CFBTJM
MEM01RES
DU MUSEUM
NATIONAL
D’HISTOIRE
NATURELLE
TOME 180
1999
Resultats des Campagnes MUSORSTOM
Volume 20
Coordonne par
Alain CROSNIER
Publie avec le concours de I'Institut frangais de Recherche scientifique
pour le Developpement en Cooperation (ORSTOM)
COM
Source : MNHN, Paris
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3 3001 00057288 2
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Mich&le Bertoncini
Sylvie Chalier
R6gis Cl£va
Jean-Fran^ois Dejouannet
Maurice Gaillard
Virginie Heros
u
Pierre Laboute
Gabrielle Gadaleta
Jacques Rebi£re
Fran^oise Theureau
Le vingtieme volume des Resultats des Campagnes Musorstom est dedie a ceux qui, discretement, out
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A tous, les divers editeurs de la serie expriment leur reconnaissance.
Source : MNHN. Paris
resultats des campagnes
Volume 20
Source
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., 1 1 pi. couleur (1994). ISBN : 2-85653-212-8.
Volume 13 : Mem. Mus. natn. Hist, nat., 163 : 1-517, 132 fig., 4 pi. couleur (1995). ISBN : 2-85653-224-1.
Volume 14 : Mem. Mus. natn. Hist, nat., 167 : 1-647, 987 fig., 3 pi. couleur (1995). ISBN : 2-85653-217-9
Volume 15 : Mem. Mus. natn. Hist, nat., 168 : 1-539, 205 fig., 6 pi. couleur (1996). ISBN : 2-85653-501-1.
Volume 16 .* Mem. Mus. natn. Hist, nat., 172 : 1-667, 432 fig., 2 pi. couleur (1997). ISBN : 2-85653-506-2.
Volume 17 : Mem. Mus. natn. Hist, nat., 174 : 1-213, 93 fig., (1997). ISBN : 2-85653-500-3.
Volume 18 : Mem. Mus. natn. Hist, nat., 176 : 1-570, 458 fig., 7 pi. 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.
ISBN : 2-85653-520-8
ISSN : 1243-4442
© Publications Scientifiques du Museum, Paris, 1999
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MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE
TOME 180
ZOOLOG1E
Resultats des Campagnes MUSORSTOM
Volume 20
Coordonne par
Alain CROSNIER
Museum national d’Histoire naturelle
Laboratoire de Biologic des Invert6br6s marins et Malacologie
55 rue Buffon
F-75005 Paris
Publie avec le concours de I'Institut frangais de Recherche scientifique pour le Developpement
en Cooperation (ORSTOM)
PUBLICATIONS SCIENTIFIQUES
DU MUSEUM
PARIS
1999
Source : MNHN, Paris
SOMMAIRE
Pages
1. La campagne MUSORSTOM 9 dans larchipel des lies Marquises
(Polynesie fran^aise). Compte rendu et liste des stations
Berirand RICHER DE FORGES, Joseph Poupin & Pierre Laboute
2. Cnidaria Anthozoa : Scleractiniaires d'eau profonde sans zooxanthelles
du Vanuatu et des iles Wallis et Futuna (cn anglais) .
Stephen D. Cairns
3 Entoproctes et Bryozoaires Cheilostomida (Pseudomalacostegomorpha et Crypto-
cystomorpha) des campagnes MUSORSTOM autour de la Nouvelle-Caledonie
Jcan-Loup D’HONDT & Dennis P. GORDON
4. Crustacea Isopoda : Bopyridae des collections MUSORSTOM recoltes
dans l'lndo-Pacifique tropical. II. Especes de la sous-famille Pseudioninae
infestant des hotes non anomuriens (en anglais) .
John C. Markham
5 Crustacea Decapoda : Revision de Pasiphaea sivado (Risso, 1816) et
des especes qui lui sont proches. Description d'un genre et cinq especes nouveaux
(Pasiphaeidae) (en anglais) .
Ken-Ichi HAYASHI
6. Crustacea Decapoda: Revision des especes du genre Parapagurus
Smith, 1879 (Parapaguridae) des oceans Indien et Pacifique (en anglais)
Rafael Lemaitre
7. Crustacea Decapoda: Albuneidae et Hippidae de l'lndo-Ouest
Pacifique tropical (en anglais) .
Christopher B. BOYKO & Alan W. Harvey
8 Crustacea Decapoda: Especes des genres Agononida Baba
& de Saint Laurent, 1996 et Munida Leach, 1820 (Galatheidae) recoltees
durant la campagne Musorstom 8 au Vanuatu (en anglais) .
Enrique MaCPHERSON
9. Crustacea Decapoda: Revision de la famille Dynomenidae
(en anglais) .
Colin L. McLay
10. Pisces Anguilliformes: Poissons-serpents d'eau profonde (Ophichthidae)
de la region neo-caledonienne (Pacifique sud-ouest) (en anglais) .
John E. McCoSKER
CONTENTS
Pages
1. The Musorstom 9 cruise in the Marquesas Archipelago. Report and list of stations
(in French) . ^
Bertrand RICHER DE FORGES, Joseph POUPIN & Pierre LABOUTE
2. Cnidaria Anthozoa: Deep-water azooxanthellate Scleractinia from Vanuatu,
and Wallis and Futuna Islands . 31
Stephen D. Cairns
3. Entoprocta and Bryozoa Cheilostomatida (Pseudomalacostegomorpha and
Cry ptocystomorpha) from the MUSORSTOM cruises around New Caledonia (in French) . 169
Jean-Loup D'Hondt & Dennis P. GORDON
4. Crustacea Isopoda: Bopyridae in the MUSORSTOM collections from the tropical
Indo-Pacific. II. Species in subfamily Pseudioninae infesting nonanomuran hosts . .. 253
John C. Markham
5. Crustacea Decapoda: Revision of Pasiphaea sivado (Risso, 1816) and related species,
with descriptions of one new genus and five new species (Pasiphaeidae) . 267
Ken-Ichi Hayashi
6. Crustacea Decapoda: A review of the species of the genus Parapagurus Smith,
1879 (Parapaguridae) from the Pacific and Indian Oceans .
Rafael Lemaitre
7 Crustacea Decapoda: Albuneidae and Hippidae of the tropical Indo-West
Pacific region .
Christopher B. BOYKO & Alan W. Harvey
8. 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 .
Enrique MACPHERSON
9. Crustacea Decapoda: Revision of the family Dynomenidae .
Colin L. McLay
10. Pisces A nguilliformes: Deepwater snake eels (Ophichthidae)
from the New Caledonia region, Southwest Pacific Ocean
John E. McCosker
379
407
427
571
Source : MNHN, Paris
[’ATS DES CAMPAGNES MUSORSTOM. VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES CA
La campagne MUSORSTOM 9 dans l’archipel
des lies Marquises (Polynesie franchise).
Compte rendu et liste des stations
Bertrand RICHER DE FORGES * Joseph POUPIN **
cfc Pierre LABOUTE*
*ORSTOM
B. P. A5, Noumea Cedex
Nouvelle-Caledonie
**Laboratoire d'Oceanographie
Ecole navale
29240 Brest naval
RESUME
La campagne MUSORSTOM 9, realisee h bord du N. O. "Atis", s’est deroulee dans les eaux de Polynesie franyaise,
dans l'archipel des lies Marquises, du 18 aout au 1 1 septembre 1997. 168 operations de dragages et de chalutages ont eu
lieu dans les zones bathyale superieure et circalittorale sur les pentes des lies et sur le sommet du haut-fond Dumont
d’Urville. Une terrasse en pente douce entoure chaque He et se termine par une brusque rupture de pente a 100 m de
profondeur. Ensuite les pentes sont tres abruptes et accidentees avec, parfois. des terrasses entre 400 et 600 m. Cet
archipel isole, situe dans 1c Pacifique central, presente une faune benthique remarquablement pauvre en especes dans tous
les groupes zoologiques.
ABSTRACT
The MUSORSTOM 9 cruise in the Marquesas Archipelago. Report and list of stations.
The Musorstom 9 cruise was carried out in the Marquesas Archipelago from 18 August to 11 September 1997.
168 samples by dredging and trawling were made in the upper-bathyal zone and in the circalittoral depths, on the slope
of the islands and on the top of the Dumont d’Urville Seamount. A terrace with a gentle slope is surrounding each island.
Deeper than 100 meters the slope is very steep with from time to time terraces between 400 and 600 meters deep.
The benthic fauna of this remote archipelago isolated in the Central Pacific is remarkably poor in all the groups.
Richer de Forges, B., Poupin, J. & Laboute, P.. 1999. — La campagne Musorstom 9 dans l’archipel des lies
Marquises (Polynesie frangaise). Compte rendu et liste des stations. In : A. Crosnier (ed.). Resultats des Campagnes
MUSORSTOM, Volume 20. Memoires du Museum national d'Histoire naturelle , 180 : 9-29. ISBN-2-85653-520-8.
10
B. RICHER DE FORGES. J. POUPIN & P LABOUTE
INTRODUCTION
L’etude des faunes littorales dc rindo-Pacifique tropical a montre quc la richesse specifique des zones liltorales
n'est pas homogene en longitude (Paul ay, 1997). On observe en effet, pour lous les groupcs sur Icsqucls les
donnees sont suffisamment fiables, une decroissance de la richesse specifique d’ouest en est dans le Pacifique
(Ekman. 1953 ; Briggs, 1974. 1996 ; Springer. 1982 ; Rosen. 1988). Ceci a etc particulicrcment bicn illustre
pour le groupe des coraux constructeurs qui servent a la fois d’abri et de nourriture a de tres nombreuses especes
d'autres groupes (VERON, 1995).
Pour la faunc bathyale. Ic petit nombre d 'etudes zoologiques portant sur les collections constitutes lors des
"Grandes Expeditions", entre 1873 et 1952, ne permettait pas de verifier si cetle decroissance de la diversite
specifique s'observait egalement. Les donnees des campagnes russes, recemment publiees, apportent quelqucs
informations sur la repartition geographique dc cettc faunc et plus particulierement du groupe des brachiopodes
(Zezina, 1997).
L'exploration du benthos profond dc 1’Indo-Ouest Pacifique, entreprise conjointement par I'ORSTOM et le
Museum national d’Histoire naturelle, essaye depuis une vingtaine d'annees de combler les lacunes des
connaissances zoologiques dans 1'Indo-Ouest Pacifique, tout en apportant des informations sur les ressources
halieutiques et en mettant en evidence des molecules actives extraites des especes profondes (Fig. I ).
L'exploration s'est deroulee en plusieurs etapes. Celle des Ties Philippines a ete relatee par FOREST (1981,
1985, 1989), celle de la Nouvelle-Caledonie et des lies Wallis et Futuna par Richer de FORGES (1990, 1993) et
Richer de Forges et Menou (1993), celle de I'archipel de Vanuatu par Richer de Forges et al. (1996).
L'ensemble des faunes recoltees lors de ces differentes campagnes permet d'observer la repartition geographique des
especes des profondeurs bathyales. II etait done particulicrcment intcressant d'cchantillonner la faune bathyale de
I'archipel des lies Marquises situees dans le Pacifique central, a I'extremite la plus pauvre du gradient de biodi versite
et, de plus, eloigne de plus de 500 km des atolls de I'archipel des Tuamotu au sud-ouesl (sans rnonts sous-marins
entre eux), de 2000 km de File Christmas au nord-ouest. de 4000 km des Ties Hawaii au nord. et de 6000 km des
lies Galapagos a I'est (Springer, 1982 ; Wheeler & Carillet, 1997). Tel fut I'objectif de la campagne
Musorstom 9 qui s'est deroulee, du 18 aout au 1 1 septembre 1997, aux Ties Marquises, en Polynesie fran^aise.
Le seul archipel qui soit plus isole que les Ties Marquises est celui des Tics Hawaii, forme d'un alignement d'TIcs
volcaniques et de rnonts sous-marins pour lesquels l'etude de la faune marine a mis en evidence un taux
d'endemisme eleve (CARLQUIST, 1980 ; Kay. 1994).
Fig. 1. — Repartition geographique des campagnes oceanographiques d'etude de la faune bathyale dc I'lndo-Pacifique
realisees depuis 1976. Philippines : Musorstom 1 (1976), Musorstom 2 (1980), Musorstom 3 (1985) ; lndonesie :
Karubar (1991) ; Nouvelle-Caledonie (plus Chesterfield et Loyaute) : MUSORSTOM 4 (1985), MUSORSTOM 5 (1986),
Musorstom 6 (1989) ; Wallis et Futuna : MUSORSTOM 7 (1992) ; Vanuatu : Musorstom 8 (1994) ; Marquises '
Musorstom 9 (1997).
Source : MNHN. Pans
CAMPAGNE MUSORSTOM 9
1 I
GENERALITIES
Origine geologique des Iles Marquises (Fig. 2). — L'archipel des Ties Marquises est situe dans le
Pacifique central (ANONYME, 1988 ; DUPON et al ., 1993; DUPON & SODTER, 1993).
Ces Ties volcaniques, habitees par des populations prehistoriques polynesiennes depuis environ 200 av. J.- C.
(Irwin, 1992 ; Panoff, 1995), furent decouvertcs par les occidentaux en plusieurs etapes : en 1595 par d’Alvaro
MENDANA, en 1775 par James COOK, en 1791 par Ingraham et par Etienne Marchand. La prise de possession
par la France, par DupetitThouars, a eu lieu le ler mai 1842.
Cet archipel est encore tres mal cartographic et hydrographie. Les Ties, volcaniques, sont constitutes de sept a
dix volcans individuels eriges sur des fonds oceaniques de plus de 4000 m de profondeur.
Parmi les Ties des Marquises, quatre presentent un facies de volcans effondres avec une caldera centrale, ouverte
vers le sud pour Ua Huka (853 m), Nuku Hiva (1208 m), Tahuata (1050 m), et vers l'ouest pour Fatu Hiva
(960 m). L'Tle d'Hiva Oa, qui culmine & 1 190 m d'altitude, est formee de la coalescence de quatre volcans. Ua Pou
prtsente un relief tres caractcristique avec sa serie de pitons phonolithiques et trachytiques culminant a 1252 m
(BROUSSE el al., 1978).
Les mesures d'ages absolus realisees sur les roches des Ties Marquises vont de 6,3 + ou - 0,2 M. A. au nord, a
Eiao, et de 1,3 + ou - 0,02 M. A. au sud, a Fatu Hiva (BROUSSE & Bellon, 1974). Cet ’’alignement” volcanique
serait du au mouvement SE-NO de la plaque Pacifique a une vitesse moyenne de 10,3 cm/an.
La superficie totale des Ties de l’archipel est d'environ 1050 km2, repartie en 10 Ties dont les plus vastes sont
Nuku Hiva, 330 km2, et Hiva Oa, 320 km2.
Environnement HYDROCLIMAT1QUE. — Les conditions hydrologiqucs regnant aux Ties Marquises ont ete
decrites a la suite des campagnes HYDROPOL realisees par le N. O. " Mcircira " entre 1986 et 1990 (ROUGERIE el al. ,
1992 ; ROUGERIE & Wauthy, 1993). Ces campagnes ont mis en evidence la presence d’eaux cotieres vertes,
riches en plancton et turbides par rapport aux eaux oligotrophes du Pacifique central.
Les eaux de surface ont une temperature comprise entre 26,7 el 28,3°C (parfois 30° dans les baies) et une
salinite de 35 a 35,8 %c. Ces eaux sont plus riches en nutriants, en phytoplancton, et en particules que celles du
reste de la Polynesie.
Le regime des pluies est tres irregulier (960 a 2880 mm/an) et les cours d’eau permanents peu nombreux
(3 rivieres de plus de 9 km de long). Le couvert vegetal originel, qui se composait essentiellcment de forets, a ete
presque totalement detruit par 1’homme (incendies, bdtail ) accentuant ainsi les phenomenes de secheresse et
d'erosion par ruissellement (HALLE, 1978).
Les Campagnes ANTERIEURES et la Biodiversite marine. — Les campagnes realisees anterieurement dans
cet archipel sont peu nombreuses et ont echantillonne principalement la taunc marine littorale, accessible a la
plongee en scaphandre autonome. Le navire "Pele" appartenant a Mrs Mariel KING a travaille aux Marquises de
septembre a octobre 1967 ; outre des recoltes littorales, il a effectue quelques dragages jusqu’a 130 m de profondeur.
La mission "Museum-IX”, realisde en fevrier-mars 1973, a permis des recoltes de poissons de coraux et de
mollusques. D’apres Plessis et Mauge (1978), la faune ichtyologique (45 families et 391 especes) ne serait pas
sensiblement differente de celle des autres Ties de Polynesie. Randall (1985) dans sa liste des poissons de
Polynesie franchise (800 especes) mentionne 353 especes signalees des Marquises. Randall (1978) estimait
qu'environ 10% des especes de poissons connues des Ties Marquises etaient endemiques.
Chevalier (1978) a donne un inventaire preliminaire des especes de coraux des Ties Marquises. II constate la
pauvrete de la faune corallienne et le faible developpement des formations recilales. On denombre 3 a 4 tois plus
d'especes de coraux dans les autres archipels polynesiens qu'aux Marquises. Veron (1995) indique dans son etude de
la biogeographie des coraux, que les Ties Marquises auraient environ 50 especes de scleractiniaires, alors que
Pichon (1985) releve 51 genres et 168 especes de scleractiniaires signales de Polynesie fran9aise. Chevalier
(1978) signalait aussi l'absence du genre Acropora aux Ties Marquises (alors que 39 especes de ce genre sont
presentes dans les autres archipels de Polynesie fran^aise) ainsi que celle des Mussidae et des Faviidae.
12
B. RICHER DE FORGES. J. POUPIN & P LABOUTE
Dans le groupe des Mollusqucs, Rehder (1968) evaluait le taux d'endemisme de la faune malacologique des
lies Marquises a 20 %, ce qui les place juste apres les lies Hawaii et I’tle de Paques
SPRINGER (1982) attire l'attention sur le fait qu'aux ties Marquises le for. taux d'endemisme dans plusicurs
g < upes est associe a I absence ou a la rarefaction de certains groupcs, comme c'cst le cas pour les scleractiniaires
J Ponpr^mnn6, m " V y ai,CU(Je signaler les campagnes du N. O. "Marara", sous la direction scientific de
e, * S '•ItS!' Hm,' *hTf’"nages avec dc poses de cas.ers e. quelques dragages (Poup.N
er a/., 990 , Poupin, 1991). Les pcches au easier ont explore la tranche bathymetrique de 100 a 1000 m, recoltant
prmcipalement des crustaces (Poupin et al., 1990 ; Poupin & Richer de Forges, 1991).
Une partie du materiel zoologique de ces campagnes a deja fait I'objct d'etudes. Le crabe de profondeur du genre
Chaceot,, qui etait la cible pr.ncipale des peches aux casiers, s'est revele etre une nouvelle espece, Chaceon poupini
Manning, 1992 (Poupin et al., 1991 ; Manning, 1992). P P
Parmi les 758 especes de crustaces decapodes repertories par POUPIN (1994 ; 1996a, b ; sous presse) pour
ensemble des arch.pels composant la Polynesie fran9aise, environ 171 sonl connues des ties Marquises.
DEROULEMENT DE LA CAMPAGNE MUSORSTOM 9
lTl^E^A'^ ~ La camPa8ne a cu lieu a bord du N. O. "Alts", au depart de Vairao situd sur h
presqu lie de 1 1 lie de Tahiti. Compte tenu du mauvais etat de la mer, trois jours et demi de route furent ndeessaires
DW ,7 r nbat d^r.^qUiT L° r" ddbU,a danS 16 n°rd dC llto d'U:1 (premiere station
destinces7ech-.m llonn^l . 7 ’ 7 °S S,aI'°nS de dragaSes Cn Peliles P™'«ndeurs (30 a 100 m,
& 7" c,rcalmo:a : ,oul en met,ant au poin' lc maldriel de P£che (Richer forges
Drofond^r nn ' r 7 7 65X3,5 rCUSS'S anS nord-esl de nie' sur des fonds rocheux de 1 50 a 450 m de
a u, h n 7S ^ 7 ' Z dC n0S d6UX dr3gUeS Waren Cel intidenI malencontreux devai. nous obLer
dan r"e“e Note Hiva o'i & "f'?' n0“ P°Ur ‘en‘er * Mrc “nstr“‘re ““ drague
i^“i=r3SfPfr
canardu^Baord,e!ais°SrchenaIdd[ro^fefDeu^^ll!'a'e|Ht Z '7 P“ttS ^ 0a p“ d“ “ions a idnas, d„
P ■ y tres accores el rocheuses el les profondeurs depassenl
Source : MNHN. Paris
CAMPAGNE MUSORSTOM 9
13
Motu Nao
ft)
HATIUTI
00
Banc
Lawson
Banc Clark
Dumont d'Urville
UA HUKA
FATU HUKU
•0
HIVA OA
Canal
du
Bordelais
139°30'
1 39c
8°30'
9°30'
10°30‘
Sud
Fig. 2. — Carte de l'archipel dcs lies Marquises (Polynesie fran^aise) montrant l'itineraire de la campagne MUSORSTOM 9.
Ua Pou (stations 1 142-1151, 1256-1265) ; Eiao (1152-1160, 1266-1280) ; Mom One (1281-1282) ; Hatutaa (1283-
1287) ; Ua Huka (1288-1297) ; Nuku Hiva (1161-1197, 1298-1307) ; Hiva Oa (1198-1239) ; Fan, Hiva (1240-1247) ;
liaul-fond Dumont d'Urville (1248-1255).
Source
14
B. RICHER DE FORGES. J POUPIN & P. LABOUTE
rapidement 2000 m et sont done au-dela des capacites de travail du N. O. "Alis". Unc escale de quelques heures a
Atuona (1300 habitants) permit aux plongeurs dc prelever des echantillons de la faune littoralc (3-27 m), en
particulier des especes fixees, spongiaires, ascidies, scleractiniaircs (PL 1216). Plusieurs dragages furent realises
dans la baie d'Atuona entre 50 et 120 m (stations 1211-1218), mettant en evidence dcs fonds de sable coquilliers a
articles d 'Halimeda. Quatre jours de travail (28-31 aout) permirent d'echantillonner egalement les pentes nord-est et
nord-ouest de 1'ile (stations 1217-1239). Un dragage ful reussi par 1110 m de profondeur rapportant dcs
octocoralliaires (pennatules), des mollusques scaphopodes et des crustaces Polychelidae (DR 1221).
L'exploration des pentes de Pile la plus au sud de Parchipel, Fatu Hiva, fut realisee le ler septembre, en une
seule journee. Les pentes y sont tellement rocheuses et accores qu'il fut tres difficile d'y realiser quelques
prelevements entre 70 et 1000 m de profondeur, bien que le tour de Pile ait etc explore dans sa totalitd
(stations 1240-1247).
Le haut-fond Dumont d'Urville, situe a egale distance des Ties de Hiva Oa et de Ua Pou fut echantillonne
le 2 septembre. II s'agit d'une sorte de dome culminant a 333 m de profondeur. Ses pentes sont assez douces mais
les fonds y sont rocheux (des echantillons de roches voleaniques ont ete rapportes. DR 1249. 1253). Un excellent
trait de chalut fut reussi par 500-600 m de profondeur (CP 1251), avec de trks nombreux oursins Cidaridae. Par
contre, au chalutage suivant, par 1000 m de profondeur, une tres forte croche provoquait la perte de la perche et une
avarie sur le cable. Comme e'est souvent le cas sur les monts sous-marins, les dragues ramenerent des sables
detritiques grossiers avec de nombreuses dents fossiles de requins et de raies, et des otolithes de poissons egalement
fossiles.
La journee du 3 septembre fut consacree aux prelevements de la pente sud de PTle d'Ua Pou (stations 1256-
1265).
Fig. 3. Lequipe scientifique embarqu£e h bord de \'"Alis’\ a I’exception de Pierre Laboute absent. De gauche a droite :
Joseph Poupin. Helmut Zibrowius, Bertrand Richer de Forges, Benoit Dayrat, Philippe Bouchet, Regis Cleva
En arrifcre-plan, le navire "Alis" mouille a Nuku Hiva.
Source : MNHN. Pans
CAMPAGNE MUSORSTOM 9
15
Les derniers jours furent utilises pour completer l'dchantillonnagc du groupe d’Tles les plus au nord, qui avait du
etre prematurement interrompu en debut de campagne : Eiao du 4 au 6 septembre (stations 1266-1280) ; Motu One
le 7 septembre (stations 1281-1282) ; Hatutaa le 7 septembre egalement (stations 1283-1287).
Cette zone presente des fonds en pente douce au nord-ouest jusqu'a 800 m de profondeur sur lesquels les chaluts
a perche ont bien fonctionne (CP 1270, 1271, 1272, 1276). La journee du 6 septembre fut decrefoe "journee de
repos" et, ce jour la, seulement deux dragages eurent lieu en zone circalittorale d'Eiao ainsi que des recoltes
littorales en plongee. Certains membres de Texpedition profiterent de cette escale pour recolter la faune
malacologique terrestre et d'eau douce de l'Tle d'Eiao.
L'Tlot Motu One est la seule formation non volcanique de I'archipel. II s'agit, en fait, dune grosse caye de sable
£rigee sur un vaste banc corallien. Deux prelevements seulement ont pu avoir lieu le 7 septembre sous le vent de
ce relief, par 450 m de profondeur. Un alizd de plus de 25 nceuds s'etant leve, les stations suivantes furent
effectuees a I’abri, au nord-ouest de l'Tle dc Hatutaa.
La journee du 8 septembre fut occupee a I'exploration de l'Tle Ua Huka. A la suite d'une forte croche par 800 m
de profondeur, fort heureusement a la fin dc la campagne, nous perdions notre deuxieme et derniere drague Waren
avec environ 500 m de cable (DW 1291 ).
Les 9 et 10 septembre furent employes a completer I'echantillonnage dans l'ouest de Nuku Hiva. La campagne
pnt fin le 10 septembre dans la matinee a Nuku Hiva, ce qui laissa Papres-midi pour reconditionner les
echantillons et changer le liquide fixateur. Les membres de la mission MUSORSTOM 9 regagnerent Tahiti par avion
le 1 1 septembre, afin de laisser le bateau aux geologues de la campagne suivante PALEOMARQ.
En complement de la campagne en mer, un atelier a terre
de quatre semaines (du 16 septembre au 19 octobre) a ete
organise sur l'Tle Ua Huka. Des recoltes dans la zone
intertidale et jusqu'a une trentaine de metres de profondeur, en
plongee el a la drague, ont ete effectuees par une equipe de
trois chercheurs.
Les dragages ont utilise une petite drague de 30 et 40 cm
de largeur, manceuvree a partir d'une embarcation a moteur
locale de 8 metres de longueur.
Quarante stations ont ete explorees (quelques-unes en
echantillonnage repete) et, au total, les operations suivantes
ont ete effectuees :
— 22 sorties a pied, en maree;
— 8 sorties de plongee en apnee entre 0,5 et 5 m de
profondeur.
— 115 traits de drague entre 5 et 32 m de profondeur.
— 3 recoltes en eau douce.
— 4 prelevements d echantillons pour la recherche dc
mollusqucs terrestres.
La liste detaillee des stations est publiee, en annexe, apres
celle des stations de la campagne MUSORSTOM 9.
Materiel et Methodes. — Pour cette campagne, nous disposions de deux dragues Waren seulement. Or des
la fin de la premiere journee de travail de la campagne, une drague fut perdue. Durant la suite de la campagne, la
seule drague restante nc fut utilisce qu'avec precaution sur des fonds plats et non rochcux. Toutes les autres opera¬
tions de dragages eurent lieu a l'aide d'une drague a roche (DR). Cette drague est habiluellemcnt employee par les
geologues pour rapporter des echantillons de roches volcaniques. Elle est constituee d'un cylindre de 40 cm de dia-
metre et de 60 cm de longueur, prolongc par un sac muni d'une cotte de maille (Fig. 5). Cet engin s'est avere assez
efficace sur les fonds tourmentes el rocheux des Ties Marquises. Des essais compares avec la drague Waren montre-
rent que la difference de recolte est seulement quantitative mais que les peuplements sont bien echantillonnes.
Fig. 4. — Lequipe de Tatelier 5 terre. De gauche a
droite : Jean Tardy, Jean Trondl£ et Rudo von
Cosel
16
B. RICHER DE FORGES. J. POUPIN & P. LABOUTE
Fig. 5. — La drague & roches (DR) est videe sur Ic pom au large de Nuku
Hiva (photo P. LABOUTE).
Le chalut a perche de 4 m a ete
maintes fois decrit (Forest, 1981). II
lut utilise des quc les fonds ctaient
suffisamment meubles, meme avec une
forte pentc. Par contre, aucun fond
permettant I'utilisation du chalut a
crevctte ne fut trouve.
Le tamisage a ete effectue dans l'eau,
sur mail les successives allant de 20 mm
a 1 mm. Le tri des residus supericurs a
3 mm a ete effectue a hord (Fig. 6) et
les refus de tailles inferieures ont ete
fixes et seront tries au laboratoirc pour
en extraire les micromolluscjues.
Certains crustaces ont fait I'objet de
dissections pour prelever les gonades
destinecs aux etudes sur la morphologie
des sperraatozoides (Albuneidae, Gery-
onidac, Pinnotheridae, Parthenopidae,
Palinuridae).
La plupart des especes de crustaces,
quelques mollusques et quelques pois-
sons ont fait I'objet de photographies en
couleurs.
COMMENTAIRES SUR LES ZONES PROSPECTEES ET LA FAUNE RECOLTEE
. LES/'rA,yEF0RMES CORALUENNES- — L'absence de veritables recifs coralliens autour des lies des Marquises a
intrigue differents auteurs (Agassiz, 1903 ; Crossland, 1927 ; Sournia, 1976 ; Brousse et al 1978 •
WAUTHY e, al., 1988 ; ROUGERIE er al., 1992), Par ailleurs, ces mernes auteurs avaient remarque que lesiles son.'
bordees d une sorte de plateforme de quelques milles dc largeur et dont la profondeur ne depassc pas 90-100 m Une
h^othese a done ete emise : ,1 s'agirait des ancicnnes constructions recifales vestige d'un ancien niveau marin La
simditude bathymetnque observee entre le rebord de ee plateau et les pen.es externes des recifs barrieres renforce
demanHP HSagTil r f°SSI'e Au cours de la ^ampagne MUSORSTOM 9, les dragages realises, a la
CO albcnneseSrf,< fn,™' nf'rmDenl bien ^ la ruPturc de pen.e (80-120 m) por.e de vieilles constructions
p'J i w t eS m°rtS rappor,ds dans lcs dragues (DW 1 173, 1 182, CP 1 191, DW 1201,
' ’ u’: . 6' , p 'r62' DW l275) Permettront, peut-etre. de verifier par datation si Page de ces coraux
correspond bien a celui de la derm&re glaciation (Bloom et al., 1974 ; Bard et al., 1996).
formn0nCHEVAfLiIER ^ Z Z* hypoth®ses expliqueraient la pauvrete des ties Marquises en especes et en
formations recifales : 1) P.solement geographique limitant Parrivee des larves, d'au.an, plus que les courants
sekc'Zne leesneTnl > V ^ mauvaises COnditions ^ologiques limitant l instal lation des larves e. ayant
selectionne les especes les plus resistantes. y
MarZZ^amvReZa/' h"^' iSSUeS dC '3 dern*re deg1aciation auraien. provoque, aux ties
d Une ,empera,ure Inf™e a 1 7°C- Provoquant la mort du recif barriere. Un phenomene
eaux et deI Z n n(w'qUe, P'atef0rme corallienne fossilc serai' a ''origine de Penrichissemen, des
eaux de la forte production planctomque. Ce. "effet d’Tle" caraclerise par une forte production primaire a ete mis
cause pour expliquer le faible developpement corallien (Sournia, 1976 ; Rougerie & Wauthy 1993)
perm-ZZotZST “T-™ 9 * dC '3 Campa«ne ”le PaleomarQ. les plough realises ont
Perm S R Laboute d observer la presence des sclerac.iniaires vivan.s suivants : Po rites lobata, P. (?) lutea.
Source : MNHN, Pahs
CAMPAGNE MUSORSTOM 9
17
Millepora platyphyllia , Pocillopora sp 1 ct sp 2, Leptoseris hawaiensis , Dendrophyllia sp., Fungia sp.,
Pavona sp., Montipora sp.
Parmi les coraux morts remontes dans les dragages pendant MUSORSTOM 9, figuraient des specimens du genre
Acropora , suppose absent de cettc region (DW 1201, 1275).
Fig. 6. — Tri du benthos sur la plage arriere, devant le paysage grandiose de l'lle d’Ua Pou. De gauche a droite
Regis Cleva, Benoit Dayrat, Philippe Bouchet, Bertrand Richer de Forges et Joseph Poupin (Photo P. Laboute).
La Richesse specifique. — L'isolement geographique et bathymetrique de cet archipel a certainement
conditionne la relative pauvrete des formations recifales entrainant celle des organismes marins qui leurs sont ordi-
nairement associes (PAULAY, 1997). II est bien connu en
ecologie theorique des ties que la biodiversite presente depend
& la fois de l’isolement, de la dimension, ct de la capacite
dispersive des organismes : "...the degree of isolation relative
to the mean dispersal distance will usually be decisive" (Mac
Arthur & Wilson, 1967).
Joseph Poupin, s'appuyant sur la 1 i tterature
(Monteforte, 1984 ; Poupin, 1996a, b. sous presse ;
Chan & Yu. 1998 ; Poupin & McLaughlin, sous-presse)
a trouve qu'environ 171 especes de crustaces decapodes et
stomatopodes ont ete signales aux ties Marquises. D’apres
ses observations pendant la campagne MUSORSTOM 9.
environ 60 especes supplementaires de crustaces ont ete
recoltees, ce qui porterait le nombre d'especes connues de
decapodes et stomatopodes a 240 especes. Les especes
nouvellement reconnues pendant MUSORSTOM 9 concernent
surtout des groupes de profondeur : des crevettes bathy-
pelagiques (Aristeidae, Benthesicymidae, Solenoceridae et
Fig. 7. — Mollusque gasteropode de la famille des
Architectonicidae, tres commun sur la
plateforme corallienne des lies Marquises.
18
B. RICHER DE FORGES, J POUPIN & P. LABOUTE
Sergestidae) ; des Stenopodidae ; des Crangonidae ; des Polychelidac ( Stereonuistis phosphorus) pas encore signales
du Pacific] ue central : des anomoures (Diogenidae, Galatheidae (Munida), Chirostylidae) ; des crabcs Dromiidac,
Dorippidae, Calappidae, Leucosiidae, Majidae, Portunidae, Xanthidae, Palicidae. Les families des Dorippidae et
Palicidae n etaient pas encore connues de cette region du Pacifique.
A 1'occasion du congres international sur les recifs coralliens qui s’cst tenu a Tahiti en 1985, une tentative
d'inventaire de la faune et de la flore marines de Polynesie franchise a etc faite (Richard. 1985). Pour le groupe des
mollusques, 262 especes connues des ties Marquises ont etc recensees a cette occasion. De nombreuses especes
supplementaires ont ete recoltees durant Musorstom 9. Sur la plateforme littorale, le groupe des gasteropodes
Architectonicidae (Fig. 7) et celui des bivalves Mytilidae sont particulierement frequents.
Le groupe des echinodermes est particulierement frequent et abondant dans les prelevements bien qu’il s’agissc
souvent de peuplcments peu diversifies d’oursins (Cidaridae, Loveniidac, Echinothuridae) et d'etoiles de mer
(Fig. 8). Bien que le groupe des crinoides ne soit pas cense s'etcndre jusqu'au Pacifique central, une tige de crinoide
pedoncule a ete recoltee aux ties Marquises sur le sommet du haut-fond Dumont d’Urville (DR 1255) et une
comatule vivante a ete rapportee de 1000 m de profondeur sur la pcnte sud d'Ua Pou (CP 1263).
Fig. 8. Exemple du conienu d un chalui b perche monirant un peuplement a dominante d'6chinodermes.
Le groupe des poissons est le seul qui semble presenter une diversitc specifique assez dlevee (une vingtaine
d especes par trait) avec des Moridae, Lophiidae ( Lophiomus ), Chaunacidae ( Chaunax ), Macrouridae,
Macrorhamphosidae, Chlorophthalmidae, Hoplichthyidae, Ogcocephalidae ( Halieutea ), Ophidiidae, ainsi que de
nombreux poissons plats Bothidae (Chascanopsetta spp.), Cynoglossidae, et Soleidae. Les stations de chalutages
au dela de 800 m de profondeur ramenerent en abondance un Hoplosthetus de couleur sombre et de petite taille
(Fig. 9). Signalons la presence, par 300 m de profondeur. dune raie de l’espece Plesiobathys daviesi (Fi«. 10).
Sur la plateforme littorale, vers 60 m de profondeur, on note I’abondance de Scorpaenidae et de Trislidae ainsi que
plusieurs petites especes de Pegasidae. "
Source : MNHN. Paris
CAMPAGNE MUSORSTOM 9
19
Pig 9 — Hoplostethus melanopus, poisson Fig. 10. — Raie recoltee & la station CP 1129 au large
b6ryciforme de la famille des Trachychtyidae recoltd d'Hiva Oa, Plesiobathys daviesi (identifiee par
h la station CP 1 195 (identifie par B. SERET.). B S£ret).
CONCLUSIONS
Le programme MUSORSTOM, debute aux lies Philippines, zone prochc du maximum de biodiversile marine de
la planete, vient, apres de nombreuses etapes intermediates, d’echantillonner I'archipel le plus eloigne vers lest,
dans la zone reputee la plus pauvre en especes de I'lndo-Ouest Pacifique. L'exploration de la faunc de profondeur des
Ties Marquises par la campagnc MUSORSTOM 9, avec des techniques efficaces ayant fait leurs preuves dans d'autres
regions de I'lndo-Ouest Pacifique et une equipc de recherche aguerrie, va permettre d’obtenir des donnees fiables sur
la diversitc specifique de cet archipel isole. L’inventaire des especes sera notablement accru dans tous les groupes
zoologiques pour lesquels il existe encore des taxonomistes. A partir de ces nouvelles bases, il sera alors possible
de tester les differentes hypotheses sur 1'origine des pcuplcments, les raisons de la pauvrete corallienne, specifique
et quantitative (isolement ou conditions environnementales defavorables) et l'appauvrissement des faunes marines
vers 1'est du Pacifique.
Une fois encore, il est demontre qu’avec des moyens relativement modestes, petit bateau, petite equipe de
recherche et campagne de courte duree, il est possible de renover totalement les connaissances sur la faune dun
archipel. Ce qui manque a la biogeographie marine dans l'lndo- Pacifique n’est pas tant des theories que des donnees
gdographiques et taxonomiques de qualite. Bien sur, l'equipe de terrain assurant la collecte, le tri et la preservation
des echantillons biologiques n'est que la partie emergente du programme MUSORSTOM qui ne saurait exister sans
les dizaincs de taxonomistes etudiant ce materiel de par le monde et surtout sans une edition efficace des resultats
scientifiques.
D'ores et deja, il est possible de confirmer que la plateforme entourant les lies correspond bien a des formations
recifales anciennes, lagon et recif barriere ; que la faune bathyalc est. lout comme la faunc littorale, beaucoup plus
pauvre en especes que celle du Pacifique ouest. L'absence supposee du groupe dcs crinoides dans le Pacifique central
est infirme.
Sur le plan quantitatif, seul le groupe des echinodermes presente de fortes concentrations d’echinides et
d'asterides. Aucun organisme pouvant faire 1’objct d'exploitation halieutique n'a ete detecte avec les engins utilises.
REMERCIEMENTS
Nous remercions vivement les personnes qui, dune maniere ou dune autre, 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 BOUCHET ; pour 1'aide logistique a Tahiti, Fran^ois-Xavier Bard,
20
B RICHER DE FORGES, J. POUPIN & P. LABOUTE
Pierre Carabasse et Raymond Guilhamat ; a bord du N. O. "Alis", le Commandant Raymond Proner a fait
le maximum pour realiser de bons prelevcmcnts dans des zones souvent non hydrographiees ; le Maitre d equipage
Loic LEGOFF a inlassablement ramende les chaluts ; les cartes sont I'oeuvre d'Yves Penvern, dessinatcur au
Centre ORSTOM de Noumea.
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mixte de Surveillance radiobiologique et biologique (SMSRB), Montlhery, 86 p.. 8 pi. couleur.
Poupin, J., 1996a. — Atlas des crustaces marins profonds de Polynesie Franyaise. Recokes du navire Marara
(1986/1996). Service mixte de Surveillance radiobiologique et biologique (SMSRB), Montlhery, 59 p.
Poupin, J., 1996b. — Crustacea Decapoda of French Polynesia (Astacidea. Palinuridea. Anomura, Brachyura). Atoll
Research Bulletin, 442 : 1-114.
Poupin, J., sous presse. — Crustacea Decapoda and Stomatopoda of French Polynesia (Dendrobranchiata. Stenopodidea
Caridea, Thalassinidea. and Stomatopoda, with additions to Astacidea, Palinuridea, Anomura, and Brachyura). Atoll
Research Bulletin, (451).
Poupin. J., Buat, P. & Ellis, T., 1991. — Les crabes profonds dcs Ties Marquises ( Chaceon sp. nov. - Decapoda -
Geryonidae). Rapport scientifique et technique. Service mixte de Controle biologique, (SMCB), Tahiti, 4. p.
Poupin, J. & McLaughlin, P., sous-presse. — Additional Calcinus (Decapoda, Anomura. Diogenidae) from French
Polynesia with three new species and a key to Indo-West Pacific species. Crustacean Research. Tokyo.
Poupin, J. & Richer de Forges, B., 1991. — New or rare crustaceans from french Polynesia (Crustacea : Decapoda).
In : P. J. F. Davie, & R. H. Quinn (eds). Proceedings of the 1990 International Crustacean Conference, Brisbane,
1 September, 1991. Memoirs of the Queensland Museum, 31 : 211.
Poupin. J., Tamarii, T. & Vandenboomgaerde. A., 1990. — Peches profondes aux casiers sur les pentes externes des lies
de Polynesie Frangaise (N/O Marara, 1986:1989). Notes et Documents d'Oceanographie du Centre ORSTOM de laluti.
(42) : 1-100.
Randall, J. E.. 1978. — Marine biological and archeological expedition to southeast Oceania. National Geographic
Society Research Reports, 1969 Projects : 473-495.
Randall, J. E., 1985. — Fishes. In : G. Richard, Faune et Flore. Premier abreg£ des organismes marins de Polynesie
Frangaise. In : B. Delesalle, R. Galzin & B. Salvat, French Polynesian coral reefs. Volume 1. Proceedings of the 5th
International Coral Reef Congress, Tahiti, 21 May- 1 June 1985 : 462-481.
Rehder, H. A.. 1968. — The marine Molluscan fauna of the Marquesas Islands. Bulletin of the American Malacological
Union Inc., Annual Reports, 1968 : 29-32.
Richard, G.. 1985. — Faune et Flore. Premier abrege des organismes marins de Polynesie Fran<;aise. In : B. Delesalle,
R. Galzin & B. Salvat, French Polynesian coral reefs. Volume 1. Proceedings of the 5th International Coral Reef
Congress, Tahiti, 27 May- 1 June 1985 : 381-518.
22
B. RICHER DE FORGES. J. POUPIN & P. LABOUTE
Richer de Forges, B., 1990. — Les campagnes d'exploration dc la faune bathyale dans la zone economique de la
Nouvelle-Caledonie / Explorations for bathyal fauna in the New Caledonian economic zone. 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., 1993. — Campagnes d'exploralion de la faune bathyale faites depuis mai 1989 dans la zone
economique de la Nouvelle-Caledonie. In : A. Crosnier (ed.). Resultats des Campagnes Musorstom, Volume 10.
Memoires du Museum national d'Histoire naturelle, 156 : 27-32.
Richer de Forges. B., Faliex. E. & Menou, J.-L.. 1996. — La campagne Musorstom 8 dans 1'archipel de Vanuatu.
Compte rendu et liste des stations, in : A. Crosnier (ed.), Resultats des Campagnes Musorstom, Volume 15.
Memoires du Museum national d'Histoire naturelle , 168 : 9-32.
Richer de Forges, B. & Laboute, P.. 1998. — La campagne Musorstom 9 aux ties Marquises. ORSTOM-Actualites,
(55): 8-14.
Richer de Forges, B. & Menou, J.-L., 1993. — La campagne Musorstom 7 dans la zone economique des ties Wallis et
Futuna. Compte rendu et liste des stations. In ; A. Crosnier (ed.), Resultats des Campagnes Musorstom, Volume 10.
Memoires du Museum national d'Histoire naturelle , 156 : 9-25.
Rosen, B. R., 1988. — Progress, problems and patterns in the biogeography of reef corals and other tropical marine
organisms. Helgoldnder Meeresuntersuchungen, 42 : 269-301.
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upwelling. Comptes Rendus hebdomadaires des Seances de V Academic des Sciences, Paris, 315 (ser. 2) : 677-682.
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E. Vigneron (eds), Atlas de la Polynesie Frangaise. Editions de l’ORSTOM : 20-21.
Sournia, A.. 1976. — Abondance du phytoplancton et absence de recifs coralliens sur les cotes des ties Marquises.
Comptes Rendus hebdomadaires des Seances de F Academic des Sciences, Paris , 282. ser. D : 553-555.
Springer, V. G., 1982. — Pacific Plate Biogeography, with special reference to shorefishes. Smithsonian Contributions
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Press, Sydney .
Wauthy, B., Rougerie, F., Charpy, L.. Rancher, J. & Thouard, M., 1988. — Formations recifales et effet d’tle par
endo-upwelling autour des ties Marquises. Notes et Documents du Centre ORSTOM de Tahiti. (37), 36 p.
Wheeler, T. & Carillet, J.-B., 1997. — Tahiti & French Polynesia. 386 p. Lonely planet. Travel survival kit.
Zezina, O. N., 1997. — Biogeography of the bathyal zone. In : A. V. Gebruk, E. C. Southward & P. A. Tyler (eds). The
biogeography of the oceans. Advances in Marine Biology. 32 : 389-426.
Source : MNHN. Paris
CAMPAGNE MUSORSTOM 9
23
ANNEXES
LISTE DES PARTICIPANTS A LA CAMPAGNE MUSORSTOM 9
Chef de mission : B. Richer de Forges
Autres participants : P. BOUCHET, R. Cleva, B. Dayrat, P. Laboute, J. POUPIN, H. Zibrowius.
LISTE DES STATIONS DE LA CAMPAGNE MUSORSTOM 9
(DW : drague Waren ; DR : drague a roche ; CP : chalut a perche)
Source : MNHN, Paris
B. RICHER DE FORGES. J. POUPIN & P. LABOUTE
CAMPAGNE MUSORSTOM 9
25
DR 1220
DR 1221
DW 1222
DR 1223
DW 1224
DW 1225
CP 1226
CP 1227
CP 1228
CP 1229
DW 1230
DR 1231
DR 1232
DW 1233
DW 1234
DW 1235
DW 1236
CP 1237
CP 1238
CP 1239
FATU H1VA
26
B. RICHER DE FORGES. J POUPIN & P. LABOUTE
CAMPAGNE MUSORSTOM 9
27
LISTE DES PARTICIPANTS A L' ATELIER A TERRE DE UA HUKA
Chef d’atelier : Rudo VON COSEL.
Autres participants : Jean TARDY, Jean Trondle.
LISTE DES STATIONS DE L' ATELIER A TERRE
Station 1. Baie de Heine, cot6 ouest de la baie. 8°55,6 S, 139°32,1 W. .
Zone intertidale, sous les rochets et cailloux (pierres rondes volcaniques plus quelques morceaux de corail et
beachrock).
Station 2. Baie de Heine. 8°55,6’S, 139°32,1'W.
Zone supralittorale, rochers et sable.
Station 3. Beiie de Heine. 8°55,5'S, 139°32,1'W.
Digue au fond de la baie. Zone supralittorale.
Station 4. Beiie de Haavei. 8°56,50'S, 139°35,63'W.
Cote est de la baie, trottoir tres battu, sur et entre des rochers.
STATION 5. Baie de Haavei. 8°56,45'S, 139°35,78'W.
Embouchure du ruisseau cote ouest, eau douce stagnante derriere une dune de sable.
Station 5bis. Baie de Haavei. 8°56,45'S, 139°35,80'W.
Embouchure cote mer, rochers et sable. Zone supralittorale.
Station 6. Baie de Haavei. 8°56,44'S, 139°35,80'W.
Dans la vallee proche du ruisseau, bas de la falaise ouest, dans la liliere. Recoltes terrestres.
Station 7. Baie de Veiipaee. 8°56,30'S, 139°34,40'W.
Digue cote est. Zone supralittorale.
STATION 8. Baie de Vaipeiee. 8°56,00'S, 139°34,50’W.
Vaipaee, riviere face au "Chez Christelle", sur el sous les cailloux. Recoltes fluviatiles.
Station 9. Baie de Vaipeiee. 8°56,30'S, 139°34,40'W.
Sable intertidal proche de l'embouchure de la riviere.
Station 10. Baie de Vaipaee. 8°56,30’S, 139°34,40'W.
Cote est, sur la rampe destinee a la mise a l'eau des bateaux.
Station 11. Beiie de Veiipaee. 8°56,30'S, 139°34,50'W.
Cote ouest, cote rocheuse. Zone intertidale.
STATION 12. Baie Hiniaehi - beiie Hinitaihava. 8°56,00’S, 139°32,80'W.
Large trottoir de rocher avec des grandes cuvettes a fonds de sable ou d algues, alimentees par la hou e.
Station 13 .Baie Vaioina. 8°52,50'S, 139°35,20’W.
Falaise et fonds rocheux, 0-5 m. Plongees.
Station 14. Baie Haahevea. 8°53,15’S, 139°35,60’W.
Falaise et fonds rocheux avec de grands rochers et des algues encroutantes, 0-3 m. Plongees.
Station 15. Baie Manihina. 8°56,40'S, 139°33,80'W.
Cotes est et ouest de la pointe rocheuse; cailloux. blocs ct cuvettes a algues. Zone intertidale.
STATION 16. Baie Haneiinamoa. 8°53,35'S, 139°35,70'W.
Falaise; fonds a cailloux, pierres et gros blocs, 0.5-3 m. Plongees.
Source : MNHN. Paris
28
B. RICHER DE FORGES. J POUPIN & P. LABOUTE
Station 17. BateHaahue. 8°53,65'S, 139°35,95'W.
Falaise; fonds a pierres et gros blocs, 0,5-3 m; plateau couvert d'algues vers 1-1,5 m. Plongees.
Station 18. Baie (Pointe) Kuiapaku (est de Manihina). 8°56,45'S, 139°33,50'W.
Cote rocheuse, trottoir avec cuvettes garnics de sable, cailloux el algues; plateforme battue; plage de galcts.
Station 19. Baie de Hane, cote ouest. 8°55,65’S, 139°32,40'W.
Dalle de corail avec prairie d'algues (surtout Halimeda), cailloux, morceaux de corail, aussi des gros blocs et
quelques patates de corail (Porites spp. etc.) isolees et peu de sable, 0-3 m. Plongees et recoltes a pied.
Station 20. Baie Hinipahue. 8°56,20’S, 139°32,90'W.
Cote est et ouest d'une langue de trottoir, grandes cuvettes et baignoires a fonds de sable, algues et cailloux;
dalle avec prairie d'algues et Iegere couche de sable. Zone intertidale.
Station 2 1 . Baie Hiniaehi. 8°56,15'S, 139°32,90'W.
Fonds de pierres et cailloux (volcaniques) avec quelques algues. Milieu battu, 0-1 m. Recoltes a pied.
Station 22. Baie de Vaipaee. 8°56,40'S, 139°34,40'W - 8°56,60’S, 139°34,25'W.
Dragages 6-10 m, sable fin gris, vaseux vers la plage, plus propre au large. 6 traits.
Station 23. Partie ouest de la baie Haamamao (a 1'est de Hokatu). 8°55,90’S, 139°31,45'W.
Cote rocheuse battue avec des cuvettes contcnant du sable et des cailloux. Prise d'un echantillon de sable.
Station 24. Baie Haahue. 8°53,60'S, 139°37,00’W.
Dragages 9-25 m, sable fin.
Station 24 bis. Baie Haahue . 8°53,60’S, 139°37,00'W.
Dragages 25-34 m, sable fin et moyen. 10 traits pour I'ensemble des stations 24 et 24 bis.
Station 25. Baie Teuahai. 8°55,70'S, 139°36,70'W.
Dragages 6-15 m, sable calcaire glossier et galets et Halimeda. 3 traits.
Station 26. Baie Manihina. 8°56,35'S, 139°33,70’W.
Cote est de la baie, sous rochers h marce haute et en zone supralitlorale.
Station 27. Baie Manihina. 8°56,50’S, 139°33,95'W.
Cote ouest de la baie. Rochers. Zone intertidale.
Station 28. Vallee de Vaipaee. 8°54,90'S, 139°34,20'W.
Sortie terrestre. Route vers le fond de la vallee, ravin humide.
Station 29. Baie de Hane. 8°55,70’S, 139°32,00’W.
Dragages 7-11 m, sable fin, mixte et grossier. 10 traits.
STATION 30. Entre Motu Hane et pte Teavatainamanu. 8°56,10’S, I39°32,00’W.
Dragages 20-30 m a la sortie de la baie, sable calcaire plus ou moins grossier. 3 trails.
Station 31. Baie Hiniaehi. 8°56,10'S, 139°32,70'W.
Dragages 4-7 m. Sable fin, cailloux et dalle avec algues. 5 traits.
Station 32. Baie Hiniaehi 8°56,I0'S, 139°32,70’W.
Dragages 12-17 m, sable fin, cailloux et algues. 4 traits.
Station 33. Baie de Manihina. 8°55,60'S, 139°33.90’W.
Dragages 15 m, sable, cailloux et patates. 2 traits avec des croches.
Station 34. Entre baie Haavei. pointe Tenoni et He au x Oiseaux (Tlot Teuaua). 8°56,80'S, 139°35,70'W (position
sommaire).
Dragages 10-15 m. Entree de la baie (6 traits) et entre tie aux Oiseaux et pte Tenoni (10 traits). Sable calcaire,
cailloux et boules d'algues calcaires.
Source : MNHN. Paris
CAMPAGNE MUSORSTOM 9
29
STATION 34 bis. Dragages 20-32 m. Entre tie aux Oiseaux, Motukeokeo et pte. Teaeopiki. 8°57:00'S,
139°36,00'W (position sommaire). Sable fin avec dc debris de coquilles de Pinna et quelques algues calcaires et
cailloux proche des Tlots. 19 traits.
Station 35. Baie Haamamao. 8°55,90’S, 139°31,20’W.
Dragages 24-25 m, sable calcaire et debris de coquilles. 3 traits.
Station 35 bis. Haavahae. 8°55,85'S, 139°30,60'W.
Dragage 20 m, sable. 1 trait.
STATION 36. Baie de Hone. 8°55,60’S, 139°32,17’W.
Platier intertidal avec algues, 0,5 m. Brossage de cailloux
Station 37. Baie Haamamao. 8°55,80'S, 139°31,00'W.
Sable et cailloux, 4-5 m. Plongees.
Station 38. Au large de la baie de Vaipaee. 8°56,70'S, 139°34,23'W - 8°56,90'S, 1 39 34,23 W.
Dragages 10-25 m, sable et algues calcaires avec debris dc coquilles. 4 traits.
Station 39. 8°56,10’S, 139°34,70'W. Dans le ravin du chemin vers Haavei.
Prise d'echantillon de litiere et de terre.
STATION 40. 8°55,80'S, 139°34,60'W. Dans un ravin plus au nord en face colonic de vacances, proche de la riviere.
Prise d'dchantillon de litiere et de terre.
Source
Source : MNHN, Paris
S DE S CAMPAGNES MUSORSTOM. VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES CA
Cnidaria Anthozoa: Deep-water
azooxanthellate Scleractinia
from Vanuatu, and Wallis and Futuna Islands
Stephen D. CAIRNS
National Museum of Natural History
NHB-163. W-329
Smithsonian Institution
Washington, D. C. 20560
USA
ABSTRACT
A total of 134 Recent species of azooxanthellate Scleractinia are reported from the Vanuatu <1 16 species) and Wallis
and Futuna (83 species) Archipelagos, all but one being new records for this region of the tropical central Pacific. The
newly reported specimens originate primarily from the MUSORSTOM 7 and 8 expeditions, including approximately
4400 specimens from 227 stations, most of these stations from deeper than 100 m. Sixteen new species and one new
subspecies arc described, and two new combinations are proposed: Asterosmilia gigas and Javania fusca. Tables of
comparison are provided for the Indo-Pacific species of Fungiacyathus ( Fungiacyaihus ); the Recent Trochocyathus
(Aplocyathus)', all species of Aulocyathus: all species of spined Deltocyathus\ and the Recent species and subspecies of
Anthemiphyllia. To facilitate comparisons of species among these taxa. three additional species having distributions
other than the Vanuatu/Wallis and Futuna region are described as new: Deltocyathus corrugatus, Anthemiphyllia
multidentata , and A. macrolobata.
The distribution and bathymetric ranges of the 134 species known from the Vanuatu/Wallis and Futuna region are
tabulated. Within the tropical central Pacific these corals show a strong affinity with those from the ridges and islands
north of New Zealand (56 species) and a lesser relationship with the Hawaiian Island fauna (24 species). Other regions in
the central Pacific are too poorly known for comparison. Beyond the tropical central Pacific, the Vanuatu/Wallis and
Futuna fauna is part of the larger Indo-Polynesian province, sharing 95 (71%) of its species with the tropical western
Pacific and 62 species (46%) with the Indian Ocean. Only seven species are found in common with the tropical eastern
Pacific and I 1 with the Atlantic Ocean. Finally, 43 species from the Vanuatu/Wallis and Futuna Archipelagos are also
known from temperate Japan (exclusive of the Ryukyu Islands) and 32 from temperate New Zealand and southern
Australia.
Examples of commensal/parasitic relationships are reported to occur with petrarcid ascothoracican crustaceans
(2 coral hosts) and acrothoracican cirripede crustaceans (8 hosts). The shells of the gastropod Xenophora ("carrier
shells") were found to be effective collectors of deep-water corals; a total of 19 coral species were found incorporated into
the shells, including three species that were found only on these shells and another five species that were otherwise very
rarely collected by conventional means.
Cairns, S. D., 1999. — Cnidaria Anthozoa: Deep-water azooxanthellate Scleractinia from Vanuatu, and Wallis and
Futuna Islands. In: A. Crosnier (ed.), Resultats des Campagncs MUSORSTOM, Volume 20. Memoires du Museum national
d'Histoire naturelle , 180: 31-167. Paris ISBN 2-85653-520-8.
32
S. D. CAIRNS
RESUME
Cnidaria Anthozoa : Scleractiniaires d'eau profonde sans zooxanthelles du Vanuatu et des
lies Wallis et Futuna.
Cent-trente-quatre especes recentes de Scleractiniares sans zooxanthelles sont recensees dans I'archi pel du Vanuatu
(116 especes) et aux lies Wallis et Futuna (83 especes). A une exception pres, il s'agit de signalisations nouvelles pour
cette region du Pacifique central tropical. Lc materiel etudie provient essentiellement des campagnes Musorstom 7 et 8 et
comprend environ 4400 specimens provenant de 227 stations, le plus souvcnt a plus de 100 m de profondeur. Seize
especes nouvelles ( Fungiacytahus sandoi, Anthemiphyllia spinifera, Caryophyllia abrupta. Oxysntilia corrugata,
O. epithecata, Trochocyathus efateensis, T. pcitelliformis, Polycyathus octuplus, Deltocyathus crassiseptum,
D. camercitus, Conotrochus asymmetros, Lochmaeotrochus gardineri, Cryptotrochus brevipalus, Flabellum arcuatile,
Truncatoflabellum vigintifarium, et Javania exserta) et une sous-espece nouvelle ( Anthemiphyllia patera costata) sont
decrites et 2 combinaisons nouvelles sont proposees ( Asterosmilai gigas et Javania fusca). Des tableaux de comparison
sont fournis pour les especes indo-pacifiques de Fungiacyathus ( Fungiacyathus ), les especes recentes de Trochocyathus
(Aplocyathus), toutes les especes d 'Aulocyathus, toutes les especes de Deltocyathus h epincs et les especes et sous-
esp£ces d' Anthemiphyllia. Afin de faciliter les comparisons, trois especes nouvelles d'origine autre que Musorstom 7 et
8 ( Deltocyathus corrugatus, Anthemiphyllia multidentata, et A. macrolobata) sont incluses dans les tableaux.
Les repartitions geographique et bathymetrique des 134 especes connues du Vanuatu et de Wallis et Futuna sont
rassembl£es dans un tableau. A I'interieur du Pacifique central tropical, la faune etudi£e ici a une forte affinite avec celle des
rides et lies se trouvant au nord de la Nouvclle-Zelande (56 especes) et une affinite moindre avec la faune des Ties Hawaii
(24 especes). Les autres regions du Pacifique central sont trop peu connues pour etre compares. Au vu de leur faune, la
region du Vanuatu et celle de Wallis el Futuna font partie de la large province biogeographique indo-polynesicnnc. Elies
ont en commun 95 especes (71%) avec le Pacifique occidental tropical et 62 (46%) avec 1’ocean Indien. Seulcs sept
especes sont connues aussi dans le Pacifique oriental tropical et onze dans 1’Atlantique. Par ailleurs. si Ion compare cette
faune & celles de quelques regions temperees du Pacifique, on releve que 43 des 134 especes recensees au Vanuatu et aux Ties
Wallis et Futuna sont connues aussi au Japan et 32 en Nouvelle-Zelande et dans l’Australie du Sud.
Les relations commensales ou parasites observees concernent les crustaces ascothoraciques Petrarcidae (deux especes
hotes) et les crustaces cirripedes acrothoraciques (huit especes holes). Des coraux de profondeur sont souvcnt incorpores
dans les coquilles des gasteropodes Xenophora. Dix-neuf especes de coraux ont ainsi ete trouv£es sur ces coquilles
"porteuses"; dont trois n’ont pas ete recoltees autrement et cinq n’ont ete que ires rarement recoltees par les engins de
capture convenlionnels.
CONTENTS
Introduction .
List of Abbreviations .
Combined List of Stations and of Specie s obtained per Station
Historical Review .
Material .
Methods .
Distribution .
Systematic Account .
Family Pocilloporidae .
Family Fungiacyathidae .
Family Micrabaciidae .
Family Oculinidae .
Family Anthemiphylliidae .
Family Caryophylliidae .
Family Turbinoliidae .
Family Guyniidae .
Family Flabellidae .
Family Gardineriidae .
Family Dendrophylliidae .
33
33
34
46
48
48
49
53
53
54
59
60
63
69
108
113
1 15
128
129
Source : MNHN, Paris
AZOOXANTHELLATE SCLERACTINIA
33
ACKNOWLEDGEMENTS
References .
INTRODUCTION
The azooxanthellate corals of Vanuatu, Wallis, and Futuna Archipelagos arc representative of the tropical
central Pacific, which, in turn, is very closely tied to that of the Indo-West Pacific fauna. For the purpose of this
study, the tropical central Pacific is defined as those islands east of the Philippines, Indonesia. New Guinea, and
Australia, and extending to the Tuamotu Archipelago in the east, the Hawaiian Islands to the north, and the
Kermadec Islands to the south. Although this vast region was and is still not comprehensively sampled, the
intensive MUSORSTOM collections provide a good basis for the deep-water coral fauna for this province, and provide
zoogeographic data for one of the last poorly-known deep-water coral faunas. It should be noted that, whereas Table
1 lists all papers pertaining to azooxanthellates in the central tropical Pacific, the MUSORSTOM collections made
very few collections shallower than 200 m or deeper than 1500 m, which reduces the number of species reported
herein.
LIST OF ABBREVIATIONS
Museums:
MNHN Museum National d'Histoire Naturelle, Paris.
MoNZ Museum of New Zealand Te Papa Tonga-rewa, Wellington (formerly the National Museum of
New Zealand: NMNZ).
NHM The Natural History Museum, London (formerly the British Museum (Natural History):
BMNH).
N M V National Museum Victoria, Melbourne.
NMNH National Museum of Natural History, Smithsonian, Washington. D. C.
NZOI New Zealand Oceanographic Institute, Wellington.
USNM United States National Museum, now the National Museum of Natural History. Smithsonian,
Washington, D.C.
Y PM Yale Peabody Museum, New Haven.
ZMB Zoologisches Museum, Berlin.
Expeditions and Vessels:
Throughout
BANZARE
CANCAP III
HURL
KARUBAR
MUSORSTOM
USGS
the paper, the names of vessels are given in italics and quotation marks.
British, Australian. New Zealand Antarctic Research Expedition, 1929-1931.
"Tydeman", Madeira-Mauritania Expedition 1978. Initiated by the Nationaal Natuur-
historisch Museum (Leiden). Named for Canaries and Cape Verde Islands.
Hawaiian Underseas Research Laboratory.
Frcnch-Indonesian expedition ( 1991 ) that collected in the southeastern Banda Sea. Named
for the Kai, Aru, and Tanimbar Islands.
Cruises organized jointly by the Museum National d'Histoire Naturelle and the Institut
Frangais de Recherche Scientifique pour le Developpcment en Cooperation (formerly:
Office de la Recherche Scientifique et Technique d'Outre-Mer, = ORSTOM).
United States Geological Survey.
Morphological Terms:
CD
GCD
GCD:LCD
Calicular diameter.
Greater calicular diameter.
Ratio of greater calicular diameter to lesser calicular diameter.
34
S. D. CAIRNS
P D Pedicel diameter.
PD:GCD Ratio of pedicel diameter to greater calicular diameter of a solitary corallum.
Sx, Cx, Px Septa, costae, or pali (respectively) of cycle designated by numerical subscript.
S x > S y In the context of a septal formula, septa of cycle x more wide than septa of cycle y.
COMBINED LIST OF STATIONS AND OF SPECIES OBTAINED PER STATION
This list provides the data for all stations mentioned in this report. Vessels and expeditions are listed
alphabetically, whereas listing of species per station follows the order of the text. If the specimen was collected as
a result of being cemented to a Xenophora gastropod shell, the species name is followed by an X in brackets.
Banzare
Stn 115. — 24.03.1931, 41°03'S, I48°42'E. 128 m, northeastern Tasmania: Anthemiphyllia multidentata.
Cancap
Stn 3.053. — 20.10.1978, 32°25'N, 16°57'W, 2850-3010 m, south of Madeira: Truncatoflabellum stabile.
Hurl
Stn 83-202. — 16.10.1983, 16°4ri2"N, 169°24’18"W, 183 m, Johnston Atoll: Madracis kauaiensis.
Stn P5-063. — 23.05.1988, 20°35.8,N, I56°03.5'W, 1020 m, Alenuihaha, Hawaiian Is: Trochocyathus
( T rochocyathus ) patelliformis.
Karubar, "Banina Jaya r
Stn 5. — 22.10.1991, 5°46'39"S, 132°20’04"E, 285-323 m, Kai Is: Trochocyathus (Trochocyathus) vasiformis.
Stn 18. — 24.10.1991. 5°17’49" S, I33°0r51ME, 205-212 m, Kai Is: Anthemiphyllia spinifera.
Stn 44. — 29.10.1991, 7°52'27"S, 132°48'24"E, 291-295 m, Tanimbar Is: Javania exserta.
Stn 49. — 29.10.1991, 7059'51"S, 132°58’50"E, 206-209 m, Tanimbar Is: Javania exserta.
Stn 86. —04.1 1.1991, 9°23’59"S, \3\014'29UE, 222-226 m, Tanimbar Is: Javania exserta.
Kimbla
Stn K7/73-37. — 23.11.1973, 39°02.0'S, 148°36.5'E, 256 m, northeastern Tasmania: Anthemiphyllia
multidentata.
MUSORSTOM 1. "Vauban"
Stn 65. 27.03.1976, 14°00.0'N, 120°19.2'E, 194-202 m, SW Luzon, Philippines: Javania exserta.
Musorstom 3, " Coriolis "
Stn 87. 31.05.1985, 14°00.6'N, 120°19.6'E, 191-197 m, SW Luzon, Philippines: Aulocyathus juvenescens.
Musorstom 7, "Alis"
Stn 494. 10.05.1992, 14°18.9'S, 178C03.0'W, 100-110 m, Futuna: Polycyathus octuplus, Heterocyathus cf.
sulcatus , Heteropsammia cochlea
Stn 495. 10.05.1992, 14°19.2'S, 178°04.3'W. 180-210 m. Futuna: Heterocyathus cf. sulcatus. Conotrochus
asym metros, Truncatoflabellum phoenix, Heteropsammia cochlea.
Source : MNHN. Paris
AZOOXANTHELLATE SCLERACTINIA
35
Stn 4%. _ 10.05.1992, I4°19.6’S, 178°04.3'W, 250-330 m, Futuna: Madracis kauaiensis. Caryophyllia
(Acanthocyathus) grayi , Oxysmilia epithecata , Trochocyathus (Trochocyathus) philippinensis, Polycyatlms
octuplus , Heterocyathus cf. sulcatus , Conotrochus asymmetros , Thalamophyllia tenuescens , Balanophyllia
desmophyllioides. Heteropsammia cochlea.
Sln 499. _ 10.05. 1992, 14°19.6'S, 178o04.6'W, 290-395 m. Futuna: Madracis kauaiensis , Madrepora porcellana ,
Caryophyllia {Acanthocyathus) grayi, Trochocyathus (Trochocyathus) maculatus, Polycyatlms octuplus,
Thalamophyllia tenuescens , Dactylotrochus cervicornis , Truncatoflabellum Vanuatu , Javan ia exserta,
Endopachys grayi.
Stn 500. — 1 1.05.1992, 14°19.5’S, 178o04.FW, 350-394 m, Futuna: Madracis kauaiensis, Madrepora porcellana,
Balanophyllia desmophyllioides.
Sln 5oi. _ 11.05.1992, 14°19.8'S, 178°06.FW, 500-530 m, Fuluna: Truncatoflabellum angustum .
Enallopsammia rostrata.
Stn 502. — 11.05.1992, 14°19.8'S, 178°06.5'W, 516-535 m. Futuna: Madracis kauaiensis, Madrepora oculata
forma tenuis, M . porcellana, Caryophyllia ( Caryophyllia ) lamellifera, Crispatotrochus ntbescens,
Truncatoflabellum angustum, Balanophyllia desmophyllioides.
Sln 504. — 11.05.1992, 14°19.6'S, 178°04.5'W. 300-390 m, Futuna: Madracis kauaiensis. Caryophyllia
( Caryophyllia ) hawaiiensis, Trochocyathus ( Trochocyathus ) maculatus. Polycyatlms octuplus, Conotrochus
asymmetros, Thalamophyllia tenuescens, Dactylotrochus cervicornis, Guynia annulata, Flabellum ( Flabellum )
pavoninum, Endopachys grayi, Heteropsammia cochlea.
Sm 505. — 11.05.1992, 14°I9.5'S, 178°04.3'W, 245-400 m. Futuna: Trochocyathus (Trochocyathus) maculatus,
Truncatoflabellum Vanuatu.
Sm 506. — 1 1.05.1992, 14°19.8'S, 178°05.0’W, 400 m, Futuna: Dendrophyllia alcocki.
Sm 507. — 11.05.1992, I4°19.6'S, 178°06.7'W. 419-425 m, Futuna: Madracis kauaiensis, Madrepora oculata
forma formosa, Crispatotrochus rubescens, Oxysmilia epithecata, Endopachys grayi.
Sm 508. — 1 1.05.1992, 14°19.5'S, 178°04.5'W, 245-400 m, Futuna: Madracis kauaiensis, Madrepora porcellana.
Trochocyathus (Trochocyathus) maculatus.
Sm 509. — 12.05.1992, 14°14.8'S, 1 78° 1 1.5'W, 200-240 m. Fuluna: Madracis kauaiensis. Madrepora porcellana.
Oxysmilia epithecata, Trochocyathus ( Trochocyathus ) maculatus, Polycyatlms octuplus. Bourneotrochus
stellulatus, Deltocyathus Stella, Heterocyathus cf. sulcatus, Conotrochus asymmetros, Thalamophyllia
tenuescens, Rhizosmilia robusta. Idiot rochus kikutii, Truncatoflabellum phoenix, T. Vanuatu, Balanophyllia
desmophyllioides, Heteropsammia cochlea.
Sm 510. — 12.05.1992. 1 4° 1 4.5'S, 1 78° 1 1.5'W, 280-370 m, Futuna: Bourneotrochus stellulatus. Deltocyathus
Stella, Heterocyathus cf. sulcatus, Asterosmilia gigas.
Sm 511. _ 12.05.1992, I4°14.0’S, 1 78° 1 1.5'W, 400-450 m. Futuna: Anthemiphyllia dentata, Crispatotrochus
rubescens, C. rugosus, Oxysmilia epithecata, Tethocyathus virgatus, Bourneotrochus stellulatus. Deltocyathus
crassiseptum, Deltocyathus Stella, Conotrochus funicolumna, C. brunneus, Rliizotrochus flabelliformis.
Stn 512. — 12.05.1992, 14° 1 3.5'S, 178°10.3'W, 210-245 m, Futuna: Madrepora porcellana, Anthemiphyllia
spinifera, Bourneotrochus stellulatus, Deltocyathus Stella, Heterocyathus cf. sulcatus. Conotrochus
asymmetros. Thalamophyllia tenuescens, Dactylotrochus cervicornis, Deltocyatlioides orientalis, hhotrochus
kikutii, Truncatoflabellum phoenix, Balanophyllia desmophyllioides, Heteropsammia cochlea.
Stn 513. — 12.05.1992, 14° 1 3.5'S, 178°10.8'W, 260-300 m, Futuna: Madrepora minutiseptum, M. porcellana.
Anthemiphyllia spinifera, Polycyatlms octuplus, Bourneotrochus stellulatus. Deltocyathus Stella.
Heterocyathus cf. sulcatus, Conotrochus brunneus, C. asymmetros. Thalamophyllia tenuescens, Dactylotiochus
cervicornis. Idiotrochus kikutii, Guynia annulata. Flabellum ( Flabellum ) pavoninum. Truncatoflabellum
phoenix , Jcivcinia exserta.
Stn 514. _ 12.05.1992, 14°13.3'S, 178°10.7'W. 349-355 in, Futuna: Madrepora porcellana. Anthemiphyllia
spinifera, Oxysmilia epithecata, Bourneotrochus stellulatus, Deltocyathus Stella. D. heteroclitus, Heterocyathus
cf. sulcatus. Thalamophyllia tenuescens, Dactylotrochus cervicornis, Truncatoflabellum phoenix. Javania
exserta , Dendrophyllia alcocki.
36
S. D. CAIRNS
Stn 515. — 12.05.1992, 14°13.5’S, 178°10.3'W, 224-252 m. Futuna: Madracis kauaiensis, Madrepora
minutiseptum , M. porce liana, Dactylotrochus cervicomis , Asterosmilia gigas.
Stn 516. — 12.05.1992, 14°13.5'S, 178°11.6'W. 441-550 m, Futuna: Polycyathus octuplus, Bourneotrochus
stellulatus , Deltocyathus Stella , Heterocyathus cf. sulcatus , Conotrochus asymmetros , Flabellum (Flabellum)
pavoninuni , Truncatoflabellum phoenix , Endopachys grayi , Heteropsammia cochlea.
Stn 517. — 12.05.1992, 14°13.4'S, 178°10.4’W, 233-235 m, Futuna: Madrepora minutiseptum.
Stn 520. — 13.05.1992, 1 4° 1 0.6'S, 176°16.7'W. 920-930 m, Wallis: Deltocyathus cameratus, Javania
lamprotichum (or station 52 1 ), Enallopsammia rostrata.
Stn 521. — 13.05.1992, 1 4° 1 1.0'S, 176°17.3'W, 890-915 m, Wallis: Javania lamprotichum (or station 520).
Stn 522. — 13.05.1992, 1 3° 1 0.7'S, 176°15.0'W, 650-765 in, Wallis: Fungiacyathus (Fungiacyathus) pusillus
pacific us, Anthemiphyllia dentata , A. patera costata , Caryophyllia (Caryophyllia) abrupta, Trochocyathus
(Trochocyathus) vasiformis , Tethocyathus virgatus, Deltocyathus suluensis, D. taiwanicus, Conotrochus
funicolumna, Enallopsammia rostrata.
Stn 523. — 13.05.1992, 13°12.0'S, 176°15.6'W, 455-515 m, Wallis: Fungiacyathus (Fungiacyathus) sandoi,
S t ephan ophyll i a complicata, Caryophyllia ( Caryophyllia ) abrupta. Crispatotrochus rubescens , C. rugosus ,
Oxysmilia epithecata, Trochocyathus ( Aplocyathus ) hastatus , Deltocyathus taiwanicus, D. crassiseptum, D.
Stella, Conotrochus funicolumna, Asterosmilia gigas , Deltocyathoides orientalis, Flabellum ( Flabellum )
pavoninuni. Truncatoflabellum mortenseni , Javania exserta.
Stn 524. — 13.05.1992, 13°I1.8'S, I76°15.6'W, 300 m, Wallis: Caryophyllia (Caryophyllia) abrupta,
Trochocyathus (Trochocyathus) maculatus , Rhizosmilia robusta , Flabellum (Flabellum) arcuatile .
Truncatoflabellum Vanuatu, Balanophyllia desmophyllioides.
Stn 525. — 13.05.1992, 1 3° 1 0.6'S, 176°14.7’W, 500-600 m, Wallis: Anthemiphyllia dentata, Caryophyllia
(Caryophyllia) marmorea, Deltocyathus taiwanicus , Conotrochus funicolumna, C. brunneus , Javania fusca.
Stn 527. — 14.05.1992, 13°24.1'S, I76°I4.6’W, 540-560 m, Wallis: Trochocyathus (Trochocyathus) vasiformis.
Stn 529. — 16.05.1992, I2°31.4'S, 176°39.6’W, 500 m, Waterwitch Bank: Deltocyathus suluensis,
D. taiwanicus, D. crassiseptum.
Stn 530. 16.05.1992, 12°32.7’S, 176°39.3'W, 580-600 m, Waterwitch Bank: Anthemiphyllia dentata, A. patera
costata. A. spinifera , Caryophyllia (Caryophyllia) rugosa , C. (C.) abrupta, Deltocyathus suluensis,
D. cameratus , Conotrochus brunneus, Lophelia pertusa, Flabellum (Flabellum) arcuatile , Javania fusca,
Enallopsammia rostrata.
Stn 532. — 16.05.1992, 12°28.9’S, 176°41.0'W. 516-530 m, Waterwitch Bank: Deltocyathus suluensis,
Conotrochus funicolumna, Flabellum ( Jlocyathus) deludens.
Stn 534. 16.05.1992, 12°23.3'S, 176°42.0'W, 440-500 m, Waterwitch Bank: Fungiacyathus (Fungiacyathus)
pusillus pacificus, Fungiacyathus (Bathyactis) margaretae , Anthemiphyllia dentata . Caryophyllia (Caryophyllia)
abrupta, C. (C.) marmorea^ Deltocyathus suluensis, Flabellum (Flabellum) arcuatile.
Stn 535. 16.05.1992. 12°29.6'S, 176°41.3'W, 340-470 m, Waterwitch Bank: Fungiacyathus (Fungiacyathus)
pusillus pacificus, Fungiacyathus (Bathyactis) margaretae, Stephanophyllia complicata, Anthemiphyllia
dentata. A. patera costata, Caryophyllia (Caryophyllia) abrupta, Trochocyathus (Trochocyathus) vasiformis,
Deltocyathus suluensis, D. taiwanicus, Conotrochus funicolumna, Asterosmilia gigas , Flabellum (Flabellum)
arcuatile.
Stn 537. — 16.05.1992, 12°30.0'S, 176°4I.0’W, 325-400 m, Waterwitch Bank: Anthemiphyllia dentata,
A. spinifera, Caryophyllia (Caryophyllia) abrupta, Deltocyathus taiwanicus, D. crassiseptum. D. cameratus,
D. Stella, Conotrochus funicolumna, Asterosmilia gigas, Javania exserta.
Stn 538. 16.05.1992. 1 2 30.8’S, 176°40.3'W, 275-295 m, Waterwitch Bank: Fungiacyathus (Fungiacyathus)
sandoi, Anthemiphyllia dentata, Caryophyllia (Caryophyllia) hawaiiensis. Flabellum (Flabellum) pavoninuni,
Truncatoflabellum phoenix, Javania exserta, Balanophyllia desmophyllioides.
Sm $39. 17.05.1992, 12°27.3'S, 177°27.3'W, 700 m. Combe Bank: Trochocyathus (Trochocyathus) discus,
Lochmaeotrochus gardineri.
Source : MNHN. Paris
AZOOXANTHELLATE SCLERACTINIA
37
Sln 540. — 17.05.1992, 12°26.7'S, 177°28.4'W, 600 m, Combe Bank: Fungiacyathus (Fungiacyathus) pusillus
pcicificiis , F. (F.) sandoi , Fungiacyathus ( Bathyactis ) margaretae , Stephanophyllia complicata , Anthemiphyllia
dentata , 4. patera costata , Caryophyllia ( Caryophyllia ) crosnieri, Deltocyathus suluensis , /). taiwanicus ,
Conotrochus funicolumna, Javania fiisca.
Sln 54i. _ 17.05.1992, 12°26.7'S, 177°28.0'W, 500-505 m. Combe Bank: Fungiacyathus (Fungiacyathus)
sandoi , Fungiacyathus ( Bathyactis ) margaretae , Anthemiphyllia spin if era, Deltocyathus suluensis ,
77. taiwanicus , D. cameratus , /7. Stella, Conotrochus funicolumna.
Sm 542. _ 17.05.1992, 12°26.4'S, 177°28.2'W, 370 m, Combe Bank: Fungiacyathus (Fungiacyathus) sandoi.
Stephanophyllia neglecta, Anthemiphyllia dentata, A. patera costata. A. spinifera , Crispatotrochus rugosus ,
Deltocyathus taiwanicus , D. cameratus, Conotrochus funicolumna, Flabellum (Flabellum) arcuatile.
Sm 546. _ 17.05.1992, 12°26.9'S, 177029.1'W, 550-552 m, Combe Bank: Fungiacyathus (Bathyactis)
margaretae, Anthemiphyllia dentata, A. patera costata. Caryophyllia (Caryophyllia) abrupta, Trochocyathus
(Trochocyathus) maculatus, Deltocyathus suluensis, D. taiwanicus, D. cameratus, Conotrochus brunneus,
Flabellum (Flabellum) arcuatile.
Stn 548. — 17.05.1992, 12°23.3’S, 177°24.4’W, 700-740 m, Combe Bank: Lochmaeotrochus gardineri.
Sm 551. — 18.05.1992, 1 2° 1 5.3’S, 177°28.rW, 791-795 m, Combe Bank: Fungiacyathus (Fungiacyathus)
stephanus, Madrepora oculata fonna tenuis, Flabellum (Ulocyathus) apertum apertum.
Sm 552. — 18.05.1992, 1 2° 1 5.7'S, 177°27.8’W, 786-800 m. Combe Bank: Fungiacyathus (Fungiacyathus)
stephanus, Madrepora oculata forma tenuis, Caryophyllia (Caryophyllia) scobinosa, Deltocyathus cameratus,
Flabellum (Ulocyathus) apertum apertum.
Sm 555. — 19.05.1992, 11°47.5'S, 178°19.2’W, 540-542 m, Tuscarora Bank: Anthemiphyllia dentata,
Trochocyathus ( Aplocyathus ) hastatus, Deltocyathus taiwanicus, D. cameratus, Conotrochus funicolumna
Sm 556. — 19.05.1992, 1 1°48.7'S, 178°18.0'W, 440 m, Tuscarora Bank: Stephanophyllia neglecta. Caryophyllia
(Caryophyllia) quadragenaria, Oxysmilia epithecata. Bourneotrochus steUulatus. Deltocyathus taiwanicus,
D. Stella , Heterocyathus cf. sulcatus , Conotrochus funicolumna.
Sm 557. — 19.05.1992, ll°48.rS, 178°18.2’W. 600-608 m. Tuscarora Bank: Stephanophyllia complicata,
Caryophyllia (Caryophyllia) abrupta, Deltocyathus suluensis, D. taiwanicus, D. cameratus, Conotrochus
funicolumna, Lochmaeotrochus gardineri.
Sm 560. — 19.05.1992, 1 1°47.0'S, 178°20.0'W, 697-702 m, Tuscarora Bank: Deltocyathus taiwanicus,
D. cameratus, Conotrochus funicolumna, Lochmaeotrochus gardineri.
Sm 564. — 20.05.1992, ll°46.rS, 178027.4’W, 1015-1020 m, Tuscarora Bank: Fungiacyathus (Bathyactis)
margaretae, Caryophyllia (Caryophyllia) ambrosia, Stephanocyathus (Stephanocyathus) regius. Aulocyathus
recidivus, Flabellum (Ulocyathus) apertum apertum.
Sm 565. — 20.05.1992, 11047.4'S, 178025.3'W. 900 m, Tuscarora Bank: Fungiacyathus (Fungiacyathus)
stephanus, Fungiacyathus (Bathyactis) margaretae, Caryophyllia (Caryophyllia) scobinosa. C. (C.) ambrosia,
Stephanocyathus (Stephanocyathus) regius, Flabellum (Ulocyathus) apertum apertum.
Sm 567. — 20.05.1992, 11°47.0'S, 178°27.3’W, 1010-1020 m, Tuscarora Bank: Stephanocyathus
(Stephanocyathus) regius, Deltocyathus cameratus, Aulocyathus recidivus, Habellum (Ulocyathus) apertum
apertum.
Sin 569. — 21.05.1992, 12°30.0’S, 176°51.2’W, 300-305 m, Waterwilch Bank: Fungiacyathus (Fungiacyathus)
sandoi , Anthemiphyllia dentata, A. spinifera, Caryophyllia (Caryophyllia) abrupta, Bourneotrochus steUulatus,
Deltocyathus Stella, D. ornatus, D. cameratus, Guynia annulata, Flabellum (Flabellum) pavoninum. Javania
exserta.
Sm 570. — 21.05.1992, 12°30.9'S. I76°51.4’W. 420-439 m, Waterwilch Bank: Fungiacyathus (Fungiacyathus)
pusillus pcicificiis, Anthemiphyllia dentata, Caryophyllia (Caryophyllia) abrupta, Deltocyathus crassiseptum ,
Heterocyathus cf. sulcatus, Conotrochus funicolumna.
Sln 571. — 21.05.1992, 12°31.3’S, 176°51.7’W, 502-508 m. Waterwitch Bank: Caryophyllia (Caryophyllia)
abrupta, Conotrochus funicolumna.
38
S. D. CAIRNS
Stn 572. 21.05.1992, 12 31.8'S, I76°52.2’W, 500-560 m, Waterwitch Bank: Fungiacyathus (Fungiacyathus)
pusillus pacificus, Anthemiphyllia dentata , Caryophyllia ( Caryophyllia ) crosnieri , Vaughanella concinna ,
Lophelia pertusa.
Stn 574. - 21.05.1992, 12°30.9'S, 176°52.3’W, 105-410 m, Waterwitch Bank: Enallopsammia rostrata
Stn 575. - 21.05.1992. I2°30.9'S, I76°52.3'W, 425 m. Waterwitch Bank: Fungiacyathus (Fungiacyathus)
pusillus pacificus, Anthemiphyllia patera costata, Deltocyathus suluensis.
Stn 578. — 22.05.1992, 13°08.2'S. I76°15.6'W. 640-730 m. North of Wallis: Anthemiphyllia patera costata,
Deltocyathus suluensis, D. cameratus, Flabellum ( Flabellum ) arcuatile , Javania fusca
Stn 581. - 22.05.1992. 13°09.9'S, I76°13.9'W, 461-550 m. North of Wallis: Fungiacyathus (Fungiacyathus)
pusillus pacificus, Canophyllia ( Caryophyllia ) crosnieri.
Stn 584. - 22.05.1992. 13°11.2'S. 176°14.3'W, 360-400 m, North of Wallis: Crispatotrochus rugosus,
Dactylotrochus cervicomis.
Stn 585. — 22.05.1992, 13 10.2'S. 1 76° 12.6'W. 415-475 m. North of Wallis: Madrepora oculata forma formosa,
Anthemiphyllia dentata, Caryophyllia (Caryophyllia) marmorea, Crispatotrochus ruhescens. Bourneotrochus
stellulatus, Stephanocyathus ( Acinocyatlius ) spiniger, Deltocyathus taiwanicus, D. crassiseptum, D. Stella.
Conotrochus funicolumna, Truncatoflabellum mortenseni.
Stn 586. — 22.05.1992, 1 3° 10.7‘S. 1 76° 1 3. 1’W. 510-600 m. North of Wallis: Madrepora oculata forma formosa
Anthemiphyllia dentata, A. patera costata, A. spinifera, Caryophyllia (Canophyllia) abrupta, Trochocyathus
( Aplocyathus ) hastatus, Bourneotrochus stellulatus, Deltocyathus crassiseptum, Conotrochus brunneus
C. asymmetros , Asterosmilia gigas.
Stn 589. — 23.05.1992, 12°16.2'S, 174°41.4'W, 400 m, Field Bank: Fungiacyathus (Fungiacyathus) sandoi,
Anthemiphyllia dentata, A. spinifera, Caryophyllia (Caryophyllia) lameUifera, Crispatotrochus rugosus,
Deltocyathus taiwanicus, D. cameratus, Dactylotrochus cervicomis. Flabellum (Flabellum) arcuatile
Balanophyllia desmophyllioides.
Stn 590. _ 23 05.1992, 12°3I.4'S, I74°18.7'W. 400 m, Field Bank: Fungiacyathus (Fungiacyathus) sandoi,
Anthemiphyllia dentata. 4. patera costata, Deltocyathus suluensis, D. taiwanicus, Conotrochus brunneus.
Stn 59 1 . — 23.05.1992, 12°31.1'S, 174°19.4'W, 320 m, Field Bank: Anthemiphyllia dentata, A. patera costata
Bourneotrochus stellulatus, Deltocyathus taiwanicus, Conotrochus brunneus
Stn 592. - 24.05.1992, I2°32.4'S, I74°22.0'W. 730-775 m. Field Bank: Javania fusca
Stn 594 - 24.05.1992 12«3 1.0'S, 174°19.9'W. 495-505 m. Field Bank: Anthemiphyllia dentata. A. patera
costata , Bourneoti ochus stellulatus , Deltocyathus suluensis , D. taiwanicus D Stella
S,n 595. 24.05.1992, 12°30.9'S, 174-18.9'W, 566-580 m. Field Bank: Antliemipbyllia patera costata,
Bourneotrochus stellulatus, Deltocyathus suluensis
S,„ 597. - 24.05.1992 12"31 4'S, 174‘18.6'W, 469-475 m, Field Bank: Fungiacyatbus (Fungiacyatbus) sandoi.
Lc tepsamnua franki. Anthem, phyll, a dentata , Deltocyathus taiwanicus , D. crassiseptum , D. cameratus
Conotrochus brunneus.
Tf5!- '74°18'4W' 7°2'708 Fidd B”k: Madr“>°r° *»»
Cary ophyllia (Cary ophyllia) ambrosia.
Safh Tiff"2' U''STS- 176°17-2'W. 35» SE of Wallis: Madrepora oculata forma formosa
l runcan, flabellum mortenseni, Balanophyllia desmophyllioides
Stn 604. -26.05.1992, 13°21.4'S, 176°08.3'W, 415-420 m, SE of Wallis: Caryophyllia (Caryophyllia) abrupta
Crispatotrochus rubescens, Bourneotrochus stellulatus, Stephanocyathus (Acinocyatlius) spiniger. Deltocyathus
crassiseptum, Conotrochus funicolumna.
S,n - 2“5:1f | 13°2!'3 S' 176°08.4'W, 335-340 m, SE of Wallis: Fungiacyathus (Fungiacyatbus)
s,eZa,us ZPd Zmfem',?aTkyU:a ^opWial abrupta. Osysmilia epithecata, Bourneotrochus
' D“'7'‘"roC*“ Leontis, FlabeUum (Fiabellum) paramount.
176"083'W- 420-«» -V SE Wallis: Madrepora oculata forma formosa.
Caryophyllia ( Caryophyllia ) abrupta , Stephanocyathus ( Acinocyatlius ) spiniger.
Source : MNHN. Paris
AZOOXANTHELLATE SCLERACTINIA
39
Stn 608. — 26.05.1992, 13°21.7'S, 176°08.5'W, 440-458 m, SE of Wallis: Caryophyllia (Caryophyllia) abrupta ,
Bourneotrochus stellulatus , Deltocyathus crassiseptiim.
Stn 609. — 26.05.1992. 13°21.5’S, 176°08.5'W, 430 m, SE of Wallis: Madrepora oculata forma formosa,
Trochocyathus ( Trochocyathus ) vasiformis.
Stn 610. — 26.05.1992, 13°21.5'S, 176°08.9'W, 286 m, SE of Wallis: Anthemiphyllia spinifera, Caryophyllia
( Caryophyllia ) rugosa , Crispatotrochus rugosus , Oxysmilia epithecata , Bourneotrochus stellulatus ,
Deltocyathus Stella , Conotrochus asymmetros , Flabellum ( Flabellum ) pavoninum .
Sln 6i8. _ 27.05.1992, 14°21.7’S, 178°00.5'W, 420-435 m, E + SE of Alofi: Stephanophyllia neglecta,
Madrepora oculata forma formosa, M. porcellana , Anthemiphyllia dentata , Trochocyathus ( Aplocyathus )
hastatus, Bourneotrochus stellulatus , Deltocyathus crassiseptiim, Conotrochus funicolumna.
Sm 619. — 27.05.1992. 14°21.8'S, 178°00.4'W, 455 m, E + SE of Alofi: Fungiacyathus ( Bathyactis ) granulosus ,
Anthemiphyllia dentata , Caryophyllia ( Caryophyllia ) abrupta. Trochocyathus ( Aplocyathus ) hastatus ,
Conotrochus funicolumna.
Sm 620. — 28.05. 1992, 12°34.4’S, 178°1 1 .0’W, 1280 m, bank SW of Combe Bank: Madrepora oculata forma
formosa , Aulocyathus recidivus , Flabellum ( Ulocyathus ) apertum apertum.
Stn 621. — 28.05.1992, 12°35.0'S, 178°11.5'W, 1280-1300 m. bank SW of Combe Bank: Fungiacyathus
(Fungiacyathus) stephanus , Madrepora oculata forma tenuis, Stephanocyathus ( Stephanocyathus ) regius ,
Stephanocyathus (Odontocyathus) coronatus, Deltocyathus rotulus , Flabellum (Ulocyathus) apertum apertum.
Stn 622. — 28.05.1992, 12°34.5'S. 178°10.9'W. 1280-1300 m, bank SW of Combe Bank: Stephanocyathus
(Odontocyathus) coronatus , Deltocyathus rotulus, Aulocyathus recidivus.
Stn 623. — 28.05.1992, 12°34.2'S, 178015.I'W, 1280-1300 m, bank SW of Combe Bank: Madrepora oculata
forma tenuis, Stephanocyathus (Stephanocyathus) regius, Stephanocyathus (Odontocyathus) coronatus,
Deltocyathus rotulus, Aulocyathus recidivus, Flabellum (Ulocyathus) apertum apertum.
Stn 625. — 29.05.1992, 1 1°52.4'S. 179°33.8’W, 425-430 m, Bayonnaise Bank: Conotrochus brunneus.
Stn 626. — 29.05.1992, 1 l°53.6’S, 179°32.0’W, 597-600 m, Bayonnaise Bank: Deltocyathus Stella.
Stn 631. — 29.05.1992, 1 1°54.0'S, 179°31.6'W. 600 m, Bayonnaise Bank: Deltocyathus suluensis.
Sm 635. — 30.05.1992, 13°49.0’S, 179°56.0’E, 700-715 m, SW of Rotumah Bank: Anthemiphyllia patera
costata, Caryophyllia (Caryophyllia) scobinosa, C. (C.) ambrosia, Stephanocyathus (Stephanocyathus) regius,
Deltocyathus cameratus.
Stn 636. — 30.05.1992, 13°39.4’S, 179°55.5'E, 650-700 m, SW of Rotumah Bank: Caryophyllia (Caryophyllia)
ambrosia, Stephanocyathus (Stephanocyathus) regius, Deltocyathus cameratus.
Sm 637. — 30.05.1992, 13°37.2’S, 179°56.0’E, 820-830 m, SW of Rotumah Bank: Madrepora oculata forma
tenuis, Stephanocyathus (Stephanocyathus) regius, Desmophyllum dianthus, Pleotrochus zibrowii.
Musorstom 8, "Alis"
Stn 956. — 20.09.1994, 20°33'S, 169°35'E, 1175-1210 m, Anatom: Fungiacyathus ( Bathyactis ) margaretae,
Stephanocyathus (Odontocyathus) coronatus, Deltocyathus cameratus, Lochmaeotrochus gardineri , Flabellum
(Ulocyathus) marcus.
Stn 958. — 20.09.1994, 20°20'S, 169°47'E, 497-570 m, Anatom: Stephanophyllia neglecta, Trochocyathus
(Trochocyathus) discus, Deltocyathus crassiseptiim, Cyathotrochus pileus.
Stn 959. — 20.09.1994, 20°20'S, 169°48’E, 436-475 m, Anatom: Stephanophyllia complicata, Anthemiphyllia
dentata, Caryophyllia (Caryophyllia) diomedeae , Oxysmilia epithecata, Trochocyathus (Trochocyathus)
vasiformis, Trochocyathus (Aplocyathus) hastatus, T. (A.) brevispina, Deltocyathus crassiseptiim, Conotrochus
brunneus, Cyathotrochus pileus, Flabellum (Ulocyathus) aotearoa, Javania fitsca, Enallopsammia rostrata.
Stn 961. — 21.09.1994, 20°18'S, 169°49'E, 100-110 m, Anatom: Trochocyathus (T.) maculatus, T. (T.) cooperi,
Rhizosmilia robusta, Heteropsammia cochlea.
Stn 962. — 21.09.1994, 20°19’S, I69°49'E, 370-400 m. Anatom: Letepsammia franki, Madrepora porcellana ,
Caryophyllia (Caryophyllia) quadragenaria, Oxysmilia epithecata, Flabellum (Flabellum) pavoninum, Javania
exserta, Balanophyllia laysanensis.
40
S. D. CAIRNS
Stn 963. — 21.09.1994, 20°20’S, 169°49'E, 400-440 m. Anatom: Fungiacyathus (Fungiacyathus) stephanus,
F. (F.) paliferus , Fungiacyathus ( Bathyactis ) margaretae , Stephanophyllia neglecta , S. complicata [X).
Madrepora porcellana, Caryophyllia ( Caryophyllia) abrupta [X], Trochocyathus (Aplocyathus) hastatus , T. (4.)
brevispina, Stephanocyathus ( Acinocyathus ) spiniger , Deltocyathus magnificus , Bourneotrochus stellulatus
[X], Tropidocyathus labidus [X], Temnotrochus kennadecensis [X], Flabellum ( Flabellum ) pavoninum ,
Flabellum ( Ulocyathus ) aotearoa, Truncatoflabellum mortenseni, Balanophyllia laysanensis.
Sin 964. — 21.09.1994, 20°19'S, 169°49’E, 360-408 m, Anatom: Madrepora porcellana, Caryophyllia
( Caryophyllia ) octonaria , Trochocyathus ( Aplocyathus ) brevispina , Deltocyathus magnificus , D. ornatus.
Rhizosmilia robusta, Flabellum ( Ulocyathus ) aotearoa, Truncatoflabellum mortenseni, Javania exserta ,
Balanophyllia desmophyllioides.
Sin 965. — 21.09.1994, 20°20'S, 169°5rE, 361-377 m, Anatom: Stephanophyllia neglecta, Caryophyllia
( Caryophyllia ) quadragenaria , Flabellum {Flabellum) pavoninum, Truncatoflabellum dens, Javania fusca,
Balanophyllia laysanensis.
Stn 967. — 21.09.1994, 20°19'S, I69°53'E, 295-334 m. Anatom: Fungiacyathus (Fungiacyathus) paliferus,
Anthem iphy Ilia dentata , Caryophyllia (Caryophyllia) rugosa , C. (C.) quadragenaria, Deltocyathus Stella ,
D. ornatus , Truncatoguynia irregularis, Truncatoflabellum mortenseni, Endopachys grayi , Heteropsammia
cochlea.
Stn 968. — 2 1 .09. 1 994, 20° 1 8'S. 1 69°53'E, 1 99-2 1 4 m, Anatom: Madrepora porcellana.
Stn 969. — 21.09.1994, 20°19'S, 169°53'E, 252-280 m, Anatom: Fungiacyathus (Fungiacyathus) paliferus,
Stephanophyllia neglecta, Madrepora porcellana, Anthemiphyllia dentata. Caryophyllia ( Caryophyllia )
hawaiiensis , C. (C.) octonaria, Trochocyathus (Trochocyathus) philippinensis , Bourneotrochus stellulatus,
Deltocyathus Stella. Heterocyathus cf. sulcatus , Truncatoflabellum mortenseni. Balanophyllia
desmophyllioides . Endopachys grayi , Heteropsammia cochlea.
Stn 970. — 21.09.1994. 20° 1 8’S, 169°53'E, 252-310 m. Anatom: Truncatoflabellum mortenseni. Balanophyllia
red i viva.
Stn 971. — 21.09.1994, 20C!9'S, 169°53'E. 250-315 m, Anatom: Rhizosmilia robusta, Flabellum (Flabellum)
pavoninum, Truncatoflabellum mortenseni, Dendrophyllia alcocki.
Stn 973. 22.09.1994, 1 9°2 1 'S, 169°27'E, 460-480 m, Tanna: Caryophyllia (Caryophyllia) crosnieri,
Trochocyathus (Trochocyathus) vasiformis, Tethocyathus virgatus, Truncatoflabellum pusillum.
Stn 974. 22.09.1994, 1 9°2 1 S, 169°28'E. 492-520 m, Tanna: Catyophyllia (Caryophyllia) crosnieri. Javania
lamprotichum, Enallopsammia rostrcita.
Stn 975. — 22.09.1994, 19°23’S, 169°29'E, 536-566 m. Tanna: Fungiacyathus (Fungiacyathus) stephanus.
Caryophyllia (Catyophyllia) crosnieri. Deltocyathus suluensis, Javania lamprotichum, Polymyces wellsi.
Stn 976. 22.09.94, 1 9°25'S, 169°27'E, 160-182 m, Tanna: Madrepora porcellana, Caryophyllia ( Caryophyllia )
octonaria, Trochocyathus (Trochocyathus) philippinensis, Heterocyathus cf. sulcatus, Aulocyathus juvenescens,
Flabellum (Flabellum) pavoninum, Endopachys grayi, Heteropsammia cochlea.
Sm 977. 22.09.1994, I9°25'S, I69°29'E. 410-505 m, Tanna: Fungiacyathus (Bathyactis) granulosus,
Anthemiphyllia dentata, Crispatotrochus rugosus, Oxysmilia circularis, Trochocyathus (Trochocyathus)
vasiformis, T. (T .) rhombocolumna, Trochocyathus (Aplocyathus) hastatus. Tethocyathus virgatus,
Deltocyathus crassiseptum, Conotrochus brunneus, Flabellum (Ulocyathus) aotearoa, Truncatoflabellum dens,
Balanophyllia gigas, Dendrophyllia alcocki, Enallopsammia rostra ta.
Stn 978. 22.09.1994, 19°23'S, 169°27'E, 408-413 m, Tanna: Stephanophyllia neglecta. Trochocyathus
(Trochocyathus) vasiformis, Tethocyathus virgatus, Deltocyathus crassiseptum. Javania exserta. Balanophyllia
crassitheca, Dendrophyllia alcocki.
Stn 980. 22.09.1994, 19°21'S, 169°25'E, 433-450 m. Tanna: Fungiacyathus (Bathyactis) granulosus,
Letepsammici franki, Deltocyathus magnificus, D. crassiseptum, Cyathotrochus pileus, Dendrophyllia alcocki
Stn 982. 23.09.1994, I9°22'S, 169 26'E, 408-410 m, Tanna: Anthemiphyllia dentata. Caryophyllia
(Caryophyllia) crosnieri, Tethocyathus virgatus, Javania fusca, Dendrophyllia alcocki.
Source : MNHN , Paris
AZOOXANTHELLATE SCLERACTINIA
41
Sln 983 _ 23.09.1994. I9°22'S, 169°28'E, 475-480 m, Tanna: Anthemiphyllia dentala, Crispatotrochus
rubescens, Oxysmilia circularis, Trochocyathus (Trochocyathus) vasiformis, Deltocyathus crassiseptum ,
Dendrophyllia alcocki..
Stn 988. _ 23.09.1994, 19°16’S. 169°24'E, 372-466 m. Tanna: Madracis kauaiensis, Fungiacyathus
( Fungiacycithus ) paliferus , Stephanophyllia neglecta, Madrepora porcellana, Anthemiphyllia spinifera.
Caryophyllia ( Cctryophyllia ) rugosa, C. (C.) abrupta, Crispatotrochus rugosus , Labyrinthocyathus limatulus ,
Dactylotrochus cervicornis, Cryptotrochus brevipalus , Flabellum ( Flabellum ) pavoninum, Truncatoflabellum
dens. T. pusillum , Javania lamprotichum , J.fusca , Balanophyllia desmophyllioides, Dendrophyllia alcocki.
S[n 990. — 24.09.1994, I8°52'S. I68°5I'E, 980-990 m, Erromango: Fungiacyathus ( Fungiacyathus ) stephanus.
Stephanocyathus ( Stephanocyathus ) regius.
Sln 992. _ 24.09.1994, 18°52'S, 168°55'E, 748-775 m, Erromango: Fungiacyathus ( Fungiacyathus ) stephanus.
Caryophyllia ( Caryophyllia ) scobinosa. Stephanocyathus ( Stephanocyathus ) regius. Deltocyathus cameratus,
Lochmaeotrochus gardineri.
S(n 996. - 24.09.1994, I8°52'S, I68°56'E, 764-786 m, Erromango: Fungiacyathus (Fungiacyathus) stephanus.
Stephanocyathus ( Stephanocyathus ) regius, Truncatoflabellum stabile.
Stn 999. — 25.09.1994. 18°49'S, 169°00'E, 80-110 m, Erromango: Trochocyathus ( Aplocyathus ) brevispina.
Sm 1001. — 25.09.1994, 18°49’S, I68°59'E, 150-250 m, Erromango: Caryophyllia (Acanthocyathus) grayi.
Stn 1002. — 25.09.1994, 18°49’S, 168°59'E, 200-300 m, Erromango: Caryophyllia (Acanthocyathus) grayi.
Sm 1003. — 25.09.1994, 18°49'S, 168°59'E, 200-327 m, Erromango: Fungiacyathus (Fungiacyathus) paliferus.
Trochocyathus (Aplocyathus) brevispina. Stephanocyathus ( Acinocyathus ) spiniger.
Sm 1004. — 25.09.1994, 18°49’S, 168°59'E, 319-350 m, Erromango: Caryophyllia (Acanthocyathus) grayi.
Trochocyathus (Aplocyathus) brevispina. Stephanocyathus ( Acinocyathus ) spiniger. Heterocyathus alternants.
Flabellum (Flabellum) pavoninum. Flabellum (Ulocyathus) aotearoa. Truncatoflabellum vigintifarium.
Stn 1005. — 25.09.1994, I8°49'S. 168°59'E, 360-450 m, Erromango: Fungiacyathus (Fungiacyathus) paliferus.
Stephanophyllia complicata, Flabellum (Flabellum) pavoninum. Flabellum (Ulocyathus) aotearoa. Endopachys
grayi.
Sm 1006. — 25.09.1994, 18°50'S, 168°57'E, 574-61 1 m, Erromango: Trochocyathus (Trochocyathus) vasiformis.
Deltocyathus vaughani, Conotrochus brunneus, Gardineria hawaiiensis . Enallopsammia rostrata.
Sin 1007. — 25.09.1994. 18°52'S. 168°56'E, 720-830 m, Erromango: Fungiacyathus (Fungiacyathus) stephanus.
Stephanocyathus (Stephanocyathus) regiu's. Deltocyathus cameratus.
Sm 1009. — 27.09.1994. 17°46'S. 168°13'E, 430-460 m, Efate: Caryophyllia (Caryophyllia) crosnieri.
Balanophyllia gemma.
Stn 1011. — 27.09.1994, I7°50’S, 168°U'E, 547-585 m. Elate: Fungiacyathus (Batliyactis) margaretae.
Trochocyathus ( Trochocyathus ) vasiformis, Deltocyathus vaughani. Gardineria hawaiiensis.
Sm 1014. — 27.09.1994, 17°54'S, I68°19'E, 495-498 m. Elate: Fungiacyathus (Batliyactis) margaretae.
Anthemiphyllia spinifera, Caryophyllia ( Caryophyllia ) diomedeae, Cryptotrochus brevipalus, Flabellum
(Ulocyathus) hoffmeisteri, Javania lamprotichum, Gardineria hawaiiensis, G. paradoxa.
Sm 1015. — 27.09.1994, I7°54'S, 168°22'E, 375-420 m, Efate: Crispatotrochus rugosus. Trochocyathus
(Trochocyathus) vasiformis, Tetliocyathus virgatus, Truncatoflabellum dens. Balanophyllia gemma.
Dendrophyllia alcocki, Enallopsammia rostrata.
Stn 1016. — 27.09.1994, I7°53’S, I68°28’E. 291-300 m, Efate: Fungiacyathus (Fungiacyathus) paliferus,
Letepsammia franki, Anthemiphyllia dentata, Caryophyllia (Caryophyllia) abrupta, Conotrochus asymmetros
[XI, Tropidocxathus lessonii, Cyathotrochus pileus, Guynia annulata. Flabellum (Flabellum) pavoninum,
Flabellum (Ulocyathus) aotearoa. Truncatoflabellum angustum. Truncatoflabellum pusillum [X|, Endopachys
grayi..
Stn 1017. — 27.09.1994, 17°53'S, 168°26'E. 294-295 m. Efate: Fungiacyathus (Fungiacyathus) paliferus.
Letepsammia franki. Deltocyathus ornatus, Conocyathus asymmetros [X], Flabellum (Flabellum) pavoninum.
Flabellum ( Ulocyathus ) aotearoa, Truncatoflabellum angustum, Endopachys grayi [X],
42
S. D. CAIRNS
Stn 1018. 27.09.1994, 17°53'S, 168°25'E, 300-301 m, Efate: Fungiacyathus (Fungiacyathus) paliferus.
Letepsammia franki , Stephanophyllia complicata.? Caryophyllia (C.) lamellifera , Crisp at otrochus rugosus .
Oxysmilia epithecatci , Deltocycithus ornatus . Bourneotrochus stellulatus (X), Heterocyathus alternatus.
Notocyathus conicus [X], Flabellum ( Flabellum ) pavoninum . Flobellum ( Ulocycithus ) aoteciroci,
Truncatoflabellum angustum. T. vigintifarium. Endopachys grayi . Dendrophyllia arbuscula.
Stn 1019. 28.09.1994, 17°38'S, 168°34'E, 397-430 m, Efate: Stephanophyllia neglecta. Trochocyathus
( Trochocyathus ) vasiformis . 7. (7.) efateensis , Trochocyathus (Aplocyathus) hastatus , BalanophylUa gemma .
Dendrophyllia alcocki.
Stn 1020. 28.09.1994, 1 7 40'S, 168°35'E, 380-391 m, Efate: Trochocyathus (Trochocyathus) efateensis.
Trochocyathus (Aplocyathus) hastatus , 7. (4.) brevispina .
Stn 1021. 28.09.1994, 17°43'S, 168°37'E, 124-130 m, Efate: Trochocyathus (Trochocyathus) maculatus.
7. (7.) cooperi . Rhizosmilia robusta . Javania exserta. Rhizotrochus typus . Dendrophyllia arbuscula.
Stn 1023. — 28.09.1994, 17°48'S, 168°49'E, 321 m, Efate: Letepsammia franki. ?Caryophyllia (C.) lamellifera.
Oxysmilia epithecata . Bourneotrochus stellulatus [X], Temnotrochus kermadecensis [X], Flabellum
(Ulocyathus) aotearoa, Dendrophyllia alcocki.
Stn 1024. 28.09.1994, 17°48'S, 168°39'E, 335-370 m, Efate: Enallopsammia rostrata.
Stn 1026. 28.09.1994, 1 7 50 S , 168°39'E, 437-504 m, Efate: Oxysmilia epithecata. Trochocyathus
(Trochocyathus) efateensis. Tethocyathus virgatus. Tropidocyathus labidus.
Stn 1028. 28.09.1994, 1 7 54 S, 168°40'E, 624-668 m, Efate: Madrepora oculata forma tenuis. Deltocyathus
suluensis , Cryptotrochus brevipalus.
Stn 1030. 29.09.1994, 1 7°5 1 'S, 168°30'E, 180-190 m, Efate: Crispatotrochus rugosus. Oxysmilia circular is.
O. corrugata. Javania exserta. BalanophylUa desmophyllioides. Dendrophyllia arbuscula.
Stn 1031. 29.09.1994, 17 '52'S, 168°33'E, 310 m, Efate: Madrepora oculata forma tenuis. Anthemiphyllia
pacifica.
Stn 1034. - 29.09.1994, 17°54'S, 168°42'E, 690-750 m, Efate: Caryophyllia (Caryophyllia) scobmosa,
Lochmaeotrochus gardineri. Cryptotrochus brevipalus.
Stn 1035. 29.09.1994, 17°56'S, 168°44'E, 765-780 m, Elate: Caryophyllia (Caryophyllia) scobinosa.
Stn 1036. — 29.09.1994, 18°01'S, 168°48'E, 920-950 m, Efate: Fungiacyathus (Fungiacyathus) stephcinus.
Stephanocyathus (Stephanocyathus) regius. Deltocyathus cameratus. Lochmaeotrochus gardineri.
Truncatoflabellum stabile.
Stn 1037. - 29.09.1994, 18°03’S, I68°54’E, 1058-1086 m, Efate: Flabellum (Ulocyathus) marcus.
T runcatoflabcllum stabile.
Stn 1042. - 30.09.1994, 16°52'S, 168°27'E, 200-260 m, Epi: Javania exserta.
Stn 1043. - 30.09.1994, 16°52'S, 168°25'E, 350-372 m, Epi: Fungiacyathus (Fungiacyathus) paliferus.
Crispatotrochus rugosus.
Stn 1047.-30.09.1994, I6°53'S, 1 68° 10'E, 486-494 m, Epi: Trochocyathus (Trochocyathus) maculatus.
Stn 1051. — 01.10.1994. 16°36'S. 167 59'E, 555-558 m, Epi: Fungiacyathus (Bathyactis) margaretae,
Caryophyllia (Caryophyllia) sp. cf. calveri.
Stn 1056. - 01.10.1994, 16°33'S, 167°55'E, 602-620 m, Epi: Caryophyllia (Caryophyllia) sp. cf. calveri.
Stn 1058. - 02.10.1994, 16°12'S, 167°20'E, 319 m. Malakula: Trochocyathus (Aplocyathus) brevispina,
Stephanocyathus (Acinocyathus) spiniger. Dendrophyllia arbuscula.
Stn 1059. -02.10.1994, I6°12'S, !67°20'E. 408-430 m, Malakula: Conotrochus funicolunma.
Stn 1060. -02.10.1994. 16°13'S, I67°20’E, 375-397 m. Malakula: Trochocyathus (Trochocyathus) vasiformis,
Conotrochus funicolunma. Flabellum (Ulocyathus) aotearoa. Truncatoflabellum dens. T. vigintifarium Javania
exserta.
Stn 1061. _ 02.10.1994, 16°14'S, I67°20'E, 458-512 m, Malakula: Bourneotrochus stellulatus, Deltocyathus
crassiseptum. D. cameratus.
Source : MNHN. Paris
AZOOXANTHELLATE SCLERACTINIA
43
Stn io65. — 02.10.1994. 16°16'S, 167°21'E. 360-419 m, Malakula: Caryophyllia (Caryophyllia) abrupta,
Caryophyllia (Acanthocyathus) grayi, Trochocyathus (Aplocyathus) brevispina, Conotrochus funicolunma,
Tropidocyathus labidus, Flabellum (Flabellum) pavoninum, Flubellum ( Ulocyathus ) aotearoa,
Truncatoflabellum angustum , T. vigintifarium, T. mortenseni, Balanophyllia desmophyllioides.
Stn io67. — 02.10.1994, 16°16'S. I67°21'E, 344-366 m, Malakula: Caryophyllia (Caryophyllia) crosnieri.
Trochocyathus ( Trochocyathus ) vasiformis , Notocyathus conicus , Cryptotrochus brevipalus, Gardineria
hawaiiensis.
S[n io68. — 02.10.1994, 16°15'S, 167°19'E, 536-619 m. Malakula: Deltocyathus crassiseptum, Alatotrochus
rubescens, Tropidocyathus labidus.
Stn 1069. - 04.10.1994, 15°32'S, 167°14'E, 122-125 m, SE of Espirilu Santo: Caryophyllia ( Acanthocyathus )
grayi, Heterocyathus cf. sulcatus.
Stn io70. — 04.10.1994, 15°36'S, 167°16'E, 184-190 m, SE of Espiritu Santo: Fungiacyathus (Fungiacyathus)
paliferus. Letepsammiafranki, Caryophyllia ( Acanthocyathus ) grayi, Trochocyathus ( Trochocyathus )
philippinensis , Aulocyathus juvenescens, Flabellum ( Flabellum ) pavoninum , Truncatoflabellum mortenseni.
Stn |071. — 04.10.1994, 15°36'S, I67°16'E, 180-191 m, SE of Espiritu Santo: Caryophyllia (Acanthocyathus)
grayi, Trochocyathus (Trochocyathus) philippinensis, Rhizosmilia robusta, Flabellum (Flabellum) pavoninum,
Truncatoflabellum mortenseni, Endopachys grayi.
Stn 1072. — 04.10.1994, I5°39'S, I67°I9'E, 622-625 m. SE of Espiritu Santo: Caryophyllia (Caryophyllia)
octonaria, Trochocyathus (Trochocyathus) vasiformis, Conotrochus funicolunma. Heteropsammia cochlea.
Sin 1074. — 04.10.1994, 15°48'S, I67°24'E, 775-798 m, SE of Espiritu Santo: Fungiacyathus (Fungiacyathus)
stephanus, Caryophyllia (Caryophyllia) crosnieri, C. (C.) scobinosa, Stephanocyathus ( Odontocyathus )
weberianus.
Stn 1077, _ 05.10.1994, I6°04'S, 167°06'E, 180-210 m, Malakula: Madracis kauaiensis, Oculina virgosa,
Madrepora porcellana, Caryophyllia (Caryophyllia) lamellifera, Trochocyathus (Trochocyathus) maculatus,
Rhizosmilia robusta, Truncatoflabellum mortenseni, Balanophyllia rediviva.
Stn 1078. — 05.10.1994, 16°03’S, 167°26'E, 194-230 m, Malakula: Rhizosmilia robusta. Rhizotrochus typus.
Stn 1080. — 05.10.1994. 15°57'S, 167°27’E, 799-850 m, Malakula: Fungiacyathus ( Fungiacyathus ) stephanus,
Caryophyllia ( Caryophyllia ) diomedeae, Stephanocyathus (Odontocyathus) weberianus.
Stn 1084. — 05.10.94, 15°50'S. 167°17'E. 207-280 m, Malakula: Madrepora porcellana, Rhizosmilia robusta,
Flabellum ( Flabellum ) pavoninum.
Sm 1085. — 05.10.1994, 15°48’S, 167°18’E, 155-161 m, Malakula: Flabellum (Flabellum) pavoninum.
Truncatoflabellum martensii, Javania exserta. Endopachys grayi.
Stn 1086. — 05.10.1994, I5°36'S, 1 67° 1 6'E, 182-215 m. Malakula: Caryophyllia (Acanthocyathus) grayi.
Trochocyathus (Trochocyathus) philippinensis, Flabellum (Flabellum) pavoninum. Truncatoflabellum
candeanum , T. martensii , Endopachys grayi.
Stn 1087. — 06.10.1994, 1 5° 10'S, 167°14'E. 394-421 m, NE of Espiritu Santo: Stephanocyathus (Acinocyatlms)
spiniger, Bourneotrochus stellulatus [X], Conotrochus funicolunma, Flabellum (Flabellum) pavoninum,
Truncatoflabellum angustum.
Stn 1088. — 06.10.1994. 15°09’S, 167°15’E. 425-455 m. NE of Espiritu Santo: Madrepora oculata forma
formosa, Caryophyllia (Caryophyllia) abrupta, Trochocyathus (Trochocyathus) discus, Conotrochus
funicolunma, Desmophyllum dianthus. Peponocyathus folliculus.
Stn 1089. — 06.10.1994, 15°08'S, 167°17'E, 494-516 m, NE of Espiritu Santo: Madrepora porcellana.
Trochocyathus (Trochocyathus) discus, Conotrochus brunneus. Desmophyllum dianthus.
Sm 1090. — 06. 10.1994, 15°08'S, 167°17'E, 470-502 m. NE of Espiritu Santo: Trochocyathus (Trochocyathus)
discus , Conotrochus brunneus.
Sm 1091. — 06.10.1994, 15°I0'S. I67°13’E, 344-350 m, NE of Espiritu Santo: Stephanophyllia complicata [X).
Caryophyllia (Caryophyllia) abrupta [X], Conotrochus funicolunma, Flabellum (Flabellum) pavoninum [X],
Flabellum (Ulocyathus) aotearoa, Truncatoflabellum angustum.
44
S. D. CAIRNS
Stn 1092. — 06.10.1994, 15°10'S, 1 67° 1 2'E, 314-321 m, NE of Espiritu Santo: Stephcinocyathus ( Acinocyathas )
spiniger , Conor rochus brunneus [X].
Sm 1094. — 06.10.1994, 15°08’S, 1 67° 1 l'E, 312-314 m, NE of Espiritu Santo: Caryophyllia { Caryophyllia )
abrupta , Deltocyathus ornatus , Heterocyathus cf. sulcatus , Truncatoflabellnm pusillum , 7. vigintifarium.
Sm 1095. — 06.10.1994, 15°07'S, 1 67° 1 l’E, 304-320 m, NE of Espiritu Santo: Caryophyllia { Caryophyllia )
rugosa , ?C. fCJ lamellifera, Crispatotrochus rugosus , Tethocyathus virgatus , Dactylotrochus Genicom is,
Dendrophyllia alcocki.
Sm 1097. — 07.10.1994, 15°05’S, 167°10'E, 281-288 m, NE of Espiritu Santo: Stephanophyllia complicate! [X],
Caryophyllia {Caryophyllia) rugosa , Oxystnilia epithecata, Bourneotrochus stellulatus [X], Deltocyathus Stella
[X], Heterocyathus cf. sulcatus, Conotrochus asymmetros [X], Guynia annulata [X], Truncatoflabellnm
pusillum, T. vigintifarium , Javania fusca.
Stn 1098. — 07.10.1994, 15°04'S, 167°10'E, 277-285 m, NE of Espiritu Santo: Truncatoflabellnm pusillum,
T. vigintifarium, BalanophyUia desmophyllioides.
Stn 1 102. — 07.10.1994, 15°03'S, 167°08'E, 208-210 m, NE of Espiritu Santo: Oculina virgosa, Flabellum
( Flabellum ) pavoninum, Truncatoflabellnm mortenseni.
Stn 1 103. — 07.10.1994, 15°03'S, 167°07'E, 163-165 m, NE of Espiritu Santo: Caryophyllia (Acanthocyathus)
grayi, Trochocyathus ( Trochocyathus ) semperi, Flabellum {Flabellum) pavoninum, Truncatoflabellnm
mortenseni.
Stn 1 106. — 07.10.1994, 15°05’S, 167° 1 l'E, 305-314 m, NE of Espiritu Santo: Fungiacyathus (Fungiacyathus)
paliferus, Caryophyllia {Caryophyllia) quadragenaria, Crispatotrochus rugosus, Bourneotrochus stellulatus [X],
Deltocyathus Stella, D. heteroclitus [X], Truncatoflabellnm vigintifarium, T. pusillum [X], Javania fusca
Stn 1107. 07.10.1994, 15°05'S, 167°15’E, 397-402 m, NE of Espiritu Santo: Trochocyathus {Trochocyathus)
vasiformis , Conotrochus funicolumna.
Stn 1 108. 07.10.1994, 15°04'S. 1 67° 1 5'E, 405-419 m. NE of Espiritu Santo: Caryophyllia {Caryophyllia)
crosnieri, Tethocyathus virgatus, Dendrophyllia alcocki.
Sm 1109. — 08.10.1994, 14°52’S, I67°18’E, 1550-1620 m, N of Espiritu Santo: Stephcinocyathus
{Stephcinocyathus ) regi us.
Stn 1110. 08.10.1994, 14°49'S, 1 67° 1 5'E, 1360 m, N of Espiritu Santo: Stephanocyathus {Stephcinocyathus)
regius.
Stn 1111. 08.10.1994, 14°51'S, 167°14’E, 1210-1250 m, N of Espiritu Santo: Trochocyathus {Trochocyathus)
pate lliform is.
Sin 1113. - 08.10.1994, 14°52'S, 167°06'E, 700-736 m. N of Espiritu Santo: Stephanophyllia complicata,
Conotrochus funicolumna, Pleotrochus zihrowii, Cryptot rochus brevipalus, Truncatoflabellnm vigintifarium.
Stn 1 1 14. — 08.10.1994, I4°52'S, I67°03'E. 647 m, N of Espiritu Santo: Pleotrochus venustus, Javania fusca.
Stn 1125. 10.10.1994, 15°57S, 1 66°38'E. 1 160-1220 m, Guyot Bougainville: Fungiacyathus (Fungiacyathus)
stephanus. Stephanocyathus ( Stephanocyathus ) regius, Stephanocyathus ( Odontocyathus ) co.ronatus,
Deltocyathus rotulus, Truncal oflabellum stabile.
Stn 1127. - 10.10.1994, 15°58'S, 166°37’E. 1052-1058 m, Guyot Bougainville: Stephanocyathus
(Stephanocyathus) regius. Deltocyathus rotulus. Truncatoflabellnm stabile.
Stn 1128. 10.10.1994, 1 6"02 S, 166°38'E, 778-811 m, Guyot Bougainville: Fungiacyathus (Fungiacyathus)
pusillus pacificus, Caryophyllia ( Caryophyllia ) diomedeae, Desmophyllum dianthus, Javania fusca.
Stn 1 129. 10.10.1994, 16°00S, I66°39'E. 1014-1050 m, Guyot Bougainville: Fungiacyathus (Fungiacyathus)
stephanus, Stephanocyathus (Stephanocyathus) regius, Deltocyathus rotulus, Flabellum (Ulocyathus) marcus
Stn I 131. - 1 1.10.1994, 15°38'S, I67°03'E, 140-175 m, S of Espiritu Santo: Rhizotrochus typus.
Sin 1132. 11.10.1994, 15°38'S, 167°02'E, 161-182 in. S of Espiritu Santo: Flabellum (Flabellum)
pavoninum.
Source : MNHN. Pans
AZOOXANTHELLATE SCLERACTIN1A
45
S,n 1 134. — I 1.10.1994, 15°39’S, 167°02'E, 230-287 m, S of Espiritu Santo: Letepsammiafranki, Caryophyllia
( Caryophyllia ) octonaria, Caryophyllia (Acanthocyathus) grayi. Flabellum (Ulocyathus) aptearoa,
Truncatoflabelium mortenseni, Balanophyllia rediviva, Endopachys grayi.
Stn ) i 35. _ 1 1.10.1994, 15°40'S, 167°02'E, 282-375 m, S of Espiritu Santo: Flabellum (Flabellum) pavoninum
[X], Truncatoflabelium angustum [X],
Nimbus
Stn 12. — 7.07.1968, 26°32'S, I53°45’E, depth unknown, Queensland: Anthemiphyllia multidentata.
Stn 55. — 1968, 26°27'S, 153°50’E, 270-272 m, Queensland: Anthemiphyllia multidentata.
NZOI (New Zealand Oceanographic Institute), "Tangaroa"
Stn C527. — 18.09.1960, 32°30.0'S, 179°12.0'W, 508 m. southern Kermadec Ridge: Anthemiphyllia
macrolobata.
Sin 192. — 23.07.1975, 29°24.8'S. 168°13.2'E, 570-578 m, Norfolk I.: Flabellum (Flabellum) arcuatile.
Stn 197. _ 25.07.1975. 32°22.9'S. 167°28.2'E, 540-544 m. southern Norfolk Ridge: Flabellum ( Flabellum )
arcuatile.
Stn 1741. — 12.05.1979, 22°43.0'S, 159°16.0'E, 328 m, Lord Howe Seamount Chain: Balanophyllia
desmophyll ioides.
Stn K803. — 22.07.1974, 29°16.0’S, 177°50.3'W. 190-140 m, Raoul I„ Kermadec Is: Madracis kauaiensis.
Stn K825. — 25.07.1974, 28°47.8'S, 177°47.8'W, 145 m, Raoul I„ Kermadec Is: Madracis kauaiensis.
Sin K826. —25.07.1974, 28°48.0’S, 177°48.0'W, 142 m. Raoul I.. Kermadec Is: Madracis kauaiensis.
Stn K830. — 26.07.1974, 29° 1 1.5’S, 177°53.0'W. 545-590 m. Raoul I„ Kermadec Is: Oxysmilia circularis.
Stn K842. — 29.07.1974, 30°10.2'S, 178°35.9'W. 325-370 m. McCauley I., Kermadec Is: Anthemiphyllia
pacifica.
Stn K858. — 30.07.1974, 30°34.2’S. I78°29.8'W, 465-501 m, Curtis I.. Kermadec Is: Oxysmilia circularis.
Stn K872. — 2.08.1974, 31°20.4’S, 178°49.2'W, 280-235 m, Esperance Rock. Kermadec Is: Anthemiphyllia
pacificci
Stn PI 15. — 31.05.1977, 31°25.9'S, 159°02.2'E, Lord Howe Is: Thalamophyllia tenuescens.
Stn U599. — 08.02.1988. 30°43.0'S, 173°16.0'E, 640-590 m. Three Kings Ridge. New Zealand: Flabellum
( Flabellum ) arcuatile.
" Townsend Cromwell"
Stn 81-01-14. — 02.02.1981. 23°15'48"N. I6I°50'12"W, 369 m, Hawaiian Is: Anthemiphyllia macrolobata.
"Toyoshio- Maru "
Stn 1 1. — 1 1.1 1.1995, 28°05'N. I29°47'40"E, 302 m, Ryukyu Is: Madrepora minutiseptum.
USGS (United States Geological Survey)
Stn 24918. — Late Pleistocene, Navaka River, Espiritu Santo: Madrepora porcellana.
Stn 25715. — Late Pleistocene, Kcre River, Espiritu Santo: Madrepora porcellana. Caiyophyllia ( Acanthocyathus )
grayi , Dendrophyllia ijirnai.
Stn 25718. — Late Pleistocene, Kere River, Espiritu Santo: Madrepora porcellana.
Stns SM242, 129a and 129b. — Late Pleistocene, Kere River, Espiritu Santo: Caiyophyllia (Acanthocyathus)
grayi.
46
S. D. CAIRNS
HISTORICAL REVIEW
No azooxanthellate Scleractinia were previously known from the Wallis and Futuna region, and only two
authors reported azooxanthellate corals from Vanuatu: Neohelia porcellana from Epi (Moseley. 1881), and
17 Late Pleistocene species from Espiritu Santo (Wells, 1984). However, many species have been reported from
other islands throughout the tropical central Pacific, which are summarized in Table 1. Although the specimens
reported in this paper are primarily deep-water in habitat, it was attempted to include references to all
azooxanthellate species in Table 1, regardless of depth. Only some of the more significant papers are briefly
discussed below.
Table 1. — Summary of azooxanthellate Scleractinia previously reported from the tropical central Pacific. (S = 0-100 m,
D = over 100 m, + = fossil).
Source : MNHN. Paris
AZOOXANTHELLATE SCLERACT1NIA
47
The first azooxanlhellate coral reported from the tropical central Pacific was Flabelluni pavoninum Lesson.
1831 from the "Sandwich Islands" (= Hawaiian Islands), which is ironic, since it is a deep-water species that was
collected before the era of deep-water dredging. The "Challenger" Expedition, which ushered in the era of deep-water
biology, collected 6 species from various localities throughout the central Pacific, including one new species
(Neohelia porcellana) from the Vanuatu archipelago (MOSELEY. 1881). The collection ol the nine species rcpoited
by Gardiner (1899a) from the Loyalty Islands, all from Sandal Bay at 73 m. is significant in that many of these
species are also found at greater depths, as reported herein. Gardiner's specimens were split between the British
Museum and the University Museum of Zoology, Cambridge.
The Hawaiian azooxanlhellate fauna is relatively well known, due to the seminal work of Vaughan (1907)
and later additions by Cairns (1984), all specimens of which are deposited at the USNM. Although a northern
outlier to the central Pacific region, the Hawaiian fauna includes many species found throughout the Indo- Pacific
(see Distribution section), as well as some apparent endemics.
Another significant paper on corals from the central tropical Pacific was that ot Wells (1954) on the Recent
corals of the Marshall Islands, in which 13 azooxanlhellate species are reported from Bikini Atoll, mostly from
depths shallower than 200 m. These specimens arc deposited at the USNM.
Keller (1981a) reported eight species from several exotic central Pacific bathyal localities, including the
Marcus-Necker Ridge (18-23°N, 157-I78°E), Hermit Atoll (1°S. 145°E), Demitri Mendeleev Seamount (5°N.
155°E), and the Hawaiian Islands. These specimens are deposited at the Institute of Oceanology. Moscow.
As the Hawaiian Islands are to the northern border of the central tropical Pacific, (he ridges and islands north of
New Zealand (north of 33°S) are to the southern central tropical Pacific. In his revision of the New Zealand
azooxanlhellate Scleractinia, Cairns (1995: Table 2) listed 76 species from the Kermadec and the Lord Howe
Islands, and the northern Colville, Three Kings, and Norfolk ridges; additional records of 12 of these species were
also reported from Chesterfield, Cook, Tonga, and Samoa Islands.
Azooxanlhellate species were reported in an incidental fashion in a number of papers published alter 1984 (see
Table 1), often as host species for symbionts, or as a part of a larger checklist of the shallow water coral fauna
48
S. D. CAIRNS
from a region. It was attempted to make Table 1 as complete as possible, but undoubtedly some references of this
nature were overlooked.
As a result of this study, 1 16 azooxanthellate species are reported from the Vanuatu Archipelago and 83 from
the Wallis and Futuna Archipelago (Table 2). Incidental records arc also reported from the Kermadec Islands
(3 species), Hawaiian Islands (2), Johnston Atoll (1), Lord Howe Seamount Chain (1), Guam (1), Chesterfield
Island ( 1 ), and the ridges and islands north of New Zealand ( 1 ).
The following azooxanthellate coral faunas have been reviewed from regions bordering the tropical central
Pacific: northern temperate Pacific (Cairns, 1994), eastern temperate Pacific (Cairns, 1994), eastern tropical
Pacific (Cairns, 1991a), southern temperate Pacific (Cairns, 1982, 1995), and western tropical Pacific (Cairns,
1989a; Cairns & Zibrowius, 1997).
MATERIAL
This study was based on the examination of approximately 4400 specimens (1042 lots) collected from
227 stations. Most of the specimens were collected by the French research expedition MUSORSTOM 7 in the Wallis
and Futuna archipelago in 1992 (Richer de Forges & Menou, 1993) and MUSORSTOM 8 in the Vanuatu
archipelago in 1994 (Richer de Forges, Faliex & Menou. 1996).
A novel source of specimens heretofore overlooked by most are those deep-water corals that have been cemented
into the shell of Xenophora carrier shells. Nineteen species and several more unidentified taxa were distinguished
from 40 Xenophora shells from the MUSORSTOM 8 expedition in the Vanuatu Archipelago, some shells bearing as
many as 6 different species. This source of specimens is considered promising because, despite numerous stations
made by the MUSORSTOM expedition in this region, of the 19 species found on these shells, three ( Fungiacyathus
variegatus, Peponocyathus folliculus , and Temnotrochus kermadec ensis) arc known from this region only as
Xenophora- collected, and another live species ( Guynia annulata, Tropidocyathus labidus, Notocyathus conicus,
Deltocyathus heteroclitus, and Trochocyathus discus) were otherwise rarely collected from this region by
conventional means. The depth ranges of the Xenophora- collected coralla are consistent with the known depth
range of the coral species, implying that bathymetric transport by the gastropod is probably negligible.
Furthermore, although most coralla cemented to Xenophora shells are dead, some were still alive when attached and
when collected.
METHODS
Species descriptions and illustrations arc provided only for those species described as new or for those for which
no adequate description previously existed. Shorter diagnoses are provided for most of the remaining species for
which new material is reported, with an indication as to where to find a more complete description.
Species synonymies are complete unless otherwise indicated with a reference to a more complete account;
however, li was attempted to include in the synonymies all references to specimens reported from the tropical
central Pacific. When possible, historical records were verified, but when material was unavailable and the
published account unclear, the synonymy entry and corresponding distribution record arc queried. In order to clarify
what historical material was examined by the author, the following convention was employed (see Matthews,
1973). The three symbols used in the synonymies, which always precede the year of publication, are:
*’ meaning this entry represents a valid species under the terms of the ICZN (1985);
v*, meaning that the author (SDC) has examined the type of this species;
v., meaning that the author (SDC) has examined this/these nontype specimens and agrees that they belong
in synonymy.
The letter v is Latin for vidimus (we have seen). The vidimus convention is not used for publications written
by the author of the cited publication, since it is self-evident that he has seen the specimens that he has previously
reported.
Source : MNHN, Paris
AZOOX AN'THELL ATE SCLERACT1NIA
49
In the "Material Examined" sections, the number of specimens examined follows the station number, followed
by the museum of deposition, and its catalog number, if any. Holotypes and paratypes arc deposited primarily at
the MNHN and NMNH.
In order to avoid erroneous depth ranges for species as a result of bathymetrically wide-ranging trawls, a
confirmed depth range is employed in this paper, which is defined as the deepest shallow to the shallowest deep
component of all trawls considered. For example, if a species was trawled at a station indicating 20-300 m and
again at a station indicating 250-500 m, the confirmed depth range is 250-300 m. a depth range within which the
species must occur.
Conventional photography was done by the author, in some cases the corallum dyed black with cloth dye and
recoated with sublimed ammonium chloride in order to improve contrast for photography.
DISTRIBUTION
Inter-regioncil comparisons. — The Vanuatu archipelago and Wallis and Futuna islands and adjacent seamounts
lie near the centre of the large Indo-Polynesian tropical marine province as defined by Briggs (1974). which
extends from the Persian Gulf to the Tuamotu Archipelago. The 1 16 azooxanthellate species listed for the Vanuatu
region (Table 2) compare closely in number to the 125 listed by Cairns & ZlBROWlUS ( 1997) lor the highly
diverse Kai Islands. Indonesia, especially considering that the shallow and extremely deep environments of the
Vanuatu region were not sampled. Indeed, 95 (71%) of the 134 species from the Vanuatu and Wallis and Futuna
regions are also known in the tropical western Pacific region, this number decreasing to 62 (46%) farther west in
the tropical Indian Ocean (Table 2). On the other hand, only seven species are found in common with the
Vanuatu/Wallis and Futuna region and the tropical eastern Pacific: five of those being cosmopolitan; one.
Endopachys grayi , being Indo-Pacific in distribution; and the seventh. Polymyces wellsi . amphi-Pacific in
distribution. Ironically, the Vanuatu/Wallis and Futuna region has more species in common with the Atlantic
Ocean (11) than the eastern Pacific (7)(see Table 2). Among these II species, 6 may be considered to be
cosmopolitan; two ( Caryophyllia ambrosia and Truncatoflabellum stabile ) have potentially continuous
distributions via the Indian Ocean; but three species ( Stephanocyathus coronatus , Vaughanella concinna , and
Peponocyathus folliculus) have apparent disjunct distributions between the central Pacific and the western or
northern Atlantic.
Because deep water temperature gradients do not always exactly correspond to shallow water gradients, the
zoogeographic boundaries of deep-water corals arc sometimes less well-defined than lor shallow-water organisms
(Briggs, 1974; Cairns, 1994, 1995). Thus, it is not considered unusual that 43 species from the Vanuatu/Wallis
and Futuna region are also known from the warm to cold temperate region ol Japan, and 32 species extend to the
temperate regions of New Zealand and southern Australia (Table 2).
Intraregional comparisons. — Within the tropical central Pacific, the Vanuatu/Wallis and Futuna
azooxanthellates have a strong affinity (56 species, Table 2) to those species found in the tropical regions
to the south, composed of the Kcrmadec Islands and Ridge, Colville Ridge, Norfolk Island and Ridge, and
the Lord Howe Islands and Seamount Chain. These submarine ridge systems north of New Zealand form a natural
conduit for the migration of corals in both directions between the New Zealand region and the Tonga Platform
and the New Caledonian region. And, whereas only 9 species are listed lor the New Caledonian region in I able 2,
unstudied MUSORSTOM collections from this region and the Loyalty Islands suggest an extremely diverse fauna
very similar to that of the Vanuatu region. The Hawaiian azooxanthellate fauna constitutes a separate province in
the tropical Pacific (Briggs, 1974), having an endemic component of 48% (CAIRNS, 1984). Nonetheless,
24 species known from the Vanuatu/Wallis and Futuna region also occur off the Hawaiian Islands ( I able 2). Other
than the Hawaiian Islands, the ridges north of New Zealand, and the region under study, azooxanthellates are poorly
known from other tropical central Pacific regions, only 17 species listed in Tabic 2: column 5, symbol X;
see also Table 1 .
50
S. D. CAIRNS
Table 2. — Geographic distribution and bathymetric ranges of the Recent azooxanthellate scleractinian species known
from the regions of Wallis & Futuna and Vanuatu Archipelagos.
Key to areas: 1-7, Tropical regions: I. Indian Ocean; 2. western Pacific; 3. Vanuatu region; 4, Wallis and Futuna region; 5 other central
Pacific islands (H. Hawaiian Islands; NZ. ridges north of New Zealand; NC, New Caledonia. Loyalty Islands, and/or Chesterfield
Islands; OI, other central Pacific islands); 6. eastern Pacific; 7. Atlantic Ocean. — 8-9, Temperate regions: 8. Japan (excluding Ryukyu
IsL); 9. New Zealand. Australia. }
Bathymetric ranges (given in meters) only for records from Vanuatu/Wallis and Futuna region.
Symbols: + fossil occurrence.
Source . MNHN, Paris
AZOOXANTHELLATE SCLERACTINIA
51
C. (Acanthocyathus) grayi (H. Milne Edwards &
Haime, 1848)
Crispatotrochus rubescens (Moseley. 1881)
C. rugosus Cairns. 1995
Labyrinthocyathus limamlus (Squires, 1964)
Oxysmilia circularis Cairns
0. corrugata sp. nov.
0. epithecata sp. nov.
Trochocyathus (T.) vasiformis Bourne, 1903
T. (T.) rhombocolumna Alcock. 1902
T. (T.) philippinensis Semper, 1872
T. IT.) maculatus Cairns, 1995
T. (T.) efateensis sp. nov.
T. (T.) patelliformis sp. nov.
T. (T.) semperi Cairns & Zibrowius. 1997
T. (T.) cooperi (Gardiner, 1905)
T (T.) discus Cairns & Zibrowius, 1997
T. (Aplocyathus) hastatus Bourne. 1903
T. (A.) brevispina Cairns & Zibrowius, 1997
Tethocyalhus virgatus (Alcock, 1902)
Polycyathus octuplus sp. nov.
Bournedtrochus s tel I ulatus (Cairns, 1984)
Stephanocyathus (S.) regius Cairns & Zibrowius,
1997
5. ( Odontocyathus ) coroncitus (Pourtales, 1867)
S. (O.) weberianus (Alcock, 1902)
S. ( Acinocyathus ) spiniger (Marenzeller, 1888)
Vaughanella concinna Gravier, 1915
Dellocyathus magnificus Moseley, 1876
D. roiulus (Alcock, 1902)
D. suluensis Alcock, 1902
D. taiwanicus Hu, 1987
D. vaughani Yabe & Eguchi. 1932
D. crassiseptum sp. nov.
D. cameraius sp. nov.
D. Stella Cairns & Zibrowius, 1997
D. heteroclitus Wells. 1984
D. o mat us Gardiner. 1899
Heterocyathus sp. cf. H. sulcatus (Verrill, 1866)
H. alternants Verrill, 1865
Conotroc/ius funicolumna (Alcock. 1902)
C. brunneus (Moseley, 1881)
C. asymmetros sp. nov.
Lochmaeotrochus gardineri sp. nov.
Aulocyathus recidivus (Dennant, 1906)
A. juvenescens Marenzeller, 1904
Desmophyllum dianthus (Esper. 1794)
Thalamophyllia tenuescens (Gardiner, 1899)
Source
52
S. D. CAIRNS
Lop he I ia pertusa (Linnaeus. 1758)
Ddctylotrochus cervicornis (Moseley, 1881)
Rhizosmilia robusta Cairns, 1993
AsterosmUici gigas (van der Horst, 1931)
TURBINOLIIDAE
Alatotrochus rubescens (Moseley, 1876)
Pleotrochus venustus (Alcock, 1902)
P. zibrowii Cairns, 1997
Tropidocyathus lessonii (Michelin, 1842)
T. labidus Cairns & Zibrowius, 1997
Cyathotrochus pileus (Alcock, 1902)
Deltocyathoides orientcilis (Duncan, 1876)
Notocyathus conicus (Alcock, 1902)
N. venustus (Alcock, 1902) sensu Wells, 1984
Cryptotrochus brevipcilus sp. nov.
Idiotrochus kikutii (Yabe & Eguchi, 1941)
Peponocyathus folliculus (Pourtales, 1868)
GUYNIIDAE
Guynia annulcita Duncan, 1872
Truncatoguynia irregularis Cairns, 1989
Temnotrochus kermadecensis Cairns. 1995
FLABELLIDAE
Flabellum (F.) pavoninum Lesson, 1831
F. (F.) circuatile sp. nov.
F. (Ulocyathus) deludens Marenzeller, 1904
F. (U.) aotearoa Squires, 1964
F. (U.)marcus Keller. 1974
F. (U.) hoffineisteri Cairns & Parker, 1992
F. (U.) apertum apertum Moseley, 1876
T run cat oflabell um stabile (Marenzeller, 1904)
T. dens (Alcock, 1902)
T. pus ilium Cairns, 1989
T. angustum Cairns & Zibrowius. 1997
T. phoenix Cairns, 1995
T. vigintifarium sp. nov.
T. mortenseni Cairns & Zibrowius, 1997
T. Vanuatu (Wells, 1984)
T. aculeatum (H. Milne Edwards & Haime. 1848)
T candeanum (H. Milne Edwards & Haime, 1848)
T. martensii (Studer, 1878)
Javania lamprotichum (Moseley, 1880)
J.fuscus (Vaughan, 1907)
J. exserta sp. nov.
Rhizotrochus typus H. Milne Edwards & Haime
1848
Source : MNHN, Pans
AZOOXANTHELLATE SCLERACTIN1A
53
NUMBER OF SPECIES IN EACH AREA: 1 = 62; 2 = 95; 3 = 1 16; 4 = 83; 5 = 72: 6 = 7; 7 = II; 8 = 42; 9= 32
SYSTEMATIC ACCOUNT
Order SCLERACTINIA
Suborder ASTROCOENIINA
Family POCILLOPORIDAE
Genus MADKAC1S H. Milne Edwards & Haime, 1849
Madracis kauaiensis Vaughan. 1907
Figs 1 a-e
? Madracis Hellena - STUDER, 1878: 636 [Not M. hellana H. Milne Edwards & Haime. 1850].
IMadracis singularis Rehberg, v‘1892: 10-11, pi. 1, figs 3-4 (Not specimen from Fiji).
Madracis kauaiensis Vaughan, v*1907: 83-84, pi. 9, figs 1-4. — Cairns. 1984: 6.
IMadracis inlerjecia Marenzeller, ‘1907: 20-21. pi. 2, fig. 3.
MATERIAL EXAMINED. — Wallis and Futuna Islands. Musorstom 7: stn 496. 1 branch (MNH'N). — Stn 499,
I branch (98543). — Stn 500, 1 branch (USNM 98544). — Stn 502. 1 branch (MNHN). — Stn 504. 6 branches (MNHN).
— Stn 507. I branch (MNHN). — Stn 508, I branch (MNHN). — Stn 509. 1 large colony and numerous branches
(MNHN). 5 branches and SEM stubs 869-870 (USNM 98541). — Stn 515. 5 branches (MNHN).
Vanuatu. Musorstom 8: stn 988. 41 branches (MNHN). — Stn 1077, 2 colonies and several branches (USNM
98549).
Hawaii Islands. HURL: stn 83-202. 1 colony (USNM 83460).
54
S. I). CAIRNS
New Zealand. NZOI: stn K803. 1 branch (USNM 93996). — Stn K825. 4 branches (USNM 93997). — Sin K826.
I branch (USNM 93998).
TYPE Locality. — 'Albatross" tin 3982: 2I°56,25,'N, I59°21,40"W (Kauai. Hawaiian Islands). 73-426 m.
Diagnosis. — Colonies flabellatc lo slightly bushy; irregularly branched; lower branches of large colonies
often anastomosing with one another. Largest corallum (Musorstom 7 stn 509) 14 cm in height. 14 cm wide,
and uniplanar. Branches circular in cross-section: basal branches up to 8 mm in diameter, distal branches 2.5-
3.0 mm in diameter. Corallites slightly elliptical in shape, their greater axis aligned with the branch axis;
corallites 1.25-1.35 mm in greater diameter, occurring homogeneously and well spaced on all branch surfaces.
Coenosteum white, bearing prominent spines up to 15 mm in height. Corallites contain 10 equal-sized septa, each
about 0.15 mm exsert and bordered internally by a small (0.05 mm wide) paliform lobe. Central region of cal ice a
broad, flat platform, connected to axial edges of septa and supporting a laterally compressed styliform to lamellar
columella, the greater axis of columella aligned with the greater axis of corallile.
Remarks. — H. Milne EDWARDS & Haime (1850) described Madracis hellana from Reunion at a depth of
37 m, for a branching species of this genus having 10-12 septa per corallile. Studer (1878) reported a second
specimen of this species from Bougainville Island (88 m). this specimen deposited at the ZMB. the first report of
this genus for the Pacific. Rehberg (1892), believing that STUDER's southwest Pacific specimen must be different
from the western Indian Ocean M. hellana, redescribed STUDER's specimen as Madracis singulars , n. sp., and
reported another specimen of his presumed new species from Viti (Fiji) at 146 m. Finally. VAUGHAN (1907)
described another decameral Pacific Madracis species. M. kauaiensis , from the Hawaiian Islands (44-541 m). The
type of M. hellana was not examined and the type of M. singulars Rehberg, 1892 (Studer’s M. hellana from
Bougainville. ZMB 1842) was not available for examination and thus their identity with the specimens reported
herein remains in doubt, but the second specimen reported by REHBERG from Fiji at 80 fathoms (= 146 m)(ZMB
601) was examined and found to be quite different from the specimens reported herein. The Fiji specimen, which
might be interpreted as a syntype of M. singular is, has thicker, blunt to clavate branches; larger calices (GCD 1.6-
2.2 mm); no paliform lobes; and a continuous, spinose ridge that encircles each calice. The specimens reported
above are. however, indistinguishable from the type series of M. kauaiensis Vaughan, 1907, and thus are identified
as such.
Distribution. — Wallis and Futuna region: Futuna; 240-516 m. Vanuatu region: Tanna and Malakula; 210-
372 m. Elsewhere: Hawaiian Islands; Johnston Island (reported herein); Kermadec Ridge (reported herein);
‘/Bougainville (Studer, 1878); ?Gulf of Aqaba (Marenzeller. 1907): 44-541 m.
Suborder FUNGIINA
Family FUNGIACYATHIDAE
Genus FUNGIACYATHUS Sars, 1872
Subgenus FUNGIACYATHUS (FUNGIACYATHUS) Sars, 1872
Fungiacyathus (F.) stephanus (Alcock, 1893)
Baihyactis stephanus Alcock, *1893: 149, pi. 5, figs 12-12a.
Bathyactis sibogae Alcock. v*1902a: 108 (in pari); v,1902c: 38 (in part: "Siboga" stn 95 and large specimen of 57 mm
GCD).
Fungiacyathus (F.) stephanus - Cairns. 1989a: 7-9, pi. 1, figs a-k. pi. 2. figs a-b (synonymy); 1994: 37. pi. I 3. figs g-j-
1995: 31-32. pi. 1, figs a-c, map 1; 1998: 369. — Cairns & Zibrowius, 1997: 68-69 (synonymy).
Source :
AZOOXANTHELLATE SCLERACTINIA
55
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 551. 2 (MNHN). — Stn 552.
3 (MNHN). — Stn 565, 1 (MNHN). — Stn 621, 1 (MNHN).
Vanuatu. Musorstom 8: stn 963. 2 (MNHN). — Sin 975, 1 (MNHN). — Stn 990, 1 (MNHN). — Stn 992. I (USNM
98539). — Stn 996, 3 (USNM 98537). — Stn 1007, 1 (MNHN). — Stn 1036, 3 (MNHN). — Stn 1074, I (MNHN). —
Stn 1080, 1 (USNM 98540). — Stn 1125, 1 (MNHN). — Stn 1129, 13 (MNHN).
Type Locality. — "Investigator" stn 133: 15°43’30"N, 81°19'30"E (off Kristna Delta, Bay of Bengal),
1240 m.
Table 3. — Characteristics of the seven Indo-Pacific species of Fungiacyathus (Fimgiacyathus).
Source : MNHN, Paris
56
S. D. CAIRNS
Remarks. — This species is more fully described and illustrated by CAIRNS (1989a. 1994). It is distinguished
from its consubgenerics by having corrugated septa and sinuous costae; small, acute paliform lobes (P2): and a
very fragile corallum, probably a result of its greater depth of occurrence (Table 3).
Distribution. — Wallis and Futuna region: Combe Bank; 795-1280 m. Vanuatu region: Anatom, Tanna.
Erromango, Efate, Espiritu Santo, and Malakula; Guyot Bougainville; 440-1160 m. Elsewhere: widespread
throughout Indo-West Pacific; 245-2000 m (Cairns & Zibrovvius, 1997).
Fungiacyathus (F.) pusillus pacific us Cairns. 1995
Fungiacyathus (F.) pusillus pacificus Cairns, *1995: 32-33, pi. 1, figs g-i. 1. map 13.
Material EXAMINED. — Wallis and Futuna region. Musorstom 7: sin 522, 3 i.MNHN). — Stn 534, 1 (USNM
98536). — Stn 535, 2 (MNHN). — Stn 540. 2 (MNHN). — Stn 570. I (MNHN). — Stn 572, 3 (USNM 98535) —
Stn 575, 1 (USNM 98534). — Stn 581, I (MNHN).
Vanuatu. MUSORSTOM 8: stn 1 128, 1 (MNHN).
Type Locality. — NZOI stn U599: 30°43'S, I73°16 E (northern Three Kings Ridge). 590-640 m.
Remarks. — Based on these specimens little can be added to the original description, except to note a
maximum size of 23.3 mm GCD (MUSORSTOM 7 stn 535). The species is distinguished by its tall S 1 -4 septal
lobes, the peripheral edges of which all extend about the same distance outward and are almost vertical, which
results in an almost cylindrical corallum with a domed surface, like a thick button. The species is compared to
other consubgenerics from this region in Table 3.
Distribution. — Wallis and Futuna region: Wallis; Waterwitch and Combe Banks; 425-650 m. Vanuatu
region: Guyot Bougainville; 778-81 1 m. Elsewhere: Norfolk. Three Kings, and Colville Ridges north of New
Zealand; 350-988 m.
Fungiacyathus (F.) sandoi sp. nov.
Figs 1 f-h
Material EXAMINED/TYPES. — Wallis and Futuna region. MUSORSTOM 7: stn 523. 2 paratypes (MNHN). —
Stn 538. holotype (MNHN) and 1 paratype (USNM 98531). — Stn 540. I paratype (MNHN). — Stn 541. 1 paratype
(MNHN). — Stn 542, I paratype (USNM 98533). — Stn 569, 5 paratypes (USNM 98532). — Stn 589, I paratype
(MNHN). — Stn 590, 3 paratypes (MNHN). — Stn 597, 1 paratype (MNHN). — Stn 605. I paratype (MNHN).
Type Locality. — Musorstom 7, stn 538: I2°30.8'S, 176°40.3’W (Waterwitch Bank), 275-295 m.
Etymology. — This species is named in honour of my friend and colleague William J. Sando (1927-1996),
noted Lower Carboniferous coral taxonomist and stratigrapher.
Description. — Corallum circular: largest specimen (Musorstom 7 stn 542) 20.2 mm in CD; holotype
15.2 mm in CD. Base slightly convex, the basal angle ranging from 140°- 1 70°. All costae equally wide and
prominent (no cycle larger than another), rounded, each ornamented with a row of coarse (0. 16-0. 1 8 mm diameter)
rounded granules. All costae project about 0.4 mm beyond calicular edge. Septa hexamerally arranged in
5 complete cycles in typical fungiacyathid fashion, the S5 easily visible in a corallum of 8 mm CD. Si consist of
3 or 4 blunt, axial trabecular spines inclined toward the fossa; peripheral to these spines is a tall septal lobe
composed of 9- 1 3 vertically oriented trabeculae, each trabeculum corresponding to a serrate ridge on the septal face.
Ihe peripheral edge of the Si lobe is vertical, changing abruptly into a low marginal shelf region consisting of 3
or 4 low spines. Seven to nine synapticular plates occur along the length of an Si, the 4th or 5th from the centre
being the tallest. S2 also consist of 3 or 4 blunt, axial trabecular spines, but these spines are longer and wider than
Source
AZOOXANTHELLATE SCLERACTINIA
57
those on the Si, some composed of 2 trabeculae; peripheral to these spines is a prominent septal lobe composed of
7-9 trabeculae, the lobe inclined slightly outward. The peripheral edge of the S2 lobe is vertical to slightly
undercut, also changing abruptly to a low marginal shelf. S3 consist of 4-6 slender, axial trabecular spines
peripheral to which is a small septal lobe composed of 3-6 trabeculae. S3 septal lobe shorter and more inclined
outward than S3 lobe, but its peripheral edge is similar. S4 consist of 5-7 small trabecular spines; S5. 3 or
4 spines. All septa planar, with straight upper and peripheral edges. A low marginal shelf of about 0.4 mm width
surrounds the corallum. The columella is an intermingling of the innermost, blunt Si-2 trabecular spines.
Remarks. — Fungiacyathus sandoi appears to be most similar to F. paliferus (Alcock. 1902), both species
having granular costae, a similar septal structure, and achieving the same size. But. F. sandoi is distinguished in
having: a convex base; coarsely granular costae; thick, blunt S2 trabecular spines (not small P2 paliform lobes);
and uniformly exsert costosepta (Table 3).
Distribution. — Wallis and Futuna region: Wallis; Waterwitch, Combe, and Field Banks; 295-600 m.
Fungiacyathus ( F.) paliferus (Alcock, 1902)
Fig. 2 a
Bathyactis palifera Alcock, v* 1902a: 108; v. 1902c: 38. pi. 5, figs 34, 34a.
Fungiacyathus sp. cf. F. Stephanas - Wells, v.1984: 207, fig. I, 3-4 (USGS 25715, USNM 71828).
Fungiacyathus fragilis - Wells, v.1984: 205-206 (in part: USGS 25718, USNM 73957).
Fungiacyathus ( F.) paliferus - CAIRNS, 1989a: 9-10. pi. 2. figs c-i, pi. 3, figs a-c (synonymy and description); 1994:
37-38, pi. 14, figs a-e (synonymy and description); 1998: 369-370. — Cairns & Zibrowius, 1997: 69-70.
MATERIAL EXAMINED. — Vanuatu. MUSORSTOM 8: stn 963, 8 (USNM 98529). — Stn 967, 3 (USNM 98530). —
Stn 969, 1 (MNHN). — Stn 988, 1 (MNHN). — Stn 1003. 5 (MNHN). — Stn 1005, 3 (MNHN). — Stn 1016, 2 (MNHN).
— Stn 1017. 2 (MNHN). — Stn 1018, 9 (USNM 98528). — Stn 1043, 1 (MNHN). — Stn 1070. 1 (MNHN). — Stn 1 106,
1 (MNHN).
Type Locality. — " Siboga " stns 153 and 98: Sulu Sea and Moluccas, 141-350 m.
Remarks. — The species was redescribcd and illustrated by Cairns (1989a. 1994); however, several well-
preserved specimens listed above allow an additional observation that the CS1-2. and their adjacent CS5, project
about 1.2 mm beyond the other costosepta as rectangular lancets.
All specimens reported above were intact coralla, not the result of asexual fragmentation. Comparisons to other
species in this region are made in Table 3.
Although the Pleistocene Vanuatu specimens reported by WELLS (1984) as F. fragilis from USGS stn 25718
are reidentified herein as F. paliferus , those he reported and illustrated from USGS stn 24918 (USNM 71837 and
73977) belong to the subgenus F. ( Bathyactis ), but arc too small to identify to species.
Distribution. — Vanuatu region: Anatom. Tanna, Erromango, Efate, Epi. and Espiritu Santo (also
Pleistocene of Vanuatu: WELLS, 1984); 190-400 m. Elsewhere: widespread throughout Indo-West Pacific;
69-823 m (Cairns & Zibrowius, 1997).
Subgenus FUNGIACYATHUS (BATHYACTIS) Moseley, 1881
Fungiacyathus (B.) margaretae Cairns, 1995
Figs 2 b-c
Fungiacyathus (B.) margaretae Cairns. *1995: 33-34. pi. 2. figs a-c.
Source :
58
S. D. CAIRNS
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 534, 9 (MNHN). — Sin 535,
4 (MNHN). — Stn 540, 21 (USNM 98524). — Stn 541, 1 (MNHN). — Stn 546, 7 (MNHN). — Stn 564, 57 (USNM
98527). — Stn 565, 56 (MNHN).
Vanuatu. MUSORSTOM 8: stn 956, 2 (MNHN). — Stn 963. 47 (MNHN). — Sin 1011, 1 (USNM 98525). — Stn 1014.
I (MNHN). — Stn 1051. 2 (USNM 98526).
Type Locality. — NZOI stn P944: 27°20.8’S, I79°20.9'W (northern Colville Ridge), 673 m.
Remarks. — Previously known from a type series of only 5 specimens: 188 additional coralla arc
reported herein, the largest specimen (MUSORSTOM 8 stn 963) 18.6 mm in CD. the same diameter as the
largest type. Although the base of this species is always concave, the MUSORSTOM specimens show that the
types had unusually concave bases. Many of these specimens also show a pronounced ridging of the C|-2
(Fig. 2 c).
Fungiacyathus margaretae is most similar to F. granulosus , small and/or damaged specimens of the latter being
difficult to distinguish from the former. In general, F. margaretae is distinguished by having a concave base (that
of F. granulosus is usually flat); higher S| septal lobes with vertical peripheral edges, and constructed of vertically
aligned trabeculae (the Si septal lobes of F. granulosus are lower and sloped at the peripheral calicular edge,
and composed of trabeculae that are inclined outward toward the peripheral edge); and having taller synapticular
plates between septa. Both species have fine or coarse granular costae, and often granular coenosteum between
the costae.
Distribution. — Wallis and Futuna region: Waterwitch, Combe, and Tuscarora Banks; 470-1015 m. Vanuatu
region: Anatom. Efate, and Epi; 440-1 175 m. Elsewhere: northern Colville Ridge; 635-673 m.
Fungiacyathus (II.) granulosus Cairns, 1989
Fungiacyathus (B.) granulosus Cairns, *1989a: 11. pi. 4. figs d-i; 1994: 39, pi. 15. figs d-e; 1998: 370. — Cairns &
Zibrowius, 1997: 71.
Material EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 619, 2 (MNHN).
Vanuatu. Musorstom 8: stn 977. 1 (MNHN). — Stn 980. 3 (USNM 98520).
Type Locality. — "Albatross" stn 5590: 4°10'50MN, 1 18°39'35nE (Celebes Sea off Sabah). 567 m.
Remarks. — See Remarks for previous species.
Distribution. — Wallis and Futuna region: Alofi; 455 m. Vanuatu region: Tanna; 433-450 m. Elsewhere:
northern Ryukyu Islands to Rowley Shoals. Western Australia; 287-640 m (Cairns & Zibrowius. 1997;
Cairns, 1998).
Fungiacyathus (B.) variegatus Cairns, 1989
Fig. 2 d
Fungiacyathus fragilis - Wells, v.1984: 205-206 (in part: USGS 24918. pi. I, figs 1-2) [Not F.fragilis Sars, 1872].
Fungiacyathus variegatus Cairns. *19898: 11-12. pi. 5. figs a-h; 1994: 38-39. pi. 15, figs a-b. — Cairns & Zibrowius.
1997: 71-72.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 963. 1 cemented to Xenophora shell (MNHN).
Type Locality. — " Albatross " stn 5113: 13°52'N, 120°5LE (Verde Is. Passage, Luzon, Philippines), 291 m.
Remarks. — This species is more fully described by Cairns (1989a) and Cairns & Zibrowius ( 1997).
Distribution. — Vanuatu region: Anatom; 400-440 m (Late Pleistocene of Espiritu Santo. Wells. 1984).
Elsewhere: Japan through Indonesia; 84-715 m (Cairns & Zibrowius. 1997).
Source : MNHN. Paris
AZOOXANTHELLATE SCLERACTINIA
59
Family MICRABACIIDAE Vaughan, 1905
Genus LETEPSAMMIA Yabe & Eguchi, 1932
Letepsammia franki Owens, 1994
Letepsammia frcinki Owens, v*1994: 586-589, figs 1-2.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 597. 1 (MNHN).
Vanuatu. MUSORSTOM 8: stn 962, 1 (MNHN). — Stn 980. 3 (MNHN). — Stn 1016. 2 (MNHN). — Stn 1017.
8 (MNHN). — Stn 1018. 18 (USNM 98519). — Stn 1023, 1 (USNM 98518). — Stn 1070. 2 (MNHN). — Stn 1134.
1 (USNM 98517).
Type LOCALITY. — "Anton Bruun " stn 390-S: 29°35'S. 31°42'E (off Durban, South Alrica), 138 m.
Remarks. — In describing the type material. Owens (1994) stated that the septa were highly perforate, which
led to an assignment of this species to Letepsammia . However, re-examination of the type series shows that,
whereas the higher cycle septa are always highly perforate, the Si of the holotype and most paratypes arc
imperforate. (The Si of some paratypes from the same lots arc highly perforate). The same is the case for the
southwest Pacific specimens reported above. The imperforate nature of the Si suggests an affinity with
Rhombopsammia . and the variation in the expression of this character casts doubt on the distinction between the
genus Rhombopsammia and Letepsammia.
The specimens reported above are very similar to the type series, except that many of the specimens contain
108 septa (18 septa per system), instead of 120 septa (20 septa per system). The largest paratype (USNM 75640)
measures 32.2 mm in CD (slightly larger than that reported by Owens, 1994), whereas the largest Pacific
(MUSORSTOM 8 stn 1018) specimen measures 32.4 mm in CD.
Letepsammia franki differs from L. formosissima in having a papillose columella, coarser septal dentition, and
a more robust corallum. It differs from L. superstes in having more septa; a larger corallum; and nondentate.
vertical axial edges of the Si.
Distribution. — Wallis and Futuna region: Field Bank; 469-475 m. Vanuatu region: Anatom. Tanna, Elate,
and Espiritu Santo; 190-433. Elsewhere: southwest Indian Ocean; 50-650 m.
Genus STEPHANOPHYLLIA Michelin. 1841
Stephanophyllia neglecta Boschma, 1923
Step ha nop hy Ilia negl ecta Boschma, v*1923: 144-145. pi. 10. figs 28-30. - Cairns. 1989a: 23-24. pi. II, figs c-j
(synonymy). — Cairns & Zibrowius, 1997: 77.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 542, 1 (MNHN). Stn 556,
5 (MNHN). — Stn 618. 1 (USNM 98523). „ „ WKU1KIV
Vanuatu MUSORSTOM 8: stn 958. 1 (MNHN). — Stn 963, I (MNHN). - Sin 965. 7 (MNHN). — Stn 969. 2 (MNHN).
— Stn 978. I (MNHN). — Stn 988. 3 (USNM 98522). — Stn 1019. 1 (USNM 98521),
Type Locality. — "Siboga" stn 260: 5°36.5'S, I32°55.2'E (Kai Islands, Banda Sea). 90 m.
Remarks. — This species was rcdescribed and compared to all congencrics by Cairns (1989a).
Distribution. — Wallis and Futuna region: Aloft; Combe and Tuscarora Banks; 370-440 m. Vanuatu region.
Anatom, Tanna, and Espiritu Santo; 280-497 m. Elsewhere: Philippines; Indonesia; 49-555 m (Cairns &
Zibrowius, 1997).
Source : MNHN , Paris
60
S. D. CAIRNS
Stephanophyllia complicata Moseley, 1876
Stephanophyllia complicate Moseley. v*1876: 558-561. text-fig.; v. 1 88 1 : 198-201, pi. 4, fig. 12. pi. 13. figs 3-5. —
Cairns. 1989a: 21. pi. 12, figs a-b; 1995: 37-38. pi. 3, fig. h, pi. 4, figs a-e. map 10 (synonymy). — Cairns &
Zibrowius, 1997: 77-78.
Stephanophyllia japonica - WELLS, v.1984: 207. pi. 1. figs 5-6 (USGS stn 24918. USNM 71839) [Not S. japonic.a Yabe
& Eguchi. 1934a].
Material EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 523, 2 (MNHN). — Stn 535.
2 (MNHN). — Stn 540. 10 (USNM 98552). — Stn 557, 9 (USNM 98551).
Vanuatu. Musorstom 8: stn 959. 22 (USNM 98553). — Stn 963, 3, including 1 cemented to Xenophora shell
(MNHN). — Stn 1005, 1 (MNHN). — Stn 1018, 1 (MNHN). — Stn 1091. 1 cemented to Xenophora shell (MNHN). —
Stn 1097. I cemented to Xenophora shell (MNHN). — Stn 1113, I (MNHN).
Type Locality. — •'Challenger stn 192: 5°42’S, 132°25*E (Kai Islands. Banda Sea), 236 m.
REMARKS. — Stephanophyllia complicata was redescribed and illustrated by Cairns (1989a. 1995). It is
distinguished from S. neglecta by having a thin, lamellar columella; a narrow marginal shelf; Si with vertical,
non-spinose axial edges: and in attaining a larger size.
Distribution. — Wallis and Futuna region: Wallis; Waterwitch, Combe, and Tuscarora Banks; 470-600 m.
Vanuatu region: Anatom. Erromango, Efate, and Espiritu Santo (also Pleistocene. WELLS, 1984); 288-700 m.
Elsewhere: Indo-West Pacific; 73-635 m (Cairns & Zibrowius, 1997).
Suborder FAVIINA
Superfamily FAVIOIDEA Gregory. 1900
Family OCULINIDAE Gray, 1847
Genus OCULINA Lamarck, 1816
Oculina virgosa Squires, 1958
Oculina virgosa Squires, v*1958: 39. pi. 5. figs 8-16, text-fig. II. — Cairns. 1995: 40. pi. 4. figs f. i. pi. 5. figs c-d.
color frontispiece (top left), color cover (synonymy and description), map 8.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 1077. 6 branches: 4 (MNHN), 2 (USNM 98550) —
Stn 1102, 1 colony (MNHN).
Type Locality. — Sandstone, Waitemata Group. The Funnel, Kaipara Harbour, Auckland. North Island
(Altonian, early Miocene).
Remarks. — This species was recently redcscribed and figured by Cairns (1995). The two lots reported above
differ slightly from New Zealand specimens in lacking Sa, all corallites having only 24 septa arranged in three
complete cycles, the New Zealand populations usually having 28 septa. This is the first report of this species
outside New Zealand waters.
Distribution. — Vanuatu region: southeast and northeast of Espiritu Santo; 180-210 m. Elsewhere-
northeastern North Island. New Zealand; 29-388 m (CAIRNS. 1995); carlv Miocene to early Pliocene. New Zealand
(Squires, 1958).
Source :
AZOOXANTHELLATE SCLERACTIN1A
61
Genus MADREPORA Linnaeus, 1758
Madrepora oculata Linnaeus, 1758
Figs 2 e-f
Madrepora oculata Linnaeus, *1758: 798. — ZlBROWius. 1974a: 762-766, pi. 2, figs 3-5 (synonymy). — Cairns, 1991a:
9-10, pi. 2, fig. j. pi. 3, figs a-d (synonymy); 1994: 18-19, pi. 3, figs f-h (synonymy); 1995: 41. pi. 5, figs e-f, pi.
6, figs a-b, map 2; 1998: 372-373, figs 1 f-i. — Cairns & Zibrowius. 1997: 79-80.
Lophohelia tenuis Moseley, *1881: 180-181, pi. 8. figs 1 1-14. — Bourne, 1903: 26.
Cyathohelia formosa Alcock, *1898: 26-21, pi. 3, figs 2. 2a.
Sclerohelia formosa - ALCOCK, 1902c: 36.
Madrepora kauaiensis Vaughan, v*1907: 81-83, pi. 8, figs 1-2.
Madrepora tenuis - FAUSTINO, 1927: 107-108, pi. 14, figs 2. 5.
Madrepora formosa - ZlBROWius, 1974b: 568-570, figs 6-9.
Material EXAMINED. — Wallis and Futuna region. Musorstom 7: forma tenuis: stn 502. 1 branch (MNHN).
— Stn 551, numerous branches (USNM 98545). — Stn 552, 19 branches (MNHN), SEM stub 874 (USNM 98548). -
Stn 598, 6 branches (MNHN). — Stn 621, 8 branches (MNHN). — Stn 623, 1 branch (USNM 98547). — Stn 637.
2 branches (MNHN); forma formosa: Stn 507, 3 branches (MNHN). — Stn 585, I branch (MNHN). — Stn 586. I branch
(USNM 98581). — Stn 601, 1 branch (MNHN). — Stn 606, 5 colonies and numerous branches (USNM 98583). —
Stn 609, 3 colonies (MNHN), SEM stub 875 (USNM 98580). — Stn 618, 3 branches (MNHN). — Stn 620, 5 branches
(USNM 98546).
Vanuatu. Musorstom 8: forma tenuis: stn 1028. 2 branches (98546). — Stn 1031. 1 branch (MNHN); forma
formosa: stn 1088, 1 branch (MNHN).
Type Locality. — Tyrrhenian Sea and off Sicily (depth not given).
REMARKS. — This widespread and variable species has been redescribed and discussed at great length
(Zibrowius, 1974a, b; Cairns, 1991a, 1994, 1995; Cairns & Zibrowius, 1997). Two forms of the species
occur in the material reported herein. One form is characterized by having rather large, uniplanar colonies with
nonanastomosing branches formed of regular, sympodially budded coral lites. The coenosteum is light beige, finely
granular, and longitudinally striate. Corallites are 3.0-3. 6 mm in CD, containing 2 or 3 cycles of slightly exsert
septa, the S3 often rudimentary or absent. The axial edges of S2 often bear laciniate paliform lobes. The fossa is
deep and the columella rudimentary. This form corresponds to MOSELEY'S (1881) Madrepora tennis ; probably
M. kauaiensis Vaughan, 1907; Madrepora oculata forma beta of Cairns (1991a); and most specimens reported by
Cairns & Zibrowius (1997) from the Philippines and Indonesia. It is herein reported as forma tenuis. The second
form is characterized by having smaller, uniplanar to bushy colonies. Colonies are formed by sympodially
branched corallites, but often result in anastomosing branches caused by a symbiosis with a eunicid polychaete
worm, which produces a gall tube along the larger branches. The coenosteum is usually white and finely granular.
Corallites are smaller (2. 2-2. 5 mm in CD), more closely spaced, containing 3 cycles of highly exsert septa, the Si
usually well developed. The axial edges of the S2 usually bear prominent paliform lobes. The fossa is shallow and
the columella well developed. This form was described as Cyathohelia formosa by ALCOCK (1898) and as the
"symbiotic form" by Cairns (1995). It is herein reported as forma formosa.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Combe, Field. Rotumah. and Aloft Banks;
350-1280 m. Vanuatu region: Efate and Espiritu Santo; 310-624 m. Both forms are equally distributed in these
regions. Elsewhere: cosmopolitan, except for off continental Antarctica; 15-1500 m (CAIRNS & ZlBROWius,
1997).
Madrepora minutiseptum Cairns & Zibrowius, 1997
Madrepora minutiseptum Cairns & Zibrowius. *1997: 82-84, figs 4a-d. 5a-b.
Source : MNHN, Paris
62
S. D. CAIRNS
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 513, 3 branches (MNIIN). — Stn 515,
3 colonies and several branch fragments (MNHN). — Stn 517, 4 colonies and many branch fragments (USNM 98549).
Ryukyu Islands. "Toyoshio-Maru" : stn 11. 2 colonies (USNM 98461).
Type Locality. — "Snellius 2" stn 4.196: 6°23’S, 120°26.5’E (southwest of Salayer, Flores Sea).
150- 200 m.
Remarks. — Nothing can be added to the description of this recently named species. It is distinctive within
the genus by having 24 rudimentary septa; small, infundibuliform corallites; and a low, papillose columella. All
colonies contain a polychaetc-induced tubular deformation that runs the length of all major branches.
DISTRIBUTION. — Wallis and Futuna region: Futuna; 233-260 m. Elsewhere: Ryukyu Islands (reported herein),
Taiwan. Indonesia; 150-302 m.
Madrepora porcellana (Moseley, 1881)
Neohelia pore ell ana Moseley. v*188I: 176-177, pi. 10, figs 7. 7a. — Pratt. 1900: 591-603, pis 62-63. — Hickson.
1903: 344.
Madrepora porcellana - Wells, v. 1984: 207, figs 1. 7-9.
Neohelia sp. cf. N. porcellana - Cairns & ZlBROWius. 1997: 84-85, figs 5c-e, g-h.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 499, I branch (MNHN). — Stn 500.
1 branch (MNHN). — Stn 502, 1 branch (MNHN). — Stn 508, 2 branches (MNHN). — Stn 509, 2 branches (MNHN). —
Stn 512, 4 branches (MNHN). — Stn 513. 5 branches (MNHN). — Stn 514. 1 colony (USNM 98569). — Sin 515
3 colonies (USNM 98575). — Stn 618. 1 branch (MNHN).
Vanuatu. Musorstom 8: stn 962. 4 colonies (USNM 98578). — Stn 963. 1 branch (MNHN). — Stn 964. 5 branches
(MNHN). — Stn 968. 1 colony (MNHN). — Stn 969. I colony (USNM 98576). — Sin 976. 1 branch (MNHN). —
Stn 988, 2 branches (MNHN). — Stn 1077, 2 branches (MNHN). — Stn 1084. 2 colonies (USNM 98577). — Stn 1089
1 branch (MNHN).
USGS: stn 25718. 6 fragments (USNM 73958). — Stn 25715, (USNM 71840). — Sin 24918 (USNM 71841) all
reported by Wells (1984).
Type Locality. Challenger" stn 177: 16°45'S, I68J07’E f Api (= Epi), Vanuatu Archipelago], I 15 m.
Remarks. — Without explanation. Wells (1984) placed this species in Madrepora, which automatically
synonymized the genus Neohelia with Madrepora , a view not adopted by Cairns & ZlBROWius (1997). The basic
dilference between the genera is that Neohelia is reputed to lack a columella, Madrepora to have one. However,
examination of many additional specimens of Neohelia shows that it usually does have a well-formed papillose
columella, even though some distal corallites sometimes lack one. Likewise, corallites o {Madrepora often have
well-developed columella, but sometimes lack them altogether. Because the presence or absence of a columella
does not appear to be a conservative generic character, even within species, I now agree with Wells (1984) in
considering Neohelia porcellana to belong to the genus Madrepora.
All of the specimens reported above are pentamerally symmetrical, having 20 septa. The hexamerally
symmetrical specimens having 24 septa reported by Cairns & Zibrowius (1997) from Indonesia are considered as
a form of M. porcellana that occurs to the west and usually in shallower water than the typical form.
This species was recently redcscribed by Cairns and ZlBROWius (1997) and more fully monographed by Pratt
(1900). A consistent characteristic of the species is a tubular cavity that runs the length of the larger branches,
which is lined by a parchment-like encrustation. Pratt (1900) suggested that the encrustation was secreted by the
coral, but Hickson ( 1903) implied that a polychaete was responsible. Polychaetes are often found in these tubes,
and it cannot be doubted that gorgonians or antipatharians often form the framework for the encrustation of
M. porcellana. However, the specimens reported herein also indicate that a thecate hydroid may be responsible for
the internal parchment-like lining ol the tubes, short branches of which project from the apertures of the lube
through the coenosteum. This complex symbiosis is a fascinating one that requires the examination of additional
well-preserved specimens.
Source :
AZOOXANTHELLATE SCLERACTINIA
63
DISTRIBUTION. — Wallis and Futuna region: Futuna and Alofi; 240-516 m. Vanuatu region: Anatom. Tanna.
Epi, Malakula, and Espiritu Santo (also Pleistocene: WELLS, 1984); 1 15-494 m. Elsewhere: Loyally Islands; New
Caledonia; Indonesia (as the hexameral form); 55-238 m.
Family ANTHEMIPHYLLIIDAE Vaughan, 1907
Genus ANTHEMIPHYLLIA Pourtales, 1878
Anthemiphyllia dentata (Alcock. 1902)
?Discotrochus investigatoris Alcock. *1893: 142, pi. 5, figs 5, 5a.
Discotrochus dentcitus Alcock, v* 1902a: 104; v. 1902c: 27. pi. 4. figs 26. 26a.
Not Anthemiphyllia dentata - Cairns, 1984: 1 1. pi. 1. figs F-G (= A. macrodentala sp. nov.).
Anthemiphyllia dentata - Zibrowius & GRYGIER. 1985: 137 (New Caledonia). — Cairns & Parker. 1992: 16-17 (in part:
only those specimens from Western Australia). — Cairns, 1994: 44, pi. 18, figs d-f (synonymy); 1995: 41-42 (in
part: not sins NZOI K842 and K872 (= A. pacifica) and C527 (= A. macrodentata ). pi. 6, figs c-g (synonymy); 1998:
374-375. — Cairns & Zibrowius. 1997: 86.
Deltocyathus andamanicus - Keller, 1982: 52 (in part: pi. 1 1=4]. figs 3-4, Dimitri Mendeleev stn 1411) [Not
D. andamanicus Alcock. 1898].
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 511, 1 (MNHN). — Stn 522.
3 (MNHN). — Stn 525. 1 (MNHN). — Stn 530, 6 (MNHN). — Stn 534. 2 (MNHN). — Stn 535, 12 (USNM 98565). —
Stn 537. 3 (MNHN). — Stn 538, 2 (MNHN). — Stn 540. I (MNHN). — Stn 542. 4 (MNHN). — Stn 546, I (MNHN). -
Stn 555. 1 (MNHN). — Stn 569, 3 (MNHN). — Stn 570. 6 (USNM 98564). — Stn 572, 2 (USNM 98563). — Stn 585,
3 (USNM 98562). — Stn 586, 1 (USNM 98566). — Stn 589, 2 (MNHN). — Stn 590. 4 (MNHN). — Stn 591. 4 (USNM
98574). — Stn 594, 3 (USNM 98561). — Stn 597, 1 (USNM 98560). — Stn 618. I (USNM 98567). — Stn 619.
1 (MNHN).
Vanuatu. MUSORSTOM 8: stn 959. 2 (MNHN). — Stn 967. 2 (MNHN). — Stn 969. 2 (MNHN). — Stn 977. 6 (MNHN).
— Stn 982. 1 (MNHN). — Stn 983, 1 (MNHN). — Stn 1016, 1 (MNHN).
Type Locality. — "Siboga" stns 95, 98. 100: Sulu Sea, 350-522 m.
Remarks. — The holotype of Discotrochus investigatoris Alcock, 1893 is probably a juvenile specimen of
A. dentata and thus would have nomenclatural priority. It has a CD of only 8 mm and 48 septa, which is
consistent with the number of septa present in an A. dentata at that size. However, the type, presumed to be
deposited at the Indian Museum, Calcutta, was not examined by the author. Until this verification can be made,
the later name of A. dentata is used.
Whereas some of the largest specimens have a full fifth cycle (96 septa), most adult coralla have 60-72 septa,
resulting from the insertion of one or two pairs of S.s in each system, respectively. The addition of S5 pairs
usually occurs in a fixed manner as follows. In most coralla a bilateral symmetry can be determined by drawing
a line through the slightly elongate or elliptical columella, dividing the corallum into two halves each containing
six half-systems (Fig. A). If the 12 half-systems are numbered clockwise, pairs of S5 first occur in half-systems
I, II. V, VIII, X, and XII. which results in a mirror image septal complement, but with two adjacent
half-systems with pairs of S5 (I and XII) and two other half-systems (V and VIII) separated by two half-systems
having no S5 at all. Note that the numbering of half-systems is somewhat arbitrary in that, if the corallum is
rotated 180°, the half-systems with pairs of S5 are then: II. IV. VI. VII. IX, and XI, but the configuration is
the same.
