'S and 167°10'-167°30'E, 200-350 m, 7-10.10.1986: 1 2 (MNHN-B
26966).
Bathus 1: stn DW 640, 21°52.33'S, 166°47.93’E, 174 m, 10.03.1993: 1 2 (MNHN-B 27088). — Stn CP 713,
21°45.28'S, 166°36.83'E. 250 m, 19.03.1993: 1 2 (MNHN-B 27089).
Bathus 3: stn DW 836, 23°02'S, 166°59’E. 295-306 m, rocks and gravel. 30.1 1.1993: 1 3 , 2 2 (MNHN-B 27090).
Bathus 4: stn DW 901. 19°02.72'S, 163°15.39'E. 297 m. 4.08.1994: 3 2 (MNHN-B 27091). — Stn CP 905.
19°02'S. 163°15’E. 294-296 m. 4.08.1994: 3 3 . 11 2 (MNHN-B 26754).
Loyalty Islands. MUSORSTOM 6: stn CP 419. 20°41.65'S, 167°03.70’E. 283 m, 16.02.1989: I 3 (MNHN-B
27092). — Stn DW 452, 21°00.30'S, 167°25.50'E. 300 m, 20.02.1989: 1 2 (MNHN-B 27093). — Stn DW 462.
21°05. 10’S. 167°26.85'E. 200 m. 21.02.1989: 1 2 6.7 x 9.1 mm (MNHN-B 26710).
Futuna Island, MUSORSTOM 7: stn CP 515, 14°13'S. 178°10'W. 224-252 m. 12.05.1992: I 2 (MNHN-B 26965).
Types. — Holotype : 12 5.4 x 7.6 mm. Mariel King Memorial Expedition, stn AH 1/4 (RDC
CB2733).
Type Locality. — North of Cape Batang Kapal. Haruku Island, Moluccas. Indonesia, 03°36'S, 128°24'E,
shell sand and rubble, 110-115 m.
Diagnosis. — Carapace (Figs 16a, 19a; MOOSA & SERENE, 1981, fig. 2a) with anterolateral tooth on each
side; tooth with truncate outer border and short, rounded upper border. Small, tooth-like tubercle between
anterolateral tooth and postorbital angle. Supraorbital borders each with three notches. Suborbital borders each with
narrow, V-shaped gap between inner and outer suborbital lobes; outer lobe much smaller than inner lobe. Dorsal
borders of cheliped propodi each with two proximal tubercles, distalmost rounded, conspicuous. Abdomen of
mature males (Fig. 16b) with two triangular tubercles on segment 3 and larger one on segment 4. Male first
pleopods (Fig. 16c) with helicoidal basal parts; each distal portion beak-like, with spinulcs along dorsal margin.
Abdomen of mature females (Fig. 16d) with one transverse ridge along each segment 1-3 (segment 3 slightly
constricted twice medially); short ridge at each outer edge of segment 4.
Discussion. — P. ambonensis was until now know from only one female specimen. Only the last pair of legs
was present, MOOSA & Serene (1981) did not give any information on the morphology of the chelipeds or of the
suborbital borders, and the only photograph given is of poor quality. The examination of the holotype (RDC
CB2733) and the presence of its distinctive characters in specimens collected from the Andaman Sea to the Coral
Sea confirms its status as a distinctive species. This is the first time the male first pleopod is described and
illustrated.
The examination of the holotype also showed that one diagnostic character given in its description, the presence
of "teeth" along the inner borders of the third maxillipeds, was a misinterpretation of the serrate nature of the
borders. Although only the right maxilliped remains in the holotype, its inner border does not have separate,
distinctive teeth as shown in the figure of the left maxilliped (Moosa & Serene, 1981, fig. 3a).
Other characters useful in its identification are the presence of a basal antennal segment that is expanded dislally
as a rounded process and a small, tooth-like tubercle between each anterolateral tooth and the postorbital angle.
Its size varies, sometimes hardly visible on one side but larger on the opposite side of the same specimen.
Source : MNHN, Paris
1ND0-WEST PACIFIC SPECIES OF PALICID CRABS
491
Fig. 16. — Parapalicus ambonensis Moosa & Serene, 1981: a-b. 2 6.3 x 8.7 mm. north of Malekula Island, Vanuatu.
Musorstom 8, stn CP 1086, 182-215 m (MNHN-B 26707): a. carapace and pereopods. dorsal surface; b. abdomen. —
c, 3 5.2 x 7.3 mm, Grand Passage, New Caledonia. Musorstom 4. stn 173, 250-290 m (MNHN-B 26706): abdomen.
— d. 3 5.2 x 7.4 mm, east of Espiritu Santo Island, Vanuatu, MUSORSTOM 8, stn DW 1093, 275 m (MNHN-B 26709):
left male first pleopod, ventral view.
The fifth pair of pereopods (P5) are short (0.8 CL), each merits with 3-4 small spines along the posterior
border, the propodus with 4-8 spines along the posterior border, and the dactylus with 2-3 spines along the
posterior border and one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
492
P. CASTRO
P. ambonensis shares several features with P. nanshaensis Dai & Xu, 1991. a species known from only one
female specimen (see below). Both species have a similar carapace shape and a cheliped merus that is armed with
only two proximal tubercles. P. nanshaensis, however, has very characteristic blunt, closed-together teeth on the
cheliped fingers (Dai & Xu, 1991. fig. 28-4) and only one slight notch on each supraorbital border. The
morphology of the suborbital borders, male first pleopods, and male abdomen of P. nanshaensis remain unknown
but the holotype was not available for examination.
P. ambonensis shares with P. armatus sp. nov. and P. trispiralis sp. nov. the presence of pointed tubercles on
the male abdomen. The dorsal borders of the chelipcds of these two species are armed with conspicuous tubercles
(P. armatus) or microscopic teeth (P. trispiralis).
Size. — Maximum size among specimens examined: 6.3 x 9.7 mm (female, MNHN-B 26754); 5.5 x 7.3 mm
(male, MNHN-B 26754).
DISTRIBUTION. — Known from the Andaman Sea (off Phuket island. Thailand), Banda Sea (off Kai Islands,
Indonesia), Haruku Island (near Ambon. Indonesia) (MOOSA & SERENE, 1981), Coral Sea (Vanuatu. Chesterfield-
Bellona Plateau. New Caledonia, and the Loyalty Islands), and Futuna Island, southwestern Pacific (Fig. 58).
Depth: known reliably between 80 and 400 ; also collected in a trawl between 400 and 440 m.
Parapalicus armatus sp. nov.
Figs 17, 19b. 57
Material EXAMINED. — Vanuatu. — Musorstom 8: stn CP 963, 20°20'S. 169°49'E. 400-440 m. 21.09.1994:
I <3 holotype 6.5 x 9.0 mm (MNHN-B 26698).
Lovaltv Islands. — Musorstom 6: stn DW 457, 21°00.42'S. 167°28.71'N, 353 m, 20.02.1989: 1 6 paratype
6.5 x 8.7 mm (MNHN-B 26699). — Stn CB 481, 21°21.85'S, 167°50.30’E. 300 m. 23.02.1989: 1 $ paratype 8.1 x
10.6 mm (MNHN-B 26700), 1 3 . 2 2 (MNHN-B 26823).
Types. — Holotype: 1 3 6.5 x 9.0 mm, MUSORSTOM 8, stn CP 963 (MNHN-B 26698).
Paratypes: I 8 6.5 x 8.7 mm, MUSORSTOM 6, stn DW 457 (MNHN-B 26699); I 2 8.1 x 10.6 mm,
Musorstom 6, stn CB 481 (MNHN-B 26700).
Type Locality. — Off south coast of Aneityum (Anatom) Island. Vanuatu. 20°20'S, 169°49'E. 400-440 m.
Diagnosis. — Carapace (Figs 17a, 19b) with small, triangular anterolateral tooth on each side; conspicuous
tubercles along posterior border. Supraorbital borders each with three notches. Suborbital borders (Fig. 17b) each
with narrow, V-shaped gap between inner and outer suborbital lobes; outer lobe rounded. Dorsal borders of cheliped
propodi (Fig. 17c) each with 3-4 rows of pointed tubercles. Meri of second and third pairs of walking legs (P3-4)
with tubercles along upper and lower edges. Abdomen of mature males (Fig. 17d) with two triangular tubercles on
segment 3, similar one on segment 4. Male first pleopods (Figs 17e-f) with helicoidal basal parts; each distal part
long, blade-like, rising from thickened portion of basal part. Abdomen of mature females with one transverse ridge
along each segment 1-3 (segment 3 slightly constricted twice medially); segment 4 with short ridge at outer edges.
Description. — Carapace (Figs 17a, 19b) wider than long (CW/CL = 1.3- 1. 4); dorsal surface covered with
coarse granules. Anterolateral borders of carapace each with very small triangular tooth covered with fine granules.
Two small, low bosses, also covered with granules, on branchial region posterior to each tooth. Larger females
with four conspicuous bosses (two large median lobes flanked by smaller one on each side) on metagastric region.
Posterior border of carapace with 8-12 protuberant tubercles with rounded tips and many small tubercles with
pointed or rounded tips. Episternal processes conspicuous, bordered by very small teeth, with anterior tubercle in
some specimens. Posterior border of carapace with scattered plumose setae.
Frontal border of carapace divided into two small lobes divided by narrow V-shaped notch; triangular or slightly
rounded tips. Borders between frontal lobes and supraorbital borders rounded, marked by slight undulation medially.
Supraorbital borders each with three distinct notches. Postorbital angles short, slightly pointed. Cornea of eyes
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
493
Fig. 17. — Parapalicus annatus sp. nov., 8 holotype 6.5 x 9.0 mm, south of Aneityum Island. Vanuatu. MUSORSTOM 8.
sin CP 963, 400-440 m (MNHN-B 26698): a, carapace and pereopods. dorsal surface; b. suborbital border: c. left
cheliped, outer surface; d, abdomen; e. left male first pleopod, ventral view; f, left male first pleopod, distal part,
lateral (inner side) view.
Source : MNHN. Paris
494
P. CASTRO
dorsoventrally flattened, much wider than base of short eye peduncle; each peduncle with two soft, distal tubercles
(ventral largest).
Suborbital borders (Fig. 17b) each with narrow, V-shaped gap between inner and outer suborbital lobes. Outer
lobe rounded, shorter than pointed inner lobe. Pterygostomial lobes project slightly ventrally, rounded (slightly
pointed in some specimens), shorter than each inner suborbital lobe.
Each basal antennal segment expanded distally as rounded process; flagellum long with conspicuous, simple
setae. Inner border of ischia of third maxillipeds sinuous to serrate but without defined teeth; surface coarsely
granular, upper borders rounded. Meri much narrower than ischia; rounded, narrow upper borders.
Dorsal and outer borders of cheliped propodi (Fig. 17c) each with 3-4 rows ol conspicuous tubercles, those
of uppermost row with pointed or broadly acute tubercles (more conspicuous in males). Distal two-thirds of
each finger with 4-5 triangular teeth (shorter in some specimens). Carpi short, outer borders with conspicuous,
broad tubercle across distal portion and pointed tubercles throughout; meri slender, outer border with pointed
tubercles.
First three pairs of walking legs (P2-4; Figs 17a, 19b) very slender, with elongate meri and filiform propodi
and dactyli. Upper and lower borders of meri with small, similarly pointed tubercles (smaller on P2); distal most
tubercle on each anterior border much wider at base, higher, with rounded tip directed distally. Anterior
and posterior borders of carpi of second and third walking legs (P3-4) with rounded tubercles; borders ol propodi
and dactyli of all walking legs (P2-4) entire (propodi of P3-4 with microscopic tubercles, proximal portion with
visible tubercles, particularly P4). Each P2 with one dorsal carina each on carpus and dactylus. Each P3 with one
dorsal carina on merus; two dorsal carinae on carpus; one dorsal, one ventral carina each on propodus and dactylus.
Each P4 with two dorsal carinae on carpus; one dorsal and one ventral carina each on propodus and dactylus. Meri
of all legs with scattered plumose setae; dorsal surface of carpi and propodi of P3-4 with rows ol plumose setae;
dorsal surfaces of dactyli of P3-4 each with one row of conspicuous plumose setae along anterior border.
First pair of walking legs (P2) much shorter and slender than second and third pairs (P3-4); third pair about as
long as second pair. Fifth pair of pereopods (P5) short (0.8 CL), very slender; each merus with several tubercles,
3-4 spines, scattered plumose setae along posterior border; each propodus with 6-8 spines along posterior border,
scattered plumose setae along anterior border; each dactylus with three spines along posterior border, one terminal
pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males (Fig. 17d) with segments 3-5 fused, two triangular tubercles on segment 3, one on
4. Male first pleopods (Figs 17e-f) with helicoidal basal parts; each distal part long, blade-like rising from
thickened portion of basal part, which has plumose setae.
Abdomen of mature females with segments 3-5 fused. One transversal ridge along segments 1-2, each bordered
by small, pointed tubercles; segment 3 of fused segment 3-5 with ridge slightly constricted twice medially; short
ridge at each outer edge of segment 4.
DISCUSSION. — P. armatus can be easily distinguished from the other known members of the genus by the
presence of conspicuous, pointed tubercles along the dorsal surface of the cheliped propodi, the posterior border ol
the carapace, and on the anterior and posterior borders of the meri of walking legs. The abundance of pointed
tubercles through the body and appendages give individuals a rough appearance. It shares with P. inanis sp. nov.
a greatly reduced anterolateral tooth on each side of the carapace; with P. ambonensis and P. trispiralis sp. nov.
pointed tubercles on the male abdomen.
Size. — Maximum size among the six specimens examined in this study: 8.1 x 10.6 mm (female paratype,
MNHN-B 26700); 6.5 x 9.0 mm (male holotype, MNF1N-B 26698).
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
495
Etymology. — From armatus, Latin for furnished with weapons, in reference to the diagnostic pointed
tubercles along the dorsal surface of the chelipeds, meri of walking legs, and posterior border of the carapace.
Distribution. — Known only from the Coral Sea (Vanuatu and the Loyalty Islands) (Fig. 57). Depth: known
reliably between 300 and 400 m; also collected in a trawl between 400 and 440 m.
Parapalicus clinodentatus sp. nov.
Figs 18, 19c, 57
Material EXAMINED. — Vanuatu. Musorstom 8: stn CP 1092, 15°10.80'S, 167°12.33’E. 314-321 m,
6.10.1994: 1 <3,2 2 (MNHN-B 26954). — Stn CP 1135, 15°40.50’S, 167°02.43'E. 282-375 m. 11.10.1994: 5 2
(MNHN-B 26955).
New Caledonia. Biocal: stn CP 105, 21o30.71'S, 166°21.72'E. 330-335 m, 8.09.1985: 1 juv. 2 (MNHN-B
28006).
Bathus 1: stn CP 670, 20°54'S, 165°53'E. 394-397 m, 14.03.1993: 1 <3 , 1 2 9.0 x 13.1 mm (MNHN-B 26752). —
Stn CP 701, 20°57.54'S, I65°35.86'E. 302-335 m, 18.03.1993: 3 3,42 (MNHN-B 26956). — Stn 707. 21°42'S.
166°35'E, 347-375 m, 19.03.1993: 5 <5,102 (MNHN-B 27000). — Stn 708, 21°43.05'S, 166°38.57'E, 550-580 m.
19.03.1993: 1 2 (MNHN-B 26957). — Stn CP 710, 21°43.16'S, 166°36.35'E, 320-386 m, 19.03.1993: 1 <3 paratype
5.9 x 8.7 mm (MNHN-B 26702), 1 2 paratype 8.5 mm x 12.0 mm (MNHN-B 26703), 13 <3 . 9 2 (MNHN-B 27001). —
Stn CP 711. 21°43.00'S, 166°35.57'E, 315-327 m, 19.03.1993: 3 d , 2 2 (MNHN-B 26958).
Bathus 2: stn CP 742, 22°33.45'S, 166°25.86'E, 340-470 m, 14.05.1993: 8 <5, 13 2 (MNHN-B 26753), 1 2 8.6 x
12.2 mm (MNHN-B 26705), 1 <3 paratype 5.7 x 8.6 mm, 1 2 paratype 8.1 x 11.3 mm (USNM), 1 <3 paratype 7.3 x
10.6 mm, 1 2 paratype 8.0 x 11.3 mm (ZRC 199.1431-1432). — Stn CP 743. 22°35.56'S. 166°26.23'E. 713-950 m.
14.05.1993: 1 <3 , 1 2 (MNHN-B 26959). — Stn CP 756. 22°22’S. 166°13'E, 672 m. 16.05. 1993: 1 <3 paratype 6.4 x
9.3 mm (MNHN-B 26704).
Halipro I: stn CP 851, 21°43'S, 166°37'E. 314-364 m, 19.03.1994: 1 <3 holotype 6.2 x 9.0 mm (MNHN-B 26701).
5 <3,5 2 (MNHN-B 27003). — Stn CC 856, 21°44'S. 166°37'E. 311-365 m. 20.03.1994: I 2 (MNHN-B 27004). —
Stn 868, 2 1 0 1 4'S, 165°55’E, 430-550 m, 23.03.1994: 1 <3 (MNHN-B 27005).
Bathus 4: stn CP 897, 20°15.93'S. I63°51.75'E. 305-350 m. 3.08.1994: 1 <3 (MNHN-B 26960). — Stn CP 899.
20°16.68'S, 1 63°50.26'E, 500-600 m, 3.08.1994: 1 2 (MNHN-B 26961). — Stn CP 946. 20°33.8I'S. 164°58.35'E.
386-430 m. 10.08.1994: 1 <3 (MNHN-B 26962).
Fiji. Musorstom 10: stn 1348, 17°30.29'S, 178°39.63'E. 353-390 m, 11.08.1998: 2 2 (MNHN-B 26751).
Bordau I: stn CP 1427, 17°16'S. 179o01'W, 364-369 m. 1.03.1999: 2 <3 . 1 2 (MNHN-B 27135).
Types. — Holotype-- 1 <3 6.2 x 9.0 mm. HALIPRO 1, stn CP 851 (MNHN-B 26701).
Paratypes : I <3 5.9 x 8.7 mm. BATHUS 1, stn CP 710 (MNHN-B 26702); 1 2 8.5 x 12.0 mm, B.ATHUS 1,
stn CP 710 (MNHN-B 26703); 1 <3 6.4 x 9.3 mm, Bathus 2, stn 756 (MNHN-B 26704); 1 2 paratype 8.6 x
12.2 mm, Bathus 2, stn CP 742 (MNHN-B 26705); 1 <3 paratype 5.7 x 8.6 mm, 1 2 paratype 8.1 x 1 1.3 mm.
Bathus 2, stn CP 742 (USNM); 1 <3 paratype 7.2 x 10.6 mm. 1 2 paratype 8.0 x 11.3 mm. Bathus 2.
stn CP 742 (ZRC 1999.1431-1432).
Type Locality. — Off east coast of New Caledonia, 21°43'S, 166°37'E, 314-364 m.
DIAGNOSIS. — Carapace (Figs 18a, 19c) with rounded anterolateral tooth on each side; tooth posteriorly bent at
a slight angle in relation to carapace. Supraorbital borders irregular, undulating, without defined notches.
Suborbital borders (Fig. 18b) each with very narrow, V-shaped gap between inner and outer suborbital lobes.
Dorsal borders of cheliped propodi each with carina-like row of irregular tubercles. Abdomen of mature males
(Fig. 18c) with two low tubercles on segment 3, similar one on segment 4. Male first pleopods (Fig. 18d) with
helicoidal basal parts; each distal part with two curved, unequal, pointed tips that cross each other. Abdomen of
mature females with one transverse ridge along each segment 1-4.
Colon Specimens that were examined only nine months after collection (MNHN-B 2675 I ) showed that some
of the tubercles on the meri and carpi of the walking legs, the meri of chelipeds, and on the dorsal surface of the
carapace (mostly along the anterior border) were orange.
496
P. CASTRO
Fig. 18. — Parapalicus clinodentatus sp. nov., <J holotype 6.2 x 9.0 mm. off east coast of New Caledonia. Halipro 1.
stnCP 851. 314-364 m (MNHN-B 26701): a. carapace and pereopods, dorsal surface; b. suborbital border;
c, abdomen; d. left male first pleopod. ventral view.
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
497
DESCRIPTION. — Carapace (Figs 18a, 19c) wider than long (CW/CL = 1.4- 1.5); dorsal surface covered with
coarse granules. Anterolateral borders of carapace each with granular, rounded to triangular tooth bent posteriorly in
relation to carapace. Two granular, rounded bosses (one high, close to carapace border plus less conspicuous one),
on branchial region posterior to each tooth. Two large but low bosses on metagastric region. Slightly protuberant,
granular tubercle at junction of posterolateral and posterior borders. Episternal processes oblong, with rounded tip.
Larger specimens with four similar but slightly smaller, granular tubercles that protrude along posterior border.
Posterior border with plumose setae.
Frontal border of carapace divided into two small lobes divided by narrow V-shapcd notch; slightly rounded tips.
Borders between frontal lobes and supraorbital borders slightly pointed medially separates straight anterior half and
slightly raised posterior half. Supraorbital borders irregular, undulating, without well-defined notches (slight notch
before postorbital angle in some specimens). Postorbital angles short, rounded (slightly pointed in some
specimens). Cornea of eyes dorsoventrally flattened, much wider than base of eye peduncle; peduncles without
well-defined tubercles.
Suborbital borders (Fig. 18b) each with very narrow, V-shaped gap between inner and outer suborbital lobes.
Outer lobe pointed, narrow, slightly shorter than pointed inner lobe. Pterygostomial lobes project slightly
ventrally, elongate, pointed, slightly shorter than each inner suborbital lobe.
Each basal antennal segment expanded distally as narrow, pointed process; flagellum long with conspicuous,
simple setae. Inner border of ischia of third maxillipeds sinuous to serrate but without defined teeth; surface
coarsely granular, upper borders rounded. Meri much narrower than ischia; narrow, pointed (in inner direction)
upper borders.
Dorsal borders of cheliped propodi (Fig. 18a) each with carina-like row of irregular tubercles; one additional row
of low tubercles along outer surface. Distal half of each finger with 4-5 triangular teeth. Carpi short, outer borders
with many pointed tubercles throughout; meri slender, outer border each with 1-2 rows of low tubercles.
First three pairs of walking legs (P2-4; Figs 18a, 19c) very slender, with very elongate meri. filiform propodi and
dactyli. Upper and lower borders of meri with small, similarly pointed tubercles (smaller on P2. slightly larger
tubercles proximally in P3 of some specimens); distalmost tubercle on each anterior border wider at base, higher,
with rounded tip slightly directed distally (particularly in P3). Anterior and posterior borders of carpi, propodi. and
dactyli of all walking legs (P2-4) entire, without tubercles (propodi of P3-4 with microscopic tubercles). Each P3
with two dorsal carinae on carpus; one dorsal carina on propodus; one dorsal, one ventral (very slight) carina on
dactylus. Each P4 with two dorsal carinae on carpus; one dorsal carina on propodus; one dorsal, one ventral carina
each on propodus and dactylus. Meri of all legs with scattered plumose setae; dorsal surfaces of carpi and propodi of
P2-4 with rows of plumose setae; dorsal surfaces of dactyli of P2-4 each with one row of conspicuous plumose
setae along anterior border.
First pair of walking legs (P2) much shorter and slender than second and third pairs (P3-4); third pair about as
long as second pair. Fifth pair of pereopods (P5) short (0.9 CL), very slender; each merus with microscopic
tubercles, scattered plumose setae along posterior border; each propodus without setae or with scattered plumose
and/or simple setae; each dactylus with 2-3 short spines along posterior border, long simple setae along anterior
border, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males (Fig. 18c) with segments 3-5 fused, two slightly raised tubercles on segment 3 (plus
low central prominence), similar one on 4. Male first pleopods (Fig. 18d) with helicoidal basal parts; each distal
part with two curved, unequal, pointed tips that cross each other.
Abdomen of mature females with segments 3-5 fused. One transversal ridge along each segment 1-2, two along
proximal portion of fused segment 3-5 (very low one may be present on 5).
498
P. CASTRO
Pig 19. — a, Parapalicus ainbonensis Moosa & Serene, 1981, 8 5.4 x 7.6 mm. Grand Passage, north of New Caledonia.
Bathus 4 stn CP 905, 294-296 m (MNHN-B 26754): dorsal view. — b. Parapalicus armatus sp. nov.: 2 paratype 8.1
x 10.6 mm. Loyalty Islands, MusORSTOM 6, stn DW 481, 300 m (MNHN-B 26700): dorsal view. — c. Parapalicus
clinodeniatus sp. nov., 2 9.0 x 12.7 mm, off southeast coast of New Caledonia, Bathus 2. stn CP 742, 340-470 m
(MNHN-B 26753): dorsal view. — d, Parapalicus inanis sp. nov., 2 paratype 4.9 x 6.2 mm. Chesterfield-Bellona
Plateau, MUSORSTOM 10, stn 285, 245-255 m, (MNHN-B 26722): dorsal view.
Discussion. — P. clinodentatus is close to P. unidentatus (Zarcnkov, 1968) since both species share several
key characters: a narrow, V-shaped gap between the inner and outer suborbital lobes, a carina-like row ol it tegular
tubercles on the dorsal borders of the chelipeds, two salient teeth on the sides of the carapace, two curved and
pointed tips that cross each other on the distal part of the male first pleopods, and relatively low tubercles on the
male abdomen. P. clinodentatus may be readily differentiated from P. unidentatus, however, by having much
longer walking legs (total length of P3 varied 2. 9-3. 3 CL in P. unidentatus whereas it is 2.7 CL in
P. clinodentatus), rounded anterolateral teeth each with its posterior edge bent downward in relation to the rest of
the carapace (triangular in shape and a straight continuation of the carapace in P. unidentatus), irregular subraorbital
borders (two clear notches in P. unidentatus). and unequal pointed tips in the male first pleopods (equal in
P. unidentatus).
Size. — Maximum size among specimens examined: 9.0 x 13.1 mm (female, MNHN-B 26752); 7.3 x
10.4 mm (male. MNHN-B 26753).
ETYMOLOGY. — From clino, Latin for bend or slant, and dens. Latin for tooth, in reference to the diagnostic
position of the anterolateral tooth, which is bent in relation to the carapace.
Source MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
499
Distribution. — Known From the Coral Sea (Vanuatu and New Caledonia) and Fiji (Fig. 57). Depth: known
reliably between 320 and 713 m; also collected in trawls between 282-375 and 713-950 m.
Parapalicus denticulatus sp. nov.
Figs 20, 21a, 57
Material EXAMINED. — Vanuatu. Musorstom 8: stn CP 1077, 16°04'S, 1 67°06'E, 180-210 m, 5.10.1994: 2 S ,
2 2 (MNHN-B 26840). — Stn CP 1117, 15°09'S. 166°53'E, 170-220 m. 9.10.1994: 1 2 (MNHN-B 26842). —
Stn DW 1128. 16°02'S, 166°38'E, 778-811 m. 10.10.1994: 1 2 (MNHN-B 26841).
New Caledonia. Bathus 1: stn 667, 20°57'S, 165°34'E. 205-212 m, 14.03.1993: 1 6.9 9 (MNHN-B 26843),
I 6 5.0 x 7.4 mm (MNHN-B 26711), I 6 paratype 5.2 x 7.6 mm (MNHN-B 26712), 1 6 holotype 5.2 x 8.0 mm
(MNHN-B 26711). 1 2 paratype 7.3 x 11.0 mm (MNHN-B 26713). — Stn CP 668, 20°57.21'S. 165°34.57'E, 205-
219 m, 14.03.1993: 15 3.9 2 (MNHN-B 27002), 1 6 (MNHN-B 26839), 1 6 paratype 5.0 x 7.6 mm. 1 2 paratype
6.8 x 10.2 mm (USNM), 1 6 paratype 5.1 x 7.4 mm, 1 2 paratype 7.3 x 10.8 mm (ZRC 1999.1433-1434). —
Stn CP 669, 20°57'S, 165°35'E. 255-280 m, 14.03.1993: 4 3.42 (MNHN-B 26756). — Stn DE 700. 20°57’S.
165°34'E, 160-222 m. 18.03.1993: 1 2 (MNHN-B 26844). — Stn CP 713, 21°45'S. 166°36'E. 250 m, 19.03.1993:
3 <3,7 2 (MNHN-B 26755).
HALIPRO 1: stn CP 863. 21°31'S. 166°20'E. 190-227 m. 22.03.1994: 2 3,39 (MNHN-B 26845).
BATHUS 4: stn CP 953, 21°45'S, I66°36'E. 220-234 m, 11.08.1994: 1 2 7.5 x 10.6 mm (MNHN-B 26715), I 2
(MNHN-B 26846). — Stn 954. 21°44'S, 166°35'E, 255-250 m. 11.08.1994: 1 2 parasitized by bopyrid (MNHN-B
96847).
Loyalty Islands. MUSORSTOM 6: sin DW 410, 20°38.05'S, 167°06.65'E. 490 m. 15.02.1989: 1 2 (MNHN-B
26848).
Types. — Holotype: 1 6 5.0 x 7.4 mm. Bathus 1. stn CP 667 (MNHN-B 2671 1).
Paratypes : 1 S 5.2 x 7.6 mm, BATHUS 1, stn CP 667 (MNHN-B 26712); 1 2 7.3 x 11.0 mm. BATHUS 1.
stn 667 (MNHN-B 26713); 1 $ paratype 5.2 x 8.0 mm, Bathus 1, stn 667 (MNHN-B 26714); I 2 paratype
7.5 x 10.6 mm, Bathus 4, stn CP 953 (MNHN-B 26715); 1 S 5.0 x 7.6 mm. 1 2 6.8 x 10.2 mm, Bathus 1.
stn CP 668 (USNM); 1 6 5.1 x 7.4 mm, 1 2 7.3 x 10.8 mm. BATHUS 1. stn CP 668 (ZRC 199.1433-1434).
Type Locality. — Off east coast of New Caledonia, 20°57.23’S, 165°34.59'E. 205-212 m.
DIAGNOSIS. — Carapace (Figs 20a, 21a) with anterolateral tooth on each side; tooth with truncate outer border
and sinuous, nearly straight upper border. Supraorbital borders each with two very slight notches. Suborbital
borders (Fig. 20b) each with broad, V-shaped gap between inner and outer suborbital lobes; outer lobe longer than
postorbilal angle. Dorsal borders of cheliped propodi (Fig. 20c) with rows of microscopic teeth: proximal tubercle
on propodus with similar microscopic teeth. Abdomen of mature males (Fig. 20d) with two rounded, elongate
tubercles on segment 3; similar but lower one on segment 4. Male first pleopods (Fig. 20e) with helicoidal basal
parts; each distal part straight, blade-like, with trigger-like process at base. Abdomen of mature females with one
transverse ridge along each segment 1-4 (slightly constricted twice medially in segment 3).
Description. — Carapace (Figs 20a, 21a) wider than long (CW/CL - 1.4- 1.5); dorsal surface covered with
coarse granules. Smooth sulcus between hepatic and branchial regions connecting with deep fissure between
anterolateral tooth and large swelling (with dorsal, rounded boss) along carapace border posterior to postorbital
angle. Anterolateral borders of carapace each with tooth covered with fine granules, truncate outer border, tip
slightly directed anteriorly, and sinuous, nearly straight upper border. One large, elongate to rounded, granular boss
close to carapace border on branchial region posterior to each anterolateral tooth. Two large, low bosses on
metagastric region. Episternal processes oblong. Posterior border of carapace with 6-8 slightly protuberant
tubercles, plumose setae.
Frontal border of carapace divided into two very broad lobes with straight margins sometimes marked by very
slight median notch; lobes divided by narrow V-shaped notch. Borders between frontal lobes and supraorbital
borders rounded, slightly raised in posterior portion. Supraorbital borders each with two very slight notches, one
before postorbital angle. Postorbital angles very short (shorter than outer lobe of suborbital border), with slightly
500
P. CASTRO
pointed lip. Cornea of eyes dorsoventrally flattened, much wider than base of short eye peduncle; peduncles without
conspicuous tubercles.
Fig. 20. — Parapalicus denticulatus sp. nov.: a-d. 3 holotype 5.0 x 7.4 mm, off south coast of New Caledonia, Bathus 1.
stn CP 667, 205-212 m (MNHN-B 26711): a, carapace and pereopods. dorsal surface; b. suborbital border; c, right
cheliped, upper surface; d, abdomen. — e, 3 paratype 5.2 x 7.6 mm, off south coast of New Caledonia, Bathus 1.
stn CP 667 (MNHN-B 26712): left male first pleopod. ventral view.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
501
Suborbital borders (Fig. 20b) each with broad, V-shaped gap between inner and outer suborbital lobes. Outer
lobe pointed (obliquely directed in some specimens), shorter than pointed inner lobe. Pterygostomial lobes nearly
straight, triangular, slightly shorter than each inner suborbital lobe.
Each basal antennal segment expanded distally as slightly pointed process; flagellum long with conspicuous,
simple setae. Inner border of ischia of third maxillipeds sinuous to serrate but without defined teeth; surface
coarsely granular, upper borders rounded. Meri much narrower than ischia; narrow, pointed (in inner direction)
upper borders.
Dorsal borders of cheliped propodi (Fig. 20c) with rows of microscopic teeth; proximal portion of propodi with
tubercle with denticulate margin. Distal half of each finger with 4-5 triangular teeth. Carpi short, meri slender,
outer borders of both with microscopic teeth throughout.
First three pairs of walking legs (P2-4; Figs 20a, 21a) very slender, with very elongate meri, filiform propodi
and dactyli. Upper and lower borders of meri with small, similarly pointed tubercles (slightly larger tubercles
proximally in some specimens); distalmost tubercle. on each anterior border wider at base, higher, with
conspicuously pointed tip directed distally in P3-4 (similar tubercle but not conspicuously pointed in P2). Anterior
and posterior borders of carpi, propodi, and dactyli of all walking legs (P2-4) entire. Each P2 with one dorsal (very
slight) carina on dactylus. P3-4 each with one dorsal carina on carpus, propodus (very slight), and dactylus. Meri of
all legs with scattered plumose setae; dorsal surfaces of propodi of P3-4 with rows of plumose setae; dorsal
surfaces of dactyli of P2-4 each with one row of conspicuous plumose setae along anterior border.
First pair of walking legs (P2) much shorter and slender than second and third pairs (P3-4); third pair about as
long as second pair. Fifth pair of pereopods (P5) short (0.8-0.9 CL), very slender; each merus with scattered
plumose setae along posterior border; each propodus with smooth borders, scattered plumose setae along anterior
and posterior borders; each dactylus with 1-4 short spines increasing in length distally along posterior border, long
simple setae along anterior and posterior borders, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males (Fig. 20d) with segments 3-5 fused, two rounded, elongate tubercles on segment 3,
similar but slightly lower tubercle on segment 4. Male first pleopods (Fig. 20e) with helicoidal basal parts; each
distal part straight, blade-like, corneus distal part; proximal, ventrally pointed, trigger-like process.
Abdomen of mature females with segments 3-5 fused. One ridge along each segment 1-2. two along proximal
portion of fused segment 3-5 (slightly constricted twice medially in segment 3).
DISCUSSION. — P. denticulatus is very close to P. inermis sp. nov. It differs from the later by the presence of
numerous, very small teeth on the dorsal surface of the propodi of the chelipeds (nearly smooth in P. inermis), a
trigger-like process at the base of a shorter and straight distal part in each male first pleopod. much lower tubercles
on the male abdomen, and slightly longer second walking legs in relation to carapace length (2. 8-2.9 CL against
3. 0-3. 5 CL in P. inermis). There is also a difference in size, adult P. denticulatus being larger than adult
P. inermis. In P. inermis the average carapace length of all specimens available for measurement was 5. 1 mm in
males (4. 1-5.3 mm, N = 22) and 6.6 mm in females (5. 1 -7.5 mm. N = 33) in contrast to 4,1 mm in males (3.7-
4.4 mm. N = 1 1) and 5.7 mm in females (4. 1-6.3 mm; N = 1 12) in P. inermis. Morphological differences (most
significantly between the male first pleopods) and the large number of morphologically similar specimens collected
in separate geographical locations indicate that P. denticulatus and P. inermis are different species and not merely
populations of different age groups that belong to only one species.
There are also some similarities with P. ambonensis. As in P. denticulatus, the anterolateral teeth of
P. ambonensis have truncate outer borders. The upper borders, however, are short and rounded (which makes the
teeth look conspicuously narrow), not long and nearly straight as in P. denticulatus. P. ambonensis can be easily
Source
502
P. CASTRO
identified by a very short second suborbital tooth, the presence of two tubercles along the dorsal border of each
cheliped propodus. and male first pleopods with a beak-like distal part (Fig. 16c).
P. denticulatus also shares with P. unidentatus (Zarenkov, 1968) the morphology of the dorsal surface ol its
carapace, which is covered (except the cervical groove and areas around the gastric portion) by granular bosses and
anterolateral borders that are each armed with a triangular tooth and a conspicuous blunt boss at the branchial
region posterior to the anterolateral tooth. The two species can be distinguished by the morphology of their male
first pleopods (much more complex in P. unidentatus, with each distal part consisting ot two pointed tips that
cross each other; MOOSA & Serene, 1981, fig. 4), the ornamentation of the cheliped propodi (carina-like row of
irregular tubercles in P. unidentatus), and the gap between the inner and outer suborbital lobes (U-shaped in
P. denticulatus, narrow and V-shaped in P. unidentatus-, Zarenkov, 1968, fig. 3B).
P. denticulatus also shares with P. trituberculatus (Chen, 1981) similarities in carapace shape and in the
presence of microscopic teeth on the dorsal borders of the cheliped propodi. They differ from each other by the
morphology of their male first pleopods (pointed distal parts that are bordered with spinules along the ventral
borders in P. trituberculatus'. Figs 28a-b), the gap between the inner and outer suborbital lobes (much broader,
almost circular in P. trituberculatus; Chen, 1981, pi. 2, fig. 1), and the relative length of the walking legs
(shorter in P. trituberculatus, P3 only 2. 1-2.4 CL and P4 2.3 CL).
SIZE. — Maximum size among specimens examined: 7.5 x 11.2 mm (female, MNHN-B 26755); 5.3 x
7.9 mm (male, MNHN-B 26756).
ETYMOLOGY. — From denticulus, the Latin diminutive of dens, or tooth, in reference to the diagnostic rows
of microscopic teeth along the dorsal surface and anterior border of the cheliped propodi.
Distribution. — Known only from the Coral Sea (Vanuatu. New Caledonia, and Loyalty Islands) (Fig. 57).
Depth: known reliably between 210 and 778 m; also collected in trawls between 160-222 and 778-811 m.
Fig. 21. — a. Parapalicus denticulatus sp. nov.: 2 paratype 7.5 x 10.6 mm, off west coast of New Caledonia. BaTHUS 4.
stn CP 953, 220-234 m (MNHN-B 26715): dorsal view. — b, Parapalicus inermis sp. nov., 2 paratype 5.3 x 7.9 mm,
Fiji, Musorstom 10. stn CP 1351. 292-31 1 m (MNHN-B 26720): dorsal view.
Parapalicus inermis sp. nov.
Figs 21b, 22, 57
Material EXAMINED. — Fiji. Musorstom 10: stn CP 1322, 17°17.10'S. 177°47.92,E, 225-285 m. 7.08.1998:
1 2 (MNHN-B 26856). — Stn CP 1323, 17°I6.10'S, 177°45.75'E, 143-173 m, 7.08.1998: I <3 paratype 4.4 x 6.8 mm
(MNHN-B 26717), 1 <3.31 2 (MNHN-B 26757), 1 <3 paratype 3.9 x 6.6 mm. I 2 paratype 5.7 x 8.3 mm (USNM). —
Source : MNHN. Paris
1ND0-WEST PACIFIC SPECIES OF PALICID CRABS
503
StnCP 1324, 1 7° 1 7.37’S, I77°47.05'E, 102-104 m, 7.08.1998: 1 <3,1 $ (MNHN-B 26858). — Sin CP 1325,
17°16.39’S, 177°49.80'E, 282-322 m, 7.08.1998: 1 2 (MNHN-B 26859). — Sin CP 1349, 17°31.07'S, 178°38'79'E.
244-252 m, 11.08.1998: 2 3 , 15 2 (MNHN-B 26850). — Sin CP 1351, 17°31.14'S. 178°39.96'E. 292-311 m.
1 1.08.1998: 1 2 paratype 5.4 x 8.0 mm (MNHN-B 26719), 1 2 paratype 5.3 x 7.9 mm (MNHN-B 26720). 7 2 (MNHN-
B 26851). — Stn CP 1355, 17°49.54'S, 178°49.39’E. 302-310 m. 12.08.1998: 1 3 holotype 4.2 x 6.4 mm (MNHN-B
26716), 1 2 paratype 5.7 x 8.4 mm (MNHN-B 26718). 4 2 (MNHN-B 26852). — Stn CP 1358, 17°48.49'S,
178°46.70'E, 80-120 m. 13.08.1998: I 2 (MNHN-B 26857). — Stn CP 1363, I8°12.39'S, 178°33.01'E. 144-150 m,
15.08.1998: 2 <J , 19 2 (MNHN-B 26849). — Stn DW 1365, 18°12.73'S. 178°32.38'E, 295-302 m, 15.08.1998: I 2
(MNHN-B 26860). — Stn DW CP 1366, 18°12.36'S, 178°33.06'E, 149-168 m, 15.08.1998: 1 3 , 6 2 (MNHN-B 26853).
— Stn CP 1370, 1 8° 1 2.32’S, 178°33.10'E, 113-123 m. 16.08.1998: 1 3 . 12 2 (MNHN-B 26854). — Stn CP 1371,
1 8° 1 2.36’S, 178°32.85'E, 135-151 m, 16.08.1998: 12 2 (MNHN-B 26855).
BORDAU 1: stn CP 1403. 16°40'S, 179°36'E. 220-224 m, 25.02.1999: 1 <3 . 4 2 (MNHN-B 27136). — Stn CP 1405,
16°39'S, 179°36'E. 180 m, 25.02.1999: 2 2 (MNHN-B 27137).
TYPES. — Holotype-. 1 6 4.2 x 6.4 mm, MUSORSTOM 10, stn CP 1355 (MNHN-B 26716).
Paratypes : 1 <J 4.4 x 6.8 mm, MUSORSTOM 10, stn CP 1323 (MNHN-B 26717): 1 2 5.7 x 8.4 mm.
MUSORSTOM 10, stn CP 1355 (MNHN-B 26718); 1 2 5.4 x 8.0 mm. MUSORSTOM 10. stn CP 1351 (MNHN-B
26719); 1 2 5.3 x 7.9 mm, MUSORSTOM 10, stn CP 1351 (MNHN-B 26720); I 3 3.9 x 6.6 mm, 1 2 5.7 x
8.3 mm, MUSORSTOM 10, stn CP 1323 (USNM).
Type Locality. — Off east coast of Viti Levu, Fiji, 1 7°49.54'S, 1 78°49.39'E. 302-3 1 0 m.
DIAGNOSIS. — Carapace (Figs 21b, 22a) with one anterolateral tooth on each side; tooth with truncate outer
border, sinuous, nearly straight upper border. Supraorbital borders each with two very slight notches. Suborbital
borders (Fig. 22b) each with broad, V- or L-shapcd gap between inner and outer suborbital lobes; outer lobe longer
than postorbital angle. Dorsal borders of cheliped propodi (Fig. 22c) nearly smooth, only with barely visible
microscopic teeth; proximal tubercle smooth. Abdomen of mature males (Fig. 22d) with two low, elongate
tubercles on segment 3; similar but slightly lower one on segment 4. Male first pleopods (Fig. 22e) with
helicoidal basal parts; each distal part blade-like, corneous, proximally curved but distally widened. Abdomen of
mature females with segments 3-5 fused; one transverse ridge along each segment 1-4 (slightly constricted twice
medially in segment 3).
Color A color photograph of a live specimen showed a red-brown carapace, light red-brown tubercles on the
meri of the walking legs, and narrow, red-brown bands across the walking legs. Some of the color shown in the
photographs remained in specimens that had been preserved in alcohol for nine months.
Description. — Carapace (Figs 21b, 22a) wider than long (CW/CL = 1.4-1. 5); dorsal surface covered with
coarse granules. Smooth sulcus between hepatic and branchial regions connecting with deep fissure between
anterolateral tooth and large swelling (with dorsal, rounded boss) along carapace border posterior to postorbital
angle. Anterolateral borders of carapace each with tooth covered with fine granules, truncate outer border, tip
slightly directed anteriorly, and sinuous, nearly straight upper border. One large, elongate to rounded, granular boss
close to carapace border on branchial region posterior to each anterolateral tooth. Two large, low bosses on
metagastric region. Episternal processes oblong. Posterior border of carapace with 6-8 slightly protuberant
tubercles, plumose setae.
Frontal border of carapace divided into two very broad lobes with straight margins sometimes marked by very
slight median notch; lobes divided by narrow V-shaped notch. Borders between frontal lobes and supraorbital
borders rounded, slightly raised in posterior portion. Supraorbital borders each with two very slight notches, one
before postorbital angle. Postorbital angles very short (shorter than outer lobe of suborbital border), with slightly
pointed tip. Cornea of eyes dorsoventrally flattened, much wider than base of short eye peduncle; peduncles without
conspicuous tubercles.
Suborbital borders (Fig. 22b) each with broad, V-shaped gap between inner and outer suborbital lobes. Outer
lobe pointed (obliquely directed in some specimens), shorter than pointed inner lobe. Pterygostomial lobes nearly
straight, triangular, slightly shorter than each inner suborbital lobe.
Each basal antennal segment expanded distally as slightly pointed process; flagellum long with
conspicuous, simple setae. Inner border of ischia of third maxillipeds sinuous to serrate but without defined teeth:
504
P. CASTRO
Fig. 22. — Parapalicus inermis sp. nov., 6 holotype 4.2 x 6.4 mm, east of Viti Levu, Fiji. MusORSTOM 10,
stn CP 1355, 302-310 m (MNHN-B 26716): a, carapace and pereopods, dorsal surface; b, suborbital border; c. right
cheliped, upper surface; d. abdomen; e, left male first pleopod. ventral view.
surface coarsely granular, upper borders rounded. Meri much narrower than ischia; narrow, pointed (in inner
direction) upper borders.
Dorsal and outer borders of cheliped propodi (Fig. 22c) nearly smooth, only with barely visible microscopic
teeth; low, rounded tubercle at proximal to each propodal-carpal joint. Distal half of each finger with 4-5 teeth
(triangular in smaller females, low and rounded in larger females). Carpi short, outer borders smooth; meri slender,
outer borders smooth.
Source : MNHN, Paris
INDOAVEST PACIFIC SPECIES OF PALICID CRABS
505
First three pairs of walking legs (P2-4; Figs 21b, 22a) very slender, with elongate meri, filiform propodi and
dactyli. Upper and lower borders of meri with small, similarly pointed tubercles (smaller on P2); distalmost
tubercle on each anterior border much wider at base, higher, with pointed tip directed distally in P3-4 (similar
tubercle but not conspicuously pointed in P2). Anterior and posterior borders of carpi, propodi. and dactyli of all
walking legs (P2-4) entire. Each P2 with one dorsal (very slight) carina on dactylus. Each P3 with one dorsal
carina on carpus; one dorsal (very slight) carina on propodus; one dorsal carina on dactylus. Each P4 with one
dorsal carina each on carpus, propodus, and dactylus. Meri of all legs with scattered plumose setae; dorsal surfaces
of propodi of P3-4 with rows of plumose setae; dorsal surfaces of dactyli of P2-4 each with one row of
conspicuous plumose setae along anterior border.
First pair of walking legs (P2) much shorter and slender than second and third pairs (P3-4); third pair about as
long as second pair. Fifth pair of pereopods (P5) short (0.8-0.9 CL), very slender; each merus with scattered
plumose setae along posterior border; each propodus with smooth borders, long plumose setae along anterior and
posterior borders; each dactylus with 1-4 spines increasing in length distally along posterior border, long, simple
setae along anterior and posterior borders, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows;
Abdomen of mature males (Fig. 22d) with segments 3-5 fused, two low, elongate tubercles on segment 3,
similar but lower on segment 4. Male first pleopods (Fig. 22e) with helicoidal basal parts; each distal part long,
blade-like, proximally curved, distally widened.
Abdomen of mature females with segments 3-5 fused. One transversal ridge along each segment 1-2. one along
proximal portion of fused segment 3-5 (slightly constricted twice medially in segment 3).
DISCUSSION. — P. inermis is very close to P. denticulatus sp. nov., which is known only from the Coral Sea
region (Vanuatu to Loyalty Islands), while P. inermis is only known from Fiji (sec discussion for P. denticulatus).
There are also some similarities with P. ambonensis on account of the presence of anterolateral teeth that have
obtuse outer borders.
P. inermis has legs that are unusually long in relation to carapace length. The total length of the second pair of
walking legs (P3) varied between 3.0-3. 5 CL, the longest in any known species of Parapalicus.
Size. — Maximum size among the females examined; 6.3 x 9.4 mm (MNHN-B 26757); the largest of the
males examined: 4.4 x 6.8 mm (MNHN-B 26717).
Etymology. — From inermis , Latin for unarmed, in reference to the smooth appearance of the outer border of
the cheliped propodi.
Distribution. — Known only from Fiji (Fig. 57). Depth: known reliably between 104 and 302 in; also
collected in trawls between 80-120 and 282-322 m.
Parapalicus elaniticus (Holthuis, 1977)
Fig. 57
Palicus elaniticus Holthuis, 1977: 185. figs 3a-f.
MATERIAL EXAMINED. — Red Sea. Gulf of Aqaba, off Elat, "La-Merkhav", stn 8. 60-80 m, C. LBWtNSOHN coll..
7.09.1966: 1 8 holotype of Palicus elaniticus Holthuis 3.5 x 4.5 mm (RMNH 30334).
Madagascar. "Vauban": stn CH 52. off Majunga (northwest coast), 15°21.0'S. 46°12.5'E. 150 m, mud. A. CROSNIER
coll., 8.11.1972: 1 2 (MNHN-B 26758).
Source : MNHN. Paris
506
P. CASTRO
Types. — Holotype of Palicus elaniticus Holthuis, 1977: 1 8 3.5 x 4.5 mm, La-Merkhav , stn 8 (RMNH
30334).
Type Locality. — Off Elat, Gulf of Aqaba, Red Sea, 60-80 m.
Diagnosis. — Carapace (HOLTHUIS, 1977, fig. 3a) with triangular, anteriorly-pointed anterolateral tooth on
each side. Supraorbital borders each with two slight notches, distalmost one followed by rounded, lobe-like
tubercle located on proximal border of postorbital angle. Suborbital borders (HOLTHUIS. 1977, fig. 3b) each with
U-shaped gap between inner and outer suborbital lobes: outer lobe much smaller than inner lobe. Dorsal bolder ot
cheliped propodi (Holthuis, 1977, fig. 3d) each with two proximal tubercles. Abdomen of male holotype
(HOLTHUIS, 1977, fig. 3D with two low tubercles on segment 3, lower one on segment 4. Male first pleopods
(HOLTHUIS, 1977, fig. 3g) with helicoidal basal parts; each distal part blade-like, bordered with small teeth and two
wing-like proximal processes. Abdomen of only mature female examined with one transverse ridge along each
segment 1-4.
DISCUSSION. — A character diagnostic of P. elaniticus is the rounded tubercle on the supraorbital border
proximal to each postorbital angle, which gives the angle a bilobed appearance (HOLTHUIS, 1977, fig. 3a).
The two specimens that were examined in this study had a notch on the middle portion of each of the supraorbital
borders. The description of the species, however, mentions that the borders lack any "distinct incisions" other than
the notch before the tubercle proximal to the postorbital angles (HOLTHUIS, 1977: 185). The distal halt of
the cheliped fingers of the female examined was armed with 3 triangular teeth, the pollex with 5. They were
absent, however, in the male holotype (Holthuis, 1977, fig. 3d).
The length of the walking legs was measured only in the Madagascar specimen (MNHN-B 26758): total length
of second and third pair of walking legs (P3-4) was 2.1 CL (P3 merus 0.6 CL, dactylus 0.5 CL; P4 merus 0.5 CL,
dactylus 0.5 CL).
Size. — Size of the only two specimens known to exist: 6.5 x 8.9 mm (female. MNHN-B 26758). 3.5 x
4.5 mm (male holotype, RMNH 30334).
Distribution. — Previously known only from the Gulf of Aqaba, Red Sea (HOLTHUIS, 1977). Its
distribution is now extended to the northwest coast of Madagascar (Fig. 57). Depth: known reliably between 80
and 150 in; also collected in a trawl between 60 and 80 m.
Parapalicus inanis sp. nov.
Figs 19d, 23, 57
Material EXAMINED. — Chesterfield-Bellona Plateau. MUSORSTOM 5: stn 280. 24°09.99'S, 159°35.75'E,
270 m, 10.10.1986: 1 S holotype 3.9 x 4.7 mm (MNHN-B 6721), 1 2 (MNHN-B 26824). — Stn 282, 24° 1 1 .55'S,
159°32.22'E, 226-230 m, 10.10.1986: 2 2 (MNHN-B 26825), 1 2 paratype 4.7 x 6.0 mm (USNM). — Stn 285.
24°09.35'S, 159°34.04’E. 245-255 m. 10.10.1986: 1 2 paratype 4.9 x 6.2 mm (MNHN-B 26722).
Bathos 2: stn DW 752, 22°22.32'S, 166°14.9'E. 330 m. 15.05.1993: 1 2 (MNHN-B 26826).
Loyalty Islands. MUSORSTOM 6: stn DW 462, 21°05.10'S, 167°26.85'E, 200 m, 21.02.1989: 1 2 paratype 4.4 x
5.8 mm (MNHN-B 26723).
Types. — Holotype : 1 $ 3.9 x 4.7 mm, MUSORSTOM 5, stn 280 (MNHN-B 26721).
Paratypes: 12 4.9 x 6.2 mm, MUSORSTOM 5, stn 285 (MNHN-B 26722); 1 2 4.4 x 5.8 mm,
MUSORSTOM 6, stn DW 462 (MNHN-B 26723); 1 2 4.7 x 6.0 mm, MUSORSTOM 5, stn 282 (USNM).
Type Locality. — Kelso Bank, Chesterfield-Bellona Plateau, Coral Sea, 24°09.99’S, 159°35.75’E, 270 m.
Diagnosis. — Carapace (Figs 19d, 23a) with small anterolateral tooth on each side; dorsal surface covered
with many small granules. Frontal lobes each with slight notch. Supraorbital borders each with two slight
notches. Suborbital borders (Fig. 23b) each without outer suborbital lobe, leaving very wide gap between inner
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
507
suborbital lobe and postorbital angle. Dorsal borders of cheliped propodi (Fig. 23c) each with two proximal
tubercles, microscopic teeth. Abdomen of mature males with two rounded tubercles on segment 3. very low one on
segment 4. Male first pleopods (Figs 23d-e) with sinuous basal parts; each distal part curved, corneous apex with
spinules. Abdomen of mature females (Fig. 23f) with segments 3-5 fused; straight or twice-constricted transverse
ridge on segment 3, segment 4 smooth or with short ridge at each outer edge.
DESCRIPTION. — Carapace (Figs 19d, 23a) wider than long (CW/CL = 1.2- 1.3); dorsal surface covered with
small granules. Anterolateral borders of carapace each with very small triangular tooth covered with fine granules.
Two bosses (more conspicuous in larger specimens) on metagastric region. Large granules on branchial region and
along posterior border (number and distribution varies among specimens). Episternal processes small, rounded
(more protuberant in larger specimens). Posterior border of carapace with very few plumose setae in some
specimens.
Frontal border of carapace divided into two small lobes divided by narrow V-shaped notch; each rounded lobe
divided into two very low lobes by slight notch. Borders between frontal lobes and supraorbital borders rounded.
Supraorbital borders each with two very slight notches (larger specimens with irregular borders without clear
notches). Postorbital angles short, slightly pointed. Cornea of eyes dorsoventrally flattened, slightly wider than
base of eye peduncle; each peduncle with two slightly granular, distal tubercles (ventral the largest).
Suborbital borders (Fig. 23b) each without outer suborbital lobe, leaving very wide gap between inner
suborbilal lobe and postorbital angle. Pterygostomial lobes straight, broad, rounded, only slightly shorter than
each inner suborbital lobe.
Each basal antennal segment expanded distally as rounded process; flagellum long with conspicuous, simple
setae. Inner border of ischia of third maxillipeds sinuous to serrate but without defined teeth; surface finely
granular, upper borders rounded. Meri much narrower than ischium; rounded, broad upper borders.
Dorsal borders of cheliped propodi (Fig. 23c) each with two proximal tubercles and microscopic teeth. Distal
two-thirds of each finger with 3-4 short teeth. Carpi short, outer borders with few. mostly microscopic tubercles;
meri slender, outer borders with scattered, low tubercles.
First three pairs of walking legs (P2-4; Figs 19d. 23a) very slender, with elongate meri, filiform propodi and
daetyli. Upper and lower borders of meri with similar, slightly pointed tubercles (smaller along anterior border of
P2); distalmost tubercle on each anterior border much wider at base, higher, rounded tip directed distally. Anterior
and posterior borders of carpi, propodi, and daetyli of all walking legs (P2-4) entire, without tubercles. Each P2
with one dorsal carina on carpus. P3-4 each with one dorsal carina on carpus, propodus (very slight), and dactylus.
Dorsal surfaces of meri of all legs and carpi and propodi of P3-4 with scattered plumose setae; dorsal surfaces of
daetyli of P3-4 each with one row of conspicuous plumose setae along anterior border.
First pair of walking legs (P2) much shorter and slender than second and third pairs (P3-4); third pair about as
long as second pair. Fifth pair of pereopods (P5) short (0.9 CL), very slender; each merus with 3 spines; each
propodus with 2-4 spines along posterior border, scattered plumose setae along anterior border; each dactylus
smooth, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of only mature male examined (holotype) with segments 3-5 fused, two conspicuous, rounded to
elongate tubercles on segment 3, one very low tubercle on segment 4. Male first pleopods (Figs 23d-e) with
sinuous basal parts; each distal part with corneous apex, tip with spinules.
Abdomen of mature females (Fig. 230 with segments 3-5 fused. One ridge along each segment 1-2; segment 3
of fused segment 3-5 with ridge straight or slightly constricted twice medially; segment 4 smooth or with very
low, short ridge at each outer edge.
508
P. CASTRO
Fig. 23. — Parapalicus inanis sp. nov.: a-c, f, 2 paratype 4.4 x 5.8 mm. Loyalty Islands. Musorstom 6. stn DW 462.
200 m (MNHN-B 26723): a. carapace and pereopods, dorsal surface; b. suborbital border; c, left chclipcd. outer
surface; f. abdomen. — d-e, 8 holotype 3.9 x 4.7 mm. Kelso Bank. Chesterfield-Bellona Plateau. Coral Sea.
Musorstom 5. stn 280. 270 m (MNHN-B 26721): d. left male first pleopod. ventral view; e, left male first plcopod.
distal part, dorsal view;
DISCUSSION. — P. inanis is unique among the known species of the genus in the absence of an outer
suborbital lobe. Like in P. armatus sp. nov., the anterolateral teeth are unusually small. It shares with
P. nanshaensis Chen, 1981 a cheliped merus that is armed with two basal tubercles.
All of the seven specimens that were examined in this study were unusually small for Parapalicus, the largest
specimen being only 4.9 x 6.2 mm (MNHN-B 26722).
Source : MNHN , Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
509
Size. — Maximum size among the six females examined: 4.9 x 6.2 mm (MNHN-B 26722); the only male
examined (MNHN-B 26721): 3.9 x 4.7 mm.
Etymology. — From inanis, Latin for empty, in reference to the empty appearance of the suborbital region
due to the absence of the outer suborbital lobe.
DISTRIBUTION. — Known only from the Coral Sea (New Caledonia and the Loyalty Islands) (Fie. 57). Depth'
200-330 m.
Parapalicus microphthalmus sp. nov.
Figs 24, 26a, 57
Material EXAMINED. — Madagascar. "Vauban": stn CH 52, off Majunga (northwest coast). 15°2 1 ,0'S,
46°12.5'E, 150 m. A. Crosnier coll., 8.11.1972: 1 3 paratypc 7.3 x 11.4 mm (MNHN-B 26725), 1 3 paratype 5.9 x
9.6 mm (MNHN-B 26726), 1 9 holotype 7.8 x 1 1.2 mm (MNHN-B 26724), 2 9 (MNHN-B 26827).
Types . — Holotype: 1 $ 7.8 x 1 1.2 mm (MNHN-B 26724).
Paratypes : 1 S 7.3 x 1 1.4 mm (MNHN-B 26725); 1 8 5.9 x 9.6 mm (MNHN-B 26726).
Type Locality. — Off northwest coast of Madagascar, 15°2 1.0'S, 46°12.5'E, 150 m.
DIAGNOSIS. — Carapace (Figs 24a, 26a) with conspicuous, triangular anterolateral tooth on each side.
Supraorbital borders sinuous, each with two notches. Cornea and eye peduncle small; peduncles with many pointed
granules. Suborbital borders (Fig. 24b) each with wide, U-shaped gap between inner and outer, dentiform,
suborbital lobes. Dorsal border of cheliped propodi each with very low carina bordered with microscopic teeth.
Abdomen ol mature males (Fig. 24c) with two low, elongate tubercles on segment 3, triangular one on segment 4.
Male first pleopods (Figs 24d-e) with helicoidal basal parts; each distal part with helicoidal, keel-like process,
curved, corneous apex. Abdomen of mature females with one transverse ridge along each segment I -4.
Description. — Carapace (Figs 24a, 26a) wider than long (CW/CL = 1. 4- 1.6); dorsal surface covered with
very coarse granules. Anterolateral borders of carapace each with conspicuous triangular tooth covered with
granules. Two bosses, also covered with granules, on branchial region posterior to each tooth; larger, more pointed
of the two bosses closer to border of carapace, giving carapace appearance of having two anterolateral teeth. Four
conspicuous bosses (two large median bosses Hanked by smaller one on each side) on metagastric region.
Episternal processes triangular, with conspicuous granules. Posterior border of carapace with several rounded
tubercles (6 protuberant tubercles in males), scattered plumose setae.
Frontal border of carapace divided into two small lobes divided by wide U-shaped notch; triangular tips. Borders
between frontal lobes and supraorbital borders rounded. Supraorbital borders sinuous, each with two notches.
Postorbital angles short, slightly pointed. Cornea of eyes small, dorsoventrally flattened, wider than base of eye
peduncle; peduncles with many small, pointed granules.
Suborbital borders (Fig. 24b) each with wide, U-shaped gap between inner and outer suborbital lobes. Outer
lobe narrow, pointed, shorter than similarly pointed (projected outwards when seen dorsally) inner lobe.
Plerygostomial lobes projecting slightly ventrally, pointed, equal in length to each inner suborbital lobe.
Each basal antennal segment expanded distally as elongate, granular process; flagellum long with conspicuous,
simple setae. Inner border of ischia of third maxillipeds slightly sinuous; surface coarsely granular, upper borders
slightly pointed. Meri much narrower than ischium; pointed, granular upper borders.
Dorsal and outer borders of cheliped propodi (Fig. 24a) each with very low. unarmed (only barely visible
microscopic teeth) carina, low proximal tubercle. Distal half to two-thirds of each finger with 3-7 short teeth.
Carpi short, outer borders with broad tubercle across distal portion, pointed tubercles throughout; meri slender,
outer borders with small distal tubercles.
First three pairs of walking legs (P2-4; Figs 24a, 26a) very slender, with elongate meri, filiform propodi and
dactyli. Upper and lower borders of meri with small, similarly pointed tubercles; distalmost tubercle on each
anterior border much wider at base, higher, pointed tip directed distally. Anterior and posterior borders of carpi,
510
P. CASTRO
propodi, and dactyli of all walking legs (P2-4) entire, without tubercles (carpi and propodi of P3-4 with
microscopic tubercles). P3-4 each with one dorsal carina each on carpus and propodus, one dorsal one ventral (very
slight) carina on dactylus. Meri of all legs with numerous plumose setae; dorsal suriaces oi carpi and propodi ot
P3-4 with rows of plumose setae; dorsal surface of dactyli of all legs each with one row of conspicuous plumose
setae along anterior border.
First pair of walking legs (P2) much shorter and slender than second and third pairs (P3-4); third pair about as
long as second pair. Fifth pair of pereopods (P5) short (0.8-0.9 CL), very slender; each merus with row of very-
low tubercles, conspicuous plumose setae along posterior border; each propodus with conspicuous plumose setae
along posterior border; each dactylus with one spine on posterior border, plumose setae along anterior border, one
terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as tollows:
Abdomen of mature males (Fig. 24c) with segments 3-5 fused, two very low tubercles (plus central
prominence) on segment 3, salient, triangular one on 4. Male first pleopods (Figs 24d-e) with helicoidal distal
parts, helicoidal. keel-like process along its length; each distal part curved, corneous (flexible but rigid) apex with
sperm channel.
Abdomen of mature females with segments 3-5 fused. One ridge along each segment 1-2, two along proximal
portion of fused segments 3-5 (most proximal one bordered by microscopic, blunt tubercles).
DISCUSSION. — The male first pleopods of P. microphthalmus are one of the most complex among the Indo-
west Pacific species of paheids. Their general morphology is closest to that of P. trispiralis sp. nov., known only
from the South China Sea and the Coral Sea.
The retina and the peduncle of the eyes are reduced in size in comparison to those of the other congeners. I'he
orbits, however, remain wide and deep.
Size. — Maximum size among specimens examined: 7.8 x 12.0 mm (female, MNHN-B 26724); 7.3 x
1 1 .4 mm (male paratype MNHN-B 26725).
ETYMOLOGY. — From mikros, Greek for small, and ophthalmos, Greek for eye, in reference to the diagnostic
small eyes.
Distribution. — Only known from off the northwest coast of Madagascar (Fig. 57). This and P . elaniticus
are the only two species of Pcirapalicus known from the Indian Ocean. Both are known from very few specimens.
Depth: 150 m.
Parapalicus nanshaensis Dai & Xu, 1991
Parapalicus nanshaensis Dai & Xu. 1991: 38. 46 [English], fig. 28.
TYPES. — Holotype: 1 5 6.5 x 8.2 mm (Dai & Xu, 1991), 7.05.1986 (Academia Sinica, Beijing).
Type Locality. — West of Nansha (= Spratly) Islands, South China Sea. 07°02'N, 108°52’E, soft muddy
bottom, 138 m.
Diagnosis (after Dai & Xu, 1991). — Carapace (Dai & Xu, 1991, fig. 28-1) with triangular anterolateral
tooth on each side. Supraorbital borders sinuous, each with one notch. Suborbital borders unknown. Dorsal border
Source MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
51 I
Fig. 24. — Parapalicus microphthalmus sp. nov.: a-b, $ holotype 7.8 x 1 1.2 mm. off northwest coast of Madagascar.
150 m (MNHN-B 26724): a. carapace and pereopods. dorsal surface; b. suborbital border. — c-e. paratype. 7.3 x
1 1.4 mm. off northwestern coast of Madagascar. 150 m (MNHN-B 26725); c. abdomen: d. left male first pleopod.
lateral (inner side) view; e, left male first pleopod. ventral view.
of chelipcd propodi (Dai & Zu, 1991, fig. 28-4) each with two low proximal tubercles; distal third of each finger
with 3-4 short, closed-together teeth. Males unknown.
Source :
512
P. CASTRO
DISCUSSION — P. nanshaensis was described from one female specimen. The description and the figures that
accompany it did not 2ive details of the morphology of the suborbital borders of the carapace that are needed lor its
proper comparison with other species. The holotype was not available for examination. No male specimens are
"' The chelipeds were described as smooth (Dai & Xu, 1991 : 46) but the dorsal border of each chcliped propodus
is armed with two basal tubercles (Dai & Xu, 1991, fig. 28-4). This is a characteristic shared with P. arnbonensis
(Fig. 16a), P. elaniticus (HOLTHUIS, 1977, fig. 3d), and P. inanis sp. nov. (Fig. 23c). P. nanshaensis can be
clearly differentiated from P. elaniticus, which has supraorbital borders each with two notches and a rounded, lobe¬
like tubercle before the postorbital angle (HOLTHUIS, 1977, fig. 3a), and from P. inanis, which has a very small
anterolateral tooth on each side of the carapace and notched frontal lobes. The carapace is similar to that ot
P. arnbonensis and P. trispiralis sp. nov. (both having triangular anterolateral teeth that are pointed in an anterior
direction; see Figs 16a, 27a) but the teeth on each finger of the chelipeds of P. nanshaensis are very dill erent:
triangular, blunt teeth that are closed together at the distal portion of each finger (Dai & Xu, 1991, I lg. 28-4),
a character unique to the genus. Furthermore, each supraorbital border has only one slight notch in P. nanshaensis,
but three in P. arnbonensis (Fig. 16a).
Size. — Size of the female holotype: 6.5 x 8.2 mm (Dai & Xu, 1991).
DISTRIBUTION. — Known only from near the Nansha (= Spratly) Islands in the South China Sea (Dai & Xu,
1991). Depth: 138 m.
Parapalicus piruensis Moosa & Serene, 1981
Figs 25. 26b, 58
Parapalicus piruensis Moosa & Serene, 1981: 34, tigs 2b. 3b, pi. 1, bg. C.
Material examined. — Indonesia. Ceram. Mariel King Memorial Expedition: stn CP I/I. Piru Bay, off Cape
Tutuhuhur, 03°15'S. 128°08’E, 59-64 m, 1.06.1970: 1 2 holotype 4.7 x 6.3 mm (RDC CB2730).
Kai Islands. Karubar: stn DW 22, 05°22'S, 133°01'E. 85-124 m, 25.10.1991: 1 juv. <3 (MNHN-B 26835)
New Caledonia. MUSORSTOM 4: Stn 203, 22°35.8'S, 167°04.8'E, 105-110 m, 27.09.1985: I juv. 2 (MNHN-B
Expedition Montrouzier: Touho, 55-60 m, 7.09.1993: 1 2 (MNHN-B 26837); 55 m: 1 2 (MNHN-B 26838).
Koumac Channel. 49-63 m, 7.10.1993: 2 d (MNHN-B 26760), 1 d CL 3.6 mm (MNHN-B 26761).
Fiji. Musorstom 10: stn CP 1358, 17°48.49'S. 178°46.70'E, 80-120 m, 13.08.1998: 2 <3,82.1 juv. 2
(MNHN-B 26759).
Types. — Holotype : 1 2 4.7 x 6.3 mm, Mariel King Memorial Expedition, stn CP I/l (RDC
CB2730).
Paratype : 1 2 3.2 x 4.2 mm (Moosa & Serene, 1981), Coronado Bay. Mindanao, Philippine Islands, sand
and mud, 46-128 m, 10.1 1.1964 (RDC CB2728).
Type Locality. — Off Cape Tutuhuhur. Piru Bay. Ceram, Indonesia. 03°15'S, 128“08'E, 59-64 m.
DIAGNOSIS. — Carapace (Fig. 26b; Moosa & Serene, 1981. fig. 2b) with small, triangular, anteriorly
pointed anterolateral tooth on each side; dorsal surface covered with very small granules. Supraorbital borders
sinuous, each with two notches. Suborbital borders each with wide, V-shaped gap between inner and outei
suborbital lobes; outer lobe much smaller than inner lobe. Cheliped propodi smooth. Abdomen of mature males
(Fig. 25a) with three very low, elongate tubercles on segment 3. Male tirst plcopods (Figs 25b-d) with sinuous
basal parts; each distal part with two blunt, slightly curved tips. Abdomen of mature females with segments 3-5
fused; one very low transverse ridge on segment 3 (constricted twice medially); low ridge at each outer edge ol
segment 4.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
513
Fig. 25. — Parapalicus piruensis Moosa & Serene, 1981, 3 CL
3.6 mm. Koumac Channel, New Caledonia, Expedition
Montrouzier, 49-63 m (MNHN-B 26721): a, abdomen; b, left
male first pleopod, lateral (inner side) view; c, apex, dorsal tip;
d, apex.
Color : Specimens from Fiji that had
been preserved for nine months after
collection had irregular brown patches
across the dorsal surface of the carapace
and broad, vertical bands of the same
color across the walking legs.
Discussion. — The specimens that
were examined in this study agree in
most respects with the description of
Moosa & Serene (1981), which was
based on only two female specimens. It
was short but it included relatively good
(but incomplete) figures and a very poor
photograph.
P. piruensis is characterized by a
carapace that is only covered with small
granules (Fig. 26b), short and acute
anterolateral teeth that are anteriorly
pointed, a very small outer suborbital
lobe, and an almost smooth male
abdomen (Fig. 25a). The total length of
the second pair of walking legs (P3) of
the specimens examined varied between
1 .9-2.0 CL (merus 0. 6-0.7 CL, dactylus
0.6 CL); total length of P4 2.0-2. 1 CL
(merus 0.6 CL, dactylus 0.6 CL).
This is the first time male specimens
of P. piruensis have been examined and
the male first pleopods described and
illustrated (Figs 25b-d). The basal part is
sinuous, not helicoidal as in most
species of Parapalicus. As in P. inanis
sp. nov. (Figs 23d-e), this appears to be the consequence of the short length of the pleopods.
SIZE. — Maximum size among the females examined: 6.7 x 7.4 mm (MNHN-B 26759); the largest ot the
males examined: 3.8 x 4.5 mm (MNHN-B 26760).
Distribution. — Moosa & Serene (1981) recorded this species from the Sulu Sea (Mindanao. Philippine
Islands) and Ceram Sea (Ceram, Indonesia). Its range (Fig. 58) is now extended to the Banda Sea (Kai Islands,
Indonesia), Coral Sea (New Caledonia), and Fiji. Depth: known reliably between 55 and 105 m, collected too in
trawls between 49-63 and 85-124 m.
Parapalicus trispiralis sp. nov.
Figs 27, 58
MATERIAL EXAMINED. — Philippine Islands. MUSORSTOM 1: stn 63. 14°00.8'N. 120°15.8'E. 191-195 m.
27.03.1975: 1 3 holotype 5.8 x 8.1 mm (MNHN-B 26727).
Loyalty Islands. Musorstom 6: stn CP 419. 20°41.65'S, 167°03.70'E, 283 m. 16.02.1989: 1 9 paratype 7.3 x
9.5 mm (MNHN-B 26728).
Source
514
P. CASTRO
Fig. 26. — a, Parapalicus microphthalmus sp. nov., $ paratype 7.3 x 11.4 mm. off northwest coast of Madagascar.
150 m, A. Crosnier coll. (MNHN-B 26725): dorsal view. — b. Parapalicus piruensis Moosa & Serene. 1981. 9 5.4
x 7.3 mm. Fiji. MUSORSTOM 10. stn CP 1358, 80-120 m (MNHN-B 26759): dorsal view. — c. Parapalicus
trituberculatus (Chen, 1981): 9 paratype 6.8 x 9.7 mm. South China Sea. stn 6028. 78 m (ZRC): dorsal view. —
d, Parapalicus unidentatus (Zarenkov, 1968): $ 7.6 x 10.9 mm. South China Sea off the Philippine Islands,
Musorstom 3. stn CP 101, 194-196 m (MNHN-B 26998): dorsal view.
Types. — Holotype : 1 <5 5.8 x 8.1 mm, Musorstom 1. stn 63 (MNHN-B 26727).
Paratype : 1 2 7.3 x 9.5 mm, MUSORSTOM 6, stn CP 419 (MNHN-B 26728).
Type Locality. — Off Luzon Island, Philippine Islands, 14°00.8'N. 120°15.8'E. 191-195 m.
Diagnosis. — Carapace (Fig. 27a) with small, triangular anterolateral tooth on each side. Wide notch between
two frontal lobes. Supraorbital borders each with three notches forming two small lobes. Suborbital borders (Fig.
27b) each with wide, V-shaped to nearly L-shaped gap between inner and outer suborbital lobes; outer lobe much
smaller than inner lobe. Dorsal borders of cheliped propodi with microscopic teeth. Abdomen of mature males
(Fig. 27d) with two conspicuous, triangular tubercles on segment 3, similar, larger one on segment 4. Male first
pleopods (Figs 27e-f) with three coils when seen ventrally: helicoidal basal part, twice-helicoidal, tusk-shaped
distal part. Abdomen of mature females with one transverse ridge along segment 3 (constricted medially); short
ridge at each outer edge of segment 4.
Description. — Carapace (Fig. 27a) wider than long (CW/CL = 1 .3-1.4); dorsal surface covered with small
granules. Anterolateral borders of carapace each with small triangular, anteriorly-pointed tooth covered with fine
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
515
granules. Two small bosses, also covered with granules, on branchial region along border of carapace posterior to
each tooth. Two conspicuous bosses on metagastric region. Episternal processes triangular. Posterior border of
carapace with six slightly protuberant tubercles (more pronounced in female paratype) with rounded tips, few
plumose setae.
Frontal border of carapace divided into two small lobes with triangular or slightly rounded tips, divided by wide
V-shaped notch. Borders between frontal lobes and supraorbital borders rounded. Supraorbital borders each with
Fig. 27. — Parapalicus irispiralis sp. nov., S holotype 5.8 x 8.1 mm. Philippine Islands, South China Sea.
MUSORSTOM 1, stn 63. 191-195 m (MNHN-B 26727): a. carapace and pereopods, dorsal surface; h. suborbital border;
c, right cheliped. outer surface; d. abdomen; e, left male first pleopod, ventral view; f. left male first pleopod. distal
part, dorsal view.
Source :
516
P. CASTRO
three conspicuous notches, forming two small lobes. Postorbital angles short, pointed. Cornea of eyes
dorsovcntrally flattened, much wider than base of eye peduncle; each peduncle with two soft. low. distal tubercles.
Suborbital borders (Fig. 27b) each with wide, V-shaped to nearly L-shaped gap between inner and outer
suborbital lobes. Outer lobe much smaller than pointed inner lobe. Pterygostomial lobes project slightly ventrally.
rounded, shorter than each inner suborbital lobe.
Each basal antennal segment expanded distally as small, slightly pointed process; flagellum long, with
conspicuous, simple setae. Inner border of ischia of third maxillipeds sinuous; surface with small granules, upper
borders rounded. Meri much narrower than ischia: rounded, narrow upper border.
Dorsal borders of cheliped propodi with row of microscopic teeth (more conspicuous in female paratype).
Distal third of each finger with 4-5 short, rounded teeth. Carpi short, outer borders each with low, broad
tubercle across distal portion, small, pointed tubercles throughout; meri slender, outer borders with small, pointed
tubercles.
First three pairs of walking legs (P2-4; Fig. 27a) very slender, with elongate meri, filiform propodi and dactyli.
Upper and lower borders of meri with similar, low tubercles; distalmost tubercle on each anterior border much
wider at base, with rounded tip directed distally. Anterior and posterior borders of carpi, propodi. and dactyli of all
walking legs (P2-4) entire, without tubercles (propodi of P3-4 with microscopic tubercles). Each P2 with one
dorsal carina on dactylus. Each P3 with one dorsal carina each on carpus, propodus, and dactylus. Each P4 with one
dorsal carina on carpus and propodus; one ventral (very low) carina on propodus; dactylus unknown. Meri ol all
legs with scattered plumose setae; dorsal surfaces of carpi and propodi of P3-4 with rows of plumose setae; dorsal
surface of dactyli of P2-3 each with one row of conspicuous plumose setae along anterior border; dactylus of P4
unknown.
First pair of walking legs (P2) much shorter and slender than second and third pairs (P3-4); third pair about as
long as second pair. Fifth pair of pereopods (P5) short (0.8 CL), very slender; each merus with row ol low
tubercles, some provided with spine on posterior border, scattered plumose setae along anterior border; each
propodus with two proximal spines on posterior border, scattered plumose setae along anterior border; dactylus
with two spines on posterior border, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males (Fig. 27d) with segments 3-5 fused, two conspicuous, triangular tubercles on
segment 3, similar but more salient one on segment 4. Male first pleopods (Figs 27e-f) with three coils when seen
ventrally (helicoidal basal part, twice-helicoidal distal part with tusk-shaped tip); corneous, flexible apex with
opening of sperm channel.
Abdomen of mature females with segments 3-5 fused. One ridge along each segment 1-2, one along proximal
portion of fused segment 3-5 (ridge very slightly constricted twice medially); short ridge at each outer edge of
segment 4.
DISCUSSION. — The description of P. trispiralis is based on only two specimens. Each of the measurements of
the length of the walking legs was based on only one specimen, except for the second pair of legs (P2).
This species is close to P. ambonensis in that both species show conspicuous, triangular tubercles on the
male abdomen. Characteristic of P. trispiralis is the presence of more conspicuous frontal lobes (formed by a
wide median sulcus) and very small but clearly demarcated supraorbital lobes. Both of these features are
characteristics of Pseudopalicus species. Nevertheless, the species clearly belongs to Parapalicus because of
its helicoidal male first pleopods and the morphology of the carapace (only one anterolateral tooth), suborbital
borders (tooth-like suborbital lobes), chelipeds (very slender and equal), walking legs (slender and long, smooth
carpi, tiliform propodi and dactyli; tubercles along borders of meri equal in size and shape), and shape of the male
abdomen (triangular).
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
517
SIZE. — Size of the female paratype: 7.3 x 9.5 mm (MNHN-B 26728); male holotypc: 5.8 x 8.1 mm
(MNHN-B 26727).
ETYMOLOGY. — From tres , Latin for three, and spiralis, Latin for coil, in reference to the diagnostic
arrangement of three coils or turns of the male first pleopods when seen ventrally.
Distribution. — Known only from the South China Sea (off Luzon Island. Philippine Islands) and the Coral
Sea (Loyalty Islands) (Fig. 58). Depth: 195-283 m.
Parapalicus trituberculatus (Chen, 1981)
Figs 26c, 28, 58
Palicus trituberculatus Chen, 1981: 465, pis 1, 2. — Dai et ai, 1986: 413, pi. 57, fig. 6.
Parapalicus marielae Moosa & Serene, 1981: 27, fig. 1. pi. 1. fig. A.
Palicus bidentatus Sakai. 1983: 630, fig. 3g-j.
Parapalicus trituberculatus - Dai & YANG, 1991: 450, pi. 57, fig. 6.
MATERIAL EXAMINED. — South China Sea. "Albatross": stn 5302, southeast of Hong Kong, 21°42'N.
1 14°50'E, 69 m, mud. 9.08.1909: 1 2 holotype of Palicus bidentatus Sakai, 1983 CW 9.2 mm (USNM" 195065).
Stn 6028, 22°00'N, 115°30'E. 78 m, soft mud. 14.04.1960: 2 6 paratypes 5.1 x 7.3 mm. 5.2 x 7.5 mm, 1 2 paratype
6.9 x 10.0 mm (ASIO); 2 6 paratypes 4.8 x 7.0 mm, 5.4 x 7.5 mm, 1 2 paratype 6.8 x 9.7 mm (ZRC 1999.1435-1437).
China. Hainan Island. 26.12.1955: 1 2 (AS1ZB).
Viet Nam. "Orlik": stn 206, Gulf of Tonkin, unknown depth. N.A. Zarenkov coll., 12.07.1960: 1 8 5.2 x
7.7 mm, 2 2 (MNHN-B 26807).
Philippine Islands. MUSORSTOM 1: stn 72, 14°11.8'N, 120°28.7'E. 127-122 m, 28.03.1976: 2 8 (MNHN-B
26808).
Indonesia. Aru Islands. Mariei. King Memorial Expedition: stn AW 1/H 1 1-12, Wokam, west coast of Wasir Island.
05°30'S, 134°12'E, 73-91 m, 15.06.1970: 1 2 holotype of Parapalicus marielae Moosa & Serene, 1981. 5.0 x 7.1 mm
(RDC CB2710).
Types. — Holotype of Palicus trituberculatus Chen. 1981: 1 8 , South China Sea off China, 8.04.1960
(Institute of Oceanology, Academia Sinica, Qingdao, China; S209B-14).
Allotype of Palicus trituberculatus Chen, 1981: 1 2 , South China Sea off China. 21°30’N. 1 15°30'E. 1 15 m.
13.04.1960 (Institute of Oceanology, Academia Sinica, Qingdao, China; S219B-23).
Paratypes of Palicus trituberculatus Chen, 1981: 1 <3,1 2 , South China Sea off China. 21°00'N. I 14°30'E,
83 m, 10.12.1959 (ASIO S143B-13); 2 8 paratypes 5.1 x 7.3 mm. 5.2 x 7.5 mm. 1 2 paratype 6.9 x 10.0 mm
(ASIO); 2 8 paratypes 4.8 x 7.0 mm, 5.4 x 7.5 mm, 1 2 paratype 6.8 x 9.7 mm (ZRC 1999.1435-1437).
Holotype of Parapalicus marielae Moosa & Serene, 1981: 1 8 5.0 x 7.1 mm. Mariel King MEMORIAL
Expedition, stn AW 1/H 1 1-12 (RDC CB2710).
Paratype of Parapalicus marielae Moosa & Serene, 1981: I 2 6.7 x 9.0 mm (Moosa & SERENE. 1981).
Same data as holotype (RDC CB271 1).
Holotype of Palicus bidentatus Sakai, 1983: 1 2 CW 9.2 mm, "Albatross" Philippine Expedition, stn 5302
(USNM 195065).
Type Locality. — South China Sea off China, 21°00'N, 1 14°30'E, 82 m.
DIAGNOSIS. — Carapace (Fig. 26c; CHEN, 1981, pi. 1) with triangular anterolateral tooth on each side.
Supraorbital borders each with two slight notches. Suborbital borders (CHEN, 1981. pi. 2, fig. I) each with
very broad, almost circular gap between inner and outer suborbital lobes. Eye peduncles with numerous, small
tubercles. Chcliped propodi (CHEN, 1981, pi. 2, fig. 2) with minute teeth. Abdomen of mature males (CHEN,
1981, pi. 2, fig. 4) with two slightly raised tubercles on segment 3, similar one on segment 4. Male first pleopods
(Figs 28a-b; Chen. 1981, pi. 2, figs 5-8; MOOSA & SERENE, 1981, fig. 1) with helicoidal basal parts; each distal
part straight, pointed, bordered with minute teeth. Abdomen of mature females with one transverse ridge along each
segment 1-4.
Source
P. CASTRO
Discussion. — P. trituberculatus is similar to
P. clinodentatus sp. nov. and P. unidentatus (Zarenkov,
1968) in the morphology of the dorsal surface of their
carapaces, which are covered by granular bosses and have
a triangular tooth and a blunt boss posterior to the
anterolateral tooth. These two species can be differentiated
from P. trituberculatus by the morphology of their male first
pleopods (more complex in P. clinodentatus and
P. unidentatus). the ornamentation of the cheliped propodi (a
carina-like ornamentation present in P. clinodentatus and
P. unidentatus), and the gaps between the inner and outer
suborbital lobes (narrow and V-shaped in P. clinodentatus and
P. unidentatus). P. trituberculatus also shares several
characteristics with P. denticulatus sp. nov. (sec contrast
between the two species in the description of P. denticulatus).
The specimens that were examined agree with the
paratypes that were available and the excellent figures given in
its description (Chen, 1981). The large boss posterior to the
triangular tooth is referred to in the description as a tooth,
hence the presence of "two teeth near the middle". The gap
between the inner and outer suborbital lobes is described as
V-shaped but it was wide and U-shaped in all specimens
examined in this study as well as in the figure given in the
original description (Chen, 1981, pi. 2, fig. 1). The total
length of the second pair of walking legs (P3) in the
specimens that were measured varied between 2. 1-2.4 CL
(merus 0.6-0.7 CL, dactylus 0.6-0. 7 CL); total length of P4
2.3 CL (merus 0.7 CL, dactylus 0.7 CL).
A comparison between the holotype of Parapalicus
marielae Moosa & Serene, 1981 (RDC CB2710) and six
paratypes of Palicus trituberculatus Chen. 1981 (ASIO; ZRC
1999.1435-1437) has clearly shown that they belong to the
same species. The descriptions of both species were published in the same year but only that of Chen (1981 )
gives an exact date: September 1981. Since the exact publication date of MOOSA & Serene (1981) could not be
determined, 31 December 1981 is adopted as its date of publication (International Code of Nomenclature, fourth
edition, 1999, Article 21.3.2). It is then concluded that description of CHEN (1981) was published first and
therefore the species should be referred to as Parapalicus trituberculatus (Chen, 1981 ). P. marielae is known from
only two specimens. The description mentioned the presence of a "tooth" on the branchial region (MOOSA &
Serene, 1981: 29). This is undoubtedly a reference to the outermost of the one or two pairs of granular bosses
present in that region of the carapace in many species of Parapalicus.
The examination of the holotype of Palicus bidentatus Sakai, 1983 has shown that it is identical to
P. trituberculatus. Sakai (1983) based his diagnosis on a flattened specimen without realizing it. Although the
anterolateral teeth of the holotype are bent upward (and hence Sakai's drawing showing two conspicuous "obtuse
processes" with rounded tips representing the anterolateral tooth and the blunt boss posterior to it), the edge of the
tront folded over the carapace, and with only three walking legs and one cheliped remaining, all characters clearly
agree with those of P. trituberculatus. The notches on the supraorbital border were incorrectly shown as forming
three conspicuous lobes ("3 dorsal ... teeth") and the frontal lobes as having four, not two, lobes (Sakai. 1983:
630, fig. 3g). Although some specimens of P. trituberculatus have slight notches on the frontal lobes, it was
difficult to verify the morphology of the folded frontal lobes in the holotype of P. bidentatus. In addition, Sakai's
specimen was collected southeast of Hong Kong, not far from the type locality of P. trituberculatus.
518
Fig. 28. — Parapalicus trituberculatus (Chen,
1981), 6 5.2 x 7.7 mm. Gulf of Tonkin, Viet
Nam. unknown depth (MNHN-B 26807):
a, left male first pleopod, ventral view;
b. left male first pleopod. distal part, dorsal
view.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
519
SIZE. — Maximum size among specimens examined: 6.9 x 10.0 mm (female paratype. ASIO); 5.4 x 7.5 mm
(male paratype. ZRC 1999.1435-1437).
DISTRIBUTION. — Known from the Gulf of Tonkin (off Viet Nam), in various locations of the South China
Sea from off south China to the Philippine Islands (Chen, 1981; Sakai, 1983, as Palicus bidentatus), and
the Arafura Sea (off Aru Islands), Indonesia (Moosa & Serene, 1981, as Parapalicus marielae) (Fig. 58). Depth:
69-122 m.
Parapalicus unidentatus (Zarenkov, 1968)
Figs 26d, 58
Palicus unidentatus Zarenkov. 1968: 763. figs 3A-T.
Parapalicus unidentatus - MOOSA & SERkNE, 1981: 32, fig. 4. pi. 1, fig. B.
MATERIAL EXAMINED. — South China Sea. "Orlik": stn 211, off northern coast of Hainan. 60-180 m.
13.07.1960: 1 3, holotype (ZMMU Ma 5444). — Stn 203: ibidem, 60 m. 11.07.1960: I 9. paratype (ZMMU
Ma 5245).
Philippine Islands. MUSORSTOM 1: sin 14, 14°00.2'N. 120°17.2'E, 190 m, 20.03.1976: 1 <J (MNHN-B 27033).
— Stn 20. 13°59.2'N, 120°20.3'E. 208-222 m, 21.03.1976: 1 5 (MNHN-B 27046). — Stn 24. 14°00.0'N, 120°18.0'E.
189-209 m, 22.03.1976: 3 9 (MNHN-B 27047). — Stn 26, 14°00.9'N. 120°16.8'E, 189 m, 22.03.1976: 1 <5.6 9
(MNHN-B 27048). — Stn 27. 13°59.8'N. 120°18.6'E, 192-188 m. 22.03.1976: 3 9 (MNHN-B 27049). — Stn 30.
14°01 ,3'N, 120°18.7'E, 186-177 m, 22.03.1976: 3 3 , 1 9 (MNHN-B 27050). — Stn 31. 14°00.0'N. 120°16.0’E. 187-
195 m, 22.03.1976: 3 9 (MNHN-B 27051). — Stn 32, 14°02.2'N. 120°17.7'E. 193-184 m, 23.03.1976: 3 9 (MNHN-B
27052). — Stn 35, 13°59.0'N, I20°18.5'E. 186-187 m, 23.03.1976: 1 9 (MNHN-B 27053). — Stn 55. 13°55.0'N.
120°I2.5’E. 200-194 m, 26.03.1976: 1 9 (MNHN-B 27054). — Stn 56, 13053.1'N, 120°08.9'E. 134-139 m.
26.03.1976: 1 3 7.2 x 10.3 mm (MNHN-B 26763), 5 9 (MNHN-B 26762), 1 3 (MNHN-B 27055). — Stn 61.
14°02.2'N, 120°18.1'E, 202-184 m. 27.03.1976: 1 9 (MNHN-B 27056). — Stn 63, 14°00.8’N. 120°15.8'E. 191-195 m.
27.03.1976: 1 3 . 2 9 (MNHN-B 27034). — Stn 64. 14°00.5'N, 120°16.3'E. 194-195 m. 27.03.1976: I 3 (MNHN-B
27035). — Stn 71, 14°09.3'N, 120°26.2'E, 174-204 m, 28.03.1976: 6 9 (MNHN-B 27036).
Musorstom 2: stn CP 1, 14°00'N, 120°19'E. 188-198 m, 20.11.1980: 1 9 (MNHN-B 27057), — Stn CP 2. 14°01'N.
120°17'E, 184-186 m, 20.11.1980: 1 9 (MNHN-B 27058). — Stn CP 10. 14°00'N. 120°18'E. 188-195 m. 21.11.1980:
3 9 (MNHN-B 27059). — Stn CP 11, 14°00'N, 120°19'E, 194-196 m. 21.11.1980: 3 9 (MNHN-B 27060). —
Stn CP 12, 14°01'N. 120°20'E, 197-210 m, 21.11.1980: 1 9 (MNHN-B 27061). — Stn CP 18. 14°00'N. 120°18'E. 1 88-
195 m, 22.11.1980: 4 6 , 1 1 9 (MNHN-B 27037). — Stn CP 19. 14°00'N. 120°16'E, 189-192 m. 22.11.1980: I 9
(MNHN-B 27062). — Stn CP 20, 14°00'N, 120°18'E, 185-192 m, 22.11.1980: 2 3,1 2 (MNHN-B 27038). —
Stn CP 21, 14°00'N, 120°17'E, 191-192 m. 22.11.1980: 2 9 (MNHN-B 27062). — Stn CP 51. 14°00'N. 120°17'E. 170-
187 m. 27.1 1.1980: 1 9 (MNHN-B 27063). — Stn CP 62. 14°00'N, 120°17'E. 186-189 m. 29.1 1.1980: 2 9 (MNHN-B
27064). — Stn CP 64. 14°01'N, 120°19'E. 191-195 m, 29.11.1980: 6 9 (MNHN-B 27065). — Stn CP 67. I4°01'N.
120°19'E, 193-199 m. 29.11.1980: 1 3 , 6 9 (MNHN-B 27039). — Stn CP 68. 14°01'N. 120°18'E. 195-199 m.
29.11.1980: l 3,7 2 (MNHN-B 27040). — Stn CP 71, 14°00'N, 120°18’E. 189-197 m. 30.11.1980: 1 9 (MNHN-B
27066). — Stn CP 72, 14°00'N, 120°18'E, 182-197 m. 30.11.1980: 2 3,3 9 (MNHN-B 27041).
Musorstom 3: stn CP 86. 14°00'N. 120°18'E, 187-192 m, 31.06.1985: 1 3 (MNHN-B 27042). — Stn CP 87,
14°00'N. 120°19'E, 191-197 m, 31.06.1985. 2 3 . 19 9 (MNHN-B 27043). — Stn CP 88. I4°0TN. 1 20°1 7'E. 183-
187 m. 31.05.1985, 1 3,2 2 (MNHN-B 27067). — Stn CP 92. 14°03'N, 1 20° 12'E. 224 m. 31.05.1985: 2 9 (MNHN-B
27068). — Stn CP 96, 14°00'N, 120°18'E, 190-194 m, 1.06.1985: 3 9 (MNHN-B 27069). — Stn CP 97. 14°00'N.
120°18'E, 189-194 m. 1.06.1985: 1 3 , 2 9 (MNHN-B 27044). — Stn CP 100, 14°00'N. 120°18’E. 189-199 m.
1.06.1985, 14 9 (MNHN-B 27070). — Stn CP 101. 14°00'N. 120°19'E, 194-196 m, 1.06.1985: 9 9 (MNHN-B 26998).
— Stn CP 103, !4°00'N, 120°18'E, 193-200 m, 1.06.1985: 1 9 (MNHN-B 27071). — Stn CP 108, 14°01'N. 120°I8'E.
188-195 m, 2.06.1985: 13,8 2 (MNHN-B 27045). — Stn CP 109. 14°00'N, 120°18'E. 190-198 m, 2.06.1985: 2 9
(MNHN-B 27072). — Stn CP 111, 14°00'N. 120°18'E. 193-205 m. 2.06.1985: 2 9 (MNHN-B 27073). — Stn CP 112.
14°00'N, 120°18'E. 187-199 m, 2.06.1985: 3 9 (MNHN-B 27074).
Semirara Islands. Musorstom 3: stn CP 120. 12°06'N, I21°I5'E. 219-220 m. 3.06.1985: 18 9 (MNHN-B 27075). —
Stn CP 121, 12°08'N. 121°18’E, 73-84 m, 3.06.1985: 1 9 (MNHN-B 27076).
Indonesia. Strait of Makassar. Corindon: stn CH 267. 01°56'S. 119°17'E, 134-186 m, 7.11.1980: 4 3.12
(MNHN-B 27032).
Tanimbar Islands. Karubar: stn CP 65, 09°14'S. 132°27'E. 176-174 m, 1.1 1.1991: 2 9 (MNHN-B 27077).
TYPES. — Holotype: 1 3 CL 6.0 mm, "Orlik", stn 213 (ZMMU Ma 5444). Paratype: 1 2 CL 7.2 mm,
"Orlik", stn 203 (ZMMU Ma 5245).
Source :
520
P. CASTRO
Type Locality. — Off northeast coast of Hainan Island, China, South China Sea, 60-180 m.
DIAGNOSIS. — Carapace (Fig. 26d; Zarenkov. 1968, fig. 3A) with triangular anterolateral tooth on each side.
Supraorbital borders each with two notches. Suborbital border (Z.ARENKOV, 1968, fig. 3B) each with narrow,
V-shaped gap between inner and outer suborbital lobes. Cheliped propodi with carina-like row of irregular tuber¬
cles; two additional rows of low tubercles along outer surface below. Abdomen of mature males (Zarenkov,
1968, fig. 36) with two slightly raised tubercles on segment 3, similar one on segment 4. Male first pleopods
(Zarenkov, 1968, fig. 3T ; Moosa & Serene, 1981, fig. 4) with helicoidal basal parts; each distal part
with two equal, pointed tips that cross each other. Abdomen of mature females with one transverse ridge along
segments 1-4.
DISCUSSION. — The chelipeds were described by Zarenkov (1968: 763) as having "two long rows... of
granules". When observed from the inner side, however, the dorsal row was in the shape of a carina-like row of
irregular tubercles. The abdomen of all but one of the mature males that were examined in this study followed die
figure of Zarenkov (1968, fig, 36). In the largest male (MNHN-B 26763). however, the tubercles arc pointed.
The morphology of their male first pleopods agrees in general terms with the schematic figure of Z.ARENKOV
(1968, fig. 3D and with the holotypc. It agrees more closely with the more detailed figure of MOOSA & SERENE
(1981, fig. 4) except that the two tusk-shaped tips as well as the corneous apex are shorter in their Figure than in
most of the many specimens that were examined. In some specimens the tips were slightly curved distally.
The material that was examined also agrees with the diagnosis of MOOSA & SERENE (1981: 32), who also
studied some of the MUSORSTOM 1 material from the Philippine Islands examined during this investigation. One
exception is their statement that there is only one "shallow excavation" on the supraorbital borders. The male
abdomen is described as having two "lobes" on segment 3 but a "triangular lobe" on segment 4. They only
examined two males, however.
The total length of the second pair of walking legs (P3) in the measured specimens was 2.7 CL (merus 0.8-
O. 9 CL, dactylus 0.7 CL); total length of P4 2.7-2.S CL (merus 0.9 CL, dactylus 0.7 CL).
P. unidentatus may be confused with P. denticulatus sp. nov. and P. trituberculatus sp. nov. because in the
three species the dorsal surfaces of the carapace are covered by granular bosses and have a triangular anterolateral
tooth with a blunt boss posterior to it. P. unidentatus may be separated from P. denticulatus by the morphology of
its male first pleopods (much simpler in P. denticulatus , which has a single blade-like distal part; Fig. 20e),
the ornamentation of the cheliped propodi (microscopic teeth in P. denticulatus ; Fig. 20c), and the gap between
the inner and outer suborbital lobes (broad and U-shaped in P. denticulatus'. Fig. 20b). P. unidentatus can be
separated from P. trituberculatus also by the morphology of its male first pleopods (only a blade-like distal part
bordered by minute teeth in P. trituberculatus'. Fig. 28a-b), the ornamentation of the cheliped propodi (microscopic
teeth in P . trituberculatus ) and the gaps between the inner and outer suborbital lobes (very broad and U-shaped in
P. trituberculatus; CHEN, 1981, pi. 2, fig. 1).
SIZE. — Maximum size among specimens examined: 9.7 x 12.5 mm (female, MNHN-B 26762); 7.2 x
10.3 mm (male, MNHN-B 26763).
Distribution. — Recorded from several locations in the South China Sea (Zarenkov, 1968; Moosa &
Serene, 1981). Its distribution is now extended to the Strait of Makassar and Arafura Sea (off Tanimbar Islands)
in Indonesia (Fig. 58). Depth: known reliably between 84 and 220 m; also collected in trawl between 73-84 m.
Genus MIROPALICUS nov.
Type Species. — Palicus vietnamensis Zarenkov, 1968. Gender: masculine.
Diagnosis. — Frontal border of carapace divided into two small lobes. Anterolateral borders each with several
rounded teeth. Dorsal surface of carapace with small granules and granular bosses. Eyes dorsoventrally flatlened,
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
521
much wider than peduncle; peduncle with small, soft tubercles. Supraorbital borders each with two very short
lobes. Suborbital borders each limited by conspicuous, rounded pterygostomial lobe (inner suborbital lobe much
reduced, compressed on inner edge of border by pterygostomial lobe), short outer lobe. Each basal antennal
segment rectangular, slender, without distal expansion. Epistome narrow (not expanded dorsoventrally), vertically
inclined; two median teeth. Chelipeds increasingly unequal with increasing body size (particularly in males),
smallest typically more slender in females. First three pairs of walking legs (P2-4) very slender, with elongate
meri, filiform (long and very narrow) carpi, propodi, and dactyli. First pair (P2) much shorter than second pair
(P3), but second pair only slightly longer than third pair (P4). Borders of meri of P2-4 with nearly equal tubercles.
Distal, posterior borders of propodus and proximal, posterior borders of dactylus of P4 with short spines. Fifth pair
of pereopods (P5) reduced (0.8-0.9 CL), very slender. Abdomen of mature males elongate (sides almost parallel to
each other), with all segments free, one transverse ridge each along some segments. Male first pleopods long,
slender; basal parts straight; each distal part thick, uniramous. Abdomen of mature females with all segments free,
one transverse ridge along each segment 1-4.
Description. — Carapace transversely subovate, much broader than long, covered with conspicuous granules,
some which are grouped into bosses. Confluence of branchial and mesogastric regions depressed, smooth. Sulcus
between hepatic and branchial regions. Anterolateral borders of carapace rounded, with rounded teeth and bosses.
Thoracic sternite 7 with reduced process (episternal process) at each outer edge and posterior to insertion of fifth
pair ot pereopods (P5), visible dorsally as narrow, rounded process.
Frontal border divided into two small lobes. Supraorbital borders each with two very short lobes. Postorbital
angles long with rounded tips. Cornea of eyes dorsoventrally flattened, much wider than base of eye peduncle
(reniform); peduncle with soft tubercles. Orbits deep. wide.
Suborbital borders very wide, each limited by conspicuous, ventrally projecting, rounded pterygostomial lobe
on inner edge (inner suborbital lobe reduced, compressed by pterygostomial lobe), rounded, short outer lobe.
Each basal antennal segment rectangular, slender; flagellum long with conspicuous setae. Epistome narrow, not
expanded dorsoventrally, vertically inclined. Border of epistome with two median teeth plus two triangular outer
processes connecting distally with pterygostomial lobe. Meri of third maxillipeds narrower than ischia.
Chelipeds increasingly unequal with increasing size (particularly in males), smallest more slender in females.
Fingers of larger cheliped with cutting edge or teeth; those of smaller cheliped with teeth, pollex becoming
flatter with increasing size. Dorsal and outer borders of cheliped propodi with conspicuous tubercles. First
three pairs of walking legs (P2-4) very slender, with elongate meri. filiform (long and very narrow) carpi, propodi,
and dactyli. Upper and lower borders of meri with equal or nearly equal tubercles: carpi entire, without teeth; row
of spines along posterior borders of propodi and dactyli of third pair of walking legs (P4). First pair of
walking legs (P2) much shorter, slender than second pair (P3); third pair (P4) much shorter than second pair but
only slightly shorter than first pair. Last two pairs of walking legs (P3-4) each with slightly broadened coxae,
having slightly Battened, thick-edged anterior and posterior borders. Fifth pairs of pereopods (P5) reduced (0.8-0. 9
CL), very slender; basis-ischia slender, smooth; many short spines along posterior border of dactyli. some
on propodi.
Abdomen of mature males elongate, with both sides almost parallel to each other, with all segments free,
segments 1-2 dorsoventrally compressed, one transverse ridge each along some segments. Male first pleopods long,
slender; basal parts straight; each distal part uniramous. Second male pleopods short, thin, slightly curved; distal
segment with blunt tip.
Abdomen of mature females broad, rounded, with all segments free, one transverse ridge along each segment I-
4. Abdomen of immature females triangular (but broader than in males), segments 1-2 dorsoventrally compressed,
segments 3-6 fused, transverse ridge along each segment 1-4.
SPECIES included. — Only one species is so far known, M. vietnamensis (Zarenkov. 1968).
SEXUAL DIMORPHISM. — Females attain a larger size than males. The chelipeds are unequal in size in both
sexes but the difference is more apparent in males. The smallest cheliped is usually more slender in females. There
is no evidence of differences in the length or shape of the walking legs or fifth pair of pereopods.
Source
522
P. CASTRO
Discussion. — The genus is erected for one species that has unique characteristics while sharing some with
Pseudopalicus and others with Parapalicus (see Table 1).
Zarenkov (1968: 763) commented on the unusual length of the second pair of walking legs (P3) ol his new
species, Palicus vietnamensis Zarenkov, 1968. He pointed out that this character was mostly present in the
American species of palicids but that there were differences in the respective suborbital borders, a reference to the
very broad suborbital border of the new species. It was placed, like most other species of palicids known at that
time, in Palicus Philippi, 1838. MOOSA and SERENE (1981) provisionally included the species in their new genus,
Parapalicus. They did not examine any specimens, but commented that the narrow abdomen of the males was very
different from the triangular abdomen of Parapalicus. which "probably be restricted to species with broad abdomen
only" (MOOSA & SERENE, 1981: 26).
Characters that Miropalicus shares with Parapalicus are the morphology of the legs (long and filiform, with
equal or nearly equal tubercles along the borders of the meri, smooth borders of the carpi, propodi, and dactyl i. and
very slender P5, with smooth, slender basis-ischia). granular carapace, frontal border with two lobes, elongate
cheliped fingers, and dorsoventrally flattened eyes that are much wider than the peduncle (reniform eyes).
Characteristic of Pseudopalicus are the narrow shape of the male abdomen, a female abdomen with all segments
free and transverse ridges along segments 1-4, general shape of the carapace (more quadrate than oval, long and
conspicuous postorbital angles), greatly reduced episternal processes, unequal chelipeds, narrow and elongate
epistome provided with teeth, and slender basal antennal segments.
Unique characters of Miropalicus are the presence of a third pair of walking legs (P4) that is only slightly
longer than the first pair (P2) (markedly longer than P2 in Pseudopalicus and Parapalicus), spines along
the posterior border of both propodi and dactyli of P4, male first pleopods with straight basal parts and thick distal
parts each with a nearly square tip, and a seemingly enlarged orbit with hardly any lobes or teeth along
both supraorbital and suborbital borders, the reduced inner suborbital lobe being dwarfed by the very large
pterygostomial lobe. The coxae of P2-4, slightly expanded into thick anterior and posterior edges, are
somewhat intermediate between those of Pseudopalicus (thin, expanded edges) and Parapalicus (smooth, rounded
edges).
ZARENKOV (1968: 765) allied M. vietnamensis (together with Palicus hatusimaensis Sakai, 1963) with
R.ATHBUN's group 2 of the western Atlantic species of Palicus (Rathbun, 1918: 184) on account ol its very long
walking legs. This group consists of five species: P. acutifrons (A. Milne Edwards, 1880), P. cursor (A. Milne
Edwards, 1880), P.floridanus (Rathbun, 1918), P. gracilis (Smith, 1883), and P. gracilipes (A. Milne Edwards,
1880). Indeed, M. vietnamensis has striking superficial similarities with P. cursor. Both species have very long
walking legs, the third walking legs (P4) are conspicuously shorter than the second (P3), and there are three
anterolateral teeth. P. cursor, however, has a triangular male abdomen with at least one conspicuous tubercle, and
the carapace has four anterior lobes, a large, triangular outer suborbital tooth, and well-developed supraorbital
lobes. There are also superficial similarities with P. gracilipes (A. Milne Edwards, 1880), but as in P . cursor, this
species has a triangular male abdomen with three tubercles similar to that of Parapalicus. Thus, similarities with
RATHBUN's group 2 are only superficial and none of these five species conform with the characters used in the
definition of Miropalicus.
Etymology. — From mints, Latin for wonderful or strange, and the generic name Palicus, in reference to the
wide orbits without conspicuous lobes, which make the eyes look unusually large, and the unusually long and
graceful walking legs.
Miropalicus vietnamensis (Zarenkov, 1968)
Figs 29, 30a, 59. 60f
Palicus vietnamensis Zarenkov, 1968: 762, fig. 2A-T.
Parapalicus vietnamensis - MOOSA & SERkNE, 1981: 26, 27 (in key).
Material EXAMINED. — South China Sea. "Orlik", Vietnam, 300-350 m: 1 $ , holotype (ZMMU Ma 5248).
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
523
Philippine Islands. Musorstom 2: stn CP 83, 13°55'N, I20°30'E 3 1 8-3">0
B 27159).
2.12.1980: I 2 (MNHN-
Indonesia. Tanimbar Islands. Karubar: sin ED 68, 08°54'S, 132°0I'E, 280-296 m. 1.1 1.1991: I 2 (MNHN-B
27160). — Sin CP 79, 09°16'S. 131°22'E, 250-239 m. 3.11.1991: 5 <3, 18 2 (MNHN-B 26765). — Sin CP 83 09o^3'S
1 3 1 °00'E, 285-297 m, 4.11.1991: 2 5 , 7 2 (MNHN-B 27157). — Sin CP 84, 09°23'S, 131°09'E. 275-246 m
4.11.1991: 3 <5. 15 2 (MNHN-B 27158). — Sin CP 85, 09°22'S, 131°14'E. 245-240 m, 4.11.1991: 1 5 7 8 x 9 7 mm
(MNHN-B 26809).
Vanuatu. Musorstom 8: sin CP 980, 19021'S, 169°25’E, 450-433 m, 22.09.1994: 1 2 (MNHN-B 27161).
New Caledonia. MUSORSTOM 4: stn 236, 22°11.30'S. 167°15.00'E, 495-550 m, 2.10.1985: 1 3 , 1 2 (MNHN-B
27162). — Stn 239, 22°14.80’S. 167°15.70'E, 470-475 m, 2.10.1985: 1 2 (MNHN-B 27163). — Stn 241. 22°09.00'S,
167°12.20'E, 470-480 m. 3.10.1985: 1 3 , 1 2 (MNHN-B 27164). — Stn 243, 22°02.80’S, 167°07 70'E 435-450 m'
3.10.1985: 1 2 (MNHN-B 27165).
Bathus 1: stn CP 656, 21°13'S, 165°53'E. 452-460 m, 12.03.1993: 3 2 (MNHN-B 27166). — Stn CP 657. 21°14'S
1 65°54'E. 490-530 m, 12.03.1993: I 3 . 1 2 (MNHN-B 27167). — Stn CP 698, 20°34'S. 164°57 E 491-533 m
17.03.1993: 2 3 (MNHN-B 27168).