Anthemiphyllia dentata is more fully described by Cairns (1995) and compared to other congenerics in
Table 4. All specimens were unattached, except for one (MUSORSTOM 8 stn 977). which was firmly attached to a
rock. I (Cairns & Parker. 1992; Cairns, 1995) had previously identified all Anthemiphyllia with S5 as
A. dentata (synonymy), but it now appears that at least three other species are involved, which consistently differ
from one another in corallum shape, basal ornamentation, and septal architecture (Table 4).
Source : MNHN, Pahs
64
S. D. CAIRNS
Table 4. — Comparison of the Recent species and subspecies of the genus Anthemiphyllia.
Source : MNHN . Paris
AZOOXANTHELLATE SCLERACTINIA
65
DISTRIBUTION. — Wallis and Futuna region: Wallis. Futuna, and Alofi; Waterwitch. Combe, Tuscarora, and
Field Banks; 295-650 m. Vanuatu region: Anatom, Tanna, and Efate; 280-475 m. Elsewhere: widespread
throughout Indo-West Pacific; 50-570 m (Cairns & Zibrowius, 1997).
Ant hemiphy Ilia multidentata sp. nov.
Figs 3 a-b
Anthemiphyllia dentata - Wells, v.1958: 264, pi. 1, figs 8-1 1. — Cairns & Parker, 1992: 16-17. figs 4e-f (in part: all
but Western Australian specimen).
Material EXAMINED/TYPES. — Australia. Off Cronulla, NSW. depth unknown, holotype (USNM 83010).
"Nimbus": stn 12. I paratype (USNM 7861 1). — Stn 55, 2 paratypes (USNM 78609).
Banzare: stn 1 15, 3 paratypes (SAM H500-501).
" Kimbla stn K7/73-37, I paratype (NMV F57153).
TYPE Locality. — Off Cronulla, New South Wales, depth unknown.
Etymology. — The species name multidentata (Latin multi, many + dens , tooth), literally bearing many
teeth, refers to the numerous trabecular teeth on all septa.
Description. — Corallum unattached, with a flat to slightly bowl-shaped base. Holotype 25.3 mm in CD and
6.5 mm in height. Costae equal in width, rounded, and finely granular. Intercostal grooves shallow; however, in
most specimens there is a very narrow ridge that bifurcates the intercostal region. No evidence of transverse
division or attachment to substrate. Corallum white.
Septa hexamerally arranged in 5 cycles, the fifth cycle often missing several pairs of septa, c.g., the holotype
has only 88 septa. Septal formula: Si>S2>S3>S4»Ss regarding size and exsertness; Si-3 all extend to the col¬
umella, whereas the S4 do only as rudimentary structures. All septa low in profile, closely fitting contour of inner
wall. Si bear 20-23 tall, slender, similarly-sized trabecular spines, the innermost (axial) spines highly compressed
perpendicular to plane of septum. S2, S3, and S4 similar in shape to Si but progressively smaller and having more
slender trabecular spines. There are actually more spines on the S3 (22-24) than the S 1-2 because the spines on the
S3 are more slender and yet the width of the septum is about the same. S5 bear 10-13 extremely slender trabecular
spines, the septa becoming rudimentary near the columella. Fossa shallow, containing a papillose columella
consisting of numerous small papillae that are indistinguishable from the inner (axial) septal spines.
Remarks. — Anthemiphyllia multidentata is similar to A. dentata. and has been reported as such at least twice
(see synonymy). However. A. multidentata differs from that species in having a significantly different septal
architecture, consisting of more uniformly-sized trabecular spines per septum, and spines that are flattened in the
plane perpendicular to the septum (Table 4). Thus far, A. multidentata is known only from southeastern Australia
at relatively shallow depths. Although not found in the study region of this report, it is included to make the name
available, and to facilitate comparisons among all Recent species (Table 4).
Distribution. — Southeastern Australia from northeastern Tasmania to New South Wales; 128-270 m.
Anthemiphyllia pacifica Vaughan, 1907
Figs 2 g-h
Anthemiphyllia pacifica Vaughan, v*1907: 79-80. pi. 7. fig. 5. — Cairns, 1984: 10-1 1.
Anthemiphyllia dentata - Cairns, 1995: 41-42 (in part: NZOI sins K842 and K872).
Material EXAMINED. — Vanuatu. Musorstom 8: stn 1031. 1 (MNHN).
Type Locality. — " Albatross " stn 3858: 21°ftl*25MN, 156°47'20”W (off Molokai, Hawaiian Islands),
225-252 m.
66
S. D. CAIRNS
Remarks. — Both Vaughan (1907) and Cairns (1984) noted that this species occurred in the free and
attached (pedicellate) forms, the types being free. The MUSORSTOM specimen reported above belongs to the
pedicellate form and is the largest specimen known, measuring 10.5 mm in CD and 7.3 mm in height, with
a basal fracture of 5.2 mm. It bears well-developed epithecal bands, characteristic of the species, and perhaps
because of its large size, contains 56 septa (4 pairs of S4). It is otherwise similar to typical Hawaiian specimens
and is compared to other congenerics in Table 4. One of the pedicellate species reported as A. dentcitci by CAIRNS
(1995) is reidenlified as A. pacifica herein.
DISTRIBUTION. — Vanuatu region: Efate; 310 m. Elsewhere: Hawaiian Islands; Kcrmadec Islands (reported
herein); 205-325 m.
Anthemiphyllia macrolobata sp. nov.
Figs 3 c-d
Anthemiphyllia dentata - Cairns. 1984: 1 1. pi. I, figs F-G: 1995: 41-42 (in part: N’ZOI stn C527).
MATERIAL EXAMINED. — Haiwaiian Islands. " Townsend Cromwell stn 81-01-14, holotype (USNM 60559).
Kermadec Ridge. NZOI: stn C527. 9 paratypes (USNM 93990).
Type Locality. — "Townsend Cromwell" stn 81-01-14: 23°15'48"N. I61°50'12"W (Hawaiian Is.), 369 m.
ETYMOLOGY. — The species name macrolobata (Greek makros. long or large + lobos, lobe), literally bearing
large lobes, refers to the large septal lobes on the S1-3.
Description. — Holotype shaped as a deep bowl, unattached, measuring 16.9 mm in CD and 10.0 mm in
height. Costae well developed, rounded, and finely granular, separated by deep intercostal grooves only at calicular
edge. No evidence of transverse division; however, the Kermadec paratypes are firmly attached to the substrate.
Corallum white.
Septa hexamerally arranged in 5 cycles, the fifth incomplete, only 1 pair of S5 occurring in each system,
resulting in 60 septa. Each Si consists of a highly exsert (3.3 mm), very wide (up to 5 mm) peripheral septal
lobe, which occasionally bears shallow indentations, but is not completely subdivided into smaller lobes.
Proximal to this lobe are 1-3 (depending on size of peripheral lobe) smaller, blunt lobes that are cylindrical in
cross section. Most proximal (axial) are 3 or 4 narrow septal spines that are flattened perpendicular to septal plane.
S2 similar in shape to Si. but slightly less exsert, having smaller peripheral lobes. S3 and S4 that are Hanked by
S5 are equally as wide as S2, but slightly less exsert and have smaller septal lobes. Ss and unllanked S4 are
smallest septa, only about half the width of an S3, bearing 10-12 narrow trabecular teeth. All septa bear relatively
low granules on their faces. Fossa deep, containing a well-developed papillose columella composed of elements
that are indistinguishable from the inner axial septal spines.
REMARKS. — As well as being reported as A. dentata twice (see synonymy), this species has been referred to
as an "undescribed species" by Cairns & Parker (1992), Cairns (1995), and Cairns & Zibrowius (1997).
Although not yet found in the study region, this species is included to make the name available, and to facilitate
comparisons among the Recent species (Table 4). A. macrolobata is unique in having large peripheral Si -3
septal lobes.
Distribution. — Hawaiian Islands; Kermadec Ridge; 369-508 m.
Anthemiphyllia patera costata subsp. nov.
Figs 3 e-h, 4 a-b
Material EXAMINED/TYPES. — Wallis and Futuna region. MUSORSTOM 7: stn 522. 1 paratype (MNHN). —
Stn 530, 5 paratypes (USNM 98558). — Stn 535. 2 paratypes (MNHN). — Stn 540. 2 paratypes (MNHN). — Stn 542,
1 paratype (MNHN). — Stn 546, 1 paratype (USNM 98557). — Stn 575, 19 paratypes (USNM 98554). — Stn 578.
Source :
AZOOXANTHELLATE SCLERACTINIA
67
2 paratypes (MNHN). — Stn 586. holotype (MNHN). and 7 paratypcs (MNHN). — Stn 590. 5 paratypes (MNHN). —
Stn 591. I paratype (MNHN). — Stn 594. 8 paratypes (MNHN), SEM stub 871 (USNM 98555). — Stn 595. 7 paratypes
(USNM 98556). — Stn 635, 5 paratypes (USNM 98559).
Type Locality. — Musorstom 7, stn 586: 13°10.7'S. 176013.1'W (north of Wallis), 510-600 m.
ETYMOLOGY. — The subspecies name costata (Latin costa, rib) refers to the presence of costae on the theca of
this species.
DESCRIPTION. — Corallum shaped as a shallow bowl, with rounded, slightly upturned edges. Holotype
8.3 mm in CD and 3.9 mm in height; largest specimen (MUSORSTOM 7 stn 594) 9.3 mm in CD. Base covered by
equal, rounded, finely granulated costae. Intercostal regions shallow and narrow on base, but deeply incised at
calicular edge, as in a lurbinoliid. No evidence of a basal scar or propagation by transverse division. Corallum
uniformly white.
Septa hexamerally arranged in 4 complete cycles (48 septa) according to the formula: S|>S2-3»S4. Si bear 7
or 8 narrow trabecular spines, the outermost peripheral spines quite small and directed outward, the fourth and fifth
from the outside usually the tallest, and the innermost (axial) 2 or 3 inclined toward and over the columella, these
spines highly compressed perpendicular to plane of septum. Si extend approximately I mm into central columella.
S2-3 only slightly less wide than the Si. usually bearing one less trabecular spine. S4 much smaller, about one-
third width of S 1.3, but equally as exsert, consisting of 2 or 3 slender trabecular spines, and a laciniate axial edge
that follows the contour of the inner theca. Fossa shallow, containing a massive (up to 4 mm in diameter),
irregularly circular, Hat columella composed of irregularly-shaped, fused elements. The columella protrudes into
each of the 6 systems, fusing with and uniting the axial edges of the S2 and their flanking S3, increasing the
columcllar diameter to over 5 mm at these points. Edges of columella undercut, being wider at the top than
the base.
REMARKS. — This subspecies is very similar to the nominate subspecies A. patera patera Pourtalcs, 1878.
both taxa having the same septal and columcllar structure and corallum shape. A patera costata differs primarily
in having a costate base (that of A. patera patera is smooth and porcellaneous), and in not attaining quite as
large a size (Table 4). A. patera patera is also known to have attached as well as free coralla. The nominate
subspecies is known only from tropical northwestern Atlantic at depths of 500-700 m (Cairns, 1979). Because
of these slight, but consistent morphological differences and the apparent geographic separation, a new subspecies
is warranted.
Anthemiphyllia patera costata is compared to other Recent congeners in Table 4, but it also bears a
resemblance to the fossil species A. catinata Wells, 1977 (Late Eocene. Tonga), as noted by WELLS (1977). Both
species are similar in corallum size and shape, costal ornamentation, and columcllar structure, but A. patera costata
differs in having more septa (48 vs 32-40). Details of septal dentition are similar, but not possible to compare due
to the poor preservation of the four known specimens of the fossil species.
Distribution. — Wallis and Futuna region: Wallis; Waterwitch, Combe. Field, and Rotumah Banks;
320-700 m.
Anthemiphyllia spinifera sp. nov.
Figs A. 4 c-j.
Discotrochus sp. Alcock, v. 1 902c: 27-28.
Deltocyathus andamanicus - KELLER, 1982: 52 (in part: pi. 1 |= 4], figs 5a-b).
Material EXAMINED/TYPES. — Wallis and Futuna region. MUSORSTOM 7: stn 512. I paratype (USNM
98571). — Sin 513, 2 paratypes (MNHN). — Stn 514, 13 paratypes: 12 (MNHN), I (USNM 98573). — Sin 530.
2 paratypes (MNHN). — Stn 537. 2 paratypes (USNM 98572). — Sin 541, I paratype (MNHN). — Stn 542, 4 paratypes
(USNM 98569). — Stn 569. 3 paratypes (MNHN). — Stn 586. 1 paratype (MNHN). — Stn 589. 1 paratype (MNHN). —
Stn 605, holotype (MNHN) and 18 paratypes (MNHN). — Stn 610. 2 paratypes, including SEM stub 872 (USNM 98570).
68
S. I). CAIRNS
Vanuatu. MUSORSTOM 8: stn 988, 1 paratype (MNHN). — Sm 1014, 1 paratype (MNHN).
Indonesia (Moluccas). KARUBAR: stn 18. 6 paralypes (USNM 96777).
" Albatross sm 5584, 4°I7’40"N, 1 18057’42"E. 532 m. 27.09.1909, 1 paratype (USNM 87605).
Type Locality. — Musorstom 7 stn 605: 13°21.3'S, 176°08.4'W (southeast of Wallis), 335-340 m.
Etymology. — The species name spinfera (Latin spinci, thorn + /<?/*, to bear) refers to the 6 costal spines.
Description. — Corallum (anthocyathus) discoidal, with a flat base, vertical edges, and circular calice.
Holotype 4.76 mm in CD and 2.34 mm in height; largest specimen (Musorstom 7 stn 513) 6.2 mm in CD and
tallest (Musorstom 7 stn 605) 3.8 mm in height. Base smooth, porcellaneous, and white, usually with a small
(2.3 mm in diameter), slightly concave scar or repaired scar of attachment in centre. Six elongate, tapered, smooth
costal spines (C3) project from the calicular edge in a specific pattern (see Remarks). Spines straight, or sometimes
curved or bent, and often broken, the longest spine 3.7 mm. Well-preserved coralla reddish brown at calicular edge,
otherwise white. One anthocaulus is known (Musorstom 8 stn 1014, Fig. 4j), consisting of a nondescript,
faintly costatc, cylindrical corallum 2.3 mm in height and 4.2 mm in diameter, bearing an anthocyathus 4.8 mm
in CD. There is an annular constriction between the anthocaulus and anthocyathus, but at this stage they are still
firmly attached. The attached anthocyathus bears 36 septa but has no evidence of costal spines.
Septal hexamerally arranged in 3 1/2 cycles, each system containing one pair of S4 inserted in a very spe¬
cific pattern (see Remarks), resulting in 36 septa. Only 2 size classes of septa exist: the larger septa, consisting of
the 12 S 1 -2 and the 6 S3 that are Hanked by S4 ( 1 8 of the 36 septa), and the smaller septa, consisting of all 12 S4
and the remaining 6 S3 that are unflanked by S4 (the remaining 18 septa), which alternate with the larger septa.
The larger septa extend to the columella, and bear of 4 or 5 equally sized, prominent, highly granular trabecular
spines, the innermost spines only slightly compressed in the plane of the septum. Innermost 2 trabecular spines of
larger septa inclined toward columella more than peripheral spines, the latter being vertical, which produces a small
notch in each septum, and altogether, a narrow circumferential demarcation about halfway between the columella
and calicular edge. Although the S 1.3 are of the same size, the S2 and accelerated S3 can be distinguished because
the columella extends slightly outward in each system to fuse these inner edges of the S2 and its adjacent
accelerated S3, as in A. patera costata. Smaller septa only about one-third width of the larger septa, bearing
3 slender trabecular teeth. Fossa shallow, containing a low. massive (up to 2 mm in diameter) papillose
columella, the elements of which are often fused. Columellar elements usually easily distinguished from inner
septal trabeculae.
Remarks. — Most specimens reported above bear six elongate costal spines (C3) associated with the 6 S3 that
are flanked by pairs of S4. one of which occurs in each half-system. Because the columella is circular, not elongate
as in A. dentata , an obvious plane of symmetry is not available for the numbering of half-systems. But, if the
12 half-systems are arbitrarily numbered in a clockwise direction beginning from the Si that occurs between the
two most closely spaced spines (Fig. A), then the costal spines invariably occur in half-systems I, III. V. VIII, X.
and XII, which are the same half-systems in which S5 pairs first occur in A. dentata . This results in two closely
spaced spines (half-systems XII and I) separated from one another by only 3 septa, or an angular separation of 40°;
and four spines (half-systems III. V, VIII. and X) separated from their adjacent dorsal spines by 5 septa, or 60°; and
the spines of half-systems V and VIII separated from each other by 80° (Fig. A). There are only three anomalies to
this pattern in the material examined: a specimen from 'Albatross" 5584 has 8 spines and thus 40 septa, the
additional costal spines occurring in half-systems IV and IX; a specimen from Musorstom 7 stn 542 has 7 spines
and 38 septa, the additional spine in half-system V; and the specimen reported by KELLER (1982), which appears to
have a seventh spine in half-system VIII.
Anthemiphyllia spin fera is easily distinguished from congeners by having costal spines. Other differences are
noted in Table 4.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch, Combe, and Field Banks; 245-
510 m. Vanuatu region: Tanna and Efate; 466-495 m. Elsewhere: Indonesia (Kai Islands, Banda Sea; south of
Sumba); Malaysia (Darvel Bay); Celebes Sea (off Mindanao); 212-534 m.
Source :
AZOOXANTHELLATE SCLERACTINIA
69
Fig. A. — Diagrammatic representation of the calice of Anthemiphyllia spinifera or Dehocyaihus heteroclitus, showing
the insertion pattern of the six costal spines (C3). Roman numerals define the 12 half-systems in the corallum.
arbitrarily beginning with the Si that bifurcates the smallest angular separation of costal spines, and proceeding in a
clockwise direction. Arabic numerals indicate the respective septal cycles. Length of costal spines and septa drawn in
proportion to calicular diameter of A. spinifera.
Suborder CARYOPHYLLIINA
Superfamily CARYOPHYLLIOIDEA Dana, 1846
Family CARYOPHYLLI1DAE Dana. 1846
Genus CARYOPHYLLIA Lamarck, 1816
Subgenus CARYOPHYLLIA (CARYOPHYLLIA) Lamarck. 1816
Caryophyllia (C.) hawaiiensis Vaughan, 1907
Caryophyllia hawaiiensis Vaughan, v*1907: 76. pi. 5. figs 4a-b. — Cairns, 1984: 11; 1995: 44-45. pi. 7. figs d-f,
map 18. — Cairns & Zibrowius. 1997: 93.
Source : MNHN. Paris
70
S. D. CAIRNS
Material EXAMINED — Wallis and Futuna region. MUSORSTOM 7: sin 504. I (MNHN). — Sin 538.
I (MNHN).
Vanuatu. MUSORSTOM 8: stn 969. 1 (USNM 98619).
TYPE LOCALITY. — " Albatross " stn 3838: 21°04'05"N. 157°10'35,,W (Molokai. Hawaiian islands), 1 68-
388 m.
Remarks. — Of the approximately 56 Recent species of Caryophyllia, only three have pentameral septal
symmetry: C. hawaiiensis , C. paucipalata Moseley. 1881, and C. crosnieri . the first species primarily pentameral,
the second primarily hexameral (the pentameral condition probably being aberrant, see Cairns. 1979). and the last
a mixture of pentameral and hexameral (see below). C. paucipalata is further distinguished from C. hawaiiensis by
having pali only before its antepenultimate cycle. C. crosnieri differs in having a denser, more robust, and
differently pigmented corallum; less exsert septa; and a deeper fossa. C. hawaiiensis was redescribed and illustrated
in detail by CAIRNS (1995).
Distribution. — Wallis and Futuna region: Futuna Island; Waterwitch Bank; 295-300 m. Vanuatu region:
Anatom; 252-280 m. Elsewhere: Hawaiian Islands; Japan; South China Sea; Philippines; Indonesia; Kcrmadec
Ridge; 85-279 m (Cairns & ZiBROWiUS, 1997).
Caryophyllia (C.) crosnieri Cairns & Zibrowius. 1997
Figs 5 a-b
Caryophyllia elongata Cairns in Cairns & KELLER. *1993: 236-237. pi. 4. figs A-B [Not Caryophyllia clavus var.
elongata Duncan. *1873: 311-312 (= C. smithii Smith & Broderip. *1828)]. — Cairns, 1995: 52, pi. 10. figs d-f,
map 14.
Caryophyllia crosnieri Cairns & Zibrowius, *1997: 89 (nom. nov.).
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 540, 1 (MNHN). — Stn 572. I (USNM
98623). — Stn 581. 1 (MNHN).
Vanuatu. MUSORSTOM 8: stn 973, 1 (MNHN). — Stn 974, 3 (MNHN). — Stn 975, 1 (MNHN). — Stn 982, 2 (USNM
98621). — Stn 1009. I (USNM 98620). — Stn 1067. 1 (MNHN). — Stn 1074, I (USNM 98622). — Stn 1108.
1 (MNHN).
Type Locality. — "Vityaz " stn 2716: 33°I7S, 44°55’E (Walters Shoal, Madagascar Plateau), 630-680 m.
Remarks. — Additional large, well-preserved specimens reported herein allow the observation that this species
is not consistently hexamerally symmetrical. Whereas most previously reported specimens were hexamerally
symmetrical (S i>S2>S4^S3. 48 septa and 12 pali), most of the specimens listed above are pentamerally
symmetrical (5:5:10:20, 40 septa and 10 pali), only the 3 specimens from MUSORSTOM 8 sins 975. 1067. and
1074 being hexamerally symmetrical (S|>S2>S4>S3, 12 pali), and the specimen from MUSORSTOM 8 stn 974
having 1 1 pali (44 septa). Furthermore, a lot reported by Cairns & Zibrowius (1997), e.g., Karubar stn 31,
contains a mixture of specimens with pentameral or hexameral symmetry.
The species may be diagnosed as having: a robust, cylindrical corallum with a broadly encrusting base;
noncostate, coarsely granular, yellowish-brown theca; pentameral or hexameral septal symmetry, usually resulting
in 40 or 48 septa, respectively; highest cycle septa equal to or wider than penultimate septal cycle; carinate septal
faces; and a very deep fossa with a well-defined columella of 3-9 slender twisted elements. The axial palar edges are
highly sinuous, and, in some specimens, form a narrow lamella on their axial margin oriented perpendicular to the
plane of the palus.
Distribution. — Wallis and Futuna region: Waterwitch and Wallis; Combe Bank; 550-600 m. Vanuatu
region: Tanna, Efate, Malakula, and Espiritu Santo; 366-536 m. Elsewhere: southwest Indian Ocean; Philippines;
Indonesia; Kermadec and Three Kings Ridges; 165-680 m (Cairns & Zibrowius. 1997).
Source :
AZOOXANTHELLATE SCLERACTINIA
71
Cary ophy Ilia (C.) rugosa Moseley. 1881
Cary ophy Ilia rugosa Moseley, v*I88l: 141-143, pi. 1, figs 8a-b. — Wells, v. 1954: 469, pi. 177, figs 5-6. —
CAIRNS, 1984: 11-12, pi. 2, figs A-B, pi. 4. fig. I; 1994: 47, pi. 20, fig. i, pi. 21. fig. a (synonymy); 1995: 43*44. pi. 6,
fig. h, pi. 7. figs a-c, map 16; 1998: 375. — Cairns & Zibrowius. 1997: 91-92.
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom 7: sin 530, I (MNHN) — Sm 610
2 (MNHN).
Vanuatu. Musorstom 8: sin 967, I (USNM 98618). — Sin 988. 1 (98617). — Sm 1095. I (USNM 98616) —
Sm 1097, 1 (MNHN).
Type Locality. — "Challenger" sins 192 and 201: Banda and Sulu Seas, 187-230 m.
Remarks. This widely distributed and frequently collected deep-water species can be characterised as having:
a small, cylindrical corallum with a transversely ridged theca; 32 octamerally arranged septa (Si>S2>Si) and
8 pali; and all septa and pali with extremely sinuous axial edges.
Distribution. — Wallis and Futuna region: Wallis; Waterwitch Bank; 286-580 m. Vanuatu region: Anatom,
Tanna, and Espiritu Santo; 288-372 m. Elsewhere: Indo-West Pacific from southwest Indian Ocean to Hawaiian
Islands; 71-581 m (Cairns & Zibrowius, 1997).
Caryophyllia (C.) octonaria Cairns & Zibrowius, 1997
Caryophyllia octonaria Cairns & Zibrowius, *1997: 92, figs 7a-b.
Material examined. — Vanuatu. Musorstom 8: stn 964, 5 (MNHN). — Stn 969, 6 (MNHN) — Sm 976
12 (USNM 98614). — Sm 1072, 2 (USNM 98615). — Stn I 134, I (MNHN).
Type Locality. — Musorstom I stn 64: 14°01’N, 120°16*E (Lubang Island, Philippines), 194-195 m.
Remarks. — Little can be added to the recent description of this species. It can be diagnosed as having:
a small, straight corallum with a well-defined, white pedicel, the upper theca being light brown; granular
costae sometimes covered with epitheca; a circular to only slightly elliptical calice; 32 octamerally arranged septa
(Si>S2-3) with only moderately sinuous axial edges; and eight very sinuous pali with carinate faces.
Distribution. — Vanuatu region: Anatom and Espiritu Santo; 182-622 m. Elsewhere: Philippines;
186-194 m.
Caryophyllia (C.) abrupta sp. nov.
Figs 5 d-e
MATERIAL EXAMINED/TYPES. — Wallis and Futuna region. Musorstom 7: stn 522. I paratype (MNHN). —
Stn 523, 8 paratypes (MNHN). — Sm 524, 2 paratypes (USNM 98607). — Stn 530, I paratype (MNHN). — Stn 534.
4 paratypes (MNHN). — Stn 535, holotype (MNHN), 14 paratypes (USNM 98609). — Stn 537, 2 paratypes (MNHN). —
Stn 546, 2 paratypes (MNHN). — Stn 557, 3 paratypes (USNM 98610). — Stn 569. 1 paratype (USNM 98611). —
Stn 570, I paratype (MNHN). — Sm 571. 2 paratypes (MNHN). — Stn 586, 4 paratypes (MNHN). — Stn 604,
5 paratypes (MNHN). — Stn 605, 2 paratypes (MNHN). — Stn 606, I paratype (MNHN). — Stn 608, 2 paratypes (USNM
98606). — Stn 619, 2 paratypes (USNM 98608).
Vanuatu. Musorstom 8: sin 963, 1 nontype cemented to a Xenophora shell (MNHN). — Stn 988, 1 (MNHN). —
Stn 1016, 1 paratype (USNM 98612), and another nontype cemented to a Xenophora shell (MNHN). — Stn 1065,
1 paratype (MNHN). — Stn 1088, 2 nontypes cemented to a Xenophora shell (MNHN). — Stn 1091, 2 nontypes
cemented to Xenophora shells (MNHN). — Stn 1094, 1 paratype (MNHN).
Source : MNHN, Pans
72
S. D CAIRNS
Type Locality. — Musorstom 7 stn 535: I2°29.6'S, 176°41.3'W (Waterwitch Bank), 340-470 m.
Etymology. — The species name abrupta (Latin abruptus , broken off, separated, sheer) refers to the process
of transverse division that separates the anthocyathus from the anthocaulus.
Description. — Anthocaulus stage unknown. Anthocyathus ceratoid. curved between 45°-90° usually in plane
of GCD: one to 7 episodes of rejuvenescence not uncommon in larger coralla. Holotype 8.4 x 7.5 mm in CD and
18.5 mm in height; largest corallum (MUSORSTOM 7 stn 619) 10.9 x 8.4 mm in CD. Basal scar circular to
slightly elliptical: 1. 8-2.1 x 1 .7-1.8 mm in diameter, displaying 8 major septa. Calice strongly compressed:
GCD.LCD = 1.1 2-/. 29- 1.37 (N=8). Costal expression variable, but 8 primary costae usually prominent, the other
24 flat and poorly defined; however, in some specimens the theca is smooth or bears uniformly small granules.
Most coralla uniformly white, but some are reddish brown near calicular edge.
Septa usually octamerally arranged in 3 size classes: 8:8: 16 (32 septa); however, two specimens of 6.6 mm CD
from Musorstom 7 stn 535 have a septal complement of 6:6:12 (6 pali) and 7:7:14 (7 pali), and the largest
specimen has an extra pair of septa resulting in 34 septa and 9 pali. Eight primary septa highly exsert (1.5-
1.6 mm), extend 3/4 distance to columella, having moderately sinuous axial edges. Secondary septa about half
width and exsertness of primaries, but have extremely sinuous axial edges. Tertiaries slightly more exsert than
secondaries, being fused to their adjacent primary septa to form low. triangular lancets. Tertiaries equal to or
slightly less wide than secondaries, becoming relatively wider with age, having slightly sinuous axial edges. Fossa
of moderate depth, containing 8 narrow P2 that form a distinct palar crown, each palus having extremely sinuous
edges, equivalent to those of the secondary septa they border. Columella fascicular, composed of 2-4 narrow
elements arranged linearly or in a rhomboid pattern.
Remarks. — There are six species of octamerally symmetrical Caryophyllia , all of which generally have
32 septa: C. rugosci Moseley. 1881; C. octopali Vaughan. 1907; C. nmbahithi Gardiner & Waugh, 1938;
C. barbadensis Cairns, 1979; C. marmorea Cairns, 1984; and C. octonaria Cairns & Zibrowius, 1997; a seventh
species has a variable symmetry that includes octameral: C. conuiformis Pourtales, 1868. C. abrupta differs from
these species by propagating by transverse division, and by having a compressed corallum. It is perhaps most
similar to C. conuiformis (a species known only from the Atlantic and the southwest Indian Ocean) in size, shape,
and septal architecture. Both species also have an abruptly terminated base: in the case of C. abrupta. due to
transverse division, in the case of C. conuiformis, probably due to budding, which results in an irregularly-shaped,
open base. C. abrupta also diflers from that species in having a very regular octameral symmetry with a well-
del ined palar crown, an elongate calice, exsert primary and secondary septa, and often prominent primary costae.
One other species of this genus is known to have transverse division: C. secta Cairns & Zibrowius, 1997,
from the Philippines and Indonesia (220-366 m), which differs from C. abrupta in septal symmetry (hexameral,
48 septa), size (up to 15.7 mm GCD), and corallum shape (straight, trochoid).
Distribution. — Wallis and Futuna region: Wallis and Alofi; Waterwitch. Combe, and Tuscarora Banks;
305-650 m. Vanuatu region: Anatom. Tanna. Efate. Malakula, and Espiritu Santo; 300-400 m.
Caryophyllia (C.) marmorea Cairns, 1984
Figs 5 c. f
Caryophyllia marmorea Cairns, *1984: 13. pi. 2. figs C-D.
Material EXAMINED. — Musorstom 7: sin 525. 1 (MNHN). — Sin 534. 1 lUSNM 98613). — Stn 585. I (MNHN),
Type Locality. — Deep Ocean Disposal Site Investigations. Hawaii, Stn HON 9-3: I9°48'N. I54°58'W (off
Hilo, Hawaii ), 337 m.
Source :
AZOOXANTHELLATE SCLERACTINIA
73
Remarks. — This species is diagnosed as having a small (CD < 4 mm), straight, attached subcylindical
corallum. The theca is light brown, granular, and glisteny; primary costae are prominent. Septa are octamerally
arranged: S i >S2>S3 (32 septa). This is the first report of this species subsequent to its original description
from Hawaii.
Distribution. — Wallis and Futuna region: Wallis; Waterwitch Bank: 475-500 m. Elsewhere: Hawaii,
Hawaiian Islands; 331-337 m.
Caryophyllia (C.) quadragenaria Alcock, 1902
Cary ophy Ilia quadragenaria Alcock, v* 1902a: 91-92; v. 1902c: 10, pi. 1. figs 4, 4a. — Cairns. 1994: 46-47. pi. 20,
figs c-h. pi. 41, figs c-d (synonymy); 1995: 45-46. pi. 7, figs g-h, map 7 (synonymy); 1998: 375. — Cairns &
Zibrowius, 1997: 93.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 556. I (MNHN).
Vanuatu. MUSORSTOM 8: stn 962, 1 (MNHN). — Stn 965, 1 (MNHN). — Stn 967. 5 (USNM 98603). — Stn 1 106,
1 (MNHN).
Type LOCALITY. — "Siboga" stns 90, 251, and 289: Makassar Strait, Banda, and Timor Seas, 54-281 m.
REMARKS. — In addition to C. quadragenaria, there are six other decamerally symmetrical Caryophyllia:
C. abyssorum Duncan, 1873; C. calveri Duncan. 1873; C. antillarum Pourtales, 1874; C. zopyros Cairns. 1979;
C. perculta Cairns, 1991; and C. solida Cairns, 1991. C. quadragenaria can be diagnosed as having: a relatively
small (GCD < 12 mm), straight corallum attached by a narrow pedicel; poorly defined costae; three cycles of
decamerally arranged septa resulting in 40 septa, the tertiary septa usually equal to or wider than the secondaries;
and a well-formed palar crown of 10 pali. The species was recently redescribed and illustrated by CAIRNS (1994.
1995).
DISTRIBUTION. — Wallis and Futuna region: Tuscarora Bank; 440 m. Vanuatu region: Anatom. Efate. and
Espiritu Santo; 314-430 m. Elsewhere: western Pacific from Japan to New Zealand: 54-385 m (CAIRNS &
Zibrowius, 1997).
Caryophyllia (C.) sp. cf, C. calveri Duncan. 1873
Figs 5 g-i
? Caryophyllia calveri Duncan, *1873: 316. — Zibrowius, v.1980: 57-59, pi. 21, figs A-L.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1051, 1 (MNHN). — Stn 1056, 11:7 (MNHN), 4 (USNM
98605).
Type Locality of C. calveri. — " Porcupine stn 24, 37°I9'N. 9°I3'W (off Portugal), 534 m.
Description. — Corallum elongate-conical and straight, having a robust pedicel (PD:GCD = 0.32-0.62) that
is firmly attached by a thin, broadly encrusting base. All specimens examined attached to small bits of pumice.
Largest specimen (MUSORSTOM 8 stn 1056) 10.9 x 9.3 mm in CD. and 3.5 mm in PD. having a basal
encrustation of 16 mm. Calice slightly elliptical: GCD:LCD = 1.08-1.19. Costae low and poorly defined, the
intercostal striae being infilled with glisteny textura. Base, pedicel, and calicular elements white; however, upper
theca a light yellow-brown.
Septal symmetry variable. Half of the specimens examined have hexameral symmetry (Si-2>S4^S.l 48 septa
and 12 pali); 5 specimens have decam eral symmetry (10:10:20. 40 septa and 10 pali); and one specimen has
nonameral symmetry (11:1 1:22, 44 septa and 1 1 pali). The 10-12 larger septa (primaries) are highly exsert
Source : MNHN, Paris
74
S. D. CAIRNS
(up to 2.1 mm), extend about 3/4 distance to the columella, and have moderately sinuous axial edges. Second
size class of septa (secondaries) about 0.7 mm exsert, 4/5 width of a primary septum, also having moderately
sinuous axial edges. Third size class of septa (tertiary septa) equal to or slightly wider than the secondaries,
and about 0.9 mm exsert, fusing with their adjacent primary septa at the calicular edge to form low, triangular
lancets. Depending on septal symmetry, 10-12 wide (1.1 -1.2 mm) pali form a well-defined crown, the pali having
highly sinuous edges and bearing prominent granules and short ridges. Fossa of moderate depth, bearing a
fascicular columella composed of an elongate field of 3-10 slender (0.5 mm diameter), tightly twisted and closely
spaced elements.
Remarks. — Of the approximately 56 Recent valid species of Caryophyllici , nine species can be characterised
as having an attached corallum, decameral or hexameral symmetry, and its highest cycle septa equal to or wider
than its penultimate cycle: C. calveri Duncan, 1873; C. atlantica (Duncan. 1873); C. polygona Pourtales, 1878;
C. lame life ra Moseley, 1881; C. arnoldi Vaughan. 1900; C. panda Alcock. 1902; C. alberti Zibrowius, 1980;
C. quadragenaria Alcock, 1902; and C. perculta Cairns, 1991, the last two species usually having decameral
symmetry. The specimens reported above from Epi are morphologically indistinguishable from specimens
identified as C. calveri by Zibrowius (1980), even to the point of having a variable septal symmetry that includes
both decameral and hexameral. C. calveri is known only from the northeast Atlantic and Mediterranean at 1 30-
1050 m (Zibrowius, 1980). This disjunct distribution would imply that the Epi Island population represents
a different species or that intermediate populations between the south Pacific and northeast Atlantic have not yet
been recorded.
Distribution. — Vanuatu region: Epi; 558-602 m.
Caryophyllia (C.) diomedeae Mare n zeller, 1904
Caryophyllia diomedeae Marenzeller. v*1904: 79-80, pi. 1, fig. 2. — Cairns, 1995: 49-50, pi. 9, figs a-d, map 3
(synonymy). — Cairns & Zibrowius, 1997: 88.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 959, 1 (MNHN). — Stn 1014. I (USNM 98604). —
Stn 1080. 1 (MNHN). — Stn 1 128, 3 (MNHN).
Type Locality. — " Albatross " stn 3358: 6°30'N, 81°44*W (off Coiba Island, Panama), 1043 m.
Remarks. — This widespread species was redescribed and discussed by Cairns (1995). It can be characterised
as having: an elongate, slender but slightly flared, straight corallum attached by a narrow pedicel; fiat to
porcellaneous theca; only slightly exsert septa; a septal formula of Si-2>S3>S4 (48 septa); and a shallow fossa.
Distribution. — Vanuatu region: Anatom, Efate, and Malakula; Guyot Bougainville; 475-799 m. Elsewhere:
widespread, including Indo-Pacific and Atlantic Ocean; 225-2200 m (CAIRNS, 1995).
Caryophyllia (C.) lamellifera Moseley, 1881
Caryophyllia lamellifera Moseley. v*1881: 140-141, pi. 1, figs 7a-b. — Cairns. 1995: 51-52, pi. 9, fig. i. pi. 10.
figs a-c, map 18 (synonymy ). — Cairns & Zibrowius, 1997: 90.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 502, 1 (MNHN). — Stn 589.
2 (MNHN).
Vanuatu. MUSORSTOM 8: stn 1077, I (USNM 98591).
Type Locality. " Challenger " stn 170: 29°55’S, 178°14'W (north of Macauley Island. Kermadec Ridge),
1 152 m.
Source :
AZOOXANTHELLATE SCLERACTINIA
75
Remarks. — Caryophyllia lamellifera is distinguished from the many other Pacific Caryophyllia, by having a
fine, transversely ridged, brown-striped theca. The species is more fully described by Cairns (1995), who also
compared it to the other two congenerics that have transversely ridged theca: C. rugosa and C. corrugata. Cairns
(1995) noted variation in septal symmetry (12-14 primary septa) and relative width of the S4 and S3 (S4 being
wider, as in the holotype, or narrower than S3, as in most other specimens). The four specimens reported above all
have 48 septa arranged as: Si-2>S3>S4. However, three other specimens from Musorstom 8 stns 1018. 1023,
1091, and 1095 (not reported above), are only tentatively assigned to this species. They differ from the typical
form in lacking transverse thecal ridges and in having S4 that are equal to or slightly wider than their S3, as in the
holotype. It is possible that both of these characters may be found to constitute intraspecific variation for
some species.
Distribution. — Wallis and Futuna region: Futuna; Field Bank; 400-516 m. Vanuatu region: Malakula: 1 80-
210 m. Elsewhere: Philippines; Indonesia; Kermadcc and Norfolk Ridges; Taupo Tablcmount; 89-1 152 m (Cairns
& Zibrowius, 1997).
Caryophyllia (C.) scobinosa Alcock, 1902
Caryophyllia scobinosa Alcock, v*1902: 8, pi. 1, figs 2, 2a. — Cairns, 1995: 52-53, pi. 10, figs g-i. pi. 11, figs a-c.
map 16 (synonymy). — Cairns & Zibrowius. 1997: 94.
Not Caryophyllia scobinosa - WELLS, v.1984: 207-209. tigs 2 (1-4) [= Asterosmilia sp. cf. A. marchadi (Chevalier,
1966)].
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 552. I (MNHN). — Stn 565,
1 (MNHN). — Stn 635, 2 (USNM 98601).
Vanuatu. MUSORSTOM 8: stn 992, 2 (MNHN). — Sin 1034, II (MNHN). — Stn 1035, 7 (USNM 98602). —
Stn 1074, 2 (MNHN).
Type Locality. — " Siboga " stns 45 and 102: Flores and Sulu Seas, 535-794 m.
Remarks. — This species was redescribed and figured by Cairns (1995). It is distinguished from the other
unattached cornute species known from this region. C. ambrosia , by having a much smaller, and more slender
coral I um (GCD < 16 mm); and only 48 septa and 12 pali.
DISTRIBUTION. — Wallis and Futuna region: Combe, Tuscarora, and Rotumah Banks; 715-900 m. Vanuatu
region: Erromango, Efate, and Espiritu Santo; 750-775 m. Elsewhere: Indo-West Pacific from southwest Indian
Ocean to Japan and Tonga; 353-1276 m (Cairns & Zibrowius, 1997).
Caryophyllia (C.) ambrosia Alcock, 1898
Caryophyllia ambrosia Alcock, v*1898: 12, pi. 1, figs 1. la. — Zibrowius, v.1980: 63-65, pi. 25, figs A-K
(synonymy). — Cairns, 1994: 48, pi. 21. figs d-h; 1995: 53-54, pi. 11, figs d-e, map 7. — Cairns & Zibrowius,
1997: 95-96.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 564. 2 (MNHN). — Sin 565.
2 (MNHN). — Stn 598, 1 (USNM 98599). — Stn 635, 2 (USNM 98600). — Stn 636, 4 (MNHN).
Type Locality. — "Investigator0 stns 104 and 176: Laccadive Sea, Arabian Sea, 1829-1957 m.
Remarks. — The regional variation of corallum size and number of septa was discussed by Cairns (1994.
1995), who also provided descriptions and illustrations of the species. The specimens reported above have 14-22
(mode = 16) primary septa and 56-88 (mode = 64) total septa, which allies them most closely with northern Indian
Ocean populations. Populations to the north (Japan), west (Indonesia), and south (New Zealand), usually have
more septa, e.g., 1 8-30 primary septa or a total of 72-120 septa.
Source : MNHN, Paris
76
S. D. CAIRNS
Distribution. — Wallis and Futuna region: Tuscarora, Field, and Rotumah Banks; 700-1015 m. Else
where: Amphi-Atlantic; Indo-West Pacific, including Japan and New Zealand; 311-2670 m (Cairns &
Zi brow I us, 1997).
Subgenus CARYOPHYLLIA (ACANTHOCYATHUS) H. Milne Edwards & Haime, 1848
Caryophyllia (A.) grayi (H. Milne Edwards & Haime, 1848)
Acanthocyathus grayi H. Milne Edwards & Haime, * 1848a: 293. pi. 9, figs 2, 2a. — UMBGROVE, 1950: 641-642. pi. 81.
figs 27-32 (synonymy). — Wells, v.1984: 209. pi. 2. figs 5-9.
Caryophyllia (A.) grayi - Cairns, 1994: 49, pi. 21, figs i-k (synonymy); 1998: 377. — Cairns & ZlBROWius, 1997: 97-
98, figs 7 c, f, i.
Material EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 496. I (USNM 98598). — Stn 499,
1 (MNHN).
Vanuatu. MUSORSTOM 8: stn 1001, 1 (MNHN). — Stn 1002, 1 (USNM 98597). — Stn 1004, 2 (USNM 98596). —
Stn 1065, 1 (MNHN). — Stn 1069, I (MNHN). — Stn 1070, 2 (MNHN). — Stn 1071. 2 (MNHN). — Stn 1086,
4 (MNHN). — Stn 1 103, 3 (MNHN). — Stn 1 134. I (USNM 98595).
USGS {Late Pleistocene of Kere River. Espiritu Santo): stn 25715. 4 (USNM 71844. 71846. 78610), WELLS, 1984;
Stn SM242, 129a and 129b, 4 (USNM 71845 and 73963), Wells, 1984.
Type Locality. — Unknown.
Remarks. — The species is characterised as having a compressed, ceratoid corallum. with a narrow pedicel and
rounded thecal edges, each of which bears several thecal spines that are circular in cross section. Costae are rounded,
not ridged, and the theca is often reddish brown. Septa are usually arranged: 14:14:28, resulting in 56 septa and
14 pali, but large coralla have 16 and even 18 primary septa, resulting in up to 72 septa (CAIRNS, 1995). The
species was redescribcd and figured by Cairns (1995) and Cairns and ZIBROWIUS (1997), the latter including a
key and discussion of most of the species in this subgenus.
Distribution. — Wallis and Futuna region: Futuna; 290-300 m. Vanuatu region: Erromango, Malakula, and
Espiritu Santo (also Late Pleistocene, Wells. 1984); 125-360 m. Elsewhere: Indo-West Pacific from South Africa
to Japan; 37-490 m (Cairns & Zibrowius, 1997).
Genus CRISPA TOTROCHUS Tenison Woods, 1878
Crisp at ot ro c h u s ruhescens (Moseley, 1881)
Cyathoceras ruhescens Moseley. *1881: 157. pi. 2. figs 8a-c. — Cairns. 1984: 15.
Cyathoceras tydemani Alcock, v* 1902a: 93-94; v. 1902c: 14, pi. 1, figs 7. 7a.
Cyathoceras diomedeae Vaughan, v*1907: 77-78. pi. 7. figs 1-2.
C rispatotrochus ruhescens - Cairns, 1994: 51, pi. 22, figs g-h (synonymy). — Cairns & Zibrowius, 1997: 103-104.
figs 10 a-c.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 502. I (USNM 98585). — Stn 507
2 (MNHN). — Stn 51 1, 1 (MNHN). — Stn 523, 1 (MNHN). — Stn 585, 5 (MNHN). — Stn 604, 1 (MNHN).
Vanuatu. Musorstom 8: stn 983, 1 (USNM 98584).
Type Locality. — "Challenger" stn 192: 5°49’15"S, 132°I4T5"E (Kai Islands, Banda Sea), 236 m.
Remarks. — This species was recently redescribcd and figured by Cairns (1994).
Source :
AZOOXANTHELLATE SCLERACTINIA
77
DISTRIBUTION. — Wallis and Futuna; 420-516. Vanuatu region: Tanna; 475-480 m. Elsewhere: western and
central Pacific: Japan; South China Sea; Philippines; Indonesia; Hawaiian and Christmas Islands: 1 10-634 m
(Cairns & Zibrowius, 1997).
Crispatotrochus rugosus Cairns, 1995
Figs 6 a-b
Crispatotrochus rugosus Cairns. *1995: 57, pi. 13, figs a-b, map 16; 1998: 378. — Cairns & Zibrowius. 1997: 104.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 511, 1 (MNHN). — Stn 523
I (MNHN). — Stn 542, 1 (MNHN). — Sin 584, 2 (USNM 98586). — Sin 589. 1 (MNHN). — Stn 610. 1 (MNHN).
Vanuatu. Musorstom 8: stn 977. 3 (MNHN). — Stn 988. 8 (USNM 98589). — Stn 1015. 3 (USNM 98590). —
Stn 1018, 6 (USNM 98587). — Stn 1030, 2 (MNHN). — Sin 1043, I (MNHN). — Stn 1095, 2 (MNHN). — Stn 1106
1 (USNM 98588).
Type Locality. — NZOI stn Q70: 26059.7’S, 159°18.9’E (Lord Howe Seamount Chain). 376 m.
Remarks. — Crispatotrochus rugosus is distinguished from the other 10-12 species in the genus by having
line, transverse thecal ridges, and Si that are larger than the S2. All but one of the specimens reported above arc
consistent with the type series, having 48 septa arranged in 4 cycles. However, the specimen from MUSORSTOM 8
stn 1043 (Figs 6 a-b) is considerably larger (CD = 29.2 x 18.3 mm, height = 37.1 mm) and has a fifth cycle of
septa, although missing 4 pairs of S5 (88 septa). This large specimen is similar to the fragmented specimen
reported by Cairns and Zibrowius (1997) that had an estimated GCD of 32 mm.
Corallum pigmentation is variable, including uniformly white, uniformly brownish-black, and white with
brown longitudinal stripes corresponding to the C 1 -2 near the calicular edge. One specimen from Musorstom 8
stn 988 contains an acrothoracican cirripede boring in its pedicel.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Combe and Field Banks; 286-455 m.
Vanuatu region: Tanna, Efate, Epi, and Espiritu Santo; 190-410 m. Elsewhere: Kermadec Islands; Lord
Howe Seamount Chain; Philippines; Malaysia; Western Australia; 142-508 m (Cairns & Zibrowius, 1997).
Genus LABYR/i\THOCYATHUS Cairns, 1979
Labyrinthocyathus limatulus (Squires, 1964)
Ceratotrochus (Ceratotrochus) limatulus Squires, v*1964: 3-5, pi. 1, figs 5-9. — SQUIRES & KEYES, v. 1967: 24, pi. 2,
figs 9-10.
Labyrinthocyathus limatulus - Cairns, 1979: 70; 1995: 58, pi. 13, figs c-f, map 17.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 988, 3: 2 (MNHN), 1 (USNM 98592).
Type Locality. — 7.2 km northeast of the Alderman Islands. Coromandel Peninsula, North Island,
New Zealand, 102 m.
Remarks. — This is the only species in the genus to have a transversely ridged theca, the ridging becoming
less apparent toward the calicular edge. It is more fully described by Cairns (1995). This is the first record of
L. limatulus outside the New Zealand region.
DISTRIBUTION. — Vanuatu region: Tanna; 372-466 m. Elsewhere: northern New Zealand region; Lord Howe
Seamount Chain; 20-508 m (Cairns, 1995).
Source : MNHN . Paris
78
S. D. CAIRNS
Genus OXYSMILIA Duchassaing. 1870
Oxysmilia circularis Cairns, 1998
Figs 6 g-h, 7 a
Oxysmilia circulars Cairns, *1998: 378, figs 2 i-k.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 977, 1 (MNHN). — Stn 983, 2 (USNM 98624). —
Stn 1030. 2 (MNHN).
Kermadec Islands. NZOI: stn K830. 1 (USNM 99293). — Stn K858. 1 (USNM 99294).
Type LOCALITY. — "Soela" stn 02/82/16: 1 8°4 1 ’S, 1 17°54'E (off Port Hcdland. Western Australia). 200-
204 m.
Remarks. — A characteristic of this species is the irregularity of S.s pair insertion, some half-systems within
the same specimen having a full two pairs of S5, some only one pair, and some no S5. A redescription of the
species is not included here since it was so recently described. A specimen from MUSORSTOM 8 stn 983 is the
largest known, measuring 29.8 x 27.9 mm in CD and 41.0 mm in height.
DISTRIBUTION. — Vanuatu region: Tanna and Efate; 190-475 m. Elsewhere: northwestern Western Australia;
Kermadec Islands (Curtis and Raoul), reported herein; 201-545 m.
Oxysmilia corrugata sp. nov.
Figs 6g-h, 7a
Material EXAMINED/TYPES. — Vanuatu. MUSORSTOM 8: stn 1030. II: holotype and 7 paratypes (MNHN),
3 (USNM 98625).
TYPE Locality. — Musorstom 8 stn 1030: 17°5rS, I68°30'E (Efate), 180-190 m.
ETYMOLOGY. — The species name corrugata (Latin corrugatus , ridged) refers to the fine file-like, regularly
corrugated thecal ridging.
Description. — Corallum straight and elongate-conical, with a slightly Hared calicc. Largest specimen
(holotype) I 1.0 x 9.6 mm in CD, 4.3 mm in PD, and 12.5 mm in height. Calice slightly elliptical: GCD:LCD =
1 .07-/. 14- 1 .27. Corallum firmly attached through a thick, robust pedicel: PD.GCD = 039-0.60-0.il. Theca
uniformly covered with well-defined, thin-edged, transverse ridges, about 6 parallel ridges per mm. Whereas thecal
ridges occasionally bifurcate and rejoin one another, they often run uninterrupted for the entire circumference of the
pedicel. Ridges cover the entire thecal surface, from base to distal peripheral edges of septa, but are often encrusted
toward the base by bryozoa, foraminifera, and sponges. Corallum white.
Septa hcxamerally arranged in 4 complete cycles: Si>S:>S3>S4 (48 septa), the complete fourth cycle attained
at a GCD of 5.7 mm or less. Si about 2.4 mm exsert, thick, having straight axial edges that penetrate into the
columella, and coarsely granular faces. S2 similar to the Si but slightly less wide and less exsert. S3 slightly less
exsert, wide, and thick than S2. S4 only slightly less exsert and wide as S3. Thus, there are four size classes of
septa, but the differences in size are minimal. Fossa of moderate depth. Columella composed of 4-6 coarse,
granular papillae, which in larger coralla fuse into a solid, elongate, massive structure. The linear shape of the
columella is sometimes modified into a cross-shape by short symmetrical outpocketings of the columella into the
two lateral septal systems.
REMARKS. — There are three other Recent species of Oxysmilia : O. rotundifolia (H. Milne Edwards & Haime.
1848); O. circularis Cairns, 1998; and O. epithecata. O. corrugata differs from the first two in having a smaller
Source :
AZOOXANTHELLATE SCLERACTINIA
79
corallum with fewer septa, a transversely ridged theca, and a better-developed columella. It is compared to
O. epithecata in the account of that species (below).
Distribution. — Vanuatu region: Efate; 180-190 m.
Oxysmilia epithecata sp. nov.
Figs 6 d-e, 7 b-g
Material EXAMINED/TYPES. — Wallis and Futuna region. Musorstom 7: stn 496, 1 paratype <MNHN),
SEM stub 873 (USNM 98626). — Stn 507, 2 paratypes (USNM 98627). — Stn 509. 1 paratype (USNM 98628). —
Stn 511, 1 paratype (MNHN). — Stn 514, 6 paratypes (USNM 98629). — Stn 523. 2 paratypes (MNHN). — Stn 556.
1 paratype (MNHN). — Stn 605, 17 paratypes (USNM 98630). — Stn 610. 1 paratype (MNHN).
Vanuatu. Musorstom 8: stn 959, I paratype (MNHN). — Stn 962, I paratype (MNHN). — Stn 1018, holotype
(MNHN). — Stn 1023, 5 paratypes (MNHN). — Stn 1026, 1 paratype (MNHN). — Sin 1097, 7 paratypes (MNHN).
Type Locality. — Musorstom 8 stn 1018: 17°53'S, 168°25'W (Efate), 300-301 m.
ETYMOLOGY. — The species name epithecata (Greek epi , on + theke , sheath) refers to the wrinkled epitheca
that covers the lower corallum.
Description. — Corallum straight and elongate-conical, with a slightly flared calice. One to four coralla
often originate from the dead calicular edge of a conspccific corallum. producing a small, bushy pseudocorallum
(Fig. 7 b). Holotype 7.1 x 6.4 mm in CD, 3.3 mm in PD, and 10.7 mm in height; largest spec¬
imen (MUSORSTOM 8 stn 1026) 11.3 mm in GCD. Calice circular as juvenile, becoming elliptical
with age (GCD:LCD = 1.07-1.34). Corallum firmly attached through a robust pedicel (PD:GCD = 0.33-0.46)
and thin, expansive, encrusting base, the base often twice the diameter of the calice. Entire base and lower
quarter to half of theca covered with fine transverse epithecal ridges, those ridges on the base (6 per mm)
more widely spaced than those on the pedicel (13 per mm). As in O. corrugata , the ridges may run uninterrupted
around the entire circumference of the pedicel or base, but, in general, arc more irregular in arrangement
and spacing. The ridges are about 10 pm wide and 30 pm tall (Figs 7 f-g). Proximal to the epithecal ridg¬
ing, the upper 3/4 to half of the theca is costate, the Ci-2 usually slightly ridged; all costae granular.
Corallum white.
Septa hexamerally arranged in 4 cycles, the fourth cycle complete only in the largest corallum: S|>S2>S;t>S4.
Coralla below 3.5 mm in GCD contain only 3 cycles of septa (24), whereas coralla with a GCD between 3.5
and 11.0 mm have an increasing number S4. the intermediate number of 36 septa being most common.
The holotype has 44 septa. Si up to 1.3 mm exsert, having straight axial edges that reach the columella.
S2 slightly less exsert (about 0.9 mm), also having straight axial edges that reach the columella. S3 less
exsert than S2 and well developed only in large coralla, where they are about half the width of an S2. their
lower axial edges sometimes fusing to the adjacent S2- S4 least exsert septa, only developed in largest of
coralla; otherwise restricted to exsert lobes at the calicular edge. Fossa deep, usually containing a deep-
seated papillose columella. Small coralla often have no columella, whereas most larger coralla have a small
columella composed of several small fused papillae, and one specimen (Musorstom 8 stn 1097) has a robust
columella.
REMARKS. — Oxysmilia epithecata is most similar to O. corrugata, both species being about the same size,
having the same number of septal cycles, and having transverse thecal ridges. O. epithecata differs in having finer,
irregular transverse ridges that are restricted to the lower corallum, and a granular costate upper theca; a less robust
pedicel; less septa at a corresponding GCD; a deeper fossa; and a less developed columella.
DISTRIBUTION. — Wallis and Futuna region: Wallis and Futuna; Tuscarora Bank; 240-455 m. Vanuatu region;
Anatom. Efate, and Espiritu Santo; 288-437 m.
Source :
80
S. D. CAIRNS
Genus TROCHOCYATHUS H. Milne Edwards & Haime. 1848
Subgenus TROCHOCYATHUS (TROCHOCYATHUS) H. Milne Edwards & Haime. 1848
Trochocyathus (T.) vasiformis Bourne, 1903
Figs 8 a-b. f
Trochocyalhus vasiformis Bourne, v*1903: 27-28. pi. 5, figs 6-7.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: sin 522, 1 (MNHN). Stn 527,
I (MNHN). — Sin 535, 2 (MNHN). — Sin 609, 1 (MNHN).
Vanuatu. MUSORSTOM 8: stn 959, 2 (USNM 98634). — Sin 973, 2 (USNM 98636). — Stn 977. 19 (MNHN), SEM
stub 883 (USNM 98635). — Stn 978, 4 (MNHN). — Stn 983. 3 (MNHN). — Stn 1006. I (MNHN). — Stn 1011,
1 (MNHN). — Stn 1015, 1 (USNM 98633). — Stn 1019. 1 (MNHN). — Sin 1067. 8 (USNM 98632). — Stn 1072.
1 (MNHN). — Stn 1 107, 1 (MNHN).
Indonesia (Mollucas). Karubar: sin 5, 2 (USNM 98637).
Type Locality. — Tutanga, Funafuti. Tuvalu Islands, 366 m.
Description. — Corallum straight, elongate-conical (ceratoid), and attached through a robust pedicel
(PD:GCD = 0.47-0.84) and an encrusting base. Largest specimen (MUSORSTOM 8 stn 977) 14.9 x 1 1.9 mm in CD
and 25 mm in height. Calice elliptical: GCD:LCD = 1.1 1-1.35. Theca thick; costae usually low. rounded, and
finely granular, but occasionally Ci-2 are slightly ridged. Theca and outer septa a light yellow-brown (beige), the
pali and columella white; however, one specimen (MUSORSTOM 8 sin 1 107) completely white.
Septa regularly hexamerally arranged in 4 complete cycles: Si-2>S.3-4 (48 septa). Si -2 exsert (up to 2.5 mm),
having straight axial edges each bordered by a small (0.4 mm wide) palus; however, the 2 Pi aligned with the
greater calicular axis (those before the principal septa) only about halt width and not as exsert as the other P I -2-
S3-4 equal in width, about 3/4 that of the S1-2; although equal in width, S3 slightly more exsert than S4.
A single, well-defined, elliptical palar crown occurs in the fossa, composed of 12 Pi -2 and 12 P3, the P3
2-3 times the width (0.9- 1.0 mm) of the P1-2 and usually rising slightly higher in the fossa. A large P3 alternates
with each smaller Pi -2, but axial edges of all pali are the same distance from the columella, which gives the
impression of a single crown, even though the pali are of three size classes and heights. Each palus bears several
prominent, finely dentate, obliquely-oriented menianes (Fig. 8f). Outer edges of P3 quite broad, occupying not only
space before the S3 but the two adjacent S4. Fossa of moderate depth. Columella an elliptical field of 10-25
slender, foliaceous elements. Columellar elements often shaped as papillae that support several horizontal to
slightly inclined, circular plates, the papillae passing through the centre of these platelets (Fig. 81). The shape of
the columellar elements is intermediate between the typical papillose elements characteristic ol Trochocyathus and
the twisted elements common to Caryophyllia .
Remarks. — This is believed to be the first report of this species subsequent to its description, most of the
specimens reported above collected in the same sector of the Pacific as the type locality (Funafuti), and at similar
depths to the types. The species is distinctive in having prominent menianes on its pali, a beige corallum, and
distinctively shaped columellar elements.
BOURNE (1903) stated in the original description that the smaller, worn syntype had only 42 septa, lacking
3 pairs of S4, but that specimen (BM 1903.12.1.5-6) actually has 44 septa, lacking pairs of S4 only in
half-systems II and IV, as counted from a principal septum.
Specimens from four stations contained coralla that were bored by acrothoracican eirripede Crustacea:
MUSORSTOM 7 stn 609, MUSORSTOM 8 stns 959, 977, and 101 1.
DISTRIBUTION. — Wallis and Futuna region: Wallis: Waterwitch Bank; 430-650 m. Vanuatu region: Anatom,
Tanna, Erromango, Efate, Malakula, and Espiritu Santo; 366-622 m. Elsewhere: Banda Sea (recorded herein);
Funafuti; 323-366 m.
Source : MNHN, Paris
AZOOX ANTHELLATE SCLERACTINIA
81
Trochocyathus (T.) rhombocolumna Alcock. 1902
Trochocyathus rhombocolumna Alcock, v*1902a: 98; v.!902c: 16, pi. 2. fig. 12. — Cairns, 1995: 60-61, pi. 13.
fig. I, pi. 14, figs a-b. map 19 (synonymy). — Cairns & Zibrowius, 1997: 1 06- 1 07.
Paracyathus temiicalyx Vaughan. v*1907: 69-70, pi. 6. figs la-b.
Material examined. — Vanuatu. Musorstom 8: stn 977, I (MNHN). — Stn 1060. I (USNM 98631).
Type Locality. — " Siboga " stn 95: 5°43.5'N, I I9°40'E (Sulu Sea). 522 rn.
REMARKS. This relatively commonly collected species is distinguished by having a transversely ridged theca
(basally); hexamerally symmetrical septa (Si>S:>S4>S3); three size classes of pali, the P2-3 of each system
arranged in a distinctive chevron pattern; and robust columellar elements. It is more fully described and illustrated
by Cairns (1995).
Distribution. — Vanuatu region: Tanna and Malakula; 397-410 m. Elsewhere: Indo-Wcst Pacific from
Southwestern Indian Ocean to Hawaiian Islands; 1 10-530 m (Cairns & Zibrowius, 1997).
Trochocyathus (T.) philippinensis Semper, 1872
Trochocyathus philippinensis Semper. v*l872: 253, pi. 20. fig. 16. — Cairns & Zibrowius. 1997: 107-108,
figs 10 d-e. — Cairns, 1998: 380.
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom 7: sin 496, I (MNHN).
Vanuatu. Musorstom 8: stn 969, 2 (MNHN). — Stn 976, 23 (USNM 98678). — Stn 1070, I (MNHN) — Sin 1071
1 (MNHN). — Stn 1086, 2 (MNHN).
Type Locality. — Pandanon, Philippines, 27-54 m.
Remarks. — Trochocyathus philippinensis is characterized as having a small straight, ccratoid corallum;
a porcellaneous theca that is pigmented chocolate-brown near the calicular edge; and a hexameral symmetry of
S|-2>S3-4- Some specimens have several thecal edge spines or slightly more prominent edge costae. The species
was more fully described and illustrated by Cairns & ZIBROWIUS (1997).
Distribution. — Wallis and Futuna region: Futuna; 250-330 m. Vanuatu region: Anatom. Tanna. Espiritu
Santo, and Malakula; 182-252 m. Elsewhere: Ryukyu Islands; South China Sea; Philippines; Indonesia;
northwestern Australia; 54-268 m (Cairns & Zibrowius, 1997).
Trochocyathus (T.) maculatus C aims, 1995
Trochocyathus maculatus Cairns. *1995: 61. pi. 14. figs c-d. — Cairns & ZIBROWIUS, 1997: 107.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 499, 2 (USNM 98641). — Stn 504,
| I (MNHN). — Stn 505, 1 (MNHN). — Stn 508, 3 (MNHN). — Stn 509, 3 (USNM 98639). — Sin 524. 4 (USNM
98640). — Stn 546, I (MNHN).
Vanuatu. Musorstom 8: stn 961, 1 (MNHN). — Stn 1021. 2 (MNHN). — Stn 1047, I (MNHN). — Sin 1077
1 (MNHN).
Type Locality. — NZOI stn PI 15: 31°25.9'S, 159°02.2’E (Lord Howe Island), 183 m.
REMARKS. — This species is distinguished from other Trochocyathus by having distinctively speckled (brown-
black) theca and septa, and by having exothecal dissepiments that cover raised costae around its base, similar to the
morphology of Rhizosmilia. It was fully described and figured by CAIRNS (1995).
Source :
82
S. D. CAIRNS
DISTRIBUTION. — Wallis and Fuluna region: Wallis and Futuna; Combe Bank; 240-550 m. Vanuatu region:
Anatom, Efate, Epi. and Malakula; 1 10-486 m. Elsewhere: Philippines; Kermadec Islands; Lord Howe Islands;
seamounts off eastern Australia; 92-183 m.
Trochocyathus (T.) efateensis sp. nov.
Figs 8 d-e
MATERIAL EXAMINED/TYPES. — Vanuatu. Musorstom 8: stn 1019, 25: holotype and 19 paratypes (MNHN),
5 paratypes (USNM 98642). — Stn 1020, 1 paratype (MNHN). — Stn 1026, 1 paratype (MNHN).
Type Locality. — Musorstom 8 stn 1019: I7°38'S, 168°34’E (Elate), 397-430 m.
Etymology. — Although this species is not thought to be endemic to Efate, it is named for the island of its
type locality.
Description. — Corallum straight, elongate-conical, having a slightly flared calice, and attached through a
slender (PD:GCD = 0.29-0.42), but solid, pedicel. Pedicel reinforced with layers of textura, which arc usually
subsequently encrusted by epizoic organisms. Calice strongly elliptical (GCD:LCD = 1.21-2.00), small coralla
being closer to circular, larger coralla becoming more elongate, sometimes slightly constricted medially. The
constriction is accentuated by an upward arching of the calicular edges as viewed from the side. Holotype
16.7 x 11.2 mm in CD. 5.7 mm in PD, and 21.6 mm in height; largest specimen (MUSORSTOM 8 sin 1028)
17.3 x 13.6 mm in CD. Costae broad, equal in width, and inconspicuous, covered with low. rounded granules.
Corallum uniformly white.
Septa hcxamerally arranged in 5 cycles, the fifth cycle never complete (Si-2>S3>S4>S5). One specimen
(Musorstom 8 stn 1019) of 13.4 mm GCD contains only 4 cycles of septa (48), but all other coralla have
between 1 and 10 pairs of S5, resulting in 50-68 septa. The holotype contains 68 septa. Order of insertion of S5
pairs quite irregular, but S5 pairs tend to be more common in end systems; some systems may contain 3 pairs of
S5 and other systems within the same corallum no S5. All septa equally exsert ( 1.3- 1.5 mm), having straight axial
edges. Si -2 quite thick: 0.8- 1.1 mm at their widest upper edges. The remaining three size classes of septa are less
thick and progressively less wide, the S5 being about half the width of an S 1 -2- All pal i have straight, thin axial
edges that border the columella, and broad, ridged peripheral edges that are separated from their corresponding septa
by narrow slits. Distal edges of all pal i bluntly pointed, not evenly rounded as in most species of the genus.
The 12 slender (0.8 mm wide) P| .2 form an elliptical crown around the columella. The 12 P3 are about twice the
width and rise higher in the fossa than the Pi-2, but, since their axial edges are in the same position relative to the
columella as those of the Pi -2, the P3 contribute to the same palar crown. P4 present only when pairs of S5 occur,
these pali about the same width of a P1-2, but recessed farther from the columella and positioned higher in
the fossa. Fossa of moderate depth. Columella papillose, composed of 9-20 robust, irregularly-shaped, granular
elements, in some coralla rhomboidal in cross section.
Remarks. — Of the 29 Recent species in the nominate subgenus of Trochocyathus, T. efateensis is most
similar to T. caryophylloides Alcock, 1902. Although both species may have 64 septa, T. caryophylloides
achieves it by having 16 sectors (three size classes of septa and two size classes of pali). whereas T. efateensis
retains a hexameral symmetry and has four size classes of septa and three size classes of pali. T. efateensis is
further differentiated by its thick Si-2, equally exsert septa, highly elliptical calice. and arched upper thecal faces.
Distribution. — Vanuatu region: Efate; 391-437 m.
Trochocyathus (T.) patelliformis sp. nov.
Figs 8 g. 9 a-c
Material EXAMINED/TYPES. — Vanuatu. Musorstom 8: stn Mil. I paratype (MNHN).
Hawaiian Islands. HURL: stn P5-063, holotype (USNM 83026).
Source :
AZOOXA NTHELL ATE SCLERACTINIA
83
Type Locality. — HURL stn P5-063: 20°35.8'N. 156°03.5*W (Alenuihaha Channel. Hawaiian Is.). 1020 m.
ETYMOLOGY. — The species name patelliformis (Latin patella, small pan + forma, shape) refers to
the patelli form shape of the corallum, which in Scleractinia includes those species having a basal angle of
80°- 1 60° .
Description. Corallum short and patellate, the thecal edges diverging at about 120° from a massive basal
attachment. Holotype 22.7 x 19.2 mm in CD, 9.0 mm in pedal attachment, and only 1 1.0 mm in height. Smaller
paralype 10.9 mm in GCD. Costae well defined, rounded, and covered with fine, spiny granules. Intercostal
grooves equal in width to costae on lower corallum, becoming wider and deeper near calicular edge. Corallum
uniformly white.
Septa hexamcrally arranged in 5 cycles, the fifth cycle incomplete (S|-2>S3>S4>S5). The holotype contains
12 pairs of irregularly inserted S5: 5 half-systems having 2 pairs of S5, 5 having no S5, and 2 half-systems having
1 pair o! S5, lor a total of 72 septa. The smaller paralype contains 8 pairs of incipient S5, also irregularly
distributed, or 64 septa. Si -2 highly exsert (up to 3.7 mm), having straight axial edges and granular faces. S3 less
exsert (about 3.3 mm) and 3/4 width of Si -2. S4 less exsert (about 2.7 mm) but almost as wide as the S3.
S5 about 1.3 mm exsert and 3/4 width 0! an S4. Two size classes and 4 crowns of pal i occur in the holotype. The
6 Si are quite large (2.2 mm wide), each having a straight axial edge that abuts the columella, but a moderately
sinuous peripheral edge, separated from its corresponding septum by a small notch. Remaining pali (P2-P4) all
about 1.7 mm wide, each cycle progressively more recessed from the columella and rising slightly higher in the
lossa. Fossa shallow. Columella composed of several finely granulated massive papillae that are fused into an
irregular, elongate mass.
Remarks. — Trochocyathus patelliformis is unique among the approximately 29 Recent, valid species in the
genus in having a patel late-shaped corallum, in having Pi that are significantly broader than the higher cycle pali,
and in having finely spinose costae. The Hawaiian holotype was diagnosed as a potentially new species shortly
after its collection in 1988 but pul aside until more specimens might be collected. The Vanuatu paralype, while
apparently a juvenile corallum, was collected from a similar depth range and is morphologically consistent with
the holotype.
DISTRIBUTION. — Vanuatu region: Espiritu Santo; 1210-1250 m. Elsewhere: Hawaiian Islands; 1010 m.
Trochocyathus (T.) semperi Cairns & Zibrowius, 1997
Trochocyathus (T.) semperi Cairns & Zibrowius, *1997: 108-109. figs lOg-h. Ilf.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 1 103. 1 (MNHN).
Type Locality. — Corindon 2 stn 251: 0°53.7'S, 1 19°29.6'E (Makassar Strait). 65 m.
Remarks. — This recently described species is distinguished from all others in the genus by having spatulate
edge spines and decameral septal symmetry (10: 10:20. 40 septa).
Distribution. — Vanuatu region: Espiritu Santo; 163-165 m. Elsewhere: Philippines; Indonesia; 38-245 m
(Cairns & Zibrowius, 1997).
Trochocyathus (T.) cooperi (Gardiner, 1905)
Tropidocyathus cooperi Gardiner. *1905: 955, pi. 93. fig. 30.
Trochocyathus cooperi - Cairns, 1994: 54, pi. 23, figs f-g. — Cairns & Zibrowius. 1997: Ill, fig. I le (synonymy).
Material EXAMINED. — Vanuatu. Musorstom 8: stn 961. I (MNHN). — Stn 1021. I (MNHN).
Type Locality. — Kolumadulu and Suvadiva. Maidive Islands. 64-70 m.
Source :
84
S. D. CAIRNS
REMARKS. — This is a very distinctive species, characterised by having transverse division, thecal edge crests,
and usually a brown mottled pigmentation at the calicular edge. It is more fully described and illustrated by Cairns
(1994).
Distribution. — Vanuatu region: Anatom and Elate; 101-124 m. Elsewhere: widespread from Maidive Islands
to northern Ryukyu Islands and the Marquesas; 25-100 m (Cairns & Zibrowius, 1997).
Trochocyathus (T.) discus Cairns & Zibrowius, 1997
Figs 9 d-e, 1 8 a
Trochocyathus (T.) discus Cairns & Zibrowius, *1997: 1 12, figs 1 1 g-h. 12 a-c.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: 539, 1 (MNHN).
Vanuatu. MUSORSTOM 8: stn 958, 1 (MNHN). — Stn 1088, 3 cemented to a Xenophora gastropod shell (MNHN). —
Stn 1089. 1 (MNHN). — Stn 1090. 1 (USNM 98643).
Type Locality. — Karubar stn 3: 5°48'S, 132°12'E (Kai Islands, Banda Sea), 278-300 m.
REMARKS. — Little can be added to the original description except to note that the corallum Irom
Musorstom 8 stn 1089 (Figs 9 d-e) is the largest yet recorded: 1 1.3 x 9.3 mm in CD and 7.9 mm in height.
The species is characterised by having a discoidal to bowl-shaped corallum; a flat base, the result of transverse
division; four cycles of septa, the S4 adjacent to the Si being wider than the S3; and having a reddish-brown
pigment to the calicular edge.
DISTRIBUTION. — Wallis and Futuna region; Combe Bank; 700 m. Vanuatu region: Anatom and Espiritu
Santo; 455-497. Elsewhere: Banda Sea; 240-278 m (Cairns & Zibrowius, 1997).
Subgenus TROCHOCYATHUS ( A PLO C YATH U S) d'Orbigny, 1849
Trochocyathus (A.) hastatus Bourne, 1903
Trochocyathus hastatus Bourne, v*1903: 29-32 (in part: pi. 5. figs 2-3; not pi. 6, figs 8-11, = Bourneotrochus stellulatus
Cairns, *1984). — Cairns. 1995: 63-64, pi. 15, figs c-h. map 21.
Stephanocyalhus ( Acinocyathus ) hastatus - Wells, v.1984: 213.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 523, I (MNHN). — Stn 555.
I (MNHN). — Stn 586, 1 (MNHN). — Stn 618. 1 (MNHN). — Stn 619, 2 (MNHN).
Vanuatu. Musorstom 8: stn 959, 7 (MNHN). — Stn 963, 3 (USNM 98645). — Stn 977. 2 (USNM 98646). —
Stn 1019. 2 (USNM 98644). — Stn 1020, I (MNHN).
Type Locality. — Tutanga, Funafuti, Tuvalu, 366 m.
Remarks. — Trochocyathus hastatus is compared to the other two Recent species in the subgenus in Table 5.
and was redescribed and illustrated by Cairns (1995). It is easily distinguished from congenerics by having
hexameral septal symmetry but only 5 costal spines. CAIRNS (1995) noted that the Si unaccompanied by
a costal spine was always significantly smaller than the other Si; however, both this septum and the opposing
principal Si (opposite to the first) are equally small, the 4 other lateral Si being significantly wider and
more exsert.
DISTRIBUTION. — Wallis and Futuna region: Wallis and Alofi; Tuscarora Bank; 435-540 m. Vanuatu region:
Anatom and Efate; 391-436 m. Elsewhere: Kermadec Islands; Funafuti; 366-710 m (Cairns, 1995).
Source :
AZOOXANTHELLATE SCLERACTINIA
85
Table 5. Comparison of the three Recent species of Trochocyathus (Aplocyathus).
Trochocyathus (A.) brevispina Cairns & Zibrowius, 1997
Figs 9 f-i
Trochocyathus (A.) brevispina Cairns & Zibrowius, *1997: 113, figs 12 d-f.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 959, 2 (MNHN). — Stn 963, 65: 56, including one
cemented to a Xenophora shell (MNHN), 9 (USNM 98649). — Sin 964, 14 (MNHN). — Sin 999 1 (US.NM 98647) —
Stn 1003, 4 (MNHN). — Stn 1004. 12: 8 (MNHN). 4 (USNM 98648). — Stn 1020. I (MNHN). — Sin 1058. 2 (MNHN)
— Stn 1065, 1 (MNHN).
Type Locality. — Karubar stn 3: 5°47'40"S, 132° 12' I F'E (Kai Islands. Banda Sea), 278-300 m.
Remarks. — Trochocyathus brevispina is compared to the other two Recent species in the subgenus in
Table 5. and was described and illustrated by Cairns & Zibrowius (1997). It is easily distinguished from other
species by its relative septal and palar sizes and by its frequent incorporation of substrate into its base. The most
commonly incorporated substrates are gastropod and bivalve fragments (Figs 9 f. i). but also include: pteropod
shells, sand conglomerate, pebbles, serpulid tubes, and other dead solitary corals.
The specimens reported above allow the observation that the costal spines are often curved or crooked, and can
reach a length of up to 8 mm. Although not as long as the 10 mm costal spines of T. longispina , the distinction
of costal spine length between these two species is not as much as previously thought and hardly warrants the
etymological distinction; however, several other characters do distinguish these two species (Table 5).
Specimens from three stations (Musorstom 8 sin 1004, 1058, and 1 103) bear elaborate, irregularly-shaped,
nodular proliferations at the base of their costal spines and sometimes around the entire calicular edge (Figs 9 g-h).
It is unknown what stimulates or irritates the coral to produce these growths.
Source :
86
S. D. CAIRNS
One aberrant specimen from MUSORSTOM 8 stn 963 of 16.5 mm GCD is septameral in septal and costal
symmetry, having 7 septal systems (54 septa, 1 hall-system lacking S4) and 7 costal spines.
Distribution. — Vanuatu region: Anatom, Erromango, Efate, and Malakula; 1 10-436 m. Elsewhere: Banda
Sea; 240-560 m (CAIRNS & ZlBROWlUS, 1997).
Genus TETHOCYATHUS Kuhn. 1933
Tethocyathus virgatus (Alcock, 1902)
Trochocyat hus ( Tethocyathus ) virgatus Alcock. v*1902a: 98-99; v. 1902c: 16-17. pi. 2, lig. 13.
Tethocyathus virgatus - CAIRNS, 1995: 65-66. pi. 16. figs c-f. map 11 (synonymy). — Cairns & Zibrowius. 1997:
1 14-115.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: sin 511, 1 (MNHN). — Stn 522, 1 (USNM
98652). o
Vanuatu. MUSORSTOM 8: stn 973. 7 (USNM 98650). — Stn 977. 5 (MNHN). — Stn 978, 2 (MNHN). — Stn 982,
2 (MNHN). — Stn 1015, 1 (MNHN). — Stn 1026, I (MNHN). — Stn 1095, 1 (MNHN). — Stn 1 108, 3 (MNHN).
Type Locality. — ", Sihoga " stns 96 and 105: Sulu Archipelago, 275 m.
REMARKS. — Tethocyathus virgatus is more fully described and illustrated by CAIRNS (1995). One ol the
differences between this species and the species of the similar genus Trochocyathus is that the edge zone of
T. virgatus periodically secretes additional nontrabecular calcium carbonate on the outside of the corallum, which,
in time, produces a very thick theca, transforming a conical corallum into a subcylindrical or cylindrical shape. The
layering of the added calcium carbonate is particularly noticeable in a basal fracture, and additional evidence of the
periodic layering is evidenced by the covering and incorporation into the theca of small epizoic organisms that
become covered and subsequently buried in the increasingly thick theca. According to Stolarski (1995), the
proper name for this deposit is textura, not epitheca, the latter term being reserved for nontrabecular calcium
carbonate that is secreted only at the calicular edge of a corallum (lappet cavity) and that does not periodically form
layers. One of the adaptive values of textura (in the basal region) is to strengthen attachment to a substrate;
however, it is suggested that another possible adaptive value of textura (in the wall region) is to guard against the
boring of acrothoracican cirripeds. to which this genus is particularly susceptible (see Zibrowius, 1976; Cairns,
1995; Cairns & Zibrowius, 1997). Coralla from two stations (Musorstom 8 stns 972 and 982) contain coralla
having the characteristic lenticular-shaped outline of acrothoracican cirripcde borings in the corallum of living
specimens.
Distribution. — Wallis and Futuna; 450-650 m. Vanuatu region: Tanna, Efate, and Espiritu Santo;
320-460 m. Elsewhere: Philippines; Indonesia; ridges north of New Zealand; 137-530 m.
Genus POLYCYATHUS Duncan, 1876
Polycyathus oc tup l us sp. nov.
Figs 1 0 a-c
Material EXAMINED/TYPES. — Wallis and Futuna region. MUSORSTOM 7: stn 494, 1 holotype (MNHN).
— Stn 496, 2 paratypes (MNHN). — Stn 499, 1 paratype (USNM 98654). — Stn 504, 7 paratypes (MNHN).
3 paratypes (USNM 98653). — Sin 509, 3 paratypes (MNHN). — Stn 513, 1 paratype (MNHN). — Stn 516. 1 paratype
(MNHN).
Solomon Islands (Pelau): short drop off, 90 m, 3 paratypes (USNM 98954).
Source :
AZOOX ANTHELL ATE SCLERACTINIA
87
Type Locality. — Musorstom 7 stn 494: 14°18.9'S, 178°03.0'W (Futuna), 100-1 10 m.
Etymology. The species name octuplus (Latin octuplus , eight fold) refers to the octameral septal
symmetry.
Description. Corallum solitary, straight, and subcylindrical, firmly attached to substrate through a thick
pedicel (PD:GCD = 0,56-0.79). Holotype 6.7 x 6.4 mm in CD. 4,4 mm in PD. and 7.5 mm in height. Calice
circular to slightly elliptical: GCD:LCD = 1.02-1.20. Costae well developed and finely granular, however, at an
early stage thin bands of epitheca are secreted around the corallum base, and as the coral increases in size the
epithecal bands thicken, eventually extending from the base to the calicular edge. Underlying costae and outer edges
oi septa chocolate brown in colour, but overlying epitheca, basal encrustation, pali and columella are white.
Septa octamerally arranged in 2 or 3 size classes. Small coralla below a GCD of 4.6 mm usually have
8 primary septa and 24 equally-sized secondary and tertiary septa (8:8:16, 32 septa). Larger coralla have up to
5 pairs of quaternary septa (e.g„ the holotype, 8:8:16:10, 42 septa) as 3 size classesr of septa: primaries,
secondaries, and tertiaries + quaternaries, the latter two of equal size. One aberrant specimen (MUSORSTOM 7
stn 504) has heptameral symmetry (7:7:14:6, 34 septa, Fig. 10 c). Primary septa about 1.2 mm exsen. having
moderately sinuous distal and axial edges. In small specimens, secondaries and tertiaries of equal exsertness (about
O. 9 mm) and width (about 3/4 that of the primaries), also having sinuous distal and axial edges. In larger
specimens containing sectors in which quaternaries arc present, the secondaries are slightly wider than the
remaining tertiaries and quaternaries. Quaternaries equal in size to tertiaries. Small pali (0.3-0.4 mm wide) occur on
axial edges ol the primary septa, whereas the pali flanking the secondary septa are twice as wide and rise slightly
higher in the fossa. Occasionally there is an accessory paliform lobe internal to the P2. When quaternary septa
occur in a sector, the flanked tertiary septum bears a palus of equal size to the P2, but slightly recessed from the
columella and also rising slightly higher than the P2. All pali arc obliquely carinate and highly sinuous, their
peripheral edges (edge adjacent to their corresponding septum) being quite contorted. Fossa shallow, containing a
discrete columella composed of a field of 10-16 slender, finely granular papillae.
Remarks. — It IS with some hesitation that this species is placed in Polycyatlms, since it does not display the
defining character for the genus, i.e., budding from a common encrusting coenosteum. It may be that all the
specimens examined are founder corallites or that they were broken from the substrate above their common
attachment. In all other characteristics, this species is characteristic of the genus Polycxathus. Five species of this
genus are known from the western Pacific: P. fulvus Wijsman-Bcst. 1970; P. Hodgson i, P. marigondoni, and
P. furanaensis, all described by Verheu and Best, 1987; and P. norfolkensis Cairns. 1995. P. octuplus differs
Irom these and all other known species (BEST et al., in press) in having primarily octameral septal symmetry.
Distribution. — Wallis and Futuna region: Futuna; 1 10-441 m. Elsewhere: Solomon Islands, Pelau; 90 m.
Genus BOURNEOTROCHUS Wells, 1984
Boiirneotrochus stellulatus (Cairns, 1984)
Figs 8 c, 10 d-g
Trochocyathus hcistatus Bourne, v* 1903: 32-37 (in part: pi. 6, figs 9-1 I).
Deltocycithus stellulatus Cairns, *1984: 15-16. pi. 3. figs C-D.
Bourneotrochus veroni Wells, v*1984: 213-214. pi. 3, figs 7-18.
Boiirneotrochus stellulatus - Cairns, 1995: 71-72, pi. 18. figs f. i. pi. 19, figs a-e, map 18. — Cairns & Zibrowius
1997: 115.
Material EXAMINED. — Wallis and Futuna region. Musorstom 7: sin 509. 4 (MNHN) and SEM stub 879
(USNM 98708). — Sin 510. 3 including 1 anthocaulus (MNHN). — Stn 511, 14 including 3 anthocauli and SEM stubs
880-881 (USNM 98704). — Stn 512, 65 including I anthocaulus (MNHN). — Stn 513, 25 including 1 anthocaulus
Source :
88
S. D CAIRNS
(USNM 98707) — Sin 514. 13 (USNM 98706). — Sin 516. 5 (USNM 98705). — Stn 556. 2 (MNHN). — Sin 569.
3 (MNHN). — Stn 585. 5 including 3 anthocauli (USNM 98710). — Sin 586. 2 (MNHN). — Sin 591. I (MNHN). —
Sin 594 2 (MNHN). — Sin 595. I (MNHN). — Sin 604. 13 including 3 anthocauli and SEM slub 878 (USNM 9871 1 ). —
Sin 605, 8 (USNM 98711). — Stn 608. 5 (USNM 98709). — Sin 610. 14 including 1 anlhocaulus (MNHN). — Sin 618.
3 anthocauli (MNHN).
Vanuatu. MUSORSTOM 8: stn 963. 2 cemented to Xenophora shell (MNHN). — Stn 969. I (MNHN). Stn 10 IS.
1 cemented to Xenophora shell (MNHN). — Stn 1023. I cemented to Xenophora shell (MNHN). — Stn 1061. I (MNHN).
— Stn 1087. 2 cemented to Xenophora shells (MNHN). — Stn 1097. 11 cemented to Xenophora shells (MNHN). —
Stn 1106. 10 cemented to Xenophora shells (MNHN).
TYPE Locality. — HON stn 9-3: 19°48’N, 154°58'W (off Hawaiian Islands), 337 m.
Remarks. — The anthocyathus of B. stellulcitus was recently redescribed and illustrated by Cairns (1995);
however, the anthocaulus stage remained unknown. Of the 215 specimens reported above, 15 are the anlhocaulus
stage, in some cases still attached to an incipient anthocyathus (Fig. 10 e). The anthocaulus is cylindrical to barrel
shaped, 1.5-2. 9 mm in height and 2. 6-3. 3 mm in maximum diameter, usually having 36 septa, as in most of
anthocyathi. The base is slightly expanded over the substrate and firmly attached to it. It has a white, porcellaneous
theca and lacks spines. The small, still-attached anthocyathus also lack costal spines at this stage.
Most anthocyathi examined contained 36 septa, one pair of Sa in each system. Because the costal spines are
associated with the C|. they are symmetrically distributed around the perimeter of the corallum (Fig. 10 d).
However, about 20% of the examined coralla lack one to three pairs of S4. resulting in a total of 30-34 septa and a
closer arrangement of the costal spines that border these systems. Some anthocyathi (e.g., MUSORSTOM 7 sin 513,
594. 604) appear to have the ability to asexually bud another anthocyathus from its calice. These "secondary"
anthocyathi are small (2. 7-3.0 mm in diameter), cylindrical coralla that have a well-developed basal scar, lack
spines, often have less than 36 septa, and appear to have a primitive epithecal wall.
Although no anthocyathus was found to have a CD larger than that previously reported (i.e., 6 mm by Cairns,
1995), some specimens (e.g.. Musorstom 7-510) have elongate costal spines up to 4 mm long, twice the
previously reported length.
Among the 21 coral species found cemented to Xenophora shells. B. stellulatus was most commonly lound.
27 coralla recorded from six Musorstom stations (see Material Examined). In all cases the calicular face was
directed upward and in some cases the corallum appeared to have been alive when collected.
Distribution. — Wallis and Futuna region: Wallis, Futuna, and Alofi; Tuscarora. Waterwitch, and Field
Banks; 240-566 m. Vanuatu region: Anatom. Efate, Malakula, and Espiritu Santo: 280-458 m [Pleistocene ol
Espiritu Santo (Wells, 1984)|. Elsewhere: Queensland; ridges north of New Zealand; Indonesia; Chesterlield
Islands; Funafuti and Tuvalu; Cook Islands; Hawaiian Islands; 263-476 m (Cairns & ZlBROWIUS, 1997).
Genus STEPHANOCYA THUS Segucnza. 1 864
Subgenus STEPHANOCYA/ HUS (STEPHANOCYATHUS) Segucnza, 1864
Stephanocyathus (S.) regius Cairns & Zibrowius. 1997
Figs 10 h. II a-c
Stephanocyathus (S.l regius Cairns & Zibrowius, *1997: 1 17-1 18. figs 14a-c.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7; stn 564. 68 (MNHN). — Sin 565,
8 (USNM 98659). — Sin 567. 14 (MNHN). — Sin 621, 5 (MNHN). — Stn 623, 2 (MNHN). — Sin 635. 24 (MNHN). —
Stn 636. 14 (MNHN). — Stn 637, 14 (USNM 98660).
Vanuatu. Musorstom 8: stn 990, 1 (MNHN). — Stn 992. 5 (USNM 98655). — Stn 996. 2 (MNHN). — Stn 1007.
2 (MNHN). — Stn 1036. 1 1 (MNHN). — Stn 1 109. 5 (USNM 98656). — Stn 1110. I (MNHN). — Stn 1125, I (USNM
98658). — Stn 1127, I (MNHN). — Stn 1129. 8. including SEM stub 882 (USNM 98657).
Source :
AZOOXANTHELLATE SCLERACTIN'IA
89
Type Locality. — "Hokuho-Maru" stn KH72-1-26: 9°27’S, 127°58.6’E (Timor Sea, south of Leti Islands).
610-690 m.
Remarks. — Little can be added to the original description; however, the microstructure of an actively
growing costal edge was examined, which revealed its irregularly-shaped tufts of calcareous fibres. The tufts
measured 11-15 pm in diameter, and consisted of elongate fibres about 0.9 pm in diameter (Figs 1 1 b-c).
DISTRIBUTION. — Wallis and Futuna region: Tuscarora. Combe, and Rotumah Banks; 700-1280 m. Vanuatu
region: Erromango. Efate, and Espiritu Santo; Guyot Bougainville; 775-1550 m. Elsewhere: South China Sea;
Philippines; Indonesia; Kermadec Islands; 563-2160 m (Cairns & Zibrowius, 1997).
Subgenus STEPHANOCYATHUS (ODONTOCYA THUS) Moseley, 1881
Stephanocyathus < O .) coronatus ( Pourtales, 1867)
Figs 1 1 d-f
Platycyathus coronatus Pourtales, v*1867: 114.
Odontocyathus coronatus - MOSELEY, v. 1881: 148-151. pi. 2, figs 4a-b. 5a-b.
Sabinotrochus flatiliseptis Alcock. v* 1902a: 103; v. 1902c: 26. pi. 4. figs 24. 24a (new synonym).
Stephanocyathus (O.) coronatus - Cairns, 1979: 109-111. pi. 20. figs 5-6. 8-9 (synonymy): 1995: 69. pi. 17. figs j-l.
pi. 18. figs a-b.
Material EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 621, 3 (MNHN). — Stn 622.
4 (MNHN). — Sin 623, 7 (USNM 98661).
Vanuatu. MUSORSTOM 8: stn 956, 2 (MNHN). — Stn 1125, I (MNHN).
Type Locality. — 30°4I’N, 77°03’W (Blake Plateau off Florida), 841 m.
Remarks. — This species was recently redescribed and illustrated based on specimens collected from
submarine ridges north of New Zealand (Cairns, 1995). The material reported herein contains several ontogenetic
suites including coralla as small as 9 mm CD, which allows the synonymy of the juvenile specimen ALCOCK
(1902a) reported as Sabinotrochus flatiliseptis . The holotype of ALCOCK's species (CD =11.6 mm) was figured by
Cairns & Zibrowius (1997: fig. 14i) and a similarly-sized corallum (CD = 9.1 mm) is figured herein (Fig. 1 1 f).
DISTRIBUTION. — Wallis and Futuna region: Combe Bank; 1280-1300 m. Vanuatu region: Anatom; Guyot
Bougainville; I 175-1210 m. Elsewhere: Selayar, Flores Sea (ALCOCK. 1902a); Lord Howe Rise; Three Kings
Ridge; Kermadec Ridge; western Atlantic; 543-1276 m (Cairns, 1995).
Stephanocyathus (O.) weberianus (Alcock, 1902)
Stephanotrochus weberianus Alcock. v*1902a: 101-102; v. 1902c: 25, pi. 3. figs 22, 22a.
Stephanotrochus sibogae Alcock, v* 1902a: 101-102; v. 1902c: 25-26, pi. 3. figs 23. 23a.
Stephanocyathus (O.) ixine Squires, v*1958: 54 (in part: " Albatross " stn 5545, pi. 8, figs 3-4).
Stephanocyathus (O.) weberianus - Cairns, 1994: 57-58, pi. 25. figs d-f (synonymy); 1995: 68-69, pi. 17, figs g-i
(synonymy). — Cairns & Zibrowius, 1997: 1 19-120. figs 14g-h.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: sin 1074. 7 (USNM 98662). — Stn 1080. 3 (MNHN).
Type Locality. — "Siboga" sin 284: 8°43.fS. 127°16.7'E (Timor Sea), 828 m.
Remarks. — The species was redescribed and illustrated by Cairns (1994) and compared to S. (O.) coronatus
by Cairns (1994. 1995). It is a relatively common deep-water species found throughout the western Pacific at
depths of 700- 1 500 m.
Source :
90
S. I). CAIRNS
Distribution. — Vanuatu region: Espiritu Santo and Malakula; 798-799 m. Elsewhere: western Pacific from
Japan to Lord Howe Seamount Chain; 206-1756 m. although most records are deeper than 700 m (CAIRNS &
Zi BROW I US, 1997).
Subgenus STEPHANOCYATHUS (ACINOCYATHVS) Wells, 1984
Stephanocyathus (A.) spiniger (Marerizeller, 1888)
Stephanotrochus spiniger Marenzellcr. *18886. 20-21.
Odontocyathus spiniger - Yabe & Eguchi. 1942: 124-125. pi. 10, figs 26-28 (synonymy).
Stephanocyathus (O.) spiniger - WELLS, v. 1984: 209, pi. 2. figs 10-13. — Cairns & Parker. 1992: 26-27. pi. 7. figs g-i
(synonymy). — Cairns, 1994: 57, pi. 25. figs a-c (synonymy); 1995: 67-68, pi. 17, figs d-f, pi. 18. fig. c
(synonymy); 1998: 381. — Cairns & Zibrowius, 1997: 118-119, figs I3f, 14d.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSOKSTOM 7: stn 585. 1 (MNHN). — Stn 604,
1 (MNHN). — Stn 606, I (MNHN).
Vanuatu. MUSORSTOM 8: sin 963, 8 (MNHN). — Stn 1003, 1 (MNHN). — Stn 1004, 2 (USNM 98664). — Stn 1058.
I (MNHN). — Stn 1087, I (USNM 98665). — Stn 1092, 1 (USNM 98663).
Type Locality. — Sagami Bay, Honshu, Japan (depth not given).
Remarks. — This commonly collected, distinctive species, characterised by having six elongate costal spines
(Ci) and highly exsert Si. has been described and figured several times in the recent past (see synonymy). It is
compared to 5. expiations, the only other Recent species in this subgenus, by CAIRNS & ZIBROWIUS (1997).
Distribution. — Wallis and Futuna region: Wallis; 420 m. Vanuatu region: Anatom. Erromango, Malakula,
and Espiritu Santo; 319-400 m. Pleistocene of Vanuatu (Wells, 1984). Elsewhere: widespread throughout Indo-
West Pacific from southwestern Indian Ocean to Japan and South Australia; 120-695 m (Cairns & Zibrowius,
1997). Neogenc of Japan (Yabe & Eguchi, 1932b); Oligocene of Victoria, Australia (Dennant, 1899; see Yabe
& Eguchi, 1942).
Genus VAUGHAN ELLA Gravier, 1915
Vaughanella concinna Gravier, 1915
Figs 1 1 g-h
Vaughanella concinna Gravier, v*1915: 10. — Zibrowius, v.1980: 104-105, pi. 52, figs A-K. pi. 53. figs A-L
(synonymy).
Vaughanella oreophila - Cairns. 1995: 70. pi. 18, figs d-e (Not V. oreophila Keller. *1981).
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 572. 1 (MNHN).
Type Locality. — 38°35'30"N, 28°05’45"W (Azores), 1250 m.
Remarks. — One specimen is reported, measuring 30.4 x 27.9 mm in CD. 30.3 mm in height, 12.3 mm in
pedicel diameter, and having 62 septa (7 pairs of S5). It is indistinguishable from coralla reported from the eastern
Atlantic (e.g„ Jean Charcot 134, USNM 48765) by Zibrowius (1980). A re-evaluation of the specimens
I (Cairns, 1995) previously reported from north of New Zealand as V. oreophila Keller. 1981 also appear to be
typical V. concinna . KELLER's (1981) type material of V. oreophila is not included as V. concinna because she
stated that her specimens did not have P3, which are prominent in all coralla of V. concinna.
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91
DISTRIBUTION. — Wallis and Futuna region: Watcrwitch Bank: 500-560 m. Elsewhere: Norfolk and Colville
Ridges; 646-757 m (Cairns, 1995); eastern Atlantic between Celtic Sea, the Azores, and Madeira; 1022-3018 m
(ZlBROWIUS, 1980).
Genus DELTOCYATHUS H. Mil nc Edwards & Haime, 1848
Deltocyathus magnificus Moseley, 1876
Fig. 1 1 i
Deltocyathus magnificus Moseley, v*I876: 552-553; v.1881: 147-148, pi. 4, figs 1-2. — Cairns & Parker, 1992:
27-28, pi. 7. figs j-1, pi. 8, fig. a. — Cairns, 1994: 56. pi. 24. figs d-e. g-h (synonymy); 1998: 381-382, fig. 4a. —
Guerriero ei al„ 1996: 986. — Cairns & Zibrowius, 1997: 126-127.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 963. 12: 8 (MNHN), 4 (USNM 98666). — Stn 964
I (MNHN). — Stn 980. 1 (MNHN).
Type Locality. — "Challenger" stn 192: 5°49'S, 132°14'E (Kai Islands. Banda Sea), 236 m.
Remarks. — Deltocyathus magnificus is one of four species in the genus to have 5 cycles of septa in its adult
state, the others being D. rotulus (Alcock. 1898), D. suluensis Alcock. 1902, and D. sarsi (Gardiner & Waugh,
1939), the last thought to reproduce primarily by fragmentation. D. magnificus is the largest of the four species
and was redescribed and illustrated by Cairns (1995) and Cairns & Parker (1992), and is distinguished from
other western Pacific species in a key published by CAIRNS & Zibrowius (1997). The Vanuatu specimens differ
from others previously reported in having only 72 septa, pairs of S5 lacking from each half-system adjacent to an
S2. Perhaps because of this deficiency, some of these specimens have a hexameral outline (Fig. 1 1 i). Normally a
corallum of 8 mm CD would have a full complement of 96 septa, but Vanuatu coralla as large as 25 mm CD have
only 72 septa. All other characters being similar, the septal number and shape differences are considered to be only
population differences.
Distribution. — Vanuatu region: Anatom and Tanna; 408-433 m. Elsewhere: western Pacific from Japan to
southeastern Australia; Western Australia; 88-1500 m (Cairns & Zibrowius, 1997).
Deltocyathus rotulus ( Alcock, 1898)
Trochocyathus rotulus Alcock, *1898: 16, pi. 2, figs 1, la.
Deltocyathus fragilis Alcock, v* 1902a: 99-100; v. 1902c: 21, pi. 2, figs 15. 15a.
Deltocyathus rotulus - Cairns, 1994: 55-56, pi. 24, figs j-k. — Cairns & Zibrowius, 1997: 125-126, figs 16 a-c.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 621, 3 (MNHN). — Stn 622,
3 (MNHN). — Stn 623, 2 (MNHN).
Vanuatu. Musorstom 8: stn 1125, 5 (USNM 98667). — Stn 1127, 1 (MNHN). — Stn 1129, II (USNM 98668).
Type Locality. — "Investigator" stn 216: North Maidive Atoll, 1408-1756 m.
Remarks. — Medium- to large-sized specimens of Deltocyathus rotulus are easily distinguished as having
5 cycles of septa; a serrate (lancetted) calicular margin, each S4 and adjacent pair of Ss projecting beyond the S1-3;
a large, undercut papillose columella; and a prominent crown of large P4. Smaller specimens (i.e., CD < 12 mm)
may be confused with juveniles of D. suluensis , both species having between 48-72 septa at this size. Juvenile
D. rotulus differ from D. suluensis by having S4 solidly attached to the S3, not just attached by trabecular
processes, as in D. suluensis ; and in having a lancetted calicular margin. It also appears to be adapted to a deeper
Source
92
S. D. CAIRNS
(i.e., cooler) environment, most commonly found between 1000- 2000 m. D. rotulus is more fully described and
illustrated by CAIRNS (1994), and included in a key to congenerics by CAIRNS & ZIBROWIUS (1997).
DISTRIBUTION. — Wallis and Futuna region: Field Bank; 1280-1300 m. Vanuatu region: Guyot Bougainville;
1050-1160 m. Elsewhere: Indo-West Pacific from Durban. South Africa to Japan; 210-1986 m (Cairns &
ZIBROWIUS, 1997).
Deltocyathus suluensis Alcock, 1902
Deltocyathus magnificus var. suluensis Alcock. v* 1902c: 20-21.
Deltocyathus formosus Cairns, *1995: 73-74. pi. 19, figs f-g.
Deltocyathus suluensis - CAIRNS & ZIBROWIUS, 1997: 125, fig. 16d. — Cairns, 1998: 382.
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 522, I (MNHN). — Stn 529.
2 (USNM 98672). — Stn 530, 4 (MNHN). — Stn 532. 3 (MNHN). — Stn 534. 17 (USNM 98674). — Stn 535. 14 (USNM
98671). — Stn 540, 74 (MNHN). — Stn 541, 3 (MNHN). — Stn 546. 1 (MNHN). — Stn 557. I (MNHN). — Stn 575.
1 (MNHN). — Stn 578, 1 (MNHN). — Stn 590. 6 (MNHN). — Stn 594. 2 (MNHN). — Sin 595, 3 (USNM 98673). —
Stn 631, I (MNHN).
Vanuatu. Musorstom 8: stn 975. 7 (USNM 98669). — Stn 1028, 3 (USNM 98670).
Type Locality. — " Siboga " Stns 95 and 100: Sulu Archipelago, 450-522 m.
Remarks. — Deltocyathus suluensis is characterised by having a relatively thin, Hat. costate, coarsely
granular base; a finely serrate (not lancetted) calicular edge, the lower cycle septa (c.g., Si-3) projecting beyond the
higher cycle septa (e.g., S4-5); 5 cycles of septa (96 ) above a CD of 18 mm; and rudimentary S5, joined to adjacent
S4 close to the columella by several slender processes. The largest known specimen (MUSORSTOM 7 stn 540) is
22.3 mm in CD. The species is described in greater detail by Cairns (1995) as D. formosus. and included in a key
to western Pacific Deltocyathus species by Cairns & ZIBROWIUS (1997), who also figured one of the synlypes.
It is one of four species in the genus that has 5 cycles of septa in the adult stage, three of which are reported
herein.
Distribution. — Wallis and Futuna region: Wallis, Waterwitch, Combe, Tuscarora, Field, and Bayonnaise
Banks; 400-650 m. Vanuatu region: Tanna and Efate; 566-624 m. Elsewhere: western Australia; Philippines to
ridges north of New Zealand; 142-565 m (Cairns & Zibrowius. 1997).
Deltocyathus taiwanicus Hu. 1987
Figs 12 a-b
Deltocyathus taiwanicus Hu, *1987: 39, pi. 1. figs 1, 4-5. 10.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 522, 11 (MNHN). — Stn 523
1 (MNHN). — Stn 525, I (MNHN). — Stn 529, 12 (USNM 98680). — Stn 535. I (MNHN). — Stn 537. I (MNHN). —
Sin 540. 5 (MNHN). — Stn 541, 2 (MNHN). — Stn 542, 6 (USNM 98679). — Stn 546, 4 (MNHN). — Stn 555.
5 (MNHN). — Stn 556. 10 (MNHN). — Stn 557, 1 (MNHN). — Stn 560. 1 (MNHN). — Stn 585, I (MNHN). — Sin 589.
4 (USNM 98678). — Stn 590, 9 (USNM 98675). — Stn 591, 22 (MNHN). — Stn 594. 19 (USNM 98677) — Stn 597
9 (USNM 98676).
Type Locality. Maanshan Mudstone (Plio-Pleistocene). Tantzu Village, Nanwan Bay, Hengchun
Peninsula, southern Taiwan.
Description op Recent Specimens. — Corallum shaped as a shallow bowl, sometimes with a flat base, but
always with upturned outer edges. Centre of base sometimes bears a concave scar or. just as often, is slightly
protuberant. Cal ice circular, but often slightly irregularly formed; margin not lancetted or serrate. Largest specimen
reported above (MUSORSTOM 7 stn 541) 15.6 mm in CD; however the fossil holotype is 19.1 mm in CD. Costae
equal in width and rounded, each bearing coarse (0.2 mm in diameter), unilinearly arranged granulations near
Source :
AZOOXANTHELLATE SCLERACTINIA
93
the centre of base, which rather abruptly grade into very fine (0.05 mm in diameter) granulations arranged 4-7
across a costa nearer the calicular edge. Intercostal furrows deeply incised only near calicular edge. Well-preserved
coralla have a light reddish-brown base.
Septa hexamerally arranged in 4 complete cycles and a portion of the fifth: Si-2>S3>S4>S.v Between a CD of
about 6.5-7.0 mm, pairs of S5 begin to appear, up to an observed maximum of 13 pairs, or a total of 74 septa,
e g-, in a specimen 12.4 mm in CD (Musorstom 7 stn 546) as well as the holotype. A specimen of CD
13.2 mm (Musorstom 7 stn 556) has 5 pairs of S5 (58 septa) and the largest specimen (CD = 15.6 mm) I I pairs
(70 septa), thus the correlation between CD and number of septa is not always direct. There also appears to be no
order in which the S5 pairs arc inserted, some half-systems in the same corallum having 2 pairs, 1 pair, or no S5.
Si -2 about 1.2 mm exsert, extending about half distance to columella, where each is separated from its broad palus
(up to 1 .7 mm wide) by a deep notch; however, the 2 Pi associated with the principal septa are noticeably smaller,
only about 0.9 mm wide. S3 about 0.7 mm exsert and half the width of the S1-2, each bordered by a P3 about
1.4 mm wide that is slightly recessed from the columella, its axial edge strongly fused to its adjacent P2.
S4 dimorphic in size. If unflanked by S5, S4 are rudimentary, well developed only at the calicular edge and
represented by discontinuous spines within the theca, joining to their adjacent S3 by 4 or 5 thin (0.15 mm in
diameter) processes well below the S3-P3 notch (Fig. 12 b). However, if S4 are flanked by a pair of S5, they are
2/3 the width of an S3 and bear pali (P4) of equal size to the P3, the axial edges of which are strongly fused to the
p3 and recessed slightly more from the columella than the P3. In this case the S5 resemble the unflanked S4 as
described above. All septal and palar faces are covered with spinose granulations. Fossa shallow, containing
a papillose columella consisting of 10-22 granular elements, each about 0.3 mm in diameter. Columellar elements
arranged in an elongate ellipse, the greater axis aligned with the 2 principal septa (by definition), which confers
a bilateral symmetry to the corallum.
Remarks. — Deltocyathus taiwanicus is very similar to I), suluensis , equal-sized coralla of both species
having a similar septal insertion pattern and number of septa, and both species being found at many of the same
stations. D. taiwanicus , however, seems to have a smaller corallum, a specimen of D. taiwanicus having a thick,
upturned calicular edge at the same CD that a D. suluensis would have a thin, flat, serrate edge, characteristic of a
juvenile corallum. D. suluensis ultimately achieves a full fifth cycle (96 septa) and a CD of 22.3 mm, whereas the
largest D. taiwanicus is 19.1 mm (holotype) and yet no specimen is known to have over 74 septa. Furthermore,
the costae of D. taiwanicus are finely granular at the calicular edge, whereas they are coarsely granular on
D. suluensis . Also, the columellar elements of D. taiwanicus appear to be coarser and more independent than those
in D. suluensis.
Although the three specimen type scries was not examined (deposited at the Taiwan Normal University,
Taipei), Hu's description and figures were considered adequate to identify the species.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch, Combe. Tuscarora, and Field
Banks; 320-697 m. Elsewhere: Plio-Pleistocenc of southern Taiwan (Hu. 1987).
Deltocyathus vaughani Yabe & Eguchi, 1932
Deltocyathus vaughani Yabe & Eguchi, * 1932a: 388-389. — Cairns, 1994: 54-55. pi. 23, figs i-j, pi. 24. figs a-c. f
(synonymy). — Cairns & Zibrowius. 1997: 122.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 1006. 1 (MNHN). — Stn 1011. 1 (MNHN).
Type Locality. — Bosyu (= Awa), Japan (depth not given).
Remarks. — The species is distinguished from others in the region by having four cycles of septa, pali or
paliform lobes before all septa, coarsely dentate costae, and equally exsert septa. The species can attain a CD of
27 mm and often has a patellate corallum. the basal angle ranging from 130° to 170°. It is fully described and
illustrated by Cairns (1994), and included in a key to western Pacific Deltocyathus in Cairns & Zibrowius
Source
94
S. D. CAIRNS
(1997). Although coarsely dentate costae are diagnostic for the species, it is interesting to note that the specimen
from MUSORSTOM 8 stn 1006 (CD = 20.4 mm) has finely granular costae.
DISTRIBUTION. — Vanuatu region: Erromango and Efate; 585-919 m. Elsewhere: western Pacific from Japan
through Indonesia; 88-1097 m (CAIRNS & ZlBROWlUS, 1997).
Deltocyathus crassiseptum sp. nov.
Figs 12 c-f
Material EXAMINED/TYPES. — Wallis and Futuna region. MUSORSTOM 7: stn 51 1, 2 paratypes (MNHN). —
Stn 523, 6 paratypes (MNHN). — Stn 529. 13 paratypes (USNM 98683). — Stn 537. 38 paratypes (MNHN). — Stn 570.
3 paratypes (MNHN). — Stn 585, 8 paratypes (USNM 98684). — Stn 586, 1 paratypc (MNHN). — Stn 597, I paratype
(MNHN). — Stn 604, 5 paratypes (MNHN). — Stn 608, 2 paratypes (MNHN). — Stn 618, 1 paratype (MNHN).
Vanuatu. MUSORSTOM 8: stn 958. 9 paratypes (USNM 98682). — Stn 959, 14 paratypes (MNHN). — Sin 977,
2 paratypes (USNM 98681). — Stn 978, 3 paratypes (MNHN). — Stn 980, holotype (MNHN). — Stn 983. 5 paratypes
(USNM 98686). — Stn 1061, 2 paratypes (MNHN). — Stn 1068, 2 paratypes (USNM 98685).
Type Locality. — MUSORSTOM 8 stn 980: 19°2rS, 169°25'E (Tanna), 433-450 m.
ETYMOLOGY. — The species name crassiseptum (Latin crassus, thick + septum, partition) refers to the thick
Si -2- The name is treated as a noun in apposition.
DESCRIPTION. — Corallum shaped as a small, shallow bowl, with a flat to slightly convex base and upturned
calicular edge. Largest specimen (holotype) 14.1 mm in CD and 5.8 mm in height. Calice circular; theca relatively
thick. Costae rounded and finely granular, the granules changing to slender spines near calicular edge. Costae
separated by deep intercostal grooves at calicular edge, near point of upward thecal inflection. In most coralla, each
intercostal groove is bisected by a low. narrow (0.1 mm wide) row of granules (Fig. 12 f). Most coralla bear
evidence of a former scar of detachment located at or near the centre of base. This scar may be pear-shaped in
outline, circular, or an irregularly-shaped depression. Circular-shaped scars may also occur in various diameters,
those of 0.8 mm diameter usually showing the traces of 6 larger and 6 smaller septa; larger scars of 1.3 mm
diameter or more showing 24 septa. In some cases there are 2 concentric scars, with different numbers of septa.
Most coralla are uniformly white, but some well-preserved coralla (e.g.. the holotype) have a light reddish-brown
colour to the calicular edge.
Septa hexamerally arranged in 4 complete cycles (S|>S2>S3>S4), for a total of 48 septa; however, the
holotype is missing one half-system, resulting in 1 1 major septa and a total of only 44 septa. Si about 2 mm
exsert, extend about half way to the columella, having straight, vertical to slightly concave axial edges. Si can be
remarkably thick, up to 1.0 mm at the calicular edge. S2 only slightly smaller than S| and equally thick. S3 about
half as exsert and wide as S1-2, and of normal thickness (i.e., 0.4 mm). S4 rudimentary in small coralla. but up to
3/4 exsertness and width of an S3 in larger coralla. Lower axial edges fuse to adjacent S4 low in fossa adjacent to
columella through 3 or 4 slender processes. Pi -3 all about 1 mm wide, the 6 Pi being the only independent pali.
forming a crown low in the fossa encircling the columella. The 6 P2 rise slightly higher in the fossa, and the
12 P3 higher still and are recessed from the columella, the axial edges of each pair of P3 fusing to its adjacent P2
in the typical deltocyathid chevron arrangement. Fossa moderately deep, containing a papillose columella
composed of many fine interconnected elements.
Remarks. — Among the approximately 24 Recent Deltocyathus species, D. crassiseptum can most easily be
distinguished by its unusually thick S1-2. Other consistent characters include its relatively small size, basal scar,
deep peripheral intercostal grooves, and the small rows of granules that bisect each intercostal groove. It also
appears to be restricted to a rather narrow bathymetric (i.e., temperature) range.
One corallum from MUSORSTOM 7 stn 585 showed evidence of a petrarcid ascothoracidan gall beneath its
columella, a symbiosis previously reported in this genus by ZlBROWlUS & GRYGIER (1985), ORYGIER (1991), and
Grygier & Nojima (1995).
Source :
AZOOXANTHELLATE SCLERACTINIA
95
Distribution. — Wallis and Futuna region: Wallis. Fuluna, and Alofi; Waierwitch and Field Banks:
420-510 m. Vanuatu region: Anatom, Tanna, and Malakula; 413-536 m.
Deltocyathus cameratus sp. nov.
Figs 12 g-i, 13 a
MATERIAL EXAMINED/TYPES. — Wallis and Futuna region. MUSORSTOM 7: sin 520. 2 paratvpes (MNHN). —
loP/oo!yPe (MNHN). — Stn 537. I paralype (MNHN). — Stn 541. 1 paratype (MNHN). — Stn 542, 4 paratypes
(I SNM 98688). — Stn 546. I paratype (MNHN). — Stn 552. I paratype (USNM 98690). — Stn 555. 1 paratype
~ „Sln 557' 3 ParatyPes (USNM 98691). — Stn 560, I paratype (USNM 98687). — Stn 567, 1 paratype
(MNHN). — Sin 569. I paratype (USNM 98689). - Stn 578, 2 paratypes (MNHN). — Stn 589. 2 paratypes (MNHN). -
Stn 597. 1 paratype (MNHN). — Stn 635, 18 paratypes (MNHN). — Stn 636. 3 paratypes (MNHN)
Vanuatu. Musorstom 8: sin 956, 4 paratypes (USNM 98693). — Stn 992, 1 paratype (USNM 98692). — Stn 1007
holotype (MNHN). — Stn 1036, 4 paratypes (MNHN). — Stn 1061. 1 paratype (MNHN).
Type Locality. — Musorstom 8 stn 1007: I8°52'S. I68°52'E (Erromango), 720-830 m.
Etymology. — The species name cameratus (Latin camerata, chambered) refers to the 24 elliptical chambers
formed by the robust S4-P3 and P2-P3 fusions.
DESCRIPTION. — Corallum shaped as a shallow bowl, with a flat or slightly convex base. Holotype 13.7 mm
in CD and 5.2 mm in height; largest specimen (MUSORSTOM 7 stn 557) 15.2 mm in CD. Calice circular but with
a jagged margin, the 12 CS3 and adjacent pairs of CSj projecting outward as short (about 0.9 mm) triangular to
rectangular lancets. Costae inconspicuous except at calicular edge, where they are separated by intercostal grooves.
Costae on base covered with a low granulation, changing to slender spines at calicular edge; no attachment scar.
Most coralla uniformly while, but some (e.g., the holotype) are a light reddish-brown in the palar region.
Septa hexamerally arranged in 4 complete cycles (S|>S2>S4>S3). In a large well-preserved corallum, Si are
about 1.6 mm exsert, extending about half distance to columella; S2 similar in shape and exsertness but only
slightly less wide. S3 about 1.3 mm exsert and half the width of an Si. S4, although less exsert than the S3 (about
1 .0 mm), arc slightly wider than the S3, the axial edges of each S4 pair solidly fused as a thick lamella to the outer
edge of the adjacent P3. This fusion reaches as high as the S3-P3 notch and is thick and solid except near the
columella, where it is perforated with several small pores. All septa uniformly thin and have straight axial edges.
Pali (Pi -3) uniform in width (about 1.2 mm) and separated from their corresponding septa by wide notches (about
0.8 mm). Axial edges of Pi and P2 solidly fused to the columella, although P2 rise slightly higher in the fossa.
Axial edges ot each pair of P3 solidly (used to their adjacent P2. this fusion being imperforate and reaching as high
as the S3-P3 notch. The fossa is shallow to nonextant. containing a well-developed papillose columella consisting
of 10-15 robust (0.3-0.6 mm in diameter), granular rods, in some coralla ornately sculptured (Fig. 13 a).
Remarks. — The well-developed lamellar fusions of the S4 to P3 and P3 to P2 serve to differentiate this
species Iront all others, as well as subdivide the corallum into 24 elliptical compartments, or chambers. In each
system there are 2 small chambers formed by each pair of S4. bisected by the S3; I slightly larger compartment
formed between each S3, bisected by the S2; and an elongate compartment between each system, bisected by an S|.
This compartmenlalization is better seen in a worn specimen in which the bisecting septa are reduced or missin«
(Fig. 12 i). At least two other species. D. pourtalesi Cairns. 1979 and D. italicus (Michelotti. 1838). both
Atlantic species, have similarly high S4-P3/P3-P2 fusions, joining at or above the notch that separates septum
from palus, but in neither case are these fusions as robust, and in both cases there are many other differences
among the 3 species.
On corallum from Musorstom 7 sin 557 showed evidence of a petrarcid ascothoracidan gall beneath its
columella.
Distribution. — Wallis and Futuna region: Wallis; Waterwitch. Combe. Tuscarora, Field, and Rotumah
Banks; 305-1010 m. Vanuatu region: Erromango and Malakula; 512-1 175 m.
Source
96
S. D. CAIRNS
Deltocyathus Stella Cairns & Zibrowius, 1997
Figs 13 b-c
Deltocyathus Stella Cairns & Zibrowius, *1997: 123-124, figs 15 f-h.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 509. 6 (USNM 98695). — Stn 510.
1 (MNHN). — Stn 511, 1 (MNHN). — Stn 512, 41 (MNHN). — Stn 513. 22 (MNHN). — Stn 514. 4 (MNHN). — Stn
516, 5 (MNHN). — Sin 523, 11 (USNM 98694). — Stn 537, 1 (MNHN). — Stn 541, I (MNHN). — Sin 556. 4 (MNHN).
— Stn 569, 6 (USNM 98696). — Stn 585, 1 (USNM 98697). — Stn 594, I (MNHN). — Stn 605. 16 (USNM 98698). -
Stn 610, 6 (MNHN). — Stn 626, 1 (MNHN).
Vanuatu. MUSORSTOM 8: sin 967. 3 (USNM 98700). — Sin 969. 4 (MNHN). — Stn 1097, 1 cemented to A enophora
shell (MNHN). — Stn 1 106, 1 (MNHN).
Type Locality. — Karubar stn 35: 5°46'45"S. 1 32° 1 1T0"E (Kai Islands, Banda Sea). 156-305 m.
Table 6. — Comparison of the spined species of Deltocyathus.
Source :
AZOOX ANTI IELLATE SCLERACTINIA
97
Remarks. Little can he added to the original description or this recently described species except to note
that in well-preserved coralla the distal and axial edges of the P3, and occasionally the Pi and P:. are dentate.
Deltocyathus siellci is most easily distinguished from other spined Deltocyathus (Tabic 6), by having short, thick,
stubby costal spines, and large, sinuous, dentate P3.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Combe. Tuscarora, Waterwitch, Bayonnaise,
and Field Banks; 240-597 m. Vanuatu region: Anatom and Espiritu Santo; 280-305 m. Elsewhere: Philippines and
Indonesia; 206-280 m (Cairns & Zibrowius, 1997).
Deltocyathus heteroclitus Wells, 1984
Figs 1 3 d-g. Fig. A
Deltocyathus heteroclitus Wells. v*!984: 210. figs 3. 1-6. — Cairns & Zibrowius. 1997: 124 (mentioned).
987(flATER,AL EXAM,NED' “ Wallis and Futuna region. MUSORSTOM 7: stn 514. 16: 12 (MNHN). 4 (USNM
V anuatu. Musorstom 8: stn 1 106. 2 cemented to Xenophora shells (MNHN).
Type Locality. — USGS stn 24918: Navaka River, Espiritu Santo, Vanuatu (Late Pleistocene).
Desc ription. Corallum shaped as a shallow bowl, the largest known specimen (Musorstom 7 stn 514)
5.4 mm in CD (exclusive of costal spines). Costae coarsely granular and not well defined. Six to eight robust
costal spines (C3) project up to 1.5 mm from the calicular edge, spines forming only in half-systems in which the
CS3 is flanked by a pair ol CS4 (see Remarks). Costal spines thick at base, circular in cross section, attenuate at
the tip, and coarsely granular. Corallum primarily white, but with a circular band of light red-brown pigmentation
in palar region.
Septa hexamerally arranged in 4 cycles, but usually only I pair of S4 occurs in each system, resulting in
36 septa (Fig. A, but see Remarks). Si only independent septa, each bordered axially by a slender (0.4 mm wide)
palus. S2 about 2/3 width ol the Si, each bearing a similarly sized palus, but rising higher in the fossa.
S3 dimorphic in size: S3 unflanked by S4 about 3/4 width of the S2. their axial edges fused to the adjacent P2, and
olten bear a small pal i form lobe. S3 flanked by S4 are the smallest of septa, seeming to rest on the centre of the
costal spine, each having a dentate upper edge and bordered by a prominent palus. P3 wide (0.6 mm), sinuous,
highly granular, and are the tallest and most recessed pali from the fossa. S4 slightly wider than flanked S3, have a
dentate distal margin, a peripheral margin that borders the costal spine, andean axial margin that fuses to the
adjacent P3 at the level of the S3-P3 notch. Fossa shallow, containing a papillose columella composed of 5-9 well-
formed (0.25 mm in diameter), finely granular elements.
Remarks. — I (Cairns, 1995: 72) questionably considered D. heteroclitus to be a junior synonym of
D. ornatus Gardiner, 1899; however, this decision was based on a misconception of the latter species and the
knowledge of only the small type series of the former. The additional Musorstom specimens and subsequent
examination of the type of D. ornatus show the two species to be distinct, as compared in Table 6. This is
considered to be the lirst report of D. heteroclitus subsequent to its original description and the first record from the
Recent, the types having been collected from the Late Pleistocene.
The pattern of costal spine insertion (as well as S4 pair insertion) in D. heteroclitus is identical to that of
Anthemiphyllia spinifera (Fig. A), the first 6 spines occurring in the "anterior” half-system of each system (i.e.,
half-systems I, III, V, VIII, X, and XII), which produces the "roughly hexagonal" calicular perimeter noted by
Wells (1984). The majority of specimens (the holotype, 4 paratypes, and 12 of the MUSORSTOM specimens) have
this complement, i.e., 6 costal spines and 36 septa. One paratype and 4 Musorstom specimens have 7 costal
spines and 38 septa, the additional costal spine occurring in half-system VI in two specimens, half-system VII in
one specimen, and half-system IX in two specimens. Two Musorstom specimens have 8 costal spines and
40 septa, one corallum being poorly preserved, the other with the additional costal spines in half-systems VI and
VII (Fig. 13 g).
Source
98
S. D. CAIRNS
Distribution. — Wallis and Futuna region: Futuna; 349-355 m. Vanuatu region: Espiritu Santo; 208-210 m
and Late Pleistocene of Espiritu Santo (Wells. 1984).
Deltocyathus ornatus Gardiner, 1899
Figs 13 h-i
Deltocyathus ornatus Gardiner, v* 1899a: 163-164, pi. 20. figs 25a-b.
Not Deltocyathus ornatus - Cairns, 1995: 72-73, pi. 19, figs d-e (=D. corrugatus n. sp.. described below in Remarks).
Material EXAMINED. — Wallis and Futuna region. Musorstom 7: sin 569, 1 (MNHN).
Vanuatu. Musorstom 8: stn 964, 2 (USNM 98703). — Stn 967. 1 (MNHN). — Sin 1017, 1 (MNHN). — Stn 1018, I
(USNM 98702). — Stn 1094, 2 (MNHN).
Type Locality. — Sandal Bay, Lifu, Loyalty Islands, 73 m.
Remarks. — Deltocyathus ornatus is compared to the other spined Deltocyathus in Table 6. It can be
characterised as having a circular calice, 12 relatively short costal spines (C3), dimorphic Pi, and an S4-P3 junction
slightly below the S3-P3 notch. Before I examined the holotype of D. ornatus (deposited at the University Museum
of Zoology, Cambridge), I (CAIRNS, 1995) incorrectly identified several specimens from New Zealand as this
species, based on the resemblance in size and costal spines. It is now clear that the New Zealand specimens differ
in having a strongly lancetted calicular margin; C3 so prominent that they give the base of the corallum a
corrugated aspect; much longer costal spines; a higher S4-P3 fusion; and a flatter corallum (Table 6). To this, as
yet unnamed, species I propose the name Deltocyathus corrugatus, new species, described and figured by CAIRNS
(1995) as Deltocyathus ornatus, and diagnosed and compared in Table 6, herein. The holotype is the specimen
figured by Cairns (1995: pi. 19, figs d-e, ex USNM 94169. now NZOI H 689) from NZOI stn P27 (28°54'36"S.
167°44T2'E. Norfolk Island. 390 m, = type locality), and the other 15 specimens from four NZOI stations reported
by Cairns (1995) as D. ornatus are considered as paratypes. The name corrugatus (Latin corrugatus , ridged) refers
to the corrugated base.
Distribution. — Wallis and Futuna region: Waterwitch Bank; 300-305 m. Vanuatu region: Anatom. Efate,
and Espiritu Santo; 295-360 m. Elsewhere: Loyalty Islands; 73 m (holotype).
Genus HETEROCYATHUS H. Milne Edwards & Haime, 1848
Heterocyathus sp. cf. H. sulcatus (Verrill, 1866)
Figs 14 a-d
Stephanoseris sulcatus Verrill, v*1866: 48.
Heterocyathus sulcatus - HOEKSEMA & Best, 1991: 231-233, figs 19-23 (synonymy).
Material examined — Wallis and Futuna region. Musorstom 7: stn 494. 4 (USNM 98717). — Stn 495.
16 (USNM 98716). — Stn 496. 1 + (3) (USNM 98715). — Stn 509. 5 (USNM 98713). — Stn 510. I (MNHN). —
Stn 512. 19 + (6) (MNHN). — Stn 513. 70 + (3): 60 + (3) (MNHN), 10 and SEM stubs 886-888 (USNM 98712 ). —
Stn 514. 2 (MNHN). — Stn 516, 2 + (2) (MNHN). — Stn 556, 1 (MNHN). — Stn 570. 1 (MNHN).
Vanuatu. MUSORSTOM 8: stn 969. 1 (MNHN). — Stn 976, 17 + (2) (USNM 98714). — Stn 1069. 10 (USNM 98718).
— Stn 1094, I (MNHN). — Sin 1097, 2 (MNHN).
Type Locality. — Ceylon (= Sri Lanka), depth not reported.
Description. — All specimens reported above arc unattached, the base usually forming a low, flattened
cylinder that completely encapsulates a small gastropod that was colonized by a sipunculid worm. If the gastropod
was small or circular in diameter, the concentric internal sipunculid tube forms a regular cylinder, whereas if the
Source : MNHN , Paris
AZOOXANTHELLATE SCLERACT1NIA
99
h^rPhd r e'0ngatf' an irregularly-shaPed base results. A cylindrical corallum of smaller diameter sits upon the
b,se wh.cb contams the coral polyp. The corallum base is up to 6.0 mm in diameter and about 2.5 mm in heigh,
^ab 2 r:„THU: y CXCeedS 4 0 mm in diamelCr and ,S alS0 “P ,0 2 5 - sipunculiZS
ateraTsu face of 1 ZT T“5 *° °f ,he C°ral lhr0ugh a pore of etlual diamcIcr >he lower
in diameter n base. Smaller efferent pores associated with the sipunculid are rare, occasionally one 0.2 mm
oundeTand ve^fiir ,?WCrhsur&ce of ,hc basc or the upper edge of the base. Costae well-developed.
smomh^ norc L " ,s 8T ’r F’ " ^ COra"a tCC,Ural dcP°si,S cover the costae resulting ,n
LcZn bunhe 0 u h rThC- °Slae °CCUr °n'y °n laleral faces oflhe corallum, including both upper and lower
base that covers he S COra I IS sm00,h- The C0S,al Width musl 8radual|y Crease i" *c region of the
the worrl ln, , -T worm ,ube- ln order 10 cover the larger circumference formed bv the housing of
Ci 7ZZ ' rf h TWS K1"'6 (ab°Ul 30 Mm) and deep' such ,hat the underlying theca cannot be sL.
1-3 light brown to chocolate-brown in colour, alternating with the white C4; however, only the S|.2 are similarly
Demented he,,Plgment °‘ "le C' St°Ppi"g at lhe calicular edgc‘ Palar and columellar elements usually also
pigmenicd, bui in some cases are white. y
mirSseof%heXamua"y arr^ged !" 4 CydeS’ bU‘ 2 C°mpleIC f0Ur,h cycle is rare- mosl uoralla missing I or more
co um^la anTh 7 h k !° 1 S|>S2>S4>S3' S< ab- '•> mm exsen. extend about half way to
columella and bordered by 1 or 2 narrow paliform lobes. S2 0.7 mm exsert. about 3/4 width of the S,. and also
the sL of th'e'sTh , ?S' nV‘7 ^ ^ “ PU'r °f Sa are ,he smallest *P*. only 0.5 mm exser, and half
the size of the S2> but if not Hanked by S4. an S3 is as wide as an S2 and often considerably thicker. S4 also
tmorph.c in size, those adjacent to Si as w.de as an S2 and fused to its adjacent Si in a lancet. S4 adjacent to Si
are smaller, only slightly wtder than an S3. Both S3 and S4 bear 2 or 3 slender paliform lobes. Septa crowded in
arrangement and bear prominent granulations on their faces. All paliform lobes slender, not lamellar, about
consi^inC.lnr inTster',a?. 7, °rientCd menianes' Fossa shallow, containing a papillose columella
const tmg of 10-15 cylindrical elements that are similarly transversely ridged as the paliform lobes. In fact there is
with particular septet 0 ? and C0'Ume"ar e'emenlS except lhal ,he former rise higher in the fossa and are aligned
Remarks. — Hoeksema & Best (1991) consolidated the 23 described species and subspecies of
Heterocyathus into three species and provided a key for their differentiation. According to their diagnoses. coralla
having a dark brown centre, alternating costal pigmentation, and a compact, cylindrical upper corallum, should be
identified as H sulcata. However. I have noted that several specimens in larger lots that were certainly conspecific
did not have a dark brown centre, and I have also noted other specimens of Heterocyathus with dark brown centres
that were otherwise very diiferent from the species described herein. The type of H. sulcata (YPM 764) from Sri
Lanka is now so damaged that it is of little use in defining the species; only about one-third of the original
corallum is present. Because of the large number of nominal species and the variation inherent in this genus these
specimens are only provisionally identified as H. sulcatus.
Specimens from six of the lots listed above, indicated with parentheses around the specimen number, differ in
laving a larger corallum (CD up to 6.0 mm. basal diameter up to 8.5 mm) and in having 2-6 sipunculid efferent
pores located circumferentially at the junction of the base and upper corallum. These are assumed to be larger
individuals ol the same species, the larger size mediated by the increased development of the sipunculid.
, inD,,SRz!fUI10N‘ ~ Wa"iS and Fl"Una region: Fuluna: Tuscarora and Waterwitch Banks: live specimens from
(dead specimens found to 550 m). Vanuatu region: Anatom. Tanna. and Espiritu Santo; 182-3P m (live
specimens). Elsewhere: See HOEKSEMA & BEST (1991).
Heterocyathus alternatus Vcrrill, 1865
Figs 14 c-f
V*'865: '49' ~ H°EKSEMA & BEST- 199l: 230-23F 12-18 (synonymy)
Source :
100
S. D. CAIRNS
MATERIAL EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1004, I (MNHN). — Stn 1018. 4 (USNM 98719).
Type Locality. — "Gaspar Straits" = Selat Gclasa, between the islands of Bangka and Belitung, Sumatra,
Indonesia.
Diagnosis (specimen from Musorstom 8 stn 1004). — Corallum cylindrical and free: 10.2 x 9.4 mm in CD
and 8.1 mm in height. Costae well defined, rounded, and finely granular (not porcellaneous); intercostal grooves
narrow and relatively deep. Base flat and not costate, but covered with fine granulations; the sipunculid tube
opening and one efferent pore also occur on the flat base. Septa hexamcrally arranged in 4 complete cycles
(Si>S2>S4>S3), although Hoeksema and BEST (1991: fig. 17) show that this species may sometimes attain
a full fifth cycle. Si highly exsert (3.7 mm), forming tall lancets with their adjacent S4. and axially bordered with
2 or 3 slender paliform lobes. S2 about 2.5 mm exsert. also forming lancets with their adjacent S4. S3 bear only
1 or 2 paliform lobes, whereas the larger S4 bear 4 or 5 lobes that merge indisti nguishably into the columella.
All paliform lobes are slender (cylindrical) and not ridged. Fossa quite deep, containing a papillose columella
composed of numerous fine elements.
Remarks. — These specimens appear to be conspecific with the diagnosis and illustrations of Hoeksema and
Best (1991). The holotypc (YPM 6828), although well preserved, appears to be a juvenile specimen.
Heterocyathus alternatus differs markedly from the species described above as H. cf. sulcatus by having a larger
corallum with more septa, granular costae (not porcellaneous) and base, more highly exsert Si and S2. nonridged
paliform lobes, a deeper fossa, and finer columellar elements.
Distribution. — Vanuatu region: Erromango and Efale; 301-319 m. Elsewhere: Indonesia; Philippines;
northern Indian Ocean; Western Australia; South China Sea (Hoeksema & Best, 1991 ).
Genus CONOTROCHUS Seguenza, 1864
Conotrochus funicolumna (Alcock, 1902)
Ceratotrochus (Conotrochus) funicolumna Alcock. v* 1902a: 93; v. 1902c: 11-12. pi. 1, figs 6. 6a.
Conotrochus funicolumna - Cairns. 1994: 58-59. pi. 24. fig. i, pi. 25. figs g-1 (synonymy). — Cairns & Zibrowius.
1997: 127. — Cairns, 1998: 385.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 511. 6 (MNHN). — Stn 522.
1 (MNHN). — Stn 523. 5 (MNHN). — Stn 525. 2 (MNHN). — Stn 532, 1 (MNHN). — Stn 535, 1 (MNHN). — Stn 537.
1 (MNHN). — Stn 540. 4 (MNHN). — Stn 541, 1 (MNHN). — Stn 542. 1 (MNHN). — Stn 555, 1 (USNM 98721). —
Stn 556, 3 (MNHN). — Stn 557, 4 (USNM 98720). — Stn 560. 1 (MNHN). — Stn 570. 12 (MNHN). — Stn 571.
2 (MNHN). — Stn 585, 3 (USNM 98724). — Stn 604, 3 (MNHN). — Stn 618, 40 (USNM 98722). — Stn 619. 14 (USNM
98723).
Vanuatu. Musorstom 8: stn 1059, 1 (USNM 98728). — Stn 1060. 1 (MNHN). — Stn 1065, 8 (USNM 98726). —
Stn 1072. 3 (MNHN). — Stn 1087. 8 (USNM 98729). — Stn 1088, 1 (USNM 98725). — Sin 1091. 1 (MNHN). —
Stn 1 107, 1 (USNM 98727). — Stn 1113, 1 (MNHN).
Type Locality. — "Siboga" stns 95 and 100: Sulu Archipelago, 450-522 m.
Diagnosis. — Corallum ceratoid (edge angle up to 35°), straight, and attached by a narrow pedicel
that is usually reinforced by an extensive secondary attachment along the lower edge of the corallum
contiguous with the pedicel. Calice circular to slightly elliptical, the largest specimen reported herein 12.5 mm in
GCD. The theca of a well-preserved corallum often bears slender, hollow, hispid granulations; worn coralla
often show longitudinal costae. Theca of well-preserved coralla sometimes longitudinally streaked with brown
pigmentation that occurs without symmetry, the streaks ranging from 1 to 8 septa in width, alternating with
white theca. Theca thick, projecting 0.5-0. 7 mm above the peripheral distal septal edges as a continuous rim;
Source :
AZOOXANTHELLATE SCLERACTINIA
101
stcreome present internally. Septa hexamerally arranged in 4 cycles (Si>S2>S3>S4), a full fourth cycle complete
in some coral a at a CD of 4.6 mm, hut larger specimens sometimes lacking several pairs of S4. Columella
a well-formed, discrete concentration of lamellar elements, often swirled in a circular manner, and occa¬
sionally fused into a more solid structure. Lamellar elements often fused to one another in a lahyrinthiform
arrangement. J
Remarks. — Conotrochus funicolumna is similar to C. bnmneus , both species secondarily attached basally
and having a similar thecal structure. C. funicolumna differs in having: a larger corallum (up to 12.5 mm CD
vs <8.0 mm for C. brunneus)-, more septa at the same CD. most C. funicolumna have 48 septa, whereas most
C. bnmneus have only 36-40 septa; a more open corallum (slightly higher edge angle); less internal stereome; and
occasionally a streaked theca, whereas that of C. brunneus is white or uniformly brown. The species is more fully
described and figured by Cairns (1994).
D'STRIbution. - Wallis and Futuna region: Wallis, Futuna, and Alofi; Waterwitch, Combe, and Tuscarora
Banks; 370-697 m. Vanuatu region: Malakula and Espiritu Santo; 350-700 m. Elsewhere: Philippines; Indonesia;
Japan; Australia; Hawaiian Islands; 88-616 m (Cairns & Zibroyvius, 1997).
Conotrochus brunneus (Moseley, 1881)
Pleurocyathus brunneus Moseley, v*1881: 159-160, pi. 2, figs la-e
C^oirochus brunneus - Cairns. 1995: 74-75. pi. 20. figs a-b (synonymy). - Cairns & Zibrowius, 1997: 127-128. fig.
1 “™IA1; ~Wallis and FuIuna re8ion Musorstom 7: stn 511. 1 (USNM 98734). — Sin 513
59oTmJ™ r^^Sln 5c°- ' ,USNM 9873"- - S,n 546' 1 <MNHN). - Stn 586. I iMNHN). - Sin
5 JO, 1 (MNHN). — Stn 591, 1 (MNHN). — Stn 597, 1 (MNHN). — Stn 625, I (USNM 98730)
, ,MVMu“ntU 8. stn 959. 2 (USNM 98732). - Sin 977. 1 (MNHN). - Sln 1006. I (MNHN). - Sin 1089,
(MNHN). Stn 1090, 1 (USNM 98733). — Sin 1092. I cemented to a Xenophora shell (MNHN).
Type Locality. — "Challenger" stn 194: 4°34'S, 129o57'30''E (Banda Island. Banda Sea). 366 m.
Remarks. — Conotrochus brunneus was redescribcd and figured by Cairns (1995), and compared to
C. funicolumna in the previous account.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch. Combe, Field, and Bayonnaise
Banks; 300-580 m. Vanuatu region: Anatom. Tanna, Erromango, and Espiritu Santo; 321-574 m. Elsewhere:
Indo-West Pacific from Madagascar to South China Sea; 97-1051 m (Cairns & Zibrowius, 1997).
c onoirocnus asymmelros sp. nov.
Figs 14 g, 15 a-e
, EXAMINED/TYPES. — Wallis and Futuna region. Musorstom 7: sin 495. 3 paratypes and SEM
98J35)- — S,n 496‘ 1 holo|yPe (MNHN). - Sin 504, 6 paratypes (MNHN). — Sin 509. 10 paratypes
(USNM 98736). — Sin 512. 3 paratypes (MNHN). — Sin 513. 6 paratypes (MNHN). — Stn 516. 3 paraivpes (MNHN) —
Stn 586, I paratype (MNHN). — Stn 610, 1 paratype (MNHN).
Vanuatu. Musorstom 8: sin 1016. 1 nontype cemented to a Xenophora shell (MNHN). — Sin 1017. I nomype
cemented to a Xenophora shell (MNHN). — Stn 1097, 1 nontype cemented to a Xenophora shell (MNHN).
Type Locality. — Musorstom 7 stn 496: 14°I9.6’S, 178°04.3'W (Futuna), 250-330 m.
Etymology. — The species name asynunetros (Greek asymmetros , without symmetry) refers to the
asymmetrical arrangement of septa within the calice.
Source :
102
S. D CAIRNS
Description. — Corallum varies from conical (edge angle = 30°) to flabellate (e.g.. the holotype: edge
angle up to 65°, face angle = 15°), depending on the corallum and its stage of thecal edge spine development.
Holotype 8.9 x 4.6 mm in CD, 7.3 mm in height, and 1.3 mm in pedicel diameter. Calicc elliptical (GCD:LCD =
1.2) to highly compressed (GCD:LCD = 2.0). the higher ratio characteristic of coralla in process of edge
spine formation, which temporarily elongates the corallum. Elliptical calices are sometimes asymmetrical in
shape, one face being almost straight, the other convex. Several more septa often originate from the convex thecal
face than the straight face. Calicular edge finely serrate, separated from the peripheral septal edges by a shallow
trough. Basal disc 0.8- 1.3 mm in diameter, in many cases augmented by a contiguous secondary thecal attachment
up to 2.0 mm in diameter. Thecal edges bear 1-3 pairs of spines (up to 2.0 mm long); short, triangular crests;
or low eversions. Transverse division not noted. Theca porcellaneous, covered with low, rounded
granules characteristic of the genus. Theca usually white, but occasionally longitudinally streaked with brown
pigment.
Septa hcxamerally arranged in 3 size classes, usually resulting in 32 or 30 septa accordingly: 6:10:16 or
6:9:15. The 6 primary septa arc up to 0.7 mm exsert, having straight axial edges that reach the columella. Primary
septa are not aligned with the greater calicular axis, as in all other known corals having an elliptical calice. Rather,
they occur on either side of the calicular axis and on the lateral faces, resulting in a secondary or tertiary septum
being aligned with the greater axis, usually one of each size class on either end of the corallum. Secondary septa
0.4 mm exsert, about 4/5 width and less thick than a primary septum. In coralla having 32 septa, all 6 systems
contain at least one secondary septum and 4 systems contain a pair of equal-sized secondary septa; in coralla having
30 septa, only 3 systems contain a pair of secondaries. There seems to be no consistent arrangement in which
systems contain pairs of secondaries, only that they are usually in contiguous systems and often occur in a system
aligned with the greater axis. Tertiary septa nonexsert and rudimentary, only about 1/4 the width of a secondary,
and quite thin. Fossa of moderate depth, containing a papillose columella composed of 6-11 granular,
interconnected papillae that arc also attached to the lower, axial edges of primary and secondary septa.
Remarks. — The unusual septal symmetry and thecal edge spines distinguish this species from all others in
the genus. In fact, I know of no other scleractinian species that possesses a pair of equal-sized secondary
septa within one system or a septal arrangement that aligns a secondary and/or tertiary septum on the greater
calicular axis.
Distribution. — Wallis and Futuna; 210-510 m. Vanuatu region: Efate and Espiritu Santo; 288-294 m.
Genus LOCHMAEOTROCHUS Alcock, 1902
Lochmaeotrochus gardineri sp. nov.
Figs 15 f-g
Material EXAMINED/TYPES. — Musorstom 7: stn 539. 17 paratypes (MNHN). — Stn 548. I paratype (MNHN).
— Stn 557, 1 paratype (MNHN). — Stn 560, 1 paratype (USNM 98739).
Vanuatu. Musorstom 8: stn 956, 9 paratypes (MNHN). — Stn 992, 3 paratypes (USNM 98738). — Sin 1034,
1 paratype (MNHN). — Stn 1036, holotype (MNHN) and 13 paratypes (USNM 98737).
Type Locality. — Musorstom 8, stn 1036: 18°0rS, I68°48’E (Efate), 920-950 m.
ETYMOLOGY. — This species is named for J. Stanley Gardiner, in recognition of his pioneering work on
azooxanthellate corals from the central and South Pacific (Gardiner, 1899a. b).
Description. — Corallum conical (ceratoid, having an edge angle of 18° -23° ). straight, and free, having a
basal disc about 1.1 mm in diameter that is occasionally reinforced with a small contiguous secondary attachment
of equal diameter. Largest specimen (holotype) 10.8 mm in CD and 17.4 mm in height; calice circular. Theca
covered with slender, hispid granulations, characteristic of the genus Conotrochus. Theca lipped with successive
Source :
AZOOXANTHELLATE SCLERACTINIA
103
circumferential growth ridges or lips, each projecting up to 0.3 mm in height, the uppermost thecal rim projecting
upward about 0.7 mm and separated from the distal, peripheral septal edges by a shallow trough. Theca relatively
thick, containing layers of internal stereome. Corallum uniformly white.
Septa hcxamcrally arranged in 4 cycles, the fourth cycle never complete: S|>S2>S3>S4. Most coralla
have I pair of S4 m each system, resulting in 36 septa; however, the holotype has 40 septa, and a smaller
specimen (CD 7.4 mm) has 42 septa. Si nonexsert. extend about 3/4 distance to the columella, having straight
vertical axial edges. Rarely (e.g„ the holotype) the S, bear a small (0.6 mm wide), lamellar Pi. separated from
its septum by a deep narrow notch. S2 about 3/4 width of the S,. S, dimorphic in size: S3 unHanked by S4 about
alf the size of the S2 but those flanked by S4 are only slightly less wide than the S2 and usually bear a broad
.2 mm wide), amcllar palus. The position of the P3 is variable. Although it is usually aligned with the Si
it is sometimes ahgned between the S3 and adjacent S2, or may be absent. Even when aligned with the Si. lower
m the fossa the P3 is usually connected to the adjacent S2. Also, in some systems having a full complement of
2 pairs of S4, only one palus is present before the S2. but this palus having connections to the 2 adjacent S3 lower
m the fossa. S4 about half width of a flanked S3. Axial edges of S2.4 very slightly sinuous. Fossa moderately
deep, containing a papillose columella composed of 0-10 interconnected cylindrical elements 0.2-0 3 mm
in diameter.
Remarks — Lochmaeotrochus gardineri is similar to but differs from the only other species described in this
genus, L. oculeus Alcock, 1902, in having: a solitary, free habit with a small basal disc (vs a quasicolonial habit
with a broad secondary attachment); circumferential thecal ridges; fewer septa at a corresponding size and a lower
maximum number of septa (most L. oculeus have 48 septa); wider and better developed pali: and a deeper fossa
It is also known from greater depths than L. oculeus.
Lochmaeotrochus gardineri is also similar to Conotrochus funicolumna, particularly in corallum size and
thecal granulation; these species also co-occur at one station. L. gardineri is distinguished by having at least
6 lamellar pali (P3), which are different in shape from its cylindrical columcllar elements, C. funicolumna only
having swirled, interconnected (labyrinthiform) lamellar columellar elements and no pali. L. gardineri also has
a narrower basal attachment, a deeper fossa, fewer septa at a corresponding CD. and a greater disparity between the
width of its Si and S2. L. gardineri also seems to be found, in general, at greater depths than C. funicolumna.
Distribution. — Wallis and Futuna region: Combe and Tuscarora Banks; 608-700 m. Vanuatu region-
Anatom. Erromango. and Efate; 750-1 175 m.
Genus AULOCYATHUS Marenzeller, 1904
Aulocyathus recidivus (Dennant, 1906)
Ceratotrochus recidivus Dennant, *1906: 159-160. pi. 6. figs 1-2.
Aulocyathus q recidivus - Cairns. 1982: 25-26, pi. 7. figs 7-9, pi. 8. fig. 1 (synonymy); 1994: 59-60, pi. 26, fig. a-b. -
Cairns & Parker, 1992: 22-24, pi. 6, figs d-e. g-h (synonymy). — Cairns & Zibrowius, 1997: 129-130.
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 564.
1 (MNHN). — Stn 620, I (MNHN). — Stn 622, 1 (MNHN). — Sin 623. 2 (MNHN).
4 (USNM 98741). — Stn 567.
Type Locality. — Off Cape Jaffa and Neptune Island. South Australia, 165-190 m.
Remarks. — Aulocyathus recidivus was more fully redcscribed and figured by Cairns (1982, 1994) and
Cairns & Parker (1992). The four known species of Aulocyathus are compared in Table 7. and the three Pacific
species illustrated by Cairns (1994: pi. 26). A recidivus is unique in having a notch between the theca and
peripheral septal edges, and is further distinguished by having a relatively large, fragile corallum with well-spaced
septa and a well-developed columella.
Source :
104
S l) CAIRNS
Table 7. — Comparison of the four known species of Aulocyathus.
Distribution. — Wallis and Futuna region: Tuscarora and Combe Banks; 1020-1280 m. Elsewhere: Indo-
West Pacific from southwest Indian Ocean to Japan; 128-1 137 m (Cairns & ZlBROWius, 1997).
Aulocyathus juvenescens Marenzeller, 1904
Fig. 15 h
Aulocyathus juvenescens Marenzeller. v* 1904. 301-302. pi. 18. fig. 17. — ZlBROWius. 1980: 107. — Cairns & KELLER.
1993: 247. — Cairns. 1994: pi. 26. figs h-i. — Cairns & Zibrowius, 1997: 130 (mentioned).
MATERIAL EXAMINED. — Philippines. MUSORSTOM 3: sin 87. I juvenile (USNM 97653).
Vanuatu Musorstom 8: sin 976. 2 (MNHN). — Stn 1070. I (USNM 98740).
Type Locality. — Pemba and Zanzibar Islands. Tanzania. 400-463 m.
Remarks. — Two adult, intact coralla are reported herein: one (MUSORSTOM 8 stn 1070) 4.4 mm in CD and
14.6 mm in height, the other (MUSORSTOM 8 stn 976) 3.7 mm in CD and 1 1.0 mm in height. Both specimens
have a smooth, porcellaneous theca with no indication of granulation or intercostal striae. The theca of the larger
specimen has streaks of brown pigment, the other is pure white. Both coralla have a finely serrate, circular
calicular margin and both have 32 septa arranged S|>S2>S3>S4; however, the size difference between the S2 and
the Hanking S3 is slight. The lower axial edges of S| bear robust tubercles; the columella is rudimentary. These
specimens resemble the syntypes of A. juvenescens in all respects except for thecal texture, the syntypes having
faint, longitudinal intercostal striae and low. glisteny granulations.
Aulocyathus juvenescens is compared to the other Recent congeners in Table 7. It is distinguished bv having
the smallest calicular diameter, least number of septa, and a porcellaneous theca.
Distribution. — Vanuatu region: Tanna and Espiritu Santo; 182-184 m. Elsewhere: Philippines; Tanzania;
196-463 m (Cairns & Zibrowius. 1997).
Genus DESMOPHYLLUM Ehrenberg, 1834
DesmophyUum dianthus ( Esper, 1794)
Mad repo ra dianthus Esper. *1794: pi. 69. figs 1-3.
Source :
AZOOXANTHELLATE SCLERACTINIA
105
Desmophyllum cristagalli H. Milne Edwards & Haime, v* 1848a: 253.
Noi Qesmophylhm, sp. cf. D. cristagalli - WELLS, 1954: 470 (= Javania exserta)
i? 7 Z ' CAIR^J"4: 26'27- P'- 9- a-d (synonymy and neo.ype designation); 1995: 77. pi 9,
g. d t, map 4 (synonymy), 1998: 385-386. — Cairns & Zibrowius, 1997: 131. figs 17g-h.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: sin 637. I (MNHN)
Vanuatu. Musorstom 8: stn 1088. 8 (MNHN). - S.n 1089. I (MNHN). - Stn 1 128. I (USNM 98742).
Tv pe Locality. — Sagarni Bay, Japan (depth noi known).
Remarks. - This ubiquitous coral has been redescribed and figured many times, often under the junior
synonym name of D. cristagalli (see Zibrowius, 1980; Cairns, 1979, 1982. 1994, 1995). But. despite its
& ziSBROWiLSStl I997)0n and abUndanCe Cerla'n reg'0nS’ “ is rarely collec,cd in "ie tropical South Pacific (Cairns
Distribution - Wallis and Futuna region: Rotumah Bank: 820-830 m. Vanuatu region: Espiritu Santo;
Guyot Bougainville; 455-778 m. Elsewhere: cosmopolitan except off continental Antarctica and northern boreal
Pacific; 35-2460 m (Cairns, 1994).
Genus THALAMOI’HYLLIA Duchassaing, 1870
Thalamophyllia tenuescens (Gardiner, 1899)
Desmophyllum tenuescens Gardiner, v* 1899a: 161-162. pi. 19 figs la-b
Desmophyllum delicatum - WELLS, v.1954: 470 (Noi Desmophyllum delicatum Yabe & Eguch, 194?)
Thalanutphylha tenuescens - Cairns. 1995: 78. pi. 21. figs g-i, map 19: 1998: 386. - Cairns & z'ibrowius. 1997: 133.
. ,77,17' 7. E*AMINED- — Wallis an<l Futuna region. MUSORSTOM 7: stn 496. 1 (MNHN) — Stn 499
Sm 5ll , USNM 987T1,'MNHN)- “ *" ^ 2 ,MNHN,‘ S,n 512‘ 3 ^SNM 98744,. _ S.n 513 1 (MNHN) -
Lord Howe Island. NZOI: stn PI 15. 2 colonies (USNM 94363).
Type Locality. — Sandal Bay. Lifu, Loyalty Islands, 73 m.
Remarks. — This species was rently rcdescribed and illustrated by Cairns (1995). The largest known
corallite, reported herein from Lord Howe Seamount Chain (NZOI PI 15). measures 24 x 15 mm in CD. 27 mm in
height, and contains three pairs of S5 (54 septa).
Distribution. — I-utuna; 240-349 ill. Elsewhere: Philippines; Indonesia; Bikini Island. Marshall Islands
(Wells, 1954); Loyalty Islands (Gardiner, 1899a); Kermadcc Islands; Lord Howe Seamount Chain (reported
herein); oil Queensland; 8-315 m (Cairns & Zibrowius, 1997).
Genus LO PH ELIA H. Milne Edwards & Haime. 1849
Lophelia pertusa (Linnaeus, 1758)
Madrepora pertusa Linnaeus, *1758: 797.
Madrepora prolifera Pallas, *1766: 307.
Lophelia prolifera - Cairns. 1979: 125-127. pi. 24. figs 1-5 (synonymy); 1991a: 17-18. pi. 6. fig. j.
Lophelia pertusa- Zibrowius, v.I980: 126-130. pi. 66. figs A-L (synonymy). — Cairns. 1995- 27-^S pi 9 figs e-i
(synonymy). ' ' 6
Source :
106
S. D. CAIRNS
MATERIAL EXAMINED. — Wallis and Fuluna region. MUSORSTOM 7: sin 530, 19 fragments: 15 (MNHN),
4 (USNM 98745). — Sin 572. I (MNHN).
Type Locality. — Not staled, but probably the fjords of Norway (Zibrowius, 1980).
Remarks. — This species, which is common in the Atlantic, is rarely collected in the Indo-Pacific.
represented in this collection only by several dead fragments.
Distribution. — Wallis and Futuna region: Waterwitch Bank; 560-580 m. Elsewhere: cosmopolitan, except
off continental Antarctica (rare in western Pacific); 60-2170 m (Cairns, 1995).
Genus DACTYLOTROCHUS Wells. 1954
Dactylotrochus cervicornis (Moseley, 1881)
Figs B, 16 a-f
Tridcicophyllici cervicornis Moseley, *1881: 183-184, pi. 10, figs 2a-d, 3a. — Bassett-Smith, 1890: 368.
Tridacophyllia priniordialis Gardiner, v* 1899a: 168, pi. 19, figs 7a-e.
Dactylotrochus cervicornis - WELLS, v.1954: 470-471. pi. 178. figs 1-3. — Cairns & Zibrowius. 1997: 131.
MATERIAL EXAMINED. — Wallis and Futuna region
MUSORSTOM 7: stn 499. 1 (MNHN). — Stn 504. I (MNHN). —
Stn 512. 2 (MNHN). — Stn 513. I (MNHN). — Stn 514,
4 (MNHN) and SEM stub 889 (USNM 98746). — Stn 515.
1 (USNM 98747). — Stn 584, I (MNHN). — Stn 589,
I (MNHN). — Sin 605, 3 (MNHN).
Vanuatu. Musorstom 8: stn 988. 3 (USNM 98749). —
Stn 1095. 1 (USNM 98748).
Guam. East Agana Bay. 7.08.1986. 128-137 m, I (USNM
96503).
Fig. B. — Diagrammatic representation of the
calice of Dactylotrochus cervicornis, showing
the placement, length, and orientation of the
calicular extensions. Thick lines represent a
vertical extension; thin lines represent a
horizontal to oblique extension. Length of
extensions drawn in proportion to calicular
diameter.
Type Locality. — Unknown.
Description. — Corallum attached through a robust
(3-6 mm in diameter) cylindrical to subcylindrical pedicel and
a thin, slightly expansive base, the base and lower pedicel
usually covered with several faint, annular epithecal ridges.
One damaged specimen (MUSORSTOM 8 stn 1095) and two
syntypes display a polycyclic base, achieved by thecal
bridging of raised costal ridges in the vicinity of the base.
Above the pedicel, at a height of 3-10 mm. the theca and
corresponding internal septa arc divided into several (up to 8)
elongate, tapered and sometimes bifurcating thecal extensions
(Fig. B). The largest thecal extensions originate from the two
thecal faces, the basal part of each extension 6-8 mm in
length and 4-6 mm wide, beyond which it bifurcates into two
smaller extensions 2-3 mm in width, which, in larger specimens, bifurcate once more. These laterally-placed,
bifurcating extensions are vertical in orientation, producing a constricted fossa. In a well-developed corallum,
between each laterally-placed extension and the greater axis may be 4 smaller, nonbifurcating extensions, each
about 5 mm long, 3 mm in width, oriented outward from the calicular edge. Finally, 2 more nonbifurcating,
upward-projecting extensions originate from the greater axis of the corallum, each about 4 mm long and 3 mm
wide. Not all coralla have all 8 digitiform projections, but the two lateral and two axial extensions are usually
Source :
AZOOXANTHELLATE SCLERACTINIA
107
present even in the smallest coralla. Largest specimen (MUSORSTOM 8 stn 988) 23.1 x 12.9 mm in CD and 30
mm m height. Theca white, covered with granules that are sometimes arranged in longitudinal rows, but costae and
intercostal striae absent.
Septal symmetry is difficult to determine. Large coralla have about 240 septa arranged in 7 size classes, each
progressive size class being narrower and originating closer to the calicular margin. Septa closely spaced, have
smooth faces and bear 2-5 elongate ridges (menianes) oriented parallel to the septal edge. The menianes arc 60-
pm in height and about 30 pm wide, alternating in position with those from the adjacent septal faces (Figs 16
d-e). thus blocking a view of the lower interseptal spaces. Fossa deep and elongate: no columella.
Remarks. — Dactylotraehus cemcornis is assumed to he a solitary coral since no stolons were detected and
only one elongate (ossa (one mouth) is present. All previous authors described the septate digitiform thecal
extensions of this species as branches, but it seems incongruous to refer to a solitary coral as having branches,
thus the term "thecal extension” is used.
The five syntypes of T. primordial is Gardiner, 1899 are deposited at the University Museum of Zoology
Cambridge the largest specimen a juvenile 10 mm in heigh, and 2.4 mm in pedicel diameter, having only
60 septa and only three septal extensions.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Field Bank; 245-400 m. Vanuatu region-
Tanna and Espintu Santo, 320-372 m. Elsewhere: Philippines; South China Sea: Guam (reported herein); Bikini
Marshall Islands; New Caledonia and Loyalty Islands; 73-400 m (Cairns & Zibrowius 1997)
Genus RHIZOSMILIA Cairns. 1978
Rhizosmilia robusta Cairns, 1993
robus,a Cairns in Cairns & Keller. *1993: 250-253. pi. 6. figs F-l. - Cairns & Zibrowius, 1997:
I (MNHNE|RIAL EXAMINED — Wall's and Futuna region. Musorstom 7: stn 509. 2 (MNHN). — Stn 524.
9 8 7 VD*1 “ 3 ‘ “s i T? M, c i^w ^ %4' 1 <MNHN>- - Stn 971. I (MNHN). - Sin 1021. 1 (USNM
HMNHN) S 07 ’ (USNM 98750)- — Sln l077- 3 (USNM 98751). — Stn 1078, I (MNHN). — Sin 1084.
Type Locali ty. 'Anton Bruun" stn 373B: 26°00'S, 33°05'E (off southwestern Mozambique), 1 35 m.
Remarks. — Because of its size, shape, and palar configuration, this species can be taken for a Caiyopltxllia
at first view, but differs from that genus in having a polycyclic base, endothecal dissepiments, and a columella
composed of irregularly shaped papillae, not twisted lamellae.
Distribution. — Wallis and Futuna; 240-300 m. Vanuatu region: Anatom, Efate, Espiritu Santo, and
Malakula; I 10-360 m. Elsewhere: southwestern Indian Ocean; Philippines; 66-202 m (CAIRNS & Zibrowius.
Genus ASTEROSM1LIA Duncan, 1867
Asterosmilia gigas (van der Horst, 1931), n. comb.
Figs 16 g-i, 17 a-b
Caryophyllia gigas van der Horst. v*1931: 4-5, pi. 2, figs 1.4. — Zibrowius, 1980: 70.
Rhizosmilia gigas - Cairns & KELLER. 1993: 251-252.
Source :
108
S. I). CAIRNS
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 510. I (MNHN). — Stn 515. 16:
10 (MNHN), 6 OJSNM 98754). — Stn 523. 4 (MNHN). — Sin 535. 2 (USNM 98753). — Stn 537. 2 (MNHN). —
Stn 586. I (MNHN).
Type Locality. — Mauritius, 183-366 m.
Description. — Corallum robust and ceratoid, either regularly curved or irregularly bent in a scolecoid
fashion. Corallum free, always with an open, broken base 1.5-2. 5 mm in diameter revealing 12 septa. Base
monocyclic, not reinforced with rootlets-like. Largest specimen reported herein (MUSORSTOM 7 sin 515) 28.2 x
26.0 mm in CD and 65 mm in height; however, the holotype (BM 1939.7.20.85 1 ) measures 39.4 x 31.6 mm
in CD and 79 mm in height. C1.3 expressed as low, rounded ridges; in some coralla C4-5 also present as faint
ridges. No intercostal striae or granules. Theca white, often covered with encrusting organisms (e.g., foraminifera)
and small buds, presumably of the same species. Buds are found on most coralla and cluster near the calicular
margin, each measuring 1. 5-3.0 mm in diameter and up to 2 mm in height and having 12 septa.
Septa hexamerally arranged in 5 cycles (Si>S2>S.3>S4>S5); however, it is not unusual for even large coralla to
lack several pairs of S5 resulting in less than 96 septa, or for some coralla to have additional pairs of S6. The
holotype, for example, has 5 pairs of S6 for a total of 100 septa. Si up to 3.5 mm exsert. having straight vertical
axial edges. S2-4 progressively and gradually narrower; however, S5 only half width of S4. Triangular lancets
corresponding to Si and S2 form at calicular margin. Wide (2. 5-3.0 mm), thin paliform lobes (P4) usually border
every S4 that is Hanked by a pair of S5, the P4 appearing as pairs in those half-systems containing both pairs of
S5; however, occasionally a paliform lobe occurs before an S3, even though pairs of S5 are present. Stereome
sometimes present in base of corallum, giving it a dense aspect. Lower portion of corallum also filled with
abundant vesicular endothecal dissepiments. Fossa of moderate depth, containing a well-developed, papillose
columella composed of irregularly-shaped, crispate elements.
Remarks. — This appears to be the first report of this species subsequent to its original description, which
was based on one specimen, and the first record from the Pacific Ocean. Both van der Horst ( 1931 ) and Cairns
& Keller (1993) noted that the broken cross section of the base of the holotype was surrounded by what appeared
to be contiguous rootlets, which is characteristic of the genus Rhizosmilia. However, all specimens reported above
propagated by asexual budding, the open base of every corallum corresponding in shape, size, and number of septa
to the buds that occur on its theca. A budding mode of reproduction is inconsistent with the polycyclic
reinforcement of a Rhizosmilia , and thus the root-like structures of the holotype arc not interpreted as such, but as
an artifact. Discounting the rootlet structures, this species belongs in the genus Asterosniilia , which often
reproduces by asexual budding. In fact, A. gigas is remarkably similar to A. profunda (Duncan. I864)(sec Cairns
& WELLS, 1987), known only from the Eocene to Pliocene of the Caribbean.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch Bank; 252-510 m. Elsewhere:
Mauritius; 183-366 m.
Family TURBINOLIIDAE H. Milne Edwards & Haimc. 1848
Genus ALATOTROCHUS Cairns. 1994
Alatotrochus rubescens (Moseley, 1876)
Platytrochus rubescens Moseley, v*1876: 553.
Sphenotrochus rubescens - MOSELEY, v.1881: 157-159. pi. 6. figs 8. 8a.
Alatotrochus rubescens - Cairns, 1994: 68-69, pi. 29, figs g-l; 1995: 84, pi. 24, figs a-b. map 14; 1997: 14, pi. I,
fig. a. pi. 4, fig. a; 1998: 390. — Cairns & Zibrowius, 1997: 141, fig. I8h.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: sin 1068, 2: 1 (MNHN), I (USNM 98755).
Source :
AZOQXANTHELLATE SCLEKACTINIA
109
type Locality - "Challenger" stn 192: 5°49T5"S, 132°14'15"E (Kai Islands. Banda Sea). 236 m (no.
136 m. as reported by Cairns (1994. 1995) and Cairns & Zibrowius (1997)). However, the "Challenger"
station list presented as Appendix II of the "Narrative" (TlZZARD el al., 1885) lists 140 fathoms (= 256 m) as the
depth lor station 192.
Remarks. This species was recently redescribed and the types illustrated by Cairns (1994).
distribution. - Vanuatu region: Malakula; 536-619 m. Elsewhere: Japan; Philippines; Banda Sea-
northwestern Australia; southern Norfolk Ridge; 180-751 m (Cairns & Zibrowius, 1997).
Genus PLEOTROC HUS Cairns, 1997
Pleotrochus venustus (Alcock, 1902)
Figs 17 d-c
Ceratotrochus venustus Alcock. v* 1902a: 92; v. 1902c: 10 pi 1 figs 5 5a
Cryp°TbroW'i c-rVlST- '"5: ^ lm Par,: °nly PL 27 • f,gS a'b;' remaining fi8urcs and Ascription pertain to
" Crypiotrochus " venustus - Cairns & Zirowius, 1997: 142-143.
Pleotrochus venustus - Cairns, 1997: 14 . pi. 1, fig. b, pi. 4, fig. b.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1 1 14. 1 (MNHN).
Type Locality. — "Siboga" stn 256: 5°26.6'S. I32°32.5’E (Kai Islands, Banda Sea). 397 m.
Remarks. — Although worn and obviously long dead when collected, the single specimen reported above is
undoubtedly conspecific with P. venustus. It represents the Hist record outside the Banda Sea and the largest known
corallum: 13.5 x 1 1.4 mm in CD and 11.7 mm in height. The species is more fully described by Cairns &
ZIBROWIUS (1997), and the species and genus are figured and discussed by Cairns (1997).
Distribution. — Vanuatu region: Espirilu Santo; 647 m. Elsewhere: Banda Sea, Indonesia- 200-397 m
(Cairns & Zibrowius, 1997).
Pleotrochus zibrowii Cairns, 1997
Figs 17 g-h
Crypu^ochus venustus - Cairns, 1995: 88-89 (in part: pi. 26g-i; not pi. 27a-b. which is Ceratotrochus venustus Alcock.
Pleotrochus zibrowii Cairns. *1997: 14-15, pi. I, fig, c, pi. 4. fig. c.
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 637. I (MNHN)
Vanuatu. Musorstom 8: stn 1113, I (USNM 98756).
T5 pe Locality. NZOI stn U584: 3126.3'S. I72°35.6'E (Three Kings Ridge. New Zealand). 1137-
1 150 m.
Remarks. — Although worn and obviously dead when collected, the two specimens reported above
are virtually identical to specimens in the type series, the larger specimen (MUSORSTOM 7 stn 637) measuring
13.4 x 12.6 mm in CD and 1 1.5 mm in height. These records represent the second collection of this recently
described species.
Distribution. — Wallis and Futuna region: Rotumah Bank; 820-830 m. Vanuatu region: Espirilu Santo:
700-736 m. Elsewhere: Three Kings Ridge, New Zealand; 1 137-1150 m.
Source :
] 10
S. D. CAIRNS
Genus TROP/DOCYATHUS H. Milne Edwards & Haime, 1848
Tropidocyathus lessonii (Michelin, 1842)
Fig. 17 c
Flabellum Lessonii Michelin. *1842: 119.
Tropidocyathus lessoni -Cairns, 1989a: 33-34. pi. 16. figs d-l (synonymy); 1994: 67. pi. 29, figs a-b (synonymy).
Tropidocyathus lessonii - CAIRNS & ZlBROWius, 1997: 146-147. — Cairns, 1997: 16. pi. 1. fig. e, pi. 4. fig. e. pi. 7. fig.
d; 1998: 390-392.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 1016, 1 (MNHN).
Type Locality. — Unknown.
Remarks. — The single specimen reported above, worn and dead when collected, measures 12.8 x 10.7 mm in
CD and 10.8 mm in height. This species is distinctive in having a rhomboidal calice (the lateral thecal faces
bulging outward) and highly developed edge crests up to 3.4 mm in height. This relatively common deep-water
species was redescribed, illustrated, and discussed by Cairns (1989, 1994, 1997).
Distribution. — Vanuatu region: Efate; 291-300 m. Elsewhere: Indo-West Pacific from southwest Indian
Ocean to Japan; 50-421 m (Cairns & ZlBROWius, 1997).
Tropidocyathus lab idus Cairns & Zibrowius, 1997
Fig. 2 d
"Tropidocyathus" labidus Cairns & Zibrowius, 1997: 148, figs 20 a-g. — Cairns, 1998: 392.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 8: stn 963, 1 cemented to a Xenophora shell
(MNHN). — Stn 1026, 1 (MNHN). — Stn 1065, I (MNHN). — Stn 1068, 2 (USNM 98758).
TYPE LOCALITY. — Karubar stn 2: 5°4700"S, 132° 1 135"E (Kai Islands. Banda Sea), 209-240 m.
Remarks. — This small, distinctive species was recently described and illustrated. All specimens reported
above were alive when collected and well preserved.
Distribution. — Vanuatu region: Anatom, Efate, and Malakula; 419-536 m. Elsewhere: Indonesia; Ryukyu
Islands; tropical Western Australia; 206-425 m (Cairns, 1998).
Genus CYATHOTROCHUS Bourne, 1905
Cyathotrochus pileus (Alcock, 1902)
Trochocyathus pileus Alcock, v*1902a: 96-97; v. 1902c: 15-16, pi. 2, figs 11. 11a.
? Cyathotrochus herdmani Bourne, *1905: 193, pi. 1. figs 2. 2a.
Tropidocyathus pileus - Cairns. 1989a: 34-35, pi. 17. figs a-h (synonymy); 1994: 68. pi. 29. figs d-e; 1995: 91. pi. 28,
figs a-c, map 1 1. —Cairns & Zibrowius. 1997: 147-148, figs 19 h-i.
Cyathotrochus pileus - CAIRNS. 1997: 16. pi. 1. figs f-g, pi. 4, fig. f; 1998: 392.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 958, 2 (USNM 98760). — Stn 959. I (MNHN). — Stn 980
1 (MNHN). — Stn 1016, I (MNHN).
Source : MNHN . Paris
AZOOXANTHELLATE SCLERACT1NIA
1 1
Type Locality. - "Siboga" sin 95: 5°43'N. I I9°40'E (Sulu Archipelago, Philippines), 522 m.
CAIRNS n989a^994ST997i Mand '"ST**™ °f ^ C0mm0nIy collected deeP'water coral can be found in
CAIRNS (1989a, 1994. 1997). Most of the specimens reported above were alive when collected and possess four
cycles oi septa.
Distribution^— Vanuatu region: Anatom. Tanna. and Elate: 300-497 m. Elsewhere: lndo-Wcst Pacific from
southwest Indian Ocean to Japan; 123-522 m (Cairns & ZIBROWIUS, 1997).
Genus DELTOCY A TH01DES Yabe & Eguchi. 1932
Deltocyathoides orientals (Duncan, 1876)
Deltocyathus orientalis Duncan. *1876: 431.
Peponocyathus orientalis - Wells, v.1984: 214
Pe!7Tr!Z f A,RNS' 29- *°-32> P'- u- <H. pi. 15. figs a-d (synonymy); 1994: 65-66.
p . 28 figs c-1, pi. 4 . Iig. I (synonymy). [Not Deltocyathus italicus var. australiensis Duncan. v*18701
Deltocyathoides oriental,. s - Cairns & ZIBROWIUS. 1997: 144. - CaIRNS. 1997: 17. pi. 1. fig. h. pi. 7. fig. f; 1998: 392.
Material examined. — Musorstom 7: stn 512. 1 (MNHN). — stn 523. I (MNHN).
I vpe Locality. — 34°12’N, 136°20'E (southeastern Honshu. Japan). 95 m.
Remarks.
Cairns (1989,
This small, commonly collected and locally abundant coral was rcdescribed and illustrated
1 994) as Peponocyathus australiensis.
by
D^TR'BLrnON, - Wallis and Futuna; 245-455 m. Late Pleistocene of Espirilu Santo. Vanuatu (Wells.
1 784). Elsewhere: Indo-West Pacific from southwest Indian Ocean to Japan; 44-635 m.
Genus NOTOCYA THUS Tenison Woods, 1880
Notocyathus conicus (Alcock, 1902)
Citharocyathus conicus Alcock, v*I902b: 1 18-1 19: v. 1902c: 22 pi 3 figs 18 18a
Notocyathus conicus - Cairns, 1989a: 28. pi. 13. figs a-i (synonymy); 1994: 64-65, pi. 28. figs a-b; 1995- 91-9-
figs c, g. map 10; 1997: 17. pi. 4. fig. j. — Cairns & Zibrowius, 1997: 143-144
Not Citharocyathus conicus - WELLS, v.1984: 214. figs 4, 2-5 [= ? Notocyathus venustus (Alcock. 1902)].
Material examined. — Vanuatu. Musorstom 8: stn 1018
Stn 1067. 2 (MNHN).
cemented to a Xenophora shell (MNHN). —
Type Locality. — " Siboga ’*' stn 95: 5°43.5'N. 1 19°40'E (Sulu Sea. Philippines). 522 m.
Remarks. — Three well-preserved specimens of N. conicus are reported herein, the largest 6.2 mm in CD and
7.8 mm in height. This species was redescribed and figured by Cairns (1989. 1994). as well as compared to the
other Recent sibling species, N. venustus (Alcock, 1902). WELLS (1984) reported the "venustus" form of
A', conicus from the Pleistocene ol Vanuatu, but the small size and poor condition of his specimens do not allow
a definite attribution between these two very similar species.
Distribution. — Vanuatu region: Efate and Malakula; 344-366 m. Elsewhere: Japan: Philippines; Indonesia;
Norfolk and Kermadee Ridges; 34-923 m (Cairns & Zibrowius, 1997).
Source :
1 12
S. D. CAIRNS
Genus CRYPTOTROCHUS Cairns, 1988
Cryptotrochus brevipalus sp. nov.
Figs 17 f, i
Material EXAMINED/TYPES. — Vanuatu. Musorstom 8: stn 988, 2 paratypes (MNHN). — Stn 1014, I paratype
(MNHN). — Stn 1028. I paratype (USNM 98762). — Stn 1034, holotype (MNHN) and 1 paratype (MNHN). — Stn 1067,
1 paratype (MNHN). — Stn 1113. 1 paratype (USNM 98763).
Type Locality. — Musorstom 8: stn 1034: 17°54'S, 168°42'E (Elate), 690-750 m.
Etymology. — The species name brevipalus (Latin brevis, short + pains, stick) is a reference to the
relatively short pali (P2) of this species. The name is treated as a noun in apposition.
Description. — Corallum a regular inverted cone, the blunt basal angle ranging from 68° to 72° (turbinate).
Largest specimen (holotype) 10.3 mm in CD and 10.0 mm in height. Calice circular; theca thick, about 0.7 mm.
Costae 0.4-0. 5 mm wide at calicular edge, separated by deep intercostal furrows 40-50% the costal width. C 1 -2
independent in origin, but C4 originate by trifurcation of C3 2. 3-2. 5 mm above base. Costae serrate, the outer edge
of each costa covered with a single row of small teeth about 0. 15 mm in diameter and height; smaller spines about
0.1 mm in height and 0.04 mm diameter project laterally from each costa into the intercostal spaces, although they
arc so small and well spaced that they do not obscure the view of the underlying theca between the costae.
Corallum uniformly white.
Septa hexamerally arranged in 4 complete cycles in all specimens examined (CD = 6.0-10.3 mm):
Si>S2>S3>S4, 48 septa. Si about 1.8 mm exsert, extend about 3/4 distance to columella, having straight, vertical
axial edges that fuse to the columella lower in fossa. S2 about 1.2 mm exsert and 2/3 with of the Si, each bearing
a prominent palus (P2). S3 about 0.9 mm exsert and 3/4 width of the S2. their lower axial edges strongly fused
to the peripheral edge of the P2 lower in fossa, but easily visible in calicular view. In fact, in some specimens
it would appear that each S3 bears a palus, a pair of which unite before their common S2 to form a V-shaped P2
(as in the case of Notocyathus); however, each P2 in this species rises higher in the fossa (usually just above
the level of the columella) as a single slender structure. S4 about 0.7 mm exsert. 2/3 width of the S3, and have
free axial edges. Septal faces covered with very small spines, the same size and shape as those occurring on the
lateral faces of the costae. Fossa of moderate depth, containing a papillose columella of 4-7 strongly
fused elements.
Remarks. — Of the two other species known in the genus, C. brevipalus is most similar to C. javanus
Cairns, 1988 (Java Sea, 585 m) in size and shape, and in having independent S4; however, differs in having serrate
costae (not granular), S3 fused to the peripheral edge of P2, and in having less prominent P2. C. brevipalus
resembles C. carolinensis Cairns, 1988 (western Atlantic, 320-338 m) in having serrate costae and S3 that fuse
with the P2, but differs in having a larger and more open corallum (higher basal angle), independent S4. and
pointed P2 that rise slightly higher in the fossa. Although not in the same genus, C. brevipalus resembles
Pleotrochus zibrowii Cairns, 1997, in size, shape, and in having S3 that fuse to the peripheral edge of the P2. The
two species have also been found at the same station; however, C. brevipalus differs in having a costae:septa ratio
of 1 (vs 2), costal trifurcation (vs independent origin), a blunt base (vs pointed), and less prominent P2.
Distribution. — Vanuatu region: Tanna, Efate, Malakula, and Espiritu Santo; 466-700 m.
Genus IDIOTROCHUS Wells, 1935
Idiotrochus kikulii (Yabe & Eguchi, 1941)
Placotrochides kikutii Yabe & Eguchi. *1941: 104; 1942: 149, pi. 9, fig. I6a-c.
Source :
AZOOXANTHELLATE SCLERACTINIA
I 13
“p, ft ^99* 390 89ar36,;37' P,‘ 718' r,gS a-b- d‘h >994: 69. pi. 30. figs a-d; ,997: 21. pi.
rig. g, pi. 7. Jig. I, 1998. 390. — Cairns & Zibrowius. 1997: 148-149.
5,
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom
98764). — Stn 513, 1 (MNHN).
7: sin 509, 6 (MNHN). — Stn 512, I (USNM
Type Locality. — Toyama Bay, Honshu, Japan (depth not given).
Remarks. — This distinctive, wedge-shaped species is well described and illustrated hy Cairns ( 1989a) and
Cairns & Zibrowius (1997).
Distribution. — Wallis and Futuna region: Futuna; 240-260 in. Elsewhere: western Pacific from Japan
through Indonesia; Western Australia; 97-645 m (Cairns & Zibrowius. 1997).
Genus PEPONOCYATHUS Gravier. 1915
Peponocyalhus folliculus (Pourtales, 1X68)
Figs 18 a-b
Stephanophyllia folliculus Pourtales, v*1868: 139.
Peponocyalhus variahilis Gravier, v*I9I5: 5; v.1920: 39. pi. 4. figs 60-73. pi. 13. fig. 202. pi 14 lies ?03-->04
Peponocyalhus folliculus - Cairns, 1979: 113-115, pi. 22. figs 1-4; 1989a: 32-33 (in part: pi. 16. figs a‘-c); "l 994-
, , co !-gS S;k, ‘synonymy and description); 1997: 30. pi. 3, fig. k. pi. 6, figs h-j. - Zibrowius, v.I980:
i 15, pi. 58, tigs A-L, pi. 59, tigs A-K (synonymy). — Cairns & Zibrowius, 1997: 146.
66-
13-
Material EXAMINED. — Vanuatu. Musorstom 8: stn 1088, 1 attached to a Xenophom shell (MNHN).
Type Locality. Bibb" stn 51: 24°I2’40"N. 8I°I9'25"W (Straits of Florida, western Atlantic), 433 m.
remarks. — The single specimen reported herein is "gourd"- or "onion-shaped", measuring 4.9 mm in
greatest diameter, but only 3.3 mm in CD, having 30 septa. The species is known to be variable in shape, the
onion-shaped form being well illustrated and discussed by Gravier (1920) and Zibrowius (1980). The specimen
was dead when collected, and basally incorporated into a Xenophom gastropod shell. Xenophom gastropods, also
called carrier shells", appear to be efficient collectors of small, otherwise rarely collected deep-water Scleractinia.
this particular shell (Fig. 18 a) having cemented eight solitary corals to its outer whorls, including: one
P. folliculus, two Caiyophyllia abrupta, three Trochocyathus discus, and two as yet unidentified species (i.e..
Gardineriidae gen. nov. sp. nov. sensu Stolarski. 1996).
distribution. — Vanuatu region: Espiritu Santo; 425-455 m. Elsewhere: Atlantic; western Pacific from
Japan through Indonesia: 50-582 m (Cairns & Zibrowius. 1997).