Bathus 2: sin CP 738, 23°02’S, 166°56'E, 558-647 m. 13.05.1993: 1 2 (MNHN-B 27169). — Stn CP 742. 22°33'S.
1 66°25'E, 340-470 m. 14.05.1993: 1 2 (MNHN-B 27170). — Sin CP 761, 22°18’S. 166°10'E. 490-500 m. 16.05.1993:
1 3 (MNHN-B 27171). — Stn CP 770, 22°09'S. 166°04'E, 400-402 m. 18.05.1993: 1 2 (MNHN-B 27172).
Halipro 1: sin CP 868, 21°14’S. 165°55'E. 430-550 m, 23.03.1994: 1 3 . 1 2 (MNHN-B 27173). — Stn CP 869
2 1 0 1 4'S, 165°55'E, 450-490 m, 23.03.1994: 2 5,12 (MNHN-B 27174).
Bathus 4: stn DW 888. 21°00'S. 164°27'E, 430-436 m, 2.08.1994: 1 2 (MNHN-B 27175). — Stn CP 899 ^0°I6'S
I63°50'E, 500-600 m, 3.08.1994: 2 5,22 (MNHN-B 26995).
Loyalty Islands. Musorstom 6: stn DW 410, 20°38.05'S, 167°06.65'E, 490 m. 15.02.1989: 1 5 (MNHN-B
27176). — Stn CP 415, 20°40.20'S. 167°03.95'E. 461 m. 15.02.1989: 1 5 (MNHN-B 27177). — Stn DW 428.
20°23.54'S. 1 66°1 2.57'E. 420 m. 17.02,1989: 1 2 10.5 x 14.8 mm (MNHN-B 26764). — Stn CP 467, 21°05 13'S
167°32.1 l'E, 575 m, 21.02.1989: 1 5 (MNHN-B 27178).
Types. — Holoppe : 1 5 CL 7.2 mm, "Orlik", off Viet Nam (ZMMU Ma 5248).
Type Locality. — Off Viet Nam coast, South China Sea, 300-350 m.
Diagnosis. — Carapace (Figs 29a, 30a, 60f; Zarenkov, 1968, fig. 2A) with three rounded to slightly
flattened anterolateral teeth on each side; teeth decrease in size posteriorly; 2-5 (usually four) rounded tubercles on
branchial region posterior to anterolateral teeth. Supraorbital borders each with two very short lobes. Suborbital
border (Fig. 29b; ZARENKOV, 1968, fig. 2B) very wide, limited by conspicuous, rounded, ventrally projecting
plerygostomial lobe (inner suborbital reduced, compressed by pterygostomial lobe), small, rounded outer lobe.
Dorsal and outer borders of cheliped propodi (Zarenkov, 1968, fig. 2A) each with three rows of tubercles, more
pronounced in slender chelipeds of females. Second pair of walking legs (P3) very long (3. 5-4.0 CL), third pair
(P4) short, only slightly longer than first pair (Figs 30a. 60f). Distal, posterior borders of propodi and proximal,
posterior borders of dactyli of P4 (Fig. 29c) with short spines. Abdomen of mature males (Fig. 29d: Zarenkov,
1968, fig. 2B ) narrow, with all segments free; one complete transverse ridge along segments 1-3 and along median
portion of segment 4. Male first pleopods (Figs 29e-f; Zarenkov, 1968, fig. 2T) with straight basal parts; each
distal part thick, with elongate, dorsal process with sperm channel, two ventral tubercles, one rounded, second
pointed. Abdomen of mature females with all segments distinct, one transverse ridge along each segment 1 -4 (very
low ridge may be present on segment 5).
Color: A male from the Loyalty Islands photographed live (Fig. 60f) had a pink carapace with irregular yellow
spots. The yellow spots on the nearly transparent walking legs and chelipeds may remain in specimens preserved
for several years. The cornea of the eyes is light blue in live specimens.
Discussion. — M. vietnamensis was placed by Zarenkov (1968) in Palicus but Moosa & Serene (1981 )
provisionally placed it in their new genus Parapalicus (see discussion of Miropalicus).
The material examined in this study agrees with the description and figures of Zarenkov (1968). There are.
however, some exceptions. The male abdomen was described as having segments 3-4 fused and crossed by two
ridges (Zarenkov, 1968, fig. 2B). This characteristic, suggestive of a juvenile specimen, was not found in any of
the specimens examined in this study. Zarenkov's specimen, one of three males he examined, was certainly not a
juvenile because of its size (CL 7.1 mm), which was close to the maximum size of the males examined in this
524
P. CASTRO
Fig. 29. — Miropalicus vietnamensis (Zarenkov, 1968). <5 7.8 x 9.9 mm. Tanimbar Islands, Aralura Sea. Indonesia,
Karubar, stn CP 85, 245-240 m (MNHN-B 26809): a, carapace, dorsal view; b, suborbital border; c. dactylus. right
fourth pereopod. dorsal view; d. abdomen; e, left first male pleopod, ventral view; f. left male first pleopod. distal
part, dorsal view.
study. The examination of the male holotype (ZMMU Ma 5248) showed that contrary to Zarenkov's drawings,
the abdomen has all segments free and the first pleopods are as illustrated here (figs 29e-f).
There are rows of conspicuous plumose setae along both borders of the propodi and the anterior borders of the
dactyli of P3-4. For the propodi. however, Zarenkov (1968, fig. 2A), shows setae only along the posterior
borders. No spines were indicated.
Characteristic of this species is the presence of 2-5 (usually four) high, rounded, granular bosses on ihc
branchial region posterior to each anterolateral tooth. Although on the outer border of the branchial region, they arc
slightly above the border proper, and can be differentiated from the larger and often flatter anterolateral teeth. The
most posterior of the bosses, the largest, is the first of typically 14 rounded bosses that cross the carapace along
the branchial and cardiac regions.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
525
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
The fifth pair of pereopods (P5) has smooth, slender basis-ischia and meri (except a few microscopic tubercles
along the posterior borders of the meri), each propodus with 2-7 spines along the posterior border, and each
dactylus with 9-15 long, thick spines along the posterior border and two along the distal part of the anterior border
in addition to a long terminal tooth.
Size. — Maximum size among specimens examined: 10.5 x 14.8 mm (female, MNHN-B 26764); 7.5 x
10.0 mm (male, MNHN-B 26765).
Distribution. — Originally known only from the South China Sea (Zarenkov, 1968), its distribution is
now extended to the South China Sea (coast of the Philippine Islands), Arafura Sea (off Tanimbar Islands.
Indonesia), and Coral Sea (Vanuatu, New Caledonia, and the Loyalty Islands) (Fig. 59). Depth: known reliably
between 245 and 575 m; also collected in trawl between 558 and 647 m.
Fig. 30. — a, Miropalicus vieuiamensis (Zarenkov, 1968): 2 10.7 x 15.5 mm, off northwest coasi of New Caledonia.
Bathus 4, stn CP 899. 500-600 m (MNHN-B 26995): dorsal view. — b. Paliculus kyusyuensis (Yokoya. 1933):
2 3.1 x 3.5 mm, Karubar, stn DW 2, 209-240 m (MNHN-B 26770): dorsal view.
Genus PALICULUS nov.
Type Species. — Palicus kyusyuensis Yokoya, 1933. Gender: masculine.
DIAGNOSIS. — Mature adults reach very small size. Frontal border of carapace divided into two small lobes.
Anterolateral borders each with 1-4 pointed teeth. Dorsal surface of carapace with tubercles, few granular bosses.
Eyes dorsoventrally flattened, much wider than peduncle; peduncle long with small, soft tubercles. Supraorbital
borders each with 1-2 lobes. Suborbital borders broad, each limited by conspicuous pterygostomial lobe, 1-2 short
lobes. Each basal antennal segment rectangular, slender, without distal expansion. Epistome narrow (not expanded
dorsoventrally), vertically inclined; two median teeth. Chclipeds nearly equal but more unequal in males. First
three pairs of walking legs (P2-4) very slender, with elongate meri and (with one exception) filiform (long and very
Source
526
P. CASTRO
narrow) carpi, propodi, and dactyli. First pair (P2) much shorter than second pair (P3), but second pair only
slightly longer than third pair (P4). Borders of meri of P2-4 with nearly equal tubercles; carpi, propodi, and dactyli
entire. Fifth pair of pereopods (P5) reduced (0.9 CL), very slender. Abdomen of mature males elongate (sides
almost parallel to each other), with all segments free, one transverse ridge along each segment 1-3. Male first
pleopods with basal parts straight; each distal part biramous. Abdomen of mature females with segments 3-5 or 4-
6 fused, one transverse ridge along each segment 1-3, one along fused segments 3-6 or 4-6.
DESCRIPTION. — Very small size (females ovigerous at less than CL 3.0 mm, reaching maximum known size
of CL 5.0 mm). Carapace subquadrate, tuberculate. Anterolateral borders of carapace straight, with I -4 pointed
anterolateral teeth. Confluence of branchial and mesogastric regions depressed, smooth; sulcus between hepatic and
branchial regions. Thoracic sternite 7 typically with process (episternal process) at each outer edge and posterior to
insertion of fifth pair of pereopods (P5), visible dorsally as rounded to narrow, pointed process (more conspicuous
in males).
Frontal border divided into two small lobes that may have recessed, pointed portion creating 4-lobe appearance.
Supraorbital borders each with 1-2 lobes. Postorbital angles long, pointed tips, directed outward. Cornea of eyes
dorsovcntrally flattened, much wider lhan base of eye peduncle (reniform); peduncle long with soft tubercles. Orbits
deep, wide.
Suborbital borders each limited by conspicuous, ventrally inclined, rounded pterygostomial lobe on inner edge
(inner suborbital lobe reduced, compressed by pterygostomial lobe) and 1-2 rounded, short outer lobes. Ridge
connecting each pterygostomial lobe with pterygostomial region with triangular tooth.
Each basal antennal segment rectangular, slender or slightly expanded; flagellum long with conspicuous setae.
Epistomc narrow, not expanded dorsoventrally, vertically inclined. Border of cpistome with no apparent teeth or
two short median teeth plus two narrow outer processes connecting distally with each pterygostomial lobe. Meri of
third maxillipeds as wide as ischia.
Chelipeds increasingly unequal with increasing size (particularly in males), smallest more slender in females.
Fingers of larger cheliped with cutting edge or teeth; those of smaller cheliped with teeth, pollex becoming flatter
with increasing size. Dorsal borders of cheliped propodi with conspicuous tubercles. First three pairs of walking
legs (P2-4) very slender, with elongate meri and (with one exception) filiform (long and very narrow) carpi,
propodi, and dactyli. Upper and lower borders of meri with nearly equal (can be thin, rounded) tubercles; carpi
entire, without teeth; borders of dactyli entire. First pair of walking legs (P2) much shorter and slender than second
pair (P3); third pair (P4) slightly shorter than second pair. Fifth pairs of pereopods (P5) reduced (0.9 CL), very
slender; basis-ischia slender and smooth; spines along posterior border of meri.
Abdomen of mature males elongate, with both sides almost parallel to each other, with segments free,
segments 1-2 dorsoventrally compressed, one complete transverse ridge along each segment 1-3, incomplete along
segment 4. Male first pleopods long, slender; basal parts straight; each distal part biramous. Second male pleopods
short, thin, slightly curved; distal segment with blunt tip.
Abdomen of mature females broad, rounded, segments 1-2 dorsoventrally compressed, segments 3-5 or 4-6
fused, one transverse ridge along each segment 1-3, proximal portion of fused segment. Abdomen of immature
females triangular (but broader than in males), segments 1-2 dorsoventrally compressed, 3-6 fused, transverse ridge
along each segment 1-4.
Species INCLUDED. — Three species are known: P. foliatus sp. nov., P. kyusyuensis (Yokoya, 1933), and
Paliculas sp., an undescribed species known from only one incomplete specimen.
Sexual Dimorphism. — As in other Indo-west Pacific palicids, females are larger in size than males. The
chelipeds are slightly unequal in size in both sexes but the difference is more apparent in males. There is no
evidence of sexual dimorphism in the relative length of pereopods.
Discussion. — Paliculus (with the exception of P. foliatus sp. nov.) shares with Parapalicus and Miropalicus
elongate, filiform walking legs (Fig. 30b). The tubercles on the meri of the walking legs are equal or nearly equal.
Similarities between Paliculus and Miropalicus include a narrow male abdomen, enlarged pterygostomial lobes
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
527
each fused to a reduced inner suborbital lobe, simple (not expanded) basal antennal segments, and a similar type of
chelipeds, the largest with a thick and high propodus, particularly in males. In fact, Zarenkov (1968: 765) placed
M. vietnamensis together with P. kyusyuensis (as Palicus hatusimaensis) in a group of two Indo-west Pacific
palicids related to group 2 ol Rathbun (1918: 184) (see discussion of Mi ropalicus). Moosa & Serene (1981:
26) provisionally placed these two species in their new genus Parapalicus but remarked on the differences between
their narrow male abdomens and the broad abdomen of Parapalicus. A similar narrow male abdomen is
characteristic of Pseudopalicus hut this genus and Paliculus differ widely in the morphologies of the carapace,
pereopods, and male first pleopods. There are also marked differences in the morphologies of the carapace and male
first pleopods ol Paliculus and Parapalicus (see below).
Besides a small size, Paliculus has several characters unique among Indo-west Pacific palicids: a subquadrate
carapace with a straight or nearly straight sides and 1-4 pointed (not truncate and short as in Rectopalicus)
anterolateral teeth on each side, and third maxillipeds where the meri is as wide as the ischia. None of these unique
characters appear to be present among described species of palicids from the western Atlantic and eastern Pacific
regions. The male first pleopods (Fig. 3 1 e) do not have an helicoidal basal part (in contrast to Parapalicus) and the
distal part is biramous but much different from any known species of Pseudopalicus.
Etymology. — Front the generic name Palicus and the diminutive suffix -ulus (Latin).
Key to the species of PALICULUS
1. Posterior border of carapace slightly pointed medially (Fig. 32a) .
. Paliculus foliatus sp. nov.
— Posterior border of carapace slightly convex, not pointed medially . 2
2. Anterolateral border of carapace with two pointed, elongate teeth. Supraorbital border with
one median lobe having one pointed tip. Suborbital border with one lobe .
. Paliculus kyusyuensis (Yokoya, 1933)
— Anterolateral border of the carapace with one pointed, elongate tooth. Supraorbital border
with one median lobe having two pointed tips. Suborbital border with two lobes .
. Paliculus sp.
Paliculus kyusyuensis (Yokoya, 1933)
Figs 30b, 31. 56
Palicus kyusyuensis Yokoya, 1933: 206, 217, fig. 70. — Sakai, 1976: 595, fig, 325b.
Cymopolia kyusyuensis - Sakai. 1939: 609, fig. 90b; 1956: 52.
Palicus hatusimaensis Sakai, 1963: 230, fig. 7c; 1965: 185, pi. 89, fig. 3; 1976: 595, pi. 205, fig. 3. — Takeda &
Miyake, 1968: 577, fig. 11; 1972: 87. — Zarenkov, 1968: 765, fig. 3 A [as Palicus contractus in caption]. —
Takeda, 1973: 55.
Parapalicus hatusimaensis - MOOSA & SERENE, 1981: 26.
MATERIAL EXAMINED. — Madagascar. Northwest coast, off Nosy Be. "Vauban": dredging 2, 12°38.5'S.
48°16.5'E, 240 m, sand, A. Crosnier coll.. 11.10.1974: I 9 (MNHN-B 26769).
Japan. Sagami Bay. 30-50 m, Manazuru Marine Laboratory staff coll., 5.08.1960: 1 9 holotype of Palicus
hatusimaensis Sakai, 1963 3.3 x 4.0 nun (SMF 24697).
Kii Peninsula. "Tansei Marti"', stn KT96-6, stn TB-3, off Tanabe, 33°39.06'N, I35°06.82’E, 180-187 m 12 05 1 996-
I 9 (CBM-ZC 4955).
Bungo Strait. Sandy bottom, T. Komai coll., 11.07.1994: I 9 (CBM-ZC 981).
Kyushu. "Soyo Marti", stn 304, off southern Miyazaki-ken, 241 in, 12.07.1928: 1 juv. 9 (KMNH).
Tsushima Islands. Stn 33, 34°16'N, I29°31.5'E, 105 m, sand and shells, 5.08.1968: 1 9 (ZLKU 13565). — Stn 35,
34°25.1'N, 1 29°59.3'E, 115 in, coarse sand and shells, 5.08.1968: 1 9 (ZLKU 12901). — Sin no. missing: 2 6.29
(ZLKU 13566).
Source :
528
P CASTRO
Amain i Islands. " Toyoshio-Maru stn 8, off Kekuroma Island, 28°04.7I'N, I29°27.38'E, sledge nei, 310 m,
E. TSUSHIDA coll., 10.1 1.1994: 3 2 (CI3M-ZC 4951). ........
Indonesia. Kai Islands. Siboga EXPED.ITIQN: stn 260, 05°36.5'S, 132°55.2'E, 90 m sand coral and shells . 6-
l8 12 1899- 1 8 tZMUC). — KARUBAR: stn DW 2. 05°47'S, 132°13'E, 209-240 m, 22.10.1991: 2 9 (MNHN-B 26770).
Tanimbar Islands. Karubar: sin DW 49, 08°00'S. 132°59'E. 210-206 m, 29.10.1991: 2 $ (MNHN-B 26771)
Chesterfield Islands. MUSORSTOM 5: stn 301, 22°06.90'S, 159°24.60'E, 487-710 m. 12.10.1986: I 5 2.8 x
3.2mm (MNIIN-B 26766). _ ........ u
New Caledonia. BlOCAL: sin DW 77, 22°15'S, 167°15'E, 440 m, 5.10.1985: I <J 2.7 x 3.1 mm, I 2 (MNIIN-B
26767); 1 2 , 1 juv. 2 3.5 x 3.6 mm (MNHN-B 26768).
Wallis Island. Musorstom 7: stn DW 610, 13°21'S, 176°09'W. 286 m, 26.05.1992: I 2 (MNHN-B 26772).
TYPES. — Holotype of Palicus kyusyuensis Yokoya, 1933: 1 2 , south of Koshiki Islands, Japan, stn 423,
243 m, 15.07.1929 (deposit unknown, possibly lost).
Holotvpe of Palicus hatusimaensis Sakai, 1963: 1 $ 3.3 x 4.0 mm, Sagami Bay, Japan (SMF 24697).
TYPE Locality. — South of Koshiki Islands, southwest of Kyushu, Japan, 243 m.
Diagnosis. — Carapace (Figs 30b, 31a; Yokoya, 1933, fig. 30; Sakai, 1963, fig. 7c; 1976, pi. 205, fig. 3;
Sakai's references as Palicus hatusimaensis) with two narrow, pointed anterolateral teeth (2-3 very small teeth
may be present posterior to anterolateral teeth) on each side. Dorsal surface ol carapace with small, rounded
tubercles (less common in smaller individuals); two raised, granular bosses on metagastric region; four salient
tubercles along posterior border. Supraorbital borders each with one short, rounded or pointed, median lobe.
Postorbital angle long, pointed outward. Suborbital border (Fig. 31b; ZARENKOV, 1968, lig. 3 A. as Palicus
contractus in caption; Takeda & MIYAKE, 1968, fig. 1 la, as P. hatusimaensis) each limited by conspicuous,
ventrally projecting ptcrygostomial lobe and broad, rounded outer lobe. One pointed or rounded tooth on each ridge
connecting pterygostomial lobe with ptcrygostomial region of carapace (Fig. 31b); conspicuous, sharp tooth on
pterygostomial region anterior to insertion of chelipcd. Dorsal and outer borders of cheliped propodi each with
2-3 rounded tubercles (Sakai, 1963, fig. 7c, as P. hatusimaensis). Abdomen of mature males narrow, with all
segments free; one complete transverse ridge along segments 1-3 (only along central portion of segment 4). Male
first plcopods (Fig. 3 le; Takeda & Miyake, 1968, fig. 1 lb-c, as P. hatusimaensis) with straight basal parts;
each distal part with elongate dorsal process lopped by toothed structure and much shorter ventral process with
rounded tips. Abdomen of mature females with segments 4-6 fused, one transverse ridge along each segment 1-3
and proximal portion of fused segment 4-6; ridges with lobed edges, particularly that along segment 3.
DISCUSSION. — Palicus kyusyuensis was described from one female specimen from southern Japan (3.2 x
4.3 mm; Yokoya, 1933: 206). The holotype, which like other specimens examined by Y. Yokoya should have
been deposited at the Zoological Laboratory, Kyushu University, Fukuoka but eventually transferred to
the Kitakyushu Museum of Natural History, Kitakyushu, appears to be lost. The original specimen lacked both
chelipeds and had only one P2 and one P5.
P. hatusimaensis Sakai, 1963 was described from one female from central Japan (Sakai, 1963: 230)
(the holotype was erroneously referred to as a male in the measurements given by SAKAI). Sakai unlortunately
failed to compare his new species with P. kyusyuensis. The examination of Sakai's holotype (3.3 x 4.0 mnv,
SMF 24697) plus additional material from Japan and other Indo-west Pacific locations lailed to show that
P. hatusimaensis is different from P. kyusyuensis.
One difference, however, is that while the frontal border of the carapace ol both P. kyusyuensis and
P. hatusimaensis was described as being bilobed, each lobe of P. hatusimaensis is recessed ventrally and topped by
a sharp denticle (Sakai, 1963, fig. 7c), giving the front a 4-lobe appearance. The morphology of the frontal border
was found to vary widely. Of the 20 specimens with complete anterior borders that were examined in this study,
12 agreed with the holotype of P. hatusimaensis in having a bilobed border with four separate tips and two had
four tips in patterns different from that of P. hatusimaensis (two broad, straight lobes each with only one pointed
tip on the inner or outer border). One female from Kyushu, southern Japan (3.8 x 4.1 mm; KMNH), one female
from the Amami Islands, southern Japan (3.0 x 3.5 mm; CBM-7.C 4951), and one juvenile female from New
Caledonia (3.5 x 3.6 mm; MNHN-B 26768) clearly had two lobes each with a pointed tip at the apex identical to
Source : MNHN, Pans
1NDO-WEST PACIFIC SPECIES OF PAUCID CRABS
529
those shown in Yokoya's figure (YOKOYA, 1933, fig. 70). One female from the Chesterfield Islands (2.8 x
3.2 mm; MNHN-B 26766) had a bilobed front, each lobe triangular but without the pointed tip shown in
Yokoya's figure, and another female from the Tsushima Islands, Japan with a single left lobe and a right lobe
with two tips (3.4 x 3.8 mm; ZLKU 13565). Finally, one male from the Tsushima Islands (2.4 x 2.6 mm;
ZLKU 13566) had a straight front without lobes.
Fig. 31. — Paliculus kyusyuensis (Yokoya, 1933), 3 2.4 x 2.6 mm, off south coast of New Caledonia. Biocal.
stn DW 77, 440 m (MNHN-B 26767): a. carapace, dorsal view; b. suborbital border; c. dactylus. right fourth
pereopod. dorsal view; d, abdomen; e. right male first pleopod, ventral view.
There are nevertheless two other differences between Yokoya's and Sakai’s species. Yokoya did not mention
in his description nor showed in the accompanying figure two characteristics of the species: two raised bosses on
the metagastric region of the carapace and four salient tubercles along the posterior border of the carapace, it is
assumed that they were overlooked by Yokoya. His drawings have been known not to be always accurate (see
Komai, 1996).
Sakai's description of P. hatusimaensis describes "four or five teeth" along each anterolateral border (Sakai,
1963: 230) but included in this number are the postorbital angle and "two or three liny teeth" posterior to the two
longer anterolateral teeth that are always present. His description therefore should have been five or six but
including small teeth posterior to the second anterolateral tooth.
Source :
530
P CASTRO
The outer suborbital lobe is shown in a figure given by Takeda & MIYAKE (1968, lig. I la, as
Palicus liatusimaensis ) as a narrow lobe. In all of the specimens that were examined in this study it was broad and
rounded like that shown by Zarenkov (1968, fig. 3 A). Furthermore, TAKEDA & Miyake’s ligure does not show
the characteristic pointed teeth on the anterior edge of the ridge that connects the pterygostomial lobe with the
pterygostomial region of the carapace. Two teeth are shown in Zarenkov's ligure. Another disagreement between
the material examined in this study and Takeda & MlYAKE's figures involves the left male first pleopod
(Takeda & Miyake, 1968, figs 1 Ic-d). It was shown as having a single distal part with a "transparent, twisted
beak at the tip". The right first pleopod of a male from New Caledonia (2.7 x 3.1 mm; MNHN-B 26767) that is
shown in Fig. 3 le and those of a male from an unknown location in Japan (2.8 x 3.2 mm; ZLKU 13566) had
a biramous distal part that consisted of an elongate dorsal process that ended in a toothed structure and a shorter
ventral process with rounded tips. Takeda & Miyake's specimens were collected in the East China Sea east and
southeast of Kyushu, Japan. It was almost identical in size to the New Caledonia and Japan specimens, 2.7 x
3.2 mm. Either these specimens were incorrectly drawn or they may represent a different species .
Even in the absence of YOKOYA's specimen, there is sufficient evidence to consider Palicus liatusimaensis
Sakai a subjective junior synonym of Palicus kyusyuensis Yokoya.
P. kyusyuensis , which is rarely collected due to the small size and fragility of individuals, shows a much wider
geographic distribution than originally recorded. It ranges from Madagascar to Japan and Wallis Island in the
southwestern Pacific (Fig. 56). It is most probably found across the Indian Ocean.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
SIZE. — Maximum size among specimens examined: 3.6 x 4.4 mm (female, CBM-ZC 4955); 2.7 x 3.1 mm
(male, MNHN-B 26767).
Distribution. — Japan (Yokoya, 1933; Sakai, 1963, 1965, 1976; Takeda & Miyake, 1968, 1972;
some records as P. liatusimaensis), South China Sea (ZARENKOV, 1968, as P. liatusimaensis), and now also
known from Madagascar, Sunda and Banda seas (Indonesia), Coral Sea (Chesterfield Islands and New Caledonia),
and Wallis Island, southwestern Pacific (Fig. 56). Depth: known reliably between 50 and 487 in; also collected in
trawls between 30-50 and 487-710 m.
Paliculus foliatus sp. nov.
Figs 32
MATERIAL EXAMINED. — New Caledonia. Expedition Montrouzier: Touho, unknown depth (less than 50 m),
09.1993: 1 2 holotype 5.0 x 5.0 mm (MNHN-B 26729).
TYPE. — Holotype: 1 9 5.0 x 5.0 mm (MNHN-B 26729).
Type Locality. — Touho, northeastern coast of New Caledonia, unknown depth.
DIAGNOSIS. — Carapace (Fig. 32a) subquadrate, with four narrow, pointed anterolateral teeth on each side.
Dorsal surface of carapace with small, rounded tubercles; conspicuously depressed area along confluence ol
branchial and mesogastric regions; two raised, granular bosses on metagastric region; posterior border medially
pointed, with four salient tubercles. Supraorbital borders each with one pointed median lobe. Postorbital angle
long, pointed outward. Suborbilal borders (Fig. 32b) each limited by conspicuous, ventrally projecting
pterygostomial lobe and broad, rounded outer lobe. One blunt tooth on each ridge connecting pterygostomial lobe
with pterygostomial region of carapace; pointed tooth on pterygostomial region anterior to insertion of cheliped.
Source : MNHN, Pans
INDO-WEST PACIFIC SPECIES OF PALIC1D CRABS
531
Dorsal borders of cheliped propodi each with one conspicuous, rounded tubercle. Borders of meri and carpi of
second to fourth walking legs (P2-4; Fig. 32c) with thin, rounded tubercles. Abdomen of mature females with
segments 3-5 fused, one transverse ridge along segment 1; ridge with lobed edges. Males unknown.
Description. — Carapace (Fig. 32a) subquadrate, as wide as long (CW/CL = 1.0 in single specimen known);
dorsal surface much depressed, with few large granules, two raised, granular bosses on metagastric region.
Confluence of branchial and mesogastric regions conspicuously depressed, granular. Anterolateral borders of
carapace each with four narrow, pointed anterolateral teeth. Posterior border pointed, with one salient, rounded
tubercle at junction with each posterolateral side, two similar but smaller tubercles on median pointed tip.
Frontal border of carapace divided into two narrow, widely separated lobes. Borders between frontal lobes and
supraorbital borders slightly folded upward medially, ending in sharp angle forming L- shaped fissure before
supraorbital border. Supraorbital borders each with one pointed median lobe. Poslorbital angles long (extend
beyond dorsal border of retracted eye), pointed outward. Cornea of eyes dorsoventrally flattened, wider than base of
long eye peduncle; each peduncle with several conspicuous, elongate, granular tubercles.
Suborbital borders (Fig. 32b) each limited by conspicuous, ventrally inclined, broad pterygostomial lobe on
inner edge (inner suborbital lobe reduced, compressed by pterygostomial lobe) and one rounded, short outer lobe.
Ridge connecting each pterygostomial lobe with pterygostomial region with blunt tooth. Pointed tooth on
pterygostomial region anterior to insertion of cheliped.
Each basal antennal segment slightly expanded outward; three rounded tubercles along expanded margin;
flagellum long, with very long, simple setae. Epistome narrow, vertically inclined, with no apparent teeth. Inner
border of ischia of third maxillipeds straight; surface granular, upper borders rounded. Meri much narrower than
ischia; two large tubercles, broad, serrated upper borders.
Fig. 32. — Paliculus foliatus sp. nov., 2 holotype 5.0 x 5.0 mm, Touho, northeast coast of New Caledonia, Expedition
Montrouzier, unknown depth (MNHN-B 26729): a. carapace, dorsal view: b. suborbital border; c. merus, left third
pereopod, dorsal view; d, dactylus, right fourth pereopod. dorsal view; e, dactylus, left fifth pereopod, dorsal view.
Dorsal and outer borders of propodus of largest cheliped with one conspicuous, rounded, anteriorly inclined
tubercle; smallest cheliped with one large, anteriorly inclined tubercle and two smaller, rounded tubercles on
Source :
532
P. CASTRO
proximal and distal borders. Fingers of both chelipeds with cutting edges. Outer borders of carpi and meri with
conspicuous, elongate, rounded tubercles.
Walking legs (P2-4) slender but with flattened, non-filiform propodi. Upper and lower borders of meri
(Fig. 32c) with thin, rounded tubercles; distalmost tubercle on each anterior border much wider at base, much
higher; similar thin, rounded tubercles on anterior borders of propodi. Anterior and posterior borders ol propodi
with short teeth (most topped by small spine on posterior border); posterior border of dactyli with 1-2 teeth
(Fig. 32d). P3-4 with one dorsal, one ventral carina each (all very slight) on carpi, propodi, and dactyli. Meri of all
walking legs with long plumose setae; dorsal surfaces of propodi of P3-4 each with one row of conspicuous
plumose setae each along anterior and posterior borders, dactyli with one similar row along anterior borders.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair slightly
shorter than second. Fifth pair of pereopods (P5) short (0.7 CL); each merus slender, low tubercles along posterior
border, each propodus with four low tubercles, three of which topped by short spine; each dactylus (Fig. 32c) with
four short, thin spines along posterior border, one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows;
Abdomen of mature females with segments 3-5 fused, one transverse ridge with lobed edges along segment 1 .
Male first pleopods and abdomen of males and immature females unknown.
DISCUSSION. — This species is described from only one small, ovigerous female. Its has a unique, depressed,
leaf-like carapace with pointed posterior border and thin, broad tubercles along the borders of the walking legs (P2-
P4). It shares more characters with Paliculus, specifically with P. kyusyuensis, than with any other genus.
It shares with P. kyusyuensis the morphologies of the supraorbital border (only one median lobe), suborbital
border (one broad, short lobe, inner lobe reduced and compressed by pterygostomial lobe), pterygoslomial region
(blunt tooth on ridge connecting the pterygostomial lobe with the pterygostomial region, pointed tooth on
pterygostomial region anterior to insertion of cheliped; Fig. 32b), female abdomen (fusion of some of the
segments, lobed transverse ridge), narrow and pointed anterolateral teeth, salient tubercles along posterior border,
long and outwardly pointed postorbital angle, and narrow epistome. Yet, there are very strong differences between
the walking legs of both species (very long and fililform in P. kyusyuensis) and body size (smaller in
P. kyusyuensis). There are also differences in the basal antennal segments (rectangular without outward expansion
in P. kyusyuensis) and maxillipeds (much longer exopod and merus, narrower anterior border in P. kyusyuensis).
Some of the outstanding differences between the two species may be accounted to different habitats.
P. kyusyuensis is an inhabitant of soft sediments in moderate depths. The habitat of P. foliatus remains unknown
but its unique shape and shorter walking legs suggest that it lives cryptically on a hard substrate.
SIZE. — Size of female holotype: 5.0 x 5.0 mm (MNHN-B 26729).
ETYMOLOGY. — From phyllon, Greek for leaf, in reference to the leaf-like shape of the sides of the carapace
and tubercles along the walking legs.
DISTRIBUTION. — Known only from New Caledonia. Depth; unknown but probably shallow water.
Paliculus sp.
Fig. 33
Material EXAMINED. — Philippine Islands. Musorstom 3: stn DR 117, 12°31'N, 120°39'E. 92-97 m.
3.06.1985: I $ 2.3 x 2.4 mm (MNHN-B 26730).
Source : MNHN. Pans
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
533
Diagnosis. — Carapace (Fig. 33) subquadrate with one small,
pointed anterolateral tooth on each side (4-5 very small pointed tubercles
present posterior to anterolateral tooth). Dorsal surface of carapace with
scattered, small, rounded tubercles; two slightly raised, granular bosses
on metagastric region; four tubercles along posterior border. Supraorbital
borders each with one conspicuous, bilobed median lobe. Postorbital
angles long and pointed outward. Suborbital border each limited by
conspicuous, ventrally projecting pterygostomial lobe and two broad,
rounded lobes. One tooth on each ridge connecting pterygostomial lobe
with pterygostomial region of carapace. Abdomen of mature females
with segments 3-6 fused and one transverse ridge along each segment 1-3
and proximal portion of fused segments 3-6. Males unknown.
DISCUSSION. — One very small, ovigerous female from the Mindoro
Strait, Philippine Islands (2.3 x 2.4 mm; MNHN-B 26730) shares many
characters with P. kyusyuensis. The specimen, however, lacks all
appendages.
As in P. kyusyuensis, the carapace is subquadrate with few tubercles
(less tubercles that are less pronounced than in P. kyusyuensis) and one
pointed anterolateral tooth on each side (shorter that the two in
P. kyusyuensis), the postorbital angles are thin, curved, and pointed
outward, each supraorbital border has one median lobe (but bilobed and
larger than in P. kyusyuensis), the suborbital borders are limited by
a conspicuous pterygostomial lobe and each has two lobes (only one in
P. kyusyuensis), there is one tooth on the ridge that connects each
pterygostomial lobe with the pterygostomial region of the carapace, and one tooth on the pterygostomial region
(all more conspicuous and salient in P. kyusyuensis). The episternal processes are narrow, pointed, and smaller
than in P. kyusyuensis.
Fig. 33. — Paliculus sp. : 2 2.3 x
2.4 mm, Mindoro Strait. Philip¬
pine Islands, Musorstom 3,
stnDR 117, 92-97 m (MNHN-B
26730): carapace, dorsal view.
Although the species appears to be new, it is not being described as such because of the lack of information on
the morphology and relative size of the walking legs. It is placed in Paliculus due to its close proximity to
P. kyusyuensis, at least as far as the morphology of the carapace is concerned.
The only specimen known is the smallest ovigerous female recorded (2.3 x 2.4 mm) for any of the species of
palicids that was examined in this study.
Distribution. — Known only from the Philippine Islands. Depth: 92-97 m.
Genus RECTOPALICUS nov.
Type Species. — Rectopalicus woodmasoni (Alcock, 1900). Gender: masculine.
DIAGNOSIS. — Frontal border of carapace divided into four pointed or two broad lobes. Carapace subquadrate or
subpyriform, border posterior to postorbital angle nearly straight; truncate anterolateral teeth. Dorsal surface of
carapace with tuberculate bosses or rounded tubercles. Eyes dorsoventrally flattened, wider than peduncle; peduncles
with many granular tubercles. Supraorbital borders each with two rectangular or rounded lobes. Suborbital borders
each with two short lobes. Each basal antennal segment rectangular, without distal expansion. Epistome vertically
inclined or expanded dorsoventrally; two triangular median teeth. Chelipeds nearly equal or unequal in both sexes.
First three pairs of walking legs (P2-4) with flattened (not filiform) carpi, propodi, and dactyli. First pair (P2)
much shorter than second pair and third pairs (P3-4), third pair as long as second pair. Borders of meri and carpi of
P2-4 with nearly equal rounded tubercles; posterior borders of propodi and dactyli with short spines. Fifth pair of
pereopods (P5) reduced (0.5-0.8 CL), thick or slender. Abdomen of mature males, elongate (sides almost parallel to
Source :
534
F. CASTRO
each other), with all segments free, one transverse ridge along each segment 1-4 (may be incomplete in segment
4). Male first pleopods with sinuous basal parts; each distal part uniramous, laterally flattened, dissimilar dorsal
and ventral tips that may have teeth. Abdomen of mature females with all segments free, one transverse ridge along
each segment 1-4.
Description. — Carapace subquadrate or subpyriform; dorsal surface covered with granules or rounded
tubercles, some grouped into bosses. Confluence of branchial and mesogastric regions depressed. Sulcus between
hepatic and branchial regions apparent. Anterolateral borders of carapace nearly straight, with short, truncate teeth,
or straight along anterior portion and conspicuously rounded posteriorly with short, truncate teeth. Thoracic
sternite 7 with much reduced process (episternal process) at each outer edge and posterior to insertion of fifth pair
of pereopods (P5), visible dorsally as rounded process.
Frontal border of carapace divided into four pointed or two broad lobes. Supraorbital borders each with two
rectangular or rounded lobes. Postorbital angles short or long and pointed. Cornea of eyes dorsoventrally flattened,
wider than base of eye peduncle; each peduncle with many granular tubercles. Orbits deep and wide.
Suborbital borders each with short inner and outer lobes. Pterygostomial lobe at each anterolateral angle of
buccal frame straight, with broad, rounded or straight border.
Each basal antennal segment thick, rectangular, and covered with tubercles; flagellum short with few long
plumose setae. Epistome vertically inclined or broad and expanded dorsoventrally. Border of epistome with two
large, triangular median teeth plus two similar outer processes fusing distally with each pterygostomial lobe. Meri
of third maxillipeds narrower than ischia.
Chelipeds nearly equal or unequal in both sexes. Fingers slender, with broadly rounded teeth. First pair of
walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair about same long as second.
First three pairs of walking legs (P2-4) with flattened (not filiform) carpi, propodi, and dactyli; anterior (dorsal) and
posterior (ventral) borders of meri with nearly equal tubercles; anterior border of carpi with rounded or slightly
pointed tubercles; posterior borders of propodi and dactyli with spines or small teeth tipped by a spine. Last two
pairs of walking legs (P3-4) each with broad coxae, having flattened, thick edged anterior and posterior borders.
Fifth pair of pereopods (P5) reduced (0.5-.08 CL), thick or slender; basis-ischia and meri with small tubercles;
spines along posterior border of dactyli.
Abdomen of mature males elongate, with both sides almost parallel to each other, with all segments free,
segments 1-2 dorsoventrally compressed, one transverse ridge along each segment 1-3 or 1-4 (incomplete in
segment 4). Male first pleopods with sinuous basal parts; each distal part biramous or uniramous. thick, laterally
flattened and with dissimilar dorsal and ventral tips; teeth may be present. Second male pleopods short, thin,
slightly curved; distal segment with blunt tip.
Abdomen of mature females broad and rounded, with all segments free, segments 1-2 dorsoventrally com¬
pressed, one transverse ridge along each segment 1-4. Abdomen of immature females, when known, triangular (but
broader than in males), segments 1-2 dorsoventrally compressed, 4-6 fused, transverse ridge along segments 1-4.
Species included. — Three species are known: R. amphiceros sp. nov., R. ampullalus sp. nov., and
R. woodmasoni (Alcock, 1900).
Sexual Dimorphism. — Males are smaller than females as in other palicids. Only one specimen of each sex
were examined in R. amphiceros and R. ampullalus ; more specimens of R. woodmasoni were available for
examination but only three were males. The cheliped propodus of the largest cheliped is thicker in the males of
R. amphiceros and R. woodmasoni, while they are only slightly thicker in the male of R. ampullalus.
Discussion. — Rectopalicus comprises three species that share conspicuously nearly straight lateral borders,
short and truncate anterolateral teeth, and spines on the propodi of the walking legs. There are also some
similarities in the male first pleopods. Two of the species, R. amphiceros and R. woodmasoni, are very similar in
most characters except in their male first pleopods, clearly uniramous in R. amphiceros but seemingly biramous in
R. woodmasoni. R. ampullalus, however, shows differences that are perhaps the result of drastic modifications to
its general body plan, its uniquely subpyriform carapace that is straight anteriorly but rounded posteriorly
(Figs 36a, 37a). The epistome is broader than that of R. woodmasoni (but not so much than in R. amphiceros,
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
535
which is slightly expanded dorsoventrally and as such it may be considered intermediate between R. ampullatus
and R. woodmasoni ), the walking legs (P2-P4) are shorter, last pair of pereopods (P5) more slender and shorter,
and the body is covered by rounded tubercles. Its male first pleopods, however, are uniramous and very similar to
those of R. amphiceros.
There are some similarities between Rectopalicus and Exopalicus new genus, which has a very different body
plan, with a carapace that is conspicuously convex and very short walking legs (Fig. 37d). Both genera share
spines along the propodi of the walking legs and (with the exception of R. woodmasoni) a uniramous, angular
male first pleopod. The similarities between Exopalicus and R. ampullatus, in particular, are more marked: short
walking legs, slender and short P5. and body covered with rounded tubercles. Their respective body plans, however,
are unique and very different from each other, in addition to differences in the epistome and third maxillipeds. The
collection of additional specimens and perhaps new species may clarify the position of both genera within the
Palicidae.
The nearly straight lateral borders and truncate anterolateral teeth of Rectopalicus are not shared with any known
species of western Atlantic or eastern Pacific palicids.
ETYMOLOGY. — From rectus, Latin for straight, and the generic name Palicus, in reference to the diagnostic,
nearly straight lateral borders of the carapace.
Key to the species of RECTOPALICUS
1 . Carapace with anterolateral borders straight anteriorly but rounded posteriorly (Figs 36a,
37a). Abdomen of males and females covered with low, rounded tubercles (Fig. 36e) .
. Rectopalicus ampullatus sp. nov.
— Carapace with nearly straight lateral borders. Abdomen of males and females not covered
by conspicuous tubercles . 2
2. Postorbital angles conspicuously pointed outward (Figs 35a-b). Spines but no teeth along
the posterior border of dactyli of walking legs (P2-4; Fig. 35c). Male first pleopods
uniramous (Figs 35d-e) . Rectopalicus amphiceros sp. nov.
— Postorbital angles not conspicuously pointed outward (Figs 8c, 37c). Teeth (which may be
topped by a spine) along posterior border of dactyli of walking legs (P2-4). Male first
pleopods with two lobes (Figs 34a-b) . Rectopalicus woodmasoni (Alcock. 1900)
Rectopalicus woodmasoni (Alcock, 1900)
Figs 8c, 34, 37c. 53
Palicus Wood-Masoni Alcock, 1900: 795. — Alcock & McArdle, 1903: pi. 67, fig. 3.
Palicus microfrons Sakai, 1963: 228, fig. 7b; 1965: 184, pi. 89, fig. 2; 1976: 594. fig. 325a. pi. 205. fig. 4.
Pseudopalicus woodmasoni - Moosa & Serene, 1981: 35 (in key).
Palicoides microfrons - TAKEDA, 1982b: 17, fig. 2.
MATERIAL EXAMINED. — Japan. Kii Peninsula. Shionomisaki, 33°26.13'N, 135°40.64'E, 160 m. S. Nagai coll..
16.01.1997: I $ (CBM-ZC 3603); 100 m, S. Yamaguchi coll., 22.09.1978: 2 9 (CBM-ZC 4262).
Ryukyu Islands. "Toyoshio-Maru": stn 8, Amami Islands, off Kekaroma Island, 28°04.71'N. 129°27.38'E, 310 m,
K. TSUSHIDA coll., , 10.il. 1994: 1 juv. 9 (CBM-ZC 4951).
Philippine Islands. MusorstomI: stn 57, 13°53.1'N, 120°I3.2'E, 107-96 m. 26.03.1976: 1 9 (MNHN-B
26778).
Thailand (Andaman Sea coast). Stn H38/D, off Phuket island, 9.05.1996: I 6 CL 5.1 mm (PMBC).
Indonesia. Kai Islands. Karubar: stn DW 22, 05°22'S, 133°01'E, 124-85 m. 25.10.1991: 1 9 (MNHN-B 26773).
New Caledonia. "Kandjar", dredgings between 22°40'-22°50'S and 167°10'-I67°30'E. 200-350 m, P. Tirard coll..
7-10.10.1986: 1 9 (MNHN-B 26780).
Smib 5: stn DW 95, 22°59.7'S, 168°19.8'E, 200 m, 14.09.1989: 1 juv. 6 5.4 x 5.7 mm (MNHN-B 26775).
Source :
536
P. CASTRO
Bathus 2: stn DW 723. 22°50.21'S, 167°26.84'E. 430-433 m, 11.05.1993: 1 <3 7.2 x 7.6 mm (MNHN-B 26777). —
Stn DW 728, 22°47.H'S. 167°28.H'E, 241-245 m, 12.05.1993: 1 2 9.4 x 10.3 mm (MNHN-B 26774), 1 2 (MNHN-B
26781).
Bathus 3: sin DW 836, 23°02'S, 166°59'E, 295-306 m. rocks and gravel, 30.1 1.1993: 1 juv. 3 5.5 x 5.6 mm. 1 2
(MNHN-B 26776).
Vanuatu. MUSORSTOM 8: stn CP 1131, 15°38.41S, 167°03.52'E, 140-175 m. 11.10.1994: I juv. 2 (MNHN-B
26779).
Types. — Holotype of Palicus woodmasoni Alcock, 1900: 1 6
9x11 mm (Alcock, 1900), Andaman Islands. Deposit unknown:
Zoological Survey of India, Calcutta ?
Holotype of Palicus microfrons Sakai. 1963: 1 $ 8.0 x 9 mm
(Sakai, 1963), Misaki, Sagami Bay, Japan, 85 m, Emperor of
Japan coll. (Showa Memorial Institute, National Science Museum.
Tsukuba).
Type Locality. — Andaman Islands, India.
Diagnosis. — Carapace (Figs 8c, 37c; ALCOCK & McArdle,
1903, pi. 67, fig. 3; Sakai, 1963, fig. 7b: 1976. fig. 325a, pi. 205.
fig. 4) subquadrate, typically with four truncate anterolateral teeth on
each side; teeth decrease posteriorly in size. Dorsal surface of carapace
covered by large granules and several granular bosses; posterior
border with six elongate tubercles (continuous in small specimens).
Supraorbital borders each with two rounded lobes, outer lobe slightly
narrower than inner lobe. Suborbital border obliquely directed, with
rectangular inner lobe and broad, proximally rounded outer lobe.
Dorsal borders of cheliped propodi with several rounded or elongate
tubercles. Dactyli of walking legs (P2-4) with small teeth along
posterior border (2-3 in P4; Fig. 8c). Abdomen of mature males with
all segments free, one complete transverse ridge along each segment
1-4 (ridge along segment 4 may be incomplete). Male first pleopods
(Figs 34a-b; Takeda, 1982b, fig. 2A) with straight basal parts;
each distal part with two lateral processes, inner one narrower.
Abdomen of mature females with all segments free and one complete
transverse ridge along each segment 1-4 (low ridge may be present
along segment 5).
DISCUSSION. — R. woodmasoni was described as Palicus woodmasoni from a single male specimen collected
in the Andaman Islands (Alcock, 1900). It is not know if the holotype is still extant. The description and
excellent figure (ALCOCK & McArdle, 1903, pi. 67, fig. 3; reproduced here as Fig. 8c) plus additional material
from the western Pacific Ocean and the Andaman Sea not far from the type locality, however, permit its
characterization as a species of Rectopalicus.
Diagnostic characters other than those given in the diagnosis are a border between the anterior lobes and each
supraorbital margin that is folded upward and slightly pointed anteriorly. Each basal antennal segment is
rectangular but thick and slightly expanded along the outer border, and a row of low, rounded tubercles along its
ventral surface. Each eye peduncle has a conspicuous, triangular tubercle on its anterior border, and a much lower
tubercle at the outer end of the tubercle. The pterygostomial lobes project ventrally, forming a Hat. triangular
structure posterior to each inner suborbital lobe. The epistome is vertically inclined, with two short, slightly
pointed median teeth flanked on each side by a large triangular outer process and a thin, rounded margin before each
pterygostomial lobe. The third pair of walking legs (P4; Fig. 37c) has short meri (0.4 CL), each armed with one
row of a few, elongate teeth along the upper border and one broad and slightly pointed distal tooth. Each dactylus is
slender (0.3 CL) and its posterior border is armed with 2-3 small spines or teeth (see below). The fifth pair of
Fig. 34. — Rectopalicus woodmasoni
(Alcock, 1900), 3 7.2 x 7.6 mm, off
south coast of New Caledonia, Bathus
2, stn DW 723, 430-433 m (MNHN-B
26777): a. left male first pleopod,
ventral view; b. left male first
pleopod, distal part, dorsal view.
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
537
percopods (P5) is thick and short (0.7 CL). The meri are slender and tuberculate, with conspicuous, rounded and
triangular tubercles (small specimens have larger tubercles, each with a short spine), plus a few short spines and
plumose setae along the posterior border; each propodus with tubercles and 4-7 spines; each dactylus with 1-2 thick
spines along posterior border, microscopic teeth, 1-2 short distal spines along anterior border, and one terminal
pointed tooth.
Two female specimens (MNHN-B 26774, Fig. 37c; MNHN-B 26773) had a frontal border with only three
lobes, the median one being entire without a sulcus.
Palicus microfrons Sakai, 1963, which was described from one female from Japan, has a subquadrate carapace
with straight lateral borders and truncate anterolateral teeth as in R. woodmasoni. Sakai (1963: 230) mentioned
that his new species was "closely related" to Palicus woodmasoni but that they differed in the "shape and arrange¬
ment of the frontal and lateral teeth " and of "features of the granules and tubercles of the dorsal surface of the cara¬
pace". These differences were most certainly based only on the examination of Alcock's description and figure
(Alcock & McArdle, 1903, pi. 67, fig. 3; reproduced as Fig. 8c) and not by the examination of the holotypc
that is presumably in Calcutta. ALCOCK mentioned that there were three anterolateral teeth in his species. The
figures of Sakai (1963, fig. 7b; 1976, fig. 325a, pi. 205, fig. 4), however, show four teeth, while those of
Takeda (1982b, figs 2B-C, as Palicoides microfrons) show five.
All specimens examined in this study had four anterolateral teeth. Tubercles along the edge beyond the last
tooth may be easily interpreted as small teeth. In a female specimen from Japan (CBM-ZC 4226) one such lubercle
appeared on the right side but none on the left. In small specimens the separation between the teeth becomes less
well defined. The number of teeth along the anterolateral borders of palicids has always been a matter of
interpretation since the postorbital angles are considered as teeth by some and small, tubercle-like teeth may not.
The anterolateral borders of the carapace of P. microfrons carapace were described by Sakai (1963, 228. fig.
7b; 1976, pi. 205, fig. 4) as being "almost straight and slightly divergent backwards". All specimens examined
during this study (which included four from Japan) had carapaces that were straight but clearly divergent on a
posterior direction, as noticed by Takeda (1982b, figs 2B-C, as Palicoides microfrons) and shown in a figure by
Sakai (1976, fig. 325a).
The only significant difference between P. woodmasoni and P. microfrons is in the fact that no mention was
made in ALCOCK's description of P. woodmasoni of the presence of spines or teeth on the dactyli of the walking
legs. Small teeth were shown in his figure (Alcock & McArdle, 1903, pi. 67, fig. 3; reproduced as Fig. 8c).
Sakai (1963: 230) described "minute spinules" on the dactyli of P. microfrons, as shown in his first figures
(Sakai, 1963, fig. 7b; 1976, pi. 205, fig. 4), but were absent in another (Sakai. 1976, fig. 325a). Either the
presence of small spines in the holotypc was overlooked by ALCOCK or the dactyli actually had no spines, a
situation that has been observed in at least some of the legs of some of the specimens that were examined.
The presence of spines or teeth (if any) on the dactyli of the walking legs is a variable character. Most
specimens examined in this study had either teeth or spines. The larger specimens tend to have teeth and the
smaller spines but this varies. For example, the largest specimen examined, a female from Japan (9.7 x 10.3 mm;
CBM-ZC 4262), had spines on the dactyli of P2-4 of the right side and P2 on the left side but no spines or teeth
on the P3-4 of the left side (only half of the dactylus of P4 remained); a smaller female from the same locality
(8.6 x 9.1 mm; CBM-ZC 4262) with four walking legs had smooth dactyli in three legs but a spine on the P2:
a small female from Japan (6.7 x 7.5 mm; CBM-ZC 3603) with three legs had teeth topped by a spine on a P2.
a very short tooth on a P3, and a short tooth topped by a spine on a P4: a large female from New Caledonia (9.4 x
10.3 mm; MNHN-B 26774) had teeth topped by spines, lacking a spine, or only single spines in the four legs that
remained.
Only one specimen from the Andaman Sea near the type locality was examined. Although the male specimen
(CL 5.1 mm; PMBC) was incomplete, the dorsal surface of the carapace, suborbital region, and abdomen agree
with the Japanese and western Pacific specimens that were examined in this study. The legs and first pleopods,
which would have provided the key pieces of evidence for the probable separation between the two species, were
unfortunately missing.
MOOSA & Serene (1981: 35) did not examine any specimens but included Palicus woodmasoni in their new
genus, Pseudopalicus , but strangely enough without mentioning Palicus microfrons. No mention of their syn-
Source : MNHN. Pans
538
P. CASTRO
onymy was made, however. There is nevertheless enough evidence to consider P aliens microfrons Sakai a junior
subjective synonym of Palicus woodmasoni Alcock, now included in the new genus Rectopalicus.
There are also close similarities between R. woodmasoni and R. amphiceros sp. nov., a species being described
from only two specimens from New Caledonia. R. amphiceros, however, has postorbital angles that are
conspicuously pointed outward and there are no teeth on the dactyli ot the walking legs, among other characters
(see description of R. amphiceros below).
TAKEDA (1982b) placed Palicus microfrons Sakai in Palicoides Moosa & Serene, 1981 based on the presence
of a "subterminal process" on the male first pleopods. Indeed, the subquadrate shape of the carapace, the presence
of truncate anterolateral teeth, and the relatively thicker P5 of R. woodmasoni are characteristics of Palicoides.
The two lateral processes of the pleopods (Figs 34a-b; Takeda, 1982b, fig. 2A) nevertheless do not bear
a close resemblance to those of Palicoides longimanus (Miyake, 1936) (Fig. 43a) or of P. whitei (Miers. 1884)
(Figs 43b-d), the later characterized by a very conspicuous, arched, and spinous process. The pleopod illustrated by
Takeda had a longer inner process than that illustrated here (Fig. 43a) and the process showed a row of setae. The
difference in the size of both specimens is small (Japanese specimen of Takeda: 6.2 x 6.5 mm; specimen
illustrated in Figs 34a-b: 7.2 x 7.6 mm, MNHN-B 26777) but it may explain the differences observed. The male
first pleopods of two juvenile males (5.4 x 5.7 mm, MNHN-B 26775; 5.5 x 5.6 mm, MNHN-B 26776) were very
thin and unspecialized into any defined structures. The male first pleopods ol R. woodmasoni are somewhat closer
to those of R. amphiceros sp. nov. (Figs 35d-e), and R. ampullatus sp. nov. (Figs 36f-g) in their general shapes.
In addition to the differences in the male first pleopods, other characters support the placement of Palicus
microfrons in Rectopalicus, not in Palicoides: 1) four lobes (inner pair rounded, narrow, and short; outer pair very
broad, rounded, and typically as high as first pair) on anterior border (only two in Palicoides), 2) segments 2-3 of
the antennae are relatively narrow and there are conspicuous setae on the flagellum (segments 2-3 very wide and
provided with long setae, few setae on flagellum in Palicoides ), 3) cornea ol eyes is dorsoventrally flattened, clearly
wider than peduncle, and there is one large distal tubercle on each peduncle (spherical, almost as wide as peduncle
and conspicuous crescent-shaped process on peduncle in Palicoides), 4) each supraorbital border has two lobes (only
one lobe in Palicoides), 5) presence of four anterolateral teeth on each side of the carapace (only two in Palicoides),
6) chelipeds are relatively short (much longer and slender in Palicoides), 7) meri ol walking legs (P2-4) have
conspicuous tubercles on the upper and lower borders (microscopic tubercles in Palicoides), 8) abdomen ol mature
females has all segments free (segments 5-6 fused in Palicoides), and 9) epistome with median teeth and outer
process (smooth in Palicoides).
Size. — Maximum size among specimens examined: 9.7 x 10.3 mm (female, CBM-ZC 4262); 7.2 x 7.6 mm
(male, MNHN-B 26777).
Distribution. — Andaman Islands (Alcock. 1900) and Japan (Sakai, 1963, 1976; Takeda, 1982b; as
P. microfrons) and now recorded from the Andaman Sea coast of Thailand, Banda Sea (Indonesia), and Coral Sea
(Vanuatu and New Caledonia) (Fig. 53). Depth: known reliably between 107 and 430 m; also collected in a trawl
between 85 and 124 m.
Rectopalicus amphiceros sp. nov.
Figs 35
Material EXAMINED. — New Caledonia. Musorstom 4: stn DW 203, 22°35.8'S. I67°04.8'E. 105-110 m,
27.09.1985: 1 6 holotype 4.6 x 4.8 mm (MNHN-B 26731). — Stn DW 204. 22°37.0'S, 167°05.7'E. 120 m,
27.09.1985: 1 juv. $ paratype 4.1 x 4.5 mm (MNHN-B 26732).
Types. — Holotype: 1 <3 4.6 x 4.8 mm, Musorstom 4, stn DW 203 (MNHN-B 26731).
Paratype: 1 juv. 2 4.1 x 4.5 mm, Musorstom 4, stn DW 204 (MNHN-B 26732).
Type Locality. — Off south coast of New Caledonia, 22°35.8'S, 167°04.8’E, 105-1 10 m.
Source : MNHN, Pans
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
539
Diagnosis. — Carapace (Fig. 35a) subquadrate, with four short, rounded anterolateral teeth on each nearly
straight side; teeth slightly decrease posteriorly in size. Dorsal surface of carapace with large granules and several
granular bosses; posterior border with four very elongate tubercles. Supraorbital borders each with two rounded
lobes. Postorbital angles very long, narrow, pointed outward so they extend only to about half of dorsal border of
retracted eye. Suborbital borders (Fig. 35b) obliquely directed, each with rectangular inner lobe and broad,
proximally rounded, outer lobe. Dorsal borders of cheliped propodi with several rounded or elongate tubercles.
Dactyl i of walking legs (P2-4) with thick spines along posterior border (two spines on P4; Fig. 35c). Abdomen of
males with all segments free and one complete transverse ridge along each segment 1-3, incomplete ridge along
segment 4. Male first pleopods (Figs 35d-e) with straight basal parts armed with teeth and one dorsal tubercle; each
distal part with two broad lateral processes, inner one narrower. Abdomen of mature females unknown.
Musorstom4, stn DW 203, 105-110 m (MNHN-B 26731): a, carapace, dorsal view; b. suborbital border;
c, dactylus, right fourth pereopod, dorsal view; d. left male first pleopod. lateral (inner side) view: e. left male first
pleopod, distal part, lateral (outer side) view.
DESCRIPTION. — Carapace (Fig. 35a) subquadrate, only slightly wider than long (CW/CL = 1 .0); dorsal surface
of covered by large granules and several granular bosses (several small, rounded bosses on branchial and gastric-
regions, one row across lower third of carapace). Anterolateral borders of carapace nearly straight, each with four
Source
540
P. CASTRO
short, rounded teeth decreasing posteriorly in size. Posterior border with four elongate tubercles and scattered
plumose setae.
Frontal border of carapace divided into four lobes, inner pair rounded, narrow and short, outer pair very broad,
rounded, shorter than first pair. Borders between frontal lobes and supraorbital borders folded upward, with almost
straight margin ending in sharp angle forming V-shaped fissure before supraorbital border. Supraorbital borders
each with two rounded lobes, outer lobe slightly narrower than inner lobe. Postorbital angles long, slender, and
pointed outward, so they only extend to about half of dorsal border of retracted eye. Cornea of eyes dorsoventrally
flattened, wider than base of eye peduncle; each peduncle with one large distal tubercle.
Suborbital borders (Fig. 35b) obliquely directed, each with rectangular inner lobe, broad, proximally rounded,
outer lobe. Pterygostomial lobes short, straight, with rounded borders. Ridge connecting each pterygostomial lobe
with pterygostomial region with row of pointed teeth. Pointed tooth on pterygostomial regions anterior to
insertion of chelipeds.
Each basal antennal segment slender and rectangular (row of rounded tubercles along ventral surface); flagellum
long, with very few, simple setae. Epistome slightly expanded dorsoventrally, with two triangular median teeth,
each flanked by triangular, narrow, pointed outer process and thin, rounded margin before pterygostomial lobe.
Inner border of ischia of third maxillipeds straight; surface granular, upper borders rounded to slightly pointed. Mcri
narrower than ischia; oblong upper borders.
Dorsal borders of chelipcd propodi each with 3-4 rows of rounded and pointed tubercles. Fingers with 3-4
broadly rounded teeth. Carpi short, outer border with pointed tubercles; meri slender, with rounded tubercles.
First three pairs of walking legs (P2-4) of holotype slender but flattened. Upper and lower borders of meri with
rounded, proximal tubercles; distalmost tubercle on each anterior border much wider at base, much higher, slender,
pointed or rounded, directed distally. Carpi of walking legs (P2-4) elongate: posterior borders of propodi and dactyli
with small spines (Fig. 35c). Each P2 with one dorsal carina. 2-4 spines along posterior border of propodus; one
minute spine along posterior border of dactylus. P3-4 each with one dorsal, one ventral carina on carpus, propodus,
and dactylus; 1-3 spines along posterior border of propodus. Meri of all walking legs with scattered plumose setae;
dorsal surfaces of propodi and dactyli of P3-4 each with one row of conspicuous plumose setae along anterior and
posterior borders.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); fourth pair slightly
longer than third. Fifth pair of pereopods (P5) short (0.8 CL); each merus very slender, with plumose setae along
posterior border; each propodus and dactylus with many long simple and plumose setae along both borders; each
dactylus with one terminal pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of males with all segments free. One complete transverse ridge along each segment 1-3 (low,
incomplete ridge along most of segment 4). Male first pleopods (Figs 35d-e) with sinuous basal parts, borders
with teeth and one tubercle on each dorsal side; each distal part laterally flattened, with one pointed dorsal edge and
one straight ventral edge bordered by minute teeth. Abdomen of mature females unknown.
DISCUSSION. — P. amphiceros is described from only one complete specimen, the male holotype. The
paratype, a juvenile female, had only two walking legs. Notwithstanding the scarcity of material and its very close
proximity to R. woodmasoni, it is described as a separate species on account of the unique male first pleopods and
the outward orientation of its postorbital angles. It can be separated from R. woodmasoni by these two characters
and the presence of shorter and slightly rounded anterolateral teeth (conspicuously truncate in R. woodmasoni), four
tubercles along the posterior border of the carapace (six in R. woodmasoni), and by having spines along the
posterior border of the dactyli of the walking legs (teeth that may be topped with spines in R. woodmasoni). Some
of these characters, however, may be present only in small specimens, the only type of specimens thus far known.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PAL1CID CRABS
541
The unique male first pleopods are laterally flattened like those of R. ampullatus but they are much more
complex. Each basal part has thick teeth and one large dorsal tubercle vary similar to that of Pseudopalicus
undulatus (Figs 14d-e). There are also similarities in their general shape with the male first pleopods of Exopalicus
maculatus (Edmondson, 1930) (Fig. 38d). It lacks, however, the large lateral teeth of E. maculates.
Size. — Size of the only two specimens known to exist: 4.1 x 4.5 mm (female paratype. MNHN-B 26732):
4.6 x 4.8 mm (male holotype, MNHN-B 26731).
Etymology. From the Latinized word for amphikeros , Greek for having two horns, in reference to the
characteristic postorbital angles, which are pointed in an outward direction as in the horns of cattle.
Distribution. — Known only from the Coral Sea (New Caledonia). Depth: 110-120 m.
Rectopalicus ampullatus sp. nov.
Figs 36, 37a-b, 61 a
Material EXAMINED. — Loyalty Islands. MUSORSTOM 6: stn DW 462, 2I°05.IO'S, 167°26.85'E 200 m
21.02.1989: I <J holotype 6.2 x 6.3 mm (MNHN-B 26782).
Futuna Island. MUSORSTOM 7: stn DW 513, I4°13'S, 178°II'W, 260-300 m, 12.05.1992: I 2 paratype 7 9 x
8.7 mm (MNHN-B 26783).
Types. — Holotype-. 1 8 6.2 x 6.3 mm, MUSORSTOM 6, stn DW 462 (MNHN-B 26782).
Paratype-. 1 $ 7,9 x 8.7 mm, MUSORSTOM 7, stn DW 513 (MNHN-B 26783).
Type Locality. — Loyalty Islands, 21°05.10'S, 167°26.85'E. 200 m.
DIAGNOSIS. — Carapace (Figs 36a, 37a, 61a) subpyriform, with anterolateral borders anteriorly straight,
posteriorly rounded; rounded portion with four short, truncate teeth on each side. Dorsal surface of carapace with
many rounded tubercles and tuberculate bosses. Supraorbital borders each with two very short lobes. Suborbital
borders (Fig. 36c) each with short, irregular inner and outer lobes. Dorsal borders of cheliped propodi with rounded
tubercles. Dactyli of walking legs (P2-4) with teeth along posterior border (two teeth in P4; Fig. 36d). Abdomen
ol males (Fig. 36e) with all segments free, one complete transverse ridge along each segment 1-3 (incomplete ridee
along each median portion of segment 4). Male first pleopods (Figs 36f-g) with sinuous basal parts; each distal
part laterally flattened with irregular dorsal tip. Abdomen of mature females with all segments free, one complete
transverse ridge along each segment 1-4 (low ridge along segment 5).
Color: The carapace of the live holotype, a male (Fig. 61a), was yellow-brown with a few symmetric, white
streaks. The meri and carpi of the second and third walking legs (P3-4) were of the same color, the remaining
portions white. A red-brown band crossed the propodi of the second walking legs (P2).
Description. — Carapace (Figs 36a, 37a, 61a) subpyriform, only slightly wider than long (CW/CL = 1.0-
1.1); dorsal surface covered with many rounded tubercles and tuberculate bosses, most conspicuous of which are on
branchial and metagastric regions. Carapace straight along anterior portion, broad posteriorly; broad portion with
four short, truncate teeth on each side (teeth, particularly first two, may be divided into separate teeth or tubercles).
Episternal processes reduced (short), granular, rounded. Posterior border with many rounded tubercles, no setae.
Frontal border of carapace divided into two broadly rounded, short lobes. Borders between frontal lobes and
supraorbital borders as three successive curved or straight, slightly folded-upward portions separated from each other
by pointed tips (first two pointed tips together with pointed lobes gives carapace a four-lobed appearance).
Supraorbital borders each with two very short, straight lobes. Postorbital angles short (not extending to dorsal
border of retracted eye) with slightly pointed tips. Cornea of eyes dorsoventrally flattened, wider than base of eye
peduncle; each peduncle with many granular tubercles (most dorsal, distal tubercles the two most conspicuous).
Anterior portion of body between frontal lobes of carapace and posterior border of meri of third maxillipeds
(Figs 36b, 37b) nearly fiat; epistome. pterygostomial lobe, and suborbital border fused together. Suborbital borders
Source :
542
P. CASTRO
(Fig. 36c) each with short inner and outer lobes, both with irregular, tuberculate borders. Pterygostomial lobe at
each anterolateral angle of buccal frame short, straight, with rectangular anterior border (broad, flat, irregular border
when seen anteriorly).
Each basal antennal segment thick, rectangular, covered with thick tubercles; flagellum short with few long
plumose setae. Epistome broad, expanded dorsoventrally, straight except concavity along median portion. Border of
epistome with two large, triangular median teeth plus two similar outer processes fusing distally with each
pterygostomial lobe. Inner border of ischia of third maxillipcds broadly triangular; surface with rounded tubercles,
upper borders straight with oblong tip. Meri as wide as ischia; straight-edged, triangular upper borders.
Chelipeds nearly equal in both sexes. Dorsal and outer borders of cheliped propodi with rows of small, rounded
tubercles. Fingers with very short, rounded teeth. Carpi short, outer borders with low, rounded tubercles; meri
slender, outer borders with low, rounded tubercles.
First three pairs of walking legs (P2-4; Figs 37a-b) slender but flattened. Anterior (dorsal) and posterior
(ventral) borders and dorsal and ventral surfaces of meri with nearly equal, rounded tubercles. Anterior borders of
carpi with rounded tubercles; posterior borders of propodi with minute, pointed tubercles, spines on P2. Posterior
borders of dactyli with teeth capped with a spine. Each P2 with one dorsal, one ventral row of tubercles on carpus;
one dorsal carina, two ventral rows of tubercles, 2-3 short spines on propodus; one dorsal, one ventral carinae,
2-3 teeth on posterior border of dactylus. Each P3 with one ventral row of tubercles on carpus; one dorsal row of
tubercles, one carina, two ventral rows of tubercles on propodus; one dorsal, one ventral carinae, 2-3 teeth on
posterior border of dactylus. Each P4 with two dorsal, one ventral rows of tubercles on carpus; one dorsal, two
ventral carinae on propodus; one dorsal, one ventral carinae, two teeth on posterior border of dactylus (Fig. 36d).
Meri of all walking legs with very few, scattered plumose setae; dorsal surfaces of propodi and dactyli of P3-4 with
one row of conspicuous plumose setae along anterior and posterior borders (setae along posterior borders of dactyli
particularly longer).
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair about same
size as second. Fifth pair of pereopods (P5) very short (0.5 CL); each merus slender, tuberculate surface; each
propodus with low tubercles; each dactylus with three short, thin spines along posterior border, one terminal
pointed tooth.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of males (Fig. 36e) with all segments free, covered with rounded tubercles. One complete transverse
ridge along each segment 1-3; ridges bordered by conspicuous, rounded tubercles. Male first pleopods (Figs 36f-g)
with sinuous basal parts; each distal part laterally flattened; ventral tip rounded, dorsal tip with blunt, pointed
processes.
Abdomen of mature females broad, rounded, covered with rounded tubercles, with all segments free, one
transverse ridge along each segment 1-4 (very slight along segment 5). Abdomen of immature females unknown.
DISCUSSION. — The unique shape of the carapace of P. ampiillatus parallels a remarkable shortening and
flattening of the anterior part of the body. Several morphological changes have taken place as part of this
readjustment. The epistome has expanded dorsoventrally (Fig. 36b). Each pterygostomial lobe is short and straight
but it has been laterally expanded to fuse with the edges of the epistome. As a result, the inner portion of the
suborbital margin is thicker and makes contact with the laterally expanded pterygostomial lobe. The third
maxillipeds, which form the ventral boundary between the flat anterior portion and the rest of the body, are folded
so that each merus is bent nearly 90° in relation to the straight ischium.
The unique shape and coarse tuberculation of R. ampiillatus is most probably correlated with a particular
habitat that may be unique among palicids. Perhaps it burrows in coarse sediments while leaving the flattened
anterior portion of the body above the bottom. Short but dorsoventrally flattened walking legs may be involved in
Source : MNHN, Pans
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
543
digging. It may also be involved in swimming short distances, as in the case of Exopalicus maculatus
(Edmondson, 1930) (see below).
Fig. 36. — Rectopalicus ampullatus sp. nov., <J holotype 6.2 x 6.3 mm. Loyalty Islands. Musorstom 6. stn DW 462.
200 m (MNHN-B 26782): a, carapace, dorsal view; b. carapace, anterior portion; c, suborbital border; d, dactylus.
right fourth pereopod. dorsal view; e. abdomen; f, left male first pleopod, lateral (inner side) view; g. left male first
pleopod. distal part, lateral (outer side) view.
Size. — Maximum size among specimens examined: 7.9 x 8.7 mm (female paratype; MNHN-B 26783); 6.2 x
6.3 mm (male holotype; MNHN-B 26782).
Etymology. — From ampulla, Latin for bottle or flask, in reference to the unique flask-shape of the body:
narrow anteriorly, wider and rounded posteriorly.
Distribution. — Loyalty Islands and Futuna Island, southwestern Pacific Ocean. Depth: known reliably
between 200 and 260 m, collected loo in a trawl between 260-300 m.
Genus EXOPALICUS nov.
TYPE Species. — Palicus maculatus Edmondson, 1930 by monotypy. Gender: masculine.
Diagnosis. — Frontal border of carapace divided into two rounded, short lobes. Carapace hexagonal. Three
triangular anterolateral teeth. Dorsal surface of carapace convex, covered with rounded tubercles and tuberculate
Source
544
P. CASTRO
bosses. Eyes dorsoventrally flattened, wider than peduncle; peduncles with many granular tubercles. Supraorbital
borders each with one short, rectangular lobe. Suborbital borders each with two short lobes. Each basal antennal
segment rectangular, without distal expansion. Epistome slightly expanded dorsoventrally; two triangular median
teeth. Chelipeds unequal in both sexes. First three pairs of walking legs (P2-4) with flattened (not filiform) carpi,
propodi, and dactyli. First pair (P2) much shorter than second pair and third pairs (P3-4), third pair as long as
second pair. Borders of men and carpi of P2-4 with nearly equal rounded tubercles; posterior border of propodi and
dactyli with short spines. Fifth pair of pereopods (P5) reduced (0.5 CL), thick. Abdomen ol mature males, elongate
(sides almost parallel to each other), all segments free, one transverse ridge along each segment 1-4 (may be
incomplete in segment 5). Male first pleopods with sinuous basal parts; each distal part uniramous, lateially
flattened, dissimilar dorsal and ventral tips with teeth. Abdomen of mature females with all segments free, one
transverse ridge along each segment 1-3 (may be low in segment 4).
DESCRIPTION. — Carapace hexagonal; dorsal surface conspicuously convex and covered with rounded tubercles,
some grouped into bosses. Confluence of branchial and mcsogastric regions depressed. Sulcus between hepatic and
branchial regions apparent. Anterolateral borders of carapace with three broad, triangular teeth. Thoracic sternitc 7
with process (episternal process) at each outer edge and posterior to insertion ol fifth pair of pereopods (P5), visible
dorsally as oblong process.
Frontal border of carapace divided into two broad lobes. Supraorbital borders each with one short, rectangular
lobe formed by two very shallow notches. Postorbital angles very short and triangular. Cornea of eyes
dorsoventrally flattened, wider than base of eye peduncle; each peduncle very short, with many granular tubercles.
Orbits deep. wide.
Suborbital borders each with broadly rounded, short inner and outer lobes. Pterygostomial lobe at each
anterolateral angle of buccal frame with slight vertical inclination and broad, rounded to slightly pointed border.
Each basal antennal segment short, rectangular; flagellum short with few long plumose setae. Epistome broad,
slightly expanded dorsoventrally. Border of epistome with two large, triangular median teeth plus two pointed outer
processes fusing distally with each pterygostomial lobe. Meri of third maxillipeds narrower than ischia.
Chelipeds unequal in both sexes. Fingers thick (very short in largest cheliped), with cutting edge and no teeth.
Inner surfaces of propodi of males with thick clump of plumose setae. First pair of walking legs (P2) shorter,
more slender than second and third pairs (P3-4); third pair about same long as second. First three pairs o! walking
legs (P2-4) with flattened (not filiform) carpi, propodi. and dactyli; anterior (dorsal) and posterior (ventral) borders
and dorsal and ventral surfaces of meri with nearly equal, rounded tubercles; anterior border of carpi with rounded or
slightly pointed tubercles; posterior borders of propodi and dactyli with thick spines or (on dactyli) small teeth
tipped by a similar spine. Last two pairs of walking legs (P3-4) each with broad coxae, having flattened, thick
edged anterior and posterior borders. Fifth pair of pereopods (P5) much reduced (0.5 CL), thick; basis-ischia and
meri with small tubercles; spines along posterior border of dactyli.
Abdomen of mature males elongate, with both sides almost parallel to each other, with all segments free,
segments 1-2 dorsoventrally compressed, one transverse ridge along each segment 1-4 (incomplete in segment 5).
Male first pleopods (Fig. 38d) with sinuous basal parts; each distal part uniramous, thick, laterally flattened, with
dissimilar dorsal and ventral tips with teeth. Second male pleopods short, thin, slightly curved; distal segment with
blunt tip.
Abdomen of mature females broad and rounded, with all segments free, segments 1-2 dorsoventrally
compressed, one transverse ridge along each segment 1-3 (may be low in segment 4).
SPECIES INCLUDED. — Only one species is known, Exopalicus maculatus (Edmondson, 1930).
SEXUAL Dimorphism. — Males are smaller than females as in other palicids. The chelipeds are unequal in
both sexes. There is a thick clump of plumose setae on the inner surfaces of the cheliped propodi in males, as in
five other species of Indo-west Pacific palicids (see "Sexual Dimorphism and Mating Behavior").
DISCUSSION. — Exopalicus shares with Rectopalicus the presence of short walking legs with spines along the
posterior border of the propodi. There are also some similarities in the epistome (slightly expanded dorsoventrally
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
545
as in R. amphiceros). male first pleopods (laterally flattened with spines, as in R. amphiceros\ Figs 35d-e), and
conspicuous rounded tubercles on the carapace and appendages as in R. ampullatus. Nevertheless, the hexagonal,
dorsally convex (almost rounded) shape of its carapace, large and triangular anterolateral teeth, and complex male
first pleopods with terminal and lateral teeth of different sizes and shapes set it apart from the subquadrate or
pyriform carapace and body plan of Rectopalicus and its simpler male first pleopod. Other characteristic features of
Exopalicus are the very short eye peduncles and postorbital angles (perhaps as a result of the rounding of the
carapace), more conspicuous episternal process, and the presence of only one very short lobe on each supraorbital
border.
The walking legs and fifth pair of pereopods are shorter in relation to carapace size in Exopalicus than in other
Indo-west Pacific palicids. The shortening of the walking legs may be an adaptation to living in hard substrate
habitats, such as coral, where swimming is very much limited.
No known species of western Atlantic or eastern Pacific palicids resembles the general body plan of
Exopalicus.
ETYMOLOGY. — From ex. Latin for out of, and the generic name Palicus, in reference to the unique triangular
outline of the conspicuously convex carapace, which sets it apart from other genera of palicids.
Fig. 37. — a-b. Rectopalicus ampullatus sp. nov., d holotype 6.2 x 6.3 mm. Loyalty Islands, Musorstom 6.
stn DW 462. 200 m (MNHN-B 26782): a, dorsal view: b. anterior view of carapace. — c. Rectopalicus woodmasoni
(Alcock, 1900), $ 9.4 X 10.3 mm. off southwest coast of New Caledonia. Bathus 2, stn CP 728. 241-245 m (MNHN-
B 26774): dorsal view. — d. Exopalicus maculatus (Edmondson, 1930), 2 9.5 x 13.1 mm. Oahu. Hawaiian Islands,
J. Hoover & D. Dickey coll.. 3-4 m (MNHN-B 26784): dorsal view.
Source :
546
P CASTRO
Exopalicus maculatus (Edmondson, 1930)
Figs 37d, 38. 56. 61b
Palicus tuberculatus Edmondson. 1925: 57, figs 8e-g. pi. 4, figs A-G; 1933: 268; 1946: 310.
Palicus maculatus Edmondson, 1930: 15. figs 6a-g, pi. 1, fig. C; 1933: 268, fig. 158c; 1946: 310. fig. 184c.
Cymopolia medipacifica Edmondson, 1962: 9, figs 4a-c.
Cymopolia maculata - Edmondson. 1962: 1 1. figs 4d-e.
Pseudopalicus sp. - Hoover, 1998: 287. unnumb. fig.
Material EXAMINED. — La Reunion. Cap Houssaye, night diving, on Millepora coral, 15 m, J. Beneteau coll..
10.1973: 1 8 5.5 x 7.2 mm, 1 2 5.6 x 6.8 mm (MNHN-B 26785).
Marshall Islands. Enewetak Atoll. Parry Island, lagoon, 3-5 m, J.S. Garth coll.. 22.07.1957: I 8 , I 2 (LACM).
Hawaiian Islands. Oahu. Waikiki, Ostergaard coll.. 11.1922: 1 8 , 1 2 (BPBM 932). — Waikiki. 1923: 1 2
(BPBM 3266). — Waikiki, 30.03.1930: 1 8 holotype of Palicus maculatus Edmondson. 1930, 5.6 x 7.2 mm (BPBM
3337). — Honolulu, Ala Moana Beach Park. Magic Island Boat Channel, rock wall, night diving, 5 m, J. Hoover coll..
7.10.1997: 1 2 (QM W24746); rock wall, night diving, 3-4 m, J. Hoover & D. Dickey coll., 17.06.1999: 1 2 9.5 x
13.1 mm (MNHN-B 26784); on boulders, night collection, 1 m, D. Takaoka coll., 1.08.1999: 1 2 9.8 x 14.0 mm
(MNHN-B 26999). — Blow Hole, 18 m, R. HOLCOM coll., 02.1996: 1 8 (QM W21847); 15 m, 01.1997: 2 8 7 I x
10.2 mm, 7.4 x 10.3 mm (QM W24745).
Molokai. Kaunakakai, Ball & C.H. Edmondson coll., 22.02.1922: I 2 (BPBM 940).
Types . — Holotype of Palicus tuberculatus Edmondson, 1925 apparently lost: 1 9 9 x 12 mm, Ocean
(= Kure) Island, 28°25'N. 178°25'W, "shallow water on the reef’ (Edmondson, 1925) (BPBM 1 134).
Holotype of Palicus maculatus Edmondson, 1930; 1 8 5.6 x 7.2 mm, Oahu, Hawaiian Islands (BPBM 3337).
Type Locality. — Waikiki reef, Oahu, Hawaiian Islands, shallow water.
Diagnosis. — Carapace (Figs 37d, 38a) with three triangular, broadly pointed anterolateral teeth on each side;
large tubercle posterior to postorbital angle, several small, rounded tubercles between the anterolateral teeth. Dorsal
surface of carapace convex, with many rounded tubercles and tuberculate bosses. Supraorbital borders each with two
shallow notches resulting in one very short lobe. Suborbital borders (Fig. 38b) each with very broad, short inner
and outer lobes. Dorsal borders of cheliped propodi with rounded tubercles. Dactyli of walking legs (P2-4) with
thick spines along posterior borders (one tooth in P4; Fig. 38c). Abdomen of mature males with all segments free,
one complete transverse ridge along each segment 1-4, incomplete ridge along median portion of segment 5. Male
first pleopods (Fig. 38d) with sinuous basal parts; each distal part with one flattened process with teeth. Abdomen
of mature females with all segments free, one complete transverse ridge along each segment 1-3 (low ridge along
segment 4).
Color. Individuals photographed live in Hawaii show a very wide range of color patterns: pink carapace with
large red dots (Fig. 61b), pink with irregular white markings (Hoover, 1998: 287, unnumbered figure), irregular
bright or light orange spots, and bright red carapace and white legs. Another was described as "dull, dark grayish
brown" (Hoover, personal communication).
Discussion. — This species was described as Palicus tuberculatus by EDMONDSON (1925). The holotype
(BPBM 1 134) is apparently lost, but three specimens identified by EDMONDSON as P. tuberculatus and mentioned
by him in the description (BPBM 940, BPBM 942) agree with the actual description. One exception, however, is
in the number of anterolateral teeth on each side of the carapace. Edmondson (1925: 57; pi. 4, figs A. E) gives
its number as five but the number, shape, and relative length vary widely among the specimens examined in this
study, even between the right and left side of the same individuals. For example, a male identified by EDMOND¬
SON (BPBM 932) had four obvious teeth and tubercles on the left side but five on the right. The anterolateral teeth
are perhaps more correctly given as three. Most, in not all of the teeth typically have a small, rounded tubercle on
one or both sides (Figs 37d, 38a). There is also a conspicuous, rounded tubercle posterior to the postorbital angle
that can be almost as large as the first tooth, particularly in small individuals.
Palicus maculatus Edmondson, 1930, which is known only from one male collected from shallow-water
coralline algae in Hawaii (5.6 x 7.2 mm; BPBM 3337), is clearly a small specimen of P. tuberculatus.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
547
Edmondson (1930: 17) distinguished
P. maculatus from P. tuberculatus by having a
smaller fifth anterolateral tooth ("as long as the
fourth, but narrower" than P. tuberculatus ),
tubercles on the "upper and lower surfaces" less
prominent, and the fingers of the chclipeds "less
deflexed and the inner borders of the hands are
free from hair". These characters fall within the
range of morphological variation of the species.
One exception, however, is the absence of the
thick clump of plumose setae that characterizes
males of P. maculatus. It is possible that the
setae were absent in the holotype after almost
seventy years of preservation or that perhaps
this is a character absent in small males.
Furthermore, EDMONDSON'S figures of the
holotype were incorrect. The holotype has four
anterolateral teeth, the second is the largest and
the last two are located on an almost straight
line posterior to it. The figure, however, shows
what might be interpreted as two rounded teeth
anterior to a large, pointed tooth and two broad,
rounded teeth posterior to it.
Edmondson's living specimen was
described as having a "broad crimson band"
across the propodi of the walking legs and
chelipeds and a large crimson spot on a de¬
pression behind each eye (Edmondson, 1930:
15, pi. 1, fig. C). The dark band across each
second pair of walking legs that is shown in
Edmondson's photograph is similar to that
shown in the holotype of R. ampullatus
(Fig. 61a). A specimen photographed live in
Hawaii (Hoover, 1998: 287. unnumbered
figure) was cryptically colored pink and with
irregular white markings on the carapace and
walking legs. A female (9.5 x 13.1 mm;
MNHN-B 26784) that had been preserved in alcohol for 18 days showed numerous, small red dots on the dorsal
surfaces of the chelipeds, meri of walking legs, and borders of the carapace.
The name of P. tuberculatus was subsequently changed by EDMONDSON (1962: 8) to P. medipacificus
(Edmondson, 1962) (as Cymopolia medipacifica) since the name was already preoccupied by Cymopalia
tuberculata Faxon. 1895, a western Atlantic species. P. maculatus Edmondson, 1930, however, takes precedence
since the name was used in 1930 for a junior subjective synonym.
All pereopods of E. maculatus are particularly short when compared with other palicids. The relative length of
pereopods 2-5 among the specimens that were measured is as follows:
Fig. 38. — Exopalicus maculatus (Edmondson. 1930), 3 7.4 x
10.3 mm. Blow Hole, Oahu, Hawaiian Islands, 15 m (QM
W24745): a, carapace, dorsal view; b. suborbital border; c,
dactylus, right fourth pereopod, dorsal view; d. left male first
pleopod, lateral (outer side) view.
Source :
548
P. CASTRO
There are 5-6 spines each along the posterior border of the propodi of P2-P3, and 7-8 in P4. Each dactylus of
the walking legs is armed with a 1-2 teeth, each capped by a thick spine (Fig. 38c). The P5 are very short, the
ischia and mcri have small but conspicuous tubercles, and the posterior borders of the meri and dactyl i are each
armed with two and one spine respectively.
E. maculatus is known only from few locations at the extreme ends of the Indo-west Pacific region: the
southwestern Indian Ocean and the north-central Pacific Ocean. Most specimens have been collected at night.
It appears that individuals hide in coral and other hard substrates and emerge at night. Their nocturnal habits (see
Hoover, 1998: 287), small size, and cryptic coloration should explain their puzzling absence in collections from
other Indo-west Pacific locations.
Size. — Maximum size among specimens examined: 8.4 x 11.1 mm (male, LACM), 9.8 x 14.0 mm (female,
MNHN-B 26999).
Distribution. — Known from the Hawaiian Islands and Kure Island (EDMONDSON, 1925, 1930, 1933,
1946, 1962; some records as P. tuberculatus or Cymopolia medipacifica ) but its range now includes the Marshall
Islands, and La Reunion, southwestern Indian Ocean (Fig. 56). Depth: "shallow water" (EDMONDSON, 1925, 1930)
to 18 m.
Genus NEOPALICUS Moosa & Serene, 1981
Neopalicus Moosa & Serfcne, 1981: 41.
Type Species. — Cyrnpolia jukesii White, 1847 by original designation (MOOSA & Serene. 1981). Gender:
masculine.
DIAGNOSIS. — Carapace subquadrate; frontal border divided into two large lobes followed by distinct notch on
each side. Anterolateral borders each with two large truncate teeth (similar but shorter teeth may be posterior to
first two). Dorsal surface of carapace with large, low granules in horizontal rows, very low bosses. Eyes spherical;
each peduncle with one conspicuous, granular, median tubercle and two smaller, granular or soft tubercles.
Supraorbital borders each with 1-2 triangular lobes. Suborbital borders with two large, triangular, dentiform lobes.
Each basal antennal segment expanded into thick, flat process or two wing-like expansions. Epistome expanded
dorsoventrally, forming broad, semicircular, nearly flat surface; thin carina-like process along median portion
without protruding median teeth. Chelipeds equal or nearly equal in males and females, slender in females, heavier
in males. First three pairs of walking legs (P2-4) with flattened (not filiform) carpi, propodi, and dactyli. First pair
(P2) shorter than second and third pairs (P3-4); third pair about as long as second pair. Mcri of P2-4 oval in shape,
borders with thick tubercles or triangular teeth. Anterior borders of propodi of P3-4 with conspicuously wide,
convex enlargement; posterior border of dactyli entire. Fifth pair of pereopods (P5) reduced (0.7-0.8 CL), thick;
rows of thick spines along posterior border of propodi. Abdomen of mature males elongate (sides almost parallel to
each other), all segments free, one transverse ridge along each segment. Male first pleopods long and slender; basal
parts sinuous; each distal part biramous. Abdomen of mature females with all segments free, one transverse ridge
along each segment.
RfiDESCRiPTiON. — Carapace subquadrate, slightly broader than long, with large, low granules arranged in
horizontal bands. Anterolateral borders straight or slightly curved, each with two truncate teeth. Confluence of
branchial and mesogastric regions depressed and smooth. Conspicuous sulcus between hepatic and branchial
regions. Thoracic stemite 7 with reduced process (episternal process) at each outer edge and posterior to insertion of
fifth pair of pereopods (P5), visible dorsally as elongate or slightly rounded process below posterolateral border of
carapace. Posterior border of carapace smooth, without protruding tubercles or plumose setae.
Frontal border divided into two rounded lobes; distinct, wide notch separates frontal lobes on each side from
folded, slightly rounded border ending in sharp angle forming V-shaped fissure at supraorbital border. Supraorbital
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
549
borders each with 1-2 triangular lobes. Postorbital angles relatively short (never extending above retracted eyes),
pointed. Cornea of eyes spherical, almost as wide as base of peduncle; each peduncle with one conspicuous,
rounded, granular, median tubercle and two smaller, granular or soft tubercles. Orbits deep.
Suborbilal border with two triangular, dentiform lobes. Pterygostomial lobe at each anterolateral angle of
buccal frame projects ventrally, sinuous edge.
Antennules short, thick, and transversely folded beneath front; interantennular septum narrow. Each basal
antennal segment expanded into thick, dorsoventrally flattened, scale-like process or two thin, wing-like, anterior
expansions. Epistome expanded dorsoventrally, forming nearly flat surface between antennular fossae and anterior
border of endostome; no median teeth, only low, thin, carina-like process along median portion, flanked at same
level by two outer lobes that connect with each pterygostomial lobe. Meri of third maxillipeds smaller, narrower
than ischia.
Chelipeds equal or nearly equal in males and females, long, slender in females and shorter, thicker in males
(very thick propodus in largest males). Fingers of chelipeds with triangular teeth (broadly rounded teeth in very
large chelipeds of largest males), pollex becoming flatter with increasing size. First pair of walking legs (P2)
shorter, more slender than second and third pairs (P3-4); third pair slightly shorter or as long as second. First three
pairs of walking legs (P2-4) with flattened (not filiform) carpi, propodi, and dactyli. Meri slightly oval (not much
more longer than wide); anterior (dorsal) and posterior (ventral) borders with thick, unequal tubercles or triangular
teeth, rows of tubercles on dorsal and ventral surfaces. Elongate (but not filiform and conspicuously slender)
propodi and dactyli; posterior borders entire. Last two pairs of walking legs (P3-4) each with slightly broadened
coxae, having slightly flattened, thick-edged anterior and posterior borders. Anterior borders of propodi of P3-4
with conspicuously wide, convex enlargement. Fifth pair of pereopods (P5) reduced (0.7-0. 9 CL), thick; basis-
ischia and meri long and tuberculate; propodi with 2-3 rows of very thick spines along posterior border; dactyli
thick and slightly curved.
Abdomen of mature males elongate, with both sides almost parallel to each other, with all segments free,
segments 1-2 dorsoventrally compressed, one transverse ridge along each segment. Male first pleopods long and
slender; basal parts sinuous; each distal part biramous. Second male pleopods short, thin, slightly curved; distal
segment with blunt tip.
Abdomen of mature females broad, rounded, with all segments free, segments 1-2 dorsoventrally compressed,
one transverse ridge along each segment. Abdomen of immature females triangular (but broader than in males),
segments 1-2 dorsoventrally compressed, segments 3-6 fused, transverse ridge along each segment 1-2 plus two
along proximal portion of fused segments 3-6.
SPECIES included. — N. contractus (Rathbun, 1902) and N. jukesii (White, 1847).
Sexual Dimorphism. — As in other palicids, females are considerably larger in size than males. The
chelipeds are equal or nearly equal in both sexes. They are slender in females but heavier in males. There is no
evidence of differences in the length or shape of the walking legs or the fifth pair of pereopods.
DISCUSSION. — The description of Neopalicus by Moosa & Serene (1981) included the most important
characters of this small and relatively homogeneous genus. As in the other Indo-west Pacific genera, however, its
description has been expanded to include several new characters, even though no new species are described here.
Neopalicus is morphologically similar to Palicoides Moosa & Serene, 1981. The four species included in these
two genera are adapted to live in coarse sediments in relatively shallow water in or near coral reefs. Two and even
three of the four species in the two genera have been repeatedly collected together from the same stations in the
Coral Sea region. There are sharp differences between these four species and the species included in the other palicid
genera (see Table 1), most of which are inhabitants of finer sediments in deeper waters. Many of these differences
are seemingly the result of morphological and behavioral adaptations to their particular habitats. Unfortunately,
nothing is known about the habits of the four species of Neopalicus and Palicoides but it is assumed that they
bury in the sand (that is, they cover themselves with sand without forming burrows) at least during part of the day.
Species of Neopalicus and Palicoides share a subquadrate carapace, large and low tubercles that do not form
bosses, and thick fifth pereopods (Figs 39, 42). They even have a similar color pattern (Figs 61 c-e). Neopalicus
550
l>. CASTRO
species are characterized by three rounded tubercles on each eye peduncle, the largest of which is round, high and
granular, while there is a very conspicuous, larger, crescent-shaped (thin, rounded, and with pointed tip) process on
each peduncle in Palicoides species (see Table 1). Another character that is diagnostic in Neopalicits is the
expansion of the anterior borders of the propodi of the second and third pair of walking legs (P3-4) as a wide and
convex enlargement. Such an expansion is absent in the legs of Palicoides. Segments 2-3 of the antennae (Fig.
42a) are expanded and provided with long setae in Palicoides, whereas they are slender in Neopalicus as in most
other Indo-west Pacific palicids (Figs 40a-c).
No described species of palicids from outside the Indo-west Pacific region show the subquadrate, fiat carapace,
and thickened fifth pereopods characteristic of Neopalicus.
Key to the species of NEOPALICUS
1 . Carapace widest at level of first anterolateral tooth, lateral borders convergent posteriorly
(Fig. 39a). Basal antennal segment thick, scale-like (Fig. 40a) .
. N. contractus (Rathbun, 1902)
— Carapace widest at level of second anterolateral tooth, lateral borders not convergent
posteriorly (Fig. 39b). Basal antennal segment with two thin, nearly equal, wing-like
processes (Figs 40b-c) . N. jukesii (White, 1847)
Fig. 39. — a, Neopalicus contractus (Rathbun, 1902), $ 14.2 x 15.7 mm. Chesterfield Islands, CORAIL 2, stn DW 65,
62 m (MNHN-B 26788): dorsal view. — b , Neopalicus jukesii (White, 1847), $ 8.1 x 9.6 mm, east coast of
New Caledonia, Lagon, stn 713, 34-35 m (MNHN-B 26996): dorsal view.
Neopalicus contractus (Rathbun, 1902)
Figs 39a, 40a, 41a-b, 49
Palictts contractus Rathbun, 1902: 126, figs 7-8. — SERfeNE, 1968: 96.
Palicus jukesi - Laurie, 1906: 430, pi. 1, fig. 12 [non Neopalicus jukesii (White, 1847)].
Cymopolia robusta Ward, 1942a: 46, pi. 4, figs 3-4; 1942b: 53. — Michel, 1964: 33; 1974: 134. — GutNOT, 1967: 280.
Palicus robustus - SerTne, 1968: 96.
Neopalicus contractus - Moosa & Serene, 1981: 44, pi. 2, fig. D.
Palicus jukesi - Garth et al., 1987: 259 [non Neopalicus jukesii (While. 1847)).
non Palicus contractus - Zarenkov, 1968: figs 3,4, E [/’. contractus in caption and English summary, P. jukesii in text;
fig. 3 A = P aliculus kyusyuensis (Yokoya, 1933), fig. 3E = N. jukesii (White, 1847)1.
Source : MNHN, Paris
INDO-NVEST PACIFIC SPECIES OF PALICID CRABS
551
MATERIAL EXAMINED. — Seychelles. ReveS 2: stn 17, 05°44.8'S, 56°39.1'E, 55 m. coralline algae, 5.09.1980:
I <3, 1 9 (MNHN-B 27149). — Stn 18, 05°43.2’S, 56°34.0'E, 50 m, shelly sand, 5.09.1980: I 9 (MNHN -13 27147). —
Stn 38, 05°03.5'S, 56°50.5'E, 44 m, coralline algae and dead coral, 13.09.1980: I 9 (MNHN-B 27145). — Stn 52.
03°52.8'S, 55°25.3'E, 60 m, sand, 16.09.1980: I 9 (MNHN-B 27144).
Maldives. Miladummadulu Atoll. Nallandu Island, 44 m, A. Agassi/, coll.. 18.01.1902: 1 3 lectotype of Palicus
contractus Rathbun, 1902 4.9 x 7.5 mm (MCZ 6699).
South Nilandu Atoll. J.S. Gardiner coll.: I 8 , I 9 (UMZ).
Sri Lanka. Gulf of Mannar. W.A. Herdman coll.: 1 3 (BMNH 1934.1. 16.164).
Philippine Islands. MUSORSTOM 3: sin DR 137, I2°03'N, 122°06'E, 56 m, 6.06.1985: 3 9 (MNHN-B 27143).
Chesterfield Islands - Chesterfield-Bellona Plateau. Chalcal 1: stn D 43, 20°4I.50'S, 158°38.4()'E,
78 m, Halimeda sand, 07.1984: 1 3 (MNHN-B 27151). — Stn D 45, 20°48.93'S, I58°30.21'E, 50 m, 07.1984: I 9
(MNHN-B 27154). — Stn D 52, 21°13.40'S, 158°49.20'E, 69 m, 07.1984: 1 <J (MNHN-B 27155). — Sin D 53.
21°19.50'S, 158°05.30'E, 60 m, 07.1984: 3 S (MNHN-B 27148). — Stn D 55, 21°23.90'S, I58°59.60'E, 55 m,
Halimeda and foraminiferans, 07.1984: 1 3 (MNHN-B 27150). — Stn D 56, 21°24.40'S, 159°08.80'E, 60 m. Halimeda
and foraminiferans, 07.1988: I 3 10.8 x 12.0 mm (MNHN-B 26789), 1 9 (MNHN-B 27139).
Corail 2: stn DW 2, 20°55.90'S, 161°40.70'E, 62 m, boulders and Halimeda, 28.07.1988, I 9 (MNHN-B 27156). —
Stn DW 18, 20°44.08'S, 160°59.92'E, 69 m, 21.07.1988: I 9 (MNHN-B 27142). — Stn DW 61, 19°14.96'S,
158°53.60'E, 54 m, Halimeda sand, 24.08.1988: I 9 (MNHN-B 27152). — Sin DW 65, 19°15.00'S, 158°40.64’E, 62 m,
Halimeda sand, 24.08.1988: 1 9 14.2 x 15.7 mm (MNHN-B 26788). — Sin DW 80, 19°11.98'S. 158°47.0I'E. 66 m.
Halimeda sand, 25.08.1988: 1 9 (MNHN- B 27141). — Stn DW 94, 19°06.00'S, I58°50.00'E, 36-53 m, boulders and
Halimeda, 27.08.1988: 1 9 (MNHN-B 27146).
Bellona Reefs. Stn 2 DE, 21°21.5'S, 158°48.0'E, 61-64 m, 19.10.1985: 1 9 (MNHN-B 27153). — Stn 7 DE,
21°21.0'S, 158°52.7'E, 56-58 m, 19.10.1985: 2 3 (MNHN-B 27140).
New Caledonia. Lagon: stn 392, 22°48.2'S, 167°02.3'E. 80 m, shell debris, foraminiferans, 22.01.1985: I 9
17.3 x 18.6 mm (MNHN-B 26786).
SMIB 5: stn DW 100, 23°22.9'S, 168°05.2'E, 80-120 m, sand, 14.09.1989: I <3 12.9 x 14.0 mm (MNHN-B 26787).
Marshall Islands. Enewetak Atoll. Stn 423, J.W. Knudsen coll., 14.08.1967: 1 3 (LACM). — Stn 429.
15.08.1967: 1 9 (LACM). — Stn 187: 1 9 (LACM).
TYPES. — Lectotype of Palicus contractus Rathbun, 1902: 1 6 4.9 x 7.5 mm, 1 S paralectotype, Maldives
(MCZ 6699).
Holotype of Cymopolia robusta Ward, 1942: 1 9 CW 16 mm (Ward, 1942), Mauritius. Deposit unknown:
Mauritius Institute, Port Louis, Mauritius ?
Type Locality. — Nallandu Island, Miladummadulu Atoll, Maldives, 44 m.
Diagnosis. — Carapace (Fig. 39a; Rathbun, 1902, fig. 7; Moosa & Serene, 1981, pi. 2, fig. D)
subquadrate, with large, low granules; lateral borders wider anteriorly than posteriorly so that widest extension of
carapace at level of first and larger of two anterolateral teeth. Supraorbital borders each with one triangular inner
lobe, one short outer lobe. Suborbital borders (Fig. 40a) each with two triangular, tooth-like lobes. Each basal
antennal segment (Fig. 40a) with thick, dorsoventrally flattened process along outer border, much shorter, elongate
process along inner border. Dorsal borders of cheliped propodi with rounded, microscopic tubercles, much more
pronounced in thick chelipeds of large males; outer borders with hook-like setae. Dorsal and ventral sides and upper
and lower borders of meri of second and third pairs of walking legs (P3-4; Fig. 39a) with conspicuous tubercles;
anterior borders of carpi of P3-4 with similar tubercles; anterior borders of propodi with conspicuously wide,
convex enlargement. Abdomen of mature males narrow, with all segments distinct; one transverse ridge along each
segment. Male first pleopods (Figs 41a-b) with sinuous basal parts; each distal part biramous, with one flattened,
anterior tip with four pointed teeth, and long, curved extension with teeth along dorsal border extending
immediately above toothed base. Abdomen of mature females with all segments distinct, one transverse ridge along
each segment (less pronounced and curved in median portion in segment 6).
DISCUSSION. — This is the first lime the male pleopod of N. contractus is described and illustrated. The figure of
the male pleopod given by Zarenkov (1968, fig. 3E) is actually that of N. jtikesii , although it is correctly
identified in the text (ZARENKOV, 1968: 765). The ventral view of the anterolateral portion of the carapace that is
identified as N. jtikesii in the text and as N. contractus in the caption (ZARENKOV, 1968, fig. 3 A, 765) docs not
552
R. CASTRO
belong to Neopalicus but to Paliculus kyusyuensis (Yokoya, 1933) instead. To add to the confusion, N. contractus
and not N. jukesii , is listed in the English summary (ZARENKOV, 1968: 766)!
Re. 4°. — a, Neopalicus contractus (Rathbun, 1902), 2 14.2 x 15.7 mm, Chesterfield Islands. Corail 2. stn DW 65.
62m (MNHN-B 26788): ventral view of anterior portion of carapace. — b-c, Neopalicus jukesii (White. 1 847)-
2 8.1 x 9.6 mm, east coast of New Caledonia, Lagon, stn 713, 34-35 m (MNHN-B 26996): b. ventral view of
anterior portion of carapace; c. anterior view of carapace.
Source : MNHN , Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
553
Although the chelipeds are long, slender, and equal in immature males, ihey become thicker with increasing
size. In the largest male examined in this study, a specimen from New Caledonia (12.9 x 14.0 mm; MNHN-B
26787), the right cheliped was greatly enlarged, much more than the chelipeds in the second largest male (10.8 x
12.0 mm; MNHN-B 26789). A large male from the Maldives (UMZ) that was not measured, also had an enlarged
right cheliped. The fingers of the enlarged cheliped were short and very thick. The propodus and fingers of the left
cheliped were missing but the segments that remain were only slightly smaller than those of the right cheliped.
The male first pleopods and other morphological characters of the largest specimens agreed with those of smaller
males.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
lhe fifth pair of pereopods (P5; Fig. 39a) is typical of Neopalicus, with thick, tuberculate segments (except
dactylus), very thick spines along the posterior border of the propodus, and a slightly curved dactylus.
Most specimens of N. contractus can be easily distinguished from N. jukesii, a close species (see Table 3).
Nevertheless, both species can be confused when there is limited material or only small specimens, as
demonstrated by repeated mistakes in the literature. The shape of the carapace is a relatively stable character but the
shape of the anterolateral teeth tends to overlap, particularly in small specimens and in large females.
The shape of each basal antennal segment is a much more reliable character. It has a conspicuous, thick, scalc-
like process as well as a smaller, inner process (Fig. 40a) in contrast to the two thin, nearly equal, wing-like
processes (Figs 40b-c) in N. jukesii. In their key to the species of Neopalicus. Moosa & Serene (1981: 41)
characterized the basal antennal segment of N. jukesii as having a "subacutely carinate" anterolateral angle in
contrast to a "bilobed process" on the anterolateral angle of N. contractus. It seems that both characters were
reversed since it is difficult to characterize the segment in N. contractus as "bilobed" (unless the inner prominence
was considered a lobe) and one of the two wing-like expansions in N. jukesii was interpreted as a carina.
Moosa & Serene (1981: 41) also included in the key they used to separate between the two species a
difference in the relative length of the dactylus of P4 in relation to the length of the propodus. the dactylus being
less than half the propodus length in N. contractus but more than half in N. jukesii. This was found not to be the
case in the many specimens that were examined in this study. The dactylus length to propodus length ratio was
found to vary between 1:1.8 to 1:1.9 in both species.
Cymopolia robusta Ward, 1942 can be identified as a subjective junior synonym of N. contractus from the
photograph that accompanied its description (Ward, 1942a). The holotype, supposedly at the Mauritius Institute,
was unavailable for examination. As previously mentioned by MOOSA & SERENE (1981: 44), the characters used
by Ward (1942a) to separate his species from N. jukesii also apply when N. contractus is compared with
N. jukesii.
Rathbun (1902) based her description of Palicus contractus on two specimens without selecting a holotype.
One of the two syntypes (<3 4.9 x 7.5 mm. MCZ 6699) is hereby designated as the lectotype and the second male
specimen as a paralectotype.
Size. — Maximum size among specimens examined: 17.3 x 18.6 mm (female, MNHN-B 26786); 12.9 x
14.0 mm (male, MNHN-B 26787).
Distribution. — Mauritius (Ward, 1942a, 1942b; as Cymopolia robusta), Maldives (Rathbun, 1902). Sri
Lanka (Laurie, 1906, as Palicus jukesi), and Ceram and Kai islands, Indonesia (Moosa & Serene. 1981). It is
here recorded for the first time from the Seychelles, Philippine Islands, the Coral Sea (Chesterfield Islands. Bellona
Reefs, and New Caledonia), and the Marshall Islands (Fig. 49). It has been repeatedly collected from coarse sand
rich in coral, coralline algae, and shell rubble. In the Coral Sea, it has been collected together with N. contractus
Source :
554
P. CASTRO
and the two species of Palicoides. Depth: known reliably between 44 and 80 m; also collected in trawls between
36-53 m and 80-120 in.
Neopalicus jukesii (White, 1847)
Figs. 39b, 40b-c, 41c, 49, 61c
Cymopolia jukesii White, 1847a: 54 Inomen nudum).
Cymopolia jukesii White, 1847b: 338, pi. 2, fig. 1; 1861: pi. 2. — Miers, 1874: 3, pi. 3, figs 4, 4a-c; 1886: 335. —
HASWELL, 1882: 138. — HENDERSON, 1893: 405.
Cymopolia carinipes Paulson, 1875: 76, pi. 9, figs 4, 4a; 1961: 79, pi. 9, figs 4, 4a.
Palicus jukesii - Bouvier, 1897: 65. — ALCOCK, 1900: 451. — Carman. 1900: 29, pi. 1, figs 9-13. — De Man, 1902:
545. — Ratiibun, 1902: 126; 1911: 240, pi. 19, fig. 9. — Laurie, 1915: 416. — Nobili, 1906a: 325. — Zarenkov,
1968: 765, fig. 3E \P. contractus in caption and English summary]; 1971; 173.
Palicus jukesi - Borradaile, 1903: 433. — SerLne, 1968: 96,
Cymopolia jukesi - GuiNOT, 1967: 280.
Palicus carinipes - HoLTHUlS, 1977: 181, fig. 2.
Neopalicus jukesi - MOOSA & SERENE, 1981: 42, fig. 7, pi. 2, fig. C.
non Palicus jukesii - Laurie, 1906: 430, pi. 1, fig. 12 |= Neopalicus contractus (Rathbun. 1902)].
non Palicus jukesii - Zarenkov, 1968: fig. iA [P. contractus in caplion = Paliculus kyusyuensis (Yokoya, 1933)].
MATERIAL EXAMINED. — Gulf of Aden. "Meteor": stn Me5-236 Ku, 1 2°21 ,2'N- 1 2°19.0'N. 43°27. rE-43°27.8'E,
35-45 m. 6.03.1987: 5 <3, 19 9, 21 juv. (SMF 24709), 3 <3 , 4 $ (SMF 24710).
Comoro Islands. Mayotte. 50 m, coarse sand, A. Crosnier coll., 09.1958: 1 <3 (MNHN-B 27216).
Madagascar. Northwest coast : Nosy Iranja. Sand, R. Plante coll., 26.08.1965: I 2 (MNHN-B 27217). —
Nosy Be. A. Crosnier coll.: 1 <3 (MNHN-B 27218).
Saya de Malha Bank. "Sealark" Expedition: 101 m, , J.S. Gardiner coll., 6.09.1905: 1 2 (UMZ).
Maldives. "Sealark" EXPEDITION: South Nilandu Atoll. J.S. Gardiner coll.: 3 <3 , 5 2 (UMZ), 2 2 (UMZ), I 2
(UMZ). — Kolumadulu Atoll, J.S. Gardiner coll.: I 2 (UMZ). — Haddumati Atoll. J.S. Gardiner coll.: I 2 (UMZ), I 2
(UMZ), I 2 (UMZ), 1 9 (UMZ), 1 2 (UMZ). — Stivadiva Atoll. J.S. Gardiner coll.: I (3,6 2 (UMZ), 2 <3,12 (UMZ),
2 2 (UMZ), 1 2 (UMZ).
Sri Lanka. Trincomalee, 1 1 m, K. Fristedt coll., 03. 1889: 1 6 , 1 2 (MZUF C.2481).
Malaysia. Tioman Island: Tekek Bay, 26.06.1985: 1 2 (ZRC 1989.3765).
Singapore. Senang Island: 1 2 (ZRC 1965.8.2.235).
Japan. Ogasawara Islands. Muko-jima Island, 27°48.52'N, 142°02.62'E, 98-99 m, T. Komai coll., 15.10.1997: 1 9
(C.BM-ZC 2004).
Philippine Islands. Musorstom 3: stn CP 142, 11°47'S, 123°02'E, 26-27 m. 6.06.1985: I 2 (MNHN-B 27215).
Indonesia. Celebes Sea. "Challenger": stn 212, 06°54'N, 122°I8'E, 18 m: I 9 (BMNH 1884.31).
Moluccas, Ternate. W. KOkenthal coll., 1894: 3 2 (SMF 4713).
Kai Islands. Danish Expedition to Kai Islands: stn 26, 05°38'S, 132°55'E, sand, 90 m, 14.04.1922: I <3 (ZMUC).
Papua New Guinea. Bismark Archipelago. New Britain Island, Ralum, 24-30 m, S. Dahl coll., 24.1 1.1897: 1 2
(ZMB 18139).
Australia. Great Barrier Beef. Sir Charles Hardy Island, 20 m, coarse sand, J.B. Jukes coll., 08.1844: 1 2 holotfpe
of Cymopolia jukesii White, 1847 9.0 x 10.9 mm (BMNH 46.80).
Torres Strait. Murray Island, A.C. H.addon coll., 1888: I <3 (BMNH 1954.9.14.91).
Chesterfield Islands - Chesterfield-Bellona Plateau. Chalcal I: sin D 52, 21°13.40'S, 158°49.20'E,
69 m, 07.1984, 1 <3 (MNHN-B 27219). — Stn D 55, 2I°23.90'S, 158°59.60'E, 55 m, Halimeda and foraminiferans,
07.1984: 1 2 (MNHN-B 27220).
Stn 4 DE, 21° 19.0'S, 158°48.0'E, 66 m, 19.10.1985: I 2 (MNHN-B 27179).
Corail 2: stn DW 61, 19°14.96'S, 158°53.60’E, 54 m, Halimeda sand, 24.08.1988: I <3 , I 2 (MNHN-B 27221). —
Sin DW 73, 19°12.H'S, 158°22.57'E, 41 m, coarse sand, 25.08.1988: 1 2 (MNHN-B 27222). — Stn DW 76,
19°12.25'S, 1 58°32.90'E, 41 m, coarse sand and Halimeda, 25.08.1988: 1 2 (MNHN-B 27226). — Stn DW 87,
I9°06.I4'S, 158°59.94’E, 31 m, coarse sand and Halimeda, 26.08.1988, 1 2 (MNHN-B 27223). — Stn CP 98,
I9°04.32'S, 158°31.66'E, 44-48 m, corals, 26.08.1988: 1 2 (MNHN-B 27224), — Stn DW 135, 19°3 1.37'S,
1 58° 19.1 4'E, 46 m, coarse sand, 30.08.1988: 1 <3 , 1 2 (MNHN-B 27182). — Stn DW 137, I9°34.00'S, 158°14.60'E,
32 m, coarse sand, 30.08.1988: 1 2 (MNHN-B 27225).
New Caledonia. Lacion: sin 517, 19°08.9'S, 163°35.0’E, 42 m, Halimeda sand, 5.03.1985: 1 2 (MNHN-B 27210).
— Stn 528, 19°3 1 ,2'S, I63°30.0'E, 47 m, fine sand, 6.03.1985: I <3 (MNHN-B 27195). — Stn 536, I9°08.8'S,
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
555
1 63°22.6'E, 61 m, shelly sand. 6.03.1985: 1 £ (MNHN-B 27200). — Sin 541. I9°06.0'S. 163°13 3'E 48-43 m
foraminiferan sand. 6.03.1985: 1 £ (MNHN-B 27211). — Stn 560, 22°43.2’S, 166°56.8’E, 48 m shelly sand"
16.07.1985: 1 <3 (MNHN-B 27181). — Stn 598. 22°19.1S, 167°06.7'E. 73-75 m, shelly sand. 5.09.1986: 1 2 (MNH.N-
B 27227). — Stn 620, 22°02.4'S. 166°56.2'E. 50-52 m, Halimeda , 6.08.1986: 1 3 (MNHN-B 27228). — Stn 626.
21°57.9’S, 166°52.5'E, 47-48 m, foraminiferan sand. 6.08.1986: 3 <3 . 1 £ (MNHN-B 27229). — Stn 627. 2l°58.9"s.
I66°50.7’E, 45-47 m, Halimeda sand, 6.08.1986: 1 <3 (MNHN-B 27205). — Stn 632. 21°57'3S, 166°49 6'E 44-45 m
foraminiferan sand. 6.08.1986: 1 6 (MNHN-B 27201). - Stn 633, 21°55.6'S. I66°48.2'E. 50 m. foraminiferan sand and
Halimeda. 6.08.1986: 1 c3 (MNHN-B 27230). — Stn 641, 21°53.0'S. 166°43.0'E, 50-52 m. Halimeda. 7.08.1986: 1 <5
(MNHN-B 27231). — Stn 682, 21°33.7'S, 166°18.6'E, 36-37 m, foraminiferan sand. 9.08.1986: 1 2 (MNHN-B 27190).
— Stn 697, 21°27.6'S, 166°10.0'E, 35-36 m, foraminiferan sand, 10.08.1986: 1 <J , 2 £ (MNHN-B 27196). 1 <3 (MNHN-
B 27191). — Stn 702, 21°26.7'S, 166°08.2'E, 37 m, foraminiferan sand, 10.08.1986: 1 <3 6.0 x 7.1 mm (MNHN-B
26814). — Stn 708, 21°23.6'S, 166°05.2'E, 34-35 m, foraminiferan sand. 10.08.1986: 1 $ . 4 2 (MNHN-B 27232). _
Stn 709, 21°22.2'S, 166°03.5'E, 39-40 m, foraminiferan sand, 10.08.1986: 3 3 . 4 £ (MNHN-B 27233). Stn 713,
21°22.6'S. 166°00.7'E, 34-35 m, coarse sand, 11.08.1986: 1 <3.12 8.1 x 9.6 mm (MNHN-B 26996). — Stn 714,'
2I°2|.0'S, 166°01 ,8’E, 37-38 m, coarse sand, 11.08.1986: 1 c3 7.7 x 9.2 mm (MNHN-B 26790). — Stn 724. 21°19.7’S.
165°57.8'E, 36-38 m, foraminiferan sand and Halimeda. 12.08.1986: 1 £ (MNHN-B 27212). — Stn 725, 2 1 0 1 8.6'S.
165°56.0'E, 41-43 m, coarse sand, 12.09.1986: 1 3 (MNHN-B 27206). — Stn 730, 21 °1 7.2'S. I65°54 5'E 40-41 m"
coarse sand, 12.08.1986: 1 <3 , 1 £ (MNHN-B 27192). — Stn 747, 21°14.7'S, 165°50.9'E, 31-34 m. foraminiferan sand
and Halimeda. 6.01.1987: 2 5 (MNHN-B 27234). — Stn 753, 21°14.9'S, 165°48.4'E, 53 m, shell debris, 7.01.1987:
1 £ (MNHN-B 27213). — Stn 754, 2 1 0 1 3. I S, 165°49.2'E, 36 m, foraminiferan sand and Halimeda. 7.01.1987: 1 3.
2 £ (MNHN-B 27186). — Stn 761, 21°13.1'S, 165°44.3'E. 41-44 m, fine sand, 7.01.1987: I 3 (MNHN-B 27202). —
Stn 772. 21 °07.7'S. 165°40.5'E, 30 m. coarse sand, 8.01.1987: 1 £ (MNHN-B 27203). — Stn 780. 2I°06.0'S.
165°39.2'E. 33 m. coarse sand, 8.01.1987: 2 £ (MNHN-B 27193). — Stn 781. 21°04.6'S. 165°37.8’E, 36 m. coarse
sand, 8.01.1987: 1 <3 . 1 £ (MNHN-B 27235). — Stn 788, 21°01.6'S, 165°34.7'E. 33 m, coarse sand. 9.01.1997- 1 £
(MNHN-B 27236). — Stn 794. 21°03.2'S. 165°30.9'E, 51 m, mud, 9.01.1987: 1 3.4 £ (MNHN-B 27207). — Stn. 795.
21 °01 ,6'S, 165°32.0'E, 31 m, coarse sand. 9.01.1987: 1 3 (MNHN-B 27184). — Stn 796. 20°59.9'S. 1 65°33. 1 E. 38 m
coarse sand. 9.01.1987: 1 £ (MNHN-B 27185). — Stn 808, 20°57.4'S, 165°29.6’E. 30 m, coarse sand. 10.01.1987: 1 £
(MNHN-B 27237). — Stn 828. 2I°50.1'S, 165°19.5'E, 28 m, coarse sand, 10.01.1987: 1 <3 (MNHN-B 27188). —
Stn 856, 20°36.9'S, 165°11.4'E, 30 m, coarse sand. 12.01.1987: 1 3 (MNHN-B 27204). — Sin 863. 20°39.4'S,
165°06.5'E, 28 m, coarse sand, 13.01.1987: 1 <3 (MNHN-B 27241). — Stn 865. 20°38.7'S, I65°04.4'E. 24 m. shelly
sand, 13.01.1987: 2 <3 , 1 £ (MNHN-B 27214); 1 £ (MNHN-B 27240). — Stn 866. 20°37.5'S. 165°02.7'E. 26 m,
foraminiferan sand. 13.01.1987: 1 <3 (MNHN-B 27197). — Stn 892, 20°18.3'S, 164°32.1'E, 26 m, coarse sand and
Halimeda. 14.01.1987: 1 £ (MNHN-B 27238). — Stn 898. 21°13.6'S. 164027.1'E, 22 m. gray sand, 14.01.1987: I 3
(MNHN-B 27239). — Stn DW 1026. 20°04.6’S, 163°47.6'E, 29 m, foraminiferan sand and Halimeda. 3.04.1988: 1 £
(MNHN-B 27194). — Stn DW 1075, 19°52.0’S, 163°58.4'E. 28 m, white sand and Halimeda. 23.10.1989: 1 £ (MNHN-B
27242). — Stn DW 1094, 19°54.4'S. 163°41.2'E, 26 m. foraminiferan sand and Halimeda. 20.10.1989: 1 £ (MNHN-B
27243). — Sin DW 1097, 19°51.7'S, 163°42.5'E, 34 m, foraminiferan sand, 24.10.1989: 1 £ (MNHN-B 27198). —
Stn DW 1110, 19°43.6'S, 163°41.8'E. 31 m, fine sand and foraminiferans. 25.10.1989: 1 £ (MNHN-B 27208) —
Stn DW 1129, 19°29.2'S, 163°48.8'E. 40 m, shelly sand, 26.10.1989: 1 <3 (MNHN-B 27244). — Stn DW 1134.
19°31.3'S, 163°34.6'E, 40 m, coarse sand and Halimeda. 26.10.1989: 1 <3 , 1 £ (MNHN-B 27187). — Stn DW 1155,
19°09.3'S, 163°15.9'E. 48 m, coarse sand, 30.10.1989: 1 <3 (MNHN-B 27199). — Stn DW 1163, 19°1 1.3'S, 163°2I.9'E.
48 m, fine sand, 30.10.1989: 2 3 (MNHN-B 27209). — Stn DW 1181, 19°23.9'S, 163°14.7'E. 45 m. coarse sand
31.10.1989: 1 3 , 1 £ (MNHN-B 27180). — Stn DW 1182, 19°27.3'S, 163°16.2’E. 48 m, coarse sand. 31.10.1989: 6 £
(MNHN-B 27183). — Stn DW 1192, 19°35.3'S, 163°24.6'E, 48 m, coarse sand, 1.11.1989: 2 <3 , 1 £ (MNHN-B 27189).
— Stn DW 1197, 19°35.6'S, 163022.1'E, 41 m. foraminiferans, 1.11.1989: 1 3 (MNHN-B 27245). — Stn DW 1205,
19°41.6'S, 163°25.6'E, 38 m, coarse sand. 2.11.1989: 1 £ (MNHN-B 27246). — Stn DW 1217. 19°51.6'S. 163°35.6'E.
30 m, coral debris, 3.1 1.1989: 1 <3 (MNHN-B 27247).
Types. — Holotype of Cymopolia jukesii White, 1847: 1 9 9.0 x 10.9 mm. Great Barrier Reef, Australia
(BMNH 46.80).
Holotype of Cymopolia carinipes Paulson, 1875: 1 c3 6 x 7 mm (Paulson, 1875), Red Sea. Material most
probably lost.
TYPE Locality. — Sir Charles Hardy Island, Great Barrier Reef, Australia, coarse sand. 20 m.
Diagnosis. — Carapace (Figs 39b. 61c; Calman, 1900, pi. 1, fig. 9; Rathbun, 191 1, pi. 19. fig. 9; Moosa
& SerEne, 1981, pi. 2, fig. C) subquadrate, with large, low granules; anterolateral borders slightly curved, with
widest extension at level of second and smallest of first two. truncate, anterolateral teeth; 2-3 shorter, smaller teeth
may be found posterior to first two (Holthuis, 1977, fig. 2a, as Palicus carinipes). Supraorbital borders each with
Source : MNHN , Paris
556
P. CASTRO
Fig. 41. — a-b, Neopalicus contractus (Rathbun, 1902): 8
10.8 x 12.0 mm, Chesterfield-Bellona Plateau, Chalcal 1,
stn D 56, 50 m (MNHN-B 26789): a, left male first
pleopod. ventral view; b, left male first pleopod, distal
part, lateral (outer side) view. — c, Neopalicus jukesii
(White, 1847), 8 7.7 x 9.2 mm, east coast of New
Caledonia, Lagon, stn 714, 37-38 m (MNHN-B 26790):
left male first pleopod, ventral view.
one short, triangular lobe. Suborbital borders
(Fig. 40b; Calman, 1900, pi. 1, fig. 10;
Holthuis, 1977, fig. 2b, as Palicus carinipes)
each with two triangular, tooth-like lobes. Each
basal antennal segment (Figs 40b-c) with two
thin, nearly equal, wing-like ventral processes
along ventral side (V-shaped in dorsal or ventral
view). Dorsal borders of cheliped propodi with
irregular, microscopic tubercles, increasing in
size to form rows of conspicuous tubercles in
large males; outer borders with hook-like setae;
dense clusters of plumose setae on inner border
of cheliped propodi in males (Holthuis, 1977.
fig. 2c, as Palicus carinipes). Dorsal and ventral
sides and upper and lower borders of meri of
second and third pairs of walking legs (P3-4)
with rounded tubercles (Calman, 1900, pi. 1,
fig. 12); anterior borders of carpi of P3-4 with
similar tubercles; anterior borders of propodi
with wide, convex enlargement. Abdomen of
mature males narrow, with all segments
distinct; one transverse ridge along each
segment (slightly more salient and wider in
segment 6). Telson of males and females with
low, elongate, and rounded tubercles (Calman,
1900, pi 1, fig. 1 1). Male first pleopods (Fig.
41c; Moosa & Serene, 1981, fig. 7) with
sinuous basal parts; each distal part with two
pointed tips (close together in most specimens
but separate in larger specimens; Zarenkov,
1968, fig. 3E; Holthuis, 1977, fig. 2c, as
Palicus carinipes), dorsal with short teeth and
ventral with small, blunt teeth. Abdomen of
mature females with all segments distinct, one
transverse ridge along each segment (less
pronounced and curved in median portion in
segment 6).
Color : A specimen from New Caledonia
photographed live (Fig. 61c) had a broad orange-red, vertical band through the middle of the carapace, a black patch
on each branchial region, and broad, orange-red bands across the walking legs.
Discussion. N. jukesii may be easily confused with N. contractus. Differences between the two species are
summarized in Table 3. Although the lateral shape of the carapace (particularly by being broader at the level of the
second anterolateral tooth in N. jukesii but at the level of the first anterolateral tooth in N. contractus) can be used
as a diagnostic feature in most specimens, variations have been observed. In some specimens the first anterolateral
tooth may be more salient than the second (although often on only one side of the carapace) or the broadest width
of the carapace is at the level of the teeth posterior to the second anterolateral tooth (see Holthuis, 1977. fig. 2a.
as Palicus carinipes). The shape of each basal antennal segment (Figs 40b-c), V-shaped in dorsal or ventral view in
contrast to a conspicuous, thick, dorsoventrally flattened process in N. contractus (Fig. 40a) is a useful diagnostic
character.
Source : MNHN, Paris
1NDO-WEST PACIFIC SPECIES OF PALICID CRABS
557
Table 3. — Morphological differences between Neopalicus contractus (Rathbun. 1902) and N. jukesii (White, 1847).
The relative length of pcreopods 2-5 among the specimens that were measured is as follows:
The fifth pair of pereopods (P5) (Fig. 39b; HOLTHUIS, 1977, fig. 2d, as Palicus carinipes ) has. like in
N. contractus, thick, tuberculate (except dactylus) segments, very thick spines along the posterior border of the
propodus, and a slightly curved dactylus.
HOLTHUIS (1977) identified his Red Sea material as Palicus carinipes (Paulson, 1875), which is clearly
a subjective junior synonym of N. jukesii. His conclusion was based on several differences, most important of
which was the presence in his specimens of a curved ridge of tubercles (subhepatic crest) on the pterygostomial
(subhepatic) region posterior to the suborbital margin (HOLTHUIS, 1977, fig. 2b). This ridge appears in the figure
that accompanies the description of P. carinipes (Paulson, 1875. pi. 9, fig. 4a; 1961. pi. 9, fig. 4a). while there
is only a simple tubercle in the figure of N. jukesii given by Calman (1900, pi. 1. fig. 10) for N. jukesii.
Calman's specimen (BMNH 1954.9.14.91) actually had a curved ridge on its pterygostomial region, even if it
was stated that "the tubercle on the underside of the hepatic region ... represented (in P. carinipes] by a curved
transverse ridge" was absent in the specimen (Calman, 1900: 31). A curved ridge of tubercles is present in both
Source : MNHN. Paris
558
P. CASTRO
N. jukesii (Fig. 40b) and N. contractus (Fig. 40a), a fact already mentioned by Rathbun (1911: 240). It becomes
more conspicuous with growth, however. A specimen (BMNH 1934.1.16.164) incorrectly identified by Laurie
(1906: 430, 431; pi. 1, fig. 12) as Palicus jukesii is actually N. contractus. LAURIE mentioned the characteristic
curved ridge of tubercles but, to complicate matters, his figure shows only a short row.
HOLTHUIS (1977: 182, fig. 2b) mentioned that another difference between his Red Sea specimens and
CALMAN's figure of N. jukesii (CALMAN, 1900, pi. 1, fig. 10) was an outer suborbital tooth described as
"narrowly triangular" and a broader inner lobe in contrast to the opposite situation shown in Calman's figure.
While most of the specimens examined in this study agreed with Calman's specimen, some did show the
arrangement described by HOLTHUIS.
HOLTHUIS unfortunately referred to the fifth pereopod (P5) of his specimens as "subchelate" on account of the
slightly curved dactylus (HOLTHUIS, 1977: 183). The same characteristic was attributed to N. jukesii based on the
figure of the type by MlERS (1874, pi. 3, fig. 4) but not to the specimen of the same species as illustrated by
Calman (1900, pi. 1, fig. 9). The dactylus, although curved, cannot articulate enough lo make contact with the
posterior border of the propodus and as such cannot be regarded as truly subchelate. As foreseen by HOLTHUIS
himself, this and the other differences that were given have proven lo be the result of individual differences and
incorrect information.
The male first pleopods of N. jukesii were described as having a "spiraled shaft" by Moosa & Serene (1981:
44). The basal part of the pleopods should be described as sinuous, not spiraled (or helicoidal) as in the case of
species of Parapalicus.
A female specimen from the Ogasawara Islands, Japan (8.0 x 8.8 mm; CBM-ZC 2004) is the first record of
Neopalicus from Japan. It has slightly higher tubercles on the eye peduncles, two conspicuous rounded tubercles
on the carpus of its only cheliped, unusually pointed postorbital angles similar to those of N. contractus , and the
upper lobe of the meri of the third maxillipeds is larger and rounder that of N. jukesii. All other characters agree
with N. jukesii. The examination of the male first pleopods would confirm if it is N. jukesii or if it belongs to a
new species.
Size. — Maximum size among specimens examined: 13.8 x 15.1 mm (female, UMZ); 7.7 x 9.2 mm (male,
MNHN-B 26790).
DISTRIBUTION — N. jukesii is known from many records from a very wide geographic distribulion (Fig. 49).
It is known from the Red Sea (PAULSON, 1875, as Cymopolia carinipes; NOBILL 1906a; Z.ARENKOV, 1971;
HOLTHUIS. 1977, as Palicus carinipes ); Amirante Islands and Saya de Malha Bank (Rathbun. 191 1); Maldives
(Alcock, 1900; Rathbun, 1902; Borradaile, 1903); Sri Lanka and Andaman Islands (Alcock, 1900);
Myanmar (Henderson, 1893); Celebes Sea, Indonesia (Miers, 1886); Ternate, Indonesia (De Man. 1902);
Ceram, Kai, and Aru islands, Indonesia (MOOSA & SERENE, 1981); Torres Snail (WHITE, 1847b; MlERS. 1874;
Haswell, 1882; Calman, 1900); Queensland, Australia (Haswell, 1882; Ward, 1942a). Its distribulion is
now extended to include the Gulf of Aden, Comoro Islands, Madagascar. Sri Lanka, Malaysia, Singapore. Japan.
Philippine Islands, Bismark Archipelago, and the Coral Sea (Chesterfield Islands, Chesterfield-Bellona Plateau, and
New Caledonia). It has been collected from the same Coral Sea stations together with N. contractus and the two
species of Palicoides. Depth: 10 m (MlERS, 1886) to 146 m (Rathbun, 191 1).
Genus PALICOIDES Moosa & Serene. 1981
Palicoides Moosa & Serfene, 1981: 45.
IYPE Species. — Cymopolia whitei Miers, 1884 by original designation (MOOSA & SERENE, 1981). Gender
masculine.
DIAGNOSIS. Carapace subquadrate; frontal border divided into two large lobes that continue to supraorbital
border as sinuous border without distinct notch. Anterolateral borders each with two large truncate teeth. Dorsal
Source : MNHN . Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
559
surface of carapace with large, low granules and very low bosses. Thick, simple setae throughout carapace and
surface of appendages. Eyes spherical; each peduncle with a conspicuous, crescent-shaped process on anterior border
and hard, rounded tubercle on ventral side. Supraorbital borders each with 1-2 triangular lobes. Suborbital borders
each with one triangular inner lobe, one broad, rectangular outer lobe. Segments 2-3 of antennae very wide and
provided with numerous long setae; flagellum relatively short. Each basal antennal segment expanded into thick,
flat process. Epistome expanded dorsoventrally. forming broad, semicircular, nearly Oat surface; thin carina-like
process across median portion with two rounded, tubercle-like central teeth. Chelipeds nearly equal in males and
females, slender in females, heavier and very long in males. First three pairs of walking legs (P2-4) with flattened
(not filiform) carpi, propodi, and dactyli. First pair (P2) shorter than second and third pairs (P3-4); third pair about
as long as second pair. Meri of P2-4 elongate, borders with small, low tubercles except distalmost one. Anterior
borders of propodi of P3-4 without wide, convex enlargement; posterior borders of dactyli entire. Fifth pair of
pereopods (P5) reduced (0.8-0.9 CL), thick; rows of thick spines along posterior border of propodus. Abdomen of
mature males elongate (sides almost parallel to each other), with all segments free or segments 4-6 fused, complete
transverse ridge along segments 1-3 (two on fused segments 4-6). Male first pleopods long and slender; basal parts
sinuous, each distal part biramous. Abdomen of mature females with segments 3-6 fused, one complete transverse
ridge along segments 1-2 (varying number of complete or incomplete ridges on fused segments).
REDESCRIPTION. — Carapace subquadrate, slightly broader than long, with large, low granules and well
defined, low bosses. Anterolateral borders straight, each with two truncate teeth followed by granular or tuberculaie
posterolateral border. Confluence of branchial and mesogastric regions depressed and smooth. Thoracic sternitc 7
with reduced process (episternal process) at each outer edge and posterior to insertion of fifth pair of pereopods
(P5), almost invisible dorsally. Posterior border of carapace smooth, without protruding tubercles. Thick, simple
setae (sometimes dark in color) throughout carapace and appendages.
Frontal border divided into two rounded lobes, each continuing without distinct notch into folded border ending
in sharp angle forming V-shaped Fissure at supraorbital border. Supraorbital borders each with 1-2 triangular lobes.
Postorbital angles long (extending just to anterior border of retracted eyes), pointed. Cornea of eyes spherical,
almost as wide as base ol peduncle; each peduncle with one conspicuous, thin, crescent-shaped process on anterior
border, several soft or granular tubercles, and one hard, rounded tubercle on ventral side. Orbits deep.
Suborbital border with one small, triangular inner lobe, one broad, rectangular outer lobe. Pterygostomial
process at each anterolateral angle of buccal frame projects ventrally, triangular or rounded edge.
Antennules short, thick, and transversely folded beneath front; interantennular septum narrow. Each basal
antennal segment expanded into thick, dorsoventrally-flattened, scale-like process. Epistome expanded
dorsoventrally, forming nearly Hat surface between antennular fossae and anterior border of endostome; two
rounded, tubercle-like central teeth on low, thin, carina-like process along median portion, flanked at same level by
two very short outer lobes that connect with each pterygostomial lobe. Meri of third maxillipeds smaller and
narrower than ischia.
Chelipeds equal or nearly equal in males and females, long and slender in females, very long (propodus and
merus becoming greatly elongated, chelipeds slightly more unequal with increasing body size), slender in males.
Fingers of chelipeds with cutting edge or triangular teeth (broadly rounded teeth in chelipeds of largest males),
pollcx becoming flatter with increasing size. Numerous simple and plumose setae along inner surfaces of cheliped
propodi and fingers of males. First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-
4); third pair slightly shorter than second. First three pairs of walking legs (P2-4) with flattened carpi, propodi, and
dactyli. Meri elongate; anterior (dorsal) and posterior (ventral) borders with small, unequal, low tubercles, rows of
low tubercles on dorsal and ventral surfaces. Elongate (but not filiform and conspicuously slender) propodi and
dactyli; posterior borders entire. Last two pairs of walking legs (P3-4) each with slightly broadened basis-ischia.
having slightly flattened, thick-edged anterior and posterior borders. Anterior borders of propodus of P3-4 without
conspicuously wide, convex enlargement. Fifth pair of pereopods (P5; Figs 42 a-b) reduced (0.8-0.9 CL), thick;
basis-ischia and meri long and tuberculate; propodi with 2-3 rows of very thick spines along posterior border;
dactyli thick and slightly curved.
Abdomen of mature males elongate, with both sides almost parallel to each other, with all segments free or
segments 4-6 fused, segments 1-2 dorsoventrally compressed, one transverse ridge along each segment (incomplete
Source :
560
P. CASTRO
and very slight in segments 5-6) or only on segments 1-3 (plus two incomplete and very slight ridges along
proximal portion of fused segment 4-6). Male first pleopods long and slender; basal parts sinuous; each distal part
biramous. Second male pleopods short, thin, slightly curved; distal segment with blunt tip. Abdomen of immature
males elongate, with varying number of fused segments.
Abdomen of mature females broad, rounded, with segments 3-6 fused, segments 1-2 dorsoventrally compressed,
one complete transverse ridge along each segment 1-2 plus two along proximal portion of fused segment 3-6.
Abdomen of immature females triangular (but broader than in males), segments 1-2 dorsoventrally compressed, 3-6
fused, complete transverse ridge along each segment 1-2 and one along proximal portion of fused segment 3-6.
SPECIES included. — P. longimanus (Miyake, 1936) and P. whitei (Miers, 1884).
FIG. 42. — a. Palicoides longimanus (Miyake, 1936), S 9.9 x 1 1.5 mm, north coast of New Caledonia. Lagon, stn 894.
12 m (MNHN-B 26997): dorsal view. — b. Palicoides whitei (Miers. 1884), 6 8.6 x 9.6 mm, Seychelles. Reves 2.
stn 50, 45-50 m (MNHN-B 26798): dorsal view.
Sexual Dimorphism. — Females are generally larger in size than males. The chelipeds are equal or nearly
equal in both sexes but they are much longer in males, increasing in length with body size. The lower surfaces ot
the chelipcd meri and fingers of the males are provided with many long simple and plumose setae. They are
particularly abundant and longer along the distal portion of the meri.
Discussion. — Although close to Neopalicus in the morphology of the carapace, species of Palicoides can be
easily differentiated by the conspicuous, crescent-shaped process on each eye peduncle. In contrast, the eye
peduncles of Neopalicus have several tubercles but never a conspicuous process. There are also signi ficant
differences in the morphology of the basal antennal segment (see discussion of Neopalicus) and abdomen. Several
segments are fused in Palicoides (4-6 in males of P. longimanus, 3-6 in females of both species), while none are
fused in Neopalicus. Other differences are listed in Table 1. Another characteristic of Palicoides is the decrease in
size of the upper borders of the meri of the third maxillipeds. It is reduced to a short, rounded or slightly pointed
protuberance.
MOOSA & SERENE (1981) mention in the diagnosis of their new genus Palicoides that the male abdomen has
"distinct segments" (MOOSA & SERENE, 1981: 45). Segments 4-6 are fused in P. longimanus and in smaller
males of P. whitei. The fusion of segments in P. longimanus, however, is mentioned in their key to the species
(Moosa & Serene, 1981: 46).
No described species of palicids from outside the Indo-west Pacific region resemble Palicoides in terms of the
quadrate, relatively flat carapace, and thickened fifth pereopods. Since the same applies to Neopalicus, it appears
that shallow-water, coarse-sediment habitats associated with coral reefs are occupied by palicids only in the Indo-
west Pacific region.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
561
Key to the species of the genus PALICOIDES
1. Two frontal lobes of carapace small, separated by narrow, shallow notch (Fig. 42a).
Supraorbital border with only one triangular lobe . P. longimanus (Miyake, 1936)
— Two frontal lobes of carapace broad, separated by wide, deep notch (Fig. 42b). Supraorbital
border with one triangular lobe and small, rounded to slightly pointed outer lobe .
. P. whitei (Miers, 1884)
Palicoides longimanus (Miyake, 1936)
Figs 42a, 43a, 50, 61 d
Cymopolia longimana Miyake, 1936: 495, fig. 1, pi. 35, figs 3-4. — Sakai, 1939: 609, fig. 90c; 1956: 52.
Palicus longimanus - SerEne, 1968: 96. — Sakai, 1976: 595, fig. 325c.
Palicoides longimanus - Moosa & SerEne, 1981: 46, fig. 8, pi. 3. fig. A.
Palicus whitei - Garth et ai, 1987: 259 [non Palicoides whitei (Miers, 1884)].
MATERIAL EXAMINED. — Japan. Yaeyama Islands, lriomotc Island, Sonai, Yaeyama Expedition. H. Ohshima coll..
08.1934: 1 3 holotype of Cymopolia longimana Miyake, 1936 9.4 x 10.9 mm (ZLKU 5771).
Indonesia. Sulawesi. Bahuluang Island, from beach sand, 10.10.1984: 1 <3 (RMNH 47901).
Moluccas, Ambon. RUMPHIUS Expedition 1: stn MO-1, 03°32’00"S, 128°12'30"E, Kasijan coll., 8.01.1973: 1 2
(MNHN-B 10006).
Rumphius Expedition 2: seagrass meadow, R. SerEne coll. (CB 1261), 26.01.1975: 1 <3 (MNHN-B 27270).
Chesterfield Islands. CORAIL 2: stn DW 41, 19°21.52'S. 158°31.87'E, 52 m. Halimeda sand. 23.07.1988: 1 <3
(MNHN-B 27271). — Stn DW 59, 19°18.50'S, 158°56.55'E, 50 m, 24.08.1988: 1 2 (MNHN-B 27272). — Sin DW 73,
19°1 1.00'S, 158°22.57'E, 41 m, 25.08.1988: 1 2 (MNHN-B 27273). — Stn DW 87, 19°06.14'S, 158°59.94'E. 31 m,
coarse sand and Halimeda, 28.08.1988: I 2 (MNHN-B 27274). — Stn DW 154, 19°52.04'S. 158°26.5'E, 35 m.
Halimeda, 1.09.1988: 1 2 (MNHN-B 27275). — Stn DW 160, 19°46.00'S. 158°23.00'E. 35-41 m. sand. 1.09.1988: 2 2
(MNHN-B 27276).
New Caledonia. Lagon: stn 1, 22°18.0’S, 166°24.6'E, 19 m. fine sand and Halimeda, 05.1984: 1 2 (MNHN-B
27248). — Stn 7, 22°24.0'S, 166°19.7'E, 14 m, white sand, 05.1984: 1 2 (MNHN-B 27249). — Stn 49, 22°18.5’S,
1 66°13.8’E, 10 m, sargassum. 05.1984: 1 2 (MNHN-B 27263). — Stn 50, 22°16.6'S, 166°12.2'E, 12 m, white sand,
05.1984: 2 2 (MNHN-B 27277). — Stn 52, 22°14'S, 166°14'E, 13 m, seagrasses, 1984: 1 3 (MNHN-B 27258). —
Stn 69, 22°22.8’S, 166°31.7'E. 13 m, shelly sand, 08.1984: 1 2 14.7 x 17.0 mm (MNHN-B 26791). — Stn 99.
22°32.6'S, 166°34.6'E, 14 m, corals, 08.1984: 1 2 (MNHN-B 27278). — Stn 180, 21°59.6'S, 166°04.5’E. 10 m, mud.
09.1984: 1 2 (MNHN-B 27279). — Stn 185, 22°04.8'S, 166°02.2'E. 15 m, coral blocks and Halimeda, 19.09.1984:
1 <3 (MNHN-B 27253). — Stn 187. 22°02.8'S, 166°01.7'E. 13 m, shelly sand, 09.1984: 2 2 (MNHN-B 27259). —
Sin 192, 22°00.6'S, 166°00.0'E, 18 m, boulders, 09.1984: 1 2 (MNHN-B 27264). — Stn 253, 22°22.FS. 166°22.9'E.
16 m. coarse sand, 10.1984: 1 2 (MNHN-B 27280). — Stn 269, 22°18.0'S. 166°18.1'E. 20 m. muddy sand and Halimeda,
11.1984: 2 2 (MNHN-B 27250). — Stn 284, 22°25.8'S, 166°24.9'E, 6 m, corals, 10.1984: 1 2 (MNHN-B 27265). —
Stn 455, 1 8°29.5’S, 163°07.9'E, 40 m, coarse sand, 28.02,1985: 1 2 (MNHN-B 27254). — Stn 483, 19°01.0'S.
163°32.2'E, 33 m, boulders and Halimeda, 2.03.1985: 1 2 (MNHN-B 27281). — Stn 550, 22°54.0'S, 166°57.5'E, 26 m,
coral debris, 15.07.1985: 2 2 (MNHN-B 27282). — Stn 554. 22°50'S, 166°54'E, 27 m, white sand and shells.
16.07.1985: 1 3 (MNHN-B 27255). — Stn 623, 22°01.0'S, 166°50.5'E, 32-40 m, 6.08.1983: 1 2 (MNHN-B 27283). —
Stn 890, 20°20.3'S, 164°35.6’E, 23 m, coarse sand, 14.01.1987: 1 6 , 1 2 (MNHN-B 27260). — Stn 891, 20°19.8'S,
164°33.5'E, 25 m, mud and shells, 14.01.1987: 1 2 (MNHN-B 27284). — Stn 892, 20°18.3'S, 164°32.1'E. 26 m, coarse
sand and Halimeda, 14.01.1987: 1 3 , I 2 (MNHN-B 27285). — Stn 894, 20°16.6'S, 164°28.0'E. 12 m, coarse sand and
Halimeda, 14.01.1987: 1 3 9.9 x 11.5 mm (MNHN-B 26997). — Stn DW 896, 20°15.2'S, 164°29.3'E, 20 m. medium-
size sand and mud, 14.01.1987: 1 <3 (MNHN-B 27251). — Stn 898 20°13.6'S, 164027.1'E, 22 m, gray sand. 14.01.1987:
1 2 (MNHN-B 27267). — Stn 899, 20°14.2'S, 164°25.2'E, 16 m, coarse sand and Halimeda. 14.01.1987: 1 2 (MNHN-B
27286). — Stn DW 921, 20°51.2'S, 164°26.6’E, 10-11 m, shelly sand. 27.04.1988: 1 2 (MNHN-B 27287). —
Sin DW 934, 20°43.0’S, 164°16.8'E, 10 m, coarse sand and Halimeda, 27.04.1988: 1 2 (MNHN-B 27269). —
Stn DW 944. 20°35.2'S, 164°11.9'E, 14-15 m, shelly sand. 28.04.1988, 1 2 (MNHN-B 27288). — Stn DW 948.
20°32.2'S, 164°08.8'E, 16 m, coarse sand and Halimeda, 28.04.1988: 2 2 (MNHN-B 27256). — Stn DW 952. 20°30.8'S.
164°07.4'E, 16-17 m, coarse sand, 28.04.1988: 1 <3 9.6 x 11.2 mm (MNHN-B 26794). — Stn DW 953, 20°3I.8'S.
164°05.8'E, 18-19 m. white sand, 28.04.1988: 1 3 (MNHN-B 27261), — Stn DW 954, 20°31.0'S. I64°03.0'E. 15-17 m,
Source
562
P. CASTRO
coarse sand. 20.04.1988: 1 <J , 1 2 (MNHN-B 27289). — Stn 985, 20°20.3'S. 163°57.9'E. 15-17 m. coarse sand and
Halimeda, 30.04.1988: 1 2 (MNHN-B 27257). — Stn DW 989. 20°18.1'S, 1 63°57. 1 E, 21 m. foraminiferan sand.
30.04.1988: 1 2 (MNHN-B 27290). — Stn DW 1008. 20°11.0'S. 163°53.4'E. 27 m, shells and foraminiferans.
2.05.1988: 1 2 (MNHN-B 27262). — Stn DW 1046. 20°05.0'S. I64°06.6'E. 6-7 m, boulders and Halimeda. 4.05.1988:
1 <3. 1 2 (MNHN-B 27291). — Stn DW 1075, 19°52.0'S. 163°58.4’E, 28 m, white sand and Halimeda. 23.10.1989: 1
2 (MNHN-B 27268). — Stn DW 1094, 19°54.4'S. 163°41.2'E, 26 m, foraminiferan sand and Halimeda. 24.10.1989: I 3
(MNHN-B 27292). — Stn DW 1163, 19°11.3'S. 163°21.9'E. 48 m, fine sand. 30.10.1989: 1 2 (MNHN-B 27213). —
Stn DW 1168, 1 9°15.9'S, 163°09.3'E. 50 m, coarse sand. 30.10.1989: 1 2 (MNHN-B 27294). — Stn DW 1181,
19°23.9'S. 163°14.7'E, 45 m. coarse sand, 31.10.1989: 1 6 . 1 2 (MNHN-B 27295). — Stn DW 1182. 19°27.3'S.
I63°16.2'E, 48 m, coarse sand, 30.10.1989: 1 3 , 1 2 (MNHN-B 27296). — Stn DW 1205, 19°41.6'S. 163°25.6'E, 38
m. coarse sand. 2.11.1989: I <3 6.2 x 6.8 mm (MNHN-B 26793). — Stn DW 1210, 19°45.5'S, I63°35'E. 31 m
3.11.1989: 1 3 (MNHN-B 27252).
Passe de Boulari: 15 m, 20.03. 1988: 1 3 11.4 x 12.5 mm (MNHN-B 26792). — llot Goeland: 10 m, 16.04.1993:
1 3 (MNHN-B 27297). — lloi Maitre: 6 m, seagrass meadow, suction pump, P. BOUCHET coll., 24.03.1993: 1 2
(MNHN-B 27300). — Bancs de I'Ouesf. I 3 (MNHN-B 27298). — Recif Above: 10 m. P. BOUCHET coll., 8.10.1992: 1 2
(MNHN-B 27299).
Loyalty Islands. Musorstom 6: stn DW 434. 20°21.21'S, 166°08.64'E, 23 m, 18.02.1989: 1 3 (MNHN-B
27266).
Marshall Islands. Enewetak Atoll: stn 429. J.W. Knudsen coll., 15.08.1967: 1 2 (LACM). — Stn 431.
16.08.1967: 2 2 (LACM). — Stn 183: 2 3. 1 2 (LACM). — Stn 184: 1 3 (LACM). — Stn 187: 1 juv„ 3. 12 (LACM).
— Stn 189: 1 juv. 5 (LACM). — Stn 722: 1 3 (LACM).
TYPES. — Holotype of Cymopolia longimana Miyake, 1936: 1 S 9.4 x 10.9 mm, Ryukyu Islands, Japan
(ZLKU5771).
Type Locality. — Sonai, Iriomote Island, Yaeyama Islands, Ryukyu Islands, Japan.
Diagnosis. — Carapace (Figs 42a, 61d; Miyake, 1936, pi. 35, fig. 3; Moosa & Serene, 1981. pi, 3,
fig. A) subquadrate, with large, low granules. Frontal border with two small lobes separated by narrow, shallow
notch. Two truncate anterolateral teeth. Supraorbital borders each with one triangular lobe. Each eye peduncle with
rounded, low, soft tubercle at distal end. Postorbital angles triangular, with rounded tips. Suborbital borders each
with one rounded to triangular inner lobe and one broad, rounded outer lobe (Miyake, 1936. fig 1 A). Segments 2-
3 of antennae very wide, thick, numerous long setae. Each basal antennal segment expanded into thick,
dorsoventrally- flattened process. Chelipeds with long carpi and meri in larger specimens of both sexes but
extremely long and increasingly (but slightly) unequal in males (Fig 42a; MlYAKE, 1936. pi. 35, figs 3-4); dorsal
borders of propodi with irregular, microscopic tubercles; outer border with hook-like setae; long, simple and
plumose setae along lower borders of cheliped propodi and fingers in males. Upper and lower borders of meri and
anterior borders of carpi of second and third pairs of walking legs (P3-4) with very small, low tubercles; meri of P2
elongate, equal or longer than half carapace length (0.5-0. 6 CL). Abdomen of mature males narrow, with segments
4-6 fused; one complete transverse ridge along segments 1-3, low ridges on proximal portion of fused segment 4-6.
Male first pleopods (Fig. 43a; Miyake, 1936, figs ID, F; Moosa & Serene, 1981, fig. 8) with sinuous basal
parts; each distal part spherical, with two pointed ventral teeth and long, toothed dorsal process. Abdomen of
mature females with segments 3-6 fused, one transverse ridge along each segment 1-2, two complete ridges along
proximal portion of fused segment 3-6.
Color: A male from New Caledonia photographed live (Fig. 6 Id) had a broad, vertical, red-brown band through
the middle of the carapace, irregular black patches along the sides of the carapace, and thin red-brown bands across
the walking legs. The chelipeds were red-brown. A female from the same area had a similar color pattern except
that the band across the carapace was solid red-brown. Many preserved specimens of both sexes had a dark dot on
the median portions of the dactyli of the chelipeds and fifth pereopods (P5).
Discussion. — The single male specimen that was used in the description of P. longimanus (Miyake, 1936,
pi. 35) was characterized by very long and slender chelipeds, hence the name of the species. MOOSA & Serene
(1981), who examined only three specimens (one male and two females, all of a relatively large size), concluded
that the chelipeds were always very long in both sexes. They used the relative length of the chelipeds to
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
563
distinguish between P. longimanus and the two other species they included in the new genus Palicoides , P. whitei
and P . ternatensis Moosa & Serene, 1 98 1 , a subjective junior synonym of P. whitei (see below).
Fig. 43. — a, Palicoides longimanus (Miyake. 1936), 3 11.4 x 12.5 mm, Passe de Boulari. New Caledonia. 15 m
(MNHN-B 26792): left male first pleopod, venlral view. — b-d, Palicoides whitei (Miers, 1884): b, 3 5.1 x 5.0 mm.
Lagon, Grand Recif, New Caledonia, stn 1094, 26 m (MNHN-B 26795): left male first pleopod (ventral view) of
immature specimen. — c, 3 8.2 x 9.7 mm, west coast of New Caledonia, Lagon, stn 815, 32 m (MNHN-B 26800):
left male first pleopod (ventral view) of intermediate-size specimen. — d. 3 CL 14.5 mm. Lagon, west coast of New
Caledonia, stn 748, 35 m (MNHN-B 26801): left male first pleopod (ventral view) of large-size specimen.
Examination of a large number of specimens of P. longimanus and P. whitei has shown that very long
chelipeds (total length far exceeding carapace length) are found in large individuals of both species (Figs 42a-b,
61e). The relative length of the chelipeds increases with body size in both sexes. A direct relationship between
chcliped length and body size is not always observed, however, perhaps the result of some chelipeds being
regenerated (Fig. 6 Id). In P. longimanus, male chelipeds become longer than carapace length at a carapace length
of approximately 9.0 mm, reaching 2.2 times carapace length (chelipcd meri 0.9 times carapace length) in the
largest male examined, 1 1.4 x 12.5 mm (MNHN-B 26792). Female chelipeds reach a length equal to carapace
length at a carapace length of approximately 1 1.0 mm but they never exceeded carapace length in any of the
females that were examined. The meri of male chelipeds were never found to be longer than carapace length, as
Source
564
P. CASTRO
stated in the key of MOOSA & SERENE (1981: 45). They also referred to only two fused segments (5-6) in the
only P. longimanus male they examined (MOOSA & Serene, 1981: 52). Adult males were actually found to have
segments 4-6 fused.
The specimens examined in this study agree with the description of the species (MIYAKE. 1936) in most
aspects. No mention was made in the description of the crescent-shaped process on the eye peduncles, a diagnostic
characteristic of the genus, although it is present in the holotype (ZLKU 5771). The tip ol the process on the left
peduncle, however, is severed. A small but not crescent-shaped process is shown in two figures (Miyake, 1936,
figs 1 A-B), one of which is reproduced by Sakai ( 1939, fig. 90c; 1987, fig. 325c). The figure of the anterolateral
portion of the carapace (MIYAKE, 1936, fig. IB) does not show the gap between the first anterolateral tooth and
the postorbital angles. The gap was found in the holotype and in most but not all of the specimens that were
examined. Another diagnostic character present in the holotype but not mentioned in the description or shown in
Miyake's figures is the presence of long setae along the lower borders of the chelipeds.
MOOSA & Serene (1981: 45) described the male first pleopods as having a "spiraled shaft". It is best described
as sinuous, but not spiral (or helicoidal) as in the basal parts of Parapalicus. The male first pleopods of the
smallest male examined in this study (6.2 x 6.8 mm; MNHN-B 26793) consisted of two flat terminal processes
similar to the male first pleopods in small males of P. whitei (see below). Unlike P. whitei. however, two minute
spines were present on opposite sides at the base of the processes.
Some variation was observed in the shape of the basal antennal segment. It was elongate or even biramous in
a few specimens but thick and dorsoventrally-llattened in the great majority of specimens that were examined.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
P. longimanus may be easily confused with P. whitei, its only congener. Differences between species are given
in Table 4. In many specimens of P. longimanus there is only a very narrow fissure between the first anterolateral
teeth and the postorbital angles, whereas it is wider in many specimens of P. whitei (MOOSA & SERENE, 1981,
fig. 10). This, however, is not an absolute character.
TABLE 4. — Morphological differences between Palicoides longimanus (Miyake, 1936) and P. whitei (Miers. 1884).
SIZE. — Maximum size among specimens examined: 14.7 x 17.0 mm (female, MNHN-B 26791); 1 1.4 x
12.5 mm (male, MNHN-B 26792).
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
565
Distribution. — Previously known only from the Ryukyu Islands, Japan (Miyake, 1936; Sakai, 1956) and
Kai (Banda Sea) and Tanimbar (Arafura Sea) islands, Indonesia (MOOSA & SERENE. 1981). It is now also known
from Ambon (Indonesia), the Coral Sea (Chesterfield Islands, New Caledonia, and the Loyalty Islands), and the
Marshall Islands (Fig. 50). It has been collected from the same Coral Sea stations together with Neopalicus jukesii
and P. whitei. Depth: 6 m to 57 m (Moosa & SERENE, 1981).
Palicoides whitei (Miers, 1884)
Figs 42b, 43b-d, 50, 61e
Cymopolia whitei Miers, 1884: 551, pi. 49, figs C. c. — GutNOT, 1967: 280.
Palicus whitei - Bouvier, 1897: 65. — Alcock, 1900: 453. — Calman. 1900: 31. pi. 2, figs 14-19. — Rathbun. 1911:
240, pi. 19, fig. 10. — Laurie, 1915: 416, 421, 473. — McNeill. 1968: 82. — Serene, 1968: 96.
Palicoides whitei - Moosa & Serene, 1981: 46, figs 9-10.
Palicoides ternatensis Moosa & Serene, 1981: 50, fig. 1 1, pi. 3, fig. B.
MATERIAL EXAMINED. — Socotra. Stn 146, I2°37.1'N, 54°18.3'E. 20-25 m, sand, under coral Goniopora
stokesii, M, APEL coll., 2.04.1999: 1 3 (SMF 25280).
Seychelles. "Alert": stn 194, 7-22 m, COPPINGER coll., 03.1882: 1 6 lectotype of Cymopolia whitei Miers. 1884
6.0 x 6.7 mm, 1 3 paralectotype 5.8 x 6.5 mm, 2 2 paralectotypes 8.4 x 9.4 mm, 7.5 x 8.4 mm (BMNH 1882.24).
Reves 2: stn 41, 04°44.0'S, 56°15.1'E. 50 m, shelly sand and Halimeda. 13.09.1980: 1 2 (MNHN-B 27303). —
Stn 47. 04°03.8'S, 55°59.5'E, 45-55 m, shelly sand, 14.09.1980: 2 3 (MNHN-B 27310). — Stn 50. 03°56.I’S.
55°40.5'E. 45-50 m. shelly sand, 15.09.1980: 1 3 (MNHN-B 27304), 1 3 8.6 x 9.6 mm (MNHN-B 26798).
Mauritius. Tombeau Bay, stn 38. 73 m, T. Mortensen coll., 8.10.1929: 1 3 (ZMUC).
Madagascar. Nosy Be. A. Crosnier coll.: 1 <J (MNHN-B 27317).
Comoro Islands. Mayotte. Lagoon, medium-size sand, 55 m, A. Crosnier coll., 09.1958: I 2 (MNHN).
Japan. Ryukyu Islands. Okinawa, 26°30.4'N, 127°52.6'E, 48.8 m, R.F. Bolland coll.. 12.06.1984: 1 3 (USNM).
— 26°30.0'N, 127°50.9'E, 64 m, R.F. Bolland coll.. 29.06.1984: 1 3 (USNM). 1 2 (USNM). — Kerama Islands.
Amuro-jima Island, "Toyoshio-maru", stn 7, 26°10.00'N, 127°18.90'E, 52 m. T. Komai coll.. 19.05.1998: I 3 (CBM-ZC
4671).
Indonesia. Hahnahera. Mariel King Memorial Expedition: stn HD i/I. Dodinga Bay, 0°49'N. 127°33'E. 31-42 m,
coral ( Acropora ) sediment and mud, 20.05.1970: 1 3 holotype of Palicoides ternatensis Moosa & Serene. 1981 10.5 x
12.3 mm (RDC CB 2724).
Saparua Island. RUMPHIUS EXPEDITION 1: stn Ip3. 03°30'I5"S. 128o41'20"E. KaSIJAN coll.. 18.01.1973: 1 3 (MNHN-
B 10005).
Australia. Torres Strait. Murray Island. A.C. HaddON coll., 1888: 2 2 (BMNH 1954.9.14.89-90).
Queensland. Great Barrier REEF EXPEDITION: stn 16. off Lookout Point, 9.03,1929: 1 (3,1 2 . — Stn 17. off
Lookout Point. 9.03.1929: 3 2 (BMNH 1950.12.1.24-27).
Chesterfield Islands, Chesterfield-Bellona Plateau. Chalcal 1: stn DC 34, 19°52.10'S, I58°20.10E.
33-37 m, Halimeda sand, 21.07.1984: 1 3 (MNHN-B 27343). — Stn DC 45, 20°48.93'S, 158°30.21E. 50 m,
23.07.1984: 1 3 . 2 2 (MNHN-B 27346). — Stn CP 14. 2I°13.50'S. 158°50.20'E, 66 m. 24.07.1984: I 2 (MNHN-B
27347). — Stn DC 53. 21°19.50'S, 158°55.30'E, 60 m. 24.07.1984: 1 2 20.5 x 23.5 mm (MNHN-B 26799), I 3
(MNHN-B 27351). — Sin DC 55. 21°23.90'S. 158°59.60'E. 55 m, Halimeda and foraminiferans. 25.07.1984: 3 2
(MNHN-B 27348).
"Vauban": stn 7 DE, 21°21.0'S, 158°52.7'E. 56-58 m. 19.10.1985: 1-3,12 (MNHN-B 27349). — Stn 14 DE,
21°46.5'S, 159°28.3'E, 42-52 m, 22.10.1985: 1 2 (MNHN-B 27350).
Corail 2: stn DW 41, 19°21.52'S, 158°31.87'E, 52 m, Halimeda. 23.07.1988: 1 juv. (MNHN-B 27352). —
Stn DW 59, 19°18.50'S, 158°56.55'E, 50 m, 24.08.1988: 1 2 (MNHN-B 27354). — Stn DW 67. I9°14.9'S.
158°36.94’E. 66 m, 24.08.1988: 1 2 (MNHN-B 27355). — Stn DW 79, 19°11.55'S. 158°43.40'E. 58 m, Halimeda sand.
25.08.1988: 1 3 , 3 2 (MNHN-B 27356). — Stn DW 93. 19°05.92'S, 158°53.00'E. 58-60 m, Halimeda. 27.08.1988:
2 3 , 1 2 (MNHN-B 27357). — Stn 94, 19°06.00'S. I58°50.00'E. 36-53 m, boulders and Halimeda. 27.08.1988: 1 3
(MNHN-B 27358). — Stn 101, 19°08.99'S, 158°26.24'E, 37 m, Halimeda sand. 27.08.1988: 1 2 (MNHN-B 27359). —
Stn DW 110, 19°08’95'S. 158°55.8?'E, 48 m, shelly sand, 28.08.1988: 1 2 (MNHN-B 27360). — Stn DW 117.
19°25.10'S, I58°31.70'E, 52 m. Halimeda sand. 28.08.1988: 1 3 16.2 x 18.9 mm (MNHN-B 26796). — Stn DW 135.
19°31 ,37'S. 1 58°19. 14'E. 46 m, coarse sand, 30.08.1988: I 2 (MNHN-B 27361). — Stn DW 136, 19°31.20'S,
158°I6.00'E. 37 m. sand and boulders, 30.08.1988: 1 2 (MNHN-B 27362). — Stn DW 154. 19°52.04’S. I58°26.50'E.
35 m, Halimeda sand. 1.09.1988: 1 2 (MNHN-B 27363). — Stn DW 155. 19°49.08'S. I58°24.85'E, 42 m. foraminiferan
sand, 01.09.1988: 1 2 (MNHN-B 27364). — Stn DW 166. 19°41.49'S. 158°25.24’E. 56 m, boulders. 2.09.1988: 1 2
(MNHN-B 27365).
Source : MNHN. Paris
566
P. CASTRO
New Caledonia. LaGON: sin 100, 22°32.6’S. 166°34.6'E. 15 m, shelly sand. 08.1984: 1 2 (MNHN-B 27318). —
Stn 239, 22°24.3'S, 166°57.8'E, 43 m. coarse sand, 10.1984: 1 2 (MNHN-B 27319). — Stn 313, 22°40.3'S,
166°50. 1'E, 30 m, coarse sand and Halimeda, 11.1984: 1 2 (MNHN-B 27331). — Sin 345, 22°46.4'S, 166°50.4'E, 39 m,
while sand and corals, 1 1.1984: 2 2 (MNHN-B 27311). — Sin 353, 22°33.5'S, 167°00.8'E. 70 m. coarse sand and corals.
11.1984: I 2 (MNHN-B 27312). — Stn 474, 18°02.4’S. 163°01.8'E, 52 m, Halimeda sand, 1.03.1985: 1 2 (MNHN-B
27332). — Stn 477, 18°51.0'S, 163°27.0'E, 50 m, Halimeda sand. 2.03.1985: 2 2 (MNHN-B 27320). — Stn 540,
19°06.2'S. 163°15.8'E, 35-40 m. Halimeda sand. 6.03.1985: 1 3 (MNHN-B 27333). — Stn 541, 19°06.0’S. 163°13.3’E,
foraminiferan sand. 48-43 m. 6.03.1985: 1 2 (MNHN-B 27321). — Stn 601. 22°18.0'S, 167°02.5'E. 47-48 m, shelly
sand, 5.08.1986: 1 2 (MNHN-B 27305). — Stn 708, 21°23.6'S, 166°05.2'E. 34-35 m, foraminiferan sand, 10.08.1986:
1 2 (MNHN-B 27313). — Stn 709, 21°22.2'S. 166°03.5'E, 39-40 m. foraminiferan sand. 10.08.1986: I 2 (MNHN-B
27334). — Stn 710, 21°24.0'S, 166°02.5'E, 30-31 m, foraminiferan sand, 10.08.1986: I 2 (MNHN-B 27335), I 3
(MNHN-B 27306). — Stn 714, 21°21.0’S, 166°01.8'E, 37-38 m, foraminiferan sand, 11.08.1986: 1 5 (MNHN-B
27307). — Sin 747, 21°14.7'S, 165°50.9'E, 31-34 m, foraminiferan sand and Halimeda, 6.01.1987: 1 3 (MNHN-B
27322), 1 2 (MNHN- B 27323). — Stn 748, 21°16.9'S, 165°49.9'E, 35 m, coarse sand and Halimeda. 6.01.1987: 1 3
CL 14.5 mm (MNHN-B 26801), 1 3,42 (MNHN-B 27324), 1 3 15.5 x 18.1 mm (MNHN-B 26797). — Stn 749,
21°18.4'S. 165°1 8.4'E, 49 m, mud and shells. 6.01.1987: 1 2 (MNHN-B 27308). — Stn 765, 21°13.8'S, 165°41.8'E.
35 m. coarse sand, 8.01.1987: 1 2 (MNHN-B 27325). — Stn 780, 21°06.0'S, 165°39.2'E. 33 m, coarse sand.
8.01.1987: 1 3 (MNHN-B 27309). — Stn 782, 21°06.rS, 165°36.7'E, 30 m, coarse sand. 8.01.1987: 1 2 (MNHN-
B 27326). — Stn 788, 21°01.6'S. 165°34.7'E, 33 m, coarse sand, 9.01.1987: I 2 (MNHN-B 27336). — Stn 794.
21 °03.2'S. 165°30.9'E, 51 m. mud, 9.01.1987: 13.22 (MNHN-B 27337). — Sin 795, 21°01.6, 165°32.0'E. 31 m,
coarse sand. 9.01.1987: 3 3 (MNHN-B 27338). — Stn 801, 2r02.0'S, 165°29.3'E, 29 m, foraminiferan sand,
9.01.1987: 1 2 (MNHN-B 27339). — Stn 815, 21°54.1'S, 165°26.9'E, 32 m. coarse sand. 10.01.1987: I 3 8.2 x
9.7 mm (MNHN-B 26800). — Stn 821, 20°51.9'S, 165°23.2E, 32 m, coarse sand, 10.01.1987: 2 3,12 (MNHN-
B 27314), 1 3 (MNHN-B 27341). — Stn 828, 20°50.1'S, 165°19.5'E, 28 m, coarse sand, 10.01.1987: 2 3 (MNHN-
B 27327). — Stn 847, 20°37.6'S, 165°13.4'E, 28 m, fine sand, 11.01.1987: 1 2 (MNHN-B 27340). — Stn 867,
20°39.0'S, 165°01.3'E, 25 m, shell debris, 13.01.1987: 1 2 (MNHN-B 27328). — Stn DW 985, 20°20.3'S. 163°57.9'E,
15-17 m, coarse sand and Halimeda, 30.04.1988: 1 3 (MNHN-B 27315). — Stn DW 1094, 19°54.4'S, 163°41.2'E. 26 m.
foraminiferan sand and Halimeda, 20.10.1989: 1 3 5.1 x 5.0 mm (MNHN-B 26795). — Stn DW 1126, 19°33.0'S,
163°46.0'E, 41 m, coarse sand, 26.10.1989: 1 3 (MNHN-B 27316). — Stn DW 1139, 19°23.6'S, 163°47 0'E 39 m
Halimeda sand, 27.10.1989: 6 2 (MNHN-B 27342).
Passe de Touho. Suction pump, 20 m, 18.09.1993: 1 2 (MNHN-B 27345).
Ildt Maitre. 22 m, P. BOUCHET coll., 28.09.1992: 1 2 (MNHN-B 27344).
Types. Lectotype of Cymopolia whitei Miers, 1884: 1 3 6.0 x 6.7 mm; paralectotypes 1 3 5.8 x 6.5 mm,
2 2 8.4 x 9.4 mm; 7.5 x 8.4 mm, "Alert", stn 194 (BMNH 1882.24).
H o!o type of Palicoides ternatensis Moosa & Serene, 1981: 1 3 10.5 x 12.3 mm, Marif.l King Memorial
Expedition, stn HD 1/1 (RDC CB2724).
TYPE LOCALITY. — Seychelles, "Alert", stn 194, exact location unknown. 7-22 m.
Diagnosis. — Carapace (Figs 42b, 61e; Miers, 1884, pi. 49, fig. C; Calman, 1900, pi. 2, fig. 14; Rathbun,
1911, pi. 19, fig. 10; Moosa & Serene, 1981. pi. 3, fig. A, as P. ternatensis-, fig. 10) subquadrate, with large,
low granules. Frontal border with two broad lobes separated by wide notch. Two truncate anterolateral teeth.
Supraorbital borders each with one triangular inner lobe and smaller, rounded to slightly pointed outer lobe. Each
eye peduncle with conspicuous, pointed, soft tubercle at distal end. Postorbital angles elongate, with pointed tips.
Suborbital borders each with two rectangular to slightly rounded lobes (Calman, 1900, pi. 2, fig. 15). Segments
2- 3 ol antennae very wide, thick, numerous long setae. Each basal antennal segment expanded into thick,
dorsovcntrally-flattened process. Chelipeds with long carpi and meri in larger specimens of both sexes but
extremely long and increasingly (but slightly) unequal in males (Figs 42b, 61e); dorsal borders of propodi with
rounded or pointed tubercles; outer border with hook-like setae; long, simple and plumose setae along lower
borders of cheliped propodi and fingers of males. Upper and lower borders of meri and anterior borders of carpi of
second and third pairs of walking legs (P3-4) with very small, low tubercles (Calman, 1900, pi. 2, fig. 18).
Abdomen ot mature males narrow, with all segments free; one complete transverse ridge along segments 1-3
(incomplete one in segment 4, very low or absent in segments 5-6). Abdomen of immature males with segments
3- 6 or 4-6 fused (Miers, 1884, pi. 49, fig. c). First pleopods of small males (Fig. 43b; Calman, 1900, pi. 2, fig.
19; Moosa & SerCne, 1981, fig. 9) with sinuous basal parts; distal part each with two flattened processes provided
Source : MNHN, Paris
1NDO-WEST PACIFIC SPECIES OF PALICID CRABS
567
with teeth. First pleopods of larger males (Figs 43c-d; Moosa & Ser£ne, 1981. fig. II, as P. ternatensis) with
sinuous basal parts; each distal part with bent, toothed tip becoming enlarged as thick, arched process with thick
teeth. Abdomen of mature females with segments 3-6 fused, one complete transverse ridge along each segment 1-2,
two complete ridges along proximal portion of fused segment 3-6.
Color: A large male from New Caledonia photographed live (Fig. 61e) had a carapace that was mostly red and
orange except a broad, dark-red band bordered by pink and orange blotches on the cardiac and intestinal regions, red
chelipeds, and broad, red bands across the walking legs. Smaller males and females from the same region had a
similar color pattern except that there was a broad band of red in the form of geometric patterns through the middle
of the carapace. Many preserved specimens of both sexes had a dark dot on the median portion of the dactyli of the
chelipeds and fifth pereopods (P5).
Discussion. — The male first pleopods of P. whitei were illustrated by Cal.man (1900, pi. 2, fig. 19) and
MOOSA & Serene (1981, figs 9a-b). Their figures, however, are of the pleopods characteristic of small males,
a situation found in those with a carapace length of up to 9.0 mm (5.1 x 5.0 mm; MNHN-B 26795) among the
specimens that were examined. Calman did not give the size of this specimen; the specimen of Moosa & Serene
had a carapace length of 9.2 mm (Moosa & SERENE, 1981: 50). Such pleopods have sinuous basal parts
(although MOOSA & Serene described them as having a flattened shaft) and bifurcated distal parts (Fig. 43b). The
ventral terminal process, the longest of the two, has rows of small teeth and its tip is dorsovcntrally flattened and
bordered by smaller teeth. As body size increases, the basal part of the pleopods becomes more distinctly sinuous,
the two terminal processes and their teeth enlarge and the flattened tip of the ventral process bends downward
(Fig. 43c). These were features used by Moosa & SERENE (1981, figs 1 la-d) to characterize their new species.
P. ternatensis, here regarded as a subjective junior synonym of P. whitei. Nevertheless, there was not much
difference between size of the only specimens examined by Moosa & SERENE: one male P. whitei (9.2 x
10.6 mm) and one male, the holotype, of P. ternatensis (10.5 x 12.3 mm; RDC CB2724). The larger specimen
identified as P. ternatensis, however, had all abdominal segments free, whereas the small male had "segments 3-5
soldered" (MOOSA & Serene, 1981: 46), a characteristic of immature males (see below). In larger males the
terminal processes of the pleopods are further enlarged (Fig. 43d). The ventral process gradually becomes thicker
and arched, with thick teeth along its proximal portion and two rows of curved teeth along the outer (ventral)
border. Whereas MOOSA & SERENE refer to the basal part of the pleopods of P. whitei and P. ternatensis as having
a "flattened shaft”, their figure of the pleopod of P. ternatensis (MOOSA & SERENE, 1981. fig. 11) clearly shows a
sinuous basal part.
Moosa & SERENE (1981) also used the relative length of the chelipeds to distinguish between the species of
Palicoides, stating that it was shorter in P. whitei than in P. longimanus and intermediate in P. ternatensis. They
based their conclusion using only a relatively small male of P. whitei and a slightly larger male of P. ternatensis
(see above). Very long chelipeds where total length exceeds carapace length, however, characterize both species
(Figs 42a-b). Chelipeds of P. whitei were longer than carapace length in all the males and females that were
examined in this study except in the small males, where cheliped length was approximately equal to carapace
length. The chelipeds of the largest P. whitei male examined in this study (16.2 x 18.9 mm; MNHN-B 26796)
were 1.3 times longer than carapace length (cheliped meri 0.4 CL) but they were relatively longer in other
specimens (15.5 x 18.1 mm [MNHN-B 26797], Fig. 6 1 e; 8.6 x 9.6 mm [MNHN-B 26798], Fig. 42b).
Differences in the relative length of chelipeds probably result from their loss and eventual regeneration in some
individuals. Female chelipeds were always longer than carapace length but only reached a length slightly longer
than carapace length (cheliped meri 0.3 CL) in the largest female examined (20.5 x 23.5 mm; MNHN-B 26799).
The number of abdominal segments also vary with increasing size. Immature males have segments 3-6 fused,
but traces of sutures become progressively evident with increasing size until all segments are evident in mature
males.
Another characteristic used by MOOSA & SERENE (1981: 46) to distinguish between the species of Palicoides
is the presence in P. longimanus of irregular granules (larger on the branchial region) while being "of about the
same sizes" in P. whitei and its subjective junior synonym P. ternatensis. This was found not to be true: larger
specimens of both species have slightly larger granules on the branchial region.
Source :
568
P. CASTRO
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Length relationship of pereopods 2-5 in Palicoides whitei
MlERS (1884) based his description of Cymopolia whitei on four specimens without selecting a holotype. One
of the four syntypes (6 6.0 x 6.7 mm, BMNH 1882.24) is hereby designated as the lectotype. A second male and
two females from the same collection are paralectotypes.
Size. — Maximum size among specimens examined: 20.5 x 23.5 mm (female. MNHN-B 26799); 16.2 x
18.9 mm (male, MNHN-B 26796).
Distribution. — Previously known from the Red Sea (Laurie, 1915), Seychelles (Miers, 1884; Rathbun,
1911), Andaman Islands, India (ALCOCK, 1900), Halmahera (Moluccas) and Kai Islands (Banda Sea), Indonesia
(Moosa & Serene, 1981, some records as P. ternatensis), Torres Strait (Calman, 1900), and Queensland,
Australia (McNeill, 1968). Its distribution is now extended to Socotra, the Comoro Islands, Madagascar,
Mauritius, Japan, and the Coral Sea (Chesterfield Islands, Chesterfield-Bellona Plateau, and New Caledonia) (Fig.
50). It is an inhabitant of coarse-sand sediments (particularly those rich in the coralline alga, Halimeda ) that is
associated with coral reefs. It has been collected from the same Coral Sea stations together with P. longimanus and
the two species of Neopalicus. Depth: 7 m (MlERS, 1884) to 70 m.
Subfamily CROSSOTONOTINAE Moosa & Serene, 1981
Crossotonotinae Moosa & Serene. 1981: 52.
Type Genus. — Crossotonotus A. Milne Edwards, 1873.
DIAGNOSIS. — Fifth pair of pereopods (P5) reduced in size but similar in shape to anterior walking legs (P2-
4); articulation to carapace at same level as walking legs. All borders of carapace with teeth or rounded to pointed
tubercles. Dorsal surface of carapace relatively flat or slightly convex except regions with high bosses or tubercles.
No episternal process overhanging each posterolateral border of carapace posterior to fifth pair of pereopods.
Epistome very narrow, not expanded dorsoventrally (vertically inclined). Abdomen of mature males and females
with all segments free, segments 1-2 wide (not dorsoventrally compressed); transverse ridges complete, incomplete,
or absent.
GENERA included. — Crossotonotus A. Milne Edwards, 1873 and Pleurophricus A. Milne Edwards, 1873.
DISCUSSION. — The subfamily was established by MOOSA & SERENE (1981) to include Crossotonotus
A. Milne Edwards, 1873, Pleurophricus A. Milne Edwards, 1873, and Manella Rathbun, 1906. Manella is here
synonymized with Crossotonotus (see below). Parapleurophrycoides Nobili, 1906, sometimes included in the
Palicidae but "discarded" by MOOSA & SERENE (1981: 52) since the description of its only species was based on a
juvenile specimen, does not belong to the Palicidae at all (see below). The two genera that remain in the subfamily
share, among several characters, a fifth pair of pereopods (P5) that is not greatly reduced in size, is similar in
morphology to the anterior walking legs (P2-4), and is located at the same level as that of the remaining walking
legs, that is, not dorsal to them as in the Palicinae. Other significant differences include the morphologies of the
outline of the carapace and in the abdomen of both sexes.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PAL1CID CRABS
569
Parapleu roph rycoides Nobili, 1906 was described for P. roseus from French Polynesia (see also Nobili. 1907;
Forest & Guinot, 1962). It was questionably included, as Pleurophrycoides [s/c], in the Palicidae by Serene'
(1968: 97) but "discarded" by Moosa & Serene (1981: 52) since the description of its only species "was a
juvenile which can not be used for the definition of the genus". The very small male holotype (1.3 x 1.5 mm;
MNHN-B 13091) is a xanthoid (P.K.L. NG, personal communication) and possibly, as stated by NOBILI (1907:
402), Balss (1957: 1662), and Takeda & SHIMAZAKI (1974: 75), a pilumnid. A very small male specimen from
the Gulf of Thailand identified as P. roseus (1.2 x 1.4 mm; ZMUC) by Rathbun (1910: 358; also see
NAIYANETR, 1980: 43; 1998: 119, as Pleurophry'coides [sic] roseus ) is very close if not identical to NOBILI's
species and therefore not a palicid either.
The shape and placement of the P5 in Crossotonotus and Pleurophricus in contrast with the rest of the palicids
led to their placement in other groups of brachyurans. Crossotonotus was placed between "Ocypodidae and
Plagusiinae" by A. Milne Edwards (1873a: 258) and de Man (1888: 344) included his new species, P. spinipes
(a junior subjective synonym of Crossotonotus compressipes A. Milne Edwards, 1873) among the Corystidae.
A. Milne Edwards (1873a: 260) placed Pleurophricus among the Oxystomata close to the Ocypodidae, while
MlERS (1879: 660) included it among the Oxyrhyncha.
All members of the subfamily appear to be restricted to hard substrates in relatively shallow water. No species
are known from outside the Indo-west Pacific region.
Key to the genera of the subfamily CROSSOTONOTINAE
1 . Carapace subquadrate, with low bosses on dorsal surface and many small ( 10 to more than
20), rounded or pointed tubercles along posterior border .
. . CROSSOTONOTUS A. Milne Edwards, '1873
Carapace subcircular, with two very conspicuous, high bosses on metagastric region and
few (5-6) larger, salient tubercles along posterior border .
. PLEUROPHRICUS A. Milne Edwards, 1873
Genus CROSSOTONOTUS A. Milne Edwards, 1873
Crossotonotus A. Milne Edwards, 1873a: 258 [82], — A. Milne Edwards. 1873b: 282. — Sakai, 1965: 186; 1976: 595
— Takeda & Shimazaki, 1974: 75. — Holthuis, 1977: 187. — Moosa & Serene, 1981: 58.
Crossonotus - HaSWEll, 1882: 95.
Manella Rathbun, 1906: 837. — Moosa & Ser£ne, 1981: 53.
TYPE Species. — Crossotonotus compressipes A. Milne Edwards, 1873 by monotypy. Gender: masculine.
Type species of Manella Rathbun, 1906: Pleurophricus spinipes De Man, 1888 by monotypy.
Diagnosis. — Frontal border of carapace divided into 2-4 lobes (innermost lobes sometimes secondarily
notched). Anterolateral borders each with pointed teeth. Dorsal surface of carapace with small granules and high
granular bosses. Eyes spherical; peduncles with soft tubercles. Supraorbital borders each with two lobes.
Suborbital borders each with two lobes. Each basal antennal segment rectangular but outer side can be slightly
expanded. Epistome very narrow, with conspicuous, triangular median teeth. Chelipeds unequal in males and
females. Walking legs (P2-5) with flattened carpi, propodi, and dactyli. First (P2) and fourth (P5) pairs shorter than
second and third pairs (P3-4). Borders of meri of P2-5 with equal or nearly equal tubercles; carpi, propodi. and
dactyli with teeth along anterior and posterior borders. P5 reduced (0.9- 1.0 CL), thick and flattened. Abdomen of
mature males elongate (sides parallel to each other), with all segments free. Male first pleopods long and slender:
basal parts sinuous; each distal part uniramous (secondary, proximal tip may be present), tip with minute teeth.
Abdomen of mature females with all segments free.
Source :
570
P. CASTRO
REDESCRIPTION. — Carapace subquadrate, slightly broader than long, with low, granular bosses. Anterolateral
borders each with a continuous row of teeth; posterolateral borders with tubercles, straight or slightly rounded,
with marked junction with convex posterior border. Confluence of branchial and mcsogastric regions depressed and
smooth; no conspicuous sulcus between hepatic and branchial regions. Posterior border with many rounded or
pointed tubercles. Thoracic sternite 7 without episternal process visible dorsally.
Frontal border divided into 2-4 pointed or rounded lobes (if four, inner pair more advanced anteriorly than outer
pair, may be secondarily lobed). Supraorbital borders each with two triangular or rounded lobes. Postorbital angles
conspicuous, with long and pointed tips. Cornea of eyes spherical, slightly wider than base of eye peduncle;
peduncle with granular tubercles at least on distal border. Orbits deep.
Suborbital border with two broad, rounded lobes usually bordered by long teeth or pointed tips. Pterygostomial
lobe at each anterolateral angle of buccal frame broad, bordered by long teeth or slightly pointed.
Each basal antennal segment rectangular, with or without small, outward expansion. Epistome very narrow,
not expanded dorsoventrally, vertically inclined. Border of epistome with two triangular, pointed median teeth (plus
two triangular, pointed outer processes) connecting distally with each pterygostomial lobe. Meri of third
maxillipeds smaller and narrower than ischia.
Chelipeds unequal in males and females. First (P2) and fourth (P5) pairs of walking legs shorter than second
and third pairs (P3-4). Walking legs (P2-5) with flattened carpi, propodi, and dactyli; anterior (dorsal) and posterior
(ventral) borders of meri with pointed teeth and rows of tubercles on dorsal and ventral surfaces; anterior and
posterior borders of carpi, propodi. and dactyli with teeth. Second and third pairs of walking legs (P3-4) each with
broad coxae, having flattened, thin-edged anterior and posterior borders.
Abdomen of mature males elongate, with both sides almost parallel to each other, with all segments free, and
no transverse ridges; first segment broad and long. Male first pleopods long and slender; basal parts sinuous; each
distal part uniramous (secondary, proximal tip may be present), tip bordered by minute teeth. Second male
pleopods short, slightly thick, slightly curved; distal segment with blunt tip.
Abdomen of mature females broad, rounded, with all segments free and no transverse ridges; first segment broad
and long. Abdomen of immature females triangular (but broader than in males), with all segments free.
Species INCLUDED. — C. compressipes A. Milne Edwards, 1873, C. spinipes (De Man, 1888). C. ceramensis
(Moosa & Serene, 1981), and C. lophocheir sp. nov.
Sexual Dimorphism. — The largest specimen known for C. spinipes is a male, the only case among Indo-
west Pacific palicids. The chelipeds are unequal in both sexes in C. spinipes but a conspicuous, dense cluster of
plumose setae is found only on the inner surface of the propodi and fingers of male chelipeds. The lower margin of
the large cheliped of males has a series of sinuous ridges, presumably a stridulating surface (Rathbun, 191 I).
DISCUSSION. — Crossotonotus was established by A. MlLNE EDWARDS (1873a) for a new species,
C. compressipes, which was described using specimens from Samoa. The genus was considered not to belong to
any known family of the Catometopa and was placed "between Ocypodidae and Plagusiinae" (A. MlLNE
Edwards, 1873a: 258).
A second species, Pleurophricus spinipes, was described from Ambon, Indonesia by DE Man (1888), who
placed it in Pleurophricus A. Milne Edwards, 1873. Rathbun (1906) identified P. spinipes from the Hawaiian
Islands but created Manella (with P. spinipes De Man as the type species) to recognize the position of the species
as a palicid based on the relative size of its walking legs: "I have separated generically De Man's species from the
type of Pleurophricus on account chiefly of the legs" (Rathbun, 1906: 837). The legs of RATHBUN’s species
(Manella spinipes Rathbun, 1906) were found to be similar to those of other palicids (that is, first and fourth
walking legs smaller than the second and third) whereas the legs of Pleurophricus cristatipes A. Milne Edwards,
1873, originally the only species in the genus, were more similar in size and therefore not considered palicid-like
(Rathbun, 1906). RATHBUN (191 1), Ward (1933), Monod (1938), Sakai (1939), Edmondson (1933, 1946.
1962) and Zarenkov (1968) followed Rathbun (1906) and placed C. compressipes and P. spinipes under
Manella. Three additional species were eventually added: M. gardineri Rathbun. 1911 . M. hrevimana Ward, 1933,
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
571
and M. ceramensis Moosa & Serene, 1981. Sakai (1965, 1974, 1976), however, regarded Manella as a synonym
of Crossotonotus , a view followed by Serene (1968), Holthuis (1977) and McNeill (1968). Moosa &
SERENE (1981: 60) examined the type material of C. compressipes and mistakenly concluded that it was different
from Manella , although they stated that the genera "are close one from another and perhaps identical". They
restricted Crossotonotus to the type material of C. compressipes that was examined (MNHN-B 3078) plus three
additional specimens examined by Ward (1933).
The figure that was published following the description of C. compressipes (A. MlLNE Edwards. 1873b: pi.
13, fig. 1) shows a rounded carapace that lacks a noticeable junction between the posterolateral borders and a
convex posterior border. This character was unfortunately used by MOOSA & SERENE (1981) to separate
Crossotonotus from Manella. In the type material of C. compressipes (MNHN-B 3078), the posterior edge of the
carapace is slightly convex and there are noticeable junctions with each posterolateral border.
Key to the species of CROSSOTONOTUS
1. Carapace subquadrate, depressed; pointed or broad, straight-edged anterolateral teeth . 2
— Carapace subcircular, dorsally convex; rounded anterolateral teeth .
. C. compressipes A. Milne Edwards, 1873
2. Dorsal border of propodus of cheliped with two rows of short tubercles . 3
— Dorsal border of propodus of cheliped with one large, rounded tubercle .
. C. lophocheir sp. nov.
3. Frontal lobes broad with tips bent upward or rounded (in smaller individuals). Inner surface
of cheliped propodi (distal portions) and fingers (proximal borders) of males with dense
clusters of plumose setae . C. spinipes (De Man, 1888)
Frontal lobes broadly rounded and only slightly protuberant. Inner surface of cheliped
propodi of males without clusters of setae . C. ceramensis (Moosa & Serene, 1981)
Crossotonotus compressipes A. Milne Edwards, 1873
Figs 44, 51
Crossotonotus compressipes A. Milne Edwards, 1873a: 259 [83J. — A. Milne Edwards. 1873b: 283, pi. 13. figs 1. la-f.
— Ward, 1933: 389. — Sakai, 1965: 186. — McNeill, 1968:81. — Serene, 1968:96. — Moosa & Serene. 1981:
60. — Fransen et al.. 1997: 152. — Muraoka, 1998: 49.
Crossonotus [sic] compressipes - Haswell, 1882: 96.
Crossotonotus taketomiensis Sakai, 1974: 94; 1976: 596, fig. 326.
Crosstonotus [sic] taketomiensis - Nagai & Nomura, 1988: 197, color photograph.
MATERIAL EXAMINED. — Japan. Ishigaki Islands, Taketomi Island. Taketomi Island. Ishigaki Islands, Ryukyu
Islands, T. SAKAI coll., 25.10.1973: 1 juv. <3 holotype of Crossotonotus taketomiensis Sakai. 1974, 5.7 x 6 1 mm
(SMF 24716).
No locality. 1 6 (SMF 24717).
Indonesia. Moluccas. Rumphius Expedition I: stn Li-1, Ambon, 03°36'06-,S. 1 28°14'58"E. Kasdan coll
15.01.1973: 1 9 (MNHN-B 10014).
Papua New Guinea. Bismark Archipelago. New Britain Island. Ralum, coral reef. S. Dahl coll 4 09 1986' 1 2
(ZMB 18167).
Australia. Queensland. Hope Islands. 12°28'S. 145°15'E, A.R. McCulloch coll.. 1905. 1 <S (AM P3740).
Low Isles. Great Barrier Reef Expedition: 1 9 9.5 x 10.5 mm (BMNH 1950.12.1.28).
New Caledonia. Noumea, baie des Citrons, under rock. 1 m, G. Paulay coll., 18.05.1999: 1 2 (MNHN-B 27138).
Unknown locations. M. Balansa coll.: 1 2 7.0 x 7.3 mm (MNHN-B 26272). — BOUGRIN ? coll 1903' I 2 (MNHN-
B 13251).
Samoa. Upolu. 1 2 lectotype 8.4 x 9.5 mm, 1 2 paralcctotype 6.4 x 7.4 mm (MNHN-B 3078). — 2 2 6.7 x
8.0 mm, 7.3 x 8.7 mm (SMF 1747). — 2 2 6.7 x 7.1 mm, 7.0 x 7.4 mm (ZMUC CRU-3623).
Source :
572
P CASTRO
Types. — Lectotype: 1 $ 8.4 x 9.5 mm, Samoa (MNHN-B 3078).
Paralectotypes: I $ 6.4 x 7.4 mm, Samoa (MNHN-B 3078).
Holotype of Crossolonotus taketomiensis Sakai, 1974: 1 6 5.7 x 6.1 mm, Taketomi Island, Ishigaki Islands,
Ryukyu Islands, T. SAKAI coll., 25.10.1973 (SMF 24716).
Type Locality. — Upolu, Samoa, unknown depth.
DIAGNOSIS. — Carapace (Fig. 44a) subcircular, dorsoventrally convex, with large, low bosses. Five to six
broad, rounded anterolateral teeth on each side of carapace, followed by 3-4 slightly smaller, rounded tubercles
along each posterolateral border; 9-14 rounded tubercles along convex posterior border separated from posterolateral
border by clear junction. Supraorbital borders each with two short, slightly rounded lobes. Four rounded to slightly
pointed lobes along anterior border; median lobes slightly broader and longer than outer lobe. Suborbital borders
(Fig. 44b) each with two pointed lobes, proximal bifurcated. Dorsal borders of cheliped propodi each with two
rows of rounded tubercles; lower border of propodus of largest cheliped straight. Dactyli of P2-4 with teeth along
anterior and posterior borders (two on posterior borders of P4); dactyli of P5 (Fig. 44c) with 3-4 spines along
posterior borders. Abdomen of mature males and females with all segments free (A. Milne Edwards, 1873b,
pi. 13, figs le, 10- Male first pleopods (Figs 44d-e) with sinuous basal parts; each distal part straight, one simple
tip bordered by conspicuous teeth.
Color: A color photograph in NAGAl and NOMURA (1988: 197) identified as C. taketomiensis, a junior
synonym of C. compressipes, shows an individual partially camouflaged against a hard substrate in an Okinawa
coral reef. The carapace of the specimen is pinkish brown, with a bright yellow spot on the cardiac region Hanked
on each side by a greenish spot. The legs and clumps of setae across the dorsal surface of the carapace are gray.
Discussion. — C. compressipes was actually described twice by A. Milne Edwards (1873a, 1873b), with
both publications appearing in the same year. Unfortunately, it has not been possible to obtain information on
which of the two was first published. The descriptions are identical except that in one of the two (A. MlLNE
Edwards, 1873b: 284) there is information on additional material from New Caledonia, which leads to belief that
it was written after the intended description (A. MlLNE Edwards, 1873a).
Two dry female specimens from Samoa that were originally deposited at the Godeffroy Museum. Hamburg and
the source of specimens used in the description, were designated by Moosa & Serene (1981) as the "holotype"
and "paratype" of C. compressipes. Since A. Milne Edwards had not selected a holotype, the two specimens
were actually syntypes. The selection of a "holotype" and a "paratype" among the syntypes by Moosa & Serene
was incorrect and therefore the two selected specimens are a lectotype and a paralectolype respectively, not a
holotype (International Code of Nomenclature, fourth edition. 1999. Article 74.1). Both specimens are small
(lectotype 8.4 x 9.5 mm, paralectotype 6.4 x 7.4 mm; MNHN-B 3078). A photograph of one of the specimens of
the type material, perhaps the lectotype, is probably shown in a photograph in MOOSA & SERENE (1981, pi. 3.
fig. D). They are similar to the measurements given by A. Milne Edwards (1873a: 259 [83]) in his description
(8x9 mm). The number of lateral teeth given in the description (10-12) agree with the number observed in the
lectotype and paralectotype but not in the case of the posterior teeth: 12 in the specimen against 6-8 in the
description. Some of the posterior teeth arc rounded and broad, while others are small and pointed and perhaps they
were not taken into account. The posterior border of the paralectotype is damaged. The carapace of a third dry
specimen from New Caledonia presumably examined by A. Milne Edwards (MNHN-B 26272) is in poor
condition. The figure given by A. MlLNE EDWARDS (1873b, pi. 13, fig. 1 ) shows a rounded carapace bordered by
similar rounded teeth, of which 12 may be interpreted as lateral. The number of posterior teeth cannot be
determined.
Two alcohol-preserved females from Samoa (6.8 x 7.3 mm, 8.0 x 8.7 mm; SMF 1747), originally at the
Godeffroy Museum as the type material, have eight lateral teeth on the left side and nine on the right. The largest
specimen has 14 uneven posterior teeth and the smallest 12, also uneven. Two other alcohol-preserved females
Irom Samoa that had also been deposited at the Godeffroy Museum (6.7 x 7.1 mm, 7.0 x 7.4 mm; ZMUC CRU-
3623). The other specimens examined are similar to the type material and the remaining Godeffroy Museum
specimens. A small female from New Caledonia (6.9 x 7.2 mm; MNHN-B 13251) had 10 lateral and nine
Source : MNHN, Paris
1NDO-WEST PACIFIC SPECIES OF PALICID CRABS
573
posterior teeth and tubercles, all rounded. A female specimen from the Great Barrier Reef (9.5 x 10.5 mm; BMNH
1950.12.1.28) had similarly shaped teeth and tubercles (nine lateral, 12 posterior).
Fig. 44. — Crossotonotus compressipes A. Milne Edwards. 1873: a-e. 9 7.3 x 8.7 mm, Upolu. Samoa (SMF 1747):
a. carapace, dorsal view; b. suborbital border; c, dactylus, right fifth pereopod. dorsal view. — d-e. <3 9.9 x 10.9
mm. Hope Islands. Queensland, Australia (AM P3740): d. left male first pleopod. lateral view (outer side) view; e. left
male first pleopod. distal end (inner side).
According to L.B. HOLTHUIS (in litt.), the Godeffroy Museum offered specimens of C. compressipes for sale
(Museum Godeffroy, Catalog V. 1874, p. 75). Subsequent issues of their catalogues (VI-IX. 1877-1884 ) did not
list any specimens so specimens were probably sold, which explains their presence in Frankfort (SMF 1747),
Copenhagen (ZMUC CRU-3623), and Berlin (ZMB 4463, lost).
C. taketomiensis was described by Sakai (1974) from two small specimens collected in the southern Ryukyu
Islands. The species was distinguished from C. compressipes mostly by the shape of the teeth along the carapace
borders. The holotype of C. taketomiensis (male. 5.7 x 6.1 mm; SMF 24716), however, is almost identical to the
lectotype of C. compressipes. Sakai (1974: 95; 1976: 597) stated that the teeth on the anterior border of
Source : MNHN. Paris
574
P. CASTRO
C. taketomiensis were rounded, in contrast to those of C. compressipes. A. MlLNE Edwards, however, described
them as rounded and they are shown as such in the figure that accompanied the description (A. MlLNE Edwards,
1873b, pi. 13, fig. 1). Although the number of teeth along the rest of the carapace borders were found to be similar
in both species, Sakai (1974: 95; 1976: 597) characterized them as "variable in size and their lips obtusely
pointed" in contrast to the rounded, similar teeth shown in A. MlLNE Edwards' figure. Sakai, nevertheless,
mistakenly used the shape of the first anterolateral tooth ("very prominent and acuminate") to contrast between the
two species. It is the second tooth, however, that is the largest of the three anterolateral teeth in (lie holotype of
C. taketomiensis and the lectotype of C. compressipes. The serration of the anterior borders of (lie meri of the
walking legs is a characteristic of the species found in both type specimens, although Sakai noted to be entire in
C. compressipes, obviously based only on A. MlLNE EDWARDS' figure. Sakai (1974: 95; 1976: 597) also used
the presence of plumose setae on the chelipeds of C. compressipes (in contrast to their absence in
C. taketomiensis). Although A. Milne Edwards (1873b: 284) mentions that setae were present on the inner
surface of the small chelipeds ("poils sur sa face interne"), no dense clusters of setae were observed in specimens of
C. compressipes, as is the case of the male of C. spinipes.
Size. — Maximum size among specimens examined: 9.5 x 10.5 mm (female, BMNH 1950.12.1.28): 9.9 x
10.9 mm (male, AM P3740).
Distribution. — Known from Japan (Sakai, 1974, 1976, both as C. taketomiensis). Solomon Islands
(Ward, 1933), Queensland. Australia (Haswell, 1882; McNeill, 1968), New Caledonia (A. MlLNE Edw-ards,
1873b), and Samoa (A. MlLNE Edwards, 1873a, 1873b). Its range is now extended to Ambon (Indonesia) and the
Bismark Archipelago (Papua New Guinea) (Fig. 51). It appears to have been collected only in shallow waier: "reef
flat at low tide" (MCNEILL, 1968: 81) and "coral reef' (Sakai, 1974: 95).
Crossotonotus spinipes (De Man, 1888)
Figs 45, 46, 51. 61 f
Pleurophricus spinipes De Man. 1888: 344, pi. 15, figs 1, la-c.
Manella spinipes - Rathbun, 1906: 837, fig. 3, pi. 7, fig. 6. — Edmondson, 1933: 268. fig. 158b; 1946: 310.
fig. 184b; 1962: 12, figs 2c, 4f. — Ward, 1933: 389. — Monod, 1938: 153, figs 26D-E, 29 (as "IManella
spinipes"). — Sakai. 1939: 610, pi. 103, fig. 3; 1956: 52. — Guinot, 1967: 280. — Zarenkov, 1968: 762. —
Yang, 1979: 34.
Manella gardineri Rathbun, 1911: 240, pi. 20, fig. 9. — Ward. 1933: 389. — Guinot. 1967: 280.
Manella brevimana Ward. 1933: 387, pi. 21. figs 7-8. — Moosa & Serene. 1981: 54, figs 12 a-b. 13 a.
Crossotonotus spinipes - Sakai, 1965: 187, fig. 25, pi. 89, fig. 4; 1976: 596, pi. 206, fig. 1. — Chen, 1975: 167.
fig. 9, pi. 2, fig. 3. — Holthuis, 1977: 187. — Dai et al., 1986: 414, fig. 229 A. — Dai & Yang, 1991: 450.
fig, 229A. — Wada, 1995: 418. pi. 118, fig. 11.
Crossotonotus gardineri - SERENE, 1968: 97.
Crossotonotus brevimanus - Serene, 1968: 97.
Crossotonotus sp. - Takeda & Shimazaki. 1974: 75, pi. 2. fig. B.
Material EXAMINED. — Red Sea. Al Sayad. Stn 23. R. DOLLFUS coll., 30.12.1928: 1 9 (MNHN-B 13255).
Port Sudan. Wingate Reef, stn SAN-166. V. Neumann coll., 5 m, on dead corals, 21.09.1992: 2 9 (SMF 24541). —
Sanganeb Atoll, stn SAN-120, V. Neumann coll., 6-10 m. on dead corals, 3.10.1992: 1 9 (SMF 24542).
Seychelles. Reves 2: stn 47, 04°03.8'S, 55°59.5'E, 45-55 m. shelly sand. 14.09.1980: 1 <3 (MNHN-B 27131).
Amirante Islands. "Sealark": stn E21. 55 m, 17.10.1905: 1 juv. 3 8.7 x 10.0 mm. 1 3 10.1 x 12.1 mm,
holotype of Manella gardineri Rathbun, 1911 (USNM 41364).
La Reunion. M. Peyrot-Clausade coll.: 1 juv. (MNHN-B 26275).
Sri Lanka. Trincomalee, south side of first bay north of harbor, 26 m, C. Koenig coll., 5.04.1970: 1 9 (USNM).
Japan. Boso Peninsula. Katsuura. Rocky shore intertidal, A. Asakura coll., 23.08.1995: 1 3 (CBM-ZC 1931 ).
Miura Peninsula. Washed up at Nahama beach, H. Ikeda coll., 05.1995: 1 9 (HSM).
Shimoda ?: 1 9 (SMF 2471 1).
Wagtt. 2 9 (SMF 24712). — From lobster nets, 10-15 m, H. Ikeda coll., 04.1980: 1 3 (HSM).
Kii Nagashima. Yamamoto coll., 12.1963: 1 3 (SMF 24713).
Source : MNHN . Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS 575
Ryukyu Islands. Okinawa, off Horseshoe Cliffs, 26°30.0'N, 127°50.9'E, 57.9 m, R.F. Bolland coll. 1 8 04 1981-
I 2 (USNM).
Miyako Islands. Irabujima Island, 12 m, in cave, S. Nakatani coll., 17.07.1999: 1 2 31.5 x 35.8 mm (MNHN-
B 26802).
Ishigaki Islands. Ohama, 27.08.1977: 1 <3 (SMF 24714).
No locality. 1 juv. 2 (SMF 24715).
Taiwan. Lan Yu (= Orchid) Island. Night diving, 5 m, S.-H. WU coll., 8.07.1997: 1 juv. 2 26.6 x 30.0 mm (NTOU)
Viet Nam. Nhatrang, R. SerEne coll., 13.08.1964: 1 3 (MNHN-B 9765).
Singapore. Sultan Shoal, 18-24 m, 12.1933: 1 2 (ZRC 1965.8.2.234).
Indonesia. Timor. SlBOGA Expedition: stn 303, Samau Island. 10°27.9'S, 123°28.7'E, 36 m. Litlwtliamnion
2-5.02.1900: 1 2 (ZMA De 203890).
Papua New Guinea. Hansa Bay. Laing Island lagoon, 4°11.62'S. 144°51.46'E. 5 m, in Pocilldpora damicornis
colony, S. De Grave coll., 25.10.1993: 1 juv. 2 (MNHN-B 27031).
Capricorn Croup. North West Island, 23°!8'S, 151°22’E, under large spreading coral in pool, outer edge of reef.
M. Ward coll., 12.1929: 1 3 holotype of Manella brevimana Ward, 1933 14.1 x 16.5 mm (AM P10636).
Chesterfield Islands - Chesterfield-Bellona Plateau. Chalcal I: stn D 51. 21°13 21'S 1 S8°4^ 50'E
55 m, 07.1984: 1 3 juv. (MNHN-B 27129).
Corail 2: stn DW 104. 19°08.95'S, 158°35.67'E. 49 m, boulders, 27.08.1988: I 3 (MNHN-B 27098). —
Stn CP 127, 19°27.73'S, 158°27.30'E, 44-45 m, boulders and corals, 29.08.1988: 1 2 (MNHN-B 27119).
New Caledonia. Lagon: stn 427, 22°41.9'S, 167°18.0'E. 60 m, boulders and corals. 25.01.1985: 1 3 (MNHN-B
27127). — Stn 428, 22°40.7'S, 167°16.4'E, 56 m. Halimeda sand, 25.01.1985: 1 juv. 2 (MNHN-B 27094). — Sin 477.
18°5 1.0'S, 163°27.0'E, 50 m, Halimeda sand and boulders, 2.03.1985: 1 2 (MNHN-B 27128). — Stn 606. 22°I2.8'S.
167°00.5'E, 46-48 m, corals, 5.08.1986: 1 3 , 1 2 (MNHN-B 27122). — Stn 625, 21°59.2'S, 1 66°53.6 E. 34-40 m
Halimeda sand and corals, 6.08.1986: 1 <3 (MNHN-B 27096). — Stn 635, 21°57.7'S. 166°44.5'E, 52-45 m. boulders and
corals. 6.08.1986: 1 juv. 2 (MNHN-B 27095). — Stn 640, 21°54.8'S, I66°45.8'E. 50-80 m, Halimeda sand. 7.08.1986:
1 2 (MNHN-B 27130). — Stn 650, 21°49.3'S, I66°37.7'E. 50 m, boulders and Halimeda. 7.06.1986: 1 2 (MNHN-B
27123). — Stn 657. 21°48.2'S, 166°33.7'E, 40-42 m. Halimeda sand. 8.08.1986: 1 2 (MNHN-B 27100). — Stn 659.
21°45.3'S, 166°33.4'E, 46-48 m. boulders and corals. 8.08.1986: 1 <J (MNHN-B 27120). — Stn 662. 21°44.0'S.
166°32.0'E, 50 m, boulders and corals, 8.08.1986: 1 3 10.6 x 12.8 mm (MNHN-B 26810), I 3 (MNHN-B 26215). —
Stn 668, 21 o40.5'S, 166°29.1’E, 40 m. boulders and corals. 8.08.1986: 1 3 (MNHN-B 27097). — Stn 769 "* 1 0 1 2 0'S
1 65°40.2'E, 39 m, coarse sand, 8.01.1987: 1 3 (MNHN-B 27099).
Expedition Montrouzier: Touho, 09.1993: 1 juv. 3 (MNHN-B 27125).
Recif Mbere. 22°19.9'S. 166°13.2'E, outer reef slope, 25-30 m. P. Bouchet coll.. 5.05.1993: 1 juv. 2 (MNHN-B
27121). — 7.01.1993: 1 3 , 1 juv. 3 , 1 juv. 2 (MNHN-B 27126).
Passe de Dumbea. Outer reef, under coral ridge, night diving, 12 m, 8.01.1999: 1 3 36.9 x 43.3 mm (MNHN-B
26803).
Norfolk Ridge. Smib 5: stn DW 100, 23°22.90’S, 168°05.20'E, 80-120 m, sand. 14.09.1989: 1 3 (MNHN-B 27124).
Unknown location. — Night diving, 20 m, P. Laboute coll., 19.11.1988: 1 2 (MNHN-B 26811).
Fiji. Great Astrolabe Reef, off north tip of Dravuni Island, stn WE 83-27. poison station. 10-15 m, 29.03.1983: 1 3
(USNM).
Types. — Holotype of Pleitropliricus spinipes De Man. 1888: 1 juv. 3 7.0 x 8.5 mm (De Man, 1888),
J. BROCK coll., Ambon. Indonesia. Deposit unknown (material lost ?).
Holotype of Manella gardineri Rathbun, 191 I: Amirante Islands, western Indian Ocean, "Sealark". stn E2I.
55 m, 17.10.1905: 1 6 10.1 x 12.1 mm (USNM 41364).
Holotype of Manella brevimana Ward, 1933: 1 S 14.1 x 16.5 mm, North West Island. Capricorn Group,
Queensland, Australia, 23°18'S, 151°22'E, under large spreading coral in pool, outer edge of reef. M. Ward coll..
12.1929 (AM PI 0636).
TYPE LOCALITY. — Ambon. Moluccas, Indonesia, unknown depth.
DIAGNOSIS. — Carapace (Figs 45a, 46, 610 subquadrate, depressed, with large, slightly swollen areas with low
bosses topped by rounded tubercles. Three broad, pointed anterolateral teeth (second typically bifurcated) on each
side of carapace, followed by 3-4 large, pointed or blunt tubercles along each lateral border. 4-6 similar but slightly
smaller, pointed tubercles along each posterolateral border; 17-24 pointed or blunt tubercles along convex posterior
border separated from posterolateral border by clear junction. Supraorbital borders each with two triangular lobes
(sometimes bifurcated). Four lobes along anterior border; tips bent upward, median lobes broader and notched with
increasing carapace size (lobes rounded, median lobes slightly longer than outer lobes, and with entire borders in
Source : MNHN. Paris
576
P. CASTRO
very small specimens). Suborbital borders (Fig. 45b) each with two broad, rectangular lobes bordered by long
narrow teeth. Dorsal borders of chcliped propodi each with two high and conspicuous rows of pointed to rounded
tubercles; cluster of abundant plumose setae on inner, distal surface of propodi and proximal border of lingers in
males (less abundant, shorter setae in largest females); propodus of largest cheliped very high, with lower border
expanded to form J-shaped proximal edge. Dactyli of P2-4 with teeth along anterior and posterior borders three on
posterior borders of P4); dactyli of P5 (Fig. 45c) with 2-3 spines along posterior borders. Abdomen of mature
males (Fig. 45d) with all segments free (all fused in juveniles). Male first pleopods (Fig. 45c, Moosa & Sere, e.
1981, figs 1 2a-b) with sinuous basal parts; each distal part straight, one simple tip bordered by minute teeth.
Color. Orange carapace and legs (Fig. 610- A light spot on the cardiac region was observed in two specimens
from New Caledonia. Red spots flanked the yellow spot of one.
Fig. 45. - Cwssoionotus spinipes (De Man, 1888), 5 10.6 x 12.8 mm, east coast of New Caledonia, Lagon, sin 662,
50 m (MNHN-B 26810): a. carapace, dorsal view: b, suborbital border; c, dactylus, right fifth pereopod, dorsal view,
d. abdomen; e, left male first pleopod, lateral (inner side) view.
DISCUSSION. — The chelipeds of C. spinipes are very unequal in both sexes, the largest having a much thicker
and higher propodus. Each finger of the largest cheliped has one cutting edge plus broad teeth in the largest speci¬
mens. The fingers of the smaller cheliped has spoon-shaped tips and short, rounded teeth along the distal halt. In
males there is a dense cluster of plumose setae at the distal end of the propodus and proximal portion of the linger
on the inner surface of both chelipeds. Shorter and less dense setae, however, are found in the largest females.
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
577
The lower border of the propodus of the largest cheliped has low undulating ridges that were referred to as
"stridulating lines" by Rathbun (191 1: 241). Their function remains unknown. Males do not seem to have a
specialized plectrum, the structure rubbed by the cheliped to emit sound. Two possibilities for the plectrum arc the
teeth along the dorsal border of the second walking leg or the tubercles on the dorsal border of the small cheliped.
Both of these structures are nevertheless also found in females.
The lourth pair of walking legs (P5) is identical in shape to the other walking legs, in sharp contrast to the
members of the subfamily Palicinac where the segments are much more slender. The P5 of C. spinipes have the
same relative width and spinulation of the remaining three pairs of walking legs. Nevertheless, they are slightly
shorter than the first pair (P2) and their relative length (0.9 CL) is the same or similar as in many other species of
the subfamily Palicinae. It appears that the widening of the P5 is an adaptation to crawling on hard substrates.
Nevertheless, at least one species of the subfamily Palicinae ( Exopalicus maculatus ) has a similar habitat but it
shares slender P5 with the other members of the subfamily.
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
The maximum size of juvenile females varies widely so that size alone does seem to determine when maturity
is reached. A female as large as 26.6 x 30.0 mm (NTOU) had the narrow abdomen characteristic of juvenile
females.
DE Man (1888) described Pleurophricus
spinipes from Ambon, Indonesia. The holotype. a
"juvenile male" that was collected by J. Brock and
deposited at the Gottingen Zoological Museum
under registration number 761. is apparently lost. It
is not in Frankfurt (SMF), where the collection of
the Gottingen Zoological Museum is now
deposited, nor in either Amsterdam (ZMA) or
Leiden (RMNH), where some of Brock's material
studied by DE Man was deposited (Fransen ei al.,
1997: 218).
The abundant material examined agrees with the
detailed description and the illustration given of the
holotype given by De Man (1888). The antero¬
lateral teeth of the holotype, however, appear much
too thin and pointed but only on the left side of the
carapace (De Man, 1888, pi. 15. fig. I): the
anterior border of the carapace has two dissimilar
lobes (De Man, 1888, pi. 15. figs 1-la).
Manella gardineri , described by RATHBUN (191 1) from five specimens collected from the western Indian Ocean
(male holotype from the Amirante Islands, 10.1 x 12.1 mm; USNM 41364), is a junior subjective synonym of
C. spinipes. RATHBUN (1911: 241) distinguished M. gardineri from C. spinipes by citing four characters:
"carapace wider, surface less hairy, projections less spinous, presence of a distinct stridulating mechanism".
Rathbun obviously based at least part of the comparison on De Man's illustration, which shows abnormally
thin anterolateral teeth but only on the left side of the carapace (see above). The "stridulating lines", the thin,
sinuous ridges observed by RATHBUN, are also evident in C. spinipes. One difference in the holotype is that its
lateral and posterolateral tubercles are thicker and rounder than usual. This feature, however, has been found in
specimens of C. spinipes from other Indo-west Pacific locations.
Fig. 46. — Crossotonolus spinipes (De Man, 1988): $ 31.5
x 35.8 mm, Ryukyu Islands, Japan, 12 m, S. Nikatani
coll. (MNHN-B 26802): dorsal view.
Source : MNHN. Paris
578
P. CASTRO
Manella brevimana was described by Ward (1933) from a relatively large male (14.1 x 16.5 mm; AM
PI 0636) from Queensland, Australia. The specimen unquestionably belongs to C. spinipes. Ward compared his
new species with M. gardineri Rathbun and P. spinipes (as M. spinipes) but the differences noted were clearly the
result of individual variation, such as the relative length and shape of the teeth, spines, and granules of the
carapace, and the length and ornamentation of the chelipeds. The species was named "brevimana" because the chela
was "not as long as the carapace", whereas that of M. gardineri was longer, "about 1 1/2 times as long as carapace"
(Rathbun, 191 1; 241). The total length of the propodus and dactylus of the largest cheliped of the holotype of
M. brevimana is the same (0.6 CL) as in the holotype of M. gardineri (USNM 41364); the total length of the
cheliped (merus, carpus, propodus, and dactylus) is 1.0 CL in the holotype of M. brevimana against 1.3 CL in the
holotype of M. gardineri. Rathbun's "length of the chela" measurement certainly referred to cheliped length,
which was misinterpreted by Ward.
The abdomen of the male holotype of M. brevimana was described as having fused segments, but the presence
of separate segments is clearly visible in the fragment still extant as well as in Ward’s figure (Ward, 1933: 389,
pi. 21, fig. 8). One significant difference is that there is no evidence of "stridulating ridges" in the large cheliped of
the holotype, as mentioned in the description. Such a structure was also reported as absent in three males from
Palau (= Belau) (Takeda & Shimazaki, 1974: 79).
A small (5.6 x 6.3 mm) male from Ceram, Indonesia, most probably a juvenile since segments 3-6 of the
abdomen were fused, was identified by MOOSA & Serene as M. brevimana. It could not be examined but it
almost certainly represents C. spinipes on account of the illustrations, particularly of the male first pleopod
(Moosa & Serene, 1981: 54. figs 12 a-b).
Size. — Maximum size among specimens examined: 31.5 x 35.8 mm (female, MNHN-B 26802); 36.9 x
43.3 mm (male, MNHN-B 26803).
DISTRIBUTION. — A very wide distribution throughout the Indo-west Pacific region, the widest known so far in
a palicid: Red Sea (Monod, 1938; Zarenkov, 1968); Seychelles, Amirante Islands, and Saya de Malha Bank
(Rathbun, 191 I, as M. gardineri ); Japan (Sakai, 1939, 1956, 1965, 1976); South China Sea (Chen. 1975:
Dai at al. , 1986; Dai & Yang, 1991); Ceram (MOOSA & SERENE, 1981. as M. brevimana) and Ambon
(De Man, 1888), Indonesia; Queensland, Australia (Ward, 1933, as M. brevimana)-. Palau (= Belau) (Takeda &
Shimazaki. 1974); Samoa (A. Milne Edwards, 1873a, 1873b); Hawaiian Islands (Rathbun, 1906:
Edmondson, 1933, 1946, 1962). Its range is now extended to La Reunion, Sri Lanka, Cocos (Keeling) Islands,
Western Australia, Viet Nam, Singapore, Timor, Papua New Guinea, the Coral Sea (New Caledonia, Chesterfield
Islands, and the Chesterfield-Bcllona Plateau), and Fiji (Fig. 51). It seems to be restricted to hard substrates. Of the
New Caledonia stations where the type of substrate was recorded, 13 of the 17 stations where the species was
collected consisted at least in part of hard bottoms such as boulders and coral. Depth: "intertidal" (Sakai. 1974:
94) and "reef flat at low tide" (MCNEILL, 1968: 81) to 146 m (RATHBUN, 1911).
Crossotonotus ceramensis ( Moosa & Serene, 1981)
Manella ceramensis Moosa & Serene, 1981: 54. figs 12 c-d, 13 b, pi. 3, fig. C.
Material EXAMINED. — Indonesia. Ceram. Mariel King Memorial Expedition: stn CP 1/8-14, Piru Bay,
03°15'S, 128°8'E, coarse sand and rubble, 26-48 m, 2.06.1970: 1 3 holotype 5.7 x 6.5 mm (RDC CB2735).
Types. — Holotype of Manella ceramensis Moosa & Serene, 1981: 1 S 5.7 x 6.5 mm. Mariel King
Memorial Expedition, stn CP 1/8-14 (RDC CB2735).
Paralyse: 1 $ 7.7 x 8.5 mm (Moosa & SerEne, 1981). Piru Bay, Ceram, 3°15’S, 128°8'E, coarse sand and
rubble. 26-48 m, Mariel King Memorial Expedition, stn CP 1/8-14, 2.06.1970 (RDC CB2737).
Type Locality. — Piru Bay, Ceram, Indonesia, 3°15'S, 128°8'E, coarse sand and rubble, 26-48 m.