Superfamily FLABELLOIDEA Bourne, 1905
Family GUYNIIDAE Hickson. 1910
Genus GUYNIA Duncan. 1872
Guynia annulata Duncan. 1872
Guynia annulata Duncan. v*1872: 32, pi, 1. figs 1-8. — Zibrowius, v,!980: 161-162. pi. 83. figs A-Q (synonymy). —
Cairns, 1989a; 42-43. pi. 21, fig. f, pi. 22, figs a-e; 1998: 392. — Cairns & Parker, 1992:' 42-43, pi 14 figs g-h
— Cairns & Zibrowius, 1997: 150 (synonymy).
Source :
S. D. CAIRNS
1 14
Material EXAMINED. — Wallis and Fuluna region. MUSORSTOM 7: sin 504, 4 attached to theca of ?dead
corallum of Conotrochus asymmetros (MNHN). — Stn 513. 1 attached to theca of dead corallum of Heterocycithus cf.
sulcatus (MNHN). — Sin 569. 1 attached to theca of dead corallum of Flabellum arcuatile, n. sp. (MNHN).
Vanuatu. Musorstom 8: stn 1016, 1 attached to base of dead discoidal coral, perhaps a Deltocyathus (USNM
98765). — Stn 1097. 1 cemented to a Xenophora shell (MNHN).
Type Locality. — Adventure Bank. Mediterranean. 168 m.
Remarks. — This extremely small, serpulid-like corallum is often found attached to living and dead coralla of
other scleractinian species (see Material Examined). Although difficult to detect in collections because of its small
size (CD about 1 mm) and its sometimes cryptic habit, its widespread distribution is becoming increasingly well
known as large collections are closely examined. The species is more fully described and illustrated by Zibrowius
(1980), Cairns (1989a), and Cairns & Parker (1992).
Distribution. — Wallis and Futuna region: Fuluna; Waterwitch Bank; 260-300 m. Vanuatu region: Efate and
Espiritu Santo; 288-300 m. Elsewhere: cosmopolitan in tropical and warm temperate regions; 28-653 m (Cairns
& Zibrowius, 1997).
Genus TRVNCA TOGUYMA Cairns, 1989
Truncatoguynici irregularis Cairns, 1989
Fig. 18 c
Truncatoguynia irregularis Cairns, * 1989a: 43. pi. 22. figs f-g. pi. 23. figs a-c, f; 1994: 70. pi. 10, lies e-fi 1995- 93-
94. pi. 29, figs g-h, pi. 30, figs a-b, map 12.
Truncatoguynia sp. Cairns. 1989a: 43. pi. 23, figs d-e.
Material examined. — Vanuatu. Musorstom 8: stn 967. 4 (one in 5 pieces) (USNM 98766).
Type Locality. - " Albatross " stn 5311: 21°33'N, 1 16°15'E (north of Pratas Island. South China Sea)
161 m.
Remarks. — This species was described and illustrated several times recently as indicated in the synonymy.
The specimens reported above pertain to the diminutive form of the species, similar to those reported from NZOI
stn C531 by Cairns (1989a. 1995). These specimen are characterised by an elongate, curved corallum up to 21
mm in length, but having a small calice of only 2.4-2.S mm GCD. resulting in a length:GCD ratio of 7 3-7 6
The basal scar is correspondingly small (1. 1-1.8 x 0.9- 1.4 mm), coralla containing onlv 24 hexamcrally arranged
septa: Si-2»S3. °
Distribution. - Vanuatu region: Anatom; 295-334 m. Elsewhere: Ryukvu Islands; South China Sea-
Norlolk and Kermadec Ridges; 80-248 m (Cairns. 1995).
Genus TEMNOTROCHUS Cairns, 1995
Temnotroclius kermadecensis Cairns, 1995
Figs 18 d-e
Temnotroclius kermadecensis Cairns. *1995: 96, pi. 31, figs a-d.
Stn ltmE2RrpLm^INvD' VaT,,U MuSORSTOM 8: s,n 963’ 1 ce“d to a Xenophora shell (MNHN) -
Mn IU2J, 2 cemented to a Xenophora shell (MNHN).
Source : MNHN. Paris
AZOOXANTHELLATE SCLERACTINIA
I 15
Type Locality.
3.7 km off Nugent Island. Raoul Island. Kermadec Ridge. 366-402 m.
*™ARKS: ~ L'Ulc can.be added 10 >he original description based on these three specimens Temnotrochia
,ransversrdT^ioVaTaDirSe « haV,1,nS el°"gate ^Pissed-cylindrical anthocyathus that originates by
transverse division. a pap.llose columella, and paltlorm lobes before the Si-2. The characteristic thecal snots are
°,en 1 ICU 1 10 s®e’ and are ev,denl on|y on the specimen from Musorstom 8 stn 963. The specimens reported
MusorstonTs Stn % "bd rePTi°r lhlS Srr'CS ^ 'nClude Specimens ,arSer lhan in lhe type series, one from
MUSORSTOM 8 stn 963 being 2.1 mm m GCD and 6.0 mm in length. All three specimens were obtained as
s*^ range!" ‘° gaStr°P°d Shc" XenoPhora> a" efficient collector of azooxanthellate corals of this
Distribution. - Vanuatu region: Anatom and Elate; 321-400 m. Elsewhere: Kermadec Ridge; 366-402
Ill.
Family FLABELL1DAE Bourne. 1905
Genus FLABELLUM Lesson, 1831
Subgenus FLABELLUM (FLABELLUM) Lesson. 1831
Flabellum (F.) pavoninum Lesson, 1831
Figs 18 g-i
Flabellum pavoninum Lesson, v* 1 83 1 : 2. — Cairns 1989a- 46-50 nl 75 fi„c „ i „i -> a r , , ,
_ IT 70‘7r Pl- 2 flgs g"’ 3I- <* «. - Cairns * z,.row,?s, 19/7" llolifl fig S ' ** lsy"”W'
F'abhelr co“1"'"" Marenrel'er. v* 1888a: 48-49. - Cairns. 1989a: 46. 47. 50. nl. 24 figs e-f I
Flabellum sp. - Cairns. 1989a: 24. pl. 24, figs e-f. H ’
? f\i\jw\hR,Alc E^M,,NfD- “ Wa,Iis and Futuna re£ion Musorstom 7: sin 504 l (MNHN) — Sin 5H
4 imnhn)- - sb » 3 ius™ - - - 3 -
Q, ^."“f^-^SORSTGM 8: stn 962. I (MNHN). - Stn 965, 12 (USNM 98871 ) _ Stn 964 5 (USNM 98X74,
1 Y pe Locality. "Sandwich Islands" (= Hawaiian Islands), no depth given.
Df,CR,IP™N (coalitum form)- — Coral lum fiabellate: angle of thecal faces, 27°-47°; angle of thecal edges
'modal, the lower 5 mm ol the corallum (the pedicel) having an edge angle of 40°-50°, above which the corallum
widens to an edge angle ol 62°-l43°. Thecal faces virtually planar, meeting in acute thecal edges that bear
a prominent, discontinuous edge crest composed of 1-4 (depending on size of corallum) thecal eversions each
evers.on an irregularly shaped discrete structure up to 2.5 mm in height. Largest specimen examined
(Musorstom 8 stn 1 102) 29.1 X 16.3 mm in CD and 22.5 mm in height. Basal disc quite small (0.9-1 0 mm
m diameter), containing 6 pro.osepta, only the smallest coralla maintaining its original attachment to substrate
Theca dull in lustre, covered with fine transversely oriented ridges, and often covered with encrusting organisms in
four cases (MUSORSTOM 7 stn 538, MUSORSTOM 8 sins 976. 1 104 and 1132) bored by acrothoracican Crustacea
( ig. 8 h). Lower 5 mm of corallum (basal disc and lower pedicel) white; remaining theca a light reddish brown,
with more intense radial stripes corresponding to S1-3.
Septa hexamcrally arranged in 5 cycles and often part of a sixth cycle (S|.3>S4>S.s>S6) in larger coralla pairs
ot particularly common in the quarter systems adjacent to the principal septa, but also occurring in random
Source :
I 16
S. D. CAIRNS
order in lateral face quarter systems; largest corallum contains 140 septa. Distal peripheral edges of S 1-3 nonexsert,
separated from thecal edge by a low notch or slight concavity, then projecting into fossa as a prominent lobe;
midaxial edges of S1-3 often slightly concave, and lower axial edges thickened and very sinuous. The two principal
Si aligned with the GCD are smaller than the other 4 Si- Distal edges of S4 not lobate and thus much narrower
than S i-3, but lower in fossa almost 3/4 width of the S1-3; axial edges straight. S5 about 1/3 width of an S4.
When pairs of S6 are present, the S5 they flank widen to 2/3 the width of an S4 and the S6 are the size of a typical
S5. Fossa deep and elongate; lower axial edges of S1-3 fuse to form a rudimentary columella.
Remarks. — Although recently redescribed and figured (Cairns, 1989a, 1994; Cairns & Zibrowius. 1997),
/- . pavonimun is redescribed above based on the abundance of well-preserved specimens of this otherwise poorly
represented species. All specimens reported above pertain to the "coalition" form of the species, as discussed by
Cairns (1994) and Cairns & Zibrowius (1997), which is distinguished from the typical form by having a
smaller corallum. a lower edge angle, and fewer septa.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch Bank; 286-455 m. Vanuatu
region: Anatom, Tanna, Erromango, Efate, Espiritu Santo, and Malakula; 161-400 m. Elsewhere: Indo-Pacific
from southwest Indian Ocean to Hawaiian Islands, forma coalitum being most common off Japan; 73-665 m
(Cairns & Zibrowius, 1997).
Flab ell um (F.) arc u at He sp. nov.
Figs 19 a-d
Flabellum ( F.) angiosiomum - Cairns. 1995: 99. pi. 32. figs d-f. map 13 [Noi Fla belli, m angiostomum Folkeson
v* 1919].
Material EXAMINED/TYPES. — Wallis and Futuna region. Musors-tom 7: stn 524, 2 paratypes (MNHN). —
Stn 530, 3 paratypes (MNHN). — Stn 534, I paratype (MNHN). — Stn 535. 2 paratypes (MNHN). — Stn 542. I paratype
(USNM 98876). — Stn 546, 2 paratypes (USNM 98877). — Stn 578, I paratype (USNM 98878). — Stn 589 I paratvne
(MNHN). r
New Zealand region. NZOI: stn 192, 9 paratypes (USNM 94322). — Sin 197, 35: 20 paratypes (NZOI). holotype
(NZOI H688), 14 paratypes (USNM 94323). — Stn U599, 1 paratype (NZOI).
Type Locality. — NZOI stn 197: 32°22.9'S, I67°28.2’E (southern Norfolk Ridge), 540-544 m.
Etymology. The species name arcuatile (Latin arcuatilis , shaped like a bow) refers to the regular crescent-
shaped arc formed by the calicular margin.
Diagnosis. — Corallum flabellate, highly compressed (GCD:LCD = 2. 1-3. 1 ). never constricted, and robust.
Corallum wider than (all: GCD: H = 1.25-1.45. Thecal edges always rounded, the lower 7-9 mm of the corallum
having an edge angle of 60°-80°. changing to a more open I25°-180° with height. Face angle = 26°-39°. Holotype
38.4 x 15.4 mm in CD and 28.3 mm in height, chosen because it represents a corallum with a full sixth cycle of
septa; largest corallum (paratype from NZOI 197, NZOI) 49.8 x 20.5 mm in CD and 33.8 mm in height. Pedicel
small: 1.1-1. 3 mm in diameter. Thecal of well-preserved specimens uniformly reddish brown. Septa hexamerally
arranged in 6 or more cycles, the sixth cycle attained at a GCD of 20-25 mm. and larger coralla having some pairs
ol S7: S|-4>S5>S6>S7- Axial edges of S 1.4 extremely sinuous.
Remarks. — Among the 23 Recent species in the subgenus, F. arcuatile is one of three species to have a
t label late corallum with rounded thecal edges, the other two species being F. knoxi Ralph & Squires, 1962
(New Zealand region south of 38°S), and F. impensum Squires, 1962 (Antarctic, 46-2200 m). F. arcuatile is most
similar to F. knoxi, but differs in having a more robust corallum, a smaller pedicel with fewer protosepta
(6 vs 12), a lesser developed columella, and a more compressed corallum.
A fuller description of Flabellum arcuatile can be found in Cairns (1995), as Flabellum angiostomum
1 (Cairns, 1998) subsequently examined the type of Flabellum angiostomum , transferred it to the genus
Truncatoflabellum, and eliminated the New Zealand records from its range, but did not at ihal time indicate the
Source :
AZOOX ANTHELLATE SCLERACTINIA
1 17
seriefo7//ahrc!lefLZealand SPeCime"S PrCVi°US'y aS F Boston, urn, which now form part of the type
OiST-Rtsu-nOM. - Wallis and Futuna region: Wallis; Waterwitch. Combe, and Field Banks; 300-640 m
. sewhere. southern Norfolk Ridge; Three Kings Ridge: 544-590 m (Cairns, 1995).
Subgenus FLABELLUM ( VLOCYATHUS ) Sars. 1851
Flabellum (U.) deludens Marenzeller, 1904
Fig. 18 f
Flabellum deludens Marenzeller, *1904: 269-272. pi. 17 figs 10 10a — Cairns lQSOa. « ...
(synonymy); 1994: 73. pi. 32. figs d-e; 1998: 395. - Cairn! & z'ibrowius. 1997: V5i- 1 56 P S‘S
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 532, I (MNHN).
Tyre Locality. — "Valdivia"* tns 185 and 203: west of Sumatra, 614-660 m.
Remarks. Only one worn specimen is reported herein, measuring 27.8 x 16.1 mm in CD, 20.6 mm in
^unusuaU n that , r ‘n T'T™* ?2 SCP'a ‘ U Pair °f rudlmCn,ar-v S? flanki"g each St Tins specimen
.. “"usual m that its C, 2 are produced as prominent, rounded ridges (Fig. 18 f). The species is redescribed and
(1997raTabley2?AIRNS ’ ^ * ,ab'C COmparing similar sP«ies is given by Cairns & ZlBROWius
Distribution. - Wallis and Futuna region: Waterwitch Bank; 516-530 m. Elsewhere: Indo-West Pacific from
northern Indian Ocean through Indonesia to Japan; 106-1035 m (Cairns & ZlBROWius, 1997).
Flabellum (U.) aotearoa Squires, 1964
Fig. 19 e
FUtyZy°Zar0a SqUireS’ V*'964: 7‘9, PL 2- figS 15'18- ~ CA,RNS’ l995: l02-103' P‘- 33, figs d-f, i. Map 16
Flabellum sp. cf. F. deludens - Wells. v.1984: 215, pi. 4. figs 8-10 |Not Flabellum deludens Marenzeller. *19041.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn *959, 3 (MNHN). — Sin 963 25 including 1 cemented m
8,fZs T,MMel(,USNMc 98882) ~ S,n *964' 4 <MNHN). - Stn "977. I (MNHN), - S,n *1004 1 (MNHN) -
Stn 1005, 2 (MNHN). — Stn *1016, 30 (MNHN). — Stn *1017. 6 (USNM 98880) — Stn *1018 148 (MNHN
i;nn :is 2 - s,n ,06°- 1 (MNHN)- - si- '«*■ 1 Sm%,!°iVsNMMS 1
1 84!^' w“atare" S,n B'26: 35°°4'S' ' 74°23-2'E ,Bay °f IS,3ndS' nCar Ca<* Bre*«- New Zealand),
Remarks. — Although previously known from the Late Pleistocene of Vanuatu (Wells 1984) this is the
first record of living specimens from this archipelago. The species was recently redescribed and illustrated by
Cairns (1995). It is characterised by having a scalloped calicular edge; a granular, hut lustrous theca: and Si 4 with
highly sinuous axial edges. Several stations reported above (indicated with an asterisk) include populations that are
diminutive in size, the coralla never exceeding 15 mm in CCD. These specimens nonetheless have a full fifth
cycle and often have highly developed edge crests (Fig. 19 e), resulting in an edge angle (crests included) of 180°.
Distribution. — Vanuatu region: Anatom, Tanna, Erromango, Efate. Malakula, and Espiritu Santo
(including Late Pleistocene, see WELLS. 1984); 287-436 m. Elsewhere: northern New Zealand and ridges north of
New Zealand; Chesterfield Islands; 130-1300 m (Cairns. 1995).
Source :
S. D. CAIRNS
I 18
Flabellum (U.) marcus Keller, 1974
Fig. 19 f
Flabellum deludens - Vaughan, v*1907: 63-65, pi. 3. fig. 5 [Noi Flabellum deludens Marenzeller. *1904],
Flabellum marcus Keller. *1974: 208-209, pi. I, fig. 5. pi. 3, figs 5-6, pi. 5. fig. 8, text-fig. I; 1981a: 31-32. pi. I
figs 7a-b. pi. 2. figs 5a-b. — Cairns, 1984: 21.
Material EXAMINED. — Vanuatu. Musorstom 8: sin 956. 7 (USNM 98883). — Sin 1037 8 (MNHN) —
Stn 1129. I (MNHN).
Type Locality. — "Vityaz" stn 6363: 21°10'N. 163°I6’E (Marcus-Necker Ridge), 1350 m.
Remarks. — The best description of this species is that of specimens from Hawaii by Vaughan (1907), as
Flabellum deludens. The specimens reported above differ from the Hawaiian populations in having a lesser edge-
angle (125°- 142° vs 180°), and a taller pedicel (up to 4 mm vs 1.5- 1.8 mm in diameter). They are otherwise quite
similar and found at a comparable depth range.
distribution. — Vanuatu region: Anatom and Efate: Guyoi Bougainville; 1050-1175 m. Elsewhere: central
Pacific, including Marcus-Necker Ridge and Hawaiian Islands; 1261-1602 m (Cairns, 1984).
Flabellum (U.) hoffmeisteri Cairns & Parker, 1992
Flabellum (U.) hoffmeisteri Cairns & Parker. *1992: 47-48. pi. 16. figs d-f (synonymy). — Cairns. 1995: 103-104.
pi. 33. figs g-h. map 22; 1998: 394-395. — Cairns & Zibrowius, 1997: 157-158.
Material examined. — Vanuatu. Musorstom 8: stn 1014. 1 (MNHN)..
Type Locality. — "Soda” stn 27: 37°59'S, 150°05'E (off Victoria. Australia). 452 m.
Remarks. — Only one poorly-preserved corallum is reported herein measuring 33.1 x 20.6 mm in CD and
22.5 mm in height. Descriptions and illustrations can be found in Cairns & Parker (1992) and Cairns (1995),
and a table comparing similar species in Cairns & Zibrowius ( 1997: Table 2).
Distribution. Vanuatu region: Efate; 495-498 m. Elsewhere: Indonesia; Western Australia; Victoria:
Tasmania; Kermadec and Colville Ridges; 1 10-646 m (Cairns, 1998).
Flabellum (U.) apertum apertum Moseley, 1876
Flabellum apertum Moseley, v*l876: 556 (in pari: * Challenger " stn 145); v.1881: 167-168 (in pari: pi. 6. figs 7a-c> -
Cairns. 1982: 44-46, pi. 13. figs 8-1 1. pi. 14. figs 1-4 (synonymy). g
Flabellum patagonichum Moseley. v*l88l: 166-167. pi. 15. figs 1-7.
Flabellum raukawaensis Squires & Keyes, v*1967: 27, pi. 4, figs 8-9.
Flabellum apertum apertum - Cairns. 1995: 104-105. pi. 35, figs a-c, map 4 (synonymy).
meH<Rnoo'0^fXAMcINED; — Wallis a,ld Futuna region. MUSORSTOM 7: stn 551. 49 (MNHN) — Stn 553
?«55SK=ra7 fs&rss ,-«» 53 ,usnm 98885)' - sm 56?- 6 1mnhni' - * ™
Type Locality. — "Challenger" stn 145: 46°40'S, 37°50'E (off Prince Edward Island), 567 m.
Remarks. — Little can be added to the redescriptions and figures of Cairns (1982. 1995). Once thought to be
iqoc, 3n fl,b0™al distribulion- il is now ^own from the warm temperate region off New Zealand
(L-aikns, Ivys), and now from beneath tropical waters.
Source :
AZOOXANTHELLATE SCLERACTINIA
Distribution. — Wallis and Futuna region: Combe and Tuscarora Banks; 795-1280 m Elsewhere circum
Subantarctic, New Zealand region to Three Kings Islands; 220-1575 m (Cairns, 1995).
Genus TRUN CATO FLAB E I. L UM Cairns.
1989
Truncato flab ell u m stabile (Marenzellcr. 1904)
Figs 19 i-j
Flabellum stabile Marenzellcr. *1904: 273-274. p|. 17. figs Pa-h — Zibrowius 1980 ISO
Truncatoflabellum stabile - Cairns. 1989a: 61. - Zibrowius & Gili, 1990: 39. .
Truncatoflabellum sp. cf. T. stabile - Cairns & Kki.lER, 1993: 264-'}65 fie IOC F
truncatoflabellum sp. A - Cairns, 1994: 79. pi. 34. figs c-e.
1 (L™ME<)8887)EX A ? n,UiV “ MUS0RST0M 8: stn "6- 1 «MNHN). - Sin 1036.
M ^ , ~T S " 1 l25’ 2 including an anthocaulus) (MNHN). — Sin 1127. I (MNHN)
Made.ra Islands. Cancap III: stn 3.053. 4 (USNM 98886).
MNHN). — Stn 1037.
Type Locality. — "Valdivia" sin 37: 16°14'0r'N. 22°38'03"W (Cape Verde Islands). 1694
111.
thecal fecl 29° 40~Ttll f 8 ° °' Stra,gh‘ thecal cdSes 59°'73°- inclination of slightly convex
f : t ; s!„;r b T fafCS "T ln an aCUlC anglc at the edScs and are very slightly ridged within 1-4 mm
8 s i 996 28 2 x .6 1 868 rn°' ^ UrgeSt Specimcn rePorled herein (Musorstom
CCD-LCD - I 36 I 67mrrn h L mm in height. Calicular margin slightly arched and finely serrate;
CCD. LCD - 1.36-1.67, GCD:H = 1.00-1.15. Basal scar small and elliptical: 3.6-6.4 x 2.6-4 I mm
Theca overlying C,.2 produced as low, rounded longitudinal ridges, perpendicular .0 which are fine growth
lines. Theca uniformly white. Anthocaulus 5.3 mm in height and 2.0 mm in pedicel diameter, having 3\-vcles
Septa hexamerally arranged in 5 complete cycles: Sl-2>S3>S4>S5, the base of an anthocyathus having only
cycles plus several pairs of S4 (24-32 septa), and a full fourth cycle present by a GCD of 1 1 mm. Si-’ only
slightly wider than S3, whereas S3 are twice .he width of the S4, and S3 are only about 1/3 the width of the si
Axial edges of ah septa straight and vertical, those of the Sl-3 thickened deep in fossa near their fusion to the
columella. Fossa deep, containing a well-developed, elongate columella 1. 8-2.0 mm wide.
Remarks. - Although smaller than previously reported specimens from the Atlantic, Mozambique, and
ra;:h'Ch:efr mm In GCD' thC specimens reporIcd herein are bclieved 10 ^ conspecific. Marenzeller
( 904) described his larger syntype as having a GCD of 60 mm, but his figure implies a corallum with a GCD of
41 mm. The syntypes of Flabellum stabile appear to be lost (Zibrowius. 1980). Among the 29 Recent species of
Tnincatoflabellum . only four lack edge spines and edge crests: T. inconstans (Marenzeller, 1904); T. trapezoideum
(Keller. 1981); 7. panpavoninun, (Alcock, 1898); and T. stabile (Marenzeller. 1904). the last three being found at
great depths. Although I previously hesitated in identifying Pacific specimens as T. stabile. I now believe them to
be the same based on the examination of specimens from the Madeira Islands (see Material Examined).
Discussions and comparisons of the nonspinose Tnincatoflabellum species are found in Zibrowius & GlLi (1990)
and CAIRNS (1994). Although they are similar in corallum morphology. T. stabile differs from: T. trapezoideum
m having a smaller basal scar; from T. paripavoninum in having a smaller basal scar, a small edee ancle, less
septa and a more robust corallum; and from T. inconstans in having less septa, a more circular cal ice and'a much
deep depth range.
Distribution. — Vanuatu region: Erromango and Efate; Guyot Bougainville; 786-1160 m. Elsewhere: Cape
Verde, Selvagems, and Madeira Islands; Mozambique; Ryukyu Islands; 964-3010 m (Zibrowius & Gili. 1990).
Source :
120
S. D. CAIRNS
Trim cato flab ell u m dens ( Alcock, 1902)
Figs 19 g-h
F label I urn dens Alcock, v*!902a: 106-107; v.!902c: 32, pi. 4, figs 30, 30a. — Cairns, 1989a: 54, pi. 28. figs g-k.
Truncatoflabellum dens - Cairns, 1995: 1 14-1 15 (in part: pi. 37. fig. g). — Cairns & Zibrowius, 1997: 170.
MATERIAL EXAMINED. — Wallis and Futuna region. MllSORSTOM 7: stn 546, 1 (MNHN). — Stn 569. 8 (USNM
98889) — Stn 585, 2 (MNHN). — Stn 597. 1 (MNHN). — Stn 605. 4 (USNM 98890). — Stn 610. 1 (MNHN).
Vanuatu. Musorstom 8: sin 965. 3 (MNHN). — Stn 977. 1 (MNHN). — Stn 988. 3 (USNM 98888). — Stn 1015.
1 (MNHN). — Stn 1060. 1 (MNHN).
Type Locality. — "Siboga" stn 95: 5°43.5'N, 1 19°40'E (Sulu Archipelago. Philippines). 522 m.
DIAGNOSIS. — Corallum small and compressed (GCD:LCD = 1.7-2. 3), the largest specimen reported herein
(Musorstom 8 stn 1060) 8.6 mm in CD. Thecal edges rounded, at a height of about 6 mm above basal scar
changing from an angle of 60°-80° to a more slender 20°-35°. At point of angular change there is usually a pair of
short edge spines or low crests, often broken or worn away, as in the case of the syntypes. Basal scar quite small
and almost circular, ranging from 1 .4-1.7 x 1.1 -1.3 mm in diameter, containing only 6 protosepta. Well-preserved
coralla often show a reddish-brown longitudinal striping. Septa hexamerally arranged in 4 cycles (Sl-2>S3»S4);
however, pairs of S4 often lacking in lateral half-systems, resulting in 40-44 total septa. Large coralla may have
additional pairs of S5 in the end half-systems, but may also be lacking pairs of S4 in the lateral half-systems.
Axial edges of Si -2 highly sinuous.
Remarks. — This species is more fully described by Cairns (1989a, 1995). It is distinctive in having a
relatively small corallum with a bimodal edge angle, and a very small, almost circular basal scar.
Distribution. — Wallis and Futuna region: Wallis; Combe. Waterwitch, and Field Banks; 286-500 m.
Vanuatu region: Anatom. Tanna, Efate, Malakula, and Espiritu Santo; 314-410 m. Elsewhere: Philippines;
Indonesia; Kermadec and Norfolk Ridges; New Caledonia; 300-555 m (Cairns & Zibrowius, 1997).
Truncatoflabellum pusillum Cairns, 1989
Fig. 20 a
Truncatoflabellum pusillum Cairns, *1989a: 71-72, pi. 37, figs a-e. — Cairns & KELLER, 1993: 265. pi. II. fig. E. —
Cairns & Zibrowius. 1997: 170.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 973, 2 (USNM 98891). — Stn 988. I (MNHN). —
Stn 1016, 1 cemented to a Xenophora shell (MNHN). — Stn 1094, 1 (MNHN). — Stn 1097. 10, including I cemented to
a Xenophora shell (MNHN). — Stn 1098. 1 (USNM 98892). — Stn 1 106, 3 attached to a Xenophora shells (MNHN).
Type Locality. — "Albatross" stn 5178: I2°43'N, 122°06'15"E (Sibuyan Sea, Philippines), 143 m.
Remarks. — The original description suffices for this species. It is characterised as having a small corallum
(usually less than 7 mm GCD) with straight, rounded thecal edges (edge angle = I4°-I8°), each edge bearing
2-4 downward projecting thecal spines. The basal scar is elliptical, up to 3.3 mm in greater diameter, bearing
12-24 septa. Septa are arranged in 3 cycles (Si-2>S3), larger specimen having pairs of S4 in their end half-systems
(i.e., usually 24-32 septa). It differs from T. dens by having a larger basal scar with more septa, a constant edge
angle, and additional pairs of edge spines.
Distribution. — Vanuatu region: Tanna and Espiritu Santo; 285-460 m. Elsewhere: Philippines; Indonesia;
Mozambique; 85-300 m (Cairns & Zibrowius, 1997).
Source :
AZOOXANTHELLATE SCLERACTINIA
121
Truncatoflabellum angustum Cairns & Zibrowius, 1997
Fig. 20 b
Truncatoflabellum dens - Cairns, 1995: 1 14 (in part, pi. 37, figs f, h; NZOI K858)[Not Flabellum dens Alcock, v*1902).
Truncatoflabellum angustum Cairns & Zibrowius. *1997: 172-173, figs 23c-f.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 501. 1 (MNHN). — Stn 502,
2 (MNHN).
Vanuatu. Musorstom 8: stn 1016, 5 (USNM 98894). — Stn 1017, I (MNHN). — Stn 1018, 11 (MNHN). —
Stn 1065, 3 (MNHN). — Stn 1087, 1 (MNHN). — Sin 1091, 1 (USNM 98893). — Stn 1 135. 3 (USNM 98895).
Type Locality. — Musorstom 3 sin 143: 1 1°28.3’N, I24°l 1.6'E (Visayan Sea, Philippines), 205-214 m.
Remarks. — Truncatoflabellum angustum was recently described and figured. In that account, specimens from
MoNZ stn BS441 , previously illustrated as T dens by Cairns (1995: pi. 37, fig. g). were incorrectly listed as
nontype specimens of T. angustum. Instead, those specimens should be considered as valid T. dens. T. angustum
is most similar to T. pusillum , but diifers by having a larger corallum, a higher edge angle, and more septa
(i.e., usually 56).
Most of the specimens reported herein and most of the paratypes differ from the holotype in having 20 highly
sinuous primary septa, not 24 Si -3.
DISTRIBUTION. — Futuna; 516-530 m. Vanuatu region: Elate, Malakula, and Espiritu Santo; 295-394 m.
Elsewhere: Philippines; Indonesia; Kermadec Islands; 195-465 m (CAIRNS & Zibrowius, 1997).
Truncatoflabellum phoenix Cairns, 1995
Truncatoflabellum sp. B. - CAIRNS, 1994: 79, pi. 33, figs i, I.
Truncatoflabellum phoenix Cairns, *1995: 115-116. pi. 37. fig. i. pi. 38. figs a-f. — Cairns & Zibrowius, 1997: 171.
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 495. I (USNM 98896). — Stn 509,
7 (MNHN). — Stn 512. 11 (USNM 98897). — Stn 513, 4 (MNHN). — Stn 514. I (MNHN). — Sin 516. 1 (MNHN). —
Stn 538, 1 (MNHN).
Type Locality. — NZOI stn C53I: 29°14,40,,S. 178°02’W (Raoul Island, Kermadecs Islands), 179 m.
Remarks. — This species is best characterised in its original description, and is distinguished from other
species by having a small corallum with parallel thecal edges, and often elongate coralla caused by incomplete sep¬
aration of successive anthocyathi. The specimens listed above differ from previously reported specimens in having
an edge angle of 0°- 10°, which results in a calicular diameter slightly greater than that of the basal scar, and a basal
scar that reveals only 12 protosepta, instead of 24 protosepta as in typical T. phoenix. All other characters
arc similar.
DISTRIBUTION. — Wallis and Futuna region: Futuna; Waterwitch Bank; 240-441 m. Elsewhere: Philippines;
Indonesia; Ryukyu Islands; Kermadec Islands; 18-421 m (CAIRNS & ZIBROWIUS, 1997).
Truncatoflabellum vigintifarium sp. nov.
Figs 20 c-f
Material EXAMINED/TYPES. — Vanuatu. Musorstom 8: stn 1004. I paratype (MNHN). — Stn 1018.
3 paratypes (USNM 98900). — Stn 1060. I paratype (MNHN). — Sin 1065, 3 paratypes (USNM 98899). — Stn 1094,
Source :
122
S. I). CAIRNS
2 paratypes (MNHN). — Stn 1097, 5 paratypes (MNHN). — Stn 1098, holotype (MNHN) and 4 paratypes (USNM
98898). _ stn 1106, 9 paratypes (MNHN). — Stn 1113, 1 paratype (MNHN).
Type Locality. — Musorstom 8 stn 1098: 15°04'S, 167°10’E (northeast of Espiritu Santo), 277-285 m.
ETYMOLOGY. — The species name vigintifarium (Latin viginti , twenty + farius, a sullix meaning
a multiplication in number of parts) refers to the 20-fold septal symmetry of the anthocyathus.
Description of Anthocyathus. — Corallum flabellatc: angle of straight, rounded thecal edges, 67°-84°; face
angle, 25°-30°. Holotype 21.1 x 9.7 mm in CD. 17.3 mm in height, and 2.9 x 2.0 mm in basal scar diameter;
largest specimen (MUSORSTOM 8 stn 1018) 26.4 x 13.4 mm in CD. Calicc elliptical: GCD.LCD = 1.95-2.40.
Thecal edges bear 2 or 3 (usually 3) pairs of elongate (up to 5 mm), attenuate spines, the first pair occurring
3-4 mm above the basal scar and additional pairs regularly spaced at 2. 5-3.0 mm intervals. Basal scar small
(2. 9-3. 6 x 2.0-2. 7 mm) and elliptical in shape (ratio of greater to lesser scar diameters 1.24-7.37-1.60), clearly
showing 12 septa (Fig. 20 c). Theca thin and transversely wrinkled in a chevron pattern. Theca longitudinally
striped with reddish-brown pigment. Anthocaulus stage unknown.
Despite the hexameral arrangement of septa in the basal scar, the septa of mature (GCD over 20 mm)
specimens as expressed at the calicular margin are arranged 20-fold: 20:20:40, resulting in 80 septa. The distal,
axial edges of the 20 primary septa are not exsert, but project well into the fossa, having straight distal axial edges
and sinuous lower axial edges. Secondary septa much narrower, reaching their greatest width near the columella
where they are about half the width of the primaries. Tertiary septa often rudimentary, usually only half the width
and length of a secondary. Fossa deep and narrow, the columella a fusion of the lower axial edges of the 20 primary
septa.
Remarks. — Among the 30 known Recent species of Truncatoflabelliwi , only one other has decamerally ar¬
ranged septa in the adult anthocyathus stage: T. formosum Cairns. 1989. T. vigintifarium differs from that species
primarily in shape, having a more flabcllate corallum characterised by a higher edge angle (67°-84° vs 37°-59°) and
a higher GCD:LCD ( 1 .95-2.4 vs 1.4- 1.8). It has a smaller basal scar and usually one more pair of edge spines.
Distribution. — Vanuatu region: Erromango, Efate, Malakula, and Espiritu Santo; 288-700 nr, however,
most records between 300 to 400 m.
Truncatoflabellum mortenseni Cairns & Zibrowius, 1997
Truncatoflabellum mortenseni Cairns & Zibrowius, *1997: 171-172, figs 22g-h.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 523, 5 anthocauli (MNHN). —
Stn 585, 1 anthocaulus (MNHN). — Sin 601, 4 anthocauli (MNHN).
Vanuatu. Musorstom 8: stn 963, 1 anthocaulus (MNHN). — Stn 964, 1 anthocaulus (MNHN). — Sin 967,
5 anthocauli and 2 anthocyathi (MNHN). — Stn 969, 5 anthocauli (MNHN). — Stn 970, I anthocaulus (USNM 98901).
— Sin 971, 3 anthocauli (USNM 98902). — Stn 1065. 2 anthocauli (MNHN). — Stn 1070, 1 anthocyathus (MNHN). —
Stn 1071, 1 anthocaulus and 1 anthocyathus (MNHN). — Stn 1077, 1 anthocaulus (MNHN). — Stn 1102. 2 anthocauli
and 1 anthocyathus (USNM 98903). — Stn 1103, 3 anthocauli and 30 anthocyathi (USNM 98904). — Stn 1134,
1 anthocaulus and 1 anthocyathus (MNHN).
Type Locality. — Mortensen's Java-South Africa Expedition stn 5: 1 1°36'N. I21°43'E (Sulu Sea),
120-122 m.
Remarks. — This species is characterised by having an anthocaulus that, even after forming a single pair of
edge spines, usually resists the tendency to transversely divide. This results in a relatively large, triangular
anthocaulus with a small pedicel (0.9- 1.1 mm in diameter), 1 pair of edge spines, and 56. 64, or 80 septa,
depending on the development of the fifth septal cycle. Most of the specimens reported above represent this
distinctive anthocaulus stage; however, some coralla represent the anthocyathus stage, which has a greater basal
scar diameter of 6-8 mm, several pairs of edge spines, and usually a full five cycles of septa (96). Anthocauli and
anthocyathi are often found at the same station.
Source :
AZOOXANTHELLATE SCLERACTIN1A
123
Distribution. — Wallis; 350-455 m. Vanuatu region: Anatom, Malakula, and Espiritu Santo; 165-400 m.
Elsewhere: Philippines; Indonesia; 50-156 m (Cairns & ZIBROWIUS, 1997).
Truncatoflabellum Vanuatu (Wells, 1984)
Flabellum Vanuatu Wells, v*1984: 215, figs 4(11-12). 5 (1).
Truncatoflabellum Vanuatu - CAIRNS, 1989a: 63, 69. pi. 36, fig. c.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 499, 1 (MNHN). — Sin 505. I
(MNHN). — Stn 509, 6 (MNHN). — Stn 524, 4 (MNHN). — Sin 605. 1 (USNM 98906).
TYPE Locality. — Kere River (Late Pleistocene), Espiritu Santo, Vanuatu.
Remarks. — This species is characterised as having a small basal scar (4. 1-4.9 mm in greater diameter)
with 12 protosepta, low edge and face angles (20°-27°, 12°-17°, respectively), and a relatively tall corallum
that bears up to five pairs of edge spines. In small to medium-sized coralla there are 16 primary septa (16:16:
24-32, = 56-64 septa), but in larger coralla there may be up to 20 primary septa (20:20:32-40. = 72-80 septa).
Theca reported herein are all smaller than those in the type series, most having 16 primary septa and
56 septa. They are the first report of this species subsequent to its original description and first report from
the Recent.
DISTRIBUTION. — Wallis and Futuna; 240-335 m. Vanuatu region: Late Pleistocene of Espiritu Santo
(Wells, 1984).
Truncatoflabellum aculeatum (H. Milne Edwards & Haime, 1848)
Flabellum aculeatum H. Milne Edwards & Haime, v* 1848a: 272, pi. 8, Tigs 3, 3a.
Truncatoflabellum aculeatum - Cairns. 1989a: 61, 64. table 6. pi. 31. figs h-l, pi. 32. figs a-c (synonymy); 1998: 399-
400. table 4. — Cairns & Zibrowius. 1997: 166-167.
Material EXAMINED. — Vanuatu. South Santo Island, coll. J.N. CARNEY, stn SDC-5, 30-50 m. II 1975,
2 (USNM 98905).
Type Locality. — Philippines (depth not given).
REMARKS. — Although not reported from the MUSORSTOM cruises in this region, probably because of its
shallow depth range, two typical specimens are reported from an independent collection made by J. N. CARNEY in
1975. The appearance of the specimens suggests a fossil age, perhaps Late Pleistocene, similar to the fauna
reported by WELLS (1984). T. aculeatum was redescribed and illustrated by Cairns (1989a).
Distribution. — Vanuatu region: Espiritu Santo; 30-50 m. Elsewhere: Philippines; Indonesia (also
Pleistocene of Talaud); Western Australia; Northern Territory; 1 1-1 15 m (Cairns. 1998).
Truncatoflabellum candeanum (H. Milne Edwards & Haime. 1848)
Flabellum candeanum H. Milne Edwards & Haime, * 1848a: 278, pi. 8, fig. 13.
Truncatoflabellum candeanum - Cairns, 1989a: 70-71, table 6. pi. 36, figs d-h (synonymy, neotype designation); 1994;
76-77. pi. 33, figs e-f. — Cairns & Zibrowius, 1997: 167.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1086. 4: 3 (MNHN), I (USNM 98907).
Type Locality. — "Albatross" stn 5369: 13°48'N, 121°43'E (Luzon. Philippines), 194 m.
Source :
124
S. D. CAIRNS
REMARKS. — Only one lot of this species is reported herein, probably because of its relatively shallow depth
range. It is distinguished from other species by its slightly scalloped calicular margin and its three pairs of flattened
edge spines. It is more fully described and illustrated by Cairns (1989a).
Distribution. — Vanuatu region: Malakula; 182-215 m. Elsewhere: western Pacific from Kyushu through
Indonesia; 70-290 m (Cairns & Zibrowius, 1997).
Truncatoflabellum martensii (Sluder. 1878)
Figs 21 a-f
Flabellum Martensii Sluder. v*1878: 630-631. pi. I. figs 4a-b; 1889: 268.
Flabellum paripavoninum - WELLS, v.1984: 214-215 (in part: USGS 25715. fig. 4 (6-7)) (Not F. paripavoninum Alcock.
1894).
Truncatoflabellum martensii - CAIRNS, 1989a: 61.
MATERIAL EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1085. 13: 10 (MNHN), 3 (USNM 98908). — Stn 1086.
4 (MNHN).
Type Locality. — "Gazelle" stn 40: 26°51.1'S, 153°29.6’E (off Brisbane, Queensland), 139 m.
Description. — Corallum flabellate: edge angle. 40°-63°; face angle. 17°-19°. Thecal faces almost planar,
meeting in straight, sharply defined (but not carinate) edges that usually bear 3 pairs of spines. Holotype (ZMB
1798) 21.7 x 10.9 mm in CD, 7.7 mm in greater basal scar diameter, and 18.6 mm in height; largest known
specimen (Pleistocene specimen figured by Wells, 1984; USNM 71858) 28.6 x 12.0 mm in CD, 8.3 mm in
greater basal scar diameter, and 22.9 mm in height. Range of greater basal scar diameter 7. 2-9. 3 mm, well-
preserved scars showing 4 complete cycles of septa. Theca dark reddish-brown overall, with more intense
longitudinal striping associated with the S i -3 Basal scar, edge spines, and septa white. Anthocaulus unknown.
Septa hexamerally arranged in 5 cycles, small coralla having less than 96 septa, larger coralla having additional
pairs of S6 in half-systems adjacent to the principal septa (See Remarks). Septa formula: Si-3>S4>S.s>S6. Axial
edges of Si -3 highly sinuous, their lower axial edges fusing into a columellar structure low in centre of fossa.
S4-6 progressively narrower and less sinuous, their axial edges not attaining the columella.
Remarks. — The number of septa per corallum is roughly a function of the GCD. the largest specimen
(Wells' Pleistocene corallum from Vanuatu) of 28.6 mm GCD having 15 pairs of S6. or 126 septa, whereas
a small anthocyathus of 16 mm GCD (e.g., MUSORSTOM 8 stn 1085) has only 64 septa. The transition to a full
fifth cycle of 96 septa occurs at a GCD of 20-22 mm, the holotype of GCD 21.7 mm still having only 88 septa,
but other specimens of similar GCD having 96 or even 104 septa. The first Sr, occur in the four
half-systems adjacent to the two principal septa, resulting in 104 septa, and are progressively inserted in
half-systems away from the edge.
This is believed to be the first report of this species subsequent to its original description from off Brisbane.
It is distinguished from other congeners by its sharply angled thecal edges, almost planar thecal faces, and
its overall reddish-brown theca.
Distribution. — Vanuatu region: Malakula; 161-182 m. Vanuatu region: Late Pleistocene of Espiritu Santo
(Wells, 1984). Elsewhere: off Brisbane, Queensland, 139 m (STUDER, 1878).
Genus JAVANIA Duncan, 1876
Javania lamprotichum (Moseley, 1880)
Desmophyllum lamprotichum Moseley, v* 1 880: 41-42, figs 1-2.
Source
AZOOXANTHELLATE SCLERACTINIA
125
J civ an i a lamprotichum - Cairns, 1984: 21, p). 4, figs D-E; 1995: 1 12. pi. 37. Tigs h-c, map 20; 1998: 403, figs 8j, m. —
Cairns & Zibrowius, 1997: 164 (synonymy).
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: sin 520 or 521 (confusion on label).
1 (MNHN).
Vanuatu. MUSORSTOM 8: sin 974, 1 (USNM 98909). — Stn 975. I (MNHN). — Sin 988. I (MNHN). — Sin 1014.
1 (USNM 98910).
Type Locality. — Unknown.
Remarks. — Of the len Recent species in this genus, J. lamprotichum is one of five that has five cycles of
septa. It is further distinguished by having a relatively large corallum, a Hared calice. and usually having a reddish-
brown theca, although the thick tectural deposits reinforcing the pedicel are usually white. Reported herein is
the largest known specimen of this species (MUSORSTOM 8 stn 975), measuring 46.8 x 36.2 mm in CD and
47.9 mm in height, the base being broken above its attachment. One corallum, from MUSORSTOM 8 stn 974,
contains several acrothoracican cirripcdc borings, a commensalism previously reported for this species by Cairns
& Zibrowius (1997) and Cairns (1998). The species was most recently redescribed and illustrated by Cairns
(1995).
DISTRIBUTION. — Wallis and Futuna region: Wallis; 890-920 m. Vanuatu region: Tanna and Efate: 466-
536 m. Elsewhere: central and western Pacific (Hawaiian Islands, Johnston Atoll. Philippines. Kermadec Ridge);
Western Australia; 191-842 m (Cairns, 1998).
Javania fusca (Vaughan. 1907) comb. nov.
Figs 20 g-i
Placotrochus fuse us Vaughan, v*1907: 66-67. pi. 4. figs 2-3.
"Placotrochus " fuscus - Cairns, 1989a: 45. 75.
Javania pachy theca Cairns, *1995: 1 12-1 13. pi. 36. figs j-1, pi. 37, fig. a, map 17. — Cairns & Zibrowius. 1997: 165,
figs 2 1 i , 22a.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 525, I (USNM 98913). — Stn 530.
2 (MNHN). — Stn 540. I (MNHN). — Stn 578. 3 (MNHN). — Stn 592. I (USNM 98912).
Vanuatu. MUSORSTOM 8: stn 959. I (MNHN). — Stn 965, 3 (USNM 98911). — Stn 982, I (MNHN). — Stn 988.
3 (MNHN). — Stn 1097, 2 (MNHN). — Stn 1 106. I (MNHN). — Stn 1 1 14. 2 (USNM 98914). — Stn 1 128. 20 (MNHN).
Type Locality. — "Albatross" sins 3886 and 3999: Kauai and Molokai Islands, Hawaiian Islands, 271 m.
Diagnosis. — Corallum small (GCD usually less than 9 mm), straight, and ceratoid to subcylindrical, with
a thickened pedicel. Calice only slightly elliptical: GCD:LCD = 1. 1-1.2; calicular edge serrate, triangular apices
corresponding to the 24 Si-3, these projections best preserved and seen in small specimens. Theca thick, covered
with numerous, very small, porcellaneous granules. Base colour of corallum white, but often pigmented with
brownish-black rings encircling the theca, and irregular pigmentation of the same colour on the distal peripheral
faces of the Si-2. Septa hexamerally arranged in 4 complete cycles: S|-2>S3»S4 (48 septa). S i -2 1.2- 1.5 mm
exsert, having slightly sinuous axial edges. S3 up to 0.5 mm exsert and only 1/2-2/3 width of the Si-2, having
moderately sinuous axial edges. S4 nonexsert, in fact, their distal edges often several mm below the calicular edge.
S4 only 1/5- 1/4 width of the S3, also having moderately sinuous axial edges. Fossa deep and lacking a columella,
as is consistent with the generic definition.
REMARKS. — This species was originally placed in the genus Placotrochus by VAUGHAN because of the
presence of a well-formed lamellar columella in one of the three syntypes (corallum number 'T' of Vaughan.
1907, USNM 20734), notwithstanding the fact that a columella could not be found in the other two syntypes.
After careful cleaning and examination, it was discovered that the columella in specimen 1 was simply a fragment
from a distal septal margin, probably from the same specimen, that had accidently become lodged in an axial
Source :
126
S. D. CAIRNS
columellar position. This fragment has now been removed from the fossa. Lacking a columella, this species
naturally falls into the genus Javania; in fact, it is the senior synonym of a species I recently described as
J. pachytheca Cairns, 1995.
VAUGHAN (1907) originally designated three syntypes of Placotrochus fuscus, two from "Albatross" stn 3999
and one from stn 3886, listing two USNM catalog numbers for the specimens: 20731 and 20732. Whereas USNM
20732 applies to the single syntype (specimen #2: Vaughan's pi. 4, fig. 2) from "Albatross" stn 3886. USNM
20731 was previously assigned to the holotype of Gardineria hawaiiensis , a species described by Vaughan on the
page previous to Placotrochus fuscus. The true catalog numbers (see CAIRNS. 1991b) for the remaining two
syntypes from "Albatross" stn 3999 are: 20733 (specimen #3. Vaughan's pi. 4, fig. 3) and 20734 (specimen #1.
unfigured by VAUGHAN).
Two of the specimens examined have acrothoracican cirripede borings: one from MUSORSTOM 8 stn 988
(Fig. 20 i), the other being one of the three syntypes (USNM 20733).
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch, Combe, and Field Banks; 600-
730 m. Vanuatu region: Anatom. Tanna, and Espiritu Santo; Guyot Bougainville; 314-778 m. Elsewhere:
Indonesia; Malaysia; northern New Zealand: Kermadec Ridge: Lord Howe Seamount Chain; Aitutaki Atoll. Cook
Islands [not Chesterfield Islands, as incorrectly reported by Cairns (1995) and Cairns & Zibrowius (1997) for
J. pachytheca ]; Hawaiian Islands; 271-1045 m.
Java nia exserta sp. nov.
Figs 21 g-i
Desmophyllum sp. cf. D. crista-galli - Wells, 1954: 470 [Not Desmophyllum cristagalli H. Milne Edwards and Haime,
1848].
Javania sp. - Cairns & Zibrowius, 1997: 165. figs 22b-c.
MATERIAL EXAMINED/TYPES. — Indonesia. Karubar: stn 30, 2 paratypes (USNM 98919). — Stn 44. holotype
(MNHN). — Stn 49, 2 paratype fragments (USNM 97498). — Stn 86, I paratype (MNHN).
Philippines. MUSORSTOM 1: stn 65. I paratype (MNHN).
Wallis and Futuna region. MUSORSTOM 7: stn 499, 1 paratype (USNM 98916). — Stn 513, 2 paratypes
(MNHN). — Stn 514, 1 paratype (MNHN). — Sin 523, 2 paratypes (MNHN). — Stn 537. 1 paratype (USNM 98915). —
Stn 538. I paratype (MNHN). — Stn 569, 1 paratype (MNHN).
Vanuatu. Musorstom 8: stn 962, 1 paratype (MNHN). — Stn 964. 1 paratype (MNHN). — Sin 978, 1 paratype
(USNM 98918). — Stn 1021, 5 paratypes (USNM 98917). — Stn 1030. 8 paratypes (MNHN). — Stn 1042, 2 paratypes
(MNHN). — Stn 1060. 3 paratypes (MNHN). — Stn 1085, 1 paratype (MNHN).
Caroline Islands (Pelau). "Short drop off". 91 m, 1 fragment of a corallum, paratype (USNM 98956).
Type Locality. — Karubar stn 44: 7°52'27"S, 132°48'24”E (south of Tanimbar Island), 291-295 m.
Etymology. — The species name exserta (Latin exsertus , project or exsert) refers to the highly exsert Si-2 of
this species.
Description. — Corallum ceratoid and straight, having a slightly flared calicular edge. Calice slightly
elliptical: GCD.LCD = 1.05-1.20. Holotype 15.6 x 13.7 mm in CD, 33.8 mm in height, and 7.2 mm in PD.
Calicular edge highly serrate, a tall, acute triangular apex corresponding to each Si-2. Calicular edge between Si -2
of small specimens straight, but with increasing size becoming slightly convex or rounded, and in the largest
specimens expressed as a low, obtuse, triangular apex corresponding to the S3. Theca porcellaneous and usually
white; however, specimens from Karubar stations pigmented a light purple-grey. Ridged C1-2 sometimes
expressed within 0-3 mm of calicular edge.
Septa hexamerally arranged in 4 complete cycles (48 septa): Si>S2»S3>S4. Si extremely exsert, in the
holotype as much as 6.5 mm, but even in coralla of moderate size. 3-4 mm. Axial edges of Si straight and
vertical, the 4 lateral Si almost meeting in centre of fossa. S2 also quite exsert but only I/2-2/3 that of the Si,
about 3/4 width of the Si. and also less thick than the S|. S3 not exsert: rudimentary or absent near calicular edge.
Source :
AZOOXANTHELLATE SCLERACTIN1A
127
increasing to about 3/4 width of S2 lower in fossa. S4 also nonexsert, 1/3- 1/2 width of the S3. Axial edges of Si-?
slightly sinuous. Fossa deep and narrow.
Remarks. Javania exseria is distinguished from the other three Recent species in the genus that have four
cycles of septa [J. cailleri (Duchassaing & Michelotti. 1864); J. pseudoalabastra Zibrowius. 1974; and J. fusca
(Vaughan. 1907)1 by its relative septal exsertness: its Si being larger than its S2, and its S3.4 being nonexsert.
Specimens from MUSORSTOM 8 stn 1085 and Karubar stn 86 contain acrothoracican cirripede borings.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch Bank; 295-455 m. Vanuatu
region: Anatom. Tanna, Elate, Epi. and Malakula; 130-408 m. Elsewhere: Philippines; Arafura Sea south of
Tanimbar Island; Pelau; Bikini, Marshall Islands; 91-291 m.
Genus KHIZOTROCHUS H. Milne Edwards & Haime, 1848
Rhizolrochus typus H. Milne Edwards & Haime. 1848
Fig. 22 a
Rhizolrochus lypus H. Milne Edwards & Haime, *1848a: 282. pi. 8, fig. 16. — Cairns. 1989a: 79-81. pi. 41. figs f-j
(synonymy); 1994: 81. pi. 35. figs a-c, pi. 40. figs h-i (synonymy). — Cairns & Zibrowius. 1997: 161. figs 22d-e.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: sin 1021. 1 (USNM 98920). — Stn 1078 1 (MNHN) —
Stn 1131, 1 (MNHN).
Type Locality. — Singapore. South China Sea (depth not given).
Remarks. — Three specimens of this common, relatively shallow-water azooxanthcllate are reported herein.
I he genus is distinguished from other genera in the region by having several cycles of discrete (free standing, not
contiguous with corallum), hollow rootlets. R. levidensis Gardiner, 1899. known from nearby Loyalty Islands at
84 m, differs in having a much smaller corallum (GCD < 6 mm) and fewer septa (< 34). R. npus was recently
redescribed and figured by Cairns (1989a).
Distribution. — Vanuatu region: Elate, Malakula, and Espiritu Santo; 130-194 m. Elsewhere: Indo-West
Pacific from Red Sea to Japan; 20-1048 m (Cairns & Zibrowius. 1997).
Rhizotrochus flabelliformis Cairns, 1989
Flabellum latum - Alcock, v. 1902c: 31 (Not F. latum Sluder, 1878].
Rhizolrochus flabelliformis Cairns. v*!989a: 81. pi. 41. figs k-l. pi. 42, figs b. d: 1995: 109-110. pi. 35. figs g-i,
pi. 36, figs a-b, map 17.
"Rhizotrochus" flabelliformis - CAIRNS & ZIBROWIUS. 1997: 161-162.
Material EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 511. 1 (MNHN).
Type Locality. — "Siboga" stn 105: 6°08'N, 12I°19 E (Sulu Archipelago, Philippines), 275 m.
Remarks. — One relatively small (GCD=24 mm), worn specimen is reported herein, still in the process of
forming its edge rootlets. It is distinguished from all other corals in this region by having two massive rootlets,
one on each corallum edge. It is best described and illustrated by Cairns (1995).
Distribution. — Futuna; 400-450 m. Elsewhere: Philippines; Indonesia; New Zealand; 228-419 m (Cairns
& Zibrowius, 1997).
Source
128
S. I). CAIRNS
Genus POLYMYCES Cairns. 1979
Polymyces wellsi Cairns, 1991
Polymyces wellsi Cairns, *1991a: 22, pi. 8. figs f, i. pi. 9, figs a-b: 1995: 108-109. pi. 35, figs d-f, map 10. — Cairns
'& Zibrowius, 1997: 160-161. — Cairns. 1998: 403-404.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 975. I (MNHN).
Type Locality. — "Johnson-Sea-Link" stn 1916: 1°18.7*S, 89°48.8'W (Espanola, Galapagos), 545-562 m.
REMARKS. — One live specimen, with tissue preserved, is reported. This species is distinctive in having four
asymmetrically arranged, contiguous rootlets, all four rootlets occurring on one side of the corallum. It is best
described and illustrated by Cairns (1995).
Distribution. — Vanuatu region: Tanna; 536-566 m. Elsewhere: Philippines; Indonesia; northwestern
Australia; northeastern New Zealand; Kermadcc Islands; Galapagos; 355- 1 165 m (Cairns & Zibrowius. 1997).
Superfamily VOLZEIOIDEA Melnikova, 1974
Family GARDINER1IDAE Stolarski. 1996
Genus GARDINER I A Vaughan, 1907
Gardineria hawaiiensis Vaughan. 1907
Gardineria hawaiiensis Vaughan. v*1907: 65-66, pi. 4. fig. 1. — Cairns, 1984: 23; 1995: 110-111, map 13. pi. 36,
figs c-f, i; 1998: 404. — Stolarski. v.1996: 348-350. figs 2F-G. 4A-I. 8A-C.
Gardineria musorstomica Cairns, v* 1989a: 82-83. pi. 42, figs c. c-g.
MATERIAL EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1006, 1 (USNM 98921). — Stn 1011. 1 (MNHN). —
Stn 1014, I (MNHN). — Stn 1067, 1 (MNHN).
Type Locality. — "Albatross" sin 3991: 22°15,24,,N. 159°23'15"W (Kauai. Hawaiian Islands). 497-541 m.
Remarks. — This species is characterised by having a thick epithecal wall, and hexamerally arranged septa in
four cycles and three size classes, the last cycle incomplete (24-44 septa). Larger specimens possess 6 P: and a
rudimentary columella. G. hawaiiensis is best described and figured by Stolarski (1996).
Distribution. — Vanuatu region: Erromango, Efate, and Malakula; 366-574 m. Elsewhere: Philippines; New
Caledonia (Stolarski, 1996); Norfolk Ridge; Kermadec Islands; Bay of Plenty. New Zealand; Western Australia;
Hawaiian Islands; 142-602 m (Cairns, 1995).
Gardineria paradoxa (Pourtales, 1868)
Fig. 22 b
Haplophyllia paradoxa Pourtales, v*1868: 140-141.
Gardineria paradoxa - Cairns, 1979: 160-161, map 46, pi. 31, figs 4-6, 10 (synonymy). — Stolarski. v.I996: 348-350.
figs 2C-E. 5A-G. — Cairns & Zibrowius, 1997: 163, figs 2 1 g-h.
Material examined. — Vanuatu. Musorstom 8: 1014, 1 (MNHN).
Type Locality. — "Bibb" stn 22: 24°14’20"N, 80°59'40"W (Straits of Florida), 692 m.
Source : MNHN, Paris
AZOOXANTHELLATE SCLERACTINIA
129
Remarks. — The single specimen reported herein measures 9.3 mm in CD and 36.6 mm in length, having
a lateral thecal attachment for the basal 17 mm. It has 40 septa (20:20:40) and is very similar to the specimen
reported by CAIRNS & ZIBROWIUS (1997) from the Banda Sea. G. paradoxa is distinguished from other congeners
by having decamerally arranged septa, a strong lateral thecal attachment, and a heavily encrusted and worn looking
corallum, even when the coral is collected alive.
Distribution. — Vanuatu region: Elate; 495-498 m. Elsewhere: Banda Sea; western Atlantic (Antilles);
91-700 m.
Suborder DENDROPHYLLIINA
Family DENDROPHYLLIIDAE Gray. 1847
Genus BALANOPHYLLIA Searles Wood. 1844
REMARKS. — There are approximately 58 valid Recent species of Balanophyllia worldwide, and the genus is
badly in need of revision (Cairns, 1995: 1 18). There are several reasons for the confused state of taxonomy in this
genus. Pirst, the range of corallum variation is poorly known for most species, several species known only from
their type specimens. In an extreme case [i.e., B. comiculcins (Alcock, 1902a)]. the species was based on one
specimen that is now lost, and no figure of the holotype or indication of the type locality was included in the
original description. Secondly, basally-broken corallites of other genera, such as Rhizopsammia, Eguchipsammia,
Cladopsammia , and even Dendrophyllia , could be mistaken for a Balanophyllia. and the juvenile stage of all of
these genera pass through a solitary Balanophyllia-Wke stage. Third, most species of Balanophyllia are provincial
in distribution, endemic to one side of an ocean basin, and thus a large number of species have been described,
making it difficult to do a comprehensive comparison of unidentified material. Although the Atlantic species have
been revised (Cairns, 1977; Zibrowius, 1980), there has been no comprehensive worldwide revision of the
genus. A subgeneric division or a good dichotomous key would alleviate some of these difficulties. At this point,
however, it is useful to compare specimens collected from a geographic region to others known from that region.
For instance, there are 10 species known from the western Atlantic, 4 from the eastern Atlantic, 5 from the
southwestern Indian Ocean, 4 from the northern Indian Ocean, 5 from eastern Australia, 3 from New Zealand,
5 from the Hawaiian Islands, 21 from the tropical western Pacific (including northwestern Western Australia).
3 from the eastern Pacific, and one from the Subantarctic. There appears to be little cross over of species between
regions. For instance, there are no species in common between the eastern and western Atlantic, and the eastern
Pacific fauna is also discrete; however, there arc several species that have wider distributions in the tropical western
Pacific and the Hawaiian Islands and/or Japan and/or New Zealand. Therefore, in identifying the specimens collected
by MUSORSTOM 7 and 8, all species known from the western and central Pacific were considered, but, even so,
I was only able to confidently identify about one-third of the specimens available as one of six of the more
common species.
Balanophyllia desmophyllioides Vaughan, 1907
Fig. 22 c
Balanophyllia desmophyllioides Vaughan. v*1907: 149-150, pi. 45. fig. 1. — Cairns & ZIBROWIUS, 1997: 177-178,
figs 23g-h.
Balanophyllia desmophylloides -CAIRNS, 1984: 26.
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 496, 2 (MNHN). — Stn 500,
4 (MNHN). — Stn 502, 1 (MNHN). — Sin 509. 17 (USNM 98924). — Stn 512. 5 (MNHN). — Stn 524, 2 (MNHN). —
Stn 538, 1 (MNHN). — Stn 589, I (MNHN). — Stn 601 . 1 (MNHN).
Source :
130
S. D. CAIRNS
Vanuatu. Musorstom 8: stn 964, 2 (MNHN). — Stn 969. 3 (USNM 98926). — Stn 988, 1 (USNM 98922). —
Stn 1030, 1 (MNHN). — Stn 1065. ) (USNM 98925). — Stn 1098, 4: 3 (MNHN). I (USNM 98923).
Lord Howe seamount chain. NZOI: stn 1741. 9 (USNM 94366).
Type Locality. — "Albatross" stn 4061: Hawaii, 44-152 m.
Diagnosis. — Corallum straight, firmly attached, and Hared distally. Calice elongate; calicular edge arched and
constricted in centre, in some cases (Fig. 22 c) producing a division of the fossa. Costae well defined; thin epitheca
often present basally. Septa hexamerally arranged in 5 cycles, coralla of GCD 11-12 having a complete Fifth cycle;
additional half-systems of septa may occur in elongate specimens (e.g.. the holotype). Si-3 equal in size, pairs of
S5 fusing high in fossa before their Hanking S4. Lower axial edges of Si .3, 5 coarsely dentate. Fossa deep;
papillose columella small, elongate, and low.
Remarks. — The species was recently redescribed and illustrated by Cairns & ZIBROWIUS (1997).
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch and Field Banks; 240-516 m.
Vanuatu region: Anatom, Tanna, Elate, Malakula, and Espiritu Santo: 190-372 m. Elsewhere: Hawaiian Islands;
Philippines; Indonesia; Chesterfield Islands (reported herein); 95-658 m (Cairns & Zibrowius, 1997).
Balanophyllia laysanensis Vaughan, 1907
Figs 22 d-e
Balanophyllia laysanensis Vaughan, v*1907: 150-151, pi. 45. figs 2a-b.
Material examined. — Vanuatu. Musorstom 8: stn 962, 2 (MNHN). — Stn 963. 1 (MNHN). — Stn 965.
I (USNM 98927).
Type Locality. — "Albatross" stn 3937: 25°52’05"N. I71046'47MW (Laysan Island. Hawaiian Islands),
238-271 m.
Remarks. — Little can be added to the Vaughan's original description. The species can be characterised as
having a straight, ceratoid corallum with a highly serrate calicular margin. The theca is highly porous and coarsely
granular; costae are poorly defined, but the C1-2 are prominent. Septa are arranged in 4 complete cycles, pairs of S4
bending toward but not quite fusing before their enclosed S3 relatively low in fossa.
This is the first report of B. laysanensis subsequent to its original description.
DISTRIBUTION. — Vanuatu region: Anatom; 377-400 m. Elsewhere: Laysan, Hawaiian Islands; 238-271 m
(= type locality).
Balanophyllia rediviva Moseley, 1881
Balanophyllia rediviva Moseley, v* 1 88 1 : 193-194. pi. 15, figs 10-12. — Cairns & Zibrowius. 1997: 181-182,
figs 25d-f (synonymy).
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 970. 1 (MNHN). — Stn 1077, 2 (USNM 98928) —
Stn I 134, 2 (MNHN).
Type Locality. — "Challenger" stn 192: 5°49'15"S. 132°14'I5"E (Kai Islands, Banda Sea), 256 m.
Remarks. — Balanophyllia rediviva was recently redescribcd and figured by CAIRNS & ZIBROWIUS (1997). It
is distinguished by having an elongate, subcylindrical corallum that often shows signs of rejuvenescence,
the rejuvenated corallum often smaller in calicular diameter than the parent. Its costae (C1-3) are also
characteristically slightly ridged.
Source :
AZOOXANTHELLATE SCLERACTINIA
131
Distribution. — Vanuatu region: Anatom, Malakula, and Espiritu Santo; 210-252 m. Elsewhere:
Philippines; Indonesia; 90-256 m (Cairns & Zibrowius, 1997).
Balanophyllia gemma (Moseley, 1881)
Thecopsammia gemma Moseley, v*1881: 195, pi. 15, figs 8a-b.
Balanophyllia gemma - Cairns & ZIBROWIUS. 1997: 179. figs 24g-i (synonymy).
Material EXAMINED. — Vanuatu. Musorstom 8: stn 1009. 2 (MNHN). — Stn 1015, 7: 5 (MNHN), 2 (USNM
98930). — Stn 1019, 2 (USNM 98929).
Type Locality. — u Challenger" sin 201: 7°03’N, 121°48'E (Sulu Sea), 187 m.
REMARKS. — Balanophyllia gemma was recently redescribed and the hololype illustrated by CAIRNS &
Zibrowius (1997). It is characterised by having a well-developed epitheca that covers most of the theca; low. equal
costae; and a shallow fossa containing a discrete, swirled columella.
Distribution. — Vanuatu region: Efate; 397-430 m. Elsewhere: Philippines; Indonesia; 137-522 m (Cairns
& Zibrowius, 1997).
Balanophyllia gigas Moseley, 1881
Balanophyllia gigas Moseley, v* 1881: 193. — Wells, v.1984: 217, figs 5 (2-3). — Cairns. 1994: 83. pi. 35. figs j- 1
(synonymy); 1995: 1 19-120, pi. 40, figs f-h, map 7 (synonymy): 1998: 404. — Cairns & Zibrowius, 1997: 182.
Material examined. — Vanuatu. Musorstom 8: stn 977. 1 (MNHN).
Type Locality. — Japan (depth unknown).
REMARKS. — Balanophyllia gigas is well described and figured by CAIRNS (1994. 1995), including an
illustration of the holotype. The single specimen reported herein measures 24.5 x 21.5 mm in CD and 54.5 mm in
height. WELLS' (1984) Pleistocene specimens from Vanuatu (USNM 71862) were re-examined and considered
conspecific. B. gigas attains the largest size of all Balanophyllia, and usually has five cycles of septa, if not some
Sr> in various half-systems.
Distribution. — Vanuatu region: Tanna; 410-505 m; Late Pleistocene of Espiritu Santo (Wells, 1984).
Elsewhere: widely distributed in western Pacific from Hawaiian Islands to New Zealand and West Australia; 90-
640 m (Cairns & Zibrowius, 1997).
Balanophyllia crassitheca Cairns, 1995
Balanophyllia crassitheca Cairns, *1995: 120-121. map 18. pi. 40. fig. i. pi. 41. figs a-b.
Material examined. — Vanuatu. Musorstom 8: stn 978, I (MNHN).
Type Locality. — 37°17.0'S, 176°51.0'E (Rangatira Knoll, northwest of White Island. Bay of Plenty, New
Zealand), 251-308 m.
Remarks. — The single record reported herein does little more than extend the known distribution slightly to
the north. The species is distinguished by having a very thick theca and crowded septa.
Distribution. — Vanuatu region: Tanna; 408-413 m. Elsewhere: northeastern New Zealand; Lord Howe and
Norfolk Islands; Kermadec Ridge; 190-508 m (Cairns. 1995).
Source :
132
S. D. CAIRNS
Genus ENDOPACHYS H. Milne Edwards & Haime. 1848
Endopachys grayi H. Milne Edwards & Haime, 1848
Fig. 22 f
Endopachys grayi H. Milne Edwards & Haime, * 1848b: 82-83, pi. I, figs 2, 2a. — ZlBROWius & GRYGIER. 1985: 137
(New Caledonia). — Cairns, 1994: 84-85. pi. 36, figs c, h, pi. 37, fig. i (synonymy); 1995: 121-122, pi. 41,
figs c-h, map 13 (synonymy); 1998: 405. — Cairns & Zibrowius, 1997: 185-186.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: sin 499. I (MNHN). — Stn 504,
I (MNHN). — Stn 507. 1 (MNHN). — Stn 516, 1 (MNHN).
Vanuatu. Musorstom 8: stn 967. 2 (USNM 98932). — Stn 969, 1 (MNHN). — Stn 976, 17 (MNHN). — Sin 1005,
I (MNHN). — Stn 1016. 2 (MNHN). — Sin 1017, 1 cemented to a Xenophora shell (MNHN). — Stn 1018. 6 (MNHN). —
Stn 1071, 3 (MNHN). — Stn 1085, 1 (USNM 98933). — Sin 1086, 6 (USNM 98934). — Sin I 134, 2 (USNM 98931).
Type Locality. — Unknown.
Remarks. — Most specimens of this commonly collected species are easily distinguished by having
unattached, cuneiform-shaped coralla w ith prominent edge crests; asexual budding; and five cycles of septa. Its wide
distribution may be due to its reproductive success, which employs two modes of asexual reproduction: transverse
division and anthoblast production (= bud shedding)(see Cairns, 1989b). In the first case, once an attached
anthocaulus reaches a height of about 7 mm and a GCD of 3. 5-4.0 mm, it forms a crested anthocyathus that
subsequently transversely divides from the anthocaulus (Fig. 22 f). The detached, free-living anthocyathus
maintains the characteristic basal scar (greater diameter 3. 5-4.0 mm) for a time but eventually heals its base, which
becomes rounded. Each anthocyathus has the potential to form numerous buds (anthoblasts), which form at the
calicular edge, usually adjacent to the edge crests. Only one other coral species is known to employ both asexual
reproduction strategies, Blastotrochus nutrix H. Milne Edw-ards & Haime, 1848. a member of a different suborder.
Distribution. — Wallis and Futuna region: Futuna; 390-441 m. Vanuatu region: Anatom, Tanna,
Erromango, Efate, Espiritu Santo, and Malakula: 181-360 m. Elsewhere: widespread throughout tropical and warm
temperate Indo-Pacific, from the southwestern Indian Ocean to the Gulf of California; 37-386 m (Cairns &
Zibrowius, 1997).
Genus HETEROPSAMMIA H. Milne Edwards & Haime. 1848
Heteropsammia cochlea (Spengler, 1781)
Madrepora cochlea Spengler. *1781: 240-248, figs A-D.
Heteropsammia cochlea - VERON & PlCHON, 1980: 416-420. in part: figs 727, 729 (synonymy and description). —
Hoeksema & Best, 1991: 234-237, figs 24-28 (synonymy!). — Cairns, 1998: 406-408.
MATERIAL EXAMINED. — Wallis and Futuna region. Musorstom 7: stn 494, 83 (MNHN). — Stn 495.
9 (USNM 98937). — Stn 496, 30 (USNM 98935). — Stn 504. 10 (MNHN). — Stn 509, 1 1 (USNM 98936). — Stn 512,
3 (MNHN). — Stn 516. 9 (MNHN).
Vanuatu. MUSORSTOM 8: stn 961, 1 (MNHN). — Stn 967, 1 (MNHN). — Stn 969. 1 (MNHN). — Stn 976. I (MNHN)
— Stn 1072. 1 (MNHN).
Type Locality. — Tranquebar, off southeastern India.
Remarks. — The species is easily distinguished by its obligate symbiotic association with a sipunculid
worm, which is housed in the base of the corallum and communicates with the environment through one large
efferent pore in the base of the corallum and several smaller pores on the lower theca. It is similar in shape to
Source : MNHN. Pans
AZOOXANTHELLATE SCLERACTINIA
133
species of Heterocyathus, which also lives with an obligate sipunculid. but differs in having a porous upper theca,
and septa that are arranged in a Pourtales plan. H. cochlea is best described and figured by Veron & PlCHON
(1980) and HOEKSEMA & BEST (1991). Shallow-water representatives of this species are assumed to be
zooxanthellate, whereas the deeper specimens must be azooxanthcllate.
DISTRIBUTION. Wallis and Futuna region: Futuna; 110-441 m. Vanuatu region: Anatom. Tanna. and
Espiritu Santo; 110-622 m. Elsewhere: widespread throughout tropical Indo-West Pacific; 9-137 m, although
depths have rarely been reported (HOEKSEMA & BEST. 1991 ).
Genus DENDROPHYLLIA Blainville, 1830
Dendrophyllia ijimai Yabe & Eguchi, 1934
Dendrophyllia ijimai Yabc & Eguchi. * 1934b: 2026. — EGUCHI. 1965: 294. 2 figs; 1968: C65 (in part: pi. C16. figs 1-2.
pi. C22. fig. D(synonymy). — Cairns & Keller. 1993: 280. fig. I3G. — Cairns. 1995: 89. pi. 38. figs c, f
(synonymy).
? Dendrophyllia subcornigera cylindrica Eguchi. *1968: C64-65. pi. C32. figs 1-2 [Not D. subcornigera subcornigera
Eguchi. 1968].
Dendrophyllia subcornigera - WELLS, v.1984: 215-216. fig. 5 (4-5) [Not D. subcornigera subcornigera Eeuchi. 1968].
Dendrophyllia sp. cf. D. ijimai - Cairns & ZlBROWlUS, 1997: 191-192. fig. 29e.
MATERIAL EXAMINED. — USGS stn 25715 ( Dendrophyllia subcornigera of Wells. 1984): figured specimen (USNM
71863). 12 branches (USNM 73976).
Type Locality. — Unknown, but presumed to be from off Japan.
REMARKS. — Dendrophyllia ijimai belongs to the "first group" of Dendrophyllia species sensu CAIRNS
(1995), this group characterised by having a robust axial coral I ile from which additional corallites bud at right
angle. WELLS (1984) correctly identified the Vanuatu Pleistocene specimens as D. subcornigera. listing D. ijimai
as a junior synonym, not realizing that D. ijimai was described earlier. The nominal subspecies of D. subcornigera
is probably a junior synonym of D. arbuscula van der Horst. 1922 (see Cairns, 1995). D. ijimai was recently
redescribed and figured by Cairns (1995).
Distribution. — Vanuatu region: Pleistocene of Espiritu Santo (Wells. 1984). Elsewhere: southwestern
Indian Ocean to Japan; 10-366 m (Cairns. 1995).
Dendrophyllia arbuscula van der Horst, 1922
Dendrophyllia arbuscula van der Horst, v*1922: 53 (in part: ,,Sibogau stn 277, pi. 8. fig. 6). — Cairns. 1994: 90-91.
pi. 38, figs i-l (synonymy); 1995: 125-126. pi. 43. figs e-f, map 15; 1998: 408-409. — Cairns & Zibrowius. 1997:
192-193. figs 29a-c (lectolype designated).
Dendrophyllia horsti Gardiner & Waugh. v*1939: 237-238. pi. 2. figs 5-6.
Material EXAMINED. — Vanuatu. Musorstom 8: sin 1018, 4 colonies (USNM 98938). — Stn 1021. 1 colony
(MNHN). — Sin 1030. 2 colonies and 2 branches (MNHN). — Sin 1058. 2 colonies and 1 branch (MNHN).
Type Locality. — "Siboga" stns 260 and 277: Banda Sea. 45-90 m.
Remarks. — Dendrophyllia arbuscula is the only species reported herein that belongs to Dendrophyllia
"group 2" sensu Cairns (1995). i.e., species having relatively small, bushy colonies with irregular branching
from a short, but robust axial corallite. It is distinctive in having a relatively shallow fossa with a well-developed
columella that is often constricted into three lobes. It w'as recently redescribed and figured by Cairns (1994. 1995).
Source :
134
S. I) CAIRNS
Distribution. — Vanuatu region: Efate and Malakula; 130-319 m. Elsewhere: Indo-West Pacific from
southwestern Indian Ocean to Japan; 2-353 m (CAIRNS. 1998).
Dendrophyllia alcocki (Wells, 1954)
Sclerhelia alcocki Wells. v*!954: 465-466. pi. 177. figs 1-2.
Dendrophyllia alcocki - ZlBROWlUS, v. 1974b: 570-573. figs 10-14. — Cairns. 1995: 126-127. pi. 43. figs g-i. pi. 44,
figs a-b, map 3 (synonymy); 1998: 408, fig. 9g. — Cairns & ZlBROWlUS. 1997: 193.
MATERIAL EXAMINED. — Wallis and Futuna region. MUSORSTOM 7: stn 506. I colony (MNHN). — Stn 514.
3 branches (USNM 98943).
Vanuatu. MUSORSTOM 8: stn 971. 4 colonies (MNHN). — Sin 977, 1 colony (MNHN). — Stn 978. 1 branch
(MNHN). — Stn 980, 1 branch (MNHN). — Stn 982. 4 branches (MNHN). — Stn 983. 1 colony and 2 branches (USNM
98940). — Stn 988, 4 branches (USNM 98942). — Stn 1015. 2 branches (USNM 98939). — Sin 1019. I branch (USNM
98941). — Stn 1023, 2 branches (MNHN). — Stn 1095, 1 branch (MNHN). — Sin 1 108, 1 branch (MNHN).
Type Locality. — Bikini Atoll, Marshall Islands, 177-243 m.
Remarks. — Dendrophyllia alcocki belongs to the "third group" of Dendrophyllia species sensu Cairns
(1995), i.e., those species having dendroid coralla produced by sympodial branching. D. alcocki is further
distinguished as having dense, spinose coenosteum (porous only near distal branch tips); three cycles of septa; and
prominent P2. It was recently redescribed and illustrated by Cairns (1995).
Distribution. — Wallis and Futuna region: Futuna; 355-400 m. Vanuatu region: Anatom. Tanna. Efate, and
Espiritu Santo; 315-475 m. Elsewhere: Indo-West Pacific from Maidive Islands to New Zealand, including the
Marshall Islands and South China Sea; 1 18-616 m (Cairns, 1998).
Genus ENA LLOPSA MM I A Michelotti, 1871
Enallopsammia rostrata (Pourtales, 1878)
Amphihelia rostrata Pourtales. v*1878: 204. pi. 1. figs 4-5.
Dendrophyllia amphelioides Alcock, v* 1902a: 112-113.
Enallopsammia rostrata - ZlBROWlUS, 1973: 44-45, pi. 2, figs 14-15. — Cairns. 1982: 57. pi. 18, figs 1-4 (synonymy);
1994: 92-93, pi. 39, figs d-f (synonymy); 1995: 127-128. pi. 44. figs c-f, map 5. — Cairns & Zibrowius, 1997:
195.
Enallopsammia amphelioides - Zibrowius. 1973: 45-46 (Tuamotu Archipelago).
MATERIAL EXAMINED — Wallis and Futuna region. MUSORSTOM 7: stn 501. 5 nonrostrate branches (MNHN).
— Stn 520, 2 rostrate branches (MNHN). — Stn 522, 3 rostrate branches (USNM 98947). — Stn 530, 1 nonrostrate
branch (MNHN). — Stn 574, 2 rostrate colonies (MNHN).
Vanuatu. MUSORSTOM 8: stn 959, 4 nonrostrate branches (USNM 98945). — Stn 974. 3 nonrostrate branches
(MNHN). — Stn 977, 3 nonrostrate colonics and 3 branches (MNHN). — Stn 1006, 2 nonrostrate branches (USNM
98946). — Stn 1015, 4 nonrostrate branches (MNHN). — Stn 1024. 1 nonrostrate colony and 2 branches (USNM
98944).
Type Locality. — "Blake" stn 2: 23°14'N, 82°25'W (Straits of Florida), 1472 m.
Remarks. — This widespread species varies in calicular diameter and the expression of the costoseptal rostrum
(see Cairns, 1982. 1995). All Vanuatu specimens lack the costoseptal rostrum and have calices of intermediate
size (2. 6-3. 5 mm GCD), whereas those from the Wallis and Futuna region represent rostrate and nonrostrate forms
(see Material Examined). There seems to be no correlation between having a rostrum and GCD, the rostrate forms
ranging lrom 2.3 (MUSORSTOM 7 stn 574) to 4,1 (MUSORSTOM 7 stn 530) mm in GCD. E. rostrata was recently
redescribed and figured by Cairns (1994, 1995). It is easily distinguished from other colonial deep-water corals
Source : MNHN, Paris
AZOOXANTHELLATE SCLERACTINIA
135
from this region by having a flabellate colony with unifacially arranged corallitcs, the corallites often having
a prominent costoseptal rostrum; and three cycles of normally inserted septa.
The corallum of one specimen from MUSORSTOM 7 sin 501 contained several acrothoracican crustacean
borings.
Distribution. — Wallis and Futuna region: Wallis and Futuna; Waterwitch Bank; 400-920 m. Vanuatu
region: Anatom. Tanna, Erromango, and Efate; 370-574 m. Elsewhere: cosmopolitan, except for eastern Pacific
and off continental Antarctica; 1 10-2165 m (Cairns & Zibrowius. 1997).
ACKNOWLEDGEMENTS
I would like to thank Bertrand RICHER DEFORCES (ORSTOM. Noumea). Philippe BOUCHET (BIMM. Museum
national d'Histoire naturclle. Paris) and Alain CROSNIER. who collected the specimens used in this study. Alain
CROSNIER provided too the support and encouragement to complete the study.
I also thank the following people, who generously loaned specimens used in the study and/or facilitated visits
to their collection: Richard Preece (University Museum of Zoology, Cambridge). Eric Lazo-Wasem (Yale
Peabody Museum), Kei Mori [Institute of Geology and Paleontology, Tohoku (Imperial) University, Sendai).
Maya Best (Nationaal Natuurhistorisch Museum, Leiden). Sheila Halsey (The Natural History Museum,
London), and Peter Bartsch (Zoologisches Museum, Berlin).
The scanning electron photomicrographs were taken in the SEM Laboratory, NMNH.
I am also grateful to Helmut Zibrowius, who carefully reviewed an early draft of this manuscript.
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142
S. D. CAIRNS
Fig. 1 a-e. — Mad rac is kauaiensis : a. MUSORSTOM 7 stn 509 (MNHN): a large colony, x 0.46. — b-e, MUSORSTOM 7
stn 509 (USNM 98541): b. branch with 3 corallites, x 15; c. coenosteal spines, x 1 15; d. calice. x 38; e, close up of
columella and paliform lobes, x 100.
Fig. I f-h. — F ungiacyathus sandoi. holotype. MUSORSTOM 7 stn 538 (MNHN). oblique, basal, and stereo calicular views,
all x 3.0.
Source :
AZOOX ANTHELLATE SCLERACTINIA
143
Fig. 2 a. — Fungiacyathus paliferus, Musorstom 8 stn 1003 (MNHN), oblique calicular view, x 3.3.
Fig. 2 b-c. — Fungiacyathus margaretae : b. MUSORSTOM 7 stn 567 (MNHN), oblique calicular view, x 3.3. —
c, Musorstom 7 stn 565 (MNHN), granular base, x 3.1.
Fig. 2d. — Fungiacyathus variegatus, MUSORSTOM 9 stn 963 (MNHN). corallum cemented to a Xenophora shell, along
with a corallum of Tropidocyathus lahidus , x 6.6
Fig. 2 e. — Madrepora oculata forma formosa, Musorstom 7 stn 609 (USNM 98580). stereo view of a calice with
asymmetrically arranged paliform lobes, x 23.
Fig. 2 f. — Madrepora oculata forma tenuis . MUSORSTOM 7 stn 552 (USNM 98548), calice lacking paliform lobes and
columella, x 16.5.
Fig. 2 g-h. — Anthemiphyllia pacifica , MUSORSTOM 8 stn 1031 (MNHN), side and calicular views of a pedicellate
corallum. x 3.9, x 4.3. respectively.
Source :
144
S. D. CAIRNS
Fig. 3 a-b. — Anthemiphyllia multidentata , hololype, Cronulla (USNM 83010), detail of septal edge dentition and stereo
calicular view, x 6.3, x 2.0, respectively.
FlG. 3 c-d. — Anthemiphyllia macrolobata , holotype, Townsend Cromwell stn 81-01-14 (USNM 60559). side and
calicular views, both x 3.1.
FlG. 3 e-h. — Anthemiphyllia patera costata: e, MUSORSTOM 7 stn 586 (MNHN), calicular view of holotype. x 5.6. —
f-h. paratype, Musorstom 7 stn 594 (USNM 98555). calice and details of septal spines, x II. x 24, x 75.
respectively.
Source : MNHN, Paris
AZOOXANTHELLATE SCLERACTIN1A
145
FlG. 4 a-b. — Anthemiphyllia patera costata, paratype. MUSORSTOM 7 stn 594 (USNM 98555), edge views of costal
granulation, x 8.5. x 17.5. respectively.
Fig. 4 c-j. — Anthemiphyllia spinifera : c-e. Musorstom 7 stn 605 (MNHN): c-d. stereo calicular and basal views of
holotypc, both x 5.0; e. 9 paratypes, x 2.0. — f-i. paratypes, MUSORSTOM 7 stn 610 (USNM 98570): f-g. calicular
views, x 5.8. x 6.6. respectively; h-i, paratypes, septal spines, x 27. x 75, respectively. — j. paratype, MUSORSTOM
8 stn 1014 (MNHN), anthocyathus attached to its anthocaulus, x 9.5.
Source :
146
S. D. CAIRNS
FIG. 5 a-b. — Caryophyllia crosnieri : a. Musorstom 8 stn 974 (MNHN). broken cOrallum revealing ridged pali and deep
fossa. X 6.7. — b. Musorstom 8 stn 973 (MNHN), calice, x 5.0.
Fig. 5 c, f. — Caryophyllia marmorea. MUSORSTOM 7 stn 525 (MNHN), side and ealieular views, x 6.7. x 15.2,
respectively.
Fig. 5 Cl-e. — Caryophyllia abrupta, holotype. Musorstom 7 stn 535 (MNHN), side and calicul ar views, x 2.6, x 6.6,
respectively.
FIG. 5 g-i. — Caryophyllia sp. cf. C. calveri , MUSORSTOM 8 stn 1056 (MNHN): g-h. side and ealieular views of corallum
with 12 pah. x 4.1, x 5.0, respectively; i. ealieular view of corallum with 10 pali. x 6.8.
Source :
AZOOXANTHELLaVTE scleractinia
147
Fig. 6 a-b. — Crispatotrochus rugosus, MUSORSTOM 8 stn 1043 (MNHN). side and calicular views, x 1.4. x 2.0.
respectively.
Fig. 6 c, f. — O.x ys mi I ia circulars , MUSORSTOM 8 stn 977 (NMNH). side and calicular views, x 1.7. x 1.9. respectively.
Fig. 6 d-e. — Oxysmilia epithecata, holotype. MUSORSTOM 8 stn 1018 (MNHN), side and calicular views, x 5.3. x 6.3,
respectively.
Fig. 6 g-h. — Oxysmilia corrugata, holotype. Musorstom 8 stn 1030 (MNHN). side and stereo calicular views, x 4. 1.
x 4.9. respectively.
Source : MNHN, Paris
148
S. D. CAIRNS
Fig. 7 a. — Oxysmilia corrugata , paratype, Musorstom 8 stn 1030 (MNHN), side view showing thecal ridges, x 4. 1.
Fig. 7 b-g. — Oxysmilia epithecata : b, paratype, Musorstom 8 stn 1023 (NMNH). cluster of 5 corallites. —
c-g, paratypes, Musorstom 7 stn 496 (USNM 98626): c, costal granules near calice. x 41; d-e. calicular and stereo
side views of same specimen, x 20, x 14.5, respectively; f-g. epithecal ridges on base of coral Lum, x 22, x 450,
respectively.
Source : MNHN, Paris
AZOOXANTHELLATE SCLERACTINIA
149
Fig. 8 a-b, I'. — Trochocyathus vasiformis , Musorstom 8 sin 977 (MNHN): a-b: side and calicular views, x 1.9. x 4. 1,
respectively; c, fracture revealing menianes on pali and twisted columellar elements, x 15.
Fig. 8 c. — Bourneotrochus stellulatus , Musorstom 7 stn 509 (USNM 98708). palar ring and columella, x 25.
Fig. 8 d-e. — Trochocyathus efateensis . holotype. Musorstom 8 stn 1019 (MNHN), side and stereo calicular views, x 2.8.
x 3.4. respectively.
Fig. 8 g. — Trochocyathus patellifonnis , holotype. HURL stn P5-063 (USNM 83026). stereo calicular view, x 2.4.
Source : MNHN. Paris
150
S. D. CAIRNS
Fig. 9 a-c. — Trochocyathus patelliformis : a-b, holotype, HURL stn P5-063 (USNM 83026), basal and side views, x 2.4.
x 2.2, respectively. — c, paratype. MUSORSTOM 8 stn 1 1 1 1 (MNHN). juvenile calice, x 4.4.
Fig. 9 d-e. — Trochocyathus discus . MUSORSTOM 8 stn 1089 (MNHN), calicular and side views, x 3.8. x 3.4. respectively.
Fig. 9 f-i. — Trochocyathus brevispina : f, i, MUSORSTOM 8 sin 963 (MNHN), bases of coralla that were originally
attached to a bivalve and a gastropod, x 2.7. x 3.0. respectively. — g. Musorstom 8 stn 1004 (MNHN), side view of
nodular proliferations on costal spines, x 2.9. — h. Musorstom 8 stn 1003 (MNHN), calicular view of nodular
proliferations on costal spines, x 2.8.
Source :
AZOOXANTHELLATE SCLERACTINIA
151
Fig. 10 a-c. — Polycyathus octuplus : a-b. holotype. Musorstom 7 sin 494 (MNHN). side and calicular views, x 6.0,
x 7.5, respectively. — c. paralype, Musorstom 7 sin 504 (MNHN), calice of septamerally symmetrical corallum.
x 7.5.
Fig. 10 d-g. — Bourneotrochus stellulatus : d-e. Musorstom 7 stn 509 (USNM 98708): d. stereo oblique calicular view.
x6.5; e. anthocaulus with incipient anthocyathus attached, x 16. — f-g. Musorstom 7 sin 511 (USNM 98704):
f, basal scar resulting from transverse division, x 1 1 ; g. costae at calicular edge, x 54.
Fig. 10 h. — Stephanocyathus regiiis . Musorstom 7 stn 509 (MNHN). juvenile corallum with 96 septa, x 2.9.
Source :
152
S. D. CAIRNS
Fig. II a-c. — Stephanocyathus regiits, Musorstom 8 stn 1129 (USNM 98657). base of rapidly growing corallum
showing progressive magnifications of tufts of fibres, x 20. x 580. x 1450. respectively.
Fig. 1 1 d-f. — Stephanocyathus coronatus : d-e. Musorstom 8 stn 1125 (MNHN). side and basal views, both x 1.2. —
f. Musorstom 7 stn 622 (MNHN). juvenile "flatiliseptis" stage, x 3.8.
Fig. II g-h. — Vaughanella concinna, Musorstom 7 stn 572 (MNHN). side and calicular views, x 1.6. x 1.7.
respectively.
Fig. Hi. — Deliocyaihus magnificus. Musorstom 8 stn 963 (MNHN). corallum with hexagonal outline, x 2.2.
Source :
AZOOXANTHELLATE SCLERACTINIA
153
Fig. 12 a-b. — Deltocyathus taiwanicus : a. Musorstom 7 sin 541 (MNHN). calicular view, x 3.2. — l>. Musorstom 7
sin 589 (USNM 98678). processes uniting S4 to adjacent S3, x 40.
Fig. 12 c-f. — Deltocyathus crassiseptum : c. f. paralype. Musorstom 8 stn 978. calicular and basal views, both x 3.5. —
d-e, holotype, Musorstom 8 stn 980 (MNHN). calicular and basal views, both x 3.3.
Fig. 12 g-i. — Deltocyathus cameratus . Musorstom 8 stn 1007 (MNHN): g-h, holotype. calicular and basal views, both
x 3.5: i. paralype, worn corallum showing calicular chambers, x 4.6.
Source : MNHN, Parts
S. D. CAIRNS
154
Fig. 13 a. — Deltocyathus cameratus, paratype, MUSORSTOM 8 stn 1036 (MNHN), specimen with ornately sculptured
columella, x 5.5.
Fig. 13 b-c. — Deltocyathus Stella , MUSORSTOM 7 stn 513 (MNHN), calicular and basal views, both x 4.7.
Fig* 1 3 d-g. — Deltocyathus heteroclitus : d. Musorstom 8 stn 1 106 (MNHN). corallum cemented to a Xenophora shell.
x 7.1. e-g, Musorstom 7 stn 514 (MNHN): e-f. calicular and basal views, both x 7.0; g. series of 9 specimens,
those in lower row having 7 or 8 costal spines, x 2.3.
Fig. 13 h-i. — Deltocyathus ornatus , holotype. Lifu (Cambridge), calicular and basal views, both x 4.0.
Source :
AZOOXANTHELLATE SCLERACTINIA
155
Fig. 14 a-d. — Heteroeyathus sp. cf. H. sulcatus , Musorstom 7 sin 513 (USNM 98712): a. side view showing sipunculid
efferent pore, x 14; b. granules of costae, x 105; c, stereo calicular view, x 1 1; d. series of 9 coralla (MNHN). x 2.0.
Fig. 14 e-f. — Heteroeyathus alternatus , MUSORSTOM 8 stn 1004 (MNHN). side and calicular views, both x 4.8.
Fig. 14 g. — Conotrochus asymmetros , holotypc, MUSORSTOM 7 stn 496 (MNHN), calicular view, x 6.5.
Source :
156
S. D. CAIRNS
Fig. 15 a-e. — Conotrochus asymmetros , paratypes: a-h. e. MUSORSTOM 7 stn 495 (USNM 98735): a. stereo calicular
view, x 14; b. inner calicular edge, x 37; e, eolumellar elements, x 35. — c-d. MUSORSTOM 7 stn 513 (MNHN), side
and calicular views ol coralla with and elliptical and an elongate calice. x 4.3. x 6.3, respectively.
Fig. 15 f-g. — Lochmaeoirochus gardineri, holotype. MUSORSTOM 8 sin 1036 (MNHN). side and calicular views, x 3 0
x 4.1, respectively.
Fig. 15 h. — Aulocyathus juvenescent. MUSORSTOM 8 stn 976 (MNHN), a longitudinally fractured and an intact specimen,
x 3.3.
Source :
AZOOXANTHELLATE SCLERACTINIA
157
Fig. 16 a-f. — Dactylotrochus cervicornis , MUSORSTOM 7 stn 514: a. side view of corallum (MNHN), x 3.2; b-e, USNM
98746: b, bifurcated extension, x 11.8; c, 3 septa viewed from above, x 51; d. cross section of several septa showing
alternation of adjacent menianes, x 74; e-f. higher magnification of meniancs and fibre arrangement on broken
surface, x 150. x 300, respectively.
Fig. 16 g-i. — Asterosmilia gigas, MUSORSTOM 7 stn 515 (MNHN): g-h. side and calicular views, x 0.8, x 1.7,
respectively; i. longitudinal section revealing dissepiments in upper corallum. x 2.5.
Source : MNHN. Paris
158
S. D. CAIRNS
Fig. 17 a-b. — Caryophyllia (= Asterosmilia) gigas , holotype (BM 1939.7.20.851), side and calicular views, x 0.65.
x 1.4, respectively.
Fig. 17 c. Tropidocyathus lessonii , MUSORSTOM 8 stn 1016 (MNHN), side view of specimen with large edge crests,
x 2.7.
Fig. 17 d-e. — Pleotrochus venustus , Musorstom 8 sin 1114 (MNHN), side and calicular views, x 3.5. x 4.0,
respectively.
Fig. 17 f. i. — Cry plot roc hits brevipalus , holotype. Musorstom 8 stn 1034 (MNHN). side and calicular views, x 4.5,
x 4.3, respectively.
Fig. 17 g-h. — Pleotrochus zibrowii, Musorstom 7 sin 637 (MNHN), side and calicular views, x 3.5, x 3.6, respectively.
Source : MNHN . Paris
AZOOXANTHELLATE SCLERACTINIA
159
Fig. 18 a-b. — Peponoeyathus folliculus, Musorstom 8 sin 108S (MNHN): a. specimen cemenied lo a Xenophora shell
( Trochocyathus discus also attached on lower shell), x 1.5; I). oblique calicular view, x 9.5.
Fig. 18 c. — Trimcatoguynici irregularis , MUSORSTOM 8 stn 967 (MNHN). 2 coralla. x 2.2.
Fig. 18 d-e. — Temnotrochus kermadecensis . Musorstom 8 stn 963 (MNHN); d. specimen cemented to a Xenophora
shell, x 1.8; e, side view, x 12.1.
Fig. 18 f. — F label I urn deludens . MUSORSTOM 7 stn 532 (MNHN). side view, x 1.4.
Fig. 18 g-i. — Flabellum pavoninuni forma coalilum (MNHN): g. MUSORSTOM 8 sin 1102. calicular view, x 1.6;
h. Musorstom 8 stn 1004. side view showing acrothoracican burrows, x 2.2; i. series of 6 coralla showing
ontogenetic development of edge spines, x 0.75.
Source :
160
S. D. CAIRNS
Fig. 19 a-d. — Flabellum arcuatile: a-b. d. holotype, NZOI sin 197 (NZOI H688). calicular edge, and side views, x 1.5.
x 2.2. x 1.3, respectively. — c. paratype. Musorstom 7 sin 535 (MNHN). broken corallum showing sinuosity of
sepial edges, x 2.5.
Fig. 19 e. — Flabellum aotearoa , Musorstom 8 sin 1018 (MNHN), side view of juvenile corallum having large edge
crests, x 3.0.
Fig. 19 f. — Flabellum marcus. Musorstom 8 sin 1037 (MNHN), side view, x 1.3.
Fig. 19 g-h. — Truncaioflabellum dens, Musorstom 8 stn 1060 (MNHN). side and calicular views, x 3.8. x 5 8.
respectively.
Fig. 19 i-j. — Truncaioflabellum stabile, Musorstom 8 sin 996 (MNHN). side and calicular views, x 1.7. x 2 1
respectively.
Source : MNHN, Pahs
AZOOXANTHELLATE SCLER ACTINIA
161
Fig. 20 a. — Truncatoflabellum pus ilium, Musorstom 8 stn 1097 (MNHN). series of 6 coralla. x 2.2.
Fig. 20 b. — Truncatoflabellum angustum , MUSORSTOM 8 stn 1018 (MNHN), series of 4 coralla. x 1.6.
Fig. 20 c-f. — Truncatoflabellum vigintifarium (MNHN): c. paratype, Musorstom 8 stn 1 106. basal scar of anthocyathus,
x 9.0. — d-f. holotype, Musorstom 8 stn 1098 (MNHN). side. edge, and calicular views, x 2.4. x 3.4, x 2.7.
respectively.
Fig. 20 g-i. — Javania fusca (MNHN): g-h. Musorstom 8 stn 1 128 (MNHN), side and calicular views, x 2.9, x 4.2.
respectively. — i. Musorstom 8 stn 988 (MNHN), corallum with an acrothoracican burrow, x 4.6.
Source :
162
S D. CAIRNS
FIG. 21 a-f. Truncatoflabellum martensii : a-c, f. Musorstom 8 stn 1085 (MNHN): a-c, side, calieular. and edge views,
x 2.2. x 2.9. x 2.4. respectively; f. scries of 8 coralla showing ontogeny of anthocyathus, x 0.8. — d-e. holotvpe
Gazelle stn 40 (ZMB 1798), side and calieular views, x 2.2, x 2.8, respectively.
Fig. 21 g-i. — Javania exserta (MNHN): g-h, holotype, Karubar sin 44 (MNHN). side and calieular views, x 2.4. x 3 4
respectively. — i. paratype, Musorstom 1 sin 65 (MNHN), side view, x 1.5.
Source :
AZOOXANTHELLATE SC LERACTINIA
163
Fig. 22 a. — Rhizotrochus typus , Musorstom 8 stn 1131 (MNHN), basal rootlets, x 2.2.
Fig. 22 b. — Gardineria paradoxa . MUSORSTOM 8 stn 1014 (MNHN). side view, x 1.6.
Fig. 22 c. — Balanophyllia desmophyllioides . Musorstom 8 stn 1098 (MNHN). calice about to undergo intratentacular
division, x 3.0.
FlG. 22 d-e. — Balanophyllia laysanensis, MUSORSTOM 8 stn 963 (MNHN), side and calicular views, x 3.7. x 5.3.
respectively.
Fig. 22 f. — Endopachys grayi . MUSORSTOM 8 stn 976 (MNHN), anthocaulus and anthocyathus, x 2.8.
Source : MNHN. Paris
164
S. D. CAIRNS
INDEX
Generic and subgeneric names used herein in combination with specific or subspecific names, or in citation, are
given in parentheses. A slash (/) separates variant spellings; spellings retained herein in first position.
Taxa of generic and species level that receive full taxonomic treatment herein are in bold.
Page numbers in bold refer to full taxonomic treatment; page numbers in italics refer to illustrations.
Index does not include the station list (pages 32-43) and the geographic distribution Table 2 (pages 48-5 1 ).
abrupt a (Cary ophy Ilia) 71-72, 113, 146
abyssorum (Caryophyllia) 73
Acanthocyathus 76
Acinocyathus 84. 9 0
aculeatum (Flabellum, Truncatoflabellum ) 123
Alatotrochus 108-109
alberti ( Caryophyllia ) 74
alcocki ( Dendrophyllia , Sclerhelia ) 47. 136
alt e mat us (Heterocyathus) 99-100, 155
ambrosia ( Caryophyllia ) 49, 75-76
amphelioides ( Dendrophyllia , Enallopsammia) 136
Amphihelia 1 06
andamanicus ( Deltocyathus ) 63, 67
angiostomum ( Flabellum ) 116, 117
angustum ( Truncatoflabellum ) 121. 161
an nu lata ( Guynia ) 48, 113-114
Anthemipliyllia 63-69, 143-144
antillarum ( Caryophyllia ) 73
aotearoa ( Flabellum . Ulocyathus) 117. 160
apertum ( Flabellum , Ulocyathus) 118-119
Aplocyathus 84-86
arbuscula ( Dendrophyllia ) 133-134
arcuatile (Flabellum) 114, 116-117,/ 60
arnoldi (Caryophyllia) 74
Asterosmiiia 1 07-108. 1 5 7
a sym metros (Co not roc h us ) 101-102, 114,
155-156
atlantica ( Caryophyllia ) 74
atlanticus (Aulocyathus) 104
Aulocyatlius 103-104, 156
australiensis (Peponocyathus) 1 1 1
Bala nop hyllia 47, 129-130, 163
barbadensis (Caryophyllia) 12
Bathyactis 57-58. 1 63
Blastotrochus 132
Bourneotrochus 84. 87-88. 149, 151
brevipal us (Cryptotrochus ) 112, 158
brevispina (Aplocyathus, Troc hocyathus)
85-86. 150
brunneus (Conotroclius . Pleurocyathus) 101
cailetti ( Javan ia) 1 27
calcar ( Deltocyathus) 96
calveri (Caryophyllia) 73-74. 1 46
cameratus (Deltocyathus) 95. 153-154
candeanum (Flabellum. Truncatoflabellum)
123-124
cardlinensis ( Cryptotrochus) 1 1 2
Caryophyllia 46. 49, 69-76. 80. 107. 1 46
caryophylloides (Trochocyathus) 82
catinata (Anthemipliyllia) 67
Ceratotrochus 77, 109
cervicornis ( Dactylotrochus, Tridacophvl I ia)
106-107, 157
circu laris ( Oxy sin ilia ) 78. 147
Citharocyathus 1 1 1
Cladopsammia 129
davits (Caryophyllia) 46. 70
coalition (Flabellum) 115, 116, 159
cochlea ( Heteropsammia, Madrepora) 132-133
Coenopsammia 46
complicata (Step ha nop hyllia) 6 0
concinna (Vaughanella) 49. 90-91. 152
conic us (Citharocyathus. N otocyatlius) 48. Ill
Conotroclius 100-102. 155-156
cooperi (Trochocyathus, Tropidocyathus) 83-84
corniculans (Balanophyllia) 129
cornuformis ( Caryophyllia) 12
coronatus ( Odontocyathus . Platycyathus,
Stephanocyathus) 49, 89, 152
corrugata (Caryophyllia) 75
corrugata (Oxysmilia) 78-79, 147- 148
corrugatus (Deltocyathus) 96, 9 8
costata (Anthemipliyllia) 64. 66-67, 68.
144-145
crassiseptum (Deltocyathus) 94-9 5,153
crassitlieca (Balanophyllia) 103
Crispatotrochus 76-77. 14 7
cristagalli (Desmophyllum) 105. 126
crosnieri (Caryophyllia) 70. 146
Cryptotrochus 112. 158
Culicia 46
Cyathoceras 76
Cyathohelia 61
Cyathotrochus 110-111
cylindrica (Dendrophyllia) 133
Cryptotrochus 109
Dactylotrochus 106-107. 15 7
delicatum (Desmophyllum) 105
Deltocyathoides 111
Deltocyathus 47, 48, 63, 67, 69. 87, 9 1 -98,
111, 114, 152-154
de lit dens ( Flabellum , Ulocyathus) 117. 118.
159
Source : MNHN, Paris
AZOOXANTHELLATE SCLERACTINIA
165
De ndrophyllia 47, 129. 133- 1 34
dens (Flabellum, Truncatoflabellum ) 120. 121.
160
dentata/dentatus (Anthemiphyllia, Discotrochus)
63-65, 66, 68
desmophyllioides/desmophylloides
( Hal an op hy Ilia) 129-130, 1 63
Desmophyllum 104-105, 124. 126
dianthus ( Desmophyllum . M adrepora) 104-105
diomedeae ( Cary op hy Ilia ) 7 4
diomedeae ( Crispatotrochus) 76
Discotrochus 63, 67
discus (Trochocyathiis) 48, 84, 1 13, 150, 159
ef at ee ft s i s ( Trochocyathiis ) 82, 149
Eguchipsammia 129
el on gala (C dry ophy Ilia) 70
Enallopsamrnia 46, 47, 134-135
Endopachys 49. 132. 163
epithecata (Oxysmilia) 79. 147-148
exserta (Javania) 126-127, 1 62
flabe lliformis (Rhizotrochus) 127
Flabellum 46, 47. 110, 1 15-1 19. 123, 124.
127, 159-160
flatiliseptis (Sabinotrochus) 89, 152
folliculus (Peponocyathus , Stephanophxllia)
48, 49, 1 13, 159
fonnosa ( Cyathohelia , Madrepora, Sclerohelia) 61, 143
formosissima (Letepsammia) 59
formosum (Truncatoflabellum) 122
formosus (Deltocyathus) 92
fragilis (Deltocyathus) 91
fragilis (Fungiacyathus) 46, 55, 57, 58
franki (Letepsammia) 5 9
frustum (Anthemiphyllia) 64
fulvus (Polycyathus) 87
Fungiacyathus 46. 48, 54-58. 142-143
funicolumna (Ceratotrochus, Conotrochus)
100-101. 103
furanaensis ( Polycyathus ) 87
fuse a (Javania. Placotrochus) 125-126. 161
gardineri ( Loc h maeotroc h us ) 102-103. 156
Gardineria 126, 128- 129. 163
gemma ( Balanop hyllia . Thecopsammia) 131
gigas (Asterosmilia, Carxophxl lia, Rhizosmilia)
107-108, 157-/58
gigas ( Balanop hyllia) 131
granulosus (Batliyactis . Fungiacyathus) 5 8
grayi (Acanthocyathus. Cary ophy Ilia) 76
grayi (Endopachys) 132. 163
Guynia 48, 113-114
Haplophyllia 128
hastatus ( Acinocyathus , Aplocyathus,
Stephanocyathus, Trochocyathiis) 84-85, 87
hawaiiensis ( Cary op hyllia ) 69-70
hawaiiensis (Gardineria) 126, 128
hellena (Madracis) 46, 53-54
heteroc litus (Deltocyathus) 48, 69, 96.
97-98. 154
Heterocyathus 98-100. 155
H eterops a mm ia 132-133
hodgsoni ( Polycyathus ) 87
hoffmeisteri (Flabellum, Ulocyatlius) 1 18
horsti (Dendrophyllia) 133
Idiot roc lius 112-113
ijimai (Dendrophyllia) 133
impensum (Flabellum) 1 16
inconstans (Truncatoflabellum) 1 19
inter jecta (Madracis) 53
investigatoris (Discotrochus) 63
irregularis (T r u nc at o guynia) 114. 159
ita liens (Deltocyathus) 95. Ill
ixine (Odontocyathus, Stephanocyathus) 89
Javania 124-127. 162
javanus (Cryptotrochus) 1 12
juvenescens (Aulocyathus) 104, 156
kauaiensis (Madracis) 53-54, 142
kauaiensis (Madrepora) 61
kermadecensis (Temnotroc hus) 48. 114-115,
159
kikutii (Idiotrochus , Placot rochides) 112-113
knoxi (Flabellum) 116
labidus (Tropidocyat hus ) 48, 110. 143
Labyrinthocyathus 11
lamellif era (Cary ophy Ilia) 7 4-75
lamprotichum ( Desmophyllum . Javania) 124,
1 25
latum (Flabellum) 127
lays anen sis (Bala nop hyllia) 130. 163
lessonii/lessoni (Flabellum. Tropidocyathus)
108. 158
Letepsammia 5 9
levidensis ( Rhizotrochus) 46, 127
li mat ulus ( Ceratotrochus. Labyrinthocyathus)
1 1
Loch maeotroc hus 102-103, 156
Ion gi spina (Aplocyathus, Trochocyathiis) 85
Loplielia/Lophohelia 61. 105-106
mabahithi (Caryophyllia) 72
macrolobata (Anthemiphyllia) 64. 66. 1 44
maculatus ( Trochocyathiis ) 81-82
Madracis 46. 53-54. 142
Madrepora 61-62. 105. 132. 143
magnificus ( Deltocyathus ) 47. 91 -92. 1 52
marcus ( Flabellum , Ulocyatlius) 46. 118. 160
margaretae ( Batliyactis . Fungiacyathus) 57.
58, 143
marigondoni (Polycyathus) 87
marmorea (Caryophyllia) 72-73. 1 46
martensii ( Flabellum, Truncatoflabellum) 124.
162
Source :
166
S. D. CAIRNS
matricidus ( Aulocyathus ) 104
minutiseptum (Madrepora) 6 1-62
mortenseni (Truncatoflabellum) 122-123
multicarincitus (Fungiacyathus) 55
multidental a (Anthemip hyllia) 64, 65, 144
musorstomica ( Gardineria) 1 28
neglecta (Stephanophyllia) 59. 60
Neohelia 46, 47. 62
norfolkensis (Polycyathus) 87
Notocyathus 48, 1 1 1
nutrix (Blastotrochus) 132
octonaria (C ary op hyllia) 71. 72
octopali (Caryophyllia) 72
oc tu plus ( Polycyathus ) 86-87. 1 5 1
oculata (Madrepora) 61. 143
oculeus ( Lqchmaeotrochus ) 103
Oculina 6 0
Odontocyathus 89-90, 152
oreophila ( Vaughanella) 90
orientalis (Deltocyathoides , Deltocyathus) 1 1 1
ornatus ( Deltocyathus ) 96. 97, 98, 154
Oulangia 46
Oxysmilia 78-79, 147- 148
pachy theca ( Javan ia) 125-126
pacifica ( Ant h ernip hyllia ) 64, 65-66. 1 43
pacifica (Caryophyllia) 46
pacific us (Fungiacyathus) 55. 5 6
paliferus/palifera (Bathyactis, Fungiacyathus)
55. 57. 14 3
panda (Caryophyllia) 74
papuensis ( Phyllangia ) 46
paradoxa ( Gardineria , Haplophyllia) 128-129.
163
paripavoninum (Truncatoflabellum) 119. 124
patelliformis (Trochocyathus) 82-83, 149-150
patera (Anthemiphyllia) 64. 67
patagonichum ( Flabellum ) 118
paucipalata ( Caryophyllia ) 70
pavoninum (Flabellum) 46, 47, 1 15-1 16. 159
Peponocyathus 48. 49, 111. 113. 159
perculta (Caryophyllia) 73. 74
pertusa (Lophelia, Madrepora) 105-106
philippinensis (Trochocyathus) 8 1
phoenix (Truncatoflabellum) 121
Phyllangia 46
pileus (Cyathotrochus, Trochocyathus.
Tropidocyathus) 110-111
Placotrochides 1 1 2
Placotrochus 125
Platytrochus 108
Pleotrochus 109, 1 5 8
Polycyathus 86-87, 1 5 1
polygona (Caryophyllia) 74
Polymyces 49, 128
porcellana (Madrepora. Neohelia) 46. 47.
62-63
pourtalesi (Deltocyathus) 95
primordialis (Tridacophyllia) 1 06- 1 07
profunda ( Asterosmilia ) 108
prolifera (Lophelia. Madrepora) 105
pseudoal abas Ira ( Javan ia) 1 27
pusillu m (Truncatoflabellum) 120. 121. 16 1
quadrage naria (Caryophyllia) 73-74
raukawaensis (Flabellum) 1 18
recidivus (Aulocyathus . Ceratotrochus) 103,
104
rediviva ( Balanop hyllia) 130-13 1
regius ( Stephanocyat hits ) 88-89. 151, 152
Rhizopsammia 129
Rhizosmilia 8 1. 10 7
Rhizotrochus 46, 127. 163
rhomb ocolumna (Trochocyathus) 8 1
Rhombopsammia 59
robust a (Rhizosmilia) 107
rostrata (Amphihelia, E n allop sammia) 46, 47.
134-135
rotulus (Deltocyathus . Trochocyathus) 91-92
rotundifolia (Oxysmilia) 78
rubescens (Alatotrochus . Platytrochus.
Sphenotrochus) 108-109
rubescens (Crispatotrochus . Cxathoceras)
76-77
rugosa ( Caryophyllia ) 71, 72, 7 5
rugosus (Crispatotrochus) 77. 147
Sabinotrochus 89
sandoi (Fungiacyathus) 55, 56-57. 142
Sclerhelia/Sclerohelia 61, 134
scobinosa (Caryophyllia) 46, 7 5
secta (Caryophyllia) 12
sentperi (Trochocyathus) 83
sibogae ( Fungiacyathus) 54
sibogae (Stephanotrochus) 89
singularis (Madracis) 46. 53-54
smithii (Caryophyllia) 70
solida ( Caryophyllia) 73
sp. A (Fungiacyathus) 55
Sphenotrochus 108
spinifera (Anthemiphyllia) 64. 67-69. 97,
145
spinifera (Acinocyathus. Odontocyathus,
Stephanocyathus Stephanotrochus) 90
stabile (Flabellum, Truncatoflabellum) 49,
119. 160
Stella (Deltocyathus) 96-97. 1 54
stellulatus ( Bourneotrochus . Deltocyathus) 84.
87-88, 149, 151
Stephanocyathus 49. 84, 88-90. 151-152
Stephanophyllia 59-60, 113
Stephanotrochus 89, 90
stephanu (Bathyactis . Fungiacyathus) 54.
55. 57
subcornigera (Dendrophyllia) 133
sulcatus (Heterocyathus, Stephanoseris) 98-99.
100, 1 14. 155
Source :
AZOOXANTHELLATE SCLERACTINIA
167
suluensis (Deltocyathus) 91, 92, 93
taiwanicus ( Deltocyathus ) 92-93, 153
Te m not roc hus 48, 114-115. 159
tenuescens (Desmophyllum. Thalamophyllia)
1 0 5
tenuicalyx (Trochocyathus) 81
teriuis ( Lophohelia, Mad repo ra) 61 , 143
Tethocyathus 86
Thalamophyllia 105
Thecopsammia 131
trapezoideum (Truncatoflabellum) 1 19
Tridacophyllia 106
Trochocyathus 48, 80-86, 87. 1 10.
149-150, 159
Tr opidocy athu s 48. 110, 158
Truncatoflabellum 49, 116, 1 19-124 .160-162
Truncatoguynia 114, 159
Tubas traea 46
typus ( Rhizotrochus ) 127, 163
Ulocyathus 117-119
Vanuatu (Flabellum, Truncatoflabellum) 123
variabilis ( Peponocyathus) 113
variegatus ( Bat hy act is , F ungiacyathus) 48.
58. 143
vasiformis ( Trochocyathus ) 80. 149
Vaughanella 49, 90-91. 152
vaughani (Deltocyathus) 93-94
venustus ( Ceratotrochus , Cryptotrochus,
Pleotrochus) 109. 15 8
venustus ( Notocyathus ) 1 1 1
veroni ( Bourneotrochus ) 87
vigintifarium (Truncatoflabellum ) 121-122.
161
virgatus (Tethocyathus , Trochocyathus) 86
virgosa (Oculina) 60
weberian us (Odontocyathus, Step h a nocy at hus
Stephanotrochus) 89-90
wellsi (Polymyces) 49. 128
willeyi (Coenopsammia) 46
Xenophora 48. 58. 60. 71. 88. 97. 111. 113. 114. 115,
120. 143, 159
zibrowii (Cryptotrochus, Pleotrochus) 109. 112.
158
zopyros ( Caryophyllia ) 73
Source : MNHN. Paris
Source : MNHN, Paris
'SULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 20 — RESULTATS D1
Entoproctes et Bryozoaires Cheilostomida
(Pseudomalacostegomorpha et Cryptocystomorpha)
des campagnes MUSORSTOM
autour de la Nouvelle-Caledonie
Jecm-Loup D'HONDT
Museum national d’Histoire naturelle.
Laboratoire de Biologic des Invertebres marins et Malacologie
U.R.A. 699 du C.N.R.S.
57, rue Cuvier, F - 75231 Paris Cedex 05
&
Dennis P. GORDON
National Institute of Water and Atmospheric Research (NIWA)
P.O. Box 14-901 Kilbirnie
Wellington. New Zealand
RESUME
Cette etude concerne la systematique des Entoproctes et des Bryozoaires Cheilostomes (infra-ordre des Pseudo-
malacostegomorphes et des Cryptocystomorphes) recueillis lors de differentes campagnes oceanographiques autour de la
Nouvelle-Caledonie. Une nouvelle espece d'Entoproctes du genre Loxokalypus est decrite, et 12 families (dont une
nouvelle), 27 genres (dont 2 nouveaux) et 40 especes (dont 16 nouvelles) de Bryozoaires Cheilostomes sont recensees.
Les nouveaux taxons de Bryozoaires crees ici sont la famille Bryopastoridae. les genres Promicroa et Lamouroiixici et le
sous-genre Henrimilnella. Une nouvelle cle de determination des genres de Cellariidae est proposee. Outre les especes
prec£dentes, une nouvelle espece du genre Himantozoiun (Cheilostomes Cellularines) est decrite. Lc genre
Pseudothyracella , prealablement connu uniquement du Paleogene du nord-ouest de I'Europe et de I'Amerique du Nord. est
represente par une nouvelle espece. actuelle. Treize genres et 19 especes sont mentionnes pour la premiere fois de la
faune neo-caledonienne.
HONDT, J.-L. D’ & Gordon, D.P.. 1999. — Entoproctes et Bryozoaires Cheilostomida (Pseudomalacostegomorpha et
Cryptocystomorpha) des campagnes MUSORSTOM autour de la Nouvelle-Caledonie. In: A. Crosnikr (ed.), Resultats des
Campagnes MUSORSTOM, Volume 20. Memoires du Museum national d'Histoire naturelle, 180: 169-251. Paris ISBN 2-
85653-520-8.
Source : MNHN. Paris
170
J.-L D'HONDT & D P. GORDON
ABSTRACT
Entoprocta and Bryozoa Cheilostomida (Pseudomalacoslcgomorpha and Cryptocystomorpha)
from the MUSORSTOM cruises around New Caledonia.
This study concerns the systematics of Entoprocta and Cheilostomatc Bryozoa (infraorders Pseudo-
malacostegomorpha and Cryptocystomorpha) collected during various cruises around New Caledonia. One new entoproct
species is described in the genus Loxokalypus, and 12 families (1 new), 27 genera (2 new), and 40 species (16 new) of
Bryozoa are recorded. The new bryozoan taxa comprise the family Bryopastoridae, the genera Lamouroiixia and Promicroa
and the subgenus Henrimilnella. A new key is provided for the identification of genera of Ccllariidae. A new species of
the buguloidean bryozoan Himantozoum is also provided. The genus Pseudothyracella , previously known only from
the Paleogene of Northwestern Europe and North America, is represented by a new, living species. Thirteen genera and
19 species are newly recorded in the New Caledonian fauna.
INTRODUCTION
Cette note s'inscrit dans le programme d'etude systematique des Bryozoaires Eurystomes (= Ctenostomcs +
Cheilostomes) marins recueillis a grande profondeur lors des campagnes MUSORSTOM autour dc la Nouvelle-
Caledonie, la zone geographique prospectee incluant les lies Loyaute et Bellona. Lc materiel correspondant, avec
notamment les types des nouveaux taxons decrits, est depose au Museum national d'Histoirc naturelle de Paris.
Les recherches sur les Ctenostomes ont revele l'existence sur place de 5 especes profondes, dont 4 nouvelles
pour la faune neo-ealedonienne, appartenant a 5 genres (4 nouveaux pour la Nouvelle-Caledonie) ct 4 families
(3 nouvelles pour la faune locale). Leur etude a etc publiee conjointement a celle de trois des Sous-Ordres de
Cheilostomes non-ascophores, les Malacosteges, les Scrupariines et en panic les Neocheilostomina (I’lnfra-Ordre
des Cellulariomorpha) par D'HONDT et GORDON (1996). Le present travail concerne les deux autres Infra-Ordres des
Neocheilostomina. les Pseudomalacosteges et les Cryptocystomorphcs. Lc Sous-Ordre des Inoviccllatina est
present en Nouvelle-Caledonie, mais ne s'y rencontre que dans les eaux superficielles (D’HONDT. 1986), cc qui
explique qu'il n’a pas ete recolte lors des campagnes MUSORSTOM.
Les quatre Sous-Ordres precites correspondent aux "Anascina" ou "Anasca" des anciens auteurs, taxon artificiel
et polyphyletique qui n'est plus actuellement utilise que par souci de commodite, par opposition au cinquicme
Sous-Ordre des Cheilostomes, celui des Ascophorina. En effet, les Malacosteges et les Scrupariines. distincts entre
eux, se differencient des autres Cheilostomes par leurs morphologie et anatomic larvaires et leur type de
metamorphose (inconnue chez les Scrupariines) ; les Inovicellatina, dont les larves n'ont pas ete decrites, sont
morphologiquement tres ditferentes des autres Bryozoaires par leur forme zoeciale ; quant aux Pseudomalacosteges,
aux Ccllularines et aux Cryptocystomorphes, qui se situent sur trois lignees evolutives morphologiquement
distinctes a 1'etat adulte, ils presentent tous trois le meme type larvaire et morphogenetique, qu'ils partagent
egalement avec le dernier des Sous-Ordres des Cheilostomes. les Ascophorina. Cette observation a incite D'HONDT
(1985) a inclure ces derniers dans les Neocheilostomina, alors que Gordon (1989) a prcfcrc les maintenir comme
Sous-Ordre distinct, considerant 1'acquisition de I'asque comme un pas evolutif important. Ce dernier auteur a
partage les Ascophorina en 4 Infra-Ordres.
Les Ascophorina profondes de Nouvelle-Caledonie sont en cours d'etude tres avancee, ayant actuellement donne
lieu a plusieurs publications : Gordon (1988 et 1993), Gordon & Braga (1994), Gordon & D'HONDT (1991).
Les aires geographiques prospectees autour de la Nouvelle-Caledonie ont etc recapitulees sur une carte publiee
dans un precedent travail (D'HONDT & Gordon, 1996), a laquelle nous invitons le lecteur a se reporter ; les
provenances des specimens etudies dans notre travail anterieur et dans celui-ci sont en effet les memes, les recoltes
de materiel ayant ete effectives lors des memes campagnes (Biocal, Chalcal 2, Biogeocal. Musorstom 4.
Smib 3, Musorstom 6, Smib 4, Calsub). Nous avons joint a cette etude la determination de quelques espcces de
Bryozoaires recueillies autour dc l'archipel des Philippines, lors des missions Musorstom 3 et Estase.
Les numeros des stations de recolte sont precedes, dans la liste ci-apres, d'un code indiquant I'instrumcnt de
prelevement : DC : drague Charcot ; DW : drague Waren ; CP : chalut a perche ; KG : carottier Usnel grande
surface ; PL : soucoupe "Cyanea".
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDON1E
171
Les abreviations utilisees pour designer les etablissements ou sont deposes les echantillons etudies sont :
MNHN : Museum national d'Histoire naturelle. Paris.
NMNZ : National Museum of New Zealand. Wellington.
NZOI : New Zealand Oceanographic Institute. Wellington.
Lorsque le lieu dc depot n'esl pas mentionne, ceci implique que l'echantillon se trouvc dans les collections du
Museum national d’Histoire naturelle.
Les noms des navires ainsi que celui des soucoupes sous-marines sont en italiques, enlre guillemets.
Dans tout le travail, les mesures indiquees ont etc effectuees sur 20 a 25 autozoecies arbitrairement choisies sur
les echantillons etudies.
LISTE DES STATIONS
Philippines
Musorstom 3
Station CP 101. — 1.06.85, 14°00’S, 120°19’E, 194-196 m : Carbasea aff. linguiformis.
Station DR 117. — 3.06.85, 12°31,3'S, 120°39’E, 97-92 m : Cellaria humilis , Paraniropora laguncula.
Station CP 139. — 6.06.85, 1 1°53'S, 122°15'E, 240-267 m : Micropora equilateralis.
Estase
Station DR 07. — 28.1 1.84. 05°56,79'N. 126°14,38'E, 890-450 m : Mesostomaria strictoramae.
Nou veil e-Caledon ie
BlOCAL
Station DW 08. — 12.08.85, 20°34'S, 166°54'E, 435 m : Melicerita alternans, Bryopastor pentagonus.
Station CP 13. — 12.08.85, 20°18’S, 167°I8’E, 3690-3740 m : Columnella magna.
Station DW 31. — 29.08.85, 23°07’S, 166°50'E, 850 m : Pseudothyracella canclelaber, Carbasea laterogranulata.
Station DW 33. — 29.08.85. 23°09,71'S. 167°10,27'E, 675-680 m : Mesostomaria strictoramae.
Station DW 36. — 29.08.85. 23°08,64'S, 167°10,99'E. 625-650 m : Mesostomaria strictoramae . Formosocellaria
magnifica, Bryopastor crass us.
Station DW 38. — 30.08.85, 22°59,74'S, 167° 1 5,3 1 'E. 360 m : Cryptostomaria alata. Melicerita (Henrimilnella)
articulata , Melicerita ( Henrimilnella ) laurifolia , Bry opastor pentagonus. Pseudothyracella candelaber.
Station DW 41. — 30.08.85, 22°45,13'S, 1 67° 1 1,74'E, 380 m : Columnella vipera , Bryopastor crassus.
Station DW 44. — 30.08.85, 22°47,30'S, 167°14,30'E, 440-450 m : Calloporidae indeterminable, Bryopastor sp.
(B. crassus ?), Bryopastor sp. (B. octogonos ?).
Station DW 46. — 30.08.85, 22°53,27'S, 167°17,41'E, 570-610 m : Concertina cultrata, Pseudothyracella
candelaber , Quadricellaria bocki , Bryopastor pentagonus.
Station DW 51. — 31.08.85, 23°05,27'S, 167°44.95'E, 700-680 m : Carbasea laterogranulata. Pseudothyracella
candelaber. Bryopastor crassus.
Station CP 52. — 31.08.85, 23°05,79'S, 167°46,54'E, 600-540 m : Columnella vipera. Mesostomaria
strictoramae, Quadricellaria bocki.
Station DW 53. — 1.09.85, 23°09,80’S, 167°42,57'E, 1005-975 m : Columnella vipera.
Station CP 54. — 1.09.85, 23°10,30'S, 167°42,55’E. 1000-950 m : Columnella vipera.
Station CP 55. — 1.09.85, 23°19,76'S, 167°30,46'E, 1 175-1 160 m : Columnella vipera.
Station CP 60. — 2.09.95. 24°01,45’S, 167°08.43'E, 1530-1480 m : Columnella vipera.
Station CP 62. — 2.09.85, 24°19,06'S. 167°48,65'E, 1395-1410 m : Columnella vipera . Cryptostomaria alata.
Station DW 65. — 3.09.85, 24°47.90’S, 168°09.09’E, 275-245 m : Cranosina coronata.
Station DW 66. — 3.09.85, 24°55,43'S, 168°21,68'E, 515-505 m : Promicroa dubitata, Lamourouxia
canal iculata, Syringotrema calobi. Cel lari idae incertae sedis. Bryopastor pentagonus.
Station CP 67. — 3.09.85, 24°55,44'S, 168°21,55'E, 500-510 m : Calloporidae incertae sedis. Lamourouxia
canaliculata.
Source :
172
J.-L. D’HONDT & D. P. GORDON
Station DW 70. — 4.09.85. 23°24,70'S. 167°53.65'E. 965-960 m : Columnella vipera , Pseudothyracella
candelaber.
Station KG 71. — 4.09.85. 22°09,85'S. 167°32,70'E, 2099 m : Larnourouxia canaliculata.
Station CP 74. — 4.09.85. 22°14,06’S, 167°29,30'E, 1300 m : Cryptostomaria alata .
Station CP 75. — 4.09.95, 22°18.65'S, I67°23.30'E, 825-860 m : Quadricellaria bocki, Cryptostomaria alata.
Station KG 76. — 5.09.95. 22°21,09’S, 167°23.00,E. 880 m : Columnella vipera.
Station DW 77. — 5.09.85. 22°15,33'S, 167°15,40'E, 440 m : Cryptostomaria alata.
Station CP 78. — 5.09.85, 22°16,26'S, 1 67° 1 5.53'E. 445-450 m : Crateropora stiliformis.
Station CP 84. — 6.09.95, 20°43,50'S, 166°54,03’E, 460 m : Smittipora fenestrata.
Station KG 103. — 8.09.85, 21°29,15'S, 166°19.98'E, 630 m : Euginoma conica.
Station CP 109. — 9.09.95, 22°10,03'S, 167°15,22'E, 495-515 m : Pseudothyracella candelaber. Cryptostomaria
alata.
MUSORSTOM 4
Station DW 150. — 14.09.85, 19°07,5'S, 163°22,1'E, 110m: Parantropora laguncula.
Station DW 151. — 14.09.85. 19°07.0’S. 163°22.0'E. 200 m : Mesostomaria strictoramae.
Station CP 153. — 14.09.85, 19°04,2’S, 163°21.2'E. 235 m : Cryptostomaria alata.
Station CP 172. — 19.09.85, 19°01,2’S. 163°16.0'E, 275-330 m : Cryptostomaria alata.
Station CC 175. — 17.09.85, I8°59.3'S, 163°17,5'E, 355 m : Melicerita ejuncida, Cryptostomaria alata .
Bryopastor pentagon us.
Station DW 187. — 19.09.89. 19°08,3'S, 163°29,3,E, 65-120 m : Nellia tenella.
Station CP 216. — 29.09.85, 22°59.5’S, 167°22.0'E. 490-515 m : Pseudothyracella candelaber.
Station DW 220. — 29.09.85, 22°58.5’S, 167°38.3'E, 505-550 m : Carbasea laterogranulata, Pseudothyracella
candelaber.
Station DW 221. — 29.09.85, 22°58,6'S, 167°36,8'E. 535-560 m : Formosocellaria magnifica.
Station DW 223. — 30.09.85, 22°57,0'S, 167°30,0'E. 545-560 m : Himantozoum crassiavicularium.
Station CP 236. — 2.10.85, 22° 1 1,3'S. 167°I5,0'E. 495-550 m : Pseudothyracella candelaber.
Station CP 238. — 2.10.85, 22° 13,0’S. 167°14.0’E, 500-510 m : Columnella vipera.
Chalcal 2
Station DW 81. — 31.10.86, 23° 19,60’ S. 168°03,40'E, 311 m : Loxokalypus pedicellinoides.
BlOGEOCAL
Station CP 205. — 8.04.87, 22°40.61'S, 166o28,01'E, 1350-1380 m : Columnella vipera.
Station KG 210. — 9.04.87, 22°44'S, 1 66°3 1 'E, 1 190 m : Euginoma conica . Bryopastor challengeri.
Station CP 214. — 9.04.87, 22°43.09'S, 166°27,19'E, 1665-1590 m : Cellaria parafistulosa. Cellaria obliquidens.
Station KG 219. — 10.04.87, 22°38,81'S, 166°33,63' E, 570 m : Euginoma conica.
Station KG 222. — 1 1.04.87, 22°44.62’S, 166°24,93’E, 1675 m : Melicerita ejuncida.
Station KG 227. — 12.04.87, 21°32.84’S. 166°23.85’E, 500 m : Euginoma conica.
Station CP 232. 12.04.87, 2 1 °33.8 1 'S. 166°27,07'E, 760-790 m : Melicerita ejuncida. Bryopastor challengeri,
Bryopastor pentagonus.
Station DW 253. 16.04.95, 21°31,75'S, 16°28,73'E, 310-315 m : ? Pseudolunularia sp.. Callopora (?) sp..
Bryopastor challengeri , Bryopastor octogonos.
Station CP 260. 17.04.87. 21°00,00'S, 166°58.34'E, 1820-1880 m : Columnella magna var. armata.
Station CP 265. — 18.04.87. 21°04,09’S, I67°00.40’E. 1760-1870 m : Formosocellaria magnifica.
Station KG 267. — 18.04.87. 21°02.20’S, 168°58,76’E, 1935 m : Cellaria tenuirostris.
Station CP 272. 20.04.87, 21'00,04'S, 166°56,94' E, 1615-1710 m : Formosocellaria magnifica
Station KG 275. — 20.04.87. 21°05,80’S, 166°53,14'E, 1959 m : Nellia tenella.
Station CP 290. — 27.04.87, 20°36,91’S. 167°03.34’E, 920-760 m : Ccllariidae indetcrmine.
Station DW 296. — 28.04.87. 20°38,35’S. I67°10,32'E. 1230-1270 m : Pseudothyracella candelaber.
Station CP 297. — 28.04.87, 20°38,64'S, 167°10,77'E, 1230-1240 m : Columnella vipera , Pseudothyracella
candelaber.
Station DW 307. 1.05.87, 2035,38'S, 166°55,25’E. 470-480 m : Ccllariidae indeterminable, Quadricellaria
bocki. Bry opastor challengeri. Bryopastor pentagonus, Bryopastor sp., Pseudothyracella candelaber
Station DW 313. - 2.05.87. 20°58,95'S. 166°59.04'E. 1640-1600 m : Columnella vipera (?). Bryopastor
challengeri.
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
173
Station KG 316. — 2.05.87, 20°48,33'S, 166°53,29'E. 1660 m : Euginoma conica.
Smib 3
Station DW 22. — 24.05.87, 22°58,85' S, 167°19,10'E, 503 m : Mesostomaria strictoramae , Mesostomaria sp.
Calsub
Station PL 09. — 27.02.89, 20°53’S, 167°03’E, 256 m : Alderina tuberosa.
Smib 4
Station DW 35. — 7.03.89, 24°54,4'S, 168°21,6'E, 520-525 m : Syringotrema calobi.
Station DW 37. — 7.03.89, 24°53,9'S, 168°22,3'E, 500-530 m : Calloporidae indeterminable, Lamourouxia
canal icu lata.
Station DW 38. — 7.03.89, 24°54,5'S, 168°22,0'E, 510 m : Calloporidae indeterminable.
Station DW 39. — 7.03.89, 24°56,2’S, 168°2I,5'E, 525-560 m : Crateropora stilifonnis, Lcimourouxia
canaliculata.
Station DW 55. — 9.03.89, 23°21,4'S, 168°04,5'E, 215-260 m : Pseudothyracella candelaber.
Station DW 60. — 9.03.89, 23°00,1'S, 167°21,6,E, 500-535 m : Quadricellaria bocki
lies Loyaute
Musorstom 6
Station DW 396. — 13.02.89, 20°48,05’S, 16°00,59,E, 1400 m : Colunmella vipera.
Station DW 421. — 16.09.89, 20°26,27'S. 166°40, 17'E, 245 m : Pseudothyracella candelaber, Crassimarginatella
spathulata.
Station KG 465. — 21.02.89, 2I°04'S, 167°32'E, 480 m : Cryptostomaria cdata.
ETUDE SYSTEMATIQUE
EMBRANCHEMENT BRYOZOA Ehrenberg, 1831
Classe EUR YSTOMATODA Marcus, 1938
Sous-Classe CHEILOSTOMONA Busk, 1852
Ordre EUCHEILOSTOMIDA d’Hondt, 1985
Sous-Ordre NEOCHEILOSTOMINA d’Hondt. 1985
Infra-Ordre CELLULARIOMORPHA Smitt, 1867
Superfamille BUGULOIDEA Gray, 1848
Famille BUGULIDAE Gray, 1848
Genre HIMANTOZOUM Harmer. 1923
Espece-Type. — Bugula mirabilis Busk, 1881.
Himantozoum crassiavicularium sp. nov.
Fig. 3 B-C, 20 A-D
Materiel EXAMINE. — Nouvelle-Caledonie. Musorstom 4 : stn DW 223. 545-560 m.
Type. — Holotype : Nouvelle-Caledonie. Musorstom 4. stn DW 223, 545-560 m (MNHN-BRY- 16404).
Diagnose. — Himantozoum dont chaque autozoecie marginale du limbe zoarial est renflee distalement de
fa^on a former une massue saillante vers l’exterieur. Cadre lateral des autozoecies marginales et centrales presentant
de chaque cote, et sur loute sa longueur, une demi-douzaine d'epines. Aviculaires des autozoecies marginales
Source :
174
J.-L. D'HONDT & D. P GORDON
(lateraux) tronconiques, des autozotcies centrales (axiaux) ovalaires. Autozotcies marginales portant unc longue
epine distale. Extremite distale interne des autozoecies marginales arrondic ct decalee proximalement.
DESCRIPTION. — Le zoarium est dresst et pluristrit, ramifit dichotomiqucment ; les branches sont triseriees
a leur base, et peuvent comporter distalement jusqu'a 7 series aulozoeciales. Les autozoecies marginales different
de celles de la region centrale du limbe zoarial. De chaque cote de ce dernier, les loges dc la scric marginale
presentent une epine laterale externe, prolongeant Tangle distal, incurvee vers 1'axc zoarial. et mesurant 0,55 mm
de long. Elies sont distalement renflees, de fa$on a former une massue saillante vers l’exterieur ; elles mesurent
0,80 mm de long, pour une largeur de 0,30 mm distalement et 150 proximalement ; elles portent proximalement
un aviculaire tronconique frontal de 0, 18 mm de long ; leur cadre autozotcial longitudinal porte habituellcmcnt
de chaque cote, sur toute sa longueur, de 5 a 7 tpines irregulieres, mais parfois moins du cote externe. Leur region
distale interne est arrondie et inerme ; elle n'est pas siluee au meme niveau que Tangle externe, mais un peu decalee
en direction proximale. Les autozoecies de la partie centrale ont une longueur de 0,60 mm et une largeur constante
de 0.20 mm ; elles portent symetriquement, de chaque cote, une epine distale oblique et actrte, plus robuste en
presence d'ovicclle ; de 5 a 7 epines Iaterales sont portees par chacun des bords longitudinaux du cadre autozoecial,
et distributes sur toute la longueur de celui-ci ; elles sont orientees irregulierement vers l'axe autozoecial.
Les autozoecies axiales portent un aviculaire proximal median long de 0,20 mm. de forme ovoi'dc regulicrc,
parfois legerement deprime au centre de sa partie distale, isodiametrique ou generalement un peu plus large
proximalement (0,18 mm) que distalement (0,16 mm). L'ovicelle n'est portee quc par unc zoecie axiale ; en forme
de casque tres peu saillant, elle ne deborde que d’unc centaine de microns en avant de Tautozoecie reproductrice : la
partie distale de celle-ci, normalement arrondie, acquiert alors une forme plus allongee et anguleuse.
Discussion. — Cette espece prtsente sur les autozoecies de la partie centrale du limbe des epines Iaterales
pointues, distributes sur toute la longueur du cadre autozotcial ; elle partage ce caractere avec Himantozoum
(Himantozouni) dissimile d'Hondt & Gordon. 1996, Himantozoum (Thaminozoum) hispidum d'Hondt & Gordon.
1996, et Himantozoum (Beanodendria) elegans d’Hondt & Gordon, 1996. trois especcs qui apparticnncnl comme
elle a la faune nto-caltdonienne. La nouvelle espece dtcrite ici n'appartient pas au sous-genre Beanodendria d'Hondt
& Gordon, 1996. car les processus spiniformes distaux ne portent pas d'aviculaires. Elle se difftrencie du sous-
genre Thaminozoum d'Hondt & Gordon. 1996, par le fait que le cadre des autozotcies axiales n'y est spinigerc que
dans sa moitit proximale, que les tpines lattrales des autozotcies marginales n'y sont porttes que par le seul bord
interne du cadre autozotcial, et par la forme difftrente des aviculaires des autozotcies axiales. Elle se distingue de
H. dissimile par le fait que le cot t externe du cadre de ses autozotcies marginales porte des tpines au meme litre
que le cott interne (les tpines externes ttant absentes chez H. dissimile), et quc les aviculaires des loges marginales
de H. dissimile sont de forme dissymttrique. tandis que ceux de H. crassiavicularium sont symttriques. Les
aviculaires lattraux de H. crassiavicularium sont identiques a ceux de Dendrobeania pseudexilis d'Hondt & Gordon,
1996, espece appartenant a un genre tres voisin. et dont le cadre autozotcial est inerme lattralemcnt.
Lclargissement distal des autozotcies marginales et Teffacement du bord distal interne de ces memes autozotcies
caracitrisenl en outre H. crassiavicularium.
ETYMOLOGIE. — Du Latin, crassus , tpais, en raison de 1'tpaisseur de Taviculaire.
Repartition. — Nouvelle-Caltdonie. Profondeur : 545-560 m.
Infra-Ordre PSEUDOMALACOSTEGOMORPH A d'Hondt, 1977
Supcrfamille CALLOPOROIDEA Norman, 1903
Famille CALLOPORIDAE Norman. 1903
Genre CRASSIMARGI NATELLA Canu. 1900
ESPECE-TYPE. — Membranipora crassimarginata Hincks, 1880.
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
175
Crassimarginatella spathulata Gordon, 1984
C rass i ma rg incite II a spathulata Gordon, 1984 : 29, pi. 3 fig. B.
Materiel EXAMINE. — lies Loyaute. Musorstom 6 : stn DW 421, 245 m.
Description. Le zoarium est encroutant. Les dimensions des autozoecies varient considerablement, presque
du simple au double : leur longueur de 0.43 a 0,85 mm, leur largeur de 0,41 a 0,52 mm. mais la plupart d’cntre
elles onl une longueur comprise entre 0.61 et 0.78 mm. L’opesie, de forme plus ou moins ovale, a bord proximal
le plus souvenl droit, a 0,62-0,78 mm de long et 0,24-0,42 mm de large. Le cryplocyste est etroit et granuleux. II
n existc pas d epines. Lovicelle arrondie a 0,24 mm de haut et 0,40 mm de large ; la presque total ite de sa surface
est occupee par une fenetre arrondie distalement, droite proximalement. Un seul aviculaire a ete observe ; long de
0,92 mm, il selargit progressivement vers son extremite distale, arrondie et legerement elargie en spatule ; la
mandibule a une largeur de 0,31 mm a sa base et de 0,12 mm a son extremite.
Repartition, — lies Kermadec, Ties Loyaute. Profondeur : 245-350 m.
Remarque. Cet echantillon n’a pu etre photographic, etant encroutant sur un galct dont il ne pouvait etre
detache.
Genr c ALDERINA Norman. 1903
Espece-Type. — Membranipora imbellis Hincks, 1860.
Alderina tuberosa (Canu & Bassler. 1929)
Membraniporidra tuberosa Canu & Bassler, 1929 : 107.
Alderina tuberosa - GORDON, 1984 : 31. pi. 4 fig. E-F. — D'HONDT. 1986 : 701.
MATERIEL examine. — Nouvelle-Caledonie. Calsub : stn PL 09, 256 m.
Description. — La longueur autozoeciale varie de 0,44 a 0,60 mm, et la largeur de 0,35 a 0, 51 mm.
L'opcsie ovale, plus etroitc distalement quc proximalement. mesure de 0,28 a 0,35 mm de long et de 0.23 a
0,32 mm de largeur maximale. Le cryplocyste est finement granuleux : il porte proximalement, de chaque cote, un
lubercule ovale, deprime a sa partie supcrieure, long de 0,06 a 0,09 mm. II n’a 6i6 observe ni ovicelle, ni
aviculaire.
DISTRIBUTION. — Philippines, Australie, Nouvelle-Zelande. Nouvelle-Caledonie. Profondeur : 10-350 m.
Genre CRANOSINA Canu & Bassler. 1933
ESPECE-TYPE. — Membranipora coronata Hincks, 1881.
Cranosina coronata (Hincks, 1881)
Membranipora coronata Hincks, 1881 : 147. pi. 10 fig. 1.
Setosellina coronata - H.ARMER, 1926 : 265-266, pi. 16 fig. 2-4.
Cranosina coronata - Winston & Heimberg, 1986 : 6, fig. 3-6. — Hayward. 1988 : 281-282. — CHIMONIDES & Cook,
1994 : 44-45.
Source :
176
J.-L. D’HONDT & D. P. GORDON
MATERIEL EXAMINE. — Nouvelle-Caledonie. Biocal, stn DW 65. 245-275 m.
Description. — Lc zoarium encroutant esi constitue d'autozoecies de formes tres variables : losangiques,
ovoides, hexagonales, fusiformes. Elies mesurcnt de 0.64 a 0,82 mm de long, 0,44 a 0,78 mm de large ; les
dimensions de l'opesie varient considerablement, de 0,51 x 0.69 mm a 0,29 x 0,39 mm ; el le peut etre circulate,
ovale, parfois a tendance triangulaire et plus ctroite proximalement quc distalement. II n'a pas etc observe
d'ovicelles. Le gymnocyste periapertural est reduit. L'aviculaire, long de 0. 11 a 0,13 mm. large de 0,18 a
0,22 mm, presente une mandibule setiforme a bord dcnticulc, orientee obliquement vers l'arriere et incurvee vers
l'axe zoarial. longue de 0,39-0,43 mm ; sa surface est granuleuse.
Remarque. — L’unique echantillon etudie n'a pas ete photographic*. ayant etc endommage en cours d 'elude ;
nous renvoyons done a I'iconographie publie par Harmer d'une part et Winston et Heimberg d'autre part, dans
les travaux cites en reference. L'espece est nouvelle pour la faunc neo-ealedonienne.
Repartition. — Espece largement distribute dans les eaux tropicales, et notamment de I'lndo-Pacifique : lies
Loyaute. Nouvelle-Caledonie, Australie, detroit de Torres, une grande partie de I'lndonesie, tie Maurice. Ceylan.
mcr de Chine. Venezuela. Profondeur : 3-275 m.
Genre PARANTROPORA Tilbrook. 1998
Espece-Type . — Parantropora penelope Tilbrook, 1998.
Parantropora laguncula (Canu & Bassler, 1929)
Fig. 3 E
Antropora marginella - HaRMER, 1926 ( pars ) : 234. pi. 14 fig. 15 {non Hincks, 1884).
Membrendoecium lagunculum Canu & Bassler, 1929 : 96. pi. 6 fig. 6-11.
Antropora lagunculum - Mawatari & Maw ATARI, 1981 : 33.
Materiel EXAMINE. — Philippines. Musorstom 3 : stn DR 1 17. 92-97 m.
Nouvelle-Caledonie. Musorstom 4 : stn DW 150. 110 m.
Remarque. — Tilbrook (sous presse) a montre que cctte espece est presenle du Japon aux Philippines et en
Indonesie. Parantropora se distingue d' Antropora Norman. 1903, par la possession de petits septules parietaux a la
place des "pore-chambers" basaux, et la presence d'aviculaires vicariants spatules et de grandcs dimensions, dont la
longueur depasse celle des autozoecies.
REPARTITION. — Japon. Philippines, Indonesie, detroit de Torres. Nouvelle-Caledonie. Profondeur : 0-1 10 m.
Genre CONCERTINA Gordon, 1986
ESPECE-TYPE. — Concertina cultrata Gordon, 1986.
Concertina cultrata Gordon. 1986
Concertina cultrata Gordon, 1986 : 27-28, fig. 1 1. pi. 2 fig. G-H.
MATERIEL EXAMINE. — Nouvelle-Caledonie. Biocal : stn DW 46, 570-610 m.
Description. — Le materiel consiste en un debris dechire de limbe bistratifie. rigide, sans ovicelle, ni
aviculaire, ni les coenozoecies caracteristiques. Les autozoecies, pentagonales ou hexagonales, a angles bien
Source : MNHN. Pahs
ENTOPROCTES ET BRYOZO AIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
177
marques, ont unc longueur dc 0,88 a 1,15 mm et une largeur dc 0,40 a 0,51 mm ; 1'opercule mesure 0.1 1 mm de
long et 0,21 mm de large.
REMARQUE. En dcpit du mauvais etat de I'echantil Ion qui nc justifiait pas son illustration, 1’un d’entre nous
(D.P.G.) 1'a identilic a une especc auparavant decrite par lui-meme, C. cultrata , de Nouvclle-Zelande et du plateau
du "Challenger", par 914-1386 m de profondeur. Ce genre est provisoirement classe dans la famille Calloporidae
sur une suggestion de Miss Patricia L. COOK (in litt.).
Genre CALLOPORA Gray, 1848
Espece-Type. — Flustra lineata Linne, 1767.
Callopora (?) sp.
Fig. 3 D. 8 H-l
Materiel EXAMINE. — Nouvelle-Caledonie. BiogLocal : stn DW 253. 310-315 m.
Description. — La petite colonie encrofltante observee nc comporte qu'une seule ovicelle, en mauvais etat de
conservation, et aucun aviculaire, ce qui empeche toute determination specifique. Les autozoecies mesurent de 0,58
a 0,64 mm dc long et de 0,51 a 0,77 mm de large. L’opesie ovale, entouree par un bourrelet granulcux ires
saillant, a 0,42-0,59 mm de long et 0,30-0,34 mm dc large. La presence de diddles bien visibles et des bases
d'une huitaine d'epines periopesiales, toutcs brisees, nous incite a classer cette espece dans lc genre Callopora.
CALLOPORIDAE incertae sedis
Fig. 20 F-G
MATERIEL EXAMINE. — Nouvelle-Caledonie : Biocal : stn CP 67. 500-510 m.
Description. — Le zoarium est encroutant, de mono- a triserie, constitue d'autozoecies regulierement
alternantes. II n'a pas etc observe d'aviculaires ni d'epines. Les autozoecies sont fusiformes, plus larges dans Icur
moitic distalc que dans leur moilic proximale ; dies portent dc chaquc cote une diddle circulaire. L'unique ovicelle
observee, hcmiglobuleuse, est moins saillantc que le bord distal de l'autozoecie qui la porte ; die presente une area
en forme de croissant bordant l'extremite distale de I'opesie . La longueur autozoeciale varic de 0.74 a 0,83 mm, la
largeur etant de 0,30-0,36 mm a l'avanl, dc 0,21 a 0.24 mm a 1'arriere. L'opcsie ovale a une longueur de 0,32-
0,36 mm et une largeur de 0,20 a 0,22 mm ; l’orifice, en forme dc D, a 0,10 mm de long et 0,14 mm de large. Le
cryptocyste est horde par un cadre zoecial eleve.
Remarque. — Les caracteres diagnostiques sont insuffisants pour pcrmettre une identification, meme au
niveau generique.
CALLOPORIDAE incertae sedis
Fig. 5 F
Materiel examine. -Nouvelle-Caledonie. Biocal : stn DW 44. 440-450 m.
Smib 4 : stn DW 37, 500-530 m. — Stn DW 38, 510 m.
Description. — Le zoarium, depourvu d'ovicelles, d'aviculaires et d'epines, est encroutant sur des tiges
d'Hydraircs ou dcs debris de coquilles. Les autozoecies dcs colonies de la campagne Smib 4 sont souvent plus
etroites proximalement, contrairement a celles de la campagne Biocal ; dies ont unc longueur variant de 0.51 a
0,80 mm, une largeur de 0,24 a 0,38 mm du cote distal ( de 0,21-0,23 du cote proximal), et presenlcnt une opesie
Source :
178
J.-L. D'HONDT & D P. GORDON
longue, de 0,24-0,31 mm de large et de 0,24-0.38 mm de long. Le cryptocyste, profond et dclimitc par un cadre
saillant, est surtout developpe proximalement a l’opesie dont il entoure la region proximate en affectant la forme
d’un croissant ; du cote proximal, il mcsure de 0,08 a 0,14 mm de long.
Remarque. — Comme la precedente, cette espece est indeterminable par insuffisance de caractcres
discriminatifs.
Genre LAMOUROUXIA nov.
Espece-Type. — Lamouronxia canaliculata sp. nov.
DIAGNOSE. — Calloporidae dressee h zoarium formant un limbe monostratifie irregulierement decoupe,
directement fixe au substrat. Paroi zoeciale laterale tres epaisse dans sa region anterieure, et renfermant une
succession lineaire de lacunes alveolaires reliees par des canalicules capillaires. Ovicelle close par l'opercule
autozoecial.
ETYMOLOGIE. — Ce genre est cree en hommage a la memoire du bryozoologue fran9ais J.V.F. LAMOUROUX.
Lamourouxia canaliculata sp. nov.
Fig. 1 A-D, 3 A, 8 A-F, 15 D-E, 16 C-D. 19 A
MATERIEL EXAMINE. — Nouvelle-Caledonie. Biocal : stn DW 66, 515-505 m. — Stn CP 67, 500-510 m. —
Stn KG 71. 2099 m.
Smib 4 : stn DW 37, 500-530 m. — Stn DW 39, 525-560 m.
Types. — Holotype : Nouvelle-Caledonie. BIOCAL, stn DW 66, 505-515 m (MNHN-BRY-16694).
Paratype : Ibidem (MNHN-BRY- 16693).
DESCRIPTION. — Le zoarium, dresse, constitue un limbe de contours irreguliers, tres rarement de cotes presque
paralleles, affectant la forme generale d'un thalle d'algue ; il atteint 2 cm de haut. Unilaminaire, il est
habituellement tri- a quadriserie, mais comportant souvent 5 series autozoeciales h l'approche dune ramification.
Une ramification est gdneralement bi- ou plus rarement triseriee a sa base ; frequemment, elle s'elargit ensuite plus
ou moins rapidement. mais elle peut parfois demeurer biseri£e sur 2 a 3 mm avant de se ramifier et que les series
autozoeciales ne divergent. Ce limbe est fixd au substrat (des Spongiaires), sans 1’intermediaire de rhizoYdes ou d'un
pedoncule, et non par sa base, mais directement par 1' une de ses faces laterales. Celle-ci s'etend alors lateralement
sur le support de fa$on a y former une breve sole encroutante ; le zoarium s'etend largement. aussi bien vers le haut
que vers le bas, de part et d'autre de ce point lateral d'insertion.
Les zoecies marginales du limbe sont soit triangulaires. soit tres generalement quadrangulaires ; dans ce dernier
cas, le cote externe est rectiligne ; les deux cotes internes convergent vers I'axe de la colonie, contribuant a faire
largement deborder lateralement vers 1'exterieur la partie distale de la loge ; le cote proximal delimite l'insertion de
la zoecie sur la region distale de l'autozoecie marginale precedente. Le cote externe mesure de 0,50 a 0.70 mm ;
celui qui sert de base d'insertion sur la zoecie proximale mesure de 0,10 a 0,16 mm ; les deux autres cotes sont de
longueur tres variable. L’autozoecie mesure de 0,67 a 0,70 mm de long et 0,38-0,40 mm de large.
Les autozoecies de la region centrale du limbe sont de forme plus reguliere et symetrique que les zoecies
marginales : linguiformes ou plus ou moins losangiques (dans ce cas a region distale arrondie) et a partie
proximale anguleuse, s'inserant sur une breve longueur (0,20 mm au maximum) a la portion distale de I'autozodcic
precddente. Leur largeur et leur longueur, tres constantes, varient respectivement de 0,39 a 0,42 mm et autour de
0,70 mm ; quclques tres rares autozoecies peuvent toutefois depasser cette taille, et alteindre alors une longueur de
0,90 mm. L'orifice est tout a fait distal, ovale, petit, large de 0,12-0,13 mm et long de 0.085-0,1 15 mm.
L’opercule est de couleur brune ; il mesure 0,26 mm de large et 0,15 mm de long ; son sclerite marginal est tres
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
179
marque. Le nombre de tcntacules, non etabli exactement, est de 1'ordre de la quinzaine. Le bord du tres etroit
gymnocyste est ome de nombreuses petites crenulations tres rapprochees et de meme taille.
La frontale autozoeciale est
membraneuse. La face basalc presente de
chaque cote une fenetre non calcifiee
bien visible, proximale a rorifice, situee
dans I'alignement d'une demi-douzaine de
tres petites perforations alignees se
succedant a faible distance des parois
interzoeciales. Ces fenetres sont ovales
et mesurent 0,60 mm de long et 0,35-
0,40 mm de large.
L'incubation s'effectue dans une
ovicelle situee a la partie distale de
1'autozoecie reproductrice qu'elle recou-
vre ; 1'autozoecie ovicellee mesure alors
0,98 mm de long. L'ovicelle est situee
dans le plan du zoarium. n'etant qu'a
peine saillante en surface ; elle n'est
done pas hyperstomiale, mais constitue
une expansion de la paroi distale de la
zoecie en reproduction ; elle porte
parfois a son extremite distale un court
aviculaire triangulaire d'orientation
transversale. De forme triangulaire, elle
mesure 0,34 mm de long et 0.37 mm de
largeur a l'avant des opesiules. La cavite
interne de l'ovicelle, en forme de cloche,
longue de 0,25 mm et large de
0,30 mm, s'elargit en arc de cercle de
0,36 mm a sa partie proximale, el
renferme dans sa concavite l'opercule (non recouvert par l'ovicelle). La partie distale de 1'autozoecie, au-dela de la
cavite d'incubation, forme une languette a extremite arrondie, partiellement occupee par une cavite alveolaire ovale,
plus grande que celles decrites ci-apres, puisque mesurant 0.06 mm de long et 0,10 mm de large.
Les parois autozoeciales des deux cotes distaux, l'interne commc l'externe, sont tout particulieremenl epaisses.
Elies renferment, de chaque cote, 5-6 cavites alveolaires successives allongees et uniseriees, et une cavite distale
impaire, communiquant entre elles, chacune avec cel le qui la precede et celle qui la suit, par des conduits
capillaires, et qui ont probablement pour fonction d'alleger la colonie ; e'est en raison de l'exislence de ce systeme
de communications capillaires que nous ne pouvons assimiler ces cavites a des dietelles. Ces cavites ont un
diametre constant de 0,06 mm et une longueur variant de 0,05 a 0.14 mm ; elles entourent aussi 1'orifice
autozoecial a la fa$on d'un collier. Limitees a la partie distale de 1'autozoecie, elles sont absentes des parois de la
region proximale, beaucoup plus fines. Les parties distales des autozoecies, celles dont les parois sont lacunaires,
recouvrent les parties les plus proximales des autozoecies precedentes. La membrane frontale recouvre directement
les cavites alveolaires ; en revanche, le conduit capillaire qui assure la communication entre deux cavites
successives traverse la cloison calcaire qui les separe.
DISCUSSION. — Cette espece, a frontale membraneuse, est un Bryozoaire du type pseudomalacostege. Elle ne
presente toutefois ni gymnocyste proximal, ni cryptocyste, ce qui la difference de la plupart des especes
actuellement rangees dans l'heterog£ne famille Calloporidae chez laquelle l'un ou l'autre au moins de ces caracteres
est present (cf. la diagnose reactualisee qu'en propose Gordon, 1984), et dont certains genres ( Ellisina .
Fig. 1. — Lcmiourouxia canaliculata : A. Quelques autozoecies observees
par transparence ; B-C. Quelques autozoecies en vue venlrale
(schematique, indiquant I'emplacement des fenetres) ; D, Detail des
cavites alveolaires se succedant a l'imerieur de la paroi. Echelle :
0,10 mm.
Source :
180
J.-L. D'HONDT & D. P. GORDON
Retevirgula) possedent comme elles des aviculaires associes aux ovicelles. Mais le genre Lamourouxia se
caracterise par ailleurs par l'existence des lacunes alveolaircs parietales communiquant par des canaux capillaires.
L'absence des "occlusor laminae" ecarte ce taxon des Chaperiidae. celle du processus epineux et de connections
interzoeciales des Hiantoporidae. les ovicelles sub-immergees des Flustridae et des Hincksinidae, les morphologies
zoariale et zoeciale des Farciminariidae ; le manque de dimorphisme zoecial et de cryptocyste les eloigne des
Bryopastoridae ; l'absence d'onychocellaires, de vibraculaires et de lout dispositif dissimulant la membrane frontale
l'ecarte des autres families de Pseudomalacosteges.
Les deux genres de Calloporidae, aux gymnocystes et cryptocystes pcu developpes, qui presentent des analogies
avec cette espece s'en ecartent par plusieurs caracteres diagnostiques. Les Retevirgula n'ont pas de dietelles, souvent
des tubes anastomotiques interzodciaux, des coenozoecies ou des aviculaires interzoeciaux. Les Ellisina en
paraissent plus proches, mais leur zoarium est encroutant, le gymnocyste et lc cryptocyste presents, les aviculaires
interzoeciaux ; elles presentent des dietelles. Ni l’un ni I'autre ne prescntent le mode dc fixation au substrat decrit
ici, ni les cavites parietales specifiqucs.
Les Calloporidae constituent une famillc assez difficile a definir et a delimiter, qui nccessite une revision
d'ensemble ; selon P.L. COOK (in litt.), les genres Cranosina , Ellisina et Lamourouxia en particulier constituent
un ensemble dont le statut est a definir, mais qui pourrait constituer une nouvelle famille. Les canaux dc la paroi
des Lamourouxia seraient, selon cet auteur (in litt. ), analogues au "basal channel network" (mais ou il n'a pas etc
observe de canaux capillaires), observe et figure par CHIMONIDES et COOK (1994) chez l'espece Cranosina spiralis
decrite dans leur publication, mais en fait il n’est pas prouve qu’il s'agisse reellement de structures homologues ;
toutefois, il faut remarquer que Cranosina spiralis n'est egalement que tres partiellement adhercnte au substrat. s’en
liberant en grande partie. Nous pouvons aussi relever la presence d'une chambre distale, qui semble associee a celle
d'une cavite avicularienne subrostrale chez ces memes trois genres. Nous sommes en disaccord avec la proposition
de CHIMONIDES et COOK de classer le genre Cranosina dans la famille Hincksinidae, celle-ci ayant un zoarium
encroutant, flustride.
Etymologie. — Du Latin, canaliculatus , avec des petits canaux, en raison de l'existence de canaux capillaires
dans l’epaisseur de la paroi.
Repartition. — Nouvellc-Calcdonie, de 500 a 2099 m.
Famille INCERTAE SEDJS (P.L. Cook, in litt.)
? Genre PSEUDOLUNULARIA Cadee, Chimonides & Cook. 1989
Espece-Type. — Pseudolunularia enigma Cadee, Chimonides & Cook, 1989.
? Pseudolunularia sp.
Materiel EXAMINE. — Nouvelle-Caledonie. Biog£ocal : sin DW 253. 310-315 m.
DESCRIPTION. — L'unique petit zoarium recueilli (4 mm de diamctre) est aplati, discoi'de et libre, forme
d'autozoecies (linguifonnes dans la partie centrale de la colonic, ovalaires et plus larges dans leur partie mediane
k la peripherie de la colonie) de taille croissante vers rexterieur, passant de 0,50 a 0,76 mm en longueur et dc 0,24
a 0,40 mm en largeur ; le bord distal des autozoecies est arrondi. L'opesie est ovale, sans opesiules, et mesure dc
0,23 a 0,39 mm de long et de 0,18 a 0,22 mm de large. En peripherie de la colonie, quelques zoecies vibraculaires
triangulaires et effilees du cote proximal, longues de 0,80 a 1,15 mm ct larges de 0,21-0,23 mm sont inserees
entre les autozoecies. Elles portent une mandibule (fouet vibraculaire) longue de 0, 14 a 2,6 mm, dont la partie
libre du rachis mesure de 1,48 a 1,60 mm ; le rachis est rectiligne ; la mandibule ne porte ni denticules ni ailes. Il
n’a pas ete observe d'ovicelles. La face inferieure des autozoecies est generalemcnt imperforee, seules les plus
peripheriques d'entre elles presentent une ou deux perforations.
Source : MNHN. Pahs
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOM IDA DE NOUVELLE-CALEDONIE
181
REMARQUE. — Si notre identification est exacte compte tenu de I'etat de l'echantillon, ce genre decrit de l'lndo-
Pacifique oriental etait encore inconnu de la faune nco-caledonienne. La colonic etudiee ici etait trop jeune pour etre
determinable, et partiellement endommagee ; aussi n'a-t'elle pas ete figuree et aucune identification specifique n'cn
sera proposee.
Famille QUADRICELLARIIDAE Gordon, 1984
Genre QU AD RICE LL ARIA d’Orbigny, 1851
Espece-Type. — Quadricellaria elegans d'Orbigny, 1851 .
Quadricellaria bocki (Silen, 1941)
Fig. 3 F
Acanthodesia bocki Silen, 1941 : 20-22, fig. 15-16.
Quadricellaria bocki - GORDON, 1984 : 24-25, pi. 1 fig. B. — D'HONDT, 1986 : 71 1.
Nelliella bocki - Mawatari, 1974 : 37-40, fig. 6.
Materiel EXAMINE. — Nouvelle-Caledonie. Biocal : stn DW 46, 570-610 m. — Stn CP 52, 540-600 m. —
Stn CP 75, 825-860 m.
Biog£ocal : stn DW 307, 470-480 m.
Smib 4 : stn DW 60. 500-535 m.
Description. — Lc zoarium, dresse et ramifie dichotomiquement, est forme de longs entre-noeuds quadriseries,
ou les autozoecies sont opposdes deux a deux, separes par des joints chitineux. Chaque entre-nocud portc sur
chacune de ses faces de 4 a 6 autozoecies rectangulaires a cotes presque paralleles (un peu plus etroites
proximalement), alternantes d'une face a 1'autre, mesurant de 0,44 a 0,76 mm de long et de 0,155 a 0,190 mm de
large. Les opesies, elles-memes de contours rectangulaires mais dont les angles proximaux sont en fait arrondis,
mesurent de 0,22 & 0,34 mm de long et de 0,14 a 0,18 mm de large. II n'a pas etc observe d'aviculaires, ni
d'ovicelles (celles-ci n'ont actuellement ete observees que sur des specimens de Nouvelle-Caledonie decrits par
D’HONDT, 1986).
Repartition. — Japon, Nouvelle-Caledonie, Kcrmadec. Profondcur : 19-860 m.
Genre NELLI A Busk. 1852
Espece-Type. — Cellaria tenella Lamarck, 1816.
Nellia tenella (Lamarck, 1816)
Fig. 3 G
Nellia tenella Lamarck. 1816 : 135. — d’Hondt. 1979 : 17.
Nellia oculata Busk, 1852 : 18, pi. 64 fig. 6. pi. 65 bis fig. 4. — Harmer, 1926 : 240-245, fig. 3-4. pi. 14 fig. 18-20. —
Osburn, 1950 : 1 19-120. pi. 13 fig. 4. — Cook, 1968 : 156-157. — Winston & Cheetham, 1984 : 257-265.
MATERIEL EXAMINE. — Nouvelle-Caledonie. Musorstom 4 : stn DW 187, 65-120 m.
Biog£ocal : stn KG 275, 1959 m.
DESCRIPTION. — Le zoarium arborescent est ramifie dichotomiquement et quadriserie ; il est constitue d’entre-
noeuds quadriseries separes par des joints chitineux cylindriques. Les autozoecies, rectangulaires, alternent d'une
Source :
182
J.-L D'HONDT & D. P. GORDON
serie longitudinalc a l'autre et sont opposees deux a deux. II n'existe pas d’epines. La surface frontale est occupce
sur les 3/4 de sa longueur par une opesie a cotes presque paralleles et a extremites arrondies. II existe une paire de
minuscules aviculaires pairs proximaux a lopesie. La longueur autozoeciale a mi-longueur des entre-noeuds est de
0.48-0,53 mm, la largeur de 0,18-0,21 mm.
REPARTITION. — Une grande partie des mers chaudes du globe : toute ITndonesie, Australie, Nouvelle-
Caledonie, Ccylan, cotes allantique et pacifique des U.S.A., Antilles, cote ouest africaine. de 0 a 150 m. La recolte
de cette espece a grande profondeur (1959 m), lors de la campagne BiOGEOCAL, est done ires surprenante, sinon
suspecte.
Famille FLUSTRIDAE Fleming, 1828
Remarque. — Famille nouvellc pour la faune neo-caledonienne.
Genre CARBASEA Gray, 1848
Espece-Type. — F lustra carbasea Ellis & Solander, 1786.
Carbasea aff. linguiformis Harmer, 1926
Carbasea linguiformis Harmer, 1926 : 249-250, fig. 7. pi. 15 fig 5.
MATERIEL EXAMINE. — Philippines. Musorstom 3 : sin CP 101. 195 m.
DESCRIPTION. — Le materiel consiste en quelques fragments de zoarium un istrati fie, formes de 5
a 7 autozoecies a cotes presque paralleles, seulement un peu plus elargies selon les cas, soil distalement, soit a
mi-longueur ; ces autozoecies sont tres epaisses, ce qui par un effet trompeur de perspective donne faussement
I’impression que le zoarium est horde par des coenozoecies laterales. Le cryptocyste est etroit, et 1’orifice distal est
subterminal. La region proximale des autozoecies est symetrique par rapport a l'axe longitudinal et s'etend, en
forme de queue de poisson, lateralement de part et d'autre de la region distale de l'autozoecie precedentc. La longueur
autozoeciale varie de 0,75 a 1,02 mm. la largeur de 0,32 a 0,41 mm. II n'existe pas d'aviculaires ni d'epines.
L'ovicelle globuleuse, en forme de ventouse medicale, longue de 0,37-0.40 mm. a une largeur maximale, dans sa
partie ampulliforme, de 0,36-0,38 mm ; elle est endotochoidale, incluse dans la partie proximale de l'autozoecie qui
la suit distalement dans la meme file longitudinale.
DISCUSSION. — La cle de determination des genres de Flustridae publiee par D’HONDT et REDIER (1977),
modifiee par D'HONDT (1983). ne laisse aucun doute sur l'appartenance de ces echanti lions au genre Carbasea.
L'absence de zoarium complet ne permet pas de determiner avec certitude a quelle espece ils appartiennent, mais le
fait qu’ils proviennent d'une colonie laciniee conduit a les attribuer a C. linguiformis Harmer, 1926, dont les
zoecies ont la meme morphologie. C. linguiformis n'est actuellement connue que par un scul echantillon de
Nouvelle-Guinee, drague a 411 m de profondeur. La presence de cette espece aux Philippines, par 195 m de
profondeur, est done tout a fait plausible.
Carbasea laterogranulata sp. nov.
Fig. 4 A, C
MATERIEL EXAMINE. — Nouvelle-Caledonie. Biocal : stn DW 31, 850 m. — Stn DW 51, 680-700 m.
Musorstom 4 : stn DW 220, 505-550 m.
TYPE. — Holotype : Nouvelle-Caledonie, BIOCAL, stn DW 31, 850 m (MNHN-BRY- 16686).
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
183
Diagnose. — Carbasea a autozoecies de grandes dimensions (environ 1 mm dc long pour 0.6 mm de large), a
zoarium monostratifie, a parois laterales cxternes rendues rugueuses par la presence sur la totalite de leur longueur
de plusieurs centaines de petites verrues granuleuses.
DESCRIPTION. — Le zoarium lacinie est dresse et unilaminaire, non ramifie, ni lobe, sans poriion basale
cncroutantc sxir un substrat ; il mesure 8 cm de long pour 6 mm dc large ; forme de 1 a 9 series autozoeciales, il
est fixe au support par un faisceau cylindrique de rhizoi’des. Les autozoecies sont de grande taille, ovoides, arrondics
distalement, tronquees proximalement, et de dimensions tres homogenes ; leur longueur ne varie que de 1,15 a
1,20 mm, leur largeur de 0,62 a 0,68 mm. L'opercule, tout a fait distal, mesure 0,12 mm de long et 0,41 mm dc
large. Les bords lateraux externes des autozoecies marginales sont ornes d'innombrables petites verrues
granuleuses, generalement un peu plus longues quc larges, de 0,02 a 0,03 mm de longueur, donnant a cette surface
laterale une apparence de rape. Le bord distal externe des autozoecies marginales est saillant vers l'exterieur.
formant un socle sur lequel vient s'inserer la partie proximale de l'autozoecie suivante.
Discussion. — Morphologiquement, cette espece rappelle Carbasea indivisa (Busk, 1852), mais les zoariums
de cette derniere ont tendance a acquerir une morphologic cupuliforme, ce qui n'est pas le cas ici. Les autozoecies dc
C. laterogranulata sont plus grandes, bien que de meme forme. La principale caracteristique de la nouvelle espece
decrite ici reside dans l'aspect tres particulier, granulcux, de rornementation des bords lateraux externes dcs
autozoecies marginales du zoarium ; C. indivisa prdsente aussi des granulations superficielles, mais cellcs-ci sont
portees par la face dorsale des autozoecies et non par les faces laterales.
ETYMOLOGIE. — Du Latin, latus, cote, et granulata , qui a des grains, en raison de la presence de nombreuses
petites verrues portees par les bords latero-externes des zoecies marginales.
Repartition. — Nouvelle-Caledonie, de 505 a 850 m.
Famille FARCIMINAR1IDAE Busk. 1852
Genre COLUMNELLA Levinsen, 1914
Espece-Type. — Columnaria borealis Levinsen, 1909.
Columnella magna (Busk, 1884)
Fig. 4 F-G
Farciminaria magna Busk, 1884 : 49-50, pi. 5 fig. 1.
Levinsenella magna - Harmer, 1926 : 402. — Hastings, 1943 : 393-394.
Columnella magna - d'Hondt. 1975 : 563; 1981a : 13. — Hayward & Cook, 1979 ; 67. fig. 9 A. — Hayward. 1981 :
29-300. fig. 6. — Gordon, 1986 : 26, pi. 2 fig. C-D.
MATERIEL EXAMINE. — Nouvelle-Caledonie. BlOCAl. : stn CP 13, 3690-3740 m (forme typique).
Biog£ocal : stn CP 260 , 1820-1880 m (var. armata ).
Description. — Le zoarium est dresse, quadriserie, et ramifie dichotomiquement. La longueur autozoecialc
varie de 1,40 a 1,95 mm. la largeur de 0.27 a 0,49 mm. Les bords lateraux de l'autozoecie sont parallels sur la
plus grande partie de la longueur, la loge ne s'elargissant qu'au niveau de l'opesie ou parlois brievcment a mi-
longueur ; il n'existe pas d'epines. L'ovicelle globuleuse a 0,60 mm de long et 0.70 mm de large ; elle est ornee de
fines lignes d'apparence granuleuse, convergeant vers 1'orifice. Les zoecies de la station CP 13 sont toutes
inermes ; la plupart de celles de la station CP 260 presentent 1'aviculaire caracteristique de la variete armata ,
decrite et figuree par BUSK (1884), differenciant cette variete de la forme typique ; cet aviculaire partiellement
dresse, long de 0,20 mm, possede une petite opesie circulate distale de 0,80 mm dc diametre, portant une petite
mandibule hdmicirculaire de 0,50 mm de long.
Source :
184
J.-L. D’HONDT & L). P GORDON
Repartition. — Ocean Atlantique occidental et oriental, sauf regions polaires et sub-polaires : Kermadec,
bassin dc Tasmanie, banc de Bellona ; Heard ; Afrique du Sud ; cote est-africaine. Profondeur : 680-5340 m.
Remarques. — Genre et espece signales pour la premiere fois de Nouvelle-Caledonic. Harmer ( 1926) a mis
en doute la validite de la variete armata. ayant aussi trouvc des aviculaires sur 1c specimen original de la forme ty-
pique. En fait la situation est plus complexe. L'aviculaire est peu frequent chez les specimens sud-africains etudies
par Hayward et Cook (1979) et chez ceux des campagnes de la "Galathea" de provenances diverses examines par
Hayward (1981), sans que cet auteur ne precise s'il existe ou non des differences en fonction des provenances geo-
graphiques. Gordon (1986) n’cn a pas observes dans son materiel des Kermadec. Dans un memo prelevement,
D’HONDT (1975) a trouve simultanement des colonies portant des aviculaires sur pratiquement toutes les autozoe-
cies, et d'autres qui en etaient completement depourvues ; dans une autre etude (D'HONDT. 1981a). les colonies
appartenaient selon les localites soit a la forme typique, soil a la variete armata \ D'HONDT en concluait qu'il etait
impossible, dans letat actuel des connaissances, de decider si la presence ou l'absence de l’aviculaire etaient des
constantes genetiques ou des caracteres reversibles lies a renvironnement. En fail, le faisceau des observations
actuelles suggererait plutot que les deux hypotheses soient simultanement plausiblcs ; dans certaines localites, la
presence ou l'absence de l'aviculaire pourraienl etre des caracteres genetiquement fixes dans la population, alors
qu’il pourrait exister une variability indi viduelle obeissant aux lois de la genetique dans d'autres populations.
Columnella vipera sp. nov.
Fig. 4E, 5 A-B.20E
MATERIEL EXAMINE. — Nouvelle-Caledonie. Biocal : sin DW 41, 380 m. — .Sin CP 52, 540-600 m. —
Stn DW 53, 975-1005 m. — Stn CP 54, 950-1000 m. — Stn CP 55. 1 160-1 175 m. — Stn CP 60, 1530-1480 m. -
Stn CP 62, 1395-1410 m. — Stn DW 70. 960-965 m. — Stn KG 76. 880 m.
Musorstom 4 : stn CP 238. 500-510 m.
Biog£ocal : stn CP 205. 1350-1380 m. — Stn CP 297. 1230-1240 m. — Stn DW 313, 1600-1640 m (?).
lies Loyaute. — MUSORSTOM 6 : stn DW 396. 1400 m.
Types. — Holotype : Biocal. stn DW 41. 380 m (MNHN-BRY- 16581 ).
Paratypes : Biocal, stn CP 54. 950-1000 m (MNHN-BRY-16593) ; stn CP 55, 1 160-1175 m (MNHN-BRY-
16584) ; stn DW 70, 960-965 m (MNHN-BRY- 16600). — BlOGEOCAL, stn CP 205. 1350-1380 m (MNHN-
BRY- 16588). — MUSORSTOM 6, stn DW 396, 1400 m (MNHN-BRY-16595).
Diagnose. — Columnella a zoecies aviculariennes localisees a la base des ramifications zoariales, sur les
aretes internes des branches (quadriseriees) ; cet aviculaire peut etre unique et alors situe a la partie proximate de
1'aretc, ou il peut exister plusieurs aviculaires sur une meme arete. Les aviculaires, non sail lants, sont inclus dans
la partie proximale d'une autozoecie ; celte region proximale est separce par une constriction du reste de
1'autozoecie, et affecte plus ou moins nettement la forme d'une tele dc reptile du genre Vipera. Mandibule
avicularicnne incluse dans le plan de la frontale. Ovicclle occupant tout l'interieur d’une coenozoecie incluse dans le
zoarium.
Description. — Le zoarium arborescent, quadriserie et ramifie dichotomiquement. est fixe au subslrat
par un faisceau de rhizoides issus d’une region discoide de 0. 10 mm de diametre de la partie proximale de certaines
des autozoecies. A la base de la colonic, ces rhizoYdes sont soudcs de fa$on a constituer un axe rigidc et rectiligne
atteignant 3 cm de haut et 3 mm de large, et cngainant les autozoecies les plus proximales. Les autozoecies,
inermes et peu calcifiees, sont linguiformes ou presque rectangulaires et alors a grands cotes sub-paralleles, parfois
tendant vers une morphologic pisciforme ; leur bord anlerieur est arrondi, le bord posterieur legerement incurve ;
dies mesurent de 0,20 a 0,30 mm de large (pour les plus larges, a la base des branches) et de 0.80 a 1,12 mm
de long ; 1'opercule hemicirculaire a 0,28 mm de long el 0,22-0,24 mm de large. Une coenozoecie pyriforme,
a parois epaisses, separe les branches nees de chaque ramification ; sa longueur varie selon les cas de 0,50
a 1.12 mm, sa largeur distalement de 0,40 a 0,50 mm, proximalcmcnt dc 0,12 a 0,16 mm ; elle est precedee par
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOST OM I DA DE NOUVELLE-CALEDONIE
185
une minuscule autozoecie. II arrive qu'a l'approche des ramifications les autozoecies aient une forme plus
irreguliere, avec un cote externe rectiligne et un cote interne formant un angle saillant vers I’axe zoarial.
L'ovicelle est renflee par rapport au plan dc la colonie ; de contours presque carres ou ampulliformes - ellc csl
alors legerement petrecie du cote proximal (de 0,60 mm distalemenl et de 0,50 mm proximalement. par exemple) -
a bords lateraux presque parallels mais a cole distal arrondi vers lav-ant, elle est fermee par l'opcrcule autozoecial.
Elle est immergee dans une coenozoecie hexagonale dont ellc occupe pratiqucmcnt tout l'interieur, el qui est situee
distalement par rapport a l'autozoecie reproductrice ; cette coenozoecie a 0.74 mm de long et 0,55 mm de largeur
maximale. L’ovicelle elle-meme a une longueur de 0,48-0,60 mm et une largeur de 0,45-0.60 mm ; sa surface
presente une ires discrete reticulation convergeant vers l'orifice. Deux zoecies aviculariennes encadrent la panic
distale de la coenozoecie incluant l'ovicelle ; dies sont allongees, quadrangulaires et plus ou moins losangiques,
longues de 0,80 a 0,82 mm et larges de 0,20 mm dans leur portion la plus renflee ; la mandibule, comparable
a celle decrite ci-apres dans le cas des aviculaires normaux, est situee tres proximalement et orientee vers la
region distale.
Chaque aviculaire porte une mandibule hemicirculaire dirigee distalement. Non saillant. situe dans le plan
de la frontale, il est inclus dans la courte region proximale dune autozoecie. qui en apparait comme un diverticule
separe par une constriction simulant un cou ; il est situe sur l'une des deux aretes internes de chacune des deux
branches issues dune ramification ; parfois seules les autozoecies les plus proximales portent de tels aviculaires,
mais il arrive souvent que plusieurs autozoecies successives d'une meme serie longitudinale presentent
leur aviculaire sur une meme arete. Les diverticules proximaux aviculiferes ont. souvent ties nettement, la forme
d'une tete de serpent dc la famille des Viperidae ; ils mesurent de 0,50 a 0.80 mm de long et dc 0,30 a 0.36 mm
de largeur maximale, la mandibule ayant une longueur de 0. 1 1 a 0.12 mm de long et de 0,22 a 0.24 mm de large.