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PAL1C1D CRABS
579
Diagnosis. — Carapace (Moosa & Ser£ne, 1981, fig. 13a, pi. 3, fig. h) subquadrate, depressed, with large,
slightly swollen areas with low bosses topped by rounded tubercles. Three very broad, straight-edged anterolateral
teeth (second divided into separate portions) on each side of carapace, followed by three slightly pointed tubercles
along each lateral border, 2-3 smaller, blunt tubercles along each posterolateral border; 15-17 rounded tubercles
along convex posterior border separated from posterolateral border by clear junction. Supraorbital borders each with
two broad, rectangular lobes with irregular borders. Four lobes along anterior border; lobes broadly round, nearly
equal in size, and entire borders. Suborbital borders each with two broad, rectangular lobes bordered by long,
narrow teeth. Dorsal borders of chcliped propodi each with two high and conspicuous rows of pointed to rounded
tubercles; distal, inner surface of propodi and proximal border of fingers smooth in male holotype; propodus of
largest cheliped very high, with lower border expanded to form J-shaped proximal edge. Dactyli of P2-4 with teeth
along anterior and posterior borders; dactyli of P5 with 2-3 spines along posterior borders. Abdomen of mature
males with all segments free. Male first pleopods (Moosa & Ser£ne, 1981, figs 12c-d) with sinuous basal parts;
each distal part straight, one simple tip bordered by minute spinules. Abdomen of mature females with all
segments free.
DISCUSSION. — Manella ceramensis was described by Moosa & Serene (1981) from a small male and a
slightly larger female from Ceram, Indonesia. It was differentiated from M. spinipes and M. brevimana (a junior
subjective synonym of Crossotonotus spinipes ) by several differences: the presence of a frontal border that is
"entire, and with obscure or very shallow median notch", wider anterolateral teeth, rounder anterolateral border,
straight posterior border with a smaller number of rounded tubercles ("teeth or lobes"), abdomen of male with
distinct segments, and a different morphology of the male first pleopod (Moosa & Serene, 1981: 57). The frontal
lobes are broad and rounded in the holotype (5.7 x 6.5 mm; RDC CB2735), not short and almost straight, and the
posterior border is slightly convex with protuberant tubercles, not almost straight, as shown in the figure of
Moosa & SerEne (1981, fig. 13b). A slightly convex border is nevertheless shown in the photograph of the
holotype (Moosa & Serene, 1981, pi. 3, fig. C). The use of the presence of distinct segments in the male
abdomen as a difference rose from the mistaken belief that the abdomen of M. brevimana had fused segments (see
discussion of C. spinipes). The differences in the male first abdomen (tip entire, small pointed "spinules" instead
of distinctive spines; see Moosa & Serene, 1981, figs 12 c-d), may be the result of the small size of the holotype.
Although drawn as having a single pointed tip, there is a small but distinctive second lobe as in other species of
Crossotonotus. One difference is that in the male holotype the inner surface of the propodi and fingers of the
chelipeds only have a few thick setae instead of the thick cluster of plumose setae in the males of C. spinipes.
This may also be a consequence of size.
The unique frontal and anterolateral borders of the holotype are distinctive enough to support considering
C. ceramensis as different from C. compressipes and C. spinipes. Nevertheless, the presumed parallel occurrence
of these characters in a female collected from the same location (the female paratype. which could not be examined)
and their absence in the many other specimens of C. spinipes that were examined in this study does not support
the possibility of individual variation. The examination of the paratype as well as of additional material may
provide additional evidence.
SIZE. — Size of the only two specimens known: 5.7 x 6.5 mm (male holotype, RDC CB2735). 7.7 x
8.5 mm (female paratype, Moosa & Serene, 1981).
Distribution. — Known only from Ceram, Indonesia (MOOSA & SERENE, 1981). Depth: 26-48 m.
Crossotonotus lophocheir sp. nov.
Figs 47
Material EXAMINED. — New Caledonia. Lagon: stn 692. 21°32.0'S, 1 66° 1 2.3'E. boulders and corals.
44-48 m. 9.08.1986: 1 2 paratype 6.0 x 6.8 mm (MNHN-B 26736).
Source : MNHN. Paris
580
P. CASTRO
Expedition Montrouzier: Touho Bank, 20°44.6'S. 165°13.7'E. suction pump, hard substrate, 15 m, 31.08.1993: 1 3
holotype 4.2 x 5.1 mm (MNHN-B 26733). — Koe Reef, 20°47.7'S. 165°15.7'E. diving, 10 m, 8.09.1993: 1 juv, 2
paratype 5.4 x 6.1 mm (MNHN-B 26734), 1 2 paratype 5.5 x 6.2 mm (MNHN-B 26735).
Types. — Holotype : 1 S 4.2 x 5.1 mm, EXPEDITION Montrouzier (MNHN-B 26733).
Paratypes: I 9 6.0 x 6.8 mm, Lagon, stn 692 (MNHN-B 26736); 1 juv. 2 5.4 x 6.1 mm. Expedition
Montrouzier (MNHN-B 26734); 1 9 5.5 x 6.2 mm, Expedition Montrouzier (MNHN-B 26735).
Type Locality. — Touho Bank, New Caledonia, 20°44.6'S, 165°13.7’E. 15 m, hard substrate.
DIAGNOSIS. — Carapace (Fig. 47a) subquadrate, depressed, with large, slightly swollen areas with low bosses.
Three broad, rounded or pointed anterolateral teeth on each side of carapace, 3-4 small, blunt tubercles along each
lateral border, 1-3 similar but slightly smaller tubercles along each posterolateral border; convex posterior border
separated from posterolateral border by clear junction, with 10-14 small, blunt or slightly pointed tubercles. Four
lobes along anterior border, inner pointed, outer lobed. Supraorbital borders each with two slightly pointed lobes.
Suborbital borders (Fig. 47b) each with two lobes bifurcated into pointed tips. Dorsal borders of cheliped propodi
each with one conspicuous, rounded tubercle. Dactyli of P2-5 with teeth or spines along anterior and posterior
borders (2-4 on posterior borders of each P4 and P5; Fig. 47c). Abdomen of mature males with all segments free.
Male first pleopods (Figs 47d-e) with sinuous basal parts; each distal part straight, two unequal tips bordered by
spines. Abdomen of mature females with all segments free.
Fig. 47. — Crossotonotus lopliocheir sp. nov., 3 holotype, 4.2 x 5.1 mm, Touho Bank, northeast coast of New
Caledonia, Expedition Montrouzier, 15 m (MNHN-B 26733): a, carapace, dorsal view; b, suborbital border;
c, dactylus, right fifth pereopod, dorsal view; d, left male first pleopod, lateral (outer side) view; e, left male first
pleopod, apex, dorsal view.
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
581
DESCRIPTION. — Carapace (Fig. 47a) subquadrate, slightly wider than long (CW/CL = 1.1-1 .2); dorsal surface
depressed, covered with small granules, large, slightly swollen areas with low bosses. Confluence of branchial and
mesogastric regions depressed, granular. Anterolateral borders of carapace each with three pointed or slightly
rounded anterolateral teeth, first rising from base of postorbital angle. Lateral borders each with 3-4 (number
increases with carapace size) small, blunt tubercles; posterolateral borders each with 1-3 similar but slightly
smaller tubercles. Posterior border separated from posterolateral border by clear junction, with 10-14 blunt or
slightly pointed tubercles, conspicuous plumose setae.
Frontal border of carapace divided into four lobes, inner lobes longer and more pointed. Borders between frontal
lobes and supraorbital borders slightly folded upward medially, ending in sharp angle forming V- shaped fissure
before supraorbital border. Supraorbital borders each with two slightly pointed lobes. Postorbital angles long
(extend to slightly beyond dorsal border of retracted eye) and pointed inward. Cornea of eyes spherical, wider than
base of long eye peduncle; each peduncle with two conspicuously long, granular tubercles on distal border.
Suborbital borders (Fig. 47b) each with one triangular inner lobe topped by two pointed teeth (innermost
longest), one broad, rounded outer lobe topped by two teeth (innermost blunt, outermost pointed and longest).
Pterygostomial lobes project slightly ventrally, slightly pointed.
Each basal antennal segment slender and rectangular; flagellum long, with very long, simple setae. Epistome
very narrow, vertically inclined, with two short, pointed median teeth, each flanked by short outer process and thin,
rounded margin before pterygostomial lobe. Inner border of ischia of third maxillipeds straight; surface granular,
upper borders round. Meri much narrower than ischia; very small, rounded upper borders.
Dorsal borders of cheliped propodi each with one conspicuous, rounded tubercle bordered by minute teeth and
1-2 small, pointed tubercles. Fingers of largest cheliped with rounded teeth, similar teeth or cutting edge in
smallest cheliped. Carpus short, each outer border with one conspicuous, rounded tubercle and several small,
pointed tubercles; meri slender, outer borders with rounded and pointed tubercles.
Walking legs (P2-5) slender but without filiform propodi and dactyli. Upper and lower borders of meri with
pointed tubercles of different sizes; distalmost tubercle on each anterior border much wider at base, much higher,
slender, pointed, directed distally (decrease in size from P2 to P5). Anterior borders of carpi of all walking legs
with small tubercles (smaller in P5); anterior and posterior borders of propodi with short teeth (thin spines along
anterior borders of P5); posterior borders of dactyli with teeth, each topped with spine (two on P2, 2-3 on P3-4.
three on P5), anterior borders with short teeth (3-4 long, curved spines along anterior border of dorsal side of each
P5; Fig. 47c). Each P2-4 with one dorsal, carina each on carpus and propodus; each P5 with one dorsal carina on
propodus. Meri, carpi, and propodi of all walking legs with plumose setae; dorsal surfaces of propodi of P3-4 each
with one row of conspicuous plumose setae along anterior and posterior borders, dactyli with one similar row
along each anterior border.
First pair of walking legs (P2) shorter, more slender than second and third pairs (P3-4); third pair slightly
shorter than second. Fifth pair of walking legs (P5) short (0.9- 1.0 CL), flattened but more slender than the anterior
walking legs (P2-4).
The relative length of pereopods 2-5 among the specimens that were measured is as follows:
Abdomen of mature males with all segments free. Male first pleopods (Figs 47d-e) with sinuous basal parts;
each distal part straight, one pointed tip divided into two unequal parts both bordered by minute teeth. Abdomen of
mature females with all segments free.
DISCUSSION. — This species is being described from only one male and three females, all of small size.
The smallest female is a juvenile; the remaining two (the smallest, 5.5 x 6.2 mm) are ovigerous. It seems to be
restricted, like C. spinipes, to hard substrates.
Source : MNHN, Paris
582
P. CASTRO
Other than the size difference (C. spinipes attains a much larger size and females become ovigerous at a much
larger size), diagnostic characteristic for C. lophocheir are the presence of a large, rounded tubercle on the dorsal
border of each cheliped propodus (two rows of relatively shorter tubercles in C. compressipes and C. spinipes),
relatively longer walking legs (P3 1. 7-2.0 CL in contrast to 1.4-1. 7 CL in C. spinipes), the large cheliped of
the male lacks the sinuous ridges, or presumed stridulating surface of C. spinipes, and the inner border of the
propodi of the male chelipeds lack a dense cluster of setae.
SIZE. — Maximum size among specimens examined: 6.0 x 6.8 mm (female); 4.2 x 5.1 mm (male holotype).
ETYMOLOGY. — From lophos, Greek for crest or ridge, and cheir, Greek for hand, in reference to the
characteristic crest-like tubercles on the dorsal border or the cheliped propodi and carpi.
DISTRIBUTION. — Known only from New Caledonia. Depth: 10-44 m.
Genus PLEUROPHRICUS A. Milne Edwards, 1873
Pleurophricus A. Milne Edwards, 1873a: 84. — Moosa & Serene, 1981: 53 (in key).
TYPE Species. — Pleurophricus cristatipes A. Milne Edwards, 1873.
DIAGNOSIS. — Carapace subcircular; frontal border divided into 2-4 lobes. Anterolateral borders with pointed or
rounded teeth. Dorsal surface of carapace with small granules and high, granular bosses, including two very high
bosses on mclagastric region. Eyes spherical, peduncles with soft tubercles. Supraorbital borders each with two
lobes. Suborbital borders each with two lobes. Each basal antennal segment rectangular. Epistome very narrow.
Chelipeds equal. Walking legs (P2-5) similar in shape; meri may be inflated, dactyli hook-like. Abdomen of
mature males elongate (sides parallel to each other), with all segments free. Male first pleopods long, slender with
basal parts straight; each distal part uniramous, tip with minute teeth. Abdomen of adult females with all segments
free.
Redescription. — Carapace subcircular, dorsal surface with granular bosses, including two very high bosses
on metagaslric region. Anterolateral borders each with three pointed or three secondarily bifurcated, slightly pointed
teeth; posterolateral borders with slightly rounded tubercles, with marked junction with convex posterior border.
Confluence of branchial and mesogastric regions depressed and smooth; no conspicuous sulcus between hepatic and
branchial regions. Posterior border with 5-6 salient, rounded tubercles. Thoracic sternite 7 without cpisternal
process visible dorsally.
Frontal border divided into 2-4 pointed or rounded lobes (if four, inner pair more advanced anteriorly than outer
pair). Supraorbital borders each with two rounded lobes. Postorbital angles short, with slightly pointed tips.
Cornea of eyes spherical, slightly wider than base of eye peduncle; peduncle with granular tubercles at least on
distal border. Orbits deep.
Suborbital border with two broad, rounded lobes. Ptcrygostomial lobe at each anterolateral angle of buccal
frame broad and pointed.
Each basal antennal segment rectangular. Epistome very narrow, not expanded dorsovenlrally, vertically
inclined. Meri of third maxillipeds smaller and narrower than ischia.
Chelipeds equal in males. Walking legs (P2-5) about same size. Meri may be inflated and dactyli hook-like.
Abdomen of mature males elongate, with both sides almost parallel to each other, with all segments free, no
transverse ridges; first segment broad and long. Male first pleopods long and slender; basal part straight; distal part
uniramous, with tip bordered by minute teeth. Abdomen of adult females with all segments free, no transverse
ridges.
Species included. — P. cristatipes A. Milne Edwards, 1873 and P. longirostris (Moosa & Serene, 1981).
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
583
Sexual Dimorphism. — Unknown.
DISCUSSION. — Pleurophricus was described for P. cristatipes A. Milne Edwards, 1873, which until now had
been known from only one specimen collected in Australia ("Nouvelle Hollandc"). The genus was placed by
A. Milne Edwards (1873a: 260) among the Oxystomata a cote des Orithya (Ocypodidae). Miers (1879: 660)
included the genus (but with some doubt) in the Oxyrhyncha and Haswell ( 1882: 23) listed it among the crabs of
Australia under the family Maiidae. The structure of the orbits, eyes, peduncles, and carapace leave no question of
its inclusion in the Palicidae, even if only two incomplete specimens have been examined.
The morphologies of the abdomen and of the male first pleopods, both absent in the holotype of P. cristatipes,
are very close to those of Crossotonotus spinipes , which would support the inclusion of Pleurophricus in
Crossotonotus A. Milne Edwards, 1873. Both genera were actually described in the same publication but placed
among different groups of brachyuran crabs. Their descriptions, however, did not include any characters that could
be used to actually differentiate between the two. The similarities were so close that the second species of
Pleurophricus that was described (P. spinipes De Man, 1888) is actually a species of Crossotonotus (see above).
Nevertheless, NOBILI (1907: 402) stated that P. cristatipes and C. spinipes (as Pleurophricus spinipes) did not
belong to the same genus. Significant differences in the shape of the carapace (subcircular in shape, fewer and
larger, salient tubercles along posterior border, two very high and conspicuous bosses on metagastric region, in
contrast to the subquadrate carapace with very low bosses and many, very small tubercles along the posterior border
in Crossotonotus) and walking legs (presumably shorter and more equal in length than in Crossotonotus).
however, support the separation of the two genera within the Crossotonotinae.
Key to the species of PLEUROPHRICUS
1. Narrow, pointed anterolateral teeth; posterior border with five tubercles (Fig. 48a).
Walking legs with narrow meri (A. Milne Edwards, 1873a, pi. 1, fig. 6) .
. P. cristatipes A. Milne Edwards, 1876
— Flattened, wider anterolateral teeth; posterior border with six tubercles (Moosa & Serene,
1981, fig. 14). Walking legs with inflated, nearly rounded meri .
. P. longirostris (Moosa & Serene, 1981)
Pleurophricus cristatipes A. Milne Edwards, 1873
Figs 48
Pleurophricus cristatipes A. Milne Edwards. 1873a: 260 [84], pi. 1, figs 6, 6b. 6c. — Haswell. 1882: 23. — Rathbun,
1906: 837. — SerTne, 1968: 97. — Moosa & Ser£ne, 1981: 60.
MATERIAL EXAMINED. — Indonesia. Kai Islands. "Alpha Helix"-, stn M-99. 05°31.6'S. I32°17.8'E. 6-18 m.
7.07.1979: 1 3 (USNM).
Australia. 1 3 holotype 9.0 x 9.0 mm (ZMUH K-347).
Types. — Holotype-. 1 3 9.0 x 9.0 mm, originally at Godeffroy Museum, Hamburg, no. 6044 (ZMUH K-
347).
Type Locality. — Australia, unknown depth.
Diagnosis. — Carapace (Fig. 48a) subcircular, with several elevated areas topped by conspicuous granular
bosses (two very conspicuous bosses on metagastric region). Three pointed anterolateral teeth (first rising from
base of postorbital angle) on each side of carapace, one large, pointed tubercle along each lateral border, 1-2
slightly smaller, pointed tubercles along each posterolateral border; straight posterior border separated from
posterolateral border by clear junction, with five blunt, salient tubercles. Four lobes along anterior border; rounded
Source : MNHN. Paris
584
P. CASTRO
median lobes slightly longer than rounded outer lobes. Supraorbital borders each with two rounded lobes.
Suborbital borders (Fig. 48b) each with two slightly pointed lobes, inner fused to pterygostomial lobe. Dorsal
borders of cheliped propodi each with two rounded tubercles (A. Milne Edwards, 1873a, pi. 1. fig. 6). First
three walking legs nearly equal in size (A. MlLNE Edwards, 1873a. pi. 1, fig. 6). Abdomen of mature males
with all segments free. Male first pleopods with straight basal parts; each distal part straight, with simple tip.
Females unknown.
Fig. 48. — Pleurophricus crisiatipes A. Milne Edwards, 1873, <3 holotype, 9.0 x 9.0 mm, Australia, unknown depth
(ZMUH K-347): a, carapace, dorsal view; b, suborbital border.
DISCUSSION. — The type material of P. crisiatipes , a male (9.0 x 9.0 mm; ZMUH K-347). is unfortunately
incomplete. While the dorsal surface and the anterior portion of the ventral surface of the carapace are complete
(Figs 48a-b), the remaining portions of the ventral surface survive as two large pieces, and the walking legs and
chelipeds are missing. The figure given in the description (A. MlLNE Edwards, 1873a, pi. 1, fig, 6) showed that
the first pair of walking legs is not reduced and that the last pair is only slightly reduced in size in comparison to
the other legs. The walking legs were described 'as having an "unequal crest" ( Crete inegale). The chelipeds were
shown as nearly identical (raising sonic doubts on the accuracy of the figure). Both were heavy, with two large
tubercles on the dorsal surface of each propodus.
One very small, soft, and incomplete (no appendages) male from the Kai Islands (Banda Sea) Indonesia (2.4 x
2.4 mm; USNM) proved to be identical to the holotype of P. crisiatipes as far as the shape of the carapace is
concerned (number, shape, and position of teeth and tubercles, morphology of the anterior, posterior, and
supraorbital borders). The coxae of the fourth walking legs, as in P. crisiatipes, were aligned with the coxae of the
anterior walking legs. The abdomen of the Banda Sea specimen had all segments free. The male first pleopods (soft
in the specimen) appeared to have a straight basal part. The distal part was straight, with a simple tip very much as
in C. spinipes and C. lophocheir. The presence of abdominal segments that were all free strongly suggests that the
male was mature.
SIZE. — Maximum size among the two specimens examined: 9.0 x 9.0 mm (male holotype; ZMUH K-347).
Distribution. — Known only from Australia ("Nouvelle Hollande"; A. Milne Edwards, 1873a) and the
Kai Islands, Banda Sea, Indonesia. Depth: 6-18 m.
Source : MNHN. Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
585
Pleurophricus longirostris (Moosa & Serene, 1981)
Marietta longirostris Moosa & Serene, 1981: 58, fig. 14.
Material EXAMINED. — Indonesia. Sunda Strait. Off Panjang Island, 16.03.1963: 1 2 4.6 x 4.6 mm (RDC
CB2712).
Types. — Holotype: 1 9 4.6 x 4.6 mm, 16.03.1963 (RDC CB2712).
TYPE Locality. — Oft Panjang Island, Sunda Strait (between Sumatra and Java), Indonesia.
Diagnosis. — Carapace (Moosa & Serene, 1981, fig. 14) hexagonal, depressed, with swollen areas with
bosses topped by rounded tubercles. Four broad, slightly pointed anterolateral teeth on each side carapace, two
large, pointed teeth along each lateral border; slightly convex posterior border separated from posterolateral border
by clear junction, six rounded, low tubercles. Two narrow, squarish lobes along anterior border. Supraorbital
borders each with two very short, rounded lobes. Suborbital borders each with two lobes, one with triangular,
pointed tip, second obliquely directed with irregular border. Dorsal border of cheliped propodi with cusp of pointed
tubercles. Meri of walking legs inflated, laterally flattened, almost circular. Dorsal borders of mcri and carpi with
thin, rounded tubercles. Dactyli short, hook-like, one spine on each posterior border. Males unknown. Abdomen of
mature females with distinct segments.
Discussion. — The species was described from one small ovigerous female collected in the Sunda Strait.
Indonesia (MOOSA & SERENE, 1981). It was included in Marietta, a junior subjective synonym of Crossotonotus
(see above). It is tentatively placed in Pleurophricus since its carapace is very similar in shape to that of
P. cristatipes (Fig. 48a). There are differences between the species, however. The most obvious is in the presence
in the walking legs of P. longirostris of inflated, almost circular meri bordered by ihin. rounded tubercles in
contrast to the thin meri of C. cristatipes (A. MlLNE EDWARDS, 1873a, pi. 1, fig. 6). They were described as
"very inflated which give the impression of a tree leaf’ (MOOSA & SERENE, 1981: 58). The flattened meri, hook¬
like dactyli, and the very long plumose setae on the meri, carpi, and particularly on the propodi suggest that the
species may be cryptic and perhaps associated with the surface of algae or another organism.
Other differences are in the shape of the anterolateral teeth. They are flattened, secondarily divided, and less
pointed (very much as in C. spinipes) in P. longirostris, while they are narrower and more pointed in P. cristatipes
(Fig. 48a). There are six rounded tubercles along the posterior border of P. longirostris but five pointed tubercles in
P. cristatipes (Fig. 48a).
The relative length and shape of the fifth pair of walking legs (P5) are unknown. They articulate to the carapace
at the same level as the other walking legs as in other members of the Crossotonotinae.
Size. — Size of female holotype: 4.6 x 4.6 mm.
DISTRIBUTION. — Only known from Panjang Island, Sunda Strait, Indonesia. Depth: unknown.
ECOLOGY AND OTHER ASPECTS OF THE BIOLOGY
OF THE INDO-WEST PACIFIC PALICIDS
Habitat. — Most species of palicids appear to be restricted to mud, sand, and other soft sediments at moderate
depths. This is the case of species of Pseudopalicus, Parapalicus, Miropalicus, Paliculus, and Rectopalicus in
the Indo-west Pacific and of Palicus in the Mediterranean, Atlantic, and eastern Pacific. Species of Neopalicus and
Palicoides are known only from coarse, biogenous sediments in shallow water around Indo-west Pacific coral
reefs. Species of Exopalicus, Crossotonotus, and apparently Pleurophricus, are inhabitants of hard bottoms in
shallow water.
Source : MNHN. Paris
586
P. CASTRO
Species of Parapalicus and Miropalicus (plus Pseudopalicus macromeles and Paliculus kyusyuensis ) have long,
slender legs that seem to be adapted to slide over muddy sediments. At least this is what has been observed in
freshly collected live specimens of Parapalicus (B. Richer deForges, personal communication). There is
no information on the microhabitats and habits of the other palicids that live on soft sediments but that
have flattened walking legs (species of Pseudopalicus and Rectopalicus ) other that the legs appear to be adaptations
for occasional swimming. This possibility is supported by observations of live P. amadaibai using its second
and third pairs of walking legs for active swimming off the sandy bottom of an aquarium (T. KOMAl, personal
communication).
The high bosses and tubercles on the carapace of most palicids do not support the possibility that any of the
species inhabiting soft sediments are long-term borrowers in the sediment itself. This, however, may not be the
case of species of Neopalicus and Palicoides , inhabitants of coarse sediments in coral reefs. Their carapaces are
relatively smooth, being devoid of high bosses. Their habits, however, remain unknown.
There is no detailed information on the microhabitats and habits of palicids living on hard substrates. Many of
the few records of Exopalicus maculatus have been from collections during night diving, a suggestion that they
hide in coral and other hard substrates during daytime.
Most of the species inhabiting soft sediments have been collected from moderate depths in the subtidal and
bathyal zones (Table 5): Pseudopalicus (30-430 m), Parapalicus (55-778 m), Miropalicus (245-575 m), Paliculus
(50-487 m), and Rectopalicus (105-430 m). Exceptions are the species inhabiting coarse sediments in coral reefs:
Neopalicus (10-146 m) and Palicoides (6-73 m). Inhabitants of hard substrates are also restricted to shallow water:
Exopalicus ("shallow water" to 18 m) and Crossotonotus ("intertidal" to 146 m). Nothing is known about the
habitat of the two little-known species of Pleurophricus but they appear to be inhabitants of hard bottoms in
shallow water (only record forP. cristatipes, 6-18 m).
Biogeography. — Information on the geographic distribution of Indo-west Pacific palicids is limited and
incomplete. The majority are deep-water, cryptic, or nocturnal species that are rarely collected or seen. Most of the
information (as well as most of the new species described here) are from the Coral Sea and southwestern Pacific,
where recent collecting effort has been more intense (Table 6). Conversely, little is known about the distribution
of palicids through most of the Indian Ocean and the central Pacific,
There is no evidence of the occurrence of any palicids in the Indian or Pacific oceans outside the tropical Indo-
west Pacific region proper. Species of Palicus , however are found in subtropical or temperate waters outside the
tropics in the Mediterranean and off the northwestern coast of Africa [P. caronii (Roux, 1830)], the eastern coast of
North America north of northern Florida and/or the eastern coast of South America south of Rio de Janeiro
[P. alternants (Rathbun, 1897), P. cursor (A. Milne Edwards, 1880), P. dentatus (A. Milne Edwards, 1880),
P.faxoni Rathbun, 1897, P. gracilis (Smith, 1883), P. obesus (A. Milne Edwards, 1880)], and the Gulf of
Calilornia [P. cortezi (Crane, 1937), P. fragilis (Rathbun, 1893), P. lucasii Rathbun, 1898 and P. zonatus
(Rathbun, 1893)]. Most of these species, however, are also found in tropical regions (see RATHBUN, 1918;
Melo, 1996).
The most widely distributed species that show a wide Indo-west Pacific distribution are inhabitants of shallow
water: Neopalicus contractus and N. jukesii (Fig. 49), Palicoides longimanus and P. whitei (Fig. 50),
Crossotonotus spinipes (Fig. 51), and Exopalicus maculatus (Fig. 56). Although shallow-water species are the
easiest to collect, some inhabitants of deeper water also show a wide Indo-west Pacific distribution. This is the
case of Pseudopalicus investigatoris , P. serripes (Fig. 52), and Paliculus kyusyuensis (Fig. 56). None of the Indo-
west Pacific species is known to cross the East Pacific barrier. The six species of palicids found in the eastern
Pacific appear to be endemics to the region.
Most species of Pseudopalicus (Figs 52-55), Parapalicus (Figs 57-58), Miropalicus (Fig, 59), and Rectopalicus
(Fig. 53) show a more restricted geographic distribution, mostly along the western Pacific Ocean. This is perhaps
best explained as a result of collection that has been mostly limited to these areas. Restriction to particular
substrates or differences in the length of larval development or the behavior of the free-living larvae, however, may
effect the geographic distribution of these species.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
587
Table 5. — Depth distribution (reliable ranges only) of Indo-west Pacific palicids.
0
100
200
Depth in meters
300 400 500
600
700
Crossonotus compressipes
Crossotonotus spinipes
Exopalicus maculatus
Pleurophricus cristatipes
Palicoides longimanus
Palicoides whilei
Crossotonotus lophocheir
Neopalicus jukesii
Crossotonotus ceramensis
Pseudopalicus serripes
Pseudopalicus investigatoris
Neopalicus contractus
Paliculus kyusyuensis
Parapalicus piruensis
Pseudopalicus amadaibai
Parapalicus tritube rculatus
Pseudopalicus sp.
Parapalicus elaniticus
Parapalicus ambonensis
Parapalicus unidentatus
Paliculus sp.
Parapalicus inermis
Rectopalicus amphiceros
Rectopalicus woodmasoni
Pseudopalicus sexlobatus
Pseudopalicus pictus
Pseudopalicus macromeles
Parapalicus nanshaensis
Parapalicus microphthalmus
Pseudopalicus declivis
Parapalicus trispiralis
Rectopalicus ampullatus
Parapalicus inanis
Pseudopalicus glaber
Pseudopalicus undulatus
Parapalicus denticulatus
M iropalicus vietnamensis
Pseudopalicus oahuensis
Pseudopalicus acanthodactylus
Parapalicus armatus
Parapalicus clinodentatus
intertidal
Source : MNHN, Paris
588
P. CASTRO
Table 6. — Geographic distribution of Indo-west Pacific palicids.
(1) SE Coral Sea includes New Caledonia, Loyalty Is, Chesterfield Is, Vanuatu.
Source : MNHN, Pans
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
589
Fig. 50. — Geographic distribution of (•) Palicoides longimanus (Miyake. 1936) and (O) P. whitei (Miers. 1884).
Fig. 51. — Geographic distribution of (•) Crossotonotus spinipes (De Man. 1888) and (O) C. compressipes A. Milne
Edwards. 1873.
Source : MNHN, Paris
590
P. CASTRO
Fig. 52. — Geographic distribution of (•) Pseudopalicus investigatoris (Alcock. 1900). (©) P. macromeles sp. nov..
(O) P. serripes (Alcock & Anderson. 1895), and (A) P. sexlobatus (Kensley, 1969).
Fig. 53. — Geographic distribution of (•) Pseudopalicus amadaibai (Sakai, 1963), (®) P. undulaius sp. nov.. and
(□) Rectopalicus woodmasoni (Alcock, 1900).
Fig. 54. — Geographic distribution of (O) Pseudopalicus oahuensis (Rathbun. 1906) and (•) P. picius sp. nov.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PAL1CID CRABS
59 I
Fig. 55. Geographic distribution of (©) Pseudopalicus acanihodactylus sp. nov., (•) P. declivis sp. nov., and
(O) P. glaber sp. nov.
Fig. 56. — Geographic distribution of (• ) Paliculus kyusyuensis (Yokoya, 1933) and (O) Exopalicus maculatus
(Edmondson. 1930).
FlG. 57. — Geographic distribution of (©) Parapalicus annatus sp. nov.. (•) P. clinodeniaius sp. nov.. (O) P. denticulatus
sp. nov., (□) P. elaniticus (Holthuis, 1977), (a) P. inanis sp. nov., (□) P. inermis sp. nov.. and (■ )
P. microphtlialmus sp. nov.
Source : MNHN, Paris
592
P. CASTRO
Fig. 58. — Geographic dislribution of (□) Parapalicus ambonensis sp. nov., (©) P. piruensis Moosa & Serene, 1981,
(A) P. trispiralis sp. nov., (O) P. trituberculatus (Chen, 1981), and (•) P. unidentatus (Zarenkov, 1968).
Fig. 59. — Geographic distribution of (•) Miropalicus vietnamensis (Zarenkov. 1968).
DIET. — Casual analyses of stomach and intestinal contents showed that palicids ingest sediment and digest
particulate organic matter in it. This applies to inhabitants of soft sediments as well as those assumed to live on
hard substrates such as Exopalicus maculatus and Crossotonotus spinipes.
Sexual Dimorphism and Mating Behavior. — The presence of more unequal and much thicker or longer
chelipeds in the males of many species plus the conspicuous presence of setae on the inner, distal surface of male
chelipeds in six Indo-west Pacific species ( Pseudopalicus serripes, Exopalicus maculatus, Neopalicus jukesii,
Palicoides longimanus, P. whitei, and C. spinipes) suggests the role of chelipeds in mating behavior. The sinuous
ridges found along the ventral border of the largest cheliped in the males of C. spinipes were suggested to serve as
a stridulating structure by RATHBUN (191 1: 241).
ROLE of the FIFTH Pair of Pereopods. — The biological significance of the reduced and slender fifth pair of
pereopods characteristic of members of the subfamily Palicinae remains a mystery. The structure of the
reproductive system of palicids is one characteristic of the less primitive families of Brachyura so that reduced P5
should not be considered a primitive, less specialized character.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
593
No observations have been made of the behavior of the P5 in live palicids. There are no differences in the
morphology or size of the fifth pair of pereopods between males and females so they do not appear to be involved
in mating behavior. It can be hypothesized that they may be involved in cleaning the carapace and/or other
appendages. The joints of the P5 are unusually mobile, and the shortening of the articles certainly increases the
overall flexibility of the appendage. The movement of the various joints alternates between the dorsoventral and
antero-posterior planes beginning with the dorsoventral plane of movement of the sternal-coxal articulation. The
total length of the P5 is surprisingly similar in most of the species that were examined, varying between 0.7-0.9
CL with very few exceptions. This length would be just enough to reach the orbits and clean them of debris. The
same relative length of the P5, however, also applies to members of the subfamily Crossotonotinae, where the P5
are broad and appear to be locomotory in function. The dactyli of P5 among members of the subfamily Palicinae
are variously provided with short teeth and/or setae, typically of the simple type. Only the rows of thick, bristle¬
like spines that are present along the posterior border of the propodi in Neopalicus and Palicoides offer
a potentially efficient cleaning surface.
WlCKSTEN (1986: 365) suggested that the reduced P5 of palicids, retroplumids, and pinnotherids may be
involved in climbing in burrows or in cleaning the carapace. There is no evidence that palicids are borrowers,
although this remains a possibility. Nevertheless, the suggested involvement of the P5 in the cleaning of the
carapace is not supported by the frequent presence of sediment or polychaete tubes on the carapace and even around
the eye (Fig. 12e).
The absence of chelate or subchelate dactyli does not preclude their use in actively carrying another organism or
object. Carrying behavior in many brachyurans, however, involves modified appendages unlike those of palicids
(see GUINOT et al, 1995). There is unfortunately no evidence of carrying behavior among palicids with reduced P5.
It is possible, however, that individuals carry shells or fragments of shells that are lost during collection.
Although still reduced in size, the P5 of members of the subfamily Crossotonotinae have the same
morphology and insertion to the carapace as the anterior walking legs. All members of the subfamily (species of
Crossotonotus and Pleurophricus) appear to be restricted to hard substrates so in these species the P5 are adapted,
possibly secondarily, to an ambulatory function. Nevertheless, the P5 of Exopalicus. w-hich also appears to be
restricted to hard substrates, remain slender as in the inhabitants of soft substrates.
Another possible explanation for the reduction in the size of the P5 is that they have no specific function. They
may thus represent vestigial structures, at least in the Palicinae. An analogous reduction in the size of the P5 is
observed among brachyuran crabs of the families Hexapodidae and Retroplumidae. The affinities between palicids
and these two families remain unknown.
AC KNO WLEDGMENTS
Without the encouragement of A. CROSNIER this work would not have been attempted; without the rich
collections of IRD (Institut de Recherche pour le Developpement, formerly ORSTOM), made available by
A. CROSNIER and B. Richer de Forges, my research would have not been possible. My hearty thanks thus go to
both of them.
Material was also made available by those who sent me valuable specimens (particularly type material), went
out of their way to provide me with additional information on the material, or welcomed me to their institutions:
A. Alspach (Forschungsinstitut Senckenberg, Frankfurt am Main. Germany), P. BERENTS (Australian Museum.
Sydney, Australia), A. BRANDT (Zoologisches Institut und Zoologisches Museum, Universitat Hamburg,
Germany), T.-Y. Chan (National Taiwan Ocean University, Keelung, Taiwan), H.-L. Chen (Institute of
Oceanology, Qingdao, China), C.H. COLEMAN (Museum fur Naturkunde, Humbolt Universitat. Berlin. Germany),
A. DAI (Academia Sinica, Beijing, China), P. Davie (Queensland Museum. Brisbane, Australia), D. ElBYE-
JACOBSEN (Zoologisk Museum, University of Copenhagen, Copenhagen, Denmark), L. ELDREDGE (Bishop
Museum, Honolulu, Hawaii), C. Fransen (Nationaal Natuurhistorisch Museum, Leiden. The Netherlands),
D. GuiNOT (Museum national d'Histoire Naturelle, Paris, France), K.-I. Hayashi (material from the Kitakyushu
Museum of Natural History, Kitakyushu & Zoological Laboratory, Kyushu University. Fukuoka. Japan),
Source :
594
P. CASTRO
J. Hoover (Hawaii Medical Library, Honolulu, Hawaii), H. IKEDA (Hayama Shiosai Museum, Miura, Japan),
M.-J. Jeng (Institute of Zoology. Academia Sinica, Taipei. Taiwan), A. JOHNSON (Museum of Comparative
Zoology. Harvard University, Cambridge, Massachusetts), D. JONES & M. Hewitt (Western Australian Museum.
Perth. Australia), T. KOMAI (Natural History Museum and Institute, Chiba, Japan), R. Lemaitre & K.J. Reed
(United States National Museum, Smithsonian Institution, Washington, D.C.), M. Lowe (Natural History
Museum, London, United Kingdom), J.W. Martin (Natural History Museum of Los Angeles County, Los
Angeles, California), K. MURAOKA (Kanagawa Prcfectural Museum of Natural History, Odawara, Japan),
K. NOMURA (Kushimoto Marine Park Center, Kushimoto, Japan), P.K.L. NG (Zoological Reference Collection.
Department of Biological Sciences. National University of Singapore), D. Platvoet (Zoologisch Museum.
Universiteit van Amsterdam. The Netherlands). R.C. PREECE and R. SYMONDS (University Museum of Zoology,
University of Cambridge, Cambridge, United Kingdom), M. VAN DER MERWE (South African Museum. Cape
Town, South Africa), and M. VANNINI (Museo di Zoologia "La Specola", Universita di Firenze, Italy). I am most
thankful to D.L. RAHAYU (Research and Development Center for Oceanology of Indonesia) and P.K.L. Ng
(National University of Singapore) for literally going out of their way to make available to me the type material of
the species described by Moos a & Serene (1981).
Suggestions by patient colleagues, although invariably resulting in giving me more things to do, yielded much
insight on the taxonomy and morphology of a group of crabs I have not yet been privileged to see alive: K. Baba
(Kumamoto University, Kumamoto, Japan), D. GuiNOT (Museum national d’Histoire naturelle. Paris), T. Komai
(Natural History Museum and Institute, Chiba, Japan), C.L. McLay (University of Canterbury, Christchurch,
New Zealand), P.K.L. Nc. (National University of Singapore). B. Richer de Forges (IRD), and M. Tavares
(Universidade Santa Ursula, Rio de Janeiro, Brazil). Comments and advise by three outstanding reviewers much
improved the manuscript: P. Davie (Queensland Museum, Brisbane), L.B. Holthuis (Nationaal Natuurhistorisch
Museum, Leiden), and P.K.L. Ng.
My very special appreciation to J.-F. DEJOUANNET, who expertly did most of the artwork, took some of the
photographs, and patiently helped to put everything together in the right place. F. Theureau drew some of the
illustrations of the species of Parapalicus and M. Dumont and R. von Cosel took some of the photographs.
M. Tortelier (IRD. Noumea, New Caledonia) prepared the maps.
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602
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FIGURE 60
a, Pseudopalicus acanthodactylus sp. nov., <J holotype, 8.9 x 10.5 mm. Norfolk Ridge, south of New Caledonia. Smib 8.
stn DW 146, 514-522 m (MNHN-B 26672): dorsal view. Photo by J.-L. MENOU, 1RD.
b, Pseudopalicus glaber sp. nov., 9 8.4 x 10.6 mm, off southeast coast of New Caledonia. SMIB 4. stn DW 57. 260 m
(MNHN-B 26812): dorsal view. Photo by P. Laboute, IRD.
c, Pseudopalicus oahuensis (Rathbun, 1906), juvenile 9 8.2 x 10.7 mm. Norfolk Ridge, south of New Caledonia.
Chalcal 2, stn DW 75, 600 m (MNHN-B 26813): dorsal view. Photo by P. Laboute, IRD.
d, Pseudopalicus invesligaioris (Alcock, 1900), 9 11.7 x 14.5 mm, Vanuatu. Musorstom 8. stn CP 1077. 180-210 m
(MNHN-B 26746): dorsal view. Photo by J.-L. MENOU, IRD.
e, Pseudopalicus pictus sp. nov., <5 holotype, 8.6 x 10.4 mm. Vanuatu. Musorstom 8, stn DW 1042, 200-265 m
(MNHN-B 26690): dorsal view. Photo by J.-L. MENOU, IRD.
f, Miropalicus vietnamensis (Zarenkov, 1968), Loyalty Islands, Musorstom 6, stn SW 410, 490 m (lost specimen):
dorsal view. Photo by P. Laboute, IRD.
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
603
Source : MNHN. Paris
604
P. CASTRO
FIGURE 61
a, Rectopalicus ampullatus sp. nov., 3 holotype, 6.2 x 6.3 mm. Loyalty Islands, MUSORSTOM 6. stn DW 462. 200 m
(MNHN-B 26782): dorsal view. Photo by P. Laboute, 1RD.
b, Exopalicus maculatus (Edmondson. 1930), Makaha, Oahu. Hawaiian Islands, 6 m: dorsal view. Photo by J. Hoover.
c, Neopalicus jukesii (White, 1847). <3 6.0 x 7.1 mm. New Caledonia, Lagon, stn 702, 37 m (MNHN-B 26814): dorsal
view.
d , Palicoides longinuinus (Miyake, 1936), 3 9.6 x 11.2 mm, New Caledonia, Lagon, stn 952, 16-17 m (MNHN-B
26794): dorsal view.
e, Palicoides whitei (Miers, 1884), 3 CL 14.5 mm. New Caledonia, Lagon, stn 748, 35 m (MNHN-B 26801).
f, Crossoionotus spinipes (De Man, 1888), 3 36.9 x 43.3 mm. New Caledonia, 12 m (MNHN-B 26803).
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
605
Source : MNHN, Paris
606
P. CASTRO
INDEX
Family and genus names are given in capital letters, species names in italics. Bold family, genus and species
names indicates that the taxa are treated in detail.
Page numbers in bold indicate the pages where the subject is treated in detail; page numbers in bold italics
refer to identification keys, italics refer to illustrations.
abdomen, terminology 442
acanthodactylus, Pseudopalicus 450; 451;
452; 453; 457; 483; 588; 591; 602
acutifrons, Palicus 488; 522
ajfinis, Palicus 450
affinities 593
altematus, Palicus 450; 586
amadaibai, Palicus 455
amadaibai, Pseudopalicus 450; 451; 455;
456; 586; 588; 590
ambonensis, Parapalicus 482; 487; 489; 489;
491; 494; 498; 501; 505; 512; 516; 588; 592
amphiceros, Rectopalicus 483; 486; 534;
535; 538; 539; 545; 588
ampullatus, Rectopalicus 534; 535; 541;
543; 545; 547; 588; 604
angustus, Palicus 488
armatus, Parapalicus 487; 488; 492; 493;
498; 508; 588; 591
bidentatus, Palicus 440; 517; 518
biogeography 586
brevimana, Manella 570; 574; 578; 579
brevimanus, Crossotonotus 574
carapace length 440
carapace width 440
carapace, terminology 441; 442
carinipes, Cymopolia 554
carinipes, Palicus 554; 557
caronii, Cymopolia 445
caronii, Palicus 439; 446; 450; 586
ceramensis, Crossotonotus 570; 577; 578;
588
ceramensis. Manella 571 ; 578
clinodentatus, Parapalicus 487; 489; 495;
496; 498; 518; 588; 591
compressipes, Crossonotus 57 1
compressipes, Crossotonotus 569; 570;
571; 571; 573; 579; 582; 588; 589
contractus, Neopalicus 549; 550; 550; 552;
556; 557; 558; 586; 588; 589
contractus. Palicus 550
cortezi, Palicus 586
cristatipes, Pleurophricus 570; 582; 583;
583; 584; 585; 588
Crossonotus 569
compressipes 57 1
Crossotonotinae 440; 444; 445; 568; 593
Crossotonotus 439; 440; 445; 568; 569; 569; 583;
585; 586; 593
brevimanus 574
ceramensis 570; 571; 578; 588
compressipes 569; 570; 571; 571; 573;
579; 582; 588; 589
gardineii 574
lophocheir 570; 571; 579; 580; 584; 588
sp. 574
spinipes 570; 571; 574; 576; 577; 579; 581;
582; 583; 584; 585; 586; 588; 589; 592;
604
taketomiensis 57 1 ; 573
CROSSTONOTUS
taketomiensis 571
cursor. Palicus 488; 522; 586
Cymopolia 444
carinipes 554
caronii 445
cyrenae 469; 472
fished 469; 472
jukesii 548; 554
kyusyuensis 527
longimana 561
maculata 546
medipacifica 546; 547
oaliuensis 455; 465
robusta 550; 553
serripes 473
white i 558; 565
Cymopoliidae 443
cyrenae. Cymopolia 469; 472
cyrenae. Palicus 450
cyrenae, Pseudopalicus 469
declivis, Pseudopalicus 450; 451; 457; 458;
461; 482; 483; 588; 591
dentatus. Palicus 450; 586
denticulatus, Parapalicus 487; 489; 499;
500; 502; 505; 518; 520; 588; 591
depressus. Palicus 488
diet 592
DORIPPIDAE 445
Source MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
607
ecology 585
elaniticus, Parapalicus 487; 489: 505; 510;
512; 588; 591
EXOPALICUS 445; 446; 448 ; 543; 585; 586; 593
maculatus 543; 544; 545: 546; 547; 586;
588; 591: 592; 604
faxoni, Palicus 450; 586
fisheri, Cymopolia 469; 472
fisher i, Palicus 450; 469
fisheri, Pseudopalicus 469
floridanus, Palicus 488; 522
foliatus, Paliculus 526; 527; 530; 531: 588
fragilis, Palicus 586
gardineri, Crossotonotus 574
gardineri, Manella 570; 574; 577
glaber, Pseudopalicus 450; 451: 457: 462;
464\ 588; 591: 602
gracilipes, Palicus 488; 522
gracilis, Palicus 488; 522; 586
granulatus, Palicus 445
habitat 585
hatusimaensis, Palicus 522; 527; 528
hatusiniaensis, Parapalicus 527
Hexapodidae 593
inanis, Parapalicus 487; 488: 494; 498: 506;
508: 512; 513; 588; 591
inermis, Parapalicus 487; 489: 501; 502: 504:
588; 591
investigatoris, Palicus 469; 473
investigatoris, Pseudopalicus 450; 451:
456: 458; 468: 469; 470: All: 479: 586;
588; 590: 602
jukesii, Cymopolia 548; 554
jukesii, Neopalicus 549; 550: 551; 552; 553;
554; 556: 565; 586; 588; 589: 592; 604
jukesii. Palicus 550; 554
kyusyuensis, Cymopolia 527
kyusyuensis, Paliculus 525: 526; 527; 527;
529; 532; 533; 552; 586; 588; 591
kyusyuensis, Palicus 525; 527
longimana, Cymopolia 561
longimanus, Palicoides 538; 560; 561: 560;
563: 567; 586; 588; 589: 592; 604
longimanus, Palicus 561
longirostris, Manella 585
longirostris, Pleurophricus 582; 583: 585;
588
lophocheir, Crossotonotus 570; 577; 579;
580: 584; 588
lucasii, Palicus 450; 586
macromeles, Pseudopalicus 450; 451: 479:
480; 482: 486; 586; 588; 590
maculata, Cymopolia 546
maculatus, Exopalicus 543; 544; 545: 546;
547: 586; 588; 591: 592; 604
maculatus. Palicus 543; 546; 547
male first pleopod, terminology 442
Manella 439; 440; 568; 569; 570
brevimana 570; 574; 578; 579
ceramensis 571; 578
gardineri 570; 574; 577
longirostris 585
spinipes 574; 579
Manella, 585
marielae. Parapalicus 488; 517:518
mating behavior 592
medipacifica, Cymopolia 546: 547
microfrons, Palicoides 535; 538
microfrons, Palicus 535; 537
microphthalmus, Parapalicus 487; 489:
509: 511:514: 588; 591
MlROPALICUS 445; 446; 448: 481; 520; 526; 585
vietnamensis 483; 521; 522; 524: 525; 527;
588; 592: 602
nanshaensis, Parapalicus 440; 487: 488: 492;
508; 510; 588
Neopalicus 440: 445; 447; 448: 548; 560; 585;
586: 593
contractus 549; 550: 550; 552; 556: 557;
558; 586; 588; 589
jukesii 549; 550: 551; 552; 553; 554; 556;
565; 586; 588; 589: 592: 604
oahuensis, Cymopolia 455; 465
oahuensis, Palicus 455; 465
oahuensis, Pseudopalicus 450; 451: 458;
460; 465; 479: 483; 588; 590: 602
obesus, Palicus 450; 586
P5, role of 592
PALICAE 444
PALICES 444
PALICI 444
PALICIDAE 439; 443
PALICIDAE, affinities 445
PALICINAE 440; 444; 445: 445; 593
PALICOIDE 447
Palicoides 440: 445; 448: 538; 549; 558; 585;
586; 593
Source MNHN, Pans
608
P. CASTRO
longimanus 538; 560; 561: 561; 563; 567;
586; 588; 589 ; 592; 604
microfrons 535; 538
tematensis 563; 565; 567
whitei 538; 560; 561 ; 563; 564; 565; 586;
588; 589 ; 592; 604
PALICULUS 445; 446; 448: 525; 585; 586
foliatus 526; 527; 530; 53/; 588
kyusyuensis 525; 526; 527; 527; 529; 532;
533; 552; 586; 588; 591
sp. 526; 527; 532; 533; 588
PALICUS 439; 440; 444; 445; 446; 450; 488; 522;
523; 585
acutifrons 488; 522
afftnis 450
altematus 450; 586
amadaibai 455
angustus 488
bidentatus 440; 5 1 7; 5 1 8
carinipes 554; 557
caronii 439; 446; 450; 586
contractus 550
cortezi 586
cursor 488; 522; 586
cyrenae 450
dentatus 450; 586
depressus 488
faxoni 450; 586
fisheri 450; 469
floridanus 488; 522
fragilis 586
gracilipes 488; 522
gracilis 488; 522; 586
granulatus 445
hatusimaensis 522; 527; 528
investigatoris 469; 473
jukesii 550; 554
kyusyuensis 525; 527
longimanus 56 1
lucasii 586
maculatus 543; 546; 547
microfrons 535; 537
oahuensis 455; 465
obesus 450; 586
robustus 550
serripes 448; 473
sexlobatus 467
sic a 488
trituberculatus 440; 486; 517
tuberculatus 450; 546; 547
unidentatus 5 1 9
velerae 450
vietnamensis 520; 522
whitei 565
woodmasoni 535
zonatus 586
Parapalicus 440; 445; 446; 448: 450; 481;
486; 522; 523; 526; 564; 585; 586
ambonensis 482; 487; 489: 489; 491 ; 494;
498 ; 501; 505; 512; 516; 588; 592
armatus 487; 488: 492; 493: 498: 508; 588;
591
clinodentatus 487: 489: 495; 496: 498: 518;
588; 591
denticulatus 487; 489: 499; 500; 502; 505;
518; 520; 588; 591
elaniticus 487; 489: 505: 510; 512; 588;
591
hatusimaensis 527
inanis 487; 488: 494; 498: 506; 508: 512;
513; 588; 591
inermis 487; 489: 501; 502; 502; 504: 588;
591
marielae 488; 517; 518
microphthalmus 487; 489: 509; 511: 514:
588; 591
nanshaensis 440; 487; 488: 492; 508; 510;
588
piruensis 487; 489: 512; 5/3; 514: 588; 592
trispiralis 487; 489: 492; 494; 510; 512;
513; 515: 588; 592
trituberculatus 487; 488; 489: 502; 514:
517; 518: 520; 588; 592
unidentatus 482; 487; 488; 489: 498: 502;
514: 518; 519; 588; 592
vietnamensis 522
Parapleurophrycoides 439; 440; 568: 569
roseus 440; 569
pictus, Pseudopalicus 450; 451: 476; 478:
479: 588; 590; 602
piruensis, Parapalicus 487; 489: 512; 5/3;
514: 588; 592
PLEUROPHRICUS 439; 440; 445; 568; 569: 582;
585; 586; 593
cristatipes 570; 582; 583: 583: 584: 585;
588
longirostris 582; 583: 585; 588
spinipes 569; 570; 574: 577
Pleurophrycoides 439
Pseudopalicus 440; 445; 446; 448: 448; 481;
516; 522; 527; 585; 586
acanthodactylus 450; 451: 452; 453: 457:
483; 588 : 591: 602
amadaibai 450; 451, 455; 456: 457: 588;
590
cyrenae 469
declivis 450; 451: 457: 458: 461: 482; 483;
588:59/
Source : MNHN, Paris
INDO-WEST PACIFIC SPECIES OF PALICID CRABS
609
fisheri 469
glaber 450; 457; 457 \ 462; 464; 588; 597;
602
investigatoris 450; 457; 456; 458; 468:
469; 470: 477; 479; 586; 588; 590: 602
macromeles 450; 457; 479; 480; 482: 486;
586; 588; 590
oahuensis 450; 457; 456; 458; 460; 465;
479; 483; 588; 590: 602
pictus 450; 457; 476; 478: 479: 588; 590:
602
serripes 450; 457; 456; 468: 470: 471; 473;
475; 477; 479; 482; 586; 588; 590; 592
sexlobatus 450; 457; 460; 467; 588; 590
sp. 450; 474; 475; 588
undulatus 450; 457; 479; 483; 485: 588; 590
woodmasoni 535; 537
Rectopalicus 445; 446; 448: 527; 533; 544;
585; 586
amphiceros 483; 486; 534; 535: 538; 539:
545; 588
ampullatus 534; 535: 541; 543: 545: 547;
588; 604
woodmasoni 468: 533; 534; 535: 535; 536:
540; 545; 588; 590
Retroplumidae 593
robusta, Cymopolia 550; 553
robustus, Palicus 550
roseiis, Parapleurophrycoides 440; 569
serripes, Cymopolia 473
serripes, Palicus 448; 473
serripes, Pseudopalicus 450; 457 ; 456; 468:
470: 471; 473; 475; 477; 479; 482; 586;
588; 590; 592
sexlobatus, Palicus 467
sexlobatus, Pseudopalicus 450; 457; 460;
467; 588; 590
sexual dimorphism 592
sica, Palicus 488
spinipes, Crossotonotus 570; 577; 574; 576;
577; 579; 581; 582; 583; 584; 585; 586;
588; 592; 604
spinipes, Manella 574; 579
spinipes, Pleurophricus 569; 570; 574; 577
taketomiensis, Crossotonotus 571; 573
taketomiensis, Crosstonotus 57 1
terminology
abdomen 442
carapace 447; 442
male first pleopod 442
peropods 441
te mate ns is, Palicoides 563; 565; 567
trispiralis, Parapalicus 487; 489: 492; 494;
510; 512; 513; 575; 588; 592
trituberculatus, Palicus 440; 486; 5 1 7
trituberculatus, Parapalicus 487; 488; 489:
502; 514: 517; 518: 520; 588; 592
tuberculatus, Palicus 450; 546; 547
undulatus, Pseudopalicus 450; 451: 479:
483; 485: 588; 590
unidentatus, Palicus 519
unidentatus, Parapalicus 482; 487; 488; 489:
498; 502; 574; 518; 519; 588; 592
velerae, Palicus 450
vietnamensis, Miropalicus 483; 521; 522;
524; 525; 527; 588; 592; 602
vietnamensis, Palicus 520; 522
vietnamensis, Parapalicus 522
whitei, Cymopolia 558; 565
whitei, Palicoides 538; 560; 561: 563; 564;
565; 586; 588; 589 ; 592; 604
whitei, Palicus 565
woodmasoni, Palicus 535
woodmasoni, Pseudopalicus 535; 537
woodmasoni, Rectopalicus 468: 533; 534;
535: 535; 536: 540; 545; 588; 590
zonatus, Palicus 586
ADDENDUM
Specimens of Palicidae collected in Fiji by the BORDAU 1 and Suva 2 expeditions were examined while this
paper was in press. The specimens are listed below. Seven of the ten species listed (printed m bold) are new records
for Fiji.
Miropalicus vietnamensis (Zarenkov. 1968); BORDAU 1. stn CP 1411. 16°05'S. '^28^390-403 m:
1 juv. 2 (MNHN-B 27650). - Stn CP 1434, 17°11'S, 178‘41'W. 400-401 m: 1 <3 (MNHN-B 27651). -
Stn CP 1481, 20°57'S. I78°45'W, 441-506 m: 1 juv. 2 (MNHN-B 27652).
Znnrre MMHM Pane
610
P. CASTRO
Palicoides whitei (Miers, 1884): Suva 2, stn DW 62, Viti Levu, W lagoon. 17°49'S. 177°I2.9'E. 32 m:
1 juv. 9 (MNHN-B 27656).
Paliculus kyusyuensis (Yokoya, 1933): BORDAU 1, stn DW 1432, 17°20'S, I78°44’W. 477-493 m: I 9
(MNHN-B 27653). — Stn CP 1507, 18°09'S, 178°38’W, 294-300 m: 1 9 (MNHN-B 27654).
Parapalicus ambonensis Moosa & Serene, 1981: BORDAU 1, stn DW 1494, 18°55'S, 178°29'W.
240-319 m: 1 S (MNHN-B 27640). — Stn DW 1498, 18°41'S, 178°28'W, 300-307 m : 2 9 (MNHN-B 27641).
— Stn CP 1507, 18°09’S, 178°38'W, 294-300 m: 1 8 ; 5 9 (MNHN-B 27642).
Parapalicus armatus Castro, 2000: BORDAU 1, stn CP 1475, 19°41'S, 178°H'W. 321-424 m: 1 9
(MNHN-B 27643).
Parapalicus clinodentatus Castro, 2000: BORDAU 1, stn CP 1445. 1 7° 10'S, I78°42'W, 350-385 in: 6 6
(1 juv.); 3 9 (1 juv.) (MNHN-B 27644). — Stn CP 1446, 1 7° 1 l'S, 178°42'W. 350-367 m: 2 juv. 9 (MNHN-B
27645). — Stn CP 1500, 18°42'S, 178°26'W, 366-389 m: 2 6 ; 5 9 (MNHN-B 27646). —Stn CP 1501,
18°40'S, 178°30'W, 350-357 m : 5 $ ; 8 9 (2 juv.) (MNHN-B 27647).
Parapalicus inermis Castro, 2000: BORDAU 1, stn CP 1404, I6°40'S, 179°36'E. 180 m: 2 3 : 4 9 (MNHN-B
27648). — Stn CP 1405, 16°39'S, 179°36'E, 180 m: 1 juv. 9 (MNHN-B 27649).
Pseudopalicus declivis Castro, 2000: BORDAU 1, stn CP 1506, 18°09'S, 178°37'W. 294-300 nr 1 <3 ■
1 juv. 9 (MNHN-B 27638).
Pseudopalicus investigators (Alcock, 1900): BORDAU 1. stn DW 1473, 19°43'S, 178°I0'W. 270-288 nr
1 juv. 9 (MNHN-B 27639).
Pseudopalicus serripes (Alcock & Anderson, 1895): SUVA 2, stn CP 45. 17°51.6'S, 177°13.3'E 35 nr
1 juv. 9 (MNHN-B 27655).
Source : MNHN, Paris
FATS DCS CAMPAGNES MUSORSTOM, VOLUME 21 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 21
RESULTAT!
Pycnogonida: Pycnogonids from French cruises to
New Caledonia, Fiji, Tahiti and the Marquesas.
New records and new species
ROGER N. BAMBER
Department of Zoology
The Natural History Museum
Cromwell Road. London SW7 5BD
U.K.
ABSTRACT
Pycnogonid material from the Museum national d'Histoire naturelle, Paris, collected during French oceanographic
campymgns8°o New Caledon, a. Fiji. Taint, and the Marquesas in 1993. .996, .9 7 and £
species recorded. Anoplodactylus batangensis (Heifer. 1938) and A. pycnosoma (Heifer 1938) are ncwlyrecorded
New Caledonia; three of four recorded species of Pallenopsis are described as new (one of whl^ ^S ris descnhed from a
from Vanuatu); a new species of Austrodecus, representing the fifth spec.es of the subgenu jji tbtde °
single female. In the light of the intermediate morphology ol an Ammothella spec, men. and that of
elsewhere. A. indica Stock is synonymized with A. appendiculata Dohrn.
RESUME
Pycnogonida: Pycnogonides provenant de campagnes frangaises en Nouvelle-Caledonie, aux
Ties Fidji a Tahiti' et aux Ties Marquises. Nouvelles mentions et nouvelles especes.
E>es pycnogonides espfeceT idenidfides, ^AnofAodactylw
SSS.tr si esfmise ep syn„nymie avec A.
Dohrn.
n,MBro r N 2000 - Pycnogonida; Pycnogonids from French cruises to New Caledonia, Fiji. Tahiti and the
Matts New fetrand eat spfc.es. ,A C—
Memoires du Museum national d'Histoire naturelle, 184: 61 1-625. Pans ISBN---856. . -----
Snurr* MNHN Pans
612
R.N. BAMBER
INTRODUCTION
Pycnogonid material from the Museum national d'Histoire naturelle, Paris, collected during French
oceanographic campaigns to New Caledonia, Fiji, Tahiti and the Marquesas in 1993, 1996, 1997 and 1998, has
kindly been made available by Alain CROSNIER and Bertrand Richer de Forges [IRD. Institut de Recherche pour
le Developpement (ex ORSTOM) et Museum national d’Histoire naturelle, Paris] and is described herein.
There has been a number of French benthic marine research campaigns in the vicinity of New Caledonia since
the late 1970s. Pycnogonid material from these surveys has been assessed and described by Stock (1991; 1997),
who recorded 67 species for the region, many from deeper waters (continental slope). Other recent studies of the
New Caledonia pycnogonid fauna were those of CHILD (1977) and Bamber (1997b), which dealt largely with
shallower-water species.
There have been numerous recent publications on the pycnogonid fauna of other islands in the Indo-west and
southern Pacific Oceans, notably by CHILD and, in shallow waters, Muller, listed in, and including Stock
(1994).
The present material comprises ten species; four are from shallow waters and include two species of
Anoplodactylus new to New Caledonia; the six species from deeper water beam trawl samples include four species
of Pallenopsis, three of which are described as new, together with a previously undescribed species of Austrodecus
of the New Caledonia-endemic subgenus Tubidecus.
Material is deposited at the Museum national d'Histoire naturelle, Paris (MNHN).
SYSTEMATIC ACCOUNT
Tire species recorded are:
Family Colossendeidae: Colossendeis minor Schimkewitsch. 1893;
Family Phoxichilidiidae: Anoplodactylus batangensis (Heifer, 1938); A. pycnosoma (Heifer, 1938);
Family Callipallenidae: Pallenopsis ( Bathypallenopsis ) richeri sp. nov.; P. (Pallenopsis) sexacentra
sp. nov.; Pallenopsis (P.) kupei Clark, 1971; P. (P.) mascula sp. nov.;
Family Ammotheidae: Achelia assimilis (Haswell, 1885 )\ Ammothella appendiculata (Dohrn, 1881);
Family Austrodecidae: Austrodecus (Tubidecus) oferrecans sp. nov.
Higher taxonomy follows that of Arnaud and Bamber (1987) except for the genus Pallenopsis which is now
considered to be a member of the Callipallenidae.
Family COLOSSENDEIDAE Hoek. 1881
Genus COLOSSENDEIS Jarzynsky, 1870
Colossendeis minor Schimkewitsch, 1893
Colossendeis minor - STOCK. 1984: 702-704. figs 1-6.
Py 825>rERIAL EXAM,NED- ~ Sncidy Islands. Zepolyf 1: 19°15.5'S, 15I°35.4’W. 618 m: I specimen (MNHN-
1 spSnUe(MNHNl?yd826rS°RSTOM * ^ ^ ^ P0U' 09°20'S’ 140°°8'W’ 85°-905 m' 03.09.1997:
Remarks. - Recorded in this region by STOCK (1991), from 1300 to 21 10 m; a widely distributed Indo-
Pacific species known also from the North-east Atlantic, with a depth range of 1000 to 4850 m The present
records are therefore the shallowest for this species.
Source : MNHN , Paris
PYCNOGONIDS FROM THE SOUTH PACIFIC
613
Family PHOXICHILIDIIDAE Sars, 1891
Genus ANOPLODACTYLUS Wilson, 1878
Anoplodactylus batangensis (Heifer, 1938)
Anoplodactylus batangensis - Child, 1992: 41-42, fig. 18.
MATERIA qui m'est imputable.£DiTiON Montrouzier: East Lagoon, Touho (Haut-Fond de Tie), 20°50S. 165°20'E.
5-7 m, 19.09.1993: 1 male, 1 female (MNHN-Py 827).
REMARKS. — A pantropical-temperate species, known from the Caribbean, Australasia, W Africa, Pacific and
Indian Oceans, from shallow water to 40 m. It has a distinctive upturned tapering or styliform proboscis without
any processes in the female. Not recorded previously from New Caledonia.
Anoplodactylus pycnosoma (Heifer, 1938)
Anoplodactylus pycnosoma- MllLLER, 1992: 173, figs 43-48.
Material examined.
1 female (MNHN-Py 828). -
(MNHN-Py 829).
— New Caledonia. South West Lagoon, Recif Ricaudy. 22°20'S. 166°25'E, 10.04.1993:
- East Lagoon. Touho (ilot de Sable), 20°50'S. 165°20’E. 5-7 m. 06.09.1993: 1 temale
REMARKS. — While isolated females of this genus are difficult to attribute to species, these specimens are
entirely consistent with HELFER's Indo-West Pacific species, recorded from the shore down to over 100 m. Not
recorded previously from New Caledonia.
Family CALLIPALLENIDAE Hilton. 1842
Genus PALLENOPS1S Wilson. 1881
Pallenopsis (Bathypallenopsis) richeri sp. nov.
Fig. 1
Pallenopsis ( Bathypallenopsis ) ?oculotuberculosis - STOCK, 1997: 406-407. fig. 7 (non P. oculotuberculosis Hilton.
1942).
MATERIAL EXAMINED. — Fiji. Musorstom 10: stn CP 1361. 18°00.00'S. 178053.71'E. 1058-1091 m.
13.08.1998: 1 female, holotype (MNHN-Py 830).
DESCRIPTION —Trunk (Fig. 1A) elongate, fully segmented, lateral processes well-separated, naked; cephalon
less than half trunk length, narrowing anterior to origin of first lateral processes where ovtgers ar.se. ocular
tubercle a medium dome, with eyes, of which anterior pair larger, and minute distolateral tubercles. Abdomen
naked, exceeding 4th lateral process but not achieving distal edge ot first coxa of 4th leg.
Proboscis cylindrical, widest at mid-point, 0.55 times total trunk length. , , .
Chelifores (Figs IB, C) with 2-arlicled scape, proximal article 1.34 times length of distal article; chela typical
of genus, fingers without teeth, longer than palm, slender, tips crossing; no setose pad at base of moveable finger.
Palps small, one-articled, ventrolateral knobs. . .
Ovmer (Fi«. IE) 10-articled, articles 4 and 5 subequal and dorsally sp.nose; articles 6 to 9 with simple spines
as figures; article 10 (Fig. IF) with 3 inner spines and larger curved terminal spine with appearance ol a terminal
claw.
Snurne MNHN Pans
614
R.N. BAMBER
FlGa'd ncheris p nov., holotype female: A. trunk, dorsal; B. cephalon, proboscis
I f? RC,Hn ’ D’ 3rd '*g; E- ov,ger: F- dlslal oviSer arIicle; propodus and claws, 4th leg. Scale line =
2 mm for A. B and D, 1 mm for C and E, 0.2 mm for F and 0,7 mm for G.
Source : MNHN, Paris
PYCNOGON1DS FROM THE SOUTH PACIFIC
615
Third leg (Fig. ID): coxa 1 short; coxa 2: 3 times length of coxa 1; coxa 3: 0.4 coxa 2; femur straight, simple,
with dorsodistal spine group, tibia 1 subequal to femur, tibia 2 longest article, tibiae with spines and setae not as
long as diameter of tibia and not denticulate, dorsally on tibia 1 and dorsally and ventrally on tibia 2; tarsus small;
propodus (Fig. 1G) with 2 unequal heel spines and 6 to 8 sole spines, the distal pair laterally adjacent: main claw
0.63 times propodus length and 4 times length of auxiliary claws.
Male unknown.
Measurements of holotype female (mm). — Trunk length: 5.85; proboscis length (ventral): 3.19; ehelifore
scape article 1: 2.19, article 2: 1.63.
Third leg: coxa 1: 0.8; coxa 2: 2.88; coxa 3: 1.08; femur: 7.2; tibia 1: 7.6; tibia 2: 10; tarsus: 0.2; propodus:
1.54; main claw: 0.97.
Etymology. — Named after Bertrand RICHER DE FORGES of IRD. who collected most of the material
available for study.
Remarks. — STOCK (1997) recorded and described a specimen (MNHN-Py 932), also a female, Irom
Vanuatu (15°53.81'S, 162°52.19'E) at 1100-1191 m depth, which he attributed tentatively to Hilton's
P oculotuberculosis, a species recently redescribed by Child (1994). The latter species is characterized by
prominent lateral tubercles on the tall ocular tubercle and article 1 of the ehelifore scape exceeding the proboscis.
The present species has a lower ("normal") ocular tubercle and the proximal scape article is far shorter than the
proboscis (0.7 times dorsal proboscis length). While the lateral tubercles on the ocular tubercle are more prominent
in Stock’s specimen than in the present specimen, they do not suggest the bifurcation lhat those of Hilton s
species do, and I conclude that the Vanuatu and Fiji specimens are conspecific and distinct from
P. oculotuberculosis. .
There arc few Pallenopsis species with only two major heel spines, although current studies ol P. tritoms
Hoek 1883 show this feature to be variable, partly with maturity (unpubl. data). The conformation of the
proboscis places P. richeri sp. nov. within Stock’s (1975) “tydemani- group”; it is distinct from P. tydeman,
Loman 1908 owing to its well-developed ocular tubercle with eyes, propodal spination. proportion of scape
articles’ and oviger supination, and from P. tritoms by the proboscis length to trunk length proportions, proportion
of scape articles and the lack of denticulate spines on the longer leg articles. Analysis of the leg article proportions
suggests closest affinity to P. californica Schimkewitsch, 1893 (see STOCK, 1975. for redescript.on), but that
species has a much longer main claw, dense long setae on the longer leg articles, and distinct oviger spination.
P. richeri sp. nov. is known from the Melanesian islands of Fiji and Vanuatu, at depths between 1058 and
1 191 m.
Pallenopsis (Pallenopsis) sexacentra sp. nov.
Figs 2-4
Material examined. — Fiji. Musorstom 10: stn CP 1358. 17°48.49'S 178046_7°'E, 80-|20 m. 13.08.1998:
1 male carrying eggs, holotype (MNHN-Py 831); 1 male and 3 females, paratypes (MNHN-Py 832).
DESCRIPTION OF Male. - Trunk (Figs 2A-B) elongate, fully segmented, with paired mid-dorsal spines at
posterior margin of trunk segments 1 to 3; lateral processes separated by just less than their own width, first 3
with paired dorsodistal spines, fourth with 3 or 4 such spines. Cephalon half length of trunk, narrowing anteriorly;
ocular tubercle dome-like, eyes pigmented, anterior pair larger than posterior pair. Abdomen about as long as 4th
lateral process, armed with paired dorsodistal spines.
Proboscis stout, shorter than cephalon, with sparse short spines on distal half.
Chelifore scape 1 -articled, exceeding proboscis. Chela (Fig. 2C) setose, moveable finger with setose basal pad.
fixed finger with crenulated cutting edge. Palps small, one-articled, ventrolateral knobs.
Oviger (Figs 3B-C) 10-articled; article 1 short; article 2: 3 times as long as wide; article 3: 0.54 times article
article 4 twice article 3, with mid-ventral indentation; article 5: 1.2 times article 4, with rows of stout setae; distal
articles (Fig. 3C) each with numerous stout setae, pointing predominantly prox.mally. No terminal claw.
Source :
616
R.N. BAMBER
Fig. 2. — Paltenopsis ( Pallenopsis) sexacentra sp. nov.: A. holotype male, trunk, dorsal; 1$. paratypc male, trunk,
dorsolateral; C, right chela. Scale line = 2 mm for A and B. 0.4 mm for C.
Source : MNHN, Paris
PYCNOGONIDS FROM THE SOUTH PACIFIC
617
Third leg (Fig. 4A) elongate; coxa 1 (also Figs 2A, B) with 4 dorsodistal tubercles each bearing short distal
seta; coxa 2 nearly 4 times coxa 1 length, with conspicuous dorsal genital pore in distal third and venlrodistal
setose tubercle ("spur"); coxa 3 longer than coxa 1 . Femur sparsely setose, without distal spur, slender mid-ventral
cement-gland tube just longer than femoral diameter; tibia I as long as femur but more slender, tibia 2 longest.
Tarsus (Fig. 4B) short, ventrodistal spines stout; propodus elongate, slightly curved. 3 heel spines progressively
larger distally, 12 to 14 small sole spines, elongate setae distally and dorsally; main claw 0.53 times length of
propodus, auxiliary claws 0.8 of main claw.
FIG. 3. — Pallenopsis (Pallenopsis) sexacenira sp. nov., paratypes; oviger of A. female and B. male (distal setation
semidiagrammatic); distal oviger articles of C, male and D, female. Scale line = 1 mm lor A and B. 0.23 mm lor C
and D.
DESCRIPTION of FEMALE. — Generally similar to male, but femur slightly stouter and without cement gland.
Oviger (Fig. 3A) much smaller (Figures 3A and B are to the same scale from similarly-sized, mature animals), of
10 articles, distal five articles (Fig. 3D) with numerous, predominantly ventral, stout setae, all pointing distally.
Measurements of holotype male (mm). — Trunk, length: 3.8; proboscis length (ventral): 1.48; chelifore scape.
1. 84.
Third leg: coxa I: 0.51; coxa 2: 1.79; coxa 3: 0.76; femur: 4.15; tibia I: 4.29; tibia 2: 4.68; tarsus: 0.1 I;
propodus: 0.87; main claw: 0.48; auxiliary claw: 0.33.
ETYMOLOGY. — from the Latin “six prickled”, referring to the mid-dorsal trunk spines.
Remarks. — A number of shallower-water (30 to 300 m) species of this subgenus have been recorded from
this region (e.g. STOCK 1983; 1991; 1997) of a similar overall morphology, including Pallenopsis spimpes
Carpenter 1907. P. dentifera Stock, 1983 and P. virgata Loman, 1908. P. spimpes is distinct in its touching first
Source : MNHN. Paris
618
R.N. BAMBER
Fig. 4. — Pallenopsis (Pallenopsis) sexacentra sp. nov., male holotype, third leg. A, entire and B. detail of distal
articles. Scale line = 2 mm for A. 0.4 mm for B.
pair ot lateral processes, and in having barbed spines on the legs. P. dentifera has a similar crenulation of the
cutting edge of the chela fixed finger as does P. sexacentra sp. nov., but is without spines on the trunk segments
or lateral processes. P . virgata has setose/spinose tubercles distally and laterally on the femur and tibia I . There are
subtle differences in the propoda! spination of these species. None, however, has the paired mid-dorsal trunk spines
shown by P. sexacentra sp. nov. nor the degree of development of dorsodistal setose tubercles on coxa 1 .
Pallenopsis (Pallenopsis) kupei Clark, 1971
Pallenopsis kupei - Clark. 1971: 430-433, fig. 2. — Child, 1995: 143.
Material EXAMINED. — Fiji. Musorstom 10: stn CP 1344, 16°45.26’S. 177°40.53'E, 588-610 m, 10.08.1998'
1 female (MNHN-Py 834).
Remarks. — A species previously recorded from the Antarctic north to northern New Zealand, at depths
between 121 and 1097 m.
Pallenopsis ( Pallenopsis ) mascula sp. nov.
Figs 5-6
Material EXAMINED. — Fiji. Musorstom 10: stn CP 1364, 18°11.95'S, 178°34.50'E. 80-86 m. 15 08 1998-
1 male, holotype (MNHN-Py 835).
Source : MNHN. Paris
PYCNOGONIDS FROM THE SOUTH PACIFIC
619
DESCRIPTION. — Trunk (Figs 5A-B) elongate, slender, fully segmented, moult suture apparent laterally on all
segments, cuticle finely reticulated; cephalon tapering anteriorly, ocular tubercle tall, with eyes, mounted at
anterior of cephalon; lateral processes long, separated by about their own width, with anterior and posterior lateral
setae and three slender dorsodistal tubercles bearing setae, central one longest, about as long as lateral process
diameter. Abdomen with basal articulation, exceeding first coxae of fourth legs and as long as iirst 2 trunk
segments, with 3 pairs of dorsal setose tubercles.
FIG. 5. - Pallenopsis (Pallenopsis) mascula sp. nov., holotype male: habitus. A lateral and B dorsal; C. oviger. Scale
line = 2 mm for A and B, 0.95 mm for C.
Source MNHN. Paris
620
R.N. BAMBER
Proboscis short, cylindrical, densely setose.
Chelifore scape without articulation, setose distally, without tubercles; chela (Fig. 6A) with rectangular,
densely setose palm, fingers anaxial. denticulate, setose cushion at base of moveable finger.
Palp conspicuous, semicircular, with basal articulation and distal seta.
Oviger (Fig. 5C) 10-articled, all but first article setose, article 2 slightly inflated with 2 distal setose tubercles,
article 3 short, slightly inflated, article 4 elongate, straight, article 5 as long as article 4 with proximally directed
setae; distal five articles heavily setose, article 10 more than twice as long as wide.
Leg (Figs 6B-C) moderately elongate, setose with most setae longer than article diameters; coxa I with lateral
setae and 4 or 5 dorsodistal setose tubercles; coxa 2 setose, without tubercles, 3 times length of coxa I ; coxa 3 half
length of coxa 2; femur with few dorsal and single dorsodistal setose tubercles, with midventral cement-gland tube
twice as long as femur diameter and mounted on large eminence; tibiae densely setose with rows of dorsal setose
tubercles, tibia 1 longer than tibia 2; tarsus short with 4 short “sole setae", 2 ventral and 1 dorsal distal spines;
propodus with 2 heel spines lying close to propodal ventral margin, about 12 sole spines, 3 dorsal setae and
5 distal setae above main claw; main claw slender, curved, two-thirds length of propodus, auxiliary claws lorn-
slender, curved, almost as long as main claw. c
Female unknown.
F,G'c6' ~Pal,en°Psis ( Pallenopsis ) mascula sp. nov.. holotype male: A. chela; B, 4th leg; C. distal articles of 3rd le»
Scale line = 0.75 mm for A and C, 2 mm for B.
Source : MNHN, Paris
PYCNOGONIDS FROM THE SOUTH PACIFIC
621
Measurements of holotype male (mm). — Trunk length: 3.42: trunk width across 2nd lateral processes: 2.35;
proboscis length: 1.15; ocular tubercle height: 0.84; abdomen length: 1.93.
Fourth leg: coxa 1: 0.57; coxa 2: 1.81; coxa 3: 0.91; femur: 2.91: tibia 1: 3.51; tibia 2: 3.2; tarsus: 0.1 I ;
propodus: 0.97; main claw: 0.66; auxiliary claws: 0.57.
ETYMOLOGY. — From the Latin for “male”, alluding both to the extremely well-developed cement-gland
mound and tube, and the dense “beard” of setae on the proboscis.
Remarks. — Of a number of species of Pallenopsis with long setae mounted on tubercles on the longer leg
articles, the new species is closest to P. cidaribatus Child, 1975, recorded from Western Australia at depths
between 100 and 141m. Both are distinct from the other species in having three slender setose tubercles, each
bearing a long distal seta, around the dorsodistal margin of the lateral processes, together with a "crown” of such
tubercles distally on the first coxa. Those of P. cidaribatus are much longer; but with only three tubercles on
coxa 1 compared with four or five in P. mascula. Further, the leg tubercles of P. cidaribatus are longer than those
of the Fiji species; conversely, the cement gland tube of P. mascula is mounted on a femoral eminance and
unusually long, at 1 mm more than twice the diameter of the femur (about as long as femoral diameter in
P. cidaribatus). P. cidaribatus has unusual paired tubercles lateral to the ocular tubercle (absent in P. mascula).
a distal oviger article no longer than wide (much longer than wide in P. mascula). chela fingers without
denticulation (heavily denticulate in P. mascula) and proportionately shorter claws on the propodus.
Family AMMOTHEIDAE Dohrn, 1881
Genus A CHELIA Hodge, 1864
Achelia assimilis (Haswell, 1885)
Achelia assimilis - Stock, 1994: 32-33, fig. 9.
MATERIAL EXAMINED. — New Caledonia. Lagoon: Recif Mbere, external slope. 22°19.9'S. 166°13.2'E.
10-15 m, 07.01.1993: 1 male (MNHN-Py 837). — East Lagoon: Touho (Haut-Fond de Tie). 20°50'S. 165°20'E. 5-7 m.
19.09.1993: 1 male (MNHN-Py 838).
Remarks. — A species which ranges from 0 to 900 m depth, widely recorded from the Indo-West Pacific,
Australia, New Zealand, the Subantarctic and coasts of Chile and Argentina; previously recorded from New
Caledonia by CHILD (1977) and STOCK (1997).
Genus AMMOTHELLA Verrill, 1900
Ammothella appendiculata (Dohrn. 1881)
Ammothea appendiculata Dohrn, 1881: 152-155. pi. 7.
Ammothella indica Stock. 1954: 113-119, figs 54-57 (new synonymy).
MATERIAL EXAMINED. — New Caledonia. Grotte Merlct. outer slope of barrier reef. 22°43.50'S. 166°41.00'E.
20-30 m, 19-21.01.93: 1 female (MNHN-Py 839).
Remarks. — The problems of distinguishing A. indica Stock from A. appendiculata (Dohrn) were
highlighted by STOCK himself in his original description of the former (loc. cit.); characters such as the density of
blunt "tubular” spines on the leg articles, and the relative proportions of the chelifore scape articles, are somewhat
subjective, while the presence or absence of blunt spines on the proximal scape article shows no congruence with
Source :
622
R.N. BAMBER
the typical expression of these other features (see, for example, Bamber. 1997a). It is also the case that the
expression of these features varies with maturity.
The present fully mature female, from typical A. indica geographic range, has slender leg articles (containing
eggs), sparsely furnished with blunt spines (an A. appendiculata character), scape article 2 flared, distally fully
enclosing the vestigial chela, and about twice the length of article 1 (an A. appendiculata character); the latter bears
a single blunt spine (characteristic of A. indica)-, lateral processes are not markedly elongate (width across 2nd
lateral processes 2.3 limes 2nd trunk segment width, compared with three times in A. indica).
With the frequency of specimens showing intermediate morphology, and the inconsistency of occurrence of the
characterizing features, I see no reason to maintain further the distinction of these two species. STOCK (1997)
recorded 1 male, as A. indica, from Recif Senez at 7 m.
Family AUSTRODECIDAE Stock, 1954
Genus AUSTRODECUS Hodgson, 1907
Austrodecus ( Tubidecus ) oferrecans sp. nov.
Fig. 7
Material EXAMINED. — New Caledonia. Halipro 2, stn BT 60, 24°52'S, 168°44'E, 1 133-1280 m
18.11.1996: 1 female, holotype (MNHN-Py 841).
Description. — Trunk (Figs 7A-B) typical of the subgenus, completely segmented, with the segment length
anterior to the lateral processes increasing from the cephalon to trunk segment 4 (thus 2nd and 3rd lateral processes
separated by about their own width); lateral processes less than twice as long as trunk diameter, and bearing
tuberculation on dorsal and lateral surfaces. Cephalon with tall, slender anterior ocular tubercle with distal
tuberculation and eyes; slender mid-dorsal post-ocular tubercle either short or (presumed) broken. Remaining trunk
segments with tall, slender (filiform) mid-dorsal trunk tubercles, each bearing subdistal seta. Abdomen reaching
distal edge of coxa 1 of 4th leg.
Proboscis (Fig. 7B) slender and tapering, nearly twice as long as trunk length, distally down-curved, annulated.
Palp (Figs 7A-B and D) of 6 articles, articles 2 to 4 bearing dense thorn-like tuberculation; article 2 longest, as
long as trunk, article 4 length 60% that of article 2, bearing 6 inner slightly-curved spines; articles 5 and 6
anaxially articulated, short, with distal setae.
Chelifores absent.
Oviger (Fig. 7C) small, of 4 articles, articles 2 and 3 with single distal setae, article 4 twice length of article 2
with single medial and 3 distal setae (spinules).
Legs (Fig. 7E) with stout, tuberculate coxae; coxa 1 armed distodorsally with 1 short and 1 long filiform,
irregular tubercle, the latter with subdistal seta; coxa 2 nearly twice length of coxa 1; coxa 3 shortest, with
irregular filiform dorsal tubercle. Femur slender, tuberculate, longer than either tibia; tibiae elongate, slender, not
tuberculate, tibia 1 longer than tibia 2 and with dorsodistal spine, tibia 2 with spine on swollen base distal of mid¬
dorsal; distal leg articles (Fig. 7F), tarsus short, rectangular, with 3 ventral spines; propodus slender, elongate,
with numerous sole setae, decreasing in number from anterior to posterior legs (23 on 2nd, 12 on 3rd). 3 or
4 dorsal spines and 3 distal spines; main claw short, blunt, one-quarter the length of the propodus; auxiliary claws
absent.
Male unknown.
Measurements of female (mm). — Trunk length: 1.34; width across 2nd lateral processes: 1 .0; ocular tubercle
length: 0.855; abdomen length: 0.23.
Third leg: coxa 1: 0.23; coxa 2: 0.4; coxa 3: 0.14; femur: 1.25; tibia I: 1.17; tibia 2: 0.80; tarsus- 0 04-
propodus: 0.46; claw: 0. 1 1 3.
Source : MNHN, Paris
PYCNOGONIDS FROM THE SOUTH PACIFIC
623
FIG
i — Austrodecus (Tubidecus) oferrecans sp. nov., holotype female: A, habitus, dorsal and. B. lateral, C ovtger;
. distal articles of palp, dorsal; E, 3rd leg; F. distal articles of 2nd leg. Scale line = I mm for A. B and E. 0.17 mm
r D and F; 0.3 mm for C.
Source : MNHN, Paris
624
R.N. BAMBER
ETYMOLOGY. — From the Anglo-Saxon ofer meaning "beyond” and raecan to reach or extend (thus "over¬
reach"), alluding to the elongate palps of this species.
Remarks. — The only described Tubidecus species with a postocular dorsal trunk tubercle on the cephalon is
A. (T.) tuberculatum Stock, 1991, but that species has a distinctly longer abdomen, more than 2 distodorsal
tubercles on the 1st coxa of legs 1 to 3, a less tuberculate palp and markedly less slender leg articles. The present
species resembles more closely A. (T.) bathyale Stock, 1991 (known only from the male), from which it is
distinguished by the postocular cephalon tubercle, the main claw and tarsus being proportionately shorter, coxa 2
being less elongate, and the very long palps [1.8 times the trunk length in A. (T.) oferrecans sp. nov., only 1.42
times in A. (T.) batliyale]. While the coxal proportion may be a gender-related morphology, these other features are
not attributable to sexual dimorphism.
In a number of respects the present species is intermediate between these two of Stock’s species; however, as
only 7 individuals of these three taxa are at present known (indeed, only 20 specimens of the five described
Tubidecus species), it is not feasible as yet to suggest that the present species and A. (T.) bathyale (recorded from
1665-1590 m) represent attenuate deeper-water phenotypes of A. (T.) tuberculatum. The present specimen is
presently considered distinct, characterized particularly by the exceedingly elongate palps.
The subgenus remains confined to the New Caledonia region.
REFERENCES
Arnaud, F. & Bamber, R.N., 1987. — The Biology of Pycnogonida. Advances in Marine Biology, 24: 1-96. Academic
Press, Inc., London.
Bamber, R.N.. 1997a. — Pycnogonids (Arthropoda: Pycnogonida) from the Cape d'Aguilar Marine Reserve, Hong Kong.
In: The Marine Fauna and Flora of Hong Kong and Southern China, IV (ed. B. Morton.), pp. 143-157. Proceedings of
the 8tli International Marine Biological Workshop: the Marine Flora and Fauna of Hong Kong and Southern China,
Hong Kong, 2-20 April 1995. Hong Kong, Hong Kong University Press.
Bamber, R.N., 1997b. — Two pycnogonids (Arthropoda) from the Coral Sea Region: a new Callipallene and a
reassessment of the Anoplodactylus cribellatus complex. Species Diversity. 2: 43-50.
Child, C.A., 1977. — On some Pycnogonida of French Oceania. Proceedings of the Biological Society of Washington,
90 (2): 440-446.
Child, C.A.. 1992. — Shallow-water Pycnogonida of the Gulf of Mexico. Memoirs of the Hourglass Cruises. 9(1): 1 -86.
Child, C.A., 1994. — Deep-sea Pycnogonida from the temperate West coast of the United States. Smithsonian
Contributions to Zoology, (556): 1-23.
Child, C.A., 1995. — Antarctic and subantarctic Pycnogonida: III, The family Nymphonidae. IV. The families
Colossendeidae and Rhynchothoraxidae, V, The families Pycnogonidae, Phoxichilidiidae, Endeididae and
Callipallenidae. including the Genus Pallenopsis. Biology of the Antarctic Seas XXIV (Ed. S.D. Cairns). American
Geophysical Union, Antarctic Research Series, 69: 1-165.
Clark, W.C., 1971. — Pycnogonida of the Antipodes Islands. New Zealand Journal of Marine and Freshwater Research.
5 (3 & 4): 427-452.
Dohrn. A., 1881. — Die Pantopoden des Golfes von Neapel und der angrenzenden Mceresabschnitte. Monographic der
Fauna und Flora des Golfes von Neapel. 3: 1-252; I-XVIII.
MOller, H.G., 1992. — Pycnogonids from Malaysian coral reefs, including descriptions of three new species. Bonner
Zoologisclie Beitrage, 43: 155-178.
Stock, J.H.. 1954. — Pycnogonida from Indo-West-Pacific, Australia and New Zealand waters. Papers from Dr Th
Mortensen s Pacific Expedition 1914-1916. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i
Kjobenhavn, 116: 1-168.
Source : MNHN. Paris
PYCNOGONIDS FROM THE SOUTH PACIFIC
625
STOCK, J.H., 1975. — Pycnogonida from the continental shelf, slope, and deep sea of the tropical Atlantic and East
Pacific. Biological results of the University of Miami deep-sea expeditions, 108. Bulletin of Marine Science. 24(4):
957-1092.
STOCK, J.H., 1983. — Pycnogonides des campagnes Corindon et Musorstom II (detroit dc Makassar et Philippines), avec
description de Pallenopsis dentifera sp. nov. Bulletin du Museum national d'Histoire naturelle, Paris, ser. 4. 5. Sect.
A, (1): 299-305.
Stock, J.H., 1984. — The deep-water Pycnogonida of the Safari cruises to the Indian Ocean. Bulletin du Museum national
d'Histoire naturelle. Paris, ser. 4. 6. Sect. A, (3): 701-709.
Stock. J.H., 1991. — Deep-water Pycnogonida from the surroundings of New Caledonia. In: A. Crosnif.r (ed. ). Rcsultats
des Campagnes MUSORSTOM, Vol. 8. Memoires du Museum national d'Histoire naturelle. Paris. (A). 151: 125-212.
Stock, J.H., 1994. — Indo-West Pacific Pycnogonida collected by some major oceanographic expeditions. Beaufortia,
44 (3): 17-77.
STOCK, J.H., 1997. — Pycnogonida collected in recent years around New Caledonia and Vanuatu. In: A. Crosnier (cd.).
Resultats des Campagnes Musorstom. Vol. 18. Memoires du Museum national d'Histoire naturelle, Paris. (A). 176:
389-409.
Source :
Source : MNHN, Paris
LTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 21 — RESULTATS
Echinodermata Crinoidea: Comatulid Crinoids
of the KARUBAR Expedition to Indonesia.
The families Comasteridae, Asterometridae,
Calometridae and Thalassometridae
Charles G. MESSING * Nadia AMEZIANE**
& Marc ELEAUME**
*Nova Southeastern University Oceanographic Center
8000 North Ocean Drive
Dania, FL 33004 USA
**Museum National d'Histoire Naturelle
Laboratoire de Biologie des Invertebrds Marins et Malacologie
55 rue Buffon, F-75005 Paris Cedex 05
ABSTRACT
Fifteen species of comatulid crinoids in eleven genera and four families collected in 180-800 m by the joint French-
Indonesian Karubar Expedition to the Kai and Tanimbar Islands, Indonesia (October 991). are described in de ail. The
specimens represent the most important collection of bathyal coma.uhds from the East Indies in the second I half of the
twentieth century. The material described herein comprises four species of Comasteridae. one Asterometridae. two
Calometridae and eight Thalassometridae. One species of calometrid (Neometra xenocladia sp. nov.) is described as new.
Five species are recorded for the first time since they were originally collected. Four thalassometnds '^onietra va .
Oceanometra annandalei, Cosmiometra phihppmensls and Stenometra casta, a) together account for 75% of .den .Bed
specimens, reflecting this family's importance to the outer shelf-upper bathyal comatulid tauna ol the > tropical Ind°-
Westem Pacific. The substantial amount of material of several species, notably .4. valuta and O
better understanding of morphological variability than previously. Statistical analyses of several supposedly diagnost c
characteristics, including aspects of calyx form and number of cirri, reveal substantial variation,
synonymy as a result. SEM studies of different ossicles have been made for the first time for 4. valuta and O. annandalei.
The preliminary results show that great morphological differences exist within the family. , , ■
Three species (SUremetra breviradia. Palaeocomatella hiwia and Cosmo, me, ra tote) are recorded from J^newan
waters for the first time. Moreover, most of the Karubar comatul.ds represent geographical and bathymetrical range
extensions.
Messing C G Am£ziane, N. & Eleaume, M„ 2000. - Echinodermata Crinoidea: Comatulid Crinoids of the
Karubar Expedition to Indonesia. The families Comasteridae Asterometridae, Calometridae and Thal^som^ nda^
In, A. CROSNIER (ed.). Resultats des Campagnes MUSORSTOM. Volume 21. Memo, res du Museum national d Hisioiu
naturelle. 184: 627-702. Paris ISBN-2-85653-526-7.
Source MNHN, Parts
628
C.G. MESSING. N. AMEZIANE & M. ELEAUME
RESUME
Echinodermata Crinoidea : Comatules recoltees lors de la campagne Karubar en Indonesie.
Families Comasteridae, Asterometridae, Calometridae et Thalassomelridae.
Quinze especes dc comatules. appartenant it 11 genres el quatre families, recoltees en octobre 1991. entre 180 et
800 m. lors de la campagne franco-indonesienne Karubar aux lies Kai et Tanimbar (Indonesie) sont decrites en detail.
Les echantillons recoltes forment la collection de comatules bathyales la plus importante faitc dans la region durant la
seconde moitie du XXeme siecle. Le materiel etudie ici comprend quatre esp&ces de Cosmateridae. une d'Asteromelridae.
deux de Calomatridae et huit de Thalassometridae. Une espece. Neometra xenocladia, est nouvclle. Cinq sont sianalces
pour la premibre fois depuis leur description. Quatre especes appartenant a la famille Thalassometridae. Aglaometm
valida, Oceanometra annandalei, Cosmiometra philippinensis et Stenomeira cristata. representent 75% des specimens
recoltes. montrant l'importancc de cette famille dans la faune des comatules de la zone bathyale superieure dc la petite
continentale dans 1 lndo-ouest Pacilique. Les recoltes abondantes de plusieurs especes, notamment Aglaometra valida et
Oceanometra annandalei, ont permis de mieux apprehender la variabilite de certains caracteres morphologiques. Des
analyses statistiques de plusieurs caractdres supposes diagnostiques, notamment la forme des calices et le nombre de
cirres, ont montre des variations substantiellcs. Ceci nous a conduit a etablir plusieurs synonymies. Des etudes au SEM
des dtlferents ossicules ont ete faitcs pour la premiere fois sur A. valida et O. annadelei. Les resultats preliminaires
montrent de grandes differences morphologiques it I'intdrieur d'une meme famille.
Trois especes. Stiremelra breviradia, Palaeocomatella hiwia et Cosmomietra iole sont signalees dans les eaux indo-
nesiennes pour la premiere lois. De plus, la plupart des especes mentionnees voient leurs repartitions geographique et
bathymdtrique etendues.
CONTENTS
Introduction .
Checklist of Comatulids .
Station list .
Species list .
Material and Methods .
Statistical analysis .
Comatulid Morphology, Measurements and Symbols ..
General Form and Orientation .
Centrodorsal .
Cirri .
Rays .
Pinnules .
Disk . .. ....
Systematic Account .
Superfamily COMASTERACEA .
Family COMASTERIDAE A.H. Clark. 1908 .
Genus APHANOCOMASTER Messing, 1995
Aphanocomaster pulcher (A.H. Clark, 1912) .
Genus PALAEOCOMATELLA A.H. Clark, 1912
1 Palaeocomatella difficilis (Carpenter, 1888) .
Palaeocomatella hiwia McKnight. 1977 .
Genus PHANOGEN1A Loven, 1866
Phanogenia sp .
Unidentified comasterid .
Superfamily TROPIOMETRACEA .
Family ASTEROMETRIDAE Gislen, 1924 .
Genus ASTEROMETRA A.H. Clark, 1907 .
Asterometra longicirra (Carpenter, 1888) .
. 629
. 631
. 631
. 632
. 634
. 634
. 635
. 635
. 636
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Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
629
Family CALOMETRIDAE A.H. Clark, 1912 .
Genus NEOMETRA A.H. Clark, 1912 .
Neometra cf. alecto (A.H. Clark, 1911) .
Neometra xenocladia C.G. Messing, sp. nov.
Family THALASSOMETRIDAE A.H. Clark, 1908
Genus AGLAOMETRA A.H. Clark, 1913 .
Aglaometra valida (Carpenter, 1888) .
Genus COSMIOMETRA A.H. Clark. 1909 .
Cosmiometra iole A.H. Clark, 1950 .
Cosmiometra philippinensis A.H. Clark, 1911 ...
Genus CROTALOMETRA A.H. Clark, 1909 .
Crolalometm rustica A.H. Clark, 1909 .
Genus OCEANOMETRA A.H. Clark, 1916 .
Oceanoinetra annandalei (A.H. Clark, 1909)
Genus STENOMETRA A.H. Clark. 1909 .
Stenometra cristata A.H. Clark, 1911 .
Slenoinetra quinquecostata (Carpenter, 1888)
Genus STIREMETRA A.H. Clark, 1909 .
Stiremetra breviradia (Carpenter. 1888) .
Biogeography .
Summary of proposed taxonomic Changes .
Acknowledgments .
References .
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INTRODUCTION
Comatulid crinoids form an ecologically important component of benthic communities in many areas ol
Recent seas. Because many species, particularly those from deeper water, arc known from only a few specimens,
their morphological variability, taxonomic relationships, behavior and habits remain poorly understood.
The comatulids dredaed by the joint French-Indonesian Karubar Expedition to the Kai and Tanimbar Islands
(Kepulauan Kai and Kepulauan Tanimbar). Indonesia (October 1991 ), represent the only major collection of these
animals taken from balhyal East Indian waters during the second half of the twentieth century. Previous important
collections include those made by MORTENSEN (Danish, 1914-1916, 1922), and the "Challenger" (British. 1872-
1876), "Siboga" (Dutch, 1899-1900), "Albatross" ( U.S. Fish Commission, 1907-1910). and "Willebrord Snelhus"
(Swedish, 1929-1930) expeditions (see A.H. CLARK, 1918, 1931, 1941. 1947, 1950; A.H. CLARK and
A.M. Clark, 1967).
The following systematic account covers all collected material except that belonging to the families
Charitometridae (superfamily Tropiometracea) and Antedonidae (superfamily Antedonacea). The former in particular
requires a major revision before the KARUBAR collection can be meaningfully identified. Identified material
includes taxa belonging to the superfamilies Comasteracea and Tropiometracea as follows; Comasteridae (4 species
in 3 genera), Asteromctridae (1 species), Calometridae (2 species in 1 genus), and Thalassomelndae (8 species in 6
genera). Specimens were collected between about 180 and 800 m, with most stations in 200-400 m and a single
station recorded as 124-850 m. We review geographic and bathymetric distributions of all species represented.
The substantial amount of material of several species, notably the thalassometrids Oceanoinetra annandalei
(AH Clark) (40 specimens) and Aglaometra valida (Carpenter) (31), permits a better understanding ol
morphological variability than previously. In particular, statistical analyses on these two species indicate that
several morphological features formerly considered as diagnostic (e.g., centrodorsal shape, number of cirri) are
extremely variable. We place several taxa in synonymy as a result. Skeletal ossicles of these two species plus a
third, Stenometra cristata (A.H. Clark) (25 specimens), have been examined using scanning electron microscopy
Source :
630
C.G. MESSING, N. AMEZIANE & M, ELEAUME
(SEM) for the first time. Future comparisons of this material with other members of the family may provide
additional diagnostic features. Perhaps more than anything else, this collection indicates how severely most
comatulid families stand in need of detailed morphological examination, character analysis, and phylogenetic
hypothesis-testing.
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
631
CHECKLIST OF COMATULIDS
STATION LIST
The following list of stations includes latitude, longitude, depth, date, gear and number of specimens of each
species collected. Figure 1 illustrates station locations. Table 1 summarizes species collected at each station.
Table I. — Distribution of comatulid species by Karubar Expedition station. X - present.
Kepulauan Kai Stations.
Station CP 05: 05°49'S, 132°18’E, 296-299 m, 22.10.1991, fish trawl: Aphmocomaster pulcher (1 specimen),
Cosmiometra philippinensis (3), Oeeanometra annandalei (1).
Station CP 09: 05°23'S, 132°29'E, 368-389 m, 23.10.1991, fish trawl: Aphmocomaster pulcher {\\).
Station CP 16: 05°17'S, 132°50’E, 315-349 m, 24.10.1991, fish trawl: Aphanocomaster pulcher (1),
Cosmiometra philippinensis (16, probably plus 6 additional), Crotalometra rustical D. Oeeanometra annandalei
(8).
Station CP 17: 05°15'S, 133°01'E, 439-459 m, 24.10.1991, fish trawl: Palaeocomatella hiwia (2).
Station DW 18: 05°18'S, 1 33°01'E, 205-212 m, 24.10.1991, dredge: Palaeocomatella difficilis (1),
Palaeocomatella hiwia (2), unidentified comasterid (1), Neometra xenocladia (2), Stenometra cristata (1),
Stenometra quinquecostata (4).
Station CP 19: 05°15'S, 133°01'E, 576-605 m, 25.10.1991, fish trawl: Aglaometra valida (4).
Station CP 20: 05°15'S, 132°59’E, 769-809 m, 25.10.1991, fish trawl: Aglaometra valida (1). Stiremetra
breviradia (1).
Station CC 21: 05°14'S, 133°00'E, 688-694 m, 25.10.1991, shrimp trawl: Aglaometra valida (23).
Station DW 22: 05°22'S, 133°01’E, 124-850 m, 25.10.1991, dredge: Phanogenia sp. (2), Neometra cf. alecto (2).
Station CP 25: 05°30'S, 132°52'E, 336-346 m, 26.10.1991, fish trawl: Crotalometra rustical 1, small broken).
Station CP 27: 05°33'S, 132°52'E, 265-302 m, 26.10.1991, fish trawl: Cosmiometra philippinensis (1, small).
Station CP 32: 05°47'S, 132°51'E, 170-206 m, 26.10.1991, dredge: Stenometra cristata 1 1, small).
Station CP 35: 06°08’S, 132°45'E, 390-502 m, 27.10.1991, fish trawl: Aphanocomaster pulcher (6).
632
C.G. MESSING, N. AMEZIANE & M. ELEAUME
Stations between Kepulauan Aru and Kepulauan Tanimbar.
Station DW 50: 07°59S, 133°02 E, 184-186 m. 29.10.1991, dredge: Asterometra longicirra (1), Neometra
xenocladia (6).
Stations south of Kepulauan Tanimbar.
Station CP 82: 09°32'S, 131°02'E, 215-219 m, 04.1 1.1991, fish trawl: Stenometra cristata (10).
Station CP 83: 09°23'S, 131°00'E, 285-297 m, 04.1 1.1991, fish trawl: Oceanometra annandalei (31), Stenometra
cristata ( 1 ).
Station CP 86: 09°26'S, 131°13’E, 223-225 m, 04.1 1.1991, fish trawl: Cosmiometra iole (4), Stenometra cristata
SPECIES LIST
The following taxonomic list includes all nominal comatulid species known from the East Indian archipelago
(the islands of Indonesia, Malaysia, New Guinea and the Philippines) at depths between about 100 and
1000 m. Superscript (i) indicates species represented in the Karubar collections; superscript (2) refers to species
dredged previously around the Kai, Aru and Tanimbar Islands, but not found during the Karubar expedition;
superscript ( ’) indicates species known from elsewhere in this region and depth range (which may be added to the
eastern Indonesian outer shelf to upper bathyal comatulid fauna in the future). An asterisk (*) indicates species
endemic to the Kai, Aru and Tanimbar Island area.
Species known chiefly from shallow water (<50 m) are not included [e.g., of A.H. Clark’s (1931) 81 records
oi Capillaster multiradiatus from the Indo-Malayan archipelago and the Philippines that include depth ranges, only
our equal or exceed 100 m], A few taxa with unknown depth ranges (e.g., Parametra ajax) have been included
because all other species in their genera occur in the 100-1000-m depth range. Pre-KARUBAR records are from
A.H. Clark (1931, 1941, 1947, 1950) and A.H. Clark & A.M. Clark (1967); other sources are noted in the list.
Family COMASTERIDAE
Aphanocomaster pulcher (A.H. Clark) >*.
Comatulides decameros (A.H. Clark)2 [includes C. australis (A.H. Clark) (Hoggett & Rowe, 1986)1,
Comissia gracilipes (A.H. Clark)2; ?C. horridus (A.H. Clark)3; C. luetkeni A.H. Clark2; ?C. pereerina
(Bell)3; C. spinosissima A.H. Clark3; C. parvula A.H. Clark3.
Palaeocomatella difficilis (Carpenter)'; P. hiwia McKnightL
Phanogenia distincta (Carpenter)2 [formerly Comaster distinctus (see Messing, 1998b)].
Family ZYGOMETRIDAE
Catoptometra magnifica A.H. Clark2.
Family EUDIOCRINIDAE
Eudiocrinus gracilis A.H. Clark2; E. indivisus (Semper)2; E. junceus A.H. Clark2; £. ornatus A.H Clark3'
E. serripinna A.H. Clark2; E. venustulus A.H, Clark2.
Family HlMEROMETRIDAE
Homalometra denticulata (Carpenter)3.
Family MARIAMETRIDAE
Mariametra vicaria (Bell)2.
Pelometra ambonensis A.H. Clark3.
Family COLOBOMETRIDAE
Alisometra longipinna (A.H. Clark)3.
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
633
Cotylometra gracilicirra (A.H. Clark)2.
Cyllometra manca (Carpenter)3.
Epimetra nympha A.H. Clark3.
Iconometra anisa (H. L. Clark)2; I. marginata (A.H. Clark)3.
Family ASTEROMETRIDAE
Asterometra cristata A.H. Clark3; A. longicirra (Carpenter)1*; A. mirfica A.H. Clark2.
Pierometra pulcherrima (A.H. Clark)2; P. trichopoda (A.H. Clark)2.
Family CALOMETR1DAE
Calometra discoidea (Carpenter)2.
Neometra acanthaster (A.H. Clark)3; N. cf. alecto (A.H. Clark)1; N. diana (A.H. Clark)2; N. multicolor
(A.H. Clark)3 (Rowe & Gates, 1995); N. sappho A.H. Clark2; N. sibogae A.H. Clark3; N. xenocladia
sp. nov.1*.
Pectinometra carduum (A.H. Clark)3; P. magnified (A.H. Clark)3.
Family THALASSOMETRIDAE
Aglaometra valida (Carpenter)1 [includes A. incerta (Carpenter), A. propinqua (A.H. Clark) and A. sulcata
(A.H. Clark)].
Cosmiometra iole A.H. Clark1; C. philippinensis A.H. Clark1; C. woodmasoni (Bell)3.
Crotalometra rustica A.H. Clark1.
Daidalometra arachnoides (A.H. Clark)2; D. eurymedon A.H. Clark3.
Oceanometra annandalei (A.H. Clark)1 [includes O. magna (A.H. Clark)].
Parametra ajax A.H. Clark3; P. compressa (Carpenter)2; P. granulata A.H. Clark3; P. lisa A.H. Clark2*.
Stenometra cristata A.H. Clark1 (includes S. snelliusi A.H. Clark); 5. quinquecostata (Carpenter)1.
Stiremetra breviradia (Carpenter)1; S. perplexa (A.H. Clark)3.
Thalassometra gracilis (Carpenter)3; T. hirsuta A.H. Clark3; T. margaritifera A.H. Clark3.
Family PENTAMETROCRINIDAE
Pentametrocrinus diomedeae A.H. Clark2; P. japonicus (Carpenter)3; P. various (Carpenter)3.
Thaumatocrinus naresi (Carpenter)3.
Family ATELECRINIDAE
Atelecrinus wyvilli Carpenter3.
Note: Specimens belonging to the following two families were collected by the Karubar Expedition but
were not examined for this paper.
Family CHARITOMETRIDAE
Chondrometra aculeata (Carpenter)3; C. robusta (A.H. Clark)3; C. rugosa A.H. Clark3.
Glyptometra angusticalyx (Carpenter)3; G. crassa (A.H. Clark)2; G. distincta (Carpenter)2; G. inaequalis
(Carpenter)3; G. investigatoris (A.H. Clark)2; G. levigata (A.H. Clark)3; G. timorensis A.H. Clark3;
G. tuberosa (Carpenter)2.
Monachometra flexilis (Carpenter)2*; M. fragilis (A.H. Clark)3; M. patula (Carpenter)2; M. robusta
(Carpenter)2.
Poecilometra acoela (Carpenter)3.
Strotometra ornatissimus A.H. Clark3; S. parvipinna (Carpenter)2; 5. priamus A.H. Clark2.
Family ANTEDONIDAE
Adelometra angustiradia (Carpenter)2*.
634
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Antedon parviflora (A.H. Clark)3.
Balanometra batanoides (Carpenter)3.
Dorometra clymene A.H. Clark3.
Erythrometra australis A.H. Clark3.
Euantedon exquisita (A.H. Clark)3; E. moluccana (A.H. Clark)-3.
Eumetra chamberlain i A.H. Clark3.
Fariometra alcyon (A.H. Clark)3; F. dione A.H. Clark3; F. explicata (A.H. Clark)3; F. nieippe (A.H.
Clark)3.
Iridometra maxima A.H. Clark3.
Nanometra clymene A.H. Clark2.
Nepiometra laevis (Carpenter)3.
Perometra pusilla (Carpenter)2*.
Psathyrometra anomala A.H. Clark3; P. mira A.H. Clark3.
Thaumatometra thysbe A.H. Clark2.
Thysanometra tenuicirra (Carpenter)2.
Tonrometra brevipes (A.H. Clark) 3.
Toxometra paupera A.H. Clark3.
MATERIALS AND METHODS
Comatulids were preserved in 70% ethanol upon colleetion and subsequently transferred to fresh 70% ethanol
or, in a few instances, dried. All specimens are deposited in the echinoderm collection ol the Museum national
d'Histoire Naturelle, Paris. Catalogue numbers are prefaced EcCh. All drawings were made with a camera lucida on
a stereomicroscope.
We used a caliper for linear measurements >1 mm, and an ocular micrometer or camera lucida for finer
measurements and those difficult to make directly. We measured the length of curvilinear structures by placing a
fine waxed cord along the structure (or the image of the structure under a camera lucida) and then straightening the
cord along a ruler. Ray length and maximum cirrus length are rounded to the nearest millimeter. Other
measurements are rounded to the nearest 0. 1 mm. Measurements are included in descriptions. In addition to standard
measurements (e.g., A.H. Clark, 1950; BREIMER, 1978; MESSING, 1995a, b, 1997), we include additional
morphological features that we hope will help clarify comatulid taxonomy and permit comparison with other taxa,
including fossils. In particular, the large numbers of specimens of the thalassometrids. Aglaometra and
Oceanometra , permit examination of morphological variation in greater detail via statistical analyses than possible
for other collected taxa.
STATISTICAL ANALYSES
Statistical analyses were performed on meristic and morphometric data of Aglaometra valida and Oceanometra
annandalei using STATlab version 3.0 (Jambu, 1991-1994). Because the great majority of specimens approach
full size and only a few are small, the data arc not skewed; fewer than 10% of datapoints ever fall outside two
standard deviations of sample means. We have therefore treated the data as normally distributed.
SOKAL & Rohlf (1995) note that, in a normal frequency distribution, p±3cr accounts for 99.73% of_vari-
ability (where p is the population mean and a is the population standard deviation). The expression (Y i - X) / s
calculated for each variate Y gives the distance of each variate to the sample mean (X), expressed as a multiple of 5
(sample standard deviation). We thus treat any data point beyond the estimated 3cr-limit as divergent and warranting
re-examination.
Descriptive statistics reported for a wide range of skeletal features include minimum and maximum values,
sample mean (A), standard deviation (5); and coefficient of variation (V = (5 x 100) / X). We calculated the mode
for cirrus traits and brachial proportions: features that generally exhibit relatively limited variation. These
quantitative additions permit us to examine morphological variation in greater detail and more thoroughly and
accurately compare putative taxa.
Source :
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
635
Pinnules.
Brachials
Synarthrial Swellings
(or Tubercles)
Interradial Ridge
Longest Cirral
Centrodorsal
COMATULID MORPHOLOGY, MEASUREMENTS AND SYMBOLS
GENERAL FORM AND ORIENTATION
Comatulids consist chiefly of five, usually branched, feathery rays that radiate from a small central body. A
central visceral mass, the disk (Fig. 2a), rests on the bases of the rays and bears both mouth and anus on its
upper, oral surface. Each ray bears an open ambulacrum, a ciliated food groove lined with lingerlike, food¬
collecting podia (extensions of the water vascular system); ambulacra converge on the central mouth. The aboral
surface of the comatulid, the side away from the mouth, consists chiefly of articulated series of calcareous
cndoskeletal pieces called ossicles (Figs 2-4). As in other echinoderms, the fine structure of the ossicles forms a
fenestrated meshwork (stereom). Because much of comatulid taxonomy is based on detailed examination of
ossicles, most observations and measurements are taken on the aboral surface. Also, because the skeleton is highly
repetitive, symbols for different skeletal parts greatly condense taxonomic descriptions. The following outline of
comatulid morphology and symbols (Figs 2-5) derives primarily from MESSING (1997; see also A.H. Clark,
1915, 1921; BREIMER, 1978). Additional orientation terms include the following: Abambulacral: away from or
opposite the surface bearing the food groove (= aboral). Adambulacral: toward the surface bearing the food
groove. Apical: toward the top (aboral end) of the centrodorsal. Basal: toward the broader, oral side of the
centrodorsal, or toward the base of a structure (= proximal). Distal: a direction or position away trom the
aboral/oral axis, or toward the tip of a structure (e.g., ray, arm, cirrus or pinnule). Proximal: a direction or
position toward the aboral/oral axis, or toward the base of a structure.
Distal
Proximal
Brachitaxis
(Division Series)
Aboral profile sinuous
Distal profile of
aboral spine concave
Proximal profile of
aboral spine convex
Aboral carinate spine
triangular in transverse
Peripheral Cirrus
Cirrus Sockets
(in columns)
Midradial Height
Obsolete Socket
Aboral Pole
Apical Cirrus
Polar Diameter
Basal Diameter
Fig. 2. — a, Lateral view of proximal portion of a thalassometrid showing two cirri and bases of three rays, with
taxonomically important features, orientations, symbols and measurements, b. Three cirrus segments (ctrra s
indicating measurements and selected morphological features.
636
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Pinnule
Muscular Articulation
Spines
Spine
Claw
Synarthry
Sockets
Cuneate
(Wedge-shaped)
Fig. 3. — a, Oblique, semi-exploded view of proximal portion of a comasterid showing radial pentagon and brachitaxes of
two rays (visceral mass not shown), with taxonomically important features and symbols, b. Portion of comatulid arm
showing distinction between cuneate and triangular brachials. W and L indicate width and length measurements (from
Messing, 1997).
CENTRODORSAL
Morphology. — The centrodorsal is the largest skeletal ossicle, lies in the center of the aboral surface,
ranges from discoidal through hemispherical to conical or cylindrical, and almost always bears numerous sockets
for the articulation of hook-like, segmented cirri (Figs 2, 3a). [NOTE: The term centrodorsal derives from the
traditional designation of the mouth-bearing surface as ventral and the opposite surface as dorsal, in agreement with
most other groups of living echinoderms. However, because crinoids generally live with mouth uppermost,
functional and anatomical landmarks of orientation conflict. As a result, we use the terms aboral and oral instead of
dorsal and ventral].
Source : MNHN, Pahs
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
637
As the centrodorsal enlarges, additional cirri are added at the margin in a double spiral pattern so adjacent
sockets are offset, not side-by-side. Some species align cirri in columns from apex to margin with two or three
columns under each radial often segregated by an interradial ridge. Small apical cirri developed when the comatulid
was small may be retained near the centrodorsal apex, or they may be lost; traces of their attachment may remain
as obsolete sockets. The aboral pole, at least a small area devoid of cirri and sockets at the centrodorsal apex,
may be flat, convex or conical, smooth, papillose or spiny. In many comatulids, apical cirri are lost and their
sockets obliterated with centrodorsal growth so that the aboral pole broadens and cirri arc restricted to the
centrodorsal margin.
Measurements. — Oral (= proximal or basal) diameter; aboral pole diameter; midradial centrodorsal height
(from midradial basal margin to apex).
CIRRI
Morphology. — Cirri are unbranched, usually hook-like or prehensile, anchoring appendages that arise from
the centrodorsal and are composed of ossicles called cirrals (Figs 2, 3a). The terminal segment forms a curved
terminal claw and the penultimate usually bears an opposing spine. Cirrals typically increase in length from
a short basal few to the longest, always in the proximal half of the cirrus, which is followed by shorter distal
segments. Middle and distal cirrals may bear an aboral carina (=keel-likc ridge), spine or transverse ridge (2b, 3a).
In some taxa with dull proximal cirrals and polished, shiny distal cirrals, a distinct transition segment has
a polished distal and a dull proximal portion.
Symbols. — Individual cirrals are indicated by an Arabic subscript counting from the base of the cirrus (c.g.,
C3, cn) (Messing, 1997).
Measurements. — Number of cirri (= functional cirrus sockets), given in Roman numerals; number of cirrals
(of longest cirrus), given in Arabic numerals; maximum cirrus length; number and length/width ratio (L/W)
of longest cirral (Fig. 2b); position of transition cirral and of initial cirral with aboral spine or tubercle
(often approximate due to gradual appearance of aboral ornamentation); number of columns of cirri adjacent to
each radial ossicle (i.e., per radial area), and number of cirri per column. For Aglaometra valida and Oceanometra
annandalei , the following were also recorded: longest cirral length and width, and (for A. valida ) number
of regenerating cirri. In those lhalassometrids bearing numerous cirri composed of numerous cirrals, the
longest cirrus and longest cirral were selected for measurement by visual estimation, rather than by measuring
every cirrus.
RAYS
Morphology. — Rays are the five, usually branched, series of ossicles and associated soft tissues that radiate
from the center of the comatulid body (Figs 2-4). A radial is the first (most proximal) ossicle of a ray and is
associated with one of the five internal radial canals of the water vascular system that arise from the circumoral
ring canal. The five radials together form a radial pentagon or circlet that lies against and is partly (rarely
completely) covered by the centrodorsal. Structures associated with the extrapolated central axis ol these ossicles
are also referred to as radial. Interradial structures are oriented between rays. Basal ossicles lie interradially
between the centrodorsal and radials but are reduced in comatulids and are not considered herein (see A.H. Clark.
1915; BREIMER, 1978, or Messing, 1997, for a fuller discussion).
Brachial ossicles follow the radials. Division series (or brachitaxes) are series of two to four
brachials between ray branches, either following a radial and including the first ossicle at which the ray branches
(axillary or axil), or following an axil and including the next. Each axil bears two articular faces distally and
may thus bear two additional brachitaxes, two unbranched arms, or one of each. Interior and exterior arms,
brachitaxes or associated structures are those closest to and furthest from, respectively, the extrapolated axis of the
preceding branching series. As examples, in Figure 4, the brachitaxis labelled IIIBr4(3+4) is exterior; it arises from
the outer side of the IIBr2 axil relative to the extrapolated axis of the IBr2 series. Also in Figure 4, the interior
margin of bn is the side closest to the extrapolated axis of the preceding axil. Rays and their various branches carry
extensions of coelomic, nervous and water vascular systems. Branching patterns are often diagnostic at generic and.
sometimes, family levels.
638
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Most articulations between successive
brachials are muscular articulations,
which contain both muscles and ligaments
that anchor in depressions (fossae) in the
articular face. Moving from the ambulacral
to the aboral surface, each muscular
articulation consists of a pair of
muscular fossae, a pair of interar-
ticular ligament fossae, a trans¬
verse fulcral ridge [typically oblique
(e.g., Figs 13d, 34a, 40b) except in the
articulation between the radial and first
succeeding ossicle in which it runs at right
angles to the oral/aboral axis of the ossicle
(e.g.. Figs 30f, 31a)], axial canal and
semicircular aboral ligament fossa.
The raised area immediately around the
axial canal is referred to as periaxial.
Syzygies are rigid, exclusively
ligamentary articulations in which ridges
(culminae) radiating from the center of
one articular face correspond with those on
the apposed face; the external appearance
resembles a perforated line (e.g., Figs 13f,
3 1 e). Flat areas between ridges are
crenellae. Syzygies represent fracture
points for arm autotomy and sometimes occur at diagnostic intervals along arms. They also typically occur
between the third and fourth ossicles of four-ossicle brachitaxes, and between the third and fourth arm ossicles.
Synarthries bear a pair of semicircular ligaments separated by a fulcral ridge oriented through the
ambulacral/abambulacrai axis of the articular face (e.g.. Figs 13e, 31b). The abambulacral (= aboral) surface around
a synarthry may be swollen as a synarthrial tubercle or swelling (Fig. 2a). Synarthries join two-ossicle
brachitaxes, the first two ossicles of four-ossicle brachitaxes, and the first two arm brachials. However, they may
be replaced by syzygies, or (in some comasterids) modified as cryptosynarthries: flat articular facets bearing
low irregular or concentric sculpture with only a trace of a fulcral ridge (Figs 2-3).
Brachial shape, when viewed aborally, varies from oblong (both lateral margins of similar length), through
cuneate (= wedge-shaped; one lateral margin obviously longer) to triangular (proximal and distal margins
converging on one side of the brachial) (Fig. 3b). In some species, brachitaxial and proximal arm ossicles have
truncated corners that leave openings called water pores between adjacent rays (Fig. 5).
Brachial ossicles may be variously ornamented with spines, knobs, tubercles, teeth, crenulae or lamellae,
scattered across the aboral surface or restricted to ossicle margins or a midaboral ridge or carina (Fig. 5).
Brachitaxial ornamentation tends to continue onto the proximal few brachials of the unbranchcd arm.
Symbols. — Each successive brachitaxis is indicated by a Roman numeral followed by Br; an Arabic number
following indicates the number of component ossicles (e.g., IBr2 is the first brachitaxis, always composed of two
ossicles in comatulids). (NOTE: The radial is always the second ossicle counting from the first axil toward the ray
base). Individual ray ossicles are indicated by br (plural, brr) followed by an Arabic subscript (e.g.. II Ibr4, the
fourth ossicle of the third brachitaxis; br i , bri, etc., for arm ossicles following the last axil). This is a slight
modification of previous usage (e.g., A.H. Clark, 1931; A.H. Clark & A.M. Clark, 1967) in which
individual division series ossicles were indicated by Br followed by an Arabic subscript (e.g., IIBrj). The change
reduces possible confusion between references to entire brachitaxes (e.g.. IIIBr2) and their component ossicles (e.g.,
IlIbQ) (Messing, 1997).
Fig. 4. — Aboral view of centrodorsal and proximal portion of a ray of
a comasterid showing abbreviations for brachitaxes.
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
639
A + sign indicates a syzygy [e.g.. IIBr4(3+4) is a second brachitaxis of four ossicles with the third and fourth
joined by syzygy; br.3+4 indicates brachials three and four joined by syzygy (Figs 2a, 3a, 4)]. Synarthries are not
usually designated by a symbol. However, they exhibit an unusual arrangement in Aphanocomaster pulcher and are
indicated in this species by one-em dashes [e.g., IIIBr4(l — 2, 3 — 4) (MESSING, 1995b|.
Measurements. — Arm number; ray length (from midradial margin of centrodorsal to tip of longest remaining
arm, which is often broken distally); width to length ratio (W/L) of various brachitaxis and brachial ossicles,
depending upon species, and diameter of br3+4. For Aglaometra valuta and Oceanometra annatidalei, lengths and
widths of specific brachitaxis and brachial ossicles and (for 0. annandalei ) total number of axils.
Exterior lateral margins
flattened and apposed
Converging lateral margin with spines
Diverging lateral margin with crenulae
Exterior lateral ma
weakly everted and
Exterior lateral margin
Midaboral ridge
Proximolateral margin
everted and dentate
Weak midaboral ridge
Interior distolateral
Interior distolateral corner
truncated (cut away)
Water pore
Exterior proximolateral margin
strongly everted and smooth
Exterior lateral margin everted,
with knobs coalesced as lamellae
Lateral margin everted and spinose
Proximolateral margin finely spinose
FIG. 5. — Aboral view of a series of brachitaxes of a thalassometrid identifying selected sculpture and ornamentation.
PINNULES
Morphology. — Pinnules are the small, segmented, unbranched appendages that arise on alternating sides of
successive brachials. They are composed of ossicles called pinnulars (Fig. 3a), give the arms their characteristic
featherlike appearance, and are the primary site of food-collection. One or more pairs of proximal oral pinnules
often differ from those succeeding (e.g., they may be longer and more flexible, more robust and spinelike, or lack
an ambulacral groove). In the Comasteridae, the distal pinnulars of oral pinnules each bear a blade or tooth that,
taken together, form a comb (Fig. 3a). At least several pairs of genital pinnules bearing gonads follow the
oral pinnules. These rarely exhibit skeletal modifications; they are usually either similar to or shorter and stouter
than the more distal, exclusively food-collecting pinnules. Relative lengths, stoutness, numbers of pinnulars and
ornamentation may be diagnostic of taxa at specific, generic and familial levels.
Symbols. — P indicates pinnules. Arabic subscript numbers and lower case letters indicate successive exterior
and interior pinnules, respectively (e.g., Pi, P2, Pa)- The interior side of an arm or brachitaxis is that side closest
to the extrapolated median axis of the preceding axil. In most species, br2 bears exterior Pi, br4 bears interior Pa
(most comatulids have br3+4 united by syzygy; the proximal ossicle of a pair joined by syzygy or synarthry never
Source :
640
C.G. MESSING. N. AMEZIANE & M. ELEAUME
bears a pinnule), brs bears exterior P2, etc. Brachitaxes of three or four ossicles bear an exterior pinnule on the first
or second ossicle, respectively, indicated by a subscript Roman numeral corresponding to the brachitaxis (e.g., Pn
on IIBr).
Measurements. — Pinnule length and number of pinnulars of various pinnules (e.g., Pi, P3, middle or distal
pinnules); length to width ratios (LAV) of specific pinnulars.
DISK
Morphology. — The comatulid visceral mass, which rests on top of the radial pentagon and proximal ossicles
of the rays, is called the disk (Fig. 2a). Its oral surface, the tegmen, is usually naked integument although it
may be invested with small calcareous pieces or polygonal plates. [NOTE: Disk and tegmen are sometimes used
synonymously as the oral surface of the visceral mass]. The tegmen bears the anus, located at the apex of a small
anal papilla or tube, and the mouth with its radiating ambulacral trunks. The mouth is usually located centrally
on the tegmen with the anus displaced to one side. However, in most Comasteridae the mouth is displaced off-
center or to the disk margin and the anal papilla is central or subcentral. Areas on the tegmen between ambulacral
grooves are interambulacral.
SYSTEMATIC ACCOUNT
Superfamily COMASTERACEA
Family COMASTERIDAE A.H, Clark. 1908
Genus APHANOCOM ASTER Messing, 1995
Aphanocomaster pulcher ( A.H. Clark, 1912)
Comaster pulcher A.H. Clark. 1912c: 22; 1918: 37, 40, 275. pi. 14 figs 14-15; 1931: 443, pi. 50. — Gislen. 1919: 14.
Aphanocomaster pulcher - MESSING, 1995b:658. fig. 1.
Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn CP 05: I specimen (MNHN-EcCh 21). —
Stn CP 09: 11 specimens (MNHN-EcCh 22). — Stn CP 16: 1 specimen (MNHN-EcCh 23). — Stn CP 35; 6 specimens
(MNHN-EcCh 20).
Description. — This description emphasizes differences between the Karubar material and the holotype
which was redescribed by MESSING ( 1995b).
Centrodorsal often almost circular rather than rounded pentagonal; diameter 4.6-6.6 mm., D/H 5.0-6.0. Aboral
pole, especially in larger specimens, depressed, its margin formed by raised edges of apical cirrus sockets; in one
specimen, pole only about half centrodorsal diameter due to numerous encroaching apical sockets.
Cirrus sockets as in holotype, but ranging from one or two high to two or three high between centrodorsal base
and aboral pole (rather than in a single marginal row). Cirri more numerous than in holotype. XXXVI1-LI (one
specimen with LXV), 13-17 (one specimen with 18-19), 11-15 mm (most specimens 1 1 mm, as in holotype).
Cirrals generally as in holotype, but with proximal segments often more elongated; ci short; C2 and C4 with L/W
2.0-2. 6; C3 longest. LAV 2. 8-3. 3. Weak transverse ridge first appearing on C3 to cs; developing into low rounded or
blunt triangular tubercle on distal cirrals as in type, or becoming sharp triangular spine as early as C6.
Radials barely visible as small interradial triangles, or slightly more exposed, but barely visible midradially, if
at all. Articulation between Ibri and Ibr2 a cryptosynarthry in dissociated specimen. Ibri very short, W/L >5.0.
Gaps between adjacent IBr series as in type. IIBr series chiefly IIBr4( 1 — 2, 3—4) (1-em dashes indicate ossicles
joined by synarthry); a few specimens with up to four IIBr2(l— 2). Variants include one each of the following on
three different specimens: one IIBr4(l — 2, 3+4); one IIBr 1 with the single large ossicle fused interiorly to the
Source : MNHN, Paris
C0MATUL1D CR1NOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
641
adjacent large llbri; one IIBr6 with the first two ossicles apparently united by synarthry, with Pu exterior on Ilbri,
IIbr3 very short, an interior pinnule on IIbr4 and an exterior pinnule on Ilbrs. IIIBr series as variable as in type:
4(1_2, 3—4), 4(1—2, 3+4), 4(1+2, 3+4), 2(1—2) and 2(1+2) apparently all present in various combinations.
Occasional !VBr2 series present, usually apparently 2(1+2). Identification of the first articulation of III and IVBr
series as syzygy or cryptosynarthry often difficult (see below). Pinnule-bearing brachitaxis ossicles lacking
strongly projecting lateral margins found in type. Brachitaxis pinnules (i.e., P|| and Pm) apparently arising more
toward ambulacral side of ossicle than in type.
Anns 32-39; mostly broken. Ray length up to 100 mm in few remaining intact cases; bn+4 diameter 0.9-
1.2 mm on arms arising from IIIBr series. Middle and distal brr generally somewhat shorter than in type; brio with
W/L up to 2.5; distal brr with W/L 0.75. Proximal ami syzygies generally as in the type (bri+2. 3+4 on arms
arising from IIIBr series, and bn+4 alone on IIBr series), but are often followed by an additional syzygy at br.s+6,
br6+7 or br7+8- At least one arm of one specimen bears bn+4, brs+6 and bn+8. On one specimen, interior arms
arising from IIBr2(l— 2) bear bri+2. 3+4- Arms arising from IIIBr series rarely bear bri+2 or bn+4 alone. Several
specimens bear one to several br2+3, usually exteriorly, with the preceding bn usually enlarged, suggesting fusion
of an original br 1 +2 (making br2+3 homologous with bn+4); bn+3 occasionally followed by bn+5. Following
syzygy between brn+16 and br22+23- Distal syzygial interval usually 4, as in type.
Pu up to 51 segments, 17 comb teeth, 19 mm; not as robust basally as in type; most pinnulars constricted and
longer than median width; comb arising gradually; no partly twisted proximal tooth. On another specimen (lacking
intact Pu), Pi up to 54 segments, 18 teeth, 20 mm; similar to Pu but slightly less robust; P2 of 42 segments.
14 teeth, 14 mm; P3-P5 similar, declining slightly in size, 34-37 segments, 14-15 teeth, -8 mm; P6 35
segments, 16 teeth, 7 mm; last comb-bearing pinnule (probably Ps) 25 segments, 1 1 teeth. 5.6 mm. On another
specimen, P4 32 segments, 16 teeth, 6 mm; P5 27 segments, 15 teeth, -4.6 mm; P6 to Ps last combed pinnule;
first combless pinnule (P7-P9) 18 segments, 5.2-6.2 mm; stouter and longer than preceding, with short pinnulars;
those beyond the base with a cluster of lateral spines. Gonads first well developed on P9-P11; present to at least
P28. Combs apparently arise gradually on brachitaxial and proximal brachial pinnules, and more abruptly on distal
combed pinnules; initial tooth partly twisted on some combs. Some specimens with dark reddish brown granules
in pinnule tissue, reminiscent of saccules but of varying size.
Disk naked; mouth central; anal papilla large, about midway between mouth and disk margin; anal
interambulacral area only somewhat larger than other interambulacral areas, as in type.
Color in alcohol : Aboral pole of centrodorsal usually darker than remainder of skeleton, blotched, spotted or
speckled with brown, usually in a radiating pattern. Cirrals often tan with white articulations. Proximal portions
of rays often with tan or brown spots, chiefly on articulations. Disk usually gray, sometimes spotted or speckled
with brown.
DISTRIBUTION. — Known only from the Kai Islands. The type specimen was recorded from Siboga station
257 in the Du-roa Straits, described by A.H. Clark (1918: 275) as "Down to 52 metres. Coral." The KARUBAR
material was retrieved from substantially deeper water: certainly 299-390 m. possibly to 502 m.
Remarks. — MESSING (1995b) redescribed the type specimen of Comaster pulcher and found that its unique
ray branching pattern, combined with the lack of the transverse initial comb tooth diagnostic of the genus
Comaster as construed by A.H. CLARK (1931) and HOGGETT and Rowe (1986). required that the species be
assigned to a new genus. MESSING (1998b) removed all other species of Comaster except the type. C. multifidus
(Muller) [= Comanthina variabilis (Bell)], to Phanogenia Loven.
The 19 KARUBAR specimens are the first known since the "Siboga" collected the holotype in 1899. Most are
larger and demonstrate an even greater variability in brachitaxes composition and proximal syzygy placement.
Despite a range of centrodorsal sizes, however, arm number exhibits a narrow range.
The preceding description chiefly points out differences between the KARUBAR material and the holotype.
However, a relatively thick epidermis makes it difficult in some cases to distinguish between syzygies and
synarthries, particularly close cryptosynarthries. To better examine problematic articulations, a small paintbrush,
trimmed short to stiffen the bristles, was used to remove this epidermis from some rays of some specimens. One
specimen and portions of two others, dissociated in 5% NaOCl solution, show that the first brachial syzygy
642
C.G. MESSING, N. AMEZIANE & M. ELEAUME
(bri+2) is often incompletely developed and appears to be derived from a synarthrial articulation. They also show
clearly that the IBr articulation is a tight synarthry or cryptosynarthry.
The combination of widely variable branching pattern, generalized pinnule combs, central mouth and putative
saccules suggests that -4. pulcher may represent a plesiomorphic form of Comasteridae.
Genus PALAEOCOMATELLA A.H. Clark, 1912
1 Palaeocomatella difficilis (Carpenter, 1888)
Aclinometra difficilis Carpenter, 1888: 93. pi. 52 fig. 2.
Actinometra pulchella (part) - Carpenter, 1888: 304, pi 52 fig 2
Pal&' R0WE!l986^2Cf5 ' A H‘ CLARK' ,9I3: ^ l918: 8‘ 2?3' P'- " figS ‘~2; 1931 : i54‘ pL 7 figS 23'24' - HoGGETT
Comissia difficilis - HOGGETT & Rowe, 1986: 123.
Material EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 18: 1 specimen (MNHN-EcCh 24).
Description. — Specimen fragmentary; arms lost beyond br2 or bry; no cirri. Centrodorsal discoidal. 4 4 mm
across. Rad.als mostly hidden by centrodorsal. IBr2 and IIBr2 series short, aborally flattened, well separated, united
y synarthry; Ibri partly hidden by centrodorsal; Ibr2 axil pentagonal with short lateral margins W/L 2 2-
Ilbr, oblong, W/L 3.3; Ilbr2 axil similar to lbr2, W/L 1.8. Seven IIBr2; arms 17; all 14 arms arising from IIBr
with bri+2. The three exterior arms with one additional remaining ossicle also have bn+a; diameter at brt+4 I 2
mm. Single Pi of 32 segments, 1 1 comb teeth; teeth tall and triangular, confluent with exterior side of pinnulars-
initial tooth not transverse. Disk naked; mouth central.
Distributon. - Previously recorded only off Jolo in the Sulu archipelago, southern Philippines, in 256-
V (A.H Clark. 1931). The Karubar specimen, if conspecific, extends the range to eastern Indonesia in
slightly shallower water (205-212 m).
Remarks. — This specimen most closely approaches Palaeocomatella difficilis (Carpenter) for the following
reasons. IBr2 and IIBr2 series united by synarthry; first brachial syzygy at br,+2; second brachial syzygy at br3+4
on a arms (three exterior) in which ossicles remain beyond bn; no initial transverse comb tooth; mouth central
HOGGETT and Rowe (1986) reassigned this species to Comissia on the basis of its central mouth and similar
comb form. However, this comb form is the most generalized and widespread in the family, and probably
represents a plesiomorphic condition. The central mouth position, characteristic of juvenile comasterids (and
Ciller' C°fI7atUI'fS) '?IS0 relamed in adul‘ coma,ulids belonging to Phanogenia, Aphanocomaster ,
S r ’ MuSS,NG (,"5a) n°ted ,hat ,hC m°U,h in lhe ^ sPec*men of Comissia luetkeni
A H. Clark, the type species ot the genus, is subcentral, a feature also found in Neocomatella spp. and Comissia
( eptonemaster) venustus (personal observations and HOGGETT and Rowe, 1986). It thus seems premature to
submerge Palaeocomatella within Comissia on this basis.
Palaeocomatella hiwia McKnight, 1977
Fig. 6
Palaeocomatella hiwia McKnighi, 1977: 94, figs 1-2.
Comissia hiwia - Hoggett & Rowe, 1986: 122? 124.
EXAMINED. - Indonesia. Karubar. Km Islands: stn CP 17-
Mn DW 18: 2 specimens (MNHN-EcCh 25).
specimens (MNHN-EcCh 26). —
5CRIPT'0N LARGE SPEC,MEN- - Centrodorsal discoidal, 6.4 mm across, D/H 4.6; aboral pole broad
ghtly concave. Cirr, crowded around margin. 1-2 high; XXXVIII, 18-20. 14-16 mm; several slender immature
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
643
cirri present; ci very short; C2 with LAV 1.3-1. 7; C3 longest, cylindrical. LAV 2.8-3.0; following cirrals rapidly
shorter, wider and compressed; C4 LAV 1.7; cs L/W 1.0; distal cirrals slightly wider than long, L/W 0.8. Aboral
distal tubercle well-developed on C4, compressed and sometimes accompanied by proximal hump on cs; following
cirrals with tall, sharply compressed, longitudinal ridge occupying more than half the length of each cirral, weaker
on 1-2 cirrals preceding penultimate; ridge crests flat, convex, sometimes finely denticulate. Opposing spine
rounded triangular in profile. Terminal claw gently curved, longer than preceding cirral.
Radials almost completely hidden by centrodorsal, perhaps visible as small interradial triangles. All brachitaxes
of two ossicles united by synarthry; IIIBr and IVBr series developed only exteriorly. IBr2 very short, partly hidden
by centrodorsal; aboral surface Battened. Ibri visible only interradially, in contact with adjacent Ibri only
proximally so that the ray bases are well separated; Ibr2 axil pentagonal, with very short lateral margins, W/L 3.0.
Ilbri short, oblong, united interiorly. W/L 3.3; Ilbr2 axil pentagonal, asymmetric (interior distal articular lace
supporting free arm not as wide as exterior face supporting IIIBr), W/L 2. 3-2. 5. Illbri longer exteriorly, W/L 2.4-
2.7; IIIbr2 similar to preceding axil but not as asymmetric, W/L 2.1. II-IVBr series more rounded aborally, with
weak midaboral synarthrial swellings. Exterior profile of IIbr2-IIIbri and that of IIIbr2-bri distinctly convex.
Arms 33; ray length = 70 mm. On exterior arms, bri oblong or slightly longer exteriorly. W/L 2.0-2. 3; br2
distinctly longer exteriorly, W/L 2.0; bn+4 arising from IIIBr series with diameter 1.2 mm. W/L 1.5; brrs-8
oblong, W/L 1.5- 1.6. On interior arms, bri+2 squarish, W/L 1. 0-1.1; brn-6 or bn oblong, W/L 1. 7-2.0.
Following brr rapidly becoming triangular, W/L 2. 0-2. 2; distal brr cuneate, W/L 1.3, longer neai arm lip. Distal
margins of distal brr finely denticulate. Syzygy at bn+2 (never bn+4) on interior arms arising from II-IVBr series;
at br3+4 on exterior arms; second syzygy br 1 3+ 1 4 or bri4+is; distal interval 4-5.
Oral pinnules fragile, mostly broken. Pi 40 segments, 13 comb teeth, 11 mm (possibly longer); proximal
several segments rhombic, wider than long; middle segments about as long as wide; comb teeth tall, triangular,
incurved. P2 32-33 segments, 10-12 teeth, 6-10 mm; middle segments with L/W up to 1.6. P3 shorter.
27 segments, 10 teeth, 7 mm. Distal comb tooth tall. Oat, broad and rounded, sometimes fan-shaped; one or two
low incipient teeth sometimes preceding comb. P4 without comb, 14 segments. 4.6 mm. Middle pinnules longer
and more robust, 15 segments, 7 mm. Distal pinnules slenderer, up to 16 segments, 7.6 mm.
Disk naked; mouth large, central; anal papilla large, adjacent to mouth. Epidermis of middle and distal arms and
pinnules filled with minute irregular (?calcareous) grains.
Description of small Specimen. — Centrodorsal discoidal, diameter 3.4 mm; projecting slightly in
interradial angles; aboral pole Bat with central low papilla. Cirri chiefly in single crowded marginal row,
occasionally two high; XXVIII, 10-11, 6.0-7.4 mm; ci short; c2 LAV 1.6-1.7; c3 longest, cylindrical with slightly
expanded distal end? LAV 2.7; following segments rapidly shorter and compressed; c5 and following cirrals as long
as wide; C4 or cs with low distal aboral swelling, increasing on following cirrals as subdistal to almost median
triangular or rounded triangular spine, sometimes longitudinally Battened on one or two distal cirrals. Opposing
spine triangular. Terminal claw gently curved, longer than preceding cirral.
Radials mostly hidden by centrodorsal; barely visible in interradial angles, but obscured by cirri. Brachitaxes al
of two ossicles united by synarthry. Battened aborally, with weak midaboral synarthrial swellings. Ibri extremely
short, partly hidden by radials, just touching at proximal corners so that ray bases are well separated; Ibi 2 axil
pentagonal with proximal margin gently concave and lateral margins short, W/L 2.8. Ilbr, oblong, slightly
thickened where united interiorly, W/L 2.7; IIbr2 axil similar to Ibr2 with distinctly diverging lateral margins, W/L
2.4. Exterior profile of IIbr2 and bri distinctly convex.
Arms 20; ray length 60 mm; bn united interiorly and usually wider than br2 so that bases of adjacent arms
(beyond bn) usually separated interiorly by distinct gap. On exterior arms, bri longer exteriorly, W/L 2.7; br2
longer and slightly longer exteriorly, W/L 1.9; br3+4 diameter 1.2 mm, W/L 1.5; brr5-6 oblong or slightly
cuneate, W/L 2.4. On interior arms, bn+2 longer exteriorly, W/L 1.6; brr3-5 oblong, W/L 2.3. Following brr
rapidly becoming triangular with finely denticulate distal margins, W/L 1. 8-2.1. Distal brr cuneate, W/L 1.0.
longer than wide near arm tip; distal margins with stronger (but still small) triangular teeth. Syzygy at bn+2 on
interior arms and bn+4 on exterior arms; second at bris+16 (one bri9+2o); distal interval chieBy 4-5.
Pinnules similar to those of larger specimen, with distinctive final comb tooth. Pi 36 segments, 12 comb
teeth, 10 mm. P4 25 segments, 9 teeth, 6 mm. P5 similar but shorter. P6 first genital pinnule, no comb.
644
C.G. MESSING. N. AMEZIANE & M. ELEAUME
11 segments, 5 mm. Middle pinnules 14 segments, 6.6 mm; pinnulars following short basal two longer than
wide. Distal pinnules 16 segments, 7.6 mm, slenderer with more elongated segments.
Disk absent. Irregular grains present as in large specimen, but fewer.
Fig. 6. — Palaeocomatella hiwia. a, cirrus, large specimen (EcCh 26); b. cirrus, small specimen (EcCh 26); e-d comb
tips showing tan-like terminal pinnular (c. Pi; d. Pi). Scale bars: a-b = 1.0 mm; c-d = 0.25 mm.
Additional POSSIBLE SPECIMENS (stn 18). — Centrodorsal discoidal; diameter 2.4 mm; cirri in sinale
marginal row, similar to those of smaller specimen above (with weaker aboral ornament), XV1I-XXI. 7-8,
2.4 mm; C3 longest. LAV 2. 3-3.0; weak, slightly longitudinally flattened, triangular aboral spine on distal one or
two cirrals. Radials hidden in aboral view; Ibri at least partly hidden by centrodorsal. Arms 10. all broken. Final
comb tooth of few remaining oral pinnules apparently tall, broad and rounded.
DISTRIBUTION. — The only previously published records are from off East Cape. North Island, New Zealand,
in 602-808 m (possibly 432-924 m), and the Kermadec Islands in 542-545 m (McKnight. 1977). The Karubar
collection, if conspecilic, extends the range to eastern Indonesia in at least 212-439 m (possibly 205-459 m).
Remarks. — The large specimen conforms closely to McKnight's (1977) original description of P. hiwia.
but is substantially larger. The type material has centrodorsal diameters of 3-4 mm; cirri XVIII-XXIX, 15-20.
10 mm long; arms 16-20; Pi of 25-30 segments with 8-9 teeth; combs extending to P3, and was collected in 432-
688 m. The smaller specimen falls well within the range of variation exhibited by the type material, but lacks the
well-developed aboral, longitudinal cirral ridge. The slight longitudinal compression of the distal cirral spines is
atypical ot other comasterids and is interpreted here as an early developmental stage of the longitudinal ridge found
in the large specimen and type material. Both Karubar specimens lack second syzygies at br4+5 or br5+6 on
interior arms as described by McKnight (1977).
The two additional small, fragmentary specimens from station 18 are very tentatively assigned here because
1 iey appear similar in several respects to the smaller of the two specimens from station 1 7. despite being collected
in substantially shallower water.
Genus PHANOGENIA Loven, 1866
Phanogenia sp.
Material EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 22: 2 specimens (MNHN-EcCh 27).
COMATUUD CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
645
DESCRIPTION. — Two small specimens, a and b, as follows. Centrodorsal diameter: a, 1.6 mm; b, 1.4 mm.
Cirri: a, XIV, 8-10, 3.4-5.0 mm; b, XXVII, 10, 6.0-6.4 mm. C3 (and sometimes C4> longest, L/W 2.9-3. 3. Arms:
a, 16; b, 14-17 (3 rays detached or lost, 1 ray broken). Ray length: a, 25 mm; b, 45 mm. Combs present to P3-P5.
sometimes one or two at intervals beyond; combs with transverse initial tooth. IBr2 united by synarthry. Each
specimen with one ray in which the articulations between the third and fourth ossicles of the II Br series and
between bri and bn. on both succeeding arms appear to be a cryptosynarthry or incompletely developed syzygy
with few, if any, culminae. Otherwise, several IIBr4(3+4) series and (in specimen a) one interior IVBr2
(articulation tight, irregularly perforated) series present, and first brachial syzygy at bri+2-
REMARKS. — MESSING (1998b) recently transferred all species previously included in Comaster Agassiz except
the type species, C. multifidus (Muller), to Phanogenia. The two Karubar specimens most closely approach
Phanogenia serrata (A.H. Clark), previously recorded only from Japanese waters [as Comaster serratus (A.H. Clark.
1931)]. A.H. CLARK’s key is misleading here because he diagnoses P. serrata as having 30 amis (p. 413. 452), but
then goes on to describe specimens with 13, 15 and 28 arms as well. Messing (1998b) removed A.H. Clark s
other very small Comaster, C. tenellus A.H. Clark, to Comanthus.
The Karubar specimens probably represent juveniles of one of several cirrus-bearing species of East Indian
Phanogenia (e.g., P. novaeguineae [Muller], P. multibrachiata [Carpenter], P.fruticosa A.H. Clark, P. schoenovi
A.H. Clark or P. distincta [Carpenter]) that may represent a growth series (perhaps including P. serrata as well).
It is thus not clear which species name should be applied.
Unidentified comasterid
MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 18: I specimen (MNHN).
DESCRIPTION. — Centrodorsal discoidal; diameter 3.8 mm. Cirri in single marginal row, XXIX. 10-11.8 mm;
C3 longest, LAV 3.1. Arms 1 1; one IIBr2; bn+2 on interior arm, br2+3 on exterior arm.
Remarks. — Possibly Palaeocomatella sp. with an abnormal exterior ami, or Capillastei sp.
Superfamily TROPIOMETRACEA
Family ASTEROMETRIDAE Gislen, 1924
Genus ASTEROMETRA A.H. Clark. 1907
Asterometra longicirra (Carpenter, 1888)
Fig. 7a
Antedon longicirra Carpenter, 1888: 103, pi. 17.
Asterometra longicirra - A.H. CLARK, 1947: 439. — A.M. CLARK. 1972. 15—
MATERIAL EXAMINED. — Indonesia. Karubar. Tanimbar Islands : stn CP 50: I specimen (MNHN-EcCh 28).
DESCRIPTION. — Centrodorsal conical with almost straight proximal sides; diameter 5.8 mm; D/H 1.1. Pole
with five small radially arranged apical tubercles. Cirri XVI. none complete; longest remaining of 62 segments.
90 mm long (tip missing; attached portion 12 segments, detached fragment 50 segments); cylindrical proximal ly.
compressed distally; ci short, wider (transversely, across articulation) and taller (along the oral-aboral axis) than
following segments; c, 3 or c, 4 to about c2o longest, LAV 1.4; distal aboral edge first projecting on c27. becoming
small sharp distally-directed spine by c3o. Spine becoming a transversely thickened, rounded car.na on shorter distal
segments.
646
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Radials almost flat, some with elongated midaboral tubercle, W/L 3. 1 . !Br2 sharply flat-sided and apposed; Ibri
with elongated midaboral ridge, W/L 2.5; ridge smooth and straight, with irregular crest or divided into two or
three tubercles; Ibr2 axil with elongated midaboral tubercle (one axil has a second smaller tubercle on aboral
midhne adjacent to one distal articulation), W/L 1.7.
Arms 10; ray length 105 mm (measured from centrodorsal apex)(arm tips lost). Brachials with lateral margins
narrowly everted and flat-sided to about bry; adjacent arms closely apposed to about bra; everted lateral margins on
brr5-9 appearing as narrow flanges in aboral view; bn and bn each with elongated midaboral tubercle, weaker than
on IBr; br3+4 very short, W/L 2.3, diameter 3.2 mm; brs to brg short and oblong, W/L 3.1. Following brachials
strongly cuneate by bri3 or bru, W/L 2.5-3.0; low narrow midaboral ridge first appearing on brn Middle
brachials laterally compressed, cuneate with stronger midaboral ridge. Distal brachials short, more strongly
compressed, carinate, W/L 1 .7; carina ranging from low and rounded to high and fin-shaped
Most pinnules broken; all strongly carinate. P, 12 segments, 7.6 mm long. P2 stouter. 13 segments. 9.6 mm.
P3 and Pa similar to P2, both of 12 segments, 10.0 and 7.6 mm, respectively. Small spine present on distal end of
carinate margin ol distal segments on P2 and following pinnules. Distal pinnules up to 20 segments 16 5 mm
long; longest middle segments with L/W to 1.7.
FlGh 1rZ^ATr°me,ra J°n^rra <EcCh 28)' een,rodorsal- bases of several cirri and bases of ihree rays.
20 mm C° ECCH 42) Sma Spedmen' centrodorsal- bases of two cirri and bases of two rays. Scale bar =
longicirra was recorded previously only from the Kai Islands in 90-256 m (A.H. Clark.
iyj|)- lhe KaRUBAR specimen from Tanimbar falls within this depth range (184-186 m).
Remarks. — Asterometra mirifica A.H. Clark, A cristata A.H. Clark, A. macropoda (A.H. Clark) and
(A uTr ,SP!3Sr)D d,flCr V6ry llUle eXCeP‘ 'n th£ CXlenl and degree 0f lhe Proximal carination"
(A.H. CLARK 1947: 433). Previous experience suggests that, when A.H. CLARK admits that two or more species
are very similar, they are likely synonymous. The Karubar specimen is identified as A longicirra because the
aboral carina 1) is not obviously as prominent or continuous as in A. mirifica or A cristata, 2) appears stronger
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
647
than described for 4. macropoda, 3) may be reduced to one or more tubercles as in A.H. Clark’s description, and
4) resembles the figure of the type in Carpenter (1888, pi. 17). Also, the single axil with two tubercles on the
KARUBAR specimen approaches most closely the three-tubercle axils described for A. longicirra. YULIN (1984:
109) describes three specimens of A. mirifica from south of Hainan Island as having a high, thin median carina-
tion, the crest of which is strongly convex in lateral view". Although the clearly convex and continuous carinae in
the accompanying figure are typical of that species, they are distinctly weaker than in the KARUBAR specimen.
A. longicirra is the senior synonym among the four names. However, we do not formally synonymize any ol
the four nominal species pending examination of additional material. The aboral spines on the cirri immediately
distinguish the KARUBAR specimen from the other ten-armed asterometrid genus, Sinometra (Yulin, 1984).
Family CALOMETRIDAE A.H. Clark, 1912
Genus NEOMETRA A.H. Clark, 1912
REMARKS. — The genus Neometra currently consists of nine species distinguished on the basis of arm number
and ornamentation, relative lengths of cirri and cirrals, and cirrus ornamentation. Examination of material referable
to eight of these (including type material of six) suggests that several may not represent distinct taxa. For
example, a comparison of syntypes of N. gorgonia A.H. Clark (USNM 35556) and N. conanimis A.H. Clark
(USNM 35559) indicates that the two are very similar and probably synonymous. Both are endemic to western
Australia have broadly overlapping bathymetric ranges (Rowe & Gates, 1995). and were originally described
from material collected together (A.H. CLARK, 1947). N. gorgonia is larger, with longer cirri and more arms.
Perhaps its diagnostic pair of flanking aboral cirral carinae develop only once specimens reach a larger size than
those identified as N. conanimis.
The specimens described below as N. cf. alecto belong to a group of imperfectly distinguishable species that
differ chiefly in the nature and strength of their brachial ornamentation. They are discussed following the
description. . , , ,
Some confusion also exists at the generic level. A.H. Clark (1947) distinguished Neometra from
Gephyrometra A.H. Clark chiefly as follows: in the former, interradial extensions of the radials are broad and
"entirely and usually rather widely separate the bases of [adjacent] IBri" (= Ibri) (p. 355); in the latter, the
extensions are undeveloped or narrow and the bases of adjacent Ibri meet distal to them. However, in several
specimens of N. multicolor , including the holotype (USNM 22619, 35452, 35465), the interradial extensions are
very narrow, and the proximolateral corners of adjacent Ibri are barely separated or actually meet beyond the radials.
Similar variations exist in a syntype of N. conanimis (USNM 35559).
We offer no formal revision of the genus due to the relatively small number of specimens examined. Data m
Rowe and Gates (1995) (e.g., first records of N. diana and N. multicolor from Australian waters; range
extensions of N. conanimis and N. gorgonia) indicate that substantial new material exists that should be examined
before the genus can be adequately revised. However, the KARUBAR collection also includes eight specimens that
differ so much from known Neometra species that they require recognition as a new species.
Neometra cf. alecto (A.H. Clark, 1911)
Fig. 8
Calometra alecto A.H. Clark, 1911a: 544.
Neometra alecto - A.H. Clark, 1947: 367.
MATERIAL EXAMINED. - Indonesia. KARUBAR. Kai Islands : stn DW 22: 2 specimens (MNHN-EcCh 30).
DESCRIPTION - Centrodorsal discoidal; diameter 2.9-3.5 mm. Pole smooth, flat or slightly conical, with
traces of obsolete sockets around margin. Cirri XI-XII, 30-44. longest 21mm. in single marginal row; all
648
C.G. MESSING, N. AMEZ1ANE & M. ELEAUME
segments short. C5-C7 longest, LAV 1.0- 1.3 (on small cirrus of 30 segments, C4-C6 longest, L/W 1.7- 1.8). Aboral
process first appearing on C4 to C6 as distally-directed spine (sometimes transversely widened). On c 1 2-c 1 5 and
following segments, spine lengthens along cirral axis as a curved, distally sharp carina, shortening to a compressed
blunt spine on distal very short cirrals; distal few cirrals about as long as broad. Opposing spine large, blunt
triangular, compressed.
Rays narrow, well separated; aboral surface at least weakly carinate beyond radials. Radials short midradially
with truncated triangular interradial extensions separating adjacent Ibrj . Rays bear two IIBr2. one IIBr2 and one free
arm, or two free arms. Lateral adambulacral margins of IBr ossicles bear thin flanges, with margins converging
distally; lateral flanges absent on IIBr2; proximal third to half of Ilbri united interiorly. In larger specimen, weak
narrow midaboral ridge strongest at synarthry on IBr series, terminating in tubercle at one or both distal margins of
Ibr2 and IIbr2 axils; one IBr2 series almost smooth. Smaller specimen (lacking mature IIBr series) with a thin,
bluntly dentate midaboral ridge running from middle of lbri to middle of axil.
Arms 12 to 18 or, possibly, 19; ray length to at least 80 mm; brr 1 -2 with midaboral ridge strongest at
synarthry (sometimes strong and dentate on smaller specimen); br2 expanded exteriorly for attachment of enlarged
base of Pi; expansion reduced on bra; bri through bra or brs oblong. Following brr cuneate, almost triangular by
about bri 3, strongly cuneate again by about br20, and gradually less cuneate distally. Distal margin of each br
following br2 with narrow, distally-directed, sometimes spout-like, spinose projection, midaboral on brri-6, offset
to longer side of following brachials, weaker and spinelike on distal brachials.
Pi up to 31 segments, 8 mm, but sometimes much smaller and weaker. First segment greatly enlarged,
convex, produced on side toward arm tip as thin curved carina; second segment much smaller, thick, short, with
converging lateral margins, W/L 2.6; following segments small, short and cylindrical. P2 =17 segments, to
14 mm long, with small spines at distal corners of distal segments; enlargement of first segment much less than
on Pi, with thin rounded carina on side of basal two segments facing arm tip. One arm with Pi and P2 similar,
small, flexible and slender. P3 similar to P2 or longer, stiffer and slightly more robust, up to = 19 segments,
15 mm long, with spines at distal corners of all but basal two segments. Following pinnules gradually decreasing
in length and number of segments; carina on basal two segments lost by P4.5; Ps 14 segments, 10 mm long;
middle pinnules of 15 segments, 10.3 mm long. Proximal pinnules taper over their entire length. Middle pinnules
more straight-sided, tapering mostly over shorter distal segments so that these pinnules appear more robust. Middle
and distal pinnules often with several short thin distal segments. Pinnules in the smaller specimen as follows: P|
25 segments, 6.5 mm; P2 and P3 = 15 segments, 12 mm (tips broken); Pg = 8.5 mm; middle pinnule (P„)
17 segments, 10.5 mm; distal pinnule to 19 segments, 10.9 mm.
No disk.
Color (in alcohol). Cirri banded brown and white; centrodorsal and brachitaxes white; brachials yellow
midaborally, purplish brown laterally, white at pinnule bases; with occasional pairs of brachials white; pinnules
banded white and pale brown.
Distribution. — Previous records extend from the Lesser Sunda Islands, Indonesia, to the Philippines and
east of Hainan I., China, in 77-208 m (A.H. Clark, 1931). The Karubar collection falls within this depth
range, but extends the distribution eastward to the Kai Islands.
Redescription of the Holotype OF N. alecto. — (USNM 27494, 10°10'40"N, I24°02'45"E, 77 m. Bohol
Straits, Philippines, "Albatross" sta. 5414, 24 March 1909). This redescription emphasizes differences between the
Karubar specimens and holotype.
Larger than the Karubar specimens. Centrodorsal diameter 5.0 mm. Cirri XV, 44, longest 27.6 mm; C6-7
longest, L/W 1.0- 1.1. Triangular interradial extensions of radials not truncated. Lateral adambulacral flanges
weakly developed and only slightly converging. Adjacent Ilbri united interiorly for most of their length, with
distolateral margins convex rather than concave as in the Karubar specimens; IIbr2 axil lacking distal tubercles.
Brachials almost triangular by brrg- 1 1 , remaining triangular to about mid-arm; distal brr short and cuneate, W/L
1.8-2.0; brr 1 -2 with a rounded synarthrial swelling; midaboral distal margins of brr2-7 flared and spinose, but not as
strongly as in the Karubar specimens. Flared distal margins subsequently slightly overlapping next brachial,
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
649
weakening and sometimes disappearing on distal brr, not spinelike as in KaRUBAR specimens. On P2. distal
margins of distal segments rimmed with spines. Middle pinnules of up to 17 segments. Distal segments of middle
and distal pinnules with stronger distal spines than in Karubar specimens. Color pattern similar to that of
KARUBAR material, but some pinnules entirely brown or white; no paired white brachials.
Fig. 8. — Neomelra cf. alecto (EcCh 30): a, cirrus (shown in two sections; the distal several cirrals of the proximal
section are viewed slightly obliquely); b. aboral view of base of one ray; c. Pi: d, P2; e. aboral view of five middle
brachials. Scale bars: a = 1.0 mm; b, e = 1.0 mm; c. d = 2.0 mm; enlarged pinnulars of c = 1.0 mm; enlarged pinnulars
of d = 0.5 mm.
REMARKS. — Differences between the holotype of N. alecto and the Karubar specimens appear largely related
to size and ornamentation. Similar variations between these and other Neometra species blur current diagnostic-
distinctions. For example, the low alternating spinelike projections on the distal brachials of the KARUBAR
specimens are better developed than in the holotype of N. alecto, but are as described in the type of
650
C.G. MESSING. N. AMEZIANE & M. ELEAUME
N. spinosissima (A.H. Clark. 1947: 365). Neometra diana (A.H. Clark) and N. sappho A.H. Clark differ from
N. alecto, N. conanimis A.H. Clark, N. spinosissima and N. multicolor (A.H. Clark) in having relatively longer
cirrals (i.e., L/W of longest cirrals about 2.0 versus 0.7- 1.1). However, the Karubar specimens span much of
this range: longest cirral LAV is 1 .0- 1 .3 on large cirri and 1 .7- 1 .8 on a smaller cirrus.
Neometra diana supposedly differs from all other species in the genus in having very tall, narrow synarthrial
swellings clearly visible in photographs of the type specimen from "Siboga" station 294 (A.H. Clark. 1947,
pi. 37, figs 195-196). However, in syntypes from the same station (USNM E476, 3 specimens), these swellings
are only weakly to moderately developed, and the IBr series are similar to those of the Karubar N. cf. alecto
minus the latter's distal axil tubercles. Further, one N. diana syntype bears a pair of distal tubercles on the axil of
a regenerating IIBr series.
Neometra xenocladia C. G. Messing, sp. nov.
Fig. 9
MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands: stn DW 18: 2 specimens (MNHN-EcCh 32). —
Tanimbar Islands: stn DW 50: 1 specimen (MNHN-EcCh 31); 5 specimens (MNHN-EcCh 33).
Types. — The specimen (MNHN-EcCh 31) collected at stn DW 50 is the holotype. The 5 other specimens
(MNHN-EcCh 33) collected at the same station are paratypes.
Description of Holotype. — Centrodorsal a slightly convex disk; diameter 4.6 mm. Cirri XIX, 29-34.
longest 21 mm, in partly double marginal row; C7-8 longest, LAV 1.0; following cirrals decreasing in length; distal
cirrals much wider than long. Proximal cirrals beyond basal three or four wider distally and weakly carinate
aborally (sometimes also orally); aboral carina gradually increasing in height, occupying entire length of cirral, and
becoming curved, distally pointed and sharp; on distal several cirrals it becomes a shorter, compressed, distally-
directed carinate spine, sometimes tonguelike or triangular. Opposing spine larger than preceding spines,
triangular, fat but somewhat compressed.
Radials short midradially; interradial extensions long, tonguelike, parallel-sided.
Remaining four rays (including one detached) bearing two IIBr2 each. Adjacent IBr2 separated, with low narrow
midaboral ridge; Ibr i short, with diverging lateral margins thinning toward distolateral corners, W/L 3.1; Ibr2 axil
with parallel or converging, thin, flanged adambulacral margins, W/L 1.8. IIBr2 ossicles about as long as those of
IBr2, with adambulacral flanges absent or narrow; adjacent Ilbri united proximally, W/L 1 .7; rounded synarthrial
swelling extending from middle of Ilbri to middle of IIbr2 axil; axil W/L 1 .5.
Anris probably 20 (4 on each of 4 remaining rays); ray length = 80 mm; brrj-2 both slightly longer exteriorly
with rounded midaboral synarthrial swelling extending from middle of bri to at least middle of br2; bn mostly
united interiorly, W/L 1.7; bn with large exterior convexity to accommodate enlarged proximal segment of Pi,
W/L 1.3; bn+4 L3 mm across, W/L 1.1; brs oblong, W/L 1.7. Following brr euneate, W/L 1.3- 1.4, with low
midaboral ridge that widens distally on each brachial and becomes offset to longer side of brachial by brr6-7 (so that
ridge appears gently zigzag or sinuous); distal margins raised and smooth, overlapping next brachial slightly or not
at all. Brachials triangular or almost so by brs, W/L 1.2; euneate again by mid-arm, W/L 1.4; low offset aboral
ridge still present. Distal brachials short, strongly euneate, with very low aboral ridge. W/L 1.6. First syzygy at
br3+a; second at bri3+i4 to brn+is; next interval five; distal interval chiefly four (some five).
Pi 34 segments, 8.4 mm; first segment greatly enlarged, projecting toward arm tip as huge. thin, convex,
lanlike expansion; second segment short, hall as wide as first, with converging lateral margins, W/L 2.5;
following segments abruptly much narrower, short and cylindrical. P2 23 segments, 8.8 mm; as thin as P| but
still and straight; expansion of first segment smaller than in Pi; second segment as in Pi; third segment squarish;
lollowing segments longer than wide, carinate (i.e., triangular in cross-section), gradually shorter and thinner
distally; middle segments with L/W 3.0; middle and especially distal segments with spines at distal corners. P3 19-
22 segments, 10.1-11.6 mm; similar to P2 but thicker, more robust, stiffer and spinelike; expansion of first
segment weaker than in P2, but second segment moderately expanded; third segment longer than wide; distal
segments with strong distal spines, max. W/L 3.6; distal one or two to several segments short. P4 similar to P3
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
651
but all tips broken. Subsequent pinnules gradually decreasing in length, similar to Pa but more strongly carinate
and tapering only near tip; P7 16 segments, 7.4 mm; P10 19 segments, 8.2 mm; carina on basal two segments
disappearing by P5-6. Distal pinnules longer, more slender and flexible than middle pinnules, but generally similar,
with several short distal segments; up to 21 segments, 1 1 mm.
Disk missing.
Color (in alcohol). — Centrodorsal and IBr series yellow, cirri banded brown and yellow, IIBr series and arms
purplish-brown, pinnules purplish-brown, some with yellow tips.
FlG 9 — Neometra xenocladia sp. nov.. holotype (EcCh 31): a. aboral view of base of one ray; b. cirrus; e. P y. d. P2;
e. Pi. Scale bars: a = 1 .0 mm; b = 1 .0 mm; c-e (lower right bar) = 2.0 mm; enlarged pinnulars = 1.0 mm.
VARIATIONS in other Specimens. — Centrodorsal discoidal or rounded pentagonal; diameter 3. 1-4.6 mm;
pole flat or slightly convex. Cirri XI-XVII, 22-42 (chiefly 30-36), longest 14-26 mm (chiefly > 21 mm), in single
or partly double marginal row. One to four successive segments between 05 and cio (e.g.. c6, c5-8, or C7-10)
652
C.G. MESSING. N. AMEZIANE & M. ELEAUME
longest, L/W 1.0- 1.4. Aboral carina sometimes terminating in a small distal spine on C7-9. Carinate spines on
distal cirrals occasionally blunt.
Radials sometimes with midradial tubercle; interradial extensions occasionally with slightly converging sides,
sometimes extremely narrow.
Midaboral ridge on IBr series sometimes low and rounded; Ibri W/L 2. 8-4.2; lateral margins of Ibri rarely with
thin, variably shaped (e.g., tongue- or ear-like), adambulacral flange extending distal to interradial extensions of
radials; Ibr2 axil with W/L 1.4- 1.8. One or two IIBr2 per ray: Ilbri W/L chiefly 1.5- 1.6 (to 1.9); Ilbri axil with
synarthrial swelling weak or strong and rounded, W/L chiefly 1.3 (to 1.7). IlIBr2 developed interiorly, when
present, similar to preceding brachitaxes but completely rounded laterally without adambulacral flanges.
Arms 10-21; ray length 50 to at least 80 mm (broken in most specimens); brri-2 oblong or longer exteriorly;
br 1 W/L chiefly 1.7 (1. 5-2.0); br2 W/L 1.2- 1.8; exterior convexity may be enormous and sometimes also present
as far as brs (but gradually weakening); br.3+4 diameter 1.2- 1.3 mm, W/L 1 .0-1.2; br.s oblong or slightly cuneate.
W/L 1.7-1. 9. Following brachials cuneate, W/L 1.3-1. 7. triangular by brio to bri2, W/L 1. 2-2.0 (chiefly 1.5-1. 7);
brachials strongly cuneate again beyond mid-arm, W/L 1. 4-2.2; distal aboral margins often thin; low midaboral
ridge or keel offset to longer side and/or gently curved by br6 to bri2- Distal brachials with midaboral ridge ranging
from virtually absent to well developed, W/L 1. 6-1.7. Second syzygy variable among specimens, chiefly between
br9+l0 and bris+19 (extremes: brs+9 and br24+25); distal interval chiefly five or six (mostly three or four in a few
specimens, occasionally up to 11).
Pi 24-35 segments (chiefly > 30), 6.4-S.4 mm; second segment with W/L 2.0-2. 5. P2 19-23 segments, 5.8-
9.7 mm (chiefly > 8.0); sometimes slightly more robust than Pi; second segment as in Pi or with weaker
expansion than on first segment; distal segments with L/W up to 5.0. One arm of one specimen with P2 similar
to Pi. P3 17-22 segments, 10-12 mm; expansion may be absent on second segment. P4 up to 15 segments,
8.3 mm. P2 through P5 to P7 increasingly wider and less tapering along most of length; tapered portion
progressively more restricted to shorter distal segments. Middle pinnules 15-19 segments, 7-8 mm (chiefly 15-16.
< 7.5 mm); expansion of basal segments persisting to between P6 and P10.
Disk (retained in one specimen) globular, covered with pavement of Hat plates.
Color (in alcohol). — Chiefly yellowish white with pale purple bands on cirri, pinnules and, sometimes,
proximal arms.
Description of a small Paratype (EcCh 33). — Centrodorsal discoidal: diameter 2.4 mm; middle of pole
low rounded conical. Cirri XIII, 22-30, longest 14.4 mm; in partly double marginal row; C5-8 longest. L/W 1.2
(a small cirrus has C5-6 longest, L/W 1.3); curved aboral carina never occupying entire length of segment. On one
small cirrus, curved carina present on only two cirrals before becoming crescentic spine; on another small cirrus,
carinac are successively straight-profiled (on proximal cirrals), triangular, and crescentic (on distal cirrals).
Radials longer midradially than in larger specimens; lateral margins of interradial projections slightly
converging; midaboral tubercle absent. Ibri oblong with converging lateral margins. W/L 2.6; Ibr2 W/L 1.4. One
IIBr2 present. Arms 1 1 ; ray length 50 mm; br 1 wider exteriorly, W/L 1 .6; br2 with large exterior convexity for Pi,
W/L 1.4; br.t+a diameter 0.9 mm, W/L 1.0, constricted at articulation; br.s and following brachials cuneate, W/L
1.6; bri and br2 with weak synarthrial swelling; br3+4 and following brachials with weaker midaboral ridge; br.s and
following brachials with slightly raised distal midaboral margins. Brachials triangular by brs, W/L 1.6; weak
midaboral ridge slightly curved (convex to longer side of br) but not offset, so ridge appears gently sinuous along
arm; distal margin at end of ridge slightly raised. Brachials strongly cuneate by about bris, W/L 1.4. Distal
brachials short, cuneate, W/L 1.3; midaboral ridge as strong as on proximal portion of arm. Syzygies at br.3+4.
brs+9 to br9+io; distal interval three to five (chiefly four).
Pi 30 segments, 5.2 mm, second segment W/L 2.0. P2 12 segments, 5.2 mm, slender and stiff; first segment
as in larger specimens; second segment not expanded, W/L 1.3- 1.5; following segments longer than wide, up to
W/L 5.0 in distal half; distalmost segment short. P3 13 segments, 6 111m. P4 similar to P3, 10 segments, 4.6 mm;
P5 13 segments, 5 mm. P9 14 segments, 5.6 mm; distal six segments shorter than proximal segments but still
longer than wide; maximum L/W of longest distal segments 3.7. Distal pinnules as in larger specimens;
15 segments, 6.7 mm.
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
653
ETYMOLOGY. — From the Greek xenos (foreign, strange) and dados (branch) because the IIIBr scries develop
interiorly rather than exteriorly as in other members of the genus.
Distribution. — Known only from the Karubar collection: Kai and Tanimbar Islands, 186-205 m (possibly
184-212 m).
Remarks. — At least three other species of Neometra develop brachitaxes beyond IIBr. Unlike N. xenodadia ,
their IIIBr series develop exteriorly (at least initially). N. acanthaster (A.H. Clark) bears up to 30 arms with
exterior IIIBr2. N. sibogae and N. gorgonia bear up to about 40 amis with IIIBr2 developed exteriorly or on both
IIBr branches. In N. sibogae, !VBr2 develop only exteriorly. In N. gorgonia, IVBr2 develop interiorly on the
exterior IIIBr. A specimen identified as N. spinosissima from Phuket. Thailand (USNM El 1622) (actually similar
to N. alecto and N. conanimis), also bears a single exterior IIIBr2 series.
N. xenodadia shares with N. diana an aboral ridge on the brachials, but lacks the latter's dislal triangular or
spinous projections. In N. diana, the IBr series is similar in shape to that illustrated above for N. cf. alecto, very
different from the short IBr series of N. xenodadia with its diverging margins.
Family THALASSOMETRIDAE A.H. Clark, 1908
Remarks. — The family Thalassometridae is in dire need of revision. Many of the features used by
A.H. Clark (1950) to distinguish species, genera and groups of genera are either inconsistently applied or widely
variable, sometimes ontogenetically. As an example, all genera but one ( Horaeometra , with uniquely expanded
genital pinnules) fall into two supposedly distinct groups: those with arms laterally compressed and carinate at
least distally, and “Pi longer and proportionally stouter than P2, though not otherwise dillerenf' (6 genera), and
those with arms aborally "rounded to the tips, never carinate; . . Pi usually much enlarged, very stout and much
broadened basally" (7 genera) (A.H. Clark. 1950: 5-6). However, the cross-sectional shape of the distal arms in
some species of Cosmiometra (group one) is essentially the same as in Oceanometra (group iwo). Parametra,
which A.H. CLARK included in the first group in his key. nevertheless includes two species that bear aborally
rounded arms with Pi more or less enlarged basally (typical of group 2). Clark (1950: 64) relered to these as
transitional between the two groups, but on this basis they would be impossible to identity using his key. As
a result of these difficulties, we identify several of the taxa in the KARUBAR collection only tentatively.
Genus AGLAOMETRA A.H. Clark. 1913
Aglaometra valida (Carpenter, 1888)
Figs lOa-c, lle-f, 12b, 13-14; Tables 2a-b
Antedon incerta Carpenter, 1884: 57, 83. pi. 54 figs 6-7 (nomen nudum); 1888: 106. pi. 18 figs 4-5.
Antedon valida Carpenter, 1888: 104, pi. 15 figs 5-8.
Crotalometra eupedata A.H. Clark, 1909c: 404; 1 9 1 2d: 209.
Crotalometra valida - A.H. CLARK, 1912c: 33. 209.
Crotalometra propinqua A.H. Clark, 1911: 549; 1 9 1 2d: 209.
Crotalometra incerta - A.H. CLARK. 1912c: 33, 209.
Crotalometra veto A.H. Clark, 1912b: 427.
Crotalometra sulcata A.H. Clark, 1912b: 427.
Aglaometra valida - A.H. Clark, 1913: 47; 1918: 163, 164, 273; 1950: 108, 109. pi. 13 figs 38-39.
Aglaometra eupedata - A.H. CLARK, 1913: 47; 1918: 163. 164; 1921: 179 fig. 225.
Aglaometra incerta - A.H. CLARK. 1913: 47; 1918: 163, 165; 1950: 108, 114.
Aglaometra vera - A.H. CLARK, 1918: 163. 165. pi. 21 tig. 54.
654
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Aglaomeira propinqua - A.H. Clark, 1918: 163. 165; 1950: 108, 116, pi. 15 fig. 47.
Aglaometra sulcata - A.H. Clark, 1918: 163. 164, 273, pi. 21 fig. 53; 1950: 109. 118, pi. 13 fig. 40.
Material EXAMINED. — Indonesia. Karubar. Kai Islands: sin CP 19: 4 specimens (MNHN-EcCh 46) — Sin
20: 1 specimen (MNHN-EcCh 45). — Sin CC 21: 15 specimens (MNHN-EcCh 43); 8 specimens (MNHN-EcCh 44).
Description. — Centrodorsal blunt (sometimes rounded or truncated) conical, with strong interradial
projections; midradial margin moderately to deeply excavated (interradial extensions and midradial concavity reduced
or absent in smaller specimens); basal diameter 3.7-7.8 mm; midradial height 2.7-6.4 mm. Aboral pole conical,
rounded or flattened, smooth, spinose or tuberculate; diameter 1. 4-4.3 mm. Cirrus sockets in two columns per
radial area, with two to four functional sockets per column, often with one or two apical obsolete sockets. At least
peripheral sockets separated midradially by flat triangular space; smaller apical sockets contiguous.
F'G„!°;,7 a-c\AS‘aome!ra valida (a-b, EcCh 43; c, EcCh 46), bases of single rays (radial and centrodorsal margin
° 0 mm m C Sh0W‘ng vana,lons ln ornamentation; d. Cosmiometra iole (EcCh 34). base of single ray. Scale bars =
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
655
Cirri XXI-XXXVIII, 51-77, 48-75 mm long; ci to C4-7 shorter than wide; next one or two cirrals squarish
(LAV = 1.0); following four to seven longer than wide; C7 or C8 usually longest, with L/W chiefly 1.5- 1 .7 (rarely
1.2-2. 2); following cirrals gradually shorter; distal cirrals short (L/W < 1.0); cirrus tapered near tip. Transitional
cirral usually cs (rarely C6-cio). Small aboral spine, triangular in lateral and cross sections; first appearing between
C7 and ci 5 (chiefly C9 to c 12); increasing in height and length over next several cirrals and rounded triangular in
cross section; becoming taller and triangular or distally curved in distal half of cirrus; not carinate. Aboral profile
of middle cirrals from proximal end of ossicle to spine tip ranges from slightly concave to straight or with low'
convex hump. Well-developed spines on distal half of cirrus sometimes with distal margins finely dentate and with
microspines on distal face. Spines weakening and sometimes becoming rounded in lateral view on smaller
segments near cirrus tip. Opposing spine tall, triangular and sharp or low and rounded, similar to preceding spines.
Terminal claw gently curved, usually longer than penultimate segment.
Fig. 11. — Proximal pinnules, a , Stenomeira cristata (EcCh 56). Pi (tip shown from 2 angles); b. Cosmiomeira
philippinensis (EcCh 37), Pi ; c, Cosmiomeira iole (EcCh 34), Pi; d, Oceanometra anncmdalei (EcCh 51), Pi;
e-f, Aglaometra valida (EcCh 46): e, Pi; f, P2- Scale bar = 1.0 mm.
656
C.G. MESSING, N. AMEZIANE & M. ELEAUME
Fig. 12. — Cirri: a, Oceanometra annandalei (EcCh 48) (interrupted at x); b, Aglaomeira valida (EcCh 44) (interrupted
at y). Scale bar: a, b = 4.0 mm; enlarged cirrals = 1.0 mm.
Radials distally concave, usually very short, but well-exposed in some specimens; often swollen midradially to
conform to excavated margin of centrodorsal (rarely with midaboral tubercle); fine spines sometimes on aboral
surface or restricted to distal margin; interradial ends swollen, smooth or with 1-2 small spines.
Brachitaxes (IBr2) and first two brachials rounded to moderately arched aborally with broad, weak to moderate
synarthrial swellings; lateral margins everted. Hat, closely apposed and weakly rugose or spinose; no aboral
carination. Ibri short, shallow U-shaped, W/L chiefly 3.3-3.9 (range: 2.5-5.0); Ibr2 axil hexagonal or pentagonal,
with short slightly diverging lateral margins, W/L chiefly 1.6- 1.9 (range: 1. 3-2.0). Fine spines (rarely crenulae)
often along proximal margin of Ibri (sometimes also bri), rarely along distal margin of axil. Fine spines, teeth or
crenulae usually only along lateral thirds of distal margins (of Ibri and br i ) and proximal margins (of Ibr2 and br2);
a few tiny aboral spines rarely present.
I wo smaller specimens (stn CC 21) with small "water pores" produced by truncation of distolateral corners of
Ibri and (interiorly) bri, and to lesser degree proximolateral corners of Ibr2 axil and (interiorly) br2. One of these
with numerous slender lateral spines developed on distolateral corners of ossicles from Ibri to br2 or bra.
Arms 10, ray length 71-154 mm; bri short, at least shallowly U-shaped (oblong in small specimen with water
pores), W/L chiefly 2. 5-3. 7 (range: 2. 2-4.0); br2 about twice as long as bri, W/L chiefly 1. 7-2.1 (range: 1 .5-3.9);
both bri and br2 longer exteriorly; br3+4 constricted at articulation, diameter chiefly 2. 7-3. 3 mm (range: 2.0-
3.8 mm), W/L 1. 4-2.1; alternating articular tubercles weak or moderately developed (absent in some smaller spec¬
imens), brs to brr9-io (brn in one small specimen) oblong. Following brachials cuncate, usually becoming almost
triangular by brri6-17; brachials strongly cuneate again beyond mid-arm; weakly cuncate in distal third of arm.
Aboral surface of bra (rarely br2 or br6) and following brachials with numerous small weak spines (rarely tuber¬
cles); slightly larger spines sometimes present away from midaboral line, but spines usually absent from sides of
brachials; lateral ornamentation of brachitaxes rarely present as Tar as brr4-5. Beyond proximal third of arm, aboral
spines recumbent, distally directed, spread along distal margin of brachials. Beyond mid-arm, distal margins of
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
657
brachials raised, especially midaborally, with longer, stronger spines. Distal brachials compressed but not carinate,
with strong midaboral spines sometimes overlapping next brachial and producing moderately serrate arm profile.
FlG. 13. — Aelaometra valida (EcCh 44) (SEM): a-b. Centrodorsal: a, lateral view; b. oral view; c. aboral view of radial
circlet showing basal rosette; d-f, ray ossicles showing lateral flattening; d. Ilbri. muscular articulation; e. IbQ axil,
synarthry; f, Hbr3, syzygy. Scale bars = 1 mm.
First syzygy at bn+4 (rarely br6+7 to brs+9): second usually at bri3+i4 to bris+16; distal interval chiefly five to
seven, occasionally longer.
Pi up to 30 short segments, 17 mm (usually 21-25, 8-12 mm), often much more robust than following
pinnules; distal margins of distal segments spinose; bases of adjacent Pi (and Pa) usually apposed and flattened.
Pinnules decreasing in length and strength from Pi to P5.7: pinnules longer thereafter. Bases of proximal pinnules
often rugose or finely spinose, with one or both lateral margins sometimes extended as irregularly spinose flanges
(flange or spines often restricted to lateral margin facing arm tip). On middle pinnules, W/L of distal segments up
to 1 4; middle pinnulars sometimes slightly broadened. Distal pinnules up to 28 segments. 19 mm; basal two
segments trapezoidal and often much broader than succeeding segments; W/L of distal segments up to 1.5 (longer
in small specimens). Middle and distal pinnules with abambulacral margin carinate and spinose (sometimes first
appearing on distal segments of P2-3); sometimes with middle segments slightly broadened.
Disk densely covered, at least along ambulacra and on anal papilla, by small rounded nodules (sometimes
peanut-shaped or flattened); nodules usually absent from disk margin.
DESCRIPTION OF A SMALLER SPECIMEN (Stn CP 20). - Centrodorsal similar to preceding, but margin only
slightly concave midradially; peripheral cirri separated only by small triangular area; basal diameter 3.7 mm;
658
C.G. MESSING. N. AMEZIANE & M. ELEAUME
D/H 2.7. Small obsolete apical sockets almost cover pole except for small rounded apex. Cirri XXI. 58, 48 mm;
c 1-4 short; cs squarish; C6-I2 longer than broad, L/W 1.5; small proximal aboral spines more rounded in cross
section than in preceding specimens, but becoming more sharply triangular on shorter middle and distal cirrals;
opposing spine taller than preceding spines.
Radials well exposed with large midaboral round knob. Brachitaxis ossicles arched aborally with trace of broad
aboral ridge, not flat-sided or apposed. Ibr i not united laterally but in contact; lateral margins sometimes slightly
everted with one or two distolateral spines, and converging distally to form large water pores. Ibr2 axil hexagonal
with short diverging lateral margins; disto- and proximolateral margins weakly everted with several strong spines.
Arms 10; ray length 88 mm; bri almost oblong, not U-shaped; slightly longer exteriorly; slightly everted. Hat-
sided and apposed exteriorly with one or few interior distolateral spines; br2 larger, longer exteriorly; interior lateral
margin slightly everted with proximolateral spine; neither flattened nor apposed exteriorly. Water pores well
developed interiorly, smaller exteriorly; brri-3 with very weak trace of midaboral ridge; brrs-g oblong. Brachials
strongly cuneate, almost triangular by bri2, remaining almost triangular to mid-arm, less cuneate distally, strongly
compressed and weakly cuneate near tip. Fine aboral spines first appear weakly on about brio; very weak on
proximal arm; better developed and distally directed on distal half of arm. Distal intersyzygial interval generally
four or five (up to eight).
Pi and Pa weakly flattened if at all; lateral extended margins of proximal pinnulars rounded and lacking spines.
Weak spines present on distal segments of proximal pinnules. Middle and distal pinnules not expanded. Pinnulars
of distal pinnules elongated.
Distribution. — A.H. Clark (1950) recorded Aglaometra valida in the narrow sense from Mindanao.
Philippines, to northern Sulawesi (Celebes), Indonesia, in 457-1165 (possibly 1264) m. Inclusion of the three
other species in the genus (see Remarks following) extends the range to the northern Philippines (Luzon) the
Kermadec Islands and northern New Zealand (A.H. Clark, 1950; McKnight, 1977). The Karubar material falls
well within this range.
Remarks. — The four species recognized by A.H. Clark (1950) form a rough size gradient from the large,
robust valida, followed in descending order by incerta, propinqua and sulcata. Most Karubar specimens are
equivalent to propinqua, but the material spans much of the genus’ known variability and a few specimens closely
approach the other three species. As a result, the four are herein synonymized. Only cirrus length previously
recorded for A. valida exceeds that of the Karubar material. Previously recorded specimens of all four species bear
fewer cm (X-XX) than the Karubar material (XXI-XXXVIII), but this simply reflects a difference of one or two
versus two to four cm per column. Many Karubar specimens also bear one to several tiny immature peripheral
cirri that may not have been included in earlier descriptions.
One feature used by A.H. CLARK to distinguish A. valida from A. incerta — variations in ihe angle of the
profile ol the proximal part of the rays — is most likely a vagary of preservation. The water pores diagnostic of
A sulcata appear on two smaller Karubar specimens that are otherwise indistinguishable from the rest or from
t e yet smaller specimen described separately above. The expansion of the basal two pinnulars of the distal
pinnules that CARPENTER (1888) used to distinguishing valida from incerta also varies.
Although CARPENTER first mentioned Antedon incerta in 1884, the species cannot be identified from his
diagnostic specific formula (which applies equally to several ten-armed thalassometrids) or his comments about its
pinnules. CARPENTER described both Antedon incerta and Antedon valida in 1888 with the latter having page
precedence. When A.H. CLARK first mentioned Aglaometra (1913), he included valida and incerta but neither
diagnosed the genus nor indicated a type species. A.H. Clark finally designated valida as the type species of
Aglaometra in 1950.
ON Variations. - A close examination of variability in skeletal characteristics of Aglaometra valida reveals
several important features critical to their diagnostic and descriptive value.
1) Different body parts exhibit different levels of variability (Tables 2a-b). Most cirrus characteristics exhibit
narrower coefficients ; of variation (V) [8.4% (position of longest cirral) to 15.0% (number of cirri/column)] than do
centrodorsal traits [18.5% (basal diameter) to 27.1% (diameter of pole)] or proximal ray ossicle dimensions [9.7%
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
659
(Ibr2 width) to 22.6% (Ibn length)]. Features of major body parts thus vary somewhat independently. Each
specimen must be interpreted as a mosaic of somewhat independent traits.
TABLE 2a — Aglaometra valid* measurements and statistics lor 19 trails: centrodorsal. err, and P (n - - 28
Abbreviations: S,,.. specimen number; S,a.. station number; D/H. ratio of basa! d.ameter to mtdrad.al height
S centrodorsal surface area [it x (basal diam./2) x midradial height]; No . Cirri, number ol cirri; Max. L max mum
As an example, centrodorsal surface [taken as the surface of a cone expressed as S = tt x (basal diameter / lx
midradial height] varies from 15.7 to 68.6 mm2 with V = 37.2%. If cirri increase m number co^p°nd,ng >
their number per unit surface area (U = number of cirri / S) should remain relatively constant. However. U vanes
from 0 44 to 1.55 cirri mm-2. Number of cirri is thus not correlated with centrodorsal surface area 1 he cm , are
concentrated on the proximal two-lhirds of the centrodorsal, leaving a naked apex (recorded as pole diaiger
Table 2a) The total surface may thus be divided into an area occupied by cm that remain relatively undo n ly
spaced, and a variable naked aboral pole. A bivariate analysis (not shown here) in which polar diameter correlates
660
C.G. MESSING, N. AMEZIANE & M, ELEAUME
inversely (r = 0.70) with cirri per unit area (U) confirms this. Centrodorsal size thus varies independently of the
number of cirri (except for the smallest specimens). Variation in aboral pole dimensions account for differences
between small and larger centrodorsals.
Table 2b. — Aglaometra valida; measurements and statistics for 16 traits: rays (n = 28). LAV, length to width ratio;
W/L, width to length ratio. Other abbreviations as in Table 2a.
2) With a few exceptions (e.g., smaller specimens may have water pores and a less excavated centrodorsal
margin), shape does not correlate closely with size. It seems reasonable to assume, at this stage, that smaller
specimens represent younger individuals. The Karubar specimens span a substantial size range that likely reflects
age heterogeneity (at least for the smaller specimen described above). However, most descriptive traits (Tables 2a-
b) reveal no distinct pattern of shape change with increasing size. For example, figure 14a illustrates the lack of
any such pattern in centrodorsal shape plotted as basal diameter versus midradial height. Also, A. valida bears only
ten arms, a possibly plesiomorphic or paedomorphic trait acquired at a young stage, and the number of cirri varies
little relative to increasing size (see above). A. valida thus appears to develop rapidly to a point beyond which size
continues to increase considerably while shape is conserved.
3) Serial components of a body part may also exhibit different levels of variability both between and within
components. For example, Ibr i varies much more in width than do the following ossicles (Fig. 14b. lop).
Regeneration cannot account for the variation because rays autotomize and regenerate at syzygies, which do not
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
661
appear in A. valida proximal to br3+4- By contrast, Ibr 1 and bri vary much less in length than does Ibn (Fig. 14b.
bottom). The distal articulation in the former two is a synarlhry, suggesting that variability may be constrained by
both structure and function.
Fin |4 _ Aelaotnetra valida : a. Plot of basal diameter versus midradial height of centrodorsal. Individual data points
show centrodorsal shape (derived from heigh,, basal diameter and apical diameter measurements . b Maxtmum
(squares) and minimum (diamonds) widths (top) and lengths (bottom) of proximal ray ossicles (lbn, Ibr., hr,. r_,
brj+4) (n = 28).
Genus COSMIOMETRA A.H. Clark, 1909
Cosmiometra iole A.H. Clark, 1950
Figs lOd. 11c, 15a
Cosmiometra iole A.H. Clark, 1950: 43, 61. pi. 4 lig. 14.
MATERIAL EXAMINED. — Indonesia. KARUBAR: Tanimbar Islands : stn CP 86: 4 specimens (MNHN-EcCh 34).
DESCRIPTION — Centrodorsal low hemispheric; diameter 5.0-6.4 mm; D/H 2.2-2.8: periphery slightly
concave radially or not excavated at all; small protruding interradial process sometimes developed as short narrow,
projecting ridge. Aboral pole irregularly convex, with several low tubercles or irregularly eroded. Cirrus sockets
crowded three across each radial area, 1-2 high, not organized in distinct columns.
662
C.G. MESSING, N. AMEZIANE & M. ELEAUME
Cirri XXVI-XXXV, 30-47. 27-36 mm; short relative to ami length for a thalassometrid; ci-3 or C4 short; C4 or
C5 slightly longer. L/W 0.7; C5 or C6 longest, slightly constricted, transitional. L/W 1. 8-2.0; following cirrals
compressed but not carinate, gradually shorter; c 1 1 or ci3 with LAV 1.0; ci sometimes with a distinct aboral knob.
Slightly subdistal aboral transverse ridge (weakly dentate, convex in cross section) first appearing on cs or cf„
increasing in size on following cirrals, initially triangular in profile, becoming a crescentic, compressed, strongly
hooked spine on shorter distal cirrals. Alternatively, the transverse ridge strongly rounded in profile (triangular in
profile on a few cirrals if at all); becoming a tall, erect or distally tilled, compressed, tab- or tongue-like process on
distal cirrals; rarely hooked and sharp; sometimes eroded, especially near cirrus tip. Opposing spine a distally tilted
triangle, no larger than preceding spines. Terminal claw gently curved, as long as or longer than penultimate cirral.
Radials barely visible midradially, if at all; smooth, with one or two round tubercles or an irregular distal
margin; corners rising to tall round tubercles; tubercles of adjacent radials meet distal to short interradial
projections of centrodorsal.
All brachitaxes (I and IIBr) of two ossicles each; moderately arched, flat-sided and laterally apposed, with
rounded synarthrial swellings and smooth, low midaboral ridges. Ossicle margins weakly to moderately everted, at
least in lateral portions (never midaborally); eversion smooth, irregular, or with weak rounded teeth, strongest on
Ibri and proximolateral margins of Ibr2 axil, weaker distally but detectable on bri in one specimen. Ossicles joined
by synarthry have lateral margins projecting laterally well beyond synarthrial ligaments, leaving a small water pore
interiorly between adjacent Ilbri and IIbr2, and between adjacent bri and br2. !Br2 series with gently diverging
lateral margins; Ibri short, shallowly concave distally, W/L 3.3-4.0; Ibr: hexagonal, W/L 1, 8-23). Ilbri slightly
longer exteriorly, W/L 3.0; libra hexagonal with straight lateral margins, W/L 1.4- 1.5. One interior IIIBr2 series
present in one specimen.
Arms 20-21; ray length 110-120 mm. Proximal brachials flat-sided to between bri2 and brao; brachials in
proximal third of arm very tall (along ambulacral-abambulacral axis); low. narrow, midaboral ridge of brachitaxes
continued on all brachials. Middle and distal brachials moderately compressed so that they appear carinate; distal
margins finely spinose by about br20, raised and sometimes with stronger spines, especially midaborally,
producing weakly serrate profile on middle and distal arms; distally projecting midaboral ridge spines sometimes
overlapping base ol next brachial; bri slightly longer exteriorly, with interior lateral margin weakly everted, W/L
2.2; br2 larger, W/L 1.3; br3+4 squarish, 1.7-2.0 mm across, W/L 1.2; brs to brrg-io slightly cuneate, W/L 1.7-
2.5. Next two or three brachials almost oblong (clearly less cuneate), sometimes slightly shorter than preceding
brachials (very short in side view; almost four times taller along ambulacral/abambulacral axis than long). W/L
1.9-2. 2. Following brachials becoming almost triangular by about brig, W/L 1.8-2. 1. Brachials cuneate again
beyond mid-arm, W/L 1.5; distal brachials short, cuneate, W/L 1.6-1. 8. Syzygies at br3+4; next at bri 3+ 1 4 to
br24+25; next interval seven to 13; distal interval mostly five or six (up to nine) articulations.
Proximal pinnules robust, of short stout segments. Pi of 15-19 segments, 9-12 mm; proximal 4-5 segments
flat, smooth, sometimes broadened and laterally carinate (side facing arm tip); following pinnulars smooth except
that carinate abambulacral lateral margin may rise to distal rounded triangular process or spine. P2 similar to
smaller Pi. 14 segments, 8.5 mm; or much smaller and narrower, 13 segments, 7.6 mm. P3 similar to P2 or
smaller, 10 segments, 5.2 mm. Following pinnules gradually shorter and weaker to about P7-8; Pr, 10-
11 segments, 5. 0-5. 6 mm; Ps 9 segments, 4.4 mm. Following pinnules gradually longer; middle pinnules Id-
15 segments, 7-8 mm; distal pinnules 17 segments, 9 mm.
Disk naked.
Color (in alcohol). — White; or proximal arms brown laterally; middle and distal syzygies and sometimes one
or two adjacent brr brown or tan; proximal pinnules banded brown and white.
Distribution. — Known only from Zamboanga in the southern Philippines (A.H. Clark. 1950) and
Tammbar Islands in Indonesia, in 225-292 m (possibly 223-365 m).
Remarks. — The Karubar material represents the only known specimens of this species apart from the two
collected by Mortensen’s Pacific Expedition in the Philippines in 1914 and described by A.H. Clark in 1950.
The specimens depart from the original description only as a result of their larger size. The description of the type
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
663
specimen, for example, cites XVII cirri, 25 mm long, of 29-35 segments, and Pi 7 mm long, of 14 segments of
which the distal are longer than broad.
FIG. 15. - Cirri: a, Cosmiomelra iole (EcCh 34); b, Cosmiomelra philippinensis (EcCh 37). Scale bar: a. b - 4.0 mm
enlarged cirrals = 1.0 mm.
Cosmiometra philippinensis A H. Clark, 1911
Figs lib, 15b; Table 3
Cosmiometra philippinensis AH. Clark, 1911: 548; 1912c; 215; 1918: 153. 155, 272. 274, pi. 23 fig. 61; 1950: 43.
52, pi. 5 fig. 16, pi. 6 fig. 22.
MATERIAL EXAMINED. - Indonesia. Karubar: Kai Islands : stn CP 05: 3 specimens (MNHN-EcCh 38). -
Stn CP 16: 16 specimens (MNHN-EcCh 37). - Stn CP 27: 1 small specimen (MNHN-EcCh 35).
DESCRIPTION. - Centrodorsal conical or truncated, rounded or blunt conical; covered with papillae, at least
basally; margin with shallow, usually V-shaped, midradial excavation; basal diameter 3.5-6.0 mm; m.drad.a
height 1 5-4 0 mm; D/H 1.3- 1.9. Aboral pole conical or rounded, smooth or papillose. Cirri in ten colun s
2-4 sockets per column; adjacent columns very closely crowded, with sockets alternating mierrad.ally; columns
separated midradially by triangular papillose space; ap.cal sockets still separated m.drad.a ly
Cirri XX-XXIX, 53-65, maximum length 37-45 mm. Cirr. compressed beyond basal few c.rrals tapered near
tip; c, to c4-6 short; c4-6 still shorter than long or squarish; next two or three cirrals longest, abrupt^ onger han
preceding L/W 1 1- 1 .8; following cirrals gradually shorter, squarish by cl0-i2; c5-7 trans.t.onal (the more proximal
cirrals usually brown; the more distal cirrals white); c,o-.3 and following cirrals becoming aboral y cannate with
slightly subdistal aboral tr.angular spine. Spines triangular in cross section, enlarging and lengthen.ng on
664
C.G. MESSING, N AMEZIANE & M ELEAUME
succeeding cirrals, with erect distal margin often bearing one or few teeth; spines becoming curved and crescentic
on shorter distal cirrals (proximal profile strongly convex, distal profile often slightly concave), smaller and
narrower (no longer with triangular transverse face) on distal few cirrals, still curved and sharp or sometimes weak
and rounded; length of spine base shorter than cirral length. Opposing spine blunt or triangular, distally directed,
much larger than preceding spines. Terminal claw gently curved, longer than penultimate cirral.
Radials hidden or visible as extremely narrow band or small midradial triangle, concave distally, covered with
fine, often spinulose, spines.
Brachitaxes all of two ossicles (I-IIIBr), gently to moderately convex aborally; lateral margins straight, flat and
apposed; Ibr 1 shallow V-shaped, W/L 3.0-4. 1; lateral portions thickened and Hared, rugose, or with spinulose
spines or papillae, often appearing spongy; ornamented area broadest proximally, continuous across Hared proximal
margin, but restricted to lateral quarters or thirds of aboral surface; Ibr2 axil hexagonal, short and wide. W/L 2.0-
2.1. Raised spongy ornamentation occupying progressively narrower lateral portion of successive ossicles,
disappearing beyond IIlbr2 axil. lIBr similar to IBr; llbri still thickened with rugose ornament exteriorly; smooth
and Hat except for few small distolatcral spines interiorly, W/L 3. 1-3.7; IIbr2 axil with spinulose thickening
stronger exteriorly, but sometimes reduced to narrow band of fine spinules, W/L 1 .4- 1 .6.
One specimen differs in having the proximal margin of Ibri slightly raised midaborally, if at all; one
IIBr4(3+4), and the IlBr series slightly Battened exteriorly, if at all. Another specimen has brachitaxes with low
synarthrial swellings and Ibr i with a weak midaboral ridge; ornament similar to that of preceding specimens,
somewhat weaker than most, but with entire aboral surface of brachitaxes and arm base ossicles bearing scattered
tiny spines (fewest midaborally).
Arms 18-20 (chiefly 20); ray length 65-105 mm. Proximal brachials more strongly convex than brachitaxes
ossicles; laterally Hat to brr4-5. Ornament similar to that of brachitaxes (ossicles weakly thickened and spinulose
laterally) present at most to bo exteriorly and brs interiorly, usually strongest at interior-distal corner of bn and
interiorly on br2; br j almost oblong, concave distally, W/L 2. 1-2.2; br2 larger, longer exteriorly, proximal margin
moderately convex, W/L 1.5-2.0; br3+4 diameter 1.5-1.8 mm, W/L I.2-1.3; br5 to brry-io short, oblong, with
slightly concave lateral margins, W/L 1. 7-2.0. Following brachials becoming cuneate (never triangular), W/L 2.3
at mid-arm. Distal brachials strongly compressed but never carinate, weakly cuneate, with longer side expanded for
pinnule base, W/L 1.0-2.0. Aboral surfaces of brachials beyond proximal fourth or third of arm developing weakly
raised distal margins and raised longitudinal striae (the latter sometimes first appearing on brr2-4) that terminate in
spines along distal margins. In distal half of arm, distal margins of brachials more strongly raised and spinose,
sometimes slightly overlapping next brachial; aboral surface covered with short, distally directed recumbent spines.
Fust syzygy at br3+4. second usually at br 1 6+ 1 7 to br23+24, but sometimes beyond br4o; distal interval four to
seven (sometimes eight or nine).
Pi 22-27 segments, 10-15 mm; proximal pinnulars short and broad; several with knob or thick rounded
carina on lateral margin facing arm tip; middle and distal pinnulars much narrower and longer than wide. Prox¬
imal broadening and carina much weaker on succeeding pinnules, usually persisting to P3 at most. P2 similar
to Pi or shorter, of 16-24 segments, 6.5-14 mm. P3 similar to P2, up to 18 segments, 10 mm. P4 usually
abruptly shorter, of 10-12 segments, 4. 5-6.0 mm (15 segments, 8.0 mm in one specimen). P6-7 usually shortest,
ot 9 segments, 3. 8-4.4 mm. Following pinnules increasing in length and strength; middle pinnules of
16 segments, 6.2-7.0 mm; distal pinnules of 15-17 segments, 7.S-9.8 mm. Most segments of middle and distal
pinnules (except basal and distal few) of about equal length, with spinose carinate margin; (L/W depends on
orientation to observer).
Disk naked.
Notes on three small Specimens (EcCh 37, 2 spec.; EcCh 38, 1 ). - Centrodorsal diameter 2.2-2.S mnv
midradial height 1.1- 1.4 mm; D/H 1.8-2.5. Cirri XVI1-XXVI, 39-42, 21-27 mm. Arms 13-17; ray length 47 to
about 55 mm. Two retain small apical cirri of 24-28 segments, 9.2-1 1.0 mm long.
Distribution. — Known from the southern Philippines, Sulawesi and the Kai Islands in 245-621 m Four of
Stat,°nS reC°rded by A H- CLARK (I931) occur in 245-320 "1. The Karubar material was collected in
299-390 m (possibly 296-502 m).
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
665
Table 3. — Cosmiometra philippinensis : measurements and statistics for 8 traits: centrodorsal, cirri and rays (n - 21).
Abbreviations as in Table 2a and b.
Remarks. — The holotypc (USNM 27497, Tagolo Point, Mindanao I., Zamboanga del Norte. 08°48'44"N.
123°27'35"E, no depth, "Albatross" stn 5523, 10.08.1909) falls well within the range of variation exhibited by the
Karubar specimens, with variations only in details of ornamentation, as follows: centrodorsal diameter about
4.2 mm, D/H 1.1; maximum cirrus length 40 mm. Aboral brachitaxis ornament consisting of fine spines,
covering lateral thirds of Ibri and best developed on lateral margins of succeeding ossicles (as in Karubar
specimens), but also occurring in row along proximal and distal margins, and reduced but present to brj. Arms 19;
ray length about 80 mm. In some other specimens identified as C. philippinensis by A.H. CLARK (USNM
E 19 176, E3214), the aboral brachitaxis ornament consists of flattened finely spinose tubercles adjacent to lateral
margins, similar to that in some KARUBAR specimens.
Genus CROTALOMETRA A.H. Clark, 1909
Crotalometra rustica A.H. Clark, 1909
Fig. 7b
Crotalometra rustica A.H. Clark, 1909b: 80, 147; 1950: 91. 92, pi. 11 figs 35-36. pi. 18 fig. 58.
Crotalometra rusticata - MCKNIGHT, 1977: 106.
666
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Material EXAMINED. — Indonesia. Karubar: Kai Islands: stn CP 16: I specimen (MNHN-EcCh 41). —
Stn CP 25: I small specimen, broken (MNHN-EcCh 42).
Description of LARGER Specimen (Stn CP 16). — Centrodorsal domelike, deeply excavated midradially;
diameter 1 1 mm; midradial height 6.5 mm; maximum interradial height 8 mm. Aboral pole flattened with weak
interradial ridges. Cirri XXVIII, 64-66, 96 mm, stout (relative to most other thalassometrids); in 10 columns of
two or three each; two columns of each radial area converging apieally around deep midradial triangular furrow
(with apical sockets not separated); columns of sockets on adjacent radial areas closely apposed interradially ; ci-4
short; C6 squarish; C7-9 barely longest, LAV 1.1; cio-18 almost as long but slightly more elongate (L/W 1.1-1.15)
due to slight narrowing of cirrus; following segments very gradually shorter; C22 transitional, squarish; following
several cirrals becoming shorter rapidly, LAV 0.6 by C28- Distal aboral margin of C23 slightly raised, becoming
enlarged over next several cirrals as a rounded process, rounded or rounded triangular in transverse section, never
sharp or carinate. Processes on short distal cirrals with convex proximal profile and concave distal profile, the latter
tilted back from distal edge of cirral and sometimes weakly dentate. Cirrus tapering dislally with aboral processes
weaker and more rounded. Opposing spine low rounded triangular. Terminal claw with blunted tip. as long as or
longer than penultimate cirral.
Radials extremely short and concave, with irregular or weakly papillose distal margins; chiefly hidden by
centrodorsal; visible to greatest degree in excavated midradial furrow of centrodorsal. Visible portion of adjacent
radials just touching distal to short interradial processes of centrodorsal.
IBr2 series strongly arched and rounded, with lateral margins deeply recessed, straight, flat and apposed; Ibri a
shallow V-shape with few small distolateral teeth. W/L 2. 8-3. 3; Ibr2 axil hexagonal or rhombic, the diverging
lateral margins forming a continuous line with the proximolateral margins, W/L 1.7; synarthrial swelling broad,
moderately developed; no midaboral ridge or carination.
Four rays with two IIBr4(3+4) each; one ray with two IIBr2; ossicles similar to those of IBr series but more
strongly arched aborally, with moderate synarthrial swelling at IIbrri-2 (absent in the one smoothly rounded IIBr2
series). Ilbri with exterior lateral margin flattened and much longer than interior margin, W/L 2.3; exterior
distolateral corner rough or with few weak teeth or papillae; interior distolateral corner with strong spine. IIbr2
widest distally, with strong, exterior articular tubercle at IIbrr2-3. Hbn+4 concave laterally, but with recessed
interior lateral margin still flattened, W/L 1.1, diameter 4.4 mm.
One IIBr2 series bears a IIIBr2 interiorly and a IIIBr4(3+4) exteriorly.
Arms 22; ray length 235 mm (probably 245 mm; tips missing); brri-2 similar to IIbrri-2 with slightly weaker
synarthrial swellings; bri sometimes with interior distolateral spine, W/L 1.6; br2 widest distally, flattened
interiorly, W/L 1.7. Exterior articular tubercle at brr2-3 weaker than on brachitaxes; br.3+4 and following brachials
with strongly concave lateral margins, W/L 1.3, diameter 3.3 mm; brr.s-io short, oblong. Brachials becoming
strongly cuneate by bru and triangular by brri8-20, with proximal and distal margins narrowly raised and with
longer lateral margins strongly concave. Proximal brachials smooth aborally; distal midaboral margins of brachials
beyond proximal fourth to third of arm linely spinose. Entire distal margin of brachials finely spinose by mid-arm,
with midaboral portion ol margins moderately raised and bearing stronger recumbent spines (not overlapping next
brachial). Distal brachials short, cuneate, with strong midaboral spines and raised midaboral distal margins slightly
overlapping next brachial. Arm profile weakly serrate.
Syzygy following br3+4 at br24+25 to br29+301 next interval I 1-24; distal interval 5-18.
Pinnules strongly carinate, ot short segments with one or more spines at distal corners. Proximal segments
ot proximal two or three pinnules with abambulacral surface flattened to conform with close lateral apposition
of rays, and with strong, sometimes spinose carinae along both ab- and adambulacral edges of lateral mar¬
gins lacing arm tip. Arms too closely apposed to examine most oral pinnules, but one P2 of 32 segments,
18 mm long.
Description of smaller Specimen (Stn 25, EcCh 42, Fig. 7B). - Features differing from the preceding :
Centrodorsal conical with triangular interradial extensions; midradial margin not excavated; diameter 5 4 mm
maximum height 4.5 mm, midradial height 4.1 mm, Aboral pole papillose; papillae derived from obsolete cirrus
Source : MNHN, Paris
C0MATUL1D CR1NOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
667
sockets, small and slender at apex, larger smooth and convex nearer apical functional sockets. Peripheral sockets
slightly separated midradially.
Cirri XXII, 55-56, 62 mm; ci-3 short; cs squarish; C6-CI2 longest, equally long, L/W 1. 1-1.3 (more
distal cirrals slightly narrower); c 1 3 transitional, slightly longer than wide, with weakly spinose distal aboral
margin; C|4 squarish with slightly raised spinose aboral distal end. Following cirrals becoming shorter, with distal
aboral margin developing into triangular spine (never taller than its basal length). Spine with slightly convex
proximal profile and slightly concave dentate distal profile; low and rounded on distal few cirrals as cirrus tapers
to tip.
Radials exposed but shorter than Ibri, with smooth midradial swelling or low rounded tubercle; swollen
laterally against adjacent radial and triangular interradial extension of centrodorsal. IBr series strongly rounded
aborally with low, smoothly rounded synarthry; lateral margins of ossicles everted and thickened but recessed and
barely apposed. Ibr 1 with recessed distolateral corners everted and bearing one or two teeth, W/L 3.1; Ibn axil
hexagonal with lateral margins recessed, everted and weakly dentate or rugose, especially at proximolateral corner.
W/L 1.7.
All IIBr4(3+4); similar to IBr series; laterally flattened and apposed interiorly, with weak rounded synarthrial
swelling. Ilbri with exterior margin strongly thickened, everted, flat-sided, with distolateral corner rugose or
weakly spinose or dentate; IIbr2 with exterior margin and proximolateral corner similar to those of Ilbri; llbn with
exterior margin smooth; libra with recessed exterior margin flared and spinose.
Arms 19 or 20; ray length 120 mm; brri-2 similar to lIbrri-2, with few exterior spines; br3+4 2.1 mm
across, W/L 1.3; brrs-9 oblong with one or a few weak spines along their slightly longer lateral margins.
Distal aboral margins weakly raised and finely spinose beyond proximal third of arm; by mid-arm. distal margins
of brachials more strongly raised, spinose, with long recumbent spines sometimes slightly overlapping next
brachial.
Pinnules similar to those of large specimen. Proximal two or three pinnules with finely spinose or dentate
adambulacral carina along lateral margin of proximal segments (side facing arm tip).
Distribution. — A.H. Clark (1931) recorded this species from north of Halmahera (Indonesia), Taiwan, the
Nicobar Islands and Madras, India, in 55 to possibly 2047 m. McKnight (1977) recorded it (as C. rusticata ) oil
northern New Zealand. ROWE and GATES (1995) cited additional material in the Western Australia Museum (Perth)
from Dampier to Rowley Shoals, northwestern Australia, in 306-508 m. The KARUBAR specimens (336-346 m,
possibly 315-349 m), fall well within this latter depth range.
Remarks. — Both specimens fit A.H. Clark's generic diagnosis of Crotalometra and appear to represent the
same species. They have been described separately to better illustrate the differences associated with their vastly
different sizes. A.H. Clark (1950) recognized three species in the genus, and says that "The only reliable feature
for specific differentiation . . . appears to be the relative production of the distal dorsal [= aboral] edge of the outer
brachials" (p. 90). However, A.M. CLARK (1967) noted that some specimens identified by A.H. CLARK as
C. sentifera A.H. Clark bear bri+2, 3+4 on arms arising from HBr4(3+4) series instead ol just br3+4 as in C. rustica
A.H. Clark and C. magnicirra (Bell). She describes two additional specimens as C. sp„ Isentifera also bearing
hr 1+2 3+4- In addition to having the first brachial syzygy at br3+4, the two KARUBAR specimens most closely
approach C. rustica in having the distal aboral edges of the middle and distal brachials "somewhat produced and
spinous" (A.H. CLARK, 1950: 91). However, A.H. Clark (1950: 96) also refered to specimens in which this
feature is very prominent and overlapping or with an "abrupt tuft or (in ot rather long spines . approaching his
diagnosis of C. sentifera.
The large specimen is, to our knowledge, the largest thalassometrid ever recorded. Arm length exceeds the
largest previously known by at least 50 mm [180 mm in Oceanometra gigantea (A.H. Clark)). The cirri are about
as long as the maximum recorded (95 mm in Leilametra necopinata A.H. Clark, which has up to 83 unusually
elongated cirrals), but they are shorter relative to arm length than in many other thalassometrids. The much shorter
cirri of Parametra spp. are an exception, but this genus is transitional with the Charitometridae, which also bear
relatively short cirri.
668
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Genus OCEANOMETRA A.H. Clark, 1916
Oceanometra annandalei (A.H. Clark, 1909)
Figs lid, 12a, 16-18. 29-38; Tables 4a-b
Crotalometra eupedata - A.H. CLARK, 1909c: 406 (part).
Crotalometra annandalei A.H. Clark. 1909d: 642.
Thalassometfa annandalei - A.H. Clark, 1911: 551.
Tlialassomeira magna A.H. Clark, 1912b: 425.
Oceanometra annandalei - A.H. CLARK, 1916: 606; 1918: 151, 153; 1950:138, 144, pi. 15 figs 45-46.
Oceanometra magna - A.H. Clark. 1916: 606; 1918: ix, 151, 274, pi. 22 fie. 57; 1921: 171, fie. 221; 1950- 138 14^
pi. 14 figs 42-43. ’
Material EXAMINED. — Indonesia. KARUBAR, Kai Islands: stn CP 05: 1 specimen (MNHN-EcCh 52). —
Sm CP 16: 8 specimens (MNHN-EcCh 51). — Tanimbar Islands: stn CP 83: 8 specimens (MNHN-EcCh 48)-
12 specimens (MNHN-EcCh 49); 12 specimens (MNHN-EcCh 50).
Description. — Centrodorsal domelike or rounded conical (sometimes apically flattened or truncated), with
narrow, usually short interradial projections, and sometimes with narrow, rarely spinose interradial ridges;
midradial margin usually excavated; basal diameter 3.5-6.8 mm; midradial height 3. 5-6.0 mm. Ahoral pole 1.4-
3.7 mm across, domelike, rounded conical, flat or irregular; papillose, spinose, rugose or tuberculate. rarely
smooth or with irregular interradial ridges. Cirri in ten columns, two per radial area, of up to five sockets each;
adjacent columns closely crowded interradially, separated midradially by Hat to deeply concave triangular area:
apical sockets contiguous midradially.
6
- 1 - 1 - 1 -
3 4 5 6 7
Centrodorsal basal diameter (mm)
Fig. 16. — Oceanometra annandalei. Plot
of basal diameter versus midradial
height of centrodorsal. Individual data
points show centrodorsal shape
(derived from height, basal diameter
and apical diameter measurements).
hexagonal, W/L chiefly 1.3- 1.5 (range: 1
Cirri XXVII-XLIX, 61-91, maximum length 42-83 mm, long and
slender; ci to C3-5 short, increasing in length; next one or two cirrals
squarish (LW = 1.0). Following cirrals in proximal third of cirrus
longer than wide; one to three cirrals between C5 and ci6 longest
(chiefly C7 in specimens from stn CP 16; chiefly C12 in specimens
from stn CP 83); L/W of longest cirral chiefly 1.4-2. 2 (extremes:
1. 3-2.3); cirrals in distal 40% of cirrus shorter than wide. Transitional
cirral C7-C13 (c7 in specimens from stn CP 16; chiefly cn-13 in
specimens from stn CP 83). Distal aboral margin first slightly raised
between c 1 5 and C27, gradually enlarging and becoming distal
triangular spine; accompanied by increasing compression and aboral
carination of cirrals. Aboral margin of middle cirrals straight, convex
or gently sinuous, sloping up to distal spine, with distal profile of
spine straight or slightly concave. Spines triangular or rounded
triangular in cross section; distal margin of spines sometimes finely
spinose or dentate on distal cirrals; spines often weaker, rounded
(sometimes still sharp, rarely hooked) on distal few segments.
Opposing spine triangular or blunt. Terminal claw gently curved,
slightly longer than penultimate cirral.
Radials short, with midradial tubercle or 1-3 spines; swollen
laterally with 0-2 spines (or tubercle); distal margin smooth, spinose
or dentate.
Brachitaxes IBr2; II and IIIBr series chiefly 4(3+4), occasionally of
2 ossicles (one specimen with two IIBr4 bearing synarthries at Ubri _
2 and IIbr3 — 4); IIIBr interior or exterior. Ibri very short, shallowly U-
or V-shaped, W/L chiefly 3. 6-5.5 (range: 3.0-7. 5); Ibr2 axil
.0-2.2). Ubri longer exteriorly, W/L chiefly 3.0-4.8 (range: 2. 1-9.0); IIbr2
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
669
larger, convex proximally, W/L chiefly 1. 4-2.0 (range: 1.2-2. 7); IIbr3+4 W/L chiefly 0.9- 1.4 (range: 0.9-6.8).
Brachitaxes strongly arched; narrow midaboral ridge low to moderately developed, smooth, spinose or irregularly
dentate, rarely with large proximal or distal tooth (ridge reduced on Ilbr; of one specimen to single distal spine or
tooth), rarely T- or Y-shaped on libra axil; proximal and distal margins of ossicles smooth, spinose or dentate;
spines or teeth often weaker midaborally. Lateral margins of IBr and IIBr series ranging from weakly everted to
Fig 17 — Oceanometra annandalei : a. aboral view of base of one ray of specimen showing elaborate ornamentation
(EcCh 51); b-d. IIBr4(3+4) series of three specimens showing variations in ornamentation (b-c, EcCh 49; d, EcCh
51). Arrows indicate bases of Pii- Scale bars = 1.0 mm.
670
C.G. MESSING. N. AMEZIANE & M. ELEAUME
thickened and flared, sometimes flat-sided and apposed, with few to many spines, teeth or papillae; strongly flared
margins sometimes forming concave pocket along lateral portion of aboral surface of ossicles; lateral
ornamentation on HBr occasionally coalesced into irregular lamellae; midaboral ridge and ornamentation on IIIBr
often weaker and sparse, with Haring of lateral margins and ornamentation often weak or absent. Scattered aboral
spines or papillae sometimes present. In a few specimens with reduced ornamentation, Ibri with one or few spines,
teeth or papillae at distolateral corners, and Ibr2 axil and IIBr series with weakly papillose, irregular lateral margins.
Arms 13-32; ray length 84-150 mm. Midaboral ridge continued on brr ] -2; Haring and ornamentation of
brachitaxes usually persisting to brr2-4 (sometimes as far as brr7-9), but lateral margins often not Hared or everted,
and ridge and ornamentation typically weaker than on brachitaxes; midaboral ridge sometimes persisting as low
swelling beyond mid-arm. Crowded low aboral papillae or fine spines first appear on brr2-4, becoming recumbent
and elongated as narrow striations, so that brachials develop spinose distal margins by brri6-20- Brachials oblong or
slightly cuneate to bi n, strongly cuneate by brri3-14- Distal brachials short, cuneate, compressed, with raised and
more strongly spinose distal aboral margins. One arm of one specimen with brrs-8 duplicating the articulations of
brri-4 (i.e., a synarthry at brr5-6 followed by bn+8).