Les aviculaires les plus proximaux d'une meme bifurcation ne sont pas situes au meme niveau sur les deux
branches opposees. Tres exceptionnellement, ces aviculaires pcuvent manquer sur certaines aretes.
DISCUSSION. — Le mode d'insertion de la zoecie avicularienne, immergee dans un diverticule proximal d’une
autozoecie, et non saillante a 1'exterieur, est unique dans ce genre. Ce diverticule a par surcroTt une forme tres
particuliere, et la forme de la mandibule avicularienne est elle-meme inhabituelle dans ce genre. La localisation des
aviculaires sur les aretes internes des branches zoariales est, elle aussi, caracteristiquc.
ETYMOLOGIE. — Le nom specifique est evocateur de la morphologie du diverticule autozoecial aviculifere,
dont les contours rappellent ccux d’une tete de reptile ophidicn du genre Vipera.
Repartition. — Nouvelle-Caledonie et ties Loyaute, de 380 a 1640 m.
Famille BRYOPASTORIDAE nov.
Diagnose. — Zoarium drcsse, en forme de baguette et non ramifie, ou articule et ramifie, portant des rhizoides
a sa base. Axe et branches a quatre faces, cylindriques, et / ou en forme de lentillcs bilaminaires ; zoecies
alternantes. Cryptocyste zoecial bien developpe, deprime ; gymnocyste absent. Opesie occupant approxima-
tivemenl la moitie de la longueur zoccialc ; chaque angle disto-lateral est parfois creuse d'une depression menagee
pour le muscle d'occlusion. Pas d'epines orales. Aviculaires vicariants, identiques aux autozoecies mais avec
de grandes mandibules, ou absents. Incubation des embryons interne dans des autozoecies. ou a 1 interieur
de zoecies femelles tres developpees ou de structures ovicelliennes endozoeciales. Ancestrula enracinee,
rcssemblant a une autozoecie, mais de plus petites dimensions.
Nous proposons I'insertion dans cette famille, connue du Maestrichtien a lepoque actuelle, des genres
suivants : Bryopastor Gordon, 1982. Dioptropora Marsson. 1887, Escharinella d’Orbigny. 1851. Monticellcirici
Voigt. 1987, Pseudothyracella Labracherie, 1975. el Thyracella Voigt, 1930 II n’est pas a exclure que les genres
actuels Acanthodesiomorpha d'Hondt. 1981a el Cookinella d’Hondt. 1981a puissent aussi trouver place dans cette
famille ; mais comme ils sont encore incompletement connus, nous avons juge premature de les y inclurc.
186
J.-L D'HONDT & D. P. GORDON
Remarques. — Les genres inclus dans la famille Bryopastoridae sont caracterises, aussi bien par leur port
dresse et leur enracincment que par leurs zoecies depourvues d'epines orales, 1'absence d'aviculaires adventifs et
d'ovicelles hyperstomiales. Les affinites du genre encroutant Hagenowinella Canu, 1900. du Maestrichtien.
chaperiide presume (GORDON, 1982), avec les bryopastorides du mcmc Maestrichtien. sont incertaines. Les
Bryopastoridae et Hagenowinella pourraient toutes deux etre issues d'ancetres onychocellidiens. Le genre
chaperiidicn ndo-zelandais Patsyella Brown. 1948, present du Miocene Superieur a l'epoque actuelle, semble nc
presenter qu'une ressemblance superficielle avec les bryopastorides - Tune au moins des especes qu’il renferme a des
epines articulees.
Etymologie. — Famille creee a partir du genre Bryopastor preexistant, dont les colonies de 1'espece-type.
recourbees a leur partie superieurc, ont la forme d'unc canne de bcrger.
Genre BRYOPASTOR Gordon, 1982
Espece-Type. — Heterocella pentagona Canu & Bassler, 1929.
Bryopastor challengeri Gordon. 1982
Fig. 4 D
Biyopastor challengeri Gordon. 1982 : 20, fig. 9 D-E ; 1986 : 40, pi. 10 fig. B-C.
Materiel examine. — Nouvelle-Caledonie. Biog£ocal : stn KG 210, 1190 m. — Stn CP 232, 760-790 m. —
Stn DW 253, 310-315 m. — Stn DW 307. 470-480 m. — Stn DW 313, 1600-1640 m.
Description. — Le zoarium est vinculariiforme, dresse et arque ; il est quadriserie, les autozodcics alternant
par paires opposees. Les fragments etudies, effiles a leur panic proximale, ne component que des autozoecies
incrmes ; ils presentent des loges fcmelles ou a joints, comparables a cellcs decrites par GORDON (1982), mais pas
d'aviculaires. La longueur autozoeciale varie de 0,78 a 0,94 mm. la largeur de 0,42 a 0,56 mm. L'opesie a des
cotes paralleles sur une partie de sa longueur et se retrecit proximalement ; pour une longueur de 0,24 mm, elle a
une largeur proximalement de 0,70-0,80 mm et distalemenl de 0,17 mm. Les orifices de passage des muscles
occluseurs sont visibles distalement a l'orifice, un de chaque cote. Le cryptocyste, developpe proximalement et
distalement a l’opesie, a une surface granuleuse. Les occlusor laminae se presentent comme deux cretes calcaires
convergeant distalement.
Repartition. — Nouvelle-Zelande (plateau du " Challenger ", a Test du detroit de Cook). Espcce nouvelle pour
la Nouvelle-Caledonie. Profondeur : de 310 a 1640 m.
Bryopastor octogonos sp. nov.
Fig. 5 C-D
Materiel EXAMINE. — Nouvelle-Caledonie. Biocal : stn DW 44, 440-450 m (echantillon erode, identifi¬
cation incertaine).
Biog£ocal : stn DW 253, 310-315 m.
Type. — Holotype : Nouvelle-Caledonie. BiOGEOCAL, stn DW 253, 310-315 m (MNHN-BRY- 16649).
Diagnose. — Bryopastor a zoarium cylindrique constitue de 8 series longitudinales d'autozoecies. Occlusor
laminae tres rapproches, delimitant une etroite fente a bords paralleles, precedant un elargissement proximal de
forme ovale. Opesie grande (0,32-0,40 mm de long) et ncttement plus longue que large, occupant environ un tiers
de la surface frontale.
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
187
DESCRIPTION. — Le zoarium, robuste et non ramifie, de section cylindrique, est forme de 8 series alternantes
d'autozoecies, les series diametralement opposees se correspondant deux a deux. Chaque autozoecie est partagee en
deux parties sensiblement d'egale longueur, une distale portant l'opesie, unc proximate correspondant
essenticllement au cryptocyste et se retrecissant graduellement en direction proximale. La longueur autozoeciale
varie de 1,1 h 1,2 mm, la largeur de 0,55-0,60 mm distalement et de 0,32 a 0.39 mm proximalement. Les limites
interzoeciales sont saillantes. L'opesie mesure 0,32-0,40 mm de long ct 0,20-0,26 mm de large. Les gros orifices
triangulaircs des muscles occluseurs sont situes en position disto-laterale par rapport a l’orifice. tres prochcs de
celui-ci. Les occlusor laminae forment des lames presque paralleles, tres rapprochees 1'une de 1'autre, ne laissant
entre elles qu'une mince fente longitudinale ; ils se touchent proximalement, avant de s'ecarter pour mcnager entre
eux un elargissement de forme ovale, allonge, deux a trois fois plus large que la fente qui les separe distalement.
II n'a ete observe ni de zoecies femelles, ni a joints. Le cryptocyste est ornc de fines granulations espacees.
DISCUSSION. — Le nombre des series autozoeciales longitudinales distingue cette espece de la plupart des autres
du genre. B. challenged ct B. tongensis sont quadriseriees, B. pentagonus quintiseriee, exceptionnellement
hexaseriee. Les deux autres especes de Bryopastor h zoarium octoseric, B. crassus et B. elongatus, ont
rcspcctivement la premiere une disposition tres differente des occlusor laminae (voir plus loin), la seconde des
opesies proportionncllement de tres petite taille et des loges presumees femelles demesurement allongees.
ETYMOLOGIE. — Du Latin, octogonos, qui a huit angles, les branches du zoarium comportant huit angles en
section transversale.
Repartition. — Nouvelle-Caledonie, de 310 a 450 m.
Bryopastor pentagonus (Canu & Bassler, 1929)
Fig. 5 G, 6 A-B
Heterocella pentagona Canu & Bassler, 1929 : 111-112, pi. 9 fig. 13-16.
Bryopastor pentagonus - GORDON, 1982 : 18-20, fig. 9 A. F.
MATERIEL EXAMINE. — Nouvelle-Caledonie. BiOCAL : stn DW 08, 435 m. — Stn DW 38. 360 ni. —
Stn DW 46, 570-610 m. — Stn DW 66. 505-515 m.
MUSORSTOM 4 : stn CC 175, 355 m.
Biog£ocal : stn 232, 760-790 m. — Stn DW 307. 470-480 m.
DESCRIPTION. — Le zoarium arborescent est constitue de 5 series alternantes d'autozoecies longues de 0,72 a
1,00 mm et larges de 0,37 a 0,41 mm distalement, 0,27 a 0,30 mm proximalement, arrondies a leur extremite
distale (nous n'avons pas observe sur notre materiel de portions a 6 series). L’opesie, qui deborde sur la moitie
proximale de 1'autozoecie, est tantot regulierement ovale, tantot a cotes presque paralleles (et dans ce cas a bords
proximal droit et distal arrondi) ; clle mesure de 0,12 a 0,16 mm de large et de 0,28 a 0,31 mm de long. Les petits
orifices des muscles occluseurs sont circulaires, et situes disto-lateralcment a l'opesie, a proximite du bord
de celle-ci. Le cryptocyste est finement granulcux.
Repartition. — Philippines, Nouvelle-Zelande. Espece nouvelle pour la faune neo-caledonienne. Profondeur :
310-790 m.
Bryopastor crassus sp. nov.
Fig. 6 C-D
MATERIEL EXAMINE. — Nouvelle-Caledonie. Biocal : stn. DW 36, 650 m. — Stn DW 41, 380 m. —
Stn DW 44, 440-450 m. — Stn DW 51, 680-700 m.
Types. — Holotype : Biocal, stn DW 36, 650 m (MNHN-BRY- 16660).
Paratype : BlOCAL, stn DW 51, 680-700 m (MNHN-BRY- 16664).
Source :
188
J.-L. D'HONDT & D. P. GORDON
DIAGNOSE. — Bryopastor a zoarium robuste de section hcxagonale. Occlusor laminae formant une fente
longitudinale large, plus ou moins nettement triangulaire. plus large proximalement que distalement.
Description. — Bryopastor a zoarium robuste (jusqua 1,4 mm de diametre) dc section hcxagonale, forme de
longues autozoecies spatuliformes de 0,99-1,30 mm de long et larges de 0,64-0,82 mm. L'opesie triangulaire,
a angle mousse dirigee distalement, est longue de 0,28-0,40 mm et large a sa base de 0,20-0,29 mm et a son
sommet de 0,035-0,050 (exceptionnellement jusqu'a 0,10) mm. Les orifices des muscles occluseurs sont situes
de part et d’autre dc la region distale de l'opesie, et debordent distalement au-dela de son cxlremite. Les occlusor
laminae , qui constituent deux lames planes se rejoignant distalement, sont plus ou moins ecartes proximalement ;
ils determinent entre eux une fente de forme nettement triangulaire, parfois presque equilaterale, et se retrecissant
graduellemcnt en direction dislale ; leur ecarlement a la base varic selon les colonies. Quelques autozoecies un peu
plus longues que leurs voisines (1,20 mm contre 1.00-1,06 mm), a opesies regulieremcnt ovales. longues de
0,60 mm et larges de 0,40 mm, sont presumees femelles (elles sont les plus nombreuses a la station DW 51).
Cryptocyste orne de denses petites granulations subglobuleuses.
DISCUSSION. — La morphologic de la fente determinee par les occlusor laminae est typique. Lc zoarium
de cette espece est par ailleurs regulieremcnt hexagonal, caracterc qui n'est connu, dans le genre Bryopastor, que
dans certaines portions de colonies de B. pentagonus , espece dont les colonies sont habituellement de section
pentagonale et dont les occlusor laminae se presentent sous la forme de cretes.
ETYMOLOGIE. — Du Latin, crassus , epais, en raison de 1'epaisseur des colonies.
Repartition. — Nouvelle-Caledonie, de 380 a 700 m de profondeur.
Bryopastor sp.
Fig. 6 E
MATERIEL EXAMINE. — Nouvelle-Caledonie. BiogLocal : stn DW 307, 470-480 m.
Description. — Les autozoecies mesurent 0.82-0,85 mm de long et 0.40-0,42 mm de large distalement, 0,30-
0,34 mm proximalement ; leur opesie a 0.30 mm de large et 0,32-0,35 mm de long. Sur Tun des fragments
zoariaux etudies existe une grande aulozoecie isolee, presumee femclle, longue de 1,24 mm et large de 0.66 mm du
cote distal, de 0,50 mm du cote proximal ; son opesie mesure 0,56 mm de long et 0,54 mm de large.
Remarque. — Vu 1'etat d erosion du materiel, I'identification generique est elle-meme douteuse.
Genre PSEUDOTHYRACELLA Labracherie, 1975
Espece-Type. — Pseudothyracella pulchella Labracherie, 1975.
Diagnose completee. — Zoarium arborescent candelabriforme, rami fie dichotomiquement, densement
recouvert de rhizoides. Zoarium dichotomiquement ramifie, chacunc des deux branches principales etant constitute
de petits entre-noeuds portant chacun distalement vers l'interieur un tres long entre-noeud non ramifie. Joints
pcletonnes. Autozoecies peu alternantes, parfois presque au meme niveau dune serie a I'autre. Zoecies femelles
beaucoup plus grandes que les autozoecies normales.
Pseudothyracella candelaber sp. nov.
Fig. 5 E, H ; 12 J ; 13 B-E ; 14 ; 18 A-D ; 19 B-D
Materiel EXAMINE. — Nouvelle-Caledonie. BlOCAL : stn DW 31. 850 m. — Stn DW 38, 360 m —
Stn DW 46, 570-610 m. — Stn DW 51. 700-680 m. — Stn DW 70. 965-960 m. — Stn CP 109, 495-515 m.
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
189
Musorstom 4 : stn CP 216, 490-515 m. — Stn DW 220, 505-550 m. — Stn CP 236, 495-550 m.
Biog£()CAL : stn DW 296, 1230-1270 m. — Stn CP 297, 1230-1240 m. — Stn DW 307, 470-480 m.
Smib 4 : stn DW 55, 215-260 m.
lies Loyaute. Musorstom 6 : stn DW 421, 245 m.
Nouvelle-Zelande. West Norfolk Ridge. Wanganella Bank : stn BS 886, 29.01.1981 : 32°35,3'S, 167°41,8'E,
437-422 m (NMNZ). — Three Kings Ridge : stn U582, 31°52,0'S, 172°26,5'E. 1058-988 m (NZOI).
Types. — Nouvelle-Caledonie. Holotype : Biocal, stn DW 46, 570-610 m (MNHN-BRY- 1 6406).
Paratypes : BiOCAL, stn DW 70, 960-965 m (MNHN-BRY-16429) ; stn CP 109, 495-515 m (MNHN-BRY-
16427).
Diagnose. — Pseudothyracella actuelle a zoarium robuste el joints chitincux pelotonnes, a autozoecies de
forme rdgulierement hexagonale mesurant environ 0,80 mm de long, a opesie distale ovale a tendance
quadrangulaire presque deux fois plus longue que large. Zoecies fcmelles a bords lateraux parallels, droits, a opesie
ovale, une fois el demie plus grande qu'une autozoecie normalc. Zoecies aviculariennes de meme longueur que les
autozoecies, mais nettement elargies proximalement, ct avec un cryptocyste en occupant la moitie de la longueur,
presque deux fois plus large que celui des autozoecies normales.
Description. — Le zoarium, dresse et ramifie dichotomiqucment, atteint 10 cm de haut ; il comporte six
series longitudinales de loges et a des contours hexagonaux, parfois presque circulates, en section transversale.
D'une partie inferieure, formee de quelques entre-noeuds successifs separes par des joints chitineux pelotonnes,
partent deux ramifications principales formant entre elles un angle tres ouvert ; chacune de ces deux ramifications
est constitute par une chaine de courts entre-noeuds "primaires", longs de 4-7 mm, eux-memes separes les uns des
autres par des joints pelotonnes. Chacun de ces entre-noeuds porte a son extremite distale, separe aussi par un joint
pelotonne, un entre-noeud "secondaire" extremement allonge et non ramifie, de longueur decroissante de la partie
proximale de la colonie (oil il atteint la longueur de 6,4 cm) vers la partie distale ; ainsi tous les entre-noeuds
"secondaires" issus d'une meme branche principal atteignent-ils sensiblement le meme niveau. Toutes les
branches "secondaires" etant dirigees vcrticalement et paraMement a l'axe du zoarium, chacune des deux branches
principales et ses ramifications se presente done comme un peigne a dents de longueur decroissante vers le haut, et
symetrique d'un peigne analogue par rapport a l’axe longitudinal ; l'ensemble de la colonie affecte done une forme
de candelabre, comme certaines especes de Bifaxarioidea. Le diamelre des entre-noeuds est de 1,4 mm.
Les entre-noeuds qui constituent la partie basale de la colonie sont plus aplatis que les autres entre-noeuds. Ils
sont recouverts par un feutrage de longs rhizoides longitudinaux, issus au nombre de un ou deux par loge des
parties proximo-frontales des autozoecies les plus inferieures. Les branches principales sont elles-memes bordees,
de chaque cote, par un epais feutrage de rhizoides.
Les autozoecies sont hexagonales ; elles alternent. mais le deealage d’une serie zoeciale a 1'autre est peu mar¬
que ; elles mesurent de 0,77 a 0,84 mm de long et de 0,38 a 0.42 mm de large ; leur opesie, de forme ovale
a tendance quadrangulaire, a 0,36-0,38 mm de haut et 0.18-0,22 mm de large ; il ny a pas de denticules
aperturaux ; l'opercule a 0,16-0,18 mm de long et 0.28-0,30 mm de large ; le cryptocyste est tres finement et peu
densement granuleux, les tubercules etant plus serres lateralement que proximalement. Les zoecies femelles sont
assez nombreuses, et exceptionnellement contigues sur deux series zoeciales alternantes ; leur longueur varie de
0,81 a 1,40 mm ct leur largeur de 0,38 a 0,60 mm a la base ; leur opesie occupe un peu plus des 2/3 de la
longueur autozoeciale, le cryptocyste, granuleux et incurve, mesurant de 0,20 a 0,45 mm de long : le bord interne
du cryptocyste est finement denticule. L'aviculaire est long de 0,81 a 0,85 mm ; il est arrondi et plus etroit
(0,35 mm) a son extremite distale, elargi dans sa partie proximale (0.68 mm) ; celle-ci. dont les contours ont une
forme sensiblement hexagonale, presente parfois des processus spini formes pointes distalement au-dessus de
l’orifice de passage du muscle de la mandibulc. La mandibule avicularienne ressemble a un opercule autozoidal.
DISCUSSION. — Le genre Pseudothyracella a ete defini par LabraCHERIE (1975) pour une espece de
Bryozoaires fossiles de l'Eocene infericur nord-aquitanien (Cabanac. Gironde), P. pulchella , genre auquel I auteur
rattachc aussi une espece nord-americaine du Paleocene d'Arkansas dont l'opesie est plus proximale que chez
I'espece-type et la nouvellc espece decrite ici, P. midwayanica (Canu & Bassler, 1920). Chez P . pulchella , les
Source :
190
J.-L. D'HONDT & D. P. GORDON
autozoecies sont moins regulierement hexagonales que chez P. candelaber , dont l’opesie est par ailleurs plus large.
Voigt (1987) a redecrit P. pulchella , dont les aviculaires et les zoecies reproductrices different de ceux de
P. candelaber , et decrit ou reetudie en outre trois autres especes fossiles du genre Pseudothyracella provenant du
Montien de Mons. Le materiel etudie par ces differents auteurs n'etant constitue que de fragments d'entre-noeuds
dissocics, sur lesquels aucun joint n’avait etc conserve, ils ne pouvaient avoir une vue d'ensemhle des colonies, et
e'est pour cettc raison que tous les ont classees dans la famille Onychocellidae. Chez P. ciplyensis Voigt, 1987,
les contours autozoeciaux sont plus parallels et les opesies petites ; P. mucronata (Mcunier & Pergens, 1886) a
une petite opesie etroite et rectangulaire bien plus large que haute et des pores frontaux ; chez P. vandenbroecki
(Meunicr & Pergens, 1886), il existe des ovicelles globulcuses saillantes.
Remarques. — Ce genre n’etait jusqu'a present connu qu’a Petal fossile et de la seule Europe occidentale ;
il est pour la premiere fois trouve dans la faune actuelle, et dans une region tres eloignee de celle d'oii des
specimens fossiles etaient connus. Ces recoltes permettent enfin de connaTtre la morphologic zoariale dcs
Pseudothyracella , que la simple etude des fragments jusqu'a present connus ne permettait pas d'imaginer, et de les
ranger dans leur famille legitime.
ETYMOLOGIE. — Du Latin, candelaber , candelabre, afin de rappeler la forme des colonies de cette espece.
Repartition. — Nouvelle-Caledonie, Ties Loyautc, Nouvelle-Zelande, de 215 a 1270 m.
Infra-Ordre CRYPTOCYSTOMORPHA Silcn, 1942
Famille ONYCHOCELLIDAE Jullien, 1882
Genre SMITTIPORA Jullien, 1882
ESPECE-TYPE. — Vincularia abyssicola Smitt, 1873.
Smittipora fenestrata sp. nov.
Fig. 7 A-D. F
Materiel EXAMINE. — Nouvelle-Caledonie. Biocal : sin CP 84, 460 m.
Type. — Holotype : Nouvelle-Caledonie. Biocal, stn CP 84, 460 m (MNHN-BRY- 16604).
Diagnose. — Smittipora a zoarium retepori forme, les orifices autozoeciaux ne s'ouvrant que sur la face
frontale. Presence de nombreuses zoecies aviculariennes de memes dimensions que les autozoecies, a mandibule
aliforme et rachis lisse.
DESCRIPTION. — Le zoarium est monostratifie, les orifices autozoeciaux etant portes par la seule face frontale
du zoarium ; la face frontale est concave (cette morphologie zoariale suggere a tort que Ic materiel etudie etail
initialement encroutant, et avait ete detache d'un substrat reteporiforme ; il n'en est rien, puisque la face dorsalc des
colonies vivantes porte des organismes epibiontes, dont des colonies encroutantes d'autres especes de Bryozoaires).
La colonie est constitute d'autozoecies et de zoecies aviculariennes, ces dernieres etant plus nombreuses sur les
bords des "mailles" determinees par le substrat. Les zoecies aviculariennes occupent dans la colonie l'cmplacement
qui aurait normalement du etre devolu a une autozoecie ; elles ont la mcmc longueur (0,46-0,60 mm) et parfois la
meme largeur (0,29-0,42 mm), mais sont generalement un peu plus etroites ; elles portent une mandibule
aliforme, dont les deux lobes, longs de 0,24 mm et sensiblemcnt en forme de quarts de cercle (de 0,14 mm de
rayon), sont symetriques par rapport a un rachis lisse qui atteint 0,40 mm de longueur. Il n'a pas ete observe
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
191
d'ovicclles. Les autozoecics onl une opesie de 0,19 a 0,22 mm dc long et de large ; ovale, elle est souvent plus
large proximalement ; son bord proximal est plus ou moins rectiligne. Le cryptocyste, granuleux, a le meme
aspect chez les autozoecies et les zoecies aviculariennes ; celui des autozoecics mesure de 0,12 a 0,20 mm de long.
DISCUSSION. — La morphologie zoariale, reteporiforme, est unique dans le genre Smittipora. Le rachis de
l'aviculaire est depourvu de dents, contrairement a S. cordiformis Harmer, 1926. Cette nouvelle espece se
rapproche, par la morphologie des deux types zoeciaux. du groupe d'especes affines, toutes encroutantes, constitue
des S. abyssicola (Smitt, 1867), S. levinseni (Canu & Bassler. 1928) et C. harmeriana (Canu & Bassler, 1929),
discute par Marcus (1953), Cook (1964, 1985), Winston et Heimberg (1986) et Hayward ( 1988).
ETYMOLOGIE. — Du Latin, fenestratus, garni de fenetres, en raison dcs nombreuses perforations du zoarium.
Repartition. — Nouvelle-Caledonie, a 460 m de profondeur.
Famille MICROPORIDAE Gray, 1848
Genre MICROPORA Gray, 1848
Espece-Type. — Flustra coriacea Johnston, 1847.
Micropora equilateralis sp. nov.
Fig. 7G, 9F, 19 E
MATERIEL EXAMINE. — Philippines. MUSORSTOM 3 : stn CP 139, 240-267 m.
Type. — Holotype : Philippines. Musorstom 3, stn CP 139, 240-267 m (MNHN-BRY- 16607).
Diagnose. — Micropora a aviculaire en forme de triangle equilateral, oricnte selon le grand axe de
l'autozoecie, distal par rapport a l'orifice, inconstant, a angle pointe distalemenl. Pas d'epines. Opesiules petites, au
nombre d'une paire, et proportionnellement tres posterieures.
DESCRIPTION. — Le zoarium encroutant est forme d’autozoecies ovales, de 0,30-0,32 mm de long et de 0,18-
0,22 mm de large, sdparees par des murailles saillantes. Elies ne presentent qu'une seule paire d'opesiules,
circulaires, proportionnellement tres posterieures, puisque situees aux 2/5 de la longueur autozoeciale en partant de
l’extremite distale. L'aviculaire, d'orientation axiale, distal a l'orifice, a pointe dirigee vers l'autozoecie suivante, est
inconstant ; il a la forme d'un triangle equilateral de 0,12 mm dc hauteur ; il fait toujours defaut en presence
d'ovicelle. L'orifice, en forme de segment de cercle, a une largeur de 0,1 1-0,12 mm et une longueur de 0,10-
0,1 1 mm chez les zoecies ovicellees, de 0,07 mm chez les zoecies non ovicellees. Le cryptocyste est finement et
uniformement perfore. Proximo-latdralement a l'orifice, le cadre lateral s'epaissit en bourrelets lateraux peu
marques. L'ovicelle, granuleuse, mesure 0,18 mm de long comme de large.
Discussion. — L'orientation et la forme dc l'aviculaire caracteriscnt cette espece. Chez les autres especes du
genre, il est soit completement absent, soil oriente plus ou moins obliquement en direction proximale ou distale.
Contrairement a plusieurs autres especes de la region indo-pacifique ( M . coriacea, M. elegans ), elle est depourvue
d'epines. Ses opesiules, dont il n'existe qu'une seule paire (contrairement a M. gracilis et a M. variperforata ), sont
petites et tres posterieures (contrairement a certaines formes de M. mortenseni a opesiules tres marquees).
ETYMOLOGIE. — Du Latin aequilateralis , pour rappeler la forme en triangle equilateral des aviculaires.
Repartition. — Philippines, a 240-267 m de profondeur.
192
J.-L. D’HONDT & I) P GORDON
Genre PROMICROA nov.
ESPECE-TYPE. — Promicroa dubitata sp. nov.
Diagnose. — Microporidae arborescente. biseriee, doni les deux series aulozoeciales sont portees par la memo
face du zoarium ; zoarium ramifie, sans joints ni aviculaires ; autozoecies a deux paires d'opesiules dislo-latcrales
successives de chaque cote ; ovicelles granuleuses finement perforees.
Le genre etant monospecifique, les diagnoses generique et specifiquc scront provisoirement con fondues.
ETYMOLOGIE. — Anagramme du nom generique Micropora.
Promicroa dubitata sp. nov.
Fig. 4 B. 7 E
Materiel EXAMINE. — Nouvelle-Caledonie. Biocai. : stn DW 66. 515-505 m.
Nouvelle-Zelande. Au nord des iles Norfolk, stn G4. 28°25’S. 1 67° 1 5’E. 831 m (NZOI).
Type. — Holotype : Nouvelle-Caledonie. Biocal, stn DW 66, 505-515 m (MNHN-BRY- 16607).
Description. — Le zoarium, arborescent et biscrie, ramifie dichotomiquement. sans joints, est constituc dc
deux series alternantes d'autozoccics pisciformes longues de 0,55-0,60 mm et larges de 0,22-0.26 mm. portees par
la meme face du zoarium et obliques 1'unc par rapport a l'autre. Chaque autozoecie porte. de chaque cote, deux
opesiules successives, longues, la plus distale de 0,045-0.050 mm et la plus proximale de 0,030-0,035 mm.
La plus distale est situee au 1/3 de la longueur autozoeciale cn partant de I'extremite distale. Le cryptocyste porte
une demi-douzaine de perforations eparses. Les bourrelets lateraux pre-aperturaux sont ires peu marques. II n 'exisle
pas d’aviculaires. Les ovicelles ont une surface granulcuse et presentent dc nombreuses perforations minuscules,
eparses sur leur surface ; elles presentent aussi une lame calcaire frontale en forme de Y. a branches ires larges ;
I'une de ces branches est axiale et se situc sur la partie proximale de I’ovicelle, les deux autres branches sont
laterales et longent le bord distal de lorifice en encadrant I'anter. L'orifice, dc forme semi-lunairc. a 0,06 mm de
long et 0,14 mm dc large. Le cote lateral des zoecies porte, presque a la limile de la face dorsale. une rangee de 4
pores arcolaires equidistants.
Discussion. — Cette espece presente les caracteres de la famille Microporidae (avec laquelle Prenant et
Bobin, 1966, ont fusionne les Calpensiidae. les criteres discriminatifs entre elles leur paraissant llous). qui reunit
des genres cncroutants. avec ou sans aviculaires, dresses, cupuliformes ou claviformes avec joints ; les ovicelles
sont absentes ou endozoeciales. le cryptocyste tres etendu et l’opesie petite, les opesiules distales etant ouvertes ou
fermees, I'operculc plus ou moins hemicirculaire.
Cette nouvelle espece ne trouve place dans aucun dcs genres preexislants, et e'est pour cette raison qu'elle est
decrite sous un nouveau taxon de rang generique. Le nombre des opesiules, le port de la colonic et a un moindre
degre rornementation dc I'ovicelle (presentant de nombreuses petites perforations, caractere qui dans ce genre n'est
presente que par une scule espece. M. santacruzana Soule, Soule & Chaney, 1995) ecartent cette espece des
Micropora. Nous limiterons la discussion ci-apres aux genres de Microporidae qui ont un port dressc. et qui
nc sont pas qu cncroutants par definition. Monsella Canu, 1900, forme d'une succession de segments separes par
des entre-nocuds, sen diflerencie en outre par la presence d'une lame calcaire frontale dclimitant des opesiules tres
allongees et extremement etroites : le genre Microporina est pourvu de joints, d'aviculaires. et d'une seule paire
d opesiules. Chez les Poricellaria , que BASSLER (1953) incluait parmi les Calpensiidae et qui appartiennent a
present a la famille Poricellariidae, le zoarium est quadriscrie, mais les autozoecies portent commc ici deux paires
d'opesiules.
Etymologie. Du Latin, dubitatus, douteux, en raison des affinites incertaines de 1'especc.
Repartition. — Nouvelle-Caledonie. Nouvelle-Zelande, a 505-83 1 m de profondeur.
Source : MNHN, Pans
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
193
Famillc ASPIDOSTOMATIDAE Jullicn, 1888
Genre CRATEROPORA Levinsen, 1909
ESPECE-TYPE. — Crateropora falcata Levinsen, 1909.
Remarque. — Genre nouveau pour la faune neo-ealedonienne.
Crateropora stiliformis sp. nov.
Fig. 9 A-E. 20 H-K
MATERIEL EXAMINE. — Nouvelle-Caledonie. Biocal : stn CP 78, 445-450 m.
Smib 4 : stn DW 39, 525-560 m.
Types. — Holotype : Nouvelle-Caledonie. Smib 4, stn DW 39, 525-560 m (MNHN-BRY- 16668).
Paratype : Biocal, stn CP 78, 445-450 m (MNHN-BRY- 16669).
Diagnose. — Crateropora presentanl un aviculaire lateral impair extremement allonge, parallele au cadre
zoecial, styliforme et incurve vers l'orifice, arrondi a son extremite. La partie distale du cadre zoecial sc renforce de
fa^on a former une paire de comes au-dessus de l'opercule, orientees cn direction proximalc. Ovicelle formant une
sorte de visiere. Presence d'epines distales chez les jeunes autozodcies.
DESCRIPTION. — Le zoarium encroutant est forme d’autozoccies plus ou moins irregulierement hexagonales,
parfois a 7 cotes ; pour la colonic de Smib 4, elles sont longues de 1,30 mm (en peripheric de la colonie)
a 0,90 mm (dans la partie centrale), larges de 0,70 a 1,20 mm ; pour celle de Biocal. elles mesurent 1,80
-1,95 mm de long el 0,74-0,84 mm de large. Chez la colonie ou les zoecies sont les plus grandes, l'ovicelle est
tres saillante et forme de chaque cote, au-dessus de l'opercule, une corne robuste dirigee vers la region proximale.
Le cadre zoecial, tres epaissi distalement, a une largeur de 0,50-0,80 mm ; il est nettement retreci lateralement et
en avant de l'orifice. L'ovicelle, subglobulcuse, se presente comme une visiere saillante a l'extremite distale de
l'autozoecie, en forme de croissant, large de 0,60 mm et longue de 0,30-0,34 mm. La region de la lace frontale
dclimitee par le bord distal et le tube opesiulaire, de forme ovale, mesure 0,20 mm de long et 0,30 mm de large.
Le cryptocyste, fmement et densement granuleux, est Finement pore, el s'enfonce vers la region distale. II n'cxiste
pas d'epines. Le tube opesiulaire forme lateralement un processus, delimitant de chaque cote, entre lui et le bord de
1’opesie, un tres court sinus marginal en forme de U. L'opercule mesure 0,23-0,24 mm de large. Sur les plus
jeunes zoecies, les bases de quclqucs cpines (3-5), cassees a leur niveau d'insertion, sont distributes distalement
au peristome ; ces epines sont absentes chez les plus grandes autozoecics. Les aviculaires sont inconstants. Une
zoecie sur trois porte un aviculaire etroit et tres allonge, styliforme, peu ou moderement incurve vers l'orifice,
longeant exterieuremenl le cadre zoecial ; il a une longueur de 0,54 a 0,19 mm et une largeur de 0,12 mm a la
base, de 0,07 mm a l'extremite. Sur une unique zoecie, il a en plus ete observe un aviculaire plus large et plus
incurve, long de 0,30 mm, en position latero-distale par rapport a l'orifice, et sensiblement perpendiculaire a l'axc
autozoecial longitudinal. Chez deux des plus jeunes autozoecics, l'aviculaire impair etait plus spatuliforme et
arrondi distalement.
Discussion. — La forme caraeteristique des aviculaires, etroits et allonges, constitue le principal caractcrc
diagnostique de cette cspece. Elle rappelle C. falcata Levinsen. 1909. par la forme generate de ses autozoecics et
par celle de l'extremite du tube polypidien en arriere de l'opesie. La forme des aviculaires evoque celle decrite
chez C. expansa Harmer, 1926. mais chez cette derniere espcce ils sont pairs, plus courts, parfois spatules.
Il n'est pas impossible que la colonie figuree par GORDON (1985) et provenant des Kermadec, sur laquelle
il n'avait pas observe d'aviculaires et qu'il avail mentionnee sous le nom de C. falcata, appartienne en fait h cette
espece.
Source :
194
J.-L. D’HONDT & L) P. GORDON
Etymologie. — Du Latin, stilus, style et formis , en forme dc, du fait de I’existence d'aviculaires etroits et
allonges.
REPARTITION. — Nouvelle-Caledonie, entre 445-450 et 525-560 m de profondeur.
Famille CELLARI1DAE Fleming, 1828
Genre CELLARIA Ellis & Solander, 1786
Espece-Type. — Cellaria sinuosa Hassall, 1840.
Cellaria parafistulosa sp. nov.
Fig. 8 G. 17 D-E
Materiel EXAMINE. - Nouvelle-Caledonie. BiogSocal : stn 214. 1665-1590 m.
Type. — Holorype : Nouvelle-Caledonie. BlOGEOCAL, stn 214, 1665-1590 m (MNHN-BRY- 164 17).
DIAGNOSE. — Cellaria a joints tubulaires, a entre-nocuds fusiformes et areolation losangique, a zoecies avicu-
lariennes en forme de D. Pore ovicellien de forme variable : arrondi aux extremites de I'entre-nocud. il prend de plus
en plus une forme de croissant en se rapprochant de la region centrale, pour delimiter chez les ovicelles mures dc la
partie centrale n'ayant pas encore pondu une plaque obturatrice hemicirculaire granuleuse, herissee en Peripherie ;
cette plaque est cassee apres la ponte et le pore ovicellien acquiert alors une forme oblique et dissymetrique.
Frontale densement granuleuse, les granulations etant plus eparses dans la region immediatement proximale a
rorifice. Frontale un peu deprimee par rapport au cadre autozoecial. Cadre autozoecial se confondant avec les
limites interzoeciales, sauf a mi-longueur oil, bien que tres rapproches, ils sont distincts sur une breve distance.
"Orifice" commun a l’ouverturc autozoeciale et a l'opesie (accolees distalement comme chez tous les Cellariidae)
prdsentant une paire proximale de condyles articulaircs de 1'operculc, et une paire de denticules distaux.
Description. — Le zoarium est cellariiforme, dresse et ramifie dichotomiquement, les entre-noeuds greles
etant separes par des joints chitineux, jaunes chez les jeunes branches, brun-noir chez les plus agees. Les entre-
noeuds sont fusiformes, renfles en leur milieu, region ou sont localisees les zoecies reproductrices. Ces dernieres
mesurent 0,42-0,44 mm de long el 0,27-0,29 mm de large, contre 0.45-0,90 mm et 0,22-0,26 mm pour les
autozoecies normales. Celles-ci sont pratiquement losangiques ; lorsqu'elles tendent vers la forme hexagonale, les
cotes lateraux sont tres courts, sinon quasiment inexistants. Leur cadre se confond avec les limites interzoeciales ;
ce cadre n'est distinct du bord zoecial que sur une tres courte distance, a mi-longueur de I'autozoecie, ou il en est
(tres peu) ecarte. Les limites interzoeciales sont surelevees par rapport a la frontale, mais sans former une muraille
verticale (comme chez C. fistulosa ). Leurs orifices ont 0,130-0,145 mm de large et de 0.80-0,90 mm de long,
et pnSsentent une paire de denticules proximaux (condyles) et une paire de denticules distaux. Les zoecies
aviculariennes sont en forme de D a cole arrondi situe en position distale ; elles ont de 0,185 a 0,195 mm
de large et de 0,115 a 0,120 mm de long ; elles portent une mandibule dont le bord proximal est droit
ou faiblement incurve et le bord distal nettement arrondi, qui mesure de 0.1 1 a 0,12 mm de large et 0,07-0,08 mm
de long. La longueur d'un entre-noeud varie de 4 a 5,5 mm. La surface frontale des autozoecies est
densement et uniformement granuleuse, sauf dans la region proximale a l'orifice ou ces granulations sont plus
clairsemees.
Les ovicelles, longues de 0,13-0,14 mm, presentent un pore de forme variable selon leur position sur la partie
renflee de l'entrc-noeud. Aux deux extremites de la partie renflee, le pore est parfaitement circulaire et symetrique,
et a 0,05 mm de diametre. Lorsqu'il est situe en position plus centrale sur I'entre-nocud. ce pore s'elargil, acquiert
Source : MNHN. Pahs
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOM I DA DE NOUVELLE-CALEDONIE
195
d'abord la forme de croissant peu ouvert et symetrique par rapport a l'axe longitudinal, puis s'clargit, et mesure
alors 0,065 mm de long et 0,08 mm de large.
Dans sa region centrale de l'cntre-noeud, le pore ovicellien est en forme de croissant tres ouvert ; il affecte la
forme dune fente en croissant, longue et etroitc, et delimite ainsi une plaque calcaire qui 1'obture presque comple-
tement ; cette plaque est hemicirculairc, a une surface densement granuleuse, les granulations situees en peripherie
lui donnant un aspect herisse. Sur quelques autozoecies de la region centrale, cclles supposees avoir pondu, cette
plaque est cassec ; l'orifice ovicellien est alors oblique et dissymetrique et mesure de 0,05-0,06 mm de diametre.
DISCUSSION. — Cette espece resscmble morphologiquement beaucoup a I'espece europeenne Cellaria fistulosa
(Linne, 1758) (Fig. 17 A-C) ; seul leur examen en microscopie electronique a balayage permet de les differencier.
L'etude d'excmplaires de C. fistulosa, recueillis en mai 1996 au large de file de Batz (Roscoff), montre que chez
eux la surface frontale n'est que faiblement granuleuse et que les granulations sont eparses ; elles sont essentielle-
ment localisees proximalemcnl et dans la region dcs angles distaux. Sur une grande partie de la longueur de la loge,
le cadre zoecial et les limites intcrzoeciales sont nettement distants les uns des autres ; lc cadre forme une fine
muraille verticale continue et bien visible qui entoure la face frontale. La region proximale a l'orifice est lisse. Les
autozoecies ont une tendance hexagonale plus marquee, les cotes lateraux etant bien developpes, sans atteindre
toutefois rimportance qu'ils peuvent avoir chez les Cellaria du "type hexagonal vrai" de Prenant et Bobin ( 1966)
chez lequel les hexagones ne sont pas en contact dans une mcme file longitudinale. La forme des pores ovicelliens
est beaucoup moins variable que chez C. parafistulosa ; le pore est ovale, a tendance parallelepipedique, lorsque
l'ovicelle est associee a un aviculaire, de forme triangulaire avec angle distal dans les autres cas ; il mesure alors
0,10 mm de long et 0,80-0,85 mm de large a sa base ; il n'a jamais une forme de croissant et ne delimite jamais de
plaque granuleuse.
ETYMOLOGIE. — Du Grec, para , pres, pour rappeler combien cette espece est proche de I’espece de LlNNE,
fistulosa.
Repartition. — Nouvelle-Caledonie, a 1590-1665 m de profondeur.
Cellaria tenuirostris Busk, 1852
Cellaria tenuirostris Busk, 1852 : 17-18, pi. 63 fig. 4. — Gordon, 1984 : 58, pi. 18 fig. B ; 1986 : 74. pi. 29
fig. C-E.
MATERIEL EXAMINE. — Nouvelle-Caledonie BlOGkoCAL : stn KG 267, 1935 m.
DESCRIPTION. — Le zoarium dresse et quadriserie est constitue d'entre-noeuds de 0,30-0,40 mm separes par des
joints peu apparents. Les autozoecies, etroites et allongees, ont 0,60-0,82 mm de long et 0.20 mm de large a
1'avant ; hexagonales a bord distal arrondi, elles se retrecissent graduellement vers leur region proximale ; leur cadre
zoecial est tres saillant. L'orifice autozoecial, long de 0,10 mm et large de 0, 14 mm, est sensiblement en forme de
D, mais son cote proximal n'est pas rectiligne, mais incurve ; il existe une paire de denticules proximaux
(condyles). La frontale est presque lisse. Situees sur une portion plus rcnflee du zoarium, les ovicelles se
presentent sous la forme de bonnets granuleux et renfles de 0,12 x 0,12 mm. avec une fente etroite proximale-
ment ; la zoecie portant l'ovicelle est un peu plus large (0,30 mm) que les autozoecies normales. Un seul
aviculaire a ete observe, large de 0,28 mm et long de 0,68 mm ; il est insere a la place habituelle d'une auto-
zoecie ; sa mandibule, triangulaire et effilee, a 0,32 mm de long et 0. 135 mm de large a sa base.
REMARQUES. — Ce materiel est conforme a la description de I'espece, mais s'en differencie par l'etroitesse
des entre-noeuds. Cellaria tenuirostris a ete illustree a plusieurs reprises par I’un des auteurs de ce travail (D.P.G.),
et nous renvoyons a l'iconographie qu'il en a publiee. Cette espece est nouvelle pour la laune de
la Nouvelle-Caledonie.
Repartition. — lies Bonin, lies Chatham, detroit de Cook, Nouvelles Galles du Sud, Nouvelle-Caledonie.
Nouvelle-Zelande. Profondeur : 100-1935 m.
Source :
196
J.-L. D'HONDT & D. P. GORDON
Cellaria humilis Moyano, 1983
Fig. 7 H. 10 A
Cellaria humilis Moyano, 1983 : 7, fig. 15-17. — Gordon, 1984 : 58, pi. 18 fig. A ; 1986 : 76.
MATERIEL EXAMINE. — Philippines. Musorstom 3 : stn DR 117, 92-97 m.
Description. — Le materiel etudie consiste en fragments non ramifies de zoarium dresses, etroits
et quadriseries. Les autozoecies, allongees, hexagonales ou pisciformes, ont unc longueur de 0,66-0,78 mm
et une largeur de 0,28-0,42 mm a leur extremite distale (arrondie) et de 0,08-0,14 mm a leur extremite proxi-
male (en forme de queue de poisson) ; leur surface est lisse. L'orifice autozoecial est tres anterieur ; long de
0,065 mm, il a une largeur de 0.075-0,14 mm et presentc unc paire de denticules proximaux (condyles) ;
le bord proximal est a peine arque vers 1'avant. Le cadre zoecial est saillant cl suit assez regulierement les limites
autozoeciales ; il est plus attenue proximalement ou il dessinc deux aretes convergeantes mais n'entrant pas
en contact. L’ovicclle globuleuse a un diametre de 0,12 mm ; elle porte un pore en croissant, tres etroit,
de 0,35-0,40 mm de haut, parallele au bord distal de l’orifice. Un rhizoide s'implantc parfois au milieu
de la face frontale ; d'autres rhizoides constituent un faisceau a la base de la colonie. Il n'a pas ete observe
d'aviculaires.
Discussion. — Les caracteres observes concordent avec ceux mentionnes par Moyano (1983) et Gordon
(1984), qui font etat de la presence d’aviculaires. Aucun de ces deux auteurs ne mentionne l'existence de condyles
peristomiaux chez C. humilis (GORDON en signale la presence dans sa definition du genre) ; les echantillons
etudies ici en presentent, mais ils sont tres discrets. Il serait done intcrcssant de les rechercher sur les specimens
etudies par Moyano et Gordon.
Repartition. — Chili, Kermadec, par 350 m. Espece nouvelle pour la faune des Philippines, a 92-97 m de
profondeur.
Cellaria obliquidens sp. nov.
Fig. 15 A-C
Materiel EXAMINE. — Nouvelle-Caledonie. Biog£ocal : stn 214. 1665-1590 m.
TYPE. — Holotype : Nouvelle-Caledonie. BiOGEOCAL, stn 214, 1665-1590 m (MNHN-BRY-1641 7 bis).
DIAGNOSE. — Cellaria a entre-noeuds cuneiformes, tres elargis distalemcnt. Orifice autozoecial a bords
proximal et distal rectilignes, presentant une paire de condyles proximaux, orientes obliquement vers l'exterieur. et
une paire de denticules distaux.
Description. — Le zoarium robuste est constitue d'entre-noeuds larges, s'elargissant regulierement du simple
a pres du triple de leur partie proximale a leur partie distale, non claviformes, pouvant comporter jusqu’a 7 series
autozoeciales en examen frontal, et separes par des joints chitincux. Les autozoecies, regulierement alternantes,
hexagonales, a cryptocyste granuleux (un peu moins densement proximalement a l'orifice), sont longues de 0,53
a 0,56 mm et larges de 0,22 a 0,26 mm. L'orifice a une forme presque rectangulaire, ses bords proximal et distal
etant droits, mais a angles arrondis ; il mesure 0,135 mm de large et 0,085 mm de long ; il presente deux paires
de condyles : deux denticules distaux larges et emousses, deux condyles proximaux un peu plus fins, orientes vers
l'exterieur. Proximalement, le bord de l'orifice presente de chaquc cote unc profonde incisure. Aucun aviculaire n'a
ete observe. Le pore ovicellien est porte directement sur le bord distal de 1'autozoccic ; il a une forme de croissant,
accentuee par la presence d'une paire d'incisurcs proximo-laterales ; la longueur est de 0,045 mm et sa largeur
de 0,085 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOM1DA DE NOUVELLE-CALEDONIE
197
DISCUSSION. — La morphologic des entrc-noeuds, la forme tres particuliere dc 1'orifice avec ses deux bords
proximal et distal rectilignes, et l'orientation des condyles proximaux caractcrisent cette especc.
ETYMOLOGIE. — Du Latin, obliquus , oblique et dens , dent, les condyles aperturaux proximaux etant orientes
obliquement vers l'cxterieur.
Repartition. — Nouvelle-Caledonie, a 1590-1665 m dc profondeur.
REMARQUES GENERALES SUR LE GENRE CELLARIA
Le reexamcn, a 1'occasion de cette etude, de nombreux specimens de Cellaria nous a rememore unc precedente
conversation avec notre collegue Hugo I. Moyano, a qui nous sommes redevables d'importantes etudes sur les
Cellariidae, et qui avait attire rattenlion de Tun des auteurs de ce travail (J.-L. d'H.) sur 1'heterogeneitc de ce genre.
Les especes regroupees dans le genre Cellaria se repartissaient en effet en deux groupes, caracterises par deux types
differents de ramifications zoariales, qui avaient deja ete analyses et discutes par BUSK (1884 : 84) : celles a joints
tubuleux ( tubulatae ) et celles a joints pelotonnes ( nodatae ), caracteres qu’il a correles avec 1'areolation frontale
(rhomboidale chez les tubulatae , hexagonale vraie chez les nodatae). Mais Hastings (1947) a toutefois montre que
cette correlation n'etait pas aussi stride que nc le pensait BUSK, que des especes presentant des joints differents
pouvaient etre plus affines entre elles qu'avec des especes du meme modcle de joints, et que les joints
ulterieurement pelotonnes et rigides se formaient de fat^on identique chez les jeunes colonies des deux types. Aussi,
bien qu’il serait a priori tres tentant de scinder le genre Cellaria en genres distincts caracterises par leur mode de
ramification zoariale, nous avons prefere y renoncer en reference aux remarques formulees par Hastings.
Genre FORMOSOCELLARIA d’ Hondt, 1981
ESPECE-TYPE. — Formosocellaria magnifica (Busk. 1884).
Formosocellaria magnifica (Busk, 1884)
Fig. 10 C
Salicornaria magnifica Busk. 1884 : 93-95, pi. 12 fig. 4, 6.
Formosocellaria abyssicola d'Hondt, 1981a : 20-22 ; 1983 : 85.
Formosocellaria magnifica - D'HONDT & SCHOPF, 1984 : 931.
MATERIEL EXAMINE. — Nouvelle-Caledonie. BlOCAL : stn DW 36. 625-650 m.
Musorstom 4 : stn DW 221, 535-560 m.
Biog£ocal : stn CP 265, 1760-1870 m. — Stn CP 272, 1615-1710 m.
Mer de Tasmanie. New Zealand Oceanographic Institute : stn U202. 28.09.1982, 33°43,7'S, 160°16,2’E, 3480-
3543 m (NZOl).
Description. — Les echantillons etudies ici correspondent parfailement aux specimens decrits par d'Hondt
(1981) sous le nom de Formosocellaria abyssicola, h la description desquels nous renvoyons, et qui ont
ulterieurement ete compares au materiel-type de BUSK prete par le British Museum. II n'a pas ete observe
d’ovicelles sur le materiel etudie ici. La colonie est fixee au substrat par un faisceau basal de rhizoides.
La bifurcation du zoarium s’effectue du meme cote de la colonie. Les condyles aperturaux sont absents.
Repartition. — Cette espece n'etait jusqu'a present connue que de I'Atlantique intertropical, par 587-4749 m
de profondeur. Les presentes recoltes ont ete effectuees entre ces limites bathymetriques et jusqu'a 535 m. mais
dans une region du globe (Nouvelle-Caledonie et mer de Tasmanie) qui en est tres eloignee, ce qui suggere
l'existence probable de localites intermediaires.
Source :
198
J.-L. D'HONDT & D. P. GORDON
Genre EUGINOMA Jullien, 1882
ESPECE-TYPE. — Euginoma vermifomiis Jullien, 1882.
Euginoma conica Gordon, 1986
Fig. 10 D
Euginoma conica Gordon, 1986 : 76, pi. 30 fig. E.
MATERIEL EXAMINE. — Nouvelle-Caledonie. BlOCAL : stn KG 103, 630 m.
Biog£ocal : stn KG 210, 1 190 m. — Stn KG 219. 570 m. — Stn KG 227, 500 m. — Stn KG 316, 1660 m.
Description. — Lc zoarium s'acheve a sa base par un tube chitineux de 2,1 mm de diametre, dent le sommet
constitue un manchon autour de la partie inferieure effilee et calcifiee de la colonic ; ce lube, tres plisse. a I'aspect
d'un peristome de Nolella gigantea. La colonic est conique, a pointe (ancestrula) dirigee vers lc bas ; ellc nc mesure
que de 1 a 2 mm de haul. Les autozoecics, pentagonales, sont alternantes, les plus grandcs mesurant 0,40 mm de
large et 0,36 mm de haut. Leur orifice, presque circulate, est en forme de D. un peu plus large (0,12 mm) que
long (0,10 mm). II n'existe pas d'aviculaires ni de dcnticules aperturaux. Le cryptocyste, finement granulcux, a lui
aussi une forme de D. Sur les echantillons etudies ici, le cadre zoecial est moins prononce proximalement que sur
la photographic publiee par GORDON ( 1986).
Repartition. — A Test du detroit de Cook, plateau du " Challenger ", entre 1457 et 3347 m. Espece nouvelle
pour la faune neo-caledonienne. a moindre profondeur (500-1660 m).
Genre SYRINGOTREMA Harmer, 1926
Esp^CE-Type. — Syringotrema auriculatum Harmer, 1926.
Syringotrema calobi sp. nov.
Fig. 10 E-G
MATERIEL EXAMINE. — Nouvelle-Caledonie. BlOCAL : stn DW 66, 510 m.
SMIB 4 : stn DW 35, 520-525 m.
Type. — Holotype : Nouvelle-Caledonie. BlOCAL, stn DW 66, 510 m (MNHN-BRY-16435).
DIAGNOSE. — Syringotrema a orifice autozoecial elargi transversalement, et de forme ovale ou en croissant.
Description. — Le zoarium, dresse et ramifie sans joints, est fixe au substrat par des rhizoides issus de sa
base ; il est quadriscrie, les files autozoeciales etant disposees en spirale autour de 1'axe. La longueur autozoeciale
varie de 0,45 a 0,56 mm. la largeur de 0,28 a 0,32 mm. L'orifice, median, est en forme de croissant peu ouvert,
parfois pratiquement ovale ; ii est large de 0,19-0,21 mm et long de 0,40 a 0,65 mm ; son bord proximal est
legerement arque, et correspond a l'extremite du tube apertural. Le cadre opesial est tres saillant et crenele ; de
forme generale losangique, mais arrondi distalement, il a une longueur de 0,28-0,31 mm et une largeur de 0,24-
0,255 mm. Il n'a pas ete observe d'ovicelles et il n'existe pas d’aviculaires. La ramification est tres ouverte ; la
zodcie axiale a la base dune ramification est un peu plus grande que les autres.
DISCUSSION. — Ce genre etait jusqu'a present monospecifiquc pour 5. auriculatum , espece indonesienne dont
l'orifice est regulierement ovale et l'opesie arrondie proximalement.
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
199
ETYMOLOGIE. — Lc nom specifique calobi est une anagramme de BlOCAL.
Repartition. — Nouvelle-Caledonie, entre 510 et 525 m de profondeur.
Genre MESOSTO MARIA Canu & Bassler, 1927
Espece-Type. — Mesostomaria strictoramae Canu & Bassler, 1927.
Mesostomaria strictoramae Canu & Bassler. 1927
Fig. 5 I, 10 B
Mesostomaria strictoramae Canu & Bassler, 1927 : 4-5, pi. 1 fig. 3 ; 1929 : 176-177. — Gordon, 1984 : 59.
pi. 18 fig. F.
MATERIEL EXAMINE. — Philippines. Estase : stn DR 07. 890-450 m.
Nouvelle-Caledonie. BlOCAL : stn DW 33, 675-680 m. — Stn DW 36, 625-650 m. — Stn CP 52. 600-540 m.
Musorstom 4 : stn DW 151, 200 m.
Smib 3 : stn. DW 22, 503 m.
Description. — Le zoarium cylindrique et non articule, dresse et ramifie dichotomiquement (bien que de fa$on
irreguliere), est fixe au substrat par dcs rhizoides, issus de la partie proximale de certaines autozoecies. Trois
rangees d'autozoecies sont visibles de chaque cote ; non alternantes, regulierement disposes cote a cote au meme
niveau, elles sont hexagonales et mesurent habituellcment de 0,58 a 0,70 mm de long et de 0,46 a 0,53 mm
(exceptionnellement 0,59 mm) de large. L'orifice, presque circulaire, a neanmoins son bord proximal presque
rectiligne ; sa largeur est normalement de 0,14-0.16 mm (exceptionnellement jusqu’a 0,20 mm), et sa longueur de
0,13-0,15 mm. II n'existe pas de denticules aperturaux, ni de mucron, ni d'aviculaires. Les zoecies ovicellees sont
plus grandes quc les autozoecies normales, ayant une longueur de 0,80-0,85 mm et une largeur de 0,78-0,80 mm ;
toutes les ovicelles observees etaient brisees, seul en restait le fond, orne d'une reticulation longitudinale ; leur
longueur est comprise entre 0,44 et 0,55 mm, leur largeur entre 0,31 et 0,37 mm ; elles semblent etre
endotochoidales et subglobuleuses. L'opercule est incline, sa region distale etant enfoncee par rapport au plan de
la frontale. La surface autozoeciale est granuleuse.
DISCUSSION. — La seule espece qui puisse etre confondue avec celle-ci, M. atlantica (Busk, 1884), presente un
zoarium aplati.
Repartition. — Philippines, Sabah ; Kermadcc. Genre et espece nouveaux pour la taune neo-caledonienne.
Profondeur : 200-890 m.
Mesostomaria sp.
Fig. 1 1 A-D
MATERIEL EXAMINE. — Nouvelle-Caledonie. SMIB 3, stn DW 22, 503 m.
Description. — Non alternantes, les autozoecies sont disposees en rangees parallels et sont situees au meme
niveau ; elles mesurent 0,58-0,66 mm de long et 0,54-0,59 mm de large ; l'orifice hemicirculaire, en position
centrale, a 0,12 mm de long et 0,22 mm de large. La surface frontale est ornee d’une granulation fine et homo¬
gene ; juste proximalement a l'opesie, le cryptocyste presente une striation longitudinale serree et dense, sur une
longueur d'une cinquantaine de microns. Les autozoecies portant des ovicelles sont un peu plus grandes ; longues
de 0,80-0,85 mm, elles sont larges de 0,78-0,80 mm. Les ovicelles, toutes partiellement detruites, semblent avoir
ete hemiglobuleuses et endotochoidales et presentent au fond une reticulation longitudinale, elles mesurent 0,44-
0,55 mm de large et 0,31-0,37 mm de long. L'opercule est incline vers l'exterieur, et sa region distale semble
deprimee par rapport au plan de la frontale. II n'existe pas d'aviculaires.
Source :
200
J.-L. DHONDT & D. P. GORDON
Genre CR YPTOS T OMA RIA Canu & Bassler. 1927
ESPECE-TYPE. — Crypto stomaria crassatina Canu & Bassler, 1927.
Cryptostomaria alata sp. nov.
Fig. 6G, 11 E-G, 16 A-B
Materiel EXAMINE. — Nouvelle-Caledonie. BlOCAL : stn DW 38, 360 m. — Stn CP 62. 1395-1410 m. —
StnCP 74, 1300 m. — Stn DW 77, 440 m. — Stn CF 109. 495-515 m.
Musorstom 4 : stn CP 153, 235 m. — Stn CP 172, 275-330 m. — Stn CC 175, 355 m.
lies Loyaute. Musorstom 6 : stn KG 465, 480 m.
TYPE. — Holotype : Nouvelle-Caledonie. BlOCAL : stn DW 77, 440 m (MNHN-BRY- 16425).
Diagnose. — Cryptostomaria a surface autozoeciale parcouruc par une fine striation longitudinale serree.
Cadre zoecial eleve, surtout autour de I'orifice de chaque cote duquel il forme souvent une aile arrondie.
DESCRIPTION. — Le zoarium est dresse, cylindrique, subquadrangulaire en section, non articule mais presentant
souvent des renflements au niveau des bifurcations ; ses branches ont une epaisseur de 0,5 a 0,8 mm. II est com¬
pose de 4 series d’autozoecies alternantes et opposees deux a deux. La longueur autozoeciale varie de 0,80 a
0,91 mm, la largeur maximale de 0,40 a 0,44 mm ; l'opesie a une longueur de 0,28-0,36 mm et une largeur dans
sa partie ddprimee de 0,40 a 0,80 mm. Lc cadre autozoecial est tres saillant lateralement, formant de chaque cote
une Crete sur environ les 2/3 distaux de la loge. et s'elevant de part et d'autre de I'orifice de fa$on a y former une aile
calcaire reticulee et arrondie a son sommet, et se resserrant ensuite proximalement a I'orifice ; chacune de ces ailes
atteint une hauteur de 0,18-0,24 mm chez les zoecics jeunes, mais sont parfois ires erodees chez les autozoecies
agees. Le cryptocyste est plus ou moins fusiforme. deprime par rapport au cadre, et horizontal dans sa partie
proximale (le cadre zoecial est done bcaucoup plus en relief que les limites interzoeciales). Les faces frontale et sur¬
tout laterales sont parcourues par des rides longitudinales sinueuses, fines et serrees, reliees par une reticulation
transversale tres discrete. L'orifice autozoecial a une largeur de 0,1 15-0.12 mm et une longueur de 0,13-0,14 mm ;
il ne presente ni denticules ni cardelles. Les aviculaires remplacent des autozoecies dans la continuity du zoarium ;
ils sont rares, losangiques et atteignant 0,70 mm de longueur ; leur opercule triangulairc a 0,10 mm de long et
0,20 mm de large ; il est situe au centre, sa region distale est renflee, et il presente un petit cryptocyste hemicircu-
laire de 0,24 mm de diametre. Des rhizoides sont parfois issus des parties proximales de zoecies, notamment dans
les parties agees ; ils finissent par former un feutrage praliquement continu a la base des colonies, et adherent au
substrat. L'ovicelle est hemiglobuleuse, en forme de petite visiere recourbee en avant de I'orifice, a bord proximal
presque droit, legerement proeminent dans sa partie centrale ; elle dissimule parfois, en partie, I'orifice autozoecial
qui est situe dans un plan oblique ; elle mesurc 0,16 mm de long et 0,1 1 mm de large ; son pore est allonge trans-
versalement. En presence d'ovicelle, les ailes lateroperistomialcs paircs debutent en arriere du pore ovicellien.
Les jeunes colonies sont fixees par des faisceaux de rhizoides sur les branches d'autres zoariums, en les
entourant d'un veritable lacis. Les peristomes sont plus saillants chez les colonies jeunes que chez les agees ;
les cretes frontales du cadre zoecial se rapprochent proximalement a I'orifice, avant de se reecarter ensuite.
Discussion. — Cette espece est la seule du genre a presenter un cadre autozoecial aussi developpc, notamment
de chaque cote de I'orifice ou il forme des ailes bien developpees ; le resserrement tres marque de ce cadre,
proximalement a I'orifice, est unique a l’interieur du genre. La forme de l'ovicelle caracterise aussi cette espece.
L'ornementation autozoeciale est tres differente de cellc presentee dans le genre voisin Stomhypselosaria , dont
Gordon (1984 : 18) a illustre la surface tuberculeuse.
Etymologie. — Du Latin, alatusy aile, en raison de la forme des cretes laterales a I'orifice de la loge.
Distribution. — Nouvelle-Caledonie, Ties Loyaute, par 235-1410 m de profondeur.
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
201
Genre MELICERITA H. Milne Edwards, 1836
REMARQUE. — D'origine britannique et ne en Belgique, Henri Milne Edwards, a signe toutes ses
publications sous le double nom Milne Edwards (sans trait d'union), probablcment pour se distinguer de son
frere William. II a choisi le nom de Milne Edwards, des 1826. pour tout ce qui concerne son etat civil
(consulter a ce sujet, la note tres bien documentee de J. Forest, 1996).
Sous-Genre MELICERITA sensu stricto
ESPECE-TYPE. — Melicerita charlesworthii Morris, 1843.
Remarques. — Nous retiendrons comme definition du sous-genre nominatif la diagnose completee qui en a
ete publiee par GORDON (1986 : 76). Ce sous-genre n'avait pas encore etc mentionne de Nouvelle-Caledonie.
Melicerita (Melicerita) ejuncida Gordon. 1986
Fig. 10 H-I
Melicerita ejuncida Gordon, 1986 : 77-78, pi. 31 fig. D-E.
MATERIEL EXAMINE. — Nouvelle-Caledonie. MUSORSTOM 4 : sin CC 175, 355 m.
Biog£ocal : sin KG 222, 1675 m. — Stn CP 232, 760-790 m.
Description. — Le zoarium est aplati, dresse et ramifie dichotomiquement, dune largeur de 0,48 a 0.84 mm.
Les autozoecies hexagonales sont disposees en quatre series transversales non alternantes, done situees
parallelement les unes aux aulres et au meme niveau ; leur longueur (0,96-0,98 mm) et leur largeur (0,37-
0,41 mm) sont peu variables. Le cadre zoecial delimite une surface de 0,60-0.70 mm de long et 0.20-0.22 mm de
large. L'ovicelle apparait comme un bombement distal a l'autozoecie, d'un diametre de 0.14 mm, portant un pore
arrondi de 0,035-0,040 mm de long et 0,080 mm de large. Le cryptocyste et les parois laterales sont granuleux.
L'opesie a sensiblement les memes dimensions que l'orifice ; en forme de D, elle a 0,17-0,20 mm de long et
0,20 mm de large & la base. Le bord proximal de l'orifice porte un large processus (0,14 mm de large), delimite par
deux profondes indentations laterales. II n'a pas ete observe d'aviculaires.
Repartition. — Nouvelle-Zelande (plateau du "Challenger" a Test du detroit de Cook). Espece nouvelle pour
la faune neo-caledonienne. Profondeur : 132-1675 m.
Melicerita (Melicerita) a Iter nans sp. nov.
Fig. 12 A-D, 13 A
MATERIEL EXAMINE. — Nouvelle-Caledonie. BlOCAL : stn DW 08, 435 m.
Type. — Holotype : Nouvelle-Caledonie. Biocal, stn DW 08, 435 m (MNHN-BRY- 16437).
DIAGNOSE. — Melicerita a zoarium forme de verticilles successifs de deux et de trois autozoecies, alternant 1 un
avec l'autre. Autozoecies reproductrices situees en position axiale dans les verticilles de 3 loges. Les orilices
autozoeciaux sont, dans le cas des verticilles a deux autozoecies, disposes obliquement et situes dans 1' angle distal
interne. Pas d'aviculaires.
Description. — Les portions de zoarium etudiees constituent de petits Elements dresses et aplatis, longuement
ovales, depourvus de joints. Chacun de ces elements est constitue par une succession reguli6re de verticilles de
deux et de trois autozoecies, situees cote a cote au meme niveau et accolees sur toute leur longueur.
Les autozoecies des verticilles, constitues de deux loges, sont longues de 0,66-0.70 mm et larges de 0.77-
0,80 mm. L’orifice autozoecial hemicirculaire, long de 0,10 mm et large de 0,16 mm, est oblique ; son rebord
Source :
202
J -L. D'HONDT & D. P GORDON
distal arrondi, dirige vers I'exterieur, est situ 6 dans I'angle distal interne, sans deborder 1'axe longitudinal ; il porte
du cote proximal, rcctiligne, deux petits condyles reunis par une lame calcaire. Le cote anterieur dc la logo dessine
une ligne brisee, avec un angle proximal bien marque, dans lequel est partiellement contenu I’opercule.
Lc cryptocyste est granuleux. II n'existe pas d'aviculaires.
Les autozodcies des verticilles de trois loges sont elles aussi accolees sur toute leur longueur et done non
alternantes. Leur longueur varie de 0,80 a 0.84 mm. leur largeur de 0,17-0,20 mm pour les laterales. 0,37-
0.41 mm pour l'axiale. Les laterales sont plus etroites dans leur partie centralc, plus saillantes vers I'exterieur a
leurs deux extremites ; leur orifice est oblique vers I'exterieur, le rebord proximal etant tourne vers l'axe zoarial.
L'axiale est de forme hexagonale, et un peu plus etroitc dans sa partie mediane qu'a ses deux extremites ; I'orifice,
central et symetrique par rapport a l'axe longitudinal, en forme de D aplati, est situe au quart de la longueur en
partant de l'extremite distale. Seules des zoecies axiales des verticilles de 3 sont susceptiblcs d'etre ovicellees ; elles
sont alors un peu plus grandes que leurs homologues non ovicellees (longueur : 0,86 mm ; largeur : 0,50 mm).
L'ovicellc, en forme dc large croissant (0,61 mm de large et 0,36 mm de long), est endotochoidale, tres faiblement
renflee en surface, et seulement visible par transparence ; elle presente une tres fine reticulation transversale du cote
proximal. Le pore ovicellien, ovale, a 0,90 mm de long et 0,24 mm de large. II n'existe pas d'aviculaires. Un
meme segment peut porter plusicurs autozoecies axiales ovicellees, separees l'une de 1'autre par un verticil lc dc
deux loges non reproductrices.
Discussion. — Cette espece se differencie de toutes les autres Melicerita connucs par sa disposition reguliere
en verticilles alternant de 2 et 3 autozoecies en vuc frontale. La presence d'une ovicellc coiffant seulement la zoccic
axiale des verticilles dc trois est unique dans le genre.
ETYMOLOGIE. — Du Latin, alternans , alternant, pour rappeler la disposition des loges en verticilles successifs,
rcspcctivement formes de 2 et 3 zoecies, se succedant en altcrnance.
Repartition. — Nouvelle-Caledonie, a 435 m de profondeur.
Sous-Genre HENRIMILNELLA nov.
Espece-Type. — Henrimilnella cirticulata sp. nov.
Diagnose. — Colonic dressee, bistratifiee, aplatie, constitute de petits elements ; autozoecies hexagonales,
groupees en series alternantes. Limites interzoeciales tres peu saillantes. Orifice hemicirculaire (incline vers
I'exterieur sur les autozoecies marginales) presentant une paire de condyles aperturaux proximaux et une paire de
denticules distaux. Cryptocyste lisse, ou tres finement granuleux. Pas d’aviculaires.
DISCUSSION. — Ce sous-genre se differencie des Melicerita (subgn. Melicerita) par la morphologic de ses
elements ; ils sont en forme de batonnets, a extremite distale arrondie et extremite proximale effilee, la possession
constante de zoecies alternantes, la discretion des limites interzoeciales, 1'existence de deux paires de condyles
aperturaux et un cryptocyste pratiquement lisse. La non-observation d’individus en reproduction ne permet pas
d'elargir la discussion, mais il n’est pas h exclure qu'une meilleure connaissance de cette espece, ainsi que de la
suivante qui en est tres affine, a partir de specimens plus complcts, justifie en fait pour elles la creation d'un
nouveau genre.
Etymologie. — Ce sous-genre est dedie a la memoire d'Henri Milne Edwards, en souvenir dc ses
rccherches anatomiques et ecologiques sur les Bryozoaires marins.
Melicerita (Henrimilnella) articulata sp. nov.
Fig. 12 F-G
Materiel EXAMINE. — Nouvelle-Caledonie. BlOCAL : stn DW 38, 360 m.
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
203
TYPE. — Holotype : Nouvelle-Caledonie. BlOCAL, stn DW 38, 360 m (MNHN-BRY- 16443).
DIAGNOSE. — Melicerita (Henrimilnella ) triseriee. Zoarium petit et en forme dc baguette.
DESCRIPTION. — Les morceaux de zoarium etudies se presentcnt comme dc petites baguettes triseriees a bords
paralleles, aplaties et non ramifiees, plus etroites et effilees a leur extremite inferieure. II n’a etd observe ni
ovicelles ni aviculaires. Les series d’autozoecies sont alternantes ; cellcs de la file axialc ont leurs orifices
symetriques par rapport a l'axe longitudinal, celles des laterales ont leurs orifices legerement obliques vers
1'extdrieur et moins distaux que les precedentcs. Le cryptocyste est plat, presque lisse. Les autozoecies axiales
mesurent de 0,52 a 0,56 mm de long pour une largcur de 0,28-0,30 mm ; les limites interzoeciales ne sont pas
saillantes ; elles sont hexagonales a bord antdrieur arrondi, un peu plus effilees a l'arricre (jusqu'a 0,16 mm).
L'orifice est tres anterieur (il est situe dans le quart ou lc cinquiemc distal de l’autozoecie) ; son bord proximal est
droit, et presentc une paire dc condyles proximaux et une paire de denticules distaux ; il est long de 0,08 mm et
large de 0,12 mm. Les autozoecies laterales ont un bord externe droit et un bord interne presentant a mi-longueur
un angle dirige vers l'axe autozoecial.
DISCUSSION. — Cette espcce se differencie de la suivante par la triseriation de son zoarium. l'emplacement de
l'orifice autozoecial et le parallelisme sur une grande longueur de ses articles.
ETYMOLOGIE. — Du Latin, articulate /, articule, chaque fragment ayant la forme d'un article d'appendicc
d'arthropodes.
Repartition. — Nouvelle-Caledonie, a 360 m de profondeur.
Melicerita (Henrimilnella) laurifolia sp. nov.
Fig. 12 H-I
MATERIEL EXAMINE. — Nouvelle-Caledonie. BlOCAL : sin DW 38, 360 m.
TYPE. — Holotype : Nouvelle-Caledonie. BlOCAL. stn DW 38, 360 m. (MNHN-BRY- 16443 bis).
Diagnose. — Melicerita (Henrimilnella) a large zoarium quadriserie, en forme de feuille, a extremites
arrondies et a cotes presque paralleles.
DESCRIPTION. — Les morceaux de zoarium etudies sont larges et aplatis, non ramifies, et ont des cotes
pratiquement paralleles et des extremites arrondies a large rayon de courbure ; leur extremite inferieure presente un
leger retrdcissement. Comme pour l'especc precedente, il n'est pas a exclure qu'il corresponde au point d insertion
d’un joint, et il serait done a priori impossible de preciser si le materiel decrit ici correspond a des portions isolees
d'un zoarium, ou a une colonie entiere ; mais la premiere hypothese semble peu probable, puisque les opercules
autozoeciaux, eux-memes chitineux, sont parfaitement conserves. Il n'existe pas d'aviculaire et aucune ovicclle n a
ete observee. Les series autozoeciales, alternantes, sont au nombre de 4 ; l'orifice hemicirculaire en est situe au
tiers de la longueur en partant de la region distale ; il presente une paire de condyles proximaux et une paire de
denticules distaux. Les autozoecies sont hexagonales ; celles appartenant aux deux series de la partie centrale du
zoarium sont longues de 0,41 a 0,44 mm, larges de 0,49 a 0,56 mm, et leur orifice, symetrique par rapport a l'axe
zoarial longitudinal, mesure 0,08 mm de long et 0.12 mm de large ; la serie marginale, de chaque cote, est formee
d'autozoecies de 0,39-0,41 mm de long et de 0,52-0.60 mm de large, dont l'orifice a les memes dimensions que
pour les autozoecies centrales mais est incline vers l'exterieur. La surface frontale est lisse et les limites
interzoeciales presque indiscemables.
DISCUSSION. — Cette espece se differencie de la precedente, dont elle partage la localite-type, par la largcur et
I'aplatissement dc ses elements, la quadriseriation dc son zoarium et la position tres anterieure dc lorilice
autozoecial.
ETYMOLOGIE. — Du Latin, laureus, de laurier, et folium , feuille, en apposition, chaque fragment ayant la
forme des outils prehistoriques connus sous le nom dc "feuilles de laurier .
Source :
204
J.-L. D'HQNDT & D. P. GORDON
Repartition. — Nouvclle-Caledonie, a 360 m dc profondeur.
CELLARIIDAE incertae sedis
Plusieurs specimens de Cellariidae figurant dans la collection etudiec etaient trop incomplets pour etre
dcterminables, meme au niveau generique ; l'un d'eux, trop erode pour pouvoir etre decrit, est simplemenl figure :
Biogeocal, stn DW 307, 470-480 m (Fig. 6 F ). Les autres feront ci-apres 1’objet d'une breve description :
1 - Biocal, stn DW 66
Fig. 12 E. 13 F-G
L'unique fragment de zoarium. non rami fie, est aplati et depourvu de joints. II presentc deux scries d'aulozoecics
sur chacune de ses faces, allernantes sur chaque face comme dos h dos. La longueur autozoeciale varie de 0,70 a
0,73 mm. la Iargeur a l'avant de 0,30 a 0,32 mm, a l'arriere de 0,20 a 0.24 mm. L'orifice hemicirculaire a
0,08 mm dc haut et 0,12 mm de large ; le bord proximal est droit ; il exisle deux condyles proximaux et deux
denticules distaux. non rcunis par unc lame calcairc. Le cote externe des autozoecies est rectiligne, le cote interne
formant a mi-longueur un angle vers 1'axe zoarial. II n'cxiste ni aviculaires ni ovicelles. La surface frontale est
finement et regulierement granuleuse.
Cet echantillon ressemble beaucoup a une Euginoma, dont il possede de nombreux caracteres. mais pas
suffisamment pour qu'il soil possible d'en faire une analyse complete. Les Euginoma ne presentent pas de
denticules aperturaux, ce qui les differencie de 1'espece etudiee ici. La forme du seul fragment possede evoque celle
rencontree chez les Henrimilnella (voir ci-dessus), identification generique que confirmerait la presence des deux
paires de denticules aperturaux, mais nous estimons preferable de ne pas donner de determination generique a partir
d'un seul et tres petit specimen.
2 - Biogeocal, stn CP 290
Le zoarium robuste a 1,2 mm de section. II presentc des autozoecies longues dc 0,96-1,20 mm et larges de
0.57-0,61 mm. La frontale est nettement depriniee par rapport aux limites interzoeciales ; le cadre frontal est peu
visible. L'orifice autozoecial est arrondi proximalement comme distalement et ne presente pas de condyles ;
l'opercule, cn forme de D, a 0,48 mm de long el 0,37 mm de large. L'ovicelle, long de 0.30 mm et large
de 0.50 mm. possede un pore en croissant large de 0,40 mm, d'une longueur totale de 0,16 mm (dont 0,10 mm
pour le pore seul). L'aviculaire rappelle beaucoup celui des Pseudorhyracella candelaber ; long de 0,60 mm et
large de 0,50 mm, il porte une mandibule de 0,60 mm de long et de 0,34 mm de large, arrondie a son
extremitd distale, et soutenue par un sclerite triangulaire. Perdu en cours de preparation, ce specimen n'a pu etre
photographic.
CLE ACTUALISEE DE DETERMINATION DES GENRES ET DES SOUS-GENRES
DE LA FAMILLE CELLARIIDAE
(N.B. - Les Henrimilnella , dont nous ignorons si les individus etudies correspondent a des zoariums complcts ou a
des colonies incompletes, sont considerees ici comme ayant un zoarium non articule).
1. Zoarium non articule . 6
— Zoarium presentant des entre-noeuds, separes soil par des fractures, des discontinuites. ou
des retrecissements des files zoeciales, soit par des joints chitineux : dans les cas ou la
ramilication n'est pas articulee, des branches adventives peuvent etre fixees par des
rhizoYdes sur certains entre-noeuds . 2
Source : MNHN, Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
205
2. Enlre-noeuds scpares par des joints chilineux ou des iractures ; zoarium cylindriquc rami lie
dichotomiquement a autozoecies alternantes . 3
— Entre-noeuds delimites par des discontinues ou des retrccissements dans les files zoe-
ciales ; parfois des ramifications latcrales adventivcs fixees par des faisceaux dc rhizoides ;
autozoecies non alternantes, etroitement accolees au meme niveau . 4
3. Joints marques par une simple fracture ; orifices autozoeciaux nettement plus longs que
larges ; bord proximal de l'orifice tres arquc ; petites zoecies aviculariennes quadrangulaires
a mandibule triangulaire . PARACELLARIA Moyano, 1969
— Entre-noeuds sdpares par des joints chitineux ; orifices autozoeciaux plus larges que longs,
ou dc memes dimensions ; bord proximal de l'orifice rectilignc ou a peine arquc ; joints
cylindriques de type tubuleux plus ou moins visibles .
. CELLARIA Ellis & Solander, 1786
4. Entre-noeuds scpares par des fractures ou des retrecissemcnts entoures ou non de tubes
chitineux . 5
— Entre-noeuds non separes par des retrccissements ou des fractures, mais certaines branches
pcuvent porter latcralement des ramifications adventives fixees par des rhizoides. Petit
aviculaire triangulaire a extremite arrondie, insere symctriquement par rapport a 1'axe
longitudinal de 1’autozoccie proximale ; orifice hemicirculaire, a deux condyles proximaux
et deux dents distalcs tres discretes . NEOCELLAR IAEFORMA d'Hondt. 1984
5. Orifices autozoeciaux plus ou moins quadrangulaires ; petits aviculaires interzoeciaux
obliques a mandibule hemicirculaire ; presence de deux condyles apcrluraux proximaux et
de deux dents distales . CELLAR IAEFORMA Rogick, 1956
— Orifices autozoeciaux dc forme variable ; gros aviculaires rhomboidaux interzoeciaux a
mandibule triangulaire ; presence dc deux condyles aperturaux proximaux relies par une
lame calcaire .... SWANOMIA Hayward & Thorpe, 1989 (= MAWSONIA Livingstone)
6. Zoarium (disjoint ?) constituc dc petits elements non ramifies, en batonnets ou ovalaircs ;
deux paires de denticulcs aperturaux, unc proximale et une distalc ; zoecies alternantes ;
orifice incline vers l'interieur sur les autozoecies marginales .
. HENRIMILNELLA subgn. nov.
— Zoarium ramifie cylindrique ou aplati ; condyles aperturaux et dents au nombre d une ou dc
deux paires, ou absents ; zoecies alternantes ou non ; orifices inclines ou non . 7
7. Ovicelles presentant des orifices lateraux en plus de l'orifice axial ; zoarium flabelliforme ;
autozoecies en files longitudinales alternantes ; aviculaires interzoeciaux spatuliformes ;
des condyles aperturaux proximaux . LA R VAPOR A Moyano, 1970
— Ovicelles ne presentant qu'un seul orifice, exceptionnellemcnt susceptible d'etre trilobe ;
zoarium cylindrique ou aplati ; autozoecies en files longitudinales alternantes ou en
couronnes radiaires ; aviculaires interzoeciaux variables, parfois absents, jamais
spatuliformes ; condyles aperturaux presents ou absents . 8
8. Base du zoarium formee par une succession de nodules alternant deux a deux, chacun
d’entre eux emettant une ou plusieurs branches zoariales ; orifice hemicirculaire ; termcn
peu saillant . DUBIOCELLARIA d'Hondt & Schopf, 1984
— Base du zoarium jamais formee d'une succession de nodules emettant chacun une ou
plusieurs branches ; orifice et developpcment du termen variables . 9
9. Orifice aulozoecial transversal, ovale ou en forme de croissant peu marque ; termen
formant des murailles saillantes . SVRINGOTREMA Harmer, 1926
— Orifice en forme de D ou de croissant bien marque ; termen gcneralement peu eleve (saul
chez les Atelestozoum) .
Source : MNHN. Pahs
206
J.-L. D’HONDT & D. P GORDON
10. Zoariuni aplati et bilaminaire, parfois foliace . 1 1
— Zoarium conique (certaines Euginoma) ou cylindrique, parfois un peu aplati mais dans
ce cas jamais completcment bilaminaire . 13
1 1 . Autozoecies planes, disposees en rangees transversales ; aviculaires vicariants ou absents ;
opesies presentant des incisions laterales de part et d'autre dune lame ; oviccllc formant un
epaississement en avant de 1'orifice . 1 2
— Autozoecies immergees au fond de profondes depressions delimitees par lc cadre du tcr-
men ; aviculaires absents ; opesie presentant un long processus proximal median ; ovicelle
inconnue . ATELESTOZOUM Harmcr, 1926
12. Pas d'aviculaires ; condyles fusionnes avcc une lame aperlurale proximale ; orifices
obliques sur tout le zoarium . PSEUDOCELLARIA Livingstone, 1928
— Des aviculaires vicariants, habiluellement portes sur les bords de la colonie ; des condyles
aperturaux proximaux et des denticules distaux ; orifices obliques sur les bords du zoarium
. ME LICE RITA H. Milne Edwards, 1836
13. Aviculaires et autozoecies de memes dimensions ; mandibules avicularienncs sensi-
blement hemicirculaires et a implantation mediane ; des condyles opesiaux proximaux ;
autozoecies disposees en spirale . STEGINOCELLARIA David & Pouyet. 1986
— Pas d'aviculaires ; denticules opesiaux presents ou absents . 14
14. Ovicelle non differenciee ; pas de denticules opesiaux ; pore ovicellien perforant la partie
distale de 1'autozoecie et separe d'elle par une Crete . EUGINOMA Jullien, 1882
— Ovicelle differenciee, soit distalement a 1'orifice comnie un bombement simulant un
bonnet, soit comme une heterozoecie en forme de pain de sucre reticule egale a au moins
la moitie de la surface d'une autozoecie nonnale . 15
15. Ovicelle en forme de pain de sucre reticule, egale h au moins la surface d'une autozoecie
nonnale ; pas de denticules opesiaux ; zoarium cylindrique ; autozoecies non alternantes
longitudinalement mais reunies en couronnes radiaires .
. FORMOSOCELLARIA d'Hondt, 1981
— Ovicelle apparaissant comme un bombement en forme de bonnet distalement a 1'orifice
autozoecial ; condyles opesiaux presents ou absents . 1 6
16. Autozoecies disposees de maniere spirale sur les branches zoariales . 17
— Autozoecies disposees en couronnes radiaires autour des branches zoariales ; condyles
opesiaux presents ou absents . CRYPTOSTOMARIA Canu & Bassler, 1927
17. Opesie a 1'extremite distale du cryptocyste ; orifice ovicellien etroit et transversal ;
denticules opesiaux presents ou absents .
. STOMHYPSELOSARIA Canu & Bassler, 1927
— Opesie au centre du cryptocyste ; orifice ovicellien large et transversal ; pas de denticules
opesiaux . MESOSTOMARIA Canu & Bassler, 1927
Remarque. — Cette cld n’inclut pas quatre genres exclusivement fossiles (les trois premiers du Cretace, le
quatrieme du Tertiaire) appartenant a la famille Cellariidae, lous les criteres diagnostiques utilisables n'y ayant pas
ete observes (denticules, pores ovicelliens, lame interdenticulaire, morphologie aviculariennc) :
- Dimorphocellaria Voigt, 1930 : zoarium articule, cylindrique ; series autozoeciales alternantes ; aviculaires
vicariants ; ovicelle endotochoidale ; existence de deux types autozocciaux, se caractdrisant par une forme differente
de 1'orifice (fun d’entre eux pouvant correspondre a des gonozoides).
- Hemistylus Voigt, 1928 : zoarium articule, cylindrique ; series autozoeciales alternantes ; orifices
autozoeciaux portes sur une seule des faces zoariales (comme chez les Euginoma).
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
207
- Escharicellaria Voigt, 1924 : zoarium bilaminaire dichotome inarticule ; aviculaires vicariants ; opesie en
forme de D allonge et presque hemicirculaire.
- Paramawsonia Androsova, 1972 : zoarium cylindrique, dont il n'est pas certain qu'il soit articulc, a zoecies
disposees en series longitudinales alternantes et aviculaires vicariants a rostre triangulaire. II n'est pas a exclure que
1'unique espece actuellement rangee dans ce genre, tel qu'il a ete dclimitc par Hayward et Ryland (1993), entre
en fait dans le genre Celia ria.
Embranchement ENTOPROCTA Nitsche, 1870
Famille LOXOKALYPODIDAE Emschermann, 1972
Genre LOXOKALYPUS Emschermann, 1972
ESPkCE-TYPE. — Loxokalypus socialis Emschermann, 1972.
Loxokalypus pedicellinoides sp. nov.
Fig. 2 A-C
MATERIEL EXAMINE. — Nouvelle-Caledonie. Chalcal 2 : stn DW 81.311 m.
Type. — Holotype : Nouvelle-Caledonie. Chalcal 2 : stn DW 81,311m (MNHN-BRY- 16706).
DIAGNOSE. — Loxokalypus a pedoncule long, fin et isodiametrique, 3 a 4 fois plus long que l'ensemble
calice-tentacules retractes.
Fig. 2. — Loxokalypus pedicellinoides : A, Zoarium. Echelle : 0,2 mm ; B. Calice. Echelle : 0,2 mm ; C. Detail de
l'omementation superficielle d'un pedoncule. Echelle : 0,1 mm.
Source :
208
J.-L. D'HONDT & D. P. GORDON
Description. — L'unique petite colonic recueillie est formee d'une douzaine de zoides separcs. issus d'une sole
basale commune ; elle ne comporte aucun stade de bourgeonncment de jeune individu. Cette sole a 0,1 I mm de
long et 0,10 mm de large. Les zoides ne presentent pas de cloison, ni entre le calice et le pedonculc. ni entre
celui-ci et la sole basale ; ils atteignent une hauteur de 0.95-1 mm. Les pedoncules, inscrcs cote a cote, ont un
diametre de 0,05 mm et une hauteur de 0,95-1 mm. Les tentacules, au nombre d'une huitaine. sont retractes a
I'interieur du calice chez tous les individus ; leur panache y a, selon les echantillons, une hauteur de 0,14
a 0,20 mm (0,29-0.32 mm avec les tentacules retractes). Ce calice, cupuli forme, a 0,10-0,1 1 mm de hauteur et
de diametre. L'etat de conservation du materiel ne permet pas d'en donncr une description plus complete.
Discussion. — Le genre Loxokalypus etait jusqu'a present monospecifique pour L. socialis, espece
uniquement connue des ties de la Reine Charlotte, sur la cote pacifique du Canada, dans la region dc Vancouver.
Les deux especes se differencient par la longueur et le diametre des pedoncules. Chez I’espece-type du genre, le
pedoncule est sensiblement de la meme longueur que Ic calice.
ETYMOLOGIE. — Du latin pedicelloides , cn raison de la ressemblance morphologiquc des individus de cettc
espece avec ceux des Pedicellina , autre genre d’Entoproctes (solitaires).
Repartition. — Nouvelle-Caledonie, a 31 1 m de profondcur.
CONCLUSIONS ET REMARQUES GENERALES
Les deux etudes que nous avons consacrees aux Bryozoaires Cheilostomes "Anascina" profonds, recoltes lors
des campagnes oceanographiques frangaises dans les parages de la Nouvelle-Caledonie (D’HONDT & Gordon,
1996; presente publication) ont permis un accroissement considerable de nos connaissances sur les faunes
balhyale et abyssale de cettc region. Les echantillons recueillis appartiennent a 25 families, dont une nouvelle pour
la science, a 53 genres dont six nouveaux (trois nouveaux sous-genres ont etc dcfinis a l'occasion de ce travail) et
90 especes dont 32 nouvelles (auxquelles il convient d'ajouter les quatre nouvelles especes collectees a faible
profondcur et decrites par D'HONDT, 1986) et dont 23 sont egalement presentes cn Nouvelle-Zelandc et / ou aux
lies Kermadec. Ce programme de recherche a permis en particulier d'ajouter a 1'inventaire faunistique local six
families, 26 genres et 43 especes d'"AnascinaM deja connues d'autres localites, notamment indo-pacifiques, mais
non encore signalees de Nouvelle-Caledonie.
Actuellement, letude systematique des Bryozoaires Cheilostomes Ascophorina des eaux profondes de Nouvelle-
Caledonie, cntreprise parallelement par les memes auteurs (Gordon, 1988, 1993 ; Gordon & Braga, 1994 ;
Gordon & D'HONDT. 1991) porte sur 37 families (dont deux nouvelles), 70 genres (dont 16 nouveaux et
quatre endemiques) et 124 especes (dont 79 nouvelles pour la science et 76 endemiques), auxquelles s'ajoutent les
1 1 especes nouvelles decrites par D’HONDT en 1986 de la zone Iittorale. Les genres Bifaxaria et Diplonotos sont
particulierement riches en especes. Ces donnees sur les Ascophorina seront ulterieurement a confronter a celles
actuellement reunies sur les "Anascina” et les Ctenostomes dans le cadre d'une synthese generate sur les
Bryozoaires Eurystomes neo-caledoniens.
En ce qui concerne les Bryozoaires Ctenostomes, les campagnes Musorstom ont permis d'ajouter a
1 invenlairc launistique local trois families, quatre genres et quatre especes des grandcs profondeurs, s'ajoutant a
celles, d eaux plus superficiellcs, signalees par D'HONDT en 1986 ; dans cettc derniere note, une nouvelle espece de
Bryozoaires Cyclostomes avait egalement ete nominee. Par ailleurs, une nouvelle espece de l'Embranchexnent des
Entoproctes, autrefois rapproche des Bryozoaires (ce qui fait que les specimens des deux groupes zoologiqucs
recoltes aux memes localites continuent a etre etudies conjointement par les specialistes) et non encore signale dc
Nouvelle-Caledonie, a ete identifiee lors de ce travail ; elle appartient a une famille (Loxokalypodidae) jusqu'alors
monogenerique et monospecifique, uniquement connue jusqu'a present de la region nord-ouest pacifique.
Plusieurs genres de Bryozoaires Eurystomes, trouves a proximite de la Nouvelle-Caledonie a l'occasion de ce
programme de recherche, n'etaient connus que de I'Atlantique : les Cheilostomes Nordgaardia et Formosocellaria ,
Source :
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
209
les Ctenostomes Aethozoon et Pachyzoon. Les especes de ces deux genres recueillies ici appartiennent cllcs-mcmcs
a la faune de l'ocean Atlantique, ce qui laisse esperer la decouverte ulterieure de localites intermediaires.
Un certain nombre de genres recoltes autour de la Nouvelle-Caledonie sont reputes cosmopolites, et leur
presence dans la faune de cette region cst parfaitement logique : Notoplites (9 especes et sous-especes recueillies
dans le cadre de ce programme), Cciberea, Comucopina, Camptoplites , Beania , Himantozoum , Membranipora. En
revanche, d'autres tres repandus dans l'lndo-Pacifique sont pauvrement represents en Nouvelle-Caledonie :
Dendrobeania , Menipea. Certains y presentent une grande diversite specifique : Himantozoum . Crateropora ,
Bryopastor. Par ailleurs, un genre fossile y a ete pour la premiere fois rctrouve a 1'etat vivant ( Pseudothyracella ).
Sont endemiques de cette region, dans 1’etat actuel de nos connaissances, les genres Candoscrupocellaria,
Candomenipea et Astoleiosalpinx definis par D'HONDT et GORDON (1996). Le nombre des espcces appartenant aux
genres Bryopastor et Crateropora est presque triple. Le genre Notoplites , represent par 1 1 especes dans les parages
de la Nouvelle-Caledonie, est celui qui y presente la plus grande diversite specifique.
Quelques-unes des especes figurant dans le materiel etudie ont une large distribution geographique. soit au
niveau mondial ( Bugulella gracilis : Irlande et Indo-Pacifique ; Scrupocellaria delilii el S. spatulata : toutes les
mers chaudes, et dans tous les cas a faible profondeur, inferieure a 100 m ) ; d'autres sont, soit uniquemenl connues
des regions australes ( Menipea patagonica a une profondeur inhabituelle, cf. D’HONDT & GORDON. 1996), soit
largement reparties dans rindo-Pacifiquc ( Euoplozoum cirratum , Notoplites scutatus , Canda clypeata , Caberea lata.
C. glabra , C. darwini , Amastigia rudis , Comucopina bella el Beania discodermiae , e’est-a-dire presque uniquemenl
des Ccllularines). n’etaient pas encore connues de la region neo-ealedonienne et les presentes recoltes permettent
d'elargir leur aire de distribution.
Si nous considerons les espcces recoltees en fonction de leurs distributions bathymetriques, les donnees peuvent
etre regroupees com me suit :
- de 0 a 100 m : 15 especes,
- de 101 a 500 m : 31 especes,
- de 50 1 a 1 000 m : 48 especes,
- de 1001 a 1500 m : 15 especes,
- de 1501 a 2000 m : 15 especes,
- de 2001 a 2500 m : 5 especes,
- de 2501 a 4000 m : 4 especes.
Ces donnees montrent que la diversite specifique est la plus riche entre 501 et 1000 m, un peu moindre de 101
a 500 m ; les recoltes entre 0 et 100 m etant tres peu nombreuses, nous ne considerons pas le nombre d'especes
recueillies dans cette tranche bathymetrique comme significatif. Lc nombre des especes est plus reduit en zone
bathyale entre 1000 et 2000 m, et semble tres faible en milieu abyssal, mais peut-etre cette faible diversite
specifique constatee n'est-elle que le reflet du faible nombre de dragages effectues a grande profondeur (trois entre
2001 et 2500 m, trois entre 2501 et 4000 m).
La plupart des especes etudiees dans ces deux travaux ont une bathymetric limitee a une ou deux des tranches
determinees ci-dessus. Toutefois une dizaine d'entre elles couvrent en Nouvelle-Caledonie trois ou quatre d'entre
clles : Crypto stomaria alata entre 0 et 1500 m ; Melicerita ejuncida , Nellia tenella et Bryopastor pentagonus entre
0 et 2000 m ; Pseudothyracella candelaber entre 101 et 1500 m ; Bnopastor challenged entre 101 et 2000 m ;
Formosocellaria magnifica entre 500 et 2000 m ; Lamourouxia canaliculata entre 500 et 2500 m ; Columnella
magna entre 1500 et 4000 m. Quatre especes ne sont presentes en Nouvelle-Caledonie que dans une tranche
bathymetric limitee, alors qu'elles sont connues dans l’lndo-Pacifique entre des Irontieres bathymetriques plus
importantes : Comucopina bella , C. moluccensis , Camptoplites lineatus , Euoplozoum cirratum.
Les 18 especes rcncontrees au-dela de 1000 m sont toutes arborescentcs ou en coussinet : huit Cellariidae,
trois Scrupocellariidae, deux Farciminariidae, une Chaperiidae, une Bugulidae, une Quadricellariidae,
une Calloporidae, une Pachyzoidae (emendation de Pachyzoontidae). Sept especes seulement ont, au cours de ce
programme, ete rencontrees a plus de 2000 m de profondeur : les Cheilostomes "Anascina" dresses Columnella
magna. espece typique de la faune abyssale (D’HONDT, 1981b), Lamourouxia canaliculata (nouvelle espece),
Penemia crassospina (genre de creation recente et a bathymetric peu etudiee) et Notoplites cassiduloides ,
Comucopina sp. et Dendrobeania pseudexilis (appartenant a des genres comportant deja plusieurs especes
210
J.-L. D'HONDT & D. P. GORDON
profondes), ainsi que le Ctenostome en coussinet Pachyzoon atlanticum. II convient d'ajouter a cettc listc
Formosocellaria magnified , recoltee vers 3500 m en mer de Tasmanie. lors d'une campagne du New Zealand
Oceanographic Institute.
Seules quelques especes sont communes aux faunes dc Nouvelle-Caledonie et des Ties Kermadec :
Crassimarginatella spathulata , Concertina cultrata , Bryopastor challenged. Melicerita ejuncida . Notoplites
longispinosus et Penemia ignota (?), qui ne sont connues que de ces deux regions geographiques, et Caberea
darwini , C. glabra , Notoplites obliquidens , Amastigia rudis , Comucopina geniculata , C. moluccensis , Bugulella
gracilis , Quadricellaria tenuis , Cellaria tenuirostris , Beania discodermiae, Alderina tuberosa ct Leiosalpinx
australis, dont la distribution geographique est plus etendue. Si 1'on excepte les especes presentes simultanement
en Atlantique et en Nouvelle-Caledonie sans localites intermediaires (. Aethozoon pellucidum , Pachyzoon
atlanticum et a un moindre degre Formosocellaria magnified), deux seulement des especes etudiees dans ces deux
travaux ont une ires large distribution geographique : l'espece abyssale Columnella magna, qui ne fait defaut que
dans les eaux polaires et sub-polaires, et une espece largement presente dans l'infra-littoral des eaux temperees et
chaudes, Nelli a tenella, dont les seules recoltes profondes ont ete effectuees en Nouvelle-Caledonie.
REMERCIEMENTS
Les photographies illustrant cette publication ont ete realisees, les uncs en Nouvelle-Zelande directement par
1'un dcs co-auteurs (D.P.G.) - grossissements indiques sur les pages de legendes les autres en France sur un
appareil different par le deuxieme auteur (J.L.D'H.) avec l’aide de Mme D. GUILLAUMIN (Centre d’Accueil de
Microscopic Elcctronique de I'Universite Pierre et Marie Curie - Paris VI) - echelles figurant sur les planches
photographiques - ; elles ont ete assemblies par Mme M.-J. D'HONDT qui a egalemcnt effectue les figures 1-2.
Nous leur adressons 1'expression de notre vive gratitude. Nous remercions sincerement Miss Patricia L. Cook
(Bendigo) et le Dr P.J. Hayward (Swansea) pour les informations qu'ils nous ont transmises et leurs remarques
constructives concemant ce travail.
Les recherches de l'un des auteurs (D.P.G.) ont ete financees par la New Zealand Foundation for Research,
Science & Technology (Contrat CO 1421).
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Source : MNHN. Paris
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PHOTOGRAPHIES
Source : MNHN. Paris
216
J.-L D’HONDT & D P. GORDON
FIGURE 3
A : Larnourouxia canaliculata , holotype (x 66).
B : Himantozoum crassiavicularium, holotype. Quelques autozoecies (area en place).
Echelle : 0,10 mm. a : aviculaire \fleche : orifice ovicellien.
C : Himantozoum crassiavicularium , holotype. Disposition des epines frontales. Echelle : 0.07 mm.
D : Callopora (?) sp. Quelques autozoecies. BiOGEOCAL, stn DW 253. Echelle : 0.10 mm.
E : Parantropora laguncula. Quelques autozoecies. Echelle : 0,10 mm.
F : Quadricellaria bocki. Ramification zoariale. Echelle : 0,10 mm.
G : Nellia tenella. Ramification zoariale. BiOGEOCAL, stn KG 275. Echelle : 0,10 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
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Source : MNHN. Paris
218
J.-L. D'HONDT & D. P GORDON
FIGURE 4
A : Carbasea laterogranulata , holotype. Portion de zoarium. Echelle : 1 mm.
B : Promicroa dubitata. Trois autozoecies et deux aviculaires (x 94).
C : Carbasea laterogranulata , holotype. Detail dc la region laterale dune autozoecic.
Echelle : 0,10 mm.
D : Bryopastor challengeri. Portion de zoarium. Echelle : 0,10 mm.
E : Columnella vipera , holotype. Ramification zoariale. Echelle : 0,10 mm.
F : Columnella niagna. Autozoecies ovicellees. Biocal, stn CP 13. Echelle : 1 mm.
G : Columnella magna. Autozoecies non ovicellees (vue frontale). BIOCAL, stn CP 13.
Echelle : 0, 10 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
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Source : MNHN. Pans
220
J.-L. D'HONDT & D. P. GORDON
FIGURE 5
A : Columnella vipera , holotype. Detail des aviculaires. Echelle : 0.10 mm.
B : Columnella vipera , holotype. Disposition des aviculaires. Echelle : 0.10 mm.
C : Bryopastor octogonos, holotype. Une autozoecie. Echelle : 0.10 mm.
D : Bryopastor octogonos, holotype. Portion de zoarium. Echelle : 1 mm.
E : Pseudothyracella candelaber, holotype. Portion de zoarium (erode). Echelle : 1 mm.
F : Calloporidae incertae sedis. Quelques autozoecies. BlOCAL, stn DW 44. Echelle : 0,10 mm.
G : Bryopastor pentagonus. Portion de zoarium. BlOCAL, stn DW 46. Echelle : 1 mm.
H : Pseudothyracella candelaber , holotype. Quelques autozoecies (erodees). Echelle : 0.10 mm.
I : Mesostoniaria strictoramae . Ornemcntation frontale dune autozoecie. BlOCAL. stn CP 52.
Echelle : 0,13 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOA1RES CHEILOSTOMIDA DE NOUVELLE-CALEDON1E
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Source : MNHN. Pahs
222
J.-L. D'HONDT & D. P GORDON
FIGURE 6
A : Bryopastor pentagonus. Quelqucs autozoecies. BlOCAL, stn DW 46. Echelle : 0, 10 mm.
B : Bryopastor pentagonus. Une autozoecie. BlOCAL. stn DW 46. Echelle : 0,10 mm.
C : Bryopastor eras sus, holotype. Une autozoecie. Echelle : 0.10 mm.
D : Bryopastor crassus, holotype. Fragment de zoarium. Echelle : 1 mm.
E : Bryopastor sp. Fragment de zoarium. BiOGEOCAL, stn DW 307. Echelle : I mm.
F : Cellariidae incertae sedis. Quelques autozoecies. BiOGEOCAL, stn DW 307.
Echelle : 0,10 mm.
G : Cryptostornaria alata. holotype. Portion dune ramification zoariale (x 27).
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
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Source : MNHN . Paris
224
J.-L. D'HONDT & D. P. GORDON
FIGURE 7
A : Smittipora fenestrata, holotypc. Portion dc zoarium. Echelle : 0,10 mm.
B : Smittipora fenestrata, holotype. Quclques zoccies. Echellc : 0,10 mm.
C : Smittipora fenestrata, holotypc. Quelques zoecies (area presente, aviculaires ouverts).
Echelle : 0,10 mm.
D : Smittipora fenestrata, holotype. Un aviculairc ferme. Echelle : 0,10 mm.
E : Promicroa dubitata, holotype. Fragment de zoarium (area presente). Echelle : 0,10 mm.
F : Smittipora fenestrata, holotype. Quelques autozoecies ct un aviculairc (opesie ovale) (x 76).
G : Micropora equilateralis, holotype. Quelques autozoecies. Echelle : 0,10 mm.
H : Celia ria humilis. Zoarium. MUSORSTOM 3, stn DR 117. Echelle : 1 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHE1LOSTOMIDA DE NOUVELLE-CALEDONIE
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Source : MNHN. Paris
226
J.-L. D HONDT & D. P GORDON
FIGURE 8
A : Lamourouxia canaliculata , holotypc. Portion de zoarium. Echelle : I mm.
B : Lamourouxia canaliculata, holotype. Region latcrale du zoarium, avec une ovicelle.
Echelle : 0,10 mm.
C : Lamourouxia canaliculata, holotype. Autozoecie ovicellee. Echelle : 0.10 mm.
D : Lamourouxia canaliculata. holotype. Autczoecies non oviccllees. Echelle : 0,10 mm.
E : Lamourouxia canaliculata. holotype. Groupe d'autozoecies (alveoles parietaux visibles).
Echelle : 0,10 mm.
F : Lamourouxia canaliculata, holotype. Bord crenele de l'opesie. Echelle : 0.010 mm.
G : Cellaria parafistulosa, holotype. Une autozoecie. Echelle : 0.10 mm.
H : Callopora (?) sp. Portion de zoarium. BlOGEOCAL, stn DW 253. Echelle : 0,10 mm.
I : Callopora (?) sp. Deux autozoecies. BlOGEOCAL, stn DW 253. Echelle : 0.10 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOM1DA DE NOUVELLE-CALEDONIE
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Source : MNHN. Paris
J.-L. D'HONDT & D. P. GORDON
FIGURE 9
A : Crateropora stiliformis , holotype. Portion dc zoarium. Echelle : 1 mm.
B : Crateropora stiliformis . holotype. Une autozoecie. Echelle : 0. 10 mm.
C : Crateropora stiliformis. holotype. Autozoecie ovicellec. Echelle : 0, 10 mm.
I) : Crateropora stiliformis , holotype. Portion de zoarium. Echelle : I mm.
E : Crateropora stiliformis , holotype. Autozoecies et aviculaires. Echelle : 0, 10
F : Micropora equilateralis , holotype (x 161 ).
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
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Source : MNHN. Pahs
230
J.-L. D'HONDT & D. P. GORDON
FIGURE 10
A : Cellar ia humilis . Une autozoecie. MUSORSTOM 3, stn DR 117. Echelle : 0,10 mm.
B : Mesostomaria strictoramae. Ramification zoariale. BlOCAL, stn CP 52. Echelle : 0,10 mm.
C : Formosocellaria magnified. Quelques autozoecies. Echelle : 0,10 mm.
D : Euginoma conica. Zoarium. Echelle : 0. 10 mm.
E : Syringotrema calobi , holotype. Portion de zoarium. Echelle : 0,10 mm.
F : Syringotrema calobi , holotype. Une autozoecie. Echelle : 0.10 mm.
G : Syringotrema calobi , holotype. Orifice autozoecial et ornementation frontale.
Echelle : 0,10 mm.
H : Melicerita ejuncida. Une autozoecie. Echelle : 0,10 mm.
I : Melicerita ejuncida. Portion de zoarium. Echelle : 0,10 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
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Source : MNHN. Pahs
232
J.-L. D'HONDT & D. P. GORDON
FIGURE 1 1
A : Mesostomaria sp. Portion de zoarium. Smib 3, sin DW 22. Echelle : 1 mm.
B : Mesostomaria sp. Detail de la photographic precedente. Echelle : 0.10 mm.
C : Mesostomaria sp. Autozoecie ovicellee. Smib 3, stn DW 22. Echelle : 0, 10 mm.
D : Mesostomaria sp. Autozoecie non ovicellee. Smib 3. stn DW 22. Echelle : 0, 10 mm.
E : Cryptostomaria alata . holotype. Ramification zoariale. Echelle : 1 mm.
F : Cryptostomaria alata , holotype. Une autozoecie de 1’axe du zoarium. Echelle : 0,10 mm.
G : Cryptostomaria alata, holotype. Autozoecie de la base d'une ramification. Echelle : 0,10 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
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Source : MNHN. Paris
234
J.-L. D HONDT & D. P. GORDON
FIGURE 12
A : Melicerita alternans , holotype. Zoarium. Echelle : 1 nun.
B : Melicerita alternans , holotype. Portion de zoarium. Echelle : 0.10 mm.
C : Melicerita alternans , holotype. Zoecies laterales. Echelle : 0. 10 mm.
D : Melicerita alternans , holotype. Une autozoecie ovicellee. Echelle : 0,10 mm.
E : Cellariidae incertae sedis. Detail de la surface frontale. BlOCAL, stn DW 66. Echelle : 0.01 mm.
F : Melicerita ( Henrimilnella ) articulata , holotype. Zoarium. Echelle : 1 mm.
G : Melicerita ( Henrimilnella ) articulata, holotype. Quelques autozoecies. Echelle : 0.10 mm.
H : Melicerita (Henrimilnella) laurifolia , holotype. Zoarium. Echelle : 1 mm.
I : Melicerita ( Henrimilnella ) laurifolia, holotype. Quelques autozoecies. Echelle : 0.10 mm.
J : Pseudothyracella candelaber, holotype. Portion de zoarium. Echelle : 10 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOM1DA DE NOUVELLE-CALEDONIE
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Source : MNHN. Paris
236
J.-L. D'HONDT & D. P GORDON
FIGURE 13
A : Melicerita alternans, holotype. Quelques autozoecies. Echelle : 0.10 mm.
B : Pseudothyracella candelaber , holotype. Portion de zoarium. Echelle : 0.10 mm.
C : Pseudothyracella candelaber , holotype. Une autozoecie. Echelle : 0.10 mm.
D : Pseudothyracella candelaber , holotype. Quelques autozoecies. Echelle : 0.10 mm.
E : Pseudothyracella candelaber , holotype. Ramification zoariale. Echelle : 1 mm.
F : Cellariidae incertae sedis. Zoarium. BlOCAL, stn DW 66. Echelle : 1 mm.
G : Cellariidae incertae sedis. Quelques autozoecies. BlOCAL, stn DW 66. Echelle : 0.10 mm.
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
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Source : MNHN. Paris
238
J.-L. D'HONDT & D F. GORDON
FIGURE 14
Pseudothyracella candelaber , holotype. Zoarium complet (x 2).
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
239
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J.-L. D’HONDT & D. P. GORDON
FIGURE 15
A : Cellaria obliquidens , holotype. Quelques autozoecies, l'une presentant un pore ovicellien (x 85).
B : Cellaria obliquidens , holotype. Orifice autozoecial surmonte d'un pore ovicellien (x 300).
C : Cellaria obliquidens , holotype. Entre-noeud (avec depart de deux ramifications) en vue apicale (x 81 ).
D : Lamourouxia canaliculata , holotype. Cavites alveolaires intra-parietales de la partie distalc de
1'autozoecie ; aviculaire distal (x 233).
E : Lamourouxia canaliculata , holotype. Ovicelle portanl 1'aviculaire distal (x 160).
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
241
Source : MNHN, Paris
wi-nv. vr wifihwi wv «■
242
J.-L. D’HONDT & D. P. GORDON
FIGURE 16
A : Cryptostomaria cilata. Quelques autozoecies. Biocal, CP 74 (x 50).
B : Cryptostomaria alata. Deux autozoecies, la plus proximale ovicellee. Musorstom 6, stn KG 465 (x 54).
C : Lamourouxia canaliculata , holotype. Interieur de la partie distalc dune aulozoccie ovicellee (x 190).
D : Lamourouxia canaliculata, holotype. Partie distale dune aulozoccie non ovicellee. surmontee
d'un aviculaire (x 380).
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMI D A DE NOU VELLE-C ALELX)N I E
243
Source : MNHN. Paris
244
J.-L. D’HONDT & D. P. GORDON
FIGURE 17
A : Cellaria fistulosa (Roscoff, pour comparaison). Portion de zoarium. avec un aviculaire
et quclques pores ovicelliens (x 54).
B : Cellaria fistulosa (Roscoff, pour comparaison). Quclques autozoecies avec pores ovicelliens ages.
Noter le relief du cadre autozoecial (x 125).
C : Cellaria fistulosa (Roscoff, pour comparaison). Aviculaire ct pore ovicellien immature,
a la limite des portions fertile et sterile de l'entre-noeud (x 195).
I) : Cellaria parafistulosa, holotype. Aviculaire et pore ovicellien age (x 46).
E : Cellaria parafistulosa , holotype. Portion de zoarium avec un aviculaire et des pores ovicelliens.
Noter le faible relief du cadre autozoecial (x 85).
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
245
Source : MNHN. Paris
246
J.-L. D’HONDT & D. P. GORDON
FIGURE 18
A : Pseudothyracella candelaber. NZOI (NIWA). Portion ramifiee du zoarium avec joints peletonnes
(x 12).
B : Pseudothyracella candelaber. NZOI (NIWA). Deux branches zoariales avec autozoecies, zoecies
femelles (tres large opesie) et aviculaires (x 13).
C : Pseudothyracella candelaber. NZOI (NIWA). Un aviculaire (noter la presence des processus
spiniformes proximaux) surmontant distalement une zoecie femelle (tres large opesie) (x 51 ).
D : Pseudothyracella candelaber. NZOI (NIWA). Un aviculaire (x 159).
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
247
Source : MNHN, Paris
248
J,L. D’HONDT & D. P. GORDON
FIGURE 19
A : Lamourouxia canaliculata , holotype. Cavites alveolaires de la paroi (x 137).
B : Pseudothyracella ccindelaber. Processus epineux proximaux submandibulaires d'un aviculairc.
NMNZ, stn U 582 (x 201).
C : Pseudothyracella candelaber. Mandibule avicularienne. NMNZ, stn U 582 (x 108).
D : Pseudothyracella candelaber. Bord interne du cryptocyste d'un zoide femelle. NMNZ, stn U 582
(x 214).
E : Micropora equilateralis , holotype. Une autozoecie ovicellee et un aviculairc (x 95).
Source : MNHN. Paris
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE 249
Source : MNHN. Paris
250
J.-L. D’HONDT & D. P. GORDON
FIGURE 20
A : Himantozoum crassiavicularium , holotype. Region distale dune autozoecie oviccllec. Echellc : 0.10 mm.
B : Himantozoum crassiavicularium, holotype. Region distale dune autozoecie non ovicellee.
Echellc : 0.10 mm.
C : Himantozoum crassiavicularium, holotype. Aviculaire d'une autozoecie laterale. Echclle : 0.010 mm.
D : Himantozoum crassiavicularium , holotype. Aviculaire d'une autozoecie axiale. Echclle : 0.10 mm.
E : Columnella viper a, holotype. Ovicelle. Echclle : 0.10 mm.
F : Calloporidae incertae sedis. Une autozoecie. Echelle : 0,10 mm.
G : Calloporidae incertae sedis. Ovicelle. Echelle : 0,10 mm.
H : Crateropora stiliformis, holotype. Cicatrice avicularienne. Echclle : 0,10 mm.
I : Crateropora stiliformis, holotype. Aviculaire cn place. Echelle : 0.10 mm.
J : Crateropora stiliformis , holotype. Region distale d'une autozoecie. Echelle : 0,10 mm.
K : Crateropora stiliformis , holotype. Ancestrula. Echelle : 0,10 mm.
ENTOPROCTES ET BRYOZOAIRES CHEILOSTOMIDA DE NOUVELLE-CALEDONIE
251
Source : MNHN. Paris
Source : MNHN. Paris
'SULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS D
Crustacea Isopoda: Bopyridae in the MUSORSTOM
collections from the tropical Indo-Pacific.
II. Species in subfamily Pseudioninae infesting
non-anomuran hosts
John C. MARKHAM
Arch Cape Marine Laboratory
Arch Cape. Oregon 97102-0105
USA
ABSTRACT
Gigantione petalomerae sp. nov. infests the dromiid crab Petalomera pulchra Miers in New Caledonia. Two species of
Pseudione show new host and geographic records: P. nephropsi Shiino. 1951. infests Metanephropsis veluiimts Chan &
Yu at Tanimbar Islands. Indonesia; P . elongata elongcita (Hansen. 1897) infests Nematocarcinus sp. in Chesterfield
Islands; both species are redescribed in detail. Pseudione tanimbarensis , sp. nov. infests Nephropsis sulcata Macpherson
at Tanimbar Islands. Indonesia. As a result of these redescriptions, the subspecies P. nephropsi atlantica Bourdon, 1971.
is considered a separate species, Pseudione atlantica Bourdon. 1971, and the variety P. elongata var. nonnalis Nierstrasz
& Brcnder a Brandis, 1931, is considered invalid.
RESUME
Crustacea Isopoda : Bopyridae des collections MUSORSTOM recoltes dans ITndo-Pacifit|ue
tropical. II. Especes de la sous-famille Pseudioninae infestant des hotes non anomuriens.
Gigantione petalomerae sp. nov. a 6t£ trouve sur le crabe dromiide Petalomera pulchra Miers en Nouvellc-Calcdonic.
De nouveaux hotes ct de nouvelles localisations gcographiques sont mentionnes pour deux especes de Pseudione :
P. nephropsi Shiino, 1951, trouv6e sur Metanephropsis velutinus Chan & Yu, aux Ties Tanimbar, en Indonesie et
P. elongata elongata (Hansen, 1897) sur Nematocarcinus sp. aux Ties Chesterfield; ces deux esp&ces sont red£crites en
detail. Pseudione tanimbarensis sp. nov. infeste Nephropsis sulcata Macpherson aux lies Tanimbar. Des consequences dc
ces redescriptions sont l'elevation au rang d'espece de la sous-espfcce P. nephropsi atlantica Bourdon. 1971. tandis que
la varidte P. elongata var. normalis Nierstrasz & Brender a Brandis, 1931, est consideree comme non valide.
Markham, J. C., 1999. — Crustacea Isopoda: Bopyridae in the Musorstom collections from the tropical Indo-
Pacific. II. Species in subfamily Pseudioninae infesting non-anomuran hosts. In: A. Crosnier (ed.), Resultats des
Campagnes Musorstom, Volume 20. Memoires du Museum national d'Histoire naturelle, 180: 253-265. Paris ISBN
2-85653-520-8.
Source : MNHN, Pans
254
J.C. MARKHAM
The extensive collections of ORSTOM. housed in the Museum national d'Histoire naturellc (designated
MNHN), contain many uncatalogued and unidentified bopyrid isopods, parasites of numerous different decapod
crustaceans. This report, one of a continuing series, deals with a small part of that material.
FAMILY BOPYRIDAE Rafinesque
SUBFAMILY PSEUDIONINAE Codreanu
Gigantione petalomerae sp. nov.
Fig. 1
“Bopyrid." — McLay, 1993: 166 [Chesterfield Islands; infesting Petalomera pulchra Miers].
Material EXAMINED. — Chesterfield Islands. Chalcal 1: stn. DC 53, 2I°19.5'S. I58°55.3'E, 60 m,
24.07.1984. Infesting Petalomera pulchra Miers, host det. C. L. McLay: I 9, holotype; 1 6 , allotype (MNHN Ep-888).
Description. — Holotype female (Fig. I A-I): Length 6.9 mm, maximal width 6.8 mm. head length 1.8 mm,
head width 2.4 mm, pleon length 2.8 mm. Body outline nearly circular, body axis distorted only 18°; all body
regions and segments distinct (Fig. I A).
Head subrectangular, about 2/3 embedded in pcrcon. Short frontal lamina not quite extended to sides of head;
small anterolateral Haps projecting slightly form each side of head. Dorsal surface produced into 2 large low lobes.
Antennae fairly large, extending beyond anterior margins of head, of 3 and 5 articles respectively. Barbula
(Fig. IB) with single terete projection on each side and triangular lobe in center. Maxilliped (Fig. 1C) suboval, its
anterior article much larger than posterior one and marginally fringed by dense setae.
Pereon largely enclosing both head and pleon, rounded on both sides but bulging outward slightly more on
long side, broadest across pereomere 4. Coxal plates well-developed on both sides of all pereomeres, those on long
sides very long and slender and reflexed over dorsal surfaces of pereomeres; those opposite somewhat broader and
angled to extend forward along sides of pereomeres. Indistinct dorsolateral bosses on long sides of some
pereomeres. Oostegitcs almost completely enclosing brood pouch, but not tightly pressed onto each other.
Oostegite 1 (Fig. ID-E) with much larger anterior plate, deep groove separating it from posterior region
externally; internal ridge bearing 5 long slender flexible projections along lateral half, nothing on medial half.
Pereopods 1-4 on long side visible dorsally along side of body, others hidden in dorsal view, all of about same size
(Fig. 1F-G) and with all articles present; pereopod 7 much more setose, its basis produced into prominent lobe
along flexor margin.
Pleon of 6 distinct pleomeres, each produced into lateral plates on both sides; lateral plates on longer sides
basally constricted into falcate points, lanceolate ones opposite extending out from slender sides of pleomeres
without constriction. Sixth pleomeres also produced into slender lateral plates. All pleonal appendages visible only
in posterior view (Fig. IH): 5 pairs of biramous pleopods (Fig. II) with deeply digitate margins; they and minute
biramous uropods tightly clustered inside cavity formed by lateral plates.
Allotype male (Fig. 1J-N): Length 4.7 mm, maximal width 1.7 mm, head length 0.7 mm, head width
1.1 mm. pleonal length 1.4 mm. All body regions and segments distinctly separated. Minute splotchy eyes near
posterolateral corners of head; tiny scattered pigment spots on dorsal surfaces of some pereomeres and of first
pleomere (Fig. 1J-K).
Head trapezoidal, broadest at posterior margin but narrower than pereomere I. Antennae (Fig. 1L) of 3 and
5 articles, respectively; antenna 1 quite reduced, but antenna 2 extending beyond margins of head on both sides.
Pereon broadest across pereomere 5, tapering slightly both ways from there. Pereopods (Fig. 1M-N) all equally
developed and about same size, with all articles distinct; propodi somewhat larger and dactyli sharper posteriorly.
Pereopods moderately covering much of ventral surface of pereon without extending beyond sides.
BOPYRIDAE OF THE TROPICAL INDO-PACIFIC
255
Pleon (Fig. IN) of 6 distinct pleomercs, tapering slightly posteriorly; margins sparsely setose. Five pairs of
irregularly lengthened uniramous flaplike pleopods, progressively smaller posteriorly. Pair of large uniramous
uropods extending prominently rearward at end of body.
Fig. 1. — Gigantione petalomerae, sp. nov., A-l. holotype female; J-N, allotype male. A. dorsal view. B. barbula. right
side. C, right maxilliped. D. right ooslegite 1. external view. E. same, internal view. F. right pereopod LG. right
pereopod 7. H. pleon, ventral view. I, right plcopod 2. J. dorsal view. K. ventral view. L. left antennae. M, lett
pereopod 1. N, right pereopod 7 and pleon in ventral view.
Scale: 2.5 mm for B-F. I; 5.0 mm for A, H. J-L; 10.0 mm for M-N; 16.7 mm for F-G.
ETYMOLOGY. — Specific name petalomerae , genitive singular of generic name of the host of the new species.
DISCUSSION. — The genus Gigantione contains 12 previously described species, as discussed by MARKHAM
(1994). Three of the species are parasites of thalassinideans. the others infesting diverse brachyurans. For now. I
am retaining the genus in the subfamily Pseudioninae, although some of its characters indicate that its placement
in the Ioninae, the typical parasites of the Brachyura, might be more appropriate. Regardless, it belongs among the
least differentiated bopyrid genera. The new species, G. petalomerae , is most similar to G. mortenseni Adkison
(1984), from the Gulf of Mexico and eastern Florida, U. S. A., w'hich is, interestingly, the only other bopyrid
species knowm to infest recent dromiid crabs. The diagnostic differences between these 2 species are that the female
of G. mortenseni has a more elaborate barbula, a proportionately longer first oostegite and coxal plates more nearly
alike on opposite sides; its male has a more extended head, a rather sharply tapered pleon, pleopods and uropods
more distinctly bilobate and the latter more extended. In presenting a new generic diagnosis of Gigantione
Source :
256
J.C. MARKHAM
(Markham. 1994), I did not mention the structure of the seventh percopods of the females, whose bases are
characteristically expanded into lobes along the flexor margins, as seen also in G. petalomerci (Fig. 1G).
The type-specimens of G. petalomercie were first recorded without identification in the report on its host,
Petalomerci pulchra , by McLay (1993), who also mentioned the presence of an unidentifiable larva attached to the
abdomen of Dromia wilsoni (Fulton & Grant). The only other recent dromiids reported to be bopyrid hosts are
those bearing G. mortenseni (see ADKISON. 1984): Dromidia antillensis Stimpson, Hypoconcha sabulosa (Herbst)
and H. spinosissima Rathbun. Housa (1963) published a record of evident bopyrid infestation of 2 Mesozoic
fossil dromiid species from Moravia, so, w-hile their parasites, lost in fossilization, remain unidentifiable, it is
evident that bopyrid infestation of dromiids is among the most ancient known.
Fig. 2. — Pseudione nephropsi Shiino, 1951, reference female. A, dorsal view. B. ventral view. C. right side of barbula.
D, right maxilliped. E, palp of same. F. right oostegite 1, external view. G, same, internal view. H. right pereopod
1. I, right pereopod 7. J, pleon in ventral view.
Scale: 1.8 mm for E, H-I; 5.4 mm for C-D; 5.9 mm for J: 10.8 mm for A-B, F-G.
Source :
BOFYRIDAE OF THE TROPICAL INDO-PACIFIC
257
Pseudione nephropsi Shiino, 1951
Figs 2-3
Pseudione nephropsi Shiino, 1951: 32-36; figs 5-6 [Type-locality Ovvase, Mie Prefecture, Japan; infesting Nephrops
japonicus Tapparone-Canefri]; 1952: 34-35, 41-42; 1972: 7. — Bourdon, 1968: 216; 1971: 375. — §adoglu,
1969: 197. — H0EG & Rybakov, 1992: 601, table 1.
MATERIAL EXAMINED. — Indonesia. Karubar: st. CC 41. Tanimbar Island, 07°45*S, 132°42’E, 401-393 m,
28.10.1991. Infesting Metanephrops ve I minus Chan & Yu: 1 9 , 1 8 (MNHN-Ep 889).
Discussion. — This is only the second
record of Pseudione nephropsi , so the host
(both species and genus) and locality records
are new. Both sexes display characters
distinctive for this species, as described by
Shiino (1951): in the female, the shape of
the head and body (Fig. 2A-B), the peculiar
short row of a few tiny processes on the
internal ridge of the first oostegite (Fig. 2G),
and the posterolaterally extended pleonal
appendages bearing prominent tubercles on
their ventral surfaces (Fig. 2J); and, in the
male, the very long slender body (Fig. 3A-B),
the proportionately small pereopods (Fig. 3D-
E). and the extended pleon produced into a
bilobate posterior margin (Fig. 3F). Some
differences between the new material and the
types are noteworthy. In the type-female, the
palp articulates with the maxilliped, the first
oostegite is less sharply pointed, and there are
no midventral depressions on pleomeres 1-3.
The type-male has all 6 pleomeres completely
distinct and discernible rudiments of some
plcopods.
Bourdon (1971) described the subspecies
Pseudione nephropsi atlantica as a parasite of
Nephropsis atlantica (Norman) from the
Congo. In presenting a tabulation of the
characters by which his new subspecies
Fig. 3. — Pseudione nephropsi Shiino, 1951. reference male.
A, dorsal view. B, ventral view. C, right antennae. D, right
pereopod 1. E, left pereopod 7. F. pleon in ventral view.
Scale: 2.6 mm for C-F; 5.0 mm for A-B.
differed from the typical one, BOURDON (1971) observed that it possibly deserved separate species status. In
particular, the lack of tubercles on the pleonal appendages of the female sets it off enough for such status.
Accordingly, I am here considering Pseudione atlantica Bourdon. 1971, a separate species and thus not using any
subspecific designation for the new material of Pseudione nephropsi.
Pseudione tanimbarensis sp. nov.
Figs 4-6
MATERIAL EXAMINED. — Indonesia. Karubar: st. CP 81. Tanimbar Islands, 09°34-54”S, l31°0r49"E, 199-
206 m, 4.11.1991. Infesting Nephropsis sulcata Macpherson. host det. T.-Y. Chan: 1 9, holotype; 1 6 (damaged),
allotype (MNHN-Ep 891).
Source :
258
J. C. MARKHAM
Fig. 4. — Pseudione tanimbarensis sp. nov., holotype female. A, dorsal view. B, ventral view. C, left side of barbula.
D. left maxilliped, external view. E. left oostegite 1. external view with pereopod attached. F. same, internal view.
G. right pereopod 7. Scale: 2.5 mm.
Description. — Holotype female (Figs 4-5). Length 5.4 mm, maximal width 3.4 mm, head length 1.1 mm,
head width 1.4 mm, pleon length 2.0 mm. Body outline elongate-oval, body axis distorted 74°; all body regions
and segments distinct (Fig. 4A-B).
Head suboval, deeply set into pereon. Fully developed frontal lamina completely covering front of head, but not
extending beyond its sides. Antennae reduced, only antennae 2 reaching margins of head, of 3 and 8 articles
respectively (Fig. 5A-B). Barbula (Fig. 5C) with small angled outer projection and broadly lobate inner projection
on each side and irregularly lobed margins in center. Maxilliped (Fig. 5D) subtriangular, its anterior article much
larger then posterior one; nonsegmented articulating palp (Fig. 5E) just lateral to anteromedial corner of maxilliped
and barely extending beyond its front, its margin and adjacent corner of maxilliped bearing sparse long setae;
pleclron (Fig. 5F) acutely angled but not prominent, its tip bearing minute setae. No eyes visible.
Pereon surrounding head laterally, doubly oval, tapering both ways from pereomere 3, then abruptly broader
across pereomere 5 and tapering from there through pleon. Small coxal plates well-developed on both sides of
pereomeres 1-4, those on long sides prominent and laterally extended. Dorsolateral projections on both sides of
pereomeres 1-3 and prominent on long sides of pereomeres 5 and 6. Oostegilcs completely enclosing brood pouch.
Oostegite 1 (Fig. 5G-H) with anterior and posterior plates equally long, latter slightly concave posteriorly;
internal ridge entire except for some tiny lobes near center. Pereopods extending laterally and visible dorsally along
sides of body, all with all articles distinct and bases lobate, their bases and ischia progressively larger posteriorly
(Fig. 5I-J).
Source :
BOPYRIDAE OF THE TROPICAL INDO-PACIFIC
259
Fig. 5. — Pseudione tanimbarensis sp. nov., holotype female. A, right antenna 1. B. left antenna 2. C, barbula. D, right
maxilliped. E, palp of same. F, plectron of same. G, right oostegite 1, external view. H. same, internal view. I. left
pereopod 1. J, left pereopod 7.
Scale: 1.2 mm for C; 1.0 mm for D, G-H; 0.4 mm for others.
Pleon greatly extended, of 6 distinct broad, anteriorly convex pleomeres; pleomeres 1-5 each produced into long
slender lateral plates on both sides, deeply separated by basal constriction on long sides but slightly overlapping
opposite. Five pairs of biramous pleopods, their endopodites lanceolate and reaching far laterally, their endopodites
falcately curved and extending rearward. Pleomere 6 heart-shape, with tiny posterior anal cone and lanceolate
uniramous pleopods similar to but larger than lateral plates and extending far posteriorly.
Allotype male (Fig. 6): Length 3-4 mm (uncertain because of damage), maximal width 0.9 mm, head length
0.4 mm, head width 0.7 mm, pleonal length unknown. All body regions and segments distinctly separated. Sides
of body nearly parallel (Fig. 6A-B).
Head suboval, broadest just before posterior margin and narrower than pereon. Antennae (Fig. 6C) of 3 and
6 articles, respectively; antennae 2 extending beyond margins of head on both sides. No eyes.
Pereon broadest across pereomere 3, but only slightly so; all pereomeres deeply separated by anterolateral
indentations. Pereopods (Fig. 6D-F) all equally developed, decreasing slightly in overall size and markedly in
dactylus size posteriorly; all carpi and meri fused, other articles distinct. Pereopods moderately covering much of
ventral surface of pereon without extending beyond sides.
Pleon of at least 3 distinct pleomeres, total number uncertain because of missing posterior region, pleon
tapering slightly posteriorly. Evidently no pleopods or midventral projections. Status of uropods unknown.
Source :
260
J.C. MARKHAM
Fig. 6. — Pseudione tanimbarensis sp. nov., allotype male. A, dorsal view. B. ventral view. C. right antennae. D. right
pereopod 1. E, right percopod 3. F, right pereopod 7.
Scale: 1.2 mm for A-B; 0.2 mm for C-F.
ETYMOLOGY. — Specific name tanimbarensis , derived from name of type locality, Tanimbar Islands.
DISCUSSION. — Pseudione tanimbarensis belongs to that small group of species infesting hosts in the family
Nephropidae, in host selection as well as morphology and probable lineage. It would be considered a third
subspecies of Pseudione nephropsi Shiino, 1951, considered above, if I were not raising the second subspecies to
specific rank as Pseudione atlantica Bourdon, 1971. Characters that Pseudione tanimbarensis shares with those two
species are, in the female: body much longer than wide; sides of head completely embedded in pereon, with frontal
lamina across whole anterior but not along sides at all; no eyes; palp on or next to anteromedial corner of
maxilliped and extending little or none beyond it; 2 pairs of lateral projections on barbula, though inner pair much
reduced; first oostegite with only short row of small lobes on internal ridge and falcately pointed posterolateral
projection; brood pouch completely enclosed; meri and carpi of pereopods fused; pleon extended far posteriorly;
pleomeres deeply separated laterally and produced into extended slender lateral plates; first pleomcrc wider than last
pereomere; reduced sixth pleomere heart-shape, plainly visible dorsally; large pleopodal endopodites curved to
extend rearward. Males of the three species share these characters: body long and slender; head suboval, much
broader than long, lacking eyes; antennae 2 extending beyond margins of head; pcreomcres and pleomeres deeply
separated laterally; first pereopods larger and with more sharply pointed dactyli than others. The female of
Pseudione tanimbarensis differs from the other two species in being much more distorted and having its coxal
plates placed laterally, not dorsally, and in having all pleopodal endopodites about the same size as their respective
Source :
BOPYRIDAE OF THE TROPICAL INDO-PACIFIC
261
endopodites (rather than being larger in P. nephropsi and smaller in P. atlantica) and differently shaped. It differs
further from P. nephropsi in having no tubercles on its pleonal appendages. The female of Pseudione
tanimbarensis differs further from that of P. atlantica in having narrower lateral projections on its barbula,
relatively shorter first oostegite and a less prominent basal carina on its seventh pereopod. The male of Pseudione
tanimbarensis differs from that of P. nephropsi by lacking pleopods, and from that of P. atlantica by having its
head completely separated from the pereon.
DISCUSSION. — This discovery of infestation of Nephropsis sulcata Macpherson brings the number of
nephropid species known to harbor bopyrid parasites to six. Pseudione nephropsi , discussed above, infests
Nephrops japonicus Tapparone-Cancfri in Japan and Metanephrops velutinus Chan & Yu north ol Tanimbar
Islands, Indonesia; P. atlantica infests Nephropsis atlantica (Norman) near the Congo. Two other species of
nephropids have been reported as bopyrid hosts, but their parasites were never named. De Man (1916) cited
infestation of Nephrops andamanicus Wood Mason near Tuvalu in the South Pacific, and BOUVIER (1925) recorded
“un Bopyrien" from Nephropsis aculeata Smith, a junior synonym of N. rosea Bate (L. B. HOLTHUIS, personal
communication), near Grenada, in the Caribbean.
Pseudione elongata elongata (Hansen. 1897)
Fig. 7
IBopyrus - Bate, 1888: 804 [Off Sibago. Philippines; infesting Nematocarcinus undulatipes Bate).
"Bopyrid" - Faxon, 1895: 159.
Cryptione elongata Hansen, 1897: 112-115. pi. 3, figs 5-5a, pi. 4. figs 1-lg [Galapagos Islands at 1618 m; infesting
Nematocarcinus agassizi Faxon]. — RICHARDSON. 1889a: 869; 1899b: 338: 1904: 87; 1905: 520-522; ligs 567-
568; 1910: 36. — Bonnier, 1900: 48. 61. 160. 221. 285-287. 332, fig. 51. — Richard. 1900: 71. — Stebbing.
1914; 4g _ van Name, 1924: 185. — Danforth, 1963: 44; 1970: 462. — SCHULTZ, 1969: 320-321, fig. 511. —
Rioja, 1971: 511. — Brusca, 1987: 273, table 1.
Pseudione elongata - NlERSTRASZ & BRENDER h BRANDIS, 1923: 72, 78; 1931: 163. — CHOPRA, 1930: 139. — SHllNO,
1951: 32; 1952: 42. — KENSLEY, 1968: 190-191. — PAGE, 1985: 192.
Pseudione elongata var. normalis NlERSTRASZ & Brender & Brandis, 1931: 163-164, fig. 29 [South of Zamboanga,
Philippines; infesting Nematocarcinus sp.]. — Shiino, 1951: 32. — KENSLEY, 1968: 190-191. — Page, 1985: 195.
Material EXAMINED. — Chesterfield Islands. MUSORSTOM 5: st. CC 384. 19°42.40'S, 158°50.80'E. 772-
776 m, 21.10.1986. Infesting Nematocarcinus sp.: 13 9, 16 8 (of which 2 immature), 1 cryptoniscan larva (MNHN-
Ep 890).
Descriptive Notes. — Female body outline and proportions (Fig. 7A-B) as in type, as well as configuration
of barbula (Fig. 7E), shape of maxilliped (Fig. 7F), first oostegite (Fig. 71). posterior dorsal margins of
pereomeres, proportions of articles of pereopods (Fig. 7K-L) and orientation of pleonal appendages. Female s
frontal lamina separated as in normalis form, maxilliped palp (Fig. 7G) shorter, internal ridge of oostegite 1
(Fig. 7J) more elaborate than in type. One of other females with maxilliped (not drawn) lacking projection next to
palp as in female drawn, so more like that of type. One female with knobs on pleopods 1 and 2 (not drawn); two
other females (Fig. 7M-N) with uropods more like those of holotype than one drawn in detail.
Male (Fig. 70-P) proportionately slightly broader than type male and with head separated as in normalis form;
matching type-male in general body shape and head shape, size and structure of antennae (Fig. 7Q-R), pereopods
(Fig. 7S-T), midventral tubercles (Fig. 7P) and pleonal appendages (Fig. 7P, U). Immature male (Fig. 7V-W) with
pereopods proportionately longer, midventral tubercles absent and pleopods greatly reduced.
One of the hosts (Fig. 7X) had an anterior opening into the swelling of the branchiostegite revealing the female
inside. In the lot examined, there were 15 infested specimens of Nematocarcinus sp. Ot these, 8 were inlested
dextrally and 7 sinistrally. Two of the parasites were missing; of the 13 females present, each was accompanied by
at least 1 male, 2 mature males were with 1 female, 1 mature and 2 immature males were with another female, and
1 mature male and a cryptoniscan larva were with yet another female.
Source :
262
J.C. MARKHAM
Fig. 7. — Pseudione elongata elongata (Hansen, 1897). A-L, reference female; M. second female; N, third female; O-U,
reference male; V-W, immature male; X, fourth female on host Nematocarcinus sp. A, dorsal view. B, ventral view.
C, right antenna 1. D, left antenna 2. E, right side of barhula. F. right maxilliped. G. palp of same. H, plcctron of
same. I, right oostegite 1, external view. J, same, internal view. K. right pereopod 1. L, right pereopod 7. M, end of
pleon in dorsal view. N, same. O, dorsal view. P. ventral view. Q. right antenna 1. R. right antenna 2. S, left
pereopod 1. T, left pereopod 7. U, end of pleon. ventral view. V, same, dorsal view. YV, ventral view. X, parasite in
host’s branchial chamber.
Scale: 0.2 mm for Q-R; 0.4 mm for S-U; 1.2 mm for C-D. K-L, V-W; 2.5 mm for G-H, O-P; 5.0 mm for A-B, E-F,
I-J, M-N; 10.0 mm for X.
Systematic Remarks. — The specimens mentioned by Bate (1888) seem never to have been identified,
though their locality and host make it likely that they represented P. elongata. Faxon (1895) cited the specimens
subsequently described as the types of P. elongata. Hansen (1897) provided a very detailed description of this
species, based on a single pair. He placed it in the new genus Cryptione, for which he gave no diagnosis, stating
only that “...it is necessary to institute ... new genera. - a result with which I am rather dissatisfied, not being
sure that they will all prove valid.” NlERSTRASZ & Brender a BRANDIS (1923) synonymized Cryptione with
Pseudione , with which action I agree, though most subsequent authors disregarded this generic transfer.
NlERSTRASZ & BRENDER a Brandis (1931) described the new variety, Pseudione elongata var. normal is, which
they considered to differ from the typical form mainly because its male had the head separated from its first
pereomere, as is typical in the genus Pseudione ; I am disregarding that varietal name, while noting that all of the
males examined here also show such a separation. RIOJA (1971), in a catalog of species from Latin America, said
that P . elongata was known from Acapulco, on Mexico's Pacific coast, but he cited no source for that record, nor
do I know of any. BRUSCA (1987), in a list of the fauna of the Galapagos Islands, called it endemic to that
Source : MNHN. Pans
BOPYRIDAE OF THE TROPICAL INDO-PACIFIC
263
archipelago, thus overlooking the record of NlERSTRASZ & BRENDER a BRANDIS ( 1931 ) from the Philippines. And
in an account of the Bopyridae of the eastern Pacific (MARKHAM, 1992). I accidentally omitted it despite its
occurrence in the Galapagos. KENSLEY (1968) described the new subspecies Pseudione elongata africana , a parasite
of Nematocarcinus longirostris Bate off southwestern South Africa. Thus this account deals with the typical
subspecies, though this is the first time it has been so cited.
AC KNOWLEDGM ENTS
Sincerest thanks are extended to Alain CROSNIER for sorting the ORSTOM collections and putting them at my
disposal, and for arranging and hosting my visit to the Museum national d'Histoire Naturclle and to Bertrand
Richer de Forges who collected many of the specimens. The financial support of the Institut Fran^ais de
Recherche Scientifique pour le Developpcment cn Cooperation (ORSTOM) is gratefully acknowledged. Colin
McLay provided information about the host of Gigantione petalomerae sp. nov. An anonymous reviewer provided
valuable suggestions that led to improvements in the manuscript. The late Mrs W. A. Markham provided
facilities at the Arch Cape Marine Laboratory, of which this is publication number 27.
LITERATURE CITED
Adkison, D. L., 1984. — Two new species of Gigantione Kossmann (Isopoda: Epicaridea: Bopyridae) from the western
North Atlantic. Proceedings of the Biological Society of Washington. 97: 761-772.
Bate, C. S., 1888. — Report on the Crustacea Macrura dredged by H. M. S. Challenger during the years 1873-1876.
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Bonnier. J., 1900. — Contribution a I’etudc des epicarides. Les Bopyridae. Travaux de la Station biologique de
Wimereux, 8: 1-476.
Bourdon, R., 1968. — Les Bopyridae des mers europeennes. Memoires du Museum national d'Histoire naturelle ,
nouvelle serie (A). 50 (2): 77-424.
BOURDON, R.. 1971. — Epicarides nouveaux pour la cote occidcntale d'Afrique £quatoriale. Bulletin de V Institut frangais
d’Afrique Noire , (A), 33: 371-391.
Bouvier. E. L., 1925. — Les macroures marcheurs. Reports on the results of dredging, under the supervision of Alexander
Agassiz, in the Gulf of Mexico (1877-78). in the Caribbean .Sea (1878-79). and along the Atlantic coast of the United
States (1880), by the U. S. Coast Survey Steamer "Blake.", Lieut. -Com. C.D. Sigsbee, U.S.N., and Commander
J.R. Bartlett. U.S.N., commanding. Memoirs of the Museum of Comparative Zoology, Harvard College, 47 (5): 399-
472, 11 pis.
Brusca, R. C.. 1987. — Biogeographic relationships of Galapagos marine isopod crustaceans. Bulletin of Marine
Science, 41 (2): 268-281.
Chopra. B., 1930. — Further notes on bopyrid isopods parasitic on Indian Decapoda Macrura. Records of the Indian
Museum, 32 (2): 113-147, pis 4-6.
Danforth, C. G., 1963. — First record of a Hawaiian shore bopyrid (Isopoda: Bopyridae). Journal of Parasitology , 49
(5): 847-850.
DANFORTH. C. G., 1970. — Epicaridea (Isopoda) of Hawaii. Bulletin of the Southern California Academy of Sciences, 69:
27-31.
DeMan, J. G., 1916. — The Decapoda of the Siboga Expedition. Part III. Families Eryonidae, Palinuridac. Scyllaridae
and Nephopsidae. Siboga-Expeditie Monograph, 39a2: 1-122, 4 pis.
Faxon, W.. 1895. — Reports on an exploration off the west coasts of Mexico. Central and South America, and off the
Galapagos, in charge of Alexander Agassiz, by the U. S. Fish Commission Steamer "Albatross”, during 1891, Lieut. -
Commander Z.L. Tanner, U.S.N., commanding. XV. The stalk-eyed Crustacea. Memoirs of the Museum of
Comparative Zoology, Harvard College, 18 : 1-292, pi. A-K + 1-56. 1 map.
264
J.C. MARKHAM
Hansen. H. J., 1897. — Reports on the dredging operations off the west coast of Central America to the Galapagos
Islands, to the west coast of Mexico, and in^the Gulf of California, in charge of Alexander Agassiz, carried on by the
U. S. Fish Commission Steamer Albatross during 1891. Lieut.- Commander Z. L. Tanner. U. S. Navy, commanding.
XXII. The Isopoda. Bulletin of the Museum of Comparative Zoology. Harvard College. 31 (5): 96-129, 6 pis.
H0EG, J. T. & Rybakov, A. V., 1992. — Revision of the Rhizocephala Akentrogonida (Cirripedia). with a list of all the
species and a key to the identification of families. Journal of Crustacean Biology. 12: 600-609.
Housa, V.. 1963. — Parasites of Tithonian decapod crustaceans (Stramberk. Moravia.). Ustred. Ustavu Geologic,
Shorn ik (Odd. Palent.), 28: 101-1 14. 2 pis.
KENSLEY. B., 1968. — Pseudione elongata africana , a new subspecies of bopyrid isopod from the west coast of the Cape
Peninsula, South Africa. Crustaceana, 15: 188-192.
Markham, J. C., 1992. — The Isopoda Bopyridae of the eastern Pacific - Missing or just hiding? Proceedings of the San
Diego Society of Natural History , 17: 1-4.
Markham, J. C, 1994. — Crustacea Isopoda: Bopyridae in the Musorstom collections from the tropical Indo-Pacific
I. Subfamilies Pseudioninae (in part), Argeiinae, Orbioninae. Athelginae and Entophilinae. In: A. Crosnier (ed.).
Resultats des Campagnes MUSORSTOM. Volume 12. Memoires du Museum national d'Histoire naturelle . 161: 225-
253.
McLay, C.L., 1993. — Crustacea Decapoda: The sponge crabs (Dromiidae) of New Caledonia and the Philippines with
a review of the genera. In: A. CROSNIER (ed.). Resultats des Campagnes MUSORSTOM. Volume 10. Memoires du
Museum national d'Histoire naturelle. 156: 1 1 1-251.
Nierstrasz H. F., & G. A. Brender a Brandis, 1923. — Die Isopoden der Siboga-Expedition. II. Isopoda Genuina.
I. Epicaridea. Siboga-Expeditie Monograph. 32b: 57-121, pis 4-9.
Nierstrasz H. F., & G. A. Brender a Brandis, 1931. — Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16.
LVII. Epicaridea II. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kpbenhavn. 91: 147-226, pi. 1.
Page, R. D. M, 1985. — Review of the New Zealand Bopyridae (Crustacea: Isopoda: Epicaridea). New Zealand Journal of
Zoology, 12: 185-212.
Richard, J., 1900. — Essai sur les Crustacds considers dans leurs rapports avec 1’ hygiene, la medecine et la
parasitologic. Lille: Le Bigot Fr&res. 85 pp.
Richardson, H.. 1899a. — Key to the isopods of the Pacific coast of North America, with descriptions of twenty-two
new species. Proceedings of the United Stales National Museum, 21: 815-869.
Richardson, H., 1899b. — Key to the isopods of the Pacific coast of North America, with descriptions of twenty-two
new species. Annals and Magazine of Natural History, (7) 4: 157-187, 260-277, 321-338.
Richardson, H., 1904. — Contributions to the Natural History of the Isopoda. Proceedings of the United States National
Museum. 27: 1-89.
Richardson, H., 1905. — A monograph on the isopods of North America. Bulletin of the United States National
Museum, 54: i-lii + 1-727.
Richardson, H.. 1910. — Marine isopods collected in the Philippines by the U. S. Fisheries steamer Albatross in
1907-8. Bureau of Fisheries Documents, (736): 1-44.
Rioja, E., 1971. — Clase IV. Los crustdceos (Crustacea). Pp. 470-554. In: Cendero, L.. editor: Zoologfa
Hispanoamericana - Invertebrados. Mexico. D. F.: Editorial Porrua, S. A.
$adoglu. P., 1969. — Variations in eye degeneration and pigment in some parasitic isopods during their life cycle.
Pubblicazioni della Stazione Zoologica di Napoli, 37 : 173-209.
Schultz, G. A.. 1969. — How to know the marine isopod crustaceans. Dubuque, Iowa: Wm. C. Brown Company, vii +
359 pp.
Shiino, S. M., 1951. — Some bopyrid parasites found on the decapod crustaceans from the waters along Mie Prefecture.
Report of the Faculty • of Fisheries. Prefectural University of Mie. 1: 26-40.
SHIINO, S. M.. 1952. — Phylogeny of the family Bopyridae. Annual Report of the Prefectural University of Mie
(Section 2, Natural Science), 1: 33-56. [In Japanese with English Summary].
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BOFYRIDAE OF THE TROPICAL INDO-PACIFIC
265
Shiino, S. M., 1972. — [The Epicaridea (list of species) from Japan). Kansai Shizenkagaku . 24: 7-10. (In Japanese).
Stebbing, T. R. R., 1914. — South African Crustacea. (Part VII of S. A. Crustacea, for the Marine Investigations in South
Africa). Annals of the South African Museum, 15: 1-55, 12 pis.
Van Name, W. G., 1924. — Isopods from the Williams Galapagos Expedition. Zoologica, 5 (18): 181-210, pis 8-19.
Source : MNHN. Paris
Source : MNHN. Paris
jUL.TATS DES CAMPAC.NES MUSORSTOM, VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DE
Crustacea Decapoda: Revision of Pasiphaea sivado
(Risso, 1816) and related species, with descriptions of
one new genus and five new species (Pasiphaeidae)
Ken- 1 chi HAY ASH 1
National Fisheries University
P.O. Box 3 Yoshimi, Shimonoseki
759-65 Japan
ABSTRACT
The study of many samples collected by Musorstom cruises, deposited in the Museum national d'Histoire naturelle, as
well as the reexamination of types and published specimens reveal that Pasiphaea sivado (Risso, 1816) and the related
species, P. propinqua de Man. 1916, P. japonica Omori, 1976, P. marisrubri Iwasaki, 1989 and P. nudipeda Burukovsky,
1993, belong to one group. All are characterized by a terminal spine on the sixth abdominal somite and a branchial
reduction. However, P. nudipeda is entirely devoid of arthrobranchia, has unarmed first pereiopods and three pairs of
spines on the posterior margin of telson and has to be separated; a new genus Alainopasiphaea is proposed for it. The
other species mentioned above, except P. marisrubri, bear three arthrobranchiae from the lourth to sixth thoracic
somites. P. marisrubri and five new species found in the Musorstom material and belonging in this group have four
pleurobranchiae from the fourth to seventh thoracic somites. On the other hand, P. propinqua , P. japonica and P. sivado
have one more, but rudimentary, pleurobranchia on the eighth somite. A key for all these species is provided.
RESUME
Crustacea Decapoda: Revision de Pasiphaea sivado (Risso, 1816) et des especes qui lui sont
proches. Description d’un genre et cinq especes nouveaux (Pasiphaeidae).
Letude de nombreuscs recoltes, rassemblees lors des campagnes Musorstom et depos6es au Museum national
d’Histoire naturelle, et le r^examen de types ou de specimens publies montrent que Pasiphaea sivado (Risso, 1816) et les
especes proches, P. propinqua de Man, 1916, P. japonica Omori. 1976, P. marisrubri Iwasaki. 1989 et P. nudipeda
Burukovsky, 1993, appartiennent & un meme groupe qui se caracterise par la presence dune epine distale sur le sixieme
segment abdominal et une reduction des branchies. Toutefois P. nudipedia est totalement depourvue d'arthrobranchies et a
des premiers pereiopodes sans epine et trois paires d epines sur le bord post6rieur du telson; il convient de la separer;
pour ellc, un nouveau genre Alainopasiphaea est propose. Les autres especes mentionnees ci-dessus, a 1'exception de
P. marisrubri , poss^dent trois arthrobranchies ^parties sur les segments thoraciques 4-6. P. marisrubri et cinq especes
nouvelles, trouvees dans le materiel Musorstom et appartenant au groupe considere ici, possedent quatre pleurobranchies
reparties sur les segments thoraciques 4-7. Enfin. P. propinqua , P. japonica et P. sivado ont une pleurobranchie supple-
mentaire, mais rudimentaire, sur le segment thoracique 8. Une c!6 d'identiheation pour toutes ces esp&ces est proposee.
Hayashi, K.-I., 1999. — Crustacea Decapoda: Revision of Pasiphaea sivado (Risso, 1816) and related species, with
descriptions of one new genus and five new species (Pasiphaeidae). In: A. CROSNIER (ed.), Resultats des Campagnes
MUSORSTOM, Volume 20. Memoires du Museum national d'Histoire naturelle , 180: 267-302. Paris ISBN 2-85653-520-8.
Source : MNHN. Pans
268
K.-I.. HAYASHI
INTRODUCTION
The genus Pasiphaea is a large group, containing nearly 60 species (Burukovsky & ROMENSKY, 1987;
BURUKOVSKY, 1996). A world wide review of the genus has not been completed, and several species have been left
unclear as to their specific status. Amongst them arc the type species of the genus, P. sivado (Risso, 1816) and
related species.
P. sivado was reported from both the Atlantic Ocean and the Indo-West Pacific region. However, specimens
from Japanese waters were shown to be a different species. P. japonica Omori, 1976, and Red Sea specimens were
referred to another different species, P. marisrubri Iwasaki, 1989. Some specimens from other localities described
under that name have not been reexamined in detail, such as those recorded in WOOD Mason & Alcock (1893)
and in KENSLEY (1977), both from the Indian Ocean.
All these species are easily distinguished by having a terminal spine on the sixth abdominal somite. Two other
species, P. propinqua de Man, 1916, and P. nudipeda Burukovsky, 1993, have such a spine. All these species share
some other important characters with one another and therefore, probably constitute a natural group: the P. sivado
species group. One of the important characters of this group is the branchial reduction, which has not been drawn
attention to as a means of distinguishing between species in the genus Pasiphaea.
Many pasiphaeids collected from various areas by several MUSORSTOM cruises are referred to seven species
belonging to the P. sivado species group and including five new species. An another species, close to this group
but entirely devoid of arthrobranchiae, seems to have to be separated from it; for this species I propose a new genus
Alainopasiphaea.
The specimen size is indicated by the carapace length (CL), not including the rostrum. The specimens examined
are preserved at the following institutions: Museum national d'Histoire naturelle, Paris (MNHN),
Forschungsinstitut Senckenberg, Frankfurt (SMF), National Fisheries University, Shimonoseki (NFU), South
African Museum (SAM). Zoologisch Museum, Amsterdam (ZMA) and Zoological Museum of the Moscow State
University (MMSU).
SYSTEMATIC ACCOUNT
Genus ALAINOPASIPHAEA nov.
Definition. — Small pasiphaeids. Rostrum short, triangular with pointed apex, arising behind anterior
margin of carapace. Carapace not carinated and smooth dorsally, with branchiostegal spine only. Abdomen not
carinated dorsally. Posterior margin of telson truncated with three pairs of spines. Mandible without palp. Fourth
pereiopod shorter than third and fifth. No arthrobranchia on third maxilliped. No arthrobranchiae but four
pleurobranchiae present on fourth to seventh thoracic somites.
ETYMOLOGY. — The genus name Pasiphaea with the prefix Alain, the first name of a French carcinologist.
Alain CROSNIER of the ORSTOM who is much involved in editing the results of the MUSORSTOM cruises.
Remarks. — The new genus is related to the genus Pasiphaea , from which it is distinguished by such
characters as the simple branchial formula, having four pleurobranchiae only and no arthrobranchiae at all, the
unarmed first pereiopod and three pairs of spines on the posterior margin of the telson.
A full set of the branchial components of the genus Pasiphaea is five pleurobranchiae from the fourth to the
eighth thoracic somites and three arthrobranchiae from the fourth to sixth thoracic somites. The reduction of the
branchial formulae is shown in some species, especially in P. sivado and related species, but then three
arthrobranchiae are always present, without exception. The posterior margin of the telson is usually armed with
eight or more spines in Pasiphaea , and is very variable in shape and armature. The completely unarmed first
pereiopod, with even no posterodistal spine on the basis, is also shown in a few species of the genus Pasiphaea.
The present genus contains a single species, Alainopasiphaea nudipeda (Burukovsky, 1993) described in detail
below.
PASIPHAEA SIVADO AND RELATED SPECIES
269
Alainopasiphaea nudipeda (Burukovsky, 1993) new comb.
Figs 1-3
Pasiphaea sp. a - DE Man, 1920: 9, pi. 1, fig. 3-3o.
Pasiphaea nudipeda Burukovsky, 1993: 35, fig. 1, 8-13; 1996, 843 (list).
MATERIAL EXAMINED. — Mozambique. "Vi!iaz"'. stn 2631, 25°28.0'S, 35°08'E. 535-490 m, 23.11.1988: 1 $
holotype (MMSU 1/84269).
Madagascar. "Vauban": Trawl 89, 21°18’S, 43°17.4’E, 620 m, 26.11.1973: 1 6 11.5 mm (MNHN-Na 13370). —
Trawl 90, 21°24.5'S, 43°13.5'E, 640-720 m, 26.11.1973: 1 ovig. 9 10.9 mm (MNHN-Na 13371).
Indonesia. Mollucas: " Siboga ": stn 148, 0°17.6’S, 129°14.5’E, 10.08.1899: 1 juv. 4.6 mm (ZMA).
DIAGNOSIS. — Shell not fragile. Rostrum small, apex not reaching anterior margin of carapace. Carapace
dorsally rounded; branchiostegal sinus developed. Abdominal somites dorsally rounded. First pereiopod unarmed on
merus, ischium and basis, posterodistal angle of basis not spiniform. Merus of second pereiopod with single spine,
ischium unarmed, basis unarmed, posterodistal spine small. Ischium of third pereiopod without spinules.
Arthrobranchiae absent. Pleurobranchiae on fourth to seventh thoracic somites. Ovigerous female about 10 mm.
DESCRIPTION. — Rostrum small and short, hardly reaching halfway between base of rostrum and anterior
margin of carapace, though apex missing in male (Figs. 1, 2c) and entirely broken in female. Branchiostegal spine
small situated just inside anterior margin of carapace. No middorsal carina on carapace. Branchiostegal sinus
distinct (Fig. 2c).
All abdominal somites dorsally smooth, without carinae and spines (Fig. 2d). Sixth abdominal somite
1.8 times as long as fifth somite and 1.9 times as long as deep; terminal spine extending straight backward;
posterolateral margin with small convexity just below spine; ventrodistal corner with small concavity near distal
end (Fig. 2e). Telson 0.6 times as long as sixth somite, dorsally with shallow groove, but almost ilat near
Source :
270
K.-I.. HAYASHI
Fig. 2. Alainopasiphaea nudipeda (Burukovsky, 1993), a-b, holotype, 9 12 mm (MMSU, 1/84269). Mozambique;
ce, i-n. ovig. 9 10.9 mm (MNHN-Na 13371); f-h, o-p. 6 11.5 mm (MNHN-Na 13370). bolh from Madagascar.
a. i, branchial chamber; b. eye, antennule and antennal scale, dorsal view; c. anterior part of body; d, abdomen; e, distal
end of sixth somite; f, telson; g, distal end of telson; h. antennal scale; j, mandible; k. maxillula; 1, maxilla; m. first
maxilliped; n. second maxilliped; o, first pleopod; p, second pleopod. Scales = 1 mm.
Source :
PASIPHAEA SIVADO AND RELATED SPECIES
271
midlength (Fig. 20; distal margin truncated with three pairs of spines, outer pair longest, without small seta on
base; inner two pairs nearly equal in length (Fig. 2g).
Eyes well developed; cornea spherical in lateral view and well pigmented (Fig. 2b-c). Stylocerite slightly
shorter than first segment of antennular peduncle (Fig. 2b). Antennal scale reaching midpoint of enlarged part of
antennular flagellum (Fig. 1), 3.8 times as long as wide, and shorter (0.88-0.92) than chela of first pereiopod;
outer margin evenly convex, and ending in small tooth; lamellar part truncated at distal end, entirely overreached by
distolateral tooth (Fig. 2b. h). Basicerite with slender spine on lower distal corner (Fig. 2c).
Mouth-parts illustrated, not apparently different from those of the genus Pasiphaea , hut slightly more simple in
structure. Mandible provided with about 10 strong teeth along mesial margin of incisor process; palp absent
(Fig. 2j). Proximal endite of maxillula small, triangular, with short simple seta; distal endite armed with six acute
teeth; endopod oblong, with stout simple seta on distal end (Fig. 2k). Maxilla composed of well developed
endopod and large scaphognathite (Fig. 21). First maxillipcd provided with elongated lamellar part, articulated
distally; no incision present on the outer margin of basal part (Fig. 2m). Second maxilliped simple pediform;
epipod and exopod absent (Fig. 2n). Third maxilliped long, reaching slightly beyond antennal scale (Fig. I); distal
segment about twice as long as penultimate segment; exopod well developed (Fig. 3a).
First pereiopod (Fig. 3b) reaching beyond antennular peduncle by chelae (Fig. 1); basis not ending in spiniform
process (Fig. 3c); ischium and merus unarmed on posterior margin; carpus short, with spine on distodorsal and
ventral ends; palm longer than fingers, with two slender movable setae on mesial margin; cutting edges toothed,
tips curved and crossing (Fig. 3d). Second pereiopod (Fig. 3e) reaching beyond antennular peduncle by chela, with
small spine at posterodistal end of basis (Fig. 3f); ischium unarmed; merus with single spine at distal 1/3 of
posterior margin; carpus with a slender spine on distoventral corner; palm as long as fingers. Third pereiopod
slender, reaching beyond anterior margin of carapace by dactylus and part of propodus; no spinules on any segment
(Fig. 3g). Fourth pereiopod shortest, reaching basial spine of second pereiopod; dactylus with long setae, propodus
with short stiff setae along posterior margin (Fig. 3h). Fifth pereiopod not reaching anterior margin of carapace;
dactylus terminally rounded with long setae (Fig. 3i).
Endopod of male first pleopod composed of three lobes, mesial lobe small, with some retinaculae in central
part; other two lobes surrounded by long plumose setae (Fig. 2o). Endopod ol male second pleopod with
appendices masculina and interna; masculina shorter than interna, and provided with eight long setae (Fig. 2p).
Uropod (Fig. I) much longer than telson; exopod much longer than endopod; outer margin ending in small spine,
not reaching end of lamella.
Branchial formula simple, consisting of only four pleurohranchiac, and six exopods from third maxilliped to
fifth pereiopod (Fig. 2a, i):
Size. — The holotype is 12 mm CL (Burukovsky, 1993). The Musorstom material are slightly smaller
than the holotype. The carapace length is 1 1.5 mm CL in male and 10.9 mm in ovigerous female. Eggs are large
and relatively few in number, 1.47 x 0.95 mm. The "Siboga" specimen is immature, its carapace length is only
4.6 mm.
Remarks. — The present species was recently described by Burukovsky (1993), based on a single female
collected from the western Indian Ocean. Although six spines on the posterior margin of the telson and the
unarmed first pereiopod were shown in the original description, the simple branchial formula was neither
mentioned nor figured by BURUKOVSKY (1993). Very recently I could examine the holotype, in which
Source :
272
K.-I.. HAYASHI
FIG. 3. — Alainopasiphaea nudipeda (Burukovsky, 1993), a-b, e. g-i, 6 11.5 mm (MNHN-Na 13370) ; c-d. f. ovig. $
10.9 mm (MNHN-Na 13371), both from Madagascar.
a, third maxilliped; b, first pereiopod; c, basal part of first pcreiopod; d, chela of first pereiopod in inner view;
e, second pereiopod; f, basal part of second pereiopod; g. third pereiopod; h. fourth pereiopod; i, fifth pereiopod.
Scales = 1 mm.
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
273
unfortunately the carapace was entirely missing, but confirmed these characters (Fig. 2a-b). The Musorstom
specimens agree well with the holotypc and show no apparent sexual dimorphism.
I could also examine the small specimen that deMan (1920) named Pasiphaea sp. a. This is a juvenile, only
4.6 mm in carapace length, and probably identical with the present species in having the following characters: The
branchial formula is the same as in the present species, only four pleurobranchiae are recognized in the branchial
chamber. The first pereiopod is entirely unarmed on the posterior margin of merus, ischium and basis. The distal
end of basis bears no spine. The merus of the second pereiopod is armed with one spine at distal third of the
posterior margin; the distal end of the basis bears a small spine.
Some minor discrepancies between the MUSORSTOM and "Siboga" specimens are present. The branchiostegal
sinus is ill defined and the spine on the basicerite is small in the " Siboga " specimen, and the ventral surface of
palm of the first pereiopod is provided with a single seta near the finger articulation, while two setae, one near the
finger articulation and the other on the midlength of palm, are observed in the MUSORSTOM specimens. Obviously
these differences depend largely on the immaturity of the "Siboga" specimen.
Distribution. — The present specimens were collected from two localities near Madagascar at depths of 620-
720 m. The holotype was obtained from off Mozambique at depths of 535-490 m. The " Sibogci " specimen was
obtained from Indonesia.
Genus PASIPHAEA Savigny, 1816
Pasiphaea sivado species group
Definition. — Small and moderately sized pasiphaeids. Rostrum short, triangular with pointed apex, arising
behind anterior margin of carapace. Dorsal margin of carapace smooth in most species but carinate anteriorly in
some species. Branchiostegal spine present. Posterior margin of telson truncate, slightly convex or slightly
concave, with four pairs of spines. Mandible without palp. Fourth pereiopod shorter than third and fifth
pereiopods. No arthrobranchia on third maxilliped. Three arthrobranchiae from fourth to sixth thoracic somites and
four or five pleurobranchiae present on fourth to seventh or eighth thoracic somites, plcurobranchia on eighth
somite, if present, always rudimentary.
Pasiphaea sivado and related species have the following characters: 1) size small to moderate. 2) carapace
usually not carinate dorsally, 3) first to fifth abdominal somites not carinated dorsally. 4) a terminal spine present
on sixth abdominal somite only, 5) distal margin of telson not forked, with four pairs of spines, 6) plcurobranchia
on eighth thoracic somite absent or rudimentary, and 7) three arthrobranchiae present on fourth to sixth thoracic
somites.
The general morphology of the species of this group resembles each other, as set out below.
Rostrum small, triangular process situated at middorsal line slightly inside carapace. Small branchiostegal
spine situated on or just inside anterior margin of carapace.
First to fifth somites smooth dorsally, unarmed. Sixth somite less than twice as long as fifth and less than
twice as long as deep. Telson shorter than sixth somite, usually with longitudinal groove dorsally. Distal margin
of telson truncate or convex, with four pairs of spines; outer pair longest, usually with small accessory seta on
their base; inner pairs shorter than outer, gradually decreasing in size.
Eye well developed; cornea spherical or semispherical in lateral view, well-pigmented. Styloccrite not reaching
or just reaching distal margin of first segment of antennular peduncle; dorsal margin ending in small point in
lateral view. Antennal scale overreaching enlarged part of antennular flagellum, about tour times as long as wide;
outer margin more or less convex; outerdistal tooth projecting beyond lamella. Basicerite armed with slender spine
on lower distal corner.
Mouth-parts of typical shape. Mandible with several strong teeth along mesial margin of incisor process.
Maxillula comparatively large; proximal endite small, obliquely truncate distally with one or lew short simple se¬
tae; distal endite with several acute teeth; endopod oblong, with stout simple seta on mesial margin near distal end.
274
K.-I.. HAYASHI
Maxilla with well developed endopod and large scaphognathite. First maxillipcd with elongated lamellar part, artic¬
ulated distally. Second maxillipcd simple, pediform; epipod and exopod absent. Third maxilliped long, reaching
slightly beyond antennal scale; distal segment about twice as long as penultimate segment; exopod well developed.
All pereiopods with well developed exopods, but no epipods. First pereiopod usually reaching beyond distal end
of antennular peduncle by entire chela; carpus short, sharply pointed on dorsal and ventral ends; palm slightly
longer than fingers, usually with two slender movable setae on mesial margin; fingers slender, their cutting edges
toothed; tips curved, crossing one another. Second pereiopod similar to first pereiopod in shape and length; carpus
with slender distoventral spine only; finger nearly as long as palm, strongly curved at distal part. Third pereiopod
slender, reaching beyond anterior margin of carapace by dactylus and a part of propodus; all segments usually
unarmed. Fourth pereiopod shortest, reaching basial spine of second pereiopod only; dactylus provided with rather
long setae on posterior margin; propodus with short stiff setae along posterior margin. Fifth pereiopod reaching
distal margin of carpus of third pereiopod; dactylus broad and rounded distally, with several long setae.
Endopod of male first pleopod with two or three lobes, mesial lobe small, with some retinaculae in central
part; outer lobe surrounded by long plumose setae. Endopod of male second pleopod with appendices masculina and
interna; masculina usually shorter than interna, with many long setae. Uropod elongate; exopod much longer than
endopod, outer margin with small distal spine, as long as or overreaching lamella.
Remarks. — The genus Pasiphaea contains many species which together show a wide range of morphological
variation. Of these the type species of the genus, P. sivado , and other related species, such as P. propinqua de Man.
1916, P. japonica Omori, 1976, and P. marisrubri Iwasaki, 1989, were reported to be very similar in shape and
size. They differ from the other members of Pasiphaea in having a terminal spine on the dorsal margin of the sixth
abdominal somite. Five other new species related to P. sivado were found in the MUSORSTOM material. They prove
to share some important characters with the above mentioned species, in addition to the terminal spine on the sixth
abdominal somite. They constitute a single natural group, the P. sivado species group, which maybe in the future
will be recognized as a distinct subgenus of the genus Pasiphaea.
The terminal spine on the sixth abdominal somite is the most apparent and useful character, by which this
species group is readily distinguished from the other species groups. This character does not easily change in shape
and size, and is less likely to be damaged than the tips of rostrum, telson and uropodal exopods, when specimens
suffer damage. In some species such as P. truncata Rathbun, 1906 and P. longitaenia Kensley, Tranter & Griffin,
1987, the dorsal margin of the sixth abdominal somite is produced as a "pointed end", not as a true spine. The
lateral aspect of the pointed end somewhat resembles that of a true spine, but spine and pointed end are easily
recognized from each other in dorsal view. The true spine is slender and needle-like, especially not broadened at
base, but the pointed end is triangular in shape with a broad base.
There are some species provided with a posterodorsal spine on other abdominal somites as well as on the sixth
somite, for example P. orientalis Schmitt, 1931, P. hoplocerca Chace, 1940 and P. semispinosa Holthuis, 1951.
They are excluded from this species group, because they do not share the above mentioned characters.
It has been emphasized that main specific characters of the genus Pasiphaea were the spination of the first and
second pereiopods and the shape of the telson end (Burukovsky & ROMENSKY, 1987; BURUKOVSKY, 1996).
They are indeed important, but the branchial differences seem to be almost neglected in this genus. The species of
this species group, such as P. sivado. P. japonica , and P. propinqua. have the same branchial formula, three
arthrobranchiae from the fourth to sixth thoracic somites and four normal and one rudimentary pleurobranchiae
from the fourth to eighth thoracic somites. This formula, however, differs from those of other remaining species,
in which the last plcurobranchia is normal, not reduced. Among the MUSORSTOM materials are five new species,
which bear different and more reduced branchial formulae.
Branchial reduction, therefore, occurs in every species of the P. sivado species group. The condition is not
uniform, though there are always three arthrobranchiae, and the species are roughly divided into two groupings:
five species bear only four pleurobranchiae and three have five pleurobranchiae, of which the last one is always
rudimentary, bearing only a few gill lamellae. It differs in form from the miniature or undeveloped gill, often seen
in the immature specimens of other large-sized species of the genus.
On the other hand, Pasiphaea species without the terminal spine on the sixth somite have three well-developed
arthrobranchiae from the fourth to sixth thoracic somites and five normal pleurobranchiae are also present on all
Source :
PASIPHAEA SIVADO AND RELATED SPECIES
275
thoracic somites. The last pleurobranch is never reduced, having usually the same shape as those of preceding
somites. Pasiphaea species with terminal spines on two or three abdominal somites, even if a spine is present on
the sixth somite, never show branchial reduction, and have a complete set of gills.
These two groupings within the Pasiphaea species group based on branchial differences coincide rather well
with a pattern of spination of the first and second pereiopods. Each basis of the first and second pereiopods always
ends in a well-developed posterodistal spine in both groupings. The mcri are armed with a series of spines, but
their arrangements are rather different from each other. There are less than ten spines on the first pereiopod and less
than 17 spines on the second pereiopod in the first grouping, while in the second grouping usually more than ten
spines on the first pereiopod and usually more than 15 spines on the second pereiopod. P. sivado is an exception,
though it has the branchial formula of the second grouping, the spination is less than ten on the first pereiopod,
and less than 15 spines qn the second pereiopod.
The basis and ischium does not show such pattern of the spination, and the presence or absence of the spine(s)
seem to be constant, though numbers are always variable. There is an exception with the ischium of P. gracilis
sp. nov., in which one female from the northern South Pacific is armed with one spine on the left side but
unarmed on the right side.
The rostrum is shown to be rather variable in shape depending on growth, or presence or absence of ellobiopsid
parasites, but its general morphology is still useful for almost all species. As pointed out by BURUKOVSKY &
ROMENSKY (1987) and BURUKOVSKY (1996), the form of the branchial sinus is important and may be constant,
though it is rather difficult to observe, because the ventral margin of carapace is often softened and folded inwards.
Pasiphaea sivado has been reported several times from various seas. Records from other areas than the Atlantic
Ocean, were partly checked and proved to be different species (Omori, 1976 and Iwasaki, 1989) and the remaining
records arc now reexamined from the specimens concerned or from the published references. Now. P. sivado s.s. is
restricted in its distribution to the northeastern Atlantic Ocean and Mediterranean Sea. Including the five new
species described here, the Pasiphaea sivado species group includes the following nine members:
P. debitusae sp. nov. (? = Pasiphaea sivado Wood Mason, 1892 and Pasiphaea sivado Wood Mason &
Alcock, 1893),
P. fragilis sp. nov.,
P. gracilis sp. nov. (? = Pasiphaea sp. P de Man, 1920),
P. japonica Omori, 1976 (= Pasiphaea aff. sivado Crosnier, 1976 and Pasiphaea sivado Kensley, 1977),
P. laevis sp. nov.,
P. marisrubri Iwasaki, 1989,
P. philippinensis sp. nov.,
P. propinquci de Man, 1916,
P. sivado (Rissq, 1816).
These species are distinguished from each other in the key presented below.
Key to the Pasiphaea sivado species group
1. Four pleurobranchiae present . 2
— Five pleurobranchiae present, though that on last thoracic somite rudimentary . 7
2. Basis of second pereiopod with 2-7 spines, excluding terminal spine . 3
— Basis of second pereiopod without spines . 4
3. Rostrum slender and short, not reaching halfway between base of rostrum and anterior
margin of carapace. Ischium of third pereiopod with 1-4 spinules .
. P. marisrubri Iwasaki, 1989
— Rostrum short, but reaching beyond halfway between base of rostrum and anterior
margin of carapace. Ischium of third pereiopod unarmed . P. philippinensis sp. nov.
Source :
276
K.-I. HAYASHI
4. Rostrum short and small. Ischium of second pereiopod with spine .
. P. debitusae sp. nov.
— Rostrum medium or long. Ischium of second pereiopod usually unarmed . 5
5. Shell fragile. Terminal spine of sixth abdominal somite extending slightly upwards .
. P. fragilis sp. nov.
— Shell not fragile. Terminal spine of sixth abdominal somite extending straight backwards
. 6
6. Branchiostegal sinus absent or obscure. Rostrum long with wide base .
. P. laevis sp. nov.
— Branchial sinus present. Rostrum medium with narrow base . P. gracilis sp. nov.
7. Sixth abdominal somite sharply carinate dorsally. Posterior margin of telson slightly
convex . P. propinqua De Man, 1916
— Sixth abdominal somite not carinate dorsally. Posterior margin of telson truncate . 8
8. Merus of first pereiopod with 1 -8 spines, merus of second pereiopod with 5- 1 5 spines . . .
. P. sivado (Risso, 1816)
— Merus of first pereiopod with 5-12 spines, merus of second pereiopod with 14-23 spines
. P.japonica Omori, 1976
Pasiphaea marisrubri Iwasaki, 1989
Fig. 4
Pasiphaea marisrubri Iwasaki, 1989: 178, figs 1-2.
Pasiphaea marisrubrae - BURUKOVSKY, 1996: 843 (list).
? Pasiphaea sivado - Balss, 1915: 17. Not Risso, 1816.
Pasiphaea sivado - C ALMAN, 1939: 185. Not Risso, 1816.
MATERIAL EXAMINED. — Central Red Sea. "Sonne": stn So-02/43-TAP, 21o14.80’N, 37°15.40’E, 0-220 m,
18.10.1977: 7 6 7.1-10.0 mm, 1 ovig. $10.8 mm, 6 $ 6.6-11.4 mm (paratypes, SMF 17545).
"Meteor": stn M5/193-Ku, 19°24.3’N, 38°31.2'E. 696-705 m. 28.02.1987: 4 $ 9.1-12.7 mm (SMF 17999).
DIAGNOSIS. — Shell moderately firm. Rostrum small, spine-like, not reaching anterior margin of carapace.
Carapace dorsally rounded; branchiostegal sinus developed. Abdominal somites dorsally rounded. Posterior margin
of telson truncate, with 4 pairs of spines. First pereiopod with 5-12 spines on merus and unarmed on basis except
for a posterodistal spine. Second pereiopod with 14-23 spines on merus, and unarmed on ischium; unarmed on
basis except for a posterodistal spine. Ischium of third pereiopod with two or three spinules on posterior margin.
Developed arthrobranchiae present on fourth to sixth thoracic somites. Pleurobranchiae from fourth to seventh
thoracic somites.
Size. — The holotype is a male, 10.0 mm in CL and the allotype an ovigerous female, 1 1.2 mm. The
smallest ovigerous female is 9.4 mm in CL. The egg size is 1.2 x 0.8 mm (Iwasaki, 1989).
Remarks. — The species is well described by Iwasaki (1989). The branchial formula of this species was
confirmed by the examination of paratypes and other materials deposited at the Forschungsinstitut Senckenberg,
Frankfurt. No trace of a pleurobranch is present on the eighth thoracic somite (Fig. 4b).
The distinctions between the present species and P. philippinensis sp. nov. are mentioned under the account of
the latter species. As shown by Iwasaki (1989), the rostrum is small and short, and the distal spine on the
basicerite is more slender (Fig. 4a). Moreover P. marisrubri always bears two or three spinules on the posterior
margin of the ischium of the third pereiopod, which is an unique character of this species (Fig. 4c-d).
Source :
PASIPHAEA SIVADO AND RELATED SPECIES
277
Fig. 4. — Pasiphaea marisrubri Iwasaki, 1989, paratypes: a. 2 9.7 mm ; b-d, 2 1 1.4 mm. both from Red Sea (SMF
17545, part).
a, anterior part of body; b, branchial chamber; c, basal part of right third pereiopod; d, basal part of left third pereiopod.
Scales = 1 mm.
References to P. sivado based on Red Sea specimens (Balss, 1915 and Calm AN, 1939) have already been
reviewed by IWASAKI (1989).
Distribution. — Only known from the Red Sea (Iwasaki. 1989).
Pasiphaea Philippine nsis sp. nov.
Figs 5-7
MATERIAL EXAMINED. — Philippines. Musorstom 2: stn CP 25. 13°39’N, 120°43'E, 520-550 m. 23.1 1.1980:
1 6 12.0 mm (MNHN-Na 13372).
Type Material. — The unique specimen, a male 12,0 mm CL (MNHN-Na 13372), is the holotype.
Diagnosis. — Shell not fragile. Rostrum moderate, not reaching anterior margin of carapace. Carapace
dorsally rounded; branchiostegal sinus obscure. Abdominal somites dorsally rounded. Posterior margin of telson
truncate, with 4 pairs of spines. First pereiopod with 9 spines on merus and 1 spine on basis excluding
posterodistal spine. Second pereiopod with 16-17 spines on merus, 1 spine on ischium and 4-5 spines on basis
excluding posterodistal spine. Ischium of third pereiopod without spinules. Developed arthrobranchiae present on
fourth to sixth thoracic somites. Pleurobranchiae on fourth to seventh thoracic somites.
Description. — Rostrum short, directed obliquely forwards, reaching halfway between base of rostrum and
anterior margin of carapace. Carapace with branchiostegal sinus hardly developed (Figs 5, 6a).
Source :
278
K.-l.. HAYASHI
Fig. 5. — Pasiphaea philippinensis sp. nov., holotype, 6 12.0 mm (MNHN-Na 13372). Philippines. Scale = 1 mm.
Sixth abdominal somite 1.7 times as long as fifth somite and 1.8 times as long as deep; ventrodistal corner
with small convexity with minute spine (Fig. 6c). Telson 0.7 limes as long as sixth somite, dorsally with
shallow groove extending for almost entire length (Fig. 6d); distal margin slightly convex with probably four pairs
of spines, outer pair longest, without small seta on base; inner two pairs nearly equal in length (Fig. 6e).
Stylocerite shorter than first segment of antennular peduncle (Fig. 6b). Antennal scale 4.0 times as long as
wide, and shorter (0.93) than chela of first pereiopod; basicerite articulated with slender spine on lower distal corner
(Fig. 6a). Mouth-parts showing typical shape of genus (Figs 6f-j, 7a).
First pereiopod with spine at posterodistal end of basis (Fig. 7b. c); ischium unarmed and merus with nine
spines on posterior margin; chela typical shape, with two movable setae on mesial margin (Fig. 7d). Basis of
second pereiopod with large spine at posterodistal end and four or five spines on posterior margin (Fig. 7e-f);
ischium with one spine on midlength of posterior margin; merus with 16 or 17 spines on posterior margin. No
spinules on ischium of third pereiopod (Fig. 7g). Fourth and fifth pereiopods typical shape for genus (Fig. 7h, i).