Pinnules becoming progressively smaller, from heavy robust Pn to small slender P5-P7. Pn up to
31 segments, 13.2 mm long. Pi up to 35 segments, 20.6 mm. P2 up to 21 segments, 7.6 mm. Short P5-P7 of
13-16 segments, 4.4-6.0 mm (P4 sometimes as short as P5-P6). Middle pinnules small and slender, becoming
longer, wider and more robust distally. Distal pinnules of 14-22 segments (chieHy 16-17). 7.0-17.5 mm long
(chiefly 7.0-10.0 mm). Proximal abambulacral surface of Pn (and to lesser degree Pi) often flattened to conform to
Hat-sided and apposed proximal brr; sometimes rugose, spinose or papillose (corresponding to lateral
ornamentation of brachitaxes). Lateral margins of basal pinnulars variably spinose; numbers of spines decreasing
from P] to P4. Lateral margin (facing arm tip) of proximal pinnulars of Pn (sometimes to P1-2) sometimes
carinate; carina smooth, spinose or dentate; following pinnules with spines only at distal corners of pinnulars.
Disk densely covered with small irregular or rounded, elongate nodules.
Distribution. — Known from the Philippines and eastern Indonesia (Kai and Tanimbar Islands, Timor). The
depth range given by A.H. Clark (1950) is 204-510 m (not including two anomalously shallow records: 55 and
124 m) with most material recorded from about 290-390 m. The Karubar specimens were collected in 297-315 m
(possibly 285-349 m).
Remarks. — A.H. Clark (1950) recognized three species of Oceanomeira : O. gigantea (A.H. Clark) from
Hawaii, O. magna (A.H. Clark) from Indonesia, and O. annandalei (A.H. Clark) from Malaysia and the
Philippines. He distinguishes O. gigantea from the other two by a midaboral ridge on Ibr2, 1 1 bro and br2, but not
on Ibri, Hbri or bri . One Karubar specimen approaches O. gigantea in having the midaboral ridge reduced to
a spine or tooth on the brachitaxes. However, because O. gigantea is currently known only from the Hawaiian
chain, we maintain it as distinct herein.
Oceanometra magna and O. annandalei present a different problem. According to A.H. Clark’s diagnoses
(1950: 138, 145), the two appear quite distinct: O. magna bears spines only along ossicle margins and midaboral
lidge, and has lateral margins of brachitaxes produced and irregularly dentate; O. annandalei bears aborally spinose
brachitaxes and arm bases, and the lateral brachitaxes margins are not produced. However, A.H. Clark's detailed
description of O. annandalei squarely contradicts his diagnosis: "the lateral edges [of Ibri] are more or less
denticulate, and there is a low, though sharp, serrate median keel"; "the lateral edges [or Ibr2 are] somewhat
denticulate; there is a sharp serrate median keel in the proximal two-thirds"; "the lateral edges [of the IIBr series] are
produced and strongly denticulate", and "The dorsal [= aboral], but not the dorsolateral, side of the fourth and
following brachials is covered with fine short spines which gradually become coarser after the proximal third of the
arm and tend to arrange themselves in longitudinal lines. The ossicles of the division series and arm bases
have strongly denticulate borders" (pp. 145-6). These are virtually identical to the description of O. magna.
A.H. Clark's description of O. annandalei never mentions "the more or less spinous dorsal surface" (p. 145) of
the brachitaxis ossicles and first two brachials found in his diagnosis. We treat the two species as synonyms.
A.M. Clark (1967) recognized another species, O. valdiviae from 1644 m off Somalia, but notes that the
"species proved very difficult to place by means of A.H. Clark’s key to the genera of Thalassomelridae"
Source : MNHN, Paris
COMATUUD CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
671
(1967: 156). It differs from 0. annandalei in bearing a more strongly excavated midradial centrodorsal margin, a
raised wedge-shaped area on the centrodorsal that separates the columns of cirrus sockets midradially, more
numerous pinnulars on the oral pinnules, and a scattering of fine rugosities all over the brachitaxes and arm
ossicles. It lacks the midaboral ridge characteristic of the brachitaxes of O. annandalei.
Oceanometra supposedly differs from Thalassometra "only in the greater number of arms; in the occurrence
of keels on one or both elements of the pairs of ossicles in the division series and first two brachials; and in
having P2 intermediate in character between Pi and P3 instead of small and similar to the following pinnules"
(A.H. CLARK, 1950: 138). The difference in arm number between the two may be ontogenetic at least in some
species. A.H. Clark also recorded midaboral carinae on the brachitaxes and proximal brachials of T. gracilis
(Carpenter) and T. hirsuta A.H. Clark (spinose in the latter), and A.M. Clark (1967) discussed several features
shared by O. valdiviae and T. attenuata A.H. Clark. It is likely that at least some species of Thalassometra and
Oceanometra will prove congeneric.
Table 4a — Oceanometra annandalei: measurements and statistics for 22 traits: centrodorsal. cirri and pinnules (n - 27).
Abbreviations: Trans. Seg., transition segment, counting from cirrus base. Other abbreviations as in Table 2a.
On Variations. — If specimen B, which bears several anomalously short proximal ray ossicles, is omitted
from consideration, coefficients of variation of all measured traits range from 9.8 (numbers of cirrals) to 30.7% (Pi
length) (Tables 4a-b). As with Aglaometra valida , the variability must be treated carefully. Figure 16 shows that,
although centrodorsal height varies less than in A. valida (see Fig. 14a), proportions based on polar and aboral
diameters and height show no clear pattern relative to centrodorsal size.
Centrodorsal surface area (S; sec Aglaometra valida above for calculations) varies from 23.5 to 53.5 mm2 with
V = 20.5%; the number of cirri per unit surface area (U) varies from 0.7 to 1.40 cirri mm-2. As with A. valida. the
wide range of the latter suggests that the number of cirri is not correlated with centrodorsal surface area. A bivariate
672
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Sp. Sia.
10
12
83
83
83
83
83
120
123
100
ISO
92
140
91
99
113
J3
A
A A
B
C
1)
83
83
16
16
16
83
83
118
141
94
140
No.
Arms
Ibr.
20
20
18
19
20
13
20
20
20
28
24
20
114' 32
W W/L
0.7
0.9
0.6
0.6
0.7
0.4
0.8
0.6
0.5
0.5
0.6
1.0
0.8
0.8
2.4
3.1
2.5
2.7
3.6
3.6
4.4
4.3
2.7
2.7
2.8
3.7
7.0
2.4
2.7
2.9
3.6
4.2
5.6
6.0
2.6 4.5
2.9
4.0
3.1
2.0
2.6
2.2
2.3
2.1
1.9
2.2
2.2
1.9
3.0 I 2.1
5.0 2.4
4.0 2.2
0.8 2.7 ' 3.5
0.5
110
122
105
136
32 0.6
24
20
26
135
84
125
22
18
20
123 19
83
135
125
122
20
16
21
0.6
3.6
2.7
3.1
7.5
4.7
5.4
0.7
0.7
0.7
0.6
2.9 4.3
3.1
3.3
2.5
4.6
5.0
4.0
0.6
0.7
0.6
0.6
2.5
2.7
3.1
2.5
0.9 2.7
4.4
4.0
5.5
4.4
3.1
Ibr.
w"
3.7
2.7
3.1
2.8
3.1
3.0
2.9
2.9
2.9
3.0
2.9
4.4
3.5
W/L
1.6
1.3
1.4
1.3
1.4
1.5
1.3
1.3
I Ibr,
W I W/L
0.7
0.9
0.6
2.0 3.0
2.6
2.2
0.6 2.5
0.7
0.5
0.7
0.6
0.6
1.5
2.1
1.5
1.8
2.6
2.6
4.0
3.0
2.1
2.3
2.1
2.2
3.1
3.3
3.2
3.2
2.4
2.2
l.l
1.5
1.5
1.5
1.5
1.9
1.7
2.7 I 1.1
2.8
3.7
2.0 3.1
1.5
2.2
2.0
3.2
3.3
1.5
1.5
0.5
0.6
1.0
0.5
0.7
0.7
0.4
0.4
0.6
0.6
0.7
0.6
0.5
0.6
0.6
0.6
1.9
0.6
1.8
2.1
2.0
2.4
2.4
2.3
2.4
2.0
3.2
2.3
2.1
3.4
3.0
3.8
Rays
I Ibr,
1.8
2.4
W W/L
2.6
1.1
4.1 1.2
2.7
4.4
3.0
1.2
1.2
1.2
4.2
4.2
1.2
4.8
4.2 | 1.1
1.1
1.3
3.4
3.1
0.0
2.5
33
1.9
2.4
2.5
1.9
2.2
2.8
3.1
2.4
0.5 i 2.2
6.5
6.2
3.3
1.3
1.9
1.1
2.2
1.1
3.6
4.4
3.6
3.8
4.8
5.5
4.2
4.6
11
1.2
1.1
3.3
1.9
2.1
1.8
2.2
2.2
1.4
1.4
1.7
1.8
1.4
llbr,.
1.9
2.5
1.5
2.1
2.1
1.8
2.6
2.2
2.3'
2.5
2.3
1.8
2.0
1.6
2.2
2.4
3.2
2.7
1.8
1.8
1.4
2.3
2.6
2.2
2.3
1.1
1.1
1.2
1.1
2.3
2.1
2.3
2.4
1.3 2.6
0.9
1.6
2.3
2 2
2.0
1.2
1.9
1.9
1.9
23)
2.0
1.9
1.8
2.1
1.9
2.7
1.4
1.9
W
1.7
2.2
1.6
2.1
2.1
1.7
2.9
1 .9 2.6
1.8 2.6
2.0
2.3
1.9
0.4
2.1
1.6
1.7
1.5
1.9
33
2.8
2.4
2.6
2.4
W/L
0.9
0.9
l.l
1.0
1.0
0.9
1.4
1.3
1.2
1.4
1.4
l.l
br.
0.7
0.0
0.5
0.5
0.6
0.6
0.6
0.6
0.6
1.5
1.3
6.8
1.3
2.3
2.1
2.0
2.7
2.4
2.3
l.l
2.3
1.5
1.6
1.1
1.1
1.2
1.3
1.0
2.4 2.1
0.8
0.5
W
1.4
1.9
2.1
1.5
2.4
1.9
1.5
1.5
0.6
0.6
0.7
2.7
2.1
W/L
2.7
2.7
3.0
3.4
3.6
2.7
3.6
3.3
2.7
2.7
3.2
2.4
5.6
3.7
2.2
2.0
0.5
0.7
0.5
0.6
0.6
0.6
0.6
0.6
0.6
0.6
2.1
3.8
3.0
4.4
2.0 3.0
1.9"
2.1
2.0
1.9
2.0
4.0
br.
1.4
W W/L
1.9
1.9
1.0
1.2
1.1
1.0
2.5
1.6
0.6
1.0
l.l
l.l
1.6
1.2
1.0
l.l
1.0
l.l
3.7
3.5
2.2
33"
"3.2"
l.l
1.0
1.0
3.3
3.5
1.0
l.l
2.1
1.7
1.9
1.7
2.7
2.1
1.8
1.9
2.0
1.9
1.9
1.6
1.8
1.7
1.0
1.8
1.3
br-
3*4
1.9
1.6
3.7
1.5
1.5
1.7
1.7
1.9
1.9
1.7
1.6
1.6
2.5
1.2
1.5
1.2
1.3
1.3
1.4
l.l
1.3
1.1
1.4
1.9
W W/L
1.7
2.2
1.5
1.5
1.4
1.5
1.4
1.7
1.7
1.9
1.4
23
l.l 1.9
1.2
1.6
1.6
1.2
1.2
l.l
l.l
l.l
1.2
1.3
1.9
1.1
1.7
1.8
1.6
1.6
1.4
2.0
2.0
0.9
0.9
1.2
1.1
1.5
1.3
1.7
1.0
1.3
1.7
1.7 1.4
1.9 1.2
0.8
1.4
1.3
1.3
1.5
1.9
Table 4b. — Oceanometra annandalei : measurements and statistics for 26 traits: rays (n = 27). Abbreviations- No Arms
arm number. Other abbreviations as in Table 2b.
Fig. 18. Oceanometra annandalei: Maximum (squares) and minimum (diamonds) widths
proximal ray ossicles (Ibr,, Ibr2, llbr,, llbr2, Hbr.3+4, br,. br2, br3+4) (n = 27).
(left) and lengths (right) of
analysis (not shown here) shows that the crowding of cirri correlates with basal centrodorsal diameter (r = 0.70,
wuh specimen B omitted), but not with aboral diameter. Simply put, number of cirri increases with centrodorsal
size. If the centrodorsal enlarges with age, the relatively narrow range of centrodorsal basal diameters (85% ran°c
rom 4.0 to 5.4 mm) suggests that the populations represented in the Karubar collection are relatively a°e
homogenous. e
Source : MNHN. Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
673
In Oceanometra annandalei, variability in the width of proximal ray ossicles remains about the same through
the brachitaxes and decreases slightly on proximal brachials of the free arm (bri-br.3+4) (Fig. 18, lei 1). Unlike
A. valida (Fig. 14b, top), the width of Ibri is not substantially more variable than that of succeeding ossicles.
The second ossicle in a brachitaxis or free arm of both species is longer than the first (e.g., Ibr2 and br2 longer
than Ibri and bri, respectively), a feature typical of most comatulids. However, in O. annandalei, the length of the
Ibr2 axil is far less variable (Fig. 18, right) than the same ossicle in 4. valida (Fig. 14b, bottom). Instead, the
broad variability found in Ibra of A. valida appears in IIbr2 and IIbr3+4 of O. annandalei. Variability in ossicle
length thus appears to be associated with the ossicle(s) immediately preceding the free arm. It seems likely that
proportions of the axil immediately proximal to the free arm (lIbr3+4 in O. annandalei) may alfect the range ol arm
movement more than the proportions of more proximal ossicles that support the viscera. Alternatively, variations
in stress on feeding arms due to different flow regimes may force variations in terminal axil proportions.
Genus STENOMETRA A.H. Clark, 1909
Remarks. — Examination of the Karubar material together with a re-examination of some previously
collected specimens forces a re-organization of Stenometra. A complete revision awaits examination ol more type
material, however. In the most current treatment of the entire genus, A.H. CLARK ( 1950) recognized lour species
based on variations in ornament of brachitaxes and proximal brachials, and on proportions ol longest cirrals. as
follows: S. diadema (A.H. Clark) and 5. quinquecostata (Carpenter) lack any eversion or swelling of, and bear few
if any spines or teeth on proximal ossicles; the longest cirrals of their slender cirri are at least twice as long as
wide. Midaboral ridges arc dentate in S. diadema and smooth in S. quinquecostata. A.H. Clark treated S. cristata
A.H. Clark as a synonym of S. quinquecostata.
In 5. dentata Gislen and 5. snelliusi A.H. Clark, the ossicles have everted and spinose or denticulate margins
and the longest cirrals have LAV < 1.5. S. dentata has strongly produced and lobale or coarsely spinose ossicles,
numerous long lateral tubercles or spines, and a "supplementary prominence on each side ot the aboral 1 idge on
some proximal brachials. S. snelliusi lacks the produced margins and brachial prominence, and has weaker spines
(A.H. CLARK, 1950: 7-8). The Karubar collection contains two clearly distinct species ol Stenometra.
Diagnostic features differ in part from those used by Clark.
Stenometra cristata A.H. Clark, 1911
Figs 1 la, 19b-c, 20a, 21, 22b, 39-41; Tables 5a-b
Stenometra cristata A.H. Clark, 1911: 553; 1912c: 210; 1918: 157.
Stenometra snelliusi A.H. Clark, 1936: 295, 313, pi. 8 fig. 9; 1950: 8. 12. pi. 1 tig. 1.
— Rowe & Gates. 1995: 165.
MATERIAL EXAMINED. — Indonesia. Karubar, Kai Islands : stn DW 18: 1 specimen WNHN-EcCh 57).
Stn DW 32: 1 small specimen (MNHN-EcCh 53). — Tanimbar Islands: stn CP 82: 10 specimens (MNHN-EcCh 55, with
branch of finely pinnate primnoid alcyonarian). - Stn CP 83: 1 specimen (no registration number). - Stn CP 86:
1 specimen (MNHN-EcCh 54); 11 specimens (MNHN-EcCh 56).
Redescription of the holotype of 5. cristata. — (USNM 27501, Malavatuan I., Verde Is. Passage.
Philippines, 13°55'55”N. 120°10’15"E, 214 m, "Albatross" stn 5275, 16.07.1908, 1 spec.) Centrodorsal with
convex sides and strong narrow interradial ridges; basal diameter about 3. 1 mm; D/H 1 . 1 . Aboral pole conical with
scattered spines and sharp papillae. Cirri in ten columns of 2-4 sockets each, separated mterrad.ally by ridge, not
separated midradially. Cirri XXIV. 68, maximum length 48 mm; slender; tapered near tip; ci-4 short; c5 longer,
with L/W 1 2- c6 longest, LAV 2.7. Following cirrals gradually decreasing in length; c2o with L/W 1.0; cirrals
strongly compressed beyond proximal short few. Oral profile moderately carinate on proximal elongate cirrals;
sharp projecting distal end of oral profile apparent from basal cirrals; distal margin ot short basal cylindrical cirrals
expanded and spinose, especially on oral side. Aboral distal margin developing sharp distally-d.rccted projection
674
C.G. MESSING, N. AMEZIANE & M. ELEAUME
by C7, increasing to long sharp distally-directed carinate spine, becoming long narrow crescentic spine on distal
cirrals; spines smaller, rounded or blunt triangular on few distal cirrals of tapered tip. Opposing spine triangular or
blunt, slightly larger than preceding spine. Terminal claw long or short.
Lateral margins of adjacent radials strongly
raised together as continuation of interradial ridge of
centrodorsal; distal corners with spine; smooth
midradially.
1 and IIBr2, well-separated laterally (except
proximal corners of Ibri in contact), strongly
rounded aborally; ossicles extending laterally
beyond articular ligaments. Ibr i shallowly V-
shaped, sometimes weakly flared laterally, often
with one or two strong distolateral spinelike teeth,
sometimes one lateral tooth; abruptly tall narrow
midaboral ridge often bidentate. Ibri axil hexagonal
with very short lateral margins; proximolateral
margins flared, lamellate and sometimes with a few
weak teeth; midaboral ridge strong, narrow, rising
to triangular proximal point like an aquiline nose,
and ending just distal to middle of axil. IIBr similar
to IBr, but ornament generally weaker. Ilbri
Fig. 19. Bases of cirri: a. Stenomelra quinquecostata
(EcCh58) showing gradual lengthening of cirrals
(Karubar, stn CP 86); b-c, Sienomeira crisicita (EcCh 56)
showing abrupt lengthening between C6 and C7 (b) and cs
and c6 (c) (Karubar. stn DW 18). Scale bar = 1.0 mm.
ODiong, united interiorly; exterior lateral margin
moderately flat-sided, with corners everted; interior
distal corner with triangular tooth.
Arms 20, ray length 85 mm; bri and following
brachials with narrow midaboral ridge, weakest on
bn, increasing to overlapping crescentic carina as
arms become compressed distally.
, f0K°,Win1 rU!CS Str°ngly Carinate' Pl '0ngeSt: foIlowin8 several pinnules successively decreasing in
^“ral 3tera marg‘n °f th'rd P'nnular abruPt|y more widely carinate than on second pinnular.
ridgesTaZrmHFkKARUBHAdSPEC1TNS' “ Ccntr°dorsal usua"y shorl- cylindrical with narrow interradial
ridges, rarely domelike; rounded conical in a small specimen without interradial ridges but with narrow interradial
projections; basal diameter 3.8-4.9 mm; midradia. height 2.1-4.0 mm; D/H 1.1-24. Aboral pole Xso“es
convex, rounded conical with two apical cirri in a small specimen
per^dtaUrla^fnw,; i617’ TT™ ‘engthu52-65 mm’ lo"g’ sle"<^ and distally tapered; in ten columns, two
mid' r . u ( y °ne ‘° ,hree) Cirri each; columns on|y rarc|y separated by narrow space
one ad'a y’ Cl 10 C5'7 short (C|-4 in smaller specimens); next cirral abruptly longer than preceding L/W 1 0-1 3-
m t |Wing C'rralS (C6'8) l0ngCSt’ L/W usual,y '-2-1.5 (extremes: ,.7-1.8); longest cirial sometimes
oTutallv bv c°,WS '7, SCneS °f 'T Ci,TalS- F°"°Wing Cirra'S gradually shor,er- becoming squarish (L/W =
2l hfrd'of cius' C Stn65; Clr^kslightly wider lba" >0"g bY or beyond mid-ciLs, very short m
with dSdends of both ^ beC°mmg sharply Carinale aboral,y and often orally
presentS weakenine^md disfl^ ^17 pr°*Ctin* dis,aIly by oral car, nation, when
sham Mv , ^nd dlSap,pearmg beyond lo"ger Pr()*lma' cirrals; sharp aboral distal margin gradually becoming
den mV y 8 ^ ^ USUaMy cresccntic on shorter dislal cirrals, sometimes with distal prS
spine 7 7"** 7' “ Up’ 0pposing Spme tr,anSuIar or blunt- ™ch larger than
Terminal claw gently curved, as long as or longer than penultimate cirral.
Had, a s short, latera margins raised or swollen against ends of interradial centrodorsal ridges usual! v with
1-2 latera, spme, tubercles or flat lamellae; abora, surface usually a, right angles to oral/abora, S of anima'l
Source : MNHN, Paris
COMATUL1D CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
675
Fig 20 — Aboral view of base of a single ray showing similar levels of ornamentation, a, Stenometra crlst°,a
(EcCh 56); b, Stenometra quinquecostata (EcCh58). Note proportionally broader, shorter IBr ossicles and midradial
knob in b. Scale bar = 1.0 mm.
F,r ?i _ Stenometra cristatci' aboral views of proximal portion of single rays showing increasing ornamentation from
F ° left to right, a. USNM 27501; b-c. Karubar. stn CP 82 (EcCh 55). Ornamentation of specimen shown in figure 20a
lies between a and c of this figure. Scale bars - 1 .0 mm.
676
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Brachitaxes strongly arched, all of two ossicles each; lateral margins projecting laterally well beyond articular
ligaments to form water pores; most ossicles from Ibra axil to br2 usually flat-sided, at least exteriorly. All !IBr2
usually present; IIIBr2, when present, interior or exterior; Ibr, united laterally, extremely short, concave distally,
W/L chiefly 4.3 (extremes: 3. 1-5.0); Ilbri short, almost oblong, united interiorly, W/L 2. 8-4.0. I-lIIbr2 axils
rhombic or hexagonal with proximal synarthrial angle strongly projecting; Ibr2 W/L 1 .3-1.9 (chiefly 1.6- 1.7);
IIbr2 W/L L4- 1.7. 1-lIIbri and bn with midaboral spine, tubercle or ridge, the latter sometimes irregular or dentate
(Ibn with up to three spines). I-IIIbr2 axils with midaboral ridge usually either smooth or dentate, often risim- to
proximal tooth, sometimes twisted distally, rarely irregularly convoluted or composed of two or three tall
coalescing knobs. Proximo- and distolateral margins of brachitaxes ossicles everted or not. usually with one to a
lew, often strong, spines or teeth; proximolateral margins of Ibr, and distolateral margins of axils more often
smooth; proximolateral margins of axils sometimes Hared and lamellate as well as dentate; distolateral corners of
- Hbri and bri (interiorly on Il-IIIbri and bri) with one to few spines. Ornamentation generally weaker on more
distal brachitaxes. One specimen with crowded aboral spines on Ibr2 axil (except midaborally).
” ■* b- s— <*«*>
lon^exteriolv Wfl 7, 9 7 t T * le°8'h "P to 95 mm' b“‘ dually incomplete; bn slightly
o ^ rr11”8 Pr“imally- W,L L4-f'7; brM W/L 1-1-1.5, diameter 1.5-
s ^s andS -nm r °^namenlall°n of brr|-2 similar to that of brachitaxes, but usually weaker; eversion,
p teeth sometimes absent; ossicles sometimes flat-sided to br3 exteriorly, br2 interiorly; br5 to about brio
Source : MNHN, Pans
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
677
short, oblong, W/L 1 .8-2.1. Following brachials shorl and cuneate, W/L 1.4-2. 1 ; distal brachials strongly
compressed and almost oblong. W/L 1. 1-1.6 (chiefly 1.4). Midaboral ridge becoming a sharp, distally curved,
carinate crest overlapping next brachial by middle of arm; sometimes offset on successive brachials or curved so
that successive crests form a sinuous carination; crests long and crescentic with finely spinose distal margin on
distal brachials; weaker on distal brachials in a few specimens.
Pinnules strongly carinate; smooth except for small spine at distal corners ot pinnulars; abambulacral carina on
proximal segments of Pi sometimes weakly dentate. Pi of up to 17 segments, 13.7 mm long (usually
13-15 segments, 8-11 mm). P2 shorter and slenderer, up to 15 segments, 10.9 mm. Following pinnules
progressively shorter to P5-6 (sometimes P7 or P« about equally short) of 7-9 segments, 4.0-6. 1 mm. Pinnules
subsequently increasingly in length; middle pinnules of 12-13 segments, 5.4-7. 5 mm. Distal pinnules ol 13-
15 segments, 6. 7-7. 6 mm long.
Disk naked.
DESCRIPTION OF A SMALL SPECIMEN (EcCh 53). — This specimen differs from the preceding as follows: cen-
trodorsal a pentagonal cylinder, diameter 2.8 mm; aboral pole flat and papillose; interradial ridges sometimes with
one or two spines near base of centrodorsal. Proximal cirrals with strongly projecting, sharp distal oral margins,
but increasing in length more gradually than preceding specimens; ci-3 short; cj with L/W 0.9; c_s with LAV 1.2,
C6 longest, with L/W 1.4. Aboral distal margin ofci sometimes strongly dentate. Brachitaxis ornamentation simi¬
lar to that of preceding specimens; e.g., midaboral ridges mostly smooth, like an aquiline nose on some axils, re¬
duced to spine on Ibri; axils with proximolateral margins flared and dentate. Arms 20, all broken at bases.
Distribution. — From Luzon, Philippines, through eastern Indonesia to Port Hedland. northwestern
Australia, in about 200 to 285 m (possibly 170-297 m), based on the type specimens of S. cristata and
S. snelliusi (A.H. Clark, 1931), additional material identified as S. snelliusi (Rowe & Gates, 1995), and
the Karubar collection.
REMARKS. — See remarks following Stenometra quinquecostatci (Carpenter), below.
T \bi E 5a — Stenometra crislata; measurements and statistics for 16 traits (max. n = 12). Abbreviations: /W one ol
the short pfnnules from P3 .0 P6; PmU, middle pinnule; Pdis„ distal pinnule. Other abbrevtat.ons as in Table 2a.
678
C.G. MESSING. N. AMEZIANE & M. ELEAUME
brachials five ZZZ SSS; oZ
Stenometra quinquecostata (Carpenter, 1888)
Figs I9a, 20b, 22a
Amedm quinquecostata Carpenter, 1888: 215, pi. 3 figs 6a-d, pi. 38 figs 1-3
I halassometra quinquecostata - A.H .Clark 1907' 360
S,en°me,ra ****»“» - A.H. Clark. 1909a: 15; 1911: 554; 1950; 8, .3, pi. 1 figs 2-3, pi. 2 figs 4. 6-7.
Material EXAMINED. - Indonesia. Karubar, Kai Islands: stn DW 18: 4 specimens (MNHN-EcCh 58)
squarish; one or ,wo ctaab from cs to =,o longest W, 7 TcLZlT'* ” ^ '° •"*"< «• « « «
length; as long as wide by ci4- stron°lv cnmnZl* u ", ,6' C following longest gradually decreasing in
at all, projecting distally on short proximal ciml ( LY°n 1^°" ^r°X'mal Clrrals; oral margin weakly carinate if
margin carinate by cl0 ZllTsuZ d^ ^ J ^ ^ ^ ™ in °"e ^-en); aboral
triangular, distally-directed spine; spines with convex proximaT ^ofile o'f 3 Wh‘Ch g™dUall.y devcloPs inl° sharp,
and crescentic. Opposing spine triangular diohii r , f L 011 ^ on very sh°rl distal cirrals; never high
curved, slightly longer than preceding cirral " Y ^ ISlinC' 3 tha" precedin8 sPines- Terminal claw gently
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
679
in one specimen), extending laterally beyond ligaments (although water pores very weak and narrow); low narrow
midaboral ridge on all brachitaxes ossicles and proximal brachials (very weak on IIBr series in two specimens). Ibri
united laterally, shallow U- or V-shaped with lateral margins sometimes weakly everted and with one or two flat
triangular teeth at distolateral corners, W/L 4.0-4.5. Ibn axil hexagonal or almost rhombic with strong proximal
synarthrial swelling; short lateral margins flat and weakly everted; midaboral ridge sometimes wider or (orked
distally, sometimes not reaching distal margin of ossicle, W/L 1.4- 1.8. Ilbri shallowly V-shaped, longer and
thickened exteriorly, with distal portion of lateral margin and lateral portion of distal margin everted aborally.
sometimes forming a sharply raised wall at distolateral corner of ossicle enclosing a concave pocket, W/L 2.7-2.9;
aboral ridges oriented diagonally, connecting ends of ridges on I and Ilbri axils. Ilbri axil hexagonal with thickened
lateral margins and weak midaboral ridge, W/L 1.4- 1 .5.
Arms 19-25; all rays broken; brri-2 similar to IIbrri-2, but weakly flat-sided and laterally everted, it at all;
aboral ridges weak; bri longer exteriorly, with a flat triangular tooth at the interior distal corner, W/L 2. 0-2. 3, br2
larger, projecting proximally. W/L 1.3- 1.5; br3+4 diameter 1.6 mm, W/L 1.2- 1.4; brr5-9 oblong, W/L 1. 8-2.0.
Brachials cuneate by bn5, W/L 1. 7-2.0; distal brachials compressed. Aboral ridge very weak on proximal brachials,
but increasing as overlapping crescentic carina on compressed distal brr.
Syzygies at br3+4 (one bn+2), br2o+2l to br3i+32; next two intervals seven to ten; distal intervals three to ten
(chiefly three to six).
Pi of 12-18 segments, 9-13 mm long, sometimes flattened proximally; sometimes with middle pinnulars
carinate; subcylindrical distally. Following pinnules shorter and becoming strongly carinate as in S. cristata.
Disk naked.
DISTRIBUTION. — Known definitely only from the Kai Islands in 212-256 m. It is not certain whether other
specimens identified as 5. quinquecostata by A.H. Clark (1950) from Sulawesi, Ternate and Timor (all eastern
Indonesia) represent this species or S. cristata.
Remarks. — We have resurrected S. cristata from synonymy with 5. quinquecostata for two reasons. Re¬
examination of the type of S. cristata (described above) indicates that it is conspecitic with one of the two species
of Stenometra collected by the KARUBAR expedition. Secondly, the other, clearly distinct, Karubar Stenometra
most closely approaches the written description of the type of S. quinquecostata (CARPENTER. 1888; A.H. Clark.
In specimens distinguished as 5. cristata , the oral margins of the cirri are more sharply carinate and ProJecl"’g-
and the cirrals following the short basal cirrals are abruptly longer. Brachitaxis ossicles are strongly arched abora y
with proximo- or distolateral margins dentate, spinose, Bared, or lamellate (or a combination), and they have
strone, often ornamented midaboral ridges. In those distinguished as 5. quinquecostata. the oral cannation on the
cirri "(except on a few proximal segments) is absent, and the proximal cirrals gradually increase in lengt .
Brachitaxis ossicles are convex aborally but not strongly arched, their lateral margins are strongly flattened and
apposed, their proximo- and distolateral margins are never ornamented as in 5. cristata, and they have low
midTheraKAR8UBAR specimens assigned to 5. cristata span a wide range of ornamentation that appears to
include 5. snelliusi as well, yet certainly represents a single taxon. We thus tentatively synonym, ze these two
species, pending comparison with the type of 5. snelliusi. Similarly, identification o the latter species as
5 quinquecostata must remain tentative pending comparison with that type material Stenometra diplax
A.H. Clark, a synonym of S. quinquecostata according to A.H. Clark (1950), is not included in the synonymy
of either Karubar species pending examination of the type material.
Genus STIREMETRA A.H. Clark, 1909a
Remarks —AH Clark (1950) distinguished Aglaometra from Stiremetra on the basis of the former s
larger size and more' broadly rounded proximal postradial ossicles. He may be correc, in distinguishing them, but
not entirely for the reasons given. Most included species of Stiremetra are known from one or few badly
680
C.G. MESSING, N, AMEZIANE & M, ELEAUME
damaged specimens, the damage usually involving complete loss of arms beyond their bases. Despite this loss
A.H. Clark (1950) placed Stiremetra (with Aglaometm) in the group of thalassometrids with arms rounded to
their tips and never carinate. This does appear to be the case in S. carnifera A.H. Clark (A M Clark 1967)
S acunr ad, a (Carpenter) (A.H. Clark, 1950), and 5. lusitanica (Carpenter) (Koehler, 1909) although
A.M. CLARK (1980) has removed the latter to Thalassometra. [NOTE: She mentioned, incorrectly, that the other
species ot Stiremetra "are said to have cirri in fifteen columns" (A.M. Clark, 1980: 192)]. A.H* Clark (1950)
describes them as having ten columns with the possible exception of 5. acutiradia which he diagnoses as having
ten columns, but notes that Carpenter’s (1888) original figure appears to have the cirri in "a single rather
crowded marginal row" (A.H, Clark, 1950: 132)]. However, in 5. breviradia (Carpenter) and 5. spinicirra
Carpenter), the distal brachials have the long, carinate, overlapping spines characteristic of A H Clark’s
firs, group of genera (Carpenter, 1888; A.H. Clark, 1950). Descriptions of 5. breviradia also note that
he distal brachials are compressed. These latter two are thus not easily identifiable using A.H. Clark’s key All
known specimens of the remaining species, 5. perple.xa (A.H. Clark) and 5. decora A.H. Clark, lack distal anus
All spee.es in the genus appear to be united by similar P,, which, with its greatly broadened proximal
portions, is much larger and stouter than the following pinnules. On this basis, they do fall into the group with
arms rounded to their tips, which A.H. Clark (1950: 6) additionally characterized as having P, "much enlarged
wh7-h T mUCHi ,°f CnCd buaSally'’- However’ <" many specimens of both Aglaometra and Oceanometra ,
which also supposedly belong to this group of genera, the pinnules gradually decrease in size and robustness from
Stiremetra breviradia (Carpenter, 1888)
Antedon breviradia , Carpenter, 1888: 110, pi. 3 figs 4a-c, 5a-c, pi. 11 fig, 5, pi. 19 figs 1-4 pi 20 figs 1-2
STxTnt ’ A H- CLARK' l909a: ,5: ,9,5: 365’ 486-487; *21. P«. 2 figs 97PJ:978. $ 44 fig. ,301;
Material EXAMINED. - Indonesia. Karubar: Kai Islands: stn CP 20: I specimen (MNHN-EcCh 47).
extenSZn,' 7 Cf "trodorsali«>nical wi,h tall, knoblike apex; margin no. concave midradially; interradial
extensions absent, basal diameter 1.9 mm; height 1.9 mm. Cirrus sockets in ten columns of two or three sockets
e“7a;“t 22Sal S0Cke,S Ta,ed SlightIy midradially by very sma11 ^angular area; apical
w- r crowded- C m XXVI1- 33, 22 mm; tapered near tip; ci-3 short; c4 squarish; c5 to at least cm longer than
directed aboraTsDine'f^ °n^est’.L/W 18; cirrals remaining about as long as wide to mid-cirrus. Sharp, distally-
n^owD tnanl r appeanng °n CB, gradually increasing in size; becoming hooked on short distal cirrals-
arrowly triangular in cross section. Some middle cirrals with distinct aboral hump proximal to spine apex- spine
sSfike with fine^ a"? ^ 7 diS‘al fCW 0n 3 few cirri’ aboral Presses low and rounded not
an I m mid 7 °" ^ °PP°Sing SPine laI1’ ^"gular and sharp on cirri with spines- low
and rounded on cm with low rounded processes. Terminal claw longer than preceding cirral. ?
distalVtevMd mSion,,hRth,r ’,hlgh ™idaboral rid*c’ highes* proximally; distal margins projecting
bn tarter W/7 mm: bf' '°n8Cr exterior|y. sometimes with weak exterior distolateral spine. W/L 1 8-
aonoS hr ’ b7-2f met,meS Wlth trace of midab°ral ndge; brr2-3 sometimes weakly fiat -sided and
FoHowfn’p h h^r!!^’ mm aCr°SS: blT5'6 °bl°ng’ W/L L6; br9 or t>r io to about br.s triangular W/L I 8
%£££$ ssres cB“bderrssed' tv? r
weaker near am'ti^Syl^^iesa^bn^a^br^^^tobH'3^3^^^!1^1^^1101^3^'^ ^ar^nafstmefimes^pinulose;
p. oyzygies at br3+4, bri3+)4 to bn4+i5; distal interval chiefly five or six (up to nine).
Source : MNHN, Pans
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
681
Pi and Pa much larger, stouter and basally broader than following pinnules; slightly flattened basally where
adjacent rays are apposed; basal segments with smoothly rounded flanges. Pi 12 segments, 5.4 mm. Following
several pinnules short, small, carinate, lacking flanges and with weak distal spines on all segments except
proximal few. P2 8 segments, 3.0 mm. P3 thinner, 9 segments, 3.0 mm. P4 similar. 8 segments, 2.6 mm.
Following pinnules more robust and increasing gradually in length. Middle pinnules of up to 12 segments,
7.6 mm. Distal pinnules of up to 14 segments, 10.0 mm. Middle and distal pinnules with all segments carinate
(except basal two) and longer than wide (except basal two and few near tips).
DISTRIBUTION. — Known previously from Fiji and the Kermadec Islands in 1 152-2285 m (A.H. Clark.
1950). The Karubar specimens extend the range to eastern Indonesia in shallower water: 769-809 m.
Remarks. — The Karubar specimen differs from Carpenter's (1888) and A.H. Clark's (1950)
descriptions of the original "Challenger" specimens only in minor respects as follows: in the latter, the aboral pole
bears papillae, there are fewer (XV-XX) cirri, and the radials are scarcely visible. However. Carpenter's figure of
a smaller specimen bearing a myzostome cyst (1888, pi. 19 fig. 2) clearly shows the radial bearing an obvious
midaboral ridge as in the Karubar specimen. The rays in the Karubar specimen are close together so that the
disk cannot be examined without damaging the specimen.
BIOGEOGRAPHY
The Karubar Expedition collected comatulids at seventeen stations over a depth range of 82 to at most
850 m. Ten stations fell between about 200 and 400 m. The collection represents a substantially smaller effort
that covered a smaller area and depth range than the previous major expeditions to the area. Yet. the collection is
surprisingly rich. The Karubar Expedition collected 22% of the 67 nominal checklist species (omitting the
unidentified families Charitometridae and Antedonidae) known from the outer shelf to the upper bathyal zone of the
East Indian archipelago ( 100 to 1000 m).
Expedition
Table 6. — Numbers of specimens of comatulids (identified to species) collected by the Karubar and previous
expeditions. Data are from A.H. Clark (1931. 1947. 1950). A plus sign and question mark (+?) indicate that
additional specimens exist but that the published references to museum holdings omit the number of specimens. Rowe
and GATES (1995) record additional specimens of Crotalometra rustica and Stenometra snelliusi from western Australia
(Western Australian Museum), but do not specify numbers of specimens.
682
C.G. MESSING. N. AMEZIANE & M. ELEAUME
No single earlier expedition collected all 15 species identified herein (Table 6). It is thus not surprising that the
Karubar material includes several geographical and bathymetric range extensions. Three species (. Palaeocomatella
hiwia, Stiremetra breviradia and Cosmiometra iole ) are recorded from Indonesian waters for the first time (Figs 23-
24). ( Stenometra crisiaia was reported previously from this area as S. snelliusi). Two of these represent substantial
range extensions. S. breviradia was previously dredged off Fiji and the Kermadec Islands (A.H. Clark, 1950).
P. hiwia was previously restricted to New Zealand waters (MCKNIGHT, 1977). By contrast, two species.
Aphanocomasler pulcher and Asterometra longicirra, remain endemic to the Kai Islands (although the latter may be
synonymous with other more widely distributed Asterometra species). Neometra xenocladia is also endemic, but is
described here as a new species (Fig. 25).
Fig. 23. — Distribution of identilied Karubar comatulids: species with broad geographic ranges.
•. Crotalometra rustics, ♦, Palaeocomatella hiwia: ■. Stiremetra breviradia: ▲, Aglaometra valida. New Zealand records
oi C. rustica and Aglaometra valida [called C. rusticata and A. incerta, respectively, by McKnight (1977)1 are
questionable.
The Karubar material extends the depth ranges of Stiremetra breviradia, Cosmiometra iole, Palaeocomatella
hiwia and P. difficiUs into shallower water and that of Aphanocomasler pulcher into considerably deeper water
(Fig. 26). For the latter, a considerable gap exists between the original record ("Down to 52 Metres."
A.H. Clark. 1918: 275) and the Karubar material (confirmed 299-390 m; possible 296-502 m). The
possibility exists that morphological differences between the two collections might be depth-related, as su^ested
lor stalked crinoids (Bourseau & Roux, 1989; Ameziane-Cominardi, 1991). However, one must treat older
bathymetric data with caution. Earlier expeditions generally recorded the depth at each station via a single lead-line
sounding. The technique took no account of local variations in sea-floor topography, which are often substantial
along insular margins where much of this sampling took place.
Source : MNHN, Parts
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
683
Fig. 24. — Distribution of identified Karubar comatulids: Thalassometridae with known ranges restricted to the East
Indian archipelago. • , Cosmiometra philippinensis\ ■. Cosmiomelra iole\ ▼ , Oceanometra annandalei ;
★ . Stenometra quinquecostata ; ♦. Stenometra cristaia.
Fig. 25. — Distribution of identified Karubar comatulids: Comasteridae, Asterometridae and Calometridae with known
ranges restricted to the East Indian archipelago. • , Aphanocomasler pulclier, A , Palaeocomaiella difficilis : ♦.
Neometra cf. alecio: ▼, Neometra xenocladia ; ■, Asterometra longicirra.
Source
684
C.G. MESSING. N. AMEZIANE & M. ELEAUME
f/Zf/f
■ <?• <?■ ^ ^ v*- 0- o- 0- 0- <3- c,. <?•
K <?• <?■ ^
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
685
Eight of the 15 identified speeies arc thalassomctrids. Four of these, Aglaometra valida, Oceanometra
annandalei, Cosmiometra philippinensis and Stenometra cristata , together account for 75% of identified specimens.
This dominance reflects an important aspect of comatulid biogeography. The East Indian archipelago supports the
richest comatulid fauna in the world. About one hundred nominal species live in shallow-water (<50 m) (MESSING.
1994, 1998a). The checklist above cites a similar number (108 species including charitometrids and antedonids) in
100-1000 m. Together, these faunas account for almost 40% of known comatulid species. Although most families
found in the region have representatives in both shallow and deeper water, the two depth ranges support
substantially different assemblages. Comasterids, himerometrids, colobometrids and mariametrids dominate the
shallow-water fauna while thalassomelrids, charitometrids and antedonids account for over fifty percent of the
deeper species (Fig. 27). Similarly, the same three latter families account for the great majority of Karubar
specimens. Of the 108 deeper-water species, only eleven occur in <50 m. Maximum richness occurs in 100-299 m
and gradually diminishes with increasing depth (Fig. 28), although this pattern may reflect collecting effort, at
least in part.
E.
w
CO
>
Q>
c
■C
4-*
a
Q1
a
0-49
50-99
100-199
200-299
300-399
400-499
500-599
600-749
750-999
1000-1199
1250-1499
1500-1999
2000+
Fig. 27. — Bathymetric distribution of comatulid species by family occurring between about 100 and 1000 m: numbers of
species plotted by depth interval (meters). The number of species in the richest depth interval is given for each
family.
The Karubar material reminds us how little we still know about tropical Indo-Western Pacific crinoid
distributions. The collection adds five species to the deeper-water comatulid fauna of the Kai Islands, bringing the
total to 37, that is, 55% of the East Indian deeper-water fauna (again, exclusive of the antedonids and
charitometrids; see the checklist). It remains unclear, however, how much additional collecting will add to the
fauna. Several species known from nearby island groups are certainly strong candidates [e.g., Thalassometra hirsuta
A.H. Clark and Cosmiometra woodmasoni (Bell) from off Timor]. Also, a tendency supposedly exists for taxa
with broad bathymetric distributions (e.g., many thalassometrids) to have wider geographic ranges (A.H. Clark.
1914; GlSLEN, 1951). However, MESSING (1998a: 187) reported that, for the belter known shallow-water
comatulid fauna, "local areas within the East Indian archipelago appear to support no more than about half the total
number recorded". Certainly such faunal variations derive at least in part from insufficient collecting, especially for
deep-water habitats. Yet, the various isolated basins within the archipelago (e.g., Celebes, Sulu. Savu and Aru)
may possibly support substantial endemism.
686
C.G. MESSING, N. AMEZIANE & M. ELEAUME
E,
w
re
£
a)
c
.c
**
Q.
0)
O
0 5 10 15 20 25 30 35 40 45
Number of Species Recorded in Each Depth Interval
Fig. 28. — Bathymetric distribution of comatulid species occurring between about 100 and 1000 m: numbers of species
plotted by depth interval (meters).
1 axonomic problems, particularly at the species level, may also cripple faunal and biogeographic assessments.
1 he Karubar material has permitted us to analyze morphological variability in greater detail than previously and,
as a result, more accurately characterize some species and their diagnostic features. High coefficients of variation
recorded lor many tealures indicate substantial sample heterogeneity. Yet, enough intergradations permit us to
synonymize several species formerly considered distinct (e.g., Aglaometra sulcata, A. incerta and /l. propinqua
under A. valida).
Because so much of comatulid structure is suspension-feeding apparatus, it follows, as with clonal organisms
(e.g., Winston, 1979; Boero, 1984; Brazeau & Lasker, 1988), that patterns of water movement might
substantially affect morphology and account for at least some of the observed variability. Among comatulids,
individuals of several nominal comasterid species (e.g., Comanthus parvicirrus , C. wahlbergii , C. suavia) in
shallow (<4 m) habitats subject to strong surge/wave action tend to have shorter more numerous arms than those
in quieter (deeper or sheltered) water (MESSING, 1994). Among stalked crinoids, Ameziane-Cominardi (1991:
45) noted that: "ll existe une relation entre la position geographique qui s’accompagne de conditions
hydrodynamiques diverses et les variations morphologiques observees au sein de ces trois taxons [Metacrimts
serratus, M. costatus et M. leva]. L'hydrodynamisme va etre un facteur limitant majeur pour les Metacrinus. Dans
un environnement calme et stable tel qu’au Sud-Ouest des Phillipines va se developper une faune aux caracteres
morphologiques constants. Alors qu’en Nouvelle-Caledonie, les specimens qui vivent dans des courants plus forts,
presentent une plus forte variability des caracteres morphologiques et un nombre de bras moindre."
However, the Karubar collection notwithstanding, we yet lack virtually all of data needed to determine what
constellation of genetic, ontogenetic, biomechanical and energetic constraints, and environmental factors generate
the variability observed within and between comatulid skeletal components.
SUMMARY OF PROPOSED TAXONOMIC CHANGES
New taxon:
Neometra xenocladia
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
687
NEW SYNONYMS:
Antedon incerta Carpenter, 1888, a synonym of Aglaometra valida Carpenter, 1888.
Crotalometra propinqua A.H. Clark, 19 1 1 . a synonym of Aglaometra valida Carpenter, 1 888.
Crotalometra sulcata A.H. Clark, 1912b, a synonym of Aglaometra valida Carpenter, 1888.
Thalassometra magrta A.H. Clark, 1912b, a synonym of Oceanometra annandalei (A.H. Clark. 1909d).
Stenometra snelliusi A.H. Clark, 1936, a synonym of Stenometra cristata A.H. Clark. 1911.
Names with changed status:
Palaeocomatella A.H. Clark, 1912a, removed from the synonymy of Comissia A.H. Clark, I909e
(Hoggett & Rowe, 1986).
Stenometra cristata A.H. Clark, 1911, removed from the synonymy of Stenometra quinquecostata Carpenter
1888 (A.H. Clark, 1950).
AC KNO WLEDGMENTS
We would like to thank the members of the Karubar Expedition, and espeeially the staff of the Museum
national d'Histoire naturelle, Paris, and of IRD (ex ORSTOM), for their care in the collection and preservation
process, which allowed us to examine material in such excellent condition. We would also like to acknowledge
Alain CROSNIER's infinite patience and support during the course of this work. The first author is also grateful to
Professor Dominique DOUMENC (Director, Laboratoire de Biologie des Invertebres Marins et Malacologie) for
inviting him to work at the Museum.
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688
C.G. MESSING, N. AMEZIANE & M. ELEAUME
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the vicinity of the Philippine Islands. Proceedings of the United States National Museum. 39 (1798): 529-563.
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Proceedings of the Biological Society of Washington, 25: 17-28.
Clark, A.H.. 1912b. — Descriptions of eleven new crinoids belonging to the families Calometridae and Thalasso-
metridae discovered by the "Siboga"in the Dutch East Indies. Zoologisclten Anzeiger, 39 (1 1/12): 420-428.
Clark. A.H., 1912c. — The Crinoids of the Indian Ocean, iii + 325 pp. Indian Museum. Calcutta.
Clark, A.H., 1913. — Notes on the Recent Crinoids in the British Museum. Smithsonian Miscellaneous Collections.
61 (15): 1-89.
Clark, A.H., 1914. — The correlation between the bathymetrical and the geographical range in the Recent crinoids.
Journal of the Washington Academy of Sciences, 4(19): 558-564.
Clark, A.H., 1915. — A monograph of the existing crinoids. Vol. 1, pt 1. Bulletin of the United States National
Museum, (82): vi + 406 pp., 17 pis.
Clark, A.H., 1916. — Six new genera of unstalked crinoids belonging to the families Thalassometridae and
Charitometridae. Journal of the Washington Academy of Sciences, 6 (17): 605-608.
Clark, A.H., 1918. — The unstalked crinoids of the Siboga Expedition. Siboga-Expeditie, (42b): x + 300, 28 pis.
Clark, A.H., 1921. — A monograph of the existing crinoids. Vol. 1. pt 2. Bulletin of the United States National
Museum, (82): xxvi + 795, 57 pis.
Clark, A.H., 1931. — A monograph or the existing crinoids. Vol. 1, pt 3. Bulletin of the United States National
Museum, (82): viii + 816, 82 pis.
Clark, A.H., 1936. — Biological results of the Snellius Expedition. 1. The Unstalked Crinoids of the Snellius
Expedition. Temminckia, 1: 295-320, pis 7-9.
CLARK, A.H.. 1941. — A monograph of the existing crinoids. Vol. I, pt 4a. Bulletin of the United States National
Museum, (82): 1-603, 61 pis.
Clark, A.H., 1947. — A monograph of the existing crinoids. Vol. 1, pt 4b. Bulletin of the United States National
Museum, (82): 1-473, 43 pis.
CLARK. A.H.. 1950. — A monograph of the existing crinoids. Vol. 1, pt 4c. Bulletin of the United States National
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Clark. A.H. & Clark, A.M., 1967. — A monograph of the existing crinoids. Vol. 1, pt 5. Bulletin of the United States
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Clark, A.M., 1967. — Crinoids of the zoological collection of the Bavarian Stale. Veroffentlichungen der Zoologischen
Staatssammlung Miinchen, 11: 151-174.
CLA24K'(2) M75 _ S°me Crinoids from lhe Indian 0cean- Bulletin of the British Museum ( Natural Histoty) Zoology,
Clark, A.M.. 1980. — Crinoidea collected by the Meteor and Discovery in the NE Atlantic. Bulletin of the British
Museum of Natural Histoty (Zoology), 38 (4): 187-210.
Source : MNHN, Paris
COMATULID CR1NOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
689
GISLEN, T., 1919. — Results of Dr. E. Mjoberg’s Swedish Scientific Expeditions to Australia. 1910-1913. XXIII.
Crinoids. Kungliga Svenska Vetenskapsakademiens Handlingar, 59 (4): 3-37.
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Swedish Deep-Sea Expedition. 2 (Zoology) (4): 51-59, I pi.
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Crinoidea), with the description of a new subfamily and a new genus. Zoological Journal of the Lmnean Society. 88.
103-142.
Koehler, R.. 1909. — Echinodermes provenant des campagnes du yacht Princesse-Alice. Resultats des campagnes
scientifiques accomplies stir son yacht par Albert Jer Prince souverain de Monaco. 34: 1-317, 32 pis.
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Forhandlingar, 1866 (9): 223-233.
MCKNIGHT, D.G., 1977. — Additions to the New Zealand crinoid fauna. New Zealand Oceanographic Institute (NZOI)
Record 's. 2 ( 1 i ): 93-112.
Messing C G 1994. — Comaiulid crinoids (Echinodermata) of Madang, Papua New Guinea, and environs. Diversity and
ecology. In: B. David, A. Guille, J.-P. Feral & M. Roux (eds), Echinoderms through Time: 237-243. Balkema.
Rotterdam.
Messing, C.G.. 1995a. — AUoeocomatella, a new genus of reef-dwelling feather star from the tropical Indo-West Pacilic
(Echinodermata: Crinoidea: Comasteridae). Proceedings of the Biological Society of Washington. 108 (3): 436-44U.
Messing CG , 1995b. — Redescription of a unique feather star (Echinodermata: Crinoidea: Comatulida: Comasteridae)
with the diagnosis of a new genus. Proceedings of the Biological Society of Washington. 108 (4): 656-661.
MESSING. C.G., 1997. - Living Comatulids. In: J. Waters, & C. Maples (eds), Geobiology of Echinoderms.
Paleontological Society Papers, 3: 3-30.
Messing C.G., 1998a. — An initial re-assessment of the distribution and diversity of the East Indian shallow-water
crinoid fauna. In: R. Mooi & M. Telford (eds), Echinoderms: San Francisco: 187-192. Balkema, Rotterdam.
MESSING C G 1998b. — A revision of the Recent Indo-West Pacific comatulid genus Comaster Agassiz. Pari 1 : The type
species of Comaster and Phanogenia Loven (Echinodermata: Crinoidea: Comasteridae). Invertebrate Taxonomy. 12:
191-209.
Rowe. F.W.E. & Gates. J., 1995. — Echinodermata. In: A. Wells (ed.), Zoological Catalogue of Australia. 33: xm +
510. CSIRO Australia, Melbourne.
Sokal . R.R.& Rohlf, F.J., 1995. — Biometry (3rd ed.). 887 pp. W. H. Freeman. San Francisco.
Winston, J.E., 1979. — Current-related morphology and behaviour in some Pacific coast bryozoans. In: Advances in
Bryozoology : 247-268. Academic Press, London.
YULIN, L„ 1984. — On the family Asterometridae (Crinoidea) of China, with a description of Sinometra acuticirra gen. et
sp.'nov. Chinese Journal of Oceanology and Limnology, 2(1): 109-1 16. pi. 1.
690
C.G. MESSING, N. AMEZIANE & M. ELEAUME
Fig. 29. Oceanometra annandalei (EcCh49) (SEM): a-c, Centrodorsals, a-b, lateral views of two different specimens;
c, oral surface of a; d-f, lateral views of adjacent pairs of cirrus socket columns separated by narrow interradial ridges
(three different specimens). Scale bars: a-b = 1.0 mm; c-f = 0.5 mm.
Source : MNHN, Paris
C0MATUL1D CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
691
FIG. 30. - Oceanometra annandalei (EcCh49) (SEM): a. Oblique view of centrodorsal showing concave oral surface,
b, oral surface of centrodorsal; c, radial circlet, oral view; d, detail of b showing central cavity, e. detail of showing
basal rosette; f, radial articular facet. Scale bars: a-c = 1.0 mm; d-f = 0.5 mm.
Source : MNHN. Pans
692
C.G. MESSING. N. AMEZ1ANE & M. ELEAUME
Fig. 31. — Oceanometra annandalei (a-e, EcCh 49; f-h. EcCh 51) (SEM): a. Ibri, muscular articular facet; b. Ibr? axil,
synarthry; c, distal view of Ibrj showing paired muscular articular facets and triangular aboral surface; d. llbri.
muscular articular facet; e. br& . syzygy; f. br4, syzygy; g. IIIbr3. note isolated patches of raised stereom between
central raised periaxial region (around lumen) and radiating culminae; h. Illbr?. showing periaxial stereom. Scale bars:
a-c = 1 .0 mm; d-f = 0.5 mm; g-h = 100 pm.
Source : MNHN, Paris
693
COMATULID CRIN'OIDS OF THE KARUBAR EXPEDITION TO INDONESIA
F.G 32. — Oceanometra annandalei (EcCh 51) (SEM): a. bn, syzygy with pinnular articulation at upper left: ^detail ol
a showing raised periaxial region around lumen at left; c. detail of a showing culm.na (ndge) d. det^l of culmma n c.
(> derail of crenella (flat area between culminae) in b; f, detail ol periaxial region in a. g. c , |
aniculation in a h detail of g. showing deep probable muscular fossa -eom; ,. detail o, g_. stereo m between
pinnular articulation and syzygial facet. Scale bars; a = 0.5 mm; b - 100 pm. c, g, i 50 pm, . P
Source :
694
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Fig. 33. — Oceanometra annandalei (EcCh 51) (SEM). br2. synarthry: a. entire ossicle; b. detail showing central lumen
and fulcral ridge; c. detail showing ligamentary fossa; d. detail of c. ligamentary stereom. Scale bars: a = 0.5 mm;
b-c = 100 |jm; d = 10 gm.
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
695
FIG 34. - Oceanometra annandalei (EcCh 51) (SEM). bn, muscular articulation: a. entire ossicle; b, detail of a. lulcral
ridge (left) and elongated lateral fossa that accommodates the ligament that unites adjacent bri.c. detail of a, ridge
between paired muscular fossae; d. detail of b. stereom in lateral ligamentary fossa; e, detail of c, muscular [ossa
stereom. Scale bars: a = 0.25 mm; b-c = 100 pm; d-e = 10 pm.
Source .
696
C.G. MESSING, N. AMEZIANE & M. ELEAUME
FlG. 35. — Oceanometra annandalei (EcCh 51) (SEM). br?, muscular articulation: a, entire ossicle; b, detail of a, lumen,
fulcral ridge and aboral ligament fossa; c, detail of b, gradation in stereom from ligamentary fossa (fine) to fulcra!
ridge (coarse); d, detail of a, fulcral ridge, ligament pit and aboral ligamentary fossa; e. detail of d, ligament pit
stereom; f, detail of a, left muscular fossa; g, detail of f, muscular fossa stereom. Scale bars: a = 0.5 mm; b. d, f =
100 pm; c = 50 pm; e, g = 10 pm.
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
697
Fig. 36. — Oceanometra aniuindalei (EcCh 51) (SEM). brsi. muscular articulation: a, entire ossicle, oblique view showing
pinnule articulation; b, detail of a. lumen and left interarticular ligament fossa; c. detail of a. pinnule articulation;
d, detail of b, interarticular ligament fossa stereom; e, detail of b, periaxial stereom; f. detail of c. periaxial stereom
of pinnule articulation fulcrai ridge; g. detail of c, pinnule articulation, stereom of fossa closest to brachial muscular
fossa. Scale bars: a = 0.25 mm; b-c = 100 pm; d-g = 10 pm.
698
C.G. MESSING. N. AMEZIANE & M. ELEAUME
Fig. 37. — Oceanomelra annandalei (EcCh 51) (SEM). P|, articular facet of basal pinnular: a, entire ossicle; b. detail of a,
lumen (center), fulcral ridge and ligament pit (upper left); c. detail of b showing fulcral ridge and ligament pit; d, detail
of a. ligamentary fossa stereom (upper right of a); e, detail of c, ligamentary stereom adjacent to ligament pit. Scale
bars: a = 0.25 mm; b = 100 pm; c = 50 pm; d-e = 10 pm.
Source : MNHN, Paris
COMATULID CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
699
F,g 38 — Oceanomeira annandalei (EcCh 51) (SEM). a-f. P2 articular facets: a, second pinnular; b, fourth pinnular;
c. eighth pinnular; d. tenth pinnular; e. 24th pinnular; f. 28th pinnular. -g-i. distal pinnule articular facets:
g. first pinnular; h, fourth pinnular; i. eighth pinnular. Scale bars: 100 pm.
Source : MNHN, Pans
700
C.G. MESSING, N. AMEZIANE & M. ELEAUME
Fig. 39. — Stenometra cristata (EcCIi 55) (SEM). Ibr2, synarthry: a, entire ossicle: b. upper right detail of a, fulcra! ridge
and ligament fossa; c. detail of b, ligament fossa stereom; d. detail of b, fulcral ridge stcrcom. Scale bars: a = 0.5 mm;
b = 100 pm; c-d = 10 pm.
Source : MNHN. Paris
C0MATUL1D CRINOIDS OF THE KARUBAR EXPEDITION TO INDONESIA
701
Fig. 40. — Stenometra cristata (EcCh 55) (SEM). Muscular articulation of brachials from a detached arm fragment:
a. middle brachial, entire ossicle; b. more proximal brachial, entire ossicle (oblique view); c. detail ol b, lei t
muscular fossa; d. detail of b. left interarticular ligament fossa; e, detail of c, muscular tossa stereom; f, detail of d.
interarticular ligament fossa stereom. Scale bars: a-b = 0.25 mm; c-d = 100 pm; e-f = 10 pm.
Source :
702
C.G. MESSING, N. AMEZ1ANE & M. ELEAUME
Fig. 41. — Stenometra cristaia (EcCh 55) (SEM). Proximal brachial (detached arm fragment), syzygy: a. entire ossicle;
b. detail of a, culmina (center) flanked by crenellae; c, detail of a, periaxial stereom; d. detail of a, crenellar stereom;
e, detail of b. culmina stereom. Scale bars: a = 0.25 mm; b = 50 pm; c-e = 10 pm.
Source : MNHN. Paris
TATS DES CAMPAGNES MUSORSTOM, VOLUME 21
RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 21 — RESULTATS DES
Ascidiacea: Plurellidae collected in the Pacific Ocean
by the cruises MUSORSTOM, KARUBAR
and the "Coral Reef Research Foundation"
Frangoise MONNIOT & Claude MONNIOT
Museum National d'Histoire Naturelle.
Laboratoire de Biologie des Invertebres marins et Malacologie
ESA 8044, CNRS
55 rue Buffon, 75231 Paris Cedex 05.
ABSTRACT
Collectings by deep SCUBA dives have brought up 4 new species of Plurellidae, a family known by 2 Plurella species
and one Microgastra. A new genus Plurascidia is also described, intermediate between the Plurellidae and the Ascidiidae
families. All these new species are compared to the 3 Plurellidae already known and to a strange deep Ascidudae. hmbrora
calsubia.
RESUME
Ascidiacea : Plurellidae recoltees dans l'ocean Pacifique lors des campagnes MUSORSTOM et
K/VRUBAR ainsi que par la Coral Reef Research Foundation.
Des plongees profondes en scaphandre autonome ont permis la recoltc de quatre especes nouvelles de Plurellidae. une
famille qui ne comprenait jusqu'k maintenant que deux especes de Plurella et une espece de Microgastra. Un genre nouveau
Plurascidia, intermediate entre les families Plurellidae el Ascidiidae est ddcnt. Les especes nouvelles sont compares aux
trois especes dej& connues de la famille ainsi qu’& une etrange Ascidiidae de grande profondeur, Ftmbrora calsubia.
INTRODUCTION
Several uncommon ascidians were collected in the tropical western and central Pacific ocean. The Plurella
species were found by diving down to 100 m deep by people from the American Coral Reel Research Foundation,
during a large biodiversity program at Palau and Philippines islands. The French MUSORSTOM cruises have
MONNIOT. F. & MONNIOT, C., 2000. — Ascidiacea: Plurellidae collected in the Pacific Ocean by the cruises
MUSORSTOM, KARUBAR and the "Coral Reef Research Foundation". In : A. CROSNIER (ed. h Resullats des Campagnes
MUSORSTOM. Volume 21. Memoires du Museum national d'Histoire naturelle. 184 : 703-721. Paris ISBN --8565. _6 7.
704
F. MONNIOT & C. MONNIOT
collected by dredging: Microgastra granosa Kott, 1973 and Fimbrora calsubia Monniot & Monniot, 1991 (oft
Vanuatu and New Caledonia). Plurascidia marquesana sp. nov. (around the Marquesas Islands). These collections
were made with the objective of an inventory of tropical Pacific invertebrates from the shore to the bathyal area.
Numerous unknown animals were collected and 4 new species of ascidians are described here.
The materiel studied here is deposited at the Museum national d'Histoire naturelle at Paris.
Family PLURELLIDAE Kott, 1973
The Plurellidae Kott, 1973 is a family in which the branchial structure is a flat tissue pierced by numerous
elongated stigmata, with longitudinal vessels and erect papillae at the crossings with the transverse vessels. Their
neural ganglion associated to the neural gland is particularly far from the prepharyngeal groove, a structure only
observed in the Phlebobranchia. Nevertheless, some original characters have to be noted. The gonads are not linked
to the gut but lie into the body wall as in Stolidobranchia. There is no dorsal tubercle but small ciliated pits at the
end of multiple short branches of the neural duct, issued from the neural gland. This last structure is characteristic
of the genus Phallusia, but the latter has a dorsal tubercle well developed.
Genus PLURELLA Kott. 1973
The Plurellidae are particularly difficult to observe as the tunic is totally impregnated with coarse sand and
some organs are included in this sandy tunic (Fig. 1). Even in samples of Plurella carefully collected by SCUBA,
the arrangement of the zooids remains unclear. The zooids cannot be separated from each other without damage, and
the sand has to be removed grain by grain to observe the heart, gonads and neural ganglion. We have tentatively
tried to follow the course of the blood vessels through the tunic by stains, to demonstrate a link between the
zooids, but without success. There is a complex network of blood sinuses, especially on the left side of the body
but no crossing from one zooid to another could be proved.
We tentatively tried to find buds but without success. The zooids are so fused by their tunics that a colonial
structure remains possible in the oviparous genus Plurella but we presently prefer to speak of agglomerated zooids
instead of colonies.
The species of the genus Plurella have all the same external shape with coalescent zooids enclosed in a tunic
embedded with coarse sediment and diverse debris. Some epibionts are settled at the colony surface or into the
spaces left open between groups of zooids, actinians for example. The siphons’ colour only varies from pale
yellow to orange (Fig. 1). Their rim is not impregnated with sand. The internal morphology needs a careful
examination to distinguish the species. A single anatomical character cannot be sufficient to define a taxon, but the
combination of several elements has to be taken into account, the number of studied samples being too small to
estimate an intraspecific variability.
The length of the siphons varies a little according to the place of the zooid, or between two groups collected in
a same station. On the other hand, the presence of one or several ovaries, the place of the tes'tis with regard to the
heart, the shape of the gut, the position of the cloacal siphon or the location of the oral tentacles can constitute
specific distinctive elements.
All species only have muscles on the right body side arranged in transverse bundles.
In a first group having only one ovary we find Plurella testacea sp. nov. collected at the Philippines and
P. monogyna sp. nov. collected at the Philippines and Palau islands for this collection. These species differ by the
place of the tentacle circle, the shape of the gut and the location of the testis related to the heart. The structure of
the tunic on the right body side is also different.
In the second group having several ovaries are P. koitae Monniot & Monniot, 1996 and P. melanesia sp. nov.
in addition to P. elongata Kott, 1973.
Source : MNHN, Paris
ASCIDIACEA: PACIFIC PLURELLIDAE
705
FlG _ a. Plurella monogyna sp. nov.; B. Plurella testacea sp. nov.; C. Plurella koixae Monnio. F. & Monnio. C
1996; D. Plurella melanesiae sp. nov.
706
F. MONNIOT & C. MONNIOT
Plurella monogyna sp. nov.
Figs I A, 2A, 3
Material EXAMINED. — Coral Reef Research Foundation coll.
Philippines. Cebu : Cabilao, 20 m (MNHN P7 Plu 17).
Bohol Sea : SW Panglao Island. 09°31.26'N, 123°41.05'E. 10 m, 16.04.1997.
Palau. Helen 's Reef: 02°52.27'N, 131°44.41'E, 18 m. 12.12.1996.
Type. — MNHN P7 Plu 17: Philippines, Cebu, Cabilao, 20 m deep.
Description. — The agglomerated individuals make thick cushions, entirely embedded with sand (Fig. 1 A).
The largest mass is 12 cm in diameter and 5 cm thick. In life the animals are half buried into the sediment. The
zooids are linked to each other by their tunic, more strongly by the left body side and the base of the cloacal
siphon. Some expansions of tunic, densely embedded with sand, arise from the base of the zooids as anchoring
parts. Spaces exist between the groups of zooids, but we could not determine their shape or extension which arc
irregular.
The oral and cloacal siphons of a same diameter are free along about I cm, erect at the colony surface. Their
naked extremity is orange in colour and dentate (Fig. I A). Each siphon has a distal muscular sphincter crossed by
very short longitudinal fibres. The cloacal siphon is shorter than the oral one but of a same diameter. It is inserted
at one third of the body length (Figs 2A, 3A-B). It has an internal velum.
The test strongly adheres to the body wall except above the right side of the body which contains the muscles.
At this level the tunic has 2 layers, one external embedded with sand and one internal thin but stiffer, which is not
linked to the external layer and no more to the body wall. Not any link other than the tunic relies the zooids. Not
any blood sinus going from one individual to the other has been found. At the level of the heart and gonads, and
around the base of the cloacal siphon the tunic and the body wall are thicker and cannot be dissociated from each
other. They are rich in blood sinuses. Dark pigment (in formalin) is present in the siphons and in the body wall
vascular system were the design is a net. There are also pigment cells in the branchial sac.
The muscles are only present on the right side on the body (Figs 2A. 3B). They are strong transverse parallel
fibres which do not reach the endostyle nor the dorsal line. The body wall is very thin on the left side, impossible
to separate from the tunic without damage. The internal side of the oral siphon is covered with numerous minute
papillae, especially developed in the space between the tentacles and the branchial sac.
The tentacles are numerous (Figs 2A, 3B), in a circle (more than 50 counted in one zooid), in 4 orders of size,
at the base ot the siphon. The prepharyngcal band is made of 2 blades, the most posterior the highest. It is pressed
against the branchial sac. Dorsally it draws a deep oblique V to join the dorsal lamina, prolonged by 2 rims on
each side of a groove. The dorsal lamina is the prolongation of the left blade, with a plain edge and ribs
corresponding to the left transverse vessels.
The neural ganglion is slightly displaced on the left side at the base of the cloacal siphon (Fig. 3B). included in
the body wall thickness. A neural gland not well defined is joined to it. A thin channel extends anteriorly inside the
body wall up to the anterior part of the branchial fold. Multiple short ramifications ended in urns open into the
cloacal cavity. The most anterior one is placed inside the dorsal prepharyngeal V.
No dorsal tubercle, as that of other Phlebobranchia ascidians, could be detected in the V delimited by the
prepharyngeal groove. The stigmata of the first row are very close to the prepharyngeal band. There are very
numerous stigmata rows. The longitudinal vessels (85 on each side in one individual) begin anteriorly to the first
row of stigmata by a papilla and extend along the whole body length. Some additional longitudinal vessels
progressively appear along the dorsal lamina at the rear part of the branchial sac. We counted 1 or 2 stigmata per
mesh. The branchial papillae are rounded above each crossing with the transverse vessels.
The branchial sac forms a high fold beginning at the indentation of the prepharyngeal band and lies on the
whole body length. The endostyle stops at the bottom of the branchial sac, down to the fold. The fold is attached
along its whole length by strips similar to those uniting the branchial sac to the body wall.
The gut forms a closed loop (Figs 2A, 3B). It is almost free in the peribranchial cavity only linked to the
branchial sac and the body wall by tew blood vessels. In the most part of the zooids, the different parts of the gut
Source : MNHN, Pans
ASCIDIACEA: PACIFIC PLURELLIDAE
707
Fig. 2. — Dissections stained by hemalum.
A. Plurella monogyna sp. nov. : external side of a specimen opened ventrally to see the siphons, tentacles, upper part o
the branchial sac, muscles, gut, gonads and heart. . . V«ntniiv tr> see ihe eut and
B, Plurella kouae Monnio. F. & Monniot C., 1996: external s.de ol a spec.men opened ventrally to see gut
C Plurella testacea sp. nov. : on the right: part of the sclerified tunic, on the left specimen with a doubled oral siphon
D. Plurella melanesiae sp. nov,: external side of a zooid opened ventrally showmg several testes close to the
3 ovaries lying at the side of the anus.
Source : MNHN. Paris
708
F. MONNIOT & C MONNIOT
arc not delimited when they are full, but sometimes it is possible to see a narrow oesophagus with a length equal
to its diameter, followed by a spherical stomach and a tubular intestine entirely folded on itself and finally curved
again, against the stomach, in a short rectum. Rarely the thin wall of the stomach is crumpled when emptied
(Fig. 3A). The anus is widely open with a thickened rim. The gut is located in the middle of the body length. The
heart forms a long arc in the posterior part of the body, crossing the whole left side from the endostyle to the base
of the cloacal siphon close to the ovary (Figs 2A. 3A-B). There is dorsally a spherical hernia containing an
inclusion (Figs 2A, 3B). The heart is partly included in the tunic and difficult to be dissociated from it.
The gonads are embedded into the tunic, with only one testis follicle and one ovary (Figs 2A, 3A-C). The testis
is located behind the gut in the internal anterior side of the cardiac arc. It has an irregular outline, either in a
compact rounded mass or more elongated with constrictions. The single sperm duct runs anteriorly along the heart
to join the ovary and is there inflated in a large ampulla (Fig. 3C). The ovary is elongated and bean-shaped, ending
in a wide oviduct (Fig. 3C).
The aggregated individuals collected at Palau only differ from those from Philippines by an oral siphon
particularly long and very large zooids reaching 8 cm.
FlG. 3. — Plurella monogyna sp. nov.: A. external dorsal side with gut, gonads and heart; B. internal side of the body;
C, detail of the testis and ovary.
Source : MNHN, Pans
ASCID1ACEA: PACIFIC PLURELLIDAE
709
Remarks. — The conglomerate form as described above has exactly the same external characters as the solitary
Microgastra specimens described by KOTT (1985). NISHIKAWA (1986) gave a very precise description of what he
named Ascidici (?) aenigmatica, which he later synonymized (1991) with Microgastra granosa Kott. 198r>.
Nishikawa in 1991 mentioned 2 different solitary forms, one collected at Oki Islands with only one ovary, and
one from Hakodate with several ovaries, which cannot belong to P. monogyna but to another dilferent Microgastra
species.
In 1985 following her description of Plurella elongata, KOTT mentions specimens from Philippines,
insufficiently described but which may also belong to Plurella monogyna.
Considering the similarity of Microgastra granosa and its synonyms and the pseudo-colonies from Palau and
the Philippines, a distinction between the genera Plurella and Microgastra is hazardous; nevertheless we retained it
but only based on the solitary condition of Microgastra opposed to a possible colonial structure ol Plurella.
Plurella testacea sp. nov.
Figs IB. 2C, 4
MATERIAL EXAMINED. — Coral Reef Research Foundation coll. X1MUM Dl , ,,
Philippines. Davao. Pakiputan strait. 07°07.43'N. 125°40.I8'E, 25 m. 29.03.1996 (MNHN P7 Plu 13).
Type. — Colony mentioned above (MNHN P7 Plu 13).
DESCRIPTION. — The pseudo-colony is embedded with coarse sand and varied debris (Fig. IB). 1 he zooids are
about 6 cm long. They are incompletely and irregularly attached to each other, leaving cavities between the zooids.
The specimen remains soft in spite of the heavy incrustation. The oral and cloacal siphons are Iree and their colour
at the rim is sulphur yellow. The tunic is thin, colourless, vitreous except on the right side where it is brown,
sclerified opaque and stiffen there is no embedded sand at this level.
The oral siphon is clearly longer than the cloacal siphon (Fig. 2C). Its edge has no well marked lobes and only
possesses a weak muscular sphincter. One of the zooids shows a doubled oral siphon (Fig. 20. The internal side
of the oral siphon is smooth. The cloacal siphon has a weak sphincter and a lobed margin. It begins at the hall
body-length and is directed anteriorly (Fig. 2C). .
A circle of about 20 tentacles in two orders of length is at the anterior 2/5 of the length ot the siphon
(Fig. 2C), without velum. The prepharyngeal groove is thin, very posterior at the base of the oral siphon and cut
in a deep dorsal V. . . , ,,
The body wall is extremely thin. Parallel transverse muscle fibres cover the half posterior part ot the right body
side (Figs 2C, 4A). On the left side the body wall is not very transparent and cannot be separated from the tunic
except in the very posterior part of the body. Blood sinuses penetrate the tunic at the level of the endostyle, ol the
gut and of the gonads. , . , . , ,
The branchial sac originates very close to the prepharyngeal groove. It is Hat except the high dorsal I old
characteristic of the Plurellidae (Fig. 4A). The longitudinal vessels begin by a papilla anterior to the first row ot
stigmata, and posteriorly bear papillae above each transverse vessel. We counted I to 2 long stigmata in a mesh.
Frequently the stigmata rows are doubled and intermediate transverse vessels appear with their papillae. At its
origin the dorsal lamina is made of two blades on each side of a short groove and it is posteriorly prolonged in a
plain membrane. The dermato-branchial strips are numerous and irregular. The endostyle is wide and Hat
The neural ganglion is included in a thicker part of the body wall, strongly linked to the tunic, at the base o
the cloacal siphon. The neural gland is not distinct, but a thin channel linked with multiple urn-shaped openings
lies from the neural ganglion to the prepharyngeal V. The last ciliated pit takes place inside the dorsal V. The heart
draws a wide arc from the endostyle to the dorsal line in the posterior part of the left body-side (Fig. 4A). Doi sally
it has a hernia which contains a large inclusion. .
The gut draws a longitudinal open arc (Figs 2C, 4A-B). The oesophagus is very short, narrow. The stomach is
oval, the intestine is long, not curved in the rectal part. The anus is wide open at the base ol the cloacal siphon.
The male and female gonads are distant from each other (Figs 2C, 4B). The massive testis is included him e
the tunic, posterior to the heart. A long and thin sperm duct joins a globular ovary and expands in a large
710
F. MONNIOT & C. MONNIOT
ampulla. The ovary is located dorsally, close to the anus facing the arc made by the gut (Fig. 4B). It is embedded
in the tunic.
REMARKS. — This species only differs from other Plurella species by internal characters which are. a brown
thickened and sclerified tunic on the right side of the body (Fig. 2C); few oral tentacles planted very anteriorly in
the siphon, far from the prepharyngeal groove: gonads made of a single testis mass posterior to the heart and a
single ovary at the side of the intestinal loop; a gut slightly curved, elongated antero-posteriorly.
Fig. 4. — Plurella testacea sp. nov.: A. internal side of the body; B, detail of the gut and ovary.
Plurella kottae Monniot & Monniot, 1996
Figs 1C. 2B. 5
MATERIAL EXAMINED. — Coral Reef Research Foundation coll.
Philippines. Mactan Island : 10°17.19'N, 124°00.15'E. 27 m, 7.02.1994 (MNHN P7 Plu 1).
Cebu: no more information.
Palau. Sea Bear: 07°16.4I'N. 134°31.43'E, 95 m.
Tsais Reef: 07°I7.90'N, 134°13.80'E. 100 m.
Papua New Guinea. Yuma Pass: I1°22.I4'S. 153°20.99'E, 40 m. 3.06.1998.
TYPE. — MNHN P7 Plu 1. Philippines, Mactan Island, 10°17.19'N. I24°00.15'E. 27 m .
Description. — In addition to the type sample, new specimens were collected in the Philippines and Palau
islands. They allow to confirm and add some details to the previous description. The individuals, 2 to 3cm long are
strongly aggregated (Fig. 1C) and lie on each other as roof-tiles with the same orientation. The siphon edge is
yellow. Both siphons are short (Fig. 5A) with pointed lobes. The tunic is deeply impregnated with sand and
Source : MNHN, Paris
ASC1DIACEA: PACIFIC PLURELLIDAE
71 1
broken shells, but thin and translucent. On the right side it is separated into two layers, one external embedded
with sediment and one internal, very thin, vitreous, soft and naked. This internal layer is not linked to the
muscular body wall except along the endostyle and the dorsal line. On the left side, the tunic cannot be dissociated
from the body wall and includes the heart and the gonads.
There are about 25 tentacles of irregular length and few other very small between them. The tentacles are
located at the base of the siphon (Fig. 2B). The internal side of the oral siphon is covered with minute papillae.
The prepharyngeal groove is close to the branchial tissue and curved in a sharp V. The body wall muscles extend
transversely on the whole right side of the body. The oral sphincter is weak, the cloacal sphincter is stronger with
short crossed longitudinal fibres.
The neural ganglion is oval at the base of the cloacal siphon. The neural gland was not seen . A group of urn¬
shaped vesicles opens anteriorly to the ganglion into the cloacal cavity, and few of them take place along the dorsal
line, the last one opens in the dorsal V.
The branchial sac has a high fold extending from the prepharyngeal V to the bottom ot the body. We counted
2 stigmata in a mesh. The longitudinal vessels possess one papilla at each crossing with the transverse vessels.
The oesophagus entrance is placed in the middle of the dorsal line. The gut forms a closed loop (Figs 2B. 5A),
only attached to the branchial sac by few thin strips; The oesophagus is very short, not well separated from
the stomach, that is only wider. The intestine is relatively long, folded on itself and a short rectum is curved
anteriorly, ended in a wide anus.
FlG. 5. _ Plurella koime Monniot F. & Monniot C. 1996: A. internal side of the body: B. ovaries
712
F. MONNIOT & C. MONNIOT
The heart draws a wide arc at about the three posterior quarters of the body length. It has a dorsal hernia.
Against this hernia, and attached to the heart, is a testis mass, below the oesophagus (Figs 2B, 5A). Three to
seven ovaries are along an antero-posterior line, with the oviducts directed dorsally (Fig. 2B). Each ovary receives
a branch of the long sperm duct with a distal extremity enlarged in an elongated seminal vesicle (Fig. 5B)
No velum was observed at the base of the cloacal siphon.
Remarks. — Among the Plurella species with several ovaries, P. kottae differs from P. melanesiae in
a shorter size of the zooids, a more elongated and folded gut and only one testis lobe. P. elongata has the cloacal
siphon directed posteriorly, a lowest branchial fold, and an open gut loop. Some material collected after 1985
identified by KOTT as Plurella sp. from Philippines corresponds to P. kottae.
Plurella melanesiae sp. nov.
Figs ID, 2D. 6
Material EXAMINED. — Vanuatu. Tongoa Island : 16°52.43’S, 168°31.88'E. 35 to 45m, dive. G. BARGIBANT/IRD
coll. (MNHN P7 Plu 12).
Type. — MNHN P7 Plu 12, Vanuatu, Tongoa Island: 16°52.43'S, 168°31.88'E, 35 lo 45m.
Description. — In the massive colony (Fig. ID), the zooids are parallel, 5 to 6 cm long, attached to each
other, but cavities of undetermined shape remain between groups of zooids. The tunic is relatively thick when
compared to the other species ol the same genus. It is embedded with coarse sand except on a part of the right side
of the body. There, it is sclerified and brown in colour. It is not attached to the muscular body wall. On the left
side of the body, facing the gut and gonads, the tunic is very thickened with encrusted sediment on the external side
only. There are no muscles at this level. There is an additional thickening of the body wall and tunic in the shape
ot an arc parallel but interior to the cardiac arc. This thickening increases towards the dorsal side, including the
cardiac hernia and the gonads. This fibrous thickening does not include sediment particles, except on the external
layer. The tunic and its incrustation extends at the base of the zooids behind the body wall, probably to anchor the
animal in the sediment.
Both siphons are orange, short, directed anteriorly with a thin wall, all damaged. Their rim is neatly toothed,
we counted 40 to 50 tentacles in a circle, about 20 of them are long. They are at a short distance from the
prepharyngeal band. The body is elongated and the cloacal siphon arises from 1/4 of the branchial sac. On the
siphons and the right body side, the body wall contains a brown-orange pigment. The musculature (Fig. 2D) is
made of parallel and dense transverse fibres on the right side starting from the anterior part of the branchial sac to
the posterior extremity of the body.
The neural ganglion is oval, slightly above the base of the cloacal siphon, in a thickening of the tunic. The
neural gland has not been seen. The neural gland channel extends up to the prepharyngeal band, with short branches
ended in urns, numerous near the neural ganglion, more sparse towards the distal extremity. The branchial sac has
the same stiucture as in others Plurella species, with a long dorsal fold. It is damaged in all specimens which
contain several kinds ot symbionts : nemerteans, Pontoniidae shrimps, amphipods, polychaetes.
The heart draws a wide half circle at 3/4 of the body length (Figs 2D. 6A, C). It has a dorsal hernia containing
an inclusion. Dorsally, applied on the cardiac hernia and inside the thickening of the tunic is the testis (Figs 2D,
6A, D). made of 2 to 6 lobes, more or less close to each other in a bush. These lobes are partly interior and
exterior to the cardiac-arc extremity, each having a thin sperm duct. The sperm ducts become parallel and then
diverge to join 2 to 4 elongated ovaries (Fig. 6E). The ovaries, included inside the tunic, are placed on a transverse
line a the base of the cloacal siphon (Fig. 2D). To each ovary is associated a dilated part of a sperm duct containing
numerous spermatozoa. The male and female papillae are joined in each gonad.
The gut forms a loop, more or less closed, very slightly attached to the branchial sac. It is placed in the first
hall of the body-length (Figs 2D, 6A, C).
Remarks. This species differs from all other Plurella by its testis divided in several lobes, each of them
having an independent sperm duct.
Source : MNHN, Paris
ASCIDIACEA: PACIFIC PLURELLIDAE
713
few
Fig. 6. — Plurella melanesiae sp. nov.: A and B. both sides of the zooid; C, internal side of a zooid ventrally opened;
D. heart diverticulum and testis; E, ovaries.
Source : MNHN, Paris
714
F. MONNIOT & C. MONNIOT
Microgastra granosa (Sluiter, 1904)
Synonymy:
Ascidia granosa Sluiter, 1904: 36.
Microgastra granosa Kott, 1985: 70. — Nishikavva. 1991: 57 and synonymy. — Monniot C. & MONNIOT
F.. 1991a:
396.
MATERIAL EXAMINED. — Ride des Loyaute. MUSORSTOM 6: st. CP 419, 20°4I.6'S-167 03.7 E. 28.’ m,
16.02.1989.
The single specimen collected in New Caledonia is too damaged to determine precisely if it corresponds to
specimens studied by KOTT or NlSHIKAWA, but all these samples are closely related. The presence of only one
testis and a single ovary, associated with other characters of the species Plttrella monogma, questions the validity
of the aggregated condition as a valuable generic character. More specimens will be necessary to solve this question
and to finally determine whether there is a true colonial state in the genus Plttrella.
Genus PLURASCIDIA nov.
DIAGNOSIS. — Solitary ascidian impregnated with sand. A dorsal tubercle and minute urn-shaped pits along
the neural gland channel. Branchial tissue flat with longitudinal vessels and papillae, and with a fold on the left
side. Male and female gonads separated linked to the gut loop. No gastric gland. Small excretion vesicles covering
the gut. Body musculature transverse.
Type Species. — Plurascidia marquesana sp. nov.
ETYMOLOGY. — Combination of two family names: Plurellidae and Ascidiidae; Gender is feminine.
Plurascidia marquesana g. nov. sp. nov.
Figs IK. 8-12
MATERIAL EXAMINED. — Marquesas Islands. MUSORSTOM 9: stn DW 1152. Eiao, 7°58.9'S. 140°43.5'W. 85-
150 m, 23.08.1997. — Stn CP 1156. Eiao, 7°59.0'S. 140°43.7'W, 80 m, 23.08.1997. — Stn DW 1186. Nuku Hiva.
8°48.1'S, 140°03.5'W. 42-45 m. 26.08.1997. — Stn CP 1187. Nuku Hiva. 8°49.2'S. 140°03.5'W, 25-30 m. 26.08.1997.
— Sin DW 1209, Hiva Oa, 9°50.2'S, 139°00.8'W. 85 m, 29.08.1997. — Stn CP 1212. Hiva Oa. 9°49.9'S. 139°02.2'W.
50-80 m, 29.08.1997. — Stn DW 1217. Hiva Oa. 9°44.5'S. 138°49.9'W, 85-87 m. 30.08.1997 (MNHN P7 Plu 2). —
Stn CP 1227. Hiva Oa, 9°44.2'S-138052.5'W. 84-85 m. 30.08.1997, — Sin DW 1230. Hiva Oa. 9°43.6'S, 139°06.6'W.
95-100 m. 31.08.1997. — Stn CP 1239. Hiva Oa. 9°42.2’S, 139°03.6'W, 89-95 m. 31.08.1997.
Type. — Marquesas Islands (Hiva Oa). MUSORSTOM 9. stn 1217. 9°44.5'S, 138°49.9’W, 85-87 m (MNHN
P7 Plu 2).
DESCRIPTION. — The individuals are solitary, ovoid, isolated, covered with coarse sand and broken shells. The
zooids are not agglomerated but live free on the soft bottom, in large populations. There is no peduncle and no
rhizoid. The oral siphon is terminal, the cloacal siphon at mid-distance of the body length. When iixed. the siphon
apertures are not visible through the sediment embedding. The individuals measure 2 to 3 cm in length and 1 .5 to
2 cm in diameter. All are mature.
The tunic is extremely thin, transparent and colourless. It adheres very slightly to the body wall and can easily
be removed except at the edge of the oral siphon. The body wall is thin, perfectly transparent (Fig. 7A ). Only
small pigment spots are placed at the base of the oral lobes. They probably correspond to sensory organs as thin
neural endings leads to them. The oral siphon has 8 large petal-like lobes separated by 8 smaller triangular lobes.
Immediately below these lobes is a muscular sphincter. Some longitudinal fibres cross it but do not extend until
the tentacular circle. More than 100 tentacles (Figs 8-9) in 3 orders of size (plus buttons) are inserted at the base of
a velum containing circular muscle fibres and a blood vessel. Between the oral tentacles and the prepharyngeal
Source : MNHN, Paris
ASCIDIACEA: PACIFIC PLURHLLIDAE
715
band, the body wall is covered with small papillae. In this thin thickness 2 longitudinal blood vessels issued Ironi
the heart extend up to the edge of the oral siphon. One of them tollows the endostyle and altcrwards the medio-
ventral line, the other runs along the neural gland channel and the dorsal line. These vessels are coated with spiral
muscle fibres.
Pig 7, — a, Plurascidia marque sana sp. nov„ free from tunic; B, Fimbrora calsubia Monniot C. & Monniot F., 1991. tree
from tunic, stained with hemalum.
Source :
pens - •'*'
716
F. MONNIOT & C. MONNIOT
Source : MNHN, Paris
ASCIDIACEA: PACIFIC PLURELLIDAE
717
The prepharyngeal band has two blades curved in a V; the posterior blade is prolonged in two parallel crests on
the sides of a furrow extending dorsally along 4 to 5 stigmata rows. These 2 crests are posteriorly prolonged by a
single dorsal lamina, with a plain margin.
The neural ganglion is rectangular and voluminous (Figs 7A, 9-10). A neural gland is associated to it with an
inconspicuous outline. The neural gland channel is straight but bears short numerous branches along its whole
length ending in small ciliated urns. A true dorsal tubercle, also urn-shaped, is placed at the bottom of the
prepharyngeal V (Fig. 10).
The dorsal part of the body wall is covered with a rectangular field of large muscular fibres, transverse and
parallel (Figs 7A, 8, 1 1C). Not any longitudinal fibre crosses them. The cloacal siphon opens in the middle of the
field. It is edged by 6 petal-like lobes and circled by a muscular sphincter. Some short radial fibres cross the
sphincter but do not overpass the siphon level (Figs 7A, 8).
FIG. 9. — Plurascidia marquesana sp. nov.: Neural area and oral siphon.
0.5 inn
Fig. 10. _ Plurascidia marquesana sp. nov.: Neural ganglion, neural gland and its multiple openings, dorsal tubercle.
dorsal blood vessel.
The branchial sac is only attached to the body wall along the endostyle and by tissular strips particularly thick
at the level of the anterior part of the dorsal lamina. The branchial sac begins against the prepharyngeal band. It is
Hat anteriorly, but a fold progressively appears on the left side and becomes higher at the bottom ol the branchial
718
F. MONNIOT & C. MONNIOT
sac (Fig. 12). It is hold by strips and a soldering at the level of the retropharyngeal line. The branchial sac is
soldered to the rectum. In a specimen of median size, we counted 48 longitudinal vessels on the left side and 52 on
the right, and about 80 rows of stigmata. The longitudinal vessels only begin at the first transverse vessel or first
parastigmatic vessel, but are not prolonged in the space between the prepharyngeal band and the first stigmata row
as they do in the Plurella species. The longitudinal vessels bear ciliated papillae at the crossings with the
transverse and parastigmatic vessels when they exist (Fig. 12). The endostyle has no peculiar characters. It ends at
the bottom of the branchial sac against the branchial fold (Figs 7A, 8A, 1 1A-B) .
The digestive tract occupies a large part of the left side. It is full in all dissected animals (Fig. 7A). The
oesophagus opens at 4/5 of the body length with a papillated internal wall. It is narrow and short. The oval
stomach has some low longitudinal folds, 4 to 5 on each side (Fig. 7A), but its wall remains thin. A closed
intestinal loop is narrowed by a neat constriction (Fig. 1 ID). A last segment, widened and with a specially thin
wall constitutes the rectum. The anus is very small on the dorsal part of the rectum (Fig. I 1C). The oesophagus,
stomach, and first intestinal loop are covered with minute vesicles, each containing a concretion, a structure
frequent in the Ascidiidae family.
Fig. II. — Phirascidia marquesana sp. nov.: A and B. left side of two specimens: C. specimen opened ventrally;
D. external side of the gut; E. internal side of the gut and gonads.
Source : MNHN, Paris
ASCID1ACEA: PACIFIC PLURELLIDAE
719
The internal side of the gut (except the rectum) is covered by the gonads (Fig. 1 IE). The testis and ovary make
a ramified network of undefined outline. The oviduct is long, wide and straight, filled with eggs running along the
oesophagus, stomach and rectum (Figs 7A, 1 ID). It opens by a large papilla with a toothed margin. The sperm
duct is dilated by the spermatozoa, attached to the oviduct and it opens at the same level.
Fig. 12. — Plurascidia marquesana sp. nov.: Posterior dorsal part ot the branchial sae.
Source : MNHN. Paris
720
F. MONNIOT & C. MONNIOT
The heart forms a large arc, first encircling the posterior part of the gut and then reaching the cndostyle. The
cardiac cells are in longitudinal lines and draw thin ridges after staining. A diverticulum in hernia placed against the
stomach contains a concretion. A complex blood system of large vessels starts from the heart. These large vessels
are lined by helicoidal muscular fibres. The largest vessels are under the endostyle and medio-dorsal, and directed
towards the gut and gonads. These vessels are similar to those described in the species Fimbrora calsubia Monniot
C. & Monniot F., 1991.
Remarks. — Plurascidia niarquesana has in common with the Plurella species an oral siphon with lobes more
or less internally folded when contracted lined interiorly with minute papillae; numerous oral tentacles in one cir¬
cle; many accessory urn-shaped openings along the length of the neural gland channel; a branchial sac with numer¬
ous longitudinal vessels, papillae, and a fold on the left side; a heart in a long tube in an arc with a hernia contain¬
ing a concretion; a musculature made of transverse parallel bundles restricted to a limited area ot the body wall.
" Many differences can be noted with the Plurella species these characters bringing Plurascidia niarquesana nearer
to the Ascidiidae : the presence of a dorsal tubercle; the longitudinal vessels which do not reach the top of the first
row of stigmata; the gonads male and female linked to the gut; the presence of renal vesicles covering the intestine.
The new species Plurascidia niarquesana , though different in aspect, possesses characters common to Fimbrora
calsubia Monniot C. & Monniot F., 1991, a deep species assigned to the Ascidiidae family. The genera Fimbrora
and Plurascidia have in common the general branchial structure, the gonads linked to the digestive tract, the struc¬
ture of the vascular system and the absence of muscles on the left side of the body. Both genera have a muscular
sphincter at the base of the oral tentacles and 2 large longitudinal blood vessels, dorsally and ventrally. in the oral
siphon. These 2 genera differ by the structure of the tunic, the branchial fold present in Plurascidia, the carnivorous
diet in Fimbrora, filter-feeding in Plurascidia. Only one dorsal tubercle in a slit exists in Fimbrora, when multiple
urn-shaped apertures are found in Plurascidia. Finally Fimbrora has an accessory gland on the stomach.
Fimbrora calsubia Monniot C. & Monniot F., 1991
Fig. 7B
Fimbrora calsubia Monniot C. & Monniot F., 1991: 383, figs 1-6. — Monniot C., 1993: 356.
The single species in this genera was collected at 1865 m off New Caledonia and at 1020 m near Tanimbar
Island in Indonesia. The anatomy of these specimens is remarkably constant and differs in numerous characters
from the Plurellidae and the new genus Plurascidia.
The tunic is not embedded with sediment but transparent, thick on the ventral side where it has a cartilaginous
consistency. The oral siphon is extremely developed (Fig. 7B), basket-like, but the mouth is narrow in the bottom
of the siphon equipped with a strong sphincter. The dorsal and anterior part of the branchial sac is attached to the
body wall, and it is in this thick tissue that the neural gland channel spreads, its single opening being located in
the dorsal V of the prepharyngeal band. The stomach with some longitudinal folds is covered by a voluminous
gland in the shape of a cauliflower (Fig. 7B). The male and female gonads lobes are separated but lined along the
internal side of the intestine, the oviduct and sperm duct parallel, long, are applied against the intestine. The gut
and gonads are included in the body wall.
Nevertheless, some common characters with the Plurellidae and the genus Plurascidia have to be emphasised,
the structure of the branchial tissue, with longitudinal vessels topped by papillae at the crossings of the transverse
and parastigmatic vessels. A dorsal lamina in a plain membrane. The heart in the shape of a long tubular arc at the
posterior part of the body. The musculature limited to one side of the body wall. Fimbrora has also in common
with Plurascidia the muscular helicoidal fibres found in the main blood vessels.
ACKNOWLEDGMENTS
We thank the Queensland Museum for the loan of Plurella specimens. We are very indebted to Todd
Newberry for his suggestions and advice regarding the English version of this paper.
Source : MNHN, Paris
ASC1DIACEA: PACIFIC PLURELLIDAE
721
REFERENCES
Kott P.. 1973. — Plurellidae a new phlebobranchiate family of Ascidiacea. Proceedings of the Linnean Society of New
South Wales, 97: 258-261.
Kott P.. 1985. — The Australian Ascidiacea. Part 1. Phlebobranchia and Stolidobranchia. Memoirs of the Queensland
Museum, 23: 1-440.
MONNIOT C., 1993. — Tunicata: Sur trois espfeces d'ascidies bathyales recoltees au cours de la campagne Iranco-
indonesienne Karubar. In: A. CROSNIER (ed.) Resultats des Campagnes MUSORSTOM. vol. II. Memotres du Museum
national d'Histoire naturelle, 158: 355-359.
Monniot C. & MONNIOT F.. 1991a. — Decouverte d’une nouvelle lignee evolutive chez les ascidies de grande profondeur:
une Ascidiidae carnivore. Comptes retidus de I’Academie des Sciences, Paris, 312, serie III. 383-388.
Monniot C & Monniot F„ 1991b. — Tunicata: Peuplements d'ascidies profondes en Nouvelle Caledonie. Diversity des
strategies adaptatives. In: A. CROSNIER (ed.), Resultats des Campagnes MUSORSTOM. vol. 8. Memotres du Museum
national d’Histoire naturelle, (A), 151: 357-448.
Monniot F. & Monniot C„ 1996. — New collections of ascidians from the western Pacific and southeastern Asia.
Micronesica, 29 (2): 133-279.
Nishikawa T., 1986. — Some Ascidians dredged around the Oki Islands, the Japan Sea. Memoirs of tlte National Science
Museum, Tokyo, 19: 175-184.
Nishikawa T„ 1991. — The Ascidians of the Japan Sea. II. Publication of tlte Seto Marine Biology Laboratory, 35 (1-3):
25-170.
Sluiter C. P., 1904. — Die Tunicaten der Siboga Expedition. Part I. Die socialen und holosomen Ascidien. Stboga
Expeditie, 56A: 1-139.
Source : MNHN. Pahs
LTATS DES CAMPAGNES MUSORSTOM, VOLUME 21 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 21 — RESULTATS DES
Pisces Gadiformes: Grenadier Fishes of
the New Caledonian Region, Southwest Pacific
Ocean. Taxonomy and Distribution,
with ecological notes
Nigel R. MERRETT
Department of Zoology,
The Natural History Museum.
Cromwell Road, London SW7 5BD,
United Kingdom
Present address: Muttlebury's Mead, Chard Street,
Thorncombe. Chard. TA20 4NB, U.K.
&
Tomio IWAMOTO
California Academy of Sciences,
Golden Gate Park, San Francisco.
CA. 94118. U.S.A.
ABSTRACT
We reported in an earlier paper the great species richness of the grenadier fauna (families Bathygadidae and
Macrouridae) from recent bathyal trawl collections made mainly during MUSORSTOM cruises in the New Caledonian region.
Here we add information from further samples to complement earlier taxonomic findings, and descriptions of 2 new
species, together with comments on species, distribution and ecology. Thus a total of 2055 specimens trom 221 samples
examined representing 20 genera and 63 species were found to have closest similarity in composition with New South
Wales. Western Australia and New Zealand. As expected, dissimilarity increased with distance from New Caledonia. Four
genera dominated in species richness: Caelorinchus (17), Hymenocephalus (8), Nezumia (5) and Veit trifossa (6),
comprising 2/3 of the total fauna. The generic make-up of the faunas closest to New Caledonia were most consistent with
that region; proportions varied radically from there in the more distant regions investigated. Bathymetrically, the
smaller trawls of the MUSORSTOM surveys collected grenadiers over a range of tows shallower than that reflected by the
Merrett, N.R. & Iwamoto, T„ 2000. — Pisces Gadiformes: Grenadier Fishes of the New Caledonian Region.
Southwest Pacific Ocean. Taxonomy and Distribution, with ecological notes. In: A. Crosnier ted.). Resultats
des Campagncs MUSORSTOM, Volume 21. Memoires du Museum national d'Histoire naturelle. 184: 723-781. Paris ISBN
2-85653-526-7.
724
N.R. MERRETT & T. IWAMOTO
commercial gear used on the Halipro 2 cruise, with a generally smaller size of fish sampled. Co-occurrence of grenadier
species within similar depth strata on the slope was remarkably high, with only two of the 63 species not represented, at
least over part of their depth range, in the upper 1600 m. Species richness peaked at 37 in both the 700 and 800 m strata,
although it did not drop below 20 across the depth range 400-1 100 m and reduced substantially only deeper than 1400 m.
RESUME
Pisces Gadiformes: Grenadiers de la region neo-caledonienne, Sud-ouest Pacifique. Taxo-
nomie, distribution et notes ecologiques.
Nous avons attire l'attention dans un travail precedent sur la grande richesse en especes de la faune des grenadiers
(families Bathygadidae et Macrouridae) rdcoltde rdcemment dans la region neo-calddonienne, au moyen de chalutagcs dans
la zone bathyale, principalement au cours des campagnes Musorstom. Ici nous donnons des informations
complementaires fournies par de nouvelles recoltes. decrivons deux nouvelles especes et fournissons des donnees sur la
distribution des especes et leur ecologie. Ainsi un total de 2055 specimens provenant de 221 recoltes, renfermant
20 genres et 63 especes, ont montre que la composition de cette faune est la plus proche de cclles de la Nouvelle-Galles du
Sud, de I'Australie de l'Ouest et de la Nouvelle-Zelande. Comme on pouvait le prevoir. la dissimilarity s'accroit avec
I'eloigncment de la Nouvelle-Caledonie. Quatre genres prddominent en ce qui concerne le nombre d'espcces:
Caelorhynchus (17). Hymenocephalus (8), Nezumia (5) et Veil trifossa (6), representant les 2/3 de la faune totale. La
composition en genres des faunes des regions les plus proches de la Nouvelle-Caledonie est homogene; elle varie par
contre nettement pour les zones plus eloignees. D'un point de vue bathymetrique, les recoltes des campagnes Musorstom
faites avec des petits chaluts ont capture des grenadiers & des profondeurs inferieures h celles observees durant la campagne
Halipro 2 faite avec un chalut commercial; de meme, les tailles des grenadiers captures alors ont did inferieures a celles des
captures faites lors d'HALlPRO 2. II n’y a que deux espbces sur les 63 recensees qui ne soient pas representees, au moins pour
une partie de leur repartition bathymetrique, dans la tranche 300-1900 m, ce qui souligne le nombre eleve d'especes de
grenadiers susceptibles de se rencontrer ensemble sur la pente, dans une meme tranche bathymetrique. La diversity
maximale en especes a culmine pour les tranches des 700 et 800 m ou 37 esp6ccs ont ete relevees; cetle diversity n'est pas
tombec <t moins de 20 entre 400 et 1 100 m et ne s'est subtantiellement reduite qu’au deli de 1400 m.
INTRODUCTION
Exploratory collections at bathyal depths of the Exclusive Economic Zone of New Caledonia, in the Southwest
Pacific, have been carried out by the Institut de Recherche pour le Dcveloppement (IRD - formerly ORSTOM)
since 1984 (Richer de Forces, 1990; Richer de FORGES & MENOU, 1993; Grandperrin et al. , 1997; Seret,
1997). The greater part of these collections were deposited in the Museum national d'Histoire naturelle, Paris,
where they were sorted into family groups by Bernard Seret and allocated to specialists to work up. Iwamoto &
MERRETT (1997) studied the astonishingly diverse grenadier fauna, families Bathygadidae and Macrouridae. and
recorded 49 species in 16 genera from the samples collected up to 1993. Now more recent collections made during
three cruises by ORSTOM off Vanuatu (Musorstom 8), New Caledonia (Bathus 1 ), and the Norfolk and Loyalty
Ridges (Halipro 2) have added 14 spp. and 4 genera, raising the total to 63 species (two listed as 'indeterminate' in
the previous paper) in 20 genera.
The New Caledonia group of islands is an emerged portion of the Norfolk Ridge, a geosyncline which extends
southwards towards New Zealand. While some of the collections of grenadiers were made in the vicinity of New
Caledonia and along the Norfolk and Loyalty Ridges, others originated from an extensive area to the west on the
Chesterfield and Bellona Banks, and Fairway and Lord Howe Rises. All these were located on the main Indo-
Australian Plate, but separated by the New Hebrides Trench from another group of samples taken around the
Matthew and Hunter volcanoes and Vanuatu Islands, also on this Plate. The tectonically complicated North Fiji
Basin and the Vitiaz Trench separated these from the most easterly group of samples, which were taken around
Wallis and Futuna Islands on the Pacific Plate (Fig. I a). The complexity of the region overall is briefly described
by Richer de Forces & MENOU (1993). Two hundred and twenty-one samples contained grenadiers from a total
ot 970 (Table I) taken between 280 and 3690 m by a variety of towed gears. All these grenadier samples were
kindly made available to us and were taken between 280 and 2105 m depth, but were concentrated in the 300-
1400 m zone.
Source : MNHN, Pans
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
725
The purposes of this paper are (1) to report on additional grenadier collections resulting from these three
cruises, (2) discuss aspects of the distribution of the overall grenadier fauna arising from this and the earlier paper
(Iwamoto & Merrett, 1997) and (3) record notes on the population ecology of a few of the more abundant
species.
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Fig. 1 a. — General chart of area.
MATERIALS AND METHODS
A variety of gear was used to collect the samples. The majority of samples containing grenadiers were taken
with a beam trawl (CP - 127 tows) and an orange roughy bottom trawl (ORBT - 65). supplemented by an
American shrimp trawl (CC - 19), a Waren dredge (DW - 7) and a Charcot dredge (DC - 3; Table 1 ). The
dimensions of these gears vary considerably and are given for the CP (mouth opening 4 m by 0.4 m) in FOREST
(1976). Richer DE FORGES (1990), for the ORBT (mouth opening 26 m by 8 m) in Grandperrin et al (1997),
for the CC (mouth opening ca 9 m by ca 1.5 m) in FOREST (1976), the DW (mouth opening 0.71 m by 0.28 m)
in RICHER DE Forges (1990) and for the DC (mouth opening 0.8 to 1.2 m by 0.35 m) in RICHER DE FORGES
(1990). The complete lists of the station data are given by RICHER DE FORGES (1990), GRANDPERRIN & LEHODEY
(1992), Lehodey et al (1993), Richer de Forges & Menou (1993) and Grandperrin et al. ( 1997). Depth ol
the capture was taken as the mid-point of the soundings recorded at the start and finish ol the tow. The variation in
Source
726
N.R. MERRETT & T. IWAMOTO
the effectiveness of the gears used in sampling active species like grenadiers (see below - pp. 730-731, table 1).
together with the lack of accurate times of arrival and departure of the gear to and from the seabed, prevented
meaningful analysis of fish density or biomass. Thus in considering species distribution, the species
presence/absence approach was the only one tenable. Catches were referred to the sample mid-depth soundings and
treated by 100 m bathymetric strata. In this connection, it should be noted that many of the ORBT hauls were
down-slope tows from off the tops of seamounts. Hence the depth limits of individual tows were sometimes in
excess of 400 m. In these cases, normalizing to mid-depth values introduced uncertain actual capture depths of
different species which should be recognized as being implicit in the results.
Table 1. — Summary of sample collections by gear and area: (a) overall collection, (b) samples containing grenadiers.
(CP - beam trawl; ORBT - orange roughy bottom trawl; CC - shrimp otter trawl; DW - Waren dredge; DC - Charcot
dredge).
(a) Overall sampling:-
Geographically, samples containing grenadiers were allocated to one degree square units, which themselves were
distributed into four regional groupings (Fig. lb). Group I catches were the most westerly in location on the Indo-
Australian Plate, taken from the area of the Chesterfield and Bellona Banks and the Fairway and Lord Howe Rises.
East of this were the catches taken on the slopes around New Caledonia and along the Norfolk and Loyalty Ridges
(Group II). Group III catches were from northeastward of the Group II stations, beyond the subduction zone where
the Indo- Australian Plate sinks below the Pacific Plate (RICHER DE FORGES, 1990), along the western and
southern slopes of the Vanuatu Islands and the Matthew and Hunter volcanoes. Considerably remote from Group
III, again in the northeasterly direction, the Group IV catches were taken in the vicinity of Wallis and Futuna
Islands on the Pacific Plate.
The samples were fixed in saline formol and later transferred into 70% industrial ethanol. Length measurements
were made on the preserved material in accordance with Iwamoto (1970) and Iwamoto & WILLIAMS (1999).
Where possible, stomachs were opened and preliminary assessment of their contents made according to broad
groupings of recognizable organisms. At the same time these specimens were sexed and ovarian maturity was
assessed according to Merrett (1970).
Institutional abbreviations follow Leviton et al. (1985) and Leviton & GlBBS (1988). The reader is referred
to Iwamoto & Merrett (1997) for discussion and explanation' of the taxonomic section. Because this second
paper is intended for use with the first, we have omitted information ip the taxonomic section that has already been
presented in the first article. Thus, the diagnoses, descriptions, and synonymies are omitted, excepl for species not
previously treated. Synonymies may be included, however, for references to recent articles. Size and distribution are
also not included in the species accounts unless additional information has been obtained.
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
727
10 12 14 16 18 20 22 24 26
Degrees latitude - S
Fig. I b. — Schematic represeniation of the 48 degree squares occupied by the 221 stations containing greanadiers by
region (I - IV). Sampling regions I (Chesterfield and Bellona Banks. Fairway and Lord Howe Ridges - 27 samples) and
II "(New Caledonia, Norfolk and Loyalty Ridges - 131) lie on the main Indo-Australian Plate and are separated from
region III (Matthew and Hunter volcanoes and Vanuatu Is. - 44) in the tectonically complicated North Fiji Basin by
the New Hebrides Trench. Region IV (Wallis and Futuna Is. - 19) alone lies on the Pacific Plate.