Fndopod of male first pleopod composed of two lobes, mesial lobe small, with some retinaculae in central part;
other lobe large, surrounded by long plumose setae (Fig. 6k). Endopod of male second pleopod of typical shape for
the group; appendix masculina with ten long setae (Fig. 61).
Branchial formula as follows:
Pleurobranchiae present on fourth to seventh thoracic somites, eighth somite with no pleurobranch.
Arthrobranchiae present on fourth to sixth thoracic somites. All pereiopods with well developed exopods, but
without epipods or podobranchiae.
Source :
PASIPHAEA SIVADO AND RELATED SPECIES
279
Fig. 6. — Pasiphaea philip pinensis sp. nov., holotype, 6 12.0 mm (MNHN-Na 13372), from the Philippines,
a, anterior part of body in lateral view; b, same in dorsal view; c. distal end of sixth abdominal somite and telson in
lateral view; d. telson in dorsal view; e, distal end of telson; f. mandible; g. maxillula; h. maxilla; i. first maxilliped;
j. second maxilliped; k. first pleopod; 1, appendices masculina and interna. Scales = 1 mm.
Etymology. — The specific name, philippinensis, refers to the type locality. Philippine waters.
SIZE. — The unique specimen, the holotype male, is 12.0 mm CL.
Remarks. — The holotype is slightly damaged, distal three segments and the distal part of merus of the left
first pereiopod, distal three segments of the right second pereiopod, and outer distal spine of the uropodal exopod
are missing. However, this species is very clear in its specific status and readily distinguished from other species.
Source : MNHN. Paris
280
K.-I.. HAYASHI
Fig. 7. — Pasiphaea philippinensis sp. nov., holotype, 8 12.0 mm (MNHN-Na 13372), Philippines,
a. third maxilliped; b, first pereiopod; c, basal part of First pereiopod; d, chela of first pereiopod in inner view;
e, second pereiopod; f. basal part of second pereiopod; g. third pereiopod; h, fourth pereiopod; i. fifth pereiopod.
Scale = 1 mm.
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
281
P. philippinensis sp. nov. is characterized by having only four pleurobranchiae and three arthrobranchiae in one
side of the branchial chamber and some spines on the basis of the second pcreiopod. The combination of these
characters has not been found in any other known species, except for P. marisrubri Iwasaki.
Four pleurobranchiae are recorded in the other five new species mentioned below, but they all have the basis of
the second pereiopod unarmed, excluding the posterodistal spine. P. marisrubri however bears four or five spines
on the posterior margin of basis of the second pereiopod, excluding the posterodistal spine. The branchial formula
was confirmed by an examination of a part of the type series. P. marisrubri shows the same branchial formula as
P. philippinensis. However, these two species arc distinguished from each other by the rostrum shape, and the
spination of the ischium of the third pereiopod. In P. marisrubri the rostrum is slender and short, not reaching
halfway between base of the rostrum and anterior margin of the carapace. On the other hand. P . philippinensis has
the rostrum slightly longer, reaching more than halfway between base ot the rostrum and anterior margin ol the
carapace (Fig. 6b). The ischium of the third pereiopod is usually armed with one to four spinules on the posterior
margin in P. marisrubri , while unarmed in P. philippinensis (Fig. 7g). The spine on the basicerite ol the second
antenna is more slender in P. marisrubri than in P. philippinensis , and one or two spinules are present on the
lower margin near the proximal part in P. marisrubri (Fig. 4a), but no such spinules are present in
P. philippinensis (Fig. 6a).
Distribution. — The Philippines at a depth of 520-550 m.
Pasiphaea debitusae sp. nov.
Figs 8-10
? Pasiphaea sivado - Wood Mason, 1892. pi. 3, fig. 6. Not Risso, 1816.
? Pasiphaea sivado - Wood Mason & Alcock, 1893, fig. 1. Not Risso, 1816.
MATERIAL EXAMINED. — Indonesia. Karubar: stn DW 08, 05°20'S. 132°3TE. 358-360 m, 23.10.1991: 1 9
139 mm (MNHN-Na 13373). — Stn CP 09. 05°23'S. I32°29’E, 368-389 m, 23.10.1991: 1 c3 13.8 mm. 2 ovig. 9
14.8 mm (MNHN-Na 13374). — Stn CP 19, 05°15’S, 133°OrE. 605-576 m, 25.10.1991: 1 ovig. 9 14.0 mm (MNHN-
Na 13375). — Stn CP 27, 05°33’S. 132°51’E, 304-314 m. 26.10.1991: 1 9 10.1 mm (MNHN-Na 13376).
Fig. 8. — Pasiphaea debitusae sp. nov., holotype. ovig. 9 14.0 mm (MNHN-Na 13375). Banda Sea. Scale - 1 mm.
Source :
282
K.-I.. HAYASHI
TYPE Material. — The ovigerous female (14,0 mm, MNHN-Na 13375) collected at the station CP 19 of
Karubar cruise is the holotypc. All the other specimens are paratypes.
Fig. 9. — Pasiphaea debitusae sp. nov., paratypes: a-b, 9 13.9 mm (MNHN-Na 13373); c-f, ovig. 9 14.8 mm (MNHN-
Na 13374, part); g-h, 6 13.8 mm (MNHN-Na 13374, part), all from Banda Sea.
a. anterior part of body in lateral view; b, distal end of sixth abdominal somite in lateral view; c, telson in dorsal view;
d. distal end of telson; e. branchial chamber; f. antennal scale; g. endopod of first pleopod; h. appendices masculina
and interna. Scales = 1 mm.
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
283
DIAGNOSIS. — Shell not fragile but not hard. Rostrum small spine-like, not reaching anterior margin ot
carapace. Carapace dorsally rounded; branchiostegal sinus obscure. Abdominal somites dorsally rounded. Posterior
margin of telson truncate, with 4 pairs of spines. First pereiopod with 4-8 spines on mcrus and unarmed on basis
excluding posterodistal spine. Second pereiopod with 9-12 spines on merus, a single spine on ischium and
unarmed on basis excluding posterodistal spine. Ischium of third pereiopod without spinules. Developed
arthrobranchiae present on fourth to sixth thoracic somites. Pleurobranchiae from fourth to seventh thoracic
somites. Ovigerous females 14-15 mm.
Description. — Rostrum small, like short spine, hardly reaching halfway between base ot rostrum and
anterior margin of carapace (Figs 8, 9a). Branchiostegal sinus hardly developed (Fig. 9a).
Sixth abdominal somite 1. 7-2.0 times as long as fifth somite and 1. 9-2.0 times as long as deep; ventrodistal
corner shallowly convex distally (Fig. 9b). Telson 0.6-0.7 times as long as sixth somite, dorsally grooved without
interruption (Fig. 9c); distal margin truncate with four pairs of spines, outer pair longest, without small seta on
base; innermost pair shortest, two intermediate pairs nearly of equal length (Fig. 9d).
Stylocerite shorter than first segment of antennular peduncle (Fig. 9a). Antennal scale 4. 1-4.8 limes as long as
wide, and shorter (0.91-0.95) than chela of first pereiopod; basicerite with slender spine on lower distal corner
(Fig. 9a). Mouth-parts showing typical shape of genus (Fig. lOa-f).
First pereiopod with spine at posterodistal end of basis; ischium unarmed; mcrus with four to eight spines on
posterior margin (Fig. lOg); palm longer than fingers, with two slender setae on mesial margin (Fig. 1 Oh). Basis
of second pereiopod ending in large spine, posterior margin unarmed; ischium with or without single spine on
posterior margin (Figs 8, lOi); merus with 9 to 14 spines on posterior margin (Fig. lOi). No spinules on ischium
of third pereiopod (Fig. lOj). Fourth and fifth pereiopods typical shape for genus (Fig. 10k-l).
Endopod of male first pleopod composed of two lobes, mesial lobe small, with some retinaculae in central part;
outer lobe surrounded by long plumose setae (Fig. 8g). Endopod of male second pleopod similar to preceding
species; appendix masculina with seven long setae (Fig. 8h).
Branchial formula same as preceding species. Pleurobranchiae present on fourth to seventh thoracic somites, no
pleurobranchia on eighth thoracic somite. Arthrobranchiae on fourth to sixth somites (Fig. 9e).
Etymology. — From the family name of Cecile Debitus, biochemist at ORSTOM. who was in charge of
the programm SMIB (Substances Marines d’Interet Biologique) from 1986 to 1996 and busily took part in the
study of the deep-sea fauna off New Caledonia, describing new molecules of pharmaceutical interest.
SIZE. — The type series is composed of one male 13.8 mm CL, three ovigerous females 14.0-14.8 mm and
two other females 10.1, 13.9 mm. Eggs are large and relatively few in number, 1.0-1.03 x 1.37-1.50 mm.
Remarks. — P. debitusae sp. nov. somewhat resembles the previous new species, P . philippinensis , in
having the same branchial formula, and a single spine on the ischium of the second pereiopod. P. debitusae ,
however, bears no spine on the basis of the second pereiopod, excluding the posterodistal spine, while lour or live
spines are present on the basis in P. philippinensis. Moreover, the rostrum is apparently shorter and smaller in
P. debitusae than in P. philippinensis.
Two males were reported from north of the Andaman Sea under the name of P. sivado by WOOD MASON
(1892) and WOOD Mason & ALCOCK (1893). The description was too brief to understand definitely the specific
status, but the entire animal and the appendices masculina and interna were figured. They have a very short and
slender rostrum and the first two pereiopods armed with a series of spines on the posterior margin. Especially some
spines are seen on the merus and one spine on the proximal segment, probably ischium, ot the second pereiopod.
These results including the locality show that the Andaman species has a high possibility of being referable to the
present species, unless it is a new species. P. marisrubri has a short rostrum too, but the armature ol the second
pereiopod is more complicated and it is restricted in its distribution to the Red Sea.
DISTRIBUTION. — Four lots of the type specimens were all collected from near Kai Islands. Indonesia, at
depths of 304-605 m during the Karubar Expedition in 1991. If the above identification of the WOOD Mason
specimens is correct, the distribution range of the species can be extended to the Andaman Sea.
Source :
284
K.-l.- HAYASH1
Fig. 10. — Pasiphaea debitusae sp. nov., paraiype, 6 13.8 mm (MNHN-Na 13374, part), from Banda Sea.
a, mandible; b, maxillula; c, maxilla; d, lirst maxilliped; e. second maxillipcd; f. third maxilliped; g, first pcreiopod;
h, chela of first pereiopod in inner view; i, second pereiopod; j, third pereiopod; k, fourth pereiopod; 1. fifth pereiopod.
Scale = 1 mm.
Source : MNHN. Paris
PAS1PHAEA SIVADO AND RELATED SPECIES
285
Pasiphaea fragilis sp. nov.
Fig. 11
MATERIAL EXAMINED. — Loyalty Islands. Musorstom 6: stn CP 438, 20°23.0’N, 166°20.rE, 780 m.
18.02.1989: 1 6 10.8 mm (MNHN-Na 13377), 1 9 10.9 mm (MNHN-Na 13378).
Type Material. — The male (10.8 mm, MNHN-Na 13377) collected at the station CP 438 of Musorstom 6
cruise is the holotype. The female collected at the same station is a paratype.
Diagnosis. — Shell fragile. Rostrum small, not reaching anterior margin of carapace. Carapace dorsally
rounded; branchiostegal sinus developed. Abdominal somites dorsally rounded. Posterior margin of telson
truncate, with 4 pairs of spines. First pereiopod with 6-9 spines on merus and unarmed on basis
excluding posterodistal spine. Second pereiopod with 9-12 spines on merus, unarmed on ischium, and unarmed on
basis excluding posterodistal spine. Ischium of third pereiopod without spinules. Developed arthrobranchiae present
on fourth to sixth thoracic somites. Pleurobranchiae on fourth to seventh thoracic somites. Specimens about
1 1 mm.
DESCRIPTION. — Rostrum short, not reaching halfway between base of rostrum and anterior margin of
carapace. Branchiostegal sinus shallow, but distinct (Fig. 1 la).
Sixth abdominal somite 1.7 times as long as fifth somite and 1.8 times as long as deep; vcntrodistal corner
with shallow convexity and with small pointed process; terminal spine angled slightly upwards (Fig. 1 lb). Telson
O. 7 times as long as sixth somite, with shallow groove extending dorsally without interruption (Fig. 1 lc); distal
margin slightly convex, probably with four pairs of spines (Fig. 1 Id).
Stylocerite shorter than first segment of antennular peduncle. Antennal scale 3. 8-4. 2 times as long as wide,
and shorter (0.89) than chela of first pereiopod; outerdistal spine overreaching lamella (Fig. I le); basicerite
with slender spine on lower distal corner. Mouth-parts not apparently different from those of other species
(Fig. 110.
First pereiopod with spine at posterodistal end of basis; ischium unarmed; merus with six to nine spines on
posterior margin (Fig. 1 1 g). Basis of second pereiopod ending in spine, posterior margin unarmed; ischium
unarmed; merus with 9 to 12 spines on posterior margin (Fig. 1 lh). No spinules on ischium of third pereiopod
(Fig. Hi).
Endopods of male first and second pleopods similar to those of preceding species (Fig. 1 1 1-m).
Branchial formula same as preceding species, pleurobranchiae from fourth to seventh thoracic somites and
arthrobranchiae on fourth to sixth thoracic somites.
ETYMOLOGY. — The specific nam e, fragilis, is based on the very fragile condition ot the shell.
Size. — The male, holotype, is 10.8 mm CL; the female, paratype. 10.9 mm.
Remarks. — P. fragilis sp. nov. belongs to the subgroup having four pleurobranchiae and three
arthrobranchiae, and is morphologically related to P. gracilis sp. nov. in the shape o! rostrum and branchiostegal
sinus. The shell is fragile and the terminal spine of the sixth abdominal somite is angled slightly upwards in
P. fragilis. Moreover, the meral spination of the first two pereiopods is slightly less in P. fragilis. Six spines in
P. fragilis and seven to nine spines in P. gracilis on the first pereiopod and 9 to 12 spines in P .fragilis and 13 or
14 spines on the second pereiopod in P. gracilis.
The spination of the distal margin of the telson is probably four pairs in the present species; in the holotype
only seven spines remains, but the left half bears four spines, their arrangement is similar to those of other
species. On the other hand, there are nine spines in the paratype, four regular spines present on the right half, five
spines on left half, in which the inner two are more slender than others.
Distribution. — The two type specimens were collected from one station in New Caledonian waters at a
depth of 780 m.
Source :
286
K.-I.. HAYASHI
Fig. 11. — Pcisiphaea fragilis sp. nov.: a-b. e-m, holotype, 6 10.8 mm (MNHN-Na 13377); c-d, paratype. 9 10.9 mm
(MNHN-Na 13378), both from Central Pacific Ocean.
a. anterior part of body in lateral view; b, distal end of sixth abdominal somite in lateral view; c, telson in dorsal view;
d. distal end of telson; e, apex of antennal scale; f. third maxilliped; g, first pcreiopod; h, second pereiopod; i, third
pereiopod; j, fourth pereiopod; k, fifth pereiopod; 1, endopod of first pleopod; m, appendices masculina and interna.
Scales = 1 mm.
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
287
Pasiphaea laevis sp. nov.
Figs 12-14
MATERIAL EXAMINED. — Indonesia. Makassar Strait. CORINDON 2: stn CH 214, 00°31'N, 1 17°50‘E, 595 m: 1 ?
11.9 mm (MNHN-Na 13379).
Moluccas . Karubar: stn CP 35, 06°08’S. 132°45’E, 390-502 m, 27.10.1991: 1 ovig. 2 12.3 mm (MNHN-Na
13380).
Type Material. — The ovigerous female (12.3 mm, MNHN-Na 13380) collected at the station CP 35 of
Karubar cruise is the holotype. The other female (1 1.9 mm, MNHN-Na 13379) is a paratype.
Diagnosis. — Shell not fragile. Rostrum long and broad based, nearly overreaching anterior margin of
carapace. Carapace dorsally rounded; branchiostegal sinus very indistinct. Abdominal somites dorsally rounded.
Posterior margin of telson truncate, with 4 pairs of spines. First pereiopod with 7-9 spines on merus and unarmed
on basis excluding posterodistal spine. Second pereiopod with 12-13 spines on merus, unarmed on ischium and
unarmed on basis excluding posterodistal spine. Ischium of third pereiopod without spinules. Developed
arthrobranchiae present on fourth to sixth thoracic somites. Pleurobranchiae from fourth to seventh thoracic
somites. Ovigerous female with carapace length of 12 mm.
Fig. |2. — Pasiphaea laevis sp. nov., holotype. ovig. $ 12.3 mm (MNHN-Na 13380). Banda Sea. Scale - 1.0 mm.
Description. — Rostrum long, with broad base, reaching or overreaching anterior margin ot carapace
(Figs. 12, 13a-b). Branchiostegal sinus indistinct, without apparent excavation (Fig. 13a, b).
Sixth abdominal somite 1.8 times as long as fifth somite and 1.8 times as long as deep; ventrodistal corner
shallowly convex, with minute spine (Fig. 13c-d). Telson 0.7 times as long as sixth somite, with groove dorsally,
but with groove almost disappearing around midlength (Fig. 13e); distal margin truncate, with four pairs ot spines,
outer pair longest, with small seta on base (Fig. 13f).
Stylocerite slightly shorter than first segment of antennular peduncle (Fig. 13a-b). Antennal scale 3.8-
4.0 times as long as wide, and as long as chela of first pereiopod; outerdistal spine rather long, entirely
Source :
288
K.-I.. HAYASHI
overreaching lamella (Fig. I3g); basiceritc with slender spine on lower distal corner. Mouth-parts showing no
apparent difference from other species (Fig. 14a-f).
First perciopod with extremely large spine on posterodistal end of basis; ischium unarmed; merus with 7 to
9 spines on posterior margin (Figs 13h, 14g); palm of typical shape with two movable setae on mesial margin
(Fig. 14h). Basis of second pereiopod ending in spine, posterior margin unarmed; ischium unarmed; merus with
Fig. 13. — Pasiphaea laevis sp. nov.: a. c, e-f, holotype, ovig. 9 12.3 mm (MNHN-Na 13380), Banda Sea; b. d. g-i.
paratype, 9 11.9 mm (MNHN-Na 13379), Indonesia.
a-b, anterior part of body in lateral view; c-d, distal end of sixth abdominal somite in lateral view; e, telson in dorsal
view; f, distal end of telson; g, distal apex of antennal scale; h, basal part of first pereiopod; i, basal part of second
pereiopod. Scales = 1 mm.
Source : MNHN, Paris
=r ts
PASIPHAEA SIVADO AND RELATED SPECIES
289
Fig. 14. — Pasiphaea loevis sp. nov.. paratype, 9 1 1.9 mm (MNHN-Na 13379). Indonesia.
, mandible; b. maxillula; c, maxilla; d, first maxilliped; e. second maxilliped; f. third maxilliped; g, first pereiopod;
.chela of first pereiopod in inner view; i, second pereiopod; j. third pereiopod; k. fourth pereiopod; I, lifth pereiopod.
Scale = 1 mm.
Source : MNHN. Paris
290
K.-I.. HAYASHI
12 or 13 spines on posterior margin (Figs 13i, 14i). No spinulcs on ischium of third pereiopod (Fig. 14j). Fourth
and fifth pereiopods same as other member of the P. sivado species group (Fig. 14k, 1).
Four pleurobranchiae and three arthrobranchiae on one side of the branchial chamber. Branchial formula same as
P. marisrubri and P.fragilis sp. nov. described above.
ETYMOLOGY. — The specific name, laevis, refers to the smooth ventral margin of carapace.
SIZE. — The ovigerous female, holotype. is 12.3 mm CL, the non-ovigerous female, 1 1.9 mm. Eggs are
numerous in number, 0.8-0. 9 x 1.2- 1.3 mm in size.
Remarks. — P. laevis sp. nov. has the same branchial formula as P. marisrubri and the preceding new species
P. frag i l is, but is distinguished from those species by the long rostrum and no branchiostegal sinus on the
carapace.
The present new species shows the same meristic value of the spination of the first two pereiopods as
P.fragilis sp. nov. and P. gracilis sp. nov. However, in latter two new species the branchiostegal sinus is more or
less defined. Moreover, the shell is distinctly fragile in P. fragilis and the rostrum and antennal scale is
comparatively shorter in P. gracilis.
Distribution. — The holotype was collected near Kai Islands, Indonesia, at a depth of 390-502 m, and the
paratype was from Makassar Strait, at a depth of 595 m.
Pasiphaea gracilis sp. nov.
Figs 15-17
? Pasiphaea sp. P - DE Man, 1920: 1 1, pi. 1, fig. 4-4g, pi. 2, fig. 4h-4p.
Fig. 1 5. — Pasiphaea gracilis sp. nov,, holotype, <3 12.3 mm (MNHN-Na 13382). New Caledonia. Scale = 1 mm.
Material EXAMINED. — Chesterfield Islands. Musorstom 5: stn DW 313, 22°24.31'S I59°32 53'E 780-
930 m, 13. 10.1986: 1 9 12.0 mm (MNHN-Na 13381).
New Caledonia. Biocal: stn CP 31, 23°08'S, 166°51E. 850 m, 29.08.1985: I <3 12.3 mm (MNHN-Na 13382) I
9 14.0 mm (MNHN-Na 13383).
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
291
Wallis and Futuna Islands. MUSORSTOM 7: stn DW 620, 12°34’S, 178°irw. 1280 m, 28.05.1992: 1 6 6.3 mm
(MNHN-Na 13384). — Stn CP 627, 11°54'S, 179031'W, 597-600 m, 29.05.1992: 3 6 8.6-10.0 mm. 1 9 9.1 mm
(MNHN-Na 13385). — Stn CP 628, 11°53’S, 179°32’W. 625-650 m, 29.05.1992: 4 6 9.6-11.8 mm, 5 9 10.0-1 1.1 mm
(MNHN-Na 13386).
Indonesia. " Siboga ”: stn 105, off Sulu Islands, 6°8'N, 121°19'E, 275 m, 4.07.1899: 1 juv. 5.8 mm (ZMA).
Type Material. — The male (12.3 mm, MNHN-Na 13382) collected at the station CP 31 of Biocal cruise
is the holotype. All the other specimens, except the " Siboga " one, are paratypes.
Diagnosis. — Shell not fragile. Rostrum small, not reaching anterior margin of carapace. Carapace dorsally
rounded; branchiostegal sinus developed. Abdominal somites dorsally rounded. Posterior margin of telson truncate,
with 4 pairs of spines. First pereiopod with 7-9 spines on merus, unarmed on basis excluding posterodistal spine.
Second pereiopod with 11-13 spines on merus, usually unarmed on ischium and unarmed on basis excluding
posterodistal spine. Ischium of third pereiopod without spinules. Developed arthrobranchiae present on fourth to
sixth thoracic somites. Pleurobranchiae on fourth to seventh thoracic somites. Females 12-14 mm.
FlG. 16. — Pasiphaea gracilis sp. nov., holotype, 6 12.3 mm (MNHN-Na 13382), New Caledonia.
a, anterior part of body in lateral view; b, distal end of sixth abdominal somite in lateral view; c. telson in dorsal view;
d, distal end of telson; e. distal apex of antennal scale; f, endopod of first pleopod; g, appendices interna and masculina.
Scales = 1.0 mm.
DESCRIPTION. — Rostrum rather variable in length, comparatively short, usually just reaching halfway
between base of rostrum and anterior margin of carapace (Figs 15, 16a). Branchiostegal sinus shallow, but distinct
(Fig. 16a).
Sixth abdominal somite 1.7 times as long as fifth somite and 1.8 times as long as deep; ventrodistal corner
with shallow convexity, with small pointed process (Fig. 16b). Telson 0.7 times as long as sixth somite, with
shallow groove dorsally without interruption; distal margin truncate with probably four pairs of spines (Fig. 16c);
outer pair longest, without small seta on base (Fig. 16d).
Stylocerite shorter than first segment of antennular peduncle. Antennal scale 3. 8-4.2 times as long as wide, and
shorter (0.85-0.95) than chela of first pereiopod; outerdistal spine slightly curved outward, overreaching lamella
(Fig. 16e). Mouth-parts not apparently different from those of other species (Fig. 17a-f).
Source :
292 K.-I.. HAYASHI
Fig. 17. — Pasiphaea gracilis sp. nov., paratype, 6 1 1.3 mm (MNHN-Na 13386), South Pacific Ocean near Wallis Island,
a. mandible; b, maxillula; c, maxilla; d, first maxilliped; e, second maxilliped; f. third maxilliped; g, first pereiopod;
h. second pereiopod; i, third pereiopod; j, fourth pereiopod; k. fifth pereiopod. Scale = 1 mm.
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
293
First pereiopod with spine at posterodistal end of basis; ischium unarmed; merus with four to ten spines on
posterior margin (Fig. 17g). Basis of second pereiopod ending in spine, posterior margin unarmed; ischium usually
unarmed; merus with 9 to 14 spines on posterior margin (Fig. 17h). No spinules on ischium of third pereiopod
(Fig. 17i). Fourth and fifth pereiopods of typical shape for the genus (Fig. 17j-k).
Endopods of male first and second pleopods similar to those of preceding species (Fig. 16f, g).
Branchial formula same as preceding species. Arthrobranchiae present on fourth to sixth somites, no
pleurobranchia on eighth thoracic somite (Fig. 9e).
Etymology. — The specific name, gracilis, refers to the slender shape of the species.
SIZE. — The type series is composed of nine males, 8.6-13.6 mm and eight non-ovigerous females, 9.1-
1 4.0 mm.
Remarks. — P. gracilis sp. nov. resembles the preceding new species, P. laevis , in having the same spination
of the first two pereiopods, and the shell not fragile but the branchial sinus is distinct and the antennal scale is
shorter than the chela of the first pereiopod in P. gracilis.
The spination of the ischium is usually consistent, but one female, 10.0 mm, from the southwestern Pacific is
an exception, which has a small spine on the left side, but unarmed on the right side as in the other specimens of
P. laevis examined.
Judging from the description given by DE MAN (1920) Pasiphaea sp. p from the "Siboga" station 105 has a
spine on the sixth abdominal somite and shows the possibility of belonging to this grouping ol the P. sivado
species group. 1 could reexamine this young female, 5.5 mm. The shell is not fragile. The branchiostegal sinus is
shallow, but distinct. There are four developed pleurobranchiae and three small arthrobranchiae in one side of the
branchial chamber. Each basis of the first two pereiopods is armed with a spine and three to four meral spines are
present on the First pereiopod, and nine to ten meral spines on the second pereiopod. Although the arthrobranchiae
are not fully grown, the branchial formula is the same as this grouping. It is rather difficult to identify this
specimen exactly, because of its small size, but it probably belongs to P. gracilis rather than to P. laevis.
Distribution. — Several localities of the South Pacific, at depths of about 600-1300 m. If the present
identification is correct, the "Siboga" specimen was obtained from off Sulu Island.
Pasiphaea propinqua de Man, 1916
Fig. 18
Pasiphaea propinqua de Man, 1916: 147; 1920: 7, pi. 1. fig. 1-lj. — BURUKOVSKY, 1996: 843 (list).
MATERIAL EXAMINED. — Indonesia. "Siboga": stn 100, 06°1 l'N, 120°37.5'E. 450 m, 29.06.1899: 1 6 17.0 mm
(holotype, ZMA 102505).
Philippines. MUSORSTOM 2: stn CP 38. 12°53.5'N. 122°26.6'E. 1650 m, 25.11.1980: 1 2 12.8 mm (MNHN-Na
13387).
Type-Material. — The male (17.0 mm, ZMA 102505) collected at the station 100 of the "Siboga"
Expedition, is the holotype.
DIAGNOSIS. — Shell not fragile. Rostrum long, probably reaching anterior margin of carapace. Carapace
dorsally carinated on anterior 1/3, rounded posteriorly; branchiostegal sinus obscure. First to fifth abdominal
somites dorsally rounded, sixth somite sharply carinated dorsally. Posterior margin of telson convex, with 4 pairs
of spines. First pereiopod with 10-12 spines on merus and unarmed on basis excluding posterodistal spine. Second
pereiopod with 16-18 spines on merus, and unarmed on ischium, and unarmed on basis excluding posterodistal
spine. Ischium of third pereiopod without spinules. Developed arthrobranchiae present on fourth to sixth thoracic
somites. Pleurobranchiae from fourth to eighth thoracic somites, that on eighth somite rudimentary. Holotype
male 17 mm, and additional female 12.8 mm.
Source :
294
K.I.. HAYASHI
Description. — Rostrum probably not short, reaching more than halfway between base of rostrum and
anterior margin of carapace (Fig. 18a). Branchiostegal sinus indistinct (Fig. 18a).
Sixth abdominal somite 1.7 times as long as fifth somite and 1.9 times as long as deep; ventrodistal corner
with shallow convexity, and with a small pointed process (Fig. 18b). Telson 0.6 times as long as sixth somite,
deeply grooved dorsally with clear longitudinal ridges lateral to groove (Fig. 18c); a rather high process proximal
Fig. 18. — Pasiphaea propinqua de Man, 1916: a. c-d, 9 12.8 mm (MNHN-Na 13387), the Philippines; b. e-i. holotype
6 17.0 mm (ZMA 102505), Sulu Sea.
a, anterior part of carapace in lateral view; b. distal end of sixth abdominal somite in lateral view; c, telson in dorsal
view; d, distal end of telson; e, branchial chamber; f, second pcreiopod; g, third pereiopod; h, fourth pereiopod;
i, endopod of first pleopod; j, appendices interna and masculina. Scales = 1 mm.
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
295
to base of groove (Fig. 18b); distal margin convex with four pairs of spines like other species of the P. sivado
species group, but with a pair of large additional seta near base of outer spines (Fig. 18d).
Stylocerite falling short of end of first segment of antennular peduncle. Antennal scale 3.6 times as long as
wide, and 1.2 times as long as chela of first pereiopod. Mouth-parts not apparently different from those of other
species.
First pereiopod with spine at postcrodistal end of basis; ischium unarmed; merus with 12 to 14 spines on
posterior margin. Basis of second pereiopod ending in spine, posterior margin unarmed; ischium unarmed; merus
with 16 to 18 spines on posterior margin (Fig. 180. Third to Fifth pereiopods similar to those of other species
(Fig. 18g-h). No spinules on ischium of third pereiopod (Fig. I8g).
Endopod of first pleopod of holotype composed of three lobes, though entirely without setae; mesial lobe
small, with a very few retinaculae in central part (Fig. 18i). Endopod of second pleopod of holotype similar to
preceding species; appendix masculina with some, probably less than ten, long setae (Fig. 18j).
Distribution of branchiae, epipods, and exopods as follows:
Eighth thoracic somite with pleurobranchia, but always rudimentary, only a very small piece, with a few short
papilla-like lamellae (Fig. 18e).
Size. — The male holotype is 17.0 mm CL. the present additional female, 12.8 mm.
Remarks. — The present species was based on a single male type, described in detail (DE Man, 1920), which
was fortunately reexamined. The branchial formula and the spination of the sixth abdominal somite are the same as
in P. sivado , having a rudimentary pleurobranchia on the eighth thoracic somite, and a well-developed spine on the
posterior end of the dorsal margin of the sixth somite (Fig. 18b).
As mentioned by DE Man (1920), P. propinqua has the carapace anteriorly carinatcd. the sixth abdominal
somite entirely carinated, and the telson with convex end. Moreover there is a well developed process at the
beginning of the dorsal groove of the telson. The groove is deeper and is better defined than that of other species by
distinct longitudinal ridges on both sides. These characters apparently separate P . propinqua from all other
members of the P. sivado species group. The present female specimen from the Philippines agrees well with the
holotype in these unique characters.
The rostrum is mostly missing in the holotype. In the additional specimen it is also broken, but some distal
parts remain attached to the rostrum base, which shows that it is probably not short, but long, extending nearly to
the anterior margin of carapace. The branch iostegal spine is almost marginal (Fig. 18a), at least the dorsal margin
of the spine is apparently continuous with the anterior margin of the carapace in both specimens examined.
In the holotype, the meri of the second pereiopods are armed with 16 spines on the right side and 18 spines on
the left side, with 17 spines on both sides in the additional material. The ischium and the basis are unarmed on
both sides, except for the posterodistal spine on the basis in both specimens.
The pleopods of the holotype have retained their shape, though they are now mostly without setae, and they do
not differ from those of the other species of the P. sivado species group.
Distribution. — Sulu Sea, at depth of 450 m (type locality). The subsequent specimen was found in one
haul from Philippine waters at a depth of 1650 m. This haul contained two pasiphaeids. both however were
thought to have been entangled in the net during the rising of the net (CROSNIER personal communication).
Source
296
K.-L HAYASHI
Pasiphaea sivado (Risso, 1816)
Fig. 19
Restricted synonymy
Alpheus sivado Risso, 1816: 93, pi. 3, l'ig. 4.
Pasiphaea savignyi H. Milne Edwards, 1837: 426.
Pasiphaea hrevirostris H. Milne Edwards, 1837: 426.
Pasiphaea sivado - H. MILNE Edwards, 1837: 426. — BELL, 1853: 312, with fig. — HELLER, 1863: 243, pi. 8, figs 4-6.
— Kemp, 1910: pi. 4, fig. 12. — Pesta, 1914: figs 22-23; 1918: 64, figs 19-20. — Zariquiey Alvarez, 1957: 4. figs
1-4, pis 1-3; 1968: 70, fig. 30. — Smaldon, 1979: 26, fig. 6. — Sivertsen & Holthuis, 1956: 29. — Omori. 1976:
254, figs 3-5. — Burukovsky & Romensky. 1987: 58 (list). — Iwasaki. 1989: 199. — Holthuis, 1993: 28, fig. 9.
— Burukovsky, 1996: 843 (list).
Pasiphaea Sivado - DE Man, 1920, pi. 1. figs 2, 2a.
Fig. 19. — Pasiphaea sivado (Risso, 1816): a, c. 8 23.9 mm (MNHN-Na 13389, part); b, d, 8 14.4 mm (MNHN-Na
13389, part); e. ovig. 9 18.6 mm (MNHN-Na 13388, part); f, 9 14.3 mm (MNHN-Na 1834, part), a-d, from West
Mediterranean Sea; e-f, from Ibero-Moroccan Gulf.
a-b, anterior part ol body in lateral view; c-d, distal part of sixth abdominal somite and telson in lateral view; e, distal
part of telson in lateral view; f, branchial chamber. Scales = 1 mm.
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
297
MATERIAL EXAMINED. — Ibero-Moroccan Gulf. "Talisman". 36°36'N. 07°26'W. Dredge. 440 m, 25.08.1883:
3 6 13.6-18.6 mm; 9 $ 13.2-17.2 mm (MNHN-Na 1834).
Balgim: stn CP 89. 34°20'N, 07°18'W. 719-724 m, 7.06.1984: 1 6 22.4 mm; 3 ovig. 2 17.9-19.4 mm (MNHN-Na
1 3388 )
Mediterranean Sea. Alboran Sea. Balgim: sin CP 119. 35°50'N. 05°13'W. 483-551 m. 13.06.1984: 2 6 14.4.
23.9 mm; 7 2 10.9-14.3 mm (MNHN-Na 13389).
DIAGNOSIS. — Shell not fragile. Rostrum not reaching anterior margin of carapace. Carapace dorsally rounded;
branchiostegal sinus shallow. Abdominal somites dorsally rounded. Posterior margin of telson truncate, with
4 pairs of spines. First pereiopod with 2-7. usually 4-5 spines on merus and unarmed on basis excluding
posterodistal spine. Second pereiopod with 9-13. usually 9-1 1. spines on merus, unarmed on ischium, and unarmed
on basis excluding posterodistal spine. Ischium of third pereiopod without spinules. Developed arthrobranchiae
present on fourth to sixth thoracic somites. Pleurobranchiae on fourth to eighth thoracic somites, that on eighth
somite rudimentary. Ovigerous females 18-20 mm.
DESCRIPTION. — Rostrum rather high, variable in length, usually reaching halfway between base of rostrum
and anterior margin of carapace. Branchiostegal sinus shallow, but quite distinct (Fig. 19a-b).
Sixth abdominal somite compressed but not carinated dorsally, 1.5- 1.9 times as long as lillh somite and 1.7-
2.0 times as long as deep; ventrodistal corner slightly convex, ending in minute spine (Fig. 19c-d). Telson 0.7-
0.8 times as long as sixth somite, with shallow and broad groove dorsally; distal margin truncated with four pairs
of spines, outer pair longest, with small spine-like seta at base (Fig. 19e).
Stylocerite almost reaching first segment of antennular peduncle (Fig. 19a-b). Antennal scale 3.6-4. 1 times as
long as wide, and slightly shorter than chela of first pereiopod. Mouth-parts showing typical shape of group.
First pereiopod with large spine at posterodistal end of basis; ischium unarmed; merus with two to seven
spines on posterior margin; palm 1.1-1. 4 times as long as finger. Basis of second pereiopod ending in large spine;
ischium unarmed; merus with 9 to 13 spines on posterior margin.
Endopod of male first pleopod composed of two lobes like other species mentioned above. Endopod of male
second pleopod of typical shape of group; appendix masculina with many long setae.
Branchial formula same as P. propinqua. Fifth pleurobranchia on eighth thoracic somite always rudimentary
(Fig. 190-
Size — Local variation in size is present, but the maximum overall size is 100 mm, usually not more than
80 mm (SMALDON, 1979). The males examined are 13.6-23.9 mm CL. the ovigerous females are 17.9-19.4 mm
CL. and non-ovigerous females are 13.2-17.2 mm. Eggs are large, 1.0 x 1.8 mm in size, and relatively few in
number.
Remarks. — P. sivado is the best known and the representative species of the genus as well as this species
group, having a pleurobranchia on the eighth thoracic somite, though it is rudimentary even in the largest male
and the ovigerous females examined, as first pointed out by KEMP (1910).
OMORI (1976) showed that the present species is distinguished from P. japonica Omori by the comparative
length of the antennal scale and the spination of the meri of the first and second pereiopods. The present Atlantic
specimens of P. sivado have the antennal scale always shorter than the chela of the first pereiopod (0.86-0.98), and
fewer spines on the meri of the first and second pereiopods (less than eight on the first, less than 14 on the
second), while the Japanese specimens of P. japonica have the antennal scale usually longer than the chela of the
first pereiopod and more than eight spines on the merus of the first pereiopod and more than 15 spines on the
merus of the second pereiopod.
As already shown by DE Man (1920), an accessory seta is present on the base of the outer pair of spines on the
telson end. It is as large as that of P. marisrubri , but may probably be more easy to detach in P. sivado (Iwasaki,
1989).
Distribution. — This well known species is now restricted to the Atlantic Ocean from Norway southwards to
the Morocco coast and the Mediterranean Sea (SMALDON, 1979).
Source
298
K.-I.. HAYASHI
Pasiphaea japonica Omori, 1976
Fig. 20
Pasiphaea sirado (sic) - Balss, 1914: 20.
Pasiphaea sivado - Kikuchi, 1932: 2. — Yokoya, 1933: 14. — KUBO, 1965: 605. with fig. — Aizawa, 1974: 28. —
Kensley, 1977: 32, fig. 10A. Not Risso, 1816.
Pasiphaea japonica Omori, 1976: 250, figs 1-8. — Burukovsky & ROMENSKY, 1987: 58 (list). — Burukovsky. 1993:
38, figs 2, 14-19; 1996: 843 (list).
Pasiphaea aff. sivado - Crosnier, 1976: 231. Not Risso, 1816.
Material EXAMINED. — Madagascar. "Vauban": Trawl 30, 12°40.0'S, 48°09.5'E, 595-605 m, 13.09.1972:
1 ovig. 9 18.3 mm; 3 9 13.0-20.0 mm (MNHN-Na 13390). — Trawl 49, 15°18.3'S, 46°10.3’E. 500-550 m, 8.11.1972:
1 8 13.8 mm (MNHN-Na 13391). — Trawl 59, 23°26.0’S, 43°29.6’E, 600-610 m, 27.02.1973: 1 8 15.3 mm; 1 ovig.
9 15.3 mm; 1 9 14.9 mm (MNHN-Na 13392). — Trawl 60, 23°36.5'S. 43°28.8'E. 710 m, 27.02.1973: 5 ovig. 9 15.0-
21.0 mm; 2 9 15.0, 16.6 mm (MNHN-Na 13393). — Trawl 65, 23°35.0’S, 43°28.6'E, 740-760 m, 29.02.1973: 2 ovig.
9 17.5 mm; 1 9 16.7 mm (MNHN-Na 13394).
"FAO 60": stn Chalutage 73/60, 21°17’S. 43°23’E. 490-530 m. 6.06.1973: 1 8 20.2 mm (MNHN-Na 13395). —
Stn Chalutage 73/70, 15°18'S, 46°15’E, 600-650 m, 25.06.1973: 1 9 15.6 mm (MNHN-Na 13396). — Sin Chalutage
73/94, 25°24'S. 46°58'E, 300 m, 12.08.1973: 1 ovig. 919.9 mm (MNHN-Na 13397). — Sin Chalutage. 22°15.7'S.
43°01.5'E. 750-810 m. 29.11.1973: 1 918.0 mm (MNHN-Na 13398).
Reunion. Le Port, P. Gu£z£ leg.: 1 9 16.5 mm (MNHN-Na 6910).
South Africa. "Meiring Naude": stn SM63, 27°10.5'S, 33°14.5’E. 140-0 m, 19.05.1976: 2 9 10.0. 14.6 mm (SAM
A 1 5 1 53). — Stn SM85, 27°59.5’S, 34°40.8’E, 550 m. Agassiz Trawl. 22.05.1976: 2 9 12.4 mm (SAM A 1 5152).
Indonesia. Karubar: stn CP 09, 05°23’S. I32°29’E, 368-389 m, 23.10.1991: 1 8 26.3 mm; 2 9 22.1. 26.2 mm
(MNHN-Na 13399).
Japan. Toyama Bay. off Miwa, set net, July 1975, N. Horii leg.: 1 ovig. 9 ca. 17 mm; 1 9 16.2 mm (NFU Cat
No. 530-2-613).
East China Sea. "Nansei Maru", 30.10.1976. Kagoshima Prefectural Fisheries Experimental Station leg.: 2 8
14.4, 15.0 mm; 3 ovig. 9 14.0-16.0 mm; 7 9 13.8-16.7 mm (NFU Cat. No. 530-2-1352).
Diagnosis. — Shell moderately firm. Rostrum short, not reaching anterior margin of carapace. Carapace
dorsally rounded; branchiostegal sinus shallow. Abdominal somites dorsally rounded. Posterior margin of telson
truncate, with 4 pairs of spines. First pereiopod with 5-12 spines on merus and unarmed on basis excluding
posterodistal spine. Second pereiopod with 14-23 spines on merus, unarmed on ischium, and unarmed on basis
excluding posterodistal spine. Ischium of third pereiopod without spinules. Developed arthrobranchiae present on
fourth to sixth thoracic somites. Pleurobranchiae on fourth to eighth thoracic somites, that on eighth somite
rudimentary. Ovigerous females 15-22 mm.
Description. — Rostrum small, reaching halfway between base of rostrum and anterior margin of carapace.
Branchiostegal sinus shallow (Fig. 20a).
Sixth abdominal somite compressed but not carinated dorsally, 1.4- 1.9 times as long as fifth somite and 1.7-
2.1 times as long as deep; ventrodistal corner uniformly convex, ending usually in minute spine (Fig. 20b).
Telson 0.6-0.7 times as long as sixth somite; dorsal groove very shallow at proximal half and prominent in distal
part; distal margin truncated with four pairs of spines, outer pair longest, with small seta on base (Fig. 20c).
Stylocerite reaching nearly to first segment of antennular peduncle (Fig. 20a). Antennal scale 3.5-4.5 times as
long as wideband 1.1 times as long as finger of first pereiopod. Mouth-parts showing typical shape of genus.
First pereiopod with large spine at posterodistal end of basis; ischium unarmed (Fig. 20c); merus with 5 to
12 spines on posterior margin; palm 1.1 times as long as finger. Basis of second pereiopod ending in large spine;
ischium unarmed (Fig. 200; merus with 14 to 23 spines on posterior margin.
Endopods of male first and second pleopods (Fig. 20g-h) similar to those of P. sivado.
Branchial formula same as in P. sivado and P. propinqua. Eighth thoracic somite with a rudimentary
pleurobranchia only (Fig. 20d).
Size. The males are 13.8-26.3 mm CL and the ovigerous females, 15.3-26.2 mm. Eggs are large, 1.0 x
1.6 mm in size, and relatively few in number.
Source : MNHN. Paris
PASIPHAEA SIVADO AND RELATED SPECIES
299
Fig. 20. — Pasiphaea japonica Omori. 1976: a-b. 2 22.1 mm (MNHN-Na 13399, pari) ; c. g-h, 8 26.3 mm (MNHN-Na
13399, part); d. ovig. 2 21.0 mm (MNHN-Na 13393. part); e-f. 2 26.2 mm (MNHN-Na 13399. part). All from
Indonesia, except d which is from Madagascar.
a, anterior part of body in lateral view; b. distal part of abdomen in lateral view; c. distal end of telson: d. branchial
chamber; e, basal part of first pereiopod; f, basal part of second pereiopod; g. endopod of first pleopod; h, appendices
interna and masculina. Scales = 1 mm.
Remarks. — From the branchial formula the present species belongs to the P. sivado and P. propinqua
groupings. From P. propinqua it clearly differs in having the dorsal surface of the sixth abdominal somite smooth.
Omori (1976) has already recorded the differences between P. sivado and the Japanese specimens of P. japonica.
I could not find any distinctive differences between the Japanese specimens and the present additional material from
Source :
300
K.-I . HAYASHI
Indonesia and western Indian Ocean, though there are several minor discrepancies between them. In the Indonesian
specimens, the rostrum and the spine of the sixth abdominal somite are comparatively small. Moreover, the
specimen size is larger (more than 22 mm) in the Indonesian specimens than in the Japanese and African
specimens (at most 21 mm).
I have examined three females from off Natal. South Africa, referred to P. sivado by KENSLEY (1977) and one
female from Reunion referred to P. aff. sivado by CROSNIER (1976). The South African females arc smaller than
the other materials examined, 10.0, 12.4 and 14.6 mm CL. The spination of the meri of the first and second
pereiopods is somewhat less in number, especially in the smaller two specimens: four to six spines on the first
and 8 to 12 spines on the second pereiopod. In the larger specimen, however, there are eight or nine spines on the
merus of the first pereiopod and 16 or 18 spines on the merus of the second. One female from Reunion is larger
than the South African specimens, 16.5 mm. The meral spines on the first and second pereiopods are slightly
numerous, eight or nine on the first and 17 to 20 on the second, as mentioned by CROSNIER (1976). These values,
however, fall within the range of spination in P. japonica rather than within that of P. sivado. The antennal scale
is slightly shorter than the chela of the first pereiopod in two smaller specimens or as long as the first chela in the
large specimens. From the distributional point of view these African specimens are referred to P. japonica. though
the large specimen has some spines relatively long, such as the outer distal spine of the antennal scale and the
terminal spine on the sixth abdominal somite.
Distribution. — This species is commercially important in Japanese waters, usually obtained from 40-120 m
in Toyama Bay, Sea of Japan, and also from the Pacific coast of the central Japan and the East China Sea (OMORI.
1976). Recently one male was reported from a far remote locality, southwestern Indian Ocean at a depth of 430-
350 m (Burukovsky, 1993). The material recorded here and specimens reported by CROSNIER (1976) and
KENSLEY (1977) partly fill the gap between these localities; Indonesia, near the Kai Islands, at a depth of 368-
389 m and Madagascan and South African waters, at a depth of 0-810 m.
ACKNOWLEDGMENTS
I thank Alain CROSNIER and Bertrand RICHER DE FORGES for their help and support during the present study.
I am also much indebted to Lipke B. Holthuis and C. H. J. M. FRANSEN of the Nationaal Natuurhistorisch
Museum, Leiden and Dirk PLATVOET of Instituut voor Systematiek en Populatiebiologie. Amsterdam for their
assistance to me while examining the "Siboga" material during my short stay in the Netherlands. Michael
Turkay of the Forschungsinstitut Scnckenberg, Frankfurt, and Barbara A. COOK of the South African Museum
sent me type and comparative material on loan. I express my sincere appreciation to Dr John C. Yaldwyn of the
Museum of New Zealand, Wellington and to an anonymous referee for reviewing the manuscript. This study was
largely supported by a fellowship made available to me by ORSTOM.
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de peche en eau profonde. In: Biologic marine et Exploitation des Ressources de l'ocean Indien occidental. Colloque
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TATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES
Crustacea Decapoda: A review of the species
of the genus Parapagurus Smith, 1879 (Parapaguridae)
from the Pacific and Indian Oceans
Rafael LEMAITRE
Department of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washington, D.C. 20560, USA
ABSTRACT
A review of the deep-water hermit crab species of the genus Parapagurus Smith. 1879 trom the Indian and Pacific
Oceans is presented based on abundant samples obtained during French expeditions to the New Caledonia region, and
supplemented with extensive material deposited in various major museums and institutions throughout the world. A total
of 14 species were found to occur in the Indian and Pacific Oceans. Of these seven are new, P. richeri sp. nov.. P. fund
sp. nov., P. stenorhinus sp. nov.. P. saintlaurentae sp. nov., P. janetae sp. nov., P . foraminosus sp. nov., and P. wolffi
sp. nov.; and three. P. abyssorum (Filhol, 1885), P. bouvieri Stebbing, 1910, and P. andreui Macpherson, 1984. include
parts of the Atlantic Ocean in their distribution. The new species are fully described and illustrated; all previously known
species are diagnosed or in the case of one obscurely defined species. P. holthuisi Lemaitre. 1989, redescribed.
Information on morphological variations is included for the most abundant species, and a key to aid in the identification
of all 14 species is given. Of the seven new species, P. richeri sp. nov. and P. furici sp. nov., were found in
the New Caledonia region but are also distributed elsewhere in the Indo-Pacific; P. saintlaurentae sp. nov. and
P. stenorhinus sp. nov., have been found exclusively in the Indian Ocean; and P. janetae sp. nov., P . foraminosus
sp. nov., and P. woljfi sp. nov., exclusively in the eastern Pacific. As result of this study, the genus now contains
17 species, of which P. pilosimanus Smith, 1879, P. nudus (A. Milne-Edwards, 1891), and P. alaminos Lemaitre. 1986,
are so far known only from the Atlantic Ocean. The bathymetric distribution of all species in the genus is summarized.
RESUME
Crustacea Decapoda: Revision des especes du genre Parapagurus Smith, 1879 (Parapagu¬
ridae) des oceans Indien et Pacifique.
Une revision des bernard-fermite du genre Parapagurus Smith, 1879, des oceans Indien et Pacitique est presentee.
Elleest basee sur les nombreuses nScoltes faites durant diverses expeditions fran?aises dans la region dc la Nouvelle-
Caledonie, complies par un abondant materiel se trouvant dans divers grands Museums et Institutions a travers
Lemaitre, R., 1999. — Crustacea Decapoda: A review of the species of the genus Parapagurus Smith, 1879
(Parapaguridae) from the Pacific and Indian Oceans. In: A. Crosnier (ed.). Resultats des Campagnes Musorstom.
Volume 20. Memoires du Museum national d'Histoire Naturelle , 180. 303-378. Paris ISBN: 2-85653-520-8.
Source : MNHN. Paris
304
R. LEMAITRE
le monde. Au total. 14 especes ont cte repertoriees dans les oceans Indien et Pacifique. Sept d'entre elles sont nouvcllcs :
P. richeri sp. nov., P. furici sp. nov., P. stenorhinus sp. nov., P. saintlaurentae sp. nov.. P. janetae sp. now. P . fora-
minosus sp. nov., et P. wolffi sp. nov.; et trois, P. abyssoriun (Filhol, 1885). P. bouvieri Stebbing. 1910. et P. andreui
Macpherson, 1984. sont <§galement presentes dans certaines parties de locean Atlantique. Les cspfcces nouvelles sont
decrites en detail et figurees, tandis qu’une diagnose est fournie pour les especes dejii connues, sauf dans le cas de
P. holthuisi Lemaitre, 1989. qui, insuffisamment definie jusqu^ present, est red6crite. Des informations sur la variability
des especes sont fournies pour les plus abondantes d’entre elles. Par ailleurs une cl 6 d'identi fication couvrant
les 14 especes est proposee. Parmi les sept nouvelles especes P. richeri sp. nov. et P. furici sp. nov. ont etc lrouv£es dans
la region n£o-caledonicnne mais egalement ailleurs dans lTndo-Pacifique; P. saintlaurentae sp. nov. et P. stenorhinus
sp. nov. n’ont ete nScoltSes que dans I'ocSan Indien, et P. janetae sp. nov., P. foraminosus sp. nov., et P. wolffi
sp. nov., que dans le Pacifique oriental. Apres cette revision, le genre Parapagurus comprend 17 especes. parmi lesquclles
P. pilosimanus Smith. 1879. P. nudus (A. Milne-Edwards, 1891), et P. alaminos Lemaitre, 1986. ne sont encore connues
que de 1'ocean Atlantique. La distribution bathymetrique de toutes les espfcces du genre est indiqu£e.
INTRODUCTION
Among the hermit crab species of the family Parapaguridae currently classified in ten genera (see Lemaitre.
1996), those of the genus Parapagurus Smith, 1879 are particularly difficult to define. The genus was revised by
Lemaitre (1989) and restricted to a group of ten very similar and morphologically variable species, including the
type of the genus, P. pilosimanus Smith, 1879. Species of this genus frequently live symbiotically with members
of the Anthozoa (zoanthids), typically have broad distributions, and are found at depths greater than 1000 m, deeper
than most other parapagurids. Morphologically, Parapagurus species are characterized primarily by the presence of
gills consisting of series of four filamentous or flattened branches arranged along the axis, reduced ocular peduncles
and corneae, considerable elongation of antennular and antennal peduncles, palm of right cheliped with rounded
mesial and lateral faces, and presence in males of well developed paired first and second pleopods (gonopods). Six
species occur in the Atlantic Ocean and have been discussed in detail by Lemaitre (1986, 1989, 1990):
P. pilosimanus , P. abyssorum (Filhol, 1885a), P. nudus (A. Milne-Edwards, 1891), P. alaminos Lemaitre, 1986,
P. bouvieri Stebbing, 1910, and P. andreui Macpherson, 1984. Of these. P. andreui and P. bouvieri have been
reported also from the southwestern Indian Ocean (Lemaitre, 1990). One species, P. latimanus Henderson, 1888,
known until now only from off New Zealand, was discussed by Lemaitre and McLaughlin (1992). However,
the remaining species of the genus, believed to be distributed exclusively in the Indo-Pacific or eastern Pacific,
have remained poorly defined.
During a study of the extensive parapagurid material obtained by various French expeditions to New Caledonia
and adjacent waters, two very abundant species of Parapagurus suspected to be new were found. Comparison with
types and numerous supplemental specimens of previously known species deposited in many museums and institu¬
tions, confirmed that the two species from New Caledonia were undescribed. The study of numerous specimens de¬
posited in museums and institutions also revealed the existence of four additional undescribed species that had been
confounded with previously known species, or in some cases misidentified. In light of the striking similarities
observed among species of this genus, and their frequent high degree of overlap in their ranges of morphological
variations, it became clear that descriptions of the new species required detailed comparisons with other species
from the Indo-Pacific and eastern Pacific regions. Given that most of the previously knowrn species were poorly
defined, it was necessary to present a review of all species of the genus from the entire Indian and Pacific Oceans.
As result of this study, 14 species of Parapagurus are now recognized from the Pacific and Indian Oceans, seven
of which arc new. Of the 14, nine are found in the Indo-Pacific, three of which are present in the New Caledonia
region, and five exclusively in the eastern Pacific. The new species are described, and those previously known arc
diagnosed. One obscurely defined species, P. holthuisi Lemaitre, 1989, is redescribed; the name of this species was
proposed by Lemaitre (1989) as a replacement for P. abyssorum Henderson, 1888, which is a junior homonym
of P. abyssorum (Filhol, 1885a). Figures are included for all species based on specimens examined during this
study or, in some cases, on published illustrations. In the case of species where abundant material has been
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
305
available, a section on morphological variations is included. A key to all the species that occur in the Pacific and
Indian Oceans is presented as an aid in the identification, and is designed to be used in combination with the
descriptive text and illustrations.
Species of Parapagurus are in most cases distinguished by a number of subtle diagnostic characters. Particularly
useful are characters derived from the ocular and antennal acicles, ambulatory legs, propodal rasp of fourth
pereopod, exopod of uropods, and telson. Some species can be separated also by marked differences in the
proportions of the segments (i.e., merus, propodus, dactyl) of the ambulatory legs or those of the left exopod of
the uropods. In addition, the relative size or proportions of other structures, such as the shield, and ocular and
antennular peduncles, can frequently be helpful in determining the identity of specimens. In contrast, other
structures, such as the mouthparts, are virtually identical in all species, and only minor differences can be observed
in the number of setae on the distal end of the endopod of the maxillule, or number of teeth of the crista dentata of
the third maxilliped. However, only a limited number of specimens have been dissected, and whether or not these
differences are stable cannot be assured. A full set of mouthparts is illustrated as an example for only one species,
P. richeri sp. nov.
The right chelipeds of Parapagurus species from the Pacific and Indian Oceans exhibit a range of morphological
variations very similar to those described for the Atlantic representatives of the genus (LEMAITRE, 1986, as the
P. pilosimanus complex). Especially prevalent on this chelipcd is variability in the dimension and sometimes
armature of segments, which is attributable to size and sexual dimorphism. Such variability makes characters
derived from this cheliped virtually useless for diagnostic purposes at the specific level. The often striking
elongation of carpus and palm is a predictable condition that has been observed in virtually every species in which
enough specimens of different sizes are available. The spines on the surfaces of carpus and palm of this cheliped are
frequently smaller, and less densely arranged, in larger males.
The general hermit crab terminology used in this study follows McLaughlin (1974), with the exception of
the condition of the fourth pereopod, where subchelate is used following the del inition ol McLaughlin (1997:
435). Terms for parapagurid morphology are used according to the definitions provided by LEMAITRE (1989),
although the second and third pereopods are here referred to as the first and second ambulator y legs respectively.
The type of gill in species of Parapagurus has previously been described as trichobranchiate (e.g., SMITH, 1882;
de Saint Laurent, 1972; Lemaitre, 1989). Recently, however, McLaughlin and de Saint Laurent (1998:
161, fig. 1) have shown that gill type is not determined by shape of the gill elements, but by the arrangement and
insertion of the elements on the rachis of the gill, and pointed out that there arc many types of true trichobranch
and phyllobranch gills. In true trichobranchiate gills the tubular elements are inserted in order or disorder around the
axis, or in regular transverse rows along the axis. Accordingly, the gills in species of Parapagurus , which consist
of series of four filamentous elements arranged around the axis (see Lemaitre, 1989: 8, tig. 2L-M), are not true
trichobranchiae. McLaughlin and DE Saint Laurent (1998) defined the term " quadriseriaV for a gill structure
equivalent to Lemaitre's (1989) "trichobranchiate" and "intermediate" conditions used for the Parapaguridae. The
term "quadriserial" is adopted here to describe the gills of Parapagurus species, and is considered a type of
phyllobranch gill.
The measurements used for different structures are defined in Fig. 1, and should be taken to the nearest
0.1 mm. In the Material examined sections, one measurement indicative of size, i.e. length of shield, is
included following the number and sex of specimens. The section on SIZE RANGE for each species is based on the
materials used for this study as well as from all available materials previously reported by Lemaitre (1986. 1989,
1990, 1997) and Lemaitre and McLaughlin (1992).
The shape of the scales on the propodal rasp of the fourth pereopod is of considerable diagnostic importance.
The shape of the scales as well as the number of rows of scales can best be observed in a ventrolateral view.
Almost invariably, the proximal portion of the rasp exhibits more rows than the distal portion; it is the number ol
rows on the distal portion that is most useful in the identification of specimens. Examples of ovate, conical , and
lanceolate scales are shown in Fig. 2 (sec also micrographs in Lemaitre. 1986: 534, fig. 5). The ovate (Fig. 2a-
b) and conical (Fig. 2c) type of scales are easily distinguished; the lanceolate scales (Fig. 2d-e) are narrow, and
terminate in curved margins tapering to a pointed or blunt end.
306
R. LEMAITRE
Fig. 1. — Diagrammatic Parapagurus , showing measurements used in text, a, shield and cephalic appendages: 1, ocular
peduncle length; 2, ocular acicle width; 3, shield length; 4, shield width, b. cheliped: 5, dactyl length (from proximal
end to tip of dactyl); 6, palm length; 7, palm width (maximum); 8. carpus length; 9. carpus width (maximum), c,
ambulatory leg: 10, dactyl length; 11, propodus length (dorsal margin, excluding proximal portion curving down to
carpus); 12, propodus height; 13, mcrus length; 14. merus height, d, propodus and dactyl of fourth pereopod: 15,
propodal rasp length; 16, propodus height, e, left exopod of uropods: 17, exopod length; 18, exopod width.
The core of the materials used for this study remain deposited in the Museum national d'Histoire naturelle,
Paris (MNHN), the National Museum of Natural History, Smithsonian Institution, Washington D.C. (USNM),
and the Zoologisk Museum, Copenhagen (ZMK). However, numerous other materials, including types, deposited
in other museums and institutions throughout the world were also used, and are abbreviated as follows:
Source : MNHN , Paris
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
307
AM. — Australian Museum, Sydney, Australia.
CBM. — Natural History Museum and Institute, Chiba, Japan.
ICM. — Instituto dc Ciencias del Mar (formerly Instituto de Investigaciones Pcsqueras). Barcelona, Spain.
IORAN. — Institute of Oceanology, Russian Academy of Sciences.
LACM. — Natural History Museum of Los Angeles County, USA.
MCZ. — Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA.
NHM. — The Natural History Museum [formerly British Museum (Natural History)], London, England.
NMV. — National Museum of Victoria, Melbourne, Australia.
NMNZ. — Museum of New Zealand Te Papa Tongarewa (formerly National Museum of New Zealand),
Wellington.
NTOU. — National Taiwan Ocean University, Keelung, Taiwan.
QM. — Queensland Museum, Brisbane, Australia.
SAM. — South African Museum, Cape Town.
SAMA. — South Australian Museum, Adelaide, Australia.
SIO. — Scripps Institute of Oceanography Invertebrate Collection, University of California, San Diego, USA.
SMF. — Senckenberg Museum, Frankfurt a. M., Germany.
ZM A. — Zoologisch Museum, Amsterdam, The Netherlands.
ZMUM. — Zoological Museum, Moscow State University.
ZRC. — Zoological Reference Collection, Department of Zoology, National University of Singapore.
Other abbreviations used are: immat, immature (sex undetermined); 6, male(s); 9, female(s); stn. station;
CP, Waren dredge; DW, beam trawl; CC, shrimp trawl; JCU, James Cook University, Townsville, Australia;
NZOI, New Zealand Oceanographic Institute (now part of National Institute of Water and Atmospheric Research),
Wellington.
The specimens examined are listed by geographic area, and within areas by cruise, vessel, and station number.
Information on stations is listed following the format of the original cruise data, that is, longitudes and latitudes
arc cited in degrees and decimals, or degrees and fractions of minutes, all depths are cited in meters.
Fig. 2. — Portion of propodal rasp of fourth pereopod, showing examples of shapes of scales referred to in text:
a-b, ovate; c, conical; d-e. lanceolate, (a, lateral view; b-e, ventrolateral view).
Source
308
R. LEMAITRE
SPECIES LIST
(Asterisk indicates species found in New Caledonia region)
P. latimanus Henderson, 1888*
P. abyssorum (Filhol, 1885a)
P. bouvieri Stebbing, 1910
P. andreui Macpherson, 1984
P. microps de Saint Laurent, 1972
P. benedicti dc Saint Laurent, 1972
P. holthuisi Lemaitre, 1989
P. riche ri sp. nov.*
P. fund sp. nov.*
P. saintlaurentae sp. nov.
P. stenorhinus sp. nov.
P. janetae sp. nov.
P. foraminosus sp. nov.
P. wolffi sp. nov.
SYSTEMATIC ACCOUNT
Family PARAPAGURIDAE Smith, 1882
Genus PARAPAGURUS Smith. 1879
Parapagurus Smith, 1879: 50. — de Saint Laurent, 1972: 101 (in part). — Lemaitre, 1989: 11, fig. 2A-C. L-M.
Type Species. — Parapagurus pilosimanus Smith, 1879 by monotypy. Gender: masculine.
DIAGNOSIS. — Eleven pairs of quadriserial gills, lacking vestigial pleurobranchiae on last thoracic somite;
gills each consisting of series of 4 long filamentous or flattened branches arranged along axis. Shield usually well
calcified. Ocular peduncles (including corneae) typically half or less than half length of shield; corneae at most
weakly dilated. Antennal peduncles and acicles distinctly overreaching ocular peduncles. Fourth segment of
antennal peduncle unarmed. Epistomial spine, when present, short and straight. Right cheliped elongate; palm
rounded mesially and laterally. Left cheliped well calcified. Ambulatory legs long, dactyls curved. Fourth pereopod
with propodal height subequal or greater than length of margin occupied by propodal rasp (e.g., Fig. Id). Second
abdominal somite with left plcuron terminating ventrally in small subtriangular lobe. Males with well developed
paired first and second pleopods; distal lobe of first pleopod subconical or subtubular; distal segment of second
pleopod slightly twisted distally, anterior face with numerous setae distally and row ot short stiff setae on lateral
margin. Females with rudimentary right second pleopod.
Distribution (Figs 47-50). — Worldwide. Depth: 82 to 5020 m.
Species. — In addition to the species reported in this study (see Species List), three others known only from
the western and eastern Atlantic are included in this genus, P. pilosimanus Smith, 1879, P. alaminos Lemaitre,
1986, and P. nudus (A. Milnc-Edwards, 1891). As previously mentioned, these three were discussed in detail by
Lemaitre (1989).
Key to species of Parapagurus from the Pacific and Indian Oceans
(Asterisk indicates species found in New Caledonia region)
1. Lateral faces of meri, carpi and propodi of ambulatory legs with spines, or with small
rounded shallow pits . 2
— Lateral faces of meri, carpi and propodi of ambulatory legs lacking spines or pits . 4
2. Lateral faces of meri, carpi and propodi with small rounded shallow pits (Fig. 41 ) .
. P. foraminosus sp. nov. (eastern Pacific)
— Lateral faces of meri, carpi and propodi lacking pits, with spines . 3
Source : MNHN, Paris
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
309
3. Anterodistal margin of branchiostegite armed wilh small spines; propodal rasp ot fourth
pereopod consisting of 2 or 3 rows of lanceolate scales .
. P. abyssorum (Atlantic; western and southeastern Pacific)
— Anterodistal margin of branchiostegite unarmed; propodal rasp of fourth pereopod
consisting of 1 row (at least distally) of ovate scales . P. nticrops (eastern Pacific)
4. Lateral faces of meri of ambulatory legs weakly calcified medially, weak calcification
more pronounced on second (Fig. 7d-f); length of ocular peduncles (including corneae)
distinctly more than half length of shield .
. P. bouvieri (southeastern Atlantic; Indo-Pacific)
— Lateral faces of meri of ambulatory legs well calcified; length of ocular peduncles
(including corneae) half or less length of shield . 5
5. Ventral margins of meri of ambulatory legs armed with spines (stronger on second leg)
. 6
— Ventral margins of meri of ambulatory legs unarmed . 7
6. Dorsal margins of carpi of ambulatory legs armed with spines (Fig. 45); propodal rasp of
fourth pereopod with 1 row of ovate scales (at least distally); shield about as broad as
long; ocular acicles terminating in simple spine .... P. wolffi sp. nov. (eastern Pacific)
— Dorsal margins of carpi of ambulatory legs unarmed; propodal rasp of fourth pereopod
with 2-4 rows of ovate scales (at least distally); shield distinctly broader than long; ocular
acicles most frequently terminating in bifid spine . P. benedicti (eastern Pacific)
7. Propodal rasp of fourth pereopod consisting of ovate scales (Fig. 2a-b) . 8
— Propodal rasp of fourth pereopod consisting of conical (Fig. 2c) or lanceolate (Fig. 2d-e)
scales . I*
8. Left exopod of uropods broad, 2.3 times or less as long as broad . 9
— Left exopod of uropods elongate, more than 2.3 times as long as broad . 1 0
9. Terminal margin of telson with rounded projections each armed with more than
12 closely-spaced spines; mesial margin of antennal acicles armed with 5-8 weak spines
(Fig. 32a-b); left exopod of uropod wilh narrow rasp (Fig. 35e-g) .
. P. stenorhinus sp. nov. (Indian Ocean)
— Terminal margin of telson with rounded projections each armed with less than 12 well-
spaced spines; mesial margin of antennal acicles armed with 1-8 distinct spines
(Fig. 19a-c); left exopod of uropod with moderately broad rasp (Fig. 23f,h) .
. P. richeri sp. nov.* (Indo-Pacific; central Pacific)
10. Propodus of second ambulatory leg at most 4.3 times as long as high, merus at most
3 times as long as high . P. janetae sp. nov. (eastern Pacific)
— Propodus of second ambulatory distinctly more than 4.3 times as long as high, merus
distinctly more than 3 times as long as high . P. holthuisi (central and eastern Pacific)
1 1. Propodus of first ambulatory leg 5 or more times as long as high . 1 2
— Propodus of first ambulatory leg less than 5 times as long as high . 13
12. Terminal margin of telson divided into 2 rounded projections by wide, frequently deep,
rounded (U-shaped) cleft; anterodistal margin of branchiostegite armed with 1 or more
small spines (Fig. 28e); antennal acicles slender, exceeding eyes by more than half length
of acicle . P. saintlaurentae sp. nov. (Indian Ocean)
— Terminal margin of telson divided into 2 rounded projections by narrow, angled (V-
shaped) cleft; anterodistal margin of branchiostegite unarmed (Fig. 24b); antennal acicles
moderately slender, exceeding eyes by half or less than half length of acicle .
. P. furici sp. nov.* (Indo-Pacific)
Source : MNHN. Parts
310
R. LEMAITRE
13. Propodi of first and second ambulatory legs more than 4 times as long as high; mesial
margin of antennal acicles unarmed .
. P. andreui (southeastern Atlantic; southwestern Indian Ocean)
— Propodi of first and second ambulatory legs less than 4 times as long as high; mesial
margin of antennal acicles armed with up to 6 small spines .
. P. latimanus* ( I ndo- Pacific)
Parapagurus latimanus Henderson. 1888
Figs 3. 47-48
Parapagurus latimanus Henderson. 1888: 91, pi. 9. fig. 2 (type locality: "Challenger", stn 167A, New Zealand). —
Murray. 1895: 597. — Alcock, 1905: 172. — Gordan, 1956: 338 (lit.). — Lemaitre, 1986: 526; 1989: 11; 1997:
575. — Lemaitre & McLaughlin, 1992: 762. fig. 9.
Parapagurus pilosimanus pilosimanus - DE Saint Laurent, 1972: 102 (in part, see Remarks).
Parapagurus pilosimanus latimanus - DE SAINT LAURENT, 1972: 103, pi. I, fig. 5.
? Parapagurus pilosimanus - Takeda, 1982: 65, unnumbered color fig. (see Remarks).
Material examined.
Japan. RV " Tansei-Maru ": off Taito-saki, Boso Peninsula. 35°07.8'N, 140°49'E, 400-416 m, 26.04.1995.
coll. T. Komai: 1 ov. 9 13.3 mm (CBM-ZC 2038).
Indonesia. "Galathea", stn 453, Makassar Strait, 3°65'S. 118°26'E. 2034 m, 24.08.1951: 1 8 6.6 mm (ZMK).
New Caledonia. Bathus 3: stn CP 844, 23°06’S. 166°45’E. 908 m, 1.12.1993: 1 8 6.6 mm, 2 ov. 9 4.0, 4.1 mm
(MNHN-Pg 5575).
Volsmar: stn DW 25, 22°22,80’S, 171°21,50'E, 940 m, 4.06.1989: 1 ov. 9 6.6 mm (MNHN-Pg 5576).
Vanuatu. Musorstom 8: stn CP 1075, 15°33’S, 167°27'E, 956-944 m, 4.10.1994: 1 8 5.6 mm (MNHN-Pg 5579).
Wallis and Futuna. MUSORSTOM 7: stn CP 564, 1 1°46.1’S, 178°27.4'W. 1015-1020 m, 20.05.92: 1 8 11.2 mm
(MNHN-Pg 5577). — Stn DW 620, 12°34.4'S. 178°11.0’W, 1280 m. 28.05.92: 1 8 10.3 mm (MNHN-Pg 5578).
Australian region. New South Wales. FRV "Kapala": stn K77-23-12. E of Broken Bay, 33°35-33'S, 152°00-02'E,
823 m, 8.12.1977: I 9 9.0 mm (AM P52737); Stn K80-20-06. between Sydney and Newcastle, 33°38'S, 152°02’E, 960-
988 m, 9.12.1980. coll. R.T. Springthorpe: 1 8 5.5 mm (AM P40411); Stn K83- 12-02. E of Eden. 37°36'S. 150°21E.
860-960 m, 26.09.1983: l 8 7.8 mm (AM P40396). — ORV "Franklin": stn SLOPE 58, 56 km ENE of Nowra,
34°43.95'S, 151°14.74'E, 1009-817 m, 22.10.1988: 1 8 7.9 mm, 1 9 7.8 mm (NMV J44913), 2 8 4.8, 4.9 mm, 1 9
6.3 mm, 1 ov. 9 7.5 mm (USNM 276125).
Tasmania. ORV "Franklin": stn SLOPE 81, 48 km ENE of Cape Tourville. 42°00.25’S, 148°43.55’E, 1264 m,
30.10.1988, coll. G.C.B. Poore: 1 9 7.2 mm (NMV J40403).
Tasman Sea. ORV "Franklin": Fro 05/89 stn 17. Lord Howe Rise, 29°42.06'S. 159°48.3rE, 2450 m, 3.05.1989: I 8
5.7 mm (AM P40432).
Great Australian Bight. "Galathea": stn 554. 37°28'S, 138°55’E, 1320-1360 m, 5.12.1951: 4 8 3.3-9.5 mm, 5 9
6. 3-7. 5 mm (ZMK); 1 9 8.2 mm (ZMK). — "Dmitry’ Mendeleev": stn DM 1373, 33°49-46'S, 127°9-27'E. 1080-1100 m,
28.02.1976: 6 8 5.6-10.0, 2 9 5.2, 7.3 mm, 2 9 ov. 7.0, 8.0 mm (AM P21966). 3 8 5.7-10.0 mm (NMV J 16194). —
FV "Saxon Progress": -120 nautical miles (222.2 km) SW of Cape Adieu. 33°58’S, 131°22’E, 1000 m. Nov 1989, coll.
D.W. LEENAN: I 8 7.0 mm [in zoanthid Epizoanthus incrustatus (Duben & Koren)] (SAM A C5823). — FV "Longra III":
stn 84, -120 nautical miles (222.2 km) SSE of Euda, 33°37’S. 129°53’E. 930-1030 m, 17.04.1990, coll. K. Gowlett-
Holmes: 1 8 14.8 mm (in zoanthid E. paguriphilus Verrill) (SAMA C5824), 1 9 11.9 mm (in zoanthid E. incrustatus)
(SAMA C5825).
New Zealand. "Shinkai Maru": cruise II, stn 77, northern Chatham Rise, 42°47.5'S, 178°22.0'E. 939-920 m,
15.11.1975: 1 8 8.9 mm (NMNZ Cr 3198). — RV "Acheron": stn BS 353, 37°30'S, I79°22'E, 1134-1207 m,
7.02.1974: 1 8 15.9 mm (NMNZ Cr 3202), 1 9 7.6 mm (NMNZ Cr 8431 ), 1 8 9.6 mm. 1 9 9.1 mm, 1 ov. 9 12.2 mm
(NMNZ Cr 8477). — RV "Tangaroa": BS 690, stn R48. 37°22.1’S, 178°26.9'E, 2027-1952 m. 18.01.1979: 1 8 8.9 mm
(NMNZ Cr 3200); BS 760, stn R1I8. -22 km E of Alderman Islands, 36°57.3’S, 176°21.5’E, 803-846 m, 24.01.1979:
2 ov. 9 1 1.5, 12.2 mm (NMNZ Cr 3199); BS 771, stn R129, -39 km E of Portland Island. Mahia Peninsula, 39°15.4’S.
178°19.3'E. 413-453 m, 26.01.1979: 1 ov. 9 1 1 .0 mm (NMNZ Cr 3201). — RV "James Cook": stn J6/1 2/8 1 , SE of East
Cape. 37°46.3'S, 178°57.4'E, 829-928 m, 16.04.1981: I <3 11.6 mm (NMNZ Cr 8484); Stn J19/9/84. Challenger
Plateau, 40°06.3'S, 167°57.9’E, 960-982 m. 13.11.1984: 1 8 8.2 mm, I ov. 9 8.3 mm (NMNZ Cr 8452);
Stn J 19/01 1/84, Challenger Plateau, 40°03.2’S, 167°53.3'E. 912-940 m, 13.11.1984: 2 8 10.3, 13.4 mm (NMNZ Cr
8447); Stn J9/42/89, 39°29.5'S. Ritchie Bank, Hawkcs Bay, 178°18.9'E, 823-870 m, 27.09.1989: I 8 15.8 mm
(NMNZ Cr 8485); Stn J9/49/89, E of Ritchie Bank, 39°32.8'S, 178°I6.5’E, 880-857 m, 29.09.1989: 1 8 10.6 mm
Source : MNHN. Pahs
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
311
(NMNZ Cr 8453). — FV " Kalinovo ": stn K/40/81, near Antipodes Islands. 40°51.5*S, 176°57.9'E, 1125-1 150 m.
28.11.1981: 2 6 8.5, 10.0 mm (NMNZ Cr 8429). — RV "Washington": stn Rock Dredge 4KD5), 34°28.8’S,
178°52.3'E. 2500 m. 12.02.1986: 1 ov. 9 12.2 mm (SIO C9546). — FV "Arrow": sin A01/37/87. Ritchie Bank. Harkes
Bay. 39°24.5'S, 178°19.9'E, 921-981 m, 26.06.1987: 1 ov. 9 10.9 mm (NMNZ Cr. 4819); Stn A01/41/87, 39°38.5’S.
178°19.9’E, 906-924 m, 26.06.1987: 1 6 13.0 mm (NMNZ Cr 4820). — FV "Cordelia": stn CO 1/23/88, N of Chatham
Islands. 42°48.rS, 176°47.7’E, 975-974 m, 16.07.1988: 2 ov. 9 11.0. 11.1 mm (NMNZ Cr 6052). — FV "Amaltal
Explorer": stn A EX 2/11/89, N of Antipodes Islands, 48°31.56'S, 179°03.irE, 660-675 m, 20.11.1989: 1 6 16.0 mm
(NMNZ Cr 8425).
Western Indian Ocean. Off Kenya. "Galathea": stn 241, 4°00'S, 41°27'E, 1510 m. 15.3.1951: 1 6 8.9 mm
(ZMK).
TYPES. — Holotype: 6 6.6 mm, New Zealand, " Challenger ", stn 167A, 18 m (depth questionable, see
Lemaitre & McLaughlin, 1992: 764), 27.06.1874 (NHM 1888:33).
DIAGNOSIS. — Shield (Fig. 3a) about as broad as long, dorsal surface well calcified; lateral projections broadly
rounded. Rostrum broadly rounded, with short mid-dorsal ridge. Ocular peduncles (including corneae) less than half
length of shield, inflated basaliy; width of cornea about same as distal width of ocular peduncle. Ocular acicles
subtriangular, terminating in simple strong spine (rarely bifid on one or both sides). Antennular peduncle
exceeding eyes by nearly entire length of penultimate segment; lateral face of basal segment with statocyst lobe
having subrectangular distal lobe armed with 2 spines, and 1 spine proximally. Antennal peduncle (Fig. 3b)
exceeding eyes by nearly entire length of fifth antennal segment; flagellum with numerous setae 1-4 flagellar
articles in length; acicle straight or weakly curved in dorsal view, exceeding distal margin of cornea by half length
of acicle, with proximal half of mesial margin armed with 1 to 6 small blunt to sharp spines or tubercles (acicle
rarely unarmed). Epistomial spine usually absent. Slernite of third maxillipeds with strong spine on each side of
midline. Left cheliped (Fig. 3c) well calcified, densely setose; carpus with irregular row of small spines or
tubercles on dorsal margin. Ambulatory legs (Fig. 3e-f) with meri, carpi and propodi unarmed except for small
dorsodistal spine on each carpus; meri each about 3.3 (first leg) or 2.9 (second leg) times as long as high. Anterior
lobe of sternite of second ambulator)' legs subsemicircular, setose, armed with small subterminal tubercle or spine.
Fourth pereopod (Fig. 3g-h) with propodal rasp consisting of 2 (rarely 3) often irregular rows of lanceolate or
conical scales. Fifth pereopod with propodal rasp less than half length of propodus. Telson and uropods (Fig. 3i-j)
asymmetrical. Terminal margin of telson divided into 2 rounded projections by angled (V-shaped) cleft; rounded
projections armed distally with moderately long, evenly-spaced corneous spines (approximately 15 to 26 left, 8 to
13 right), spines on left side frequently extending anteriorly nearly to midlength of lateral margin of telson. Left
exopod (Fig. 3i) of uropod elongate, about 2.8 to 3.0 times as long as broad; with broad rasp.
Size Range. — Males, SL 3.3 to 16.0 mm. Females 5.2 to 1 1.9 mm. Ovigcrous females 4.0 to 13.3 mm.
Variations. — The spines on the dorsomedian surface of the palm of the right cheliped can be arranged
irregularly, or frequently in more or less straight rows.
Habitat. — Usually found living in shelters formed by zoanthids, probably species of Epizoanthus [e.g.
E. paguriphilus Verrill, E. incrustatus (Duben & Koren)].
DISTRIBUTION (Figs 47-48). — Indo-Pacific: Japan; Indonesia; New Caledonia region; southern Australia; New
Zealand. Western Indian Ocean: off Kenya. Depth: 400 to 2500 m.
Affinities. — This species resembles P. andreui and to a lesser extent P. pilosimanus. The three are usually
found living in shelters associated with zoanthids (Epizoanthus spp.). There are differences in distribution and
morphology among the three. Parapagurus latimanus occurs in the Indo-Pacific, P . andreui in the southeastern
Atlantic and southwestern Indian Ocean, and P. pilosimanus only in the Atlantic. Morphological characters that
differentiate P. latimanus from P. andreui can be found in the antennal acicles (armed in P. latimanus , unarmed in
P. andreui ); the scales of the propodal rasp of the fourth pereopod (lanceolate in P. latimanus , conical in
P. andreui ); and the meri of the ambulatory legs (2.9 to 3.3 times as long as high in P. latimanus , 3.5 to 4 times
as long as high in P. andreui). The shape of the telson can serve to separate P. latimanus from both P. andreui and
Source :
312
R. LEMAITRE
Fig. 3. — Parapagurus latimanus Henderson, 1888, "Eltanin", stn 2198, 43°48,S, 174°24’W: 6 7.5 mm (USNM 256845).
a, shield and cephalic appendages; b, right antennal peduncle, lateral view; c, left chela and carpus (most setae
omitted); d, right chela and carpus (setae omitted); e, right first ambulatory leg, lateral view; f. right second
ambulatory leg, lateral view; g, propodus and dactyl of right fourth pereopod. lateral view; h. propodal rasp of same
(setae omitted), ventrolateral view; i, left exopod and endopod of uropods, dorsal view; j, telson, dorsal view.
Scales equal 3 mm (a), 1 mm (b), 2 mm (c,i), 3 mm (d), 5 mm (e-f), 1 mm (g.j). and 0.5 mm (h). (From Lemaitre
& McLaughlin, 1992).
Source : MNHN. Paris
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
313
P. pilosimanus. The terminal margin of the telson is divided into two asymmetrical rounded projections by a broad
cleft in P. I at i man us and P. andreup the terminal margin is divided into two symmetrical rounded projections by a
narrow cleft in P. pilosimanus.
Remarks. — De Saint Laurent (1972: 103) questionably included the western Indian Ocean in the
distribution of Parapagurus pilosimanus pilosimanus. based on material from the Galathea expedition collected at
stn 241, off Kenya. That material has been examined and found to represent P . latimanus.
Takeda (1982) published a color photo of a specimen identified as Parapagurus pilosimanus. However, as
previously mentioned, that species occurs only in the Atlantic Ocean (LEMAITRE, 1989). Takeda's specimen has
not been available for examination, and from the photograph alone, the identity of the specimen cannot be
determined with certainty. It is questionably assigned to P. latimanus.
Parapagurus abyssorum (Filhol. 1885a)
Figs 4-6, 47, 49-50
Parapagurus pilosimanus - SMITH, 1884: 354 (in part); 1886: 607 (in part). — A. MiLNE-EDWARDS & Bouvier. 1892a: 1
(in part); 1892b: 204 (in part). (See Remarks).
Pagurus abyssorum Filhol, 1885a: 152, fig. 1; 1885b: 131, fig. 41. (See Remarks).
Parapagurus abyssorum - HENDERSON, 1888: 87 (in part), not pi. 9, fig. 2 (= Parapagurus holthuisi Lemaitre, 1989). —
Wood-Mason & Alcock, 1891: 199. — Alcock, 1894: 242. — Murray, 1896: 388 (in part). — Gordan, 1956:
337 (lit.). (See Remarks).
Parapagurus abyssorum var. scabra Henderson, 1888: 89. pi. 9, fig. 3. — MURRAY, 1895: 257. ALCOCK. 1 905: 17_. —
Gordan, 1956: 338 (lit.).
Parapagurus pilosimanus Var. Scabra - A. MiLNE-EDWARDS & BOUVIER, 1892a: 2.
Parapagurus scabra - A. Milne-Edwards & Bouvier, 1892a: 13.
Parapagurus pilosimanus var. abyssorum - A. Milne-Edwards & Bouvier. 1892a: 13; 1892b: 205; 1899: 55, pi. 1.
fig. 1; 1900: 191, pi. 24, figs 4-6. — Alcock, 1905: 172. — Gordan, 1956: 338 (lit.).
IParapagurus pilosimanus - Porter, 1906: 129. — Haig, 1955: 17. (See Remarks).
Parapagurus pilosimanus Var. Abyssorum - NOBRE, 1931: 201, fig. 110; 1936: 126, lig. 103.
Parapagurus pilosimanus scaber - DE Saint Laurent. 1972: 102, pi. 1, fig. 3; 1985: 475. — Miyake, 1978: 72 (lit.);
1982: 196 (lit.). (See Remarks).
Parapagurus scaber - Lemaitre, 1986: 533, figs 1G-H, 3F-J. 41-J. 5G-H. 6A-C.K-L. 7D.H-1. 8F-G, 9C.
Parapagurus abyssorum - LEMAITRE, 1989: 30, figs 5D-E, 12-14; 1990: 220. - Ingle, 1993: 19.
Not Parapagurus pilosimanus abyssorum - Faxon, 1895: 68 (= Parapagurus foraminosus sp. nov., see Remarks under that
new species).
Fig. 4. — Parapagurus abyssorum (Filhol, 1885a): 9 holotype figured by Filhol (1885a: 132. fig. 1).
314
R. LEMAITRE
Fig. 5. — Parapagurus abyssorum (Filhol, 1885a): a-c, 9 11.2 mm ."Challenger", stn 68. North Atlantic (NHM 1888:33);
d. North Atlantic, a. shield and cephalic appendages; b, left chela and carpus (setae omitted); c, right chela and carpus
(setae omitted); d. telson. dorsal view.
Scales equal 5 mm (a-c), and 2 mm (d). (a-c. from Lemaitre, 1989; d, from Lemaitre, 1986).
Material EXAMINED. — North Atlantic. 'Talisman": stn 148, 42°23'N, 21°15’W. 4010 m. 24.08.1883: 1 9.
holotype of Pagurus abyssorum.
"Challenger": stn 68, 38°03’N, 39°I9'W, 3915 m. 24.06.1873: 1 9 11,2 mm. holotype of Parapagurus abyssorum
var. scabra (NHM 1888: 33).
Tasman Sea. "Galathea": stn 575, 40°H'S, 163°35'E. 3710 m, 19.12.1951: 1 6 17.7 mm (ZMK).
Southeastern Pacific. "Elianin": cruise 21, stn 233, 39°5FS. 96°52’W, 3603-3621 m, 7.12.1965: 2 6 6.6.
1 1.3 mm (USNM 155046).
Types. — Pagurus abyssorum. Holotype: 9 figured by FlLHOL (1885a: 152, fig. 1), North Atlantic,
"Talisman ", stn 148, 42°23’N. 21°15’W, 4010 m, 24.08.1883. Not located.
Parapagurus abyssorum var. scabra. Holotype : 9 1 1,2 mm. North Atlantic, " Challenger ", stn 68, 38°03’N.
39°19'W, 3915 m, 24.06.1873 (NHM 1888: 33).
Source : MNHN. Paris
PARAPAGUR1DAE FROM THE PACIFIC AND INDIAN OCEANS
315
Diagnosis. — Shield (Fig. 5a) about as broad as long, dorsal surface well calcified; lateral projections broadly
rounded. Rostrum broadly subtriangular, rounded distally and with low mid-dorsal ridge. Antcrodistal margin of the
branchiostegite armed with small spines. Ocular peduncles (including corneae) less than half length of shield,
inflated basally; width of cornea about same as distal width of ocular peduncle. Ocular acicles subtriangular,
terminating in strong simple spine. Antennular peduncle exceeding distal margin of cornea by nearly entire length
of penultimate segment; lateral face of basal segment with statocyst lobe having subrectangular distal lobe armed
with 1 or more small spines, and 1 spine proximally. Antennal peduncle exceeding distal margin of cornea by
nearly entire length of fifth antennal segment; flagellum with few setae about 1 flagellar article in length or less;
acicle nearly straight in dorsal view, exceeding distal margin of cornea by half or more than half length of aciclc,
mesial and dorsomcsial distal margin armed with 5-25 small spines. Epistomial spine usually absent. Sternite of
third maxillipeds with strong spine on each side of midline. Left cheliped (Fig. 5b) well calcified, with moderately
dense setation; carpus with tubercles or spines on lateral and dorsal faces. Ambulatory legs (Fig. 6a-b) very
spinose; dactyls each with dorsal row of small spines; meri, carpi, and propodi each armed on mesial, lateral,
view; e, left exopod of uropods, dorsal view.
Scales equal 10 mm (a), 3 mm (b), 5 mm (c), 2 mm (d), and 1 mm (e).
316
R. LEMAITRE
dorsal, and ventral faces with numerous small spines (less numerous in small specimens SL < 7.0 mm. e.g..
Fig. 6b); meri each about 4.1 (first leg) or 3.5 (second leg) times as long as high. Anterior lobe of stcrnite of
second ambulatory legs (Fig. 6c) subsemicircular, setose, armed with 1-5 small subterminal spines. Fourth
pereopod (Fig. 6d) with propodal rasp consisting of 2-3 irregular rows of lanceolate scales. Fifth pereopod with
propodal rasp less than half length of propodus. Telson (Fig. 5d) and uropods asymmetrical. Terminal margin of
telson divided into 2 rounded projections by shallow, rounded (U-shaped) cleft; rounded projections armed distal ly
with short corneous spines. Left exopod (Fig. 6e) of uropod elongate, 2.2 to 2.8 times as long as broad; with
broad rasp.
Size Range. — Males, SL 7.4 to 17.7 mm. Females 7.3 to 13.5 mm. Ovigerous females 8.0 to 13.8 mm.
Distribution (Figs 47, 49-50). — North Atlantic, including northeastern coast of the United States; in the
eastern Atlantic, from the Azores to Cape Verde Islands. In the western and southeastern Pacific known only from
single specimens, one from the Tasman Sea and one from approximately 1930 km west of Chile. Depth: 2500 to
4360 m.
AFFINITIES. — This species is similar to Parapagurus microps in the spination of the lateral faces of the meri,
carpi, and propodi of the ambulatory legs. The two, however, can easily be distinguished primarily by the anterior
margin of the branchiostegite, and the propodal rasp of the fourth pereopod. as well as overall size (shield length)
and geographical distribution. The anterodistal margin of the branchiostegite is armed with small spines in
P. abyss or uni, whereas the anterodistal margin lacks spines in P. microps. The propodal rasp of the fourth
pereopod has two or three rows of lanceolate scales in P. abyssorunr, the rasp has one row (at least distally) of
ovate scales in P. microps. In addition, P. abyssorum reaches a much larger size, with shield length up to
17.7 mm; specimens of P. microps rarely reach more than 8 mm in shield length. The distribution of
P. abyssorum is the broadest so far known for species of the genus, and includes both sides of the Pacific (Figs
49-50) and Atlantic Oceans. In contrast, P. microps is so far known only from the central eastern Pacific (Fig. 50).
Remarks. — As discussed by LEMAITRE (1989: 34), Parapagurus abyssorum Henderson, 1888 was found to
be a junior homonym of Parapagurus abyssorum (Filhol, 1885a). The specimen figured twice by FlLHOL (1885a:
152, lig. 1; 1885b: 131, Fig. 41) from " Talisman ", stn 148, is considered the holotype of Pag ur us abyssorum
Filhol, 1885a, and is reproduced herein (Fig. 4). A replacement name, Parapagurus holthuisi Lemaitre, 1989,
was given for HENDERSON'S (1888) taxon, so far known from the central and eastern Pacific. However,
the "Challenger” material used by Henderson in the description of his taxon came from widely separate localities
in the Pacific and Atlantic Oceans. In addition to P. holthuisi. Henderson's material represents P. janetae
sp. nov. and possibly other as yet undetermined species of Parapagurus (see Remarks under P. holthuisi and
P. janetae sp. nov.).
Some of the specimens used by Smith (1884. 1886) in his report of Parapagurus pilosimanus, were found by
Lemaitre (1989) to be P. abyssorum (Filhol. 1885a).
As pointed out by Lemaitre (1989). A. Milne-Edwards & Bouvier (1892a, 1892b) viewed Smith's
(1879) Parapagurus pilosimanus as a highly variable, cosmopolitan species, and considered HENDERSON'S (1888)
P. abyssorum a synonym of Smith's taxon. However, A. Milne-Edwards & Bouvier retained HENDERSON’S
abyssorum as a variety for those specimens that occurred at great depths (3650-4060 m) which differed significantly
from the typical form. These reports by A. Milne-Edwards & Bouvier (1892a, 1892b) are referable, in part, to
P. abyssorum (Filhol. 1885a) and to the Atlantic P. pilosimanus Smith, 1879.
Lemaitre (1989) questioned the reports by Wood-Mason & Alcock (1891) and Alcock (1894) of
Parapagurus abyssorum from the Indian Ocean. However, in view of the broad distribution of this species, those
reports are here considered correct.
Porter (1906) reported Parapagurus pilosimanus Smith, 1879, from Los Vilos, Chile, in the eastern Pacific.
PORTER, however, expressed some doubt in his identification, which was later questioned by Haig (1955) because
the specimens did not come from deep-water. Furthermore, Smith's species is now known to occur only in the
Atlantic (Lemaitre. 1989). Unfortunately Porter's material has not been available for examination and he did
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
317
not provide sufficient information to ascertain the identity of his material. If Porter's material is indeed of a
species of Parapagurus , it most likely could represent any of three species known to occur in the area. i.e.
p. abyssorum (Filhol. 1885a), P. holthuisi Lemaitre, 1989, or P. janetae sp. nov.
In Haig's (1955) aforementioned publication on the Anomura from Chile, she included Parapagurus
pilosimanus from off Port Otway and Juan Fernandez Islands, based on HENDERSON'S (1888) "Challenger" material
reported as Parapagurus abyssorum Henderson, 1888 (= P. holthuisi). Haig (1955) considered HENDERSON'S
(1888) P. abyssorum a synonym of Smith's (1879) P. pilosimanus. Examination of Henderson's (1888)
material, however, has shown that the single specimen obtained by the "Challenger" from Port Otway actually is
P. janetae sp. nov., and the specimens from Juan Fernandez Islands represent P. holthuisi.
Based on the literature, GORDAN (1956) listed Parapagurus abyssorum. and P. abyssorum var. abyssorum , and
as such refer to P. holthuisi Lemaitre, 1989, and P. abyssorum (Filhol, 1885a) respectively.
DE Saint Laurent (1972) included the Indo-Pacific in the distribution of Parapagurus pilosimanus scaber [=
P. abyssorum (Filhol, 1885a)], but did not list any specimens or indicate exact localities. Miyake (1978, 1982),
based on DE Saint Laurent (1972), listed P. p. scaber from Japan, without examining any specimens. Although
P. abyssorum could possibly occur in Japan, no specimens have been found from this region in any of the
collections examined during this or previous studies.
The specimens here reported from the Tasman Sea ("Galathea", stn 575, ZMK) and southeastern Pacific
("Eltanin". stn 233, USNM 155046), appear in all respects to be conspecific with Parapagurus abyssorum (Filhol,
1885a), previously known only from the Atlantic (Lemaitre, 1989). The finding of specimens at such widely
separate localities gives this species a very broad distribution, even by the standards ot Parapagurus species. The
minor morphological variations seen in the Tasman Sea specimen can be attributed to its large size (shield length
17.7 mm, the largest known for this species). When compared with other available material, the morphology of
both the Tasman Sea and southeastern Pacific specimens fall well within the range of variations documented tor
this species by Lemaitre [1986 (as P. scaber Henderson, 1 888), 1989].
Parapagurus bouvieri Stebbing, 1910
Figs 7, 47, 49
Parapagurus bouvieri Stebbing. 1910: 357. pi. 17 (Crustacea pi. 43). — Balss. 1924: 768. — Gordan. 1956: 338. -
Fuller, 1958: 164. — Kensley, 1974: 65. — Lemaitre 1986: 526; 1989: 11; 1990: 223, fig. 2.
Parapagurus pilosimanus - Barnard, 1950: 450. fig. 83a-b. — KENSLEY, 1969: 153; 1974: 65; 1977: 161.
(Not Parapagurus pilosimanus Smith, 1879).
Parapagurus pilosimanus bouvieri - DE Saint Laurent, 1972: 103. pi. 1, fig. 4. — KENSLEY. 1981: 34. — Macpherson.
1983a: 12; 1983b: 472.
MATERIAL EXAMINED. — South Africa. S.S. "Pieter Fame": stn 153. Buffalo River, NW 1/2W. 19 miles. 549 m:
1 ov. 9 6.3 mm (SAM A1524).
Australia. Queensland. MV "Iron Summer": shot 1-7, off Moreton Island, 27°13'S. 153 00 E, 500-540 m
2-3.10.1982: 1 ov, 2 8.2 mm (QM W16517); shot 2, 27°19'9"S. I53°53'47"E, 600 m. 10.05.1983: 1 <S 15.2 mm (QM
WI4337V shot 3 27°12'83"S, 153°52'87"E, [no depth recorded], 10.05.1983: 1 6 12.1 mm (QM W14333).
New South Wales. FRV "Kapala": stn K77-23-10. 33°11'S. 152°24'E. 732 m. 7.12.1977: 5 d 8.5-12.1 mm (AM
P52735). — RV "Tangaroa": NZOl stn U219, 32°59'S, 152°33.5'E, 381-444 m. 9.10.1982. colls. W. Ponder &
R. Springthorpe: 1 9 3.9 mm (AM P40417).
Great Australian Bight. "Endeavour" EXPEDITION 1909-14: S of Eucla, 33°30’S. 129°28’E, 823 m: 1 6 13.1 mm (AM
P52736).
Southwestern Indian Ocean. South Africa. "Galathea": stn 197, off Durban, 29°57 S, 31°26E, 495 m. 14.02.
1951: 3 6 4.2-7.0 mm, 1 9 7.8 mm (ZMK).
Southeastern Atlantic. Angola. "Galathea": sin 101, 8°50’S, 12°32’E, 990 m. 12.12.1950: 1 6 6.7 mm (ZMK).
South Africa. "Pickle": stn 1483, off Cape Peninsula, 34°06’S, 17°53’E. 247 m, 13.12.1929: 2 6 12.0, 12.2 mm
(ZMK).
Types. _ Lectotype: ov. 9 6.3 mm (selected by Lemaitre, 1990: 223), South Africa, S.S. "Pieter Faure ",
stn 153, Buffalo River, NW 1/2W. 19 miles, 549 m (SAM A 1524).
318
R. LEMAITRE
FIG. 7. — Parapagurus bouvieri Stcbbing. 1910. a-h. southeastern Atlantic: a-c, 8 8.2 mm; d, 8 10.8 mm; e-f, ov. 9
10.0 mm; i-j. 8 12.1 mm, Australia, Queensland (QM W 14333). a. shield and cephalic appendages; b, left chela and
carpus (setae omitted); c. right chela and carpus (setae omitted); d. left first ambulatory leg, lateral view; e, merus of
left first ambulatory leg, lateral view; f, merus of left second ambulatory leg, lateral view; g, propodus and dactyl of
left fourth pereopod, lateral view; h. propodal rasp of same (setae omitted), ventrolateral view; i, left exopod of
uropods, dorsal view; j. telson, dorsal view.
1 990)SCalCS CqUal 3 mm <a"b)> 5 mm (C f)’ 1 mm (g)' 0 5 mm (H)* 1 mm (i)’ and °'5 mm (j)' (a'g* fr°m Lema,tre’
Source : MNHN. Paris
PARAPAGUR1DAE FROM THE PACIFIC AND INDIAN OCEANS
319
DIAGNOSIS. — Shield (Fig. 7a) about as broad as long, dorsal surface well calcified or weakly calcified medi¬
ally; lateral projections broadly rounded. Rostrum broadly subtriangular, rounded distally; with short mid-dorsal
ridge. Ocular peduncles more than half length of shield, weakly inflated basally; width of cornea about same or
slightly more than distal width of ocular peduncle. Ocular acicles subtriangular, terminating in strong simple spine
(rarely bifid). Antennular peduncle exceeding distal margin of cornea by half length of penultimate segment; lateral
face of basal segment with statocyst lobe having subrectangular distal lobe armed with 1 or 2 spines, and I spine
proximally. Antennal peduncle exceeding distal margin of cornea by at most half length of tilth segment;
flagellum with setae 1 to 2 flagellar articles in length; acicle weakly curved in dorsal view, exceeding distal margin
of cornea by at most half length of acicle, mesial margin armed with 5 to 10 small spines. Epistomial spine
usually present. Sternite of third maxillipcds with strong spine on each side of midline. Left chelipcd (Fig. 7b)
well calcified, densely setose; carpus with irregular rows of small spines on dorsal margin. Ambulatory legs
(Fig. 7d-f) with meri, carpi and propodi unarmed except for small dorsodistal spine on each carpus; meri each
about 3.5 (first leg) or 2.9 (second leg) times as long as high, with lateral and mesial faces weakly calcified
medially (weak calcification more pronounced on second leg; Fig. 7d-e). Anterior lobe ol sternite of second
ambulatory legs subsemicircular, setose, armed with small subdistal spine. Fourth pereopod (Fig. 7g-h) with
propodal rasp consisting of 2 or 3 rows of conical scales. Fifth pereopod with propodal rasp less than half length
of propodus. Telson (Fig. 7j) and uropods asymmetrical. Terminal margin of telson divided into 2 rounded
projections by shallow, rounded (U-shaped) cleft; rounded projections armed distally with alternating short and
long corneous spines (approximately 15 to 20 left, 10 to 17 right). Left exopod (Fig. 7i) of uropod elongate, about
3.0 times as long as broad; with broad rasp.
Size Range. — Males, SL 4.0 to 15.2 mm. Females 3.9 to 11.0 mm. Ovigerous females 6.3 to 12.2 mm.
Color (from Barnard, 1950: 451, as Parapagurus pilosimanus). — "Body pinkish, basal joints ot chelipeds
with reddish patches, 2nd and 3rd legs red, with a conspicuous white band along the upper and lower margins,
cornea dark crimson, antenna I pink with white band along upper margin of last peduncular joint, antenna 2 pink."
VARIATIONS. — The weakly calcified area on the lateral and mesial faces of the meri of the ambulatory legs
usually can be recognized by a dark, brownish coloration. The area is often slender, and occasionally is absent on
the first leg.
Habitat. — Usually found living in shelters tonned by zoanthid species, probably Epizoanlhus sp.
Distribution (Figs 47, 49). — Southeastern Atlantic and southwestern Indian Ocean: off Angola to South
Africa, and northward to off Natal. Western Pacific; Australia. Depth: 247 to 990 m.
REMARKS. — This species can be distinguished from all others in the genus by the weak calcification (usually
marked by a dark, brown area) present on the lateral and mesial faces of the meri of the ambulatory legs, and the
greater development of the ocular peduncles. In P. bouvieri, the length of the ocular peduncles (including corneae)
is distinctly more than half the length of the shield, whereas in all other species of Parapagurus the ocular
peduncles are half or less than half the length of the shield. The reduction of the ocular peduncles in P. bouvieri
has not been as strong as in other species of the genus, and is a condition that can perhaps be attributed to the
relatively shallower depth range at which this species lives compared to other species in the genus (Fig. 47). In
other crustaceans, a reduction of eyes with increasing depth has also been documented (e.g.. MENZ1ES et a /., 1973,
Marshall, 1979), and is an evolutionary trend that evidently has occurred independently in many groups.
Parapagurus andreui Macpherson, 1984
Figs 8-9, 47. 49
Parapagurus andreui Macpherson. 1984: 81, figs 24-27. - Lemaitre, 1986: 526; 1989: 11; 1990: 221. fig. 1. —
Lemaitre & McLaughlin, 1992: 763.
320
R. LEMAITRE
MATERIAL EXAMINED. — Southeastern Atlantic. Valdivia 1: stn P-5, Valdivia bank. 25°34.5'S, 6°04’E. 930-
933 m, 17.05.1983: 1 6 14.4 mm, I ov. 9 14.8 mm, paratypes (USNM 240164).
Western Indian Ocean. La Reunion. "Marion Dufresne cruise MD32, stn CP 103, 20°41.6'S, 54°56.8'E. 2950-
2970 m, 29.08.1982: 2 6 12.2, 14.2 mm (MNHN-Pg 5646).
South Africa. " Galathea ": stn 190, off Durban, 29°42’S, 33°19,E. 2720 m, 3.02.1951: 1 6 (damaged) 9.5 mm (ZMK).
Types. — All from Valdivia Bank. Holotype : ov. 2 14.0 mm, Valdivia I: stn P-4, 25°32'S, 6°06.9'E, 904-
959 m (ICM-D 204/1991). Paratypes : 1 6, 1 ov. 2. Valdivia 1: stn P-4, 25°32'S, 6°06.9'E, 904-959 m (ICM).
— 4 6, 4 ov. 2 (ICM): 1 6 14.4 mm. 1 ov. 2 14.8 mm (USNM 240164), Valdivia 1: stn P-5, 25°34.5,S,
6°04'E, 933 m, 17.05.1983, — 1 c? , Valdivia 1: stn P-9, 25°35'S, 6°09.3’E, 922 m (ICM). — 4 6,3 2,
3 ov. 2, Valdivia 1: stn P-10, 25°29.3’S, 6°07.5*E, 900-915 m (ICM).
Fig. 8. — Parapagurus andreui Macpherson, 1984, southeastern Atlantic: a. shield and cephalic appendages: b. left chela
and carpus (setae omitted): c, right chela and carpus (setae omitted).
Scale equals 4 mm. (From Lemaitre, 1990).
Diagnosis. — Shield (Fig. 8a) about as broad as long, dorsal surface well calcified or with weakly calcified
areas medially: lateral projections broadly rounded. Rostrum broadly subtriangular, rounded distally: with short
mid-dorsal ridge. Anterolateral margin of branchiostegite unarmed. Ocular peduncles less than half length of shield,
inflated basally; width of cornea about same as distal width of ocular peduncle. Ocular acicles subtriangular,
terminating in simple strong spine (rarely bifid). Antennular peduncle exceeding distal margin of cornea by nearly
entire length of penultimate segment: lateral face of basal segment with statocyst lobe having subrectangular distal
lobe armed with 1 or 2 spines, and 1 spine proximally. Antennal peduncle exceeding distal margin of cornea by
Source : MNHN. Pans
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
321
Fig. 9. — Parapagurus andreui Macpherson, 1984, western Indian Ocean, La Reunion, "Marion Dufresne", stn CP 103:
6 14.2 mm (MNHN-Pg 5646). a, left first ambulatory legs, lateral view; b. left second ambulatory leg, lateral view;
c, coxae and sternites of ambulatory legs, ventral view; d. propodus and dactyl of left fourth pereopod. lateral view;
e, propodal rasp of same (setae omitted), ventrolateral view; f, telson, dorsal view; g-h. left (g) and right (h) exopod
of uropods, dorsal view. Scales equal 5 mm (a-c), 1 mm (d), 0.5 mm (e), and 2 mm (f-h).
Source
322
R. LEMAITRE
entire length of fifth antennal segment; flagellum with setae 1 to 2 flagellar articles in length; acicle nearly straight
or weakly curved in dorsal view, exceeding distal margin of cornea by half or more length of acicle, mesial margin
unarmed. Epistomial spine usually absent. Sternite of third maxillipeds with strong spine on each side of midline.
Left cheliped (Fig. 8b) well calcified, densely setose; carpus with row of small spines on dorsal margin.
Ambulatory legs (Fig. 9a-b) with meri, carpi and propodi unarmed except for small dorsodistal spine on each
carpus; propodi each about 4 or more (first leg) or 4.5 (second leg) times as long as high; meri each about 4 (first
leg) or 3.5 (second leg) times as long as high. Anterior lobe of sternite of second ambulatory legs (Fig. 9c)
subsemicircular, setose, unarmed or armed with small subterminal spine. Fourth percopod (Fig. 9d-e) with
propodal rasp consisting of 2 or 3 rows of conical scales. Fifth pereopod with propodal rasp less than half length
of propodus. Telson and uropods (Fig. 9f-h) asymmetrical. Terminal margin of telson divided into 2 rounded
projections by angled (V-shaped) cleft; rounded projections armed distally with numerous short corneous spines
(approximately 25 or more left, 15 or more right). Left exopod (Fig. 9g) of uropod elongate, about 2.6 times as
long as broad; with broad rasp.
SIZE Range. — Males. SL 9.5 to 21.0 mm. Females 13.1 to 20.0 mm. Ovigerous females 13.0 to 14.4 mm
or more (MACPHERSON, 1984).
Habitat. — Usually found living in shelters formed by species of Epizoanthus.
DISTRIBUTION (Figs 47, 49). — Southeastern Atlantic: southern Angola to Namibia, including the Valdivia
Bank. Southwestern Indian Ocean: La Reunion to South Africa. Depth: 406 to 2970 m.
Affinities. — See P. latimanus.
Parapagurus microps dc Saint Laurent, 1972
Figs 10-11, 47, 50
Parapagurus microps de Saint Laurent, 1972: 104, figs 1, 13, pi. 1, fig. 8. — LEMAITRE, 1989: 1 1.
Material EXAMINED. — Eastern Pacific. Albatross ": stn 4742, 0°04'S, 1I7°07'W. 4243 m, 15.11.1905: I 6
8.7 mm, holotype (USNM 168305). — Stn 4717, SW of Galdpagos Islands, 5°irS, 98°56'W, 3938 m, 13.01.1905: 1 6
5.8 mm, 1 9 6.1 mm, paratypes (USNM 168306), 1 6 6.1 mm (USNM 276115). — Stn 4721, SW of Galapagos Islands,
8°07'30"S, 104°10'W, 3812 m. 15.01.1905: 1 9 7.0 mm, 1 ov. 9 5.4 mm (USNM 276116).
S of Galapagos . FS ''Sonne". DlSCOL 2 Expedition: stn So64-83, trawl 4, 7°08.83’S-88°30. 1 3'W to 7°10.65'S-
88°29.00’W. 4261 m, 16.09.1989: 1 <5 7.7 mm (SMF 23873); Stn So64-115, trawl 5, 7°01 .22,S-88°23.07'W to
7°08.68,S-88°20.37,W, 4196 m, 20.09.1989: 5 6 4.0-7.0 mm (SMF 23874). — Discol 3 Expedition: stn So77-41, trawl
7, 7°08.58'S-88°26. 1 0’W to 7o05.70'S-88o27.00’W, 4154 m, 5.02.1989: I 6 4.9 mm (SMF 23875).
Peru. "Akademik Kurchatov ": 4th cruise, stn 271, 17°42.0’S. 78°59.2'W, 3080-2710 m, 20.10.1968: 2 6 5.0.
6.7 mm (ZMUM Ma-4979).
Types. — Holotype : 6 8.7 mm, central eastern Pacific, "Albatross", stn 4742, 0°04'S, 1 17°07'W, 4243 m,
15.11.1905 (USNM 168305). Paratypes : 1 6 5.8 mm, 1 9 6.1 mm, SW of Galapagos Islands, " Albatross ",
stn 4717, 5°1 FS, 98°56'W, 3938 m, i3.01. 1905 (USNM 168306).
Diagnosis. — Shield (Fig. 10a) longer than broad, dorsal surface well calcified; lateral projections broadly
rounded. Rostrum broadly rounded; with low, sometimes inconspicuous mid-dorsal ridge. Ocular peduncles less
than half length of shield, inflated basally; width of cornea about same as distal width of ocular peduncle. Ocular
acicles subtriangular, terminating in strong simple spine. Antennular peduncle exceeding distal margin of cornea
by full length of penultimate segment; lateral face of basal segment with statocyst lobe having subrectangular
distal lobe armed usually with 1 small spine, and 1 spine proximally. Antennal peduncle (Fig. 10b) exceeding
distal margin of cornea by half length of fifth antennal segment; flagellum with scattered setae about I to
2 flagellar articles in length; acicle nearly straight in dorsal view, exceeding distal margin of cornea by 0.3 to 0.5
length of acicle, mesial margin unarmed or with 1-3 small spines proximally. Epistomial spine usually present.
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
323
Fig. 10. — Parapagurus microps dc Saint Laurent, 1972. a-g. holotypc 8 8.7 mm . W of Galdpagos Islands. Albatross .
stn 4742 (USNM 168305); h-i, 8 6.1 mm. SW of Galapagos Islands, " Albatross ", stn 4717 (USNM 276115).
a. shield and cephalic appendages; b. right antennal peduncle, lateral view; c. left cheliped (setae omitted); d, right
chela and carpus (setae omitted); e. telson, dorsal view; f-g, left (0 and right (g) exopod of uropods, dorsal view;
h. male left first pleopod; i. male left second pleopod.
Scales equal 1 mm (a-b,f-g), 3 mm (c-d), and 0.5 mm (e.h-i). (a,d, from de Saint Laurent. 1972).
Sternite of third maxillipeds with strong spine on each side of midline. Left cheliped (Fig. 10c) well calcified, with
sparse setation; palm and carpus with numerous small spines on dorsal surfaces. Ambulatory legs (Fig. 1 la-e)
with dactyl having ventromesial row of 13 to 16 minute spinules, and dorsal and dorsomcsial distal rows of setae,
meri, carpi, and propodi each armed on mesial, lateral, dorsal, and ventral faces with numerous small spines and
sharp tubercles (more dense in larger specimens SL > 8 mm, Fig. lla,c). Anterior lobe of sternite of second
ambulatory legs (Fig. Ilf) subsemicircular, setose, with short blunt or sharp subterminal spine. Fourth pereopod
(Fig. 1 lg-h) with propodal rasp consisting of 1 row of ovate scales. Fifth pereopod with propodal rasp less than
half length of propodus. Telson and uropods (Fig. lOc-g) asymmetrical. Terminal margin of telson divided into
2 rounded projections by shallow, rounded (U-shaped) cleft; rounded projections armed distally with short corneous
spines. Left exopod of uropod (Fig. 100 broad, about 2 times as broad as long, usually paddle-shaped; with narrow
rasp.
Source
324
R. LEMAITRE
Fig. II. — Parapagurus microps de Saint Laurent. 1972. a-d, g-i. holotype 8 8.7 mm, W of Galapagos Islands,
” Albatross ", stn 4742, (USNM 168305); e. paratype 9 6.1 mm, SW of Galdpagos Islands, "Albatross", stn 4717.
(USNM 168306). a. left first ambulatory leg, lateral view; b. dactyl of same, mesial view; c, left second ambulatory
leg, lateral view; d, dactyl of same, mesial view; e, left second ambulatory leg, lateral view; f. coxae and sternites of
ambulatory legs, ventral view; g, propodus and dactyl of left fourth pereopod. lateral view; h. same (setae omitted),
ventrolateral view.
Scales equal 5 mm (a-d), 2 mm (e), and 1 mm (f.g.h).
Size Range. — Males, SL 5.8 to 8.7 mm. Females 6. 1-7.0 mm. Ovigerous female 5.4 mm.
Habitat. — Gastropod or scaphopod shells.
Distribution (Figs 47, 50). — Central eastern Pacific. Depth: 2710 to 4261 m.
Affinities. — See P. abyssorum.
Parapagurus benedicti de Saint Laurent, 1972
Figs 12-15, 47, 50
Parapagurus sp. indet. - Rathbun. 1904: 162; 1910: 162.
Parapagurus armatus - Reinhart, 1944: 56, fig. 7A. — REISCHMAN, 1959: 409 (nomen nudum, see Remarks).
? Parapagurus pilosimanus - Makarov, 1938: 223, fig. 74; 1941: 139; 1962: 212, fig. 74. — Vinogradov, 1950: 240
(key), pi. 27, fig. 113. — Birshtein & Vinogradov. 1951: 60. — Kobjakova, 1958: 221 (list). — Birshtein &
Zarenkov, 1972: 442 (see Remarks).
Source : MNHN. Pans
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
325
Parapagurus pilosimanus benedicti de Saint Laurent. 1972: 103, pi. 1. Fig. 6. — McLaughlin. 1974: 371. figs 100-101.
— Hart, 1982: 108, fig. 38.
Parapagurus benedicti - LEMAITRE, 1989: 1 1.
Not Parapagurus arrnatus - Ross, 1967: 306 [= Sympagurus trispinosus (Balss, 191 1), see Remarks).
MATERIAL EXAMINED. Type Material. — Northeastern Pacific. Holotype: ov. 9 8.5 mm (left chcliped
abnormal, see Variations and abnormalities), " Albatross ", stn 5699, California, SW of Point Sur. 36°30N, 122°00W,
2195 m, 27.04.1911: (USNM 168304).
Probable paratypes (see Remarks). " Albatross ": stn 2839, California, Channel Islands, North ol San Clemente Island.
33°08'00MN, 1 1 8°40’00"W, 757 m, 8.05.1888: 1 8 8.7 mm (USNM 18992). — Stn 2860. Canada, British Columbia.
Queen Charlotte Island. Kunghit Island. S of Cape St. James, 51°23’00"N 130°34'00"W. 1602 m, 31.08.1888: 1 8
9.4 mm (USNM 18993). — Stn 2923. California. San Diego, 32°40'30,,N, 117°3r30"W. 1503 m, 19.01.1889: 1 8
8.2 mm, 3 ov. 9 6.0-7. 5 mm (USNM 18994). — Stn 2928, California, San Diego, 32°47'30,’N, I18°10'00"W. 763 m.
23.01.1889: 6 8 5. 2-8.1 mm, 1 ov. 9 6.1 mm (USNM 18995). — Stn 3074, Washington. Sea Lion Rocks, 47°22,00’,N,
125°48’30"W, 1604 m, 29.06.1889: 4 8 6.0-9. 8 mm, 5 ov. 9 6. 0-7. 2 mm (USNM 18996). — Stn 3340, Alaska, Alaska
Peninsula, S of Chirikof Island, 55°26,00,,N. 155°26'00"W, 1271 m, 29.08.1890: 16 6 3.7-6.4 mm. 5 9 3.9-4.9 mm.
8 ov. 9 4.2-5. 8 mm (USNM 18997). — Stn 4337, California, 15.6 mi SW of Pt. Loma Light at San Diego, 1128-
1244 m. 10.03.1904: 1 8 4.6 mm (USNM 168894). — Stn 4354, off California, 15.6 mi. SW of Pt. Loma Light at San
Diego, 1214-1189 m. 14.03.1904: 2 ov. 9 7.5, 10.0 mm (USNM 168893). — Stn 4382, California, 5.4 mi. SW of
S Point of North Coronado Island. 1174-1218 m. 18.03.1904: 1 d 9.7 mm (USNM 168895). — Stn 5693, off
California, W of San Nicolas Island, 33o13'30"N, 120°04'30MW. 825 m. 6.04.1911: 3 8 6.1-9.7 mm. 1 9 6.6 mm.
5 ov. 9 5. 5-7. 5 mm (USNM 168896).
Other Material examined. — Northeastern Pacific. Gulf of Alaska, "Vitjaz". 45th cruise: stn 6093. 57 51 N,
148°57'W, 1540-1340 m, 7.05.1969: 2 8 3.5, 3.5 mm. 1 ov. 9 5.0 mm (IORAN DecPa087-089). — Stn 6094. 57°44’N,
148°37'W, 2400 m, 7.05.1969: 1 8 3.3 mm (IORAN DecPa092).
Pereyra coll .Oregon, off Columbia River, : haul 1. 1555 m, 17.05.1962: 3 8 6. 3-7. 6 mm (USNM 216296), 1 9
5.7 mm, 2 ov. 9 6.3, 7.3 mm (USNM 216297), 2 8 7.2, 8.5 mm (USNM 216299); Haul 2, 1463 m, 7.05.1962: 1 ov. 9
5.5 mm (USNM 216303). — Stn 23A. 45°50'N. 124°52'30"W, 1097 m. 9.06.1962: 2 <5 6.6. 6.7 mm (USNM 216310).
— [no stn], 1920 m. 10.06. 1962: 2 8 7.0. 7.8 mm (USNM 216300), 1 ov. 9 6.0 mm (USNM 216301). — 45°37'06"N.
124°54'36"W, 1372 m, 15.05.1963: 3 <3 6.0-8. 2 mm, 1 9 6.6 mm (USNM 216308). — 45°54'N, 125°05'W. 1555 m,
15.05.1963: 2 8 6.6, 6.7 mm, 1 9 6.7 mm, 3 ov. 9 5. 1-6.1 mm (USNM 216311). — Haul 1. 1372 m. 28.08.1963: 3
8 7 0-8 5 mm, 3 ov. 9 5.4-6.9 mm (USNM 216298). — 1646 m. 2.09.1963: 1 8 6.9 mm (USNM 216302). — [no stn
or locality], 1555 m, 8.09.1963: 2 ov. 9 5.2, 5.5 mm (USNM 216304). — 45°43'N, 125°13'W. 1929 m. 28.05.1964: 1
8 9.9 mm (USNM 216309), 3 8 5.2-9. 1 mm, 2 9 7.6, 9.0 mm, 5 ov. 9 5. 2-9.0 mm (MNHN-Pg 5648); 45°55'N,
125°09'W. 1646 m, 29.05.1964: 4 8 5.2-8.1 mm (USNM 216305). 5 8 4.9-6.7 mm. 2 9 5.5. 6.0 mm (all parasitized)
(USNM 216306). 7 <3 5. 1-9.5 mm, 2 9 6.1, 7.1 mm, 7 ov. 9 5. 5-6.6 mm. 3 damaged (USNM 216307).
Northwestern USA to Mexico. " Albatross stn 3344. off Cape Elizabeth, Washington, 47°20’00"N, 125°07'30"W,
1520 m, 21.09.1890: 1 9 4.9 mm (USNM 18998). — Stn 5687. Mexico, Baja California. S of Cerros Island,
27°39'15"N, 1 15°16'W. 878 m. 23.04.1911: 1 9 5.3 mm (USNM 216312). — Stn 5690, Mexico. E of Guadalupe Island.
29°29'N, 1 16°18'W, 2014 m, 24.04.1911: 1 8 8.2 mm (USNM 276117).
[Vessel unknown]: 40 miles W of Punta Banda. Baja California, Mexico, stn P-137-60. 2067-2085 m. 13.02.1960,
coll. Parker & Yonge: 1 9 8.5 mm (LACM 60-291.1). — off Point Arguello. California, stn P-273-60. 34°46’N,
121°40.6'W, 2012 m, 9.11.1960: 1 9 6.4 mm. 1 ov. 9 4.6 mm (LACM 60-292.1).
TYPES. — Holotype-. ov. 2 8.5 mm (left cheliped abnormal) "Albatross", stn 5699, California. SW of Point
Sur, 36°30'N, 122°00'W, 2195 m, 27.04.191 1 (USNM 168304). Paratypes, see above Type material. All the
types are from the northeastern Pacific.
Diagnosis. — Shield (Fig. 12a) broader than long, dorsal surface usually well calcified; lateral projections
broadly rounded. Rostrum broadly subtriangular. rounded distally; with low mid-dorsal ridge. Ocular peduncles less
than half length of shield, inflated basally; width of cornea about same as distal width of ocular peduncle. Ocular
acicles (Fig. 13b-c) subtriangular, terminating most frequently in strong bifid spine (occasionally simple, rarely
trifid). Antennular peduncle exceeding distal margin of cornea by half or more length of penultimate segment;
lateral face of basal segment with statocyst lobe having subrectangular distal lobe armed with 1 -3 small spines, and
1 spine proximally. Antennal peduncle (Fig. 13a) exceeding distal margin of cornea by about 0.6 length of fifth
antennal segment; flagellum with scattered short setae about I flagellar article in length or less; second segment
with dorsolateral distal angle produced into strong, broad multifid spine usually with somewhat concave dorsal
Source :
326
R. LEMAITRE
face; first segment with 1-4 spines on lateral face; acicle weakly curved in dorsal view, exceeding distal margin of
cornea by 0.3 to 0.5 length of acicle, mesial margin armed with 4-12 small spines (2 or 3 spines in small
specimens SL 4.0 mm). Epistomial spine usually present, simple or occasionally bifid. Sternite of third
maxillipeds with strong spine on each side of midline. Left cheliped (Fig. 12c) well calcified, with dense setation
on carpus and chela; palm armed with irregular rows of spines on dorsolateral and dorsomesial faces; carpus with
numerous spines on dorsal surface. Ambulatory legs (Figs 12e, 14a-d) with dactyls each having ventroniesial row
of 8 to 12 minute spinules, and dorsal and dorsomesial distal rows of setae; lateral and mesial faces of segments
smooth, except for scattered short setae; first leg with ischium and merus (Fig. 14a,c) armed with irregular rows of
small spines on ventral margins; second leg with merus armed with irregular row of small spines (less distinct and
numerous than on first leg) on ventral margin (Fig. 14b.d), ischium unarmed or with a few small blunt spines on
Fig. 12. — Parapagurus benedicti de Saint Laurent, 1972. a-b.c, NE Pacific; c, ov. 9 10.0 mm, NE Pacific, " Albatross ",
stn 4354, (USNM 168893); d, holotype ov. 9 8. 5 mm, NE Pacific, " Albatross ", stn 5699 (USNM 168304). a, shield
and cephalic appendages; b. right cheliped (setae omitted); c, left cheliped (setae omitted); d, left cheliped of
holotype (setae omitted); e, left second ambulatory leg, lateral view.
Scales equal 3 mm (a-b,e), 5 mm (c), and 2 mm (d). (a,b,e, from McLaughlin, 1974).
Source : MNHN, Paris
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
327
Fig. 13. — Parapagurus benedicti tie Saint Laurent, 1972, NE Pacific, a-b, 9 7.3 mm, off Columbia River (USNM
216297); c, ov. 9 5.5 mm (USNM 216304). a, right antennal peduncle, lateral view; b-c, ocular acicles and rostrum,
dorsal view.
Scale equals 1 mm.
Fig. 14. — Parapagurus benedicti de Saint Laurent. 1972, ov. 9 10.0 mm, NE Pacific, off California, " Albatross ",
stn 4354, (USNM 168893): a-b, ischium and merus of right first (a) and second (b) ambulatory legs, lateral view;
c-d. same of first (c) and second (d) ambulatory legs, ventral view; e. coxae and sternites of ambulatory legs, ventral
view (ischia shown for first legs).
Scale equals 3 mm.
Source :
328
R. LEMAITRE
ventral margin distally. Anterior lobe of sternile of second ambulatory legs (Fig. I4e) with anterior lobe
subsemicircular, setose, with short subterminal spine (occasionally bifid). Fourth pereopod (Fig. 15a-c) with
propodal rasp consisting of 2 to 4 rows of ovate scales. Fifth pereopod with propodal rasp less than half length of
propodus. Telson and uropods (Fig. 15d-f) asymmetrical. Terminal margin of lelson divided into 2 rounded
projections by shallow, rounded (U-shaped) cleft; rounded projections armed distally with short corneous spines.
Left exopod (Fig. 15e) of uropod about 2. 1 times as long as broad, with broad rasp.
Fig. 15. — Parapagurus benedicti de Saint Laurent, 1972. a-b, 6 7.6 mm, NE Pacific, off Columbia River (USNM
216296); c-f, ov. 9 10.0 mm, NE Pacific, off California, "Albatross", stn 4354 (USNM 168893). a. propodus and
dactyl of left fourth pereopod, lateral view; b, same (setae omitted), ventrolateral view; c, propodus and dactyl of left
fourth pereopod, lateral view; d. telson, dorsal view; e-f, left (e) and right (0 exopod of uropods.
Scales equal 1 mm.
Size Range. — Males, SL 3.7 to 13.4 mm. Females 3.9 to 8.5 mm. Ovigerous females 4.2 to 10.0 mm.
Color (from Hart, 1982: 108). — "Carapace shield opaque, the sides deep red. posterior wine-red with an
opaque, white elongated triangle in the cardiac area and whitish areas on either side of this; pubescence white.
Abdomen orange and white-red. Right cheliped with cream pubescence; ischium orange; merus scarlet; carpus pale
pink and white; hand white with ventral part of fingers pink. Left cheliped similar but fingers orange dorsally and
ventrally. Walking legs orange and scarlet. Eyestalk orange and scarlet; cornea dark brown. Antennal flagellum
orange."
Habitat. — Gastropod shells with or without actinian entirely covering shell.
Distribution (Figs 47, 50). — Northeastern Pacific: from Alaska to Baja California; possibly northwestern
Pacific (see Remarks). Depth: 757 to 2400 m.
Source :
PARAPACURIDAE FROM THE PACIFIC AND INDIAN OCEANS
329
Variations and Abnormalities. — The ocular acicles in this species most frequently terminate in a bifid
spine (Fig. 12a) or less frequently a multifid spine (Fig. 13b). However, as much as 25% of the specimens in a
series can have acicles terminating in a simple spine (DE Saint LAURENT, 1972; McLaughlin, 1974).
The holotype of this species is abnormal in that its left chcliped (Fig. 1 2d) is similar to the right cheliped in
size, armature, and proportions of the segments. In other respects, the external morphology of the specimen
appears normal. A similar condition was reported in a species of the family Paguridae, Pagurus protuberocarpus
McLaughlin. 1982, and attributed to a regenerative duplication of the right cheliped on the left side
(McLaughlin, 1982; Lemaitre et al„ 1982).
Remarks. — Several authors have reported or cited Parapagurus pilosimanus from various localities east of
Kamchatka and the Kuril Islands, in the northwestern North Pacific (e.g. Makarov, 1938, 1941, 1962;
VINOGRADOV, 1950; BlRSHTEIN & Vinogradov, 1951; Kobjakova. 1958; BlRSHSTEIN & Zarenkov. 1972).
However, as previously mentioned, P. pilosimanus is considered to occur only in the Atlantic (LEMAITRE, 1989).
The specimens reported by these authors from the northwestern North Pacific have not been examined, and their
identity cannot be established with certainty based on the information included in their publications. No reports of
other Parapagurus species arc known from the norhwcslern North Pacific, north of Japan. The specimens reported
by these authors might represent P. benedicti, the only species so far known to occur in the extreme northeastern
portion of the North Pacific (see Fig. 50). It is possible that this species might have an amphi-Pacific distribution
similar to that known for some other species of hermit crabs in the region (see McLaughlin, 1974).
In her description of Parapagurus pilosimanus benedicti, DE SAINT LAURENT (1972) listed only the holotype
for her taxon. Evidently many other specimens were used which were labeled and catalogued as paratypes in
museums such as the National Museum of Natural History, Smithsonian Institution. Washington. It is unclear
whether DE SAINT LAURENT intended to designate as paratypes all. or part of this material.
As previously mentioned, DE Saint Laurent's (1972) subspecies Parapagurus pilosimanus benedicti was
elevated to specific rank by Lemaitre ( 1989). DE Saint Laurent (1972) indicated that the southern distribution
of this taxon included the Gulf of Panama, presumably based on her examination of material from the "Albatross".
McLaughlin (1974) and Hart (1982) followed de Saint Laurent and included the Gulf of Panama in the
distribution of this taxon. However, reexamination of the " Albatross " material used by DE SAINT LAURENT has
shown that no specimens of P. benedicti were obtained in the Gulf of Panama. The southernmost record so far
known for this species is off Baja California. Mexico.
McLaughlin (1974: 378) discussed the confusion with the name "Parapagurus armatus". This name was used,
but never published, by J. E. BENEDICT in labeling several lots in the collections of the National Museum of
Natural History, Smithsonian Institution. In studies of rhizocephalan parasites, REINHARD (1944) examined the
specimens labeled by J. E. BENEDICT, and used the name "Parapagurus armatus" in a legend lor his figure (fig. 7A)
of the host of his species Angulosaccus tenuis Reinhard, 1944. No diagnosis was given for the hermit crab host,
so the name became a nomen nudum according to the International Code of Zoological Nomenclature. Reischman
(1959) followed Reinhard (1944) and used the same name for the host in his report of the same rhizocephalan.
McLaughlin (1974) also concluded correctly that ROSS' (1967: 306) reference to Parapagurus armatus from
Pemba Strait, near Zanzibar, did not refer to the eastern North Pacific P. pilosimanus benedicti. but she was unable
to determine which species ROSS' report could represent. Ross’ use of the name "Parapagurus armatus" was based
on studies of actinians from the "Valdivia" Expedition by CARLGREN (1928a, 1928b) who used the name
Parapagurus armatus var. trispinosus for specimens of the hermit crab associated with Isadamsia cancrisocia
Carlgren, 1928a, collected at stn 246, Pemba Strait. It appears that CARLGREN intended to refer to Parapagurus
arcuatus var. trispinosa Balss, 191 1. for the hermit crab specimens associated with his specimens of /. cancrisocia.
and mistakenly used "armatus" rather than "arcuatus". Balss' hermit crab specimens from "Valdivia", stn 246 are
of Sympagurus trispinosus (Balss, 1911), a species distributed in the Indo-Pacific (Lemaitre, 1994. 1996).
Parapagurus benedicti can be separated from all other species of the genus known to occur in the eastern
Pacific, by the spines present on the ventral margins of the meri of the first and second ambulatory legs
(Fig. 14a-d); and the dorsolateral distal angle of the second antennal segment (Fig. 13a), which is distinctly more
strongly developed than in other species of the genus, and terminates in a multifid spine with a slightly concave
330
R LEMA1TRE
dorsal face. The bifid or less frequently multifid condition of the ocular acicles of P. benedicti also is a useful
diagnostic character. However, as mentioned under "Variations and Abnormalities", the ocular acicles are variable
and can terminate in a simple spine in a significant number of specimens.
Parapagurus holthuisi Lemaitre, 1989
Figs 16-18, 47, 49-50
Parapagurus abyssorum Henderson. 1888: 87 (in part), pi. 9. fig. 2. — Murray, 1895: 1129 (in part); 1896: 388 (in
part). — Gordan, 1956: 337 (lit.). [Not Parapagurus abyssorum (Filhol, 1885a); see Remarks].
IParapagurus pilosimanus - Haig. 1955: 17. — Porter. 1906: 129. [See Remarks under Parapagurus abyssorum (Filhol.
1885a)].
Parapagurus pilosimanus abyssorum - DE Saint Laurent, 1972: 103 (in part). ?not pi. 1. fig. 7 (see Remarks).
Parapagurus holthuisi Lemaitre, 1989: 34. [Replacement name for Parapagurus abyssorum Henderson, 1888. junior
homonym of Parapagurus abyssorum (Filhol, 1885a)],
Material EXAMINED. — Eastern Pacific. "Challenger", stn 300, W of Valparaiso, Chile. 33°42'S. 78°18'W
2515 m, 17.12.1875: <3 14.9 mm , hoiotype (NHM 1888:33). 5 S 11.8-13.3 mm. I 9 7.8 mm, 2 ov. 9 8.1. 11.4 mm .
paratypes (NHM 1888:33). — Stn 3374. Galapagos Islands. 2°35'N. 83°53-W, 3334 m, 3.03.1891: I ov. 9 6.3 mm
(USNM 42625). — Stn 4647, off Peru, 4°33'S. 87°42'30"W. 3667 m. 9.1 1. 1904: 10 8 5.9-10.0 mm. 6 9 5. 1-7.0 mm
(USNM 2761 14); 2 8 8.8, 9.4 mm. 2 ov. 9 5.7. 6.0 mm (MNHN-Pg 5649).
"Challenger". Chile. Juan Fernandez Island, [no stn number]. 21 15 m: 1 8 13.3 mm, 3 ov. 9 10.0-10.4 mm (USNM
15298). — Off Juan Ferntmdez Island, [no stn number], 2115 m: 3 8 8.5-12.8 mm (ZMK). — Off Valparaiso, [no stn
number or other data]: 1 9 1 1.2 mm, 1 ov, 9 10.7 mm (USNM 156411).
Central Pacific. Magellan Rise. POSSE Expedition, "Alvin", dive 1816, 7°00'N, 177°00'W. 3150 m 17 03 1987
coll. K. Smith; Id 12.2 mm (SIO C9941).
Types. — Hoiotype : 8 14.9 mm (Hoiotype of Parapagurus abyssorum Henderson. 1888), "Challenger",
stn 300. W of Valparaiso, Chile, 33°42'S, 78°18'W. 2515 m, 17.12.1875 (NHM 1888:33). Paratypes : 5 8 1 1.8-
13.3 mm, I 9 7.8 mm. 2 ov. 9 8.1. 1 1.4 mm. same data as hoiotype (NHM 1888:33).
Redescription. — Shield (Fig. 16a) about as long as broad; dorsal surface usually well calcified, with
scattered short setae; anterior margin weakly concave; lateral projections broadly rounded; anterolateral margin
sloping. Rostrum broadly subtriangular, rounded distally, overreaching lateral projections; often with
inconspicuous low mid-dorsal ridge. Anterodistal margin of branchiostegite (Fig. 16b) unarmed, setose.
Ocular peduncles (including corneae) distinctly less than half length of shield, each with rows of setae dorsally ;
peduncles inflated basally, slightly constricted medially. Ocular acicles subtriangular. terminating in strong simple
spine; separated basally by slightly less than basal width of one aciclc.
Antennular peduncles slender, long, exceeding distal margins of corneae by 0.8 or more length of penultimate
segments. Ultimate and penultimate segments with scattered setae. Ultimate segment nearly twice as long as
penultimate. Basal segment with ventromesial distal spine; mesial face unarmed; "lateral face with statocysl lobe
having subrectangular distal lobe armed with 1-3 small spines, and 1 spine proximally.
Antennal peduncles (Fig. 16b) exceeding distal margins of cornea by about 0.8 or more length of fifth
segments. Fifth segment with scattered setae on lateral and mesial margins. Fourth segment with scattered setae.
Third segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced,
terminating in strong multifid spine; mesial margin with spine on dorsodistal angle. First segment unarmed on
lateral lace; ventromesial angle produced, with row of small spines. Antennal acicles weakly curved in dorsal view,
setose; exceeding distal margin of cornea by about half or less length of aciclc; mesial margin armed usually with
4 to 8 (range 1 to 10) small spines. Flagellum distinctly overreaching extended right cheliped. with sparse short
setae less than 1 flagellar articles in length.
Mandible, maxilla, and first and second maxillipeds typical of species in genus (e.g. Fig. 20). Maxillule
(FigM6c-d) with external lobe of endopod weakly developed, internal lobe with long terminal seta and
4 subterminal setae. Third maxilliped with crista dentata consisting of about 19 corneous-tipped teeth. Sternite of
third maxilliped with spine on each side of midline. Epistomial spine usually present.
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
331
Fig. 16. — Parapagurus holthuisi Lemaitre, 1989. NE Pacific, off Peru, "Albatross", stn 4647 (USNM 2761 14): a-b.e-f,
h-j, <J 8.9 mm; c.d.k.l, <3 9.6 mm; g, ov. $ 7.0 mm. a, shield and cephalic appendages; b, right antennal peduncle
and anterolateral margin of branchiostcgite, lateral view; c, left maxillule, internal view; d. distal end of endopod of
same; e, left cheliped (setae omitted); f, right cheliped (setae omitted); g. right cheliped (setae omitted); h, telson,
dorsal view; i-j, left (i) and right exopod of uropods. dorsal view; k. male left first pleopod. mesial view; 1, male
second pleopod, anterior view.
Scales equal 2 mm (a-b), 1 mm (c-d.k-l), 3 mm (e-g). and 0.5 mm (h-j).
Chclipeds markedly dissimilar, each with dorsal surfaces of carpus and chela covered with moderately dense
setation; proportions of carpus and chela influenced by size and sexual dimorphism (see Variations). Right cheliped
(Fig. 16f-g) with fingers bent inwards at tips, each terminating in small corneous claw; with tufts of setae on
dorsal and ventral surfaces; cutting edges each with irregularly-sized calcareous teeth; culling edge of dactyl also
Source :
332
R. LEMAITRE
with distal row of small, closely-set, corneous teeth. Dactyl set at oblique angle to palm, with dorsomesial and
mesial rows of small spines proximally. Palm and carpus each with numerous small spines and tubercles on dorsal
and ventral surfaces (spines and tubercles usually less numerous on ventral surfaces). Merus with small tubercles
on dorsal, dorsolateral and ventral surfaces; mesial surface smooth, with ventromesial row of spines. Ischium with
dorsal and ventromesial row of spines. Coxa with I or 2 spines on ventrodistal margin and ventromesial row of
setae.
Left cheliped (Fig. 16e) slender. Fingers each terminating in small corneous claw; dorsal and ventral surfaces
with scattered tufts of short setae; cutting edge of dactyl with row of minute, closely-set, corneous teeth distally;
cutting edge of fixed finger with row of regularly-spaced, small, evenly-sized calcareous teeth. Palm with
dorsomesial row of small spines; dorsolateral face with small spines. Carpus with irregular rows of small spines
dorsal ly; lateral face with scattered small spines or tubercles. Merus with row of short, stiff setae dorsally;
ventromesial margin with row of spines. Ischium armed with blunt spine or tubercles dorsally, and ventromesial
row of spines. Coxa usually with 2 small spines on ventrodistal margin, and ventromesial row of setae.
Ambulatory legs (Fig. 17a-d) similar Irom right to left (except slightly longer segments on right), slender,
long, distinctly overreaching right cheliped. Dactyl 1.3 to 1.5 times as long as propodus; with dorsal and
ventromesial distal row of setae; ventromesial margin with row of about 8 or more minute corneous spinules.
Merus, carpus, and propodus each with short stiff setae on dorsal margin (setae on propodus usually arranged in
short transverse rows). Propodus 6 or more times as long as high. Carpus with dorsodistal spine. Merus of first
ambulatory leg about 4.5 times as long as high; with longitudinal row of short setae on lateral face. Ischium with
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
333
1 to 3 small spines on ventral margin (first leg), or unarmed (second leg). Coxae of ambulatory legs (Fig. I7e)
unarmed or with 1-3 small spines ventromesially. Anterior lobe of sternite of second ambulatory legs (Fig. 17e)
subsemicircular, setose, armed with simple subterminal spine.
Fourth pereopod (Fig. 18a-d) semichelate. Dactyl subtriangular; shorter than length of propodal rasp,
terminating in corneous claw; with ventrolateral row of small, closely-set, corneous spines. Propodal rasp with
1 row of ovate scales at least distally (Fig. 18b) in small to medium size specimens (SL < 9.0 mm), or with 2 or
3 rows of ovate scales (Fig. 18d) in large specimens (SL > 10.0 mm). Carpus with row of long setae on dorsal
margin. Merus with setae on dorsal and ventral margins.
Fifth pereopod (Fig. 18e) chelate; propodal rasp forming subtriangular area less than half length of propodus.
Telson and uropods (Fig. 16h-j) asymmetrical. Left exopod (Fig. 16i) about 2.4 as long as broad; rasp
moderately broad. Telson without lateral indentation, and scattered setae dorsally; terminal margin divided into
2 rounded projections by shallow, often broad, rounded (U-shaped) cleft; margins of rounded projections each armed
with 6 to 15 short to moderately long corneous spines, each margin occasionally with additional 7 or 8 much
shorter subdistal spines dorsally.
Fig. 18. — Parapagurus holthuisi Lemaitre, 1989. a,b,e, 6 8.9 mm. E Pacific, off Peru, "Albatross", stn 4647 (USNM
276114); c-d, ov. $ 10.9 mm. "Challenger", Juan Ferndndez Island (USNM 15298). a. propodus and dactyl of left
fourth pereopod, lateral view; b, same (setae omitted), ventrolateral view; c, propodus and dactyl of left fourth
pereopod, lateral view; d, same, ventrolateral view; e. propodus and dactyl of left fifth, lateral view.
Scales equal 1 mm.
Size Range. — Males, SL 5.9 to 14.9 mm. Females 5.1 to 1 1.2 mm. Ovigerous females 8.1 to 1 1.4 mm.
Variations. — Males with palm and carpus of right cheliped (Fig. 16f) longer than broad: palm as much as
1.3 times as long as broad, carpus as much as 2.2 times as long as broad. Females with palm (Fig. 16g) about as
long as broad, carpus about 1 .5 times as long as broad.
Habitat. — Has been found living in anthozoans (actinians).
Distribution (Figs 47, 49, 50). — Central Pacific: Magellan Rise. Eastern Pacific: Galapagos Islands; off
Peru; and off Chile, including Juan Fernandez Island. Depth: 21 15 to 3667 m.
Source
334
R. LEMAITRE
AFFINITIES. — This species is similar to P. saintlaurentae sp. nov.; however, the Iwo differ in several
characters. The scales of the propodal rasp of ihe fourth pereopod are ovate in P. hothuisi , whereas the scales are
lanceolate or conical in P. saintlaurentae sp. nov. The terminal margin of the tclson is divided into two rounded
projections by a shallow cleft in P. holthuisi (Fig. 16h); the telson is divided by a deep cleft in P. saintlaurentae
sp. nov. (Fig. 31f,i). The antennal scales are usually armed with four or more strong spines, and exceed the distal
margin of the cornea by half or less than half the length of the acicles in P. holthuisi (Fig. 16a); the acicles are
usually unarmed or with one or two weak spines, and exceed the distal margin of the corneae by more than half the
length of the acicles in P. saintlaurentae sp. nov. (Fig. 28a-b).
REMARKS. — As previously mentioned, Parapagurus holthuisi Lemaitre, 1989, is a replacement name
proposed by LEMAITRE (1989) for P. abyssorum Henderson, 1888, a junior homonym of P. abyssorum (Filhol.
1885a). The " Challenger " material used by HENDERSON (1888) to describe his P. abyssorum included specimens
from the Atlantic (Bermuda; Sierra Leone; Tristan da Cunha). western Pacific (Banda, Indonesia; Philippines; north
of Papua New Guinea; Yokohama, Japan), and eastern Pacific (west of Valparaiso, and Port Otway, Chile).
HENDERSON indicated that his description was based on a male specimen from west of Valparaiso (stn 300); thus,
that male must be considered the holotype of his taxon and therefore of P. holthuisi. Examination of
HENDERSON'S single specimen obtained from Port Otway (stn 304) and deposited in the Australian Museum,
Sydney (AM G.1653), has revealed that it actually represents the new species P. janetae , described herein. It has
not been possible to examine Henderson's material from the western Pacific, and it could represent one or more
of the species of Parapagurus that occur in that region.
De Saint Laurent (1972, pi. 1, fig. 7) included a photograph of a right cheliped of a specimen of
Parapagurus pilosimanus abyssorum. She did not, however, provide locality information for the specimen, and
from the appendage alone it is not possible to determine whether it actually is of P. holthuisi.
Parapagurus richeri sp. nov.
Figs 19-23, 47-48
Parapagurus pilosimanus nudus - DE Saint LAURENT, 1972: 102. ?pl. 1, fig. 2 (in part, see Remarks).
Material EXAMINED. — South China Sea. Off Tungsha Tao (Pratas Island), RV "Fisheries Research /", trawled,
[depth unknown], 3.04.1995. coll. Ding-An Lee: 1 8 7.1 mm (NTOU P-1995-4-3).
Philippines. "Albatross": stn 5450. E coast of Luzon, 13°23'15"N, 124°00'30"E, 746 m, 4.06.1909: 4 8 4.8-
5.4 mm, 2 $ 4.8, 5.4 mm (USNM 276138).
Indonesia. "Siboga": stn 211. 5°40.7'S. 120°45.5'E, 1158 m. 25.09.1899: 1 8 3.9 mm (ZMA). — Stn 241.
4°24.3'S, 129°49.3'E. 1570 m, 1.12.1899: 1 8 3.6 mm (ZMA). — [stn unknown]: 1 ov. 9 7.3 mm (ZMA).
"Albatross": stn 5582, Borneo, 4°19'54,,N. 118°58’38”E, 1628 m. 26.09.1909: 1 9 5.5 mm (USNM 276144). —
Stn 5605, Celebes, Gulf of Tomini, 0°21'33"N, 121o34'10"E, 1183 m, 16.11.1908: 2 (dismembered) 3.9, 6.0 mm
(USNM 276132). — Stn 5606, 0o16'28"N, 12l°33'30”E, 1525 m. 17.11.1909: 1 8 4.6 mm, 1 ov. 9 4.5 mm (USNM
276133). — Stn 5613, Celebes, Gulf of Tomini, 0°42'00”S. I2lo44'00"E, 1375 m, 20.11.1909: 1 9 4.2 mm (USNM
276134). — Stn 5648. Buton Strait, 5o35'00"S, 122°20'00ME, 1022 m, 16.11.1909: 2 9 4.2. 5.8 mm (USNM 276135).
— Stn 5651, Celebes, Gulf of Bone, 4°43,50"N, 121°23,24,,E, 1280 m, 17.12.1909: 1 9 5.5 mm (USNM 276136). —
Stn 5660, Flores Sea, 5°36,30”S, 120°49’00”E, 1266 m, 20.12.1909: 2 8 3.7, 5.7 mm (USNM 276137).
New Caledonia. Biocal: stn CP 05, 21°16’S, 166°44’E, 2340 m. 11.08.1985: 4 8 3.7-5. 1 mm (USNM 276139).
— Stn CP 17, 20°35'S, I67°25'E, 3680 m, 14.08.1985: 4 8 2.2-5.4 mm, 1 9 2.8 mm, 1 ov. 9 3.9 mm (MNHN-Pii
5615). — Stn CP 23, 22°46'S, 166°20’E. 2040 m, 28.08.1985: 8 8 2.1-4.3 mm. 1 9 2.8 mm, 1 ov. 9 3.1 mm (MNHN-
Pg 5616). — Stn CP 26, 22°39.45’S, 166°27.41’E, 1618-1640 m. 28.08.1985: 5 8 3.4-6. 1 mm, 1 9 3.4 mm. 3 ov. 9
3. 1-4.2 mm (MNHN-Pg 5617). — Stn CP 27, 22°05.52’S, 166°25.90'E. 1850-1900, 28.08.1985: 3 8 2.2-3.7 mm, 1 9
5.4 mm. 1 ov. 9 4.1 mm (MNHN-Pg 5618). — Stn CP 57, 23°43.26'S. 166°58.06’E, 1490-1620 m, 1.09.1985: 4 8 3.3-
3.7 mm, 2 9 3.0, 3.4 mm, 1 ov. 9 3.7 mm (MNHN-Pg 5619). — Stn CP 60, 24°01.45'S, 167°08.43'E, 1480-1530 m,
2.09.1985: 2 8 1.5. 5.1 mm (MNHN-Pg 5620). — Stn CP 62, 24°19.06’S. 167°48.65'E, 1395-1410 m. 2.09.1985: 1 9
3.4 mm (MNHN-Pg 5621). — Stn CP 63, 24°28.69'S, 168°07.72’E, 2160 m. 2.09.1985: 1 8 3.2 mm, 1 9 3.3 mm
(MNHN-Pg 5622). — Stn CP 72, 22°09.02'S, 167°33.18'E, 2100-2110 m, 4.09.1985: 1 9 4.3 mm (MNHN-Pg 5623).
Biogeocal: stn CP 214, 22°43.09'S, 166°27.19'E, 1665-1590 m. 9.04.1987: 9 8 3.1-5.4 mm, 2 9 2.5, 3.3 mm.
1 ov. 9 3.3 mm (USNM 276140). — Stn CP 216, 22°50.67'S, 166°22.75'E, 2175-2250 m, 10.04.1987: 1 ov. 9
Source : MNHN. Paris
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
335
3.8 mm (NMNH-Pg 5624). — Stn CP 250. 2I°24.63'S, 166°28.21E, 2350 m, 15.04.1987: 1 <3 4.6 mm (MNHN-Pg
5625). — Sin CP 260, 21°00.00'S, I67°58.34'E, 1820-1980 m, 17.04.1987: 5 <3 2.8-4.3 mm, 4 9 3.0-4.0. 7 ov. 9
3 0-4.0 mm (MNHN-Pg 5626). — Stn CP 265. 21°04.09'S. 167°00.40’E, 1760-1870 m. 18.04.1987: 1 8 4.0 mm.
1 ov. $ 3.9 mm (MNHN-Pg 5627). — Sin CP 266, 2l°04.85'S, 167°57.14'E, 2100-1990 m. 18.04.1987: 3 <3 2.7-
3.3 mm, 2 ov. 9 3.6. 4.3 mm (MNHN-Pg 5628). — Sin 272. 21°00.04'S. 166°56.94'E. 1615-1710 m. 20.04.1987:
10 (3 3.0-6. 1 mm, 2 9 3.3, 3.7 mm. 5 ov. 9 3.4-4.0 mm. 1 immat. 1.7 mm (MNHN-Pg 5629). — Stn 272, 21°00.04'S,
1 66°56.94'E, 1615-1710 m, 20.04.1987: 1 <3 4.9 mm (MNHN-Pg 5614). — Stn 273, 21°01.53’S, I66°57.41'E. 1920-
2040 m, 20.04.1987: 6 <3 2. 7-4.6 mm, 2 9 4.5, 4.6 mm, 4 ov. 9 3. 1-4.8 mm (MNHN-Pg 5630); 2 <3 3.7. 5.2 mm
(USNM 276141). — Stn 317, 20°48.12’S, 166°53. 16E, 1630-1620 m. 2.05.1987: 2 8 2.4, 3.7 mm, 3 9 2.5-4.0 mm.
3 ov. 9 3. 3-5.0 mm (MNHN-Pe 5631). — Stn CP 321, 2ri2.00’S, 166°59.85'E, 2190-2205 m, 3.05.1987: 3 <3 2.7-
4.4 mm, 2 9 3.3, 4.1 mm, 2 ov. 9 4.2.4.5 mm (MNHN-Pg 5632). — Stn CP 329, 21°09.05'S. I66°40.08'E. 2315-
2310 m, 4.05.1987: 1 <3 3.2 mm (MNHN-Pg 5633). — Stn 341. 21C,29.73'S. 166°47.37'E. 2334 m. 6.05.1987: I <3
3.7 mm, 1 9 4.0 mm. 2 ov. 9 3.3. 4.5 mm (MNHN-Pg 5634).
BATHUS 3: stn CP 844. 23°06'S. 166°45'E. 908 m, 1.12.1993: 1 8 4.5 mm (MNHN-Pg 5635).
Halipro 1: stn CC 856, 21°44'S, 166°37'E. 311-365 m, 20.03.1994: 1 ov. 9 4.1 mm (MNHN-Pg 5636).
Vanuatu. Musorstom 8: stn CP 956, 20°33'S. 169°35'E, 1175-1210 m. 20.09.1994: 1 <3 3.9 mm (MNHN-Pg
5639). — Sin 1109. 14°52'S, 167°I8'E, 1550-1620 m. 8.10.1994: 1 9 3.7 mm. I ov. 9 5.1 mm (MNHN-Pg 5640). —
Stn CP 1110. 14°49'S, 167°15'E, 1360 m. 8.10.1994: 1 ov. 9 4.0 mm (MNHN-Pg 5641). — Stn CP 1111. 14°51 S.
167°14'E, 1210-1250 m, 8.10.1994: 1 ov. 9 5.1 mm (MNHN-Pg 5642). — Stn CP 1125. I5°57'S. I66°38'E, 1160-
1220 m. 10.10.1994: 3 c3 5.6-5.9 mm, 2 ov. 9 5.7-6.2 mm (MNHN-Pg 5643). — Stn 1126. 15°58'S. 166°39'E. 1210-
1260 m. 10.10.1994: I <3 4.7 mm (MNHN-Pg 5644).
Wallis and Futuna. Musorstom 7: stn DW 620. 12°34'S. I78°11'W, 1280 m, 28.05.1992: 1 <3 6.6 mm (MNHN-
Pg 5637). — Stn CP 621, 12°35'S, 178°11'W, 1280-1300 m, 28.05.1992: 1 8 4.5 mm, 1 ov. 9 4,6 mm (USNM
276142). — Stn CP 623, 12°34 S. 178°15'W. 1280-1300 m, 28.05.1992: 4 8 2.1-2.6 mm (MNHN-Pg 5638).
Western Pacific (other). Kermadec Trench (NE of New Zealand). "Galalhea": stn 668, 36°23'S. I77°41'E.
2640 m, 29.02.1952: 2 <3 3.1, 4.3 mm. 1 ov. 9 4.3 mm (ZMK CRU-3389).
Magellan Rise. Posse Expedition: "Alvin", dive stn 1816. 7°00'N. 177°00'W. 3150 m. 17.03.1987: 1 ov. 9
11.0 mm (SIO C9942). — R/V "Atlantis II". stn KLS 137, 7°I2.8'N, 176°07'W. 3150 m, 40' otter trawl. 18.03.1987:
3 <3 6.6-11.8 mm (SIO C9339).
Australian region. Queensland. ORV "Franklin". Cidaris 1: stn 30-4. 1 7° 1 9. l'S. 147°11.2'E, 1403 m.
12.05.1986: 1 8 8.6 mm (QM W16498), 1 ov. 9 6.1 mm (QM W16507). — Stn 32-2. 17°05.9'S. 147°11.9'E, 1517 m,
13.05.1986: 1 <3 7.5 mm (QM W16509). — Stn 33-1, 16°58.7'S. 147°11.4'E. 1564 m, 13.05.1986: 1 <3 6.5 mm (QM
W16510). — Stn 35-3, 16°50.8'S, 147°10.8'E. 1609 m. 14.05.1986: 1 8 4.2 mm (QM W16511). — Stn 37-1,
I7°01.7'S, 146°58.9'E, 1405-1500 m, 14.05.1986: 2 ov. 9 5.8, 6.2 mm (QM WI6495). — Stn 41-2, 17°33.3'S.
146°60'E, 1026-1056 m. 15.05.1986: 1 <3 5.8 mm (QM W16504).
New South Wales. R V "Tangaroa". NZOI cruise U214, off Newcastle. 2984-3058 m. 8.10.1982: 3 ov. 9 4.2-5. 1 mm
(AM P40408).
ORV "Franklin": stn SLOPE 58, 56 km ENE of Nowra. 34°43.95'S. 151°14.74'E, 817 m. 22.10.1988: 1 c5 8.2 mm
(NMV J401 13), 1 9 5.5 mm (NMV J16199).
Tasman Sea. Lord Howe Rise. ORV "Franklin": sin FRO 589-32, 27°I1.97'S. 160°37.80'E. I960 m, 7.05.1989:
1 ov. 9 3.5 mm (AM P52738). — Stn FRO 589-33, 27°13.34'S. I60043.41'E. 1989 m. 7.05.1989: 1 c3 4.5 mm (AM
P39445). — Sin FRO 589-35, E of Gifford Guyot. 26°51.57'S. 159°48.72'E. 2500 m. 8.05.1989: 1 c3 5.7 mm. 1 ov. 9
6.1 mm (AM P52739).
Western Indian Ocean. Cape Town to Durban. "Galathea": stn 178, 35°07'S, 30°35'E, 4470 m, 23.01.1951: 2 8
4.8. 7.2 mm, 2 ov. 9 4.9, 5.2 mm (ZMK CRU-3386). — Stn 190, 29°42'S, 33°19'E. 2790 m, 3.02.1951: I c3 3.2 mm
(ZMK CRU-3387). — Stn 198, 30°22'S. 34°27’E. 2765 m, 15.02.1951: 1 c3 3.5 mm (ZMK CRU-3388).
Types. — Holotype: 3 4.9 mm. New Caledonia. Biogeocal, stn 272, 21°00.04'S. 166°56.94'E. 1615-
1710 m. 20.04.1987 (MNHN-Pg 5614). Paratypes : All the others specimens mentioned above.
Description. — Shield (Fig. 19a) about as long as broad; dorsal surface usually well calcified, with rows
of short setae posteriorly on each side of midline; anterior margin weakly concave; lateral projections broadly
rounded; anterolateral margin sloping. Rostrum broadly subtriangular, rounded distally, slightly overreaching
lateral projections; with low mid-dorsal ridge. Anterodistal margin of branchiostegite (Fig. 19d) rounded, unarmed,
setose.
Ocular peduncles (including corneae) about half length of shield, each with dorsal longitudinal row ol setae;
peduncles inflated basally. Ocular acicles subtriangular, terminating in strong spine (rarely bifid); separated basally
by slightly less than basal width of one acicle.
Source :
336
R. LEMAITRE
Fig. 19. — Parapagurus richeri sp. nov., New Caledonia. a,d, Biogeocal stn 272, holotype 8 4.9 mm (MNHN-Pg 5614);
b, Biogeocal stn CP 266. paratype 8 4.6 mm (MNHN-Pg 5628); c, Biocal stn CP 17, paratype 8 3.6 mm (MNHN-Pg
5615); e-f, Biogeocal stn 272, 8 5.4 mm (MNHN-Pg 5629). a, shield and cephalic appendages; b-c, anterior
portion of shield and cephalic appendages (setae omitted); d. right antennal peduncle and anterolateral margin of
branchiostegite, lateral view; e, male first pleopod, mesial view; f. male second pleopod. anterior view’.
Scales equal 1 mm (a-c), and 0.5 mm (d-f).
Antennular peduncles slender, long, exceeding distal margins of corneae by at least 0.25 length of penultimate
segments. Ultimate and penultimate segments with scattered setae. Ultimate segment nearly twice as long as
penultimate. Basal segment with ventromesial distal spine; mesial face unarmed; lateral face with statocyst lobe
having subrectangular distal lobe armed with 2 small spines, and 1 spine proximally.
Antennal peduncles (Fig. 19d) exceeding distal margins of corneae by about half length of fifth segments. Fifth
segment with setae on lateral and mesial margins. Fourth segment with scattered setae. Third segment with strong
ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong multifid
spine; mesial margin with spine on dorsodistal angle. First segment with small spine on lateral face; ventromesial
angle produced, with row of small spines. Antennal acicles (Fig. 19a-c) nearly straight in dorsal view, setose;
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
337
exceeding distal margin of cornea by 0.2 or more length of acicle; mesial margin usually armed on proximal half
with 1-3 spines, occasionally with up to 8 spines (Fig. 19b). Flagellum distinctly overreaching extended right
cheliped; articles with setae less than 1 to 2 flagellar articles in length.
a
Fig. 20. — Parapagurus richeri sp. nov., New Caledonia, BlOGEOCAL stn 272, paratype 6 5.4 mm (MNHN-Pg 5629). Left
mouthparts, internal view: a. mandible; b. maxillule; c, distal end of endopod ot same; d, maxilla; e, first
maxilliped; f, second maxilliped; g, third maxilliped. Scales equal 1 mm (a,b,d,e-g), and 0.5 mm (c).
Mandible (Fig. 20a) as figured. Maxillule (Fig. 20b-c) with external lobe of endopod weakly developed, internal
lobe with long seta. Maxilla (Fig. 20d) with endopod exceeding distal margin of scaphognathite. First maxilliped
(Fig. 20e) with endopod exceeding exopod in distal extension. Second maxilliped (Fig. 20f) without distinguishing
characters. Third maxilliped (Fig. 20g) with crista dentata consisting of about 14 small corneous teeth; basis with
1 tooth mesially. Sternite of third maxilliped with spine on each side of midline. Epistomial spine usually present.
Chelipeds markedly dissimilar, each with dorsal surfaces of carpus and chela covered with moderately dense
setation. Right cheliped (Fig. 21c-e) with proportions of carpus and chela influenced by size and sexual
dimorphism (see Variations). Fingers bent inwards at tips, each terminating in small corneous claw; with few tufts
of setae on dorsal and ventral surfaces; cutting edges each with irregularly-sized calcareous teeth; cutting edge of
dactyl also with distal row of small, closely-set, corneous teeth. Dactyl set at slightly oblique angle to palm, with
338
R. LEMAITRE
dorsomesial and mesial row of small spines proximally. Palm and carpus each with numerous small, closely-set,
spines and tubercles on dorsal surface; ventral surfaces of palm and carpus also with spines and tubercles but less
numerous or sometimes few and scattered. Merus with small tubercles on dorsal, dorsolateral and ventral surfaces;
mesial surface smooth, with ventromesial row of spines. Ischium with dorsal and ventromesial row of spines.
Coxa with 1 or 2 spines on ventrodistal margin and ventromesial row of setae.
Fig. 21. — Parapagurus richeri sp. nov. a,c. New Caledonia, Biogeocal stn 272. holotype 6 4.9 mm (MNHN-Pg 5614);
b, ORV "Franklin", stn 30-4, 6 8.6 mm (QM W 16498); d, New Caledonia. BIOCAL stn CP 17, 6 5.2 mm (MNHN-Pg
5615); e. New Caledonia, Biocal stn CP 26, ov. 9 3.1 mm (MNHN-Pg 5617). a-b, left chelipeds (setae omitted in b);
c-e, right chelipeds (setae omitted in d,e).
Scales equal 2 mm.
Left cheliped (Fig. 2 1 a-b) slender. Fingers each terminating in small corneous claw; dorsal and ventral surfaces
with scattered tufts of short setae; cutting edge of dactyl with row of minute, closely-set, corneous teeth distally;
cutting edge of fixed finger with row of regularly-spaced, small, evenly-sized calcareous teeth. Palm unarmed
except for few small dorsomesial tubercles. Carpus armed with an irregular row or several rows of small spines or
tubercles dorsally. Merus unarmed except for row of bristles on dorsal margin. Ischium with small, blunt setose
tubercles on dorsal margin; usually with small spine on ventromesial margin proximally. Coxa usually with 2
small spines on ventrodistal margin, and ventromesial row of setae.
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
339
Fig. 22. — Parapagurus richeri sp. nov., New Caledonia: a-d,e. BlOGEOCAL stn 272, holotype 6 4.9 mm (MNHN-Pg
5614); e,f, BlOCAL stn CP 17, paratype 6 3.6 mm (MNHN-Pg 5615). a. first ambulatory leg, lateral view; b. dactyl of
same, mesial view; c, second ambulatory leg, lateral view; d, dactyl of same, mesial view; e, lirst ambulatory leg.
lateral view; f. second ambulatory leg, lateral view; g, sternite of second ambulatory legs, ventral view.
Scales equal 1 mm (a-f), and 0.5 mm (g).
Source : MNHN. Paris
340
R. LEMAITRE
Ambulatory legs (Fig. 22a-f ) similar from right to left (except slightly longer segments on right), slender,
long, distinctly overreaching right chelipcd. Dactyl about 1.7 to 2.1 times as long as propodus; with dorsal and
ventromesial distal row of setae; ventromesial margin with row of about 8 to 10 minute corneous spinules. Merus.
carpus, and propodus each with short setae on dorsal margin; carpus with small dorsodistal spine. Ischium usually
with small vcntrodistal tubercle (first leg) or unarmed (second leg). Coxa with ventrodistal margin usually armed
with 2 small spines (first leg) or unarmed (second leg). Anterior lobe of sternite of second legs (Fig. 22g) setose,
armed with simple subterminal spine.
Fourth pereopod (Fig. 23a-c) semichelate. Dactyl subtriangular, shorter than length of propodal rasp,
terminating in corneous claw; with ventrolateral row of small, closcly-set, corneous spines. Propodal rasp with
I row of ovate scales at least distally; rasp frequently with 2 or 3 short irregular rows of ovate scales proximally
(specimens SL > 5.0 mm; Fig. 23c). Carpus with row of long setae on dorsal margin. Merus with setae on dorsal
and ventral margins.
Fifth pereopod (Fig. 23d) chelate; propodal rasp forming subtriangular area less than half length of propodus.
Fig. 23. — Parapagurus richeri sp. nov. a.d.e.f.g. New Caledonia. Biogeocal stn 272. holotype <5 4.9 mm (MNHN-Pg
5614); b-c, Queensland. Australia. ORV "Franklin", stn 37-1, paratype ov. 9 6.2 mm (QM W16495); h-i. New
Caledonia, Biogeocal stn 272, paratype 9 3.9 mm (MNHN-Pg 5629). a. propodus and dactyl of left fourth pereopod,
lateral view; b. propodus and dactyl of left fourth pereopod. lateral view; c, same (setae omitted), ventrolateral view;
d. propodus and dactyl of left fifth pereopod, lateral view; e, telson, dorsal view; f-i. left (f.h) and right (g.i) exopods
of uropods, dorsal view. Scales equal 0.5 mm.
Telson and uropods (Fig. 23e-i) asymmetrical (weakly asymmetrical in specimens living in scaphopod shells).
Left exopod (Fig. 23f,h) relatively short, broad, often paddle-shaped, with moderately broad rasp (see Variations).
Telson without lateral indentations; with scattered setae dorsally, and rows of long setae laterally; terminal margin
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
341
divided into 2 rounded projections by wide, shallow, unarmed rounded (U-shaped) cleft; rounded projections each
armed distally with few (usually 7 or 8) moderately long, well-spaced corneous spines.
Size Range. — Males, SL 1.5 to 1 1.8 mm. Females 2.5 to 5.8 mm. Ovigerous females 3.1 to 1 1.0 mm.
VARIATIONS. — Right cheliped (Fig. 21c-e): in females and small males (SL < 3.0 mm) the length/width ratio
of the palm varies slightly from about 0.9 to 1. In large males (SL > 3.0 mm) the lenglh/width ratio of the palm
varies from about 1.1 to 1.6. The length of the carpus increases with size more in males than in females.
Left cheliped (Fig. 21a-b): the armature of the dorsal margin of the carpus varies from one to several irregular
rows of spines.
Ambulatory legs (Fig. 22a-f): the length/height ratio of the propodi varies from 3.4 to 5.4 (first leg), and 3.1
to 5.3 (second leg). The length/height ratio of the meri varies from 3.4 to 3.9 (first leg), and 2.8 to 3.3 (second
leg).
Uropods (Fig. 23f-i): the left exopod is usually about twice as long as broad, but can vary from about 2 to
2.3 limes as long as broad; the anterior margin varies from subsemicircular to broadly rounded.
Habitat. — Gastropod shells (often with anthozoan polyp); occasionally scaphopod shells.
Distribution (Figs 47-48). — Central and western Pacific, including the New Caledonia region; South China
Sea; southwestern Indian Ocean. Depth: 31 1 to 4470 m.
Etymology. — The species name is given in honor of Bertrand Richer de Forges, esteemed colleague
carcinologist from ORSTOM, in recognition of his outstanding contributions to the success of numerous French
expeditions in New Caledonia and other poorly known Pacific regions.
Affinities. — Parapcigurus richeri sp. nov. is most similar to P. nudus (A. Milne-Edwards, 1891), from the
Atlantic (see Lemaitre, 1986, 1989), and to a lesser extent to P. stenorhinus sp. nov. The three share, among
other characters, the presence of a broad left uropodal exopod that often has a subsemicircular anterior margin (e.g.
Figs 23 f, 35e). Parapcigurus richeri sp. nov. can be distinguished from P. nudus by subtle differences in the dactyls
of the ambulatory legs, and the left exopod of the uropods. The dactyls of the ambulatory legs are longer relative to
the propodi in P. richeri sp. nov. than in P. nudus ; the dactyls vary from 1.7 to 2.1 times as long as the propodi
in the former, and are 1.6 times or less as long as the propodi in the latter. The left exopod ot the uropods is
narrower in P. richeri sp. nov. than in P. nudus\ the exopod is 2 to 2.3 times as long as broad in the former,
whereas the exopod is usually less than 2 times as long as broad in P. nudus.
P. richeri sp. nov. can be differentiated from P. stenorhinus sp. nov. by several characters: the rounded
projections of the terminal margin of the telson, which are each armed distally with less than ten well-spaced
spines in the former (Fig. 23e), and more than ten closely-spaced spines in the latter (Fig. 35d); the armature of the
mesial margin of the antennal acicles, with one to three distinct spines in the former (Fig. 19a-c), and five to eight
spines in the latter (Fig. 32a-b); and the rasp of the left uropodal exopod, moderately broad in the former
(Fig. 23f,h), and narrow in the latter (Fig. 35e,g).
Parapcigurus richeri sp. nov. and P. furici sp. nov. are superficially similar. The two are sympatric in
distribution (Fig. 48), and are often captured in abundant numbers in the same sample. They can be distinguished
by a number of characters, such as the number of rows and shape of the scales on the propodal rasp of the fourth
pereopod (one row of ovate scales in P. richeri sp. nov., two rows of conical scales in P. furici sp. nov.); armature
of the antennal acicles (armed with distinct spines in P. richeri sp. nov., usually unarmed or at most weakly armed
proximally in P. furici sp. nov.); degree of slenderness of the ambulatory legs; and armature of the terminal margin
of the telson.
Remarks. — De Saint Laurent (1972) considered that the distribution of Parapagurus pilosimanus nudus
included the Indo-Pacific and the Atlantic. However, when LEMAITRE (1989) elevated P. p. nudus to specific rank
he determined that this taxon occurred only in the Atlantic. Examination of the Indo-Pacilic specimens used by
DE SAINT LAURENT (1972) in her report of P. p. nudus has shown that she confounded that subspecies with
the two new species P. richeri and P. stenorhinus.
Source :
342
R. LEMAITRE
Parapagurus furici sp. nov.
Figs 24-27, 47-48
Parapagurus pilosimanus - ALCOCK, 1901: 218; 1902: 133. 273, fig. 67 (See Remarks).
MATERIAL EXAMINED. — Japan. " Albatross' ': stn 4975, 33°2r30”N, 135°38'50”E. 1302-997 m, 31.08.1906:
1 8 8.3 mm (USNM 276130). — Stn 4980, Kobe to Yokohama. 34°09’N, 137°55’E. 927 m, 1.09.1906: 2 8
7.0,9.5 mm (USNM 276131). — Stn 5082, 34°05’N, 137°59'E, 1210 m, 20.10.1906: 3 9 8.7-11.5 mm (USNM
276129).
RV "Tansei-Maru": off Taito-saki, Boso Peninsula, 35°07'N. 1 40C55'E, 1351-1454 m, 24.04.1995, coll. T. KOMAi:
1 8 7.2 mm (USNM 276118).
South China Sea. Taiwan . RV " Fisheries Research l", off Tungsha Tao (Pratas Island), trawled. 1520 m,
25.04.1996, coll. Ding-An Lee: 1 8 7.0 mm (NTOU 1996-4-25).
Philippines. RV'Fishery Researcher /", stn 5-95, Lagonoy Gulf, 13°21.32'N, 124°12.26’E. 1037-1100 m,
24.09.1995: 1 8 8.3 mm (ZRC 1998.54), 1 9 6.3 mm (ZRC 1998.5).
Indonesia. Celebes Sea. " Siboga ": stn 208, 5°39'S, 122°12'E, 1866 m, 22.09.1899: 1 8 7.0 mm (ZMA). —
Stn 210a, 5°26’S, 121°18'E, 1944 m. 24.09.1899: 1 8 1.2 mm (ZMA).
New Caledonia. Biocal: stn CP 05, 21°16.49'S, 166°43.56'E. 2340 m, 11.08.1985: I ov. 9 5.7 mm (MNHN-Pg
5581) . — Stn CP 23. 22°45.84'S, 166°20.33’E, 2040 m, 28.08.1985: 1 8 5.5 mm, 3 ov. 9 4.2-6.1 mm (MNHN-Pg
5582) . — Stn CP 26, 22°39.66’S. 166°27.41E, 1618-1640 m. 28.08.1985: 3 8 4.2-5.7 mm, 1 9 3.7 mm, 1 ov. 9
4.9 mm (USNM 276128). — Stn CP 27, 22°05.52‘S. 166026.41’E. 1850-1900 m. 28.08.1985: 2 8 4.3, 5.2 mm, 4 9
3. 4-4. 3 mm, 1 ov. 9 3.7 mm (MNHN-Pg 5583). — Stn CP 30, 23°08.44'S, 166°40.83'E, 1140 m, 29.08.1985: 1 ov. 9
4.2 mm (MNHN-Pg 5584). — Stn CP 57. 23°43.26'S, 166°58.06’E, 1490-1620 m. 1.09.1985: 3 8 3.3-5.2 mm, 2 9
3.9, 4.8 mm, 3 ov. 9 3.6, 4.6 mm (MNHN-Pg 5585). — Stn CP 62, 24°19.06’S, 167°48.65,E, 1395-1410 m,
2.09.1985: 2 8 4.0, 4.5 mm, 1 ov. 9 6.9 mm (USNM 276127). — Stn CP 63, 24°28.69’S, 168°07.72'S, 2160 m,
2.09.1985: 3 8 3.2-4.9 mm (MNHN-Pg 5586). — Stn CP 72, 22°09.02'S. 167°33.I8’E, 2100-2110 m, 4.09.1985: 2 8
4.09, 6.7 mm, 2 9 4.0, 4.5 mm, 4 ov. 9 4.8-6.0 mm (USNM 276143).
Biogeocal: stn CP 214, 22°43.09’S, 166°27.19'E, 1665-1590 m, 9.04.1987: 1 8 3.2 mm, I ov. 9 4.6 mm (MNHN-
Pg 5587). — Stn CP 243, 21°27.35’S, 166°25.76'E, 1820 m. 15.04.1987: 1 8 5.8 mm, 1 ov. 9 5.2 mm (MNHN-Pg
5588). — Stn CP 250, 21°24.63’S, 166028.21'E, 2350 m, 15.04.1987: 2 ov. 9 5.4, 5.5 mm (MNHN-Pg 5589). —
Stn CP 260, 2 1 mOO'S. 167°58.34'E, 1820-1980 m, 17.04.1987: 2 8 4.6, 6.7 mm, 2 9 5.5, 6.1 mm, 4 ov. 9 5.2-
6.0 mm (MNHN-Pg 5590). — Stn CP 265, 21°04.09’S, 167°00.40’E, 1760-1870 m, 18.04.1987: 3 8 3.0-7.0 mm,
3 ov. 9 4.9-5. 7 mm (MNHN-Pg 5591). — Stn CP 266. 21°04.85'S, 167°57.14’E, 2100-1990 m. 18.04.1987: 2 8 6.0.
6.1 mm, 1 9 5.1 mm, 1 ov. 9 4.5 mm (MNHN-Pg 5592). — Stn 272, 21°00.04’S, I66°56.94'E, 1615-1710 m,
20.04.1987: 2 9 3.7. 5.7 mm (MNHN-Pg 5593). — Stn 273, 21°01.53'S. 166°57.41’E, 1920-2040 m, 20.04.1987: 7 c*
3.0-7. 1 mm, 4 9 4.0-4.6 mm, 4 ov. 9 4.2-4.8 mm (MNHN-Pg 5594). — Stn CP 283, 21°22.25’S, 166°31.07’E, 2375-
2370 m, 26.04.1987: 3 8 5.8-6.6 mm. 1 ov. 9 4.3 mm (MNHN-Pg 5595). — Stn DW 296, 20°38.35'S. 167°I0.32'E,
1230-1270 m, 28.04.1987: 1 9 3.0 mm (MNHN-Pg 5596). — Stn CP 317, 20°48.12’S. 166°53.16’E, 1630-1620 m,
2.05.1987: 1 8 5.5 mm, 4 ov. 9 5. 1-6.3 mm (USNM 276127). — Stn CP 321, 21°12.00,S, 166°59.85’E, 2190-2205 m,
3.05.1987: 1 8 5.5 mm (MNHN-Pg 5597). — Stn CP 329, 21°09.05’S, 166°40.08’E. 2315-2310 m, 4.05.1987: 1 8 6.7
mm (MNHN-Pg 5580); 1 8 5.4 mm (MNHN-Pg 5598). — Stn CP 336, 21°12.22’S, 166022.51'E, 2370-2380 m,
5.05.1987: 1 8 5.1 mm (MNHN-Pg 5599). — Stn CP 341, 21°29.73’S, 166°47.37*E, 2334 m, 6.05.1987: 2 8 4.7,
6.6 mm, 1 ov. 9 5.1 mm (MNHN-Pg 5600).
Bathus 1: stn CP 651, 21°41'S, 166°40'E, 1080-1 180 m. 1 1.03.1993: 4 8 4.6-5.2 mm. 2 9 5.9, 6.1 mm (MNHN-Pg
5601). — Stn CP 660, 21°10’S, 165°53'E, 786-800 m, 13.03.1993: 1 ov. 9 6.0 mm (MNHN-Pg 5602).
Bathus 3: stn CP 822, 23°19'S, 167°57’E, 950-980 m, 29.11.1993: 1 ov. 9 5.6 mm (MNHN-Pg 5603). —
Stn CP 844, 23°06'S, 166°45’E. 908 m, 1.12.1993: 10 8 6.4-6.6 mm, 8 ov. 9 3.7-5.6 mm (MNHN-Pg 5604).
Halipro 1: stn CC 856, 21°44’S, 166°37,E. 311-365 m. 20.03.1994: 1 ^ 6.1 mm (MNHN-Pg 5605). — Stn CH 876.
23°10'S, 166°49'E, 870-1000 m, 31.03.1994: 2 8 5.6. 6.5 mm. 3 ov. 9 4.6, 5.4 mm (MNHN-Pg 5606).
Vanuatu. MUSORSTOM 8: stn CP 956, 20°33’S, 169°35'E, 1175-1210 m, 20.09.1994: 3 8 3. 9-6.0 mm, 2 9 2.9-
5.2 mm, I ov. 2 3.7 mm (MNHN-Pg 5608). — Sin DW 987, 19°23'S, 169°35'F„ 1050-1040 m. 23.09.1994: I ov. 2
5.3 mm (MNHN-Pg 5609). — Stn CP 1037, 18°03'S, 168°54'E, 1058-1086 m, 29.09.1994: 1 2 7.5 mm (MNHN-Pg
5610). — Stn CP 1076, 15°53’S, 167°30'E, 1100-1191 m. 4.10.1994: 1 8 10.5 mm (MNHN-Pg 561 1 ). — Stn CP 1125,
15°57'S, 166°38'E, 1160-1220 m, 10.10.1994: 4 8 6.4-10.1 mm (MNHN-Pg 5612). — Stn CP 1126, 15°58'S,
166°39'E, 1210-1260 m, 10.10.1994: 5 8 3.5-6.5 mm, 6 ov. 2 5. 2-7.5 mm (MNHN-Pg 5613).
Wallis and Futuna. MUSORSTOM 7: stn CP 623, 12°34'S, 178°15'W, 1280-1300 m, 28.05.1992: I 2 4 0 mm
(MNHN-Pg 5607).
Tasman Sea. ORV "Franklin" (coll. J.K. Lowry): sin FRO 589-17. 29°42.06'S. 1 59°48.3 1 E. 2450 m. 3.05.1989:
1 8 8.8 mm, 1 ov. 2 9.6 mm (AM P39450). — Stn FRO 589-32, 27°11.97'S, 160°37.80’E, I960 m, 7.05.1989: 2 8
Source :
PARAPAGURIDAE PROM THE PACIFIC AND INDIAN OCEANS
343
7.8, 9.5 mm, 1 6 5.5 mm, 2 ov. 9 7.0. 7.1 mm (AM P39448). — Sm FRO 589-35, 26°51.57’S, 159°48.72’E. 2500 m,
8.05.1989: 2 6 5.8, 9.2 mm, 5 6 4. 6-6.3 mm, 5 ov. 9 4. 9-6. 7 mm (AM P39449).
Arabian Sea. "Investigator", [sin unknown], 1289-2195 m. [dale unknown): 1 6 7.9 mm (AM P2623).
Types. — Holotype : 6 6.7 mm, Biogeocal, sin CP 329, 21°09.05'S, 166°40.08’E, 2315-2310 m. 4.05.
1987 (MNHN-Pg 5580). Paratypes : All Ihe others specimens mentioned above.
Fig. 24. — Parapagurus furici sp. nov., New' Caledonia, Biogeocal stn CP 329: a,b,f-h, holotype 6 6.7 mm (MNHN-Pg
5580); c-e,i,j, paralype 6 5.4 mm (MNHN-Pg 5598). a. shield and cephalic appendages; b. right antennal peduncle
and anterolateral margin of branchiostegite, lateral view'; c. left maxillule, internal view; d, distal end of endopod of
same; e. left third maxilliped, internal view; f. telson, dorsal view; g-h, left (g) and right (h) exopods of uropods,
dorsal view; i, male first pleopod, mesial view; j. male second pleopod. anterior view'.
Scales equal 1 mm (a,b,e), 0.5 mm (c.g-j,f). and 0.25 mm (d).
Source : MNHN. Paris
344
R. LEMAITRE
DESCRIPTION. — Shield (Fig. 24a) about as long as broad; dorsal surface usually well calcified, with rows of
short setae posteriorly on each side of midline; anterior margin weakly concave; lateral projections broadly rounded;
anterolateral margin sloping. Rostrum broadly triangular, rounded distally, slightly overreaching lateral
projections; with low mid-dorsal ridge. Anterodistal margin of branchiostegite (Fig. 24b) rounded, unarmed, setose.
Ocular peduncles (including corneae) about half length of shield, each with dorsal longitudinal row of setae;
peduncles inflated basally. Ocular acicles subtriangular, terminating in strong spine; separated basally by slightly
less than basal width of 1 acicle.
Antennular peduncles slender, long, exceeding distal margins of corneae by at least 0.23 length of penultimate
segments. Ultimate and penultimate segments with scattered setae. Ultimate segment nearly twice as long as
penultimate. Basal segment with ventromesial distal spine; mesial face unarmed; lateral lace with statocyst lobe
having subrectangular distal lobe usually armed with 1 small spine, and 1 spine proximally.
Fig. 25. — Parapagurus furici sp. nov.: a-b. New Caledonia, Biogeocal, stn CP 329, liolotype 6 6.7 mm (MNHN-Pg
5580); c. New Caledonia, Biogeocal, stn 273, paratype ov. 9 4.8 mm (MNHN-Pg 5594); d. Tasman Sea,
ORV "Franklin", sin FRO 589-35, 6 9.2 mm (AM P39449). a. left cheliped; b-d. right chelipeds (setae omitted
in c-d). Scales equal 3 mm (a, b, d), and 2 mm (c).
Antennal peduncles (Fig. 24b) exceeding distal margins of corneae by about half length of fifth segments. Fifth
segment with setae on lateral and mesial margins. Fourth segment with scattered setae. Third segment with strong
ventromesial distal spine. Second segment w'ith dorsolateral distal angle produced, terminating in strong multifid
Source .
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
345
spine; mesial margin with spine on dorsodistai angle. First segment with or without small blunt spine on lateral
face; ventromcsial angle produced, with row of small spines. Antennal acicles nearly straight in dorsal view,
setose; exceeding distal margin of cornea by 0.3 or more length of acicle; mesial margin usually unarmed, or at
most with 1-3 small spines on proximal half. Flagellum distinctly overreaching extended right cheliped; articles
with setae less than 1 to 2 flagellar articles in length.
Mandible, maxilla, and first and second maxillipeds typical of species in genus (e.g. Fig. 20). Maxillule
(Fig. 24c-d) with external lobe of endopod weakly developed, internal lobe with long seta. Third maxilliped
(Fig. 24e) with crista dentata consisting of about 10 small corneous teeth; coxa and basis each with 1 tooth
mesially. Sternite of third maxilliped with spine on each side of midline. Epistomial spine usually absent.
Chelipeds markedly dissimilar, each with dorsal surfaces of carpus and chela covered with moderately dense
setation. Right cheliped (Fig. 25b-d) with proportions of carpus and chela influenced by size and sexual
dimorphism (see Variations). Fingers bent inwards at lips; each terminating in small corneous claw; with few tufts
of setae on dorsal and ventral surfaces; cutting edges each with irregularly-sized calcareous teeth; cutting edge of
dactyl also with distal row of small, closely-set, corneous teeth. Dactyl set at slightly oblique angle to palm, with
dorsomesial and mesial row of small spines proximally. Palm and carpus each with numerous small spines and
tubercles on dorsal surface; ventral surfaces of palm and carpus also with spines and tubercles but less numerous or
sometimes few and scattered. Merus with small tubercles on dorsal, dorsolateral and ventral surfaces; mesial surface
with scattered spines or tubercles, with ventromcsial row of spines. Ischium with dorsal and ventromcsial row of
spines. Coxa with 1 or 2 spines on ventrodista! margin and ventromesial row of setae.
Left cheliped (Fig. 25a) slender. Fingers each terminating in small corneous claw; dorsal and ventral surfaces
with scattered tufts of short setae; cutting edge of dactyl with row of minute, closely-set, corneous teeth distally;
cutting edge of fixed finger with row of regularly-spaced, small, evenly-sized, calcareous teeth. Palm unarmed ex¬
cept for few small dorsomesial and sometimes dorsolateral tubercles. Carpus armed with irregular row of small
spines dorsally. Merus unarmed except for row of bristles on dorsal margin, and 1 or 2 small spines on ventromc¬
sial margin. Ischium with small, blunt setose tubercles on dorsal margin; usually with small spine on ventrome¬
sial margin proximally. Coxa usually with 2 small spines on ventrodistal margin and ventromesial row of setae.
Ambulatory legs (Fig. 26a-f) similar from right to left (except slightly longer segments on right), slender,
long, distinctly overreaching right cheliped. Dactyl about 1.7 to 1.9 times as long as propodus; with dorsal and
ventromesial distal row of setae; ventromesial margin with row of up to 12 minute corneous spinules. Merus,
carpus, and propodus each with short setae on dorsal margin; segments distinctly more slender in large specimens
(SL > 6.0 mm, e.g. Fig. 26a-f); carpus with small dorsodistai spine. Ischium usually with small ventrodistal and
ventroproximal spine (first leg) or unarmed (second leg). Coxa with ventromesial margin usually armed with 1 or
2 small spines (first leg) or unarmed (second leg). Anterior lobe of sternite of second legs (Fig. 26g) setose, armed
with simple subterminal spine.
Fourth pereopod (Fig. 27a-d)) semichelate. Dactyl subtriangular, shorter than length of propodal rasp,
terminating in corneous claw; with ventrolateral row of small, closely-set. corneous spines. Propodal rasp with
2 irregular rows of conical scales (Fig. 27b), or occasionally with 1 row at least distally (Fig. 27d). Carpus with
row of long setae on dorsal margin. Merus with setae on dorsal and ventral margins.
Fifth pereopod (Fig. 27e) chelate; propodal rasp forming subtriangular area less than half length of propodus.
Telson and uropods (Fig. 24f-h)) weakly asymmetrical. Left exopod (Fig. 24g) about 2.6 times as long as
broad; rasp moderately broad. Telson without or at most weakly marked lateral indentations; with scattered setae
dorsally, and rows of long setae laterally; terminal margin divided into 2 rounded projections by narrow, angled
(V-shaped) cleft; rounded projections each armed distally with 15 or more closely-spaced corneous spines.
Size Range. — Males, SL 3.0 to 10.5 mm. Females 2.9 to 1 1.5 mm. Ovigerous females 3.6 to 9.6 mm.
Variations. — Right cheliped: in males the length/width ratio of the palm varies from about 1.2 to 1.7
(Fig. 25b, d); in females the palm is usually about as long as wide or slightly wider than long (Fig. 25c). The
length of the carpus increases with size more in males than in females. Ambulatory legs (Fig. 26a-l). the
length/height ratio of the propodi varies from 5.2 to 7.9 (first leg), and 5.0 to 7.0 (second leg). The length/height
ratio of the meri varies from 4.0 to 5.1 (first leg), and 3.4 to 4.4 (second leg).
Source :
346
R LEMAITRE
Fig. 26. — Parapagurus fund sp. nov., New Caledonia: a-d,g, Biogeocal stn CP 329, holotype 6 6.7 mm (MNHN-Pg
5580); e-f, Biogeocal stn 273. paratype ov. 9 4.8 mm (MNHN-Pg 5594). a. left first ambulatory leg. lateral view;
b, dactyl of same, mesial view; c, left second ambulator)' leg. lateral view; d. dactyl of same, mesial view; e. left first
ambulatory leg. lateral view; f. left second ambulatory leg. lateral view; g, sternite of second ambulatory legs,
ventral view.
Scales equal 3 mm (a-d), 1 mm (e.f), and 0.5 mm (g).
Habitat. — Usually found living in shelters formed by zoanthids.
Distribution (Figs 47-48). — Western Pacific, including the New Caledonia region; Arabian Sea. Depth: 31 1
to 2500 m.
Etymology. — The species is dedicated to Pierre Furic, captain of the French research vessels
N.O. "Vauban" and N.O. "A lis" during many MUSORSTOM deep-sea expeditions. The success of these expeditions
was largely due to his expertise.
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
347
Affinities. — As previously mentioned, P.furici sp. nov. and P. richeri sp. nov. arc superficially similar but
can be distinguished by differences in the armature of antennal acicles, slenderness of ambulatory legs, number of
rows and shape of scales of the propodal rasp of the fourth pereopod, and armature of the terminal margin of the
telson (sec Affinities under P. richeri sp. nov.).
Compared to other Parapagurus species, the degree of slenderness of the ambulatory legs exhibited in specimens
of P.furici sp. nov. is often quite striking. The length/width ratios of the meri and propodi of these appendages in
this new species frequently serve as useful characters for identification of specimens.
Fig. 27. — Parapagurus furici sp. nov.. New Caledonia: a-b,e, BlOGEOCAL stn CP 329, holotype 6 6.7 mm (MNHN-Pg
5580); c-d, Biogeocal stn CP 260, paratype 6 4.6 mm (MNHN-Pg 5590). a, propodus and dactyl of left fourth
pereopod, lateral view; b, same (setae omitted), ventrolateral view; c, propodus and dactyl of left fourth pereopod,
lateral view; d, same (setae omitted), ventrolateral view; e. propodus and dactyl of left fifth pereopod, lateral view.
Scales equal 0.5 mm (a,b,e), and 0.25 mm (c.d).
REMARKS. — ALCOCK (1901) reported many specimens collected on the "Investigator" as Parapagurus
pilosimanus. ALCOCK (1902) mentioned and depicted the same taxon in a general account of the deep sea fauna
from Indian Seas based on samples obtained during the cruises of the "Investigator". One of ALCOCK's specimens,
a male collected in the Arabian Sea and deposited in the Australian Museum (AM P2623), has been examined; it
represents the new species P.furici. The remaining specimens need to be located and studied in order to determine
whether they also represent this or possibly other species.
Parapagurus saintlaurentae sp. nov.
Figs 28-31, 47, 49
Parapagurus pilosimanus pilosimanus - DE Saint Laurent, 1972: 102 (in part, see Remarks).
Parapagurus pilosimanus scaher - DE SAINT LAURENT, 1972: 102 (in part, see Remarks).
Material EXAMINED. — Indian Ocean.
Mozambique Channel. " Galathea ": stn 217, 14°20'S, 45°09'E, 3390 m. 27.02.1951: 2 6 12.7, 15.5 mm (ZMK
CRU-3390, 3391).
N of Madagascar. "Galathea": stn 231, 8°52'S, 49°25’E, 5020 m. 7.03.1951: 1 ov. 9 11.9 mm (ZMK CRU-3392).
— Stn 232, 9°03'S, 49°22’E, 4930 m. 8.03.1951: 1 6 9.5 mm, 1 ov. 9 10.0 mm (ZMK CRU-3393, 3394). — Stn 233,
7°24'S, 48°24'E, 4730 m, 9.03.1951: 2 6 1 1.6, 12.1 mm, 2 9 7.5, 10.1 mm, 2 ov. 9 9.8, 10.1 mm (ZMK CRU-3395).
Source
348
R LEMAITRE
— Stn 234, 5°25’S, 47°09’E. 4820 m, 10.03.1951: 6 8 5.4-14.7 mm. 4 9 S.3-9.7 mm. 5 ov. 9 10.4-12.1 mm (ZMK
CRU-3396). — Stn 235. 4°47'S. 46°19'E. 4810 m. 11.03.1951: 8 13.0 mm (ZMK CRU-3397); 24 8 9.1-13.7 mm.
10 9 9.1-11.6 mm, 32 ov. 9 8.8-10.7 mm (ZMK CRU-3398. 3399, 3400), 4 8 12.5-13.4 mm. 1 9 1 1.2 mm. 3 ov. 9
10.0-10.7 mm (USNM 276123).
SE iles Glorieuses. Benthedi: stn BENT 90-CH, 1 1 °44S. 47°30’E. 3700 m. 4.04.1977: 1 9 7.8 mm (MNHN-Pg
5645).
Fig. 28. — Parapagurus saintlaurentcie sp. nov., Indian Ocean, " Galathea ", stn 235: a.e, holotype 8 13.0 mm (ZMK CRU-
3397); b, paratype 8 12.4 mm (ZMK CRU-3398); c, paratype 9 1 1.1 mm (ZMK CRU-3398); d. paratype 9 1 1.0 mm
(ZMK CRU-3398); f-j, paratype 8 12.2 mm (ZMK CRU-3398). a, shield and cephalic appendage; b. anterior portion
of shield and cephalic appendages; c-d. ocular acicles, dorsal view; e. right antennal peduncle and anterolateral
margin of branchiostegite, lateral view; f, left maxillule, internal view; g, distal end of endopod of same; h, left third
maxilliped, internal view; i, male left first pleopod, mesial view; j. male left second pleopod, anterior view.
Scales equal 3 mm (a,b), 0.5 mm (c,d,g), 2 mm (e,h), and 1 mm (f,i,j).
Source : MNHN. Paris
PARAPAGUR1DAE FROM THE PACIFIC AND INDIAN OCEANS
349
Kenya. "Galathea": stn 238. 3°23'S, 44°04'E, 3960 m. 13.03.1951: 5 <J 6.3-9.2 mm. 4 9 5.1-1. 9 mm, 3 ov. 9‘
7. 2-8.0 mm (ZMK CRU-3401).
SW of Sri Lanka (Ceylon). "Galathea": stn 279, 1°00'N. 76°17'E. 4320 m. 8.04.1951: 4 <5 6.0-12.0 mm,
1 ov. 9 9.2 mm (ZMK CRU-3402). — Stn 282, 5°32'N, 78°41'E. 4040 m. 11.04.1951: 1 6 8.5 mm, 1 9 12.1 mm
(ZMK CRU-3403).
Types. — Holotype: 6 13.0 mm ."Galathea", stn 235, N of Madagascar. 4°47'S, 46°19'E. 4810 nt,
1 1.03.1951 (ZMK CRU-3397). Paratypes : All the others specimens mentioned above.
Description. — Shield (Fig. 28a) about as long as broad; dorsal surface usually well calcified, with scattered
short setae; anterior margin weakly concave; lateral projections broadly rounded; anterolateral margin sloping.
Rostrum broadly subtriangular, rounded distally, overreaching lateral projections; lacking or with inconspicuous
low mid-dorsal ridge. Anterodistal margin of branchiostegite (Fig. 28e) armed with 1 or more small spines, setose.
Ocular peduncles (including corneae) distinctly less than half length of shield, each with few setae dorsally;
peduncles inflated basally. Ocular acicles (Fig. 28a-d) subtriangular. usually terminating in strong spine (rarely
bifid or trifid on one or both sides); separated basally by slightly less than basal width of one acicle.
Antennular peduncles slender, long, exceeding distal margins of corneae by at least 0.8 length of penultimate
segments. Ultimate and penultimate segments with scattered setae. Ultimate segment nearly twice as long as
penultimate. Basal segment with ventromesial distal spine; mesial face unarmed; lateral face with statocyst lobe
having subrectangular distal lobe armed with 1-3 small spines, and 1 spine proximally.
Antennal peduncles (Fig. 28e) exceeding distal margins of corneae by at least 0.2 length of fifth segments.
Fifth segment with scattered setae on lateral and mesial margins. Fourth segment with scattered setae. Third
segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating
in strong multifid spine; mesial margin with spine on dorsodistal angle. First segment unarmed on lateral lace;
ventromesial angle produced, with row of small spines. Antennal acicles (Fig. 28a-b) nearly straight in dorsal
view, frequently very long and slender (Fig. 28b). sparsely setose; exceeding distal margin of cornea by half or
more length of acicle; mesial margin unarmed or with I or 2 small spines (occasionally with up to 5 small, well
spaced spines). Flagellum distinctly overreaching extended right cheliped. with sparse short setae less than
1 flagellar articles in length.
Mandible, maxilla, and first and second maxillipeds typical of species in genus (e.g. Fig. 20). Maxillule
(Fig. 28f-g) with external lobe of endopod weakly developed, internal lobe with long terminal seta and 4
subterminal setae. Third maxillipcd (Fig. 28h) with crista dentata consisting of about 20 small corneous teeth;
coxa and basis each with I tooth mesially. Sternite of third maxilliped with spine on each side of midline.
Epistomial spine usually absent.
Chelipeds markedly dissimilar, each with dorsal surfaces of carpus and chela covered with moderately dense
setation; proportions of carpus and chela influenced by size and sexual dimorphism (see Variations). Right cheliped
(Fig. 29c-e) with fingers bent inwards at tips, each terminating in small corneous claw; with tufts of setae on
dorsal and ventral surfaces; cutting edges each with irregularly-sized calcareous teeth; cutting edge of dactyl also
with distal row of small, closely-set. corneous teeth. Dactyl set at oblique angle to palm, with dorsomesial and
mesial row of small spines proximally. Palm and carpus each with numerous small spines and tubercles on dorsal
and ventral surfaces (spines and tubercles usually less numerous on ventral surfaces). Merus with small tubercles
on dorsal, dorsolateral and ventral surfaces; mesial surface smooth; with ventromesial row of spines. Ischium with
dorsal and ventromesial row of spines. Coxa with 1 or 2 spines on ventrodistal margin and ventromesial row of
setae.
Left cheliped (Fig. 29a-b) slender, more so in males than in females. Fingers each terminating in small
corneous claw; dorsal and ventral surfaces with scattered tufts of short setae; cutting edge of dactyl with row of
minute, closely-set, corneous teeth distally; cutting edge of fixed finger with row of small regularly-spaced and
evenly-sized calcareous teeth. Palm with dorsomesial row of small spines; dorsolateral face with small spines.
Carpus with irregular row of small spines dorsally; lateral face with scattered small spines or tubercles. Merus with
row of setae dorsally; ventromesial margin with row of spines. Ischium armed with blunt spines or tubercles
dorsally, and ventromesial row of spines. Coxa usually with 2 small spines on ventrodistal margin and
ventromesial row of setae.
Source :
350
R. LEMAITRE
Fig. 29. — Parapagurus sainilaurentae sp. nov., Indian Ocean, " Galathea ", stn 235: a,c, holotype 6 13.0 mm (ZMK CRU-
3397); b,d, paratype 6 13.5 mm (ZMK CRU-3398); e, paratype 9 10.7 mm (ZMK CRU-3398). a-b, left chelipeds
(setae omitted); c-e, right chelipeds (setae omitted). Scales equal 5 mm.
Ambulatory legs (Fig. 30a-f) similar from right to left (except slightly longer segments on right), slender,
long, distinctly overreaching right cheliped. Dactyl about 1.5 times as long as propodus; with dorsal and
ventromesial distal row of setae (missing in holotype. Fig. 30a-b,e-f); ventromesial margin with row of about 7 to
15 minute corneous spinules. Meri, carpi, and propodi each with small low tubercles bearing short stiff setae on
dorsal margin. Meri and carpi each usually with dorsal margin ridge-like (e.g., Fig. 30c-d); merus 4.7 (first leg) or
3.9 (second leg) times as long as high. Carpus with rounded or often somewhat subglobular dorsodistal region
marked by shallow oblique depression and with 1-3 small spines (Fig. 30c-d). Ischium with vcntrodistal row of
small spines (first leg), or small setose distal tubercle (second leg). Coxae of legs (Fig. 30g-h) each with
ventromesial margin forming a lobe proximally, lobe often strongly produced and armed with 1 or more small
spines. Anterior lobe of sternite of second legs (Fig. 30g-h) setose, armed with simple subterminal spine.
Fourth pereopod (Fig. 31a-d) semichelate. Dactyl sublriangular; shorter than length of propodal rasp, termi¬
nating in corneous claw; with ventrolateral row of small, closely-set, corneous spines. Propodal rasp usually with
2 (Fig. 31b) or occasionally 3 (Fig. 31a) irregular rows of lanceolate or conical scales. Carpus with row of long
setae on dorsal margin. Merus with setae on dorsal and ventral margins.
Fifth pereopod (Fig. 3 le) chelate; propodal rasp forming subtriangular area less than half length of propodus.
Telson and uropods (Fig. 3 1 f-i) weakly asymmetrical. Left exopod (Fig. 3 1 g) elongate, about 2.6 times as long
as broad; rasp moderately broad. Telson without or at most with weak lateral indentations, and scattered setae
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
351
dorsally; terminal margin divided into 2 rounded projections by wide, often deep rounded (U-shaped cleft); rounded
projections each armed distally with moderately long corneous spines (about 15 left, 12 right).
pIG 3Q — Parapagurus saintlaurentae sp. nov., Indian Ocean, "Galathea‘\ stn 235: a-g, hojotype 6 13.0 mm (ZMK CRIJ-
3397); h, paratype 6 13.6 mm (ZMK CRU-3398). a. left first ambulatory leg, lateral view; b. dactyl of same, mesial
view; c, carpus of same, lateral view; d, same, dorsal view; e, left second ambulatory leg, lateral view; f, dactyl ot
same, mesial view; g-h, coxae and sternites of ambulatory legs, ventral view.
Scales equal 5 mm (a,b,e,f-h), and 3 mm (c.d).
Size Range. — Males, SL 5.4 to 15.5 mm. Females 5.7 to 12.1 mm. Ovigerous females 7.2 to 12.1 mm.
VARIATIONS. — The fingers of the right cheliped in large specimens (SL >13.0 mm) often leave a wide hiatus
when closed (Fig. 29d). In males (Fig. 29c-d), the length/width ratio of the palm varies from 1.4 to 1.7; in females
(Fig. 29e), the palm is usually about as long as wide. The length of the carpus increases with size more in males
than in females.
Habitat. — Found living in zoanthids, or actinians that secrete a chitinous carcinoecia.
DISTRIBUTION (Figs 47, 49). — Known so far only from the Indian Ocean, north ol Madagascar, and
southwest of Sri Lanka (Ceylon). Depth: 3390 to 5020 m.
Etymology. — The specific name is given in honor of the eminent French carcinologist, Mme Michele de
Saint Laurent, in recognition of her many important contributions to our knowledge of decapod crustaceans in
general, and of parapagurids in particular.
AFFINITIES. — See P. holthuisi.
Source
352
R. LEMAITRE
Fig. 31. — Parapagurus saintlaurentae sp. nov., Indian Ocean: a-b,e-h. "Galathea", sin 235. hololype 6 13.0 mm (ZMK
CRU-3397); c-d, " Galathea " sin 217. paralype 6 15.5 mm (ZMK CRU-3390); i. Benthedi sin BENT 90-CH. paralype
9 7.8 mm (MNHN-Pg 5645). a, propodus and dactyl of left fourth pereopod, lateral view; b. same (setae omitted),
ventrolateral view; c, propodus and dactyl of left fourth pereopod, lateral view; d, same (setae omitted), ventrolateral
view; e, propodus and dactyl of left fifth pereopod. lateral view; f, telson, dorsal view; g-h, left (g) and right (h)
exopod of uropods. dorsal view; i, telson. dorsal view. Scales equal 1 mm.
Remarks. — Examination of the Indo-Pacific material used by de Saint Laurent (1972) in her report of
P. p. pilosimanus and P. p. scaber , has shown that she confounded it with three species. De Saint Laurent's
Indo-Pacific material of P. p. pilosimanus actually contains specimens of P. saintlaurentae sp. nov. and, as
previously mentioned, specimens of P. latinianus. De Saint Laurent's Indo-Pacific material of P. p. scaber
contains specimens of the new species P. saintlaurentae and P. stenorhinus.
Parapagurus stenorhinus sp. nov.
Figs 32-35, 47. 49
Parapagurus pilosimanus scaber - DE Saint LAURENT, 1972: 102 (in part, see Remarks).
Parapagurus pilosimanus nudus - DE Saint LAURENT, 1972: 102 (in part, see Remarks under Parapagurus richer i sp. nov.).
MATERIAL EXAMINED. — Indian Ocean.
South Africa. "Galathea": stn 192, off Durban., 32°00’S, 32°41’E, 3530 m, 5.02.1951: I 6 6.4 mm (ZMK CRU-
3405).
N of Madagascar. "Galathea": stn 232, 9°03'S, 49°22'E, 4930 m, 8.03.1951: 1 ov. 9 5.5 mm (ZMK CRU-3404).
Kenya. "Galathea": stn 238, 3°23'S. 44°04'E. 3960 m, 13.03.1951: 9 6 3.6-7.5 mm. 8 9 3.3-6.0 mm, 2 ov. 9 3.7.
5.8 mm (ZMK CRU-3406. 3407).
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
353
SW of Sri Lanka (Ceylon). " Galathea stn 280. 1°56,N. 77°05’E, 4465-4530 m, 9.04.1951: 1 9 5.6 mm (ZMK
CRU-3408).
Fig. 32. — Parapagurus sienorhinus sp. nov.. Indian Ocean, "Galathea": a,c stn 232, holotype ov. $ 5.5 mm (ZMK CRU-
3404); b, stn 192. paratype 6 6.4 mm (ZMK CRU-3405); d-h, stn 238, paratype 6 4.9 mm (ZMK CRU-3405).
a, shield and cephalic appendages; b. anterior portion of shield and cephalic appendages (setae omitted); c, right
antennal peduncle and anterolateral margin of branchiostegite. lateral view; d. maxillule, internal view; e. distal
portion of endopod of same; f. left third maxilliped. internal view; g, male left first pleopod, mesial view; h, male
left second pleopod, anterior view.
Scales equal 1 mm (a-d,f-h), and 0.25 mm (e).
Source
354
R. LEMAITRE
TYPES. — Holotype: ov. 9 5.5 mm, '' Galathea ", stn 232, N of Madagascar, 9°03'S. 49°22'E, 4930 m,
8.03.1951 (ZMK CRU-3404). Paratypes : All the others specimens mentioned above.
Description. — Shield (Fig. 32a) about as long as broad; dorsal surface usually well calcified except for small
irregularly-shaped areas; with scattered short setae; anterior margin weakly concave; lateral projections broadly
rounded; anterolateral margin sloping. Rostrum broadly subtriangular, rounded distally, overreaching lateral
projections; with low mid-dorsal ridge. Anterodistal margin of branchiostegite (Fig. 32c) rounded, unarmed, setose.
Ocular peduncles (including corncae) about half length of shield, each with dorsal longitudinal row of setae;
peduncles inflated basally. Ocular acicles subtriangular, terminating in strong spine; separated basally by slightly
less than basal width of one acicle.
Antennular peduncles slender, long, exceeding distal margins of corneae by at least 0.25 length of penultimate
segments. Ultimate and penultimate segments with scattered setae. Ultimate segment nearly twice as long as
penultimate. Basal segment with ventromesial distal spine; mesial face unarmed; lateral face with statocyst lobe
having subrectangular distal lobe armed with 1 or 2 small spines, and 1 spine proximally.
Antennal peduncles (Fig. 32c) exceeding distal margins of corneae by about half length of fifth segments. Fifth
segment with few setae on lateral and mesial margins. Fourth segment with scattered setae. Third segment with
strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong
multifid spine; mesial margin with spine on dorsodistal angle. First segment with lateral face unarmed or with
small spine; ventromesial angle produced, with row of small blunt spines. Antennal acicles nearly straight in
dorsal view, setose; exceeding distal margin of cornea by about half length of acicle; mesial margin armed with
5 to 8 small spines (distalmost often very low, Fig. 32a-b). Flagellum distinctly overreaching extended right
eheliped; articles with setae less than 1 to 2 flagellar articles in length.
Mandible, maxilla, and first and second
maxillipeds typical of species in genus (e.g.
Fig. 20). Maxillule (Fig. 32d-e) with external
lobe ofendopod weakly developed, internal lobe
with long seta. Third maxilliped (Fig. 32 f)
with crista dentata consisting of about 15 small
corneous-tipped teeth; coxa and basis each with
1 tooth mesially. Sternite of third maxilliped
with spine on each side of midlinc. Epistomial
spine usually present.
Chelipeds markedly dissimilar, each with
dorsal surfaces covered with sparse setation.
Right eheliped (Fig. 33b-c) with palm slightly
longer than broad (about 1.2 times) in males,
or about as long as broad in females; spines and
tubercles on palm and carpus larger in small
(SL < 3.5 mm; Fig. 33c) individuals. Fingers
bent inwards at tips, each terminating in small
corneous claw; with few tufts of setae on dorsal
and ventral surfaces; cutting edges each with
irregularly-sized calcareous teeth; cutting edge
of dactyl also with distal row of small, closely-
set, corneous teeth. Dactyl set at slightly
oblique angle to palm, with dorsomesial and
mesial row of small spines proximally. Palm
and carpus each with numerous small spines
and tubercles on dorsal surface; ventral surfaces
of palm and carpus also with spines and
Fig. 33. — Parapagurus stenorhinus sp. nov., Indian Ocean,
"Galathea": a-b, stn 232, holotype ov. 9 5.5 mm (ZMKCRU-
3404); c. stn 238, paratype 9 3.3 mm (ZMK CRU-3407).
a, left eheliped; b-c, right chelipeds (setae omitted).
Scales equal 1 mm (a-b), and 0.5 mm (c).
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
355
tubercles but less numerous, tubercles on palm scattered (some often arranged in 2 median rows). Merus with
scattered small tubercles on dorsal, dorsolateral and ventral surfaces; mesial surface smooth; with ventromesial row
of spines. Ischium with dorsal and ventromesial row of spines. Coxa with 1 or 2 spines on ventrodistal margin
and ventromesial row of setae.
Left cheliped (Fig. 33a) slender. Fingers each terminating in small corneous claw; dorsal and ventral surfaces
with scattered tufts of short setae; cutting edge of dactyl with row of minute, closely-set, corneous teeth; cutting
edge of fixed finger with row of regularly-spaced, small, evenly-sized calcareous teeth. Palm unarmed except for
small setose tubercles on dorsomesial margin. Carpus armed with irregular row of small setose spines or tubercles
dorsally. Merus with dorsal margin unarmed except for row of bristles; ventromesial margin with small spine
distally and another proximally. Ischium with dorsal margin setose; with small ventromesial spine distally. Coxa
usually with 2 small spines on ventrodistal margin and ventromesial row of setae.
Fig. 34. — Parapagurus stenorhinus sp. nov., Indian Ocean, " Galathea ", stn 232, holotype ov. 9 5.5 mm (ZMK CRU-
3404): a, left first ambulatory leg, lateral view; b, dactyl of same, mesial view; c. left second ambulatory leg, lateral
view; d. dactyl of same, mesial view; e, coxae and sternites of ambulatory legs. Scales equal 2 mm (a-d), and I mm (e).
Ambulatory legs (Fig. 34a-d) similar from right to left (except for slightly longer segments on right), slender,
long, distinctly overreaching right cheliped. Dactyl about 1.8 times as long as propodus; with dorsal and
356
R. LEMAITRE
ventromesial distal row of setae; ventromesial margin with row of about 8 to 10 minute corneous spinules. Merus,
carpus, and propodus each with short bristles on dorsal margin; carpus with small dorsodistal spine; merus about
3.6 (first leg) or 2.9 (second leg) times as long as high. Ischium usually with small ventrodistal tubercle (first leg)
or unarmed (second leg). Coxa (Fig. 34e) with ventrodistal margin usually armed with 1 or 2 small spines (first
leg) or unarmed (second leg). Anterior lobe of sternite of second leg (Fig. 34e) setose, unarmed or with simple
subterminal spine.
Fourth pereopod (Fig. 35a-b) semichelate. Dactyl subtriangular, shorter than length of propodal rasp,
terminating in corneous claw; with ventrolateral row of small, closely-set, corneous spines. Propodal rasp with
1 row of ovate scales at least distally; rasp sometimes with 2 short irregular rows of rounded scales proximally.
Carpus with row of long setae on dorsal margin. Merus with setae on dorsal and ventral margins.
Fifth pereopod (Fig. 35c) chelate; propodal rasp forming subtriangular area less than half length of propodus.
Telson and uropods (Fig. 35d-f) asymmetrical. Left exopod (Fig. 35e) usually broad, paddle-shaped, about
2 times as long as broad; with narrow rasp. Telson without or at most with weakly marked lateral indentations;
with scattered setae dorsally; terminal margin divided into 2 rounded projections by wide unarmed, rounded
(U-shaped) cleft; rounded projections each armed distally with moderately long corneous spines (about 19 left,
13 right).
Fig. 35. — Parapagurus stenorhinus sp. nov., Indian Ocean. "Galathea": a-f, stn 232, holotype ov. 9 5.5 mm (ZMK CRU-
3404); g, stn 238, paratype 6 1 A mm (ZMK CRU-3406). a, propodus and dactyl of left fourth pereopod, lateral view;
b, same (setae omitted), ventrolateral view; c, propodus and dactyl of left fifth pereopod, lateral view; d, telson,
dorsal view; e-g, left (e,g) and right (f) exopod of uropods. dorsal view.
Scales equal 0.25 mm (a-d,g), and 0.5 mm (e-f).
Source
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
357
Size Range. — Males, SL 3.1 to 7.5 mm. Females 3.3 to 6.0 mm. Ovigerous females 3.7 to 5.8 mm.
Habitat. — One of the specimens examined was lodged in gastropod shell, another in a scaphopod shell. The
remaining specimens examined were without housing.
DISTRIBUTION (Figs 47, 49). — Indian Ocean, from off eastern Africa to southwest of Shri Lanka (Ceylon).
Depth: 2400 to 4930 m.
Etymology. — The specific name is a compound used as an adjective, derived from the Greek stenos, narrow,
and rhine , rasp, and refers to the narrow rasp of the left uropodal exopod found in this species.
Affinities. — See P. riche ri sp. nov..
REMARKS. — Several Indo-Pacific specimens identified by DE SAINT LAURENT (1972) as Parapagurus
pilosimanus scaber have been found to represent the new species P. stenorhinus.
Parapagurus janetae sp. nov.
Figs 36-38, 47. 50
Parapagurus abyssorum Henderson, 1888: 87 (in part, see Remarks). — MURRAY. 1895: 1 140 (in part). [Not Parapagurus
abyssorum (Filhol, 1885a)].
? Parapagurus pilosimanus - PORTER, 1906: 129. — Haig, 1955: 17. [See Remarks under Parapagurus abyssorum (Filhol,
1885a)].
MATERIAL EXAMINED. — Eastern Pacific.
Galapagos Islands. " Albatross stn 2807. 0°24’S, 89°06'W, 1485 m, 4.04.1888: I $ 4.8 mm (USNM 276120);
1 8 4.5 mm, 2 ov. $ 5.1, 5.2 mm (USNM 276121).
Chile. Port Otway, Golfo de Penas, ’'Challenger": stn 304, 46°53’15MS, 75°12’W. 82 m, 31.12.1875: 1 $ 5.7 mm
(AM G. 1653).
TYPES. — Holotype : 9 4.8 mm, " Albatross ", stn 2807, Eastern Pacific. Galapagos Islands, 0°24'S, 89°06'W.
1485 m, 4.04.1888 (USNM 276120). Paratypes : All the others specimens mentioned above.
Description. — Shield (Fig. 36a) about as long as broad; dorsal surface usually well calcified, with rows of
short setae posteriorly on each side of midline; anterior margin weakly concave; lateral projections broadly rounded;
anterolateral margin sloping. Rostrum broadly subtriangular, rounded distally, slightly overreaching lateral
projections; with low mid-dorsal ridge. Anterodistal margin of branchiostegite (Fig. 36b) rounded, unarmed, setose.
Ocular peduncles (including corneae) about half length of shield, each with long setae dorsally; peduncles
inflated basally. Ocular acicles subtriangular, terminating in strong simple spine; separated basally by about basal
width of one acicle.
Antennular peduncles slender, long, exceeding distal margins of corneae by half or more length of penultimate
segments. Ultimate and penultimate segments with scattered setae. Ultimate segment nearly twice as long as
penultimate. Basal segment with ventromesial distal spine; mesial face unarmed; lateral face with statocyst lobe
having subrectangular distal lobe armed with 1 or 2 small spines, and 1 spine proximally.
Antennal peduncles (Fig. 36b) exceeding distal margins of corneae by about half length of fifth segments. Fifth
segment with setae on lateral and mesial margins. Fourth segment with scattered setae. Third segment with strong
ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong multifid
spine; mesial margin with spine on dorsodistal angle. First segment with lateral face unarmed; ventromesial angle
produced, with row of small spines. Antennal acicles weakly curved in dorsal view, setose: exceeding distal margin
of cornea by 0.2 or more length of acicle; mesial margin armed on proximal half with 2-5 small spines. Flagellum
distinctly overreaching extended right cheliped; articles with setae less than 1 to 2 flagellar articles in length.
Mandible, maxilla, and first and second maxillipeds typical of species in genus (e.g. Fig. 20). Maxillule
(Fig. 36c) with external lobe of endopod weakly developed, internal lobe with long seta. Third maxilliped
358
R LEMAITRE
(Fig. 36d) with crista dentata consisting of about 13 small distal corneous teeth and 2 large partially fused
corneous teeth proximally; coxa and basis each with 1 tooth mesially. Sternite of third maxilliped with spine on
each side of midline. Epistomial spine present.
Fig. 36. — Parapagurus janetae sp. nov., eastern Pacific, " Albatross ", stn 2807: a-b,c-f, holotype 2 4.8 mm (USNM
276120); c-d, paratype ov. 2 5.2 mm (USNM 276121). a, shield and cephalic appendages; b, right antennal peduncle
and anterolateral margin of branchiostegite, lateral view; c, left maxilliped, internal view; d, left third maxilliped,
internal view; e, left cheliped (most setae omitted); f, right cheliped (most setae omitted).
Scales equal 1 mm (a-b,d-f), and 0.5 mm (c).
Chelipeds markedly dissimilar, each with dorsal surfaces of carpus and chela covered with moderately dense
setation. Right cheliped (Fig. 36f) with proportions of carpus and chela not markedly different in male and female
specimens examined. Fingers bent inwards at tips, each terminating in small corneous claw; with tufts of setae on
dorsal and ventral surfaces; cutting edges each with irregularly-sized calcareous teeth; cutting edge of dactyl also
with distal row of small, closely-set, corneous teeth. Dactyl set at slightly oblique angle to palm, with
dorsomesial and mesial rows of small spines proximally. Palm and carpus each with numerous small spines and
Source : MNHN. Pans
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
359
tubercles on dorsal surface; ventral surfaces of palm and carpus also with spines and tubercles but less numerous
than on dorsal surfaces. Merus with small setose tubercles dorsally, and scattered small spines or tubercles
dorsolaterally and on ventral surface; mesial surface smooth; with ventromesial row of spines. Ischium with blunt
spines dorsally. and ventromesial row of spines. Coxa usually with 1 or 2 small spines on ventrodistal margin and
ventromesial row of setae.
Left chelipcd (Fig. 36e) slender. Fingers each terminating in small corneous claw; dorsal and ventral surfaces
with scattered tufts of short setae; cutting edge of dactyl with row of minute corneous teeth fused distall y; cutting
edge of fixed finger with row of regularly-spaced, small, evenly-sized calcareous teeth. Palm with 1-3 small spines
or tubercles dorsomesially, and small spines dorsolaterally and laterally. Carpus armed with row of small spines
Fig. 37. — Parapagurus jane toe sp. nov., eastern Pacific, "Albatross” , stn 2807: a-e. holotype 9 4.8 mm (USNM
276120); f-g, paratype 6 4.5 mm (USNM 276121). a, left first ambulatory leg, lateral view; b, dactyl of same, mesial
view; c, left second ambulatory leg. lateral view; d. dactyl of same, mesial view; e. coxae and sternites of ambulatory
legs, ventral view; f, male left first pleopod, mesial view; g. male left second pleopod, anterior view.
Scales equal 1 mm (a-e), and 0.5 mm (f,g).
Source : MNHN. Paris
360
R. LEMAITRE
dorsally. Merus unarmed except for row of bristles on dorsal margin; with ventromesial row of spines. Ischium
with small, low setose tubercles on dorsal margin; with 2 small spines on ventromesial margin (one distally, one
proximally). Coxa usually with 2 small spines on ventrodistal margin and ventromesial row of setae.
Ambulatory legs (Fig. 37a-d) similar from right to left (except for slightly longer segments on right), slender,
long, distinctly overreaching right cheliped. Dactyl about 1.7 times as long as propodus; with dorsal and
ventromesial distal row of setae; ventromesial margin with row of about 5 to 9 minute corneous spinules. Meri,
carpi, and propodi each with short setae on dorsal margin, and smooth lateral and mesial laces; propodi about 3.8
to 4.0 (first leg) or 3.8 to 4.3 (second leg) limes as long as high; carpi each with small dorsodistal spine; meri
about 3.3 to 3.5 (first leg) or 2.4 to 3.0 (second leg) times as long as high. Ischia unarmed except for setae
distally. Coxae (Fig. 37e) with ventromesial margin unarmed or at most with minute spine proximally. Anterior
lobe of sternite of second legs (Fig. 37e) setose, armed w-ith simple subterminal spine.
Fourth pereopod (Fig. 38a-b) semichelate. Dactyl subtriangular, shorter than length of propodal rasp,
terminating in corneous claw; with ventrolateral row of small, closely-set, corneous spines. Propodal rasp with
1 row of ovate scales at least distally. Carpus with row of long setae on dorsal margin. Merus with setae on dorsal
and ventral margins.
Fifth pereopod (Fig. 38c) chelate; propodal rasp forming subtriangular area extending at most to about
midlength of propodus.
Fig. 38. — Parapcigurus janetae sp. nov., eastern Pacific, " Albatross ", stn 2807: a-f. holotype 9 4.8 mm (USNM
276120); g. paratype 6 4.5 mm (USNM 276121). a, propodus and dactyl of left fourth pereopod, lateral view;
b. same (setae omitted), ventrolateral view; c, propodus and dactyl of left fifth pereopod, lateral view; d. telson,
dorsal view; e-f, left (e) and right (0 exopod of uropods, dorsal view: g, telson. dorsal view. Scales equal 0.5 mm.
Source : MNHN. Pans
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
361
Telson and uropods (Fig. 38d-g) weakly asymmetrical. Left exopod (Fig. 38e) elongate, about 2.9 times as
long as broad; with moderately broad rasp. Telson without or at most with weakly marked lateral indentations;
with scattered setae dorsally, and few long setae laterally; terminal margin divided into 2 rounded projections by
unarmed, rounded (U-shaped) cleft (very shallow in females, deep in only known male; Fig. 38g); rounded
projections each armed distally with 8 to 14 short to moderately long corneous spines.
Size Range. — Only known male, SL 4.5 mm. Females 4.8, 5.7 mm. Ovigerous females 5.1, 5.2 mm.
Habitat. — According to Henderson (1888: 89), the specimen obtained during the "Challenger" expedition
from Port Otway, was found living in a gastropod shell ( Trochus sp.).
Distribution (Figs 47, 50). — Eastern Pacific: Galapagos Islands, and coast of Chile (Port Otway, Golfo de
Penas). Depth: 82 to 1485 m.
ETYMOLOGY. — The specific name is given in memory of Janet HAIG, whose quiet but prolific career while
associated with the Allan Hancock Foundation contributed greatly to our knowledge of anomuran crustaceans from
throughout the world in general, and the eastern Pacific in particular.
AFFINITIES. — To some degree, this new species resembles Parapagurus riche ri sp. nov. and P . nudus. All
three are more or less similar in size, have propodal rasps on the fourth pcreopods consisting of a single row ol
ovate scales distally, and the antennal acicles are similarly armed with spines. However, P. janetae sp. nov. can be
separated from the other two by the presence of a distinctly elongated left uropodal exopod, which is about
2.9 times as long as broad (Fig. 38e); the exopod is at most 2.3 times as long as broad in P. richeri sp. nov.
(Fig. 23f,h), and about 2 times as long as broad in P. nudus. In addition, the carpus of the left cheliped in
P. janetae sp. nov. is armed dorsally with strong spines (Fig. 36e); the carpus is armed with one or more irregular
rows of small spines in P. richeri sp. nov. (Fig. 21a-b), and is unarmed in P. nudus. So far P. janetae sp. nov. is
known only from the southeastern Pacific, P. richeri sp. nov. from the Indo-Pacific and central Pacific, and
P. nudus from the Atlantic.
REMARKS. — The single specimen collected during the "Challenger" Expedition at stn 304, Port Otway,
Chile, and reported by HENDERSON (1888) and MURRAY (1895) as Parapagurus abyssorum, actually represents
P. janetae sp. nov. The relatively shallow depth of 82 m recorded at "Challenger", stn 304 was attributed to an
error by HENDERSON (1888: 89); however, Parapagurus species have occasionally been collected at depths similar
to that recorded for stn 304, particularly at higher latitudes.
Parapagurus foraminosus sp. nov.
Figs 39-42, 47, 50
Parapagurus pilosimanus abyssorum - Faxon, 1895: 68. [Not Parapagurus abyssorum (Filhol, 1885a); see Remarks].
Parapagurus abyssorum - DE Saint LAURENT. 1972: 103 (in part; see Remarks). [Not Parapagurus abyssorum (Filhol,
1885a)].
MATERIAL EXAMINED. — Eastern Pacific.
Gulf of California. Farallon Basin, stn VSS-54. 23°58.4’N, 108°59.5'W. 2798 m, 8.03.1959. coll. Wisner: 1 6
7.0 mm, 2 9 6.7, 10.0 mm, 3 ov. 9 7.9-9.4 mm (LACM 59-286.1). — Off Cape San Lucas, stn P-41-59, 22°32.5'N.
109°40.8'W, 2780-2807 m, 22.03.1959, coll. Parker: 2 6 9.5. 10.3 mm (LACM 59-284.1). — Stn VSS-17, 22°35.6'N,
1 10°06.5'W, 2634-2663 m, 26.03.1959. colls Parker & Lemche: 1 9 7.8 mm. 1 ov. 9 1 1.5 mm (LACM 59-285.1).
Gulf of Panama. "Galathea": stn 739, 7°22’N, 79°32'W, 915-975 m. 15.05.1951: 1 ov. 9 6.3 mm (ZMK CRU-
3409).
Baja California to Ecuador. "Albatross": stn 2793, off Ecuador, Galera Point, 1°03'N. 80°15W, 1306 m.
3.03.1888: 9 c? 4.0-8.9 mm. 3 9 6.4-8.2 mm. 8 ov. 9 6.0-9.6 mm (USNM 18999). — Stn 2807. Galapagos Islands.
San Cristobal Island. 0°24'S. 89°06'W. 1485 m, 4.04.1888: 1 6 8.2 mm (USNM 276122); 6 6 6.3-8.4 mm. 5 9 5.8-
8.8 mm (USNM 19000). — Stn 2986. Mexico, off Baja California. 28°57'N, 1 18°14'30"W, 1251 m. 28.02.1889: 1 6
7 9 mm (USNM 276109). — Stn 3362, Cocos Island. 5°56'N. 85°10’30''W. 2149 m. 26.02.1891: 1 6 6.2 mm (USNM
Source :
362
R LEMAITRE
21667). — Stn 3363, Cocos Island. 5°43’N, 85°50’W, 1789 m, 26.02.1891: 7 8 5. 1-9.1 mm. 2 9 5.6. 9.0 mm. 3 ov.
9 7. 9-9. 6 mm (USNM 21668). — Stn 3364. Cocos Island, 5°30'N, 86°08,30’,W, 1650 m. 27.02.1891: 1 8 (damaged)
(USNM 42623). — Stn 3366. Cocos Island. 5°30'N, 86°45’W, 1951 m, 27.02.1891: 2 8 8.6, 9.5 mm, 2 8 4.5, 7.8 mm
(USNM 21669).— Stn 3371, off Colombia and Panaml 5°26’20,,N, 86°55'W. 1408 m. 1.03.1891: 6 8 7.6-9.9 mm, 5 9
6. 7-9.0 mm, ov. 9 7. 0-9. 6 mm (MCZ 4524). — Stn 3375. off Colombia. Malpelo Island, 2°34'N, 82°29'W, 2197 m,
4.03.1891: 1 <3 8.8 mm (USNM 21670). — Stn 3376. off Colombia, Malpelo Island, 3°09'N, 82°08’W. 2070 m,
4.03.1891: I 8 5.2 mm. 1 9 6.1 mm (USNM 42626): 1 8 4.6 mm, 1 9 3.0 mm (USNM 42627). — Stn 3380, off
Colombia, Malpelo Island. 4°03’N, 81°3rw, 1644 m, 5.03.1891: 1 8 6.4 mm (USNM 21671). — Stn 3392, Gulf of
Panama, SE of Azuero Peninsula, 7°05’30MN, 79°40’W. 2323 m, 10.03.1891: 1 8 6.0 mm. 1 9 3.7 mm (USNM 21672).
— Stn 3393, Gulf of Panama, SE of Azuero Peninsula, 7°15,00MS, 79°36,00MW. 1866 m, 10.03.1891: 2 8 8.2, 10.0
mm, 2 9 7.9, 8.9 mm (USNM 21673). — Stn 3400, E off Galapagos Islands, 0°36'S, 86°46’W, 2418 m. 27.03.1891: I
8 8.6 mm, 1 ov. 9 8.7 mm (USNM 21674); 3 8 6.0-10.3 mm. 2 9 7.3, 9.2 mm (MCZ 20.020). — Stn 3407, N of
Galapagos Islands, San Salvador Island, 0°04'S, 90°24'30"W, 1619 m, 3.04.1891: fragments of specimen (USNM
21675). — Stn 3413. Galapagos Islands, Culpepper Island, 2°34’, 92°06'W. 2487 m, 5.04.1891: 1 8 10.4 mm (USNM
21676); 1 8 9.2 mm, 2 9 10.4. 12.5 mm, 1 ov. 9 11.5 mm (MCZ 4526). — Stn 3429, off Mexico, Mazatian,
Fig. 39. — Parapagurus foraminosus sp. nov., eastern Pacific, "Albatross" : a.c.g.h, stn 2807. holotype <3 8 2 mm
!£SNM 276122); b. stn 2807, paratype <5 7.9 mm (USNM 19000); d-f, stn 5685. paratype <J 8.5 mm (USNM
^ / 6 1 1 _ ) . a, shield and cephalic appendages; b, ocular acicles and rostrum, dorsal view; c, right antennal peduncle and
anterolateral margin ol branchiostegite, lateral view; d. left maxillule, internal view; e. distal end of endopod of
same; f, left third maxilliped, internal view; g, left cheliped (setae omitted); h. right cheliped (setae omitted)
Scales equal 2 mm (a), 1 mm (b-d,f), 0.25 mm (e), and 2 mm (g,h).
Source : MNHN. Pahs
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
363
22°30'30"N, 107°01'W, 1681 m, 19.04.1891: 2 6 6.4 mm (parasitized), 9.2 mm (USNM 21677). — Stn 3431. 23°59'N,
108°40'W, 1820 m, 20.04.1891: 1 6 11.0 mm, 1 9 10.4 mm (USNM 21678). — Stn 3432, E of Baja California,
Cerralvo Island, 24°22,30”N, 109°03,20,,W, 2599 m, 20.04.1891: 1 ov. 9 7.8 mm (USNM 21679). — Stn 5676, Baja
California. 25°3ri5"N, 113o29'30MW. 1180 m, 17.03.1911: 7 6 6.7-9.4 mm, 4 9 7.8-9.9 mm. 11 ov. 9 7.9-9.7 mm
(USNM 276113). — Stn 5685, Baja California, S of Abreojos Point, 25°42’45"N, 113o38’30MW. 1180 m. 22.04.1911:
5 6 7. 0-8. 7 mm, 5 9 7.8-9.3. mm, 4 ov. 9 6.7-9.6 mm (USNM 276112); 3 6 7. 8-8.7 mm, 1 9 8.7 mm, 2 ov. 9 8.0,
9.2 mm (MNHN-Pg 5647). — Stn 5686, Mexico, SW of Abreojos Point, 26°14’N. 1I4°00'W( 1701 m, 22.04.1911: 1 6
8.9 mm (USNM 276111). — Stn 5690, Mexico, E of Guadalupe Island, 29°29'N. 116°18’W, 2014 m, 24.04.1911: 1 6
8.1 mm (USNM 276110).
Types. — Holotype : cJ 8.2 mm, " Albatross ", stn 2807, Eastern Pacific. Galapagos Islands, San Cristobal
Island, 0°24’S, 89°06'W, 1485 m, 4.04.1888 (USNM 276122). Paratypes : All the others specimens mentioned
above.
Description. — Shield (Fig. 39a) about as broad as long or at most slightly broader than long; dorsal surface
usually well calcified, with rows of short setae posteriorly on each side; anterior margin weakly concave; lateral
projections broadly rounded; anterolateral margins sloping. Rostrum broadly subtriangular, rounded distally,
overreaching lateral projections; with low mid-dorsal ridge. Anterodistal margin of branchiostegite (Fig. 39c)
rounded, unarmed, setose.
Ocular peduncles (including corneae) less than half length of shield, each with dorsal row of setae; peduncles
inflated basally; width of cornea about same as distal width of ocular peduncle. Ocular acicles subtriangular,
terminating in strong simple or bifid spine (Fig. 39a-b); separated basally by slightly less than basal width of one
acicle.
Antennular peduncles slender, exceeding distal margins of corneae by half or more length of penultimate
segments. Ultimate and penultimate segments with scattered setae; ultimate segment nearly twice as long as
penultimate. Basal segment with ventromesial distal spine; mesial face unarmed; lateral face with statocyst lobe
having subrectangular distal lobe armed with 1 or 2 small spines, and I spine proximally.
Antennal peduncles (Fig. 39c) exceeding distal margins of corneae by about 0.6 length of fifth antennal
segments. Fifth segment with setae on lateral and mesial margins. Fourth segment with scattered setae. Third
segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced into strong,
usually bifid spine; mesial margin with spine on dorsodistal angle. First segment with lateral face unarmed or with
1-3 small blunt spines; ventromesial angle produced, with row of small spines. Acicles weakly curved in dorsal
view, exceeding distal margin of corneae by 0.3 to half length of acicle; mesial margin armed with 3 to 8 small
spines on proximal half. Flagellum overreaching extended right cheliped; with setae about 1 to 2 flagellar articles
in length.
Mandible, maxilla, and first and second maxillipeds typical of species in genus (e.g. Fig. 20). Maxillule
(Fig. 39d-e) with external lobe weakly developed, internal lobe with 1 long terminal and 2 subterminal setae.
Third maxilliped (Fig. 39f) with crista dentata consisting of about 17 small corneous teeth; coxa and basis each
with strong sharp mesial tooth. Epistomial spine usually present. Sternite of third maxillipeds with strong spine
on each side of midline.
Chelipeds markedly dissimilar. Right cheliped (Fig. 39h) with dorsal surfaces of carpus and chela covered with
dense setation. Fingers bent inwards at tips, each terminating in small corneous claw; with tufts of setae dorsally
and ventrally; cutting edges each with irregularly-sized calcareous teeth; cutting edge of dactyl with distal row ot
small, closely-set, corneous teeth. Dactyl about as long as mesial margin of palm; set at slightly oblique angle to
palm; with numerous small spines on dorsomesial and mesial margins. Fixed finger with few small spines on
dorsal face proximally, lacking spines distally. Palm and carpus each densely covered with small (often minute)
spines on dorsal surfaces; with scattered small spines on ventral surfaces. Merus with dorsal row of short bristle¬
like setae; with small spines or tubercles on lateral and ventral surfaces, and ventromesial row of spines. Ischium
with dorsal and ventromesial row of spines. Coxa with row of spines on ventrodistal margin and ventromesial row
of setae.
Left cheliped (Fig. 39g) well calcified, with dense setation on carpus and chela. Fingers each terminating in
short corneous claw; dorsal and ventral surfaces with scattered tufts of short setae; cutting edge of dactyl with row
Source :
364
R. LEMAITRE
of minute, closely-set, corneous teeth; cutting edge of fixed finger with row of small, evenly-spaced, calcareous
teeth interspersed with small corneous teeth. Palm with irregular rows of small spines on dorsolateral and
dorsomesial faces. Carpus with irregular rows of small spines dorsally; ventral face unarmed. Merus with row of
short bristle-like setae on dorsal margin; with few small spines on lateral and ventral faces. Ischium with small
setose tubercles dorsally; ventromesial margin with small spine proximally and another distally. Coxa with row of
spines on ventrodistal margin and ventromesial row of setae.
Ambulatory legs (Fig. 40a-d) similar from right to left (except slightly longer segments on right). Dactyl
about 1.6 times as long as propodus; with ventromesial row of 2 to 8 minute spinules. and dorsal and dorsomesial
distal rows of setae. Meri, carpi, and propodi each with lateral faces with shallow pits more numerous on second
leg (Figs 40a,c, 41). Propodi about 5 times (first leg) or 4.5 times (second leg) as long as high; with short
transverse rows of bristle-like setae on dorsolateral and dorsomesial faces. Carpi each with small dorsodistal spine;
dorsal margins with short, bristle-like setae often set at bases of small spines or tubercles. Meri each about 3.7
(lirst leg) or 3 (second leg) times as long as high; dorsal margins with short, bristle-like setae; with longitudinal
Source : MNHN. Pans
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
365
Fig. 41. — Parapagurus foraminosus sp. nov., eastern Pacific, Cocos Island. "Albatross", stn 3363, paratype 6 9.4 mm
(USNM 21668): ischium, merus, carpus, and portion of propodus, of left second ambulatory leg, showing pits, lateral
view (x 4.2).
row of setae on ventral half of lateral face. Merus and ischium of first leg each with row of small spines on ventral
margins; merus and ischium of second leg unarmed or with few small blunt spines. Coxa with ventrodistal margin
unarmed or with small spines. Anterior lobe of sternite of second legs (Fig. 40e) subsemicircular, setose, with
short subterminal spine.
Fourth pereopod (Fig. 42a-d) semichelate. Dactyl subtriangular, shorter than length of propodal rasp,
terminating in corneous claw; with ventrolateral row of small corneous spines. Propodal rasp consisting ot 1
or 2 rows of ovate scales. Carpus with long setae on dorsal margin. Merus with setae on dorsal and ventral
margins.
Fifth pereopod (Fig. 42e) chelate; propodal rasp occupying subtriangular area less than half length of propodus.
Telson and uropods (Fig. 42f-h) asymmetrical. Left exopod (Fig. 42g) of uropod about 2.3 times as long as
broad, with broad rasp. Telson lacking or at most with weakly marked lateral indentations; with scattered setae
dorsally, and long setae laterally; terminal margin divided into 2 rounded projections by rounded (U-shaped) cleft;
rounded projections each armed distally with long, often curved, corneous spines (usually about 10 to 15,
occasionally up to 26 on left).
Size Range. — Males, SL 4.0 to 1 1.0 mm. Females 3.0 to 10.4 mm. Ovigerous females 5.8 to 1 1.5 mm.
Variations. — Small individuals (SL < 5.0 mm) usually exhibit less numerous pits on the lateral faces of
meri, carpi and propodi of the ambulatory legs, pits which are often more clearly visible by tilting the appendage
to a ventrolateral view.
Habitat. — Found in gastropod shells often completely covered by an actinian.
DISTRIBUTION (Figs 47, 50). — Eastern Pacific: Baja California to Ecuador, including Cocos Island and
Galapagos Islands. Depth: 915 to 2807 m.
AFFINITIES. — This new species shares only general similarities with other Parapagurus species having ovate
scales on the propodal rasp of the fourth pereopod. The dense spination on the dorsal surfaces of the merus, carpus
and chela of the right cheliped resembles that of P. microps. In both species the dorsal surfaces are covered with
dense, small (often minute) spines. However, P. foraminosus sp. nov. differs from P. microps and all other
congeners, in having many shallow pits on the lateral faces of the meri, carpi, and propodi of the ambulatory legs
(Fig. 41).
Source :
366
R. LEMAITRE
| - 1 a,b,e,g,h
I - 1 c,d
Fig. 42. — Parapagurus foraminosus sp. nov.. eastern Pacific, "Albatross": a-b,e-h, stn 2807. holoiype 6 8.2 mm
(USNM 276122); c.d. stn 2807, paratype 6 7.2 mm (USNM 19000). a. propodus and dactyl of left fourth pereopod.
lateral view; b. same (setae omitted), ventrolateral view; c, propodus and dactyl of left fourth pereopod, lateral view;
d, same (setae omitted), ventrolateral view; e, propodus and dactyl of left fifth pereopod, lateral view; f. telson. dorsal
view; g-h. left (g) and right (h) exopod of uropods, dorsal view.
Scales equal 0.5 mm (a-b,e-h,). and 1 mm (c-d).
Remarks. — The eastern Pacific material used by de Saint Laurent (1972) in her report of Parapagurus
pilosimanus abyssorum actually represents this new species.
Examination of the material reported by Faxon (1895) as Parapagurus pilosimanus abyssorum has shown that
it represents P. foraminosus sp. nov.
Parapagurus wolffi sp. nov.
Figs 43-47. 50
Parapagurus abyssorum - Garth & Haig, 1971: 5. (See Remarks). [Not Parapagurus abyssorum (Filhol. 1885a)).
Material EXAMINED. — Peru. "Anton Bruun ", cruise 11, stn 169, 8°46'S, 80°44'W, 3909-3970 m 211 1965*
1 ov. 9 4.6 mm (LACM 65-335.1).
Types. — The sole specimen known of this species is the one above which is the holotype.
Source : MNHN. Pahs
PARAPAGUR1DAE FROM THE PACIFIC AND INDIAN OCEANS
367
DESCRIPTION OF HOLOTYPE. — Shield (Fig. 43a) about as long as broad; dorsal surface well calcified, with
longitudinal rows of short setae on each side of midline, and small setose tubercle on each lateral margin medially;
lateral projections broadly rounded; anterolateral margin sloping; anterior margin weakly concave. Rostrum broadly
subtriangular, rounded distally, slightly overreaching lateral projections; with low mid-dorsal ridge. Anterodistal
margin of branchiostegite (Fig. 43b) rounded, unarmed, setose.
Ocular peduncles (including corneae) less than half length of shield, each with long setae dorsally; peduncles
inflated basally, constricted distally near corneae. Ocular acicles subtriangular, terminating in strong simple spine;
separated basally by about basal width of one acicle.
Antcnnular peduncles slender, long, exceeding distal margins of corneae by full length of penultimate
segments. Ultimate and penultimate segments with scattered setae. Ultimate segment about twice as long as
penultimate.
Fig. 43. — Parapagurus wolffi sp. nov., eastern Pacific. "Anion Bruun ", cruise 11, stn 169. holotype ov. $ 4.6 mm
(LACM 65-335.1): a, shield and cephalic appendages: b. right antennal peduncle and anterolateral margin of
branchiostegite, lateral view; c. right cheliped. dorsal view; d, left cheliped, dorsal view. Scales equal 1 mm.
368
R. LEMAITRE
Basal segment with ventromesial distal spine; mesial face unarmed; lateral face with statocyst lobe having
subrectangular distal lobe armed with 1 small spines, and 1 spine proximally.
Antennal peduncles (Fig. 43b) exceeding distal margins of corncae by about half length of fifth segments. Fifth
segment with scattered setae on lateral and mesial margins. Fourth segment with scattered setae. Third segment
with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in
strong spine; mesial margin unarmed (right) or with small blunt spine (left) on dorsodistal angle. First segment
with lateral face unarmed; ventromesial angle produced, with row of small spines. Antennal acicles weakly curved
in dorsal view, densely setose mesially; exceeding distal margin of cornea by about half length of aciclc; mesial
margin unarmed. Flagellum distinctly overreaching extended right cheliped, very setose; articles with setae I to
4 flagellar articles in length.
Mandible, maxilla, and first and second maxillipeds typical of species in genus (e.g. Fig. 20). Maxillule with
external lobe of endopod weakly developed, internal lobe with long seta. Third maxilliped with crista dentata
consisting of about 17 small corneous teeth; basis with 1 tooth mesially. Stcrnite of third maxilliped with spine
on each side of midline. Epistomial spine present.
Chelipeds markedly dissimilar, each with dorsal surfaces of carpus and chela covered with moderately dense
setation. Right cheliped (Fig. 43c) with moderately dense setation. Fingers bent inwards at lips, each terminating
in small corneous claw; with tufts of setae on dorsal and ventral surfaces; cutting edges each with irregularly-sized
calcareous teeth; cutting edge of dactyl also with distal row of small, closely-set, corneous teeth. Dactyl set at
oblique angle to palm, with dorsomesial and mesial rows of spines proximally. Palm and carpus each with
numerous sharp or blunt spines on dorsal surface; ventral surfaces of palm and carpus also with sharp or blunt
spines but less numerous than on dorsal surfaces. Merus with small setose tubercles dorsally, and small spines or
tubercles dorsolatcrally and on ventral surface; mesial surface smooth; with ventromesial row of spines. Ischium
with small spine dorsally, and ventromesial row of spines. Coxa with I small spine vcntrodistally and
ventromesial row of setae.
Left cheliped (Fig. 43d) slender. Fingers each terminating in small corneous claw; dorsal and ventral surfaces
with scattered tufts of short setae; cutting edge of dactyl with row of minute corneous teeth fused distally; cutting
edge of fixed finger with row of regularly-spaced, small, evenly-sized, calcareous teeth. Palm with small setose
tubercles dorsomesially. Carpus moderately setose; with irregular row of small spines dorsally; dorsodistal margin
with small spine laterally setae dorsally. Merus with setae on dorsal margin; with ventromesial and ventrolateral
row of spines. Ischium with small spine dorsally; with 2 small spines on ventromesial margin (one distally. one
proximally). Coxa unarmed, except for ventromesial row of setae.
Ambulatory legs (Figs 44-45) similar from right to left (except slightly longer segments on right), slender,
long, distinctly overreaching right cheliped. Dactyl about twice (first leg) or slightly more than twice (second leg)
as long as propodus; with dorsal and ventromesial distal row of setae; ventromesial margin with row of about 3 or
4 minute corneous spinules. Meri, carpi, and propodi each with numerous setae on dorsal margin, and smooth
lateral and mesial faces. Propodi about 3.2 (first leg) or 3.3 (second leg) times as long as high. Carpi each with
row of spines dorsally (Fig. 45, and small dorsodistal spine. Meri about 3.1 (first leg) or 2.6 (second leg) times as
long as high; each with ventral row of small spines (spines stronger on first leg; Fig. 45). Ischia unarmed except
for dorsal and ventral setae. Coxae (Fig. 44e) with ventromesial margin unarmed, except for row of setae. Anterior
lobe of sternite of second legs (Fig. 44e) setose, armed with simple subterminal spine.
Fourth pereopod (Fig. 46a) semichelate. Dactyl subtriangular, shorter than length of propodal rasp, terminating
in corneous claw; with ventrolateral row of small, closely-set, corneous spines. Propodal rasp with 1 row of ovate
scales at least distally. Carpus with row of long setae on dorsal margin. Merus with setae on dorsal and ventral
margins.
Fifth pereopod (Fig. 46b) chelate; propodal rasp forming subtriangular area extending at most to about
midlength of propodus.
Telson and uropods (Fig. 46c-e) asymmetrical. Left exopod (Fig. 46d) broad, paddle-shaped, about 2 times as
long as broad; with narrow rasp. Telson with weakly marked lateral indentations, and scattered setae dorsally;
terminal margin divided into 2 rounded projections by unarmed, shallow, rounded (U-shaped) cleft; rounded
projections each armed distally with 18 (left) and 10 (right) corneous spines.
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
369
Fig. 44. — Parapagurus wolff i sp. nov., eastern Pacific, "Anton Bruun ", cruise 11. stn 169, holotype ov. 9 4.6 mm
(LACM 65-335.1): a. left first ambulatory leg. lateral view; b, dactyl of same, mesial view; c, left second ambulatory
leg, lateral view; d, dactyl of same, mesial view; e, coxae and stemite of ambulatory legs, ventral view.
Scales equal 1 mm.
SIZE. — Only the holotype is known, an ovigerous female, SL 4.6 mm.
Habitat. — Unknown.
Distribution (Figs 47, 50). — Eastern Pacific: known so far only from the type locality, off Peru. Depth:
3909 to 3970 m.
Etymology. — The name of this species is dedicated to Torben Wolff, Zoological Museum, University of
Copenhagen, in recognition of his important contributions to our knowledge of the deep-sea fauna, and for his
enthusiasm in caring for the rich samples obtained during the Danish "Galcithea" expedition.
Affinities. — This species is clearly distinguished from most others in the genus by the presence of spines on
the dorsal margin of the carpi of the ambulatory legs (Fig. 45). Two other species of Parapapagurus also have
spines on the dorsal margin of the carpi, P. cibyssorum and P. microps. However, the latter differ from P. wolffi
sp. nov. in several other characters, the most visible being the armature of the lateral faces of the meri, carpi, and
propodi of the ambulatory legs; the lateral faces are unarmed in P. wolffi sp. nov., whereas they are armed with
numerous spines in P. cibyssorum and P. microps.
Source : MNHN, Paris
370
R. LEMAITRE
Fig. 45. — Parapagurus wolffi sp. nov., eastern Pacific, "Anion Bruun", cruise 11. stn 169, holotype ov. 9 4.6 mm
(LACM 65-335.1): Merus and carpus of left ambulatory legs, lateral view: a, first leg; b, second leg.
Scale equals 1 mm.
Fig. 46. — Parapagurus wolffi sp. nov., eastern Pacific. "Anton Bruun", cruise 11, stn 169, holotype ov. 9 4.6 mm
(LACM 65-335.1): a. propodus and dactyl of left fourth pereopod, lateral view; b, propodus and dactyl of left fifth
pereopod. lateral view; c, telson. dorsal view; d-e. left (d) and right (e) exopods of uropods. dorsal view.'
Scales equal 0.5 mm.
Source : MNHN. Paris
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
371
In both P. wolffi sp. nov. and P. benedicti , the ventral margins of the meri of the ambulatory legs are armed
with spines; however, these two species differ markedly in the condition of the terminal spine of the ocular acicles,
armature of antennal acicles and left cheliped, number of rows of scales of the rasp of the fourth pereopods, and
shape of the left exopod of the uropods.
The shape and rasp of the left cxopod of the uropods in P. wolffi sp. nov. is very similar to that of
P. stenorhinus sp. nov. The two differ in the armature of the antennal acicles (unarmed in P. wolffi sp. nov.,
armed with spines in P. stenorhinus sp. nov.), and the armature of the carpi of the ambulatory legs (armed with
spines in P. wolffi , unarmed in P. stenorhinus sp. nov.).
BATHYMETRIC AND GEOGRAPHIC DISTRIBUTION OF PARAPAGURUS SPECIES
As previously mentioned. Parapagurus species usually live at depths greater than most other parapagurids, with
the exception of Tylaspis anomala Henderson, 1885, ranging from 3680 to 4344 m. and Probeebei mirabilis
Boone, 1926, ranging from 1145 to 4775 m (Lemaitre, 1998). The bathymetric distribution of all Parapagurus
species is summarized in Figure 47. With the exception of P. bouvieri , the depth range of all species
includes depths of 1000 m or more, and the majority are most frequently collected at mid to lower level depths
Fig. 47. — Bathymetric distribution of species of Parapagurus Smith, 1879 from the world, showing overall range (thin
line) and most frequent range (black bar) for each species. The most frequent range was calculated from a station depth
frequency histogram, and selecting a range that included 80% of station depths symmetrically around the mode. Depth
data includes that used in this study as well as from published sources (e.g. de Saint Laurent, 1972; Macpherson.
1983, 1984; Lemaitre, 1986, 1989, 1990, 1997; Lemaitre & McLaughlin, 1992). Asterisk: species found in New
Caledonia; star: species known exclusively from the Atlantic Ocean.
Source : MNHN. Pans
372
R LEMAITRE
Fig. 48. — Geographical dislribulion of species of Parapagurus Smith, 1879 found in New Caledonia region (large circle).
In some cases symbols indicate more than one station. A "?" indicates exact locality not available.
Fig. 49. — Geographical distribution of Indo-Pacific species of Parapagurus Smith, 1879 not found in New Caledonia
region. For additional localities of P. hohhuisi and P. abyssorum see Fig. 50, and LEMAITRE (1989: 29, fig. 12). In
some cases symbols indicate more than one station.
Source : MNHN. Paris
PARAPAGL'RIDAE FROM THE PACIFIC AND INDIAN OCEANS
373
(1000 to 3000 m) of the continental slope. Six species, P. abyssorum , P. microps , P. holthuisi , P. saintlaurentae
sp. nov., P. stenorhimis sp. nov., and P. wolffi sp. nov., are so far known to occur exclusively below 2000 m; of
these, P. saintlaurentae sp. nov., is found at 5020 m. which is the greatest depth recorded among all parapagurids.
Only two species, P. pilosimanus and P. janetae sp. nov., have been collected in continental shelf depths
(< 200 m), the former at 102 m off the northeastern coast of the United States (Lemaitre, 1989), the latter at
Fig. 50. — Geographical distribution of eastern Pacific species of Parapagurus Smith, 1879. For additional localities of
P. holthuisi and P. abyssorum see Fig. 49, and Lemaitre (1989: 29, fig. 12). In some cases symbols indicate more
than one station.
Source : MNHN, Paris
374
R. LEMAITRE
82 m in the Gulf of Penas, southern Chile. These exceptionally shallow collections (both at high latitudes) are
rare, and may be attributable to the upward motion of deep cold water masses that would allow movement of
individuals to much less deep water habitats.
In general, most species of Parapagurus are broadly distributed in one or two ocean basins (Figs 48-50). Of the
14 species that occur in the Pacific and Indian Oceans, three have been found in the New Caledonia region,
P. latimanus , P. richeri sp. nov., and P. furici sp. nov. (Fig. 48); two are primarily distributed in the western
Indian Ocean and range into the southeastern Atlantic, P. bouvieri and P. andreui , and two have so far been found
only in the Indian Ocean, P. saintlaurentae and P. stenorhinus (Fig. 49). The distribution of P. abyssorum is
particularly broad; it has been found abundantly in the North Atlantic (see LEMAITRE, 1989), and sparsely in the
western and eastern Pacific (Figs 49, 50).
Seven species of Parapagurus are distributed in the eastern Pacific, P. benedicti , P. microps , P. holthuisi ,
P. abyssorum , P. janetae sp. nov., P. foraminosus sp. nov., and P. wolffi sp. nov. (Fig. 50). Of these.
P. holthuisi has been discovered living in a seamount on the central Pacific (Fig. 49), and P. abyssorum is known
from a single locality in the southeastern Pacific (Fig. 50). A species found abundantly in the region is
P. benedicti , known so far exclusively from the northeastern Pacific (Alaska to Baja California, Mexico), but
probably ranges also to the northwestern Pacific. The distribution of P. microps appears to include only the
tropical and subtropical latitudes of the eastern Pacific.
ACKNOWLEDGMENTS
This study was possible thanks to various expeditions sponsored by the French government, and the outstand¬
ing collecting efforts of French scientists, particularly Bertrand Richer de FORGES. The Parapagurus specimens
obtained during these expeditions proved indispensable in allowing me to complete this review.
A special debt of gratitude is owed to Alain CROSNIER for his unselfish commitment to the study of the
New Caledonia crustacean fauna. His help, hospitality, encouragement, and editorial advice while working with
him at the Museum national d'Histoire naturelle, are gratefully acknowledged.
At the Paris museum, I am also thankful to Nguyen NGOC-Ho, for her patience and care in processing
numerous specimens; and Daniele GUINOT, for placing museum type specimens and other comparative material
at my disposal.
Thanks are extended to many other colleagues and staff from other museums and institutions throughout the
world, all of which were instrumental in providing numerous specimens and information. These include: Keiji
Baba (Kumamoto University, Japan); Penny B. Berents, Jim Lowry, and Shane AHYONG (Australian
Museum, Sydney); Tin-Yam Chan (National Taiwan Ocean University, Keelung); Peter J. F. Davie (Queensland
Museum, Brisbane); George E. Davis and Todd Zimmerman (Natural History Museum of Los Angeles County);
Ardis B. JOHNSTON (Museum of Comparative Zoology, Harvard University); Tomoyuki KOMAI (Natural History
Museum and Institute, Chiba)); Miranda Lowe (The Natural History Museum, London); Enrique MACPHERSON
(Instituto de Ciencias del Mar, Barcelona); Patsy A. McLaughlin (Western Washington University); Peter K. L.
Ng (National University of Singapore); Dirk PLATVOET (Zoologisch Museum, Amsterdam); Gary C. B. POORE
(National Museum of Victoria, Melbourne); Michael Turkay and Andreas Allspach (Senckenberg Museum,
Frankfurt); Michelle van der Merwe (South African Museum); Waldo Wakefield (formerly at Scripps
Institution of Oceanography); Rick Webber (Museum of New Zealand Te Papa Tongarewa); Torbcn Wolff
(Zoological Museum, Kobenhavns University); Dmitry Zhadan (Zoological Museum, Moscow State
University); and Wolfgang ZEIDLER (South Australian Museum, Adelaide).
Reviews of the manuscript by Patsy A. McLaughlin and Alan Harvey were very useful. The help of Marco
Retamal (Universidad de Concepcion, Chile) in obtaining some references is gratefully acknowledged. Last but
not least, I thank Rose A. Gulledge for her invaluable help in preparing the maps and graphs.
Source :
PARAPAGURIDAE FROM THE PACIFIC AND INDIAN OCEANS
375
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ESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS [
Crustacea Decapoda: Albuneidae and Hippidae
of the tropical Indo-West Pacific region
Christopher B. BOYKO
Department of Invertebrates
American Museum of Natural History
Central Park West at 79th St.
New York, NY 10024, USA
Department of Biological Sciences
University of Rhode Island
Kingston. RI 02881, USA
&
Alan W. HARVEY
Department of Invertebrates
American Museum of Natural History
Central Park West at 79th St.
New York, NY 10024, USA
(Current address: Department of Biology
Georgia Southern University
Statesboro, GA, 30460. USA)
ABSTRACT
Based primarily on samples collected during French expeditions to New Caledonia and nearby regions, two new
species of the sand crab family Albuneidae Stimpson, 1858 are described from the tropical Indo-West Pacific Ocean:
Albunea hohhuisi , from Tanzania, Madagascar, and Indonesia, and Austrolepidopa Caledonia, from New Caledonia.
Two closely related, and often synonymized, species of Albunea: A. microps Miers, 1878 and A. elioti Benedict, 1904 are
found to be distinct. Several important diagnostic morphological features, not previously described in the Albuneidae. are
discussed. In addition, we provide diagnoses for three Indo-West Pacific species of mole crabs in the family Hippidae
Latreille, 1825, including the very similar Hippa pacifica Dana. 1852 and H. celaeno (de Man. 1896). An annotated list of
the 37 species of Hippoidea reported from the Indo-West Pacific region is provided, along with a diagnostic key to
these species.
RESUME
Crustacea Decapoda: Albuneidae et Hippidae de l'lndo-Ouest Pacifique tropical.
En se basant, au depart, sur les r£coltes faites durant diverses expeditions fran^aises au large de la Nouvelle-Caledonie
et dans des regions voisines, deux nouvelles especes appartenant k la famille des Albuneidae Stimpson, 1858 sont
BOYKO, C. B. & Harvey, A. W., 1999. — Crustacea Decapoda: Albuneidae and Hippidae of the tropical Indo-West
Pacific region. In: A. CROSNIER (ed.). Resultats des Campagnes MUSORSTOM, Volume 20. Memoires du Museum national
d'Histoire naturelle , 180: 379-406. Paris ISBN 2-85653-520-8.
Source : MNHN, Paris
380
C. B. BOYKO & A. VV. HARVEY
ctecrites do I'lndo-Ouest Pacifique: Albunea holthuisi sp. nov., connue de la Tanzania, de Madagascar et de l’lndonesie et
Austrolepidopci Caledonia sp. nov.. connue de la Nouvelle-Caledonie. Le reexamen de deux especes d'Albunea. proches
et souvent mises en synonymic. A. microps Miers, 1878 et A. elioti Benedict. 1904 a montr6 qu'elles sont distinctes.
Plusieurs caracteres morphologiques distinctifs, qui n’avaient pas encore 6te decrits chez les Albuneidae, sont
discutes. Des diagnoses pour trois espcces de la famille des Hippidae Latreille, 1825, comprenant les especes Hippo
pacifica Dana, 1852 et H. celaeno (de Man, 1896), tres proches I’une de 1’autre. sont publiees. Enfin. une liste commentee
des 37 especes d’Hippoidea signalees de I'lndo-Ouest Pacifique est proposee, accompagnce dune cle d'identi fication de
ces especes.
INTRODUCTION
Crustaceans in the anomuran superfamily Hippoidea Latreille, 1825 are specialized burrowing crabs that live in
sandy habitats in shallow waters, predominantly in the tropics. Historically, there has been little systematic work
in either hippoid family (i.e., the Albuneidae and the Hippidae) on a regional or worldwide basis. In the
Albuneidae, there has been no comprehensive study of any genus outside of Schmitt's (1942) review of
Blepharipoda Randall. 1839 and EFFORD's (1971) revision of Lepidopa Stimpson. 1858. Serene's (1979) review
of Pamlbunea Serene, 1979 was by his own admission incomplete, as he did not examine specimens of all the taxa
during the study. For Hippa Fabricius. 1787, the largest genus in the Hippidae, there have been regional reviews
(Efford, 1972; Haig, 1974b). A list of Hippa species has been prepared (Haig et al. , 1986). but the genus has
not been reviewed comprehensively since the work of DE Man (1896; 1898). EFFORD (1976) summarized
worldwide distributional data for the genus Emerita Scopoli, 1777. None of the other seven genera in these
families have been well studied systematically.
Even less is known about the phylogenetic relationships among the species and genera in the Hippoidea.
EFFORD (1969) proposed a phylogeny for the albuneid genera and later (1971) presented a preliminary tree for the
genus Lepidopa , but these were based on relatively few morphological characters. Tam et al. (1996) presented a
molecular phylogeny for American species of Emerita. We plan to generate a comprehensive phylogeny for the
Hippoidea, but this first requires better understanding of the taxonomy for the species involved.
The Hippoidea is represented in the tropical Indo-West Pacific region by 37 described species. In the present
paper we summarize the current state of taxonomic knowledge regarding the tropical Indo-West Pacific species of
Hippoidea, describe two new species of albuneids, present new information on species ranges and biology, and
discuss the morphological differences that allow the separation of two closely related species of Hippa. In addition,
we provide a dichotomous key to facilitate the identification of Indo-West Pacific hippoids.
MATERIALS AND METHODS
Since 1976, ORSTOM (Institut frani^ais de Recherche scientifique pour Ic Developpement en Cooperation) and
the Museum national d’Histoire naturelle, Paris, have participated in a joint study of the bathyal fauna of the
southwest Indo-Pacific. Several cruises have been undertaken but they have been mostly interested in the deep¬
water environment and few have collected Hippoidea which live in shallow water, rarely down to 225 m.
The cruises that have collected Hippoidea are:
- Corindon 2 in the Makassar Strait. Indonesia, from 29 October to 12 November 1980 (see Moosa, 1984).
- Smib 6, which from 28 February to 12 March 1990 explored the "grand passage" north of New Caledonia
(see Richer de Forges, 1993).
In addition, collections were made by the MONTROUZIER EXPEDITION, which from 23 August to 5 November
1993 explored the lagoon and reef environments of two sites on North Caledonia: Touho on the east coast and
Koumac on the west coast (see Bouchet, 1994).
Other collections made by ORSTOM scientists in New Caledonia and Madagascar were also examined.
All this material is deposited at the Museum national d'Histoire naturelle (MNHN).
Source :
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
381
Additionally, wc examined material from the following institutions: the Academy of Natural Sciences of
Philadelphia, Pennsylvania, USA (ANSP), the American Museum of Natural History, New York, USA (AMNH),
the British Museum (Natural History) (now The Natural History Museum), London, England (BMNH), the
Museum of Comparative Zoology, Massachusetts, USA (MCZ), the Musee Royal de PAfrique Centrale, Tervuren.
Belgium (MRAC), the Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van Natuurlijke Historie),
Leiden, The Netherlands (RMNH), the University Museum of Zoology, Cambridge, England (UMZC), the
National Museum of Natural History, Washington, D.C., USA (USNM), the Western Australian Museum, Perth,
Australia (WAM), and the Zoological Museum, University of Copenhagen, Denmark (ZMUC). The type
specimens are deposited as specified in the species entries.
In the lists of material examined, measurements are given for carapace length (CL), as measured from the
midlinc of the anterior margin (including rostrum, if any) to the midline of the posterior concavity. The names of
cruises are in all capital letters, whereas the names of vessels are in both italics and quotation marks. In the lists of
synonyms, asterisks indicate those publications that refer to materials re-examined by us during the present study.
For illustrations, we first captured specimen images on a Macintosh™ computer with a digital camera
connected to a Wild MB dissecting microscope. These images were then prepared for publication using the
programs Adobe Photoshop™ and Adobe Illustrator™. We attempted to accurately record the position and size of
setae in these drawings but, for clarity of presentation, excluded plumules of plumose setae.
SYSTEMATIC ACCOUNT
HIPPOIDEA Latreille, 1825
ALBUNEIDAE Stimpson, 1858
Remarks. — During the course of this study, we encountered several diagnostic morphological features that
have not been described previously for albuncids. For example, the carapace (Fig. 1) bears a broad mat of very
Setal field
Fig. 1. — Diagrammatic albuneid carapace,
based on Albunea microps , showing setal
field behind front, and 1 1 setose carapace
grooves discussed in species accounts.
short, dense, simple setae just behind the front. This mat,
hereafter called the setal field, varies in shape and extent across
genera and species, but appears to be relatively invariant within
species.
The carapace of albuneids typically possesses numerous
transverse, setose grooves. Although carapace grooves (CG) have
been scarcely mentioned by previous authors, we have identified
at least eleven major grooves (numbered 1-11 in Fig. 1) the
homology of which can be recognized across albuneid genera.
Variability in the presence and the degree of fragmentation of
specific grooves, in the anterior-posterior displacement of
individual fragments, and in the texture of the grooves (e.g..
smooth, crcnulate) tends to be conserved within species, and thus
carapace grooves are useful in recognizing species.
The median element of CGI forms the posterior margin of the
setal field and also of the front. In some genera, the curved lateral
elements of CGI are often displaced posteriorly (as in Fig. 1).
The metagastric region contains the short, anterior CG2 and the
longer, posterior CG3. CG4 spans the width of the carapace and
marks the border of the metagastric and mesogastric regions. CG5
is a fairly short groove that occurs medially in the mesogastric
region. CG6 corresponds to the cervical groove in other
Source :
382
C. B. BOYKO & A. W. HARVEY
Anomura. CG7 is usually divided into 2 well-separated lateral fragments, but in some genera (e.g.. Austrolepidopa
Efford & Haig, 1968 and Lepidopa ), CG7 merges medially with CG6. CG8-1 1 are relatively short medial grooves
arranged anteriorly to posteriorly in the cardiac region.
In several genera of albuncids, nearly transparent and presumably decalcified “windows” can be found on various
locations on the crab, including lateral and mesial surfaces of the second through fourth pereopods and on the dorsal
surface of the first and second abdominal somites. Such decalcified windows have not been reported previously for
any albuneid. Similar “windows” occur on the mesial surface of the merus in several genera in the anomuran
family Porcellanidae Haworth, 1825, and on the mesial surface of the coxa of the first pereopods in the hippid
genus Emerita , but are lacking in Hippa and Mastigochirus Stimpson, 1858 (Harvey, in press).
As is typical in decapod crustaceans, albuneid females have gonopores on the coxae of the third pereopods,
whereas males have gonopores on the coxae of the fifth pereopods. However, in some albuneid genera (e.g.,
Austrolepidopci and Lepidopa), males also have a small pore on the coxa of the third pereopod (hereafter referred to
as the "male pore"), in a position analogous to that of the female gonopore. The precise nature and function of this
male pore is unknown.
In albuneids, females have well developed uniramous pleopods on abdominal somites II- V, whereas pleopods
are thought to be completely lacking in males (e.g., MARTIN & ABELE, 1986). However, we have found, in
specimens with well developed pores on the fifth pereopods, rudimentary to small pleopods on abdominal somites
II- V in several albuneid genera (e.g., Austrolepidopa and Lepidopa). The existence of the “male pore” and of small
pleopods in males were not previously recognized in albuneids, which we suspect has led to the incorrect sexing of
crabs and contributed to a probably erroneous impression that males are exceedingly rare or even absent in many
species of albuneids. For example, the holotype of Austrolepidopa trigonops Efford & Haig, 1968 was described as
an immature female with incompletely developed pleopods and gonopores, but we have examined this specimen
and determined that it is in fact a male. In those species in which the male pore occurs, it is always smaller than
gonopores of same-sized females; likewise, the pleopods of females are always much more developed than those of
males. Males are most reliably recognized by the presence of a gonopore on the fifth pereopod and the rudimentary
degree of development of the pleopods. In small specimens, however, the presence or absence of the male gonopore
is a more reliable indicator of sex than is pleopod development because both males and females have small pleopod
buds as juveniles.
The presence in albuneids of ocular acicles has been controversial. Typically considered to be restricted to
hermit crabs, ocular acicles were first reported in albuneids by McLaughlin (1980), although she subsequently
Fig. 2. Diagrammatic anomuran eyes, showing relationship between proximal ocular peduncle segments (PS), distal
ocular peduncle segments (DS). ocular acicles (AC) and ocular plates (OP): A, Sympagurus dimorphus (Studer, 1883),
a typical hermit crab; B, Albunea sp. undescribed, a typical albuneid.
Source :
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
383
concluded that the structures on albuneids were "not acicles, but rather, calcified portions of the hippoid ocular
plate" (McLaughlin, 1983: 615). Nonetheless, Martin and Abele (1986: 578) scored the Albuneidae as having
ocular acicles, remarking that they were "unsure about the differences between small ocular acicles and pieces of the
ocular plate". The distinction is straightforward. In decapod crustaceans, the ocular peduncle typically consists of
a proximal segment and a distal segment; in rare cases, one or both of these segments may be secondarily
subdivided (Powar, 1969). The ocular plate is the medial calcified acron to which the ocular peduncles
are attached. Based on muscle site attachments, Powar (1969) clearly established the homology of the proximal
and distal ocular segments, as well as the ocular plate, across the Decapoda. True ocular acicles are not part of
the ocular plate, but rather are spinous or platelike anterodorsal extensions of the proximal segment of the ocular
peduncle (Fig. 2).
In some taxa (c.g., the diogenid hermit crab genus Dardanus Paulson, 1875, and many albuneids), the ocular
plate is exposed, that is, not covered by the anterior carapace. In some albuneids, this exposed ocular plate is also
deeply indented medially, perhaps consistent with the hypothesis that the ocular plate represents the fusion of
primitively paired basal eyestalk segments (Powar, 1969). This deeply indented ocular plate gives an impression
superficially similar, but clearly not homologous, to the ocular acicles on the proximal ocular segment of hermit
crabs, which we presume to be the source of confusion among earlier workers.
As observed by previous workers, the third maxilliped
contains a number of morphological characters for
distinguishing among albuneid genera and species. In most
anomurans, the basis bears a medial row of teeth, termed
the crista dentata. Among the albuneids, only two aberrant
genera, Blepharipoda and Lophomastix Benedict. 1904, arc
thought to possess a crista dentata (Martin & Abele,
1986; MCLAUGHLIN & LEMAITRE. 1997). However, we
have observed a very reduced crista dentata on several
species of Albunea. Other than the third maxilliped. the
mouthparts of albuneids tend to lack distinguishing
features and so are not described herein.
The shape of the dactylus of the pereopods, particularly
the third pereopod, has been used widely to distinguish
among species of albuneids. To describe the complex
shape of this segment, we use several shorthand terms to refer to important landmarks (Fig. 3). The "base" of the
dactylus is the ventroproximal angle, and the "heel" corresponds to the dorsoproximal angle, which is often
strongly produced. The dorsal margin is almost always concave, sometimes smoothly so; in most species,
however, the dorsal margin forms a distinct angular incision proximally, the apex of which is herein referred to as
an "indent." The dactylus terminates in a "tip," which in hippoids is at least somewhat rounded and lacking in a
corneous nail.
Fig. 3. — Pereopod II dactyl, lateral view, of Albunea
holthuisi sp. nov., showing terms used in species
accounts for landmarks on pereopod dactyli.
Genus ALBUNEA Weber, 1795
Albunea microps Miers, 1878
Figs 1, 4
Albunea microps White, 1847: 129 (nomen nudum)*. — MIERS, 1878: 328-329. pi. 5, figs 12-13*. — HENDERSON, 1888:
40*. — Ortmann, 1896: 224-225 (list). — Borradaile, 1904: 751*. — Gordon. 1938: 187. fig. 3c*. — Serene &
Umali, 1965: 95-97. pi. 6, figs 1-6. text fig. 12c. — Thomassin, 1969: 140-143 (part), text figs 2. 3b. — Miyake.
1978: 154-155, fig. 60b.
Albunea [sp.] Gordon, 1938: 189-190, fig. Id*.
Not Albunea microps - THOMASSIN, 1969: 140-143 (part), pi. 2, figs 1-9 (= Albunea elioti Benedict. 1904).
Source :
384
C. B. BOYKO & A. W. HARVEY
Material EXAMINED. — Philippines. Sulu Archipelago. "Sooloo Islands" |- Sulu Archipelago], coll, unknown:
1 8 11.3 mm (BMNH 1937.6.7.3).
"Challenger": stn 212, 06°54’N. 122°18,E, 3-6 m, 30.01.1875 : 1 6 9.5 mm (ZMUC 2715)
Indonesia. "Gloria Maris": stn 549, east coast of Rouw Island. Aoen Group, west Geelvinck (- Cenderawasih)
Bay. "New Guinea" (= Irian Jaya), coll. A. J. Ostheimer, 23.02.1956: I 6 1 1.2 mm (RMNH 23641).
CORINDON 2: stn DR 293. 02°37.7’S, I17°49.4'E, Sulawesi. Makassar Strait. 45 m. 11.11.1980: 1 ov. 2 12.1 mm
Andaman Islands. Port Blair, coll, unknown: 2 8 6.2-6. 7 mm (BMNH 1956.1.14.20).
Maidive Islands. Mahlosmadulu Atoll. 37 m, coll. J. S. Gardiner: 2 juv. 9 4.6-4.7 mm (UMZC).
Oman Muscat, 12-27 m, coll, unknown: 1 8 7.8 mm (BMNH 1901.4.20.10).
Seychelles. Mahe, coll. Mission Zoologique MRAC-ULB. July-Aug. 1972: 2 9 6.0-10.2 mm (MRAC 53.604).
Tanzania. Stn 650, dredged in shell and sponge, about 1.5 mi (= 2.5 km) east of Puopu Island, northwest Zanzibar.
13-16 m coll. A. J. OSTHEIMER III, 20.02.1957: 1 9 12.8 mm (ANSP CA4647). — Dredged in muddy sand, around Bawi
and Change Islands, off Zanzibar City, Zanzibar, 11-29 m, coll. A. J. Ostheimer III. 27.02.1957: 1 9 10.9 mm (ANSP
CA4646).
Fig. 4. — Albunea microps Miers. 1878, ov. 9 12.1 mm (MNHN-Hi 201) (A. B, D); 8 , holotype, 11.3 mm (BMNH
1937.6.7.3) (C). A, carapace, dorsal view; B. eyes, dorsal view; C, telson of male, dorsal view; D, telson of female,
dorsal view. Scale = 0.75 mm (B), 1.0 mm (C), 2.0 mm (D), and 4.0 mm (A).
Source : MNHN. Paris
ALBUNE1DAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
385
Type. — Holotype : <5 11.3 mm, “Sooloo Islands” [= Sulu Archipelago, Philippines] (BMNH 1937.6.7.3).
DIAGNOSIS. — Carapace (Fig. 4A) slightly longer than wide, covered with strongly setose grooves. Anterior
margin with about 9 teeth on either side of ocular sinus. Setal field with thick anterolateral projections and truncate
anteromedial margin. CGI with lateral elements separated, displaced posteriorly from median element; CG4 frag¬
mented, median element displaced anteriorly; CG5 entire; CG6 and CG7 united; CG 1 1 present. Rostrum present,
exceeding posterior margin of ocular plate by about half length of ocular plate. Ocular plate subquadrate (Fig. 4B).
Ocular peduncles (Fig. 4B) dorsovcntrally flattened and oblong in shape, rounded at tip. approximated along distal
2/3 of mesial margin; lateral margin convex except at concave tip; mesial margin convex. Cornea on lateral
margin at tip. Dactyli of pereopods II and III with heels smoothly rounded. Dactylus of percopod IV sinuous from
base to tip; indent slight. Telson of male (Fig. 4C) broadly triangular, inflated dorsally, broadly rounded at tip,
strongly calcified proximally; large decalcified windows on either side of thin medial calcified strip; long thin setae
medially and along anterior margin of windows. Telson of female (Fig. 4D) flattened, ovate, longitudinal row of
short, thin setae medially.
Fig. 5. — Albunea elioti Benedict, 1904, 9, holotype, 16.2 mm (USNM 26169) (A, B. E); 6, 16.5 mm (USNM 281472)
(C, D). A. carapace, dorsal view; B, eyes, dorsal view; C, telson of male, dorsal view; D, telson of male, lateral view;
E. telson of female, dorsal view. Scale = 0.75 mm (B), 2.0 mm (E). 3.0 mm (C. D). and 4.0 mm (A).
386
C. B. BOYKO & A. W. HARVEY
DISTRIBUTION. — Widely distributed in the Indo-West Pacific from the east coast of Africa to Indonesia and the
Philippines and north to southern Japan; 3-45 m depth.
REMARKS. — Albunea microps has been confused with A. elioti Benedict, 1904, previously known only from
the holotype female, from Samoa (USNM 26169, 16.2 mm ; Fig. 5A, B, E). Gordon (1938) first suggested that
A. elioti was perhaps the same as A. microps , and other authors (SERENE & Umali, 1965; THOMASSIN, 1969;
Miyake, 1978) have treated her suggestion as definitive. From our examinations of the holotype of A. elioti and
two additional male specimens we discovered from Tonga (USNM 281472, 16.5 mm ; Fig. 5C, D) and
Madagascar (MNHN-Hi-88, 10.4 mm), we have determined that this species is distinct from A. microps. The
shape of both the telson of the male and the eyes can be used to tell the two species apart. The telson of the male
of A. microps (Fig. 4C) is heavily calcified and somewhat inflated proximally, but partially decalcified distally and
narrowing to a produced tip. An oblique row of long setae is present just proximal to the demarcation line between
the calcified and decalcified regions on each side of the median line. In A. elioti. the telson of the male is narrowly
triangular (Fig. 5C), fully calcified, with short thick setae on a strong medial ridge (Fig. 5D), and lacks the oblique
row of setae. The mesial margins of the ocular peduncles are strongly convex in A. elioti (Fig. 5B) but only
slightly so in A. microps (Fig. 4B). In addition, the cornea of A. elioti is more posterolaterally displaced from the
tip of the ocular peduncle.
At least some of the Madagascar specimens described by THOMASSIN (1969) as Albunea microps are
clearly A. elioti (e.g., his pi. 2, figs 1-9). Unfortunately, Thomassin’s material cannot be located in MNHN
(NGOC-HO, personal commun.), and we have encountered no specimens of A. microps from Madagascar.
However, because we have recorded A. microps from Zanzibar, we do suspect that the species is also present in
Madagascar.
The specimen of A. microps collected during the CORINDON 2 cruise is the first report of an ovigerous female
for this species.
Albunea holthuisi sp. nov.
Figs 6-7
? Albunea steinitzi - THOMASSIN, 1969: 143-146, pi. 3, figs 1-8, text fig. 3 (not A. steinitzi Holthuis, 1958 - see remarks).
MATERIAL EXAMINED. — Madagascar. NW coast: 13°37.7’S, 47°49.6'E, near Nosy Be, 25 m, coll, unknown:
1 6 8.1 mm (MNHN-Hi 202). — Nosy Be, Andilana beach, intertidal, coll. A. Crosnier, Sept. 1959: 1 9 8.1 mm
(MNHN-Hi 203). — 13°38.3’S. 42°49.6'E, near Nosy Be. 34 m. coll. A. Crosnier :U8 mm (MNHN-Hi 204).
Tanzania. Boat unknown: stn 651, dredged grass and shell, 1.5 mi WSW Ros Nungwa, north Zanzibar. 15 m, coll.
A. J. Ostheimer III. 20.02.1957: 1 6 7.7 mm (ANSP CA4644). — Same data: 1 9 6.7 mm (ANSP CA4645).
Indonesia. " Gloria Maris": stn 522, west side of Samberbaban Bay, "Japen” (= Yapen) Island. "West New Guinea"
(= Irian Jaya), coll. C. T. Abbot . 14.02.1956: 1 6 1 1.3 mm (RMNH 23703).
Corindon 2: stn B 255, 01°56.5'S, 1 19°17.3’E, Sulawesi, Makassar Strait, 13 m, 6.11.1980: 1 9 7.1 mm (MNHN-Hi
205). — Stn B 256. 01°56.5’S, 119°17.2'E, Sulawesi, Makassar Strait, 24 m, 6.11.1980: 1 6 4.9 mm ; 1 9 5.4 mm
(MNHN-Hi 206).
TYPES. — Holotype: 6 8.1 mm. Madagascar, 13°37.7'S. 47°49.6'E, near Nosy Be, 25 m (MNHN-Hi 202).
Allotype: 9 8.1 mm, Madagascar, Nosy Be, Andilana beach, intertidal (MNHN-Hi 203). Paratypes: All the other
specimens mentioned above, except MNHN-Hi 206 (see remarks).
Etymology. — The specific name is given in recognition of Dr Lipke HOLTHUIS. Nationaal Natuurhistorisch
Museum, who has contributed immeasurably to the study of Crustacea, including the Hippoidea.
Diagnosis. — Carapace slightly longer than wide, covered with lightly setose grooves. Anterior margin with
about 9 teeth on either side of ocular sinus. Setal field with narrow lateral elements and concave anterior margin.
CGI with separate posterior lateral elements; CG4 with short element on either side of median with missing
elements at midline and between median and laterals; CG5 present only as short lateral elements; CG6 and CG7
separate; CG8 complete; CGI 1 present. Rostrum present, not reaching posterior margin of ocular plate. Ocular
Source : MNHN, Paris
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
387
plate triangular. Ocular peduncles dorsoventrally flattened and oblong in shape, tapering at tip, approximated along
distal 2/3 of mesial margin; lateral margin convex except slightly concave at tip; mesial margin convex. Cornea at
tip. Dactylus of pereopod II with heel slightly produced, low and rounded. Dactylus of pereopod III with heel
slightly projecting, acute. Dactylus of pereopod IV sinuous from base to tip, with slight indent. Telson of male
spatulate; tip broadly truncate; dorsal surface with elevated median longitudinal ridge bearing short thick setae
proximally and long thick setae distally. Telson of female flattened and spatulate, longitudinal row of short, thin
setae medially.
Fig. 6. — Albunea holthuisi sp. nov., 6, holotype, 8.1 mm (MNHN-Hi 202) (A, B); 6, paratype. 7.7 mm (ANSP
CA4644) (C); 6 , paratype, 8.0 mm (MNHN-Hi 204) (D, E). A, carapace, dorsal view; B, eyes, dorsal view; C, left
antennule, lateral view; D, left antenna, lateral view; E, left third maxilliped. lateral view. Scale = 0.75 mm (B) and
2.0 mm (A, C-E).
Description. — Carapace (Fig. 6A) slightly wider than long. Anterior margin concave on either side of ocular
sinus, becoming convex laterally; 8-11 large spines on concave region; ventral row of long plumose setae
submarginally. Rostrum a small acute tooth, not reaching to proximal margin of ocular plate. Ocular sinus
smoothly concave and unarmed. Frontal region smooth; setal field broad posteriorly, narrowing anteriorly, with
narrow lateral elements and concave anterior margin. CGI parallel to anterior margin of carapace, sinuous, slightly
crenulate, divided into medial fragment and curved lateral elements that are displaced posteriorly. Mesogastric
region smooth; CG2 short; CG3 broken into 6 short elements approximately equally spaced between posterior
388
C. B. BOYKO & A. W. HARVEY
elements of CGI; CG4 fragmented into four elements with gap at midline and between median and lateral
elements. Hepatic region smooth, with short setose groove at median of lateral margin. Epibranchial region
roughly triangular, smooth; posterolateral margin with short row of setae. Metagastric region smooth; CG5
present only as short lateral elements directly posterior to median elements of CG4; CG6 slightly crcnulate.
strongly concave medially and sloping out to convex lateral thirds; CG7 transverse and separate from CG6. Cardiac
region smooth; CG8 uninterrupted; CG9 present only as lateral short lines; CG10 present in two fragments,
separated by length of single fragment; CGI 1 present. Branchial region with numerous short, transverse rows of
setae. Posterior margin deeply and evenly convex, with submarginal groove interrupted medially. B ranch iostegite
with short anterior submarginal spine; anterior region with scattered short transverse lines ventral to linea
anomurica, with many short rows of setae and covered with long plumose setae ventrally; posterior region
membranous with numerous, irregular fragments, and covered with long plumose setae.
Ocular plate (Fig. 6B) triangular, with shallow median indentation. Ocular peduncle (Fig. 6B) elongate, with
distally convex lateral margins, tapering to rounded distolateral cornea; mesial margins approximated along entire
length; mesial and ventral margins of segment with sparse row of long plumose setae; tuft of plumose setae at
proximal lateral ventral angle; ventral surface with oblique row of long plumose setae from proximal lateral angle
almost to distal mesial margin.
Antennule (Fig. 6C) with segment III narrow proximally, expanding distally to twice proximal width; plumose
setae on dorsal and ventral margins; dorsal exopod flagellum with 76-104 segments, long plumose setae on dorsal
and ventral margins; ventral endopod flagellum short, with 2-3 segments, plumose setae on dorsal and ventral
margins. Segment II medially inflated from dorsal view, plumose setae on dorsal and ventral margins and scattered
on lateral surface. Segment I wider than long, unarmed; lateral surface with long plumose setae dorsally and on
dorsal and ventral margins.
Antenna (Fig. 6D) with segment V about 3 times longer than wide, long plumose setae on dorsal and ventral
margins; flagellum 7-segmented, long plumose setae on dorsal, ventral, and distal margins. Segment IV expanded
distally, long plumose setae on dorsal, ventral, and distal margins, and simple setae on dorsolateral margin.
Segment III with long plumose setae on ventral margin. Segment II short, widening distally, plumose setae on
margins; antennal acicle long, thin, exceeding base of segment V by about 1/2 length of segment V, long plumose
setae on dorsal margin. Segment I rounded proximally, flattened ventromesially, long plumose setae on margins;
lateral surface with acute spine dorsally, with low, semicircular dorsolateral lobe ventrodistal to spine.
Maxilliped III (Fig. 6E) with dactylus rounded at tip. long plumose setae on margins and lateral surface.
Propodus with longitudinal median row of plumose setae on lateral surface, margins with plumose setae. Carpus
slightly produced onto propodus, lateral surface with row of plumose setae ventromedially; plumose setae on
margins. Merus unarmed, plumose setae on margins. Basis incompletely fused with ischium; weak crista dentata
of 2-3 teeth. Exopod 2-segmented, proximal segment small, distal segment styliform, tapering, approximately 1/3
length of merus, plumose setae on margins; flagellum absent.
Pereopod I subchelate. Dactylus (Fig. 7A) curved and tapering; lateral and mesial surfaces smooth; dorsal
margin with long plumose and short simple setae, short simple setae on ventral margin. Propodus (Fig. 7A)
lateral surface with numerous short, transverse rows of setose rugae; dorsal margin unarmed; ventral margin
distally produced into acute spine; cutting edge lacking teeth, lined with long plumose setae; lateral, mesial and
ventral margins with long setae. Carpus (Fig. 7A) with dorsodistal angle produced into small corneous spine;
dorsal and distal margins with long plumose setae; lateral surface with distal rugose area, few transverse setose
ridges on distal 2/3 of surface; mesial surface smooth with scattered rows of long plumose setae, margins with
long plumose setae. Merus unarmed; lateral surface with scattered transverse rows of long plumose setae, margins
with long plumose setae; mesial surface with few short rows of setae. Basi-ischium incompletely fused, unarmed.
Coxa unarmed.
Pereopods II-IV with dactyli laterally compressed and dorso ventrally expanded.
Pereopod II dactylus (Fig. 7B) smooth; base to heel concave, heel with smoothly rounded low spur, heel to tip
acutely indented and narrow, tip acute, tip to base broadly convex; lateral surface smooth, several small tufts of
short setae in roughly straight line across medioproximal surface, several widely spaced submarginal tufts of short
setae dorsodistally; mesial surface smooth, ventral margin with long plumose setae, dorsal margin with short
Source :
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO- WEST PACIFIC REGION
389
plumose setae, patch of long plumose setae at base (not illustrated). Propodus dorsal surface smooth, ventral
margin inflated and rounded; oblique row of long plumose setae on distal margin of lateral surface; distal and
ventral margins with long plumose setae; dorsolateral surface a narrow, oblique, flattened shelf, short setae on
dorsal margin and long plumose setae on ventral margin; mesial surface with elevated, curved, setose ridge from
ventral junction with dactylus almost to ventral proximal junction with carpus. Carpus slightly produced, gently
rounded; lateral surface nearly smooth, with irregular, broken row of rugae and submarginal elevated ridge
ventrally, rugae and ridge with long plumose setae, margins with long plumose setae; mesial surface smooth, long
plumose setae on margins and in scattered patches on surface. Merus with medial decalcified area on lateral surface,
Fig. 7. — Albunea holthuisi sp. nov., 6 , holotype, 8.1 mm (MNHN-Hi 201) (A-G); $, allotype. 8.1 mm, MNHN-Hi 203
(H). A, left chela, lateral view; B, left pereopod II dactyl, lateral view; C. left pereopod III dactyl, lateral view; I), left
pereopod IV dactyl, lateral view; E, abdomen, dorsal view; F. telson of male, dorsal view; G, telson of male, lateral
view; H, telson of female, dorsal view. Scale = 1.0 mm (H), 1.5 mm (F. G), and 2.0 mm (A-E).
390
C. B. BOYKO & A. W. HARVEY
long plumose setae on lateral margins; mesial surface nearly smooth, with few setae. Basi-ischium incompletely
fused and unarmed. Coxa with 1 small spine on anterior margin.
Pereopod III dactylus (Fig. 1C) with base to heel concave, heel produced in short, acute spur, heel to indent
nearly straight, indent broadly concave, tip acute, tip to base smoothly convex to straight; lateral surface smooth,
dorsodistal margin with tufts of short setae, ventromesial margin with long plumose setae, dorsal margin with
short simple and plumose setae; mesial surface smooth, plumose setae proximally at junction with propodus.
Propodus weakly inflated; lateral surface smooth, long plumose setae distally, simple setae on margins, long
plumose setae on ventral margin, dorsolateral surface narrow, oblique, flattened; mesial surface with scattered long
setae on and near distal margin. Carpus produced dorsodistally, exceeding proximal margin of propodus by about
1/3 length of propodus, broadly rounded; dorsolateral margin unarmed; lateral surface slightly rugose dorsodistally,
many short and 2 longer rows of setae ventrally; mesial surface smooth, long plumose setae on margins and
scattered on surface. Merus smooth, dorsal and ventral margins unarmed, long plumose setae, distolateral margin
with long plumose setae; lateral surface with decalcified area anteriorly; mesial surface smooth. Basi-ischium
incompletely fused and unarmed. Coxa with 1 small spine on anterior margin. Female with large gonoporc on
median mesial surface of coxa, surrounded with short plumose setae; male without pore.
Pereopod IV dactylus (Fig. 7D) with base to tip convex to straight, tip acute, tip to base convex distally,
becoming broadly concave proximally; lateral surface smooth, ventral margin with long plumose setae, dorsal
margin with short simple setae; mesial surface with median decalcified area, demarcated ventrally by longitudinal
elevated ridge with row of long plumose setae, setose punctae ventral to decalcified window. Propodus expanded
dorsally and ventrally, ventral expansion exceeds ventral margin of dactylus, margin with long plumose setae,
dorsal expansion with row of long plumose setae medially; lateral and mesial surfaces smooth. Carpus not
produced dorsodistally; lateral and mesial surfaces smooth, dorsal margin with short simple and long plumose
setae, ventral margin with short plumose setae. Merus with lateral surface with scattered short transverse rows of
setae, dorsal and ventrodistal margins with long plumose setae, slightly rugose ventrodistally, with short setae;
mesial surface with large decalcified "window” proximoventrally. Basi-ischium incompletely fused and unarmed.
Coxa unarmed.
Pereopod V reduced, slender. Coxa of male with large mesioproximal gonoporc.
Abdomen (Fig. 7E) with somite I approximately as wide as long, widest posteriorly; dorsal surface with
anterior margin concave, posterior margin concave, with submarginal row of short setae, small transverse
decalcified submedial "windows". Somite II dorsal surface with submarginal transverse ridge anteriorly, tuft of
setae at posterolateral angle, extending onto pleura posteromesially, posterior margin with indistinct punctate
submarginal groove laterally; pleura expanded and directed slightly anteriorly, margins finely toothed, lateral
margins rounded, anterior and lateral margins with long plumose setae, posterior margin with short setae. Somite
III similar to somite II, but narrower, shorter, and lacking anterior submarginal ridge, small tuft of short thick
setae on posterolateral angle; pleura thinner and shorter than on somite II, directed anterolaterally, with setae as in
somite II, anterolateral angle acute, dorsal surface obliquely flattened anterolaterally. Somite IV similar to somite
III, but thinner and shorter; dorsal surface with thick setae posterolaterally; pleura thinner and shorter than on
somite III, directed laterally, dorsal surface obliquely flattened anterolaterally, margins with long plumose setae.
Somite V wider than somite IV, lateral margins with plumose setae; pleura absent. Somite VI subequal to somite
V in width but longer, dorsal surface with short oblique rows of setae laterad to midline anteriorly, lateral margins
with long plumose setae; pleura absent.
Females with uniramous, paired pleopods on somites II-V, males lacking pleopods.
Uropods lacking distinctive features.
Telson of male (Fig. 7F-G) spatulate, truncate posteriorly, weakly calcified, except for large triangular anterior
plate, margins with long plumose setae; median longitudinal groove very short, restricted to anterior of calcified
plate, calcified plate with thick elevated medial ridge (Fig. 7G) covered with short thick simple setae, small tuft of
setae at anterolateral margin. Telson of female (Fig. 7H) ovate, longer than wide, slightly truncate posteriorly,
dorsal surface smooth, with median longitudinal groove anteriorly, row of setose punctae lateral to midline from
median of longitudinal groove almost to posterior margin; margins with long plumose setae.
Distribution. — Known from Madagascar, Tanzania, and Irian Jaya and Makassar, Indonesia; 0-34 m depth.
Source :
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
391
REMARKS. — Albunea holthuisi is most similar to A. steinitzi Holthuis, 1958. Albunea holthuisi
differs from A. steinitzi in that CG8 is entire, and CGI 1 is present; additionally, the telson of the male has a
pronounced ridge, and has shorter medial setae. Albunea symmysta (Linnaeus, 1758) sensu stricto differs from both
of these species by having laterally inflated ocular peduncles, a strongly spurred heel on the dactylus of the third
pereopod, and a distinctively flat and broadly triangular male telson with only a few short setae on either side of
the midline.
No records in the literature can be ascribed definitively to A. holthuisi. However, given the excessively wide
range of variability that has been incorrectly attributed to A. symmysta, it is possible that some published records
assigned to "A. symmysta" may in fact be A. holthuisi. Many of the specimens published as "A. symmysta " that
we have been examined are actually referable to several other species of Albunea, both described and undescribed
(BOYKO, unpublished). Unfortunately, without direct examination of reported specimens, identities of most
published records of A. symmysta cannot be confirmed or corrected.
Such problems with prior identifications of Indo-West Pacific species of Albunea by various authors make
it difficult to accurately determine the geographic range of these species. We have examined the female holotypc
of Albunea steinitzi (RMNH 1 1847) from the Red Sea but have yet to see any specimens of A. steintizi from
Madagascar or nearby. Although SERENE and Umali (1965) identified A. steinitzi from the Philippines, we are
hesitant to accept those records as SERENE and UMALl's (1965) written description and illustrations are not
detailed enough to permit unambiguous identification of their specimens. We have also seen no specimens
referable to this species among a large series of Philippine Albunea examined. Haig (1974a) included A. steintizi
as part of the Western Australia anomuran fauna, but all specimens that we have seen from that region belong
to several undescribed species in this genus. At this time, A. steinitzi can only be reported with certainty from
the Red Sea (HOLTHUIS, 1958; Lewinsohn, 1969), Pakistan (Tirmizi, 1978), and probably from Oman
(Hogarth, 1988).
The specimens reported by Thomassin (1969) from Madagascar as Albunea steinitzi may belong to the present
species, but his illustration of the telson of the male in text fig. 4 suggests a rounded, produced tip, which does
not agree with the morphology of the telson of A. holthuisi, nor with that of A. steinitzi. It is possible that
THOMASSlN's figures are inaccurate, or that his specimens were of a separate, as yet unrecognized, species.
However, his specimens cannot be located in MNHN (NGOC Ho, personal commun.).
Because the two Indonesian specimens from CORINDON 2 stn B 256 (MNHN-Hi 206) are highly decalcified and
immature, we excluded them from the type series of Albunea holthuisi. although there is little doubt that they
belong to this taxon.
The Madagascar and Tanzanian female specimens of A. holthuisi have an elongate patch of striate substance on
the coxae of the third pereopods, superficially resembling a Botiyllus-iype tunicate. Isabella GORDON (unpublished
notes; BMNH) suggested that they might be “male secretions or spermatophores,** which was later confirmed by
SUBRAMONIAM (1984) in a detailed description of the structure of the spermatophore in A. symmysta. We have
found sperm ribbons protruding from the male gonopores in specimens of several species in the genus Albunea
(Boyko, unpublished). Subramoniam and Panneerselvam (1985) subsequently alluded to the “spermatophoric
ribbon attached onto the pleopodal regions” of females of A. symmysta, but did not provide further details
concerning their location or extent. We can now give the specific location of sperm ribbon deposition as the third
and sometimes fourth pereopod coxae of females in this species and others in the genus (Boyko, unpublished). As
yet we have not found similar sperm packets attached to the pereopods of females in any other albuneid genus.
Genus AUSTROLEPIDOPA Efford & Haig. 1968
Austrolepidopa Caledonia sp. nov.
Fig. 8-9
Material EXAMINED. — New Caledonia. Smib 6: stn DW 107. 19°07.6'S, 163°30.2'E. Grand Passage, 205 m,
2.03.1990: 1 6 9.2 mm (MNHN-Hi 207); 1 9 11.5 mm (MNHN-Hi 208). — Stn DW 109. I9°05.7'S. 163°29.7'E.
225 m, 2.03.1990: 1 9 8 mm (MNHN-Hi 209).
Source :
392
C. B. BOYKO & A. W. HARVEY
Types. — Holotype : 6 9.2 mm, New Caledonia, Smib 6, stn DW 107, 19°07.6'S, 163°30.2*E, Grand
Passage, 205 m (MNHN-Hi 207). Allotype : 9 1 1.5 mm, same data as the holotype (MNHN-Hi 208). Paratype :
9 8 mm, New Caledonia, SMIB 6, stn 109, 19°05.7’S, 163°29.7’E, 225 m (MNHN-Hi 209).
Etymology. — The specific name is given after the type locality. New Caledonia, and is treated as a noun in
apposition.
DIAGNOSIS. — Carapace longer than wide, covered with strongly setose grooves. Setal field with straight
anterior margin and narrow lateral elements directed posteriorly. CGI with contiguous posterior lateral elements;
CG4 nearly entire, with median section slightly displaced anteriorly; CG5 absent; CG6 merged with CG4 laterally
and with CG7 medially to form two separate hybrid grooves; CG8 complete; CGI 1 absent. Anterior margin of
carapace with finely toothed lobe lateral to ocular sinus. Rostrum absent, rostral region truncate anteriorly, finely
toothed. Ocular plate completely concealed dorsally by rostral region. Ocular peduncles dorsoventrally flattened and
triangular in shape, separated by slightly more than length of ocular peduncle, lateral margin slightly convex,
mesial margin slightly concave. Cornea not visible. Dactylus of pereopod II with heel slightly produced, low and
rounded. Dactylus of pereopod III with heel slightly projecting, rounded. Dactylus of pereopod IV deeply concave
from base to tip, with smoothly rounded indent. Telson of male and female similar; flattened and ovate, medially
with tufts of short, thin setae in paired longitudinal rows.
Fig. 8. — Austrolepidopa Caledonia, sp. nov., 2, allotype, 11.5 mm (MNHN-Hi 208) (A, B); 2, paratype, 8.0 mm
(MNHN-HI 209) (C-E). A. carapace, dorsal view; B, eyes, dorsal view; C, left antennule, lateral view; D, left antenna,
lateral view; E. left third maxilliped, lateral view. Scale = 1.0 mm (B). 1.5 mm (C-E), and 4.0 mm (A).
Source :
ALBUNEIDAE AND H1PPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
393
Description. — Carapace (Fig. 8A) aboul as wide as long. Anterior margin dentate between ocular peduncles,
submarginal ventral row of long plumose setae. Rostrum absent, rostral area truncate, overreaching base of ocular
peduncles and exceeded by anterolateral lobes. Ocular sinus concave, dentate. Anterolateral lobe broadly triangular,
dentate on margin, with mesial margin convex and lateral margin concave. Frontal region smooth, setal field with
anterior and posterior margins subequal in length, narrow lateral elements directed posteriorly, straight anterior
margin. CGI sinuous, slightly crenulate, bearing short setae, connected to posterior lateral elements. Mesogasiric
region smooth, CG2 absent; CG3 present only as short lateral grooves; CG4 nearly entire, median clement
slightly displaced anteriorly. Hepatic region rugose anteriorly and anteromesially, otherwise smooth, with short
rugose and setose lateral spine present on anterolateral margin. Epibranchial region roughly triangular, smooth,
posterior lateral margin with short row of setae. Metagastric region smooth; CG5 absent; CG6 slightly crenulate,
median concave element merging with CG7 to form hybrid groove, lateral fragments of CG6 connecting with
CG4; CG7 transverse, merging with median third of CG6. Cardiac region smooth; CG8 uninterrupted; CG9
absent; CG10 present as oblique grooves almost meeting in median; CGI 1 absent. Branchial region with 8-9
short, transverse rows of setae. Posterior margin deeply and evenly convex, with short lateral submarginal groove.
Branchiostegite unarmed, covered with long golden plumose setae, anterior region with many short rows of setae,
posterior region well calcified dorsally, membranous ventrally, with numerous irregular fragments.
Ocular plate completely concealed by front of carapace (Fig. 8B). Ocular peduncle (Fig. 8B) triangular (almost
ovate in smallest specimen), broadly separated, margins without setae, cornea not apparent but ocular pigment
visible in mesiodistal area.
Antennule (Fig. 8C) with segment III narrow proximally, expanding distally to twice proximal width, produced
distoventrally, simple setae on dorsal margin and few long plumose setae on distoventral margin; dorsal exopod
flagellum with 67 segments (only 1 specimen with intact flagella), long plumose setae on dorsal and ventral
margins; ventral endopod flagellum short, usually of 3 segments (proximal pair sometimes fused), plumose setae
on dorsal and ventral margins. Segment II with plumose setae on dorsal and ventral margins, and scattered on
lateral surface. Segment I wider than long, small tubercle on proximoventral margin, lateral surface with long
plumose setae dorsally and ventrally, and on dorsal, ventral and distal margins.
Antenna (Fig. 8D) with segment V about twice longer than wide, long plumose setae on dorsomesial margin,
and long simple setae on dorsolateral margin; flagellum with 7-8 segments, long simple setae on dorsal, ventral,
lateral and distal margins. Segment IV with long plumose setae on dorsomesial margin, and long simple setae on
dorsolateral margin. Segment III with long plumose setae on ventral margin, and scattered long simple setae on
proximodorsal margin. Segment II short, widening distally. with long simple setae on dorsolateral margin;
antennal acicle short, rounded, exceeding base of segment IV by about 1/4 length of segment IV, long plumose
setae on dorsal margin and long simple setae on lateral margin. Segment I rounded dorsally, flattened ventrally.
long plumose setae on margins, lateral surface rugose and with long setae dorsally, produced ventrally into oblong
flattened plate.
Maxilliped III (Fig. 8E) with dactylus with rounded tip, long simple setae on margins and lateral surface.
Propodus with longitudinal median row of simple setae, margins with simple setae. Carpus produced nearly to
distal end of propodus; lateral surface and margins with plumose setae. Merus inflated, unarmed, with plumose
setae on margins. Basis incompletely fused with ischium, without crista dentata. Exopod 2-segmented. proximal
segment small, distal segment styliform, tapering, approximately 1/2 length of merus, plumose setae on margins;
flagellum 1 -segmented, almost reaching distal end of merus.
Pereopod I subchelate. Dactylus (Fig. 9A) curved and tapering; lateral and mesial surfaces smooth, dorsal
margin with long plumose and short simple setae, ventral margin with short simple setae. Propodus (Fig. 9A)
lateral surface with numerous short, transverse rows of setose rugae, dorsal margin unarmed, ventral margin
distally produced into acute spine, cutting edge lacking teeth, lined with long plumose setae; lateral, mesial and
ventral margins with long setae. Carpus unarmed, dorsal and distal margins with long plumose setae; lateral
surface with few transverse rows of setae; mesial surface smooth, with scattered rows of long plumose setae,
margins with long plumose setae. Merus unarmed; lateral surface with scattered transverse rows of long plumose
setae, margins with long plumose setae; mesial side with few short row's of setae. Basi-ischium incompletely
fused, unarmed. Coxa unarmed.
Source :
394
C. B. BOYKO & A. W. HARVEY
Fig. 9. — Austrolepidopa Caledonia sp. nov., 9, allotype, 11.5 mm (MNHN-Hi 208) (A-E, G); <3, holotype, 9.2 mm
(MNHN-Hi 207) (F). A, left chela, lateral view; B, left pereopod II dactyl, lateral view; C, left percopod III dactyl,
lateral view; D. left pereopod IV dactyl, lateral view; E, abdomen, dorsal view; F, telson of male, dorsal view;
G, telson of female, dorsal view. Scale = 2.0 mm (A-D, F, G), and 4.0 mm (E).
Pereopods II-IV with dactyli laterally compressed and dorso ventral ly expanded.
Pereopod II dactylus (Fig. 9B) smooth, base to heel slightly concave, heel with smoothly rounded low spur,
heel to tip broadly indented and wide, tip rounded, tip to base broadly convex; lateral surface smooth, few setae
along dorsal margin between heel and tip; mesial surface smooth, ventral margin with long plumose setae, dorsal
margin with short plumose setae and a patch of long plumose setae between heel and base (not illustrated).
Propodus dorsal surface smooth, ventral margin inflated and rounded, oblique open row of long plumose setae on
lateral surface, distal and ventral margins with long plumose setae; dorsolateral surface a narrow, oblique, flattened
shelf, short setae on dorsal margin and long plumose setae on ventral margin; mesial surface with curved row of
simple setae from ventral junction with dactylus almost to ventral proximal junction with carpus. Caipus slightly
inflated and produced, gently rounded; lateral surface nearly smooth, with irregular, broken row of rugae and
submarginal elevated ridge ventrally, rugae and ridge with long plumose setae, margins with long plumose setae;
mesial surface smooth, submarginal and marginal rows of long plumose setae dorsally. Merus lateral surface fully
Source :
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
395
calcified, long plumose setae on margins; mesial surface with row of long plumose setae below dorsal margin and
row of setal patches 1/3 from ventral margin. Basi-ischium incompletely fused and unarmed. Coxa unarmed.
Pereopod III dactylus (Fig. 9C) with base to heel slightly concave, heel low and rounded, heel to tip broadly
concave, tip rounded, tip to base smoothly convex; lateral surface smooth, dorsal margin with few tufts of setae;
mesial surface smooth, with plumose setae proximally between heel and junction with propodus, ventral margin
with long plumose setae, dorsal margin with short simple and plumose setae. Propodus not much inflated; lateral
surface smooth, dorsolateral surface a narrow, oblique, flattened shelf, long plumose setae distally, simple setae on
margins, long plumose setae on ventral margin; mesial surface with scattered long setae on and near distal margin.
Carpus produced, nearly reaching distal margin of propodus, broadly rounded and inflated distally, dorsolateral
margin unarmed; lateral surface covered with numerous rows of short, simple setae forming a setal mat,
increasingly prominent distally; mesial surface smooth, long plumose setae on margins. Merus smooth, ovate,
dorsal and ventral margins unarmed, with long plumose setae, laterodistal margin with long plumose setae; lateral
surface fully calcified; mesial surface smooth. Basi-ischium incompletely fused and unarmed. Coxa unarmed.
Female with large gonopore on ventral surface of coxa, lacking setae, male with similar but smaller pore.
Pereopod IV dactylus (Fig. 9D) with base to heel straight, heel to tip broadly concave, tip rounded, tip to base
evenly convex, heel and blade of dactylus subequal in length; lateral surface smooth, ventral margin with long
plumose setae, dorsal margin with short simple setae; mesial surface smooth, plumose setae proximally. Propodus
expanded dorsally and ventrally; ventral expansion equals ventral dactylus margin, numerous short simple setae at
margins; dorsal expansion with row of long plumose setae; lateral and mesial surfaces smooth. Carpus not
produced dorsodistally; lateral and mesial surfaces smooth, dorsal margin with short simple and long plumose
setae, ventral margin with short plumose setae. Merus with scattered short transverse rows of setae on lateral
surface, dorsal and ventrodistal margins with long plumose setae, ventrodistal angle slightly expanded, ventral
surface fully calcified, smooth. Basi-ischium incompletely fused and unarmed. Coxa unarmed.
Pereopod V reduced, slender. Coxa of male with large mesiodistal gonopore.
Abdomen (Fig. 9E) with somite I approximately as wide as long, widest posteriorly, dorsal surface with
anterior margin straight, small submarginal decalcified spots anteriorly, posterior margin slightly concave,
submarginal row of short setae on elevated ridge, open row of setae anterior to ridge. Somite II anterior margin
straight, with tuft of setae at posterolateral angle; pleura expanded and directed slightly anteriorly, lateral margins
angled anteriorly and rounded posteriorly, anterior and lateral margins with long plumose setae, posterior margin
with row of short setae, becoming submarginal posteromesially. Somite III similar to somite II, but shorter;
pleura thinner and shorter than those of somite II, directed anteriorly, with setae as in somite II, anterolateral angle
acute, dorsal surface obliquely flattened anterolaterally. Somite IV similar to somite III, but thinner and shorter;
with posterior row of setae interrupted in median of posterior margin; pleura thinner and shorter than on somite III.
directed laterally and slightly anteriorly, dorsal surface obliquely flattened anterolaterally, margins with long
plumose setae. Somite V narrower than somite IV; pleura about 2/3 as long as on somite IV, directed posteriorly
and laterally, with setae as in somite IV, except with posterior row setae terminating at posterolateral angle of
somite. Somite VI subequal to somite V in width but longer, dorsal surface with short oblique rows of setae
laterad to midline posteriorly and medially, anterolateral margins with scattered plumose setae; pleura absent.
Females with uniramous, paired pleopods on somites II- V, males with small pleopod buds on somites II-V.
Uropods lacking distinctive features.
Telson of male (Fig. 9F) ovate, slightly wider than long, smoothly rounded posteriorly, dorsal surface with
6 short transverse rows of setae laterad to midline in median 1/3 of segment, rugose near anterolateral angle, with
short setae, margins with long plumose setae. Telson of female (Fig. 9G) similar to that of male, with larger
rugose areas anterolaterally.
Distribution. — Known from only New Caledonia; 205-225 m depth.
REMARKS. — Austrolepidopa Caledonia is most closely related to the west Australian A. trigonops Efford &
Haig, 1968, of which we have examined the holotype (WAM 62-62) and one of the paratypes (WAM 72-62).
Besides the considerable differences in geographic location and bathymetric distribution {A. trigonops ranges from
1 1 to 36 m), A. trigonops differs from A. Caledonia in having larger anterolateral spines, deeper ocular sinuses and
Source :
396
C. B. BOYKO & A. W. HARVEY
less anteriorly projected sctal field. The carapace grooves also differ between the two species; for example, in
A. trigonops , the median section of CG4 in the metagastric region is broken into four short elements, and CG8 is
interrupted medially and less produced laterally.
As mentioned earlier, there has been some confusion regarding sexual characteristics in this genus. For
example, the holotype of A . trigonops is not a female, as identified by EFFORD and Haig (1968), but rather a
male. In this genus, males not only possess gonopores on the coxae of the fifth pereopods, but also have small
but distinct pores on the coxae of the third pereopods, as well as small pleopod buds on abdominal somites 2-5.
Females have gonopores on only the third pereopod coxae and long, well developed plcopods on abdominal
somites 2-5.
The present record from 225 m is the greatest depth reported for any species in the Albuneidae, considerably
exceeding the previous record, 151m for a specimen o { Albunea synunysta (Linnaeus, 1758) from the Philippines
(USNM 68613; previously unpublished). Austrolepiclopa Caledonia appears to live in a habitat at least partially
composed of pteropod ooze, as evidenced by the small Limacina- type pteropods (Mollusca) found adhered to the
holotype (P. Mikkelsen, personal commun.).
Family HIPPIDAE Latreille, 1825
Genus HIPPA Fabricius, 1787
Remarks. — Systematic literature on the genus Hippo is extensive but confusing, and in many cases it is
impossible to determine which species are referred to without direct examination of specimens. As it is beyond the
scope of this study to resolve these problems, we give only select references in the lists of synonyms for species
in this genus.
Hippa pacifica (Dana, 1852)
Fig. 10
Remipes pacificus Dana, 1852: 407-408; 1855: pi. 25. figs 7a-g. — DE Man, 1896: 462, 476-478; 1898: 705. pi. 33,
figs 53, 53a-c; 1902: 690.
Remipes testudinarius - Miers. 1878: 316-318 (in part), probably pi. 5. fig. I (not R. testudinarius Latreille, 1806 =
Hippa adactyla Fabricius, 1 787).
Hippa pacificus - Thomassin. 1969: 157-160, pi. 7. text figs 7c, 8c, 10.
Hippa pacifica - Efford. 1972: 119-121. — Haig, 1974b: 181-183. fig. 3 (and references therein). — Bauchau, 1985:
313-314, pi. 4, pi. 5b-c.
Hippa pacific [sic] - SUN & Wang, 1996: 28-29, fig. 2.
MATERIAL EXAMINED. — New Caledonia. Islets Sainte-Marie, southwest lagoon, intertidal, coll. Bargibant.
14.02.1988: 1 ov. 9 16.4 mm (MNHN-Hi 211). — Poe beach, intertidal, coil. Danigo: 2 9 10.7-12.5 mm ; 2 ov. 9
16.5- 17.9 mm (MNHN-Hi 215). — Vata cove, intertidal, coll. Rudo von Cosel, 3.11.1993: 1 8 14.6 mm ; 2 ov 9
18.5- 20.1 mm (MNHN-HI 216).
Montrouzier Expedition. Touho Bay: intertidal, 8.09.1993: 2 ov. 9 13.4-16.4 mm (MNHN-Hi 212). — Intertidal.
9.09.1993: 14 8 5.5-11.8 mm ; 3 9 12.9-16.5 mm ; 2 ov. 9 15.4-16.1 mm (MNHN-Hi 213). — Intertidal, 9.09.1993;
1 9 12.0 mm ; 1 ov. 9 18.9 mm (MNHN-Hi 214).
Loyalty Islands. Niri Bay, intertidal, June 1986: 17 8 5.0-11.6 mm ; 20 9 6.1-18.7 mm ; 1 3 0v 9 12 2-
20.2 mm (MNHN-Hi 210).
Hawaii. Flag Poles, Kailua Beach, intertidal, coll. F. Mak. K. Bunney. Y. Kim. 27.05.1996: 6 8 7.3-12.3 mm ; 6 9
1 1.0-15.3 mm ; 1 ov. 9 13.7 mm (A. Harvey personal coll.).
Types. Syntypes: Hawaii and Fiji Islands. 1 specimen. Sandwich [= Hawaiian] Islands (MCZ 1406; not
examined for this study). Whereabouts of syntypes from Fiji Islands unknown.
Diagnosis. — Carapace (Fig. 10A) covered with wavy, transverse grooves. Frontal margin (Fig. 1 0B) with
lour rounded lobes, mesial pair slightly less projecting than lateral pair, separated by a shallow concavity, rarely
Source : MNHN. Paris
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
397
with a minute median denticle. Lateral margin of carapace (Fig. IOC) with submarginal row of 30-40 slightly
elongate, setose pits. Antenna (Fig. 10D) normally with two-segmented flagellum. Dactylus of second and third
pereopods with concave dorsal margin, indent obtuse. Telson (Fig. 10E) with lateral margins slightly convex,
smoothly converging distally (modified from Haig, 1974b).
FlG. 10. — Hippa pacifica (Dana, 1852), 9 12.1 mm (MNHN-Hi 210). A. carapace, dorsal view; B, front, dorsal view;
C. carapace, lateral view; D. left antenna, lateral view; E, telson. dorsal view. Scale = 0.75 mm (D), 1.5 mm (B). and
3.0 mm (A, C, E).
DISTRIBUTION. — One of the most widely ranging anomurans; known from Tanzania eastward to Australia,
north to China and across to the Hawaiian Islands (Haig, 1974b; SUN & Wang, 1996). It is also found in the
east Pacific from the upper Gulf of California south to Panama as well as Socorro, Clipperton, Cocos, and
the Galapagos Islands (Efford, 1972).
Source : MNHN. Paris
398
C. B. BOYKO & A. W. HARVEY
Remarks. — One of the key diagnostic features of Hippa pacifica is the two-segmented antennal flagellum.
Interestingly, in several of the New Caledonian specimens one of the antennal flagella has the normal two
segments and the other has only one segment. In a more problematic specimen from Niri Bay (MNHN-Hi 210),
both antennae are 3-segmented, and the frontal margin closely resembles that of H. celaeno . but in most other
respects this specimen is characteristic of H. pacifica. Additional material may eventually show this specimen to
represent a species distinct from H. pacifica or H. celaeno.
Fig. 1 1. — Hippa celaeno (de Man. 1896). 9 11.0 mm (MNHN-Hi 218). A, carapace, dorsal view; B. front, dorsal view;
C. carapace, lateral view; D. left antenna, lateral view; E. telson. dorsal view. Scale = 0.75 mm (D). 1.5 mm (B), and
3.0 mm (A. C. E).
Hippa celaeno (de Man, 1896)
Fig. 11
Remipes celaeno de Man. 1896: 462, 483-488; 1898: 705-706, pi. 33, figs 55, 55a-e; 1902: 690.
Hippa celaeno - HOLTHUIS, 1958: 42. — Lewinsohn, 1969: 173-174 (and references therein). — Haig, 1974b: 183-185.
fig. 4 (and references therein). — Bauchau, 1985: 314-315, pis 6. 8a. 9c.
Material EXAMINED. — New Caledonia. MONTROUZIER Expedition. Touho Bay: intertidal 8 09 1993*
1 ov. 9 9.9 mm. (MNHN-Hi 217). — Intertidal, 9.09.1993: 1 6 5.9 mm ; 8 ov. 9 10.9-12.32 mm.(MNHN-Hi 218). '
types. — Indonesia: Makassar and Amboina. 3 syntypic lots are in the RMNH (D164I. D1751. D2618, see
Fransen et all.< 1997) but they were not examined.
DIAGNOSIS. — Carapace (Fig. 1 1A) covered with wavy, transverse grooves. Frontal margin (Fig. 1 IB) with
four dentate, rounded lobes, mesial pair as projecting as lateral pair, separated by a deep concavity. Lateral margin
Source :
ALBUNEIDAE AND HIPP1DAE OF THE TROPICAL INDO-WEST PACIFIC REGION
399
of carapace (Fig. 1 1C) with submarginal row of 20-30 slightly elongate, setose pits, row sharply diverging from
margin posteriorly. Antenna (Fig. 1 ID) with onc-segmented flagellum. Dactylus of second and third pereopods
with concave dorsal margin, obtuse indent. Telson (Fig. 1 IE) with lateral margins parallel proximally, slightly
concave and with a noticeably constricted tip (modified from Haig, 1974b).
DISTRIBUTION. — Known from the Red Sea eastward to New Caledonia and south to Queensland. Australia
(Haig, 1974b).
Remarks. — According to Haig (1974b), Hippo celaeno is one of only two species in the genus with a one-
segmented antennal flagellum and less than 30 submarginal lateral pits, the other being H. picta (Heller, 1862).
Haig (1974b: 184) also remarks that "Hippo celaeno may be easily recognized by the abrupt concavity
of the anterior portion of the lateral margin, and by the strong divergence from that margin of the last
few setiferous pits". However, this may be an oversimplification. As mentioned under Hippo pacifica. the number
of segments in the antennal flagellum is somewhat variable; indeed, Haig's (1974b) key distinguishes
these two species in part by the number of segments "normally" present. In addition, we find that the number
of submarginal setose pits in both species is variable and somewhat size dependent, which is trouble¬
some since the larger species (H. pacifica) is also the species with the greater number of pits. Likewise,
the degree of concavity in the anterolateral margin of the carapace is variable, and overlaps considerably in these
two species.
Thus, the number of antennal segments and number of lateral carapace pits seem to be helpful but not infallible
in distinguishing Hippo celaeno and //. pacifica. More consistent characters include the degree of divergence of the
lateral pits from the posterior lateral margin of the carapace, the relative depth of the median indentation of the
frontal margin, and the shape of the telson. Hippo celaeno also has a relatively more slender carapace than same¬
sized specimens of H. pacifica , but this is probably too variable a character to be reliable.
Genus EMERITA Scopoli, 1777
Emerita austroafricana Schmitt. 1937
Emeritci austroafricana Schmitt, 1937: 25-29, pi. 3 top. — EFFORD. 1976: 172-173.
MATERIAL EXAMINED. — South Africa. Durban, intertidal, coll, unknown: 1 9 26.0 mm (USNM 71446). —
Durban, intertidal, coll, unknown: 1 9 (damaged) (AMNH 1155). — Near Durban, Natal, intertidal, coll, unknown,
Dec. 1961: 1 ov. 9 21.4 mm (USNM 267801).
Tanzania. Dar Es-Salaam. intertidal, coll. H. Sarroon, 1965: 1 ov. 9 25.7 mm (MRAC 51.623).
Madagascar. Toamasina, intertidal, coll. H. Bluntschli. June 1931: 1 ov. 9 29.3 mm (AMNH 17537). —
East coast, region of Pangalanes, intertidal, coll. A. Kienfr, Sept. 1970: 1 9 26.2 mm (MNHN-Hi 219).
TYPE. — Holotype : 9 26.0 mm, South Africa, Durban Bay, Natal, intertidal (USNM 71446).
Diagnosis. — Carapace cylindrical, covered with transverse grooves that extend across midline. Antenna
segment II with mediodistal spine short, less than twice length dorsodistal spine. Pereopod I with dactylus deeply
ovate, with 2 distodorsal, 1 distal, and 4-5 distoventral corneous-tipped spines, distoventral spines occupying half
to 2/3 ventral margin; carpus with pronounced anteromesial spine.
Distribution. — Known from Madagascar and Mozambique south to Durban and Natal, South Africa
(EFFORD, 1976). The above record from Dar-Es-Salaam represents an approximately 1000 km northward range
extension.
REMARKS. — Scarcely more is known about this species since EFFORD (1976) summarized its distribution and
commented that it is one of the least known members of the genus.
400
C. B. BOYKO & A. W. HARVEY
Annotated list of the tropical Indo-West Pacific species of the superfamily Hippoidea
(Taxa listed in bold-faced text are discussed in the main body of the paper).
ALBUNEIDAE
Albunea elioti Benedict, 1904 (see under A. microps in text).
Albunea holthuisi sp. nov.
Albunea madagascariensis Thomassin, 1973, known only from Madagascar.
Albunea microps Miers, 1878.
Albunea speciosa Dana. 1852, known with certainty only from Hawaii (SERENE. 1973). The identification of a
specimen from the Maldives by BORRADAILE (1904) is questionable and needs reexamination.
Albunea steinitzi Holthuis, 1958 (see under A. holthuisi sp. nov. in text).
Albunea symmysta (Linnaeus, 1758), known from the west coast of Africa north into the Red Sea and cast to
the Philippines and Japan (Thomassin, 1969). However, many of these records are not A. symmysta sensu stricto
(Boyko, unpublished) and the true range of this species is not completely known.
Albunea thurstoni Henderson, 1893, known from India and the Red Sea (Lewinsohn, 1969). Thomassin’ s
( 1 969) specimens from Madagascar are referable to A . madagascariensis (see Thomassin, 1 973).
Austrolepidopa Caledonia sp. nov.
Austrolepidopa schmitti Efford & Haig, 1968, known from Queensland, Australia.
Austrolepidopa trigonops Efford & Haig. 1968, known from Western Australia (see under A. Caledonia in
text).
Leucolepidopa sunda Efford, 1969, known only from the holotype collected in the Sunda Strait between
Sumatra and Java.
Paralbunea dayriti (Serene & Umali, 1965), known from the Philippines and China (Sun & Wang, 1996).
Haig (1974a) cited this species in a list of Western Australia anomurans, but we have seen no specimens from
that region.
Paralbunea manihinei Serene, 1979, known only from the Seychelles.
Paralbunea mariellae (Serene, 1973), known from Indonesia. Haig (1974a) cited this species in a list of
Western Australia anomurans (as Albunea undescribed sp.), but we have seen no specimens from that region.
Paralbunea paradoxa (Gordon, 1938), known from Singapore and the Philippines (see SERENE & UMALI
1965).
Stemonopa insignis Efford & Haig, 1968, known only from the holotype collected in Western Australia.
Zygopa nortoni Serene & Umali, 1965, known only from the Philippines.
HIPPIDAE
Emerita austroafricana Schmitt, 1937.
Emerita emeritus (Linnaeus, 1767), known from the west coast of India eastward to Vietnam and southward to
Sumatra and Java. Indonesia (Efford, 1976).
Emerita holthuisi Sankolli, 1965, known from the southern Red Sea along the coast to the southern west coast
of India (EFFORD, 1976).
Emerita karachiensis Niazi & Haque, 1974, known only from Karachi. Pakistan. A probable synonym of
E. holthuisi (unpublished data).
Hippa adactyla Fabricius, 1787, known from the west coast of Madagascar to Australia, eastward to the
Marquesas Islands, and northward to Japan (Haig, 1974b).
Hippa admirabilis (Thallwitz, 1892), known from New Guinea and Indonesia (DE Man, 1 896).
Hippa alcimede (de Man, 1902). known only from Java, Indonesia. A possible synonym of H. hirtipes ; the
allometric features DE Man used to distinguish the smaller H. alcimede from H. hirtipes are consistent with size-
related allometric changes within other hippie! species (unpublished data).
Source :
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
401
Hippa australis Hale, 1927, known from South and Western Australia (Haig, 1974b).
Hippa celaeno (de Man, 1896).
Hippa granulatus (Borradaile, 1904) (not in key), known from the Maldives. This species is a possible
synonym of H. alcimede (see Haig et al, 1986).
Hippa hirtipes (Dana, 1852), known from a few records in Indonesia and Papua New Guinea (Bauchau, 1985).
Hippa indica Haig, Murugan & Balakrishnan Nair, 1986, known only from the southwest coast of India.
Hippa marmoratus (Jacquinot, 1846) (not in key), known only from Raffles Bay, Australia (Haig 1974b), is a
possible synonym of H. pacifica (see Haig et al., 1986). For date see Clark & Crosnier (in press).
Hippa ovalis (A. Milne Edwards, 1862), known from the east coast of Africa eastward to Papua New Guinea
(Bauchau, 1985).
Hippa pacifica (Dana, 1852).
Hippa picta (Heller, 1862), known from the Red Sea (Holthuis. 1958).
Hippa truncatifrons (Micrs. 1878), known from Japan and China (Haig et al, 1986).
Mastigocliirus gracilis (Stimpson, 1858), known only from China (SUN & Wang, 1996).
Mastigocliirus quadrilohatus Miers, 1878, known from India, the Philippines, Western Australia and
Queensland, Australia (Haig, 1974b).
Key to the tropical Indo-West Pacific species of the superfamily Hippoidea
1. Pereopod I dactylus subchelate . Albuneidae ... 2
— Pereopod I dactylus simple . Hippidae ... 19
2. Abdominal somite V with pleura . 3
— Abdominal somite V without pleura . 6
3. Antenna with 3 flagellar articles . Leucolepidopa sunda Efford, 1969
— Antenna with more than 3 flagellar articles . Austrolepidopa Efford & Haig, 1968 ... 4
4. Ocular peduncles triangular in shape, rostral area margin dentate . 5
— Ocular peduncles ovate, rostral area margin smooth . A. schmitti Efford & Haig. 1968
5. Setal field projecting almost to base of ocular sinus, CG8 entire .
. A. Caledonia sp. nov.
— Setal field projecting anteriorly on level with base of lateral carapace spine, CG8 broken
. A. trigonops Efford & Haig, 1968 (see under A. Caledonia sp. nov. in text)
6. Antennal segment I with spine . Albunea Weber, 1795 ... 7
— Antennal segment I without spine . 14
7. Pereopod III dactylus heel acute . 8
— Pereopod III dactylus heel rounded . 1 0
8. CG 11 present . A. holthuisi sp. nov.
— CG 1 1 absent . 9
9. Telson of male spatulate, dorsoventrally compressed . A. symmysta (Linnaeus. 1758)
— Telson of male ovate with median setose ridge .
. A.steinitzi Holthuis, 1958 (see under A. holthuisi sp. nov. in text)
10. Ocular peduncle mesial margin straight . 1 1
— Ocular peduncle mesial margin convex . 13
1 1 . Ocular peduncle lateral margin convex . A. thurstoni Henderson. 1 893
— Ocular peduncle lateral margin concave . 12
12. Rostrum not exceeding distal margin of ocular plate . A. speciosa Dana. 1852
— Rostrum exceeding distal margin of ocular plate . . A. nmdagascariensis Thomassin, 1973
Source
402
C. B. BOYKO & A. W. HARVEY
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
Lateral indentation of cornea approximately 1/6 length of ocular peduncle .
. A.microps Miers, 1878
Lateral indentation of cornea approximately 1/2 length of ocular peduncle .
. A. elioti Benedict, 1904 (see under A. microps in text)
Ocular peduncles fused . Zygopa nortoni Serene & Umali, 1965
Ocular peduncles separate . 15
Ocular peduncles equal to or greater than carapace length .
. Stemonopa insignis Efford & Haig, 1968
Ocular peduncles less than carapace length . Paralbunea Serene, 1979 ... 16
Pereopod III dactylus heel acute . 17
Pereopod III dactylus heel rounded . 1 8
Pereopod IV dactylus heel acute . P. paradoxa (Gordon, 1938)
Pereopod IV dactylus heel rounded . P. dayriti (Serene & Umali, 1965)
Carapace nearly smooth . P. manihinei Serene, 1979
Carapace coarsely granular . P. mariellae (Serene, 1973)
Antennal flagellum at least as long as carapace, densely covered ventrally with long
plumose setae . Emerita Scopoli, 1777 ... 20
Antennal flagellum considerably shorter than carapace . 2 2
Pereopod I dactylus more than twice as long as wide, dorsal margin lacking spines .
. E. holthuisi Sankolli, 1965
Pereopod I dactylus less than twice as long as wide, dorsal margin armed with spines. 2 1
Pereopod I dactylus with spines along distal two-thirds or more of ventral margin .
. E.austroafricana Schmitt, 1937
Pereopod I dactylus with spines along distal half of ventral margin .
. E. emeritus (Linnaeus, 1767)
Pereopod I dactylus nearly as long as carapace, mulliarticulatc .
. Mastigochirus Miers, 1878 ... 23
Pereopod I dactylus less than one-fourth length of carapace, nonarticulated .
. Hippo Fabricius, 1787... 24
Frontal margin of carapace with 4 teeth: lateral margins of carapace unarmed .
. M. quadrilobatus Miers, 1878
Frontal margin of carapace with 3 teeth; lateral margins of carapace with several spines
. M. gracilis (Stimpson, 1858)
Frontal margin of carapace straight or slightly concave; lateral surface of carapace finely
punctate; dactylus of pereopods II and III with anterior margin nearly straight
. H. indica Haig, Murugan & Nair, 1986
Frontal margin of carapace with 1-3 median lobes; lateral surface of carapace with
submarginal row of pits or striations; dactylus of pereopods II and III with anterior
margin concave . 2 5
Frontal margin of carapace with 2 or 3 median lobes, lateral frontal lobes well developed;
lateral surface of carapace with submarginal row of setose pits . 2 6
Frontal margin of carapace with 1 broad median lobe, lateral frontal lobes weakly
developed; lateral margin of carapace with series of oblique striations or rows of pits 33
Carapace lacking Fine transverse grooves, Finely punctate . 2 7
Carapace covered with Fine tranverse grooves . 2 8
Source : MNHN, Paris
ALBUNEIDAE AND HIPPIDAE OF THE TROPICAL INDO-WEST PACIFIC REGION
403
27. Carapace with posterolateral margins oblique . H. hirtipes (Dana, 1852)
— Carapace with posterolateral margins nearly parallel . H. alcimede (de Man, 1902)
28. Lateral lobes of carapace front greatly exceed median lobes . 2 9
— All frontal lobes about equally projecting . 3 0
29. Dactylus of pereopods II and III with anterior margin cut into right angles; antennal
flagellum with 3-6 articles . H.adactyla Fabricius, 1787
— Dactylus of pereopods II and III with anterior margin obtuse; antennal flagellum with
1-2 articles . H. admirabilis (Thallwitz, 1892)
30. Antennal flagellum with 1 article; lateral margin of carapace with less than 30 pits in
submarginal row . 3 1
— Antennal flagellum with 2-3 articles; lateral margin of carapace with 30 or more pits in
submarginal row . 3 2
31. Submarginal row of pits diverges from lateral margin of carapace posteriorly; frontal
lobes broadly rounded, sinuses between lobes subcqual in depth .
. H.celaeno (de Man, 1896)
— Submarginal row of pits does not diverge from lateral margin of carapace posteriorly;
frontal lobes triangular, median sinus shallower than lateral sinuses .
. . . H. picta (Heller, 1862)
32. Lateral margin of carapace with 30-40 pits; antennal flagellum usually with 2 articles;
median frontal lobes separated by shallow sinus, rarely with minute median denticle .
. H. pacifica (Dana, 1 852)
— Lateral margin of carapace with 45-55 pits; antennal flagellum with 3 articles; median
frontal lobes separated by pronounced, broadly rounded lobe .
. H. ovalis (A. Milne Edwards, 1862)
33. Dorsal antennular flagellum with 19-26 articles, gradually tapering from base to tip .
. H. truncatifrons (Miers, 1878)
— Dorsal antennular flagellum with 32-40 articles, tapering only near tip .
. H. australis Hale, 1927
ACKNOWLEDGMENTS
We would like to thank the following for loans of specimens: NGUYEN Ngoc-Ho, Museum national d'Histoire
naturclle, Paris; Gary Rosenberg, Academy of Natural Science of Philadelphia; Charles H.J.M. Fransen,
Nationaal Natuurhistorisch Museum, Leiden; Niel BRUCE, Zoological Museum, University of Copenhagen;
Rudy JOCQUE, Musce royal de l'Afrique Centrale, Tervuren; Rafael LEMAITRE, National Museum of Natural
History. Washington; Miranda Lowe , The Natural History Museum, London; Jane GRIFFITH, Western Australian
Museum, Perth; and Ray J. SYMONDS, University Museum of Zoology, Cambridge.
The manuscript benefitted greatly from the thoughtful comments and suggestions of Darryl Felder and Ray
Manning. Special thanks are extended to Alain Crosnier and Bertrand Richer de Forges, both from
ORSTOM. for providing the MUSORSTOM materials.
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ESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS I
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
Enrique MACPHERSON
Centro de Esludios Avanzados de Blanes (CSIC)
Camf de Santa Barbara s/n
17300 Blanes (Girona), Spain
ABSTRACT
Galatheid crustaceans of the genera Agononida Baba & de Saint Laurent, 1996 and Munida Leach. 1820 collected in
Vanuatu, during the MUSORSTOM 8 cruise (September-October, 1994) have been studied. The collection contains
8 species of the genus Agononida and 25 belonging to the genus Munida. Two species are described as new: A. alisae and
M. congesta. A. alisae , close to A. callirhoe (Macpherson, 1994), can be distinguished easily by the spines of the
carapace and the antennal peduncle. M. congesta is close to M. miliaris Henderson, 1855, but is distinguished by the
shape and the spines of the chelipeds.
RESUME
Crustacea Decapoda : Especes des genres Agononida Baba & de Saint Laurent, 1996 et Munida
Leach, 1820 (Galatheidae) recoltees durant la campagne MUSORSTOM 8 au Vanuatu.
Les especes des genres Agononida Baba & de Saint Laurent, 1996, et Munida Leach, 1820. recoltees a Vanuatu
(Campagne Musorstom 8, septembre-octobre. 1994) sont au nombre de 8 et 25, respectivement. Deux especes (A. alisae
el M. congesta) sont nouvelles. A. alisae , proche de A. callirrhoe (Macpherson. 1994) se differencie facilement par
1'armature de la carapace et du pedoncule antennaire. M. congesta est proche de M. militaris Henderson. 1885, mais se
differencie par la forme et 1'armature des cheli pedes.
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. CROSN1ER (ed.). Resultats
des Campagnes MUSORSTOM. Volume 20. Memoires du Museum national d'Histoire naturelle , 180: 407-426. Paris ISBN
2-85653-520-8.
Source : MNHN. Pans
408
E. MACPHERSON
INTRODUCTION
During September-October of 1994 the cruise MUSORSTOM 8 was carried out in the Vanuatu waters (RICHER
DE Forges et al.t 1995) and numerous representatives of the genera Agononida Baba & de Saint Laurent. 1996
and Munida Leach. 1820 were collected. The study of these specimens revealed the presence of 8 species of the
genus Agononida and 25 species belonging to the genus Munida. One species of each genus has been considered
here as new.
The colour patterns of some species ( Agononida fortiantennata . A. squamosa, Munida masi, M. rhodonia,
M. rubrodigitalis and M. congesta) have been included in their remarks and descriptions. These patterns have been
described from colour slides of material obtained during the cruise by J.L. MENOU. As it has been pointed out in
other works (e.g. Rice & de Saint Laurent, 1985; Macpherson & de Saint Laurent. 1991;
MACPHERSON, 1994) the colouration can be a great help in species identification, although some species can show
some colour variability. The colour variations observed in some species (e.g. M. rhodonia, M. rubrodigitalis , see
below) recommend to use colour pattern with caution, especially when the ground colours arc the same. However,
the existence of two colour forms in Agononida squamosa (one form uniformly red and a second form with yellow
and purple spots, see below), without any additional and clear morphological difference, emphasizes the necesity of
complementary studies, e.g. genetics, mating behaviour, to clarify the validity and importance of the colour
pattern as a taxonomic character in this group (BRUCE, 1975; Knowlton, 1986).
The types of the new species and other material have been deposited in the collections of the Museum national
d'Histoire naturelle, Paris. Measurements given are of carapace length (CL), excluding rostrum, and the
terminology used mainly follows previous papers (Zariquiey- Alvarez. 1952; Macpherson & de Saint
Laurent, 1991; Macpherson, 1994; Baba & de Saint Laurent, 1996).
LIST OF STATIONS
The abbreviations of the gears used are: DW = Waren dredge, CP = Beam trawl. CC = Otter trawl. CAS = trap.
Station DW 958. — 20.09.1994, 20°20’S, 169°47'E, 497-570 m: A laurentae.
Station DW 959. — 20.09.1994, 20°20'S, 169°48,E. 436-475 m: A squamosa.
Station CP 961. — 21.09.1994, 20°18'S, 169°49’E. 100-1 10 m: M. clinata. M. gordoae.
Station CP 962. — 21.09.1994, 20°I9'S, 169°49'E, 370-400 m: M. rogeri, M. tyche.
Station CP 963. — 21.09.1994, 20°20’S, 169°49’E, 400-440 m: A alisae, A. squamosa. M. runcinata.
Station DW 965. — 21.09.1994, 20°20’S. 16905LE, 361-377 m: M. notata.
Station DW 966. — 21.09.1994. 20°19’S, 169°52'E, 128-150 m: M. clinata, M. elegantissima.
Station CP 971. — 21.09.1994. 20°19’S. 169°53'E, 250-315 m: M. sao, M. semoni.
Station DW 972. — 22.09.1994, 19°22'S, I69°28’E, 487-507 m: -4. ocxrhoe.
Station CP 973. — 22.09.1994, 19°2LS, 169°27'E, 460-480 m: A. squamosa.
Station CP 974. — 22.09.1994, I9°2TS. 169°28’E, 492-520 m: A incerta, A. ocyrhoe , A. squamosa,
M. rubrodigitalis, M. tuberculata.
Station CP 975. — 22.09.1994, 19°23’S, 169°29'E, 536-566 m: A. incerta, M. congesta, M. rhodonia.
Station DW 978. — 22.09.1994, 19°23’S, 169°27’E, 408-413 m: M. leptitis. M. runcinata.
Station CP 980. — 22.09.1994, 19°2TS, 169°25’E, 433-450 m: A laurentae, A. squamosa, M. idyia.
Station CP 982. 23.09.1994, 19°22'S, 169°26'E, 408-410 m: A. laurentae, A. squamosa, M. leagora.
Station CP 983. 23.09.1994, 19°22'S, 169°28'E, 475-480 m: A. laurentae, M. leptitis, M. rubrodigitalis.
Station CP 984. — 23.09.1994, 19°20'S, 169°26'E, 480-544 m: 4. incerta, A. ocyrlwe. A. squamosa.
M. rubrodigitalis.
Station CP 985. —23.09.1994. 19°22’S, 169°26'E, 536-563 m: M. rubrodigitalis. M. tuberculata.
Station DW 986. — 23.09.1994, I9°21'S, 169°31'E. 602-648 m: M. rhodonia.
Source : MNHN . Paris
AGONONIDA AND MUNIDA FROM VANUATU
409
Station DW 988. — 23.09.1994, I9°I6’S, 169°24'E, 372-466 m: M. rufiantennidata. M. runcinata.
Station DW 989. — 23.09.1994. 19°14'S. 169°20'E. 650-669 m: 4. incerta.
Station CP 990. — 24.09.1994. 18°52'S. 168°51'E. 980-990 m: 4. eminens, M. microps.
Station CP 991. — 24.09.1994. 18051'S, 168°52'E. 910-936 m: 4. eminens. M. microps.
Station CP 992. — 24.09.1994. 18°52'S. 168°55'E. 748-775 m: M. microps. M. rosula.
Station CP 993. — 24.09.1994. 18°48'S, 168°54'E. 780-783 m: 4. eminens. M. microps. M. rosula.
Station CP 994, — 25.09.1994, I8°47'S. 168°56'E, 641-649 m: 4. incerta.
Station CC 996. — 25.09.1994, 18°52'S, 168°56'E, 764-786 m: 4. eminens. M. rosula.
Station CP 1001. — 26.09.1994. 18°49'S, I68°59'E, 764-786 m: M. masi.
Station CP 1007. — 26.09.1994, 18°52’S, 168°56'E. 720-830 in: 4. eminens. M. microps. M. rosula.
Station CP 1008. — 26.09.1994, 18°53'S, 168°53'E, 919-1000 m: 4. eminens. M. microps.
Station DW 1014. — 27.09.1994. I7°54’S. I68°19'E. 495-498 m: M. microps.
Station CP 1017. — 27.09.1994, 17°53’S, I68°26'E, 294-295 m: M. notata.
Station CP 1018. — 27.09.1994, 17°53'S. 168°25’E. 300-301 in: M. notata.
Station DW 1021. — 28.09.1994, I7°43'S, 168°37’E, 124-130 in: M. clinata.
Station CP 1024. — 28.09.1994, 17°48'S, 168°39'E, 335-370 in: M. notata.
Station CP 1025. — 28.09.1994, 17°49'S, I68°39'E, 385-410 m: 4. alisae. 4. squamosa. M. runcinata.
Station CP 1026. — 28.09.1994, 17°50'S, 168°39'E. 437-504 m: 4. squamosa. M. sao.
Station CP 1027. — 28.09.1994, I7°53'S. 168°39'E. 550-571 in: 4. incerta. 4. normani. M. rhodonia.
M. tuberculata.
Station CP 1028. — 28.09.1994, 17°54'S. 168°40'E, 624-668 in: 4. incerta. 4. normani. M. congesta.
M. leviantennata. M. rhodonia.
Station DW 1029. — 28.09.1994, 17°53'S. I68°34'E. 324-360 m: 4. squamosa.
Station CC 1033. — 29.09.1994, 17°54'S. 168°40'E. 650-691 in: 4. incerta.
Station CC 1034. — 29.09.1994, 17°54'S, 168°42'E, 690-750 m: 4. eminens. M. militaris.
Station CP 1035. — 29.09.1994, I7°56'S, 168°44'E, 765-780 m: 4. eminens, M. rosula, M. runcinata.
Station CP 1036. — 29.09.1994, 18°01'S, 168°48'E, 920-950 m: 4. eminens. M. microps.
Station CP 1037. — 29.09.1994, 18°03'S, 168°54'E, 1058-1086 m: M. microps.
Station DW 1042. — 30.09.1994, I6°52'S, 168°52'E. 200-260 m: M. tyche.
Station DW 1043. — 30.09.1994, I6°52’S, 168°25'E, 350-372 m: M. runcinata.
Station DW 1045. — 30.09.1994, 16°54’S, I68°20'E, 459-488 in: 4. incerta. M. rhodonia.
Station CP 1047. — 30.09.1994. 16°53'S. 168°10'E, 486-494 m: 4. incerta. M. pagesi. M. rhodonia.
M. rubrodigitalis, M. sacksi.
Station CP 1049. — 1.10.1994, 16°39'S, 168°02'E. 469-525 in: 4. squamosa. M. idyia. M. rhodonia.
M. rubrodigitalis.
Station CP 1050. — 1.10.1994, 16°39'S, 168°01'E, 541-577 m: 4. incerta, M. rhodonia.
Station CP 1051. — 1.10.1994, 16°36’S, 167°59'E. 555-558 m: M. leviantennata. M. rhodonia.
Station CP 1055. — 1.10.1994. 16°30'S, 167°55'E, 572-580 m: M. rhodonia.
Station CC 1056. — 1.10.1994, I6°33'S. 167°55'E, 602-620 in: 4. incerta. M. rosula.
Station DW 1060. — 2.10.1994, 16°13'S, 167°20'E, 397-375 m: 4. squamosa. M. idyia, M. leagora.
Station DW 1061 .— 2, 10. 1994, 16°14'S. 167°20'E, 458-512 m: M. sao.
Station DW 1065. — 2.10.1994, 16°16'S, 167°2I'E, 360-419 m: M. notata. M. runcinata.
Station DW 1067. — 2.10.1994, 16°16'S, I67°21'E. 344-366 in: M. leagora.
Station DW 1070. — 4.10.1994, 15°36'S, 167°16'E. 184-190 m: M. sao.
Station CP 1071. — 4.10.1994, I5°36’S, I67°16'E, 180-191 m: M. semoni.
Station CP 1073. — 4.10.1994, 15°45'S, I67°22'E, 630-650 in: 4. incerta, M. leviantennata. M. rhodonia.
M. rosula.
Station CP 1075. — 4.10.1994, 15°53'S. 167°27'E. 944-956 in: M. microps.
Station CP 1076. — 4.10.1994, 15°53'S. 167°30'E, 1 1 00- 1191 m: 4. fortiantennata. M. microps.
Station CP 1077. — 5.10.1994, I6°04'S, 167°06'E, 180-210 in: M. elegantissima. M. semoni.
Source :
410
E. MACPHERSON
Station CP 1078. — 5.10.1994, 16°03’S. 167°26'E, 194-230 m: M. semoni.
Station CP 1080. — 5.10.1994, I5°57'S. 167°27'E. 799-850 m: 4. eminent. M. microps, M. rosula.
Station CAS 1081. — 5.10.1994, 15°55'S, I67°27'E, 908 m: M. microps.
Station CP 1083. — 5.10.1994, 15°5PS, 167°19'E, 397-439 m: A. squamosa. M. leagora.
Station CP 1086. — 5.10.1994, 15°36'S, 167°16’E, 182-215 in: M. sao.
Station CP 1087. — 6.10.1994, I5°10'S, 167°14'E. 394-421 m: A. alisae. A. squamosa. M. idyia,
M. runcinata.
Station CP 1088. — 6.10.1994, I5°09'S, I67°I5'E, 425-455 m: A. squamosa, M. idyia. M. leviantennata.
M. rubrodigitalis, M. sao.
Station CP 1089. — 6.10.1994, I5°08'S. 167°17'E, 494-516 m: A. incerta, M. rhodonia, M. rubrodigitalis.
M. sao.
Station CP 1090. — 6.10.1994, I5°08'S, 167°17'E, 470-502 m: A. incerta. M. rhodonia. M. rubrodigitalis.
Station CP 1091. — 6.10.1994, 15°10'S, 167°13'E, 344-350 m: 4. incerta. M. notata.
Station CP 1095. — 6.10.1994, 15°07'S, 1 67° 1 1'E, 304-320 m: M. sao.
Station DW 1097. — 7.10.1994, I5°05'S, 167°10'E, 281-288 m: 4. incerta. M. notata.
Station CP 1098. — 7.10.1994, 15°04’S, I67°10'E. 277-285 m: M. sao.
Station DW 1 100. — 7.10.1994, 15°04'S, 167°09'E, 258-265 m: M. leptitis.
Station CP 1 102. — 7.10.1994. 15°03'S, I67°08'E. 208-210 m: M. semoni.
Station CP 1 107. — 7.10.1994. 15°05'S. 167°15'E, 397-402 m: 4. squamosa. M. sao.
Station DW 1 108. — 7.10.1994, 15°04’S, 167°15'E, 405-419 m: 4. squamosa.
Station CP 1111. — 8.10.1994, 14°51'S, I67°14'E, 1210-1250 m : M. leviantennata, M. typhle.
Station CP 1 1 14. — 8.10.1994, 15°52'S, I67°03'E. 647 m: 4. incerta. M. militaris.
Station CP 1 1 18. — 9.10.1994, 15°08'S, 166°53'E, 191-248 m: M. semoni.
Station CP 1 1 19. — 9.10.1994, I5°08'S. 166°53'E, 254-300 m: M. semoni.
Station CP 1 124. — 9.10.1994, 15°0rS, 166°56'E, 532-599 m: 4. incerta. M. rhodonia. M. sacksi.
Station CP 1 125. — 10.10,1994. 15°57'S, 166°38'E. 1 160-1220 m: M. microps.
Station CP 1126.— 10.10.1994, 15°58'S, 166°39'E. 1210-1260 m: M. microps.
Station CP 1 129. — 10.10.1994, 16°00'S. 166°39'E, 1014-1050 m: 4. fortiantennata. M. microps.
Station CP 1131. — 11.10.1994. 15°38'S. 167°03'E, 140-175 m: M. tyche.
Station CP 1132. — 11.10.1994, 15°38’S. 167°02'E, 161-182 m: M. tyche.
Station CP 1 133. — 1 1. 10.1994. 15°38'S, I67°03'E, 174-210 m: M. tyche.
Station CP 1 135. — 1 1.10.1994, 15°40’S. I67°02’E, 282-375 m: M. masi. M. runcinata. M. sacksi.
Station CP 1 136. — 1 1.10.1994. 15°40'S, 167°01'E, 398-400 m: 4. squamosa. M. idyia. M. sao.
Station CP 1 137. — 1 1.10.1994. 15°41'S, I67°02'E, 360-371 m: 4. squamosa. M. idyia. M. leviantennata.
M. runcinata , M. sao.
Station CP 1 138. — 1 1.10.1994, I5°44'S, I67°04'E, 462-492 m: 4. incerta, M. leagora.
Station CP 1139. — 11.10.1994. 15°47'S. 167°08'E. 235-251 m: M. masi.
SYSTEMATIC ACCOUNT
Agononida alisae sp. nov.
Fig. 1
Material EXAMINED. — Vanuatu. Musorstom 8: stn 963, 400-440 m: 13 6 10.2 to 13.3 mm; 24 ov 9 10 2 tc
14.5 mm; 3 $ 1 1.6 to 12.7 mm; 1 juv. 6.2 mm. — Stn 1025, 385-410 m: 1 ov. 9 10.4 mm; 2 9 7.5 and 8 4 mm -
Stn 1087, 394-421 m: 1 <3 8.2 mm; 2 ov. 9 10.3 and 13.3 mm.
Types. The male of 12.6 mm from the Stn 1076 has been selected as holotype, the other specimens are
paratypes.
Source :
AGONONIDA AND MUNIDA FROM VANUATU
41 1
Fig. 1. — Agononida ciliscie sp. nov., 6, 12.6 mm, holotype from stn 1076 : 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, right cheliped, dorsal view; f, right First walking leg, lateral view; g. dactylus of right first walking,
lateral view.
Source : MNHN. Paris
412
E. MACPHERSON
Description. — Carapace, excluding rostrum, slightly longer than wide; few secondary striae between main
striae. One epigastric spine behind each supraocular spine; one protogastric spine behind each epigastric spine was
observed in one specimen. Three longitudinal rows of spines on carapace surface. Median row of 6 spines: first
two on median mesogastric region, third to fifth on cardiac region, sixth spine on posterior ridge. Lateral rows
each of 2-3 spines on branchiocardiac boundary. Frontal margins somewhat oblique. Anterolateral spine
pronounced, situated at anterolateral angle of carapace, not overreaching sinus between rostrum and supraocular
spines. Second marginal spine before cervical groove clearly smaller than preceding one. Branchial margins with
3 spines, decreasing in size posteriously. Rostrum nearly horizontal, feebly sinuous, about half as long as
remaining carapace. Supraocular spines more slender than rostrum, not overreaching end of corneae.
Thoracic sternites with numerous arcuate striae. Second to fourth abdominal segments with 4 spines on
anterior transverse ridge; median pair of spines larger than lateral spines; posterior ridge of fourth segment with
strong median spine. Gonopods in males absent from First abdominal segment.
Eye moderately large, maximum corneal diameter about 1/2 length of anterior border of carapace between bases
of anterolateral spines.
Basal antennular segment (distal spines excluded) not overreaching cornea; distomesial spine longer than
distolateral; two spines on lateral margin, proximal short, distal long, not exceeding distolateral spine.
Distomesial prolongation of first antennal segment well developed, clearly overreaching distal spines of
antennular peduncle and nearly reaching rostral tip; distomesial spine on second segment reaching end of third
segment, with well developed spine on its base; third segment unarmed.
Ischium of third maxi Biped about 2 times length of merus, measured along extensor margin, distoventrally
bearing long spine. Flexor margin of merus with median spine, extensor border with small distal spine.
Chelipeds squamous; palm with mesial row of well-developed spines. Fingers of chelipeds unarmed, fixed
finger bifid distally and with a longitudinal keel reaching tips.
Walking legs squamous. First walking leg more than 3 times carapace length; merus clearly longer
than carapace length, with one row of dorsal spines and one row of ventral spines; carpus only distal spines;
propodus about 1/2 merus length; dactylus of walking legs about 2/3 propodus length, without spinelets on
ventral border.
Remarks. — The new species is closely related to A callirrhoe (Macpherson, 1994) from New Caledonia,
Loyalty Islands and Chesterfield Islands. The two species have spines on the gastric, cardiac and branchiocardiac
regions, the fourth abdominal segment with one spine on the posterior ridge, the first antennal segment with an
unusually prolonged process, the distomesial spine longer than the distolateral on the basal antennular segment and
the thoracic sternites with numerous striae. The new species differs from A. callirrhoe in the following constant
characters:
Carapace armature quite different: two mesogastric spines in A. alisae , one in A. callirrhoe ; two cardiac
spines in A. callirrhoe , three in the new species.
.Second segment of antennal peduncle with one long distomesial spine reaching end of antennal peduncle,
with small spine on its base in A. callirrhoe. This spine is clearly shorter in the new species, only reaching end of
third antennal segment and with well developed spine on its base.
Etymology. — The name refers to the R/V “ Alis ", on which the cruise MUSORSTOM 8 was carried out.
Distribution. — Vanuatu, between 385 and 440 m.
Agononida eminens (Baba. 1988)
Munida eminens Baba, 1988: 95, fig. 35; 1994: 11. — MACPHERSON, 1994: 456, fig. 72; 1995: 392.
Agononida eminens - Baba & de Saint Laurent. 1995: 442 (list). — Macpherson, 1997: 600.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 990, 980-990 m: 2 9 12.4 and 13.6 mm. — Sin 991 910-
936 m: 2 d 13.0 and 15.0 mm; 3 9 9.8 to 14.6 mm. — Stn 993. 780-783 m: 1 9 15.8 m. — Stn 996, 764-786 m: 1 9
Source :
AGONONIDA AND MUNI DA FROM VANUATU
413
15.4 mm. — Stn 1007, 720-830 m: 1 d 16.7 mm; 1 9 18.0 mm. — Stn 1008, 919-1000 m: 2 d 13.4 and 14.2 mm. —
Stn 1034, 690-750 m: 1 d 20.0 mm. — Stn 1035, 765-780 m: 2 d 18.2 and 19.7 mm. — Stn 1036, 920-950 m: 2 d
12.8 and 18.8 mm; 1 9 14.9 mm. — Stn 1080, 799-830 m: 1 ov. 9 18.1 mm.
DISTRIBUTION. — Philippines, Indonesia, eastern Australia, New Caledonia, Loyalty Islands, Chesterfield
Islands, Wallis and Futuna Islands, between 564 and 1000 m. The present material was collected at 690-1000 m.
Agononida fortiantennata (Baba, 1988)
Fig. 3 a
Munida fortiantennata Baba, 1988: 101, fig. 37. — Macpherson, 1993: 428.
Agononida fortiantennata - Baba & DE Saint Laurent, 1996: 442 (list).
MATERIAL EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1076, 1100-1191 m: 4 d 11.5 to 14.2 mm; 5 9 8.3 to
13.7 mm. — Stn 1129, 1014-1050 m: 1 9 14.0 mm.
Remarks. — The species was only known from the holotype (female of 21.7 mm. including rostrum)
collected in the Molucca Sea, off west coast of Halmahcra in 763 m (Baba, 1988) and additional specimens caught
in the Philippines, southwest of Luzon, between 750 and 925 m (Macpherson, 1993). The specimens collected
in Vanuatu agree quite well with the holotype description. However, one male (CL, 1 1.5 mm) from Stn 1076 has
a well developed mesogastric spine; this spine is absent in the holotype and other material from Vanuatu.
COLOUR. — Carapace and abdominal segments reddish. Chelipeds and walking legs light red, without
conspicuous transverse bands; terminal part of chcliped lingers red.
Distribution. — Previously known from the type locality (Molucca Sea. 763 m) and southwest of Luzon at
750-925 m. The present material has been collected at 1014-1191 m.
Agononida incerta (Henderson. 1888)
Munida incerta Henderson, 1888: 130, pi. 13, fig. 4a. — Baba, 1988: 106; 1994: 12. — Macpherson, 1994: 478.
fig. 74; 1995: 394.
Agononida incerta - Baba & de Saint Laurent. 1996: 442 (list). — Macpherson, 1997: 600.
MATERIAL. EXAMINED. — Vanuatu. MUSORSTOM 8: stn 974. 492-520 m: 1 ov. 9 22.8 mm; 1 juv. 7.7 mm. —
Stn 975, 536-566 m: 3 d 25.2 to 29.1 mm; 1 9 22.5 mm. — Stn 984, 480-544 m: 1 ov. 9 20.3 mm. — Stn 989. 650-
669 m: 1 d 12.6 mm. — Stn 994, 641-649 m: 1 d 25.2 mm. — Stn 1027, 550-571 m: 5 d 20.4 to 26.8 mm; 3 ov. 9
18.3 to 22.7 mm. — Stn 1028, 624-668 m: 3 d 21 A to 32.3 mm; I ov. 9 23.4 mm. — Sin 1033, 650-691 m: I ov. 9
21.1 mm. — Stn 1045, 459-488 m: 1 9 17.0 mm. — Stn 1047, 486-494 m: 6 d 10.3 to 22.3 mm; 10 ov. 9 15.3 to
18.8 mm; 6 9 9.6 to 16.6 mm. — Stn 1050, 541-577 m: 3 d 19.2 to 22.6 mm; 2 9 12.7 and 17.9 mm. — Stn 1056,
602-620 m: 4 d 17.6 to 22.0 mm; 1 ov. 9 17.8 mm; 1 9 18.1 mm. — Stn 1073, 630-650 m: Id 12.0 mm. —
Stn 1089, 494-516 m: 8 6 8.5 to 22.4 mm; 5 ov. 9 13.6 to 16.2 mm; 5 9 7.0 to 19.3 mm. — Stn 1090, 470-502 m:
470-502 m: 5 d 15.3 to 22.2 mm; 2 9 17.3 and 17.8 mm. — Stn 1091, 344-350 m: 1 9 7.0 mm. — Stn 1097. 281-
288 m: 1 9 8.7 mm. — Stn 1114, 647 m: 1 9 23.3 mm. — Stn 1124, 532-599 m: 5 d 1 1.0 to 18.2 mm; 2 9 10.3 and
12.8 mm. — Stn 1 138, 462-492 m: 1 ov. 9 19.8 mm.
DISTRIBUTION. — Known from east African coast, Japan, Philippines, Indonesia, eastern Australia, New
Caledonia, Loyalty Islands, Chesterfield Islands. Wallis and Futuna and Kiribati between 17 and 720 m. The
material from Vanuatu has been collected between 281 and 691 m.
Agononida laurentae (Macpherson, 1994)
Munida laurentae Macpherson, 1994: 483. figs 25, 92.
Agononida laurentae - Baba & DE Saint Laurent. 1995: 442 (list).
Source :
414
E. MACPHERSON
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 980. 433-450 m: 2 8 10.6 and 13.6 mm. — Stn 982. 408-
410 m: 1 ov. 9 14.0 mm. — Stn 983. 475-480 m: 3 9 6.4 to 1 1.4 mm.
Distribution. — Known from New Caledonia. Loyalty Islands, Chesterfield Islands, Matthew and Hunter
Islands, between 260 and 610 m. The material from Vanuatu has been collected at 408-480 m.
Agononida normani (Henderson. 1885)
Munida Normani Henderson. 1885: 408.
Munida normani - HENDERSON. 1888: 129. pi. 13. fig. 5. — Baba. 1988: 83 (key). — MACPHERSON, 1994: 500; 1995:
400, fig. 20.
Agononida normani - Baba & DE Saint Laurent. 1996: 442 (list).
Material EXAMINED. — Vanuatu. Musorstom 8; stn 1027. 550-571 m: 2 9 11.2 and 12.0 mm. — Stn 1028,
624-668 m: 1 8 8.2 mm.
Remarks. — The specimens from Vanuatu agree with the types and additional material from New Caledonia,
Wallis and Futuna. However, the cardiac spines are absent in the present material, whereas there arc 0-5 spines in
the types and other material previously collected.
Distribution. — The species has previously been cited in Fiji, New Caledonia and Wallis and Futuna area
between 320 and 600 m. The present material was caught at depths between 550 and 668 m.
Agononida ocyrhoe (Macpherson, 1994)
Munida ocyrhoe Macpherson, 1994: 503. figs 35, 79; 1995: 402. fig. 21.
Agononida ocyrhoe - Baba & de Saint Laurent. 1996: 442 (list).
Material EXAMINED. — Vanuatu. Musorstom 8: stn 972, 487-507 m: I juv. 5.5 mm. — Stn 974. 492-520 m: 2
8 1 1.4 and 13.5 mm; 2 ov. 9 12.5 and 14.6 mm. — Stn 984, 480-544 m: 3 8 8.5 to 21.2 mm.
Distribution. — The species has previously been cited in New Caledonia, Chesterfield Islands and Wallis
Islands between 420 and 650 m. The material from Vanuatu has been collected at 480-544 m.
Agononida squamosa (Henderson, 1885)
Figs 3 b-c
Munida squamosa Henderson. 1885: 409; 1888: 131, pi. 13, figs la-b. — Baba. 1988: 133 (in part); 1994: 12. —
Macpherson, 1993: 425, fig. lh-1; 1994: 537. fig. 96; 1995: 406.