RESULTS
A total of 2055 specimens representing 20 genera and 63 species of grenadiers occurred in the 221 samples
examined from 48 different degree squares in the area (Table 2). A little more than hall the species richness was
contributed by the genera Caelorinchus (17 species), Hymenocephalus ( sensu lato) (8), Ventrifossa (6) and
Nezumia (5). Forty-two (66%) of these species occurred entirely or in part within the 400-800 m depth range
(Table 2). A little more than half the species (35) were represented in the catches by eight specimens or less, while
14 species were represented by >32 specimens (Fig. 2). Nineteen species were found to be undescribed hitherto
(30% - Iwamoto & MERRETT (1997) and see below). Representation of new species in the catches was not
restricted to individual records (IWAMOTO & MERRETT, 1997 - their Fig. 3). Only four of these were represented
by singletons in the overall collection, reflecting the novelty of the deep ichthyofauna of the area. Indeed, the most
abundant species sampled was new ( Hymenocephalus megalops - 390 specimens from 26 stations; fable 2).
together with another, Ventrifossa paxtoni, (88 specimens, 29 stations) and a third. Lucigadus acrolophus,
(26 specimens, 7 stations).
Table 2 (following pages). — Synopsis of grenadier representation, giving distribution per area, depth and frequency of
capture. [Distr type - Indicates species regional affinities discussed in the section on zoogeography - A, endemic;
B, broad northern tropical; C, southern temperate; D. widespread, global or multiple-ocean-basin representatives.
Tot No, total number of specimens collected per area; F, number of lows from which Tot No' were sampled (No.
hauls/area given in these columns provides a measure of the relative effort per area, from Table I).]
728
N.R. MERRETT & T. IWAMOTO
AREA I
Chesterfield, Bellona,
Fairway, Lord Howe Rise
AREA II
New Caledonia, Norfolk Ridge,
Loyalty
51
Source : MNHN, Pahs
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
729
No. hauls/area
SPECIES
Bathygadidae
Balhygadus cottoides
spongiceps
Gadomus intronger
sp indet.
Macrouridae
Caelorirchus acutirostris
analiroslris
argentalus
celaenostomus
cingulatus
cylindricus
kermadecus
melanobranchus
mycterismus
AREA III
Matthew. Hunter.
Vanuatu
Distr Tot Depth Lat
type No (m) Deg S
C
D
B 3 980-1191 15-22
B 1 399 15
B 5 541-1220 15-16
B 15 398-566 15-19
C
B 4 460-571 16-19
A
C
A
C
C 2 775 18
B
B
A 17 532-830 15-18
C
C
c
c
D
A
597
33
AREA IV
Wallace,
Futuna
Long F Tot Depth
Deg E No (m)
148
TOT
Areas
l-IV
Lat Long F
Deg S Deg W
116
12
53
5
1
3
14
30
10
11 178 1 12
1
90
5
1
7
2
1
30
4
31
1
1
1
14
3
8
3
12 174 1 3
11 179 1 1
11-12 174-178 4 32
10
6
49
11-12 176-179 2 390
11-12 174-179 10 355
3
4
1
1
26
13 176 1 108
1
12 177 1 7
11-12 1 77-178 5 22
2
26
1
2
6
12 174 1 46
11-12 177-178 3 6
1
2
11-12 177-178 3 51
208
1
12 177 2 27
1
11-12 177-179 6 48
12 177 1 43
11
11-12 176-179 10 113
88
10
2055
63
730
N.R. MERRETT & T. IWAMOTO
Frequency categories
Fig. 2. — Species representation by specimen frequency category (specimens per species in the overall collection).
Sampling.
Grenadiers were recovered from only 22.8% of the samples (221 from 970) taken from all gears (Table I ).
Largely this discrepancy could be accounted for by the high frequency of DW tows, which rarely caught grenadiers
in the overall survey. Most of the grenadiers occurred in the ORBT (62.5%), CC (61.3%) and CP (43.6%) hauls.
By area, only the smaller gears (CC, CP and DW) were used throughout; the large ORBT was not used in areas III
and IV (Table 1 ).
A bathymetric plot of the 221 samples containing grenadiers indicates the dramatic effect that the sampling
'window' (in this case, gear type and size) has on the ecological implications of the samples (Fig. 3). The smaller
nets collected grenadiers over a range of tows shallower than that suggested by the commercial gear (ORBT). The
smaller nets sampled grenadiers from the 200-300 m stratum, with good representation down to 700 m. The ORBT
used on the Halipro 2 cruise first collected these fishes in the 500-600 m stratum. Although relatively few ORBT
tows were made at depths shallower than 600 m, sufficient were made to suggest that the lack of grenadier
representation was an artefact. In the 600-700 m stratum 10 hauls lacked grenadiers, while two contained them
(Fig. 4), but in the 1000-1 100 m stratum the situation was reversed, with 14 samples in which grenadiers were
represented and two in which they were not.
The size range of individual grenadiers sampled by the variety of gears was strikingly different (generally small
specimens captured from the small nets vs large from the ORBT). This fact, together with the frequent ontogenetic
smaller-shallower trend in these fishes (cf. discussion of such sampling limitations in MERRETT et al., 1991)
suggests a ready explanation lor this sampling discrepancy. Extrusion through the large mesh size of the 30 m
ORBT trawl probably accounted for a substantial loss of small specimens and the large bobbins used on the
groundline possibly caused escapement beneath the net. This trend was confirmed qualitatively on the Halipro 2
cruise by the inclusion of a small ring net (ca 0.5 m diameter) inside the wings of the ORBT. which collected
a variety of much smaller fishes than were retained on the meshes of the ORBT. In contrast, while the smaller
gears would have sampled the smaller specimens well, large grenadiers could well avoid the on-coming mouth of a
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
731
small net towed at low speed. Taken separately, these two sets of data provide totally different views of the
grenadier fauna of the region. Together, they confirm yet again the limitations influencing the validity of
inferences drawn from deep-sea sampling (cf. ANGEL, 1977).
0 5 10 15 20 25
Frequency
Fig. 3. — Distribution of samples containing grenadiers by 100 m depth strata, showing the difference between the small
net catches and the commercial ORBT used on the Halipro 2 cruises (n = 221).
o 5 10 15
Haul frequency
Fig. 4. — Trawl haul frequency by 100 m strata during the Halipro 2 cruises, divided into those hauls containing
grenadiers and those without (n = 104).
Source
732
N.R. MERRETT & T. IWAMOTO
Distribution - inter-area.
Species' depth distributions are given in Table 2, together with their geographic locations reflected by the
catches. These relate to the regional groupings. Thus the two series of collections on the Indo-Australian Plate
(areas I and II) were represented by 30 and 51 species respectively, whilst areas III and IV, on the Pacific Plate,
yielded 33 and 16 species respectively (Table 3a). Table 3a also shows the percentage contribution to the overall
grenadier species richness of the region, together with an inter-area comparison and the number of species unique to
each. The situation it reveals is interesting, although confused. Areas I and III lacked many species found in area II,
where 25% were unique in the catches there. Whether or not this reflects sampling disparity alone is debatable.
Area IV yielded mostly widespread species and several species also found in Hawaii (e.g. Caelorinchus cingulatus ,
Hymenocephalus aterrimus, Mataeoceplialus acipenserinus, Nezumia propinqua, Sphagemacrurus pumiliceps,
Trachonurus sentipellis, and Ventr fossa atherodon). Whilst area II had high numbers of co-occurring species with
area I (26) and areas III (27), co-occurrence was much reduced between areas I and III (16). Co-occurrence between
these areas individually and area IV was rather similar (area I - 10. II - 14, III - 14) and high in relation to overall
species representation in IV.
Table 3. — (a), The number of grenadier species co-occurring among the areas sampled, together with the total numbers
of species and their proportions of the whole per area and the number of species unique to each; (b), subdivided by
genus.
(a) OVERALL
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
733
a
500
900
Q.
a>
O
1300
1700
2100
10
20
Species frequency per stratum
30
All other genera
a
Ventrifossa
Hymenocephalus
Caelorinchus
— t
40
b
500
900
e
w>
E
21300
1700
40 60
Percent/stratum
100
Caelorinchus
Hymenocephalus
Nezumia
Ventrifossa
Rest
Fig. 5. — (a), Overall species bathymetric distribution by 100 m strata (assuming congruity of occurence across strata),
with the dominant genera indicated; (b), As a percentage distribution, with the number of collections per stratum
indicated down the right axis.
Source : MNHN, Paris
734
N.R. MERRETT & T. IWAMOTO
Looking at the more speciose genera (Table 3b - Caelorinchus , Hymenocephcilus, Nezumia and Ventrifossa),
widespread species were not common. Indeed. 8 species of Caelorinchus occurred solely in area II, where a total of
17 species were encountered and far more than in any other area. The Hymenocephalus were peculiar as a group;
adelscotti and kuronumai, for example, were each confined to a single area and. while adelscotti was hitherto
undescribed (Iwamoto & Merrett, 1997), the record of kuronumai extended its known range from the Japan
region. Hymenocephalus gracilis and H. longibarbis could both have been expected to occur in areas I-III. but
instead co-occurred only in I-II and II-III, respectively (Table 2). The geographic distribution of species of Nezumia
and Ventrifossa were more regular in their occurrence.
Distribution - bathymetric.
Co-occurrence of grenadier species within similar depth strata on the slope was remarkably high, with only two
of the 63 species not represented, at least over part of their depth range, in the upper 1600 m. Assuming congruity
in occurrence across the strata in which the shallowest and deepest representatives of each genus were caught.
Fig. 5a shows the potential species richness within the strata sampled. Thus, grenadier species richness peaked at
37 in both the 700 and 800 m strata, although it did not drop below 20 across the depth range 400-1 100 m and
reduced substantially only deeper than 1400 m (and shallower than 400 m). In depths down to 900 m most of this
diversity was contributed by co-occurring species of Caelorinchus, Hymenocephalus (sensu lato), Ventrifossa and
Nezumia (Fig. 5b). Excluding the single collection in the 200 m stratum from consideration, more than 70% of
the grenadiers occurring across the 300-600 m band were representatives of these four genera, although their
combined influence on grenadier stratum diversity decreased steadily to their disappearance in the 1500 m stratum.
In the two richest strata (700 and 800 m), seven species of Caelorinchus in each level co-occurred with five
(700 m) and six (800 m) each of Hymenocephalus and Ventrifossa, and four of Nezumia (Fig. 5a). This
exceptional degree of co-occurrence may be compared with that commented upon by MERRETT & Marshall
(1981, p. 228) off northwest Africa. In that region 18 out of 26 grenadier species sampled from the depth range
261-6059 m were found to occur in soundings shallower than 1600 m, with high incidence of sympatry only in
the genus Nezumia (4 species over the range 600-1200 m). Nevertheless, the maintenance of such an association of
closely related species prompted MERRETT & MARSHALL (1981) to consider niche availability. While their data
were similarly preliminary in nature to ours, they observed indications of niche separation in the centres of
abundance in depth distribution and trophic relations. The present data from the New Caledonian region indicate far
greater potential competitive overlap among species of the four genera mentioned above and over a narrower depth
range (Fig. 5). Such dietary evidence as was gathered from this collection, however, provides little insight into
trophic separation among this assemblage. Further sampling is necessary to establish whether differences in
bathymetric centres of abundance might alleviate interspecific competition. Meanwhile, the situation found around
New Caledonia, as off northwest Africa, further supports the view of GRASSLE & Sanders ( 1 973) that in general
a low degree of competitive exclusion exists among deep-sea species.
ZOOGEOGRAPHY
Regional distributions of grenadiers.
Many grenadier species are known for their widespread distributions and such an element can be identified
among the slope-dwelling grenadier fauna of New Caledonia. Other rather ill-defined components can also be
recognised to distinguish four groupings among the overall assemblage. Thus there is an endemic component
(species designated 'A' in Table 2). a broad, northern, tropical component ('B'), a southern, temperate component
CC’), and a widespread, global or multiple-ocean-basin component ('D').
The endemic category is small and may not hold up with extensive sampling in other areas. For now. the
species that can be included in this category include Caelorinchus cylindricus, C. melanobranchus, C. sereti.
Hymenocephalus nesaeae , Kumba musorstom, Lucigadus acrolophus, Nezumia aspidentata, N. cliveri , and
Ventrifossa vinolenta.
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
735
Category B comprises the following species found in tropical waters to the west and north, especially tropical
Australia, Indonesia, Philippines, and the East China Sea to southernmost Japan: Caelorinchus acutirostris,
C. anatirostris , C. argentatus, C. cingulatus, C. platorhynchus, C. semaphoreus, Gadomus introniger,
Hymenocephalus kuronumai, H. megalops, H. nascens, Kumba punctulata, Lucigadus microlepis, Malaeocephalus
adustus, Malaeocephalus sp., Pseudonezumia parvipes, and Ventrifossa atherodon.
Category C includes species found in temperate waters to the south and west, particularly in waters off New
Zealand and southern Australia, although three species have ranges extending to southern Africa: Bathygadus
cottoides, Caelorinchus celaenostomus , C. kermadecus, C. mycterismus , C. spathulatus, C. supernasutus,
C. trachycarus, Coryphaenoides dossenus, C. striatums, Hymenocephalus adelscotti, H. longibarbis, Nezumia
coheni, Trachonurus gagates, and Ventrifossa pcixtoni. The ties to the New Zealand fauna (see below) are seen in
the presence of the six species of Caelorinchus, all of which were taken south of New Caledonia on one or more of
three oceanic ridges that form a connection with New Zealand (C. mycterismus and C. trachycarus arc also known
from southeastern Australia). Bathygadus cottoides, Coryphaenoides dossenus and C. striatums arc found across the
Indian Ocean to subtropical to temperate waters off southern Africa.
The last category (D) includes numerous widespread species that are broadly distributed across one or more
oceans. Some of these, when they arc closely studied, may be shown to consist of two or more species, each of
which has a narrower distribution. These species are: Bathygadus spongiceps, Cetonurichthys subinflatus.
Cetonurus crassiceps, Coryphaenoides rudis, Haplomacrourus nudirostris, Hymenocephalus aterrimus, H. gracilis,
Kuronezumia bubonis, Macrosmia phalacra, Malaeocephalus laevis, Malaeocephalus acipenserinus, Nezumia
spinosa , N. propinqua, Pseudonezumia pusilla, Sphagemacrurus pumiliceps, Squalogcidus modificatus ,
Trachonurus sentipellis, Ventrifossa johnboborum, V. macropogon, V. nigrodorsalis.
80
Common lo NC
Total species
NA SA WA NC NSW NZ PI/I K/P O J H EP
Region
Fig. 6. — Global regional comparison of grenadier species frequency of occurrence, indicating ihe frequency of species
common to ihe New Caledonian region (arrowed). (NA, North Atlantic; SA. South Africa; WA. Western Australia:
NC, New Caledonia; NSW. New South Wales; NZ. New Zealand; PI/I. Philippine Is/lndonesia; K/P. Kyushu/Palau
Ridge; O, Okinawa Trough; J. Japan; H, Hawaii; F.P, Eastern Pacific.)
Figure 6 shows the representation of known species of grenadier by region and those in common occurrence
with New Caledonia. There are 58 species found in New South Wales (NSW), of which 25 co-occur with
736
N.R. MERRETT & T. IWAMOTO
New Caledonia (Table 4). (This total in common could be greatly increased with the inclusion of Queensland, but
the data are far from adequate from this poorly sampled region of Australia.) New Zealand, another of the few well-
sampled areas within the New Caledonian region, has 65 species, with only 20 in common. Almost all those
species in common (within our category C, see above) are those with southerly distributions and mostly to the
south of New Caledonia (e.g. Bathygadus cottoides, Caelorinchus celaenostomus, C. kermadecus , C. mycterismus,
C. spatludatus, C. supernasutus, C. trachycarus, Coryphaenoides striatums, C. dossenus, Trachonurus gagates). It
is noteworthy that in the entire eastern Pacific region, however, there are only two species found in common with
New Caledonia. The eastern Pacific grenadier fauna is very distinct as will be seen below. There is a high frequency
of grenadier representation in many regions, often totalling around 60 (overall histograms in Fig. 6), of which the
number co-occurring with New Caledonia (solid histograms - Fig. 6) is variable. As would be expected, those
closest to New Caledonia have the highest number of species in common, although even more remote regions
(e.g. North Atlantic) share the most widespread species (Fig. 6). While the Hawaiian assemblage shares only one
more species in common than does that in the North Atlantic, a greater proportion are common than occur in the
Kyushu/Palau Ridge, Okinawa Trough or Japan areas (Table 4). New South Wales and Western Australia share the
most species in common with New Caledonia (25), although in percentage terms the shared species in Western
Australia are more (67% cf. 57%).
Table 4. — Global regional comparison of grenadier genera and species known distributions, with reference to
comparisons with ihe New Caledonian region and the dominant genera found Ihere. (NA. North Atlantic; SA. South
Africa; WA, Western Australia; NC, New Caledonia; NSW. New South Wales; NZ, New Zealand; PI/I. Philippine
Is/Indonesia; K/P. Kyushu/Palau Ridge; O. Okinawa Trough; J. Japan; H. Hawaii; EP, Eastern Pacific.)
At the generic level, regional comparisons show some interesting trends (Table 4), The total number of
grenadier genera in the Atlantic and the Indo-southwest Pacific, as indicated from sampling in six broad regions, is
rather similar at 20-21 genera. The remainder of the Pacific area, however, have reduced and less consistent
numbers of genera represented. At the specific level, Caelorinchus is the most species-rich genus of grenadier with
well over 100 species, of which many are still undescribed. It is well represented in most regions, although low
species frequency in the Atlantic and eastern Pacific suggest that this is predominantly an Indo-West Pacific group
(Table 4). At least 17 species of Caelorinchus occur around New Caledonia to comprise 28% of the grenadier
assemblage there. Similar high frequencies are apparent in the Australian, New Zealand, Philippines/Indonesian and
Japanese regions. Yet there is a similarly low species representation in the eastern Pacific and Hawaii as occurs in
the Atlantic. Surprisingly, only three species of Cotyphaenoides were represented in the New Caledonian region,
but that number could increase with more sampling on the continental rise below 2000 m. While the
representation by region is varied worldwide, Coryphaenoides contrasts with Caelorinchus by being proportionally
better represented in the North Atlantic and the eastern Pacific (Table 4). On the other hand, the eight species of
Hymenocephalus in New Caledonia is the highest number recorded from any region other than the Philippines and
Source . MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
737
Indonesia (9 spp.) (Table 4). While five species of Nezumia were collected in the New Caledonian region, and this
is not a high representation, three of these were new (Table 4). In percentage terms, this genus contributes more to
the overall assemblage in the North Atlantic, South Africa and central (Hawaii) and eastern Pacific (Fig. 7). The
taxonomy of Ventrifossa is complicated by the variability of the suite of diagnostic characters used. Nevertheless,
six species are recognized from New Caledonia, two of which were recently described as new, and this comprises a
larger group than in any other region surveyed (except Western Australia also with six).
100%
80%
Region
□
Others
Nezumia
Hymenocephalus
Coryphaenoides
Caelorinchus
Fig. 7. — Global regional percentage comparison of grenadier species by main genera occurring in New Caledonia.
(NA, North Atlantic; SA, South Africa; WA, Western Australia; NC, New Caledonia; NSW, New South Wales; NZ,
New Zealand; PI/I, Philippine Is/Indonesia; K/P, Kyushu/Palau Ridge; O, Okinawa Trough; J. Japan; H. Hawaii;
EP. Eastern Pacific.)
While regions northward and eastward of New Zealand have reduced generic representation relative to the
remainder (Table 4), it is there where the proportion of the'grenadicr assemblage contributed by the five genera
Caelorinchus , Coryphaenoides , Hymenocephalus , Nezumia and Ventrifossa together is greatest (Fig. 7). These five
dominant genera together comprise 62% of the total grenadier fauna of the New Caledonian region. The eastern
Pacific is noteworthy for its high representation of Nezumia and Coryphaenoides, more than in any other area, yet
Ventrifossa is absent. The 21 species of Coryphaenoides in the eastern Pacific is 45% of the total grenadier fauna
there (Fig. 7). In no other region of commensurate size is there such a dominance of one genus. Yet the Okinawa
Trough, a much smaller area, holds in the genus Caelorinchus a similar 45% of the total of 35 grenadier species.
The species total of this group of five genera rises through 66% in the Philippine and Indonesian regions,
however, to 78-86% peripherally around the North Pacific, in the Kyushu/Palau Ridge region, the Okinawa
Trough, Japan and the eastern Pacific. Centrally, in Hawaii, the proportion (68%) is more akin to the range in
further westerly and southerly regions. When making such inter-area comparisons, however, it must be stressed
that sampling strategies have not been consistent worldwide. Since grenadier species, like other deep demersal
groups, are constrained between various species-specific bathymetric limits, overall sampling coverage is
738
N.R. MERRETT & T. IWAMOTO
important. While the majority of grenadier species are encountered over slope depths to around 2500 m, complete
coverage is not obtained to at least 4000 m (cf. Merrett, 1992). Thus in this comparison, the apparent low
proportion around New Caledonia of species of the genus Coryphaenoides, generally the deepest living of the five
dominant genera treated, may result more from poor sampling coverage below about 1500 m than from absolute
scarcity. This can be confirmed only by further sampling.
Few data are available from these collections to examine inter-area bathymetric distributions at the generic
level. The most speciose genus, Caelorinchus , offers some evidence (Fig. 8), but such variation that does occur
may be due to sampling limitations (e.g., the lack of using the ORBT in areas III and IV where species of large
adult size like C. kermadecus, C. supernasutus and C. trachycarus may have been present deep but not sampled).
200
]
500
E.
f 800
Q>
Q
■HBBaBnBaHHnSB
555
~
_
1100
— . ' IT
Area IV
Area III
Area II
Area I
Frequency per stratum
Fig. 8. — Vertical distribution of the species of Caelorinchus by area.
POPULATION BIOLOGY
A summary of population biology parameters obtained from the samples is given in Table 5. While not
extensive, it provides preliminary data on the reproductive biology and diet of the more abundant species. In
addition. Figs 10, 1 1,19, 20, 24, 28 and 29 give the length-frequency distributions of some of the more abundant
species, individually commented upon in the species accounts below.
Less can be said about the more infrequently represented species. Results of the examinations are given in
Table 5. Certain preliminary remarks, however, can be made concerning resource partitioning in the co-occurring
genera Caelorinchus, Hymenocephalus ( sensu lato), Ventrifossa and Nezumia. While adult size, based upon
attainment of ovarian maturity, may partition species into preferred prey-size categories, there is little evidence here
to suggest that intrageneric trophic segregation occurs. Most species seem to feed benthopelagically on a mixed
diet. More detailed work using larger samples is needed for any greater clarification.
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
739
Table 5. — Summary of reproductive and feeding data from the dominant species from the New Caledonian region.
(* - macrosmatic males; $ - several generations of ripening ova present; S/NS - sediment present/absent;
G. no. - gonad examination sample size; S. no. - stomach contents examination sample size; E - number of
stomachs found empty.)
M:Fsex Size at Stomach contents material S/NS
Copepod NS
Ostracod. potychaete NS
Copepod, isopod & polychaete S
Crustacean (galathaeid), squid. NS
Copepod. polychaete, fish (Bregmaceros sp ) S
Decapod crustacean, ?myctophid otoliths NS
polychaete, ’squid beak, fish lens.’bryozoan mat S
Decapod crustacean NS
Fish
Crustacean NS
Crustacean (euphausiid) NS
Crab zoea, euphausiid/decapod. isopod & copepod S
Crustacean (copepod) NS
NS
Polychaete, copepod, amphipod, isopod, ophiuroid, S
Crustacean NS
Crustacean (copepod) NS
Polychaete, lamellibranch, pteropod. crustacean, echinoderm. NS
Decapod crustacean, amphipod, fish NS
Decapod crustacean, squid NS
Crustacean ('spider crab'), polychaete NS
NS
SYSTEMATIC ACCOUNT
Family BATHYGAD1DAE
Genus BATHYGADUS Gunther, 1878
Bathygadus cottoides Gunther. 1878
Fig. 9 a
Bathygadus cottoides - IwaMOTO & Merrett, 1997: 479-480 (in part; most spec. B. spongiceps\ see below)
Material examined. — 17 specimens.
Norfolk Ridge. Halipro 2: stn BT 43, 25°41.75'S, 167°1 1.59'E. 1030-1320 m, 15.11.1996: 3 specimens (CAS
90556).
Lord Howe Rise. Halipro 2: stn BT 102, 24°32.97'S. 161°53.65'E, 1060-1130 m. 26.11.1996: 1 specimen
33 mm HL, 133+ mm TL (CAS 90749). - Stn BT 104, 25°23.0rS. 162°35.40'E, 1118-1124 m. 27.11.1996:
1 specimen 38.2 mm HL, 178+ mm TL (CAS 90835). — Stn BT 105, 25°42.93'S. 162°47.98'E. 1200-1218 m.
27.11.1996: 1 specimen 27.2 mm HL (CAS 90845).
Kermadecs. West of Kermadec Trench. 30°32.00'S. 178°22.00'W, 971 m depth over 1100 m soundings. 5.12.1976:
5 of 1 1 specimens 29-38.5 mm HL, 150+ -177+ mm TL (NMNZ P10850). — Esperance Rock: 2 specimens 28-33 mm
HL, 142+ -150+ mm TL (NMNZ P20066).
New Zealand. East coast of North Island. 40°85'S, 177°02'E. 1353-1339 m, 03.1967: I specimen 30.5 mm HL.
186 mm TL (NMNZ P34691, ex NZOI E750). — Challenger Plateau. 39°59.70'S, 167°38.20'E. 1175-1181 m.
740
N.R. MERRETT & T. IWAMOTO
19.03.1984: 2 of 17 specimens 35-39 mm HL, 145+ -214 mm TL (NMNZ P23477). — RV Tangaroa 9406/229, Chatham
Rise. 42°39.65'S. 172°17.66'W, 1505-1524 m. 24.06.1994: 1 specimen 39 mm HL. 260+ mm TL (NMNZ P3475I).
Remarks. — Iwamoto & Merrett (1997) misidentified most of tlicir specimens of B. sp. cf. spongiceps
with B. cottoides in their treatment of the grenadiers of New Caledonia. The second author (TI) has subsequently
examined numerous specimens of B. cottoides in the Museum of New Zealand taken from the Kermadecs (the type
locality) and the New Zealand EEZ. These specimens were all small, with characters similar to those recorded by
IWAMOTO & ANDERSON (1994) from southern Africa. B. cottoides is a small species (probably not more than
about 30 cm TL) unlike B. sp. cf. spongiceps , which attains a considerable size (more than 45 cm TL).
B. cottoides has fewer pectoral fin rays (il0-il4, rarely il5, cf. i 1 5-i 1 8 in B. sp. cf. spongiceps). fewer pyloric
caeca (8-12 cf. 18-27), broader suborbital width (16-19% HL cf 14-17%), and a darker body.
FlG Wales a'iSS: illustrations of (a) Bathygadus cottoides Gunther, 1878 (AMS 1.29745-007) from off New South
Wales, Australia, (b) Bathygadus cf. spongteeps (CAS 98532) from off Lord Howe Rise. Scale = 25 mm.
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
741
Bathygadus sp. cf. spongiceps
Figs 9 b, 10
Bathygadus cottoides - Iwamoto & Merrett, 1997: 479 (in part; 3 spec. From CAS 90556 and 1 spec, from CAS 90835
are B. cottoides, all others listed are B. spongiceps).
Material examined. — 14 specimens.
Norfolk Ridge. Halipro 2: stn BT 41, 25°47.77'S. 1 67° 1 1.52'E. 1317-1383 m. 14.11.1996: 1 specimen 81 mm
HL, 445+ mm TL (BMNH 1997.5.21:31). — Stn BT 43, 25°41.75'S, 1 67°1 1 .59E, 1030-1320 m, 15.11.1996:
3 specimens (CAS 90556). — Stn BT 55. 25°02.85'S, 168°45.42'E, 1098-1480 m, 17.11.1996: 1 specimen 85 mm HL.
450+ mm TL (BMNH 1997.5.21:44), 1 specimen (CAS 90667), 1 specimen 70 mm HL, 415 mm TL (MNHN 1998-245).
— Stn BT 56. 25°02.92’S. 168°46.04'E, 1083-1196 m, 17.11.1996: 1 specimen (NMNZ P.34251). 1 specimen (NMNZ
P.34269). — Stn BT 59, 24°58.35'S, 168°42.31'E, 1312-1520 m, 18.11.1996: 1 specimen 62 mm HL, 282+ mm TL
(CAS 90748). — Stn BT 66, 24°43.92'S, 168°28.78'E, 885-1450 m, 19.11.1996: 1 specimen 73 mm HL, 420 mm TL
(CAS 90685). — Stn BT 75, 24°13.17'S, I67°36.52'E, 1128-1150 m, 21.11.1996: 2 specimens (CAS 90992).
Loyalty Ridge. Halipro 2: stn BT 23, 25°04.95'S, 170°11.7'E 1105-1360 m, 10.11.1996: 1 specimen (CAS
90993), 1 specimen (MNHN 1998-251).
Lord Howe Rise. Halipro 2: stn BT 95, 23°56.86'S, 162°07.57’E, 1224-1233 m, 25.11.1996: 1 specimen 92 mm
HL, 505+ mm TL (MNHN 1998-246).
See Iwamoto & Merrett, 1997: 479 (all specimens in Materials examined except CAS 90556 and CAS 90835).
0-30 31-60 61-90 91-120
H L groups (mm)
Fig. 10. — Length-frequency (HL, head length mm) distribution of Bathygadus cf. spongiceps by sex (n = 47).
REMARKS. — The correct identification of these specimens is uncertain although they are closely similar to
B. spongiceps , which has been recorded from Philippine and Indonesian waters. The main character distinguishing
this species and B. spongiceps is the dentition of the lower jaw, which in B. spongiceps is in a very narrow band,
reduced anterolaterally on the jaw to an irregular double row. In the New Caledonian species the band in that region
is about four irregular series wide. Also, B. spongiceps appears to be much softer bodied and has fewer pyloric
caeca (15-21 cf. 18-27). A closer study of this species is necessary. Bathygadus sp. cf. spongiceps was frequently
taken during the HALIPRO 2 cruise. On the Norfolk Ridge it was captured in 15 trawls, on the Lord Howe Rise six
trawls, on the Loyalty Ridge one trawl. Capture depths ranged between 885 m and 1520 m. The species was never
taken in numbers, however, the most being nine specimens, with one to five most frequent. The large trawl
742
N.R. MERRETT & T. IWAMOTO
deployed by the RV Tangaroa probably accounted for its frequent capture and larger size than in previous
ORSTOM cruises. The largest adult size was attained by females in these samples (Fig. 10). The distribution of
the species appears to extend through New Zealand and Australia. The northern extent is uncertain as there remains
a question as to synonyms.
Genus GADOMUS Regan. 1903
Gadomus introniger Gilbert & Hubbs, 1920
Gadomus introniger - Iwamoto & Merrett, 1997: 480-481. Fig. 4 (Matthew & Hunter Islands).
Material examined. — 4 specimens.
Vanuatu. Musorstom 8: stn CP 990, 18°51.63'S, I68°50.98'E, 980-990 m, 24.09.1994: 1 male specimen 59 9 mm
HL. 339+ mm TL (MNHN 1995-995).
Norfolk Ridge. Halipro 2: stn BT 42, 25°41'S, 167°I1'W, 1132-1160 m, 15.11.1996: I specimen 68 7 mm HL
453 mm TL (CAS 90683).
Indeterminate specimens:
Vanuatu. MUSORSTOM 8: stn CP 1075. 15°53.8I’S. 167°30.42'E, 1100-1191 m, 4.10.1994: I specimen 24 9 mm
HL. 142 mm TL (MNHN 1995-952).
New Caledonia. Bathus 1: stn CP 660, 21°10.48'S. 165°53.19’E, 790 m. 13.03.1993: 1 male specimen 26 8 mm
HL. 151 mm TL (MNHN 1995-810).
Description (not including the two indeterminate specimens). — Counts : ID. 11,9; IP. i 1 8-i 1 9; V. 8; GR-1
(outer) 5+23.
Remarks. — The larger two of our four specimens (CAS 90683 and MNHN 1995-995) agree well with our
concept of G. introniger, as does one of similar size recorded by Iwamoto & MERRETT (1997: 480; MNHN 1997-
656) from the Matthew and Hunter Islands. The recent two specimens, however, have gill rakers that are sharply
tipped (in contrast to bluntly tipped in MNHN 1997-656). The two juveniles we list as indeterminate differ from
G. introniger in having a broader internasal (23-26% HL), larger orbit (24-28% HL), narrower suborbital (12%
HL), longer upper jaw (61% HL). longer barbel (59-76% HL), and longer gill rakers (18-19% HL). The high
pyloric caeca count (60) of Iwamoto & Merrett’s (1997) indeterminate specimen from the Chesterfield and
Bellona Plateau contrasts with the 20 in MNHN 1997-656 and the 24 of the Halipro 2 specimen (counts of the
other specimens were not taken). A good size series would probably help in resolving the identity of these
problematical specimens.
Family MACROURIDAE
Subfamily MACROURINAE
Genus CAELORINCHUS Giorna, 1810
Caelorinchus acutirostris Smith & Radcliffe, 1912
Caelorinclius acutirostris - Iwamoto & Merrett, 1997: 485-486. Fig. 6a (New Caledonia, Philippines. Australia).
Material examined. — 2 specimens.
(MNHN lC995e820)a' BA™US '' ^ ^ 657’ 2'°14-45'S’ l65°5493'E' 490-530 m. 12.03.1993: 1 specimen 25 mm HL
(MNVHNUr99U5-8M70).ORSTOM * ^ ^ "36' ,5°4a62'S' 167°01-60'E- ^8-400 m, 11.10.1994: 1 specimen 43 mm HL
Remarks. This is an upper-slope species of tropical seas known from New Caledonia (22°S), Vanuatu.
nlippines, and Queensland, Australia (09°53'S). It was not taken during Halipro 2. The Bathus 1 capture is
the deepest record.
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
743
Caelorinchus anatirostris Jordan & Gilbert, 1904
Caelorinchus anatirostris - Iwamoto & MERRETT, 1997: 486-489. Fig. 6c (New Caledonia, Chesterfield & Bellona
Plateau).
Material examined. — 7 specimens.
Vanuatu. Musorstom 8: stn CP 1050, 16°39.20'S, 168°01.30'E. 541-577 m. 1.10.1994: 2 specimens 85-86 mm
HL, 330+-331+ mm TL (MNHN 1995-888). — Stn CC 1056, 16°33.1 I S. I67°55.64'E. 602-620 m, 1.10.1994:
I specimen 107 mm HL, 355+ mm TL (MNHN 1995-835), 1 specimen 67 mm HL, 265+ mm TL (MNHN 1997-698). —
Stn CP 1124, 15°57.63'S, 166°38.43'E, 1160-1120 m, 9.10.1994: 1 specimen 120 mm HL. 450+ mm TL (MNHN
1995-878).
New Caledonia. Bathus 1: sin CP 671, 20°51.18’S, I65°28.I7'E, 450-470 m, 14.03.1993: 2 specimens 76-
83 mm HL, 300-300+ mm TL (MNHN 1995-827).
Remarks. — These specimens are slightly larger than ones previously examined from the region and expand
the range of several measurements and counts in the species. The maximum size is now known to exceed 45 cm
TL and 120 mm HL. Other increases in range of counts are seen in scale rows below mid- 1 D. (3-5), lateral line
scales over distance equal to predorsal length (30-34), and number of pyloric caeca (30-32). Changes in the range of
mensural characters include: preoral length (28-37% HL), distance orbit to preopercle (32-39% HL), length upper
jaw (21-31% HL), height ID. (38-53% HL), and posterior nostril (8-11% HL). Additionally, the largest specimen
(MNHN 1995-878, 120 mm HL) has a completely naked nasal fossa, whereas all other specimens have a mostly
scaled nasal fossa.
Caelorinchus argentatus Smith & Radcliffe, 1912
Caelorinchus argentatus - Iwamoto & MERRETT, 1997: 489-490, Fig. 6b (New Caledonia, Australia).
Material examined. — 15 specimens.
Vanuatu. Musorstom 8: stn CP 973, 19°21.30'S, I69°27.03'E, 460-480 m, 22.09.1994: 1 specimen 53 mm HL
(MNHN 1995-980). — Stn CP 974, 1 9°2 1 .51 ’S, 169°28.26'E, 492-520 m, 22.09.1994: 4 specimens 52-77 mm HL
(MNHN 1995-864). — Sin CP 975. 19°23.60'S, 169°28.93'E, 566-536 m, 22.09.1994: 1 specimen 66 mm HL (MNHN
1995-863). — Stn CP 980. 19°21.02’S, 169°25.22'E, 450-433 m. 22.09.1994: 3 specimens 34-93 mm HL (MNHN
1995-833). — Stn CP 982, 19°21.80'S. 169°26.47’E, 408-410 m, 23.09.1994: 1 specimen 54 mm HL (MNHN 1995-
872). — Stn CP 983, 19°2I.61'S, 169°27.76’E, 480-475 m, 23.09.1994: 1 specimen 63 mm HL (MNHN 1995-849). —
Stn CP 984, 1 9° 1 9.62'S , 169°26.43'E. 480-544 m, 23.09.1994: 3 specimens 62-87 mm HL (MNHN 1995-887). —
Stn CP 1136, I5°40.62'S, 167°0I.60'E. 398-400 m. 11.10.1994: 1 specimen 64 mm HL (MNHN 1995-870).
Remarks. — Our specimens of Caelorinchus argentatus from the Vanuatu region differed somewhat from
those taken on earlier cruises. They, in fact, agreed more with the type specimens from the Sulu Sea in having a
slightly shorter snout (39-42% HL cf 44-45% in New Caledonia material), shorter preoral (31-35% cf 36-38%),
slightly wider interorbital space (25-27% cf. 24-25%), slightly larger orbit (27-28% cf. 24-26%), shorter barbel (9-
1 1% cf. 1 1-13%), higher first dorsal fin (47-54% cf. 33-44%), and fewer lateral line scales over a distance equal to
predorsal length (40-43% cf. 47-48%). In addition, the Vanuatu specimens were decidedly darker overall, although
the difference was a matter of degree rather than that of pattern. We previously (IWAMOTO & MERRETT, 1997:
490) attributed the differences found between New Caledonian and Sulu Sea specimens as reflecting individual
variation and small sample size. Our current thinking on this matter has not changed.
Caelorinchus argentatus is apparently a tropical species and was not taken during the Halipro 2 cruise to
regions south of New Caledonia. In previous MUSORSTOM cruises, the species was never laken south of 22°S and
no deeper than 460 m. The specimen from CP 975 extends the maximum depth range to 536-566 in.
Caelorinchus celaenostomus McMillan & Paulin. 1993
Caelorinchus celaenostomus - Iwamoto & MERRETT, 1997: 490-493, Fig. 7 (Norfolk Ridge).
744
N.R. MERRETT & T. IWAMOTO
Material examined. — 9 specimens.
Loyaltv Ridge. Halipro 2: stn BT 3, 23°3 1 . 1 5'S, 169°36.54'E, 685-831 m, 6.11.1996: 1 specimen 172 mm HL.
555+ mm TL (MNHN 1998-237).
Norfolk Ridge. Halipro 2: stn BT 32, 25°19.93'S, 168°56.56'E, 697-1340 m. 12.11.1996: 1 specimen 143 mm
HL, 520 mm TL (CAS 90982), 1 specimen (MNHN 1998-322). — Stn BT 34. 25°23.03'S. 168°56.47'E, 643-1233 m.
12.11.1996: 1 specimen 116 mm HL, 320 mm TL (CAS 90670), 1 specimen 149 mm HL, 515+ mm TL (BMNH
1997.5.21:35). — Sin BT 35, 25°23‘S, 168°56.99'E, 640-740 m. 12.11.1966: 1 specimen (MNHN 1998-235),
1 specimen 116 mm HL, 310 mm TL (CAS 90984), 1 specimen 127 mm HL. 410+ mm TL (BMNH 1997.5.21:39). — Stn
BT 91, 23°13.86'S, 167°57.19'E, 650-660 m. 23.11.1996: I specimen 188 mm HL, 710 mm TL (BMNH 1997.5.21:63).
Remarks. — The three CAS specimens were previously recorded by Iwamoto & Merrett ( 1997: 490-493).
Caelorinchus celaenostoinus attains at least 83 cm TL. it was taken occasionally off the Norfolk Ridge and once
on the Loyalty Ridge. It is distributed in New Zealand waters north of Cook Strait (about 41°30'S), and although
found off the Challenger Plateau, it does not occur on the Chatham Rise (McMillan & Paulin. 1993). The
northern record is from HALIPRO 2 station BT 91 at latitude 23°S. Depth range 606-1383 m.
Caelorinchus cingulatus Gilbert & Hubbs, 1920
Caelorinchus cingulatus - Iwamoto & Merrett, 1997: 493-495, Fig. 8 (New Caledonia, Loyalty Islands, Wallis & Futuna
Islands, Australia).
Material examined. — 5 specimens.
Vanuatu. MUSORSTOM 8: stn CP 973, I9°21.30’S, I69°27.03'E, 460-480 m, 22.09.1994: 1 specimen 55 mm HL
(MNHN 1997-697). — Stn CP 1027, I7°53.05'S, I68°39.35’E, 550-571 m, 28.09.1994: 2 specimens 22-49 mm HL
(MNHN 1995-847). — Stn CP 1049. 16°39.43'S, 168°02.97'E. 469-525 m, 1.10.1994: I specimen 55 mm HL (MNHN
1995-852).
New Caledonia. Bathus 1: stn CP 702, 20°55.97'S. 165°34.67'E. 591-660 m, 18.03.1993: 1 specimen 54 mm HL
(MNHN 1995-815).
Remarks. — This species was not collected during Halipro 2 and presumably is more tropical in distribution
than many taken during that cruise. It is now known from off New Caledonia, Loyalty Islands. Vanuatu, Wallis
and Futuna Islands, and the east coast of Australia, where it has been recorded as far south as 28°S near the
Queensland/New South Wales border.
Caelorinchus cylindricus Iwamoto & Merrett, 1997
This species is known only from the holotype taken on Seamount B on the Norfolk Ridge southeast of New
Caledonia in 550-920 m. No specimens were collected during Halipro 2, although Seamount B was sampled
during that cruise.
Caelorinchus kermadecus Jordan & Gilbert, 1904
Fig. II
Caelorinchus kermadecus - Iwamoto & MERRETT. 1997: 496-499, Fig. 10 (Norfolk and Loyalty Ridges). — Grandperrin
et al, 1997: 117 (New Caledonia; list).
Material examined. — 45 specimens.
Norfolk Ridge. Halipro 2: stn BT 02, 23°17.54'S, 167°51.36'E, 1025-1049 m, 5.11.1996: 1 specimen 94 mm
HL, 305+ mm TL (CAS 90959), 1 specimen 101 mm HL (CAS 90690), 3 specimens 103-163 mm HL. 325+-560+ mm TL
(MNHN 1998-2234). — Stn BT 38. 24°55.69’S, 168°19.9'E. 856-1450 m, 13.11.1996: 1 specimen 136 mm HL. 410+
mm TL (MNHN 1998-323). — Stn BT 41 , 25°47.77'S. 167°1I.52’E, 1317-1383 m, 14.11.1996: 1 specimen 146 mm HL,
525+ mm TL (BMNH 1997.5.21:61). — Stn BT 42, 34°29'S. 167°25.08'E, 1132-1160 m, 15.11.1996: 1 specimen (CAS
90985). — Stn BT 64. 24°41.54'S. 168°40.88'E, 800-1240 m. 19.11.1996: 1 specimen 128 mm HL 462 mm TL (CAS
91028). Stn BT 66, 24°43.92'S, 168°28.78'E, 885-1450 m, 19.11.1996: 1 specimen 163 mm HL. 500+ mm TL (CAS
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
745
uncat.). Stn BT 69, 24°40.34'S, 168°08.9'W, 943-1080 m, 20.1 1.1996: 2 specimens (MNHN 1998-236). — Stn BT
75, 24°13.17'S, 167°36.52'E, 1128-1150 m. 21.11.1996: 1 specimen 77 mm HL, 200+ mm TL (CAS 90615). — Stn BT
76. 24°05.43'S, 167°34.29'E, 936-940 m, 21.11.1996: 3 specimens (CAS 90743), 16 specimens (NMNZ P 34019)
Loyalty Ridge. Halipro 2: stn BT 14, 24°18.66'S, 170°00.6'E, 909-1160 m. 8.11.1996: 1 specimen 119 mm HL.
412+ mm TL (CAS 90985), 1 specimen (MNHN 1998-234). — Stn BT 15. 24°36.28'S, I70°03.85'E 900-1005 m
8.11.1996: 4 specimens 121-170 mm HL, 430+-595+ mm TL (MNHN 1998-316). — Stn BT 18, 24°34 42'S 1 70° 1 62T
1050-1290 m, 9.1 1.1996: 1 specimen 120 mm HL, 390+ mm TL (CAS 90986).
Lord Howe Rise. Halipro 2: sin BT 97, 24°00.16'S, 161°46.22'E, 964-1031 m. 25.11.1996: 3 specimens (NMNZ
P.34016), 3 specimens (NMNZ P.34017).
Remarks. — Caelorinchus kermadecus was one of the most often-captured species of grenadier during the
Halipro 2 cruise (Fig. 11), and it was one that caused considerable identification problems because of (he amount
of variation seen in snout length and scale spinulation. A few specimens were confused with the closely similar
C. mycterismus , which has a generally longer, more-upturned snout (45-55% HL cf. 41-47% in C. kermadecus).
shorter first dorsal fin height (28-36% HL cf. 36-43%), shorter upper jaw (15-25% cf. 25-30%), and shorter
postorbital length (25-30% cf. 34-36%). The orbit goes into the postorbital length about 1.2- 1.5 times in
C. kermadecus but only 0.9- 1.3 times in C. mycterismus. Also, the anal fin in C. kermadecus is dark posteriorly,
whereas in C. mycterismus it is usually paler over the posterior third. The species is one of the members of the
genus attaining large adult size (58+ cm TL).
40
61-90 91-120 121-150 151-180 181-210
H L groups (mm)
Fig. 11. — Length-frequency (HL, head length mm) distribution of Caelorinchus kermadecus by sex (n = 77).
Caelorinchus melanobranchus Iwamoto & Merrett, 1997
No specimen oi this interesting species of small adult size was collected during recent MUSORSTOM cruises.
Caelorinchus mycterismus McMillan & Paulin, 1993
Fig. 12
Caelorinchus mycterismus McMillan & Paulin, 1993: 827-829, Fig. 6 (holotype Challenger Plateau. New Zealand, 846-
875 m: 30 paratypes).
746
N.R. MERRETT & T. IWAMOTO
Material examined. — l specimen.
Loyalty Ridge. Halipro 2: stn BT 23. 25°04.95'S, 1 70° 1 1 .7'E, 1105-1360 m, 10.11.1996: 1 specimen 152 mm
HL, 455+ mm TL (CAS 90994).
Diagnosis. — Underside of head completely scaled; nasal fossa scaled; snout 45-5 1 % HL; anterolateral margin
of snout incompletely supported by bone; body scales with broad spinules in about 5-7 more or less parallel rows,
the median row highest, formed of 4-7 closely adpressed spinules. flanked above and below by 3-6 shorter, lower
rows, some of which consist of only I or 2 small, narrow spinules at anterior end of exposed field, large spinules
with high transverse buttresses interconnected in some to form transverse ridges running perpendicular to spinule
rows; light organ lacking external fossa; anal fin dusky to blackish anteriorly, paler at posterior end. Scales below
2D. 5-7; height ID. 28-36% HL.
Fig. 12. — Caeloriiicluis mycterismus McMillan & Paulin, 1993. Lateral view of head and trunk of holotype (NMNZ
P.23003). From McMillan & Paulin, 1993, fig. 6. Used with permission of the ASIH.
Remarks. — This species, which attains large adult size (49+ cm TL), is reported to he abundant off New
Zealand, especially around North Island and the Challenger Plateau, in depths of 833-1 150 m (McMillan &
Paulin, 1993: 829). The Halipro 2 specimen is the northernmost record of the species. Caelorinchus
mycterismus is most likely to be confused with C. kermadecus\ see Remarks under that species for a comparison.
Caelorinchus parallelus Gunther, 1 877
Caelorinchus parallelus - IWAMOTO & MERRETT. 1997: 500-502, Fig. 12 (Chesterfield & Bellona Plateau. Japan). —
Grandperrin et a!., 1997: 1 17 (New Caledonia; list).
Material examined. — 5 specimens.
Vanuatu. Musorstom 8: stn CC 996, 18°52.4I'S. 168°55.73'E. 764-786 m, 24.09.1994: 2 specimens tMNHN
1997-669).
Norfolk Ridge. Halipro 2: sin BT 2, 23°17.54'S, 167°51.36'E. 1025-1049 m, 5.11.1996: 1 specimen 65 mm HL.
191+ mm TL (CAS 204499). — Stn BT 90. 23°16.98'S. I68°08.38'E, 975-987 m. 23.11.1996: I specimen 109 mm HL,
360+ mm TL (BMNH 1997.5.21.41).
Lord Howe Rise. Halipro 2: stn BT 104, 25°23,01'S, 162°35.4E. 1118-1124 m, 27.11.1996: I specimen
103 mm HL. 349+ mm TL (CAS 90987).
Remarks. — We had previously recorded the species from two specimens collected off the Chesterfield and
Bellona Plateau. These additional specimens from Vanuatu, the Norfolk Ridge, and the Lord Howe Rise extend the
range latitudinally and to the east. The small HALIPRO 2 specimen from the Norfolk Ridge was collected along
with several larger specimens of C. kermadecus and was initially confused with that species. The absence of scales
Source : MNHN. Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
747
on the nasal fossa, the relatively extensive naked areas along the suborbital shelf, the wider spacing between scales
on the underside of the head, the crestlike rows of spinules on the dorsal surface of the snout and between the
orbits, and the paler fins of that specimen differentiate it from C. kermadecus.
Caelorinchus plalorhynchus Smith & Radcliffe, 1912
Caelorinchus plalorhynchus - Iwamoto & Merrett, 1997: 502-503, Fig. 13 (Loyalty Islands).
Material examined. — l specimen.
Loyalty Ridge. Halipro 2: stn BT 3, 23°31.15’S, 169°36.54'E, 685-831 m, 6.11.1996: 1 specimen 70 mm HL.
220+ mm TL (MNHN 1998-233).
Remarks. — A single juvenile of this rarely-caught species was taken during the Halipro 2 cruise.
Caelorinchus semaphoreus Iwamoto & Merrett, 1997
Caelorinchus semaphoreus Iwamoto & Merrett, 1997: 503-505, Fig. 14 (New Caledonia and Qld. Australia; 420-445 m).
This species is known from 30 specimens collected at three sites off Queensland, Australia, and one specimen
taken off New Caledonia. It was not represented in current collections.
Caelorinchus sereti Iwamoto & Merrett, 1997
Caelorinchus sereti Iwamoto & Merrett, 1997: 505-507 (New Caledonia. Loyally Islands. Chesterfield & Bellona
Plateau).
Material examined, — 29 specimens.
Vanuatu. Musorstom 8: stn CP 992, 18°52.34'S. I68°55.16'E. 775-748 m. 24.09.1994: I specimen 124 mm HL
(MNHN 1995-831). —Sin CP 994. I8°47.72'S, 168°56.I7'E, 649-641 m, 24.09.1994: 4 specimens 27-97 mm HL
(MNHN 1995-841), I specimen 104 mm HL (MNHN 1995-845). — Stn CC 996, 18°52.41'S. 168°55.73'E. 764-786 m.
24.09.1994: 1 specimen 116 mm HL (MNHN 1995-832), I specimen 122 mm HL (MNHN 1995-840). 1 specimen
1 12 mm HL (MNHN 1995-844), 1 specimen 94 mm HL (MNHN 1995-861), I specimen 125 mm HL (MNHN 1995-879).
I specimen 89 mm HL (MNHN 1995-884). — Stn CP 1007. 18°51.97'S. 168°55.92’E, 720-830 m, 25.09.1994:
I specimen 115 mm HL (MNHN 1995-883) and 2 specimens 44-94 mm HL (MNHN 1995-965). — Sin CP 1074.
15°48.42'S, 1 67°24.27'E, 775-798 m. 4.10.1994: 1 specimen 141 mm HL (MNHN 1995-876). — Stn CP 1124.
15°01.7'S. 1 66°56.5 1 E. 532-599 m, 9.10.1994: 1 specimen 19 mm HL (MNHN 1997-693).
New Caledonia. Bathus 1: stn CP 658, 21°13.45'S, 165°55.12'E. 515-580 m. 12.03.1993: 3 specimens 16-
83 mm HL (MNHN 1995-809). — Sin CP 671. 20°51.I8'S, 165°28.17'E, 450-470 m, 14.03.1993: 2 specimens 18-
83 mm HL (MNHN 1997-690).
Stn unknown: 5 specimens 57.2-95.1 mm HL, 165+-325+ mm TL (MNHN 1995-825). — Sin unknown: 2 specimens
53.5-88 mm HL (MNHN 1995-988).
Remarks. — Caelorinchus sereti was one of the more frequently encountered species, having been captured in
14 trawls. The smallest specimens (less than about 60 mm) we examined were juveniles; they had three saddle
marks posteriorly, the first behind and below the origin of the second dorsal fin. Their snout was more acute,
sharper pointed, and relatively longer than in adults; the scale covering was also thinner and more scattered,
especially on the snout surfaces, and the nasal fossa was for the most part naked (scaled in adults).
The species has not been taken south of New Caledonia and so far appears to be confined to the immediate
vicinity of New Caledonia, Loyalty Islands, Vanuatu, and the Chesterfield and Bellona Plateau, in depths of 412-
830 m.
Caelorinchus shcherbachevi Iwamoto & Merrett, 1997
Caelorinchus shcherbachevi Iwamoto & Merrett. 1997: 507-509. Fig. 13b (Loyally Ridge).
Caelorinchus sp. n. - Grandperrin el al., 1997: 117 (New Caledonia; list).
748
N.R. MERRETT & T. IWAMOTO
Material examined. — 2 specimens.
Loyalty Ridge. Halipro 2: sin BT 13. 24°18.03'S, 169°57.5'E, 805-835 m, 8.1 1. 1996: I specimen 122 mm HL
435+ mm TL (paratype, CAS 90995), 1 specimen 132 mm HL. 550 mm XL (MNHN 1998-318).
REMARKS. — This rarely-caught species is known from only four specimens taken in two trawls on the
Loyalty Ridge. One of the HALIPRO 2 specimens is a paratype. previously recorded by IWAMOTO & MERRETT
(1997: 507).
Caelorinchus spathulatus McMillan & Paulin, 1993
Caelorinchus spatliulata McMillan & Paulin, 1993: 832-833, Fig. 9 (New Zealand).
Caelorinchus spathulatus - IWAMOTO & Merrett, 1997: 509-511, Fig. 16 (New Caledonia, Chesterfield and Bellona
Plateau. New Zealand, Australia [Qld.] in 550-825 m).
No specimen of this species was collected during recent cruises.
Caelorinchus supernasutus McMillan & Paulin. 1993
Fig. 13
Caelorinchus supernasutus McMillan & Paulin, 1993: 833-835, Fig. 10 (holotype NMNZ P.22641. Bay of Plenty.
New Zealand. 606-673 m). — Grandperrin et at., 1997: 1 17 ( New Caledonia; listed).
Material examined. — l specimen.
Norfolk Ridge. Halipro 2: stn BT 35, 25°23'S, I68°56.99'E, 640-740 m. 12.11.1996: 1 specimen 136 mm HL.
440 mm TL (MNHN 1998-231).
Diagnosis. — Snout extremely long, 50-55% HL; underside of head completely scaled; nasal fossa scaled;
anterolateral snout margin completely supported by bone; a small, prominent dermal window' of light organ before
anus extending anteriorly to about midway between anal fin origin and pelvic fin insertion; body scales with 5-8
divergent rows of spinules; anal fin dusky, paler along base.
Fig. 13. — Caelorinchus supernasutus McMillan & Paulin. 1993. Lateral view of head and trunk of holotype (NMNZ
P.22641). From McMillan & Paulin, 1993, fig. 10. Used with permission of the ASIH.
Description (one specimen). — Counts : ID. 11,8; P. 115; scales below ID. 8, mid-base ID. 5, 2D. 7.5.
lat- line 55. Measurements (in mm, percent HL in parentheses): snout 71 (52); preoral 67 (49); interob. 22 (16);
orb. 32 (24); suborb. 15 (11); postorb. 33 (24); orb.-preop. 34.5 (25); up. jaw 28 (21); barbel II (8)-
isthm.-A. 67 (49); pre-A. 193 (142); len. V. 31 (23).
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
749
Scales small, armed with 5-8 (or more) short, slightly divergent or subparallel, comblike rows of erect
spinules.
Size. — To at least 64 cm TL.
DISTRIBUTION. — Restricted to northern New Zealand (north of 37°S) and the Norfolk Ridge (to about 25°S),
in about 640-750 m.
Remarks. See the original description for details and illustration. Caelorinchus supemasutus is apparently
common off North Island, New Zealand (McMillan & Paulin, 1993: 835) but is probably marginal to the
New Caledonia EEZ, where the current specimen is first recorded from the region. The species attains relatively
large adult size ( ca 64 cm TL). It is similar to C. sereti in many fundamental respects, and will key out to that
species in IWAMOTO & Merrett's (1997: 484-485) key to the species of Caelorinchus from New Caledonia.
However, C. supemasutus has among other characters a considerably longer snout (50-55% HL cf. 38-46% in C.
sereti) and body scales with spinules more erect and forming more comblike ridges, the ridge rows less divergent
than in C. sereti.
Caelorinchus trachycarus Iwamoto, McMillan & Shcherbachev, 1999
Fig. 14
Caelorinchus sp. n. Iwamoto & Merrett - Grandperrin el al., 1997: 1 17 (New Caledonia; list).
Caelorinchus trachycarus Iwamoto, McMillan & Shcherbachev. 1999: 49-54 (New Zealand. Norfolk Ridge, Australia, in
622-1350 m).
Caelorinchus trachycarus - Iwamoto & Williams, 1999: 166-167 (Southern Australia to New Zealand in 1015-1234 m).
Material examined. — l specimen.
Norfolk Ridge. Halipro 2: stn BT 77. 24°II.38'S, I67°45.52'E, 1349-1350 m, 21.11.1996: 1 paratype 107 mm
HL, 41 2 mm TL (CAS 90983).
Fig. 14. — Caelorinchus trachycarus Iwamoto, McMillan & Shcherbachev, 1999. Lateral view of head and trunk of
paratype (AMS 1.18712-015; 350+ mm TL) from off southern Australia. Scale = 25 mm. From Iwamoto & Williams,
1999, fig. 24. Used with permission of (he California Academy of Sciences.
750
N.R. MERRETT & T. IWAMOTO
DIAGNOSIS. — Underside of head scaled; nasal fossa naked; anterolateral snout margin incompletely supported
by bone; scales of head ridges markedly spiny and coarsely modified; body scales with numerous (8 or more)
parallel spinule rows, the median row complete to edge of scale and slightly larger than incomplete, small, short,
lateral rows of spinules; light organ lacking external fossa; anus immediately anterior to anal fin; fins black;
overall color dark, generally swarthy with purplish tinge.
Description (of one specimen). — Counts: ID. 11,8; P. i , 1 7 - i , 1 8 ; scales below ID. 5, mid-base ID. 4.5. 2D.
5.5, lat. line 30. Measurements (in mm, percent HL in parentheses): snout 47 (44); preoral 43 (40);
internasal 22 (21); interorb. 26 (24); orb. 28 (26); suborb. 19 (17); postorb. 33 (31); orb.-prcop. 35 (32);
up. jaw 24 (22); barbel 7 (7); V.-A. 42 (39); isthm.-A. 68 (64); pre-A. 167 (156); body depth 55 (51);
ID. -2D. interspace 26 (24); ht. ID. 47 (44); len. P. 38 (36); ten. V. 38 (36).
SIZE. — To about 50 cm TL.
DISTRIBUTION. — Common off New Zealand and the southern coast of Australia, from New South Wales to
Western Australia.
Remarks. — The single Halipro 2 specimen from the Norfolk Ridge represents the northernmost record of
the species. In the New Caledonian region the species most closely resembles C. kermadecus, but it can be readily
distinguished from that species by its naked nasal fossa, much darker overall color with purplish tinge in unsealed
areas, wholly black fins, and much coarser and more spiny head ridges, and the presence of many more parallel and
lower spinule rows on body scales.
Genus CETONURICHTHYS Sazonov & Shcherbachev, 1982
Head inflated, broad (interorbital width about one-third HL), soft, lacking stout ridges, covered with small, fine
scales, including lower jaw and over lower branchiostegal rays. Snout high, tipped with two small tubercular
scales; suborbital region deep, about 17-20% HL; jaws short (24-28% HL), inferior. Pcriproct small; anus about
midway between anal and pelvic fin bases; no discernible light organ. Pelvic fins posterior to vertical through
origin of first dorsal and pectoral fin bases. Body scales small, about 11-13 below ID. origin; lateral line
interrupted into a series of dashes. Only one species.
Cetonurichthys subinflatus Sazonov & Shcherbachev, 1982
Fig. 15
Cetonurichthys subinflatus Sazonov & Shcherbachev, 1982: 9, Fig. 3 (holotype, ZMMGU P.15296, and two paralypes,
ZMMGU P.15297, northwest coast of Australia. 1600-1700 m). — Grandperrin et at., 1997: 1 17 (New Caledonia;
list).
Material examined. — 1 specimen.
Norfolk Ridge. Halipro 2: stn 68. 24°48.20'S, I68°17.12'E, 1316-1564 m. 19.11.1966; 1 specimen 71.2 mm
HL. 372+ mm TL (MNHN 1998-239).
DIAGNOSIS. — Pelvic fin rays 8 or 9; P. i 1 6-i 1 8; scale rows below ID. 11-14. below 2D. 10-13; interorbital
29-33% HL; second spinous dorsal ray about 70-98% HL.
Description (of Vanuatu specimen). — Counts: ID. 11,9; P. i 1 6/i 1 6; V. 8/9; scale rows ID. about 13 or 14,
below mid- ID. 8.5, below 2D, 10, lateral line over pre-ID. length about 50.
Measurements: Total length 372 mm (tip partly missing), HL 71.2 mm. The following in millimeters
followed in parentheses by percent HL: snout 28 (39); preoral 23 (32); internasal 21 (29); interorb. 21 (29); orb.
16 (22); suborb. 14.3 (20); postorb. 28 (39); orb.-preop. 27.5 (39); up.jaw 19 (27); barbel 7 (10); body depth 62
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
751
(87); pre-A. 115 (162); pre-vent 104 (146); V.-A. 24 (34); isthm.-A. 52 (73); ht. ID. 70+ (98); len. P. 41 (58);
len. V. 40 (56).
See Sazonov and Shcherbachev (1982) for a good description of this species and diagnosis of the genus.
Fig. 15. — Cetonurichthys subinflatus Sazonov & Shcherbachev. 1982. Diagrammatic views of paratype (Z1L P.15297)
from off northwestern Australia: (a) lateral view of head and trunk; (a1) dorsal view of snout; ( b ) lateral view of entire
fish. Scale for (a) and (a') = 25 mm.
General features of fish seen in Fig. 15. Head large, rounded contours, without stout or sharp ridges; snout
broad, moderately pointed, tipped with two distinctly separated tubercular scales; mouth small, subinferior, rictus
restricted by lip folds at posterior angle. Periproct region moderately broad, but no external indication of light
organ; anus located about equidistant from insertion of pelvic fins and origin of anal fin. No large naked papillae or
free neuromasts, nor connecting naked lines along suborbital shelf. Neuromasts under head small, black.
752
N.R. MERRETT & T. IWAMOTO
Entire underside of head scaled, including fully scaled mandibular ramus, interopercle, preopercle, subopercle,
and opercle. Extent of squamation similar to that in Cetonurus, with no naked margins. Gular and branchiostegal
membranes heavily covered with small scales. Lateral line incomplete, consisting of a series of dashed grooves.
Scales over most of head and body covered with spinules that are very slender, conical, upright, bristly, with
recurved tips, not in any distinct or readily discernible arrangement, but perhaps in somewhat parallel rows. Scales
along second dorsal fin slightly enlarged, but not strikingly so as in Cetonurus and Trachonurus. Premaxillary
teeth band broad, short; greatest width about one-third length of band. Mandibular band similarly broad or even
broader, and short but tapered posteriorly (greatest width about twice into length).
Fins well developed; first dorsal high, the second spinous ray armed along leading edge with small, sharp,
spaced serrations. Outer pelvic ray weak, extending posteriorly to about fifth anal ray. Pelvic fins posteriorly
placed on abdomen, about under first dorsal fin; distance between pelvic and anal fins short; anal fin origin behind
vertical through hind margin of first dorsal.
Size. — To about 37 cm TL.
Distribution. — Known previously only from the type series (3 specimens) taken in 1600-1700 m off the
northwest coast of Australia and two specimens recorded by Iwamoto & Williams (1999) from the same region
in 1460-1500 m. The current specimen extends the range into the southwestern Pacific.
Remarks. — The Halipro 2 specimen is only the sixth known of the species. More trawling at depths
greater than 1500 m along the continental slopes ol Australia, New Caledonia, and Indonesia may capture
additional specimens that would help in understanding the distribution and biology of the fish.
Genus CETONURUS Gunther, 1878
Head large, globose, broad and deep, interorbital space 37-53% HL, suborbital deep. 22-31% HL. Mouth
interior, 25-38% HL; chin barbel small. Head fully scaled, including lowermost rays of branchiostegals and often
over gular membrane; scales along anterior portion of second dorsal fin enlarged, with larger, coarser spinules.
Course ol lateral line usually marked by free neuromasts, generally lacking grooved scales. Pelvic fin with 8-13
rays, origin usually slightly anterior to origin of first dorsal and pectoral fins. Periproct broad, immediately
anterior to anal fin; anus in middle of periproct and thus removed slightly from anal fin. (Characters mostly from
Sazonov & Shcherbachev. 1985.) Two species. Circumglobal in middle to low latitudes, but absent along
Pacific coast of the Americas.
Cetonurus crassiceps (Gunther, 1878)
Fig. 16
Coryphaenoides crassiceps Gunther, 1878: 25 (n. of Kermadec I., Challenger Stn 170, 520 fathoms and Stn 171
600 fathoms).
Macrurus (Cetonurus I crassiceps - GUNTHER, 1887: 143.
Cetonurus crassiceps - Paulin e, al. , 1989: 129 (in key). - Grandperrin et al, 1997: 1 17 (New Caledonia; list).
Material examined. — 17 specimens
mmSSSSS25’ ,5#5763'S- l66°38'43'E- n6°-1220 "• l010 l"4: 1 ma'e *■— 746
mmIH°t'aS ‘SP BT ,2’ 24°2102's' 170°01.02'E, 1140-1188 m, 8.11.1996: 2 specimens 69-82
(CAS 90602)' ' sPecimcn <CAS 90694), 1 specimen (NMNZ
(MNHN 1998-244)BT 6' 24 33 85 S' ?° 06 47 E‘ 1019-'200 8.11.1996: 1 specimen 100 mm HL. 405+ mm TL
275+mm1T,Rrr8AeR Q07LJ^°,2: S‘n BT 32' 2fl9'S’ 168°56'W' 69?-|34° m- '2-11.1996: I specimen 51 mm HL.
-75+ mm TL (CAS 90744). 1 specimen (MNHM 1998-252), 1 specimen (MNHN 1998-253). 2 specimens (NMNZ
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
753
P.34270). — Sin BT 44. 25°43'S. I67°I2'W, 1033-1200 m, 15.1 1.1996: 2 specimens 79-82 mm HL. 320+-350+ mm TL
(CAS 90603). — Sin BT 95, 23°56.86'S, 162°07.57'E. 1224-1233 m. 25.11.1996: 1 specimen (MNHN 1998-255).
Lord Howe Rise. Halipro 2: stn BT 104, 25°23'S, 162°5'W, 1118-1124 m, 27.11.1996: 2 specimens 44-48.6 mm
HL, 190+-237+ mm TL (CAS 90673).
DIAGNOSIS. — Scales on gular and branchiostegal membranes few. isolated; orbit 17-23% HL; scales below
ID. 11-14 (usually 12-13), below 2D. 9-13 (usually 10-11).
DESCRIPTION. — Counts : ID. 11,8-10; P. i 15-i 1 8; V. 10-11; GR-I (inner series) 2+11. Measurements (in
percent HL): snout 36-38; prcoral 31-37; orb. 18-26; suborb. 23-28; inlerorb. 43-48; postorb. 45-51; orb. -preop.
45-51; up.jaw 30-33.
Fig. 16. — Cetonurus crassiceps (Gunther, 1878). A 13 inch [about 32.5 cm] specimen taken north of the Kermadec
Islands. H.M.S. Challenger , stn 170. in 520 fm (951 m]. From G0NTHER, 1887, pi. XXXVII.
REMARKS. — Cetonurus crassiceps was taken in considerable numbers in several trawls during the Halipro 2
cruise. Again, the large trawl used during that cruise probably accounted for the high numbers taken, as opposed to
none on cruises previous to Musorstom 8 and Halipro 2.
On casual observation, this species can be easily mistaken for Squalogadus modificatus , with which it is
sometimes taken, owing to the similarity of their large, globose heads. C. crassiceps can be immediately
distinguished by its high, short, first dorsal fin followed by a long, low second dorsal. Squalogadus, in contrast,
has only one long, low dorsal fin. SAZONOV & SHCHERBACHEV (1985) recorded Cetonurus glohiceps , the second
member of the genus, from New Zealand, but we found no specimen corresponding to that species in our New
Caledonia collections. The two authors provided data in confirmation of the differences between the two species.
Our specimens agree well with their description of C. crassiceps , especially in terms of the smaller orbit; rougher,
somewhat larger scales; smaller, more-rudimentary chin barbel; naked margin along the anal fin; and general head
shape. We found that scale-row counts below the first dorsal fin were too difficult to make to be useful for us.
754
N.R. MERRETT & T, IWAMOTO
Also, the distribution of teeth in the jaws, which they considered distinguishing, did not appear to be so with our
New Caledonian specimens and three C. globiceps from Walters Shoals in the southwestern Indian Ocean (CAS
66484).
Genus CORYPHAENOIDES Gunner. 1765
Coryphaenoides dossenus McMillan, 1999
Fig. 17
Coryphaenoides (Coryphaenoides) sp. indet. - IWAMOTO, 1986: 355 (Mozambique, 696-960 m).
Coryphaenoides ( Coryphaenoides ) rudis - McCann & McKnight. 1980: 35, Figs 16-18 (in part; New Zealand)
7oQ7en^M Sp- n ; PAULIN,e' aL' 1989: l29' Rg' 56 30b key to New Zealand fishes). - Grandperrin ,, ah,
1997: 117 (New Caledonia; list).
Coryphaenoides sp. - Okamura in Amaoka el ah, 1990; 184. Fig. 126 (New Zealand, 100-763 m). — Gomon cl al
1994: 355, Fig. 315 (Australia).
Coryphaenoides sp. 1 - SHCHERBACHEV & Iwamoto, 1995: 309-312, Fig. 8 (synonymy; distr.. 696-1600 m)
Coryphaenoides dossenus McMillan, 1999: 482-486, Figs 1-2.
specimen 50 mm HL
Material examined. — 3 specimens.
Vanuatu. Musorstom 8: stn CP 1129, I6°00.73'S, 166°39.94'E, 1014-1050 m 10 10 1995
(MNHN 1995-986).
Norfolk Ridge. Halipro 2: stn BT 38. 24°55.69'S. 168°19.9'E. 856-1450 m, 13.11.1996: I specimen (NMNZ
?;C97l:5.2T:54" 23°4a°2'S’ l68°03-,5'E- ^-HOO m. 23.11.1996: 1 specimen 156 mm HL^femalc- tBMNII
DIAGNOSIS. Head profile low, followed by highly arched nape in adults; pelvic fin rays 8, rarely 9; snout
low, rounded, barely protruding in adults, preoral length 7-11% HL; upper jaw long, 40-45% HL. extending well
past vertical through hind end of orbit; barbel long, 21-34% HL; inner gill rakers of first arch 11-13 total;
premaxillary teeth in broad band, mandibular teeth in several rows; body scales covered with small, closely packed
sptnules in parallel to slightly convergent rows.
Size. — To at least 85 cm TL.
DISTRIBUTION. - From the southeastern Atlantic to southern Africa, Indian Ocean. Australia, New Zealand
and the New Caledonian region. Depth range about 700-1600 m.
Remarks — This species attains large adult size. It is quite abundant off the New South Wales coast of
Australia, and extensive collections are housed in the Australian Museum in Sydney. It has already been well
described numerous times (e.g., GOMON el al ., 1994; SHCHERBACHEV & Iwamoto. 1995) and well illustrated
(mcludmg color photographs), but it awaited a scientific name by Peter McMillan. Only a single juvenile was
collected oil \ anuatu, but two large adults were captured off the Norfolk Ridge during Halipro 2
Coryphaenoides rudis Gunther, 1878
Fig. 18
Macrurus ( Macrurus ) rudis - G0NTHER, 1887: 131, pi. 27.
Nemaionurus macrocephalus Maul, 1951: 17 (Madeira).
Macrourus paradoxus Smith & Radcliffe in Radcliffe 1912- 1 is l if, nl 7S Pin i n i ,•
IWAMOTO & Sazonov, 1988: 52-53 (se. Pacific) ’ g' ‘ (eaStem Palawan' PhlllPP'nes). -
Material examined. — 3 specimens.
(MNLHNdunHcat7 RiSC' HAUPR° * ^ BT 25°29 08'S- 1 63° 1 3. 1 1 6 E. 1315-1357
27.11.1996: 1 specimen
Source : MNHN, Paris
GRENADIERS PISHES OF THE NEW CALEDONIAN REGION
755
Norfolk Ridge. Halipro 2: stn BT 77, 24° 1 1.38'S. I67°45.52'E, 1349-1350 m, 21.11.1996: 1 specimen 175 mm
HL, 785+ mm TL (MNHN 1998-317). — Stn BT 82, 23°51.5'S, 1 68°25.98'E, 1846-1862 m, 22.11.1996: I specimen
257 mm HL, 1 180 mm TL (NMNZ P.34027).
Fig. 17. — Coryphaenoides dossemis McMillan, 1999. Paratype (CAS 71488, 57 cm TL) from east of New Zealand in
1200 m depth. Fins and scales partially reconstructed. Scale = 25 mm. From Shcherbachev & Iwamoto, 1995,
fig. 8. Used with permission of the California Academy of Sciences.
DIAGNOSIS. — Pelvic fins with 9 or 10 rays (rarely 8 or 11); snout low, scarcely protruding; orbits small,
about 1.5 into snout, 1.8- 1.9 into interorbital width, 5-6 into HL; upper jaw extends beyond vertical through hind
margin of orbits; premaxillary teeth in narrow band flanked by enlarged outer series, mandibular series 2-3 teeth
wide narrowing to one irregular row; outer gill slit small, 4-9% HL; inner gill rakers on first arch about 10 total;
barbel 10-23% HL; underside of head fully scaled.
DESCRIPTION (of MNHN uncat., Halipro 2 Stn BT 82). — Counts : ID. 11,10; P. i21; V. I 1; GR-I (inner)
1 + 1+7, GR-II (outer/inner) 2+6/ 1 + 1+7; scales below ID. about 10, below mid-ID. 7.5. below 2D. 6.5. lat.line
39 over distance equal to predorsal length.
Measurements'. 257 mm HL, 1 18 mm TL. The following in percent HL: snout 29; preoral 1 1 : internasal 20;
interorb. 32; orb. 18; suborb. 14; postorb. 58; orb. -preop. 54; up.jaw 42; barbel 17; depth 109; lit. ID. 52; len.
V. 38.
Size. — To more than 120 cm TL.
Distribution. — Widespread in tropical to subtropical waters of the Pacific, Indian, and Atlantic oceans.
Depth range about 980-2380 m.
REMARKS. — Coryphaenoides rudis has already been well described and illustrated in previous publications (see
synonymy). Our three large specimens taken during Halipro 2 agree well with these descriptions. MNHN uncat.
is a female with large ovaries, but small eggs.
756
N.R. MERRETT & T. IWAMOTO
F|C. IS. - Coryphaenoides ruclis Gunther, 1878. MMSU P-16528 (155 mm HL) from Nazca Ridge in 980 m. Seale =
“5 mm- From Iwam°to & Sazonov, 1988. fig. 28. Used with permission of the California Academy of Sciences.
Coryphaenoides striatums Barnard, 1925
COnCT\1tlS'riTr\S '.1wamoto & MERRETT, 1997: 511-512, Fig. 17 (New Caledonia). - Grandperrin e, ai,
1997: 1 17 (New Caledonia; list).
Material examined. — 2 specimens.
Norfolk Ridge. Halipro 2: stn BT 74, 24°44.9'S, 167°43.1'E, 1213-1246 m, 20.11.1996- 1 specimen 80 mm HL
‘ S 9084°'- ^ S“ BT M’11J8'S- 167“45-52’E' 1349-1350 m, 21.1 1. * TipTcll, ?NMNZ
Remarks. — A widespread species known from the southeastern Atlantic, Indian Ocean, southern Australia
New Zealand, and New Caledonia.
Genus HAPLOMACROURUS Tiunov, 1980
Haplomacrourus nudirostris Trunov, 1980
Haplomacrourus nudirostris - Iwamoto & Merrett, 1997: 513-514. Fig. 18 (Wallis & Futuna Islands).
Material examined. — 2 specimens
( 1 997^1 V ,Single •iuv®nile from the Wains and Futuna Islands was recorded by Iwamoto & Merrett
Halipro 2 crule6 C° eCl'°nS had tW° additional juveniles; no specimens were collected during the
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
757
Genus HYMENOCEPHALUS Giglioli, 1884
Hymenocephalus adelscotti Iwamoto & Merrett, 1997
Hymenocephalus adelscotti - Iwamoto & Williams, 1999: 176. Fig. 31a (Western Australia off Shark Bay, 680-691 in).
This species was not represented in recent Musorstom collections, but one specimen was recently recorded
from the eastern Indian Ocean by IWAMOTO & Williams (1999). The two peculiarly disjunct records (Fiji and
Western Australia) may be indicative of a much broader range for the species.
Hymenocephalus aterrimus Gilbert, 1905
Hymenocephalus aterrimus - Iwamoto & MERRETT, 1997: 516-517. Fig. 20a (New Caledonia, Norfolk Ridge. Wallis &
Futuna Islands).
Hymenocephalus ( Papyrocephalus ) aterrimus - Grandperrin et al., 1997: 1 17 (New Caledonia; list).
Material examined. — 5 specimens.
Vanuatu. Musorstom 8: stn CP 990, 18°51.63'S, 168°50.98'E, 980-990 m, 24.09.1994: I specimen (MNHN 1995-
999). — Stn CP 1125, I8052.41'S, 168°55.73'E, 764-786 m, 10.10.1994: 1 specimen (MNHN 1997-673).
Norfolk Ridge. Halipro 2: stn BT 76, 24°05.43'S, 167°34.29'E. 936-940 m. 21.11.1996: 2 specimens 15-38 mm
HL, 59-128+ mm TL (CAS 90611).
Loyalty Ridge. Halipro 2: stn BT 25, 25°16.64'S, I70°23.69'E, 1100-1348 m, 11.1 1.1996: I specimen 32 mm
HL, 140+ mm TL (MNHN 1998-315).
Remarks. — The species is broadly distributed, but is rarely caught in the area.
Hymenocephalus gracilis Gilbert & Hubbs, 1920
No specimens of this species were collected during recent cruises.
Hymenocephalus kuronumai Kamohara, 1938
No specimens of this species were collected during recent cruises.
Hymenocephalus longibarbis (Gunther, 1887)
Hymenocephalus longibarbis - Iwamoto & Merrett, 1997: 520-521, Fig. 21b (New Caledonia. Norfolk Ridge). —
Grandperrin et al., 1997: 1 17 (New Caledonia; list).
Material examined. — 20 specimens.
Vanuatu. MUSORSTOM 8: stn CP 1033, 17°54.75'S, 168°40.66'E, 650-691 m, 29.09.1994: 2 specimens (MNHN
1995-956). — Stn CP 1050, 16°39.20'S, 168°01.30'E. 541-577 m, 1.10.1994: I specimen (MNHN 1995-860). —
Stn CP 1052, 16°32.37'S, 168°00.29'E, 561-564 m, 1.10.1993: I specimen (MNHN 1995-957). — Sin CP 1073
1 5°45.70'S, 1 67°22.24'E, 630-650 m, 4.10.1994: 1 specimen (MNHN 1997-694). — Stn CP 1114, 14°52.39'S.
167°03.40'E, 647 m, 8.10.1994: 2 specimens (MNHN 1997-671).
New Caledonia. Bathus 1: stn CP 657, 21°14.45'S, I65°54.93'E. 490-530 m: 4 specimens (MNHN 1995-816). —
Stn CP 658, 21°13.45'S, 165°55.I2'E, 515-580 m, 12.03.1993: 2 specimens (MNHN 1995-826). — Stn CP 708.
21°43.05’S. 166°38.57'E. 550-580 m, 19.03.1993: 3 specimens (MNHN 1995-813). — Stn CP 709. 20°57.54 S.
165°35.86'E, 302-335 m, 18.03.1993: 4 specimens (MNHN 1997-687).
REMARKS. — This species was not taken during the Halipro 2 cruise, possibly indicating a distribution
concentrated to the north of the area fished. It is also possible that individuals were able to pass through the mesh
of the larger nets used during that cruise.
758
N.R. MERRETT & T. 1WAM0T0
Hymenocephalus megalops Iwamoto & Merrelt, 1997
Fig. 19
Hymenocephalus megalops Iwamoto & Mcrrett, 1997: 521-523. Fig. 22a (New
Plateau, Norfolk Ridge, Wallis & Futuna Islands, Australia).
Caledonia, Chesterfield & Bellona
Material examined. — 1 83 specimens.
Musorstom 8: SIn cp 980. I9°21 ,02'S, 1 69°25.22'E. 450-433 m. 22.09.1994: 2 specimens (MNHN
'^95-993h Stn CP 994, 18°47.72'S, 168°56.17'E. 649-641 m, 24.09.1994: 31 specimens (MNHN 1997-707). _
noanro07, 18°51.97'S, 1 68°55.92'E, 720-830 m, 25.09.1994: 1 specimen (MNHN 1997-705). — Sin CP 1028.
0 «,S- 1 68°40.42'E, 624-668 m. 28.09.1994: 9 specimens (MNHN 1995-962). — Sin CP 1033 I7°54 75~S
iS fo^6,Sf! m‘ 29 09-'f4: 1 spccimen (MNHN ,997-708»- - Stn CP 1034. 17-54.85-S. 168^42 04 f! 690
750 m, 29.09.1994: I specimen (MNHN 1995-974). — Stn CP 1050, 16°39.20'S. 1 68c01 ,30'E, 541-577 m I 10 1994
S£‘rirHNJ9^53)' - Stn CP I052.16“32.37'S, 168‘00.29'E, 561-564 m. 1. 10.1993: 20 specimens
i_ c.H^p9,9nJ?5 .■.oTT^.F^i^53’ l6°29-3S’ 167°58.70'E, 536-519 m, 1.10.1994: 7 specimens (MNHN 1997-706)
1 A90?'95 s' ,67°57-44'E' 522‘527 m: 2 specimens (MNHN 1995-984). — Stn CC 1056. 16°33.l I'S,
1 10 i qq2 V m: 31SP"lme"S'MNHN l995-989»- - Sln CP 1073, 15°45.70'S, I67°22.24'E, 630-650 m
's^ecimet'MTHr^-^T ^ ~ ** * 'V*** '6™A°'E' 647 8 *01994:
1995-M2Laled°nia‘ BATHUS I: ^ CP 7°9, 2°°57-54'S' 165°35-86'E, 302-335 m, 18.03.1993: 4 specimens (MNHN
50-
40
&
C
0)
=3 30 -
CT
0>
20 -
10 -
Fig. 19.
H L groups (mm)
- Length- frequency (HL, head length mm) distribution of Hymenocephalus megalops by sex (n = 384).
Female=72
Male=52
lndet=260
and in number abundant SpCCieS °f grCnadier in our samPles- both in number of individuals
and n number of captures. That none were taken during Halirro 2 is probably accounted for by its having
distribution to the north ol the sampling area. As with other species of small size, the larger mesh size in the
.raw, usee, du„„g that cru.se may a, so have been a fac.or ia its absence ,he ca.cl.es. I, eX7y li„s sexua'
Source MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
759
maturity in the range 13-22 mm HL (Table 5). One large sample contributed most to the length-frequency
analysis, but the overall indication is that the population peaks broadly over the range 20-26 mm HL classes
(Fig. 19). In common with many of the species encountered in these collections, the males of H. megalops are
macrosmatic and the females have ovaries containing several generations of ripening ova. Such features of
presumed mate location and reproductive style are well known and are commented on elsewhere (MERRETT. 1994).
The only stomach examined contained digested crustacean remains and lacked sediment, to indicate pelagic rather
than benthic feeding.
Hymenocephalus nascens Gilbert & Hubbs, 1920
Fig. 20
Hymenocephalus nascens - 1WAMOTO & MERRETT, 1997: 523-525 (New Caledonia. Norfolk Ridge. Chesterfield & Bellona
Plateau, Wallis & Futuna Islands).
Material examined. — 147 specimens.
Vanuatu. MUSORSTOM 8: stn CP 992, 18°52.34'S, 168°55.16’E, 775-748 m, 24.09.1994: II specimens (MNHN
1995-866). — Stn CP 993. 18°48.78'S, 168°54.04'E, 780-783 m. 24.09.1994: 34 specimens (MNHN 1995-969), — Stn
CP 994, 1 8°47.72'S, 168°56.17’E, 649-641 m. 24.09.1994: 20 specimens (MNHN 1995-873). — Stn CC 996.
18°52.41'S, 168°55.73'E, 764-786 m, 24.09.1994: 61 specimens (MNHN 1995-839). — Stn CP 1007. 18°51.97'S.
168°55.92'E, 720-830 m, 25.09.1994: 7 specimens (MNHN 1997-709). — Stn CP 1028. I7°54.0rs. 1 68°40.42E, 624-
668 m, 28.09.1994: I specimen (MNHN 1997-711). — Stn CP 1033. 17°54.75'S, 168°40.66'E. 650-691 m,
29.09.1994: 5 specimens (MNHN 1997-710). — Stn CP 1034, I7°54.85'S, 168°42.04'E. 690-750 m. 29.09.1994: I
specimen (MNHN 1997-712). — Stn CP 1073, I5°45.70'S. 167°22.24'E. 630-650 m, 4.10.1994: 2 specimens (MNHN
1995-851).
New Caledonia. Bathus 1: stn CP 660, 21°10.48'S. 165°53. 19E, 786-800 m, 13.03.1993: 2 specimens (MNHN
1995-823). — Stn CP 663, 20 58.66'S, 165 38.27’E, 755 m, 13.03.1993: 1 specimen (MNHN 1995-824). — Stn CP
709, 20°57.54'S, 165°35.86'E, 302-335 m, 18.03.1993: 2 specimens (MNHN 1997-703).
Remarks. — Hymenocephalus nascens was another relatively abundant species in the catches, although it was
not taken during the HAL1PRO 2 cruise. The majority of the population sampled were in the head length range lb-
28 mm of a unimodal catch distribution (Fig. 20a). Ovarian maturity was evidently attained at least by the 16 mm
HL class and breeding had occurred in some females in the 24 mm HL class, which were already spent (Fig. 20b).
More than one generation of ova were present in the ovaries, suggesting a likely batch-spawning strategy. Males
were found to be macrosmatic. Four out of the five stomachs in which contents were found contained digested
Crustacea and no sediment. Identification of material in one of these suggested that the prey was either a decapod or
a cuphausiid.
Hymenocephalus nesaeae sp. nov.
Fig. 21
Material examined. — 3 specimens.
Vanuatu. Musorstom 8: stn CP 1008, 18°53.29'S. 169°52.65'E, 919-1000 m, 25.09.1994: holotype 30.7 mm HL.
143+ mm TL (MNHN 1995-869). — Stn CP 990, 18°51.63'S, 168°50.98'E, 980-990 m, 24.09.1994: 1 paratype 28.6
mm HL, 130 mm TL (MNHN 1995-999). — Stn CP 1125, 18°52.41'S. I68°55.73'E. 764-786 m, 10.10.1994: 1 paratype
19.9 mm HL. 97 mm TL (MNHN 1997-673).
Diagnosis. — Pelvic fin rays 13-14; barbel absent; orbits 4-5 in head length, 1.2- 1.6 into interorbital width;
suborbital width 1.3- 1.7 into orbit; orbit-to-preopercle distance 42-50% HL; upper jaw 55-58% HL; body color
brown, scale pockets prominent, no trace of silvery pigmentation.
DESCRIPTION. — Counts: ID. 119-10; P. il3-il5; V. 13-14; GR-I (outer/inner) 19-21 / 5+20-21. C.R-II (4-
5)+(20-21) / (4-5)+( 19-21); scales over distance equal to predorsal length 18-19; pyloric caeca about 8 ( 1 spec.).
Measurements (in percent of HL): snout 27-33; preoral 13-14; internasal 24-26; inlerorb. 31-34; orb. 22-26.
760
N.R. MERRETT & T. IWAMOTO
FlG;.2,°. Length-frequency (HL. head length mm) distribution of Hymenocephalus nascens: (a) by sex (n = 340)-
Ki&^vSJSe?86 ^ = 73)' F‘ = imma'Ure: FH = y°lk and Ch0ri- formation; FI../V /egg nlration;'
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
761
Head moderately broad, greatest width about 1.2 into greatest body depth; snout slightly protruding, lacking
a thin flexible nasal flap. Mouth large, upper jaw extending well beyond vertical of posterior edge of orbit.
Suborbital region vertical except for narrow, tapered dorsal shelf. Head covering transparent except over opercular
region.
Luminescent tissue (ventral striae) relatively limited and not prominent; extending posteriorly as a narrow band
along both sides of isthmus below gill covers, passing dorsal to pelvic fin base, expanding medially and
posteriorly behind pelvic bases but not joining along median line and falling well short of periproct region;
anterior lens on chest small, scarcely discernible without magnification; posterior lens before periproct
considerably larger.
Fin rays all fine and slender, none thick and well-developed; outer ray of pelvic fins prolonged, extending well
beyond anal fin origin, about to base of eighth ray; spinous dorsal ray slightly produced, hairfine at distal tip.
Body color uniformly dark brown with large scale pockets prominently marked. No trace of lateral band nor
suggestion of there having been any silvery tissue. Dark pigmentation uniformly covers all of trunk and tail, with
abdomen and chest forward to isthmus black. Most of opercular bones and inner wall of suborbital bones black;
lips, jaws, gular and branchiostegal membranes black; most of roof of mouth black, but longue pale. Dorsal fin
dark dusky, most of pectoral, pelvic and anal tins dark dusky. Leading edge and median nasal ridge of snout, and
interorbital space at top of orbits black.
Fig. 21. — Hymenocephalus nesaeae sp. nov. Lateral view of head and trunk of holotype (MNHN 1995-869), with fins
partially reconstructed. Scale = 25 mm.
Size. — To more than 140 mm TL.
ETYMOLOGY. — From the Greek, Nesaie, one of the Nereids, a sea nymph.
Distribution. — So far known only from three hauls made off Vanuatu in depths between 764 m and
1000 m.
Remarks. — Hymenocephalus nesaeae has the characters of high pelvic ray count, absence of a chin barbel,
and uniiormly dark body color that are similarly found in H. aterrimus, but H. aterrimus has a much smaller orbit,
broader head, deeper suborbital, a thin anterodorsally pointed snout tip, and a more uniformly black head.
H. nascens has similar pelvic ray counts and no barbel, but the body in that species has a prominent broad lateral
stripe over a lighter ground.
762
N.R. MERRETT & T. IWAMOTO
The presence of another species of Hymenocephalus in waters in and around the New Caledonian region brings
the total known from Musorstom collections to eight, making it comparable to the Philippines-Indonesian
region in terms of the diversity of the genus.
Genus KUMBA Marshall. 1973
Kumba musorstom sp. nov.
Fig. 22
Kumba punctulata ( non Iwamoto & Sazonov) - Grandperrin el al., 1997: 1 17 (New Caledonia; listed).
Material examined. — l specimen.
Norfolk Ridge. HALIPRO 2: stn BT 55. 25°02.85'S. 168°45.42'E, 1098-1480 m, 17.11.1996: holotype, female.
72.6 mm HL. 425 mm TL (MNHN 1998-232).
Diagnosis. — Pelvic fin rays 12-13, outer ray large, greatly elongated; spinous second ray of first dorsal fin
elongated and with few tiny denticulations along leading edge; naked areas on snout extend dorsally beyond lateral
nasal angles; snout length greater than inlcrorbital width; upper jaw length about half head length; barbel about
17% HL; no external sign of light organ.
DESCRIPTION OF HOLOTYPE. — Counts: ID. 11,10; P. i22/i21; GR-I (outer/inner) 9/3+1 1, GR-I1 3+11/ 3+10;
scales below ID. 9, below mid-base ID. 7, below 2D. 10.5, lat. line 37.
Measurements (in mm, percent HL in parentheses): snout 19.7 (27); preoral 12.9 (18); internasal 1 1.4 (16);
interorb. 14.9 (21); orb. 15.0 (21); suborb. 1 1.4 (16); postorb. 41.3 (57); orb.-preop. 33.8 (47); up.jaw 35.3 (49);
barbel 12.0 (17); gill-slit 14 (19); pre-A. 108 (149); V. -A. 32 (44); depth 63 (87); 1D.-2D. 10.5(14); ht. ID. 63
(87); len. P. 43 (59); len. V. 126 (174).
General features of fish seen in Fig. 22. Head relatively compressed, deeper than wide, greatest width about 1.2
into greatest head depth, head length almost 6 times into total length. Snout rounded in profile, protruding slightly
beyond mouth. Interorbital region incised anteriorly by two grooves on either side of median nasal ridge, and
posteriorly by a shallow median depression anterior to origin of nape. Suborbital region almost vertical, widi little
development of a ridge. Mouth large, lips thick, upper jaw extends slightly beyond vertical through hind margin of
orbit. Chin barbel slender, length about equal to horizontal diameter of orbit.
Underside of head almost entirely naked, including snout, suborbital, ventral margins of preopercle and most of
interopercle (excepting small scale patch at posterior end), and lower jaw. Areas under snout, suborbital and lower
jaw peppered with small open pores. Dorsal surface of snout naked to level of anterior nostril, thence posteriorly in
two thin strips at bottom of grooves running on either side of median nasal ridge. Ridges surrounding nasal fossa
scaled, as well as upper portion of suborbital. Anus immediately before anal fin; periproct small, about equal in
area to nasal fossa. No external evidence of light organ. Holotype a female with large ovaries; small eggs
developed.
Dorsal fin with a greatly elongated spinous ray extending as a thin filament well beyond adjacent rays; leading
edge of prolonged ray smooth except for two tiny teeth about midway along length. Second dorsal fin low
throughout, begins close behind first dorsal, the interspace between fins about half length of base of first dorsal.
Pectoral fin well developed, the longest rays extending to over 8- 10th anal rays. Pelvic fin small except for stout,
greatly prolonged outer ray, which is longer than preanal distance and filamentous at distal tip. First dorsal origin
posterior to that of pectoral origin, which is posterior to that of pelvic origin. Anal fin begins approximately under
posterior l/4th of First dorsal base.
Small teeth in upper jaw in long broad band, outer series of slightly enlarged spaced conical teeth. Lower jaw
teeth in narrow band about three teeth across.
Scales of body almost entirely lost, a few scales along anterior origin of lateral line and over nape. These with
characteristic (for genus) narrow, comblike rows of slender spinules. Rows somewhat divergent. Head scales all
small, and characteristic of genus (comblike divergent rows of spinules).
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
763
Overall color of body and tail brownish wilh bluish tinge over abdomen and chest. Head generally paler, ventral
surfaces white overlain with smudgelike marks. Lips blackish along mouth opening but jaw surfaces otherwise
almost immaculate. Mouth whitish on roof, darker over other surfaces, blackish over oral valves and gums. Gill
cavity black; gill arches and rakers dark. Branchiostegal membrane black but pale over rays. Opercle and subopercle
blackish; most of other parts of head surfaces gray to whitish. Area around orbit and nostrils brownish. Fins all
brownish to gray-brown.
Fig. 22. — Kumba musorstom sp. nov. Lateral view of holotype (MNHN 1998-232) from Norfolk Ridge in 1098-1480 m
depth. Scale = 25 mm. Drawn by Alison Schroeer.
Size. — To at least 43 cm TL.
Etymology. — The name derives from the acronym, Musorstom, for the series of exploratory cruises
carried out in the Indo-Wcst Pacific region jointly by the Institut fran$ais de Recherche Scientifique pour
le Developpement en Cooperation (ORSTOM - now IRD) and the Museum national d'Histoire naturelle
(MNHN). The name is treated as a noun in apposition.
Distribution. — Known only from the Norfolk Ridge south of New Caledonia in 1098-1480 m.
Remarks. — This new species most closely agrees with K. calvifrons Iwamoto & Sazonov, 1994 in terms of
overall shape, extent of naked areas on snout, and size of upper jaw, snout and interorbital. The two species differ
radically in a number of features, with K. calvifrons having pelvic fin rays fewer (8), outer ray not extremely
elongated, barbel shorter (9% HL), serrations on spinous ray of first dorsal fin well developed, spinous ray not
greatly prolonged, dermal windows of light organ developed, and scales smaller.
764
N.R. MERRETT & T. IWAMOTO
The spinous dorsal ray in Kumba gymnorhynchus Iwamoto & Sazonov, 1994 also lacks serrations, hut that
ray is much shorter than in the new species. Numerous other features differ between the two, including: pelvic fin
ray count (10-11 in K. gymnorhynchus), internasal width (22-24% HL), interorbital width (26-27%), orbit
diameter (25-26%), postorbital length (48-51%), upper jaw length (40-43%), barbel length (about 7%), outer
pelvic ray length (63-67%), and naked area on snout not extending beyond lateral nasal angles.
Kumba punctulata, which has been taken close to the type locality of K. musorstom, is a much smaller species
with 9-10 pelvic fin rays and many differing proportions in head measurements.
Kumba punctulata Iwamoto & Sazonov, 1994
Kumba punctulata - IWAMOTO & MERRETT, 1997: 526, Fig. 23 (Norfolk Ridge).
Material examined. — 3 specimens.
Vanuatu. Musorstom 8: stn CP 975, 19°23.60'S, I69°28.93’E, 566-536 m, 22.09.1994: I specimen 21 mm HL
(MNHN 1997-721). — Stn CP 1027, I7°53.05'S, 168°39.35'E, 550-571 m, 28.09.1994: 1 specimen 16 mm HL (MNHN
1997-674). — Stn CP 1050, 16°39.20’S. 168°01.30'E, 541-577 m, 1.10.1994: I specimen 15 mm HL (MNHN 1997-
715).
Remarks. — The three Vanuatu specimens were a surprise, as only the holotype was present in previous
collections reported by Iwamoto & Merrett (1997) from the New Caledonian area. Only six specimens of this
rare fish are known.
Genus KURONEZUMIA Iwamoto, 1974
Kuronezumia Iwamoto, 1974: 509 (as subgenus of Nezumia).
Diagnosis. — Branchiostegal rays 7; snout rounded in profile, almost entirely covered with small, uniform,
finely spinulated scales. Suborbital region vertical, without an angular midlateral ridge, the region covered with
small scales and lacking a row of enlarged, modified scales. Upper jaw 30-44% of HL. Jaw teeth in broad villiform
bands, bands in lower jaw short and broad. Gill rakers 8-1 1 total on outer side of second arch. Small patches of
scales on branchiostegal rays in most species. Body scales small, adherent, densely covered with long slender
spinules. Anus tar removed from anal fin, closer to pelvic bases. Anterior dermal window of light organ usually
small, situated between pelvic fin bases and separated from anus by a broad scaly area. Colour overall grey to
brown to swarthy; fins dusky to blackish, naked membranes blackish to dark grey. Abdominal vertebrae 12-13
(After Sazonov & Iwamoto, 1992:64-65).
Remarks. — Shcherbachev et at. (1992) recently reviewed the genus, recognizing seven species, some of
which had been variously included in the genera Nezumia, Ventrifossa, Macruroplus, Lionitrus and Mac, -aunts.
Only one species is known from the New Caledonian region, but two are known from New Zealand and three from
Australia.
Kuronezumia bubonis (Iwamoto, 1974)
Fig. 23
Nezumia (Kuronezumia) bubonis Iwamoto, 1974: 507-516, Figs 1-3 (holotype: CAS 27872 from Gulf of Mexico)
Nezumia bubonis - Paxton et al„ 1989: 328 (New South Wales & Victoria. Australia)
Kuronezumia bubonis - Paulin et al., 1989: 130 (New Zealand; in key).
Material examined. — i specimen,
Vanuatu. Musorstom 8: stn CP 1080, 15°57.30'S, I67°27.73'E, 799-850 m 5 10 1994-
182+ mm TL (MNHN 1997-716).
I specimen 32.8 mm HL.
Source . MNHN , Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
765
Diagnosis. — Head completely and uniformly covered with small, finely spinulated scales; a large scaly
swelling of light organ between pelvic fins; V. usually 1 1-13; scales below 2D. 12-15.
Description (of one specimen). — Counts: ID. 11,12; IP. i21/i23; V. 11/10; GR-I (outer/inner) 5/2+1+7,
GR-I1 l-t-1+7/1+8; scales 2D. about 14.
Measurements : snout 8.8 (27); preoral 4.9 (15); internasal 5.9 (18); interorb. 8.0 (24); orb. 10.8 (33); suborb.
5.6 (17); postorb. 15 (46); orb-preop. 11.7 (36); up. jaw 1 1.6 (35); barbel 6.8 (21); V.-A. 10 (30); body depth 28
(8); depth over A. 21 (64); ht. ID. 35 (107); len. P. 1 (64); len. V. 18 (55); gill slit 6.0 (18).
Size. — A species of large adult size to about 73 cm TL.
Distribution. — Widespread, from central western Atlantic. Indian Ocean, southwestern Pacific, and Hawaii.
Depth range about 600-1300 m.
Remarks. — The single juvenile from Vanuatu represents the first record of the species from the
New Caledonian region. See Shcherbachev et al. (1992) for more references and comparisons with other
members of the genus.
Fig. 23. — Kuronezumia bubonis (Iwamoto, 1974). Lateral view of head and trunk of paratype, BPBM uncat., from
Hawaii. Scale = 25 mm. From Iwamoto, 1974, tig. 2. Courtesy of the California Academy of Sciences.
Genus LUCIGADUS Gilbert & Hubbs, 1920
Lucigadus acrolophus Iwamoto & Mcrrett, 1997
Lucigadus acrolophus Iwamoto & Merrett, 1997, 526-528, Fig. 24a (New Caledonia. Norfolk Ridge)
766
N.R. MERRETT & T. IWAMOTO
Material examined. — 6 specimens.
Vanuatu. MUSORSTOM 8: sin CP 980, 19°21.02'S. I69°25.22'E. 450-430 m, 22.09.1994: 3 specimens 23-28 mm
HL (MNHN 1995-992). — Sin CP 1137, 15°41.52'S, I67°02.67'E. 360-371 m, 11,10.1994: 1 specimen 13 mm HI
(MNHN 1997-675).
Sin unknown but probably from Musorstom 8: 2 specimens 26-34 mm HL (MNHN 1995-967).
Remarks. — This small species is known only from New Caledonia, Vanualu. and the Norfolk Ridge north
of 22°54'S, in depths of 360-450 m.
Lucigadus microlepis (Gunther, 1878)
Fig. 24
Lucigadus microlepis - IWAMOTO & Merrett, 1997: 528-530, Fig. 24b (New Caledonia. Fiji, Wallis & Futuna Islands,
Norfolk Ridge, Indonesia).
Material examined. — 21 specimens.
New Caledonia. Bathus 3: stn CP 833, 23°02.75'S, 166°58.23’E. 441-444 m. 30.11.1993: 3 specimens (MNHN
1995-817).
Vanuatu. Musorstom 8: stn CP 973, 19°2L30’S, I69°27.03'E. 460-480 m, 22.09.1994: 4 specimens 10-23 mm
HL (MNHN 1995-867). — Stn CP 974, 19°21.51'S, 169°28.26'E. 492-520 m. 22.09.1994: 3 specimens 17-22 mm HL
(MNHN 1995-976). — Stn CP 980, 19°2I.02'S, 169°25.22'E, 450-430 m, 22.09.1994: 1 specimen 18 mm HI (MNHN
1995-700). — Stn CP 983. 19°21.6rS, 169°27.76'E. 480-475 m, 23.09.1994: 1 specimen 15 mm IIL (MNHN 1997-
676). — Stn CP 984. 19°19.62'S. 169°26.-43’E, 480-544 m, 23.09.1994: 7 specimens 14-22 mm HL (MNHN 1995-973).
— Stn CP 1026. 1 7°50.35'S, 168°39.33'E, 437-504 m, 28.09.1994: 1 specimen 17 mm HL (MNHN 1995-850)
Norfolk Ridge. Halipro 2: stn BT 94, 23°36.7'S, 167°41.66'E, 448-880 m, 24.11.1996: 1 specimen 25 mm HL
147 mm TL (CAS 90700).
Remarks. — This species is quite common compared to its congener Lucigadus acrolophus. It is widespread
from Indonesia to Queensland, Australia (personal data, TI), Fiji, Wallis and Futuna Islands, New Caledonia,
Norfolk Ridge, and Vanuatu, in depths ranging from 418-600 m in the Pacific, but only 204-289 m in Indonesian
seas. No captures have been recorded south of 25°S.
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
767
The majority of the Lucigadus microlepis sampled occurred in the 13-19 mm HL group (Fig. 24). Its small
adult size was confirmed by ovarian maturation being observed within the range 18-20 mm HL. Males were seen
to be macrosmatic and the ovaries of females contained more than one generation of ripening ova. A variety of
crustacean prey was found in the stomachs of two of the nine specimens examined (Table 5). These two stomachs
were free of sediment, while another contained sediment but no identifiable food items.
Genus MACROSMIA Merrctt, Sazonov & Shcherbachev, 1983
Diagnosis. — A macrourine with seven branchiostegal rays; olfactory organ massive, about 4-5 times in head
length; head squamation reduced, mostly lacking on snout and underside of head, no scaled ridges on head and
snout; second spinous ray of first dorsal fin weakly serrated; pelvic fins anteriorly placed; anal fin origin below
hind margin of tirst dorsal fin; chin barbel developed; anus within small pcriproct immediately anterior to anal fin;
no light organ; swimbladder well developed with two gas glands and retia; pyloric caeca 15-18. (After MERRETT et
al. , 1983.)
Remarks. — A small, rarely caught, deep-living species. Monotypic.
Macrosmia phalacra Merrett, Sazonov & Shcherbachev. 1983
Fig. 25
Macrosmia phalacra Merrett. Sazonov & Shcherbachev, 1983: 554-560, Figs 1-3 (holotype ZMMGU P-15390. Ninety
East Ridge, Indian Ocean, I7°04.8'S, 88°09.0'E, 1650-1660 m; paratype from Canary Passage, E. North Atlantic,
I 654- 1 699 m).
Material examined. — i specimen.
Vanuatu. Musorstom 8: stn CP 1125, 15°57.63'S, I66°38.43'E, 1160-1220 m. 10.10.1994: I female 27.9 mm HL
207+ mm TL (MNHN 1997-724),
Diagnosis. — As for genus.
Description (one specimen). — Counts: ID. 11,9; P. i20/i2l; V. 12/11; GR-I (outer/inner) 8 / 2+1+7. GR-II
1 + 1+9 / 2+8; scales below ID. 5 or 6, below mid- ID. 4, below 2D. 6, lat.Iine scales over distance equal to
predorsal length about 29.
Measurements (in mm followed by value in percent of HL): snout 7.8 (28); preoral 4.7 (17): inlernasal 4.1
(15); post, nostril 4.5 (16); interorb. 6.0 (2); orb. 8.0 (29); suborb. 3.9 (14); postorb. 13.9 (50); orb.-preop. 10.6
(38); up.jaw 1 1.5 (41); barbel 5.9 (21); outer gill-slit 4.8 (17); body depth 22 (79); depth over A. origin 19 (68);
ht. ID. 26.5 (95); len. P. 16 (57); len. V, 26 (93); 1D-2D. interspace 8 (29).
Snout blunt, in this specimen appearing to have been somewhat pushed in during capture or preservation:
mouth (premaxillary) extends posteriorly to below hind third of orbit. Head compressed, jaws lateral, suborbilal
vertical and rounded, without sharp ridges. Snout, suborbital, and anterior half of interorbital space naked; naked
region extends to lower part of preopercle and onto entire lower jaw. Pores of cephalic lateral line system well
developed, especially along ventral margin of suborbital, preopercle, and along mandibular rami. Posterior nostril
hugely macrosmastic, diameter about equal to that of pupil. Gill rakers and filaments relatively well developed;
rakers tubercular, armed with coarse spinules. Filaments fairly long, about 3 mm. Periproct region relatively
narrow, with no indication of a light organ. Ovary large, eggs of various sizes ranging from about 0.1 mm to
more than 0.4 mm in diameter.
Snout entirely naked ventrally and dorsally to about midorbit, but some small spinulated scales at lateral angles
of snout and posteriorly to below nostrils, continuing onto suborbital shelf to below midorbit. Scales on
preopercle, opercle, and subopercle thin, non-spinulated and appearing almost non-imbricate. Lateral line well
developed. Body scales small, none enlarged.
768
N R. MERRETT & T. IWAMOTO
Teeth in wide bands, 4-5 teeth wide in both jaws, outermost in premaxillary very slightly enlarged, none in
mandibular band enlarged; some mandibular teeth with arrowhead-shaped tip.
Head all black; abdomen black, fins all black or nearly so; upper part of trunk, nape and (ail swarthy; mouth
and gill cavities black.
Fig. 25. — Macrosmia phalacra Merrelt, Sazonov & Shcherbachev, 1983. Holotype, ZMMGU P.15390 (composite of
right and left sides of fish; 11ns and scale pattern reconstructed). Scale = 25 mm.
Size. — A species of small adult size, to about 22 cm TL.
Distribution. — Eastern North Atlantic, Indian Ocean off Ninety Degree East Ridge, and Vanuatu in
southwestern Pacific. Depth range 1 160-1699 m.
Remarks. — Merrett et al. (1983) give a detailed description and discussion of this unusual wide-ranging
fish based on two male specimens. Our Vanuatu specimen agrees closely with theirs, which came from the central
Indian Ocean and the North Atlantic (27°N). A few differences were found in proportional measurements
and counts. Our specimen had 2+1+7 inner gill rakers on the first arch compared with 2+0+10 and 1 + 1+9 in
the type specimens. Scale row count below the first dorsal was 5 or 6 cf. 7 or 8, and below the second dorsal 6
cf. 10 in the holotype. The snout was shorter in our specimen (28% HL cf. 32 and 34%; preoral 17% cf. 21
and 23%), the orbit diameter slightly longer (29% cf. 27 and 21%), and the upper jaw longer (41% cf. 34 and
36%). With so few specimens available for comparison, we cannot properly assess the significance of these
dilfcrences. That our specimen was a maturing female may account for these discrepancies in counts and
measurements.
The enormous ollactory organ (Fig. 25) in the female of this species supports the statement of Merrett et al.
( 1 983: 553) that the condition is not a matter of sexual dimorphism, but one that has a phylogenetic basis.
Source : MNHN , Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
769
Genus MALACOCEPHALUS Gunther, 1887
Malacocephalus laevis Lowe, 1 843
Malacocephalus laevis - Iwamoto & Merrett, 1997: 532, Fig. 25 (Norfolk & Loyally Ridges. Chesterfield & Bellona
Plateau, Wallis & Futuna Islands).
Material examined. — 3 specimens.
Vanuatu. Musorstorm 8: stn CP 994, 18°47.72'S, I68°56.17'E. 649-641 m. 24.09.1996: 1 specimen 85 mm HL
(MNHN 1995-836). — Stn CP 1028, 17°54.01'S, 168°40.42'E, 624-668 m, 8.09.1996: 1 specimen 80 mm HL (MNHN
1995-838), 1 specimen 56 mm HL (MNHN 1995-963).
Remarks. — This circumglobal species was previously recorded by Iwamoto & Merrett (1997: 53) from
five specimens collected off the Wallis and Futuna Islands, Chesterfield and Bellona Plateau, and the Norfolk
Ridge. It is now known from three specimens from Vanuatu. None was collected during HALIPRO 2.