Agononida squamosa - Baba & DE SAINT LAURENT, 1996: 442 (list). — MACPHERSON, 1997: 603.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 959, 436-475 m: 1 8 15.2 mm. — Stn 963, 400-440 m:
1 8 14.5 mm; 1 9 7.6 mm. — Stn 973, 460-480 m: 1 8 13.2 mm. — Stn 974, 492-520 m: 2 8 14.4 and 16.5 mm. —
Stn 980. 433-450 m: 14 8 8.2 to 15.9 mm; 1 ov. 9 11.3 mm; 4 9 8.0 to 14.8 mm; 3 juv. 6.5 to 6.6 mm. — Stn 982,
408-410 m: 1 ov. 9 1 1.6 mm. — Stn 984, 480-544 m: 2 8 14.2 and 14.5 mm. — Stn 1025, 385-410 m: I 8 9.5 mm;
3 ov. $ 13.8 to 14.7 mm. — Stn 1026, 437-504 m: 2 8 14.7 and 15.3 mm; I ov. 9 11.8 mm. — Stn 1029, 324-
360 m: 1 ov. 9 12.6 mm. — Stn 1049. 469-525 m: 1 8 11.4 mm; 3 ov. 9 15.4 to 18.5 mm. — Sin 1060. 375-397 m:
1 8 15.0 mm; 1 ov. 9 10.2 mm. — Stn 1083, 397-439 m: 3 8 11.4 to 14.2 mm; 1 ov. 9 11.9 mm. — Stn 1087, 394-
421 m: 1 8 6.3 mm; 1 ov. 9 8.7 mm. — Stn 1088. 425-455 m: 1 8 15.0 mm; 3 ov. 9 1 1.8 to 14.2 mm. — Stn 1107,
397-402 m: 1 8 11.3 mm; 2 ov. 9 12.0 and 12.8 mm. — Sin 1 108, 405-419 m: I 8 12.9 mm. — Stn 1 136, 398-400 m:
1 ov. 9 12.1 mm; 2 9 10.0 and 12.2 mm. — Stn 1137, 360-371 m: 5 <3 11.6 to 16.2 mm; 6 ov. 9 9.8 to 14.2 mm;
3 juv. 6.4 to 6.8 mm.
Remarks. — The colour pattern of the ovigerous female (CL, 12.6 mm) from Stn 1029 (Fig. 3c) agrees with
the figure and description ot the specimens from Loyalty Islands, having numerous yellow and purple spots on
Source : MNHN. Pans
AGONON/DA AND MUNIDA FROM VANUATU
415
the carapace, and red and whitish bands on the pereiopods (MaCPHERSGN, 1994). However, one male (CL,
13.2 mm) from Stn 973 (Fig. 3b) has the carapace, abdominal segments and pereiopods uniformly red, without
bands and spots. This clear difference in the colour patterns suggests the existence of two forms or species.
However, no other clear morphological differences have been observed and, as has been pointed out in
the Introduction, a revision of the species using complementary techniques is strongly recommended.
DISTRIBUTION. — Japan, Indonesia, Admiralty Islands, northeastern Australia, New Caledonia, Loyalty Islands
and Wallis Islands, between 176 and 752 m. The specimens from Vanuatu were collected at 324-525 m.
Genus MUNIDA Leach, 1820
Munida clinata Macpherson, 1994
Munida clinata Macpherson, 1994: 457, fig. 11; 1995: 391; 1997: 605.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 961, 100-110 m: 2 ov. 9 7.8 and 8.7 mm. — Stn 966.
128-150 m: 1 6 5.5 mm; 1 ov. 9 8.8 mm. — Stn 1021, 124-130 m: 1 ov. 9 5.4 mm.
DISTRIBUTION. — Previously known from the Philippines, Indonesia, New Caledonia, Chesterfield Islands and
Futuna Island, between 28 and 245 m. The specimens from Vanuatu were collected at 124-150 m.
Munida congesta sp. nov.
Figs 2, 3 d. 4 a
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 975, 536-566 m: I 6 7.6 mm; 1 ov. 9 10.2 mm . —
Stn 1028, 624-668 m: 1 9 12.6 mm.
Types. — The ovigerous female of 10.2 mm from the Stn 975 has been selected as holotype, the other
specimens are paratypes.
ETYMOLOGY. — From the Latin, congestus , dense, thick, in reference to the massive chelipeds.
DESCRIPTION. — Carapace as long as wide. Main striae on posterior part of carapace interrupted in cardiac
region. Few secondary striae present. Intestinal region without striae. Three-five pairs of epigastric spines; one
branchial anterior spine on each side.
Frontal margins somewhat oblique. Lateral margins slightly convex. Anterolateral spine well developed,
situated at anterolateral angle, reaching level of sinus between rostrum and supraocular spine. Second marginal
spine before cervical groove clearly smaller than preceding one. Branchial margins with 5 spines.
Rostrum about half as long as carapace. Supraocular spines more slender than rostrum, reaching to its
midlength, but not to end of corneae, slightly divergent.
Thoracic sternites without striae.
Second abdominal segment with one row of 7-8 spines on anterior border. Second to fourth segments with one
transverse stria.
Eyes large, maximum corneal diameter about half the distance between bases of anterolateral spines.
Basal segment of antennule reaching end of cornea; distomesial spine shorter than distolateral; two spines on
lateral margin, proximal short, located at midlength of segment, distal long, overreaching distal spine.
First segment of antennal peduncle with strong distomesial spine, not reaching end of second segment; second
segment with two distal spines, mesial longer than lateral, not overreaching antennal peduncle.
Ischium of third maxilliped slightly longer than rnerus, measured along extensor border, distoventrally bearing
spine. Merus bearing two well developed spines on flexor margin, proximal spine clearly longer than distal.
Extensor margin unarmed.
Source :
416
E. MACPHERSON
Fig. 2. Munida congesta sp. nov., ovig. 9. 10.2 mm. holotype from stn 975 : 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, right cheliped, dorsal view; f. right first walking leg, lateral view; g, dactylus of right first walking,
lateral view.
Source : MNHN, Pans
AGONONIDA AND MUNIDA FROM VANUATU
417
Fig. 3. — a, Agononida fortiantennata (Baba, 1986), male 11.9 mm, stn 1076; b. Agononida squamosa (Henderson,
1885), male 13.2 mm, stn 973; c, Agononida squamosa (Henderson. 1885), ovigerous female 12.6 mm. stn 1029;
d, Munida congesta sp. nov., ovigerous female 10.2 mm, holotype, sin 975.
Source :
418
E. MACPHERSON
Chelipeds squamous. Merus armed with strong distal spines, distomcsial spine strong, clearly longer than
others. Carpus and palm with some spines on dorsal side and marginal borders. Fixed finger unarmed exccpl small
distal spine; movable finger with small proximal spine.
Second pereiopod twice carapace length; merus slightly shorter than carapace, about twice propodus length,
with dorsal row of spines, increasing in size distally, ventral border wilh one long distal spine. Carpus with two
well developed distal spines. Propodus with some movable spinules along ventral border. Dactylus slightly shorter
than propodus, with row of movable spinules along ventral margin.
COLOUR. — Holotype : Ground colour of carapace, abdominal segments and pereiopods orange, with some red
spots, more numerous on gastric and branchial regions of carapace and fourth and fifth abdominal segments.
Telson and uropods white. Rostrum and supraocular spines reddish, tips white. Merus of chelipeds with transverse
red bands; carpus and palm red, fingers with proximal and distal parts whitish, median part red. Walking legs with
some transverse red bands, terminal half of dactylus whitish (Fig. 3d).
Paratype (male CL 7.6 mm, Stn 975) with less conspicous red spots on the carapace, distal part of merus,
carpus and proximal part of palm of chelipeds whitish (Fig. 4a).
Remarks. — The new species resembles Munida militaris Henderson. 1885 from Indonesia, Fiji,
New Caledonia and Wallis and Futuna area (Baba & MACPHERSON, 1991; Macpherson, 1994; 1995). Both
species have five spines on the branchial margin, second abdominal segment armed with spines along the anterior
ridge, intestinal region without striae, one transverse stria on the second and third abdominal segments, eyes large,
thoracic sternites without striae and distolateral spine of the basal antennular peduncle longer than the distomesial
spine. However, they can be distinguished by the following characters:
— The carapace is more massive in the new species than in M. militaris. The carapace is always as long as
wide in M. congesta, whereas in M. militaris the carapace is always longer than wide.
— The chelipeds are shorter and more massive in the new species than in M. militaris. The palm of the
chelipeds is more than 2 times longer than high in M. militaris , being less than 2 times in M. congesta.
This difference is quite clear comparing specimens of similar size and sex.
— The fixed finger of the cheliped has one well developed basal spine in M. militaris. This spine is absent in
the new species.
— The colour patterns are different. The carapace and abdominal segments have numerous large red spots in the
new species, these spots are absent in M. militaris. However, as it has been pointed out in the Introduction, the
differences in the colour patterns between species should be considered with caution.
M. congesta is also close to a group of species ( M . cundrostris Henderson, 1885; M. masoae Macpherson,
1995; M. punctata Macpherson, 1997; M. rhodonici Macpherson. 1994; M. rosula Macpherson. 1994 and
M. spissa Macpherson, 1995) from the Western Pacific and the Indian ocean waters, which shares some characters
(e.g. five spines on lateral branchial margins, eyes large, second abdominal segment with a row of spines on
anterior ridge). However, it is easily distinguished, among other characters, by the size of the distal spines of the
basal antennular peduncle. The distolateral spine is longer than the distomesial one in M. congesta , whereas in the
above mentioned species the distal spines are subequal in size.
Distribution. — Vanuatu, between 536-566 and 624-668 m.
Munida elegantissima de Man, 1902
Munida elegantissima - Baba. 1988: 94 (references); 1989: 131. — Macpherson, 1994: 465; 1995: 391.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 966, 128-150 m: 2 6 7.6 and 8.5 mm; 1 ov. 9 10.3 mm;
1 9 8.9 mm; 1 juv. 3.2 mm. — Stn 1077. 180-210 m: 1 ov. 9. 9.3 mm.
Distribution. — The species has previously been cited from the Eastern Indian Ocean, Malay Archipelago,
Indonesia, Philippines, Japan, Western and Eastern Australia, New Caledonia, Bellona Island and Futuna Island,
between 20 and 440 m. The specimens from Vanuatu were collected at 128-210 m.
Source :
AGONONIDA AND MUNIDA FROM VANUATU
419
Munida gordoae Macpherson, 1994
Munida gordoae Macpherson, 1994: 469, fig. 18.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 961, 100-110 m: 1 ov. 9 4.8 mm; 1 9 4.7 mm.
DISTRIBUTION. — New Caledonia, Loyalty Islands, Matthew and Hunter Islands and Chesterfield Islands,
between 80 and 283 in. The present material was collected at 100-1 10 m.
Munida idyia Macpherson, 1994
Munida idyia Macpherson, 1994: 477, fig. 23.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 980, 433-450 m: 1 6 7.2 mm. — Stn 1049, 469-525 m:
1 9 10.4 mm. — Stn 1060, 375-397 m: 1 6 9.8 mm. — Sin 1087, 394-421 m: 1 6 9.1 mm; 3 ov. 9 8.7 to 10.2 mm;
19 11.2 mm. — Stn 1088, 425-455 m: 1 ov. 9 10.2 mm. — Stn 1136, 398-400 m; 1 6 13.2 mm; 2 ov. 9 9.5 and
9.7 mm; 2 9 7.6 and 1 1.3 mm. — Stn 1 137, 360-371 m: 2 6 7.8 and 9.5 mm; 1 9 6.8 mm.
DISTRIBUTION. — Previously known from New Caledonia, in 485 m depth. The specimens from Vanuatu
were caught between 360 and 525 m.
Munida leagora Macpherson. 1994
Munida leagora Macpherson, 1994: 485, figs 26. 76.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 982, 408-410 m: 2 6 9.4 and 9.6 mm. — Stn 1060. 375-
397 m: 1 ov. 9 12.6 mm. — Stn 1067, 344-366 m: 1 ov. 9 10.7 mm; 1 9 6.3 mm. — Stn 1083, 397-439 m: 2 6 6.3
and 12.9 mm; 5 ov. 9 8.3 to 14.2 mm; 2 9 8.6 and 10.1 mm. — Stn 1 138, 462-498 m: 1 ov. 9 12.2 mm.
Distribution. — Previously known from New Caledonia, Loyalty Islands and Chesterfield Islands, between
265 and 580 m. The present material was collected at 344-498 m.
Munida leptitis Macpherson, 1994
Munida leptitis Macpherson, 1994: 487, fig. 27; 1995: 394, fig. 14; 1997: 607.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 978. 408-413 m: 1 ov. 9 4.4 mm. — Stn 983, 475-480 m:
3 6 3.3 to 6.5 mm. — Stn 1 100, 258-265 m: 1 9 4.3 mm.
Distribution. — New Caledonia, Loyalty Islands, Wallis and Futuna Islands and Indonesia between 21 and
440 m. The specimens from Vanuatu were collected at 258-480 m.
Munida leviantennata Baba, 1988
Munida leviantennata Baba, 1988: 111, figs 41, 42; 1994: 12, fig. 5. — Macpherson. 1994: 491; 1995: 395; 1997:
608.
MATERIAL examined. — Vanuatu. Musorstom 8: stn 958, 497-570 m: 1 9 9.6 mm. — Stn 1028, 624-668 m:
1 6 13.2 mm. — Stn 1051, 555-558 m: 1 ov. 9 12.4 mm. — Stn 1073, 630-650 m: 4 6 12.0 to 13.2 mm. —
Stn 1088, 425-455 m: 2 6 9.8 and 13.3 mm. — Stn 1111, 1210-1250 m: 1 9 10.9 mm. — Stn 1137, 360-371 m: 1 6
1 1 .5 mm.
DISTRIBUTION. — Philippines, Indonesia, eastern Australia, New Caledonia, Chesterfield Islands and Wallis
Islands, between 300 and 660 m. The present material was obtained between 360 and 1250 m.
Source : MNHN. Pahs
420
E. MACPHERSON
Fig. 4. — a, Munida congesta sp. nov., male 7.6 mm, paratype, stn 975; b, Munida masi Macpherson, 1994, male
8.8 mm, stn 1001; c, Munida rhodonia Macpherson, 1994, ovigerous female 10.3 mm, stn 1027; cl, Munida
rhodonio Macpherson, 1994, female 9.0 mm, stn 1027; e, Munida rubrodigitalis Baba, 1994, male 6.4 mm, stn 985.
Source : MNHN. Paris
AGONONIDA AND MUNI DA FROM VANUATU
421
Munida masi Macpherson, 1994
Fig. 4 b
Munida masi Macpherson, 1994: 495. fig. 31.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1001, 150-250 m: 1 d 8.8 mm; 1 9 5.8 mm. — Stn 1 135,
282-375 m: 1 d 9.4 mm; 1 ov. 9 7.9 mm. — Stn 1039, 464-472 m: 1 6 7.1 mm.
Remarks. — The species was only known from the holotype (male of CL, 10.6 mm) collected in
New Caledonia, between 250-290 m. The specimens collected in Vanuatu fully agree with the holotype
description.
COLOUR. — Ground colour of carapace and abdominal segments red. Rostrum and supraocular spines red.
Cheiipeds reddish, with some white spots along lateral margins of articles. Walking legs reddish, with some
transverse deep red bands.
Distribution. — Previously known from New Caledonia between 250-290 m. The specimens from Vanuatu
were collected between 150-472 m.
Munida microps Alcock, 1894
Munida microps Alcock, 1894: 326. — Baba, 1988: 122 (references); 1994: 13. — Macpherson, 1994: 496. fig. 32;
1995: 397; 1997: 608.
Material examined. — Vanuatu. MUSORSTOM 8: stn 990. 980-990 m: 4 d 10.4 to 1 1.3 mm; 2 ov. 9 10.7 and
13.6 mm; 1 9 9.7 mm. — Stn 991, 910-936 m: 4 d 6.0 to 16.8 mm; 3 ov. 9 11.5 to 13.7 mm; 1 9 9.3 mm. —
Stn 992, 748-775 m: 12 d 6.5 to 18.8 mm; 2 ov. 9 11.0 and 16.4 mm; 4 9 7.7 to 15.4 mm. — Stn 993, 780-783 m:
Id 1 1.0 mm. — Stn 1007, 720-830 m: 1 d 10.2 mm; I ov. 9 14.4 mm. — Stn 1008, 919-1000 m: 6 d 13.4 to
17.9 mm; 6 ov. 9 11.3 to 16.3 mm. — Stn 1014, 495-498 m: 1 9 14.6 mm. — Stn 1036. 920-950 m: 4 d 6.6 to
12. mm; 2 ov. 9 10.3 and 15.6 mm. — Stn 1037. 1058-1086 m: 8 d 6.7 to 15.2 mm; 1 ov. 9 12.6 mm; 9 9 8.5 to
12.3 mm. — Stn 1075, 944-956 m: 4 d 9.7 to 19.4 mm; 6 ov. 9 13.5 to 15.8 mm; 1 9 8.3 mm. — Stn 1076, 1 100-
1191 m: 26 d 6.8 to 15.6 mm; 1 1 ov. 9 11.0 to 13.7 mm; 10 9 9.0 to 14.1 mm. — Stn 1080, 799-850 m: 23 d 14.2
to 20.2 mm; 2 ov. 9 9.8 and 10.3 mm; 7 9 10.0 to 18.1 mm. — Stn 1081. 908 m: 2d 19.1 and 19.8 mm; 2 ov. 9 13.0
and 16.5 mm. — Stn 1125, 1160-1220 m: 2 d 8.4 and 9.3 mm; 1 9 6.5 mm. — Stn 1126. 1210-1260 m: 7 d 8.6 to
11.0 mm; 1 ov. 9 11.6 mm; 1 9 8.0 mm. — Stn 1129, 1014-1050 m: 1 9 13.3 mm.
DISTRIBUTION. — The species has been previously cited in Arabian Sea, Maldives Islands, Philippines.
Indonesia, southeastern Australia, New Caledonia, Chesterfield Islands, Wallis and Futuna Islands, between 686
and 1240 m. The specimens from Vanuatu were collected at 495-1260 m.
Munida militaris Henderson, 1885
Munida militaris - Baba & Macpherson, 1991: 539, fig. 1 (synonymies and references). — Macpherson, 1994: 496;
1995: 399, fig. 16.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1034. 690-750 m: I d 20.8 mm. — Stn I 1 14. 647 m:
2 ov. 9 12.8 and 17.8 mm.
Distribution, — The species is known from Indonesia, New Caledonia, Fiji and Wallis and Futuna area,
between 183 and 1280 m. The present material was collected at depths between 647 and 750 m.
Munida notata Macpherson, 1994
Munida notata Macpherson, 1994: 500, figs 34,78; 1995: 402.
Source : MNHN. Pans
422
E. MACPHERSON
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 965, 361-377 m: 1 6 8.6 mm; 1 ov. 9 8.3 mm. —
Stn 1017, 294-295 m: 1 6 7.0 mm; 6 ov. 9 6.9 to 9.7 mm. — Stn 1018, 300-301 m; 1 ov. 9 7.8 mm; 2 9 6.4 and
7.0 mm. — Stn 1024, 335-370 m: 2 8 6.8 and 8.4 mm. — Stn 1058, 319 m: 1 6 6.4 mm. — Stn 1065. 360-419 m:
1 9 9.7 mm. — Stn 1091, 344-350 m: 1 9 8.8 mm. — Stn 1097. 281-288 m: 2 6 6.1 and 8.0 mm; 1 ov. 9 7.5 mm.
Distribution. — Previously known from New Caledonia, Loyalty Islands, Chesterfield Islands, Wallis and
Futuna Islands, between 59 and 850 m. The present material was caught at depths between 281 and 419 m.
Munida pagesi Macpherson, 1994
Munida pagesi Macpherson. 1994: 507, fig. 37.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 1047. 486-494 m: 1 6 16.8 mm; 1 ov. 9 14.8 mm.
DISTRIBUTION. — The species has previously been cited in New Caledonia and Loyalty Islands, between 250
and 600 m. The specimens from Vanuatu were obtained at 486-494 m.
Munida rhodonia Macpherson, 1994
Figs 4 c-d
Munida rhodonia Macpherson, 1994: 517, figs 13a, 43, 81.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 975. 536-566 m: 3 6 8.0 to 10.6 mm; 3 ov. 9 8.7 to
12.1 mm; 1 9 12.4 mm. — Stn 986, 602-648 m: 1 6 10.8 mm. — Stn 1027, 550-571 m: 1 6 6.8 mm; 2 ov. 9 10.3
and 12.3 mm; 5 9 8.6 to 12.5 mm. — Stn 1028, 624-668 m: 1 ov. 9 11.2 mm. — Stn 1045, 459-488 m: 1 6 6.7 mm.
— Stn 1047, 486-494 m: 3 6 7.5 to 9.2 mm; 1 ov. 9 12.1 mm; I 9 7.8 mm. — Stn 1049, 469-525 m: 1 6 9.8 mm. —
Stn 1050, 541-577 m: 1 ov. 9 14.3 mm; 1 9 8.3 mm. — Stn 1051. 555-558 m: 1 6 1 1.0 mm; 3 9 7.9 to 13.5 mm. —
Stn 1055, 572-580 m: 2 ov. 9 8.3 and 11.4 mm. — Stn 1073, 630-650 m: 1 6 6.8 mm; 2 ov. 9 12.4 and 13.0 mm. —
Stn 1089, 494-516 m: 4 9 6.2 to 11.0 mm. — Stn 1090. 470-502 m: 1 6 10.4 mm; 1 ov. 9 10.6 mm. — Stn 1 124.
532-599 m: 5 6 7.7 to 10.0 mm; 4 ov. 9 9.3 to 14.2 mm.
Remarks. — The colour pattern of the specimens photographed in Vanuatu generally agrees with the type des¬
cription. However, the female (CL, 9.0 mm) from Stn 1027, have some red spots on gastric, cardiac and branchial
regions and a median white band on the second to fourth abdominal segments. These red spots and white band arc
less conspicuous in an ovigerous female (CL, 10.3 mm) from the same station. Both specimens have the distal
part of the rostrum and supraocular spines white. These spots and bands are absent in the specimens from New
Caledonia. No other morphological differences between New Caledonia and Vanuatu specimens were observed.
Distribution. — Previously cited in New Caledonia, Loyalty Islands and Chesterfield Islands, between 475
and 705 m. The material examined here was collected at 459-668 m.
Munida rogeri Macpherson, 1994
Munida rogeri Macpherson, 1994: 518, fig. 44.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 962, 370-400 m: 1 6 4.6 mm.
Distribution. — Previously known from New Caledonia, Loyalty Islands and Chesterfield Islands, between
245 and 390 m. The material from Vanuatu was collected at 370-400 m.
Munida rosula Macpherson, 1994
Munida rosula Macpherson, 1994: 521, figs 45, 82; 1995: 404.
Source : MNHN. Paris
AG0N0N1DA AND MUNIDA FROM VANUATU
423
Material EXAMINED. — Vanuatu. Musorstom 8: stn 992, 748-775 m: 4 6 8.4 to 14.4 mm; 4 ov. 9 14.4 to
16.0 mm; 6 9 7.3 to 12.3 mm. — Stn 993, 780-783 m: 1 6 13.3 mm. — Stn 996, 764-786 m: 2 6 10.0 and 14.8 mm;
3 9 9.6 to 15.3 mm. — Stn 1007, 720-830 m: 1 9 16.5 mm. — Stn 1035, 765-780 m: 1 d 11.0 mm; 4 ov. 9 14.5 to
17.0 mm; 1 9 10.6 mm. — Stn 1056, 602-620 m: 3 d 12.2 to 15.4 mm. — Stn 1073, 630-650 m: 6 6 8.0 to 14.4 mm;
1 ov. 9 15.7 mm; 1 9 10.4 mm. — Stn 1080, 799-850 m: 3 6 11.7 to 16.8 mm; 3 ov. 9 13.4 to 16.7 mm; 5 9 8.1 to
17.7 mm.
Distribution. — The species has previously been cited in New Caledonia, Loyalty Islands, Chesterfield
Islands and Wallis and Futuna area, between 465 and 860 m. The specimens from Vanuatu were caught at 602-
850 m.
Munida rubrodigitalis Baba, 1994
Fig. 4 e
Munida rubrodigitalis Baba, 1994: 13, fig. 6. — MACPHERSON, 1997: 610.
Munida sp. - MACPHERSON, 1994: 558, figs 13b, 90.
Material examined. — Vanuatu. Musorstom 8; stn 974. 490-520 m; 3 6 9.6 to 13.6 mm; 3 ov. 9 13.2 to
13.4 mm; 1 9 9.3 mm. — Stn 983, 475-480 m: 2 6 10.0 and 11.2 mm; 2 ov. 9 12.9 and 13.0 mm; 1 9 5.9 mm. —
Stn 984, 480-544 m: 3 6 9.5 to 11.1 mm; 3 ov. 9 12.2 to 12.5 mm; 19 11.6 mm. — Stn 985, 536-563 m: 1 6
6.4 mm. — Stn 1047, 486-494 m: 1 6 11.7 mm; 1 ov. 9 11.8 mm; 2 9 10.2 and 12.0 mm. — Stn 1049, 469-525 m:
1 ov. 9 11.5 mm; 1 9 8.8 mm. — Stn 1088, 425-455 m: 3 6 7.8 to 10.2 mm. — Stn 1089, 494-516 m: 1 ov. 9
13.3 mm. — Stn 1090, 470-502 m: 1 ov. 9 14.3 mm.
Remarks. — The specimens from Vanuatu generally agree with the type description and the specimens from
New Caledonia and Indonesia. However, the specimen photografed (male, CL. 6.4 mm, Stn 985) has a
longitudinal median white band from the epigastric region to the fourth abdominal segment. This band is absent in
the New Caledonia material.
Distribution. — The species has previously been cited in Eastern Australia. Indonesia, New Caledonia and
Loyalty Islands, between 285 and 503 m. The specimens from Vanuatu were obtained at 425-563 m.
Munida rufiantennulata Baba, 1969
Munida rufiantennulata Baba, 1969: 23, fig. 7; 1988: 128; 1989: 131. — MACPHERSON, 1994: 523. figs 46, 83; 1997:
610.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 988, 372-466 m: 2 6 4.2 and 8.3 mm.
Distribution. — Previously known from Japan, Philippines, Indonesia. New Caledonia, Loyally Islands,
Chesterfield Islands, Matthew and Hunter Islands, between 205 and 610 m. The material examined here was
collected at 372-466 m.
Munida runcinata Macpherson, 1994
Munida runcinata Macpherson, 1994: 525, fig. 47; 1995: 405, fig. 19.
Material EXAMINED. — Vanuatu. Musorstom 8: stn 963, 400-440 m: 6 6 6.5 to 8.7 mm; 6 ov. 9 7.4 to
9.7 mm. — Stn 978, 408-413 m:2d 7.7 and 8.5 mm; 2 ov. 9 6.1 and 6.8 mm; 1 9 6.6 mm. — Stn 988, 372-466 m:
1 ov. 9 7.8 mm. — Stn 1025, 385-410 m: 9 6 6.7 to 9.5 mm; 3 ov. 9 7.6 to 9.3 mm; 2 9 6.6 and 7.1 mm. —
Stn 1035. 282-375 m: 1 6 6.3 mm; 1 ov. 9 7.8 mm. — Stn 1043, 350-372 m: 1 ov. 9 8.1 mm. — Stn 1065, 360-
419 m: 3 6 7.2 to 8.3 mm; 3 9 5.8 to 8.9 mm. — Stn 1087, 394-421 m: 1 6 6.3 ram; 1 ov. 9 8.7 mm. — Stn 1 135,
282-375 m: 1 6 6.3 mm; 1 ov. 9 8.7 mm. — Stn 1137, 360-371 m: 1 6 10.2 mm; 1 ov. 9 8.2 mm; 3 9 6.6 to 7.7 mm.
Distribution. — Previously cited in New Caledonia, Loyalty Islands, Wallis and Futuna Islands, between
245 and 500 m. The specimens from Vanuatu were collected between 282 and 466 m.
Source :
424
E. MACPHERSON
Munida sao Macpherson, 1994
Munidci sao Macpherson, 1994: 529, fig. 49.
MATERIAL EXAMINED. — Vanuatu. MUSORSTOM 8: stn 971. 250-315 m: 1 ov. 9 7.5 mm. — Stn 1026. 437-
504 m: 1 8 6.3 mm; 3 ov. 9 11.4 to 16.0 mm; 1 9 5.7 mm. — Stn 1061, 458-512 m: 2 8 8.6 and 9.4 mm. —
Stn 1070, 184-190 m: 3 8 5.8 to 9.9 mm; 1 9 7.4 mm. — Stn 1086. 182-215 m: 6 8 6.0 to 7.4 mm; 3 9 4.7 to
7.9 mm. — Stn 1088, 425-455 m: 1 8 14.3 mm; 6 ov. 9 10.0 to 12.6 mm. — Stn 1089. 494-516 m: 1 8 14.2 mm. —
Stn 1095, 304-320 m: 1 ov. 9 7.8 mm. — Stn 1098, 277-285 m: 2 8 5.8 and 9.0 mm; 1 ov. 9 6.3 mm; I 9 6.0 mm.
— Stn 1 107, 397-402 m: 2 8 12.3 and 15.1 mm; 1 ov. 9 1 1.6 mm. — Stn 1 136. 398-400 m: 9 8 8.3 to 15.4 mm; 6 ov.
9 8.8 to 13.0 mm; 1 9 10.4 mm. — Stn 1137, 360-371 m: 6 8 9.3 to 12.1 mm; 2 ov. 9 10.4 and 10.5 mm; 1 9
13.0 mm.
Distribution. — Known from New Caledonia, between 165 and 260 m. The material from Vanuatu was
obtained from 1 82-5 16 m.
Munida sacksi Macpherson, 1993
Munida sacksi Macpherson, 1993: 438. fig. 6; 1994 : 438 (key).
MATERIAL EXAMINED. — Vanuatu. MUSORSTOM 8: stn 1047, 486-494 m: 1 ov. 9 11.5 mm. — Stn 1124. 532-
599 m: 1 9 10.4 mm. — Stn 1 135, 282-375 m: 1 9 5.6 mm.
Distribution. — Previously known from the Philippines and New Caledonia, between 300 and 550 m.
The present material has been collected at 486-599 m.
Munida semoni Ortmann, 1894
Munida semoni - Macpherson & Baba, 1993: 411, fig. 17 (references). — Macpherson, 1994: 530; 1995: 405.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: stn 971, 250-315 m: 1 9 7.8 mm. — Stn 1071, 180-191 m:
1 8 9.3 mm; 1 9 7.6 mm. — Stn 1077, 180-210 m: 2 8 7.4 and 8.9 mm; 1 ov. 9 9.5 mm; 1 9 10.2 mm. — Stn 1078.
194-230 m: 1 8 8.4 mm. — Stn 1102, 208-210 m: 2 8 7.3 and 8.2 mm. — Stn 1118, 191-248 m: 1 9 6.3 mm. —
Sin 1 1 19, 254-300 m: 16 8 5.4 to 8.3 mm; 6 ov. 9 6.1 to 7.6 mm; 1 juv. 3.1 mm.
Distribution. — The species has been cited in Indonesia, New Caledonia and Futuna Island, between 245 and
440 m. The specimens from Vanuatu were caught at 180-315 m.
Munida tuberculata Henderson, 1885
Munida tuberculata - MACPHERSON, 1994: 547, fig. 58 (references); 1995: 408.
Material EXAMINED. — Vanuatu. MUSORSTOM 8: sin 974, 492-520 m: 1 8 3.8 mm. — Stn 985, 536-563 m:
1 9 3.9 mm. — Stn 1027. 550-571 m: 2 8 3.6 and 3.8 mm.
Distribution. — The species has previously been cited in Fiji, New Caldonia, Matthew and Hunter Islands,
Wallis and Futuna area, between 420 and 650 m. The material studied here was collected at 492-571 m.
Munida tyche Macpherson, 1994
Munida tyche Macpherson, 1994: 549, fig. 59; 1995: 408, fig. 22.
MATERIAL EXAMINED. — Vanuatu. MUSORSTOM 8: stn 962, 370-400 m: 1 8 12.3 mm. — Stn 1042, 200-260 m:
3 8 6.3 to 8.5 mm; 2 ov. 9 6.1 and 7.7 mm; 1 9 6.0 mm. — Stn 1131, 140-175 m: 4 8 4.4 to 7.1 mm; 1 ov. 9
Source : MNHN. Pans
AGONONIDA AND MUNI DA FROM VANUATU
425
10. 1 mm; 1 8 5.7 mm; 2 juv. 3.2 and 3.4 mm. — Stn 1 132, 161-182 m: 1 8 6.0 mm. — Stn 1 133, 174-210 m: 1 8
9.2 mm; 1 9 7.2 mm.
Distribution. — Known from New Caledonia, Chesterfield Islands and Futuna Island, between 200 and
440 m. The material from Vanuatu was obtained from 140-400 m.
Munida typhle Macpherson, 1994
Munida typhle Macpherson, 1994: 549, fig. 60.
MATERIAL EXAMINED. — Vanuatu. Musorstom 8: stn 1 1 1 1. 1210-1250 m: 1 ov. $ 1 1.2 mm.
DISTRIBUTION. — Previously known from New Caledonia, between 1395 and 1470 m. The specimen from
Vanuatu was collected at 1210-1250 m.
ACKNOWLEDGEMENTS
I am very grateful to A. CROSNIER and B. Richer de FORGES of ORSTOM for their support and help and for
making this interesting material available to me. Thanks are also due to M.K. WiCKSTEN (Texas A&M
University) and K. Baba (Kumamoto University) for their improvements to the manuscript. The species have
been illustrated by J. MACPHERSON and J.L. MENOU of ORSTOM took the colour photographs.
REFERENCES
ALCOCK. A., 1894. — Natural History notes from H.M. Indian Marine Survey Steamer "Investigator", commander
R.F. Hoskyn, R.N., commanding. Series II, No. 1. On the results of deep-sea dredging during the season 1890-91
(continued). Annals and Magazine of Natural History \ (6) 13: 321-334.
Baba, K., 1969. — Four new genera with their representatives and six new species of the Galatheidae in the collection of
the Zoological Laboratory, Kyushu University, with redefinition of the genus Galathea. OHMU, Occasional Papers of
the Zoological Laboratory of the Faculty • of Agriculture, Kyushu University \ 2(1): 1-32.
Baba, K., 1988. — Chirostylid and Galatheid Crustaceans (Decapoda: Anomura) of the "Albatross" Philippine
Expedition, 1907-1910. Researches on Crustacea, Special Number 2: v + 203 pp.
Baba, K., 1989. — Anomuran Crustaceans obtained by dredging from Oshima Strait, Amami-Oshima of the Ryukyu
Islands. Memoirs of the National Science Museum, Tokyo, 22: 127-134.
Baba, K., 1994. — Deep-sea Galatheid crustaceans (Anomura: Galatheidae) collected by the ’Cidaris I' expedition off
central Queensland. Australia. Memoirs of the Queensland Museum, 35: 1-21.
Baba, K., & Macpherson, E., 1991. — Reexamination of the type material of Munida militaris Henderson. 1885
(Crustacea: Decapoda: Galatheidae), with the selection of a lectotype. Proceedings of the Biological Society of
Washington. 104: 538-544.
Baba, K., & de Saint Laurent, M., 1996. — Crustacea Decapoda : Revision of the genus Bathymunida Balss. 1914, with
six new related genera (Galatheidae). In : A. Crosnier (ed.), Rcsultats des Campagnes Musorstom, VoI. 15. Memoires
du Museum national d'Histoire naturelle, 168: 433-502.
BRUCE, A.J., 1975. — Coral reef shrimps and their colour patterns. Endeavor, 34: 23-27.
Henderson, J. R., 1885. — Diagnoses of the new species of Galatheidea collected during the "Challenger" Expedition.
Annals and Magazine of Natural History , (5) 16: 407-421.
Henderson, J. R., 1888. — Report on the Anomura Collected by H.M.S. Challenger During the Years 1873-76. Report of
Scientific Results of H.M.S. Challenger, Zoology, 27: vi + 221 pp., 21 pis.
Source : MNHN. Paris
426
E. MACPHERSON
KNOWLTON, N., 1986. — Cryptic and sibling species among the decapod Crustacea. Journal of Crustacean Biology . 6:
356-363.
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., 1995. — Crustacea Decapoda : Species of the genera Munida Leach, 1820 and Paramunida Baba, 1988
(Galatheidae) from Wallis and Futuna. In:: A. Crosnier (ed.), Resultats des Campagnes Musorstom, Vol. 15.
Memoires du Museum national d'Histoire naturelle. 168: 387-421.
Macpherson, E., 1997. Crustacea Decapoda: Species of the genera Agononida Baba & de Saint Laurent, 1996 and Munida
Leach, 1820 (Galatheidae) from Karubar Cruise. In: A. Crosnier & P. Bouchet (eds.), Resultats des Campagnes
MUSORSTOM, Volume 16. Memoires du Museum national d'Histoire naturelle, 172: 597-612.
Macpherson, E.. & Baba, K., 1993. — Crustacea Decapoda : Munida japonica Stimpson, 1858, and related species
(Galatheidae). In : A. CROSNIER (ed.). Resultats des Campagnes Musorstom, vol. 10. Memoires du Museum national
d'Histoire naturelle, 156: 381-420.
Macpherson, E., & de Saint Laurent, M., 1991. — Galatheid crustaceans of the genus Munida from French Polynesia.
Bulletin du Museum national d'Histoire naturelle. Paris , (4), sect. A. 13 (3-4): 373-422.
Ortmann, A., 1894. — Crustaceen. In : Semon, R.. Zoologische Forschungsreisen in Australien und deni malayischen
Archipel. Denkschriften der Medizinisch-Naturwissenschaftlichen Gesellschaft zu Jena. 8: 3-80, pis. 1-3.
Rice, A. I., & de Saint Laurent, M., 1986. — The nomenclature and diagnostic characters of four north-eastern Atlantic
species of the genus Munida Leach : M. rugosa (Fabricius). M. tenuimana G.O. Sars, M. intermedia A. Milne Edwards
and Bouvier. and M. sarsi Huus (Crustacea Decapoda Galatheidae). Journal of Natural History, 30: 143-163.
Richer de Forges, B., Faliex, E., & Menou, J.L.. 1995. — La campagne Musorstom 8 dans I'archipel de Vanuatu.
Compte rendu et liste des stations. In:: A. Crosnier (ed.), Resultats des Campagnes Musorstom, vol. 15. Memoires
du Museum national d'Histoire naturelle, 168: 9-32.
Zariquiey Alvarez, R., 1952. — Estudio de las especies europeas del gen. Munida Leach 1818. Eos, 28: 143-231.
Source : MNHN , Paris
ESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS [
Crustacea Decapoda: Revision
of the Family Dynomenidae
Colin L. McLAY
Department of Zoology
University of Canterbury
Christchurch, New Zealand
ABSTRACT
The Dynomenidae are a group of small, uncommon, primitive crabs, which are often associated with corals. They
inhabit depths down to around 500 m. between latitudes 40°N and 40°S. All genera and species are revised and rcdescribed,
and the genus Dynomene Desmarest, 1823 is divided into two additional genera. As a result, there are thirteen known
species belonging to five genera: Dynomene Desmarest. 1823 \D. hispida Guerin-Meneville, 1832, D. prciedator A. Milne
Edwards, 1879. D. pugnatrix de Man, 1889, D. filholi Bouvier, 1894. and D. pilumnoides Alcock, 1900],
Hirsutodynomene gen. nov. [H. spinosa (Rathbun. 1911), and H. Ursula (Stimpson. 1 860)]. Metadynomene gen. nov.
[M. devaneyi (Takeda, 1977), M. tanensis (Yokoya, 1933). and M. crosnieri sp. nov.], Acanthodromia A. Milne
Edwards. 1880 [A. erinacea A. Milne Edwards, 1880, and A. margarita (Alcock, 1899)], and Paradynomene Sakai. 1963
[P. tuberculata Sakai, 1963]. A key is provided to identify these species. In addition nine fossil genera, dating from the
Upper Jurassic, are known: Stephanometopon Bosquet. 1854, Dromiopsis Reuss. 1859, Palaeodromites A. Milne
Edwards, 1865. Cyamocarcinus Bittner. 1883, Graptocarcinus Roemer, 1887. Cyclothyreus Remes, 1895.
Gemmellarocarcinus Checchia-Rispoli. 1905, Glyptodynomene Van Straclen, 1944. Trachynotocarcinus Wright &
Collins, 1972. Some extinct species have also been placed in the genus Dynomene. The definition of the family
Dynomenidae given by Alcock (1901) is updated and expanded in order to allow fossil species to be more accurately
determined. Because of overlap with the Dromiidae, there has been some uncertainty about true family affinities of some
fossils. Although these genera are in need of revision, this is not undertaken in this paper.
The status of Dynomene pilumnoides is established as a valid species. D. pugnatrix brevimana Rathbun. 1911 is
synonymized with D. pugnatrix de Man. 1889, D. granulobata Dai. Yang & Lan. 1981 is a synonym of D. hispida, while
D. sinensis Chen. 1979, D. tenuilobata Dai, Yang & Lan, 1981, and D. huangluensis Dai, Cai & Yang, 1996 are all
synonyms of D. praedator. Dynomenids are reported from Australia for the first time in D. pilumnoides, and
Hirsutodynomene spinosa. The status of Metadynomene tanensis (Yokoya, 1933) is established as a widespread
Pacific species and shown to be part of the fauna of Japan, where it has been confused with D. praedator.
Paradynomene tuberculata , previously known from Japan and New Caledonia, is now recorded from the Gulf of Aden.
Indian Ocean. P. tuberculata as well as D. praedator and H. spinosa, are reported from Guam. The Atlantic Ocean and
the Indo-Pacific share genera of dynomenids but not species. The biogeographic history of dynomenids is interpreted
McLay, C. L., 1999. — Crustacea Decapoda: Revision of the Family Dynomenidae. In : A. Crosnier (ed.), Resultats
des Campagnes MUSORSTOM, Volume 20. Memoires du Museum national d'Histoire naturelle, 180: 427-569. Paris ISBN
2-85653-520-8.
Source : MNHN . Paris
428
C. L. McLAY
in the light of their present distribution and in relation to plate tectonics. Ancestral dynomenids are assumed to have
been tethyan crabs and D. filholi and Acanthodromia erinacea, two insular Atlantic species, are shown to be tethyan
relicts. By contrast. Hirsutodynomene Ursula from the eastern Pacific, seems to be a species of quite recent origin.
In redescribing the species particular attention is paid to some new characters: setae, gills, epipods and gill cleaning
mechanisms, the subchelate structure of the last pereopods and the male pleopods. This work was undertaken using a
scanning electron microscope. Differences in the gross appearance of setae can be used to separate species and there are
substantial differences in setal structure at the microscopic level. The standard branchial formula for dynomenids is shown
to be nineteen gills plus seven epipods. There is little variation in gill numbers but substantial variation in gill shape
between species. Although dynomenid gills are often said to be "transitional" they are arranged as in phyllobranchs but
with the epibranchial part divided into varying numbers of lobes which gives them a trichobranch-like appearance.
Acanthodromia has gills which are almost identical to the phyllobranchs of the Dromiidae but which retain the
"dynomenid notch" on each side which, in cross section, give each gill plate a violin shape. The gill cleaning mechanism
in dynomenids is complex, being carried out by no less than eight appendages (long setae on the posterior margin of the
scaphognathite and the seven epipods) as well as stiff setae on the posterior hypobranchial wall of the gill chamber. In
eubrachyurans only three appendages (maxillipodal epipods) are used.
In dynomenids the last pereopod is very reduced (on average less than one-third the length of the fourth pereopod) and
carried in a horizontal position alongside the posterolateral carapace margin above the base of the preceding pereopod.
They are not. as it has been commonly described, carried subdorsally. Using a scanning electron microscope it was
revealed that this limb is sexually dimorphic: in males the dactyl has the normal shape of a tiny claw, but in females the
dactyl is a flattened plate, bearing five to sixteen spines which are opposable to an extension of the propodus. In both
males and females the propodal extension is armed with spines but in Hirsutodynomene , Metadynomene and
Paradynomene . females have a significantly larger number of spines, which are armed with tiny teeth. Males of three
species have an additional small spine on the outer margin of the dactyl. This is a character, previously only known
amongst the Dromiidae. which suggests that the last pereopod of dynomenids may have evolved from a camouflage-
carrying limb. This limb appears to be vestigial and it is difficult to know what its function may have been amongst the
dynomenid ancestors. However its most likely former role appears to be as a cleaning appendage, but certainly not for
carrying pieces of camouflage as it is found amongst the dromiids and homolids.
All dynomenids, except Acanthodromia, lack an effective abdominal locking mechanism and both sexes have five
pairs of pleopods. The female has vestigial, uniramous first pleopods followed by four pairs of normal biramous
pleopods, while the male has the normal first two pairs of pleopods as well as three pairs of rudimentary pleopods on
segments three to five. These rudimentary pleopods can be uniramous or bifid. In Metadynomene tanensis 17% of females
were gynandromorphs with small male first pleopods but the remaining pleopods were normal.
The diet of dynomenids seems to consist of food obtained by sieving fine sediment or perhaps coral mucus. The
bunches of stiff setae on the inner margins of the cheliped fingers and third maxillipeds are probably used to separate fine
organic lragments. Most of their gut contents are unidentifiable soft organic material along with small amounts of
chopped chitinous fragments perhaps coming from hydroids or other crustaceans. Dynomenids appear to be deposit
feeders.
Dynomenids have a broadcast reproductive strategy, with indirect development, laying small eggs (mean diameter =
0.49 mm) which probably produce planktonic larvae. Dynomenid larvae have never been reported in plankton samples.
Males are on average 19% larger than females which become sexually mature at 5-8 mm CW for small species, or 9-13 mm
CW for large species. Egg numbers increase logarithmically with body size. Given the sister group relationship with
homolodromiids (which have very abbreviated development) it is implied that dynomenids and dromiids evolved from
ancestors which had large eggs and perhaps a brooding strategy. This conclusion is contrary to accepted w'isdom. but it is
the most parsimonious answer. Some dromiids have retained the brooding strategy but others have independently evolved
a broadcast strategy. The evolution of such a strategy in both these families is probably related to their colonization of
the shallow' water habitat. Both dynomenids and dromiids are mostly crabs of the continental shelf w'hereas
homolodromiids are crabs of the continental slope.
Using morphological characters the phylogenetic relationships of the Dynomenidae are examined. Both the
Dynomenidae and the Dromiidae are monophyletic, sharing significant apomorphies. The resemblance of some
dynomenids and dromiids is shown to be the result of convergent evolution within these families. The Homolodromiidae
are also monophyletic but are defined almost exclusively by plesiomorphies. Monophyly of the Dromiacea de Haan,
1833 is supported by morphological characters with the Dynomenidae and Dromiidae together being the sister group of
the Homolodromiidae. The ancestor of these three families was probably a camouflage carrying crab, using both of the
last two pairs of pereopods. A controversial aspect of the sister group relationships of the dromiaceans is the need to
assume that in dynomenids the fourth pereopod has reverted to a locomotory role and the fifth pereopod became a cleaning
limb. Monophyly of the Podotremata Guinot, 1977 is also supported. This analysis suggests that camouflage-carrying
behaviour has evolved independently in the Dromiidae (and probably in the Homolodromiidae) and the Homolidae!
Dromiids carry pieces of sponges or ascidians as well as shells, using the last two pairs of pereopods, while homolids
carry sponges or anemones, using only the last pair of pereopods. The ancestor of the Dromiacea and Archaeobrachyura
was probably an inhabitant of deeper waters and not a camouflage carrying crab.
Source : MNHN. Paris
FAMILY DYNOMENIDAE
429
RESUME
Crustacea Decapoda: Revision de la famille Dynomenidae.
Les Dynomenidae forment un groupe de petits crabes primitifs, pcu communs, souvent associes au corail. Ils habitent
des profondeurs n’excedant pas les 500 m, k des latitudes comprises entre 40°N et 40°S. Tous les genres et toutes les
espEces sont r«5vis6s et redEcrits. Le genre Dynomene Desmarest, 1823, est divisE et deux nouveaux genres sont crEEs.
Finalement, les treize espEces connues sont reparties en cinq genres : Dynomene Desmarest, 1823 [D. hispida Guerin-
Meneville, 1832; D. praedator A. Milne Edwards, 1879; D. pugnatrix de Man, 1889; D. filholi Bouvier. 1894;
D. pilumnoides Alcock, 1900]; Hirsutodynomene gen. nov. [H. spinosa (Rathbun, 1911) et H. Ursula (Stimpson, I860)];
Metadynomene gen. nov. [M. devaneyi (Takeda, 1977); M. tanensis (Yokoya, 1933) et M. crosnieri sp. nov.];
Acanthodromia A. Milne Edwards, 1880 [A. erinacea A. Milne Edwards, 1880 et A. margarita (Alcock. 1899)] et
Paradynomene Sakai, 1963 [P. tuberculata Sakai, 1963]. Une clef permet d'identifier les diverses especes. Neuf genres
fossiles attribues aux Dynomenidae sont connus du Jurassique superieur : Stephanometopon Bosquet, 1854; Dromiopsis
Reuss, 1859; Palaeodromites A. Milne Edwards, 1865; Cyamocarcinus Bittner, 1883; Graptocarcinus Roemer. 1887;
Cyclothyreus Remes, 1895; Gemmellarocarcinus Checchia-Rispoli. 1905; Glyptodynomene Van Straelen, 1944;
Trachynotocarcinus Wright & Collins, 1972. Enfin, quelques espEces Eteintes ont ete rapportees au genre Dynomene.
La definition originale de la famille dcs Dynomenidae, donnee par ALCOCK (1901), est mise a jour et Etendue afin de
pouvoir y accueillir les especes fossiles. Les veritables affinites de certains fossiles sont incertaines et difficiles
& interpreter en raison de confusions possibles avec les Dromiidae. Bien qu'une revision de ces genres fossiles s'avEre
nEcessaire, il n'a pas ete possible de l'entreprendre dans le cadre du present travail.
Dynomene pilumnoides est etablie comme une espece valide. D. pugnatrix brevimana Rathbun. 1911. est mise en
synonymic avec D. pugnatrix de Man. 1889. D. granulobata Dai, Yang & Lan, 1981. est synonyme de D. hispida, tandis
que D. sinense Chen, 1979, D. tenuilobata Dai, Yang & Lan, 1981. et D. huangluensis Dai, Cai & Yang, 1996, sont
synonymes de D. praedator. Pour la premiere fois, des Dynomenidae sont signales d'Australie, & savoir D. pilumnoides et
Hirsutodynomene spinosa. Le statut de Metadynomene tanensis (Yokoya, 1933) est bien etabli : il s'agit d’une espece
largement repandue dans le Pacifique et qui, au Japon, avait EtE confondue avec D. praedator. Paradynomene tuberculata,
auparavant connue du Japon et de Nouvelle-Caledonie, est signalee du golfe d'Aden. P. tuberculata ainsi que D. praedator et
H. spinosa sont signalees de Guam. L'Atlantique et 1'Indo-Pacifique partagent des genres mais non des especes de
Dynomenidae. L’histoire biogeographique des Dynomenidae est tracee a la lumiere de leur distribution actuelle et des
phEnomEnes lies k la tectonique des plaques. Les reprEsentants ancestraux des Dynomenides sont supposes avoir ete
des formes de la Tethys. D. filholi ainsi qu Acanthodromia erinacea, deux especes insulaires de l’Atlantique, apparaissent
comme des reliques tEthysiennes. En revanche, Hirsutodynomene Ursula, du Pacifique oriental, serait une espece d'origine
trEs recente.
Lors de la description des especes, nous avons considere de nouveaux caractEres (soies, branchies, Epipodites, meca-
nisme de nettoyage des branchies, structure subchEliforme du dernier pereiopode, plEopodes males) et complete nos
observations par leur etude en microscopie Electronique a balayage. Visibles k l’ceil nu et au binoculaire, les differences
dans l'apparence des soies, qui peuvent etre utilisees pour sEparer les espEces, se rEvElent k l’Echelle microscopique comme
representant des structures trEs varices et bien distinctes. La formule branchiale standard pour les branchies est de
19 branchies plus sept Epipodites. Il y a peu de variations dans leur nombre; mais de substantielles variations dans leur
forme distinguent les especes. Bien qu'elles soient souvent qualifies comme Etant dun type intermediate, les branchies
dynomeniennes sont disposes comme des phyllobranchies k la difference que la partie epibranchiale est divisEe en
un nombre varie de lobes, ce qui leur donne une apparence de trichobranchies. Acanthodromia a des branchies presque
identiques aux phyllobranchies des Dromiidae mais conservant Yencoche dynomenienne de chaque cote, ce qui, en coupe,
donne k chaque branchie la forme d’un violon. Chez les Dynomenidae le mecanisme de nettoyage des branchies est
complexe, etant realise par non moins de huit appendices (longues soies sur le bord posterieur du scaphognathite et des
sept epipodites) aussi bien que par des soies raides sur la paroi hypobranchiale posterieure de la chambre branchiale. Chez
les Eubrachyura, seuls trois appendices (Epipodites des maxillipEdes) sont utilises.
Chez les Dynomenidae. la demiere paire de pereiopodes est tres reduite (en moyenne inferieure au tiers de la longueur du
quatrieme pErEiopode) et disposEe horizontalement le long du bord postErolatEral de la carapace, au-dessus du pErEiopode
prEcEdent. L’observation en microscopie Electronique a rEvElE que cet appendice est sexuellement dimorphique : chez
le male, le dactyle a la forme normale dune petite pince tandis que, chez la femelle, le dactyle est une piece aplatic qui
porte de cinq ct seize Epines opposables k une expansion du propode. Dans les deux sexes, I'expansion du propode est
armEe d'Epines; mais, dans les genres Hirsutodynomene, Metadynomene et Paradynomene, les femelles ont un nombre
plus ElevE depines, elles-memes armEes de tres petites dents. Le male de trois especes a une Epine supplEmentaire sur le
bord externe du dactyle, ce qui Etait jusqu'alors un caractere connu seulement chez les Dromiidae et suggere que le dernier
pErEiopode des Dynomenidae pourrait provenir de Involution d'un appendice destinE a porter des pieces de camouflage.
Le dernier pErEiopode se prEsente bien comme un appendice vestigial, et il est difficile de savoir quelle fonction il pouvait
assumer chez les Dynomenidae ancestraux. Son role le plus probable serait celui d’un appendice nettoyeur, et certainement
pas celui d’un appendice transportant un objet pour le camouflage, comme c'est le cas chez les Dromiidae et les Homolidae.
Source : MNHN. Paris
430
C. L. McLAY
Tous les Dynomenidae, a l'exception des Acanthodromia, sont dcpourvus d'un appareil de maintien dc I'abdomen. Les
deux sexes portent cinq paires de pleopodes. La femelle possede une premiere paire de pleopodes unirames. vestigiaux,
suivie de quatre paires de pleopodes normalemcnt birames, alors que 1c male a les deux paires normales de pleopodes
sexuels ainsi que trois paires de pleopodes vestigiaux sur les segments 3 a 5. Ces pleopodes vestigiaux peuvent etre
unirames ou bifides. Chez Metadynomene tanensis 17% des femalles sont gynandromorphes, avec de petils premiers
pleopodes de type male tandis que les autres pleopodes sont normaux.
L'alimentation des Dynomenidae parait consister en nourriture obtenue en filtrant les sediments fins ou peut-etre
le mucus des coraux. Les touffes de soies raides sur le bord interne des doigts des chelipedes et sur les troisiemes
maxillipedes sont probablement utilisees pour separer les fragments organiques. L'estomac contient en majorite du
materiel organique mou non identifiable en meme temps que de petites quantity de fragments durs de chitine provenant
probablement d'hydroides ou de crustaces. Les Dynomenidae se presentent comme des detritivores.
Les Dynomenidae ont une reproduction avec dSveloppement non abrege, indirect, et avec de petits ceufs (diametre
moyen = 0.49 mm) qui se d^veloppent certainement en larves planctoniques. De telles larves dynomeniennes n'ont jamais
ete trouvees dans le plancton. Les males sont en moyenne 19% plus grand que les femellcs; ces dernieres deviennent
sexuellement matures entre 5-8 mm de largeur de carapace pour les especes de taille peu elevee, entre 9-13 mm pour les plus
grandes especes. Le nombre des ceufs augmente de fa^on logarithmique avec les dimensions du corps. Etant donne
la relation de groupe-fr&re avec les Homolodromiidae, il est probable que ('ensemble que constituent les Dynomenidae et
les Dromiidae a evolue h partir d’ancetres ayant eu de gros ceufs. Cette conclusion est contraire a ce qui est communement
accepte. mais elle est la r6ponse la plus parcimonieusc. Cet ensemble (Dynomenidae et Dromiidae) a peut-etre eu une
strategic de protection de la ponte. Certains Dromiidae ont conserve une strategic de protection dc la pontc. mais d'autres
ont, independamment. developpe une strategic de dissemination. L'dvolution d'une telle strategic chez ces deux families
est probablement liee a leur colonisation dun habitat dans des caux peu profondes. Aussi bien les Dynomenidae que
les Dromiidae sont essenticllement des crabes de la plate-forme continentale, tandis que les Homolodromiidae sont
des crabes du talus continental.
Sur la base des caract£res morphologiques, les relations phylog£n6tiques des Dynomenidae sont analyses.
Dynomenidae et Dromiidae sont monophyletiques. partageant des apomorphies signi ficati ves. La ressemblance de
certains Dynomenidae avec les Dromiidae serait le resultat d'une evolution convergente h 1’interieur de ces families. Les
Homolodromiidae sont egalement monophyletiques mais ils sont presque exclusivement definis par des plesiomorphies.
La monophylie des Dromiacea de Haan, 1833, est supportee par des caractdres morphologiques, les Dynomenidae et
Dromiidae formant, ensemble, le groupe-frere des Homolodromiidae. L'ancetre de ces trois families etait probablement un
crabe qui se camouflait en utilisant ses deux dernieres paires de panes. Les relations de groupe-frere pour les families de
Dromiacea n^cessitent d’admettre une reversion pour la quatrieme paire de panes des Dynomenidae puisqu'elle a retrouve un
role locomoteur, tandis que la cinqui&me paire devenait un appendice nettoyeur. La monophylie des Podotremata Guinot,
1977, est corrobor^e. Cette analyse suggere que le comportement de camouflage par maintien d'un objet au-dessus du corps
a evolut§ independamment chez les Dromiidae (et probablement chez les Homolodromiidae) et chez les Homolidae. Les
Dromiidae portent des Sponges ou des ascidies ainsi que des coquilles, en se servant de leurs deux dernieres paires de
pereiopodes, tandis que les Homolidae transportent des eponges et des anemones, en se servant seulement de leur derniere
paire de pereiopodes. L'ancetre des Dromiacea et des Archaeobrachyoures <5tait probablement un habitant des eaux
profondes et n'dtait pas un crabe se camouflant grace au transport d'un objet au-dessus du corps.
TABLE OF CONTENTS
Introduction .
History of the DYNOMENIDAE .
Fossil DYNOMENIDAE .
Material examined .
Terminology and Presentation .
Morphology of the DYNOMENIDAE .
Carapace .
Setae .
Antennules, Antennae and Orbits .
Mouthparts . .
Gills and Epipods .
Pereopods .
Female sternal Sutures .
431
432
434
435
435
439
439
441
431
431
443
449
454
Source : MNHN, Paris
FAMILY DYNOMENIDAE
431
Abdomen . 455
Pleopods . 456
Uropods . 462
Phylogenetic Relationships . 462
Family DYNOMENIDAE . 467
Key to the species of the family Dynomenidae . 469
Genus DYNOMENE Desmarest, 1823 . 471
Dynomene hispida Guerin-Meneville, 1832 . 473
Dynomene praedator A. Milne Edwards, 1879 . 481
Dynomene filholi Bouvier, 1894 . 488
Dynomene pilumnoides Alcock, 1900 . 494
Dynomene pugnatrix de Man, 1889 . 500
Genus HIRSUTODYNOMENE gen. nov . 504
Hirsutodynomene spinosa (Rathbun, 1911) . 505
Hirsutodynomene Ursula (S timpson, 1860) . 510
Genus METADYNOMENE gen. nov . 516
Metadynomene devaneyi (Takeda, 1977) . 517
Metadynomene tanensis (Yokoya, 1933) . 521
Metadynomene crosnieri sp. nov . 530
Genus ACANTHODROMIA A. Milne Edwards, 1880 . 534
Acanthodromia erinacea Milne Edwards, 1880 . 535
Acanthodromia margarita (Alcock, 1899) . 539
Genus PARADYNOMENE Sakai, 1963 . 542
Paradynomene tuberculata Sakai, 1963 . 543
Discussion . 549
Habitat . 549
Diet . 549
Reproductive Strategy . 550
Depth Distribution . 550
Biogeography . 552
ACKNOWLEDGEMENTS . 556
References . 557
Index . 567
INTRODUCTION
Dynomenids are a small group of uncommon primitive crabs living in tropical and warm parts of the Atlantic,
Indian and Pacific Oceans. The shallow water species are often associated with reef-building corals, but other
species live in deeper waters of the continental shelf. The maximum depth record for a dynomenid is 540 m (for
Acanthodromia erinacea A. Milne Edwards, 1880). The first species to be described was Dynomene hispida Guerin-
Meneville, 1832 which was based on a specimen collected from Mauritius, in the Indian Ocean. The family name,
Dynomenidae Ortmann, 1892, was not erected until much later, but the type species for the family is D. hispida.
Until now the family has included three genera: Dynomene Desmarest, 1823, Acanthodromia A. Milne Edwards,
1880, and Paradynomene Sakai, 1963. By far the majority of species have been assigned to the genus Dynomene.
In this paper I describe one new dynomenid species, redescribe all of the previously known species, and provide
a key for their identification. This work is based on the examination of almost 600 specimens. It began with
the collections from New Caledonia, Philippines, Indonesia and Madagascar, held by the Museum national
Source :
432
C. L. McLAY
d'Histoire naturelle, Paris, but also included specimens from most of the world's major museums. In the literature,
dynomenids have been reported from coastal areas bordering all tropical seas except for Australia. In this paper
I provide the first records of these crabs from Australian waters. Also many new records for the Pacific islands are
included.
Besides the customary features used to describe these crabs, I also include some details of their structure that can
only be determined using a scanning electron microscope. This has been particularly helpful in ascertaining the
nature of the subchelate mechanism of the reduced fifth pereopods and has revealed that in dynomenids these limbs
are sexually dimorphic. The scanning electron microscope has also been very helpful in elucidating the fine
structure of setae, gills and male pleopods. Some information about the diet and reproductive strategy is also
included.
Despite their occurrence in shallow tropical waters the larval stages of dynomenids have never been collected in
the field. The only information comes from larvae dissected from advanced stage eggs of Acanthodromia erinacea
(see RICE, 1981). The lack of information about larval development, which could be useful in aiding the determi¬
nation of phylogenetic relationships, is a major gap in our knowledge of this family.
This work follows on from a study of the Dromiidae (McLay, 1993) which are regarded as the sister group of
the Dynomenidae. With the recent completion of a major review of the Homolodromiidae (GuiNOT, 1995)
the opportunity is taken to explore the phylogenetic relationships between all three families of the Dromiacea
sensu DE Haan.
HISTORY OF THE DYNOMENIDAE
The following is a chronological list of recent genera and species (where appropriate the valid name is shown in
brackets):
Genus DYNOMENE Desmarest, 1823
D. hispida Guerin-Meneville, 1832 (type species of the genus).
D. latreillii Eydoux & Souleyet, 1842 [= D. hispida Guerin-Meneville, 1832].
D. Ursula Stimpson, 1860 [= Hirsutodynomene Ursula (Stimpson, I860)].
D. praedator A. Milne Edwards, 1879.
D. pugnatrix de Man, 1889.
D. filholi Bouvier, 1894.
D. margarita Alcock, 1899 [= Acanthodromia margarita (Alcock, 1899)].
D. pilumnoides Alcock, 1900.
D. platyarthrodes Stebbing, 1905 [= Speodromia platyarthrodes (Stebbing, 1905)].
D. pugnatrix brevimana Rathbun, 1911 [= D. pugnatrix de Man. 1889].
D. spinosa Rathbun, 1911 [= Hirsutodynomene spinosa (Rathbun, 191 1)].
D. actaeiformis (Stebbing, 1921) [= D. pilumnoides Alcock, 1900].
D. tanensis Yokoya, 1933 [= Metadynomene tanensis (Yokoya, 1933)].
D. devaneyi Takeda, 1977 [= Metadynomene devaneyi (Takeda, 1977)].
D. sinensis Chen, 1979 [= D. praedator A. Milne Edwards, 1879].
D. granulohata Dai & Yang, 1981 [= D. hispida Guerin-Meneville, 1832].
D. tenuilobata Dai & Lan, 1981 [= D. praedator A. Milne Edwards, 1879].
D. huangluensis Dai, Cai & Yang, 1996 [= D. praedator A. Milne Edwards, 1879].
Genus ACANTHODROMIA A. Milne Edwards, 1880
A. erinacea A. Milne Edwards, 1880 (type species of the genus).
A. margarita (Alcock, 1899).
Genus MAXILLOTHRIX Stebbing, 1921
Maxillothrix actaeiformis Stebbing, 1921 (type and only species of the genus) [= Dynomene
pilumnoides Alcock, 1900].
Source : MNHN. Paris
FAMILY DYNOMENIDAE
433
Genus PARADYNOMENE Sakai, 1963
P. tuberculata Sakai, 1963 (type and only species of the genus).
Genus HIRSUTODYNOMENE gen. nov.
H. spinosa (Rathbun, 1911) (type species of the genus).
//. Ursula (S timpson, 1860).
Genus METADYNOMENE gen. nov.
M. devaneyi (Takeda, 1977) (type species of the genus).
M. tanensis (Yokoya, 1933).
M. crosnieri sp. nov.
The following species have been described and at least initially assigned to the genus Dynomene Desmarest,
1823. The first dynomcnid to be described was Dynomene hispida Guerin-Mcneville. 1832 based on a specimen
collected from Mauritius Id, east of Madagascar. The second species to be described was D. latreillii Eydoux &
Souleyet, 1842 from a specimen collected from Hawaii during the voyage of the " Bonite ". A. Milne Edwards
(1879) and later Peyrot-Clausade & Serene (1976) showed that the preceding species was in fact a synonym of
D. hispida. Some years later, the third species, D. Ursula Stimpson, 1860, was collected from Cape San Lucas,
Baja California, by John Xantus. The fourth species, D. praedator A. Milne Edwards, 1879, was discovered at
New Caledonia by M. Balansa. The fifth species, D. pugnatrix de Man. 1889, also came from Mauritius.
The only species of Dynomene known from the Atlantic, D. filholi Bouvier, 1894, was collected from the Cape
Verde Ids during the voyages of the "Talisman" and " Travailleur ". The seventh species was D. margarita Alcock.
1899, collected from the Andaman Sea during deep-sea surveys by the " Investigator ". The eighth species,
D. pilumnoides Alcock, 1900 was first collected from the Laccadive Archipelago, India. The next name added to
Dynomene was the sub-species D. pugnatrix brevimana Rathbun, 1911, from Providence Id. followed by
D. spinosa Rathbun, 1911 from Coetivy, Seychelles. Both of these specimens were collected during the Percy
Sladen Trust Expedition to the Indian Ocean, 1905. D. spinosa is the last species of this genus whose type locality
is in the Indian Ocean. Clearly collecting in the Indian Ocean area played an important role in the early
development of knowledge about this family of crabs, contributing ten of the eleven known species.
The second phase in the development of our understanding of the dynomcnids resulted from investigations in the
greater Pacific Ocean. The remaining species have type localities in the Pacific Ocean: Dynomene tanensis Yokoya,
1933, came from Tanegasima Id. Japan, and D. devaneyi Takeda, 1977, was collected by the submersible, "Star ll".
from Hawaii. The last four species have only recently been described from the coasts of China or
Taiwan: D. sinensis Chen, 1979, D. granulobata Dai & Yang, 1981. D. tenuilobata Dai & Lan. 1981, and
D. huangluensis Dai, Cai & Yang, 1996. Thus there have been 15 species or sub-species of dynomcnid crabs
assigned to the genus Dynomene. In addition two species of the Dromiidae have been erroneously placed in this
genus: Dynomene depressa Brocchi, 1875 (= Dromidia spongiosa Stimpson. 1858). and Dynomene platyarthrodes
Stebbing, 1905 [= Speodromia platyarthrodes (Stebbing, 1905)).
The second dynomenid genus to be erected was Acanthodromia A. Milne Edwards, 1880. for A. erinacea Milne
Edwards, 1880, collected from Guadeloupe, Atlantic Ocean, by the United States coast survey steamer "Blake".
Probably through an oversight when creating the family name Dynomenidac. Ortmann (1892) did not include this
genus in his new family. A second species, A. margarita (Alcock, 1899), was collected from the Andaman Sea.
Indian Ocean, by the " Investigator ", which was initially placed in the genus Dynomene.
The third genus Maxillothrix Stebbing, 1921, was initially placed in the family Xanthidae. But the only species
to be assigned to this genus, M. actaeiformis Stebbing. 1921. based on a specimen from the Natal coast, was later
found to be a synonym of Dynomene pilumnoides Alcock, 1900. Two authors, Serene (1968) and Takeda
(1977) used the name Dynomene actaeiformis (Stebbing, 1921) in faunal lists.
Finally, the monospecific Parody nomene Sakai, 1963 was created for P. tuberculata Sakai, 1963 from Sagami
Bay, Japan.
Thus prior to the commencement of this work, the family Dynomenidae contained a total of 16 species
belonging to 3 genera. Herein I synonymize all four of the recently described Dynomene species from China and
Source :
434
C. L. McLAY
Taiwan, and create two new genera for some of the species of Dynomene : Hirsutodynomene gen. nov. for
D. spinosa and D. Ursula , and Metadynomene gen. nov. for D. devaneyi, D. tanensis and a new species,
M. crosnieri sp. nov., from the Glorieuses Ids, north of Madagascar. In recent years new collections have extended
the ranges of known species without adding greatly to the number of species. The net result of all these changes is
that the family Dynomenidae now contains five genera and thirteen species.
FOSSIL DYNOMENIDAE
Dynomenids have an extensive fossil record, dating from the Upper Jurassic (GLAESSNER, 1980), and there has
been much discussion about the placement of genera in this family or in other families. The fossil record does not
provide a clear answer to the question about the relationships of these crabs. Some of the difficulties arise because,
while it may be easy to define the essential features of extant species, which justify their inclusion in the family, it
is not so easy when it comes to fossils. Here, we have only carapace shape, development of teeth, and a few
grooves incised on the dorsal carapace surface. For fossils, the definition of the family has to be based on a different
and very limited set of characters and it is likely that the range of fossil species included is much wider than for
modern species. For the extant species most are relatively small crabs whereas many of the fossil specimens arc
quite large and probably do not belong in the Dynomenidae. There is no reason to think that there has been a
decrease in crab size during their evolution. I have examined some of the fossils which have been assigned to this
family and have attempted to include a set of carapace characters which both modern and fossil species share.
The following chronological list of fossil genera is based on Balss (1957: 1606) and Glaessner (1969), but
modified after Wright & Collins (1972) and Collins etal. (1995):
DYNOMENE Desmarest, 1823 (type species D. hispida Gucrin-Meneville, 1832. by subsequent monotypy
of Guerin-Meneville, 1832).
STEPHANOMETOPON Bosquet. 1854 (type species Stephanometopon granulatum Bosquet. 1854 by
monotypy).
DROMIOPSIS Rcuss, 1859 (type species Brachyu rites rugosus von Schlotheim, 1820, by subsequent
designation of BEURLEN, 1928).
PALAEODROMITES A. Milne Edwards, 1865 (type species P. octodentatus A. Milne Edwards, 1865, by
monotypy).
CYAMOCARCINUS Bittner, 1883 (type species C. angustifrons Bittner, 1883, by monotypy).
GRAPTOCARC/NUS Roemer, 1887 (type species G. texanus Roemer, 1887, by monotypy).
CYCLOTHY REUS Remes, 1895 (type species C. strambergensis Rcmes, 1895, by subsequent designation
Of BEURLEN, 1928).
GEMMELLAROCARCINUS Checchia-Rispoli, 1905 (type species G. loerentheyi Checchia-Rispoli,
1905, by monotypy).
GLYPTODYNOMENE Van Straelen, 1944 (type species G. alsasuensis Van Straelen, 1944, by
monotypy).
TRACHY NOTOCARCINUS Wright & Collins, 1972 (type species Trachynotus sulcatus Bell, 1863, by
monotypy).
There are a total of nine extinct fossil genera. Only one extant genus, Dynomene has fossil representatives.
With the revision of this genus, undertaken herein, it is likely that some of the fossil species assigned to
Dynomene will be able to be placed in the new genera. It is beyond the scope of the present report to examine the
validity of all the fossil species and until this is done the fossil record has only limited value in helping to reveal
the phylogenetic relationships of the dynomenids.
Source :
FAMILY DYNOMENIDAE
435
MATERIAL EXAMINED
The following abbreviations are used for material examined at or borrowed from museum collections:
AMS - Australian Museum, Sydney.
ANSP - Philadelphia Academy of Natural Sciences.
BMNH - Natural History Museum, London.
BPBM - Bernice P. Bishop Museum, Honolulu.
LACM - Natural History Museum, Los Angeles County.
MCZ - Museum of Comparative Zoology, Harvard University.
MNHN - Museum national d’Histoire naturelle, Paris.
MZUF - Museo Zoologico de "La Specola", Firenze, Italy.
NTOU - National Taiwan Ocean University, Keelung, Taiwan.
QM - Queensland Museum, Brisbane.
RMNH - Nationaal Natuurhistorisch Museum. Leiden (formerly Rijksmuseum van Natuurlijke Historic).
SMF - Natur-Museum und Forschung Institut Senckenberg, Frankfurt.
UGM - University of Guam.
USNM - United States National Museum, Smithsonian Institution, Washington.
WAM - Western Australian Museum. Perth.
ZMB - Museum fur Naturkunde, Humboldt Universitat, Berlin.
ZMUC - Zoologisk Museum, Copenhagen.
ZRC - Zoological Reference Collection. Department of Zoology, National University of Singapore.
When in the lists of material examined, no place of deposit is mentioned, this means that the material is at the
MNHN.
TERMINOLOGY AND PRESENTATION
Measurements were made, using vernier calipers under a binocular microscope, to an accuracy of 0.1 mm.
Measurements of chelipeds and abdomen used for determining relative growth in Metadynomene tanensis were made
as follows: cheliped propodus length was measured along the inferior margin from the joint with the carpus to the
lip of the fixed finger; cheliped propodus depth was measured from the superior to the inferior margin at the w idest
point; abdomen width was measured as the greatest width of the penultimate segment. To determine the relative
sizes of the articles of the last two pairs of pereopods their lengths w'erc measured along the superior margin of each
limb. The length of each article was divided by the total length and converted to a percentage.
Carapace dimensions are given as carapace width (CW) x carapace length (CL) e.g. 1 cJ 9.8 x 8.0 mm.
Carapace width includes any anterolateral teeth and was measured across the widest point. Carapace length includes
any rostral teeth and was measured to the posterior margin in the mid-line.
The description of each species follows the format: carapace shape and ornamentation, tomentum, anterolateral
margin, frontal margin and orbital region including antennule and antenna, subhepatic area, third maxill ipeds and
female sternal sutures 7/8. This is followed by the gill formula and shape of the gill plates. The cheliped is dealt
with separately, followed by the first three pairs of walking legs, and then the reduced last pair of legs. Finally, the
abdomen size and shape, and the five pairs of male and female pleopods.
The tomentum is an important feature of these crabs and is described in terms of the length of the setae, their
density, and their distribution, especially on the surface of the carapace. Setal size is expressed either as the length
ratio of long setae/short setae, or as a ratio of CW. Setal density is expressed in terms of the degree to which the
carapace surface is obscured in dorsal view. Setae were investigated using a scanning electron microscope and the
results arc dealt with in terms of setae shape and the distribution of setules along the shaft.
The major carapace grooves are named according to Glaessner (1969) and are treated as follows: frontal
groove, cervical groove, cardiac groove, and pleural suture.
Source :
436
C. L. McLAY
frontal groove
supraorbital notch
postorbital corner
anterolateral teeth
cervical groove
submedial pit
branchial area
posterolateral tooth
crescent-shaped cardiac groove
branchial groove
posterolateral border
frontal groove
2nd article
1st article of antennule
.v.V'*';
antero
lateral
teeth
epistome
subhepatic
area
4th article
3rd article
2nd article
1 st article of antenna
exopod
superior margin
dactyl
carpus
propodus
carpal spur
merus
spines
on inner
margin
propodus
Fig. 1. — Selected figures illustrating the terminology used to describe crabs of the family Dynomenidae : a-b. e based on
Dynomene hispida , c-d based on D. praedator : a, dorsal view of right half of carapace; b. ventral view of right orbital
area; c, outer face of right cheliped; d. dorsal view of right chelipcd; e, posterior view of terminal articles of right
fourth pereopod.
Source : MNHN, Paris
FAMILY DYNOMENIDAE
437
sperm aperture
propodal spines
dactyl
lateral dactyl spine
channel carrying
second pleopod
endopod
propodus
propodal extension
propodal spines
basal article
dactyl spines
exopod
telson
propodus
dactyl
sixth abdominal segment
Fig. 2. — Selected figures illustrating the terminology used to describe crabs of the family Dynomenidac : a based on
Dynomene hispid a, b based on D. filholi\ c-e based on Meicidynomene tanensis , f-h based on M. devaneyi (reprinted
from Takeda, 1977, Pacific Science, vol. 31 (1) by permission of the publisher); i based on D. praedator : a. ventral
view of telson and terminal segments of female abdomen; b. ventral view of telson and terminal segments of male
abdomen; c, lateral view of tip of male fifth pereopod; d. apical view of lip of male fifth pereopod; e. lateral view of
tip of female fifth pereopod; f, posterior view of male left first pleopod; g, anterior view of tip of male left first
pleopod; h. posterior view of male left second pleopod; i. tip of second pleopod.
Source : MNHN. Paris
438
C. L. McLAY
Normally, the anterolateral margin begins in close proximity to, and at the same level as, the postorbital
corner. Usually the anterolateral margin is clearly defined and it may be adorned with up to five well developed teeth
or only a few small irregular granules. For the separation of Dynomene hispida from D. praedator it is necessary to
distinguish "teeth" from "granules". Teeth have a broad base which narrows apically into a clearly defined acute or
subacute tip which may be nacreous, and a different colour from the carapace itself (as in D. hispida), while
granules do not narrow apically and lack the clearly defined tip, instead terminating bluntly (as in D. praedator). The
teeth may be equidistantly spaced or irregular but the end of the anterolateral margin is usually marked by the last
tooth and the remaining posterolateral carapace margin angles towards the posterior margin.
The frontal margin is V-shaped, continuous, except for a small supraorbital notch followed by several small
spines near the postorbital corner and further spines on the suborbital margin. The medial end of the suborbital
margin is often terminated by a stout spine which is visible dorsally after setae have been removed. The orbits arc
normally clearly exposed dorsally.
The gills arc treated as follows: numbers of pleurobranchs, arthrobranchs, podobranchs and epipods are given for
each thoracic segment and are summarized in the gill formula "no. gills + no. epipods" on each side of the thorax.
Dynomenid gills are essentially phyllobranchiatc with additional dorsal lobes: each plate consists of an anterior
(smaller) and posterior (larger) part joined about the central axis, which contains the afferent and efferent vessels,
and separated by varying numbers of lobes arranged along the dorsal margin. Gill plates show a wide variation in
shape even between species of the same genus. For most species I have been able to study the gill structure using a
scanning electron microscope to capture cross-sectional views. The gills selected for this work were either
pleurobranchs or arthrobranchs taken from the pereopod two or three. Since gill shape varies along the axis of each
gill (plates tapering off to nothing at each end), I chose to study the gill plates from about halfway along each gill
in order to obtain a standardized cross section that could be compared between species.
The first article of the antennule is large, filling a large part of the ventral orbital region, and the antennular
flagellum is tucked in under the supraorbital margin at the base of the eyestalk. The four articles of the antenna,
which are all mobile, are assumed to correspond to coxa, fused basis-ischium, merus and carpus. The excretory or¬
gan opens into a beak-like structure on the medial margin of the first article or coxa. In most genera the urinal
opening is clearly visible in ventral view, but in Acanthodromia the antennal coxa is not beaked, and the urinal
opening is on the medial margin, concealed against the base of the antennule. The distolateral corner of the second
article has a fixed exopod which curves over the base of the eyestalk. The length of the antennal flagella is ex¬
pressed as a proportion of the CW. Thus the orbit is defined above by the carapace margin, which continues around
the postorbital corner to form part of the suborbital margin, the rest being formed by the distal margins of the sec¬
ond antennal and the first antennular articles, both of which are moveable. The left and right orbits are separated by
the epistome which is firmly joined to the carapace above. The eyes can be wholly withdrawn inside the orbits.
The mouthparts are not dealt with in any detail except for the position of the bases of the third maxillipeds and
the number and arrangement of teeth on the crista dentata. Female sternal sutures 7/8 end wide apart on low
tubercles behind the bases of the second walking legs and show little variation between species.
The pereopods fall naturally into three groups: firstly, the chelipeds which are used for feeding, secondly, the
first three pairs of legs which are used for walking, and thirdly, the last pair of legs which are so reduced as to be
almost vestigial. Proceeding distally, the shape and disposition of granules and spines are described, with particular
attention paid to the ornamentation of the outer (or exterior) faces and the angular margins of each cheliped article.
The inner superior border of the carpus usually has a well developed "spur", more prominent in males, which
contributes to the unusual shape of the carpal article in dynomenids. Cheliped fingers are usually stout, curved,
hollowed out internally and bearing weakly developed teeth on the outer margins and tips. The space between the
fingers is usually filled with long coarse bunches of obliquely angled setae.
Ihe first three pairs of walking legs usually decrease in length posteriorly. The arrangement of spines on each
article, from merus to propodus, are described, concentrating on the superior border or dorsal surface of each article.
On the dactyl particular attention is paid to the small spines on the inferior margin. Presumably, these small spines
are used to provide some grip on the substrate which is often dead coral. The ratio of the length (including any
spines) of the second walking leg (i.e. the third pereopod) merus to its width and to the CL are given. This
character is useful for separating some species.
Source : MNHN. Paris
FAMILY DYNOMENIDAE
439
The size of the last pair of legs is given in terms of how far it extends along the meral article of the preceding
limb (i.e. the fourth pereopod). This leg is subchelate but the mechanism appears to be obsolete because it is
incapable of grasping anything. The subchelate structure is sexually dimorphic and detailed comparison of male and
female limbs has been made using the scanning electron microscope.
I treat the abdomen as consisting of six segments and a telson. All abdominal segments bear appendages and are
freely moveable: their shape and surface are described beginning with the most anterior segment. In dynomenids,
both males and females have five pairs of pleopods on the abdomen. The first pleopod is vestigial in the female
while the last three are rudimentary in the male. Uniramous uropods arc inserted at the posterior border of the last
abdominal segment and just in front of the telson. In dynomenids the uropod plates are large (relative to other
dromiaceans), and their size is assessed in terms of what proportion of the margin of the last abdominal segment is
excluded from the lateral margin. This proportion is a sexually dimorphic feature, larger in females than in males.
Width of the telson is measured across the widest part at its base and the length is the maximum length measured
along the mid-line.
MORPHOLOGY OF THE DYNOMENIDAE
This section includes comparison of the major morphological features of dynomenids and a discussion of them
in relation to the other podotremes. Where appropriate characters are indicated to be either plesiomorphic or
apomorphic.
CARAPACE
Carapace width is distinctly greater than length (ratio 1.2 to 1.3) in all species of Dynomene and
Hirsutodynomene , while in Metadynomene width is approximately equal to length and in Acanthodromia and
Paradynomene width is distinctly less (ratio 0.9 to 0.95) than length. In most species the lateral carapace margin is
clearly defined and usually bears teeth. The exception is Acanthodromia where the anterolateral margin is poorly
defined. The number of anterolateral teeth ranges from none, in Acanthodromia , D. praedator and M. devaneyi to six
irregular teeth in P. tuberculata. Most dynomenids have four small anterolateral teeth.
In most dynomenids the carapace surface is smooth (or only minutely granulated) and gently undulating, but in
some species there are spines or areolae. All the Dynomene species have a smooth carapace, species of
Hirsutodynomene and Acanthodromia are spiny to varying degrees, while in Paradynomene the carapace is strongly
areolate and granular.
The frontal margin in most dynomenids is without teeth. The margin is V-shaped, centered on the epistome,
sweeping backwards and laterally above the eyes. The only exception is found in Paradynomene tuberculata which
has a tri-dentate rostrum resembling some dromiid crabs (for further discussion see below under P. tuberculata).
A short frontal groove extends posteriorly from above the epistome, separating firstly, the left and right
supraorbital margins, and secondly, a pair of rounded protuberances, whereupon it separates into two slightly
divergent grooves which dwindle out posteriorly. Between these divergent grooves, there may be an elongate medial
ridge. Further back are found two laterally-directed grooves: the first groove (cervical) arises from a small submedial
pit and runs anterolaterally on to the branchial region where it may be joined by one of the divergent branches of
the frontal groove. The second groove extends across the mid-line, initially running almost directly lateral but
splits into an anterior branch which follows the first groove for a short distance, while the second branch curves
posterolaterally bordering the anterior cardiac region. In effect the groove crossing the mid-line connects two
crescent-shaped grooves with the second branch joining the branchial groove (if present) which runs towards the
lateral carapace margin meeting it just in front of the last anterolateral tooth. The posterior cardiac region may or
may not be well defined. The other grooves which may be evident are to be found under the anterolateral margin of
the carapace: the pleural suture arises near the base of the antenna, curving around under the branchial region,
giving off a short cervical groove which ascends towards the base of the first anterolateral tooth, and then running
towards the base of the last anterolateral tooth where it may meet the branchial groove if it is evident.
Source :
440
C. L. McLAY
Fig. 3. — a , Dynomene hispida Guerin-Meneville, 1832, 6 11.6 x 9.2 mm. Somalia. Gesira, stn 12. intertidal coral
(MZUF): setae on right posterolateral corner of carapace. — b. Dynomene praedator A. Milne Edwards. 1879. 6 9.6 x
7.8 mm. Somalia. Gesira. stn 19. intertidal coral: setae on left posterolateral corner of carapace. — c-d, Dynomene
pilumnoides Alcock. 1900. 9 12.8 x 10.3 mm. New Caledonia. Volsmar, stn DW 7. 400 m: c. short setae from
left posterolateral corner of carapace; d. setules on long setae from left posterolateral corner of carapace.
— e. Dynomene filholi Bouvier. 1894. 9 10.0 x 8.7 mm. Cape Verde Id, Cancap, stn 7.125, 85-130 m: short setae
from right posterolateral corner of carapace. — f, Hirsutodynomene spinosa (Rathbun. 1911), 6 14.2 x 10.8 mm.
Cocos Keeling Ids. 0-37 m (WAM 139-94): short setae from left posterolateral corner of carapace. (All pictures taken
with scanning electron microscope.)
Source : MNHN. Paris
FAMILY DYNOMENIDAE
441
Compared to homolodromiids the dynomenids have a well developed and calcified carapace on which the lateral
margins are well defined and grooves are often evident. In this respect their carapace is very similar to that found
amongst dromiids and this overlap has lead to difficulties in assigning fossil material to the correct family.
SETAE (Figs 3 a-f, 4 a-f)
All dynomenids have setae of two sizes: short and long. Most short setae are at least slightly curved near the
tip, but in some species they arc bent almost at right angles. The long setae can also be curved and assume
various shapes, but they are never consistently bent at right angles. The short setae may be sparse or dense enough
to completely obscure the body surface. In some species the long setae arc arranged in clumps or tufts on the
carapace and these may be associated with irregularities in the carapace surface. The shafts of both short and long
setae can be divided into four regions: a bare basal region without setules, a region of small sparse setules,
a region of larger dense setules, and finally a bare apical region. The percentage of the length occupied by each
region varies between species. In Dynomene his pi da, D. praedator , and Metadynomene tanensis there are no
differences, between short and long setae, in the proportions of the shaft occupied by the four regions. Also the
short setae are not bent at right angles and the long setae arc not arranged in clumps on the carapace. Clumps of
long carapace setae are found in Dynomene filholi, D. pilumnoides , Hirsutodynomene spinosa, H. Ursula . and
Paradynomene tuherculata. The short setae are bent in D. pilumnoides, and H. spinosa (but not in H. Ursula).
The setae of most dynomenids have varying arrangements and sizes of setules along their length, but very unusual
setae arc found on D.fdholi and Paradynomene tuherculata where there is a marked difference between the structure
of short and long setae. The proximal 60% of short setae in D. filholi have the normal radiating setules arranged
around the shaft, but this is followed by a brush border of long fine setules along only one side of the shaft.
In P. tuherculata the proximal 40% is bare but the rest of the shaft is feather-like, bearing a row of fine setules
on opposite sides. None of the other dynomenids have setae which even closely resemble these aberrant
forms, although DE Man (1889) reported "Fedcrhaar" (feather-like setae) in D. pugnatrix. The reasons for these
differences are not clear. The setal characteristics are useful in the recognition of such genera as Hirsutodynomene
and Metadynomene , but in Dynomene the setal differences are much greater and hence are useful in recognizing the
different species. The setae of the dromiid Dromia personata (Linnaeus, 1758) bear a close resemblance to many of
the dynomenid setae (see JACQUES, 1989. her Fig. 3. 4).
ANTENNULES, ANTENNAE AND ORBITS
The antennules are composed of three articles plus flagellae. The First article is largest, about as long as the
greatest width and trapezoidal, remaining articles are much smaller, and the third article is longer than wide.
Antennules are very active during feeding. First article of antenna is wider than long, usually beaked medially
enclosing the urinary opening, second article is longer than wide bearing a fused exopod. third and fourth articles
together are as long as exopod, terminating in a flagellum whose length can be from 23% to 60% of carapace
width. The beaked first antennal article is a feature shared with members of the Homolodromiidae and Dromiidae.
However, in Acanthodromia the first antennal article is not beak-shaped and the urinary opening is on the medial
margin, concealed against the first article of the antennule.
The orbits of dynomenids are well formed and separated by the epistome which is joined to the rostrum of the
carapace. In dorsal view orbits are obliquely arranged and clearly exposed dorsally. There are well developed supra-
and suborbital margins, usually armed with spines or tubercles, which form a well defined cavity which can
accommodate the whole of the eyestalk when it is folded away. The gap between the suborbital margin and the
epistome is filled by the first article of the antennule and the first two articles of the antenna, thereby covering the
base of the eye stalk. The second article of the antennule articulates at a right angle so that the rest of the
appendage is folded horizontally above the eye stalk and under the supraorbital border. In a similar way. the fourth
article of the antenna is angled so that the flagellum is directed laterally. However, the antennal flagellum is too
long to allow the appendage to be entirely folded into the orbit.
MOUTHPARTS
During this study attention has only been paid to the third maxillipeds of dynomenids. Other authors have
dealt more or less with all six appendages: see BOUVIER (1896, fig. 23) and A. Milne Edwards & Bouvier
442
C. L. McLAY
(1900, pi. VII, figs 1-18) for mouthparis of Dynomene filholi , and Stebbing (1921, pi. 14) for mouthparts of
D. pilumnoides (as Maxillothrix actaeiformis). Note that on Stebbing's figure of the third maxilliped the epipod
is missing. The maxillae and maxillipeds of Acanthodromia erinacea were figured by A. MlLNE Edwards and
Bouvier (1902, pi. Ill, figs 6-10). Note that their third maxilliped is also shown without an epipod. ORTMANN
(1892, pi. 26, fig. 3i) figured the second maxilliped of D. praedator and CHEN (1979, figs 1, 4) figured the
external features of the third maxilliped (as D. sinensis). The epipods of the maxillipeds have an important role in
gill cleaning (see below).
F c tw\nH,?U,0Adr°me,ne “rs"la (StimPson- l860>- 6 134 x >0-3 mm. Mexico. Espiritu Santo Id. "Velero",
stn intertidal: a. long and short setae from left posterolateral corner of carapace; b, short setae from left
posterolateral corner ot carapace; c, setules on lip of long setae from left posterolateral corner of carapace. —
d, Metadynomene tanensts (Yokoya, 1933). <3 16.5 x 15.8 mm. New Caledonia, Smib 3. stn DW 25. 437 m: setae
rom right posterobteral corner of carapace. — e-f, Paradynomene tuberculata Sakai. 1963, 6 22.0 x 22.8 mm
^aZ ™ ?d°w’ S^IB 3> SlnrM‘ 246 m: e< l0ng setae from righl Posterolateral corner of carapace; f. short setae from
ight posterolateral corner of carapace. (All pictures taken with scanning electron microscope.)
Source :
FAMILY DYNOMENIDAE
443
In dynomenids the third maxillipeds are operculiform. the coxal articles are separated by tip of sternum, the
basis-ischial articles are fused but with joint still visible, the medial margins of ischia are parallel (or slightly
diverging) and close together, the meral article is square or oblong and smaller than the preceding article, followed
by a three-articled setose palp, consisting of carpus, propodus and dactyl which is folded along medial margin of
merus. The palp grasps food material from the chelipeds passing it on to other mouthparts.
Most dynomenids have a true crista dentata - a crest-like row of corneous teeth on the inner margin of the
ischium of the third maxilliped. The crista dentata is found in palinurids, nephropids, astacids, thalassinids, some
anomolans, and some podotremes. Working in a coordinated way with the mandibles it is used to grasp and tear
food items before they enter the mouth. In dynomenids the teeth are usually of even size, but in some species they
tend to increase in size distally. However, none of them have teeth such as is found in some thalassinids and
astacids where the crest is curved and terminates in a large hooked tooth. The number of teeth varies from 5-8 in
species of Dynomene and Hirsutodynomene , to 12-13 in Metadynomene and Paradynomene . A crista dentata is
absent in Acanthodromia. Amongst the podotreme crabs the crista dentata is present in homolodromiids, dromiids
and homolids as well as the dynomenids. The dromiids and dynomenids are the only decapods which have
operculiform third maxillipeds with a crista dentata. All other decapods with a crista dentata have pediform third
maxillipeds. The true crista dentata is absent from the other brachyurans.
GILLS AND EPIPODS (Figs 5 a-f, 6 a-f, 7 a-b)
In his classic work on the origin of crabs, Bouvier (1896) compared the gills of Dynomene filholi and
Acanthodromia erinacea with those of Homarus vulgaris. Although there were some errors made in the interpreta¬
tion of the gills (see below under the treatment of these species), the branchial formula was given as 20 gills +
7 epipodites on each side. Subsequently this branchial formula has been assumed to be typical of all dynomenids.
The general features of the number of gills and epipods in dynomenids are illustrated by considering the
arrangement found in the type species of the family. In Dynomene hispida there are 19 gills and 7 epipods present
on each side. There are six podobranchs, ten arthrobranchs and three pleurobranchs. The first thoracic segment has
no gills and there is only the epipod of the first maxilliped extending into the anterior end of the branchial
chamber. The second segment has a single arthrobranch and a podobranch on the epipod of the second maxilliped
which lies anterior to the arthrobranch. The third segment has the same number of gills as the second but the
epipod of the third maxilliped lies posterior to the arthrobranch. The fourth segment has two arthrobranchs and
a podobranch on the epipod which lies between the two arthrobranchs. The same pattern is repeated for segments
five through seven except that these segments have an additional pleurobranch. The eighth thoracic segment has
no gills or epipods. The podobranchs are attached to the epipods and the hypobranchial margin of each gill is
armed with long setae identical to those on the epipod itself. Thus the podobranchs themselves must also have a
cleaning role since they overly the bases of the larger gills. The epipods (Fig. 60 function as gill cleaners and are
either flattened plates or elongate lobes which bear long setae. In Paradynomene tubercidata these setae (Fig. 7e)
have the following structure: the proximal third is smooth, followed by a section covered with digital scales,
which are almost identical to those on the hypobranchial setae (see below), and towards the end of the setae these
scales are replaced on one side by two rows of closely spaced, short, curved pegs with a channel between them,
while for a short distance, digital scales continue on the other side unchanged until they too are replaced by pegs.
Near the end of the setae there are two spiralling rows of closely spaced spines on opposite sides of the setal axis.
The likely function of these scales is to dislodge debris as the epipods move between the gills. Epipod size is
related to the size of the associated gill(s) and all of them extend as far the dorsal limit of the branchial chamber.
They have an intimate association with nearby gills with their setae often penetrating the gaps between the gill
lamellae. The first epipod (on the first maxilliped) is an elongate plate with few setae, not associated with any
gill, but capable of extending back over the epibranchial surface of the first three arthrobranchs. Its role is
probably to keep the anterior part of the branchial chamber and the anterior arthrobranchs free of debris. The second
epipod is much more setose and lies anterior to the first arthrobranch, cleaning only the anterior face of this gill.
The third epipod (on the third maxilliped), which is the longest, lies between the second and third arthrobranchs
and cleans the adjacent faces of these gills. Similarly with the fourth epipod which lies between the third and
fourth arthrobranchs. Thus the anterior and posterior faces of the third arthrobranch (the largest gill) are cleaned by
444
C. L. McLAY
different epipods. The fifth epipod (associated with the second percopod) lies between the fifth and sixth
arthrobranchs, cleaning the posterior and anterior faces (respectively) of these gills as well as the posterior face of
the first pleurobranch. In the same way the sixth and seventh epipods clean the same faces of the remaining
arthrobranchs and pleurobranchs. Apart from the third arthrobranch, all the other gills are only cleaned on one or
other side: there is no cleaning limb between the first and second, fourth and fifth, sixth and seventh, eighth and
ninth arthrobranchs, and the posterior face of the tenth arthrobranch cannot be cleaned by the eighth epipod because
it is absent. It is unclear how these gill surfaces are kept clear of debris, but the setose margin of each podobranch
may contribute to this task. The epipods can move in a vertical direction and clean the adjacent gill surfaces, right
from the hypobranchial to the epibranchial margins. When they are against the body wall their setae extend under
the gill and could help to clean the hypobranchial surface. The podobranchs are cleaned by long setae on the base
of the epipod to which the gill is attached.
The epibranchial surface of the posterior gills is cleaned by several long flexible setae extending from the
posterior border of the scaphognathite. These setae reach as far as the second pleurobranch (on the third percopod).
In Metadynomene tanensis the margin of the scaphognathite carries a dense fringe of short plumose setae with two
very long, stout setae (Fig. 70 inserted on the posterior border. These setae are armed with stiff, acute selules for
almost their entire length. The setules project at about 45° from the setal axis, with those on the proximal half
directed towards the base, while those on the distal half are directed towards the tip. The setules become denser
distally. In adult dynomenids there are normally two or three such long scaphognathite setae.
Besides the epipods and long scaphognathite setae there is another gill cleaning mechanism in dynomenids.
This is best developed in Paradynomene tiiberculata where the hypobranchial wall of the posterior half of the gill
chamber is covered by a dense field of long setae. These setae (Fig. 7c) have a range of lengths, arc arranged in
clumps and project from the body wall into the hypobranchial surface of the gills. Each seta has a complex
structure: the proximal hall is smooth, followed by a section where opposite sides of the seta are covered by
apically-dirccted digital scales, separated by intervening smooth areas. At about 80% of the length of the seta the
digital scales on one side are replaced by closely-spaced short, curved pegs arranged as marginal rows with a
channel between them, while the digital scales on the other side continue unchanged. These scales give the setae a
comb-like appearance. Comparison with the setae on the epipods (see above) shows that the digital scales are
identical and it seems likely that they must also have a cleaning role. Since the setae are fixed, the gills have to
move about in order to dislodge debris. It may be that the epipods produce gill movement or perhaps cause the
long setae to move from side to side. Besides Paradynomene tiiberculata , hypobranchial setae are also well
developed in Metadynomene tanensis and M. devaneyi , but in Hirsutodynomene and all species of Dynomene there
arc only a few of these setae present. Their status in Acanthodromia is unknown. This kind of gill cleaning
mechanism has only been reported from the dromiid Cryptodromiopsis larraburei (Rathbun, 1910) (Bauer, I98K
as Dromidia larraburei). Tufts of setae arising from the body wall beneath the gills have also been observed in
some other species of the Dromiidae (McLay, unpublished). Pereopodal epipods are greatly reduced in dromiids
and it may be that these hypobranchial setae take over the role of cleaning gills in the posterior half of the
branchial chamber. In Dromia erythropus (George Edwards, 1771) the hypobranchial setae (Fig. 7d) have a unique
structure: the proximal 80% of each seta is smooth but approaching the tip there are three or four isolated, apically
directed acute spines followed by a series of separate paired rows of comb-like pegs, increasing in number distally
and spiralling around the setal axis. Near the tip these pegs are transformed into two closely spaced continuous
rows of acute spines which continue the spiral right to the end. In profile, the distal region of the setae appear to
have three or lour rosettes of these acute spines. In his review of decapod grooming Bauer (1989) did not report
any setae which resemble those found in D. erythropus. The digital scales on the hypobranchial setae of
P. tiiberculata are similar to those found on the setiferous epipods, or setobranchs, of dcndrobranchiate and
candean shrimps, as well as achelate and homarid lobsters (Bauer, 1981, 1989), but they are very different from
the long barbed setae found on the maxillipedal epipods of the portunid crab. Cronins tumidulus. Instead of
bearing scales, these setae have a single row of recurved hooks (Bauer. 1989. his Fig. 12 c-d).
Dynomenids, like other Brachyura, have an epipodal gill cleaning mechanism but this is supplemented by the
scaphognathite setae over the anterior epibranchial surface and by the body wall setae attending to the posterior
hypobranchial gill surface. Compared to the more derived Brachyura. gill cleaning in the Dynomenidae is much
Source :
FAMILY DYNOMENIDAE
445
Fig. 5. — a , Dynomene hispida Gu£rin-M6neville, 1832. 9 14.0 x 10.8 mm. Hawaii. Oahu, (BPBM 658): transverse
section through gill. — b. Dynomene hispida Guerin-Meneville, 1832. 9 10.3 x 8.9 mm. Cocos Keeling Ids. (WAM
751-89): apex of gill. — c. Dynomene filholi Bouvier, 1894, 9 10.0 x 8.7 mm. Cape Verde Ids, Cancap. sin 7.125.
85-130 m: transverse section through gill. — d, Dynomene pugnatrix de Man, 1889, 6 9.8 x 7.2 mm, Mauritius
(SMF 4857): transverse section through gill. — e, Hirsutodynomene spinosa (Ralhbun, 1911). 9 11.3 x 8.9 mm.
Madagascar, Tulcar, stn 14-11-2, 5 m (MNHN-B 22077): transverse section through gill. — f. Hirsutodynomene
spinosa (Rathbun, 1911), 6 14.2 x 10.8 mm, Cocos-Keeling Ids., 0-37 m, (WAM 139-94 ): lateral view of whole
gill. (All pictures taken with scanning electron microscope.)
Source : MNHN. Paris
446
C. L. McLAY
Fig. 6. — a, Hirsutodynomene Ursula (Stimpson, 1860), 9 ovig. 19.4 x 14.7 mm. ARGOSY 34 (USNM 247230):
transverse section through gill. — b. Hirsutodynomene Ursula (Stimpson. 1860), 9 ovig. 15.0 x 12.2 mm. Ecuador.
La Plata Id. Askoy", stn 80 (LACM): lateral view of whole gill. — c, Metadynomene tanensis (Yokoya, 1933).
9 19.2 x 18.9 mm. New Caledonia, Musorstom 4, stn 215. 485-520 m: transverse section through gill. —
d , Acanthodromia erinacea A. Milne Edwards, 1880, 9 16.7 x 17.7 mm. Yucatan, " Blake ", stn 166,^275 m
(MCZ6509): transverse section through gill. — e. Parody nomene tuberculata Sakai, 1963. 6 21.0 x 21.9 mm.
New Caledonia, Smib 8. sin DW 189, 400-402 m: transverse section through gill. — f, Paradxnomene tuberculata
Sakai. 1963, 9 21.5 x 21.2 mm, Loyalty Ids, Musorstom 6, stn DW 406. 373 m (MNHN-B 25249): cpipod and
podobranch from left third pereopod. (All pictures taken with scanning electron microscope.)
Source : MNHN, Paris
FAMILY DYNOMENIDAE
447
more complex with the task being divided amongst eight appendages (second maxilla plus the seven epipods) as
well as the body wall hypobranchial setae, compared to only three appendages (the maxillipeds). At least in part,
this must be regarded as a consequence of having more gills. Besides the podobranchs, which are always small,
dynomenids have thirteen large gills versus only eight or nine in other Brachyura. In order to transform the method
of gill cleaning seen in dynomenids into that seen in more derived Brachyura. it is necessary to remove all the
pereopodal epipods, elongate the epipod of the first maxilliped to clean the hypobranchial region, and remove the
epipod of the third maxilliped from between the second and third arthrobranchs, and elongate it so that it covers the
cpibranchial gill surface. This is sufficient to clean the reduced number of gills and no change in the relative size
of the second maxilliped epipod is necessary. Since gills are lost from the posterior part of the thorax, the field of
setae on the hypobranchial wall is no longer necessary. The relationships between gill structure and gill cleaning
mechanisms have been explored by SUZUKI & McLay (1998). Dynomenids have retained several plesiomorphic
gill and gill cleaning characters.
In cross section the gills are basically violin-shaped, with afferent and efferent vessels in the 'body' of the gill
and a notch on each side. Dynomenids show a great deal of variation in the shape of their gills, chiefly in the
number of lobes (or filaments) on the cpibranchial surface. Bouvier (1896: 26, footnote) has already noted the
variation in number of filaments along the length of each gill. Therefore in order to make comparisons between
species it is necessary to standardize the point at which a cross section is taken and the gills which are used.
Comparisons were made using a section across an arthrobranch or pleurobranch from the first two pereopods
at approximately half the length from the point of attachment. Lobes on the gills are arranged in rows length¬
wise, but they do not always lie in the same plane in cross section. This means that while the hypobranchial
half of each gill is composed of a series of plates, the lobes do not always correspond exactly to each pair
of plates. Furthermore, the lobes on the posterior side of the gill are usually longer than those on the anterior
side.
In the type species Dynomene hispida and in D. praedator the gill cross section shows a pair of plates (or
flattened lobes) on the epibranchial surface. In D. pugnatrix an additional short median lobe is added to make three.
In D. pilumnoides there are four long lobes while in D.filholi there are six lobes decreasing in size medially. [My
interpretation of the gills of D. filholi differs from that of Bouvier (1896) who gave the number of filaments as
being eight because he included the portion of the lateral notch as a filament, whereas I have treated them as part
of the ’body' of the gill.] In both species of Hirsutodynomene there are six lobes decreasing in size medially.
In Metadynomene the number of lobes increases from four (M. tanensis ), five (M. devaneyi) to six (M. crosnieri).
The gills of Paradynomene tuberculata have four lobes. In both species of Acanthodromia the epibranchial
extensions consist of two flattened lobes. Thus within the Dynomenidae we have gills ranging from the multi-
lobcd trichobranchiate-like condition seen in Hirsutodynomene and D.filholi , through I), hispida and D. praedator
in which the number of lobes is reduced to only two (which are flattened) and Finally to the phyllobranchiate-Iikc
condition seen in Acanthodromia. But even in Acanthodromia the distinctive lateral notch is evident on each
lamella of the gill so that they still differ from the condition found in dromiids, such as Epigodromia ,
Hypoconcha, and Conchoecetes , where the gill plates are rounded in outline, and not interrupted by notches.
Dynomenid gills show little variation in numbers but great variation in shape, while gills of the Dromiidae show
greater variation in numbers but almost no variation in shape.
Bouvier (1896, figs 19, 23) compared the gill structure of Dynomene filholi which has several rows of
"filaments" with Acanthodromia erinacea which has only two plates. He made comparisons with Homarus
vulgaris as well as Homolodromia paradoxa , Dicranodromia ovata , and D. mahieuxii. Like the dynomenids, the
latter two homolodromiid species have different numbers of epibranchial "filaments". Bouvier regarded
D. mahieuxii as being a little less primitive because it has fewer filaments which are more plate-like and he
suggested that these gill plates might have originated by the concrescence of several short "filaments". In making
the same comparison between the two dynomenids Bouvier discussed the interesting problem of how a species
such as D.filholi could have evolved a more advanced crab-like form and yet still retain such primitive homarid-
like gills. He considered that D. filholi , more than any other dromiacean, showed close links with the homarids.
BOUVIER argued that D. filholi cannot be considered as deriving from Dicranodromia ovata (Bouvier used
" Acanthodromia ovata ", but this must be an error for " Dicranodromia ovata”) which is, in many respects, more
448
C. L. McLAY
Fig. 7. — a. Epigodromia areolata (Ihle, 1913), 8 14.2 x 12.1 mm. New Caledonia. " Vauban ", stn DW 1 147, 210 m.
28.10.1989: transverse section through gill. (Dromiidae). — b, Hypoconcha parasitica (Linnaeus, 1763). 8 23.1 x
23.3 mm, Florida, off Panacea, coll. M. Wicksten: transverse section through gill. (Dromiidae).
— c. Paradynomene tuberculata Sakai. 1963. 8 22.0 x 22.8 mm. New Caledonia, Smib 3. stn 14, 246 m: cleaning
setae from hypobranchial wall of right gill chamber. — d. Dromia erythropus (George Edwards, 1771), 8 81.5 x
62.7 mm, St. Croix, Virgin Ids (USNM 72355): cleaning setae from hypobranchial wall of right gill chamber.
(Dromiidae). — e, Paradynomene tuberculata Sakai, 1963, 9 21.5 x 21.2 mm. Loyalty Ids. Musorstom 6,
stn DW 406, 373 m (MNHN-B 25249): cleaning setae on epipod from left third pereopod. — f, Metadynomene
tanensis (Yokoya, 1933), 9 16.2 x 15.3 mm. New Caledonia, Smib 8, stn DW 198. 414-430 m: long setae from
posterior margin of the left scaphognathite. (All pictures taken with scanning electron microscope.)
Source : MNHN . Paris
FAMILY DYNOMENIDAE
449
primitive. He believed that other dynomcnids are probably at a more advanced evolutionary state than Dynomene
praedator , and it’s possible that, as yet undiscovered species of "Acanthodromies", might be more primitive than
D. ovata, so that it is possible to consider the genus Acanthodromia as being the link which connects the genus
Dynomene to the primitive dromiaceans which originated from the homarids. This hypothesis, which now' seems
rather unlikely, is largely based on the resemblance of dynomenid gill structure to that found in homarids. Clearly
Bouvier considered phyllobranchiate-likc gills as being derived from the trichobranchiate condition. The tendency
to evolve phyllobranchiate-like gills seems to have occurred independently in the Homolodromiidae, Dynomenidae
and Dromiidae since the last common ancestor of the three groups must have had multi-lobcd gills.
PEREOPODS
Anterior pereopods: The first pereopods of dynomcnids are longer than walking legs, chelate and sexually
dimorphic, being larger in males. The meral article is long and trigonal in section, while the carpal article is dis¬
tinctive in usually having a prominent sharp spur on the inner superior border. This feature is absent in
Metadynomene, where there are three small tubercles, and in Acanthodromia , where the whole limb is spinous.
The fixed finger is straight and is armed with from two to eight small teeth often increasing in size distally. The
number of teeth on the dactyl is usually less than on the opposing finger. The tips of both fingers are usually den¬
tate. In Dynomene , Hirsutodynomene and Acanthodromia the dactyl is down-curved so that the fingers gape, but in
Metadynomene and Paradynomene the dactyl is essentially straight so there is little gap and the fingers touch for
about half their length. Both fingers are hollowed out internally with a bunch of stiff setae inserted near the base
of each finger and projecting across the gap between them. In Dynomene and Hirsutodynomene these setae are very
well developed, forming a screen or sieve behind the outer dentate margins. Observations on D. praedator show
that these setae help to sift out food particles which are passed to the third maxillipcds (sec below).
The second through fourth pereopods are well developed walking legs of similar size to the first pereopods or
slightly smaller. They tend to decrease in length posteriorly. Plane of movement of the sternal-coxal articulation
is anterior-posterior, for the ischial-basis articulation plane of movement is dorso- ventral, the meral-ischial
articulation has a small anterior-posterior plane of movement, the carpal-meral articulation is also dorso-ventral,
the propodal-carpal articulation moves anterior-posterior, and finally the propodal-dactyl articulation is dorso-
ventral. Dactyli are long and curved with their inner margins usually armed with two to six small spines which are
probably used to grasp the substrate on which the crabs live. There is no clear pattern of variation in number of
spines between the genera. The only exception is Dynomene pugnatrix which has ten dactyl spines. This may
indicate that it lives in association with a different kind of host or habitat. At least some of the other shallow
water species are known to live among corals and coral debris.
Fifth pereopod (Figs 8 a-f, 9 a-f, 10 a-d, 1 1): Perhaps the most distinctive feature of the Dynomenidae is
the very reduced last pair of pereopods which are carried alongside the posterolateral corners of the carapace, above
the base of the fourth pereopods. The reduced leg is directed anteriorly. It is commonly stated that dynomcnids,
like dromiids, carry their last legs in a dorsal or subdorsal position, but this is not correct. This limb cannot rise
above the level of the carapace margin because it is too short and relatively immobile and so cannot be described
as dorsal or subdorsal. A more accurate description would be that the limb is "horizontal". Most of the scope for
movement in this limb is attributable to the coxo-basal article because the remaining joints are scarcely moveable.
The basis-ischium articles are fused in all species, and in Acanthodromia erinacea and Paradynomene tuherculata
the merus is also fused to the preceding articles. The inferior distal margin of the merus is hollowed out as in the
previous three limbs. However, it is the anterior distal border of the propodus which is extended to form the fixed
finger of the subchelate mechanism and when the limb lies in its natural position the dactyl is almost ventral, the
opposite of the cheliped. Observations on live D. praedator show that these limbs have a very restricted scope for
movement: they are only capable of moving in an anterior-posterior plane above the bases of the preceding
pereopods. Most of the time they simply lie alongside the posterolateral corner of the carapace, but when the crab
moves using its pereopods, they often move at the same time as though they are part of the coordinated pattern of
limb movements. The last legs cannot reach above on to the posterolateral corner of the carapace or beneath the
crab into the abdominal cavity. Thus they are not capable of carrying a piece of camouflage or performing a
grooming role.
450
C.L. McLAY
Fig. 8. — a. Dynomene praedator A. Milne Edwards. 1879, 6 9.6 x 7.8 mm, Somalia. Gesira, stn 19, intertidal coral
(MZUF): propodal spines from left fifth pereopod. — b. Dynomene praedator A. Milne Edwards, 1879, 9 ovig. 9.6 x
7.5 mm, Glorieuses Ids. intertidal: dactyl spines from right fifth pereopod. — c, Dynomene filholi Bouvier, 1894, 9
10.0 x 8.7 mm. Cape Verde Ids. Cancap, stn 7.125, 85-130 m: dactyl and propodal spines from left fifth pereopod.
— d-e, Dynomene pilumnoides Alcock, 1900, 9 12.8 x 10.3 mm, New Caledonia, VOLSMAR, stn DVV 7, 400 m:
d. dactyl spines from left fifth pereopod; e. propodal spines from left fifth pereopod. — f. Hirsutodynomene spinosa
(Rathbun. 1911), 9 23.8 x 17.8 mm. Cocos Keeling Ids, 0-28 m (WAM 723-89): propodal spine from left Fifth
pereopod. (All pictures taken with scanning electron microscope.)
Source : MNHN, Paris
FAMILY DYNOMENIDAE
45!
Fig. 9. — a-b. Hirsutodynomene Ursula (Stimpson, 1860), 9 ovig. 15.0 x 12.2 mm, Ecuador, La Plata Id. "Askoy",
stn 80 (LACM): a. dactyl spines from right fifth percopod; b, propodal spines from right fifth pereopod. —
c, Hirsutodynomene spinosa (Rathbun. 1911), 9 23.8 x 17.8 mm. Cocos Keeling Ids, 0-28 m (WAM 723-89):
dactyl spines from left fifth pereopod. — d, Metadynomene tanensis (Yokoya, 1933), <3 16.5 x 15.8 mm.
New Caledonia, Smib 3, stn DW 25, 437m: lateral view of tip of left fifth pereopod. — e. Metadynomene tanensis
(Yokoya, 1933), 9 16.2 x 15.3 mm. New Caledonia, Smib 8. stn DW 198. 414-430 m: propodal spines from right
fifth pereopod. — f, Acanthodromia erinacea A. Milne Edwards, 1880, 9 9.5 x 11.5 mm, Yucatan, " Albatross ",
stn 2354, 238 m (USNM 9547): dactyl from right fifth pereopod with marginal spines. (All pictures taken with
scanning electron microscope.)
Source :
452
C. L. McLAY
Fig. 10. — a, Acanthodromia erinacea A. Milne Edwards, 1880, 9 9.5 x 11.5 mm, Yucatan, Albatross stn 2354.
238 m (USNM 9547): propodal spines from right fifth pereopod. — b-c, Paradynomene tuberculata Sakai, 1963,
9 21.5 x 21.2 mm. Loyalty Ids, Musorstom 6, stn DW 406, 373 m (MNHN-B 25249): b, dactyl from right fifth
pereopod with marginal spines; c, propodal spines from right fifth pereopod. — d. Paradynomene tuberculata Sakai,
1963, 6 22.0 x 22.8 mm. New Caledonia. Smib 3, stn 14, 246 m: lateral view of tip of left fifth pereopod.
(All pictures taken with scanning electron microscope.)
Comparison of the last two pereopods for males of all species shows that on average the fifth pereopod is only
31.2% of the length of the fourth pereopod (Fig. 1 1). Comparing individual articles between these limbs shows
that there are no differences for most articles except for the coxae and dactyli. On average the coxae of the last limb
occupy 19.6% of the length but on the preceding leg it is only 10.8%. For the dactyli the reverse is true with the
dactyli of the fourth pereopods occupying 17.0% while those of the fifth only occupy 5.3%. Thus except for the
articles at each end of the limb the fifth pereopod is just a scaled down version of the preceding limb. The coxa
carries the male gonopore, and perhaps cannot be reduced by the same amount as the whole limb and still be
functional, while the dactyl is part of a novel subchelate structure. The last pereopod of dynomenids is not reduced
as much as found in the cymonomid Elassopodus stellatus Tavares, 1993, where both of the last two pairs of
pereopods are reduced to tiny stumps, almost concealed by the abdominal segments. It seems that there is scope
for a lot of redundancy in the posterior pereopods of decapods.
Source : MNHN, Paris
FAMILY DYNOMENIDAE
453
Dynomene hispida
Hirsutodynomene Ursula
jlmJQjoxllJL f j jn,H,r .1 I I
Dynomene praedator 40
30 ■
CX B-l M Cp Pr D P5/P4
Hirsutodynomene spinosa
PA P5
cp Pr D P5/P4
Dynomene filholi
■XJCL
40 •
i I -ljarfi.lLii.fi, in. I ,
CX B-l M Cp Pr D P5/P4
P4 P5 Metadynomene tanensis
CX B-l M Cp Pr D P5/P4
P4 P5 Dynomene pugnatrix 40
30
■ .1 1 111 -Uurin.r.fi.iJ .
CX B-l M Cp Pr o P5/P4
Metadynomene devaneyi
P4P5
Cp Pr D P5/P4
Dynomene pilumnoides
CX B-l M Cp Pr D P5/P4
Acanthodromia erinacea
bJUtJL
CX B-l M Cp
Paradynomene tuberculata
P5/P4
CX B-l M Cp Pr D P5/P4
Fig. II. — Relative size of articles of the last two pereopods of Dynomene hispida. D. praedator, D. filholi.
D. pugnatrix, D. pilumnoides, Hirsutodynomene Ursula, H. spinosa. Metadynomene tanensis, M . devaneyi.
Acanthodromia erinacea and Paradynomene tuberculata. From left to right, the proportion of total limb length (fourth
pereopod followed by fifth pereopod) is shown for coxa (Cx), basis-ischium (B-I). merus (M), carpus (Cp).
propodus (Pr), dactyl (D), and the ratio of length of fifth to fourth pereopod (P5 9P4). Note that in A. erinacea and
P. tuberculata the basis-ischium and merus arc fused. All measurements made on males except for D. pilumnoides.
D. pugnatrix, and A. erinacea.
The last pereopod is subchelate. BOUVIER (1896) stated that Acanthodromia erinacea had a fifth pereopod of the
same form as Homolodromia paradoxa but their resemblance is only superficial and closer examination shows that
they are in fact very different. The subchelate mechanism in dynomenids involves the dactyl being opposed to a
distal extension of the propodus which bears toothed or untoothed spines. The structure of the subchelate
mechanism of the fifth leg is sexually dimorphic. The propodal extension is better developed in females
(resembling the condition found in Homolodromia , thus accounting for Bouvier's observation) than it is in males
(resembling the condition found in Dicranodromia . see GUINOT. 1995. her fig. 3 A-H. for details). The most
obvious difference is in the dactyl: in males the dactyl has the conventional claw shape but in females the dactyl is
modified as a flattened plate bearing 5-16 typically untoothed spines. In females there is usually more spines on
the dactyl than on the propodus. These spines are hooked and only have a few small proximal teeth or none at all.
The surface of the dactyl spines is usually concave and crenulate. Other differences between male and female
dynomenids are that in females of all species the propodal spines bear small teeth, but these arc only found in
males of D. hispida, D. praedator and in both species of Hirsutodynomene. The number of propodal spines also
differs: while both sexes of Dynomene have a similar number (approx. 5) of spines, in Metadynomene .
Source
454
C. L. McLAY
Hi rsutodynomene, and Paradynomene the females have a significantly larger number (8-16) of spines. In summary
we find that in females there are opposable rows of usually toothed spines whereas in males the usually untoothed
propodal spines are opposed by a claw-like dactyl.
In some males (Dynomene filholi , Metadynomene tanensis , and Paradynomene tuberculata) a dorsal or lateral
spine is found on the claw-like dactyl (see Figs 7d. 8d). No dactyl spine is found in males of Hi rsutodynomene and
the condition in Acanthodromia is unknown. This spine resembles a similar spine found on the dactyli of the last
pereopods of some primitive members of the Dromiidae (Tunedromia McLay, 1993, Dromidiopsis Borradaile,
1900, and Laiiridromia McLay, 1993) where it helps the animal to grasp and hold its piece of sponge camouflage.
Both sexes of these dromiid genera have the spine but it is only present in male dynomenids. In these dromiids the
spine projects from the surface of the dactyl, but in the dynomenids the spine lies in a depression on the surface
and cannot be functional. It seems to be an apomorphic vestigial structure which indicates a close relationship
with the dromiids and suggests that the last pereopod of dynomenids may be derived from a camouflage-carrying
limb.
What is the function of the reduced last pereopods? STIMPSON (1860) stated that " . they fill, apparently no
office in the economy of the animal, except when in place, they fill up neatly the chink between the carapax and
the stouter walking feet." STIMPSON clearly believed that this limb is redundant. The small size and lack of
mobility of the fifth legs suggest that the limb is vestigial. Reduction has proceeded furthest in Acanthodromia
and Paradynomene where we find the greatest number of fused articles. It would seem that it only has a function in
male dynomenids because it carries the gonopore in its coxal article. The subchelate tip does not appear to be
functional because the dactyl is largely immovable. Therefore the subchelate tip must represent some past rather
than present role. The sexual dimorphism makes it difficult to imagine what this role might have been and why it
needed to be different in the two sexes. The fifth pereopod is sexually dimorphic in some of the Scyllaridae
(Holthuis, 1985), Thaumastochelidae (Holthuis, 1974) and Palinuridae where the females have subchelate
limbs which are used to clean the abdomen and brood. STEWART et ai (1997) suggest that the subchelate fifth
pereopods in female Ibacus peronii are involved in the fertilization and manipulation of eggs, as they arc attached
to the pleopods, and subsequently used to clean and groom the egg mass. However the structure of the tip of the
limb consists of a well developed dactyl opposed to a strong, simple, spine-like propodal extension. In
dynomenids the structures on the dactyl and propodus are much more elaborate. Among the Homolidae, GuiNOT
and Richer DE Forges (1995: 307, 469) reported that the merus of the last pereopod is shorter in females than in
males of all species of Homologenus.
The dynomenid female limb has a structure which resembles some kind of cleaning or grooming appendage
which is typical of many anomolans but there the resemblance ends because the structure of anomolan fifth
pereopods is quite different (Bauer, 1989; Pohle, 1989). Anomolan fifth pereopods have a well developed
subchclate tip, not sexually dimorphic, and can be inserted into the branchial chamber to clean the gills, see for
e.g. the Porcellanidae (Fleischer et ai, 1992), but dynomenids have a closely fitting carapace which would deny
these limbs access to the gills. The dynomenid spine structure is somewhat similar to the propodal setae found on
the fifth pereopods of carideans such as Palaemon and Betaeus which are used for body grooming (Bauer. 1989).
Furthermore similar setae are found in axiid thalassinideans, astacid and cambarid crayfish, and nephropid lobsters
(Bauer, 1981). In none of these cases arc the limbs sexually dimorphic. BAUER (1989: 61) points out that no
grooming fifth pereopod has ever been described for a brachyuran but a former body or gill grooming function
seems the most likely role for the fifth pereopod in dynomenids. It should be noted that one other feature normally
accompanies decapod cleaning fifth pereopods: in order for these limbs to be fully functional, and access
all the areas in need of cleaning, the sternite of this limb must be mobile and not attached to the preceding
sternite. Once the last sternite is attached, as it is in the Brachyura, the last pereopods can no longer perform a
cleaning function.
FEMALE STERNAL SUTURES
Apart from the thoracic sternal suture 7/8 in females (see below) which is always evident, several other sutures
mark the boundaries of thoracic sterna in some genera. In Metadynomene sutures 3/4, 4/5 (faintly), and 5/6 (very
strong) are visible. Hirsutodynomene is the same as the previous genus except that the suture 5/6 is not apparent.
Source : MNHN. Pahs
FAMILY DYNOMENIDAE
455
In Pa r adynomene , A can t hodrom ia and most species of Dynomene (suture 4/5 is faint in D . hispida) only the
suture 3/4 is evident. Sternum 3 separates the bases of the third maxillipeds and its separation from sternum 4 is
always deeply marked. The suture 4/5 is only faintly evident in a few species, while suture 5/6 is strongly marked
by a semi-transparent band which only occurs in Metadynomene. The median line, where sutures from each side
meet, is not apparent in any dynomenids. Guinot (1979: 80) recognized four categories of thoracic sterna based
on the interruption of sutures 4 to 7. Unlike sternitreme crabs, most sternal sutures are absent in dynomenids, so
they do not conform to GuiNOT's classification. If we regard the absence of many sternal sutures as representing
the apomorphic condition, then dynomenids must be regarded as having very derived sterna. In the
Homolodromiidae only the 7/8 suture and traces of 6/7 arc visible (GUINOT, 1995: 174). In the Dromiidae the
structure of the sternum is very much distorted by the more anterior position of the spermathccae with the result
that the seventh and eighth sterna occupy much of the ventral surface of the cephalothorax. The sternum of
dromiids represents the derived condition with both homolodromiids and dynomenids retaining the plesiomorphic
condition where the suture is very short.
Female sternal structure 7/8: The sternal spermathccae are separate from the gonopores and lie at the
boundary between the seventh and eighth sternites. The length of the sternal sutures, which mark the suture
between these sternites, depends upon the proximity of the spermatheca to the gonopore. In dromiids the length of
the sternal sutures 7/8 is variable: the spermatheca can lie between the bases of the first percopod, requiring very
long sternal sutures 7/8, or between the bases of the fourth pereopods, requiring only very short sternal sutures
7/8. Thus the spermathccae can be anterior or posterior to the female gonopore. Also in dromiids these sutures can
end apart or together. In dynomenids the sternal sutures 7/8 are always short, usually ending just below or slightly
behind the female gonopore on the third pereopod, and they always terminate apart. The sutures lie very close to
the coxae of the adjacent pereopods. In Dynomene and Hirsutodynomene the sutures end on low tubercles. In
Metadynomene the sutures lie in a shallow, V-shaped groove, below a prominent parallel medial ridge, concealed
by a dense layer of long soft setae originating from the adjacent coxa of the fourth pereopod. In Acanthodromia the
sutures end beneath a curved over-hanging lip without a setal covering. In Paradynomene the sutures are almost
completely covered by the coxae and setae of the fourth pereopod. Close proximity of the spermathccae to the
female gonopore ensures fertilization of the eggs when they are laid. In this respect dynomenids are very similar to
the homolodromiids (GUINOT, 1995).
A feature of the sternal sutures 7/8 of dromiids is that in mature females they are often covered with a dark
gelatinous layer which closes the spermathccae. This substance is probably produced by the male and could act as
a sperm plug, preventing other males from inseminating the female. In homolodromiids GUINOT (1995) reported
that many females had the broken off tips of male second pleopods blocking the entrance to the spermathecae and
perhaps functioning as a different kind of sperm plug. In all the dynomenid females I have examined, I have never
seen sperm plugs of cither of the above kinds. This may imply that dynomenids have a mating strategy different
from dromiids and homolodromiids.
ABDOMEN
Dynomenids have an abdomen of six free segments with no segments fused. Segments increase in length and
breadth posteriorly with margins fringed with long setae. The telson is much wider than long, with the anterior
margin angled to accommodate the uropods and the posterior margin broadly rounded. Compared to other
dromiaceans, the uropods are well developed, visible externally and often completely excluding the penultimate
segment from reaching the lateral margin, especially in females. The abdomen and pleopods provide a protected
chamber where the eggs are incubated until they hatch. No dynomenids provide parental care for their larvae.
Unlike the Dromiidae, most of the Dynomenidae have no effective abdominal locking mechanism and the
abdomen in both males and females is simply curled under the cephalothorax and held loosely in position by its
own musculature. There are differences between dynomenid genera in the nature of abdomen-restricting structures.
In Dynomene males and immature females there is a small rounded sternal tubercle at the lateral margins, below
the articulation of the first walking legs, and adjacent to the uropods when the abdomen is in its natural position
(see also Guinot, 1979: 125-126). These tubercles simply restrict sideways movement of the abdomen and they
456
CL. McLAY
disappear in mature females where the abdomen occupies all the ventral surface. The same arrangement as in
Dynomene is found in Hirsutodynomene. However Metadynomene has small spines (can be bifid) or ridges on the
coxae of the second and third pereopods, adjacent to the margins of the telson and penultimate abdominal
segments, which restrict lateral movement. In Paradynomene small rounded granules cover the coxal articles ol the
pereopods, leaving the surface of the abdominal cavity smooth. The margins of the abdomen are neatly surrounded
by many granules (several on each coxa) which restrict sideways movement. In Acanthodromia the abdomen of
mature females is confined between tuberculate coxa! projections on pereopods and under projections on the coxae
of the third maxillipeds. Use of the pereopod coxae to restrain the abdomen resembles the situation found amongst
many dromiids and use of maxilliped spines resembles the condition found in the Poupiniidae. Clearly, the
abdomen locking mechanism of Acanthodromia is very different from that found in the other dynomcnid genera.
There is a gradation from abdomen maintaining mechanisms as found in Dynomene + Hirsutodynomene where
sternal tubercles are used, through Metadynomene , using coxal ridges or spines on the second and third pereopods,
and Paradynomene using coxal granules, to the abdominal locking mechanism found in Acanthodromia which uses
well developed coxal projections on the third maxillipeds and first three pereopods. (See also below under
Uropods.)
Observations of live Dynomene praedator show that the abdomen is not always held closely against the
sternum and when the crab moves, it often makes "flicking" movements similar to those seen in for e.g.
porcellanids. By themselves these abdominal movements would tend to propel the crab backwards but they occur
when the crab moves both forwards and backwards. Locomotion is achieved using the three pairs of walking legs.
The motor pattern causing abdominal movements is probably a vestige of the past when the abdomen was
involved in locomotion.
In mature females the abdomen covers the entire sternum and coxae of all pereopods with the telson covering
the proximal half of the third maxillipeds. In males the abdomen is not quite so broad and the telson only extends
as far as the bases of the third maxillipeds. While mature dynomcnid females are clearly recognizable by their wide
abdomens, immature female and male abdomens are not greatly different. Therefore abdomen width is not
as reliable a method of sexing specimens as it is in eubrachyurans. The relative size o f Metadynomene tanensis
male and female abdomens is shown in Fig. 28b. In this species females seem to have a pubertal moult at a
CW of around 1 1.0 mm and males have a pubertal moult around 15.0 mm. In both sexes the pubertal moult is
not terminal.
In order to ensure sperm transfer, the female abdomen must be flexible enough to expose the posteriorly placed
spermathecae. Males only need short pleopods, but because their abdomen must fit inside that of the female, it
must be relatively short and/or flexible, so that it can be folded or curled to allow the pleopods to come into
contact with the spermathecal openings and deposit the sperm.
PLEOPODS (Figs 12 a-f, 13 a-f, 14 a-e)
Dynomcnid crabs are unusual in having five pairs of pleopods in both sexes. The first pair of female pleopods
are uniramous and reduced in length, and do not carry eggs, while the remainder are normal biramous egg-bearing
limbs. In males the first pair are a semi-rolled lube, with an oval apical plate surrounded by setae, while the
second pair are needle-like with an exopod on the basis, and the last three pairs of pleopods are rudimentary.
Dynomenid male pleopods have been previously illustrated as follows:
Dynomene hispida : PEYROT-CLAUSADE & SERENE (1976, text-fig. 1, pi. 5. A-B, F), GUINOT (1979. fig. 60
e-f), Dai et al. (1986, fig. 11, 2-3), and Dai & Yang (1991, fig. 11, 2-3). As D. granulobata , Dai, Yang &
Lan (1981. figs 13-14), Dai et al. (1986, fig. 12, 1-2), and Dai & Yang (1991. fig. 12, 1-2).
D. praedator : Chen (1979. fig. 1, 5-6) (as D. sinensis). And Dai. Yang & Lan (1981. figs 8-9), Dai et al.
(1986. fig. 12, 3-4), Dai & Yang (1991, fig. 12, 3-4) (as D. tenuilobata).
D. filholi : Monod (1956, figs 84-88).
D. pilumnoides : STEBBING (1921, pi. 14) (as Maxillothrix actaeiformis).
Metadynomene devaneyi: Takeda (1977, text-fig. 1 A-C).
In M. tanensis , some of the sexually mature females have their first pair of pleopods developed as in males
rather than being vestigial, but in all other respects they appear to be normal (see Discussion under this species).
Source :
FAMILY DYNOMENIDAE
457
Fig. 12. — a-c, Dynomene hispida Guerin-Meneville, 1832, 6 11.6 x 9.2 mm. Somalia. Gesira. stn 12. intertidal coral
(MZUF): a. tip of right second pleopod; b. inner surface of exopod and base of left second pleopod showing location
of secretory tegumental gland; c, close up of secretory tegumental gland shown in previous figure. —
d. Dynomene praedator A. Milne Edwards, 1879. 6 10.8 x 8.5 mm, Somalia. Gesira. stn 14. intertidal coral (MZUF):
tip of right second pleopod. — e-f. Dynomene pilumnoides Alcock. 1900. 6 23.5 x 19.0 mm. New Caledonia.
Smib 3, stn 18, 338 m: e, tip of right first pleopod, note that the apical plate has been curled and deformed by
processing for the SEM; f. tip of right second pleopod. (All pictures taken with scanning electron microscope.)
Source :
458
C. L. McLAY
Fig. 13. — a-b, d: Hirsutodynomene spinosa (Rathbun, 1911), 8 19.6 x 14.5 mm, Western Australia, Exmouth Gull,
intertidal (AMS-P191 18): a. tip of left first pleopod; b, right second pleopod; d, tip of left second pleopod. —
c. e-f. Metadynomene tanensis (Yokoya, 1933), 8 16.5 x 15.8 mm. New Caledonia. Smib 3, stn DW 25. 437 m:
c, tip ol second pleopod; e, tip ol left first pleopod. note that the apical plate has been deformed by processing for
the SEM; f, subdistal spines from second male pleopod. (All pictures taken with scanning electron microscope.)
I he first male pleopod consists of two articles: the base of the proximal article forms a flattened plate, lying
beside the genital opening of the coxa, while the distal article is narrower and forms a semi-rolled tube accessed by
the second pleopod on the medial side. I he distal article ends in an oval-shaped flattened plate, borne on the medial
corner, which is surrounded by a dense fringe of long setae. The aperture for sperm delivery lies at the base of the
plate, amongst the setae. The second pleopod is about as long as the first, and is borne on a sternal plate which is
produced anteriorly at the corners. The basal article of the second pleopod bears an exopod and an endopod.
Source : MNHN, Paris
FAMILY DYNOMENIDAE
459
The proximal article of the exopod is very short while the distal article is longer, narrowing to a blunt point and
has a setose lateral margin. The proximal article of the endopod is broad but very short while the distal article is
much longer and narrows quickly to a needle-like shaft. When both pleopods are in their natural position, the
extended coxa ("penis"), bearing the gonopore, opens between the plates formed by the proximal article of the first
pleopod (above) and the flattened basal articles of the second pleopod (below). The exopod of this pleopod, lying
in front of and below the genital opening, has an important role in channeling the sperm from the genital opening
into the base of the first pleopod. Meanwhile, a conical swelling on the extension of the sternal plate at the base
of the second pleopod, closes the posterior limit of the chamber into which the sperm is delivered. The action of
the spines on the anterior surface of the needle-like part of the second pleopod may propel the spermatophores into
the first pleopod where they are delivered through the setose tip into the female spermatheca. The oval medial plate
on the first pleopod may help to guide the sperm into the spermatheca. Thus the exopod of the second pleopod, far
from being vestigial, may be an integral part of the sperm delivery process. Near the base of the endopod of the
second pleopod is the opening of a prominent tegumental gland. The role of this accessory sex gland may be to
provide extra seminal fluid aiding sperm transfer. Minigawa (1993) has reported the presence of secretory glands
in both the first and second pleopods of Ranina ranina. These glands normally secrete mucopolysaccharides and
could also be present in the first male pleopod of dynomenids, but to establish this would require sectioning and
staining. It is interesting to note that in Chionoecetes opilio (Majidae) rosette type accessory sex glands only
appear in mature crabs and are concentrated in the proximal region of the first pleopod, with ducts leading to the
ejaculatory canal (BENNINGER et al., 1995).
Table 1. — Morphological variation in male second pleopods.
In most groups of the Brachyura the male pleopods show wide variation and differences between species
which have proved taxonomically valuable. However there is little variation in structure of the first pleopods
of different dynomenid species. Under the normal light microscope the second male pleopods simply appear
to be needle-like, but examination at higher magnifications, using the scanning electron microscope, reveals
some fine detail and variation in structure. The fine structure of the second pleopods of eight out of the thirteen
species (representing four of the genera) have been examined (see Table 1). The basic plan for dynomenid pleopods
consists of a shaft bearing a row of tiny inset spines running along the length and ending with two or three
Source
460
C. L. McLAY
terminal spines. The main sources of variation are the number of subterminal spines, their disposition to
each other, and their direction. The number of spines ranges from 5 to 24. which can be arranged in a straight line
or variously curving around the shaft axis, and in some cases the spines overlap so that two adjacent spines can
be side by side. In most cases the spines are directed apically but in two species (Dynomene praedator and
Metadynomene tanensis) they are curved towards one side. The differences between the genera are not dramatic:
in Dynomene there tend to be fewer spines (usually directed apically) than in the other genera, whereas
in Metadynomene there are a large number of quite curved spines. In the cases of two pairs of species which,
for other reasons, are believed to be closely related, viz. Dynomene filholi - D. pilumnoides , and
Hirsutodynomene spinosa - H. Ursula , the only difference between them is in the number of subterminal spines.
The species of Hirsutodynomene have a unique arrangement of spines and it is the only genus in which spines
overlap. In spite of its dromiid-like features, Paradynomene tuberculatci has pleopods which are typical of
dynomenids (see Discussion under this species). The second pleopods of two dromiids, Stimdromia lamellata
(Ortmann, 1894) and Epigodromia gilesii (Alcock. 1899), examined in the same way, show no evidence of
ornamentation. The distal part of the second pleopod of Dicranodromia felderi has minute scattered spinules
(Martin, 1990, his fig. 3g). As far as is presently known, the dynomenids are the only dromiaceans with
ornamented second male pleopods.
The tube (i.e. "penis") carrying sperm to the base of the pleopods is well developed in podotremc
crabs. Dromiid males have a long soft penis extending from the coxal article but in dynomenids this is
absent. Instead the corner of the coxal article itself is extended to carry the sperm to the pleopods (sec Fig. 14 f).
Thus the dynomenids could be said to have a calcified "penis" and they share this character with
the homolodromiids (GuiNOT, 1995), although the shape is a little different. Using the implied vertebrate
analogy, the name "penis" is not really very accurate since it is not this structure which is responsible
lor introducing sperm into the female. The so-called "penis" of all crabs is analogous to the vas deferens
and should perhaps be called the "sperm duct". The first male pleopod should be referred to as the pleopod
or "penis".
Male dynomenids have rudimentary pleopods on segments three to five. In the species that have been examined
closely, the last three pairs of pleopods are biramous. Only Dynomene praedator has uniramous pleopods.
The exopod is usually longer and connected to the basal article by a joint. However, the other article is not
jointed and appears to simply be an extension of the basal article. If it is regarded as representing the endopod,
then we must assume that the joint has been lost as a result of fusion. In Metadynomene tanensis both articles
are about the same length and fused to the base. The presence of rudimentary male pleopods is also found
in homolodromiids and some dromiids. GuiNOT (1995) recorded rudimentary pleopods in both Homolodromia
A. Milne Edwards, 1880, and Dicranodromia A. Milne Edwards, 1880 where they varied in size between species,
and sometimes asymmetrically on each segment, but in all cases they were uniramous. Dromiids with rudimentary
male pleopods include Sphaerodromia Alcock. 1899, Exodromidia Stebbing, 1905, and some species of
Dromia Weber, 1795 where they are symmetrical and uniramous (McLay, 1993). Retention of these pleopods in
males must be regarded as a plesiomorphic character.
The first female pleopod is vestigial and consists of a proximal calcified basal article, attached to the sternum,
and a distal article which is soft and flexible, bearing long marginal setae and narrowing to a blunt tip. This
pleopod does not carry eggs and when the abdomen is closed it overlies the sternal suture 7/8 which harbours the
spermatheca. Their proximity to the spermathecae may mean that these pleopods have some role in ensuring that
eggs from the coxal gonopore, and sperm from the spermathecae, come into contact with each other. Since
possession of vestigial first pleopods by females is a character of all podotremes, a similar role could be
hypothesized for these pleopods in other families where there is close proximity of spermathecae, gonoporcs and
first pleopods. This could be true in primitive dromiids like Sphaerodromia and Eodromici for example, but in
more derived dromiids the spermathecae are moved to a much more anterior position in front of the gonopores.
making this liaison unlikely. The other four pleopods in female dynomenids are biramous. the basal article is very
reduced, and both the exopod and endopod consist of six articles. The endopods have long filiform setae for ego
attachment while the exopods have dense fine setae (as on the first pleopod) along the margins for brood
protection.
Source :
FAMILY DYNOMENIDAE
461
Fig. 14. — a, Paradynomene tuberculata Sakai. 1963, 6 22.0 x 22.8 mm, New Caledonia, Smib 3. stn 14. 246 m: tip of
second pleopod. — b, Dynomene praedator A. Milne Edwards, 1879. 6 10.8 x 8.5 mm. Somalia, Gesira, stn 14.
intertidal coral (MZUF): male right fifth pleopod. — c , Dynomene pilumnoides Alcock, 1900. 6 23.5 x 19.0 mm.
New Caledonia, Smib 3, stn 18, 338 m: male left fifth pleopod. — d, Hirsutodytiomene spinosa (Rathbun. 1911).
6 19.6 x 14.5 mm. Western Australia, Exmouth Gulf, intertidal ( AMS-P191 18): male right fifth pleopod. —
e, Metadynomene tanensis (Yokoya, 1933), 6 16.5 x 15.8 mm. New Caledonia. Smib 3, stn DW 25, 437 m: male
left fifth pleopod. — f , Hirsutodytiomene Ursula (Stimpson, 1860), 6 13.4 x 10.3 mm, Mexico, Espiritu Santo Id.
" Velero ", stn 638-37, intertidal: coxal article of right fifth pereopod. (All pictures taken with scanning electron
microscope.)
Source :
462
C. L. McLAY
UROPODS
Compared to other dromiaceans the uropods are very well developed in dynomenids and usually sexually
dimorphic. In Dynomene and Hirsutodynomene uropods are larger in females than in males. Uropods in both male
and female Metadynomene fill the entire margin between the telson and penultimate abdominal segment and in
Paradynomene about half the margin. In some dromiids the uropods are reduced and in some cases vestigial or
absent. When present they have a role in the abdominal locking mechanism but this is not true in dynomenids
because, for the most part, they do not have effective means of locking their abdomen. Compared to dromiids,
dynomenid uropods are plesiomorphic.
Uropods are the uniramous remnant of a biramous limb which formed part of the tail-fan in a distant ancestor.
Reduction of this appendage is associated with reduction in the size of the whole abdomen and its folding beneath
the eephalothorax. There is an obvious trade off between the development of an abdominal locking mechanism and
reduction of the uropods in most crab-like decapods. Dynomenids occupy an intermediate stage in the course of
this transition: there is only minimal restraint of the abdomen. However homolodromiids are an exception:
uropods are always rudimentary, represented by only small ventral lobes (GUINOT, 1995), but they normally
lack an abdominal locking mechanism (except in the case of Dicranodromia felderi Martin, 1990 where the
margins of the telson are held under flanges on the coxae of the chelipeds). In these crabs the abdomen is only
loosely held.
Homolodromiidae Dynomenidae Dromiidae
Dromiacea
Fig. 15. — Cladogram showing the assumed relationships between the families of the Dromiacea. Numbers refer to
apomorphic and plesiomorphic characters mapped on to this hypothesis (see text).
PHYLOGENETIC RELATIONSHIPS
I do not intend to closely examine the question of brachyuran monophyly. In their analysis of the relationships
of replant Decapoda, Scholtz & Richter (1995) list the morphological apomorphies of the Brachyura as: a
fossa orbito-antennularis surrounds the eyestalk and antennule, third maxilliped operculate or semi-operculate,
orientation of the cheliped fingers so that the dactyl is external, all thoracic sternites fused to form the sternum, an
Source : MNHN . Paris
FAMILY DYNOMENIDAE
463
abdomen which is reduced, ventrally flexed and sexually dimorphic, and uropods are reduced or absent. BAUER
(1989) considered the use of setiferous maxillipedal epipods for gill cleaning to be a strong character confirming
brachyuran monophyly. 1 agree with this suggestion, although the more elaborate nature of podotreme gill
cleaning, with the retention of several plesiomorphic characters, does make the explanation more complex.
Proposals to remove some podotreme families from the Brachyura do not seem to be worthy of serious
consideration since the arguments are usually based on the occurrence of plesiomorphic larval characters and if
followed would create even greater mayhem than currently exists. I accept the argument, by Scholtz & Richter,
that evolution has proceeded faster in the adult than in the larval characters so that some podotreme families show
a mosaic of plesiomorphic and apomorphic characters. The morphological apomorphics listed above and
spermatological apomorphies (see Jamieson, 1994; Jamieson et ai, 1995) seem to provide convincing evidence
of brachyuran monophyly. The questions about the relationships within the Brachyura seem much more
interesting.
G U I NOT (1978) divided the Brachyura into three major groups: Podotrcmata, Hetcrotrcmata and
Thoracotremata. The Podotremata were divided into two groups containing the following extant families:
Dromiacea, including the Homolodromiidae. Dynomenidae and Dromiidae, and the Archaeobrachyura, including
the Poupiniidae, Latreil 1 idae, Homolidae, Cyclodorippidae, Cymonomidae and Raninidae. The phylogenetic
relationships between some of these groups has been explored by GUJNOT et al. (1994) using characters based on
spermatozoal ultrastructure. They were able to differentiate between apomorphic and plesiomorphic characters of
the Podotremata, Heterotremata + Thoracotremata as well as the Brachyura as a whole. Within the Podotremata
they provide evidence of monophyly of this group. Using morphological characters I have carried out a similar
exercise. 1 first of all map the plesiomorphic and apomorphic characters of the Dromiacea (Fig. 15) and then
examine the implications of these characters for the Archaeobrachyura (Fig. 16). I assume that the most
parsimonious interpretation of characters is correct. The main question here is whether or not the Podotremata are
monophyletic.
If we assume that the more crab-like Dynomenidae and Dromiidae are sister groups and together are the sister
group of the less crab-like Homolodromiidae, then we can display their relationships as in Fig. 15. The characters
(Al - 19) shared by the ancestor of these three families must be: 1) carapace longer than wide, 2) carapace lacking
a margin, 3) branchiostegite membranous, 4) first antennal article beak-like, 5) pediform third maxillipeds,
6) crista dentata present, 7) propodus of second and third pereopods with a distal spine and a row of spines on the
inner margin of the dactyl, 8) fourth and fifth pereopods reduced, subdorsal, subchelate, 9) propodal and dactyl
spines on last two pairs of pereopods as found in Dicranodromia , 10) fifth pereopod with a spine on the outer
margin of the dactyl, 1 1) abdomen large and only loosely folded under the body. 12) well developed uropods and no
abdominal locking mechanism, 13) five pairs of pleopods in both sexes (first pair vestigial in the female and
last three pairs rudimentary in the male), 14) short female sternal sutures 7/8, 15) needle-like second male
pleopods without spines, 16) calcified coxal sperm lube carrying sperm to the base of the second pleopod,
17) twenty trichobranchiate-like notched gills, 18) long setae on posterior margin of scaphognathite, and
19) seven epipods.
Assumption 9) (above) implies that the dromiacean ancestor was well equipped to carry camouflage. It is
assumed that the last two pairs of pereopods resembled those found in Dicranodromia (Homolodromiidae) but
camouflage-carrying is unknown in this genus (or this family for that matter) (GuiNOT et al.. 1995). Since the
structure of these limbs is very similar to that found in Sphaerodromia (Dromiidae), which does carry pieces of
sponge, 1 predict that camouflage-carrying homolodromiids will be found. Thus I assume that the dromiacean
ancestor was a camouflage crab and that the particular kind of camouflage behaviour is an apomorphy of the
Dromiacea.
The shared characters (B 1 - 7) of the crab-like ancestor of the Dynomenidae and Dromiidae are: 1 ) carapace
margin present, 2) carapace wider than long, 3) operculiform third maxillipeds, 4) abdomen reduced and folded
more tightly under the body, 5) rudimentary abdominal locking mechanism, and 6) development of hypobranchial
cleaning setae on the inner wall of the branchial chamber. An additional character (7) shared by all dynomenids and
some of the primitive dromiids (e.g. Sphaerodromia ) is the presence of an oval apical plate on the tip of the first
male pleopod. The presence of this plate may be correlated with the fact that the females have posteriorly located
464
C. L. McLAY
spermathecae and consequently very short sternal sutures 7/8. The apical plate may help to ensure efficient transfer
of sperm into the spermathecae.
The apomorphies (D 1 - 5) of the Dynomenidae are: 1 ) the lack of distal propodal spines on the second to fourth
pereopods, 2) development of a unique vestigial fifth pereopod carried horizontally. 3) sexually dimorphic chelate
structure of the fifth pereopod, 4) reversion of the fourth pereopod to being a fully developed walking leg and
5) development of a row of spines on the second male pleopod. Perhaps the most controversial point here is the
implied reversion of the fourth pereopod to being a fully functional walking leg. Acanthodromia lacks a beak-like
first antennal article, characteristic of the Dromiacea, and a crista dentata. These must be regarded as secondary
modifications.
The apomorphies (El - 9) of the Dromiidae are: 1) usually only two pairs of pleopods in the male (but some
species of Sphaerodromia , Eodromia and Dromia for e.g. have retained the ancestral three pairs of rudimentary
pleopods on abdominal segments 3-5), 2) soft tube-like extension of coxal article carrying sperm to the base of
first pleopod. 3) spermathecae often located anterior to female genital openings and consequently the sternal
sutures 7/8 are much longer, 4) gills are phyllobranchiate, 5) gill numbers reduced because there are only three or
four epipods (although Sphaerodromia species have up to seven epipods), 6) loss of long setae on posterior margin
ot scaphognathite (still present in Sphaerodromia ), 7) abdomen reduced, 8) uropods reduced (sometimes absent),
and 9) a well developed locking mechanism for the abdomen involving the bases of the first two pereopods and
sometimes the uropods. It is apparent that there are exceptions to several of these characters some of which have
evolved within the Dromiidae. Strictly speaking, the only apomorphies, shared by all members of the family, are
the possession of phyllobranch gills and a well developed abdominal locking mechanism. Since Sphaerodromia
obviously provides several exceptions to the above list of characters, it could be proposed that this genus should
shifted to the Homolodromiidae, but this would require the assumption that phyllobranchiate gills and the coxal
abdominal locking mechanism had evolved independently within this family. By themselves these assumptions are
not necessarily unreasonable because, after all. phyllobranch gills and an abdominal locking mechanism have
evolved independently in Acanthodromia (Dynomenidae), so why couldn't this have also occurred in the
Homolodromiidae? GuiNOT (1979, p. 256) noted that the homolodromiid thoracic cndophragmal skeleton is of a
unique type and is different from the dromiid + dynomenid skeleton. This skeletal difference provides the strongest
evidence for retaining Sphaerodromia in the Dromiidae (GuiNOT, pers. comm.). This hypothesis is more
parsimonious but it requires the assumption that the species in this genus have retained several plesiomorphic
characters. Further aspects of the dromiid-homolodromiid relationship are discussed by GuiNOT (1995: 168-185).
Finally, the Homolodromiidae: the only apomorphic characters (Cl - 5) which this group has are 1 ) reduction
of uropods (GuiNOT, 1993), 2) loss of the long setae from the scaphognathite, 3) presence of well developed
abdominal pleurae, 4) possession of a very elongate telson, and 5) development of a spine-bearing distal propodal
extension on the last two pairs of legs. However, both of the first two are shared with the Dromiidae. It is difficult
to know whether the unusually elongate telson, which forms the floor of the sterno-abdominal cavity, and the
abdominal pleurae are apomorphies or plesiomorphies. Along with the membranous branchiostegite, they may
well have been features of the dromiacean ancestor and therefore plesiomorphies. The development of a spine¬
bearing distal propodal extension on the last two pairs of legs might be regarded as an apomorphy, but it only
occurs in Homolodromia while Dicranodromia retains the assumed ancestral condition. The semi-crab-like
Homolodromiidae can only be defined by a combination of plesiomorphic characters and synapomorphies. It is
interesting to note that Jamieson et a! (1995) concluded that "Homolodromia displays a remarkable mixture of
dromiid and dynomenid spermatozoal features while lacking any distinctive apomorphy.... ". Therefore the
morphological and spermatological features of the Homolodromiidae are in close agreement.
The question of monophyly of the Podotremata is more difficult to decide. Arc the Dromiacea the sister group
of the rest of the Brachyura (see Fig. 16 b) or are they the sister group of only the Archaeobrachyura (see
Fig. 16a)? II the first alternative is true then the Dromiacea and each of the major groups within the
Archaeobrachyura must be independently derived from the brachyuran line and all these crab-like animals must be
paraphyletic. Another possibility is that the Archaeobrachyura are monophyletic and are the sister group
Source : MNHN. Paris
FAMILY DYNOMENIDAE
465
Homolodromiidac Dvnomenidac Dronnidae Poupiniidae Latrcllidae Homolidac Cyclodorippidac Raninidac
Eubrachyura
Fig. 16. — Cladogram showing two different relationships between the families of the Podotremata: a. the Podotremata
are assumed to be monophyletic (modified after Guinot et a I ., 1994); b. the Podotremata are assumed to be
paraphyletic (modified after SCHOLTZ & Richter, 1995). Numbers refer to apomorphic and plesiomorphic characters
mapped on to these hypotheses (see text).
of the Eubrachyura. Since it is beyond the scope of this work to examine the detailed relationships within the
Archaeobrachyura, I confine my attention to the case represented in Fig. 16b. JAMIESON (1994) and JAMIESON et
al. (1995) have argued that spermatologically the Podotremata Guinot. 1977 is monophyletic and its constituent
groups, the Dromiacea de Haan, 1833 and Archaeobrachyura Guinot. 1977 are also monophyletic.
The sperm data seem to support the monophyletic hypothesis depicted in Fig. 16a. If this is true then we must
examine the suite of ancestral dromiacean characters to see which of them could be ancestral to all of the
Podotremata and which could be apomorphies of the Dromiacea (see Fig. 16a). The ancestral characters (FI - 14)
would include: 1) carapace longer than wide, 2) carapace lacking an anterolateral margin, 3) branchiostegite
membranous, 4) third maxillipeds pediform, 5) crista dentata present. 6) abdomen large and only loosely folded
466
C.L. McLAY
under the body, 7) uropods well developed, no abdominal locking mechanism, 8) five pairs of pleopods in both
sexes, 9) short female sternal sutures 7/8, 10) twenty trichobranchiate-like notched gills. 1 1 ) long setae on the
posterior margin of the scaphognathite, and 12) seven epipods. To these can be added 13) the coxal position of the
genital apertures in both sexes and 14) separate spermathecae in the female sternum. The remaining characters (G 1
- 7): 1) the beak-like first antennal article, 2) propodus of second and third pereopods with a distal spine and a row
of spines on the inner margin of the dactyl, 3) fourth and fifth pereopods reduced and subdorsal with 4) propodal
and dactyl spines as found in Dicranodromia , 5) fifth pereopod with a spine on the outer margin of the dactyl, 6)
needle-like second male pleopods without spines, 7) calcified coxal sperm tube carrying sperm to the bases of the
second pleopod, must be apomorphies of the Dromiacea. The assumption of monophyly implies, amongst other
things, that phyllobranchiate gills, and the mechanisms for locking the abdomen have been independently evolved
in the Archaeobrachyura and Dromiacea, and that the ancestor of the Podotremata had their last two pairs of legs as
walking legs and not reduced. Therefore this ancestor did not carry camouflage materials. Camouflage behaviour
evolved independently in the Dromiacea (e.g. Dromiidae, see McLay, 1993) and in the Homoloidea. In the
Homoloidea camouflage probably only occurs in the Homolidae (for details see GuiNOT et al.. 1995). Furthermore
the hypothesis implies that the uropods have been reduced or lost independently in the Homolodromiidae and the
Dromiidae and that operculate third maxillipeds were independently evolved in the dynomenid-dromiid line and the
Homoloidea.
In a tentative phylogenetic analysis, SCHOLTZ & Richter (1995) have argued that the dromiaceans (sensu
Borradaile, 1907) are not a monophyletic assemblage (see Fig. 1 6b) and that the homolids have a closer
relationship with the "higher" brachyurans. In their view the homolodromiids (with trichobranchiate gills, narrow
cheliped sternite, and an elevated third maxilliped sternite) are the sister group of all other brachyurans, and the
homolids (with phyllobranchiate gills, wide cheliped sternite, and non-elevated third maxilliped sternite) are the
sister group of the dromiids and the "true" brachyurans. This latter group share the homolid characters as well as
having truly operculiform third maxillipeds and an elongate gill-cleaning first maxilliped epipod. Thus SCHOLTZ
& RICHTER (1995) tentatively propose that the Podotremata are not monophyletic but are paraphyletic. This
would imply that some or all of the characters (FI -14) are plesiomorphies for the Dromiacea, Archaeobrachyura
and the Eubrachyura and therefore characters of the ground pattern of the Brachyura.
The points of conflict between these two hypotheses are. amongst other things, different interpretations of the
origin of phyllobranchiate gills, and operculate third maxillipeds. Given the variation in gill structure within the
Dynomenidae it does not seem to be necessary to assume that phyllobranchiate gills only evolved once.
Phyllobranchiate gills are found in Acanthodromia and all the Dromiidae but it is clear that both the dynomenids
and dromiids must be derived from an ancestor with multi-lobed gills. By the same token operculate third
maxillipeds could well have been independently derived from pediform appendages with a crista dentata. SCHOLTZ
& Richter (1995) make the presence of a crista dentata the apomorphic condition of the Eureptantia Scholtz &
Richter, 1995. It should be noted that there arc many examples amongst the Brachyura where the crista dentata is
absent. It has been lost in some Dynomenidae (e.g. Acanthodromia ), in all Cyclodorippidae (e.g. Tymolus ,
Xeinostoma , and Krangalangia ), Latreillidae (e.g. Latreillia ), Raninidae (e.g. Lyreidus , Ranina , Raninoides), and
all of the Eubrachyura. The polarity of this character depends upon what assumptions are made about the ancestral
decapod, and whether the absence of a crista dentata in natants is ancestral or derived. Scholtz & Richter (1995)
argue that the absence of the crista dentata in some Achelata Scholtz & Richter, 1995, (e.g. Scyllarus ),
thalassinids (e.g. Callianassa) and anomolans should be regarded as being "secondary". Loss of the crista dentata is
a synapomorphy of many groups within the Eureptantia.
The paraphyletic hypothesis also assumes that camouflage carrying was ancestral to all the Brachyura whereas
the monophyletic hypothesis assumes that it is a synapomorphy of only a few of the podotrematous families.
McLay (1991: 465) put forward an hypothesis about how the camouflage-bearing limbs of dromiids might
have evolved from walking legs. This hypothesis needs to be modified in the light of the hypothesis presented
above that the ancestor of the Dromiacea was a camouflage-carrying crab. Thus the argument presented by McLay
(1991) should be applied to the dromiacean ancestor rather than the dromiid ancestor. Although further analysis of
this complex question is required, the weight of evidence seems to favour monophyly of the Podotremata.
Source : MNHN, Paris
FAMILY DYNOMENIDAE
467
Which ever hypothesis is accepted, there is always going to be a problem with interpreting the evolution of
the fourth pereopods. Using the characters of the last two pairs of pereopods by themselves it would be natural to
group the homolodromiids and dromiids together because they have reduced fourth and fifth pereopods and to group
the dynomenids and the homolids together because they have normal fourth pereopods, used for walking, and only
the fifth pereopods reduced. But this grouping overlooks the fact that these modified limbs have different roles. In
homolids the last pair of pereopods are subdorsal and used to carry anemones but in dynomenids they are
horizontal, vestigial and probably had a cleaning function. In homolodromiids and dromiids the probable reason for
having both pairs of limbs reduced is because of their camouflage carrying role (this has yet to be confirmed for
homolodromiids). Given the hypothesized Homolodromiidae - Dynomenidae + Dromiidae link (see Fig. 15) it is
most parsimonious to assume that their ancestor had both of the last two pairs reduced, but this requires that in
the ancestral dynomenids the fourth pereopods reverted to a locomotory role and that the fifth pereopod became a
cleaning limb. Provided that we derive the homolids from an ancestor with four normal walking legs, there is no
great difficulty in hypothesizing that only the last pair of pereopods was modified for the specialized task of
carrying anemones. Pereopodal grooming in decapods involves several different limbs and it seems reasonable to
regard each different case as apomorphic.
Wherever we place the dynomenids amongst the extant groups, there is always going to be a problem with
interpreting the evolution of the last two pairs of pereopods. Perhaps the reason for the apparently uncomfortable
position of the dynomenids is that their closest ancestors or sister group arc in fact extinct, and are to be found
somewhere amongst the numerous "prosopid" species which have been described. It would be very helpful if we
knew something about the limbs of the extinct dynomenid species. Unfortunately, in most cases, we only have
information about their carapace and know nothing about their pereopods.
Another aspect of the phylogeny of the Dromiacea (i.e. Homolodromiidae, Dynomenidae and Dromiidae)
which warrants discussion is the conflict between sperm, and 18S rRNA data, and the accepted allocation of
genera to families (Jamieson et al., 1995). Briefly, in a parsimony analysis using PAUP. Homolodromia kai
(Homolodromiidae). Paradynomene tuberculata and Metadynomene tanensis (= Dynomene aff. devaneyi)
(Dynomenidae), Stimdromia lateralis and Dromidiopsis edwardsi (Dromiidae) do not show a relationship which
matches their familial position (JAMIESON et al ., 1995). Furthermore an analysis based on I8S rRNA suggests
that the dromiid Hypoconcha arcuata has anomuran affinities rather than being linked to another dromiid,
Cryptodromiopsis antillensis (= Dromidia antillensis) (Spears et al ., 1992). The dromiids are certainly a
morphologically diverse group, more so than the dynomenids, but the apomorphies listed above seem to provide
convincing evidence that the Dromiidae are in fact a monophyletic group. It may well be that "the sperm never
lie” (as claimed by some), but spermatological data certainly can be ambivalent and open to as many
interpretations as conventional morphological characters.
Finally, it has been suggested by some decapod palaeontologists (e.g. Wright & Collins. 1972.
and GLAESSNER, 1980) that the family Xanthidae was derived from amongst the Dynomenidae. Admittedly,
some extant dynomenids do resemble some xanthids (e.g. Pilumnus and Panopeits), in their chelipeds and
the conformation of their carapace (sec Discussion below under Hirsutodynoniene Ursula ), but it should be clearly
apparent from the above arguments that any resemblance of xanthids and dynomenids must be convergent and not
evidence of a close relationship. It may be significant that at least some members of each of these families inhabit
corals, so that their similarities may be attributable to colonization of the same habitat.
Family DYNOMENIDAE Ortmann, 1892
Dynomenidae Ortmann 1892: 541; 1898: 1155. — Alcock 1899: 127; 1901: 34. — Stubbing. 1905: 58. — Rathbun.
1937 : 51. — Balss, 1957: 1605. — Glakssner. 1969: R487. — Wright & Collins, 1972: 48. — Takf.da. 1973:
80. — Sakai, 1976: 28. — Guinot, 1993: 1226.
Dynomeninae A. Milne Edwards & Bouvier. 1899: 9; 1902: 22.
468
C. L. McLAY
Carapace shape usually wider than long, but can be longer than wide, generally moderately convex, commonly
subcircular, ovoid or may be oblong. Surface may be smooth, spinous or areolate and is usually densely covered
with setae. Lateral carapace margin usually well defined and armed with distinct teeth. Frontal groove well marked,
split in two posteriorly, cervical, postcervical and branchial grooves evident. Frontal carapace margin broadly
triangular, continuous, and usually without rostrum or teeth. Eyestalks short, eyes protected by well defined
orbits. Sternal sutures 7/8 of female end well apart on low tubercles behind bases of second walking legs.
Antennule can be concealed inside the orbit at the base of the eyestalk. Antennal flagella shorter than carapace
width. All articles of antenna moveable, first article (urinal) usually beaked medially and second article has an
exopod firmly fixed. Third maxillipeds opercular, completely covering the buccal cavern, separated at their bases
by a plate at the same level as the sternum; basis and ischium of endopod fused but joint always marked by a
shallow groove. Percopods include chelipeds, three pairs of walking legs, and reduced last pereopods. Chelipeds
equal, stouter than walking legs, last pair of legs very reduced, dactyl rudimentary, forming an obsolete subchelate
mechanism with an extension of the propodus. Gills usually 19 (including 6 podobranchs) + 7 epipods. Gill
structure basically phyllobranchiate but the plates are very variable in shape with different numbers of epibranchial
lobes.
Abdomen of six segments and telson, folded loosely under the thorax, uropods large, an effective abdominal
locking mechanism usually absent. Both sexes have five pairs of pleopods, first pair vestigial in female, last three
pairs rudimentary in the male. Male first pleopods very uniform in structure, consisting of a stout, setose semi-
rolled tube with an apical plate, second pair needle-like bearing tiny inset spines, termination with two or more
stouter spines.
Type Genus. — Dynomene Desmarest, 1823 designated by Ortmann (1892). Peyrot-Clausade &
Serene (1976) attribute the first use of the latinized name Dynomene to Desmarest (1825), but Manning &
Holthuis (1981) have shown that it should be attributed to DESMAREST ( 1823) where a latinized version was
used in the index to his article in the "Dictionnaire des Sciences Nature lies”.
DISCUSSION. — The above definition of the family Dynomenidae encompasses all the characters listed by
Ortmann (1892). When he defined the family, Ortmann was chiefly interested in separating it from the
Dromiidae, and Homolidae. ORTMANN regarded the following characters as being primitive: 1) incomplete
connection of the pterygostomial region and the epistome, 2) margin of carapace clearly defined. 3) fifth pcrcopod
small and simple with rudimentary dactyl. 4) uniramous uropods present, and 5) four mastigobranchs (i.e. epipods)
and four pleurobranchs on the pereopods, while on all the thoracic segments there are six rudimentary
podobranchs. Other characters which he regarded as important, but not necessarily primitive, were that the eyes
could be completely withdrawn into the orbits, the antennule could be folded away into a groove between eyestalk
and supraorbital margin, and finally the third and fourth articles of third maxillipeds are slightly widened while the
fifth, sixth and seventh articles arc significantly smaller. In fact all of these characters, except for the reduced fifth
pereopods and presence of four epipodites, are shared with the Dromiidae Dc Haan, 1833. Thus it is not surprising
that some dromiids have initially been described as dynonienids (see below).
Perhaps through an over-sight, ORTMANN (1892) did not include Acanthodromia Milne Edwards, 1880 in his
new family. A. MILNE Edwards (1880) originally placed Acanthodromia in his family "Dromiens" which
included Dromici Weber, 1795, Dromidia Stimpson, 1858, and Dicranodromia Milne Edwards, 1880. He considered
that Acanthodromia should be placed between Dromia and Dynomene. ALCOCK (1899) placed Acanthodromia in
the Dromiidae along with Dromia and Arach nodro mia. However. ALCOCK (1901) was the first to put
Acanthodromia in the Dynomenidae, along with Dynomene , and to provide a formal definition of the family. He
expanded the family definition so as to include Acanthodromia and simply commented that it differed from
Dynomene in that its carapace was longer than wide, convex, and closely covered with spines instead of setae.
ALCOCK also noted that dynomenid gills are phyllobranchiae but sometimes showing the transition from tricho-
to phyllobranchiae. A. Milne Edwards & Bouvier (1899: 10) believed the branchial formula to be the same as
that of Homarus vulgaris. Wright & COLLINS (1972) suggested that Acanthodromia should be placed in the
fossil family Prosopidae Von Meyer, 1860. The proposal is discussed further in the section on this genus.
Source : MNHN, Paris
FAMILY DYNOMENIDAE
469
Stebbing (1905) added Dynomene platyarthrodes , from South Africa, with the idea that it was intermediate
between Dynomene and Acanthodromia , believing that the characters of the front, the orbits, and the antennae were
sufficiently similar to Dynomene filholi to justify inclusion. He did not modify or provide a definition of the
family. However Balss (1938) noted that the fourth pereopods were subchelate and the female sternal sutures 7/8
extended as far as the chelipcd segment and realized that D. platyarthrodes in fact belonged to the Dromiidae.
BARNARD (1947) erected the genus Speodromia Barnard, 1947, for this crab (see McLay, 1993).
In her review Rathbun (1937) essentially restated the family definition of ALCOCK (1901) except that she
noted that the gills were phyllobranchiate, eliminating any reference to the gills being "transitional". This is true
of both species of Acanthodromia , but the other dynomenids are different. BALSS (1957) gave the same features as
ALCOCK (1901).
GLAESSNER (1969) gave a family definition which highlighted the features preserved in fossil dynomenids,
concentrating on the orbits, carapace shape and incised grooves, and added the intercalated lateral platelets (i.e. the
uropods) and the essential character of the reduced last pair of legs, even though these are most unlikely to ever be
preserved in a fossil. The same definition was repeated by Wright & Collins (1972). Carapace grooves have had
little importance in the description of modern species but they assume more importance in fossil species.
The last genus to be added was Paradynomene Sakai, 1963, but Sakai never modified the definition of the
Dynomenidae to accommodate this new form, and Sakai (1976) simply repeated the definition of Rathbun
(1937). The only substantial change that is necessary, is to include a very arcolate carapace surface. Takeda
(1973) noted that the dynomenids are distinguished from the Dromiidae by having an epipod on each of the first
three pairs of walking legs and only the last pair of legs small and subdorsal.
Therefore, the modern definition of the Dynomenidae owes a lot to ALCOCK (1901). I have added the character
of three rudimentary pleopods in males because it seems to be true of all dynomenids so far examined. It is
interesting to note that the same condition is found in some dromiids: Sphaerodromia Alcock. 1899, Eodromia
McLay, 1993, and Exodromidia Stebbing, 1905 (see McLay, 1993).
Cano (1893) described a zoea larva which he assigned to ? Dynomene Dcsmarest, but Williamson (1965)
stated that it was more likely that this larva belonged to Blepharipoda Randall or a closely allied genus of the
Albuneidae. The only dynomenid larva known is a pre-zoea dissected from late stage eggs of Acanthodromia
erinacea by RICE (1981).
The genera of the Dynomenidae have enjoyed a fairly stable existence. Only four generic names have been used
to group the species in this family. Dynomene Desmarest, 1823 was the first to be established followed by
Acanthodromia Milne Edwards, 1880, Maxillothrix Stebbing, 1921, and Paradynomene Sakai, 1963. Both
Acanthodromia and Paradynomene are very distinctive and consequently have not caused any taxonomic problems.
However species have been added to Dynomene in a rather haphazard way. without reference to the generic
definition, and some revision is required if all three genera are to have equal status. Maxillothrix was shown by
Odhner (1925) to be a junior synonym of Dynomene (see Discussion below).
Identification keys to genera and/or species can be found in ALCOCK (1901), Sakai (1936. 1965, 1976),
Rathbun (1937), and Dai & Yang (1991). Peyrot-Clausade and Serene (1976) give a key to five Indo-
Pacitic species of Dynomene. The characteristics which they used in their key were: carapace surface smooth,
spinous, tuberculate or granulate, number and size of teeth on anterolateral border, tomentum length and
clumping, spines on anterior border of P2-4, length/width ratio of P3 merus, presence of spines on borders of
orbit, presence of a toothed lobe on the cheliped carpus. Below I present a key to all known species of extant
dynomenids.
Key to the species of the family DYNOMENIDAE
1. Carapace width less than length; surface largely devoid of setae, strongly tuberculate or
densely covered with long sharp spines . 2
— Carapace width greater than length; surface setose to varying degrees, not strongly
tuberculate and without long spines . 4
Source :
470
C.L. McLAY
2. Carapace surface densely granulated and strongly tuberculate; rostrum tridentate. median
tooth on a lower level, lateral teeth at beginning of supraorbital margin; anterolateral
margins with six irregular teeth . Paradynomene tuberculata Sakai, 1963
— Carapace surface densely covered with long spines; rostrum terminated by a strong spine;
anterolateral margins with numerous spines . 3
3. Supraorbital spines near corner of the orbit arc long and bent posteriorly; fourth
abdominal segment with a small, median, pearl-like lobe only partially divided by a
short median groove; a similar smaller lobe on the fifth segment .
. Acanthodromia erinacea A. Milne Edwards, 1880
— Supraorbital spines near corner of orbit are short, blunt and not bent posteriorly; fourth
abdominal segment with a pair of large, smooth pearl-like median lobes separated by a
groove and occupying almost the entire width of the segment; fifth segment
spinous . Acanthodromia margarita (Alcock. 1899)
4. Carapace width only slightly greater than length (ratio < 1.10). densely covered with
short, soft setae which give the surface an uneven, undulating appearance, with
transverse troughs; no long setae on the carapace; dactyl of PI not strongly curved;
margins of fingers touching for about half their length; less than five spines on inferior
margin of P2-P4 dactyli . 5
— Carapace width much greater than length (ratio > 1.10); long and short setae present on
carapace; dactyl of PI strongly curved; fingers touching only at the tips; five or six
spines on inferior margin of P2-P4 dactyli . 7
5. Anterolateral carapace margin without teeth but interrupted by a faint notch mid-way
between postorbital corner and where the branchial groove meets the margin .
. Metadynomene devaneyi (Takeda, 1977)
— Anterolateral carapace margin with teeth . 6
6. Three well developed, unequal (posterior margin of second tooth extended and may bear
two smaller denticles) and blunt anterolateral teeth; strong posterolateral tooth behind
branchial groove; suborbital margin shelf-like, projecting and easily visible
dorsally . Metadynomene tanensis (Yokoya, 1933)
— Four tiny subacute anterolateral teeth, first pair separated from second pair by a blunt
swelling, similar posterolateral tooth behind branchial groove; suborbital margin not
projecting, scarcely visible dorsally . Metadynomene crosnieri sp. nov.
7. Carapace surface areolate, granulate and spinous (especially near margins) under the
surface tomentum . . . 8
— Carapace surface smooth or only minutely granulated; tomentum may be sparse or dense
in which case the setae are short and bent at right angles near the lip . 9
8. Tomentum consists of a dense cover of filiform long setae, arranged in clumps
associated with areolae or spines, and a dense understory of short serrated setae bent at
right angles near the tip; projection on inner carpal margin of chelipcd consists of
a sharp spine; suborbital margin bears small acute spines .
. Hirsutodynomene spinosa (Ralhbun, 1911)
romentum consists of a sparse cover of long and short, slightly clumped, serrate setae,
which are unbent; projection on inner carpal margin of cheliped is a broad blunt lobe;
suborbital margin bears small blunt granules .
. Hirsutodynomene Ursula (S timpson, I860)
9. Anterolateral teeth absent or only represented by two or three small granules not
terminated by a sharp tooth . Dynomene praedator A. Milne Edwards, 1879
— Anterolateral teeth present, well developed, and sharply pointed . 10
Source : MNHN , Pans
FAMILY DYNOMENIDAE
471
10. Carapace tomentum sparse, setae filiform, surface not obscured; ratio of length of
antennal flagellum to CW > 0.60; carpus and propodus of PI smooth; ratio of length of
merus of P3 to CL > 0.7 . Dynomene pugnatrix de Man, 1889
— Carapace tomentum not sparse, setae serrate, long setae may be arranged in clumps; ratio
of length of antennal flagellum to CW < 0.60; carpus and propodus of PI granulated;
ratio of length of merus of P3 to CL < 0.7 . 11
1 1. Carapace tomentum consists of dense short setae, bent at right angles, obscuring the
surface, and fifteen to seventeen tufts of long (> 0.2 x CW) setae; ratio of length to
width of merus of P3 > 2.0 . Dynomene pilumnoides Alcock, 1900
— Short setae not obscuring carapace surface, long setae may be arranged in clumps but
length < 0.2 x CW; ratio of length to width of merus of P3 < 2.0 . 12
12. Carapace surface smooth, coarse serrate setae, longer setae arranged in about twenty
clumps, ratio of CW to CL approx. 1.2; notch present in supraorbital margin; no spines
on postorbital margin; cervical groove branching off subhepatic groove; granules on
carpi of P2-P4 not arranged in rows . Dynomene filholi Bouvicr, 1894
— Carapace surface minutely granulated, coarse serrate setae, longer setae not arranged in
clumps, ratio of CW to CL approx. 1.3; no notch in the supraorbital margin; five small
acute spines around postorbital margin; no cervical branch from the subhepatic groove;
granules on carpi of P2-P4 arranged in three rows .
. Dynomene hispida Guerin-Meneville, 1832
Genus DYNOMENE Desmarest, 1823
Dynomene Desmarest, 1823: 252, pi. (18) fig. 2; 1825: 133. pi. 18. fig. 1. — Latreille, 1825: 273; 1829: 69.
Dynomene Desmarest, 1823: 422; 1825: 442. — Jarocki, 1825: 26. — Berthold. 1827: 258. — H. Milne Edwards,
1837: 179. — Stimpson, 1858: 226. — A. Milne Edwards. 1879: 1; 1899: 90. — Alcock. 1899: 133; 1901:35.—
Ortmann. 1898 : 1 155. — Stebbing. 1905: 58. — Rathbun, 1937: 54. — Balss. 1938: 6. — Sakai, 1936: 43;
1965: 12; 1976: 29. — Barnard, 1947: 371; 1950: 336. — Takeda, 1973: 80; 1977: 31. — Manning & Holthuis,
1981: 23. — Dai & Yang, 1991: 31.
Dynomena Eydoux & Souleyet. 1842: 239 (err.).
Mcixillothrix Stebbing, 1921: 456 (type species Maxillothrix actaeiformis Stebbing, 1921, a subjective junior
synonym of Dynomene pilumnoides Alcock, 1900, by monotypy, gender feminine).
Diagnosis. — Carapace shape wider than long, moderately convex, commonly subcircular. Surface may be
smooth or sparsely granulate, covered with coarse setae, which may short or long, and often arranged in tufts.
Lateral carapace margin always well defined and armed with distinct small teeth or granules. Frontal groove well
marked, split in two posteriorly; cervical, postcervical and branchial grooves usually evident. Frontal carapace
margin broadly triangular, continuous; no rostrum or teeth. Eyestalks short; eyes protected by well defined orbits.
Sternal sutures 7/8 of female end well apart on low tubercles behind bases of second walking legs.
Antcnnule can be concealed inside orbit at base of eyestalk. Antennal flagella shorter than carapace width. All
articles of antenna moveable; first article (urinal) always beaked medially and second article with an exopod firmly
lixed. Third maxillipeds opercular completely covering buccal cavern, separated at their bases by a plate at same
level as sternum; basis and ischium of endopod fused but joint always marked by a shallow groove. Crista dentata
present. Chelipeds equal, stouter than walking legs; dactyl strongly curved; fingers gaping basally. Last pair of
legs very reduced; dactyl rudimentary, forming an obsolete subchelate mechanism with an extension of propodus.
Gills usually 19 (including 6 podobranchs) + 7 epipods. Gills variable in shape.
Abdomen of six segments and telson folded loosely under thorax; uropods large. No effective abdominal
locking mechanism. Lateral movement of abdomen restricted by small sternal tubercle, at base of each of first
walking legs, which lies alongside each uropod. In both sexes, five pairs of pleopods; first pair vestigial in
female; last three pairs rudimentary in male. First male pleopods very uniform in structure, consisting of a stout.
Source
472
C.L. McLAY
setose semi-rolled tube with an apical plate; second pair simple, needle-like, with varying numbers of subterminal
spines.
Fig. 17. — a. Dynomene hispida Guerin-Meneville, 1832, 6 9.2 x 7.6 mm, Madagascar, Tulear, sin 13-3, reef flat
(MNHN-B 22087): dorsal view of whole crab, setae removed from right half of carapace, right second pereopod
missing. — b, Dynomene praedator A. Milne Edwards, 1879, 6 10.4 x 8.2 mm, New Caledonia. lie des Pins,
stn 585, 43 m: dorsal view of whole crab. — c, Dynomene filholi Bouvier, 1894, 9 ovig. 10.4 x 8.0 mm. Gulf of
Guinea, Annobon Id. 35-55 m (MNHN-B 22093): dorsal view of whole crab. — d, Dynomene pilumnoides Alcock.
1900, 9 ovig. 12.7 x 10.6 mm. Sulu Archipelago, 99-108 m, (MNHN-B 10374): dorsal view of whole crab, left
fourth pereopod and right fifth pereopod are missing. — e. Hirsuiodynomene spinosa (Rathbun, 1911), 6 17.5 x
14.0 mm. Gorong, East of Seram, Rumphius II (MNHN-B 9906): dorsal view of whole crab, some setae removed from
right half of carapace. — f. Hirsuiodynomene Ursula (Stimpson, 1860), 9 15.0 x 12.3 mm, Galapagos, Hood Id,
Hancock Galapagos Expedition, stn 30-33 (ex USNM 68313 but gifted to MNHN, Paris): dorsal view of w'hole crab.
Source :
FAMILY DYNOMENIDAE
473
Type Species. — A genus without included nominal species; type species Dynomene hispida Guerin-
Meneville, 1832, by subsequent monotypy, gender feminine.
Other Species. — Dynomene filholi Bouvier, 1894, D. pilumnoides Alcock, 1900, D. praedator A. Milne
Edwards, 1879, D. pugnatrix de Man, 1889 (including D. pugnatrix brevimana Rathbun, 1911).
DISCUSSION. — There has been considerable confusion over the authorship of the generic name Dynomene
because at the outset it was sometimes used in the vernacular and other times in the latinized form. In
DESMAREST's (1823, 1825) texts (and on the plates) the name Dynomene is given in the vernacular (French). In
the indices to these two papers (the second paper being only a new version of the first), the name, however, is
given in latin. In the 1823 index the latin names are italicized and the vernacular names are in roman type, e.g.
"Langouste (Voy. Palinurus)". This is confirmed in the 1825 index, that starts with "Nota. Les noms latins dcs
genres sont en italiques". Other uses of the latin Dynomene are found in Jarocki's (1825) general zoology book
and Berthold's (1827) German translation of Latreille's (1825) "Families naturclles du Regne Animal". (I am
indebted to Prof. L. B. Holthuis for the preceding information.) Thus Desmarest ( 1823: 422) is the author of
the genus Dynomene which was first used in the index to this publication.
There has also been confusion about the authorship of the type species of the genus Dynomene. Like many
authors before them, PEYROT-CLAUSADE and SERENE (1976) attributed the name Dynomene hispida to
Desmarest (1825) but they point out that it was Guerin-Meneville who first used the full latinized form in
his ” l cono graphic du regne animal de G. Cuvier", published between 1829-1844. MANNING and HOLTHUIS (1981)
state that livraison 22 ot this series was published in July 1832. The latinized form was first used on pi. 14 of
Guerin-Meneville ( 1 832).
Maxillothrix Stebbing, 1921 was erected for a supposed new species of the Xanthidae, collected from the Cape
Region, South Africa. But ODHNER (1925: 85) established that Maxillothrix , type species Maxillothrix
actaeiformis Stebbing, 1921, is not a xanthid, but in fact a dynomenid. which he recognized as being a species of
Dynomene. Subsequently, BARNARD (1947) placed M. actaeiformis in synonymy with Dynomene pilumnoides
Alcock, 1900.