Genus MATAEOCEPHALUS Berg, 1898
Mataeocephalus acipenserinus (Gilbert & Cramer, 1897)
Mataeocephalus acipenserinus - Iwamoto & Merrett. 1997: 533-534, Fig. 26 (Chesterfield & Bellona Plateau. Loyalty
Islands, Wallis & Futuna Islands).
Material examined. — 12 specimens.
Vanuatu. MUSORSTOM 8: sin CP 991, 18°51.6S, I68°52.19'E, 936-910 m. 24.09.1996: 2 specimens 30.3-42.7 mm
HL, 135-182+ mm TL (MNHN 1995-994). — Stn CP 992, I8°52.34’S, 168°55.16'E. 775-748 m, 24.09.1996:
2 specimens, 39.0-52 [snout missing] mm HL, 160-230+ mm TL (MNHN 1997-681). — Stn CP 996, 18°52.41'S.
168°55.73'E, 764-786 m, 24.09.1996: 6 specimens, 31.5-49.6 mm HL. 120+-210+ mm TL (MNHN 1995-881). —
Stn CP 1125, 15°57.63’S, 166°38.43'E, 1160-1220 m. 10.10.1996: 1 specimen (MNHN 1997-680).
Wallis and Futuna Islands. MUSORSTOM 7: stn CC 554, I2°I3.08'S. 177°28.00'W. 820-795 m, 18.05.1992:
1 specimen 44.4 mm HL. 176+ mm TL (MNHN 1995-667).
REMARKS. — Mataeocephalus acipenserinus was captured in small numbers in Vanuatu and other areas within
the New Caledonian region. It was rarely taken in quantity in any haul, although six specimens were captured in
Stn CP 996. Iwamoto & Merrett (1997: 61-62) recorded 1 1 specimens captured at eight stations from previous
Musorstom cruises. No specimens were collected during Halipro 2. suggesting that the species may be confined
to waters north of about 21°S. In a juvenile (30.3 mm HL) from MNHN 1995-994, the trunk and tail had
a banding pattern much as that seen in juveniles of Malacocephalus laevis.
Mataeocephalus adustus Smith & Radcliffe, 1912
Fig. 26
Mataeocephalus adustus Smith & Radcliffe in Radcliffe, 1912: 126-127. pi. 28, Fig. 3 [holotype USNM 72943, from
off Celebes in 805 fathoms (1472 m)].
Material examined. — 2 specimens.
Vanuatu. Musorstom 8: stn CP 1037, 18°03.70’S, 168°54.40'E, 1058-1086 m, 29.09.1996: I specimen 43.0 mm
HL. 183+ mm TL (MNHN 1995-854). — Stn CP 1076, 15°53.81'S, 167°30.42'E, 1100-1191 m, 4.10.1994: I specimen
61.8 mm HL. 307+ mm TL (MNHN 1995-846).
770
N.R. MERRETT & T. IWAMOTO
Diagnosis. — V. 7; P. i 17-i 19; underside of head mostly scaled; periproct region close before anal-fin origin,
no separate dermal window anteriorly; spinous second ray of first dorsal fin greatly elongated fto as much as twice
length of head), serrations along leading edge rudimentary or absent.
Description (two specimens). — Counts : ID. 11,9; GR-I (outer/inner) 0 / 7-9 total. GR-II 1+7 / 1+7; scales
below ID. 7.5-8, mid- ID. 4.5-5.0, 2D. 6.5, lat.line 34-37.
Measurements (in percent HL): snout 37-39; preoral 30-35; inlernasal 21-25; interorb. 20-22; orb. 23-24;
suborb. 15-16; postorb. 40-41; orb-preop. 38-40; up. jaw 24-27; barbel 5-7; gill slit 8-9; V.-A. 34-35; body depth
63; depth over A. 49; ht. ID. 107-157; len. P. 52-64; len. V. 53-55.
F'°a2n!'~f; “jocephalus aduf's S,mi'h & Radcliffe. 1912. MNHN 1995-846 from off Vanuatu, (a) Lateral view of head
and trunk, (a I dorsal view of head; (b) diagrammatic lateral view. Scale for (a) and (a') = 25 mm.
Head large, slightly inflated in fresh specimens with most surfaces broadly rounded. Body slender, greatest depth
about 1 .6-1.7 into head length, about eight into total length. Snout in dorsal view triangular anterior to nostrils,
sharp creases dorsally behind leading edge, shallowly triangular in lateral view, projecting well beyond small
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
771
mouth; tipped with two small tubercular scales. Head ridges smoothly rounded in fresh specimen, but suborbital
ridge acute and forming relatively sharp angle between dorsal and ventral surfaces. Gill arches greatly restricted
dorsally and ventrally, with outermost gill slit small; no rakers on outer side of first arch; rakers on inner side
tubercular, with rather long spinules. Chin barbel short, almost rudimentary; gill membranes broadly united across
isthmus.
Premaxillary teeth band short, broad, falling well short of end of rictus, outer series of teeth scarcely enlarged;
mandibular band longer, tapered to one row posteriorly.
Body scales with short, needlelike spinules in quincunx or slightly convergent pattern. Scales end abruptly
under gill cover. Underside of head uniformly scaled except for thin naked strips along suborbital and prcopercle
ridges, along median ventral line of snout, and along ventral margins of suborbital and preopercle; entire
interopercle naked. Mandibular rami broadly naked but a patch of scales on middle at posterior end.
First dorsal fin high with greatly prolonged second spinous ray; weak serrations proximally along leading edge
of ray or edge smooth. Origins of first dorsal and pelvic fins about on same vertical, origin of pectoral fin in
advance of that vertical; origin of anal fin well behind vertical through hind margin of first dorsal fin. Outer pelvic
fin ray slightly elongated, extending just beyond anal fin origin.
Anus separated by a short gap from anal fin origin, much closer to anal fin origin than to pelvic fin bases.
Periproct region small, about diameter of nasal fossa, narrow, raised; light organ small.
Mouth dark. Fins all blackish. Ground color on body and tail swarthy to gray-brown, abdomen darker. Opercle
and branchiostegal membranes blackish; gular membrane somewhat paler.
Remarks. — Only two specimens of this rare species of Mataeocephalus were captured. The species is close
to an undescribed species recorded as Mataeocephalus sp. by Iwamoto & Merrett (1997: 62) from the Loyalty
Islands and the Chesterfield and Bellona Plateau, but that species has only six branchiostegal rays, a vent that is far
removed from the anal fin and closer to the pelvic fin bases, a shorter snout (32-33% HL), and lanceolate scale
spinules. Both species have an almost completely scaled underside of head and relatively short snout for
a Mataeocephalus. Mataeocephalus adustus can be distinguished from M. acipenserinus by its shorter preoral
length (28-35% cf. 38-40% in M. acipenserinus), longer postorbital length (39-41% cf. 31-35%), longer orbit to
preopercle (38-41% cf. 30-36%), and longer upper jaw (24-29% cf. 21-23%) [additional measurements for
M. adustus provided by Y. I. SAZONOV, ZMMGU].
Mataeocephalus sp.
Mataeocephalus sp. Sazonov & Shcherbachev MS - Iwamoto & MERRETT, 1997: 62-63, Fig. 7 (6 spec.. Loyalty Islands
and Chesterfield and Bellona Plateau; 412-970 m).
Material examined. — 20 specimens.
Vanuatu. Musorstom 8: stn CP 992, 18°52.34’S, 168°55.I6'E. 775-748 m. 24.09.1994: I specimen 44 mm HL
(MNHN 1997-679). — Stn CP 993, 18°48.78'S, 168°54.04'E, 780-783 m. 24.09.1994: 1 specimen 33 mm HL (MNHN
1995-969). — Stn CP 996, 18°52.41'S, 168°55.73'E, 764-786 m, 24.09.1994: 12 specimens 30-46 mm HL (MNHN
1997-678). — Stn CP 1008. I8°53.29'S, 169°52.65'E, 919-1000 m, 25.09.1994: 3 specimens 17-32 mm HL (MNHN
1995-978). — Stn CP 1034, I7°54.85'S. 168°42.04'E, 690-750 m, 29.09.1994: I specimen 41 mm HL (MNHN 1995-
874). — Stn CP 1074, 15°48.42'S, 167°24.27'E, 775-798 m. 4.10.1994: 1 specimen 51 mm HL (MNHN 1995-856).
— Stn CP 1125, 18°52.41'S, 168°55.73'E, 764-786 m. 10.10.1994: 1 specimen 39 mm HL (MNHN 1997-677).
Remarks. — The species was not recorded from Halipro 2, but was captured seven times in Vanuatu waters.
Genus NEZUMIA Jordan, 1904
Nezumia aspidentata Iwamoto & Merrett, 1997
This rarely caught species was not represented in current collections.
772
N.R, MERRETT & T. IWAMOTO
Nezumia cliveri Iwamoto & Merrett, 1997
Nezumia cliveri Iwamoto & Merrett, 1997: 539-540, Fig. 28c (Norfolk & Loyalty Ridges).
Material examined. — l specimen,
Norfolk Ridge. Halipro 2: stn BT 42, 25°34.29'S, 167°25.08'E, 1132-1160 m, 16.11.1996: 1 specimen (CAS
90751).
Remarks. — This species is known only from three specimens taken off the Norfolk and Loyally Ridges.
Nezumia coheni Iwamoto & Merrett, 1997
Nezumia coheni Iwamoto & Merrett. 1997: 540-541, Fig. 28b (New Caledonia, Norfolk & Loyally Ridges, Kermadec
Islands, Australia).
Material examined. — 5 specimens.
Loyalty Ridge. Halipro 2: stn BT 15, 24°36.28'S. I70°03.85'E, 900-1005 m, 8.11.1996: 1 specimen 52 7 mm
HL. 305+ mm TL (MNHN 1998-230).
Norfolk Ridge. Halipro 2: stn BT 46. 25°41.99'S, 167°25.6'E. 1230-1268 m. 15.11.1996: 1 specimen 57 1 mm
HL. 331 mm TL (BMNH 1997.5.21.45). — Stn BT 55. 25°02.85'S, 168°45.42'E, 1098-1480 m, 17.1 1.1996: 1 specimen
66.2mm HL. 361 mm TL (MNHN 1998-324). — Stn BT 75, 24°13.I7'S, 167°36.52'E. 1128-1150 m, 21.1 1.1996
1 specimen 51.5 mm HL. 302+ mm TL (paratype, CAS 90614). I specimen (NMNZ P.34253).
Lord Howe Rise. Halipro 2: stn BT 104, 25°23.01'S, 162°35.4'E, II 18-1124 m. 27.11.1996: 1 specimen
(paratype, CAS 90668).
Remarks. — Known from relatively small numbers in New Caledonian waters, but fairly common off east
coast of Australia.
Nezumia propinqua (Gilbert & Cramer, 1897)
? Nezumia propinqua - Iwamoto & Merrett. 1997: 541-542, Fig. 29a (New Caledonia. Norfolk & Loyalty Ridges,
Chesterfield & Bellona Plateau, Wallis & Futuna Islands). — Grandperrin et al„ 1997: 1 17 (New Caledonia; list)^
Material examined. — 25 specimens.
Vanuatu. Musorstom 8: stn CP 970, 20°18.56'S. 169°53.19'E, 25-310 m, 21.09.1994: 1 specimen (MNHN 1997-
702). - Sin CP 991, 18°51.26'S, 168°52.19'E. 936-910 m, 24.09.1994: 2 specimens (MNHN 1995-991). —
Stn CP 992, 18°52.34'S, 1 68°55. 1 6'E, 775-748 m, 24.09.1994: 1 specimen 124 mm HL (MNHN 1995-970). _
Stn CP 993, 1 8°48.78'S. 168°54.04'E, 780-783 m, 24.09.1994: 1 specimen 33 mm HL (MNHN 1997-695). _
"6‘ 18°^2'41 S’ 1 68°55.73'E, 764-786 m, 24.09.1994: 4 specimens (MNHN 1997-682). — Sin CP 1008.
18 53.29'S, 169°52.65'E. 919-1000 m, 25.09.1994,: 2 specimens (MNHN 1997-696). — Stn DW 1012, 17°53.24's'
1 683 1 3.09’E, 775-740 m, 27.09.1994: 1 specimen (MNHN 1996-482). — Stn CP 1033, 17°54.75'S, 168°40 66'E 650-
‘ specimen (MNHN 1995-958). -Stn CP 1034, 17°54.85’S, 168°42.04'E, 690-750 ‘m.
29.09.1994: I specimen (MNHN 1995-975). - Stn CP 1035, 17°56.02’S, 168°44.06'E, 765-780 m 29 09 1994-
specimen (MNHN 1997-699). - Stn CP 1036, 18°01.00'S, I68°48.20'E, 920-950 m, 29.09.1994: 1 specimen
(MNHN 1995-968). — Stn CC 1056, 16°33.H S, I67°55.64'E. 602-620 m, 1.10.1994: 4 specimens (MNHN 1997-692).
Stn CP 1080. 15 57.30'S, 167°27.73'E. 799-850 m, 5.10.1994: 2 specimens (MNHN 1995-858)
(MNHN 1997-e686)ia MUS°RST°M 7: S,n CP 623' l2°34 02'S- '78015'W, 1300-1280 m. 28.05.1992: 1 specimen
Bathus 1: Stn CP 660, 21°10.48'S, 165°53.19'E. 786-800 m, 13.03.1993: 1 specimen (MNHN 1997-688)
27^LH,O6TI'tl?cASP9R0°7«)M" “T ‘°2- 24"32-97'S' '61"53 65'E' 1060'1,3° *»•'»* 1
Remarks. — Widespread in the Pacific and Indian Oceans; fairly common off New Caledonia.
Nezumia spinosa (Gilbert & Hubbs, 1916)
Ne7ltnTWSa ' IWAM0T° & MERRETTl 199?; 52'545' Fl§- 29b Norfolk & Loyally Ridges. Wallis & Futuna Islands,
Source : MNHN , Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
773
Material examined. — I specimen.
Norfolk Ridge. Halipro 2: stn BT 35, 25°23'S, 168°56.99'E, 640-740 m, 12.11.1996: 1 specimen 45 mm HL.
250+ mm TL (CAS 90698).
Remarks. — A widespread Indo-West Pacific species, uncommon in New Caledonian waters.
Genus PSEUDONEZUMIA Okamura, 1970
Pseudonezumia parvipes (Smith & Radcliffe, 1912)
This species was not represented in recent collections.
Pseudonezumia pusilla (Sazonov & Shcherbachev, 1982)
Pseudonezumia pusilla - IWAMOTO & MERRETT. 1997:547-548, Fig. 30b (1 spec.. Loyalty Islands).
Material examined. — l specimen.
Vanuatu. MUSORSTOM 8: stn CP 1125, 15°57.63'S, 166°38.43'E, 1160-1220 m, 10.10.1994: 1 specimen 31.7 mm HL
(MNHN 1997-725).
REMARKS. — The Vanuatu specimen is only the second known from the New Caledonian region, the first having
been collected on the Loyalty Ridge. The species is uncommon but widespread in the tropical Indo-Pacific.
Genus SPHAGEMACRURUS Fowler. 1925
Sphagemacrurus pumiliceps (Alcock, 1894)
Sphagemacrurus pumiliceps - IWAMOTO & Merrett, 1997: 549-550, Fig. 31a (New Caledonia, Loyalty Islands.
Chesterfield & Bellona Plateau, Wallis & Futuna Islands). — Grandperrin et al, 1997: 1 17 (New Caledonia; list).
Material examined. — 24 specimens.
Vanuatu. MUSORSTOM 8: stn CP 990. 18°51.63'S, 168°50.98'E, 980-990 m, 24.09.1994: 2 specimens 24-32 mm
HL (MNHN 1995-997). — Stn CP 991, 18°51.26'S, I68°52.19'E, 936-910 m. 24.09.1994: 1 specimen 32 mm HL
(MNHN 1997-701). — Stn CP 992, 1 8°52.34'S, 168°55.16'E, 775-748 m. 24.09.1994: 1 specimen 33 mm HL (MNHN
1997-683). — Stn CP 996, 18°52.4I’S, 168°55. 19E. 764-786 m, 24.09.1994: 4 specimens 27-34 mm HL (MNHN
1995-977). — Stn CP 1008. 18°53.29'S, 169°52.65'E, 919-1000 m, 24.09.1994: 3 specimens 15-33 mm HL (MNHN
1997-717). — Stn CP 1037, I8°03.70'S, I68°54.40'E, 1058-1086 m, 29.09.1994: 1 specimen 30 mm 1IL (MNHN
1995.982). — Stn CP 1075, 15°53.26'S, 167°27.21'E, 956-944 m, 4.10.1994: 1 specimen 37 mm HL (MNHN 1995-
859).
New Caledonia. Bathus 1: stn CP 660. 21°10.48'S, 165°53.19'E, 786-800 m. 13.03.1993: 4 specimens 20-
34 mm HL (MNHN 1997-689).
Norfolk Ridge. Halipro 2: stn BT 55, 25°02.85’S, 168°45.42'E, 1098-1480 m, 17.11.1996: 3 specimens 30-
38 mm HL. 162+-275 mm TL (MNHN 1998-240), I specimen 35 mm HL. 203+ mm TL (CAS 90754). — Stn BT 58.
25°04. 13’S, 168°45.21'E, 1303-1500 m, 17.11.1996: 1 specimen 31.5 mm HL. 250 mm TL (CAS 90750).
Lord Howe Rise. Halipro 2: stn BT 104, 25°23.01'S. 162°35.4'E, 1118-1124 m, 27.11.1996: 2 specimens 18-
27 mm HL. 138-1 161+ mm TL (CAS 90854).
Remarks. — Sphagemacrurus pumiliceps is a widespread Pacific and Indian Ocean species taken on numerous
occasions (but never in quantity) in the Vanuatu and New Caledonian regions. It was captured only three times
during HALIPRO 2, possibly owing to the larger trawl used during that cruise.
Genus TRACHONURUS Gunther. 1887
Trachonurus gagates Iwamoto & McMillan, 1997
Fig. 27
Trachonurus gagates Iwamoto & McMillan, 1997: 255-259, Figs 1-2 (Australia & New Zealand).
774
N.R. MERRETT & T. 1WAMOTO
Material examined. — l specimen.
Lord Howe Rise. Halipro 2: stn BT 104. 25°23.01'S, 162°35.4'E, 1118-1124 m. 27.11.1996: 1 specimen,
56 mm HL, 395+ mm TL (CAS 90666).
DIAGNOSIS. — Grooved lateral line absent; least number of scale rows from pelvic fin base to gill cover 11-14;
snout bluntly rounded, scarcely protruding beyond mouth; chin barbel 4-8% HL; suborbital width 13-15% HL;
color dark brown to blackish overall.
Size. — Attains about 48 cm TL.
Distribution. — Australia, New Zealand, New Caledonia EEZ off Lord Howe Rise. Depth range 801 to
1240 m.
a
Fig. 27. — Trachonorus gagaies Iwamoto & McMillan, 1997: (a). Lateral view of head and trunk of holotype (AMS
1.24059-009; 70.2 mm HL, 430+ mm TL) from off New South Wales, Australia, in 978 m depth; (b). Paratype, CAS
82134 (40 cm TL), from off New South Wales, Australia, in 1150 m. Scale = 25 mm.
Remarks. See original description for ligure and detailed description and comparisons with other species.
Trachonurus gagaies is relatively common at appropriate depths off almost the entire coast of Australia and New
Zealand. The Halipro 2 specimen is the first and only record from the New Caledonia region, where it is
apparently at the northern limit of its range. Off eastern Australia the species is known from off Queensland as far
north as latitude 18°S, but off Western Australia, its range extends northward only to 33° 1 6'S.
The number of Trachonurus species represented in the New Caledonian region is yet to be fully resolved, as
IWAMOTO & MERRETT (1997: 552) have recorded four specimens that may represent an undescribed species,
similar to or identical with a species found in Australian waters.
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
775
Trachonurus sentipellis (Gilbert & Cramer, 1897)
Trachonurus sentipellis - IwamOTO & MERRETT, 1997: 551-55, Fig. 31b (Hawaii. New Caledonia. Loyalty Islands.
Chesterfield & Bellona Plateau. Wallis & Futuna Islands). — GRANDPERRIN et al.. 1997: 1 17 (New Caledonia; list).
Material examined. — 6 specimens.
Vanuatu. MusCRSTOM 8: stn CP 990. 18°51.63'S, 168°50.98'E. 980-990 m, 24.09.1994: I specimen (MNHN 1995-
880). — Stn CP 991, 18°51.26'S. 168°52.19'E. 936-910 m. 24.09.1994: I specimen (MNHN 1995-875). —
Sin CP 1125. 18°52.41'S, 168°55.73'E. 764-786 m, 10.10.1994: I specimen (MNHN 1995-902). — Stn CP 1129.
1 6°00.73'S. 166°39.94'E, 1014-1050 m. 10.10.1994: 1 specimen (MNHN 1995-857).
Loyalty Ridge. Halipro 2: stn BT 28. 25°23.65'S, 170Q29.86’E. 960-1011 m, 11.11.1996: 1 specimen 78 mm
HL. 415+ mm TL (CAS 90867).
Norfolk Ridge. HaLIPRO 2: stn BT 75, 24°13'S, I67°36'E, 1128-1150 m. 21.09.1996: 1 specimen 66 mm HL.
375+ mm TL (CAS 90745).
REMARKS. — The species is relatively uncommon off New Caledonia and rarely taken in numbers. The
Loyalty Ridge specimens are the farthest south T. sentipellis has been captured.
Genus VENTRIFOSSA Gilbert & Hubbs, 1920
Ventrifossa atherodon (Gilbert & Cramer, 1897)
Ventrifossa atherodon - Iwamoto & Merrett, 1997: 554-556, Fig. 33a (New Caledonia. Chesterfield & Bellona Plateau.
Norfolk & Loyalty Ridges. Wallis & Futuna Islands).
Material examined. — 18 specimens.
Vanuatu. MUSORSTOM 8: stn CP 993, 18°48.78'S, 168°54.04'E. 780-783 m, 24,09.1994: 2 specimens 50.4-
59.0 mm HL, 215+-310 mm TL (MNHN 1995-834). — Stn CC 996. 18°52.4TS, I68°55.73'E. 764-786 m, 24.09.1994:
5 specimens 56.6-68.7 mm HL, 280+-320+ mm TL (MNHN 1995-903), I specimen 28.7 mm HL. 152+ mm TL (MNHN
1997-714). _ Stn CP 1028, 1 7°54.0 1 'S, 168°40.42’E, 624-668 m, 28.09.1994: 1 specimen 62.7 mm HL (MNHN
1995-904). — Stn CP 1033, 17°54.75'S, I68°40.66'E. 650-691 m, 29.09.1994: 2 specimens 57.5-61.0 mm HL. 312+-
307+ mm TL (MNHN 1995-901), I specimen 22.7 mm HL, 120+ mm TL (MNHN 1997-719). — Stn CP 1035.
17°56.02'S, 1 68°44.06'E. 765-780 m, 29.09.1994: 5 specimens 31.7-52.4 mm HL (MNHN 1995-990). — Stn CP 1036.
18°01.00'S, 168°48.20'E, 920-950 m, 29.09.1994: 1 specimen 53.0 mm HL. 290+ mm TL (MNHN 1995-865).
REMARKS. — This species was captured in several trawls off Vanuatu, but it was absent during the Halipro 2
cruise even though it was taken off the Norfolk and Loyalty Ridges on previous cruises.
Ventrifossa johnboborum Iwamoto, 1982
Ventrifossa johnboborum - Iwamoto & Merrett, 1997: 556-557. Fig. 34 (New Caledonia. Norfolk and Loyalty Ridges,
Chesterfield and Bellona Plateau, Wallis and Futuna Islands). — Grandperrin et al., 1997: 117 (New Caledonia; list).
Material examined. — 14 specimens.
Vanuatu. Musorstom 8: stn CP 996, 18°52.4’S, 168°55.73'E. 764-786 m. 24.09.1996: 1 female specimen. 36 mm
HL, 153+ mm TL (MNHN 1997-685). — Stn CP 1035, 17°56.02'S. 168°44.06'E, 765-780 m, 29.09.1994: 1 specimen
26 mm HL. 130+ mm TL (MNHN 1997-722).
New Caledonia. Bathus 1: stn CP 663, 20°58.66’S, 165°38.27’E. 755 m, 13.03.1993: 1 male specimen 56.2 mm
HL, 256+ mm TL (MNHN 1995-811) [large Diaphus lantemfish in stomach].
Loyalty Ridge. Halipro 2: stn BT 07, 23°41.49'S 169°54.4'E. 897-994 m. 7.11.1996: 1 specimen 104 mm HL,
400+ mm TL (MNHN 1998-238). — Stn BT 13. 24°18.03'S, 169°57.5’E. 805-835 m, 8.11.1996: I specimen 95 mm HL.
485+ mm TL (MNHN 1998-248). — Stn BT 17, 24°34.35'S, I70°11.45'E, 935-936 m. 8.11.1996: I specimen 103 mm
HL, 500+ mm TL (MNHN 1998-321). — Stn BT 18, 24°34.42'S, I70°12.62'E. 946-988 m, 9.11.1996: 1 specimen
84 mm HL (CAS 90688). 1 specimen (MNHN 1998-249).
776
N.R. MERRETT & T. IWAMOTO
Norfolk Ridge. Halipro 2: sin BT 62. 24°41.63'S, 168°38.98'E, 720-1048 m. 18. 1 1.1996: I specimen 89 mm
HL. 450+ mm TL (MNHN 1998-241), 3 specimens (CAS 90672). — Sin BT 64, 24°41.54'S. I68°40.88'E. 800-1240 m,
19.11.1996: 1 specimen (CAS 90669).
Lord Howe Rise. Halipro 2: sin BT 101. 24°16.82'S, 161°45.05'E. 970-1063 m. 26.11.1996: 1 specimen (CAS
90665).
Remarks. — A widespread species, taken in 10 trawls during Halipro 2, but never more than four specimens
in a trawl.
Ventrifossa macropogon Marshall. 1973
Venirifossa macropogon - IwAMOTO & Merrett, 1997: 557-559. Fig. 35 (New Caledonia, Norfolk & Loyalty Ridges). —
Grandperrin el al„ 1997: 1 17 (New Caledonia; list).
Material examined. — 2 specimens.
Norfolk Ridge. Halipro 2: stn BT 35, 25°23'S, 168°56.99'E, 640-740 m. 12.11.1996: 2 specimens 45-52 mm
HL, 250+-305+ mm TL (MNHN 1998-254).
Ventrifossa nigrodorsalis Gilbert & Hubbs, 1920
Fig. 28
Ventrifossa nigrodorsalis - IWAMOTO & MERRETT, 1997: 559-562, figs. 36, 37a (38 spec.. New Caledonia. Norfolk &
Loyalty Ridges, Loyalty Islands, Chesterfield & Bellona Plateau).
11-20 21-30 31-40 41-50 51-60 61-70
H L groups (mm)
Fig. 28. Length-frequency (HL, head length mm) distribution of Venirifossa nigrodorsalis by sex (n = 92).
Material examined. — 32 specimens.
Vanuatu. Musorstom 8: stn CP 975, I9°23.60'S, 169°28.93'E, 566-536 m, 22.09.1994: I specimen 44,3 mm HL.
232+ mm TL (MNHN 1995-966) and I specimen 35.0 mm HL. 187+ mm TL (MNHN 1995-985). — Stn CP 994.
I8°47.72'S, 1 68°56. 1 7'E, 649-641 m, 24.09.1994: 1 specimen 19.1 mm HL 112 mm TL (ex MNHN 1995-873)
Stn CC 996, 18°52.41'S. 168°55.73'E, 764-786 m, 24.09.1994: 4 specimens 52.4-64.5 mm HL, 251+-344+ mm TL
(MNHN 1995-903). -Stn CP 1007, 18°51.97'S, 168°55.92'E. 720-830 m, 25.09.1994: 4 specimens 21.4-55 6 mm
HL, 108+-281+ mm TL (MNHN 1995-885). — Stn CP 1033, 17°54.75'S, 168°40.66’E, 650-691 m, 29.09.1994:
1 specimen 48.9 mm HL, 255+ mm TL (MNHN 1995-521). — Stn CP 1035, 17°56.02'S, 168C44.06'E. 765-780 m,
29.09.1994: 1 specimen 66.4 mm HL, 325+ mm TL (ex MNHN 1995-990). — Stn CP 1053, I6°29.3'S 167°58.70'E,
536-519 m, 1.10.1994: 1 specimen 43.6 mm HL. 232+ mm TL (MNHN 1995-868). — Stn CP 1054, 16°27.95’s!
Source : MNHN , Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
111
167°57.44'E, 522-527 m, 1. 10.1994: 1 specimen 40.7 mm HL. 232+ mm TL (MNHN 1995-848). 1 specimen 23.5 mm
HL 145 mm TL (MNHN 1995-983). — Stn CP 1055. 16°30.H'S, 167°55.13'E. 572-580 m. 1.10.1994: 1 specimen
44.2mm HL. 145+ mm TL (MNHN 1995-871). — Stn CP 1056, 16°33.H'S, 167°55.64'E, 602-620 m. 1.10.1994:
4 specimens (MNHN 1995-842). — Stn CP 1124, 15°01.7'S, 166°56.5rE, 532-599 m. 9.10.1994.
New Caledonia. Bathus 1: stn CP 657, 21°14.45 S, 165°54.93'E. 490-530 m. 12.03.1993: 1 specimen 46.9 mm
HL, 257+ mm TL (MNHN 1995-819). — Stn CP 660, 21°10.48,S. 165°53.19 E. 786-800 m, 13.03.1993: 2 specimens
54.3-60.1 mm HL, 255+-309+ mm TL (MNHN 1995-821). — Stn CP 671, 20°51.18'S. 165°28.17'E. 450-470 m.
14.03.1993: 6 specimens 27.1-160.2 mm HL, 115+-332+ mm TL (MNHN 1995-814). — Stn CP 708. 21°43.05'S.
166°38.57'E, 550-580 m, 19.03.1993: 1 specimen 53.8 mm HL, 300+ mm TL (MNHN 1995-822).
Wallis and Futuna Islands. Musorstom 7: stn CP 623, 12°34.2’S, 1780I5.1'W, 1300-1280 m. 28.05.1992:
1 specimen 38.9 mm HL. 225+ mm TL (ex MNHN 1995-811).
Remarks. — Another widespread, abundant species of the western Pacific. The overall length-frequency
distribution of specimens among the total collection (Fig. 28) indicates a uinmodal pattern, with no apparent
sexual dimorphism in size over the range sampled.
Venlrifossa paxtoni Iwamoto & Williams, 1999
Ventrifossa species (NSW) - Iwamoto & Merrett, 1997: 564-565, Fig. 33b (12 spec.. New Caledonia, Chesterfield and
Bellona Plateau).
Venlrifossa paxtoni Iwamoto & Williams, 1999: 228-231, Fig. 55.
MATERIAL EXAMINED. — 59 specimens.
Vanuatu. Musorstom 8: stn CP 975, 19°23.60'S, 169°28.93'E, 566-536 m. 22.09.1994: 1 specimen 63 mm HL.
327 mm TL (MNHN 1995-966). — Stn CP 991, 18°51.26'S, 168°52.19'E, 936-910 m, 24.09.1994: 1 specimen 86 mm
HL. 485 mm TL (MNHN 1995-882), 1 specimen 63.1 mm HL, 327+ mm TL (MNHN 1995-996). — Stn CP 994.
1 8°47.72'S, 168°56.17'E, 649-641 m, 24.09.1994: 1 specimen (MNHN 1997-718). — Stn CC 996. 18052.41'S.
168°55.73'E, 764-786 m. 24.09.1994: 4 specimens (MNHN 1997-581). — Sin CP 1007, 18°51.97'S. I68°55.92'E,
720-830 m. 25.09.1994: 4 specimens (MNHN 1995-885). — Stn CP 1033, 17°54.75'S. 168°40.66’E, 650-691 m,
29.09.1994: 1 specimen (MNHN 1997-720). — Stn CP 1035, 17°56.02'S. 1 68°44.06'E, 765-780 m, 29.09,1994:
1 specimen (MNHN 1997-723). — Stn CP 1049, 16°39.43'S, 168°02.97'E, 469-525 m, 1.10.1994: 1 specimen (MNHN
1995-985). — Stn CP 1050, 16°39.20'S, 168°01 ,30'E, 541-577 m, 1.10.1994: 1 specimen (MNHN 1995-961).
— Stn CP 1052, 16°32.37'S, 168°00.29'E, 561-564 m, 1.10.1993: 3 specimens (MNHN 1995-905). — Stn CP 1053,
1 6°29.3'S, 1 67°58.70'E, 536-519 m, 1.10.1994: 1 specimen (MNHN 1995-868). — Stn CP 1054. 16°27.95'S.
1 67°57.44’E, 522-527 m: 1 specimen (MNHN 1995-848), 1 specimen (MNHN 1995-983). — Stn CP 1055, 16°30.H'S.
167°55.13'E, 572-580 m, 1.10.1994: 1 specimen (MNHN 1995-871). — Stn CC 1056. 16°33. 1 1 S. I67°55.64'E. 602-
620 m, 1.10.1994: 4 specimens (MNHN 1995-842). — Stn CP 1080. 15°57.30'S. 167°27.73'E. 799-850 m, 5.10.1994:
1 specimen 64.6 mm HL. 300+ mm TL (MNHN 1995-886). — Sin CP 1124, I5°0I.7'S. I66°56.51'E. 532-599 m.
9.10.1994: 6 specimens (MNHN 1995-843).
New Caledonia. Musorstom 7: stn CP 623, 12°34.02'S, 178°15'W, 1300-1280 m. 28.05.1992:1 specimen
(MNHN 1997-713).
Bathus 1: stn CP 657, 21°14.45'S, 165°54.93’E, 490-530 m. 12.03.1993: I specimen (MNHN 1995-819). —
Stn CP 660, 21°10.48'S, 1 65°53. 19 E, 786-800 m, 13.03.1993: 2 specimens (MNHN 1995-821). — Stn CP 671.
20°5 1 . 1 8'S, 165°28. 17'E, 450-470 m, 14.03.1993: 6 specimens (MNHN 1995-814). — Stn CP 702, 20°55.97'S.
1 65°34.67'E, 591-660 m, 18.03.1993: 5 specimens (MNHN 1995-818). — Stn CP 708. 2I°43.05'S. I66°38.57'E. 550-
580 m. 19.03.1993: 1 specimen (MNHN 1995-822).
Norfolk Ridge. Halipro 2: stn BT 2, 23°I7.54’S, 167°51.36'E. 1025-1049 m, 5.11.1996: 1 specimen 72 mm HL
385 mm TL (CAS 90617), I specimen 70 mm HL (CAS 90689). — Stn BT 73. 24°52.96'S, 167°37.41E. 1043-1102 m.
20.11.1996: I specimen 63 mm HL 315+ mm TL (MNHN 1998-247). — Stn BT 76. 24°05.43'S, 167°34.29'E. 936-
940 m, 21.11.1996: 2 specimens 62-79 mm HL, 340-415+ mm TL (CAS 90684). — Stn BT 90. 23°16.98'S.
168°08.38'E, 975-987 m. 23.1 1.1996: I specimen 78 mm HL, 380+ mm TL (BMNH 1 997.5.2 1 .37).
Loyalty Ridge. Halipro 2: stn BT 17. 24°34.35'S, 1 70° 1 1 ,45'E, 935-936 m. 8.11.1996: 2 specimens 76-76 mm
HL, 32S+-375+ mm HL (MNHN 1998-319). — Stn BT 18. 24°34.42'S, 1 70°12.62'E, 946-988 m, 9.1 1.1996: 1 specimen
(NMNZ P.34258).
REMARKS. — This newly described species was surprisingly abundant in these current collections, but not well
represented in previous ones reported in IWAMOTO & MERRETT (1997). It was captured on 15 occasions off
Vanuatu, six times off New Caledonia, four times off Norfolk Ridge, and twice off the Loyalty Ridge.
Source :
778
N.R. MERRETT & T. IWAMOTO
Ventrifossa vinolenta Iwamoto & Merrett, 1997
No representative of this species was found in current collections.
Subfamily MACROUROIDINAE
Genus SQUALOGADUS Gilbert & Hubbs. 1916
Squalogadus modificatus Gilbert & Hubbs, 1916
Fig. 29
Squalogadus modificatus - Iwamoto & Merrett. 1997: 482-483. Fig. 5 (New Caledonia; note that the station numbers
have been confused in the Material Examined: MNHN 1994-928 is from Stn CP 72, MNHN 1994-927 is from
Stn CP 60).
Material examined. — 15 specimens.
Norfolk Ridge. Halipro 2: stn BT 32, 25°19.93'S, 168°56.56'E, 697-1340 m. 12.11.1996: 1 specimen 105 mm
HL, 340+ mm TL (CAS 90746). — Stn BT 33, 25°19.30'S, 168°57.56'E, 699-1280 m, 14.11.1996: 1 specimen (NMNZ
P.34254). — Sin BT 41, 25°47.77'S, 167°1 1.52'E, 1317-1383 m. 14.11.1996: I specimen 110 mm HL. 325+ mm TL
(BMNH 1997.5.21.23). — Stn BT 47, 25°38.43'S. 167°38.36'E. 1441-1467 m, 15.11.1996: 2 specimens. 49-70 mm
HL, 135+-I95+ mm TL (BMNH 1997,5.21:3-4). — Stn BT 67, 24°46.21'S, I68°14.99'E. 978-1330 m. 19.11.1996:
2 specimens 70-80 mm HL, 220+-250+ mm TL (BMNH 1997.5.21:5-6).
Loyalty Ridge. Halipro 2: stn BT 16. 24°33.85'S, 170°06.47’E, 1019-1200 m, 8.11.1996: 1 specimen 107 mm
HL, 280+ mm TL (MNHN 1998-242). — Stn BT 23, 25°04.95'S, I70°11.7'E. 1105-1360 m, 10.11.1996: 1 specimen
(MNHN 1998-250). — Stn BT 25, 25°16.64'S, 170°23.69'E. 1100-1348 m, 11.11.1996: 1 specimen 128 mm HL.
360+ mm TL (BMNH 1997.5.21.72), 2 specimens 115-125 mm HL. 330+-342+ mm TL (CAS 90618). 3 specimens
(NMNZ P.34069).
31-60 61-90 91-120 121-150 151-180
H L groups (mm)
Fig. 29. — Length-frequency (HL. head length mm) distribution of Squalogadus modificatus by sex (n = 64).
Source : MNHN, Paris
GRENADIERS FISHES OF THE NEW CALEDONIAN REGION
779
REMARKS. — Squalogadus modificatus was captured in 29 trawls during the HALIPRO 2 cruise; four of those
trawls were on the Loyally Ridge, 19 on the Norfolk Ridge, and five on the Lord Howe Rise, in depths ranging
from 697 to 1520 m. Although most captures ranged between one and eight adult specimens, on one occasion 79
specimens were collected. In contrast, the species was taken only twice during all previous ORSTOM cruises, and
those captures consisted of two small juveniles. This is an obvious testimony to the effectiveness of large trawls
with high mouth openings, such as used on the RV “ Tangaroa ”, in capturing species that probably hover well
above bottom and thus evade smaller nets that fish tightly on the bottom. Squalogadus modificatus is probably
distributed throughout the New Caledonian region. On a worldwide basis, it is found almost circumtropically
(excluding the Pacific coasts of the Americas).
Squalogadus modificatus from the HALIPRO 2 cruise suggested a bimodal length frequency distribution with
females attaining the larger adult sizes (Fig. 29).
ACKNOWLEDGMENTS
We echo here our gratitude expressed earlier (Iwamoto & MERRETT, 1997) to a wide variety of people and
organizations who assisted us throughout the complete project. Pivotal in this help was Bernard SERET and his
colleagues at the MNHN, who made the material available to us and provided generous hospitality during visits to
Paris to examine the material. We thank the NATO International Scientific Exchange Programmes for their
continued support from Collaborative Research Grant (no. 940670) that enabled us to travel to the institutions at
which the material was housed. Bernard SERET also arranged for us, among others, to participate in the Halipro 2
survey of the RV “ Tangaroa ", which was funded by the ZoNeCo programme of evaluation of the marine resources
of the economic zone of New Caledonia. We therefore also wish to thank the cruise leader. Rene GRANDPERRIN,
and all the partners of that programmme. Moreover, the captain, scientific complement and crew of the "Tangaroa"
made our participation both most productive and enjoyable and none more so than the NIWA representatives on the
cruise, Malcolm Clark and Dianne Tracey. Wc thank them all.
For the loan of other specimens and assistance during visits associated with this study we express our gratitude
to: D. HOESE, M. McGROUTHER and J. PAXTON (AMS); A. WILLIAMS, A. GRAHAM and P. LAST (CSIRO
Munro Ichthyological Collection); M. GOMON (MOV); O. CRIMMEN (NHM); P. McMillan (NIWA);
R. McPhee, C. Paulin, C. Roberts and A. Stewart (NMNZ); J. Johnson and R. J. McKay (QM).
Travel funds to enable TI to participate in the Halipro 2 cruise were kindly provided by the Lindsay Research
Fund of the CAS, while his visit to the NMNZ was made possible by support from the New Zealand Foundation
for Research Science and Technology contract MNZ603 to Dr Clive ROBERTS (Programme Leader). Museum of
New Zealand Te Papa Tongawera.
We appreciate the help given by R. Hall (University of London) from his experience in the complex location
of Tectonic Plate boundaries in the New Caledonia region.
The American Society of Ichthyologists and Herpetologists kindly granted us permission to use Figs 13 and 14
(both previously published in Copeia ), as did the California Academy of Sciences for permission to use Figs 15.
18, 19 and 24 (all previously published in the Proceedings of the California Academy of Sciences). We thank them
both.
Finally we thank Alison E. SCHROEER for drawing Fig. 22 of Kumba musorstom.
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Source : MNHN, Paris
LTATS DES CAMP AGNES MUSORSTOM. VOLUME 21 — RESULTATS des campagnes musorstom. volume 21 — RESULTATS DES c
Pisces Pleuronectiformes: Flatfishes from
New Caledonia and adjacent waters.
Genus Arnoglossus
Kunio AM AO K A
Laboratory of Marine Zoology
Faculty of Fisheries
Hokkaido University
Hakodate, Hokkaido
041-8611, Japan
&
Eiji MIHARA
Resources Management Section. Muroran Branch
Hokkaido Hakodate Fisheries Experimental Station
Funami-cho, Muroran. Hokkaido
051-0013, Japan
ABSTRACT
Species of the bothid genus Arnoglossus collected from waters around New Caledonia are reviewed. Seven species,
including two new species, two new zoogeographical records and three species already recorded from the region were
identified, being Arnoglossus septemveniralis sp. nov. and A.nigrifrons sp. nov.. A. tenuis. A. elongatus. and
A. macrolophus, A. japonicus and A. polyspilus, respectively. Arnoglossus septemveniralis sp. nov., described from
ten specimens collected between 230-315 m off southern New Caledonia, is easily separable from all other members of
the genus in having seven pelvic rays on both sides. Arnoglossus nigrifrons sp. nov., described from two specimens
collected from 300-315 m on the Chesterfield Plateau and northwest of New Caledonia, is characterized by a rounded upper
head profile, several anterior dorsal fin rays elongated in males, gill rakers without serrations and a darkened head region.
Arnoglossus tenuis, collected from 10-16 m off New Caledonia, was previously known from southern Japan to the South
China Sea, and A. elongatus. from 250-350 m off New Caledonia, previously only from the Madura Sea and northwestern
Australia. Arnoglossus macrolophus was collected from relatively shallow waters (49-92 m) off New Caledonia, and
A. japonicus and A. polyspilus from deeper waters (210-385 m) off New Caledonia, the Loyalty Islands and Chesterfield
Plateau.
Amaoka. K. & Mihara. E„ 2000. — Pisces Pleuronectiformes: Flatfishes from New Caledonia and adjacent waters. -
Genus Arnoglossus. In: A. CROSNIER fed.), Resultats des Campagnes MUSORSTOM, Volume 21. Memoires du Museum
national d'Histoire naturelle. 184: 783-813. ISBN-2-85653-526-7.
* This paper was presented at the Indo-Pacific Fish Conference. Noumea (ORSTOM), 1997.
784
K. AMAOKA & E. M1HARA
RESUME
Pisces Pleuronectiformes : Poissons plats de la Nouvelle-Caledonie et des eaux avoisi-
nantes. - Genre Amoglossus .
Les espfcces du genre Amoglossus de la famille Bolhidae, recoltees autour de la Nouvelle-Caledonie, sont etudiees.
Sept espfeces dont deux sont nouvelles ( Amoglossus septemventralis sp. nov. and A.nigrifrons sp. nov.), trois sonl
mentionnees pour la premiere fois de la region (A. tenuis, A. elongatus et A. macrolophus) et trois dtaient ddja connues de
la region (A. macrolophus, A. japonicus et A. polyspilus ), ont ete identifies. Amoglossus septemventralis sp. nov.,
decrit d'apres 10 exemplaires recoltes entre 230 et 315 m de profondeur au sud de la Nouvelle-Caledonie. se distingue
ais6ment de toutes les autres especes du genre par la presence de 7 rayons pelviens sur chacune de ses faces. Amoglossus
nigrifrons sp. nov., decrit d'apres deux specimens icoltds entre 300 et 315 m sur le plateau des lies Chesterfield, au nord-
est de la Nouvelle-Caledonie. se caracterise par une tete a profil supcrieur bombe, plusieurs rayons anterieurs de la dorsale
allonges chez les males, des arcs branchiaux non dentelds et des zones sombres sur la tete. Amoglossus tenuis, recolte par
10-16 m de profondeur au large de la Nouvclle-Caidonie, n'etait auparavant connu que du Japon du Sud a la mer dc Chine
meridionale et A. elongatus, recolte entre 250 et 350 m, que de la mer de Madura et du nord-ouest dc I'Australie.
Amoglossus macrolophus a 6i recolte dans des eaux relativement peu profondes (49-92 m ) au large de la Nouvelle-
Caledonie. tandis que A. japonicus et A. polyspilus ont ete trouves dans des eaux plus profondes (210-385 m) au large de
la Nouvelle-Caledonie, des lies Loyautd et sur le plateau des Ties Chesterfield.
INTRODUCTION
Members of the genus Amoglossus are characterized by an elliptical body, eyes separated by a ridge or
extremely narrow concave space without any sexual dimorphism, no rostral or orbital spines, cycloid or feebly
ctenoid scales on the ocular side of the body, and the caudal skeleton with four undivided plates. At least
37 nominal species are known from shallow to deep waters of the Indian, Pacific and Atlantic Oceans, except off
the east coast of North and South America, and in the Mediterranean Sea.
In a revisional study of New Caledonian flatfishes, seven species of Amoglossus from coastal to deeper waters
were found. They included two new species, A. septemventralis sp. nov. and A. nigrifrons sp. nov., and two new
zoogeographical records, A. tenuis and A. elongatus. The remaining three species, A. macrolophus (as
A. tapeinosoma), A. polyspilus and A. japonicus were listed from the region by RlVATON (1989), RlVATON &
Richer de Forges (1990), and Kulbicki, Randall & Rivaton (1994).
In this paper descriptions and comments are given for each species, and an identification key provided for all of
them.
MATERIALS AND METHODS
Information on the cruises where the samples studied here were collected can be found in:
— Richer de Forges, Chevillon, Laboute, Menou & Tirard (1988) for Corail 2;
— Richer de FORGES (1990) for Chalcal 1 and 2, MUSORSTOM 4, 5 and 6, Smib 1 and 4;
— Richer de Forges (1991) for Lagon;
— Lehodey, Richer de Forges, Nauges. Grandperrin, Rivaton (1992) for Beryx 1 h
— Richer de Forges (1993) for Smib 5 and 6;
— Richer de Forges & Chevillon (1996) for Bathus 1 and Halipro 1.
Specimens were deposited in the Museum national d'Histoire naturelle (MNHN), Paris, Laboratory of Marine
Zoology, Faculty of Fisheries, Hokkaido University (HUMZ) and National Museum of New Zealand (NMNZ).
Specimens were fixed in 10 % formalin and preserved in 75 % ethanol. Abbreviations for other institutions follow
Leviton et al. (1985).
Methods for measurements followed Amaoka, Mihara & Rivaton (1993). Gill rakers were counted as upper
limb + lower limb. Vertebrae were counted as abdominal vertebrae + caudal vertebrae.
Source :
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
785
In the lists of material examined, the capital letters preceding the station number refer to the gear used:
DC: Charcot dredge; DW: Waren dredge, CP: beam trawl; CC: shrimp otter trawl; BT: benthic fish trawl.
SYSTEMATIC ACCOUNT
Genus ARNOGLOSSUS Bleeker, 1862
Arnoglossus Bleeker, 1862: 427 (type species by original designation. Pleuronecies arnoglossus Schneider, 1801).
Anticitharus Gunther, 1880: 47 (type species by original designation, Anlicitharus polyspilus Gunther, 1880).
Caulopsetia Gill, 1893: 124 [type species by original designation. Pleuronecies scaplius (Forster) Schneider, 1801).
Scidorhombus Tanaka. 1915: 567 (type species by original designation, Scidorliombus pallidus Tanaka. 1915).
Diagnosis. — Body elliptical. Rostral and orbital spines absent. Interorbital space with narrow bony ridge or
narrowly concave in both sexes. Mouth moderate or rather large in size, upper jaw length about 2. 1-2.9 in head
length. Dentition more or less equally developed on either side. Scales ctenoid with short ctenii or cycloid on
ocular side. Lateral line developed only on ocular side. Pelvic fin on ocular side originating near tip of isthmus;
base on ocular side longer than that on blind side. Four caudal plates (i.e., parhypural, hypurals, and hypural +
epural) not splitted distally.
Fig. 1. — Diagrammatic illustration of caudal-fin type
with bifurcated rays. Scale bar = 10 mm. Another
type with rays branched into more than two is
shown in Fig. 2.
Remarks. — In most of the above characters,
Arnoglossus closely resembles Psettina. However,
Psettina has ctenoid scales on the ocular side with slender,
long ctenii (vs. scales with short ctenii or cycloid) and a
relatively small mouth, more than 2.7 in head length (vs.
less than 2.9).
Comparative Material examined. — Arnoglossus
oxyrhynchus: FAKU 33193-33198, 33208-33212.
holotype and paratypes, 6 males and 5 females, 150.3-
194.1 mm SL, Mimase, Kochi Pref., 10 December 1958;
FAKU 33726. paratype, female, 186.9 mm SL. Mimase.
7 December 1959.
A. polyspilus: BMNH 1879.5.14:56, syntype, female,
190.0 mm SL, Kei Island, 236 m.
A. elongatus : BMNH 1930.10.16.1, syntype, male,
75.0 mm SL. 7°02.6'S, 1 15°23.6’E, Madura Sea, 100 m.
A. yamanakak NSMT P.45060. 45063, male and
female, 111.7-118.8 mm SL, 28°57.6'N. 124°06.0'E, East
China Sea, 82 m, trawl. 30 October 1986. R.V. "Yoko-
maru", NSMT P.45062, female, 120.9 mm SL,
28°56.5'N, 124°52.9'E, East China Sea, 90 m, trawl, 30
October 1986, R.V. "Yoko-maru", NSMT P.45070, male.
104.5 mm SL, 29°I3.3'N, 124°14.6'E, East China Sea, 74 m, trawl. 22 October 1986, R.V. "Yoko-maru".
Key to New Caledonian species of Arnoglossus
Al. Pelvic fin rays 7 . A. septemventralis sp. nov.
A2. Pelvic fin rays 6.
786
K. AMAOKA & E. MIHARA
B 1. Anterior dorsal fin rays greatly elongated, length of elongated fin rays greater than head
length (Fig. 8). Dorsal and anal fins with a large dark spot posteriorly at base (Fig. 7) ..
. A. macrolophus Alcock, 1889
B2. Anterior dorsal fin rays not or only moderately elongated, length of anterior fin rays
less than head length. Dorsal and anal fin bases without distinct spot posteriorly.
Cl. Middle caudal fin rays bifurcated (Fig. 1).
Dl. Scales almost always cycloid on ocular side. Some anterior teeth of upper jaw
enlarged and widely spaced. Only second dorsal fin ray elongated in males .
. A. japonicus Hubbs, 1915
D2. Scales almost always ctenoid on ocular side. Teeth of upper jaw gradually
becoming larger and more widely spaced anteriorly. Second to third or fourth dorsal
fin rays elongated in both sexes (Fig. 13) . A. elongatus Weber, 1913
C2. Middle caudal fin rays with more than two branches.
El. Dorsal fin rays less than 94. Anal fin rays less than 71. Scales in lateral line less
'han 59 . A. tenuis Gunther, 1 880
E2. Dorsal fin rays more than 100. Anal fin rays more than 80. Scales in lateral line
more than 70.
FI. Gill rakers serrated on each margin. Anterior head margin, snout and chin regions
pale brownish . A. polyspilus (Gunther, 1 880)
F2. Gill rakers not serrated. Anterior head margin, snout and chin regions conspic¬
uously darkened (Figs 17-18) . A. nigrifrons sp. nov.
Fig. 2. — Illustration of Arnoglossus septemventralis sp. nov., holotype. male, 86.0 mm SL, from off southern
New Caledonia (NMNZ P. 29090).
Source : MNHN, Pans
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
787
Arnoglossus septemventralis sp. nov.
Figs 2-6; Tables 1-2
MATERIAL EXAMINED. — 10 specimens (4 males and 6 females).
New Caledonia. Chalcal 2: sin CP 26, 23°18.15'S, 168°03.58'E, 296 m, 31.10.1986: 1 6 , paralype. 69.8 mm
SL (HUMZ 142680); 2 <3 and 1 2 . paralypes. 56.8-69.8 mm SL (MNHN 1995-1171 to 1173). — Stn CP 27. 23°15.29'S.
168°04.55'E, 289 m, 31.10.1986: 1 2, paralype, 90.0 mm SL (HUMZ 142679); 1 2, paratype. 85.1 mm SL (MNHN
1995-1170).
Smib 5: stn DW 103, 23°17.4'S, 168°04.8'E, 300-315 m. 14.09.1989: 1 2, paratype. 74.6 mm SL (MNHN 1995-
1 169).
BERYX 11: stn CP 16, 24°47.r-46.9'S, \6%°0&.T-0&.S'E, 240-250 m, 16.10.1992: I 6, holotype. 86.0 mm SL
(NMNZ P.29090). — Stn CP 44, 23°41 .3'-40.5'S, 168°00.6'-00.5'E, 230-250 m, 20.10.1992: 2 2. paralypes. 52.9-
56.4 mm SL (NMNZ P.29046).
Types. — Holotype: NMNZ P.29090 (Beryx 11: stn 16, 24°47.r-46.9'S, 168°08.7'-08.8'E. 240-250 m).
Paratypes: All the other specimens cited above.
Diagnosis. — Pelvic fins with 7 rays; body shallow, depth 2.66-2.93 in SL: gill rakers serrated (Fig. 4A);
dorsal fin rays 95-97; anal fin rays 73-79; scales in lateral line 78-84.
Pig 3 — Arnoglossus septemventralis sp. nov., holotype. male, 86.0 mm SL, from off southern New Caledonia (NMNZ
P.29090).
DESCRIPTION. — Data for holotype given first, followed in parentheses by ranges for paratypes and averages,
including holotype, for proportional data. Counts and proportional measurements as percent of SL shown in
Tables 1 and 2.
Head length 3.51 (3.41-3.78, 3.63) in SL; body depth 2.70 (2.66-2.93, 2.77). Snout length 4.80 (4.49-5.34,
4.84) in head length; upper eye diameter 3.10 (2.75-3.24, 2.97); lower eye diameter 3.14 (2.71-3.18. 2.99).
interorbital width 49.00 (31.40-64.00, 46.82); upper jaw length 2.75 (2.53-2.92, 2.72) on ocular side, 2.63 (2.40-
2.82, 2.61) on blind side; lower jaw length 2.04 (1.79-2.01, 1.95) on ocular side, 1.96 (1.69-1.90, 1.84) on blind
side; depth of caudal peduncle 2.92 (2.70-3.08, 2.93); length of longest dorsal fin ray 2.36 (2.18-2.40. 2.30);
Source : MNHN. Paris
788
K. AMAOKA & E. MIHARA
length of longest anal fin ray 2.19 (2.10-2.38, 2.20); pectoral fin length 1.78 (1.65-1.83. 1.74) on ocular side,
2.72 (2.85-3.25, 3.00) on blind side; pelvic fin length (except elongated ray) 1.96 (2.06-2.49, 2.14) on ocular side
in males, (2.70-3.04, 2.84) in females, 2.75 (2.64-3.23, 2.96) on blind side; length of third pelvic fin ray 1.37
(1.79-2.23, 1.84) on ocular side in males, (3.00-3.25, 3.14) in females; length of pelvic fin base 4.02 (3.49-4.19,
3.83) on ocular side, 9.80 (7.68-9.69, 8.59) on blind side; length of middle caudal fin ray 1.30 (1.28-1.36, 1.33);
lateral line curve width 2.21 (1.84-2.15, 2.03).
Body elongated, elliptical, deepest slightly anterior to midpoint; dorsal and ventral contours gently arched.
Caudal peduncle depth 24.3-27.5 % of body depth. Head large; upper profile with a slight notch anterior to upper
eye. Snout long, length 52.7-67.2 % of upper eye diameter. Rostral and orbital spines absent. Eyes large, separated
by narrow bony ridge; upper eye diameter 81.5-103.8 % of upper jaw length on ocular side; lower eye in advance
of upper. Ocular side nostrils anterior to dorsal margin of lower eye; anterior nostril tubular with a flap posteriorly;
blind side nostrils small, near dorsal fin origin, similar in shape to those on ocular side.
Mouth large, oblique; maxilla extending posteroventrally to below anterior part of lower eye; anterior tip of
lower jaw projecting slightly beyond tip of upper jaw when mouth closed. No distinct knob at mandibular
symphysis. Dentition more or less equally developed on both jaws; teeth on upper jaw sharp, uniserial, becoming
larger and more widely spaced anteriorly. Gill rakers on first arch moderate in size, dorsal margins serrated
(Fig. 4A). Ocular side scales very small, with short ctenii (Fig. 4B); snout, anterior parts of both jaws, and
interorbital region naked; cycloid scales on blind side. Lateral line curved above pectoral fin on ocular side; lateral
line absent on blind side.
FlG- 4. — First gill arch (A) and a scale (B) from ocular side of
Arnoglossus septemvemralis sp. nov., paratypes, females
(A: 90.0 mm SL, HUMZ 142679; B: 74.6 mm SL, MNHN
1995-1169).
side, opposite side of vent.
Dorsal fin origin on blind side, anterior to
interorbital region. Anal fin origin at a point
equal with vertical through posterior margin of
head. Pectoral fins unequal; length on ocular
side 1.53-1.84 times that on blind side, much
less than head length. Pelvic fins with 7 rays;
origin of ocular side fin slightly posterior to tip
of isthmus; third ray of ocular side fin in male
elongated, longer than adjacent rays; that in
female subcqual to adjacent rays (Figs 5-6);
pelvic fin base on ocular side longer than that
on blind side, space between fourth and fifth ray
on ocular side opposite to first ray on blind
side. Tip of isthmus posterior to and ventral to
posterior margin of lower eye. All fin rays
simple, except caudal fin rays. Caudal fin rays
with more than two branches, except upper- and
lowermost two simple rays.
Vent on blind side, immediately dorsal to
first anal fin ray. Urogenital papilla on ocular
Coloration in alcohol : Ocular side ground color light brown; snout, both jaws, area in front of eyes and
interorbital region darkened; a series of dark rings along dorsal and anal margins of body; a few obscure dark spots
on lateral line. Blind side uniformly pale brown. Dorsal and anal fins with series of obscure dark blotches; ocular
side pectoral fin with three dark cross bands; distal half of fifth to seventh rays of ocular side pelvic fin darkened.
Sexual dimorphism-. This species shows sexual dimorphism in length of the ocular side pelvic fin (Figs 5-6).
Distribution. — All specimens were collected off southern New Caledonia, at depths between 230-315 m.
Etymology. Named lor the seven pelvic fin rays, which constitute the singly most important diagnostic
character of this species.
Source :
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
789
Fig. 5. — Diagrammatic illustration of anterior body parts showing sexual dimorphism in male (A) and female (B)
of Arnoglossus sepiemveniralis sp. nov. Scale bars = 10 mm.
>
■S e
”
o o
3 '>
<s> a
I"
51
o
20
15
10
5
50 60 70 80
Standard length (mm)
90
100
Fig. 6. — Relationship between SL and 3rd pelvic fin ray length on ocular side in percent ol SL in Arnoglossus
sepiemveniralis sp. nov.
REMARKS. — Arnoglossus septemventralis sp. nov. is easily separable from all congeners by the presence ot
seven rays in the pelvic fin.
According to Amaoka & IMAMURA (1990), the genus Arnoglossus can be divided into two groups of species,
based on numbers of dorsal (less than vs. greater than 100) and anal fin rays (less than vs. greater than 80).
Arnoglossus septemventralis belongs to the group with lower counts, thereby resembling A. andrewsi Kurth.
1954, A. armstrongi Scott, 1975 and A. marisrubri Klausewitz & Schneider, 1986, and also shares with the latter
three species, a large number of lateral line scales. It differs from A. andrewsi and A. armstrongi in having a
shallower body (2.66-2.93 in SL vs. 2.3-2.4 in A. andrewsi, 2.4 in A. armstrongi) and larger upper eye (2.75-3.24
Source
790
K. AMAOKA & E. MIHARA
in head length vs. 5. 0-5. 4, 4.2-4. 9) (KURTH, 1954; SCOTT. 1975), and from A. marisrubri in having a lower
number of gill rakers (8-9 on lower limb vs. 1 1-13 in A. marisrubri) and serrated gill rakers (vs. not serrated)
(KLAUSEWITZ & SCHNEIDER, 1986). Furthermore, it differs from A. andrewsi in having a higher number of the
oeularside peetoral fin rays (1 1-13 vs. 10) (KURTH, 1954).
Table 1. — Frequency distributions of selected meristic characters of Arnoglossiis septemventralis sp. nov. Counts for
holotype included in boldfaced numbers; counts from blind side shown in parentheses.
Table 2. — Proportional measurements as percent of standard length in Arnoglossus septemventralis sp. nov. Averages
include measurements from holotype. O. ocular side; B, blind side.
* except elongated third ray.
Source : MNHN, Paris
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
791
Arnoglossus macrolophus Alcock, 1889
Figs 7-9; Tables 3-4
Arnoglossus macrolophus Alcock, 1889: 280, pi. 18, fig. 2. — ALCOCK, 1890: 433; 1896: 327. — Weber. 1913:
432. — Norman, 1927: 21, fig. 3. — Fowler. 1928: 90. — Arai & Amaoka, 1996: 360. figs 1-4.
Bothus ( Arnoglossus ) tapeinosoma - Weber & BEAUFORT, 1929: 127 (not Platophrys ( Arnoglossus I tapeinosoma
Bleeker, 1866).
Arnoglossus tapeinosoma - Norman, 1934: 185, fig. 131. — Fowler, 1934: 63, fig. 18. — MUNRO. 1955: 260,
pi. 49; 1958: 131, pi. 13. — Baoshan, 1962: 970, fig. 754. — Abraham, 1963: 67. — Menon & Yazdani, 1963: 157.
— Shen, 1966: 175, fig. 42a-b; 1983: 29. fig. 41. — Dor. 1970: 25. I fig. — Amaoka, 1971: 28, pi. 3A. —
Wongratana. 1975: 28. — Kotthaus, 1977: 5. fig. 434. — Rivaton & Richer de Forges. 1990: 25, 60. — Amaoka.
Okamura & YOSH1NO, 1992: 262, fig. 5.
Material EXAMINED. — 18 specimens (10 males and 8 females).
New Caledonia. Lagon: stn 610, 22°I0.0-S, 166°58.0'E. 49 m. dredge. 5.08.1986: 1 9, 69.9 mm SL (MNHN
1995-1096). — Stn 742. 22°13.9'S, 167°02.8’E. 78 m, dredge. 13.08.1986: I 9, 60.0 mm SL (MNHN 1995-1106). —
Stn 833, 20°49.8’S. 165°17.7'E, 52-70 m, dredge. 11.01.1987: 6 6 and 3 9 , 60.2-85.2 mm SL (MNHN 1995-1097 to
1 105); 2 6 and 2 9 . 65.2-82.3 mm SL (HUMZ 142653-142656).
Bathus I: stn CP 680, 20°48.03'S. 165°17.98'E, 86-92 m. 15.03.1993: 6 . 65.6 mm SL (MNHN 1995-1 107); I 6
and 1 9, 37.8-56.3 mm SL (HUMZ 142657-142658).
DIAGNOSIS. — Anterior dorsal fin rays greatly elongated in both sexes, first dorsal fin ray longer than head
length in specimens greater than 60 mm SL (Fig. 8); dorsal and anal I ms with a large dark spot posteriorly at base
of fin rays (Fig. 7).
Fig. 1.— Arnoglossus macrolophus Alcock. 1889. male, 65.6 mm SL, from off northern New Caledonia (MNHN 1995-
1 107).
Description. — Ranges for proportional data given first, followed by averages. Counts and proportional
measurements as percent of SL shown in Tables 3 and 4.
Head length 3.74-4.20, 3.98 in SL; body depth 2.43-2.78, 2.60. Snout length 3.65-4.33, 3.98 in head length;
upper eye diameter 3.47-4.14, 3.79; lower eye diameter 3.32-4.14, 3.82; interorbital width 22.29-50.50. 32.66;
upper jaw length 2.30-2.51, 2.41 on ocular side, 2.54-2.78, 2.68 on blind side; lower jaw length 1.75-1.94. 1.84
on ocular side, 1.90-2.06, 1.98 on blind side; depth of caudal peduncle 2.27-2.66. 2.47; length of lust dorsal lin
Source
792
K. AMAOKA & E. MIHARA
ray 0.33-0.84, 0.59 in males, 0.77-1.61, 0.95 in females; length of longest dorsal fin ray except anterior elongated
rays 1.68-1.96, 1.84; length of longest anal fin ray 1.64-1.95, 1.79; pectoral fin length 1.47-1.75, 1.62 on ocular
side, 2.18-2.56, 2.35 on blind side; pelvic fin length 1.65-2.97, 2.10 on ocular side in males, 2.43-2.79, 2.58 in
females, 2.29-3.06, 2.60 on blind side; length of pelvic fin base 3.24-4.04, 3.51 on ocular side, 7.13-9.57, 8.18
on blind side; length of middle caudal fin ray 1.18-1.33, 1.27; lateral line curve width 1.82-1.99, 1.89.
Body elongated, elliptical, deepest slightly anterior to midpoint. Caudal peduncle depth about 24.3-27.9 % of
body depth. Head small; upper profile with a large notch anterior to upper eye. Snout moderate, length 87.8-
107.8 % of upper eye diameter. Rostral and orbital spines absent. Eyes moderate, separated by narrow bony ridge,
upper eye diameter 56.2-69.5 % of upper jaw length on ocular side; lower eye slightly in advance of upper. Ocular
side nostrils anterior to dorsal margin of lower eye; anterior nostril tubular with a flap posteriorly; blind side
nostrils small, near origin of dorsal fin, similar in shape to those on ocular side.
Mouth large, oblique; maxilla extending to below anterior part of lower eye; anterior tip of lower jaw
projecting slightly beyond tip of upper jaw when mouth closed. Small ventrally directed knob at mandibular
symphysis. Dentition more or less equally developed on jaws of both sides; teeth on upper jaw sharp, uniserial;
lower jaw teeth uniserial, nearly similar to anterior teeth of upper jaw in size and spacing. Gill rakers on first arch
slender, not serrated, absent on upper limb. Scales large, deciduous, with short ctenii on ocular side, cycloid on
blind side. Lateral line curved above pectoral fin on ocular side; lateral line absent on blind side.
Dorsal fin origin on blind side, anterior to upper eye; anterior 3-5 rays greatly elongated in males, length 1 .2-
3.0 times head length, anterior 2-3 rays greatly elongated in females, length 0.6- 1.3 times head length (Fig. 8).
Anal fin origin at a point equal with vertical through posterior margin of head. Pectoral fins unequal, length of
ocular side fin 1.3- 1.6 times that of blind side. Pelvic fins with 6 rays; ocular side pelvic fin rays longer in males
than those in females (Figs 8-9); approximately fourth ray of ocular side fin opposite first ray of blind side fin. Tip
of isthmus ventral to vertical through posterior margin of lower eye. Caudal fin rays with more than two branches,
except upper- and lowermost two simple rays.
Fig. 8. — Diagrammatic illustration of anterior body parts showing sexual dimorphism in male (A) and female (B)
of Amoglossus macrolophus. Scale bars = 10 mm.
Source :
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
793
Vent on blind side, immediately dorsal to first anal fin ray; urogenital papilla on ocular side, opposite side of
vent.
Table 3. — Frequency distributions of selected meristic characters of Arnoglossus macrolophus. Counts from blind side
shown in parentheses.
Table 4. — Proportional measurements as percent of standard length in Arnoglossus macrolophus. O, ocular side;
B, blind side.
* except anterior elongated rays.
Coloration in alcohol : Ocular side ground color dark brown; a few obscure dark blotches on lateral line; a series
of blotches along dorsal and anal margins. Blind side uniformly pale brown. Dorsal and anal lins with series of
dark spots basally and large dark spot at base posteriorly; ocular side pectoral tin and pelvic tin on both sides with
Source :
794
K. AMAOKA & E. MIHARA
darker margins in males than females, pectoral and pelvic fins on ocular side with dark margins or spots in females
(Figs 7-8); caudal fin with a pair of obscure blotches proximally and five blotches distally along posterior margin.
Sexual dimorphism'. The specimens showed sexual dimorphism in lengths of the ocular side pelvic fin and the
elongated anterior dorsal fin rays, the number of elongated dorsal fin rays and coloration of the pectoral and pelvic-
fins (Figs 8-9).
J2
3
o
o c
c- «=
° o
3 >
m 2.
&■ «
JC S
c
©
F 14---
12 -
30 40 50 60 70
Standard length (mm)
80
90
Fig. 9. — Relationship between SL and pelvic fin length on ocular side in percent of SL in Arnoglossus macrolophus.
Distribution. — Widely distributed from the Red Sea (Kotthaus, 1977) and Persian Gulf (Norman. 1927).
through the northern Indian Ocean (Alcock, 1889; Abraham, 1963) and Malay Archipelago (Weber, 1913) to
southern Japan (Amaoka, Okamura & Yoshino, 1992) and New Caledonia (RlVATON & RICHER DE FORGES.
1990), in depths of 18-141 m.
Remarks. — Arnoglossus macrolophus has two unique characters, the greatly elongated anterior dorsal fin
rays and the large dark spots posteriorly on both the dorsal and anal fin bases. The species is easily separable from
all other congeners by these two characters.
In A. macrolophus , the several anterior rays of the dorsal fin have been reported as greatly elongated in males,
but only slightly longer than the other rays in females. However, those of New Caledonian specimens are greatly
elongated and longer than the head length also in females. This is likely to represent geographical variation in this
species, since other characters do not differ significantly from those of specimens collected elsewhere (Amaoka,
Okamura & Yoshino, 1992; Arai & Amaoka, 1996).
Arnoglossus japonicus Hubbs, 1915
Fig. 10; Tables 5-6
Arnoglossus japonicus Hubbs, 1915: 454, pi. 25, fig. 2. — Norman, 1934: 192, fig. 139. — Okada & Matsubara
1938: 421, pi. 104, fig. 1. — Kuroda, 1951: 389; 1962: 1. — BOhlke, 1953: 139. — Matsubara. 1955: 1258. —
Amaoka, 1969: 194, figs 78-79; 1973: 145, figs 1-7; 1984: 350, pi. 369-D. — Shen, 1983: 30, fig. 42.
— Gloerfelt-Tarp & Kailola, 1984: 273, 358, 1 fig. — Rivaton, 1989: 155, 163. — Nakabo, 1993: 1 173. I fig
— Kulbicki, Randall & Rivaton, 1994: 37.
Arnoglossus oxyrhynchus - Rivaton & Richer df. Forges: 1990: 25. 60 (not Arnoglossus oxyrhynchus Amaoka, 1969).
Material EXAMINED. — 29 specimens (1 1 males, 8 females and 10 juveniles).
New Caledonia. Musorstom 4: stn CP 154, 19°02.60'S, 163°17.80'E. 275 m, 14.09.1985: 1 6 and 3 2 80 I-
105.8 mm SL (MNHN 1995-1150 to 1153).
Bathus I: stn CP 712, 21°44.26'S, 166°35.34'E, 210 m. 19.03.1993: I <J . 97.9 mm SL (HUMZ 142671)- 1 2
106.0 mm SL (MNHN 1995-1154). — Stn CP 713, 21°45.28'S, 166°36.83'E, 250 m. 19.03.1993: 1 2 106 0 mm SL
(MNHN 1995-1 155).
Loyalty Islands. Musorstom 6: stn CP 419, Lifou Island. 20°41.65'S, 167°03.70'E, 283 m, 16 02 1989-
6 juveniles, 46.4-49.8 mm SL (MNHN 1995-1146 to 1149, HUMZ 142668-142669). — Stn CP 455. 21°00.65'S.
Source
PISCES PLEU RONECTIFORMES : GENUS ARNOGLOSSUS
795
I67°26.08'E, 260 m, 20.02.1989: 2 juveniles, 47.3-48.7 mm SL (MNHN 1995-1143 to 1144). — Stn DW 456,
21°00.71'S, 167°26.35'E, 225-240 m, 20.02.1989: 1 juvenile, 49.0 mm SL (MNHN 1995-1145).
Chesterfield Plateau. Chalcal 1: stn CP 5, 19°29.10'S. I58°37.63'E, 290 m. 16.07.1984: 2 <5, 101.8-
125.5 mm SL (MNHN 1995-1133 to 1134). — Stn CP 10. 20°00.20'S, 158°46.60'E. 225 m. 22.07.1984: 2 6 and 1 2 ,
48.8-51.4 mm SL (MNHN 1995-1140 to 1142); 1 6 and 1 9 , 50.9-51.9 mm SL (HUMZ 142666-142667). — Stn CP 11,
20°04.40'S, 158°47.41'E, 300 m, 22.07.1984: 3 3, 57.0-115.8 mm SL (MNHN 1995-1135 to 1137).
Corah. 2: stn CP 131, 19°25.49'S, 158°37.96'E. 217 m, 29.07.1984: 1 <3 and 1 juvenile, 45.3-58.9 mm SL (MNHN
1995-1138 to 1139).
Musorstom 5: stn DW 350, 19°34.00'S, 158°35.30'E, 280 m, 17.10.1986: 1 2, 125.7 mm SL (HUMZ 142670).
DIAGNOSIS. — Middle caudal fin rays bifurcated (Fig. 1 ); ocular side scales mostly cycloid; some anterior teeth
enlarged and widely spaced; second dorsal fin ray elongated in males; gill rakers serrated on dorsal margins.
Fig. 10. — Arnoglossus japonicus Hubbs, 1915. female. 125.7 mm SL, from Chesterfield Plateau (HUMZ 142670).
DESCRIPTION. — Ranges for proportional data given first, followed by averages. Counts and proportional
measurements as percent of SL shown in Tables 5 and 6.
Head length 3.39-4.1 1, 3.73 in SL; body depth 2.44-3.04, 2.74. Snout length 4.00-4.67, 4.38 in head length;
upper eye diameter 3.39-4.21, 3.73; lower eye diameter 3.39-4.21, 3.70; interorbital width 25.70-1 19.00, 45.05;
upper jaw length 2.11-2.80, 2.33 on ocular side, 1.79-2.73, 2. 14 on blind side; lower jaw length 1.76-2.12, 1.89
on ocular side, 1 .59-2.05, 1 .74 on blind side; depth of caudal peduncle 2.77-3.76. 3.30; length of longest dorsal fin
ray 1.78-2.20, 2.01; length of longest anal fin ray 1.75-2.20, 1.95; pectoral fin length 1.79-2.48. 2.02 on ocular
side, 3.32-4.52, 3.88 on blind side; pelvic fin length 3.1 1-3.58, 3.35 on ocular side, 2.73-3.19, 2.97 on blind side;
length of pelvic fin base 2.95-4.03, 3.41 on ocular side, 6.76-9.34, 8.04 on blind side; length of middle caudal fin
ray 1.28-1.64, 1.46; lateral line curve width 2.28-2.63, 2.41.
Body elongated, elliptical, deepest slightly anterior to midpoint; dorsal and ventral contour gently arched.
Caudal peduncle depth 20.0-25.9 % of body depth. Head large; a slight notch anterior to ventral margin of upper
eye. Snout long, length 73.8-93.8 % of upper eye diameter. Rostral and orbital spines absent. Eyes moderate,
separated by narrow bony ridge; upper eye diameter 56.0-72.7 % of upper jaw length on ocular side; lower eye in
advance of upper. Ocular side nostrils anterior to dorsal margin of lower eye; anterior nostril tubular with a flap
posteriorly; blind side nostrils small, ventral to origin of dorsal fin, similar in shape to those on ocular side.
Source :
796
K. AMAOKA & E. MIHARA
Mouth large, oblique; maxilla extending to below middle part of lower eye; anterior tip of upper jaw projecting
slightly beyond tip of lower jaw when mouth closed. Small ventrally directed knob at mandibular symphysis.
Dentition more or less equally developed on both jaws; teeth on upper jaw sharp, uniserial, some anterior teeth
enlarged and widely spaced; lower jaw teeth uniserial, nearly equal to anterior teeth of upper jaw in size and
spacing. Gill rakers on first arch serrated, absent on upper limb. Scales large, deciduous, cycloid and weakly
ctenoid with short ctcnii on ocular side, cycloid on blind side. Lateral line curved above pectoral fin on ocular side;
lateral line absent on blind side.
Table 5. — Frequency distributions of selected meristic characters of Arnoglossus japonicus. Counts from blind side
shown in parentheses.
Table 6. — Proportional measurements as percent of standard length in Arnoglossus japonicus. O. ocular side; B, blind
side; juv., juveniles.
Dorsal fin origin on blind side, anterior to ventral margin of upper eye; second ray elongated more than adjacent
rays in mature males, not elongated in females and juveniles. Anal fin origin at a point equal with vertical through
basal part of pectoral fin. Pectoral fins unequal, length of ocular side fin 1.71-2.12 times that of blind side. Pelvic
Source : MNHN. Paris
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
797
fins with 6 rays; approximately fourth ray of ocular side fin opposite first ray of blind side fin. Tip of isthmus
extending posterior to and ventral to lower eye. Caudal fin rays bifurcated except upper- and lowermost three
simple rays.
Vent opens on blind side, immediately dorsal to origin of anal fin. Urogenital papillae on ocular side, opposite
side of vent.
Coloration in alcohol'. Ocular side ground color light brown. Blind side uniformly pale yellowish-white. Dorsal
and anal fins with series of dark spots.
Sexual dimorphism: This species shows sexual dimorphism in length of the second dorsal fin ray.
Distribution. — Distributed from the southern coast of Japan (Hubbs. 1915) to Taiwan (Shen, 1983), the
northwestern coast of Australia (Gloefelt-Tarp & Kailola, 1984) and the Coral Sea (Rivaton, 1989), in
depths of 86-300 m.
Remarks. — This species closely resembles A. yamanakai and A. oxyrhynchus in having an elongated
second dorsal fin ray in males, gill rakers with serrated margins and bifurcated middle caudal fin rays. However,
it differs from them in having mostly cycloid scales on the ocular side (vs. scales always feebly ctenoid).
Arnoglossus japonicus further differs from A. yamanakai in having the dorsal fin with a series of dark spots
(vs. dorsal fin with a black band along distal margin from posterior part of head to middle of body in males)
(Fukui, Yamada & Ozawa, 1990) and from A. oxyrhynchus in having uniserial upper jaw teeth (vs. biserial
on at least posterior half of blind side upper jaw), a lower number of lateral line scales (70-80 vs. 87-94).
and the pectoral and pelvic fins without significant markings (vs. pectoral and pelvic fins with blackish
markings).
This species has been reported as having cycloid scales on either side of the body. However, the present
specimens have some weak ctenoid scales with short spinules in addition to cycloid scales on the ocular side.
Arnoglossus elongatus Weber, 1913
Figs 11-13; Tables 7-8
Arnoglossus elongatus Weber, 1913: 431, fig. 79. — Norman, 1934: 189, fig. 136. — Gloerfelt-Tarp & Kailola.
1984: 273, 358. 1 fig.
Bothus ( Arnoglossus ) elongatus: WEBER & BEAUFORT. 1929: 125, fig. 29.
MATERIAL EXAMINED. — 6 specimens (5 males and I female).
New Caledonia. MUSORSTOM 4: stn CC 173, 19°02.50'S, 163°18.80'E, 250-290 m, 17.09.1985: 1 6 . 98.8 mm SL
(HUMZ 142676).
Bathus 1: stn CP 669. 20°57.28'S, 165°35.30'E. 255-280 m, 14.03.1993: 3 <3, 66.4-88.0 mm SL (MNHN 1995-
1 165 to 1 167); 1 2 , 84.5 mm SL (HUMZ 142677).
HALIPRO 1: stn CC 856, 21°43.08'S, 166°37.76'E, 350 m, 20.03.1994: 1 <3, 96.1 mm SL (MNHN 1995-1164).
Diagnosis. — Middle caudal fin rays bifurcated (Fig. 1); dorsal fin with a dark band on distal margin in males
(Fig. 13); second and third, sometimes fourth dorsal fin rays elongated in both sexes (Fig. 13).
Description. — Ranges for proportional data given first, followed by averages. Counts and proportional
measurements as percent of SL shown in Tables 7 and 8.
Head length 3.86-4.12, 3.95 in SL; body depth 2.67-3.15, 2.83. Snout length 4.27-5.09, 4.78 in head length;
upper eye diameter 2.92-3.30, 3.11; lower eye diameter 2.92-3.25, 3.09; interorbital width 26.67-54.50, 40.01:
upper jaw length 2.38-2.55, 2.45 on ocular side, 2.20-2.38, 2.28 on blind side; lower jaw length 1.89-1.98, 1.93
on ocular side, 1 .74- 1.88, 1 .82 on blind side; depth of caudal peduncle 2.70-3.29, 3.09; length of longest dorsal fin
ray 1.68-1.89, 1.80; length of third dorsal fin ray 1.07-2.33, 1.64 in males, 2.66 in a female; length of longest
anal fin ray 1.73-1.83, 1.77; pectoral fin length 1.46-1.66, 1.57 on ocular side, 3.04-3.73, 3.38 on blind side;
pelvic fin length 3.00-3.21, 3.10 on ocular side, 2.58-2.76, 2.76 on blind side; length of pelvic fin base 3.04-
798
K. AMAOKA & E. MIHARA
3.35. 3.21 on ocular side, 8.28-9.08, 8.59 on blind side; length of middle caudal fin ray 1.28-1.39, 1.34; lateral
line curve width 1.98-2.21, 2.09.
Fig. 11, — Arnoglossus elongaius Weber, 1913, male. 98.8 mm SL, from off north of New Caledonia (HUMZ 142676).
Body elongated, elliptical, deepest slightly anterior to midpoint; dorsal and ventral contours gently arched.
Caudal peduncle depth 21.9-24.7 % of body depth. Head large; upper profile with a slight notch anterior to upper
eye. Snout long, length 59.0-77.3 % of upper eye diameter. Rostral and orbital spines absent. Eyes large, separated
by narrow bony ridge; upper eye diameter 72.5-84.8 % of ocular side upper jaw length, lower eye in advance of
upper. Ocular side nostrils anterior to dorsal margin of lower eye; anterior nostril tubular with a flap posteriorly;
blind side nostrils small, near dorsal fin origin, similar in shape to those on ocular side.
Mouth large, weakly curved obliquely;
maxilla extending to below anterior part of
lower eye; anterior tips of both jaws nearly on
same vertical line when mouth closed. Small
knob at mandibular symphysis. Dentition more
or less equally developed on both jaws; teeth on
upper jaw sharp, uniscrial, becoming larger and
more widely spaced anteriorly; lower jaw teeth
small and close-set. Gill rakers on first arch
moderate in size, serrated (Fig. 12A). Ocular
side scales large; with short ctcnii (Fig. I2B)
except those on anterior lateral line and around
pectoral fin base; snout, upper jaw. anterior
parts of lower jaw, and interorbital region
naked; cycloid scales on blind side. Lateral line
curved above pectoral fin on ocular side, lateral
line absent on blind side.
Dorsal fin origin on blind side, anterior to
inlerorbital region; second and third, sometimes
5 mm
1 mm
Fig. 12. — First gill arch (A) and a scale (B) from ocular side of
Arnoglossus elongaius Weber, 1913, female, 84.5 mm SL
(HUMZ 142677).
Source : MNHN. Paris
PISCES PLEURONECT1FORMES: GENUS ARNOGLOSSUS
199
fourth rays elongated more than adjacent rays in both sexes; third ray in males longer than that in females (Fig.
13). Anal fin origin equal with vertical through posterior margin of head. Pectoral fins unequal; length of ocular
side fin 1.90-2,47 times that of blind side, much less than head length. Pelvic fins with 6 rays, origin of ocular
side fin at tip of isthmus; base of ocular side fin longer than that of blind side; fourth ray of ocular side fin
opposite first ray of blind side fin. Tip of isthmus at a point equal with vertical through posterior part of lower
eye. All fin rays simple, except caudal fin rays. Caudal fin rays bifurcated, except upper- and lowermost three
simple rays.
Table 7. — Frequency distributions of selected meristic characters of Arnoglossus elongaius. Counts from blind side
shown in parentheses.
Table 8. — Proportional measurements as percent of standard length in Arnoglossus elongaius, including syntype.
O, ocular side; B, blind side.
* except 3rd dorsal fin ray
Source
800
K. AMAOKA & E. MIHARA
Vent on blind side, immediately dorsal to first anal fin ray. Urogenital papilla on ocular side, opposite side of
vent.
Coloration in alcohol: Ocular side ground color light brown; a few obscure dark spots on lateral line. Blind side
uniformly pale brown. Dorsal and anal fins with series of obscure dark blotches; males with a dark band on distal
margin of dorsal fin, from region above posterior part of head to middle of body (Fig. 13); caudal fin with a pair of
obscure dark spots proximally.
Sexual dimorphism: This species shows sexual dimorphism in lengths of the anterior elongated dorsal fin rays
and dorsal fin coloration (Fig. 13).
Fig. 13. — Diagrammatic illustration of anterior body parts showing sexual dimorphism in male (A) and female (B)
Arnoglossus elongatus. Scale bars = 10 mm.
Distribution. — Indonesia: Madura Sea (Weber, 1913), northwestern Australia (Gloefelt-Tarp &
Kailola, 1984) and New Caledonia, in depths of 100-350 m.
Remarks. — This species very closely resembles A. yamanakai Fukui, Yamada & Ozawa, 1988, in having
bifurcated middle caudal fin rays and a dark band in males on the distal margin of the dorsal fin (Fig. 1 3). However,
this species differs from A. yamanakai in having several elongated anterior dorsal fin rays in both sexes (Fig. 13)
(vs. only second ray elongated and only in males), a higher number of lateral line scales (74-80 vs. 63-68), larger
eyes (upper eye diameter 2.92-3.30 in head length vs. 3.31-3.92 in adult specimens, lower eye diameter 2.92-3.25
vs. 3.31-3.95), and longer dorsal (1.68-1.89 vs. 2.02-2.47) and anal fin rays (1.73-1.83 vs. 2.02-2.48).
Arnoglossus elongatus also resembles A. polyspilus in having several elongated anterior dorsal fin rays and
with the upper head profile rounded. It differs from the latter in the bifurcated middle caudal fin rays (vs. middle
caudal fin rays with more than two branches) and its longer dorsal fin ray (1.68-1.89 in head length vs. 2.00-2.70).
This is the first record of A. elongatus from waters off New Caledonia.
Arnoglossus tenuis Gunther, 1880
Figs 14-15; Tables 9-10
Arnoglossus tenuis Gunther, 1880: 55. — Jordan & Seale, 1905: 528. — Jordan & Hubbs, 1925: 295. — Wu, 1932:
89. — Norman, 1934: 187, fig. 134. — Fowler, 1934: 64. — Wu & Tang, 1935: 392. — Okada & Matsubara,
1938: 421; 1955: 1257. — Kuromuma, 1940: 213. — Kuroda, 1951: 388. — Zheng, 1963: 970, fig. 755. — Zang
& Wang, 1963: 517, fig. 385. — Shen, 1966: 176, figs 39-41; 1983: 27. fig. 39. — Amaoka. 1969: 188. 386. figs
74-75; 1974: 153, figs 1-2; 1984: 335, pi. 369-C. — Nakabo, 1993: 1172, 387, 1 fig.
Scidorhombus pallidus Tanaka, 1915: 567. — Matsubara, 1955: 1255.
Arnoglossus aspilos - Chen & Weng, 1965: 41, fig. 26 (not Rhombus aspilos Bleeker, 1851).
Source : MNHN, Paris
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
801
MATERIAL EXAMINED. — 32 specimens (16 males and 16 temales). AAn01noc j
New Caledonia. Baie de Saint Vincent: 22°04.9'S, 166°05.2'E, 13 m, otter trawl (shrimps), 20.08.1985: 2 6 and
1 9 61.7-77.9 mm SL (MNHN 1995-1108 to 1110); 1 <3 and I $ , 66.0-78.9 mm SL (HUMZ 142659-142660).
21°58.5'S, 166°01 .0'E, 10 m, otter trawl (shrimps), 18.10.1989: 4 <3 and 4 9. 62.7-68.8 mm SL (MNHN ! 995-1 1 14 to
1121); 1 9, 59.1 mm SL (HUMZ 142662). — 21°58.5'S, 166°01.0’E, 10 m, otter trawl (shrimps), 21.11.1989: 3 9,
57.1-60.0 mm SL (MNHN 1995-1111 to 1113); 1 <3, 64.6 mm SL (HUMZ 142661).
Lagon: stn 967, 20°21.6'S, 164°06.7'E. 12-16 m, beam trawl, 29.04.1988: 7 <3 and 4 19, 49.2-60.8 mm SL (MNHN
1995-1122 to 1132); 1 <3 and 2 9 , 46.6-54.2 mm SL (HUMZ 142663-142665).
DIAGNOSIS. — Body shallow, depth 2.43-2.87 in SL; ocular side pelvic fin blackish, except basal portion in
males; dorsal fin rays 84-94; anal fin rays 63-71; lateral line scales 54-59.
Fig.
I4. _ Arnoglossus tenuis Gunther, 1880, male. 77.9 mm SL. from off southern New Caledonia (MNHN 1995-1109).
DESCRIPTION. — Ranges for proportional data given first, followed by averages. Counts and proportional
measurements as percent of SL shown in Tables 9 and 10.
Head length 3.86-4.61, 4.09 in SL; body depth 2.43-2.87, 2.64. Snout length 4.38-5.28 4.7 1 in head length,
upper eye diameter 3.57-4.28, 4.01; lower eye diameter 3.71-4.52, 4.08; mterorbital width I -)-67-56.00, . 6- ,
upper jaw length 2.42-2.85, 2.61 on ocular side, 2.56-2.96, 2.80 on blind side; lower jaw length 1.90-2 10. .99
on ocular side, 1.97-2.25, 2.10 on blind side; depth of caudal peduncle 2.08-2.72, 2.43; length of longest dorsal in
ray 1.45-1.85, 1.63; length of longest anal fin ray 1.42-1.71, 1.59; pectoral fin length 1 -52-1.80 L68 on ocular
side, 2.18-2.75, 2.46 on blind side; pelvic fin length 2.29-2.89, 2.64 on ocular side m males, 2.74-3.4 1 ,3.05 in
females, 2.57-3.41, 3.10 on blind side; length of pelvic fin base 3.16-4,05, 3.60 on ocular side 6.04-9.85. « -8
on blind side; length of middle caudal fin ray 1.10-1.29, 1.20; lateral line curve w,dth 1.65-2.03. 1.84.
Body elongated, elliptical, deepest at midpoint; dorsal contour strongly arched, ventral one weakly arched
Caudal peduncle depth 24.5-29.6 % of body depth. Head small; a large notch anterior lo upper eye. Snout short,
length 68.2-95. 1 % of upper eye diameter. Rostral and orbital spines absent. Eyes small, separated by narrow bony
ridge, upper eye diameter 60.0-73.3 % of upper jaw length; lower eye slightly in advance of upper. Ocular side
nostrils anterior to interorbital area; ocular side anterior nostril tubular with a flap posteriorly; blind side nostri s
small, ventral to dorsal origin, similar in shape to those on ocular side.
Mouth large, oblique; maxilla extending to below or slightly beyond anterior margin of lower eye; anterior tip
of lower jaw projecting slightly in advance of tip of upper jaw when mouth closed. Small ventrally directed knob
Source
802
K. AMAOKA & E. MIHARA
at mandibular symphysis. Dentition more or less equally developed on both jaws; upper jaw teeth sharp, uniserial:
lower jaw teeth uniserial, nearly equal to middle teeth of upper jaw in size and spacing. Gill rakers on first arch
slender, not serrated, absent on upper limb. Scales large, deciduous, with short ctenii on ocular side, cycloid on
blind side. Lateral line curved above pectoral fin on ocular side; lateral line absent on blind side.
Dorsal fin origin on blind side, anterior to ventral margin of upper eye. Anal fin origin at a point equal with
vertical through pectoral fin base. Pectoral fins unequal, length of ocular side fin 1.31-1.64 times that of blind
side. Pelvic fins with 6 rays; ocular side pelvic fin slightly longer in males than in females (Fig. 15);
approximately fourth ray of ocular side fin opposite first ray of blind side fin. Tip of isthmus posterior to and
ventral to lower eye. Most caudal fin rays with more than two branches, except upper- and lowermost three or four
simple rays.
Vent on blind side, immediately dorsal to first anal fin ray. Urogenital papilla on ocular side, opposite side of
vent.
Coloration in alcohol: Ocular side ground color dark brown; several obscure dark spots on lateral line. Blind
side uniformly pale brown. Ocular side pelvic fin blackish, except basal portion in males; females with dark distal
margin.
Sexual dimorphism: This species shows sexual dimorphism in length and coloration of the ocular side pelvic-
fin (Fig. 15).
Distribution. — Distributed from the southern coast of Japan (Jordan & Hubbs, 1925) to the South China
Sea (Gunther, 1 880) and New Caledonia, in depths of 7- 16 m.
_ra
o
o c
° 2
3 >
CO ®
S e
_c ^
tt w
c
®
12
10 -
8 -
• males
o females
40
•O. . .o . . . r.<b
o 00 °
50 60 70
Standard length (mm)
- 1
80
Fig. 15. — Relationship between SL and ocular side pelvic fin length in percent of SL in Arnoglossus tenuis.
Table 9. — Frequency distributions of selected meristic characters of Arnoglossus tenuis. Counts from blind side shown
in parentheses.
Gill rakers
0+8 0+9 1+8 1+9 1 + 10 2+8 2+9 2+10 3+8 3+9 3 + 10 4+8
6 4 2 6 1 3 3 12 2 11
Source : MNHN, Paris
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
803
Table 10. _ Proportional measurements as percent of standard length in Arnoglossus tenuis. O. ocular side; B. blind
side; M, males; F, females.
REMARKS. — This species closely resembles A. aspilos in having a shallow body and relatively low numbei
of lateral line scales, although the latter count is somewhat higher than that of A. aspilos (54-59 vs. 46-48)
(Norman, 1934).
Sexual dimorphism has not previously been reported in this species. However, New Caledonian specimens
showed sexual dimorphism in both the length and coloration ol the ocular side pelvic I in.
This is the first record of A. tenuis from the Coral Sea.
Arnoglossus polyspilus (Giinlher, 1880)
Fig. 16; Tables 11-12
Anticitharus polyspilus Gunther, 1880: 48, pi. 22, fig. A. Weber. 1913: 433 „
Arnoglossus polyspilus - Norman, 1927: 20; 1934: 190. fig. 137. -Okaoa & MatsubaRA 1938. 421. -KaMOHARA.
1951: 6; 1952: 78; 1958: 62; 1964: 81. — MATSUBARA, 1955: 1257. — Chen & WenG. 1965_ 42 fig 27
AMAOKA, 1969: 191, figs 76-77. - Shew, 1983: 29, fig. 40. - Gloefelt-Tarp & Kailola, 1984. 73 358, fig. -
Rivaton, 1989: 145, 155, 163. — Rivaton & Richer de Forges. 1990: 25. 60. — Nakabo, 139.'. 1 1 73. I fig.
Kulbicki, Randall & Rivaton, 1994: 37.
Bothus (Anticitharus) polyspilus - Weber & BEAUFORT. 1929: 124.
Material examined. — 12 specimens (7 males and 5 females).
New Caledonia. Musorstom 4: stn CP 192, 18°59.30'S, 163°25.00'E. 320 m.
170.2 mm SL (MNHN 1995-1 160 to 1 163); 1 $ , 66.0 mm SL (HUMZ 142674).
19.09.1985: 2 3 and 2 9 , 63.0-
Source
804
K. AMAOKA & E. MIHARA
Loyalty Islands. Musorstom 6: stn DW 392, Ouvea Island. 20°47.32'S, I67°04.60'E. 340 rn, 13.02.1989: I ?,
52.5 mm SL (HUMZ 142673). — Stn CP 409. Lifou Island. 20°41.05'S, 167°07.25'E, 385 m. 15.02.1989' 1 $'
167.9 mm SL (HUMZ 142672).
Chesterfield Plateau. Chalcal 1: stn CP 4, 19°33.90'S. 158°37.90'E, 350-370 m, 16.07.1984: 1 9 . 181 3 mm
SL (MNHN 1995-1156). — Stn CP 5. 19°29.I0'S, 158°37.63'E. 290 m, 16.07.1984: 3 <3. 72.1-113.1 mm SL (MNHN
1995-1157 to 1159).
Musorstom 5: stn DW 335, 20°03.24'S. 158°45.35'E, 315 m, 15.10.1986: I <3, 171.0 mm SL (HUMZ 142675).
Diagnosis. — Upper profile of head rounded; gill rakers with serrated margins; dorsal fin rays 109-1 16;
anal fin rays 86-91.
Fig. 16. — Arnogiossus polyspilus (Gunther, 1880), female. 181.3 mm SL, from Chesterfield Plateau (MNHN 1995-
1 156).
Description. — Ranges for proportional data given first, followed by averages. Counts and proportional
measurements as percent of SL shown in Tables 1 1 and 12.
Head length 3.57-4.05, 3.87 in SL; body depth 2.42-2.83, 2.64. Snout length 4.15-4.72, 4.46 in head length;
upper eye diameter 2.95-4.08, 3.60; lower eye diameter 3.00-4.15, 3.64; intcrorbital width 22.55-41.50. 30.88;
upper jaw length 2.31-2.62, 2.48 on ocular side, 2.18-2.50, 2.34 on blind side; lower jaw length 1.82-2.02. 1.93
on ocular side, 1 .67- 1 .89, 1 .78 on blind side; depth of caudal peduncle 2.58-3.00, 2.77; length of longest dorsal fin
ray 2.00-2.22, 2.1 1; length of longest anal fin ray 1.85-2.19, 2.03; pectoral fin length 1.77-1.95. 1.83 on ocular
side, 2.84-3.93, 3.41 on blind side; pelvic fin length 2.47-3.78, 3.24 on ocular side. 2.59-3.66, 3.17 on blind side;
length of pelvic fin base 3.10-3.99, 3.49 on ocular side, 5.55-7.01, 6.24 on blind side; length of middle caudal fin
ray 1.32-1.51, 1.42; lateral line curve width 1.48-1.87, 1.72.
Body elongated, elliptical, deepest slightly anterior to midpoint; dorsal contour more strongly arched than
ventral one. Caudal peduncle depth 22.7-26.9 % of body depth. Head large; a slight notch anterior to ventral
margin ol upper eye. Snout long, length 64.5-96.2 % of upper eye diameter. Rostral and orbital spines absent.
Eyes separated by narrow bony ridge, upper eye diameter 57.8-87.3 % of ocular side upper jaw length; lower eye
slightly in advance of upper. Ocular side nostrils anterior to dorsal margin of lower eye. anterior nostril tubular
with triangular flap posteriorly, and short flap anteriorly; blind side nostrils ventral to dorsal fin origin, anterior
nostril tubular with short flap posteriorly.
Source :
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
805
Mouth large, oblique; maxilla extending to below anterior part of lower eye; anterior tips of both jaws nearly
on same vertical line when mouth closed. Small ventrally directed knob at mandibular symphysis. Dentition about
equally developed on both jaws; teeth on upper jaw sharp, uniserial, becoming larger and more widely spaced
anteriorly; lower jaw teeth uniserial, similar to anterior teeth of upper jaw in size and spacing. Gill rakers on first
arch moderate in size or slender, serrated, usually small rakers on upper limb. Scales large, deciduous, with short
ctenii on ocular side, cycloid on blind side. Lateral line curved above pectoral tin on ocular side, lateral line absent
on blind side.
Table 11. — Frequency distributions of selected meristic characters of Amoglossus polyspilus. Counts from blind side
shown in parentheses.
Caudal fin rays
2+13+2 2+12+2
1 1 1
Scales in lateral line
70
1
71
0
72
2
73
0
74
0
75
1
76
2
77
o
78
2
79
1
80
1
Vertebrae
10+30
1
10+31
7
10+32
4
Gill rakers
0+7
1
0+8
0+9
1
1+8
6
1+9 2+8 2+9
Table 12. — Proportional measurements as percent of standard length in Amoglossus polyspilus. O, ocular Side;
B, blind side.
Source :
806
K. AMAOKA & E. M1HARA
Dorsal fin origin on blind side, anterior to ventral margin of upper eye; anterior 3-7 rays slightly elongated in
some adult males, scarcely so in females and young males. Anal fin origin at vertical through posterior margin of
head. Pectoral fins unequal, length of ocular side fin 1.61-2.22 times that of blind side. Pelvic fins with 6 rays;
approximately third ray of ocular side fin opposite first ray of blind side fin. Tip of isthmus extending posterior to
and ventral to lower eye. Caudal fin rays with more than two branches, except upper- and lowermost two simple
rays.
Vent on blind side, immediately dorsal to first anal fin ray. Urogenital papilla on ocular side, opposite side of
vent.
Coloration in alcohol : Ocular side ground color light brown. Blind side uniformly pale yellowish-white. Dorsal
and anal fins with series of dark spots; caudal fin with a pair of obscure dark spots proximally.
Sexual dimorphism : This species shows sexual dimorphism in lengths of the third to seventh dorsal fin rays.
Distribution. — Timor Sea (Gunther, 1880), Kei Islands, Mergui Archipelago (Norman, 1927), south
coast of Java (Weber & Beaufort, 1929), southern coast of Japan (Kamohara, 1951), Taiwan (Chen &
WENG, 1965) and Coral Sea (Rivaton, 1989), in depths of 290-385 m in the Coral Sea and 90-252 m elsewhere.
Remarks. — Arnoglossus polyspilus closely resembles A. nigrifrons sp. nov. in having the upper head
profile rounded and in having large numbers of dorsal and anal fin rays, but differs from the latter as discussed in
that species account (see below).
Arnoglossus nigrifrons sp. nov.
Figs 17-20; Tables 13-14
I 9 , paratype, 45.9 mm SL
Material examined. — 3 specimens (1 male and 2 females).
New Caledonia. Smib 6: stn DW 121. 18°57.08'S, 163°25.06'E. 315 m. 3.03. 1 990
(HUMZ 142678).
Chesterfield Plateau. Chalcal I: stn CP 5. 19°29.I0-S, 158°37.63'E, 290 m, 16.07.1984: I 2. damaged spec¬
imen (HUMZ 152143). - Stn CP 11, 20°04.40'S, 158°47.41'E, 300 m, 22.07.1984: 1 <3, holotype. 119.8 mm SL
(MNHN 1995-1 168).
Fig. 17. — Illustration of Arnoglossus nigrifrons sp. nov., holotype, male, 119.8 mm SL, from Chesterfield Plateau,
west of New Caledonia (MNHN 1995-1168).
Source :
PISCES PLEURONECTIFORMES: GENUS ARNOCLOSSUS
807
Types . — Holotype: MNHN 1995-1168 (Chesterfield Plateau. Chalcal I, stn CP 11. 20°04.40'S,
158°47.41'E, 300 m).
Paratype: (HUMZ 142678 (New Caledonia. Smib 6, stn DW 121, 18°57.08'S, 163°25.06'E. 315 m).
DIAGNOSIS. — Upper profile of head rounded, with a very slight notch anterior to interorbital region (Figs 17-
18); anterior third to fifth rays of dorsal fin elongated more than adjacent rays in males; first to about tenth rays of
dorsal fin, snout, both jaws and area in front of both eyes, and interorbital region darkened (Figs 17-18); gill rakers
not serrated (Fig. 19A); dorsal fin rays 107-1 15; anal fin rays 88-91; scales in lateral line 79-83.
FlG. is. — Arnoglossus nigrifrons sp. nov., holotype, male. 119.8 mm SL, from Chestcrlield Plateau, west ol
New Caledonia (MNHN 1995-1168).
DESCRIPTION. — Data for holotype given first, followed in parentheses by proportional data lor paratype.
Counts and proportional measurements as percent of SL shown in lable 13.
Head length 3.88 (3.70) in SL; body depth 2.72 (2.98). Snout length 5.15 (4.96) in head length; upper eye
diameter 3.22 (2.58); lower eye diameter 3.22 (2.95); interorbital width 30.90 (41.33); upper jaw length 2.58
(2.76) on ocular side, 2.60 (2.76) on blind side; lower jaw length 2.10 (2.10) on ocular side, 2.01 (1.94) on blind
side; depth of caudal peduncle 2.97 (3.02); length of longest dorsal fin ray 2.24 (2.10); length of longest anal hn
ray 2.07 (1.97); pectoral fin length 1.78 (2.07) on ocular side, 3.02 (-) on blind side; pelvic lin length 3.32 (3.26)
on ocular side, 3.32 (3.44) on blind side; length of pelvic fin base 3.15 (4.13) on ocular side, 6.18 (6.89) on blind
side; length of middle caudal fin ray - (1.43); lateral line curve width - (1.75).
Body elongated, elliptical, deepest slightly anterior to midpoint, depth 1.23-1.43 length ol head; dorsal and
ventral contours gently arched. Caudal peduncle depth 23.6-26.6 % of body depth. Head large; upper profile with a
very slight notch anterior to interorbital region. Snout short, length 52.1-62.5 % ol upper eye diameter. Rostial
and orbital spines absent. Eyes large, separated by narrow bony ridge; upper eye diameter 80.0-106.7 % ol ocular
side upper jaw length, lower eye slightly in advance of upper. Ocular side nostrils anterior to dorsal margin of
lower eye; anterior nostril tubular with a flap posteriorly; blind side nostrils small, ventral to dorsal fin origin,
similar in shape to those on ocular side.
Mouth large, oblique; maxilla extending to below anterior part of lower eye; anterior tips of both jaws nearly
on same vertical line when mouth closed. Dentition about equally developed on both jaws; upper jaw teeth shaip.
uniserial, becoming larger and more widely spaced anteriorly, some anterior teeth canines; lower jaw teeth
uniserial, similar to anterior teeth of upper jaw in size and spacing. Gill rakers on hrst arch moderate in size, not
808
K. AMAOKA & E. MIHARA
serrated, absent on upper limb (Fig. 19A). Scales small, with short ctenii on ocular side (Fig. 19B), cycloid on
blind side; snout, anterior parts of both jaws, areas in front of both eyes, and interorbital region and pectoral fin
naked. Lateral line curved above ocular side pectoral fin; lateral line absent on blind side.
Dorsal fin origin on blind side, anterior to
interorbital region; anterior 3-5 rays elongated
more than adjacent rays in male, not so in
female (Fig. 20). Anal fin origin at vertical
through posterior margin of head. Pectoral fins
unequal, length of ocular side fin 1.71 times that
of blind side, much less than head length. Pelvic
fins with 6 rays, origin of ocular side fin at tip
of isthmus, approximately third ray of ocular
side fin opposite to first ray of blind side fin;
base of ocular side fin longer than that on blind
side. Tip of isthmus posterior to and ventral to
lower eye. All fin rays simple, except caudal fin
rays. Caudal fin rays with more than two
branches, except upper- and lowermost two
simple rays.
Vent on blind side, immediately dorsal to
first anal fin ray. Urogenital papilla on ocular
side, opposite side of vent.
Coloration in alcohol: Ocular side ground
color pale brown; snout, both jaws, and area in front of eyes, and interorbital region darkened; a few obscure dark
spots on lateral line. Blind side uniformly pale brown. Dorsal fin darkened anteriorly for about ten rays; dorsal and
anal fins with series of obscure dark blotches.
Sexual dimorphism : This species shows sexual dimorphism in lengths of the anterior third to fifth dorsal fin
rays (Fig. 20).
Fig. 19. — First gill arch (A) and a scale (B) from ocular side of
Arnoglossus nigrifrons sp. nov. (A: female, HUMZ 152143,
86.6 mm SL; B: holotype, male, 119.8 mm SL, MNHN 1995-
1168).
Fig. 20. Diagrammatic illustration of anterior body parts showing sexual dimorphism in male (A) and female (B)
Arnoglossus nigrifrons sp. nov. Scale bars = 10 mm.
Source
PISCES PLEURONECTIFORMES: GENUS ARNOGLOSSUS
809
Distribution. — The specimens were collected from the Chesterfield Plateau and northwest of New
Caledonia, in depths of 290-315 m.
ETYMOLOGY. — Named for the darkened areas on the head and anterior dorsal fin rays.
Table 13. — Proportional measurements as percent of standard length and counts of selected characters for Arnoglossus
nigrifrons sp. nov. O, ocular side; B. blind side.
REMARKS. — Arnoglossus nigrifrons sp. nov. closely resembles A. polyspilus (Gunther, 1880) and
A. capensis Boulengcr, 1898, in having several somewhat elongated anterior dorsal fin rays in the males and with
the upper head profile rounded with a very slight notch, but dillcrs from both by the darker coloration of the
anterior dorsal fin, snout, both jaws and the area anterior to the orbital region. It also clearly differs from
A. polyspilus in having slender and elongate gill rakers without serrated margins (vs. strong and serrated gill
Source :
810
K. AMAOKA & E. M1HARA
rakers) (Fig. 19A) (Amaoka, 1969, fig. 77C) and a smaller lower jaw (Table 14). and from A. capensis in having
a ridged and non-scaled interorbital region (vs. concave and scaled), larger numbers of dorsal and anal fin rays and
lateral line scales, a smaller number of gill rakers, shallower body, larger upper eye and upper jaw, and shorter
ocular side pectoral fin (Table 14) (NORMAN, 1934).
Table 14. — Comparison of proportional measurements and counts between Amoglossus nigrifrons sp. nov.,
A. polyspilus and A. capensis.
ACKNOWLEDGMENTS
We wish to express our sincere thanks to Bernard Seret, IRD (ex ORSTOM)/Museum national d'Histoire
naturelle, Paris, and Jacques RlVATON, IRD, Noumea, New Caledonia, who loaned specimens from the IRD
collection, and Graham Hardy, Thames, New Zealand for their critical reading of the manuscript. Special thanks
arc due to the former Prof. Tamotsu IWAI and to Izumi Nakamura, Kyoto University; Nigel R. MERRETT,
Ethelwynn Trewavas and Jim Chambers, British Museum, Natural History; Keiichi Matsuura, Department
of Zoology, Natural Science Museum, Tokyo, for loans of type specimens, and to Bertrand RICHER DE FORGES
and Michael Kulbicki, IRD, Noumea, for collecting specimens. This study was supported by grants from the
Ministry of Education (No. 0745423 1 ) and the Ito Foundation for the Advancement of Ichthyology.
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Remerciements aux rapporteurs / Acknowledgements to referees
La Redaction ticnl a remercier les experts ext<Srieurs au Museum national d'Histoire naturelle dont les nonts suivent, d'avoir bien voulu contribucr.
avec les rapporteurs de I'Etablissement, a revaluation des manuscrits (1997/2000) :
The Editorial Board acknowledges with thanks the following referees who , with Museum referees, have reviewed papers submitted to the Mdmoires
du Musduni (1997/2000):
Source MNHN, Paris
aciu.vt: d’imprimer
F.N NOVEMBRF. 2000
SUR I.F.S PRESSES
DE
LIMPRIMERIE F. PAILLART
A ABBEVILLE
Date tie distribution : 30 novembre 2000.
Depot 16 gal : novembre 2000.
N" d’ impression : 1 1 165.
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