Dynomene hispida Guerin-Meneville, 1832
Figs 3 a, 5 a-b, 11,12 a-c, 17 a, 18 a-g
Dynomene hispide Desmarest, 1823: pi. (18). fig. 2; 1825: 432, pi. 18. fig 2. — LATREILLE, 1829: 69.
Dynomene hispida Guerin-Meneville, 1832: 10, pi. 14, fig 2. — GRIFFITH, 1833: 175, pi. 14, fig. 2. — H. Milne
Edwards, 1837: 180; 1848 (in Cuvier): 180, pi. 14. fig. 2. — Lamarck, 1838: 482. — A. Milne Edwards. 1879:
5, pi. 12, figs 1-9, pi. 13, figs 10-15. — Richters, 1880: 158. — Miers, 1884: 13. — De Man. 1888: 408. —
Ortmann, 1892: 543; 1894: 33. — Alcock, 1901: 74 (list). — Nobili. 1907: 378. — Rathbun. 1911: 195. — Ihle,
1913: 92 (list). — Bouvier, 1915: 38. — Balss, 1922: 105; 1938: 7. — Edmondson, 1925: 30; 1933: 265,
fig. 141; 1946: 269, fig. 165. — Sakai, 1936: 43, pi. 8, fig 3; 1940: 29 (list); 1965: 660, text-fig. 1130; 1976:
29, pi. 6. fig. 4. — Buitendijk, 1939: 227. — Miyake. 1939: 198 (list); 1983: 195 (list). — Horikawa. 1940: 28.
— Ward, 1942: 71. — Lin, 1949: 12. — Tweedie. 1950: 106. — Guinot, 1967: 241 (list); 1985: 448 (list). —
Suzuki & Kurata. 1967: 95. — Serene. 1968: 36 (list); 1973: 119. — Takeda. 1973: 80; 1977: 35 (list). —
Peyrot-Clausade & SerLne, 1976: 1340, fig. I, pi. 5 A-B.F. — PEYROT-CLAUSADE, 1977: 212; 1981: 750; 1984:
114. — Guinot. 1979: 125, pi. 21. figs 8-9. — Chen, 1980: 118, text-fig. 1. pi. 1, fig. 3. — Highsmith. 1981:
369. — Baba, 1986: 310. — Dai, Yang, Song & Chen, 1986: 28. pi. 3-4, text-fig. II. 1-4. — Garth. Haig, &
Knudsen, 1987: 241. — Nagai & NomurA, 1988: 92. — Nagai, 1989: 43. — Dai & Yang, 1991: 32. text-fig 1 1.
pi. 3, fig. 2. — Poupin, 1996a: 24 (list).
Dynomena latreillii Eydoux & Souleyet, 1842: 239. pi. 3, figs 3-5. — Takeda, 1977: 35 (list).
Dynomene granulobata Dai, Yang & Lan 1981: 119, figs 10-14. — Dai, Yang. Song & Chen, 1986: 29,
text-fig. 12, 1-2, pi. 3, fig. 3. — Dai & Yang, 1991: 33, text-fig. 12, 1-2. pi. 3, fig. 3.
Not Dynomene hispida - De Man, 1902: 689 [= Hirsutodynomene spinosa (Rathbun, 1911)].
Not Dynomene hispida - Yokoya. 1933: 95, text-fig. 37 \= D. pilumnoides Alcock, 1900|.
MATERIAL EXAMINED. — New Caledonia. No locality, probably intertidal, 1873: 1 6 15.2 x 11.3 mm;
1 9 ovig. 13.7 x 10.1 mm (MNHN-B 22086). — No locality, probably intertidal. M. Balansa coll., 1873: 7 6 7.7 x
Source
474
C. L. McLAY
6.1 - 12.0 x 8.9 mm; 6 9 8.2 x 6.6 - 10.9 x 8.5 mm; 1 9 ovig. 13.0 x 10.0 mm (MNHN-B 22091). — No locality, leg.
A. Milne Edwards, probably part of the M. Balansa collection: 1 6 13.0 x 10.4 mm; 1 9 7.6 x 6.0 mm (ZMB 4324).
Mauritius. Port Louis, probably intertidal. M. Thirioux coll., 1913, E. BOUVIER det.: 1 9 ovig. 12.9 x 9.9 mm
(MNHN-B 22088). — No date or locality: 1 9 10.3 x 8.4 mm, dry (ANSP-CA331 5, originally part of the collection of
Gu£rin-M£neville, No. 209).
Madagascar. West coast. Tulear, stn 13-3, external reef Hat. M. PEYROT-CLAUSADE coll., 1968, R. Ser£ne det. :
1 6 9.2 x 7.6 mm (MNHN-B 22087) (see Peyrot-Clausade & Ser£ne. 1976 and Peyrot-Clausade, 1984).
Somalia. (M. Vannini coll.). Gesira, 18 km south of Mogadishu: stn 1, on live Pocillopora sp., low tide level,
1981: 1 9 8.3 x 6.6 mm. — Stn 2, on live Pocillopora sp., low tide level, 1981: I 9 4.4 x 3.6 mm. — Stn 3, on dead
Pocillopora sp., low tide level, 1981: 1 6 6.5 x 5.2 mm. — Stn 4, on dead Pocillopora sp.. low tide level, 1981: I 9
ovig. 1 1.7 x 8.7 mm. — Stn 5, on dead Pocillopora sp., low tide level, 1981: 2 9 3.6 x 3.3, 3.9 x 3.3 mm. — Sin 6, on
dead Pocillopora sp., low tide level, 1981: 1 9 5.4 x 4.7 mm. — Stn 1 1, on dead Pocillopora sp., low tide level. 1981:
2 6 6.0 x 5.3. 8.5 x 6.8 mm; 1 9 3.9 x 3.2 mm. — Stn 12, on dead Pocillopora sp.. low tide level: 3 6 6.7 x 5.4 -
1 1.6 x 9.2 mm. — Stn 13, on dead Pocillopora sp., low tide level: 1 6 7.6 x 6.0 mm; 2 9 3.9 x 3.3, 7.2 x 5.9 mm. —
Stn 15, on dead Pocillopora sp., low tide level, 1981: 1 6 9.7 x 7.9 mm; I 9 7.5 x 6.4 mm; 1 9 ovig. 10.0 x 7.8 mm.
— Stn 16, on dead Pocillopora sp.. low tide level, 1981: 1 6 8.8 x 7.0 mm. — Stn 17, on dead Pocillopora sp.. low tide
level, 1981: 1 9 8.0 x 6.6 mm. — Stn 18. on dead Pocillopora sp., low tide level. 1981: 1 9 3.9 x 3.4 mm (all Somalia
material from MZUF. see Vannini, 1985).
Aldabra. Takora. Seaward Cove, low tide level. J. D. Taylor coll.. 5.01.1967: I 9 I 1.7 x 9.0 mm (BMNH).
Cocos Keeling Islands. No locality, no depth, C. A. GlBSON-H ILL coll., 1941. M. Tweedie det.: 1 9 9.3 x
7.6 mm (ZRC 19646121 1). Sec Tweedie (1950). — Horsburgh Id, 0-37 m, 9.02.1989: 1 6 5.7 x 4.9 mm (WAM 375-
89). — Home Id. ocean side, no depth, 21.02.1989: 1 6 13.8 x 10.9 mm; 1 9 10.7 x 8.3 mm (WAM 751-89).
Northwest New Guinea. Salawatti Id, no depth, no date: 1 6 6.0 x 5.0 mm (ZMB 5139).
Elizabeth Reef (east of Australia). Tasman Sea, 29°58'S, 159o05.TE, no depth. 12.1987: 1 9 9.0 x 7.5 mm
(AMS-P 39155).
Middleton Reef (east of Australia). Tasman Sea, 29°29.5'S, 159°05.TE. intertidal, 9.05.1987: I 9 ovig
15.6 x 12.3 mm (QM W 1 3033).
Samoa. Pago Pago, no locality, no depth. 1924: 1 <3 14.0 x 1 1.0 mm (BPBM 2388).
French Polynesia. Tuamotu Ids: Marutea, Maitutaki. eastern reef, no locality, probably intertidal. M. Seurat
coll., 1905. G. Nobili det.: I 6 8.5 x 6.7 mm (MNHN-B 22089) (see Nobili , 1907). — Austral Ids. Marotiri. 27°35'S.
144°25'W, 9-5 m, D. M. Devaney coll.. 20.02.1971: 1 9 ovig. 13.5 x 10.6 mm (BPBM 71.201).
Howland Island. Whipp Exp., 0°48’N, 176°38'W. no locality, no depth, 1924: 2 6 9.1 x 7.6, 13.0 x 10.0 mnv
I 9 ovig. 10.6 x 8.4 mm (BPBM 2353).
Johnston Island. 16°45'N, I69°30’W, no locality, no depth, C. Edmondson coll., 1923: 2 6 8.5 x 6.8, 10.0 x
8.5 mm; 1 9 ovig. 11.5 x 9.6 mm (BPBM 1363).
Hawaii. No locality, no depth. 1836: 1 unknown sex. 7.9 x 6.0 mm (MNHN dry collection no. 23, tvpe of
Dynomena latreillii Eydoux & Souleyet. 1842). — No locality, no depth, no date: I 9 ovig. 1 0.1 x 8.0 mm (AMS-P
5478). — Honolulu, no depth, Th Mortensen coll., April 1915: 2 6 9.3 x 7.6 (damaged). 10.8 x 8.8 mm; 1 9 15.0 x
10.5 mm (ZMUC - unregistered).
Oahu. Waikiki: no depth, 1928: 1 9 ovig. 14.1 x 10.9 mm (BPBM 1587). — No depth, 1928: I 9 I 1.5 x 10.2 mm
(BPBM 3024). — No depth, no date: 1 6 10.2 x 8.0 mm (BPBM 2883). — No depth, C. Edmondson coll.. 1930: I 9
8.4 x 7.1 mm (BPBM 3131). — No depth, 24.04.1942: 1 9 ovig. 12.6 x 10.0 mm (USNM 182729). — No depth.
2.05.1942: I 9 1 1.7 x 9.4 mm (USNM 182729). — No depth, 29.05.1942: I 9 4.3 x 3.9 mm (USNM 182729).
Oahu, Waikiki Reef (C. Edmondson coll.): no depth, 1921: 3 6 1 1.4 x 9.0 - 15.1 x I 1.6 mm; 4 9 ovig 9 7 x 7 7 -
10-6 x 8.6 mm (BPBM 572). — No depth, 16.02.1922: 4 6 8.6 x 6.5 - 12.2 x 9.8 mm; 2 9 10.0 x 7.9, 14.0 x 10 8 mnv
1 9 ovig. 12.5 x 9.7 mm (BPBM 658).
Oahu, Waialee: no depth. C. Edmondson coll., 6.07.1921: 1 6 9.8 x 7.9 mm; I 9 ovig. 14.3 x 11.6 mm (BPBM
Oahu, Kahala Bay (C. Edmondson coll.): no depth. 7.03.1930: 16 6 5.8 x 4.7 - 12.4 x 10.0 mm; 11 9 5 2 x 42 -
10.8 x 8.3 mm; 4 9 ovig. 6.5 x 5.4 - 8.9 x 7.1 mm (BPBM 3168). — No depth, 1913: 2 6 5.7 x 4.7, 10.7 x 8.6 mm
(BPBM 3554). No depth. 05.1931: 7 6 9.7 x 7.8 - 12.3 x 9.7 mm (BPBM 3414). — No depth. 28.06.1934: 9 6
8.3 x 6.8 - 11.5 x 9.6 mm; 2 9 7.0 x 5.8, 10.8 x 8.6 mm; 7 9 ovig.) 9.3 x 7.5 - 10.5 x 8.6 mm (BPBM 3780)
Oahu, Pearl and Hermes Bay: no depth, 02.1928: 1 6 10.0 x 8.3 mm; I 9 10.8 x 8.5 mm (BPBM 3043)
Oahu. Barbers Point: no depth. 1.09.1936: 1 6 12.7 x 9.9 mm; 2 9 10.7 x 8.6, 12.5 x 10.3 mm (BPBM 4234)
Oahu. Rabbit Id: no depth. 13.1 1.1936: 2 6 10.1 x 8.2, 10.5 x 8.6 mm; 1 9 12.2 x 9.5 mm (BPBM 4255)
Oahu, Kawela Bay: no depth. C. Edmondson coll.. 15.07.1935: 4 6 10.7 x 8.8 - 13 0 x 10 5 mm- 5 9 ovig 9 9 x
7.7 - 13.0 x 10.3 mm, (BPBM 4038). — No depth. 10.07.1937: 5 6 9.9 x 8.1 - 15.0 x 11.5 mm; 6 9 9 0 x 7 ^nSx
11.1 mm (BPBM 4312).
Oahu, no locality no depth. 10.01.1924: 2 6 11.3 x 8.9. 12.6 x 10.2 mm (ZMUC). - No locality, no depth, no
date. - 6 5.6 x 4.6, 13.2 x 10.1 mm; 2 9 8.2 x 6.5, 11.2 x 9.1 mm; 3 9 ovig. 10.0 x 8.0 - 13.8 x 10.8 mm (BPBM
Source : MNHN, Paris
FAMILY DYNOMENIDAE
475
2186). — No locality, no depth, 1932: 5 8 8.0 x 6.5 - 12.7 x 10.0 mm; 2 9 ovig. 8.4 x 6.6, 10.5 x 8.5 mm,
C. Edmondson coll. (BPBM 3601). — No locality, no depth: 3 9 6.4 x 5.2 -10.2 x 8.3 mm (BPBM 3683). —
No locality, no depth, 1973: 4 8 7.7 x 6.0 - 13.9 x 10.8 mm; 6 9 5.2 x 4.3 - 12.7 x 10.0 mm; 2 9 ovig. 9.7 x 8.1,
11.2 x 9.1 mm (BPBM 510491). — No depth, 1973: 1 d 8.0 x 6.7 mm; 1 9 6.4 x 5.5 mm; 1 9 ovig. 11.6 x 9.3 mm
(BPBM 510492). — No locality. 12.2 m, 03.1996: 1 c? 6.8 x 5.2 mm (QM).
Oahu. Paile Point: no depth, 7.07.1952: I 8 11.1 x 8.5 mm; 1 9 ovig. 9.0 x 7.0 mm (BPBM 5804).
Oahu, Maili Point: no depth, 13.05.1953: 1 8 10.0 x 8.6 mm; I 9 8.5 x 6.9 mm; I 9 ovig. 1 1.0 x 9.6 mm (BPBM
6055). — No depth, 10.07.1953: 2 9 ovig. 9.3 x 7.6 . 10.0 x 7.8 mm (BPBM 5900).
Oahu, Kahe Point: no depth, S. Coles coll., 6-07.1977: 1 juv. 2.0 x 1.7 mm; 2 9 3.6 x 3.1. 5.0 x 4.2 mm (BPBM
1977.554).
Ocean Island (= Kure Id), 80 km NW of Midway Id: no locality, no depth, 1923: 1 9 4.0 x 3.3 mm (BPBM 1 133).
Taiwan. Nan-Wan, Pingtung County, no locality. 6 m, from coral Seriatopora hystrix Dana, M. S. Jkng coll.,
5.12.1985: 1 9 7.7 x 6.4 mm (NTOU). — Wan-li-Fong, low tide level, PlNG-Ho-Ho coll., 2.06.1992:
1 c5 6.6 x 5.2 mm; 1 9 ovig. 10.4 x 8.2 mm (NTOU).
Types. — Dynomene hispida Guerin-Meneville, 1832: according to Peyrot-Clausade and Serene (1976)
the male 14.0 x 1 1.5 mm, from Mauritius was probably considered the holotype by DESMAREST. The specimen is
part ol the dry collection held by the Museum national d'Histoirc naturelle, Paris, registration number MP-B 245.
Dynomena latreillii Eydoux & Souleyet, 1842: holotype is a small, mounted, dry specimen 7.9 x 6.0 mm,
collected from Hawaii, 1836, held by the Museum national d'Histoire naturelle, Paris, registration number
MP-B 235. As the specimen is mounted on a stub it is not possible to determine the sex.
Dynomene granulobata Dai, Yang & Lan, 1981: holotype is a male 6.2 x 5.4 mm, collected from Dongdao,
Xisha Ids, South China Sea, 04.1965, held by the Beijing Natural History Museum, registration number 65079.
Description. — Carapace wider than long (CW/CL = 1.3 approx.), broadly rounded in outline but frontal and
posterior margins truncated, surface minutely granulated and quite convex. Carapace surface and pereopods covered
with coarse, plumose setae of two lengths: short setae clothing surface, but interspersed with slightly longer (0.08
x CW) setae which also fringe limbs. Setae not arranged in clumps. Density of setae not sufficient to completely
obscure body surface. Structure of setae identical for both sizes: proximal 20% mostly smooth, followed by a
region occupying about 40% with very short sctules arranged in closely spaced bands, then a region occupying
about 25% where setules increase rapidly in size distally forming a dense bunch, and finally the distal 15% is
smooth, slightly curved or angled, and narrows to an acute tip.
A shallow frontal carapace groove separates a pair of low rounded protuberances, and then divides into separate
grooves which gradually become more faint. Just in front of cardiac region two laterally-directed grooves originate:
first groove (cervical) arises separately from small pits curving anteriorly on to branchial region, while second
groove extends across mid-line and initially runs almost directly towards lateral margin but then splits into an
anterior portion which follows the first groove for a short distance, while the second portion curves
posterolaterally, bordering anterior cardiac region. In effect groove crossing mid-line, connects two crescent-shaped
grooves. Mid- way along cardiac groove begins a faint branchial groove which runs towards base of last tooth on
lateral margin. Posterior cardiac area outlined by a faint groove. Anterolateral carapace margin begins at level of
postorbital corner, is evenly convex and bears four distinct, sharply pointed, equidistant teeth, first two of which
are directed anteriorly and last two directed more laterally. Near beginning of posterolateral border there is another
smaller tooth which lacks a sharp spine. Thus lateral margin has five teeth in total. Posterior carapace margin
recessed in order to accommodate first segment of abdomen which is visible dorsal ly.
Frontal margin continuous without orbital notch, V-shaped, ventrally-directed. joined to epistome (which
separates orbits). Supraorbital margin not projecting, continuous above orbits; towards postorbital corner are
about live small acute spines and another five spines continued on suborbital margin, which is essentially
straight. Orbits clearly exposed dorsally.
First article of antennule large, filling a large part of ventral region; distal margin obliquely angled and not
continuous with distal margin of second antennal article. Remainder of antennule folded into orbit. First article of
antenna moveable, wider than long, medially beaked; inferior tooth well developed, blunt; superior tooth above
opening of antennal gland is scarcely produced. Second article wider than long, distal margin widest, to which
exopod is fixed, curving over base of eycstalk and becoming broader but terminating as a sharp point.
Source :
476
C. L. McLAY
Third anlennal article longer than wide, and attached to remaining distal border of second article, slotting in behind
exopod, and together with small fourth article just matches length of exopod. Remaining antennal articles directed
laterally, extending about as far as postorbital corner, and can be partially folded under supra-orbital margin. Ratio
of length of antennal flagella to CW = 0.24. Eyestalk can be completely folded into orbit, and the cornea is well
developed, occupying all of tip. Epistome broadly triangular, surface concave; dorsal arm, joined to tip of carapace,
very elongate and narrow; lateral arms shorter and thicker. Joint between epistome and carapace marked by a suture
which can be clearly seen on inner surface of orbit.
Subhepatic area slightly convex. A groove begins near base of antenna, curving round under branchial region
without a cervical branch and meeting lateral carapace margin just anterior to tooth on posterolateral border and
connecting with branchial groove. Third maxillipeds operculiform; bases widely separated by tip of sternum.
Crista dentata has only five or six small, distal ly placed teeth on each side. Female sternal sutures 7/8 short,
ending wide apart on low tubercles just behind bases of second walking legs.
The branchial formula is 19 gills and 7 epipodites on each side:
Gills are unequal (anterior half larger) violin-shaped plates, marginally notched, and joined together along
central axis which carries afferent and efferent blood channels. Epibranchial corners thickened and bluntly pointed.
Towards base of each arthrobranch and pleurobranch there are pairs of elongate epibranchial lobes which increase in
length proximally. A transverse section near base shows two plates separated, on epibranchial surface, by two
lobes, while a section mid-way shows only two plates. Hypobranchial setae at posterior end of branchial chamber
poorly developed. Posterior margin of scaphognathite with two long setae. Hypobranchial margin of podobranchs
bears same setae as on epipod.
Cheliped only slightly longer than first leg. Merus trigonal; inner face smooth and fitting closely against
pterygostomial region of carapace; borders with small granules; outer face with a subtcrminal groove separating
a thickened ridge on which there are three larger granules. Outer face of carpus convex with several small granules,
two more prominent distal tubercles; inner superior border with a flattened, distomedially directed, granulated
spur which abuts against distal inner surface of propodus thereby restricting closure of cheliped against frontal
area. In a similar way, inferior carpal margin is produced as a smooth obtuse flange fitting against merus when
limb is withdrawn. These two structures give carpal article an unusual shape. Transverse section of propodus
decreases in area distally; outer and superior faces with 6-7 rows of small granules, inner and inferior faces smooth.
Fixed finger almost straight with two large teeth; moveable finger curved with a single, large tooth opposite first
tooth on fixed finger; both fingers, thick, hollowed out internally, touching only at tips which are without
interlocking teeth. Just below teeth on fixed finger is a distinct pit in which several long setae arc inserted with a
similar group of setae on inner margin. Larger groups of long stiff setae, inserted near base of dactyl and fixed
finger, are directed across space between the two fingers. On dorsal surface of dactyl there are several small distal
granules.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of meri of first two
legs and anterior face third leg merus smooth and nacreous, inferior distal margin hollowed out to accommodate
carpal article. Superior border of meri of these legs with a row of four or five small teeth and scattered granules,
and a shallow subterminal restriction; length of merus of second leg about 1 .5 times its width and equal to about a
third of CL. Carpi inflated, dorsal surface bearing three longitudinal rows of granules, and produced distally to
overhang base of propodi. Dorsal surface of propodi granulated. Dactyli curved, inferior margin armed with 5-6
small spines, tip brown and subacute.
Source :
FAMILY DYNOMENIDAE
All
wMmi
Fig. 18. — Dynomene hispida Guerin-Meneville, 1832: a-g, $ ovig. 12.9 x 9.9 mm, Port Louis, Mauritius.
M. Thirioux coll. (MNHN-B 22088): a. dorsal view of right half of carapace; b. ventral view of right orbital area;
c, outer face of left cheliped; d. dorsal view of left cheliped; e. posterior view of terminal articles of right fourth
pereopod; f, posterior view of terminal articles of right fifth pereopod; g. ventral view of telson and terminal
segments of female abdomen.
Source : MNHN , Paris
478
C. L. McLAY
Last pair of legs greatly reduced, lying along posterolateral border of carapace, reaching only as far as two
thirds along meral article of preceding limb; borders of articles unarmed. Last pair of legs subchelate, sexually
dimorphic: female with well developed distal extension of propodus which opposes dactyl, male with only weakly
developed propodal extension. Female propodal extension bearing four, unequal, stout, acute, spines each lined
with tiny flattened teeth along almost entire inner surface. Female dactyl as long as propodal extension, bearing
five stout, acute spines whose inner surface is unarmed. Male propodal extension bearing five unequal spines the
four largest of which are lined with tiny teeth along most of inner surface. Near its base the propodal extension
bears an area of rasp-like teeth which are present on only one side of limb (these structures are not present in
female). Note that there is no opposing area of similar teeth on base of dactyl. Male dactyl longer than propodal
extension and ending in a single acute claw.
All abdominal segments freely moveable, increasing in length and breadth distally; surface smooth; margins
unarmed but fringed with long setae. Telson much wider than long, anterior margin angled to accommodate
uropod, posterior margin broadly rounded. In female uropod plates are large, filling all of space between last
abdominal segment and telson. entirely excluding last abdominal segment and telson from reaching lateral margin
of abdomen. In male last abdominal segment reaches lateral margin but only occupies about a quarter of the
length. No effective abdominal locking mechanism: abdomen only loosely held against sternum in all sizes of
both sexes (see GuiNOT, 1979, pi. 21, figs 8-9). In males and immature females, there is a small rounded tubercle
at lateral margin of sternum, between first walking legs, adjacent to uropods. but this simply restricts sideways
movement of abdomen. In mature females this tubercle disappears and abdomen occupies all of ventral surface,
covering entire sternum and coxae of all pereopods with telson covering proximal half of third maxillipcds. In
male abdomen not quite so broad and telson only extends as far as bases of third maxillipeds.
Five pairs of pleopods in female, first pair vestigial, remainder biramous. First male pleopod a semi-rolled
tube ending in a curved apical plate surrounded by long setae. Second male pleopod with an exopod on basis,
needle-like distally, armed with a series of five tiny, acute, inset spines directed terminally and ending in two
hooked spines. Third to fifth male pleopods rudimentary and biramous; exopod longer and connected to basal
article by a joint.
Colour. — The coloured figure by Desmarest [1823, pi. (18), fig. 2] is almost certainly not based on a
living specimen but created by artistic license. Nagai and Nomura (1988: 92) have a picture which shows that
the body and legs are dark brown or black, fringed with light brown setae. Some patches of light blue in limb
joints, and bluish and pale pink patches on the carapace. Specimens from Somalia have light blue antennae. Baba
(1986) contrasts D. hispida with D. pilumnoides saying that its colour is dark blue or brown.
Geographic Distribution. — Type locality: Mauritius, east of Madagascar. Other localities in the Indian
Ocean are Somalia, Madagascar, Coetivy (Seychelle Ids). Aldabra. Salomon (Chagos Archipelago), and Cocos
Keeling Ids. Indonesian localities include Kupang, Timor and Ambon. In the Pacific Ocean D. hispida has been
recorded from Salawali Id (north west New Guinea). New Caledonia, Elizabeth Reef (Tasman Sea), Lord Howe Id,
Aranuka & Apamama (Gilbert Ids), Xisha Ids (Taiwan), Ryukyu Ids (Japan). Encwetak Atoll. Hawaii, Samoa,
Howland Id, Johnston Id, Ocean Id. French Polynesia (Marutea-Vaitutaki, Tuamotu, Moorea). This is a widespread
Indo-Pacific, shallow water species, inhabiting coral reefs and rocky bottoms, whose distribution is very similar to
that of Dynomene praedator.
Depth. — Most of the recorded specimens of D, liispida come from intertidal and shallow subtidal coral reefs
and rocks. The deepest reliable depth records are from Acropora humilis at 30 m. La Reunion (Ribes. 1978) and
from dead Acropora sp. at 24 m, Tulcar Reef. Madagascar (Peyrot-Clausade, 1981).
Size. — The maximum size for males is greater than for females: 19.0 x 14.0 mm (Sakai. 1976) and 15.6 x
12.3 mm (this study). The smallest ovigerous female is 6.5 x 5.4 mm recorded from Oahu. Hawaii (this study).
Records of ovigerous females (mostly from Hawaii) extend from January to July. Despite living in tropical waters
D. hispida seems to be a seasonal breeder. Newly laid eggs were found on females throughout the period January
to July, suggesting that the egg bearing season extends beyond July. The only females with eggs ready to hatch
were recorded in May and June. The smallest ovigerous females (CW = 8 mm) carried around' 70 eggs and the
Source : MNHN. Pahs
FAMILY DYNOMENIDAE
479
largest ovigerous females (CW = 14 mm) carried around 900 eggs. The average egg diameter was 0.44 mm.
indicating that there is a planktotrophic larvae (larval stages have not been described). This is to be expected from
the very wide distribution of D. hispida which occurs throughout the Indo-Pacific area.
DISCUSSION. — Different authors have attributed the name of this species to Latreille (e.g. WARD, 1942),
Desmarest (e.g. Peyrot-Clausade & Serene, 1976), or H. Milne Edwards (e.g. Guinot, 1979).
However, MANNING and HOLTHUIS (1981) point out that while the first latinized use of the generic name
Dynomene was by Desmarest (1823), the first latinized use of the specific name hispida was by Guerin-
Meneville ( 1832). Thus this species should be known as Dynomene hispida Guerin-Meneville, 1832.
The above description is based upon the same material (MNHN-B 22086) used by A. Milne Edwards ( 1 879)
who gave the first detailed description of D. hispida. The specimens examined arc of very similar size to the
original dimensions given by A. Milne Edwards (1879, pis 12-13) who illustrated some of the features of a
male specimen, but his figs 14 and 15 clearly show a fifth leg which must belong to a female and not a male as is
implied by the plate caption.
A. Milne Edwards (1879) was the first to treat Dynomena latreillii Eydoux & Souleyet, 1842 as a synonym
of D. hispida Guerin-Meneville, 1832, stating that the supposed differences were due to the fact that it was based
on a juvenile specimen. The type specimen of D. latreillii is not in fact a juvenile (7.9 x 6.0 mm, although it is
not possible to determine the sex because the ventral surface has been glued to a stub) and was collected from
Hawaii in 1836 by EYDOUX and Souleyet, two doctors on the corvette "La Bonite". In their original description
of D. latreillii they list the differences between this species and D. hispida. Among these differences are: les
bords latero-antcrieurs sont courbes, entiers, sans aucune trace de dents,... Les orbites, ovalaires, n'ont pas de dents
a 1'angle externe et sont ouverts a Tangle interne." I have examined this specimen and there are in fact four
anterolateral teeth and small spines around the supra- and suborbital margins. Thus it agrees well with D. hispida
where the anterolateral carapace margins have small well defined, acute teeth and there are small acute spines on
the orbital margin. The characters highlighted by EYDOUX & SOULEYET are in fact typical of Dynomene
praedator A. Milne Edwards, 1879 and both these species are known from Hawaii. Therefore it was essential to
check the type of D. latreillii. In treating it as a synonym of D. hispida , A. MlLNE EDWARDS (1879) pointed out
that the type specimen was small and stated that in D. hispida anterolateral teeth "..les bords anterieurs ne sont pas
aussi nettement decoupes qu'ils le deviennent par les progres de Tage." (i.e. become more distinct with age). I have
examined a large series of D. hispida specimens from Hawaii and even when small the anterolateral teeth are
distinct and easily seen, provided that the setae are cleared away. Therefore. I agree with Peyrot-Clausade and
Serene (1976) wdio supported the synonymy of A. MlLNE Edwards. It is important to clarify these differences
between D. hispida and D. praedator so as to avoid confusion and the erection of unnecessary new names.
After examining material from Hawaii and Mauritius (the type locality of D. hispida) Ward (1942) disputed
the synonymy ol D. latreillii and D. hispida. He argued that the Hawaiian species should remain distinct but it is
possible that his specimens from Hawaii were in fact D. praedator. I have examined many of the collections made
by Edmondson and deposited in the Bishop Museum, Hawaii, and found that they contain both D. hispida and
D. praedator , sometimes mixed together and sometimes mis-identified. Thus the records of Edmondson (1925,
1933, 1946) may well be for either D. hispida or D. praedator but it is clear that both species occur in Hawaiian
waters.
Dynomene granulohata Dai et al. (1981) w'as described for a small male specimen from the Xisha Ids, Taiwan.
The illustrations oi this animal closely resemble D. hispida. The distinctive features alluded to by the authors
relate to the cheliped and second pleopod. However the chclipcd features are allometric, changing with size, while
the distal characters of the second pleopod (sec Dai et al .. 1981. figs 13-14, Dai et al., 1986, fig. 12, 1-2. and
Dai & Yang. 1991, lig. 12, 1-2) are the same as found in D. hispida. The habitat of D. granulohata, "coral reefs
in shallow waters", is the same as that of D. hispida. For these reasons I think that D. granulohata is a synonym
of D. hispida.
A detailed discussion of the gills and epipods of Dynomene hispida is given above (see Morphology of the
Dynomcnidae). The branchial formula is 19 gills + 7 epipods. The gill structure of D. hispida is especially
interesting since they show both phyllobranchiate and "trichobranchiate" shapes. A section mid-way along an
arthrobranch or pleurobranch shows a pair of plates surrounding the afferent and efferent blood vessels as seen in
Source :
480
C.L. MCLAY
the phyllobranchiate Brachyura. The only differences are that the plates are bluntly pointed, as well as being
thickened, at their epibranchial corners and mid-way along each side is a notch. The thickening probably helps to
keep the lamellae separated. However, a section near the base of the gill shows the same two plates separated by a
pair of elongate epibranchial lobes thus giving the gill a "trichobranchiate-like" appearance. The podobranchs bear
only elongate lobes and their hypobranchial margins bear the same cleaning setae as found on the cpipods to
which the gills are attached. Thus within one species and one gill we have both kinds of gill structure present. As
in all dynomenids, D. hispida has an epipodal gill cleaning mechanism aided by two long setae extending from the
posterior end of the scaphognathite. The field of hypobranchial setae on the posterior body wall of the branchial
chamber seen in some other dynomenids, e.g. Paradynomene tuberculata , is scarcely developed in D. hispida .
Previous work by Peyrot-Clausade and SERENE (1976, text-fig. 1, pi. 5. A-B. F) showed the general
features of the male pleopods including light microscope photo-micrographs. Further details were provided by
Dai et al., (1986, fig. 1 1, 2-3) and Dai and Yang (1991, fig. 1 1, 2-3). This study presents the first scanning
electron microscope pictures of the male pleopods of D. hispida and these confirm the observations of Dai and
Yang (1991): there is a well developed curved apical plate on the tip of the first pleopod and there are five
subterminal and two terminal spines on the second pleopod. The subterminal spines are arranged in a series which
curves around to the inferior margin of the pleopod. Each spine is apically directed and lies in a triangular
depression. The terminal spines are larger than the others, unequal and curved sharply upwards at their tips. Near
the base of this article, beside the exopod, is the opening of a secretory gland which may provide additional
seminal fluid to aid sperm transfer. Minigawa (1993) has reported similar tegumental glands in the first and
second pleopods of Ranina ranina. The last three pairs of pleopods in I), hispida are rudimentary and biramous
with the exopod being longer. The endopod lacks a joint and appears to be an extension of the basal article. This
condition is found in most of the other dynomenids.
A. Milne Edwards (1879) illustrated examples of the long and short plumose setae found on the carapace.
His figures (pi. 12, figs 8-9) show the fine setules forming the dense distal band on the long setae as being similar
to more proximal setules, while those forming the dense band on the short setae are shown as being much stouter.
Furthermore, the tip of the short setae is shown as being angled while that of the long setae is shown as being
straight. Examination with a scanning electron microscope confirms most of the features illustrated by A. MILNE
Edwards but shows that stout distal setules are found on both setal types and the tips of the both setae can be
angled. There is a great deal of variation in setule development between setae.
In both females and males the tip of the last leg is fashioned into an obsolete subchelate mechanism but there
are significant differences between the sexes. The female has a dactyl which bears several unarmed spines while the
male has only a simple dactyl. The female propodal extension has four toothed spines while the male has five
spines on which the teeth are fewer and smaller, and there is an area of rasp-like teeth near the base of the propodal
extension. Perhaps these structures are a vestige of what was a grasping or cleaning limb and perhaps the female
has lost the rasp-like teeth but still has better developed remnants of the claws. It does not seem sensible to
hypothesize that formerly these limbs had different roles in males and females. What their exact role was is still a
matter for speculation.
Marco Vannini kindly supplied the D. hispida specimens from the coast of Somalia and data about their
habitat. Whole coral heads (n=l 19) of Pocillopora sp. were enclosed in plastic bags, removed and the fauna
extracted. A total of 36 dynomenid specimens were collected: 22 D. hispida and 14 D. praedator. Of the 19 corals
inhabited by dynomenids only 2 were alive, the others being encrusted and more or less dead. Despite being
relatively common, the dynomenids did not co-occur with the typical decapod inhabitants of live Pocillopora sp.
such as Trapezia spp.. Cynio spp., Alpheus lottini and Synalpheus charon. Perhaps these data indicate that more
recently evolved brachyurans have replaced the ancient dynomenids from the more productive live-coral habitat.
Furthermore, D. hispida occurred in 13 corals and D. praedator in 7 corals so that in only one case did the two
species occur together, suggesting that there may be some competitive exclusion between these two species.
Edmondson (1946) found that Hawaiian D. hispida are very abundant in crevices of porous rocks. Gut
contents of two specimens (CW = 13.5 and 15.0 mm) from Hawaii included mostly fine organic particles, sand
grains, a few cut lengths of tubular hydroidf?) and a chitinous fragment that may have come a decapod leg. The
groups of stiff setae on the cheliped fingers may act as a sieving device for collecting food particles.
Source :
FAMILY DYNOMENIDAE
481
Massive corals from Enewetak Atoll contained relatively few dynomenids: Goniastrea retiformis , only one
D. hispida among 36 decapods from 18 coral heads; Pontes lutea , no dynomenids amongst 77 decapods from
43 coral heads (HlGHSMITH, 1981). Of 2722 Brachyura and Anomura (except pagurids), collected from seven
stations on the barrier reef of Moorea, Peyrot-Clausade (1977) found only three specimens of D. hispida
(approx. 0.11%). The fauna was dominated by xanthid crabs (approx. 70%), and galatheids (approx. 17%).
Peyrot-Clausade (1981) recorded D. hispida from clumps of dead Acropora from 1.5-24 m, Tulear Reef,
Madagascar, and RlBES ( 1978) found only single specimens in Pocillopora damicornis , Favia stelligera , Acropora
clathra, A. humilis, A. variabilis, and Oulophyllia crispa from La Reunion. Above is also listed a Taiwan
specimen collected from Seriatopora hystrix . These data suggest that /_). hispida inhabits a variety of corals but is
comparatively rare in some reef communities.
Dynomene praedator A. Milne Edwards, 1879
Figs 3 b, 8 a-b, 11, 12 d, 14 b, 17 b, 19 a-g
Dynomene praedator A. Milne Edwards, 1879: 8. pi. 14. figs 20-26. — MlERS, 1884: 13. — Df. Man, 1888: 409. —
Ortmann, 1892: 543, pi. 26, fig. 3. — Alcock. 1901: 75 (list). — Rathbun. 1911: 196. — Ihle, 1913: 92 (list). —
Balss, 1938: 7. — Miyake, 1938: 194, text-fig. 4, 2; 1939: 198 (list); 1983: 195 (list). — Buitendijk, 1939: 221,
pi. 7, fig. 4. — Tweedie, 1947: 30; 1950: 106. — LlN, 1949: 12. — GuiNOT, 1967: 241 (list). — SERENE, 1968: 36
(list). — Takeda, 1973: 81; 1977: 35 (list). — Monod & SerLne, 1976: 25 (list). — Miyake, 1983: 195 (list). —
Guinot, 1985: 448 (list). — Garth, Haig & Knudsen, 1987: 241. — Rodgers & Olerod, 1988: 302. — Poupin,
1996a: 24 (list).
Dynomene sinense Chen. 1979: 9, fig. 1; 1980: 119, pi. 1, fig. 1. — Dai & Yang, 1991: 31 (key).
Dynomene sinensis - Odinetz, 1983: 208. — GUINOT, 1985: 448 (list).
Dynomene sp. - Naim, 1980: 55.
Dynomene tenuilobata Dai, Yang & Lan, 1981: 1 18, figs 5-9. — Dai, Yang, Song & Chen, 1986: 29. text-fis. 12, 3-4,
pi. 3, fig. 4. — Dai & Yang, 1991: 33, text-fig. 12. 3-4, pi. 3, fig. 4.
Dynomene huangluensis Dai, Cai & Yang, 1996: 234, fig. 1.
Dynomene hungluensis - Dai, Cai & Yang, 1996: 251 (err.).
? Dynomene sp. - Calm an, 1909: 703.
Not Dynomene praedator - Sakai, 1976: text-fig. 17. — Nagai & Tsuchida, 1995: 108, pi. 1. fig. 2 [= Metadynomene
tanensis (Yokoya, 1933)].
MATERIAL EXAMINED. — Somalia. Gesira, 18 km south of Mogadishu: stn 20, habitat unknown, low tide level.
G. Chelazzi coll., 1980: 1 9 ovig. 9.0 x 7.0 mm (MZUF).
M. Vannini coll.: stn 7, on dead Pocillopora sp., low tide level, 1981: 1 9 6.0 x 5.0 mm. — Stn 8, on dead
Pocillopora sp., low tide level. 1981: 1 6 9.5 x 7.2 mm (bopyrid isopod in right gill chamber, Gigantione sp. now). —
Stn 9, on dead Pocillopora sp.. 1981: 1 6 6.7 x 6.1 mm. — Stn 10. on dead Pocillopora sp.. low tide level, 1981: 1 9
8.7 x 6.7 mm. — Stn 14, on dead Pocillopora sp.. low tide level. 1981: 1 6 10.8 x 8.5 mm. — Stn 15, on dead
Pocillopora sp., low' tide level, 1981: 1 9 10.0 x 8.0 mm. — Stn 19, reef locality, low tide level, 1981 ; 3 6 5.9 x 5.1 -
9.6 x 7.8 mm; 4 9 5.1 x 4.2 - 8.0 x 6.4 mm. (All Somalia material from MZUF. see Vannini. 1985).
Glorieuses Islands. Intertidal zone, low tide level. A. Crosnier coll.. 29.01.1971: 1 6 8.7 x 6.7 mm; I 9 ovig.
9.6 x 7.5 mm.
Madagascar. Nosy Be: stn 899, no location, low intertidal. 02.1962: 1 6 12.0 x 9.6 mm (MNHN-B 6863). —
Stn 961. no location, low intertidal. 02.1962: 3 6 5.1 x 4.4 - 12.4 x 9.5 mm; 2 9 8.7 x 7.2, 1 1.0 x 8.0 mm; 1 9 ovig.
9.0 x 7.5 mm (MNHN-B 6864). — Intertidal zone. 03.1971: 2 9 9.1 x 7.3, 9.5 x 7.7 mm (MNHN-B 6903). —
Madirokely. intertidal zone, no date: 1 6 13.5 x 10.7 mm.
Seychelles (Aldabra). Reef West. A. J. Bruce coll.. no date: 1 <3 9.8 x 8.0 mm; I 9 7.8 x 6.5 mm; I 9 ovig.
8.2 x 6.6 mm.
Reunion. On Acropora sp., no locality, 10 m, no date: I S 6.8 x 6.0 mm. — On coral, no locality, 20 m. no date:
1 (5 5.0 x 4.2 mm. — On Pocillopora sp., no locality, 10 m, S. RlBES coll., no date: I 9 5.0 x 4.1 mm.
Cocos Keeling Islands, No locality, no depth. C.A. Gibson-Hill coll., 1941. M. W. F. TWEEDIE det.: 1 o’ 8.0 x
6.4 mm; 1 9 6.2 x 5.1 mm. (ZRC 19646129-10). See Tweedie (1950). — NW end of N. Keeling Id, 0-28 m. 23.02.1989:
1 6 5.9 x 4.8 mm; 2 9 6.7 x 5.4, 7.0 x 5.7 mm; 1 9 ovig. 6.9 x 5.5 mm (WAM 137-94).
Christmas Island. No locality, no depth, C.A. Gibson-Hill coll., 1940, M. W. F. Tweedie det.: 5 <3 6.1 x 5.0 -
7.9 x 6.4 mm; 1 9 6.7 x 5.3 mm; 2 9 ovig. 6.5 x 5.0. 7.0 x 5.8 mm (ZRC 19646121-8). — No locality, no depth,
C.A. Gibson-Hill coll., 1947, M. W. F, Tweedie det.: 2 6 7.3 x 6.1, 7.5 x 6.1 mm; 1 9 ovia. 10.0 x 7.9 mm (ZRC
1970.1.20.51-54). See Tweedie (1947).
Source
482
C. L. McLAY
Indonesia. Moluccas. Rumphius 2: stn 902 (26). Gorong, cast of Scram, on Porites . 1975: 1 6 5.4 x 4.3 mm. —
Amboina. No details (see DE Man, 1888). J. Brock coll.. 7.09.1885: 1 6 9.8 x 7.9 mm (SMF 163). — North Celebes.
Lembek Strait, no depth, no date: 1 9 9.8 x 7.9 mm (USNM 122953).
Mariana Islands (coll. H. T. CONLEY). Guam. Piti Reef. 13°27’N. 144°47'E, among rocks, 1.5 m, 22.07.1993:
2 6 8.5 x 6.5, 13.0 x 10.3 mm (UGM). — Ibidem, among rocks, 1 m, 1.08.1993: 1 9 7.8 x 7.0 mm (UGM).
Solomon Islands. Bougainville. Tiop, no depth, coll. H. Schoede. 4.11.1909: 10 S 4.1 x 3.6 - 9.3 x 7,6 mm.
14 9 4.3 x 3.6 - 7.5 x 6.2 mm (ZMB 14407).
New Caledonia. Exact locality unknown, probably intertidal, M. Balansa coll.. 1873: 1 9 (probably the 9
paratype) 10.2 x 8.1 mm (MNHN-B 7029). — Exact locality unknown, probably intertidal. M. Balansa coll., 1873:
6 6 6.5 x 5.4 - 9.6 x 7.5 mm; 2 9 6.5 x 5.5. 9.8 x 7.3 mm (MNHN-B 22075).
Lagon: stn 585, Tie des Pins. 22°46.0’S, 167°32.0’E, 43 m. B. Richer de Forges coll., 18.07.1985: I 6 10.4 x
8.2 mm; 1 9 5.7 x 4.2 mm.
Recif Mbere: 22°19.9’S. 166°13.2’E, 10 m. P. Bouchet coll., 5.05.1993: 2 6 6.2 x 5.1, 8.7 x 6.8 mm; 4 9 5.0 x
4.1 - 10.4 x 8.1 mm.
Operation Montrouzier. Koumac, recif Infernet, B. Richer de Forges coll., 7.10.1993: 1 6 8.7 x 7.1 mm.
French Polynesia. Society Ids. Tahiti. No details, 2 6 9.8 x 8.1, I 1.2 x 9.3 mm; 2 9 ovig. 10.2 x 8.3, 1 1.6 x
9.1 mm. (SMF 4855). Ortmann (1892) reported 2 6 from Tahiti, but under Dynomene hispida he also reported 2 6 and
2 9 from Oshima, Japan, so it is possible that he confused the material of these two species. — Moorea. On algae
{Amphiroa foliacea). O. Naim coll., 1978. : 1 9 ovig. 1 1.6 x 8.9 mm (MNHN-B 20203) (sec Naim. 1980). — Associated
with Pocillopora damicornis and P. elegans, O. Odinetz coll., 1981. D. GuiNOT det. in 1982 as Dynomene sinensis. : 1 <3
11.2x9.1 mm (MNHN-B 17090) (see Odinetz, 1983).
McDonald Volcano. 28°58'S, 140°16’W. approx. 50 m. B. Richer de Forges coll.. 19.05.1979: 3 9 4.3 x 3.9 -
4.4 x 4.0 mm.
Line Islands. Whipp Expedition: Washington Id. 4°43'N, 160o21’W. no depth, 1924: 1 6 5.0 x 4.3 mm (BPBM
2359). — Christmas Id. 1°5FN, 157°23’W. no depth. 1924: 1 6 11.1 x 8.2 mm (BPBM 2313).
Johnston Island. 16°45’N, 169°30'W, northwest side of outer reef, F. M. Bayer coll.. 28.08.1947: 3 9 4.8 x 4.0
- 5.7 x 4.5 mm (USNM 176603).
Hawaii. Oahu Id. Kawela Bay: no depth, C. Edmondson coll., 17.07.1935: 2 S 8.0 x 6.6, 9.0 x 7.0 mm (BPBM
4038). — No depth, 03.1936: 3 6 7.0 x 5.7 - 10.0 x 8.1 mm, 1 9 10.0 x 8.0 mm (BPBM 4212). — No depth,
10.07.1937: 2 6 8.6 x 7.2, 10.7 x 8.4 mm; 3 9 7.6 x 6.4 - 10.4 x 8.4 mm (BPBM 4312).
Oahu Id. Black Point: no depth, C. Edmondson coll., 12.07.1937: 1 6 9.2 x 7.3 mm (BPBM 3844). — No depth,
L. R. Woodward coll., 1937: 1 6 5.6 x 4.5 mm (USNM 175882).
Oahu Id, Waikiki Reef: no depth. C. Edmondson coll.. 1921: 1 6 10.6 x 8.8 mm, 1 9 ovig. 12.0 x 9.6 mm (BPBM
572). — No depth, C. Edmondson coll.. 1922: 1 6 12.0 x 10.2 mm (BPBM 739). — Off Waikiki, 5 m, 1.01.1945: 2 6
6.0 x 5.0, 6.6 x 5.7 mm; 2 9 5.2 x 4.0, 7.7 x 6.6 mm; 1 9 ovig. 9.7 x 7.3 mm (BPBM 5096).
Oahu Id. Kahala Bay: no depth, C. Edmondson coll., 7.03.1930: 10 6 6.2 x 5.5 - 10.3 x 8.4 mm; 8 9 6.4 x 5.2 -
9.8 x 7.4 mm; 3 9 ovig. 9.4 x 7.4 - 9.5 x 7.7 mm (BPBM 3168). — No depth, C. EDMONDSON coll., 05.1931: I 9
ovig. 10.3 x 8.1 mm (BPBM 3414). — No depth, C. Edmondson coll., 28.06.1934: 5 6 7.0 x 5.7 - 12.0 x 9.8 mm; 1 9
8.6 x 7.3 mm; 2 9 ovig. 8.2 x 6.8, 8.6 x 7.2 mm (BPBM 3780).
Oahu Id. Kahe Point: no depth, S. Coles coll., 06-07.1977: 2 6 4.6 x 4.0, 4.8 x 4.0 mm; 7 9 4.5 x 3.8 (parasitic
isopod attached to plcopods) - 7.0 x 5.6 mm (BPBM 1977.554).
Oahu Id, no locality, no depth, C. Edmondson coll., 1932: 1 6 9.2 x 7.4 mm; 1 9 ovig. 9.4 x 7.5 mm (BPBM
3601). — No locality, no depth, W.A. Bryan coll., no date: 1 6 11.0 x 9.3 mm (BPBM 184). — No depth, 1973: 1 6
9.0 x 7.2 mm (BPBM 510492).
Kauai. Anahola Bay, 15 m. 7.09.1959: 1 6 7.6 x 6.1 mm; 1 9 8.8 x 7.1 mm (BPBM 6819).
Hawaii, no locality, no depth. B. Degener coll., 21.09.1929: I 9 5.6 x 4.4 mm (USNM 108395).
Japan. Kuroshima Ids. In dead coral branches, inner reef, 1992: 1 9 10.3 x 8.0 mm; 1 9 ovie. 9.1 x 7.3 mm. — No
locality, coll. M. Osawa, 1993, 1 9 10.1 x 7.8 mm (collection of C. McLay).
Types. — Dynomene praedator A. Milne Edwards, 1879: holotype is a male 13.0 x 10.0 mm, collected by
M. Balansa from the intertidal, New Caledonia, 1873, but it probably no longer exists (see Discussion below
under this species). However there is a paratype female 10.2 x 8.1 mm, from the same collection, and held at the
Museum national d'Histoire naturelle, Paris, registration number MNHN-B 7029.
Dynomene sinense Chen, 1979: holotype is a female 9.2 x 7.5 mm. collected from Shenhang Dao, Xisha Ids,
South China Sea, 13.04.1958. held by the Beijing Natural History Museum, registration number IOAS-C00801.
A paratype male collected from Shi Dao, Xisha Ids, 6.04.1958, is held at the same institution, registration
number IOAS-C00802.
Dynomene tenuilobata Dai, Yang & Lan, 1981: holotype is a male 6.4 x 5.6 mm, collected from Jinyindao,
Xisha Ids, South China Sea, 5.12.1974. held by the Beijing Natural History Museum, under number 74073.
Source : MNHN. Paris
FAMILY DYNOMENIDAE
483
Dynomene huangluensis Dai, Cai & Yang, 1996: holotypc not designated from among the five specimens
(4 males and 1 female) collected from three different reefs of the Nansha Ids, South China Sea (6°56’N, 1 13°35’E -
10°50’N, 1 14°10’E). Specimens deposited in the Bejing Natural History Museum.
Description. — Carapace wider than long (CW/CL = 1.25 approx.), broadly rounded in outline but frontal
and posterior margin truncated, surface minutely granulated and evenly convex. Carapace surface and pereopods
covered with coarse, plumose setae of two lengths: short setae clothing surface, but interspersed with longer setae
(0.08 x CW) which also fringe limbs. Setae not arranged in clumps. The setae give this crab a yellowish colour
but their density is not sufficient to completely obscure body surface. Structure of setae identical in both sizes:
proximal 65% of shaft with many short setules, then a region occupying about 10% where there are
approximately a dozen long setules projecting almost at right angles to setal axis, and finally the distal 25%
which is smooth, slightly curved, and narrows to an acute tip.
A shallow frontal carapace groove separates a pair of low rounded protuberances, and then divides into separate
grooves which gradually become more faint. Just in front of cardiac region two laterally-directed grooves originate:
the lirst groove (cervical) arises separately from small pits curving anteriorly on to branchial region, while second
groove extends across mid-line and initially runs almost directly towards lateral margin but then splits into
an anterior portion which follows the first groove for a short distance, while the second portion curves
posterolaterally bordering the anterior cardiac region. In effect the groove crossing the mid-line, connects
two crescent-shaped grooves. No branchial groove is evident. Anterolateral carapace margin begins at level
ol postorbital corner, evenly convex and usually adorned with small granules, although there are two
(sometimes three) larger blunt granules interrupting margin at equidistant intervals and near beginning of postero¬
lateral border there is a small group of similarly prominent granules. Development of granules on anterolateral
margin is quite variable among individuals collected from the same site and often differs between the left and
right sides of the carapace of individual crabs. But there are never well defined, acute teeth as found in
D. hispida. Posterior carapace margin is recessed in order to accommodate first segment of abdomen which is
visible dorsally.
frontal margin continuous, V-shaped, ventrally-directed, joined to cpistome (which separates orbits).
Supraorbital margin not projecting, continuous above orbits without a notch, towards postorbital corner are a few
minute granules which are continued on suborbital margin, which is bluntly produced as a small lobe, making the
border sinuous. Orbits clearly exposed dorsally.
First article of antennule large, filling a large part of ventral region; distal margin almost adjacent to distal
margin ol second antennal article, the two ol them continuing the line of the suborbital margin. Remainder of
antennule folded into orbit. First article of antenna moveable, wider than long, medially beaked; inferior tooth well
developed, blunt; superior tooth above opening of antennal gland is scarcely produced. Second article wider than
long; distal margin longest, to which is fixed exopod curving over base of eyestalk and becoming broader. Third
antennal article longer than wide, and attached to remaining distal border of second article, slotting in beside
exopod, and together with small fourth article just matches length of exopod. Remaining antennal articles directed
laterally, extending about as far as postorbital corner, and can be partially folded under supra-orbital margin. Ratio
of length ol antennal flagella to CW = 0.23. Eyestalk can be completely folded into orbit, and cornea well
developed, occupying all of tip. Epistome broadly triangular, surface concave; dorsal arm, joined to tip of carapace,
very elongate and narrow; lateral arms shorter and thicker.
Subhepatic area slightly convex with a few minute granules. A groove begins near ventromedial corner of
carapace, curving round, without a cervical branch, under branchial region and meeting lateral carapace margin just
anterior to tubercle on posterolateral border. Third maxillipeds operculiform, bases widely separated by tip of
sternum. Crista dentata has only five or six small, distally placed teeth on each side. Female sternal sutures 7/8
short, ending wide apart on low tubercles just behind bases of second walking legs.
There arc 19 gills and 7 cpipodites on each side. Distribution of gills and epipods as for D. hispida. Gills are
unequal (anterior halt larger) violin-shaped plates, marginally notched, and joined together along central axis which
carries aflerent and efferent blood channels. Epibranchial corners thickened and bluntly pointed. Towards base of
each arthrobranch and pleurobranch there are pairs of elongate epibranchial lobes which increase in length
proximally. A transverse section near base shows two plates separated, on epibranchial surface, by two lobes,
484
C. L. McLAY
while a section mid-way shows only two plates. Setae on inner posterior wall of branchial chamber poorly
developed. Posterior margin of scaphognathite with two long setae. Hypobranchial margin of podobranchs bear
same setae as on cpipod.
Cheliped only slightly longer than first leg. merus trigonal, inner face smooth and fitting closely against
pteryogostomial region of carapace; borders with small granules; outer face has a subterminal groove separating a
thickened ridge without granules. Outer face of carpus convex with several small granules, two more prominent
distal tubercles; inner superior border with a flattened, distomedially directed, unornamented spur which abuts
against distal inner surface of propodus thereby restricting closure of cheliped against frontal area. In a similar
way, inferior carpal margin produced as a smooth obtuse flange fitting against merus when limb is withdrawn.
These two structures give carpal article an unusual shape. Transverse section of propodus decreases in area distal ly;
outer and superior faces covered in small granules; inner and inferior faces smooth. Fixed finger of female almost
straight with two small teeth; moveable finger curved with a single, larger tooth opposite first tooth on fixed
finger; both fingers hollowed out internally, touching only at tips which may be faintly divided into three or four
blunt, interlocking teeth. In mature male (CW = 12.0 mm) chelipeds more massive and teeth are similar to female
except that near point of articulation with dactyl there is a tooth-like projection directed distally. This projection
seems to occur in males with CW > 10.0 mm. Left and right chelipeds are sometimes different in number of
teeth. Just below teeth on fixed finger is a distinct pit in which several long setae arc inserted with a similar group
of setae on inner margin. Small groups of long stiff setae, inserted mid-way along dactyl and fixed finger, are
directed across space between the two fingers.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate; both faces of mcri of first two
legs and anterior face third leg merus smooth and nacreous; inferior distal margin hollowed out to accommodate
carpal article. Superior border of meri of these legs with several minute granules, a shallow subterminal
restriction; length of merus of second leg about three times its width and equal to about half CL. Carpi inflated
and produced distally to overhang base of propodi. Dactyli curved, inferior margin armed with 5-6 small spines, tip
brown and subacute. Borders of all leg articles minutely granulated.
Last pair of legs greatly reduced, lying along posterolateral border of carapace, reaching only as far as mid-way
along meral article of preceding limb; borders cf articles unarmed. Last pair of legs subchelate, sexually
dimorphic: female with well developed distal extension of propodus which opposes dactyl, male with only weakly
developed propodal extension. Female propodal extension bearing four, unequal, stout, acute, spines each lined
with tiny flattened teeth along almost entire inner surface. Female dactyl as long as propodal extension, bearing
fourteen unequal, stout, hooked spines (arranged asymmetrically around perimeter of the dactyl) whose inner
surface is wrinkled and mostly devoid of tiny teeth. Male propodal extension bearing five unequal hooked spines
the four largest of which are lined with many tiny flattened teeth along most of inner surface. Male dactyl longer
than propodal extension and ending in a single acute claw.
All segments of abdomen freely moveable, increasing in length and breadth distally, surface smooth, margins
unarmed but fringed with long setae. Telson much wider than long, anterior margin angled to accommodate
uropod, posterior margin broadly rounded. In female uropod plates large, filling almost all space between last
abdominal segment and telson. excluding most of last abdominal segment and telson from reaching lateral margin
of abdomen. In male last abdominal segment reaches lateral margin but only occupies about a quarter of the space.
No effective abdominal locking mechanism: abdomen only loosely held against sternum in all sizes of both sexes.
In males and immature females, there is a small rounded sternal tubercle between first walking legs, adjacent to
uropods, but this simply restricts sideways movement of abdomen. In mature females this tubercle disappears and
abdomen occupies all of ventral surface, covering entire sternum and coxae of all pereopods with telson covering
proximal half of third maxillipeds. In male, abdomen not quite so broad and telson only extends as far as bases of
third maxillipeds.
Five pairs of pleopods in female, first pair vestigial, remainder biramous. First male pleopod a semi-rolled
tube with a very small apical plate surrounded by long setae. Second male pleopod with an exopod on basis,
needle-like distally, armed with a scries of five tiny, curved, acute, inset spines and ending in three larger spines.
Terminal spines slightly curved at their tips and last two form a cheliped-like structure. Third to fifth male
pleopods uniramous and rudimentary.
Source : MNHN. Pahs
FAMILY DYNOMENIDAE
485
Colour. — Light yellowish-brown body and limbs, fingers white. Antennae same colour as the body, not
light blue as in Dynomene hispida.
Fig. 19. Dynomene praedator A. Milne Edwards, 1879: a-g, 6 13.5 x 10.7 mm. Nosy Be. Madagascar. A. Crosnier
coll.: a. dorsal view ot right half of carapace; b. ventral view of right orbital area; c, outer face of righi cheliped;
d. dorsal view’ of right cheliped; e. posterior view of terminal articles of right fourth pereopod; f. posterior view of
terminal articles ot right fifth pereopod; g. ventral view of telson and terminal segments of male abdomen.
Source : MNHN, Paris
486
C. L. McLAY
Size. — The maximum size for males is 13.5 x 10.7 mm, for females 12.0 x 9.6 mm, while the smallest
ovigerous female is 6.5 x 5.0 mm. Sakai (1976) gives the maximum size for males as 16.0 x 13.0 mm. but this
is probably based on the misidentification of Metadynomene tanensis (Yokoya. 1933). Ovigerous females have
been recorded from January to June with eggs ready to hatch in January and new eggs in June so the breeding
season is obviously longer than 6 months. Mean egg diameter = 0.46 mm. The smallest ovigerous female (6.5 x
5.0 mm) carried 50 eggs and the largest ovigerous female (12.3 x 9.6 mm) carried around 900 eggs. Dynomene
praedator has a very similar reproductive strategy to D. hispida. The larval stages of these crabs are unknown.
Geographic Distribution. — In the Indian Ocean Dynomene praedator is known from Somalia,
Madagascar, Reunion, Coetivy, Aldabra Id, Cocos Keeling Ids, and Christmas Id. There arc Indonesian records
from Pelokang and Postiljon Ids, North Celebes and Moluccas (Amboina, Obilatu Id). In the Pacific Ocean:
New Caledonia (type locality), Lord Howe Id. Solomon Ids (Bougainville Id), Samoa, Mariana Ids, Ryuku Id,
Japan, Xisha Ids, Taiwan, Gilbert Ids, Aranuka, Marshall Ids, Majcru, Viti Levu, Fiji Ids, Tuvalu, Ellice Ids, Niue
Id, Enewetak Atoll. Washington Id, Christmas Id, Johnston Id, Hawaii, Moorea. Tahiti. Rapa Id. The distribution
of D. praedator includes all of the Indo-West Pacific Ocean. This study reports new records of this species from the
coast of Africa (Somalia), Aldabra, Rapa Id, Mariana Ids, Johnston Id, Washington Id. Christmas Id (Pacific), and
Hawaii. Although Edmondson collected many specimens of this species (see Material Examined) he never
published their occurrence from Hawaii. This is at least partly attributable to the uncertainty about the validity of
the name Dynomene latreiUii Eydoux & Souleyel, 1842 (sec Discussion under D. hispida).
Depth. — Depth range is intertidal to approx. 50 m, with most specimens coming from the intertidal or
shallow subtidal depths.
Discussion. — A. Milne Edwards (1879) described this species using a male and a female specimen, but
he did not designate a type specimen. The measurements given first are those of the male and it is this specimen
which is figured, so the male should be regarded as the type. The eight specimens which remain are contained in
two tubes (MNHN-B 7029 and B 22075). The description given above is based on the female specimen. CW =
10.2 mm, CL = 8.1 mm (MNHN-B. 7029) which has the label " Dynomene praedator A. M. Edwards,
Nouvelle Caledonie. Coll. A. Milne Edwards, 1903". These dimensions are very close to those given by
A. Milne Edwards (1879) for the female specimen that he mentions so it is likely that this is the female
paratype. The other tube (MNHN-B 22075) labeled " Dynomene praedator A. M. Edwards N. Caledonie Balansa
1873. 1900" contains 7 specimens (5 males and 2 females). The largest male is CW = 9.6 mm and the largest
female is CW = 9.8 mm. This large male is smaller than the one mentioned by A. MlLNE Edwards (1879)
which measured 13x10 mm (figured on his plate 14). The figures in plate 14 show a dismembered thorax so it is
possible that the type male no longer exists. In fact, apart from the largest male and female, most of the original
material from New Caledonia consisted of immature specimens. One other specimen, a female of CW = 9.6 mm
(MNHN-B 22076), may also belong to the New Caledonian series collected by M. Balansa but the tube includes
a label stating "Balss det." and there are no other details available.
Dynomene sinensis Chen, 1979 was erected for a small male and a small female collected from the Xisha Ids.
Later, Dynomene tenuilobata Dai et al. , 1981 was erected for a small male from the same area. Then again
Dynomene huangluensis Dai, Cai & Yang, 1996 was created for one small female and four small males from the
Nansha Ids (10°50'N, 1 14°10'E), South China Sea. The characters used to separate these species from each other
and from other Dynomene species, especially D. praedator , were number of lobes on the carapace anterolateral
border, proportions of the cheliped manus and male pleopods (see CHEN, 1979, fig. 1, 5-6; Dai et al ., 1981,
figs 8-9; Dai et al. , 1986, fig. 12, 3-4; Dai & Yang, 1991, fig. 12, 3-4 and Dai, Cai & Yang, 1996, fig. I).
All the cheliped characters are sexually dimorphic and allometric. changing with size, and therefore subject to
variation within each species and therefore clearly unsuitable for species definition. The overall appearance of these
three species is very close to that of D. praedator and the male pleopods of D. tenuilobata and D. huangluensis are
identical to those described above. Furthermore, the development of lobes or granules on the anterolateral margin
of D. praedator is variable and, as noted above, it is the absence of distinct teeth on the anterolateral margin that
Source : MNHN, Paris
FAMILY DYNOMENIDAE
487
separates this species from D. hispida. All three species from the Xisha and Nansha Ids must be regarded as
synonyms of D. praedator.
When studying the dynomenids from the Palau Ids, Takeda (1973) recorded both Dynomene hispida and
D. praedator . He noted that in D. hispida the anterolateral teeth are more prominent, regular and equidistant...."
than in D. praedator , where "..the anterolateral border is provided with five obtuse teeth...". Furthermore, he
mentions that "In the smaller specimens from the Ryukyu Ids those teeth are spine-tipped". It is likely that these
small specimens may be D. hispida rather than D. praedator.
In identifying Dynomene praedator it is important to realize that the tomentum does not completely obscure
the body surface. Sakai (1976) synonymized Metadynomene tanensis (Yokoya, 1933) with D. praedator but
this was based on a misinterpretation of the nature of the tomentum. Further problems have arisen because of the
somewhat inaccurate description and figures given by A. MlLNE Edwards. This led Chen (1979) to describe
3 new species, Dynomene sinensis , from the Xisha Ids, China. CHEN recognized the similarity of her specimens
to D. praedator but believed that there were differences in the anterolateral margin, merus of the third walking leg,
and dentition of the chelipcd fingers. A close examination of a series of eight specimens from the type locality of
D. praedator shows that the arrangement of tubercles on the anterolateral carapace margin is variable on the two
sides of the carapace and also between specimens. It is not easy to ascertain the supposed differences in respect of
the merus of the third walking leg, but cheliped dentition is variable: in the female, the dactyl typically has two
teeth and the fixed finger only one, but these may be located proximally or distally; in males, the dactyl usually
has one tooth, but the fixed finger can have either one or two teeth (these can be different between left and right
limbs) and these may be located proximally or distally. The basal tooth on the fixed finger (well illustrated by
A. Milne Edwards, 1879, pi. 14, fig. 3) only occurs in males and does not develop until the size reaches about
CW = 10.0 mm. The redescription given above, based on the same material as used by A. Milne Edwards
(1879), shows that there are in fact no significant differences between the Chinese and the original specimens from
New Caledonia. The cheliped dentition characters are variable and in some cases attributable to ontogenetic
variation. Therefore D. sinensis must be regarded as a synonym of D. praedator.
The branchial formula is the same and the gill structure of Dynomene praedator is very similar to that of
D. hispida with most of each arthrobranch and pleurobranch consisting of violin-shaped plates on each side of the
gill axis but with epibranchial lobes separating the plates proximally. For other features of D. praedator not
illustrated here, see Ortmann (1892, pi. 26, fig. 3i) for the second maxilliped and Chen (1979, fig. 1, 4) and
Dai, Cai & Yang (1996, fig. 1,1) for the third maxilliped (as D. sinensis and D. huangluensis respectively).
As in D. hispida , the posterior margin of the scaphognathite bears two long setae which extend back over the
epibranchial surface ol the gills, and the hypobranchial margin of each podobranch is setose.
The original description of Dynomene praedator by A. MlLNE Edwards (1879) did not include a comparison
with the setae of D. hispida. The setae of D. praedator differ in several ways: there is no proximal smooth region
at the base ot the setae and the distal "dense band" consists of only a dozen or so long stout setules directed almost
at right angles. In other respects the two species are similar.
There are some differences in the male pleopods between Dynomene praedator and D. hispida : both species
have five inset subterminal spines on the second pleopod but in D. praedator these spines are curved instead of
straight; there are three larger curved, terminal spines in D. praedator , with the last tw'o forming a "pince-like"
structure, but in D. hispida there are only two hooked spines which do not form a "pince"; finally, the rudimentary
last three pairs of pleopods are uniramous in D. praedator but biramous in D. hispida.
Detailed illustrations of sexual dimorphism of the last leg of Dynomene praedator have not previously been
published. In most respects the structure of the obsolete subchelate mechanism in D. praedator is similar to that of
D. hispida except that the female has a much larger number of spines on the dactyl while in the male the teeth on
the propodal spines are better developed and the area of rasp-like teeth near the base of the propodal extension is
absent. Observations of living D. praedator from Hawaii (sec below) suggest that the last leg is carried
horizontally above the bases of the preceding legs and is only capable of a very restricted range of movements. The
limb cannot be placed in a subdorsal position over the posterolateral corner of the carapace and cannot reach under
the legs or under the abdomen. When the other legs move the last legs also move in an anterior-posterior direction
Source :
488
C. L. McLAY
to a very limited extent. There is no evidence that the last legs are capable of carrying pieces of camouflage or that
they could be used for grooming or cleaning.
Dynomene praedator has been collected from reefs on corals such as Pocillopora damicornis , P. elegans ,
Pocillopora sp., Acropora sp., Porites sp., and from the crustose alga Amphiroa folicicea. The specimens collected
by B. RICHER DE FORGES from the McDonald volcanic seamount came from near the top of this active volcano
where there are no corals and only fresh volcanic rocks and gravels. For discussion of the co-occurrence of
D. praedator and D. hispida in Pocillopora corals sec Discussion under the latter species.
A male specimen from Somalia contained a bopyrid parasite which is a new species of Gigantione
(Daniel ADKISON, pers. comm.). It is similar to the bopyrid specimens from Petalomera pulchra Miers, 1884
(from the Chesterfield Ids, see McLay, 1993: 166) which are being described by John Markham as a new species
of the same genus. Both these new species arc similar to G. mortenseni Adkison, 1984 which is known from
Cryptodromiopsis antillensis, Hypoconcha sabulosa , and H. spinosissima.
With the help of Ron Holcom, who video-taped Dynomene praedator in its natural habitat and in an aquarium
in Hawaii, I have been able to study the feeding behaviour of this dynomenid. During feeding the antennules are
especially active and are periodically cleaned by the third maxillipeds. There are two modes of feeding: firstly,
grazing algal covered rocks using the chelipeds to pick up food items, and secondly sifting through sand to remove
organic material. The latter feeding method is performed while the crab hangs upside-down from a rock or coral
with its third maxillipeds very close to the sand. One chcliped is used to shovel sand to the outer maxillipeds
which pick it up and sift out organic material using the setose palps and inner margins. Then the sand is pushed
aside using the other cheliped. In this way sand is moved across the mouth-field and the food extracted from it.
Examination of the stomach contents of four Dynomene praedator from Hawaii, ranging in size from 8.6 x 7.3 to
12.0 x 9.8 mm (two males and two females, BPBM 3780 and 4312), revealed unidentifiable soft organic material
and sand grains. The stomach contents of crabs, caught at a different time, confirm the observed feeding behaviour
and suggest that separation of organic material from sand (deposit feeding) is the main mode of feeding.
Dynomene filholi Bouvier, 1894
Figs 3 e, 5 c, 8 c, 11, 17 c. 20 a-g
Dynomene filholi Bouvier, 1894: 6; 1896: 57, figs 22-23. — A. Milne Edwards & Bouvier, 1900: 5, pi. 3 (col.),
fig. 3, pi. 8, figs 1-18. — ALCOCK, 1901: 75 (list). — Ortmann, 1899. pi. 1 19, fig. 11. — Ihle, 1913: 92 (list). —
Balss, 1921: 47. — Bouvier, 1922: 50. — Monod, 1956: 76. figs 84-88, 873. — Forest & Guinot, 1966: 48. —
Manning & Holthuis. 1981: 23. — Fransen, 1991: 93.
MATERIAL EXAMINED. — Cape Verde Islands. "Talisman": no stn number, ilot Branco. 60 m. July 1883: I 6
14.6 x 12.0 mm (used as the basis for the description and illustrated by A. Milne Edwards and Bouvier. 1900, see
Discussion below); 2 9 6.3 x 5.1, 7.2 x 5.9 mm (one of these specimens was listed as a male by A. Milne Edwards and
Bouvier, 1900) (MNHN-B 22080); 1 9 4.0 x 3.6 mm (ZMUC). — Stn 103, near la Praya. red coral banks, 275-150 m,
23.07.1883: 1 9 8.7 x 7.0 mm (used for describing the color) (MNHN-B 22087). — Stn 107, about 16°56'N, channel
between Saint Vincent and Saint Antoine, 75 m, 29.07.1883: 6 6 3.6 x 3.3 - 9.4 x 7.8 mm; 1 9 4.9 x 4.0 mm; 1 9
ovig. 5.8 x 4.9 mm (MNHN-B 22081); 2 6 5.4 x 4.8. 6.8 x 5.0 mm (MNHN-B 22083); 1 9 6.4 x 5.2 mm (ZMUC).
1FAN (Institut Francis d'Afrique Noire): N Maio Id, 42 m, J. Cadenat coll.. 1 1.06.1955: 1 9 3.8 x 3.6 mm. (MNHN-
B 22079).
Cancap: stn 6.069. 15°52'N, 13°00’W, 76-90 m, 13.06.1982: 1 6 7.5 x 5.8 mm. — Stn 7.125. 16°36,N, 24°36'W,
85-130 m. 1.09.1986: 1 9 10.0 x 8.7 mm (RMNH) (see Fransen. 1991).
Guinea Bissau. "Gazelle": I0°60’N, 17°I6'W, 274 m, 1.08.1874: 1 6 8.5 x 7.0 mm (ZMB 16722).
Gulf of Guinea, Annobon Island. "Calypso": stn 52, 1°27.5’S. 5°36.5’E, 35 m. 13-06-1956: 1 6 6.8 x
5.9 mm (MNHN-B 22085) (reported as a female by Forest & Guinot, 1966). — Stn 107, 1°26.15'S, 5°35.40’E. 60 m,
4.07.1956: 1 9 5.6 x 4.5 mm (MNHN-B 22082) (see Forest & Guinot, 1966: 48).
Annobon 3: Drague 2, 1°28.40’S, 5°35.50'E. 40 m, A. Crosnier coll.. 11.12.1965: 1 megalopa 3.2 x 3.2 mm
(MNHN-B 22090). — Drague 3, 1°25.30'S. 5°39.00’E, 52 m, A. Crosnier coll., 11.12.1965: 2 6 8.4 x 7.0. 8.6 x
6.8 mm; 2 9 8.0 x 6.6. 9.2 x 7.6 mm; 1 9 ovig. 10.5 x 8.2 mm. (MNHN-B 22092).
Annobon 5: chalutage au sud de file Annobon, 1°28.50'S, 5°37.50'E, 35-55 m, F. Poinsard coll., 16.06.1967: 3 6
6.0 x 5.0 - 10.2 x 7.7 mm; 1 9 8.0 x 6.5 mm; 1 9 ovig. 10.4 x 8.0 mm. (MNHN-B 22093)
Source : MNHN. Paris
FAMILY DYNOMENIDAE
489
Gulf of Guinea, Principe Island. "Calypso": stn 86, 1°35'N, 7°28'E, 45 m, 26-06-1956: 2 6 9.4 x 7.3,
10.9 x 8.6 mm; 3 9 6.9 x 5.4 - 9.2 x 7.0 mm (MNHN-B 7028). — Stn 95, 1°38.35’S, 7°21.35'E, 35 m, 1956: 1 6
12.3 x 9.9 mm (MNHN-B. 22084) (see Forest & Guinot, 1966: 48).
Types. — The specimen dissected by Bouvier in 1894 for describing the gills cannot be found and very
likely has disappeared. So I designate as neotype the male 14.6 x 12.0 mm collected near ilot Branco, Cape Verde
Islands, 60 m, in July 1883, registered at the Paris Museum under MNHN-B 22080, which was chosen by
A. Milne Edwards and Bouvier amongst the 28 specimens collected by the "Talisman" at the Cape Verde
Islands, as the basis of the first complete description of the species.
DESCRIPTION. — Carapace wider than long (CW/CL = 1.2 approx.), broadly rounded in outline but frontal and
posterior margins truncated, surface smooth and quite convex. Carapace surface and pereopods covered with coarse,
plumose setae of two lengths: short setae clothing surface, but interspersed with slightly longer setae (0.07 x CW)
which also fringe limbs and tend to be arranged in clumps, especially on carapace where there are about twenty
distinct tufts. Density of setae is not sufficient to completely obscure body surface. Structure of short and long
setae is different. In short setae the proximal 35% of shaft has very short setules, then a region occupying about
25% where long, stout setules are directed almost at right angles to shaft, forming a dense bunch, then the next
30% which bears a brush of long fine setules on only one side, and finally the distal 10% which is smooth,
slightly curved, and narrows to an acute tip. In long setae the proximal 25% is sparsely setose, 70% is covered
with small setules which distally increase in density, but not in size, and last 5% is smooth, slightly curved and
narrows to an acute tip.
A shallow frontal carapace groove separates a pair of low rounded protuberances, and then divides into separate
grooves which gradually become more faint. Just in front of cardiac region two laterally-directed grooves originate:
the first groove (cervical) arises separately from small pits curving (slightly sinuously) anteriorly on to branchial
region, while the second groove extends across mid-line and initially runs almost directly towards lateral margin
but then splits into an anterior portion which follows the first groove for a short distance, while the second
portion curves posterolaterally, bordering anterior cardiac region. In effect the groove crossing the mid-line,
connects two crescent-shaped grooves. Mid-way along cardiac groove begins a faint branchial groove which runs
towards base of last tooth on lateral margin. Posterior cardiac area is outlined by a faint groove. Anterolateral
carapace margin begins at level of postorbital corner, evenly convex and bears four distinct, broad-based,
equidistant teeth, each ending in a short spine; first two teeth directed anteriorly and last two directed more
laterally. Near beginning of posterolateral border there is another smaller tooth without a terminal spine. Posterior
carapace margin is recessed in order to accommodate first segment of abdomen which is visible dorsal ly.
Frontal margin continuous, V-shaped, ventrally-directed, joined to epistome (which separates the orbits).
Supraorbital margin not projecting, continuous above orbits with a small notch closer to postorbital corner,
without granules; suborbital margin essentially straight, but terminating as a subacute tooth. Orbits clearly
exposed dorsally.
First article of antcnnulc large, filling a large part of ventral region; distal margin obliquely angled and not
continuous with distal margin of second antennal article. Remainder of antennule folded into orbit. First article of
antenna moveable, wider than long, medially beaked; inferior tooth well developed, blunt; superior tooth above
opening of antennal gland is smaller. Second article wider than long; distal margin widest, to which is fixed the
exopod curving over base of eyestalk and becoming broader and terminating bluntly. Third antennal article is
longer than wide, and attached to remaining distal border of second article, slotting in behind exopod. and together
with small fourth article just matches length of the exopod. Remaining antennal articles are directed laterally,
extending well beyond postorbital corner, and can be partially folded under supra-orbital margin. Ratio of length of
antennal flagella to CW = 0.45. Eyestalk can be completely folded into orbit, and cornea is well developed,
occupying all of tip. Epistome broadly triangular, surface concave; dorsal arm. joined to tip of carapace, very
elongate and narrow; lateral arms shorter and thicker. Joint between epistome and carapace marked by a narrow
suture.
Subhepatic area slightly convex. A groove begins near base of the antenna, curving round under branchial
region, giving off a cervical groove which passes under base of second anterolateral tooth, and meeting lateral
490
C. L. McLAY
carapace margin jusi anterior to tooth on posterolateral border and connecting with branchial groove. Third
maxillipeds operculiform; bases widely separated by tip of sternum. Crista dentata has six or seven well developed,
distally placed teeth on each side. [Bouvier (1896, fig. 23) and A. Milne Edwards and Bouvier (1902, pi. 8,
figs 1-18) figure some of mouthparts.] Female sternal sutures 7/8 short, ending wide apart on low tubercles just
behind bases of second walking legs.
Branchial formula 19 gills + 7 epipods on each side. There is no podobranch on fifth pereopod. In cross section
gills have the lateral margin deeply notched, dividing gill into a hypobranchial plate (containing efferent vessel)
and an epibranchial lobe (anterior lobe longer). Between these marginal lobes are two pairs of lobes, first similar
and second much shorter than marginal lobes. Thus the epibranchial surface shows six rows of blunt lobes,
decreasing in size medially, which arc arranged above the afferent blood vessel.
Cheliped stout (especially in male) only slightly longer than first leg; merus trigonal, inner face smooth and
fitting closely against pterygostomial region of carapace, borders with a few small granules, outer face has a
subterminal broad, restriction which separates a thickened ridge on which there are three small blunt granules.
Outer face of carpus convex with three small granules, two more prominent tubercles on distal margin, inner
superior border with a flattened, distomedially directed, spur which abuts against proximal inner surface of
propodus thereby restricting closure of cheliped against frontal area. In a similar way. the inferior carpal margin is
produced as a smooth obtuse flange fitting against merus when limb is withdrawn. These two structures give the
carpal article an unusual and distinctive shape. Transverse section of propodus decreases in area distally; outer and
superior faces with very small granules which tend to be arranged in two or three longitudinal rows; inner and
inferior faces smooth. Fixed finger almost straight with six teeth increasing in size distally; moveable finger
curved with four teeth, first mid-way and separated from the remainder which are on tip; both fingers, thick,
hollowed out internally, touching only at tips where teeth interlock. Just below proximal teeth on fixed finger arc
two distinct pits in which several long setae are inserted with a group of similar setae on inner margin. Groups of
long stiff setae, inserted near base of dactyl and fixed finger, are directed across space between the two fingers.
Collectively, these setae form a sieving screen.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of meri of first
two legs and anterior face third leg merus smooth and nacreous, inferior distal margin hollowed out to
accommodate carpal article. Superior border of meri of these legs with two or three small granules, length of
merus of second leg about 1.7 times its width and equal to almost a half of CL. Dorsal surface of carpi bearing
three small granules on anterior margin and two on posterior margin, and produced distally to overhang base of
propodi. Dorsal surface ol propodi smooth. Daclyli curved, inferior margin armed with 5-6 small spines, tip
brown and subacute.
Last pair ol legs greatly reduced, lying along posterolateral border of carapace, reaching only as far as two-
thirds along meral article of preceding limb; borders of articles unarmed. Last pair of legs subchelate, sexually
dimorphic: female with well developed distal extension of propodus which opposes dactyl, male with only weakly
developed propodal extension. Female propodal extension bearing four, unequal, stout, acute, spines each lined
with tiny flattened teeth along almost the entire inner surface. Female dactyl as long as propodal extension,
bearing eight unequal, stout, hooked spines (arranged asymmetrically around perimeter of dactyl) whose inner
surface is concave, wrinkled and devoid of teeth. Male propodal extension bearing two unequal hooked spines
without teeth. Male dactyl longer than propodal extension, bearing a single spine on lateral margin and ending in
an acute claw.
All segments of abdomen freely moveable, increasing in length and breadth distally; surface smooth; mar¬
gins unarmed but fringed with long setae. Tclson much wider than long, anterior margin angled to accommodate
uropod, posterior margin broadly rounded. In female uropod plates are large, filling about two thirds of space
between last abdominal segment and telson, excluding most of last abdominal segment and telson from reaching
lateral margin of abdomen. In male last abdominal segment occupies about a half of length. No effective
abdominal locking mechanism: abdomen only loosely held against sternum in all sizes of both sexes. In mature
female it occupies all the ventral surface, covering coxae of all pereopods with telson covering proximal half of
third maxillipeds. In male the abdomen is not quite so broad and telson only extends as far as bases of third
maxillipeds.
Source :
FAMILY DYNOMENIDAE
491
Fig. 20. — Dynomene filholi Bouvier. 1894: a-g. 8 12.3 x 9.9 mm. Principe Island. Gulf of Guinea, " Calypso " (MNHN-
B 22084): a, dorsal view of left half of carapace; b. ventral view of left orbital area; c. outer face of left cheliped;
d. dorsal view of left cheliped; e, posterior view of terminal articles of left fourth pereopod; f. posterior view of
terminal articles ol left fifth pereopod; g. ventral view of telson and terminal segments of male abdomen.
oic
1 mm f
d
1 mm
| , ••
Source : MNHN. Paris
492
C. L. McLAY
Five pairs of pleopods in female, first pair vestigial, remainder biramous. Five pairs of pleopods in male, last
three pairs rudimentary. First pleopod a semi-rolled tube ending in a well developed, curved apical plate surrounded
by long setae. Second male pleopod with an exopod on the basis, needle-like distally, armed with a series of
ten tiny, acute, closely spaced spines decreasing in size distally, and ending in one larger terminal spine. Last
three pairs of pleopods biramous, exopod longer and connected to basal article by a joint (see MONOD, 1956.
figs 84-88).
Colour. — A. Milne Edwards and BOUVIER (1900) describe the colour of live Dynomene filholi as
yellowish-pink with red. notably along the frontal border and on certain parts of the anterior legs. In alcohol these
colours are completely lost.
Geographic Distribution. — Manning and Holthuis (1981) make the interesting observation that
Dynomene filholi is an insular West African species, known so far from the Cape Verde Ids (type locality) and the
Gulf of Guinea Ids, Principe, and Annobon. It has not been recorded from the mainland.
Depth. — Depth range of the material examined is 35-275 m. The depth of 1477 m given by Bouvier (1922)
is probably incorrect and Manning and HOLTHUIS (1981 ) note that most of the records for Dynomene filholi are
from between 23 and 75 m. They doubt the "Talisman" record of 275-150 m but the " Gazelle " specimen (ZMB
1672) apparently came from 274 m. Thus the depth range for this species is from 35-275 m. Most of the material
has come from bottoms of coralline algae and red coral with sand and rock.
Size. — The size range of Dynomene filholi material examined was from 3.6 x 3.3 mm to 14.6 x 12.0 mm
lor males, 3.8 x 3.6 mm to 10.5 x 8.2 mm for females and 5.8 x 4.9 mm to 10.5 x 8.2 mm for ovigerous
females. However Manning and Holthuis (1981) report an overall size range for both sexes of 3-16 mm and
7-12 mm for ovigerous females.
Ovigerous females of Dynomene filholi have been reported in May (" Pillsbury " material), June, July and
December (herein). A female 5.8 x 4.9 mm carried 30 eggs while a female 10.4 x 8.0 mm carried around
300 eggs. Egg size was 0.5 mm diameter. Ovigerous females collected in December carried eggs which were
newly laid. The small size of ovigerous females suggests early maturity and small egg size indicates indirect
development. Having a planktonic larval stage and an insular distribution suggests that larvae must be retained
in the system of ocean currents surrounding these Atlantic islands. It is interesting to note that D. filholi has
its breeding season during the second hall ol the calendar year whereas the very similar Indo-Pacific species,
D. hispida and D. praedator, both carry eggs during the First half of the year.
DISCUSSION. — The name Dynomene filholi was first used by BOUVIER (1894) in the context of describing
the gills to a Seance (10 November 1894) of the Societe Philomalhique de Paris, and then in his classic
"Sur Vorigine Homarienne des crabes..." paper (1896) where the carapace (his fig. 22), mouth appendages, a gill
and male pleopod of the fifth abdominal segment (his fig. 23) are figured. But it was not until A. MlLNE
Edwards and Bouvier (1900) published their monograph on the "Travail l eur" and "Talisman" material that
a substantial formal description and a list of material examined was published. However, even although Bouvier
(1894) only described the branchial apparatus, his use of the name Dynomene filholi must be regarded as the first
valid example.
Clearly Bouvier dissected the specimen (or the specimens) that he used for his 1894 paper. Since none
remain, no types are now available and it was necessary to designate a neotype.
In their paper published in 1900, A. Milne Edwards and Bouvier refer to "Je grand male qui nous a servi
de type,..." and again on page 9 they state "Ce dernier, qui est represente dans la planche VIII. a servi de type pour
notre description;..". These authors mention the measurements of this male (14.6 x 12.0 mm) and the place where
it was collected (Cape Verde Islands, Hot Branco. 60 m). This male specimen is registered under MNHN-B 22080,
along with two females. It is this male that I designate as neotype.
According to the list published by A. Milne Edwards and Bouvier, 28 specimens of D. filholi were
collected by the "Talisman" from 4 stations. Only 14 remain at the Paris Museum. One from the station 107 (not
Source .
FAMILY DYNOMENIDAE
493
examined by me) is held by the Museum of Comparative Zoology, Harvard, registration number MCZ 6559 and
2 others are held by the Zoologisk Museum at Copenhagen. Of the original 28 specimens, 1 1 are missing. But it
may be these were dissected by Bouvier for his 1894 and 1896 papers and discarded.
A. Milne Edwards and Bouvier (1900, pi. 8, figs 16-17) figure examples of the short setae, covering the
surface, and the long setae arranged in "bouquets" on the carapace as well as fringing the abdomen. The electron
microscope pictures confirm their results and add some more precise detail to the description and figures. D. filholi
differs from the two preceding species in having dissimilar short and long setae. The short setae are unique
amongst the dynomenids in having a subterminal brush of fine sctulcs. Presumably these are specialized sensory
setae.
Bouvier (1896, fig. 23) and A. Milne Edwards and Bouvier x( 1900, pi. 7, figs 1-18) describe and illustrate
the second maxilla, and all three maxillipeds of D. filholi. They also show an important feature of the second
maxilla: the presence of three long denticulate setae on the posterior margin of the scaphognathite which probably
have a role in cleaning the epibranchial surface of the gills. The presence of these setae is a primitive character
because they arc absent in the more derived Brachyura where gill cleaning is carried out by the long setose epipods
of the maxillipeds. In D. filholi these epipods are well developed and their cleaning role is supplemented by the
epipods on the pereopods. Three long scaphognathite setae are also found in D. pilumnoides but in D. hispida and
D. praedator, there are only two setae.
A. Milne Edwards and Bouvier (1900) state that the branchial formula of D. filholi is the same as in
Dicranodromia mayheuxii A. Milne Edwards, 1883 (the correct spelling is D. mahieiixii , see GuiNOT, 1995. 236)
with the addition of an epipod and podobranch on the fourth pereopod. However this is incorrect: in Dynomene
filholi there are no gills on the fifth pereopod and there is only one arthrobranch on the third maxilliped. The
branchial formula is 19 gills + 7 epipods. The gill structure of D. filholi is very different from the preceding
species. A. Milne Edwards & Bouvier (1900, pi. 8, fig 18) provide an accurate cross section of the middle
region of an anterior arthrobranch. The gill can be divided into two halves: a phyllobranchiate-like hypobranchial
half, containing the efferent vessel, and a trichobranchiate-like epibranchial half, with six lobes arranged above the
afferent vessel. As noted by Bouvier (1896) the shorter epibranchial lobes tend to disappear towards the tip of the
gill. A. Milne Edwards & Bouvier (1900) state that, in cross section, there are eight lobes but they counted
the corners of the hypobranchial plate as "filaments".
A. Milne Edwards & Bouvier (1900) were the first to recognize, in a dynomcnid species, that the sub-
cheliform last pair of legs are sexually dimorphic: the extension of the propodus is much better developed in the
female than in the male. In Dynomene filholi the last leg of the male is scarcely subcheliform. All other
dynomenids also show this dimorphism. A small number (four) of spines, bearing many flattened teeth, on the
female propodus is in keeping with the other species of Dynomene. However, the male shows some differences:
there are only two spines on the propodus and these do not bear any teeth (in D. hispida and D. praedator there arc
live spines with many flattened teeth), while there is a lateral spine on the side of the dactyl which is unarmed in
the other species o( this genus. A similar dactyl spine is found in male Metadynomene tanensis and male
Paradynomene tuberculata have a spine on the dorsal margin of the dactyl. These dactylar spines arc reminiscent of
those found in certain genera of the Dromiidae such as Dromidiopsis Borradaile, 1900, Tunedromia McLay. 1993.
and Lauridromia McLay, 1993 where they are used, along with other spines, to assist in securing the sponge
carried by the last two pairs of legs over the crab. However, in these dynomenids the spines are closely flattened
against the surface of the dactyl so that they could not function in the same way as in the dromiids. These spines
indicate a common ancestral relationship.
The figures of the first two male pleopods of Dynomene filholi by A. MILNE Edwards and Bouvier (1900,
pi. 7, 13-14) lack detail, but these pleopods are figured again by MONOD (1956, figs 84-88). The tip of the first
male pleopod bears a curved apical plate surrounded by long setae as is found in most other dynomenids. However,
MONOD's figures show that the second pleopod is armed with twice the number of subterminal spines seen in
D. hispida and D. praedator , but there is only one long terminal spine. MONOD (1956, fig. 88) is inaccurate in
the number of subterminal spines because it shows more than are there. Bouvier (1896, fig. 23, V) includes a
figure of one of the fifth pair of rudimentary male pleopods showing that it is biramous. There are two unequal
lobes shown but one lobe (endopod) is an extension of the penultimate article while the other lobe (exopod) is
Source :
494
C. L. McLAY
separated by a joint. If the shorter inner lobe is regarded as the endopod then it must be assumed that fusion has
occurred and the joint has been lost. The same situation is found in D. hispida and D. pilumnoides .
Examination of the stomach of a Dynomene filholi male 10.9 x 8.6 mm (MNHN-B 7028) from Principe Id.
Guinea Gulf, revealed unidentifiable soft particulate organic fragments and some soft calcareous granules. The
groups of stiff setae on the cheliped fingers may act as a sieving device for collecting food particles.
Dynomene filholi is most similar to D. pilumnoides (see Discussion below under the latter species). Since
D. filholi is the only species of this genus inhabiting the Atlantic it is interesting to speculate about its origins.
It seems to be a reasonable assumption that species of the genus Dynomene originated in the Tethys Sea so that
the ancestors of D. filholi were Tethyan crabs. A southern colonization route for these crabs could have been
available as early as the Upper Cretaceous (90-80 mybp) or sometime thereafter. D. pilumnoides has been recorded
from the coast of Natal although no further south than this. At least at present there does not seem to be
a dispersal route via the Cape because it is blocked by the local oceanic circulation pattern. This self-contained
circulation pattern seems to have been in existence for a considerable time because there is a suite of endemic
South African dromiid genera and species (see McLay, 1993) which have been isolated perhaps since the Upper
Cretaceous or Palaeocene (65 mybp). This interpretation requires that the Atlantic colonization by Dynomene
must have been during the late Mesozoic or very Early Tertiary.
Dynomene pilumnoides Alcock. 1900
Figs 3 c-d, 8 d-e. 11.12 e-f, 14 c, 17 d. 21 a-g
Dynomene pilumnoides Alcock. 1900: 133; 1901: 35, pi. 1, fig. 2. — Stebbing, 1905: 58 (list). — Barnard. 1947:
371: 1950: 337, fig. 65 a-c. — Sakai, 1965: 12, pi. 6, fig. 2; 1976: 29, pi. 6, fig. 3. — Guinot. 1967: 242 (list). —
Serene, 1968: 37 (list). — Peyrot-Clausade & Serene. 1976: 1344 (key). — Takeda, 1977: 35 (list). — Serene &
Vadon, 1981: 121. — Kensley, 1981: 37 (list). — Miyake, 1983: 11. pi. 4, fig. 2, 195 (list). — Baba, Hayashi. &
Toriyama, 1986: 310, fig. 163. — Garth, Haig & Knudsen, 1987: 241. — Nagai. 1989: 43.
Maxillothrix actaeiformis Stebbing, 1921: 457, pi. 14 (Crust, pi. 109).
Dynomene hispida - Yokoya, 1933: 95, text-fig. 37. Non Guerin-Meneville, 1832.
Dynomene actaeiformis - SerLne, 1968: 37 (list). — Takeda, 1977: 35 (list).
MATERIAL EXAMINED. — Madagascar. "Vauban" (A. Crosnier coll.). N.W. coast : 12°41.50’S. 48°17.00'E.
dredge. 160-170 m, 1.08.1973: I 9 14.6 x 12.0 mm; 1 9 ovig. 13.7 x 12.0 mm (MNHN-B 6913). — IV coast : Dredge,
about 18°50'S, no depth. 24.02.1973: I 6 10.0 x 8.2 mm (MNHN-B 6914). — S.E. coast : sin 72, 25°09.0’S. 17°14.2'E,
80-85 m, 3.03.1973: 1 9 8.7 x 7.4 mm; 1 9 ovig. 10.0 x 8.3 mm. (MNHN-B 6904). — Fort-Dauphin. 90 m. no date*
I <5 8.1 x 6.8 mm; 1 9 6.9 x 6.2 mm (MNHN-B 6858).
Reunion. No locality, no depth, S. Ribes coll., no dale: 1 6 1 1.8 x 10.1 mm (MNHN).
Australia. New South Wales. Port Stephens. 32°42’S, 152°6’E, no depth, no date: 1 6 17.2 x 14.1 mm (small
balanomorph barnacle on dorsal surface of carapace); 1 9 ovig. 14.8 x 12.0 mm (AMS-P 42233). — Crowdy Head
3 1 °54'S, 153°00'E. 100 m, K. J. Graham coll.. 17.08.1977: 3 6 8.0 x 6.5 - 22.8 x 17.4 mm; 1 9 14.4 x 1 1 4 mm
(AMS-P 26583).
Indonesia. Danish Expedition Kei Islands: stn 3, 5°32'S, 132°36'E, 245 m, Th. Mortensen coll . U 03 1922
1 6 10.0 x 8.9 mm (ZMUC).
Philippines. " Pele " (B. R. WILSON coll.). Sulu Archipelago. Pearl Bank: 3.2 km and 349° from Zal Id. 18 m,
22.02.1964: 1 9 ovig. 9.5 x 7.5 mm (MNHN-B 10376). — 6.4 km and 212° from Zal Id. 90 m, 22.02.1964: 1 9 7.8 x
6.6 mm (MNHN-B 10375). — 14.4 km and 242° from Zal Id, 99-108 m, 22.02.1964: 1 9 14.6 x 1 1.7 mm; I 9 ovig.
12.7 x 10.6 mm (MNHN-B 10374). — 4 km and 182° from Zal Id, 90 m, 22.02.1964: 1 6 9.2 x 7 8 mm (MNHN-
B 10483).
Musorstom 1: stn 57, 13°53.10’N, 120°13.20IE, 107-96 m, 26.03.1976: 1 6 7.2 x 5.8 mm (see SerLne & Vadon
1981 who reported the specimen as a female).
Japan. Honshu Mie-Ken Wagu: 34°04.00'N, 136°5I.30'E. no depth, no date: 2 <3 24.4 x 19 6 28 6 x ->2 1 mm
(SMF 17127).
East ol Toshima: 34°59.5’N, 139°36.3'E, 93-95 m, 1991: I 6 6.4 x 5.6 mm. — 34°19.79'N 1 39°0 1 37'F 134 m
coll. M. Osawa, 1993: 1 9 7.1 x 6.0 mm. ’
Hawaii Islands. "Albatross": stn 3823, south coast of Molokai. Lae-0 Ka Laau Light 21°0'),10"N 157015'45MW
142-406 m. 1.04.1902: 1 d 6.9 x 5.7 mm (USNM).
New Caledonia. Lagon: stn 393 bis, 22°46.00'S, 167°4.00'E, 284 m. 22.01.1985: 1 6 8.0 x 6.6 mm.
Source : MNHN, Paris
FAMILY DYNOMENIDAE
495
Musorstom 4: stn 164, 18°33.2’S, I63°13.0’E, 255 m, 16.09.1985: I 9 19.2 x 15.5 mm. — Stn 227. 22°46 00’S
167°20.00'E, 300 m. 30.09.1985: 1 9 5.0 x 4.5 mm.
CHALCAL 2: stn DW 69, 24°43.70'S, 168°07.90'E, 260 m. 27.10.1986: 1 9 4.8 x 4.0 mm. — Stn DW 70,
24°46.00'S, 168°09.00'E, 232 m. 27.10.1986: 1 juv. 5.4 x 4.5 mm; I 6 5.6 x 4.6 mm. — Stn DW 84, 23°23.80'S
1 68°07.00'E, 170 m, 31.10.1986: 2 6 6.3 x 5.2, 7.2 x 6.0 mm.
Smib 3: stn 18. 23°41.50’S, 167°59.40’E, 338 m, 23.05.1987: 1 6 23.5 x 19.0 mm.
Smib 4: stn DW 40, 24°46.20’S, 168°8.70’E, 260 m. 7.03.1989: 1 juv. 4.2 x 3.8 mm.
Volsmar: stn DW 7, 22°26.00'S, 17r44.10'E, 400 m, 1.06.1989: 2 9 4.6 x 4.2, 12.4 x 10.3 mm.
Smib 5: stn DW 94, 22°I9.60’S, 168°42.80'E, 275 m, 12.09.1989: 1 9 ovig. 20.7 x 17.6 mm.
Smib 8: stn DW 163, 24°49.10’S, 168°08.90’E, 310-460 m, 28.01.1993: I 6 13.2 x 10.6 mm; I 9 7.8 x 6 5 mm —
Stn DW 175. 23°14.10'S, 168°00.40’E, 235-240 m. 29.01.1993: 1 9 10.3 x 9.0 mm.
Lagon, rdcif Larcgnere, 12-16 m, 3.05.1993: 1 9 7.5 x 6.2 mm.
Bathus 3: stn CH 801, 23°39.00’S, 168°00'E, 270-300 m, 27.11.1993: 1 9 9.8 x 8.9 mm. — Stn DW 836, 23°02,S.
1 66°59'E, 295-306 m, 30.1 1.1993: 1 9 6.7 x 5.9 mm.
Bathus 4: stn DW 943, 20°I2.28’S, 164°30.58'E, 347-316 m, 09.08.1994: 1 9 8.2 x 6.8 mm.
Smib 10: sin DW 209, 24°49’S, 168°09’E, 329-560 m. 10.01.1995: 1 6 27.0 x 22.0 mm.
Loyalty Islands. MUSORSTOM 6: stn DW 423. 20°20.85’S. 166°40.50,E, 280 m. 16.02.1989: 2 9 10.1 x 8.6,
10.3 x 8.7 mm. — Stn DW 451. 20°59.60'S, 167°24.50’E. 330 m. 20.02.1989: 1 6 9.7 x 8.2 mm. — Sin DW 472
21°8.60'S, 167°54.70’E, 300 m, 22.02.1989: 1 6 8.1 x 7.0 mm.
Types. — Dynomene pilurnnoides Alcock, 1900: holotype is a male 11.0 x 10.0 mm, collected by the
" Investigator ", from 1 1°27’N, 73°1.00'E, off Kiltan Id, Laccadive Ids, 90-54 m. held by the Indian Museum,
Calcutta, registration number 9000/6.
Maxillothrix actaeiformis Stebbing, 1921: holotype not designated from amongst the four specimens, based
on the number of carapaces in the container (at least two males and one female according to Stebbing,
1921), collected by the SS "Pieter Fcutre" from off the coast of Natal, located NW by N 12.6 km from the
Umhlangakulu River, north of Durban (approximate coordinates 29°44'S, 31°5’E), stn PF- 12348, 90 m,
4.04.1901, held by the South African Museum, registration number SAM-A839. This type material is in poor
condition (Liz HOENSON pers. comm.). Two syntypes are held by the British Museum, registration number
1928. 12. 1. 10. According to the index card there is one specimen and fragments of another but the material in
the container is just a lot of fragments (Miranda Lowe, pers. comm.). Thus all of the original material is in very
poor condition.
Description. — Carapace wider than long, ratio of CW/CL 1.20-1.25, broadly rounded in outline but frontal
and posterior margins truncated, surlace smooth, quite convex, with a few minute granules in branchial area.
Carapace surface and pereopods densely covered with setae of two kinds: short plumose setae, bent at right angles
near the tip, clothing surface, but interspersed with longer filiform setae (6 x length of short setae and 0.20-0.25 x
CW) which also fringe limbs and arranged in clumps on carapace where there are about fifteen to seventeen distinct
groups (each with up to about four setae) which tend to be associated with rounded surface elevations. Density of
setae completely obscures body surface but most of this is attributable to short rather than long setae. Structure of
short and long setae is different. In short setae proximal 45% of shaft has very short setules (at end of which setae
arc sharply angled), then a region occupying about 45% where long, stout setules are directed almost at right
angles to shalt, forming a dense bunch, and finally the distal 10% of setae which is smooth, slightly curved, and
narrows to an acute tip. In long setae almost entire length is covered with small dendritic setules, all about same
size, and setae are acutely tipped.
A shallow frontal carapace groove separates a pair of low rounded protuberances, and then divides into separate
grooves which gradually become more faint. Just in front of cardiac region two laterally-directed grooves originate:
first groove (cervical) arises separately from small pits curving (slightly sinuously) anteriorly on to branchial
region, w'hilc second shallower groove extends across mid-line and initially runs almost directly towards lateral
margin but then splits into an anterior portion which follows the first groove for a short distance, while the
second portion curves posterolaterally, bordering anterior cardiac region. In effect the groove crossing the mid-line,
connects two crescent-shaped grooves. Branchial groove not evident. Posterior cardiac area not defined.
Anterolateral carapace margin begins just below level of postorbital corner, is evenly convex and bears four
distinct, broad-based, equidistant teeth, each ending in a small, acute spine, and accompanied by a tuft of long
496
CL. McLAY
setae. First two teeth directed anteriorly, third directed anterolaterally, and last directed more laterally.
A posterolateral tooth, smaller than preceding anterolateral teeth, marks beginning of convergent posterolateral
border alongside which lies the reduced last leg. Posterior carapace margin is recessed in order to accommodate first
segment of abdomen which is visible dorsally.
Frontal margin continuous, V-shaped, ventrally-directed, joined to epistome (which separates orbits). Supra¬
orbital margin not projecting, continuous above orbits, interrupted by a distinct notch closer to postorbital corner
which is without granules; suborbital margin has a few small granules and angles towards an acute tooth (visible
dorsally when setae are removed) then drops sharply into a notch before ending in a much smaller, blunt tooth at
its inner corner. This tooth abuts second article of antenna. Orbits clearly exposed dorsally.
First article of antcnnulc large, filling a large part of ventral orbital region; distal margin obliquely angled and
not continuous with distal margin of second antennal article. Remainder of antcnnulc folded into orbit. First article
of antenna moveable, wider than long, medially beaked; inferior tooth well developed, blunt; superior tooth, above
opening of antennal gland, is smaller. Second article wider than long; distal margin widest, to which is fixed the
exopod curving over base of eyestalk and becoming broader and terminating bluntly. Exopod has a tuft of setae on
its lateral margin. Third antennal article longer than wide, and attached to remaining distal border of second article,
slotting in behind exopod, and just matching length of exopod. Fourth antennal article smaller, as long as wide;
remaining antennal articles directed laterally, extending well beyond postorbital corner, and can be partially folded
under supra-orbilal margin. Ratio of length of antennal flagella to CW = 0.33. Eyestalk can be completely folded
into orbit, and cornea is well developed, occupying all of tip. Epistome broadly triangular, surface slightly
concave; dorsal arm. joined to tip of carapace, very elongate and narrow; lateral arms shorter and thicker. Joint
between epistome and carapace marked by a narrow suture.
Subhepatic area smooth, very convex. A groove begins near base of the antenna, curving round under branchial
region and meeting lateral carapace margin just anterior to last tooth at beginning of posterolateral border. A short
cervical groove branches off and ascends towards gap between first and second anterolateral teeth, branching, with
one branch meeting first anterolateral tooth ventrally and the other passing around behind second tooth. Third
maxillipeds operculiform, bases widely separated by tip of sternum. Crista dentata has six or seven well developed,
distally placed teeth on each side and a granulated border on outer margin. Female sternal sutures 7/8 short, ending
wide apart on low tubercles just behind bases of second walking legs.
Gill formula 19 gills + 7 epipods on each side, as found in Dynomene hispida. There is no podobranch on last
pereopod. In cross section gills have the lateral margin deeply notched, dividing it into a hypobranchial plate
(containing efferent vessel) and an epibranchial lobe (anterior lobe longer). Between these marginal lobes are a pair
of lobes, similar to marginal lobes. Thus the epibranchial surface shows four rows of blunt lobes, decreasing in
size from anterior to posterior side, which are arranged above afferent blood vessel. Hypobranchial setae poorly
developed. Posterior margin of scaphognathite with three long setae. Hypobranchial margin of podobranchs bears
same setae as on epipod.
Cheliped stout, much longer than first leg and stouter in the male, only slightly longer and stouter than
first leg in the female; merus trigonal, inner face smooth and fitting closely against pterygostomial region
of carapace, borders granulate, superior border has a subterminal broad, restriction which separates a thick¬
ened distal ridge, on which there are three small granules, from a row of three to five similar granules on superior
border. Inner inferior margin of merus has an acute lateral spine distally which is especially prominent in large
(CW > 20 mm) males where the spine forms a tooth which may itself be granulated. Outer face of carpus convex
with six small granules, two more prominent acute tubercles on distal margin, inner superior border with a
flattened, distomedially directed, spur (granulated in large males) which abuts against proximal inner surface of
propodus thereby restricting closure of cheliped against frontal area. In a similar way. inferior carpal margin is
produced as a smooth obtuse flange fitting against merus when limb is withdrawn. These two structures give
carpal article an unusual and distinctive shape. Outer face of propodus with two or three very small granules;
superior face with three parallel rows of small granules; inner and inferior faces smooth, except that there is a
small proximal spur on the inner propodal face. Fixed finger almost straight with six teeth increasing in size
distally (large males have an additional basal tooth and some of the other teeth can be rudimentary); moveable
Source : MNHN. Paris
FAMILY DYNOMENIDAE
497
-t^A*
1 cm a-b, e, g
g
Fig. 21. — Dynomene pilumnoides Alcock. 1900: a-g, 9 14.6 x 11.7 mm. Sulu Archipelago. B. R. Wilson coll.
(MNHN-B 10374): a. dorsal view of right half of carapace; b, ventral view of right orbital area; c. outer face of right
cheliped; d. dorsal view of right cheliped; e. posterior view of terminal articles of right fourth pereopod; f. posterior
view of terminal articles of right fifth pereopod; g, ventral view of telson and terminal segments of female abdomen.
Source : MNHN, Pahs
498
C.L. McLAY
finger curved with four teeth, first mid-way and separated from remainder which arc on tip; both fingers, thick,
hollowed out internally, touching only at tips where the teeth interlock. Just below proximal teeth on fixed finger
are two distinct pits in which several long setae are inserted with a similar group of setae on inner margin. Groups
of long stiff setae, inserted mid-way along dactyl and fixed finger, are directed across space between the two fingers
to form a screen.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of meri of first two
legs and anterior face third leg merus smooth and nacreous, inferior distal margin hollowed out to accommodate
carpal article. Superior border of meri of these legs with several small granules, length of merus of second leg
about 2. 0-3.0 x width and equal to about half of CL. Anterior and posterior dorsal margins of carpi bearing several
small granules, and produced distally to overhang base of propodi. Dorsal surface of propodi smooth. Dactyli
curved, inferior margin armed with 4-5 small spines, tip dark brown and subacute.
Last pair of legs greatly reduced, lying along posterolateral border of carapace, reaching only as far as a half to
two-thirds along meral article of preceding limb; borders of articles unarmed. Last pair of legs subchelate, sexually
dimorphic: female with well developed distal extension of propodus which opposes dactyl, male with only weakly
developed propodal extension. Female propodal extension bearing six, unequal, stout, hooked, spines each lined
with marginal rows of 12 - 20 tiny flattened, acute teeth along middle region of inner surface, remainder of spine
marked by transverse ridges. Female dactyl as long as propodal extension, bearing thirteen unequal, stout, hooked
spines (arranged asymmetrically around perimeter of the dactyl) whose inner surface is smooth, devoid of teeth but
strengthened by a longitudinal ridge. Male propodal extension bearing five unequal hooked spines, one of which is
submarginal, all devoid of teeth along inner surface. Male dactyl longer than propodal extension and ending in a
single acute claw.
All segments of abdomen freely moveable, increasing in length and breadth distally; surface smooth; margins
unarmed but fringed with long setae. Anterior margin of second segment sinuous, medial region convex, lateral
margins produced as a flange which fits over posterior margin of first segment preventing forward slippage of
abdomen. Subsequent segments not overlapping with preceding segments. Telson much wider than long, anterior
margin angled to accommodate uropod. posterior margin broadly rounded. In female uropod plates large, filling
about two-thirds of space between last abdominal segment and telson, excluding most of last abdominal segment
and telson from reaching lateral margin of abdomen. In male last abdominal segment occupies about a half of
length. No effective abdominal locking mechanism: abdomen only loosely held against sternum in both sexes but
in males its lateral movement is restricted by presence of a small tubercle on sternum adjacent to each of the
second pereopods. In mature female it occupies all the ventral surface, covering coxae of all pereopods with telson
covering proximal half of third maxillipeds. In male abdomen is not quite so broad and telson only extends as far
as bases of third maxillipeds.
Five pairs of pleopods in female, first pair vestigial, remainder biramous. First male pleopod a semi-rolled
tube with a small apical plate surrounded by long setae. Second male pleopod with an exopod on basis, needle-like
distally, armed with a series of fifteen tiny, straight, acute, inset spines and ending in two larger spines. Terminal
pair of spines are slightly curved at their tips. Third to fifth male pleopods rudimentary and biramous, exopod
longer and connected to basal article by a joint.
COLOUR. — Sakai (1965a) described the colouration of Dynomene pilumnoides as being yellowish red.
Miyake (1983) shows the body and legs as light reddish-orange fringed with yellowish setae, and the eyestalks as
being ied. The picture of Baba, Hayashi, and Toriyama (1986) shows the body and legs to be yellowish-brown
fringed with dark brown setae.
Geographic Distribution. — Dynomene pilumnoides has been recorded from Natal (as Maxillothrix
actaeiformis Stebbing, 1921), Madagascar, Laccadive Ids, Philippine Ids, Japan. Australia, New Caledonia
Loyalty Id. Norfolk Ids, Enewetak Atoll, Marshall Ids and Hawaii. The material reported here from Australia,
Norfolk Id, New Caledonia and Loyalty Ids represent new records and the specimen collected by the USFC
Steamer " Albatross " in 1902, from Hawaii, has languished unrecognized in the collection of the U. S National
Museum for more than 90 years. This is a widespread Indo-Pacific species.
Source : MNHN. Pans .
FAMILY DYNOMENIDAE
499
Depth. — Depth range of the material examined is 18-400 m, although most records are from 100-300 m.
The specimens reported by Baba et al (1986) came from the Okinawa Trough, 362-540 m, but given the other
more precise depth records, it does not seem justifiable to conclude that the lower depth limit for this species is
540 m. The lower depth limit is probably around 400 m. This species has been collected from rocky bottoms
covered in lithothamnion algae, sponges, and corals. Dynomene pilumnoides does not seem to be associated with
particular corals, although this could be the result of most specimens being collected from deeper water by dredge
and separated from their host. The range obviously includes depths at which corals do not occur.
SIZE. — The maximum size for males is 28.5 x 22.1 mm, for females 20.7 x 17.6 mm, and the smallest
ovigerous female is 9.5 x 7.5 mm. This female carried around 590 eggs while the largest ovigerous female with a
complete brood ( 14.8 x 12.0 mm) carried about 1600 eggs. Mean egg diameter was 0.49 mm. The breeding season
may be tentatively estimated as extending from February to September although no ovigerous females have been
collected during the months from April to July. The maximum size for Dynomene pilumnoides is somewhat
larger than for the preceding Dynomene spp. and the size of the smallest ovigerous female is also correspondingly
larger. The larval stages of D. pilumnoides are unknown.
One female specimen (7.5 x 6.2 mm) collected in 1993 from a reef in the New Caledonia lagoon shows some
peculiar features: there are no gonopores on the coxae of the second walking legs and the only indication of its sex
is the presence of sternal sutures 7/8, terminating in the spermathecal openings, and five pairs of pleopods.
However all pleopods are remarkably poorly developed, perhaps indicating that it has been parasitized.
Discussion. — The exact status of Dynomene pilumnoides has been one of the main problems in understan¬
ding the relationships between the species of Dynomene , especially those from the Indian Ocean. ALCOCK's type
specimen was a male 1 1.0 x 10.0 mm Irom 54-90 m off the Laccadive Ids. Although he created a new species for
this single specimen, ALCOCK (1900) suggested that it could well prove to be a variety of D. hispida. Both spe¬
cies have a similar carapace shape, although the CW/CL ratio tends to be larger (approx. 1 .30) for D. hispida , and
the anterolateral carapace margins have four teeth, but the teeth are much smaller in D. hispida. The features
which clearly distinguish D. pilumnoides from D. hispida are the presence of clumps of long filiform setae which
are much longer than the surface covering of short setae (D. hispida carapace setae mostly short and not clumped),
a notch in the supraorbital margin (no notch), postorbital corner without spines (five small acute spines), and only
a single spine on the suborbital margin (five small acute spines). D. pilumnoides grows to a larger size and is
usually collected from 100-300 m depth whereas D. hispida is a small dynomenid which occurs in shallow waters,
often intertidally. It seems likely that ALCOCK did not have a specimen of D. hispida at his disposal which would
have allowed a detailed comparison with D. pilumnoides. The differences between the species listed by ALCOCK
(1900) are not diagnostic. However, the differences listed by ALCOCK ( 1901 ) are more useful and here he withdrew
the suggestion that D. pilumnoides might be a variety of D. hispida. ALCOCK’s description is seriously wrong in
only one aspect: his statement that there are no rudimentary pleopods on the third to fifth abdominal segments in
the male. (I his feature is present in all dynomenid males although they are sometimes difficult to see clearly wi¬
thout special preparation.) Also his illustration does not show the longer carapace setae arranged in clumps.
Other features of Dynomene pilumnoides , not illustrated here, can be found in Stebbing (1921) (as
Maxillothrix actaeifonnis): mandible, second maxilla, second and third maxillipeds, and the first male pleopod.
Odhner (1925) pointed out that Maxillothrix is a junior synonym of Dynomene and therefore should not be
placed amongst the Xanthidae as Stebbing had proposed. The synonymy of M. actaeifonnis and D. pilumnoides
was first recognized by BARNARD (1947).
The generic name Maxillothrix Stebbing. 1921 was created to reflect the presence of three long, unequal setae
on the scaphognathite of the second maxilla. In fact similar setae had already been reported by Bouvier ( 1896) for
Dynomene filholi (see above) but Stebbing had placed his new genus amongst the Xanthidae, comparing the
second maxilla with that found in Nursia , so the apparent uniqueness of these setae is understandable. Stebbing
(1921) noted that the setae were reminiscent of the long scaphognathite spine in Axius longispina and he drew
attention to the two long terminal setae on the second maxilla of Homola andamanicus (ALCOCK. 1901, pi. A).
These setae probably have a role in cleaning the epibranchial surface of the gills.
Source :
500
C. L. McLAY
Barnard (1947, and 1950) continued to suggest that Dynomene pilumnoides is probably a synonym of
D. hispida, but this cannot be correct because the suborbital margin is visible dorsally (suborbital margin straight
and not projecting), CW/CL = 1.17-1.2 (CW/CL = 1.3), and the longer setae on carapace are clumped (setae not
clumped). The specimens available to Barnard were collected from 90 m depth, whereas D. hispida is an
intertidal-shallow water species. The confusion about the status of the name D. pilumnoides Alcock, 1900 arose,
at least in part, because of the errors and omissions of the original descriptions of ALCOCK and Stebbing (for
Maxi doth rix actaeiformis).
Barnard (1950: 336) gave the branchial formula for Dynomene pilumnoides as 18 gills + 3 epipods. But this
is incorrect because he missed the two arthrobranchs on the third maxillipcd and overlooked the epipods on the
pereopods. The branchial formula is 19 gills + 7 epipods. The gill structure of D. pilumnoides more closely
resembles that of D.filholi than of D. hispida and D. praedator. The epibranchial surface of the gills is made up of
four rows of elongate lobes which radiate out from the afferent vessel. The epipods and hypobranchial setae are
similarly developed in all these species. Another character which D. pilumnoides and D. filholi share, is the
presence of three long setae on the posterior margin of the scaphognathite. The other species of Dynomene only
have two setae. In D. pilumnoides the hypobranchial margin of each podobranch carries long cleaning setae as
found in D. hispida and D. praedator.
The subchclate structure of the last pairs of legs of females shows several differences from the other species of
Dynomene : the propodal extension of the female bears six spines (four in the other species), and the teeth on the
inner surface of these spines are fewer in number (12-20 teeth) and arranged in marginal rows in the middle region
(rather than scattered along most of the spine). The dactylar spines of the female are strengthened by a longitudinal
ridge. Male D. pilumnoides , along with D. filholi , lack the teeth on the propodal spines which arc found in the
other two species.
The second male pleopod of Dynomene pilumnoides is similar to that of D. filholi in having a large number
(15) of subterminal spines. Also the rudimentary last three pairs of pleopods are biramous in both species.
Examination of the stomach contents of a Dynomene pilumnoides female 14.6 x 12.0 mm (MNHN-B 6913)
from Madagascar revealed mostly soft amorphous material with a few sand grains and some chitinous brown tube¬
like fragments that may be hydroid in origin. It may be that this crab uses its chelae to spoon up sediment from
which organic fragments are separated by the groups of setae on the cheliped fingers.
Dynomenid crabs very rarely have epizoites on their exoskeleton, perhaps because of the dense layer of setae.
However, one large Dynomene pilumnoides male 28.5 x 22.1 mm (SMF 17127) from Japan, had a small
polychacte tube on the right dorsal side of the carapace. It may be possible for fouling organisms to colonize
larger crabs because of their longer intermoult intervals.
Dynomene pugnatrix de Man. 1889
Figs 5 d. 1 1 , 22 a-g
Dynomene pugnatrix de Man, 1889: 444, pi. 10, fig. 13. — ALCOCK, 1901: 75 (list). — IHLE, 1913: 92 (list). — GuiNOT,
1967: 242 (list). — Ser£ne, 1968: 36 (list). — Takeda. 1977: 35 (list).
Dynomene pugnatrix brevimana Rathbun, 1911: 196. — GuiNOT, 1967: 242 (list). — SERENE, 1968: 37 (list).
Material EXAMINED. — Mauritius: No details, no depth, no date: I 6 9.8 x 7.2 mm, type specimen (SMF
Providence Island. Percy Sladen Trust Expedition: stn D4, 9°14.00’S. 51°02.00'E. 90-140 m. 4.10.1905:
1 9 ovig. 6.3 x 4.8 mm. type specimen of D. pugnatrix brevimana (USNM 41047).
Types. — Dynomene pugnatrix de Man, 1889: holotype is a male 9.8 x 7.2 mm, from Mauritius, held by the
Natur-Museum Senckenberg, Frankfurt, registration number SMF 4857.
Dynomene pugnatrix brevimana Rathbun, 1911: holotype is an ovigerous female 6.3 x 4.8 mm, collected by
the R/V " Sealark ", Percy Sladen Trust Expedition, from stn D4, 9°I4.00’S, 51°02.00'E. Providence Id. north
ot Madagascar, 90-140 m, 4.10.1905, held at the Smithsonian Institute, Washington, registration number
USNM 41047.
Source :
FAMILY DYNOMENIDAE
501
Description. — Carapace wider than long, ratio of CW/CL approx. 1.30, squarish in outline, surface convex,
minutely granular. Carapace surface and percopods sparsely covered with two kinds of setae of varying lengths up
to 0.3 x CW. There are short stiff setae and longer "feathered" setae. The density of setae does not completely
obscure body surface. Microscopic details of setae not examined.
Frontal carapace groove very faint, extending only a short distance from frontal margin. The normal pair of
prominent rounded protuberances, separated by frontal groove, absent. Just in front of cardiac region two laterally-
directed grooves originate: first groove (cervical) crosses mid-line and runs directly anterolateral on to branchial
region and mid-way along its length there is a short groove in the direction of median frontal margin. The second,
shallower groove extends across mid-line and initially runs almost directly towards lateral margin but then splits
into a short anterior portion which follows the first groove for a short distance, while the second portion curves
posterolatcrally, bordering anterior cardiac region. In effect this second groove connects two crescent-shaped
grooves. No branchial groove evident and posterior cardiac area only faintly defined. Anterolateral carapace margin
begins below level of postorbital corner, initially it is a horizontal margin directed laterally, with two or three
small granules, until meeting two small, subacute anterolateral teeth close together at corner. Be'hind these teeth
margin runs posteriorly and has three similar equidistant teeth, first directed almost anteriorly, next two directed
laterally, and finally there is a smaller submarginal tooth part-way along convergent posterolateral margin
alongside which lies the reduced last leg. This gives a total of six teeth on the right-hand side, but on the left-hand
side there is only one initial tooth, giving the more normal pattern for this genus of five teeth (the last of which
is treated as being posterolateral). In the case of this species, absence of the branchial groove makes this
distinction rather arbitrary. Posterior carapace margin recessed in order to accommodate first segment of abdomen
which is visible dorsally.
Frontal margin continuous, V-shaped, ventrally-directed, joined to epistome (which separates orbits).
Supraorbital margin not projecting, continuous above orbits, interrupted by a distinct notch closer to postorbital
corner, following margin minutely granulated; suborbital margin with two small granules. Orbits clearly exposed
dorsally.
First article of antennule large, filling a large part of ventral orbital region, distal margin obliquely angled and
not continuous with distal margin of second antennal article. Remainder of antennule folded into orbit. First article
of antenna moveable, wider than long, medially beaked, inferior tooth well developed, blunt, superior tooth above
the opening of antennal gland slightly smaller. Second article wider than long, distal margin widest, to which is
fixed the exopod curving over base of eyestalk and becoming broader and terminating bluntly. Third antennal
article longer than wide, and attached to remaining distal border of second article, slotting in behind exopod, and
just matching length of exopod. Fourth antennal article smaller, as long as wide, remaining antennal articles
directed laterally, extending well beyond postorbital corner, and can be partially folded under supra-orbital margin.
Ratio of length of antennal flagella to CW = 0.67. Eyestalk can be completely folded into orbit, and cornea well
developed, occupying all of tip. Epistome broadly triangular, surface slightly concave; dorsal arm, joined to tip of
carapace, very elongate and narrow; lateral arms shorter and thicker. Joint between epistome and carapace is marked
by a narrow suture. Antennae, antennules and epistome fit closely together.
Subhepatic area convex, minutely granulated. A groove begins near base of antenna, curving round under
branchial region and meeting lateral carapace margin just anterior to last tooth at beginning of posterolateral
border. A short cervical groove branching off and ascending towards first anterolateral tooth. Third maxillipeds
operculiform, bases widely separated by tip of sternum. Crista dentata has five or six small, blunt, distally placed
teeth on each side. Female sternal sutures 7/8 short, ending wide apart on low tubercles just behind bases of
second walking legs.
No information about gill formula due to insufficient material, but structure of an arthrobranch taken from the
cheliped is as follows: in cross section, the anterior and posterior margins of the gill are notched, dividing it into
hypobranchial plates and epibranchial plates which end bluntly. The epibranchial plate on the anterior margin is
larger and between these plates is a single row of short elongate lobes.
Cheliped slender, slightly longer than first leg. Merus trigonal, inner face smooth and fitting closely against
pterygostomial region of carapace, borders granulate, outer face has a subterminal shallow, restriction which
separates a thickened distal ridge devoid of granules from a pair of small subacute granules, preceded by a row of
502
C. L. McLAY
several smaller granules on superior border. Inner inferior margin of merus has an acute lateral spine. Outer face of
carpus convex, smooth, inner superior border with a distomedially directed, sharp spur which abuts against
proximal inner surface of propodus thereby restricting closure of cheliped against frontal area. In a similar way.
inferior carpal margin is produced as a smooth, obtuse, flange fitting against merus when limb is withdrawn.
These two structures give carpal article an unusual and distinctive shape: inner face very narrow and outer face
much broader. Surface of propodus smooth; fixed finger almost straight with three teeth at tip; moveable finger
strongly curved with a single, blunt proximal tooth and three teeth at lip. Both fingers, thick, hollowed out
internally, touching only at tips where last three teeth interlock. Just below proximal teeth on fixed finger are two
distinct pits in which several long setae arc inserted.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of mcri of first two
legs and anterior face third leg merus smooth and nacreous, inferior distal margin hollowed out to accommodate
carpal article. Superior border of meri of these legs with five well developed spines in a row, increasing in size
distally, separated by a gap from a single distal spine, and five well developed spines on posterior margin; length
of merus of second leg about 2.4 x its width and equal to about three-quarters of CL. Dorsal surface of carpi
bearing a row of five acute spines, and produced disially to overhang base of propodi. Dorsal surface of propodi
with a row of four small spines. Dactyli curved, inferior margin armed with 10 small spines, tip brown and
subacute.
Last pair of legs greatly reduced, adorned with long setae distally, lying along posterolateral border of carapace,
reaching only as far as one third along meral article of preceding limb; borders of articles unarmed. Legs sub¬
chelate, sexually dimorphic: female with well developed distal extension of propodus which opposes dactyl; male
with only weakly developed propodal extension. Structural details of dactyl and propodus have not been
investigated.
All segments of abdomen freely moveable, surface smooth, margins unarmed but fringed with setae. Second
segment narrowest, anterior margin sinuous, medial region convex, lateral margins produced as a flange which
fits over posterior margin of first segment (which is the shortest) preventing forward slippage of abdomen.
Subsequent segments increasing in length and breadth distally, not overlapping with preceding segments. Telson
much wider than long, anterior margin angled to accommodate uropod. posterior margin broadly rounded. In male,
uropod plates large, filling about three-quarters of the space between last abdominal segment and telson, excluding
much of last abdominal segment and telson from reaching lateral margin of abdomen. No effective abdominal
locking mechanism: abdomen only loosely held against sternum. Abdomen extends as far as bases of the third
maxillipeds.
hive pairs ol pleopods in female, first pair vestigial, remainder biramous. Five pairs of pleopods in male, first
pleopod a semi-rolled tube ending in a curved, blunt apical plate surrounded by long setae, second plcopod needle¬
like with an exopod on the basis, remaining pleopods rudimentary. Microscopic details of second male plcopod
unavailable.
Colour. — Unknown.
Geographic distribution. — The type locality is Mauritius and the other specimens reported by Rathbun
(1911) as Dynomene pugnatrix brevimana came from Providence Id. both of which are in the vicinity of
Madagascar, although D. pugnatrix is not known from Madagascar itself. Thus IX pugnatrix is a western Indian
Ocean species. It is interesting to compare this species with D. fillioli which is an insular South Atlantic species.
DEPTH. — The depth range for Dynomene pugnatrix is 90-140 m (based on the specimens reported by
Rathbun (191 1 ), as D. pugnatrix brevimana.
SIZE. — Only three specimens of Dynomene pugnatrix are known: one male, 9.8 x 7.2 mm (the type
specimen), and two ovigerous females, one of which is 6.2 x 5.0 mm. The size of the other ovigerous female
reported by Rathbun (191 1) is unknown. It is evident that this species reaches sexual maturity at a small size as
do D. luspida and D. praedator. The specimen of D. pugnatrix examined carried 30 eggs but some eggs may have
been lost because the brood did not fill the entire abdominal space. Egg diameter is 0.4 mm. These reproductive
aspects are similar to the other Dynomene species.
Source :
FAMILY DYNOMENIDAE
503
FlG. 22. — Dynomene pugnatrix de Man, 1889: a-g, 6 9.8 x 7.2 mm, type specimen. Mauritius (SMF 4857): a, dorsal
view of right half of carapace; b. ventral view of right orbital area; c, outer face of right cheliped; d. dorsal view of
right cheliped; e. posterior view of terminal articles of right fourth pereopod; f, posterior view of terminal articles of
right fifth pereopod; g. ventral view of telson and terminal segments of male abdomen.
Source
504
C.L. McLAY
DISCUSSION. — De Man (1889) illustrated the type specimen of Dynomene pugnatrix from Mauritius
showing the left cheliped missing and stating that it was a female, but in fact the specimen is a male. De Man
stated "I judge from the shape of the telson, whose last segment is shaped like a half circle and rounded, that this
is a female" (translated from German). Apart from the pleopods, the dynomenid telson and abdomen as a whole is
sexually dimorphic only in size, not in shape. It is also possible that he judged the sex on the basis of the size of
the abdomen which, in dynomenid males, is much larger than might be expected.
Since the report of the original specimen only two further specimens (both ovigerous females) have been
recorded. These were described by RATHBUN as a subspecies, Dynomene pugnatrix brevimana Rathbun, 1911.
Both specimens are a little smaller than de Man's type. The differences alluded to by Rathbun (191 1)
are 1) greater carapace width/length ratio, 6.3/4.8=13 vs 9.75/8.25= 1.2 (as reported by DE Man, 2) palm
shorter in relation to the fingers, 3) presence of a few spinules on the upper edge of the palm, and 4) the wrist
and chelae have a few setae. A comparison of RATHBUN's specimen reveals some inconsistencies and inaccuracies.
My measurements of the carapace of the type are 9.8 x 7.2 mm which gives a ratio of 1.36 which is much
closer to the ratio of Rathbun's specimen. De Man described the cheliped as "The length of the hand is more or
less three-quarters of the length of the carapace. Hand laterally compressed and more or less twice as long as high
and a little longer than the fingers..." (translated) and because he reported his type as being a female, Rathbun
was correct in noting the difference in relative cheliped size. But in fact the type is a male and the difference in
cheliped size is simply a result of sexual dimorphism. The cheliped carpus and propodus were described as "The
carpus and the hand are entirely smooth and without setae except the fingers" (translated). Inspection of DE Man's
type shows that there arc in fact setae on the wrist and chela and they were not detected or illustrated. The only
difference between these specimens is the presence of spinules on the palm of Rathbun's females. There are
no spinules on DE Man's type but this does not seem very important given the variability seen in
other dynomenid species. I think that D. pugnatrix brevimana Rathbun, 1911 should be synonymized with
D. pugnatrix de Man, 1889. All three known specimens come from the same general area in the vicinity
of Madagascar.
In his original illustration of the male type of Dynomene pugnatrix , DE Man (1889) showed setae concen¬
trated in the anterolateral regions of the carapace and along the margins of the walking legs as well as on the
reduced last pair of legs. Furthermore he figured two kinds of setae from the legs: short, stiff "Kammhaar" (comb¬
like) setae and "Federhaar" (feather-like") setae (1889, pi. 10. fig. 13e-f). Due to the rarity of this species I was
unable to verify DE Man's observations using the electron microscope. The setae of D . pugnatrix appear to be
rather different from those of the other species of this genus.
The structure of the gills of Dynomene pugnatrix differs from that found in D. hispida only in that there is a
single row of epibranchial lobes between the plates. The branchial formula is unknown.
Genus HIRSUTODYNOMENE nov.
Diagnosis. Carapace much wider than long, moderately convex, commonly subcircular. Surface
sparsely spinous (especially in anterobranchial region), areolate, covered with coarse setae, which are short and
long, and arranged in tufts. Lateral carapace margin always well defined and armed with distinct teeth.
Frontal groove well marked, split in two posteriorly; cervical, postcervical and branchial grooves usually evident.
Frontal carapace margin broadly triangular, continuous; no rostrum or teeth. Eyestalks short, eyes protected by
well defined orbits. Sternal sutures 7/8 of female end well apart on low tubercles behind bases of second
walking legs.
Antennule can be concealed inside orbit at base of eyestalk. Antennal flagella shorter than carapace width. All
articles ol antenna moveable; first article (urinal) always beaked medially; second article has an exopod firmly
fixed. Ihird maxillipeds opercular, completely covering buccal cavern, separated at their bases by a plate at same
level as sternum; basis and ischium of endopod fused but joint always marked by a shallow groove. Crista dentata
present. Chelipeds equal, stouter than walking legs; dactyl strongly curved; fingers gaping basally. Last pair of
Source : MNHN. Paris
FAMILY DYNOMENIDAE
505
legs very reduced; dactyl rudimentary, forming an obsolete subchelate mechanism with an extension of propodus.
Gills usually 19 (including 6 podobranchs) + 7 epipods. Gill structure basically phyllobranchiate but plates are
very variable in shape.
Abdomen of six segments and telson folded loosely under thorax; uropods large; no effective abdominal
locking mechanism. Lateral movement of abdomen restricted by small sternal tubercle, at base of each of first
walking legs, which lies alongside each uropod. Both sexes have five pairs of pleopods; first pair vestigial in
female; last three pairs rudimentary in male. Male pleopods uniform in structure; first pair consist of a stout,
setose semi-rolled tube with an apical plate; second pair needle-like with numerous subdistal spines, some of
which overlap, sinuously arranged around the axis.
Type Species. — Dynomene spinosa Rathbun, 1911.
Other Species. — Dynomene ursula Stimpson, 1860.
Etymology. — Hirsutodynomene is a combination of the latin hirsutus, meaning shaggy, alluding to the
tomentum of these species, and the genus Dynomene. Gender is feminine.
DISCUSSION. — This new genus is erected for two species originally assigned to Dynomene. They clearly
stand out amongst the other species because of their "shaggy" appearance, resulting from long, stiff setae, and
their spiny, areolate carapace. The distribution of these two species does not overlap in the Pacific and they are
clearly sister species.
Hirsutodynomene spinosa (Rathbun, 1911)
Figs 3 f, 5 e-f, 8 f, 9 c, 1 1, 13 a-b, d, 14 d. 17 e, 23 a-g
Dynomene spinosa Rathbun, 1911: 196, pi. 17, fig. 1. — Ihle. 1913: 92 (list). — Balss, 1935: 115; 1938: 7. —
Miyake, 1939: 198 (list). — Ward, 1942: 71. — Holthuis, 1953: 3. — Morrison, 1954: 13. — Guinot, 1967: 242
(list); 1985: 448 (list). — SerLne, 1968: 36 (list). — Takeda, 1973: 81; 1977: 35 (list). — Peyrot-Clausade &
Serene, 1976: 1343, pi. 2 A. — Chen, 1980: 1 19, pi. 1, fig. 2. — Dai, Yang & Lan. 1981: 1 17, tex- fig. 1-4. —
Peyrot-Clausade. 1981: 750; 1984: 1 14. — Dai, Yang, Song & Chen, 1986: 27. pi. 3, 1. — Garth^ Haig &
Knudsen. 1987: 241. — Dai & Yang, 1991: 31, pi. 3. fig. 3. — Poupin, 1996a: 24 (list).
Dynomene hispida - DE Man, 1902: 689. (Not Guerin-Meneville, 1832).
Material examined. — Madagascar. Tulear, stn 14- 1 1-2. pente externe, 5 m, M. Peyrot-Clausade coll.,
1968: 1 9 1 1.3 x 8.9 mm (MNHN-B 22077) (see Peyrot-Clausade & SerLne, 1976; and Peyrot-Clausade, 1984).
Glorieuses Islands. Grande Glorieuse. Intertidal zone, A. Crosnier coll., 30.01.1971: 1 6 16.4 x 14.3 mm
(MNHN-B 6899).
Australia. W coast: Exmouth Gulf, Bundegi Reef, 21°53’S, 1 14°22’E, 3 m. under rock. N. Coleman coll..
14.08.1972: 1 6 19.6 x 14.5 mm (AMS-P 19118).
Southeast Queensland , Flinders Reef, 26°59'S, 153°29’E, 6-20 m, 10.03.1989: 1 9 27.7 x 20.4 mm (QM W 16277).
Tasman Sea , Middleton Reef, 29°29.5’S, 159°06.2’E. no depth, P. Filmer-Sankey coll., 5.12.1987: 1 6 28.7 x
21.3 mm (10 poecilasmid barnacles on chelipeds) (AMS-P 39242). — No depth, P. Hutchings coll., 8.12.1987:
1 9 ovig. 32.3 x 24.4 mm (AMS-P 39245). — Elizabeth Reef, 29°57.2’S, 159°0I.2,E, 12 m. dead coral rubble,
R. Springthorpe coll., 10.12.1987: 1 9 19.5 x 14.8 mm (AMS-P 38264).
Cocos Keeling Islands. Horsburgh Id. 0-37 m, 9.02.1989: 1 6 14.2 x 10.8 mm (WAM 139-94). — N end of
West Id, 0-30 m, 21.02.1989: 1 9 13.0 x 10.3 mm (WAM 138-94.). — NW end of West Keeling Id, 0-28 m,
23.02.1989, F. Wells coll.: 1 9 23.8 x 17.8 mm (WAM 723-89).
Indonesia. Moluccas. Amboina: no details, J. Brock coll., 7.09.1885, J. G. DE Man (1888) det.: I 6 17.8 x
13.7 mm (SMF 164). — Seram, Gorong Id. Rumphius 2, stn GO 3, no depth. 27.01.1975, T. Monod & R. SerLne coll,
and det. as Dynomene pilumnoides : 1 6 17.5 x 14.0 mm (MNHN-B 9906). — Ternate: no details, W. Kukenthal coll.,
1894: 19 18.5 x 14.3 mm (SMF 4856) (See DE Man, 1902 who reported this specimen as Dynomene hispida).
— Timor, Atapupu, coral reef, leg. "Gazelle", no depth, no date: 1 6 24.3 x 18.8 mm (ZMB 5138).
Vietnam. Institut oc6anographique, Nhatrang (coll, and det. R. Serene): stn 1060, no locality, no depth. 1950:
1 <3 18.0 x 14.3 mm (ION. 9265) (ZRC 1970.6.20.1). — Stn. 87, no locality, no depth. 1950: 1 6 (dry) 7.4 x 6.0 mm
(ION 1556) (ZRC 1970.6.20.2).
Source :
506
C. L. McLAY
Mariana Islands. Asuncion Id, 19°40'N, I45°24'E. 1-6 m along rock wall in holes and corals. P. Schupp coll.,
7.06.1992: I 9 24.2 x 18.0 mm (UGM).
Guam , Piti Lagoon, 13°27'N, 144°47’E, 1.0-2. 5 m among dead coral, 20.09.1992: 1 6 20.3 x 15.9 mm. — 1-2 m
under rubble, H. T. Conley coll., 04.1997: 1 6 10.2 x 8.1 mm. — Turnon Bay, 11 m on dead finely branched coral.
R. K. Kropp & J. 11. Dominguez coll., 7.1 1.1984: 1 9 4.4 x 3.9 mm (UGM)
Japan. Kurashima Ids, Yaeyama Id. Okinawa, in dead coral branches, inner reef, M. Osawa coll., 1993: 1 6 22.2 x
17.0 mm.
Line Islands. Whipp Expedition: Palmyra Id, 5°52'N, 162°6'W. no depth, 1924: 1 6 10.5 x 8.6 mm (BPBM 2297).
French Polynesia. Tuamotu Ids, Raroia Atoll. Homohomo Id. 16°3'S, 142°23'W, under rocks near shore in
pavement pool zone. J. P. E. Morrison coll., 21.07.1952: 1 9 25.0 x 19.8 mm (USNM 94559) (see Holthuis, 1953).
TYPES. — Dynomene spinosa Rathbun, 1911: holotype is a male 24.7 x 19.6 mm, collected by the R/V
"Sealark", Percy Sladen Trust Expedition, from 7°08.00'S, 56°16.00'E, Coetivy Id. 1905, held at the Smithsonian
Institution, Washington, registration number USNM 41048 (note that there are two dry specimens in this lot. but
the larger specimen is the holotype).
Description. — Carapace wider than long, ratio of CW/CL approx. 1.25-1.30, broadly rounded in outline but
frontal and posterior margins truncated; surface convex, areolatc, granulate, and spinous. There are about twenty
five to thirty distinct areolae, more medial areolae smooth or minutely granulate, more lateral areolae adorned with
larger acute granules, and those above anterolateral margin are adorned with short, acute spines. Behind branchial
groove is a laterally directed region of granules, grading into spines towards last marginal tooth. Carapace surface
and pereopods covered with setae of two kinds: short setae, bent at right angles near tip, clothing surface, but
interspersed with longer filiform setae ( 6 x length of short setae and 0.20-0.25 x CW) which fringe limbs and arc
arranged in clumps on carapace where there arc about twenty five distinct tufts, each with up to seven setae, which
tend to be associated with areolae. The density of setae almost completely obscures body surface but most of this
is attributable to short setae which, in some places, are separated by narrow areas lacking setae. Structure of short
and long setae are different. In short setae proximal 40% of shaft is erect and lacks ornamentation, then a region
occupying about 55% where setae are bent at right angles and long, stout setules radiate from shaft only on
external side, forming a dense bunch, and finally distal 5% which is smooth, slightly curved, and narrows to an
acute tip. In long setae proximal 95% is covered with small setules which distally increase in density, but not in
size, and last 5% is smooth, slightly curved and narrows to an acute tip.
A narrow frontal carapace groove separates a pair of prominent rounded protuberances, and then divides into
separate grooves which diverge and then curve back medially. Between these grooves is an elongate granulate
ridge. Just in front of cardiac region two laterally-directed grooves originate: first groove (cervical) arises separately
from small pits and runs directly anterolateral on to branchial region and mid-way along their length they are
joined by grooves running back from frontal groove The second, shallower groove extends across mid-line and
initially runs almost directly towards lateral margin but then splits into an anterior portion which follows first
groove for a short distance, while second portion curves posterolaterally, bordering anterior cardiac region. In effect
groove crossing mid-line, connects two crescent-shaped grooves. A faint branchial groove is evident and posterior
cardiac area is defined. Anterolateral carapace margin begins at level of postorbital corner, is evenly convex and
bears four distinct, broad-based, equidistant teeth, each ending in a well developed, upwardly curved, acute spine;
first tooth directed anterolaterally and remainder directed laterally. Each anterolateral tooth has an associated tuft of
long setae. A posterolateral tooth, which is smaller than preceding anterolateral teeth, marks beginning of
convergent posterolateral border alongside which lies the reduced last leg. Posterior carapace margin is recessed in
order to accommodate first segment of abdomen which is visible dorsally.
Frontal margin continuous, V-shaped, ventrally-directed. joined to epistome (which separates orbits).
Supraorbital margin not projecting, continuous above orbits, interrupted by a distinct notch closer to postorbital
corner, lollowed by four or five acute spines; suborbital margin with three similar spines followed by an acute
tooth (visible dorsally when setae arc removed) and then descending to a much smaller tooth at its inner corner.
Orbits clearly exposed dorsally.
First article of antennule large, filling a large part of ventral orbital region; distal margin bearing a dense fringe
of longer setae, obliquely angled and not continuous with distal margin of second antennal article. Remainder of
antennule folded into orbit. First article of antenna moveable, wider than long, medially beaked; inferior tooth well
Source :
FAMILY DYNOMENIDAE
507
Fig. 23. — Hirsutodynomene spinosa (Rathbun, 1911): a-g, 6 16.4 x 14.3 mm, Glorieuses Ids (MNHN-B 6899):
a, dorsal view of right half of carapace; b, ventral view of right orbital area; c, outer face of right cheliped; d. dorsal
view of right cheliped; e. posterior view of terminal articles of right fourth pereopod; f, posterior view of terminal
articles of right Fifth pereopod; g, ventral view of telson and terminal segments of male abdomen.
Source : MNHN, Paris
508
C. L. McLAY
developed, blunt, superior tooth above opening of antennal gland is smaller. Second article wider than long, distal
margin widest, to which is fixed the exopod curving over base of eyestalk, becoming broader, terminating bluntly
and bearing longer setae. Third antennal article longer than wide, and attached to remaining distal border of second
article, slotting in behind exopod, and just matching length of exopod. Fourth antennal article smaller, as long as
wide; remaining antennal articles directed laterally, extending well beyond postorbital corner, and can be partially
folded under supra-orbital margin. Ratio of length of antennal flagella to CW = 0.45. Eyestalk can be completely
folded into orbit, and cornea is well developed, occupying all of tip. Epistome broadly triangular, surface deeply
concave; dorsal arm, joined to tip of carapace, very elongate and narrow; lateral arms shorter and thicker. Joint
between epistome and carapace is marked by a narrow suture.
Subhepatic area smooth, very convex. A groove begins near base of antenna, curving round under branchial
region and meeting lateral carapace margin just anterior to last tooth at beginning of posterolateral border. A short
cervical groove branches off and ascends towards first anterolateral tooth. Third maxillipeds operculiform,
bases widely separated by tip of sternum. Crista dentata has five or six well developed, distally placed teeth
on each side. Female sternal sutures 7/8 short, ending wide apart on low tubercles just behind bases of second
walking legs.
Branchial formula 19 gills + 7 epipods on each side as found in Dynomene hispidci. No epipod or podobranch
on last pereopod. In cross section gills have lateral margin deeply notched, dividing gill into a hypobranchial plate
(containing efferent vessel) and an epibranchial lobe. Between these marginal lobes are two pairs of lobes, first
similar and second much shorter than marginal lobes. Thus epibranchial surface shows six rows of blunt lobes,
decreasing in size medially, which are arranged above afferent blood vessel. These lobes and hypobranchial plate
are distally thickened, maintaining spaces between adjacent rows. Hypobranchial setae on posterior wall of gill
chamber poorly developed. Posterior margin of scaphognathite bears two long setae. Hypobranchial margin of
podobranchs bears same setae as on epipod.
Cheliped stout, slightly longer than first leg. Merus trigonal, inner face smooth and fitting closely against
pterygostomia! region of carapace, borders granulate, outer face has a subterminal broad, restriction which
separates a thickened distal ridge on which there are three large acute spines from a pair of similar spines preceded
by a row of three smaller acute granules on superior border. Inner inferior margin of merus has an acute lateral
spine. Outer face of carpus convex with six large, acute granules, two more prominent acute spines on distal
margin; inner superior border with a distomedially directed, but dorsally curved, sharp spur which abuts against
proximal inner surlace of propodus thereby restricting closure of cheliped against frontal area. In a similar way.
inferior carpal margin is produced as a smooth, obtuse, flange fitting against merus when limb is withdrawn.
These two structures give carpal article an unusual and distinctive shape: inner face very narrow and outer face
much broader. Outer and superior faces of propodus with about a dozen prominent granules which form acute
spines on superior face; inner and inferior faces smooth, except that there is a small proximal granule on inner
propodal face. Fixed finger almost straight with four or five proximal teeth increasing in size distally; moveable
finger strongly curved also with four or five teeth; first tooth large, blunt, mid-way along margin, rest increasing
in size distally where last two teeth interlock. Both fingers, thick, hollowed out internally, touching only at lips;
a group of stiff setae is inserted proximally on each finger and these curve towards tips. These setae fill gap
between the fingers and form a screen.
First three pairs of walking legs decreasing in length posteriorly. Mcri elongate, both faces of meri of first two
legs and anterior face third leg merus smooth and nacreous, inferior distal margin hollowed out to accommodate
carpal article. Superior border ot meri of these legs with three small spines, in a row, separated by a gap from two
larger distal spines, and three large distal spines on posterior margin, length of merus of second leg about 1.6 x its
width and equal to about half of CL. Dorsal surface of carpi bearing three longitudinal rows each of five acute
spines, and produced distally to overhang base of propodi. Dorsal surface of propodi with three similar rows each
of two or three spines. Dactyli curved, inferior margin armed with 4-5 small spines, tip brown (in some cases the
whole dactyl is black) and subacute.
Last pair of legs greatly reduced, lying along posterolateral border of carapace, reaching only as far as half way
along meral article of preceding limb, borders of articles unarmed. Last pair of legs subchelate, sexually
dimorphic: female with well developed distal extension of propodus which opposes dactyl, male with only weakly
Source : MNHN. Paris
FAMILY DYNOMENIDAE
509
developed propodal extension. Female propodal extension bearing eight, unequal, stout, hooked, acute, spines, the
four largest lined with tiny, striated flattened and conical teeth along almost their entire inner surface, the
remaining four more sparsely covered with teeth. Female dactyl as long as propodal extension, bearing sixteen
unequal, stout, hooked spines (arranged asymmetrically around perimeter of dactyl) whose concave inner surface is
wrinkled and mostly devoid of tiny teeth. Male propodal extension bearing five unequal curved spines the four
largest of which have lateral rows of about six tiny teeth. Male dactyl longer than propodal extension and ending
in a single acute claw.
All segments of abdomen freely moveable, surface smooth, margins unarmed but fringed with long setae.
Second segment narrowest, anterior margin sinuous, medial region convex, lateral margins produced as a flange
which fits over posterior margin of first segment (which is shortest) preventing forward slippage of abdomen.
Subsequent segments increasing in length and breadth distally. not overlapping with preceding segments. Telson
much wider than long, anterior margin angled to accommodate uropod, posterior margin broadly rounded. In
female uropod plates large, filling about two thirds of space between last abdominal segment and telson, excluding
most of last abdominal segment and telson from reaching lateral margin of abdomen. In male last abdominal
segment occupies about a half of length. No effective abdominal locking mechanism: abdomen only loosely held
against sternum in both sexes; sideways movement restricted by small sternal tubercle beside telson. In mature
female abdomen occupies all of ventral surface, covering coxae of all pereopods with telson covering proximal half
of third maxillipcds. In male abdomen not quite so broad and telson only extends as far as bases of third
maxillipeds.
Five pairs of pleopods in female, first pair vestigial, remainder biramous. First male pleopod a semi-rolled
tube with a small apical plate surrounded by long setae. Second male pleopod with an exopod on the basis, needle¬
like distally, armed with a series of sixteen tiny, straight, acute, inset spines and ending in two larger straight
spines. Subterminal spines unevenly spaced, ninth and tenth spines overlap, and follow a sinuous path. Third to
fifth male pleopods rudimentary and biramous, exopod longer and jointed to basal article.
COLOUR. — Preserved specimens have a dense covering of light brown setae and the dactyli (or only their tips)
of the first four pereopods are black or dark brown.
Geographic distribution. — In the Indian Ocean Hirsutodynomene spinosa is known from Madagascar.
Mauritius, Glorieuses Ids, Coctivy Id (the type locality, Western India). Chagos Archipelago. Cocos Keeling Ids,
and Western Australia. Records from Indonesia include Timor, Ternate, and Moluccas. In the Pacific this species is
known from Southeast Queensland, Middleton Reef, Elizabeth Reef, Palau, Marshall Ids, Vietnam, Mariana Ids,
Xisha Ids, Japan, Encwetak Atoll, Palmyra Id, Tuamotu, Raroia, Marquesas Ids. The material examined in
this study establishes new records for Australia, Middleton and Elizabeth Reefs (Tasman Sea), Mariana Ids, Japan,
and Palmyra Id. It should be noted that although Balss (1935) included this species in his report on the col¬
lections made by the Hamburg Museum Expedition to South Western Australia, 1905. the material cited did
not come from Western Australia but from the Marquesas and Palau Ids. H. spinosa is clearly a widespread
Indo-Pacific species.
DEPTH. — Hirsutodynomene spinosa has been collected from intertidal habitats and to a depth of approxima¬
tely 15 m. H. spinosa is an inhabitant of shallow water coral reefs.
Size. — The maximum size for males is 28.7 x 21.3 mm. for females 32.3 x 29.4 mm. Only one ovigerous
female (32.3 x 29.4 mm) has been recorded in December from Middleton Reef. Tasman Sea and it carried about
3800 eggs with a diameter of 0.5 mm. Although H. spinosa is one of the larger dynomenids, it evidently produces
eggs of similar size to the other species and probably has planktotrophic larvae.
DISCUSSION. — Hirsutodynomene spinosa (Rathbun, 191 1) was first described by RATHBUN on the basis of
three males collected from Coetivy Id by the Percy Sladen Trust Expedition of 1905. The first female was reported
by BALSS (1935) from the Marquesas Ids.
RATHBUN (1911) referred to the shorter setae of Hirsutodynomene spinosa as being "club-shaped" and the
bunches of setae as being long and slender. Microscopic examination shows that the distal portion of the short
510
C. L. McLAY
setae is bent at right angles and ornamented, on the outer side, with sharp sctules which might give the setae a
club-shaped appearance. The long setae are clothed for almost their entire length with much smaller setulcs.
The gills of Hirsutodynomene spinosa arc similar to those of Dynomene filholi. On the cpibranchial surface
there are sequential rows of six lobes along the length of the gill, except towards the tip where the smaller medial
lobes are lost. In prepared material the lobes tend to be clumped together, even interlocking, but in live animals
the lobes are probably free, kept apart by their thickened tips. As in the species of Dynomene, the hypobranchial
setae in the posterior region of the branchial chamber of H. spinosa are poorly developed. The posterior margin of
the scaphognathite bears two long setae and the hypobranchial margin of each podobranch carries cleaning setae as
found in H. Ursula.
The male pleopods of Hirsutodynomene spinosa are similar to those of the species of Dynomene. The tip of
the first plcopod bears a dense ring of long setae surrounding an oval apical plate. The second pleopod has a large
number of subterminal spines, not all evenly spaced, which follow a sinuous path along the shaft and it ends with
two straight terminal spines. Also the last three pairs of pleopods are biramous and the connection of the exopod
with the basal article is marked by a joint.
The two species in this genus have non-overlapping distributions with Hirsutodynomene spinosa being a
widespread Indo-West Pacific species and H. Ursula being restricted to the eastern Pacific. The main differences
between these species (see Table 2) are discussed below under H. Ursula.
The habitat of Hirsutodynomene spinosa seems to be dead coral branches and rubble. Morrison (1954)
recorded H. spinosa from the inshore, more pooled area of the leeward outer reef at Raroia, Tuamotu along with
hermit crabs, Cryptodromia canaliculata , Pachygrapsus plicatus. Micippoides angustifrons, Thalamita pieta.
Eriphia sebana and several xanthids. Peyrot-Clausade (1981 ) recorded ii from dead clumps of Acropora sp. on
Tulear Reef, Madagascar where (along with Dynomene hispida) it made up only about 1-2% of the anomuran and
brachyuran fauna. On both the Tulear and Reunion reefs H. spinosa was always less common than D. hispida
which had a greater depth range (Peyrot-Clausade, 1984). Although both these species are shallow water
dynomenids, H. spinosa grows to a much larger size.
Examination of the gut contents of a male Hirsutodynomene spinosa 14.2 x 10.8 mm from the Cocos Keeling
Ids revealed a stomach packed almost entirely with sand grains and a small amount of aggregated amorphous
organic material. There were no recognizable animal or plant fragments. Food is probably obtained by using the
spooned cheliped Fingers and their stiff setae to sift out fine particulate organic material from coral sediments.
Hirsutodynomene Ursula (Stimpson, 1860)
Figs 4 a-c, 6 a-b, 9 a-b, 1 1, 14 f, 17 f. 24 a-g
Dynomene Ursula Stimpson, 1860: 239. — A. Milne Edwards, 1879: 9, figs 16-19. — ALCOCK. 1901: 74 (list). —
Rathbun, 1937: 54, pi. 12, figs 1-4. — Schmitt. 1939: 25. — Garth, 1946: 349. pi. 61, figs 5-6; 1948: 16; 1961:
121 (list); 1965: 6; 1966: 5; 1991: 125. — Birkeland et ai, 1975: 67. — Takeda. 1977: 35 (list). — Prahl &
Alberico, 1986: 98 (list). — Prahl, 1986: 96. — Rodriguez de la Cruz, 1987: 113. — Villalobos-Hiriart et ai.
1989: 53 (list). — Correa-Sandoval, 1991: 2. — Lemaitre & Alvarez-Leon, 1992: 50. — Aguilera & Guzman,
1992: 4 (list). — Hendrickx, 1995: 127 (list); 1997: 29, fig. 39 a-c. — Vargas et ai, 1996: 99 (list).
MATERIAL EXAMINED. — Galapagos. Allan Hancock Galapagos Expedition: stn 30-33. Hood Id, Gardner Bay,
no depth. W. L. Schmitt coll., 26.01.1933, M. J. Rathbun id.: 1 6 11.7 x 9.6 mm; 1 $ 15.0 x 12.3 mm (USNM
68313). — Charles Id, no depth. W. L. Schmitt coll., 27.01.1933, M. J. Rathbun id.: 2 6 9.1 x 7.4 mm, 11. 1 x
8.6 mm (USNM 68314).
Mexico. Espiritu Santo Id. " Velero ": stn 638-37, San Gabriel Bay, shore, 7.03.1937: 1 6 13.4 x 10.3 mm;
I 9 13.0 x 10.3 mm (LACM). — Stn 1110-40, 2-4 m, 14.02.1940. J. Garth id.: 1 2 7.2 x 5.7 mm (LACM).
Panama. Secas Ids. Stn 252-34, on Porites coral. 22.02.1934, M. J. Rathbun id.: I 9 ovig. 10.9 x 88 mm
(LACM).
"Velero": stn 867-38, shore, 2.03.1938: 8 6 8.4 x 6.6 - 12.7 x 10.0 mm; 5 2 ovig. 9.5 x 7.5 - 12.3 x 9.8 mm
(LACM).
Ecuador. La Plata Id. "Askoy": stn 80. 12.04.1941. J. Garth id.: 3 6 7.5 x 6.3 - 19.4 x 15.7 mm; 1 2 5.0 x
4.4 mm; 1 2 ovig. 15.0 x 12.2 mm (LACM).
Argosy 34: no depth, J. Garth coll., 5.03.1963: 1 2 ovig. 19.4 x 14.7 mm (USNM 247230).
Source : MNHN . Pans
FAMILY DYN0MEN1DAE
51 1
Types. — Dynomene Ursula Stimpson, 1860: holotype is a male 15.2 x 12.7 mm, collected by
Mr J. XANTUS from 22°50.00’N, 109°55.00'W, Cape St. Lucas, Baja California, and according to the original
paper is held by the Smithsonian Institute, Washington, but RATHBUN (1937) stated that the specimen is not
extant. However there is a syntype held by the Museum of Comparative Zoology. Harvard, registration number
MCZ 1378.
Description. — Carapace wider than long, ratio of CW/CL approx. 1.25-1.30, broadly rounded in outline but
frontal and posterior margins truncated; surface convex, areolate, sparsely granulate, and spinous. There are about
twenty to twenty five distinct areolae, more medial areolae smooth or minutely granulate, more lateral areolae
adorned with larger acute granules, and those above anterolateral margin adorned with short, acute spines. Behind
branchial groove is a laterally directed region of granules ending at base of last marginal tooth. Carapace surface
and pereopods covered with setae of two kinds: short erect serrate setae clothing surface, but interspersed with
longer serrate setae (3 x length of short setae and 0.05-0.07 x CW) which also fringe limbs and arranged in clumps
on carapace where there are about twenty distinct tufts, each with up to five setae, which tend to be associated
with areolae. Density of setae does not completely obscure body surface. Structure of short and long setae
differs. In short setae the proximal 23% of shaft lacks ornamentation, followed by a region of 45%
where tiny selules arranged in bands, then a region occupying about 22% where long, stout setules radiate from
all sides of shaft, forming a dense bunch, and finally the distal 5% which is smooth, and narrows to
an acute tip. The whole seta may be slightly curved but is not bent at right angles. In long setae
the proximal 15% is smooth, following 80% covered with small setules which increase distally in density and size
(but not reaching size of selules on short setae), and the last 5% is smooth, slightly curved and narrows to
an acute tip.
A narrow Irontal carapace groove separates a pair of prominent rounded protuberances, and then divides into se¬
parate grooves which diverge and then curve back medially. Between these grooves is an elongate granulate ridge.
Just in front of cardiac region two laterally-directed grooves originate: first groove (cervical) arises separately from
small pits and runs directly anterolateral on to branchial region and mid-way along their length they are joined by
grooves running back from frontal groove. Second shallower groove extends across mid-line and initially runs al¬
most directly towards lateral margin but then splits into an anterior portion which follows the first groove for a
short distance, while the second portion curves posterolaterally, bordering anterior cardiac region. In effect groove
crossing mid-line, connects two crescent-shaped grooves. A faint branchial groove is evident and posterior cardiac
area is defined. Anterolateral carapace margin begins at level of postorbital corner, evenly convex and bears
four distinct, broad-based, equidistant well developed teeth, each ending bluntly; first tooth directed anteriorly,
second anterolaterally, and remainder directed laterally. Each anterolateral tooth has an associated tuft of long
setae. A posterolateral tooth marks beginning of convergent posterolateral border alongside which lies
the reduced last leg. Posterior carapace margin recessed in order to accommodate first segment of abdomen
visible dorsally.
Frontal margin continuous, V-shaped, ventrally-directed, joined to epistome (which separates orbits).
Supraorbital margin not projecting, continuous above orbits, interrupted by a distinct notch closer to postorbital
corner, followed by three small blunt granules before corner; suborbital margin with three larger blunt granules,
lirst two visible dorsally, second and third (tooth-like) closer together but separated by a deep notch with third
terminating the suborbital margin. Orbits clearly exposed dorsally.
hirst article of antennule large, filling a large part of ventral orbital region; distal margin bearing a dense fringe
of longer setae, obliquely angled and not continuous with distal margin of second antennal article. Remainder of
antennule folded into orbit behind second antenna. First article of antenna moveable, wider than long, medially
beaked; inferior tooth well developed; blunt, superior tooth above opening of antennal gland smaller. Second
article about as wide as long; distal margin widest, to which is fixed the exopod curving over base of eyestalk,
becoming broader, terminating bluntly and bearing longer setae. Third antennal article longer than wide, and
attached to remaining distal border of second article, slotting in behind exopod. Fourth antennal article smaller, as
long as wide, together with third article just matching length of exopod; remaining antennal articles directed
laterally, extending well beyond postorbital corner, and partially folded under supraorbital margin. Ratio of length
of antennal flagella to CW = 0.45. Eyestalk can be completely folded into orbit, and the cornea is well developed.
Source :
512
C. L. McLAY
1 mm a-d
1 mm e
1 mm f
1 cm g
Fig. 24. — Hirsutodynomene Ursula (Stimpson, 1860), 9 ovig. 19.4 x 14.7 mm. La Plata Island, Ecuador (USNM
247230): a, dorsal view of right half of carapace; b. ventral view of right orbital area; c. outer face of right cheliped;
d. dorsal view of right cheliped; e, posterior view of terminal articles of right fourth pereopod; f, posterior view of
terminal articles of right fifth pereopod; g, ventral view of telson and terminal segments of female abdomen.
Source : MNHN. Paris
FAMILY DYNOMENIDAE
513
occupying all of tip. Epistome broadly triangular, surface deeply concave; dorsal arm, joined to tip of carapace,
very elongate and narrow; lateral arms shorter and thicker. Joint between epistome and carapace marked by a
narrow suture.
Subhepatic area very convex with a few scattered granules. A groove (pleural suture) begins near base of
antenna, curving round under branchial region and meeting lateral carapace margin just anterior to last tooth at
beginning of posterolateral border. A short cervical groove branches off and ascends towards first anterolateral
tooth before dividing into two branches, one curving towards postorbital corner and the other curving beneath
second anterolateral tooth. Third maxillipeds operculiform, bases widely separated by tip of sternum. Crista dentata
has seven or eight well developed, distally placed teeth on each side. Female sternal sutures 7/8 short, ending wide
apart on low tubercles just behind bases of second walking legs.
Branchial formula 19 gills + 7 epipods on each side. There is no podobranch or epipod on last pereopod. Gill
structure is variable. In cross section the anterior of each pair of arthrobranchs show maximum development of
epibranchial lobes: there are a series of six elongate lobes developed above afferent vessel, two narrow, short
median lobes flanked by longer lobes, with wider and longer lobes on the outside. The short median lobes are lost
towards the tip of gill. At base of outer lobes there is a deep notch on each side dividing off the hypobranchial
plate containing the efferent vessel. On posterior arthrobranchs, and all pleurobranchs, there are only four
epibranchial lobes and the median lobes are lost towards tip of gill. Hypobranchial setae at posterior end of
branchial chamber poorly developed. Posterior margin of scaphognathite with two long setae. Hypobranchial
margin of podobranchs bear same setae as on epipod.
Cheliped stout, slightly longer than first leg. Merus trigonal; inner face smooth fitting closely against ptcry-
gostomial region of carapace, borders granulate; outer face has a subterminal broad, restriction which separates a
thickened distal ridge, on which there are three spines, from a pair of similar spines preceded by a row of six
smaller granules on superior border. Inner inferior margin of merus has a small blunt lateral spine. Outer face of
carpus convex with six small blunt granules, two more prominent blunt spines on distal margin; inner superior
border with a distomedially directed, but dorsally curved, blunt spur which abuts against proximal inner surface of
propodus thereby restricting closure of cheliped against frontal area. In a similar way. inferior carpal margin is pro¬
duced as a smooth, obtuse, flange fitting against merus when limb is withdrawn. These two structures give carpal
article an unusual and distinctive shape: inner face very narrow and outer face much broader. Outer and superior
faces of propodus with about a dozen small blunt granules, inner and inferior faces smooth, except that there is a
small proximal granule on inner propodal face. Fixed finger almost straight with four or five teeth increasing in
size distally, second tooth placed midway and prominent, last three teeth dividing up tip of finger; moveable Finger
strongly curved bearing four teeth, first blunt, second more acute directed towards tip and last two interlocking
with teeth of fixed finger. Both fingers, thick, hollowed out internally, touching only at lips; a group of stiff setae
inserted proximally on each finger and these curve towards tips filling gap between fingers.
First three pairs of walking legs decreasing in length posteriorly. Mcri elongate; both faces of meri of first two
legs and anterior face third leg merus smooth and nacreous; inferior distal margin hollowed out to accommodate
carpal article. Superior border of meri of these legs with a row of three or four small spines increasing in size
distally, separated by a gap from two small distal spines; no spines on distal posterior margin; length of merus of
second leg about 1.2 x its width and equal to about 0.4 x CL. Dorsal surface of carpi armed with seven or eight
small spines not arranged in rows, and produced distally to overhang base of propodi. Dorsal surface of propodi
armed with four or five small spines. Dactyli curved, inferior margin armed with 4-5 small spines, tip brown (in
some cases whole dactyl is black) and subacute.
Last pair of legs greatly reduced, lying along posterolateral border of carapace, reaching only as far as half way
along moral article of preceding limb; borders of articles unarmed. Last pair of legs subchelate, sexually
dimorphic: female with well developed distal extension of propodus which opposes dactyl, male with only weakly
developed propodal extension. Female propodal extension bearing ten, unequal, stout, hooked, acute, spines with
lateral rows of 8-10 tiny, flattened teeth proximally, separated by a striated ridge. Distally the striae of this ridge
grade into a surface that is densely convoluted. Female dactyl as long as propodal extension, bearing thirteen
unequal, stout, hooked spines (arranged asymmetrically around perimeter of dactyl) whose concave inner surface is
wrinkled or convoluted and mostly devoid of tiny teeth except for a few small blunt marginal teeth on distal
514
C. L. McLAY
two-thirds. Male propodal extension bearing six unequal curved spines which have tiny scattered marginal teeth.
These spines are not arranged in the typical manner for dynomenids: instead of lying in an asymmetrical row
around the perimeter, they are very disorganized, facing in different directions and leaning at different angles. Male
dactyl longer than propodal extension and ending in a single acute claw.
All segments of abdomen freely moveable, surface smooth, margins unarmed but fringed with long setae.
Second segment narrowest; anterior margin sinuous, medial region convex, lateral margins produced as a flange
which fits over posterior margin of first segment (which is shortest) preventing forward slippage of abdomen.
Subsequent segments increasing in length and breadth distally, not overlapping with preceding segments. Telson
much wider than long, anterior margin angled to accommodate uropod, posterior margin broadly rounded. In
female uropod plates are large, filling about ninety percent of space between last abdominal segment and telson.
excluding most of last abdominal segment and telson from reaching lateral margin of abdomen. In male last
abdominal segment occupies about three quarters of peripheral margin. No effective abdominal locking
mechanism: abdomen only loosely held against sternum in both sexes, sideways movement restricted by small
sternal tubercle beside telson or uropods. In mature females sternal tubercles absent; abdomen occupies all of
ventral surface, covering coxae of all pereopods with telson covering proximal half of third maxillipeds. Male
abdomen not quite so broad and telson only extends as far as bases of third maxillipeds.
Five pairs of pleopods in female, first pair vestigial, remainder biramous. First male pleopod a semi-rolled
lube with a small apical plate surrounded by long setae. Second male pleopod with an exopod on basis, needle-like
distally, armed w'ith a series of twenty prominent, straight, acute, spines and ending in two larger curved spines.
Subterminal spines unevenly spaced, seventh, eighth, and ninth spines form an overlapping group as do thirteenth
and fourteenth spines, and follow a sinuous path. Third to fifth male pleopods rudimentary and biramous, exopod
longer and jointed with basal article.
COLOUR. — In his original description Stimpson (1860) noted that the color of Hirsutodynomene nrsula is
more or less reddish or crimson, the setae a light golden color, and the ambulatory legs have sharp, black, curved
unguiles (claws). A. Milne Edwards (1879) gave the colour as being crimson, and in places carmine. Rathbun
(1937) noted that the dactyli of the last pair of legs are without pigment.
Geographic Distribution. — The type locality for Hirsutodynomene insula is Cape San Lucas, Baja
California. Rathbun (1937) states that the type specimen is not extant. The range extends from the Gulf of
California in the north to La Plata Id, Ecuador in the south, and offshore it occurs on the Galapagos Ids and
Clipperton Id. Thus H. ursula is an eastern Pacific species.
DEPTH. — The depth range for Hirsutodynomene ursula is 0-99 m (rarely 125 m). Most specimens have been
collected from rocks and dead coral, and some have been recorded as coming from Porites sp. and Pocillopora sp.
but it is not clear whether these were living or dead colonies. There is no strong evidence of dependence on living
corals. H. ursula has a much greater depth range than H. spinosa which has not been recorded deeper than 1 5 m.
Size. — The maximum sizes for Hirsutodynomene ursula are 27.2 x 20.4 mm for males, and 21.8 x 16.5 mm
for females. The smallest ovigerous female examined was 9.5 x 7.5 mm, carrying around 120 eggs, and the largest
ovigerous female, 19.4 x 14.7 mm, carrying around 2800 eggs. Mean egg diameter was 0.45 mm. This egg size
suggests that this species has planktotrophic larvae. Ovigerous females have been mainly collected during
December to April with one female from Clipperton Id collected in August (Garth, 1965). Concentration of
breeding in the early part of the calendar year is similar to the Dynomene species.
DISCUSSION. — GARTH (1946) remarked upon the remarkable similarity of Hirsutodynomene ursula to the
xanthids: "With no other Galapagos species of Brachyura is the field collector so likely to be misled as to identity
as with D. ursula. Unless he notices that the fourth pair of walking legs are reduced to minute size and carried
dorsally, he will believe himself to have found a species of Pilumnus ." Indeed, initial sorting of the Galapagos
collections resulted in all the dynomenid specimens being placed with the Xanthidae. An untrained observer would
be easily misled by the black pereopod dactyli. (It should be noted here that dynomenids do not in fact carry their
last pereopods "dorsally". They are carried horizontally).
Source : MNHN , Paris
FAMILY DYNOMENIDAE
515
The main differences between the two species in this genus (see Table 2) are that in Hirsutodynomene spinosa
there are about 12 spines (and associated areolae) on the carapace (only about 6 in H. Ursula ), the suborbital
margin has 5 short acute spines (only blunt granules), the carpal projection on the cheliped is a sharp spine (a
blunt lobe), the carapace is densely covered with setae (sparsely covered), and the short setae are bent at right
angles near the tip (short setae not bent).
Other differences between Hirsutodynomene spinosa and H. Ursula are seen in the last pair of legs: both have
a similar number of propodal spines in the female but those of H. spinosa are densely covered with teeth while in
H. ursula the teeth are confined to the proximal lateral margins and the rest of the spine has a roughened,
convoluted surface. In both species these spines are opposed by concave, wrinkled dactylar spines with a few small
teeth but H. spinosa has more spines. The males have a similar number of propodal spines but in H. ursula these
spines are very disorganized and arranged irregularly around the distal perimeter. There arc no differences in the
male dactyli. Commenting about the last pair of legs of H. ursula , STIMPSON (1860) said they are "... not
prehensile, since the animal does not cover itself with a foreign body like the Dromiae; and they fill, apparently
no office in the economy of the animal, except when in place, they fill up neatly the chink between the carapax
and the stouter walking feet." This is the earliest speculation about the role of the last leg of a dynomenid.
Stimpson clearly believed that this limb is vestigial and redundant.
Rathbun (1937) describes the two kinds of setae on the surface of Hirsutodynomene ursula: "...the first kind
very short, clavate, or even pedicellate, and densely crowded; the second long and nearly as thick as the first, but
fusiform, with pointed extremities, and sparsely distributed over the surface, generally in groups of three or four,
of unequal lengths." The short erect setae are in fact plumose with a dense distal zone of long setules and most of
the shaft only sparsely covered with short setules. The long setae are mostly covered with short setules. These
long setae are very similar in H. ursula and H. spinosa but the short setae show several differences in their shape
and setule ornamentation.
The gill structure of Hirsutodynomene ursula is similar to that of H. spinosa but shows more variation
between gills. The anterior arthrobranchs of H. ursula are the same, but the other large gills show a reduced
number of median lobes. These are more phyllobranchiate-like because more of the gill consists of flattened
plates. The gills of H. spinosa are more trichobranchiale-like in having a large number of elongate epibranchial
lobes. Both species show the pattern seen in other dynomenids of a tendency to lose epibranchial lobes towards the
tip of the gills. Hypobranchial setae at the back of the gill chamber are poorly developed and there are two long
setae on the posterior margin of the scaphognathite. Setae on the hypobranchial margin of each podobranch aid the
cleaning role of the epipods.
The second male pleopod of Hirsutodynomene ursula has a larger number of subterminal spines than
H. spinosa and these spines are longer, and not inset into the surface of the pleopod. In both species these spines
are unevenly distributed along the pleopod, but more so in H. ursula . where there arc two overlapping groups.
Both species have the last three pairs of pleopods biramous and rudimentary but in H. ursula the articles are of
similar length, whereas in H. spinosa the exopod is longer. The junction of the exopod and basal article is marked
by a joint in both species.
The two species have mutually exclusive distributions: Hirsutodynomene ursula in the Eastern Pacific and
along the coast of the Americas, and H. spinosa across the Western Pacific and the Indian Ocean. The western¬
most locality for H. ursula is Clipperton Id (109°W), while the eastern-most locality for H. spinosa is the
Tuamotu Ids(142°W).
The stomach of a male Hirsutodynomene ursula , 13.4 x 10.3 mm, from Espiritu Santo Id, was almost en
tirely filled with sand grains and some amorphous aggregates of organic material. There were no recognizable
plant or animal fragments present. Food is probably obtained by using the spooned cheliped fingers and their
stiff setae to sift out fine particulate organic material from coral sediments. The contents are identical to
those found in H . spinosa (see above). In the body cavity, beside the stomach, two encysted nematodes were
observed.
On the Galapagos Ids, Garth (1991) found that in a sample (n = 169) of 20 species of rocky shore Brachyura
from Sullivan Bay, Isla Santiago, Hirsutodynomene ursula was common, making up 5.3% of the total. This
species is the only dynomenid known from the Galapagos Ids.
516
C. L. McLAY
Table 2. — Comparison of the two species of Hirsutodynomene.
Genus METADYNOMENE nov.
DIAGNOSIS. — Carapace as wide as long or slightly wider than long, moderately convex, subcircular; surface
smooth. Tomentum composed of uniformly short, soft setae, which accentuate unevenness of carapace forming
transverse troughs. Lateral carapace margin well defined and marked by indentations or armed with distinct teeth.
Frontal groove well marked, split in two posteriorly; cervical, postcervical and branchial grooves usually evident.
Frontal carapace margin broadly triangular, continuous, no rostrum or teeth Eyestalks short; eyes protected by
well defined orbits. Sternal sutures 7/8 of female end well apart on low tubercles behind bases of second walking
legs.
Antennule can be concealed inside orbit at base of cyestalk. Antennal flagella shorter than carapace width. All
articles of antenna moveable; first article (urinal) always beaked medially; second article has an exopod firmly
fixed. Third maxillipeds opercular, completely covering buccal cavern, separated at their bases by a plate at same
level as sternum; basis and ischium of endopod fused but joint always marked by a shallow groove. Crista dentata
present. Chelipeds equal, stouter than walking legs; fingers not strongly curved and touching for at least half their
length. Last pair of legs sexually dimorphic, very reduced; dactyl rudimentary, forming an obsolete subchelate
mechanism with an extension of propodus. Gill structure basically phyllobranchiate but plates have a variable
number of epibranchial lobes.
Abdomen of six segments and telson folded loosely under thorax; uropods large, no effective abdominal
locking mechanism. Sideways movement restricted by small spines or ridges on coxae of second and third
pereopods, adjacent to the margins of telson and penultimate abdominal segments. Both sexes have five pairs of
pleopods; first pair vestigial in female: last three pairs rudimentary in male. Male pleopods uniform in structure;
first pair consist of a stout, setose semi-rolled tube with an apical plate; second pair needle-like, bearing a row of
curved spines on anterior surface.
Type Species. — Dynomene devaneyi Takeda, 1977.
Other Species. — Dynomene tanensis Yokoya. 1933, Metadynoniene crosnieri sp. nov.
ETYMOLOGY. — Metadynoniene is a combination of meta, meaning after , and the genus Dynomene . Gender is
feminine.
DISCUSSION. — This new genus is erected for a group of three species, two of which were originally assigned
to Dynomene. The third species is newly described herein. All of these species are characterized by having a
Source : MNHN. Pans
FAMILY DYNOMENIDAE
517
carapace about as wide as long, or only slightly wider than long, densely covered with short, soft setae which give
the surface an uneven, undulating appearance with transverse troughs, and chelipeds with fingers not strongly
down-curved and touching for about half their length. All the known species are substantially larger than the other
dynomenids and occur in deeper waters. Metadynomene crosnieri sp. nov. is known only from the type locality,
M. devaneyi (Takeda, 1977) from the type locality and Marquesas Islands, whereas the third species, M. tanensis
(Yokoya, 1933), is widespread in the Pacific.
Fig. 25. — a, Metadynomene devaneyi (Takeda, 1977) 6 21.2 x 20.1 mm, holotype, S. E. Oahu. Hawaii. " Star II",
367 m (BPBM-S 8509): dorsal view of whole crab, left second pereopod and right fourth pereopod not shown. —
b. Metadynomene tanensis (Yokoya, 1933). 9 13.5 x 12.5 mm. New Caledonia, Smib 8, 305-355 m: dorsal view of
whole crab. — c , Metadynomene crosnieri sp. nov., 6 23.2 x 22.7 mm, Glorieuses Ids, BENTHEDI, 330-440 m
(MNHN-B 22510): dorsal view of whole crab, setae removed from right half of carapace, right cheliped, left third
pereopod and left fifth pereopod are missing. — d. Paradynomene tuberculata Sakai. 1963, 6 21.5 x 22.3 mm,
Loyalty Ids, MUSORSTOM 6, stn DW 406, 373 m: dorsal view of whole crab.
Metadynomene devaneyi (Takeda, 1977)
Figs 11, 25 a, 26 a-c
Dynomene devaneyi Takeda, 1977: 31, figs 1-3; 1978, fig. 1. — McLay, 1991: 465, fig. 4b.
Not Dynomene devaneyi - GuiNOT, 1993: 1227 (= Metadynomene crosnieri sp. nov.).
MATERIAL EXAMINED. — Hawaii. "Star II Submersible ": S.E Oahu, off Makapuu Point. Kaiwi Channel, 21°18'N.
157°39'W, 367 m, 28.02.1974, collected from precious coral ( Corallium sp.) beds: 1 6 21.2 x 20.1 mm. holotype
(BPBM-S 8509); 1 9 15.6 x 15.2 mm. allotype (BPBM-S 8510).
Source :
518
C. L. McLAY
Marquesas Islands. Musorstom 9: stn CP 1290, Ua Huka, 8°53’S, 139°38'W, 341-344 m, 8.09.1997: I 9 18.4 x
18.3 mm. — Stn CP 1306, Nuka Hiva, 8°55.2'S, 140°14'W, 283-448 m, 10.09.1997: 1 6 29.0 x 28.3 mm; 1 $ ovig.
25.0 x 25.3 mm.
Types. — Dynomene devaneyi Takeda, 1977: holotype is a male 21.2 x 20.1 mm. collected by the
submersible "Star 1 1" from 21°18'N, 157°39’W, Southeastern Oahu, off Makapuu Point. Kaiwi Channel, Hawaii,
367 m, 28.02.1974, held by the Bernice P. Bishop Museum. Honolulu, registration number BPBM-S 8509. The
same institution holds a paratype male 22.7 x 21.6 mm (BPBM 1975.77. lot no. 2), and an allotype female
15.6 x 15.2 mm (BPBM-S 8510). A paratype male 21.0 x 20.0 mm, is held by the National Science Museum,
Tokyo, registration number NSMT Cr. 5075 (ex BPBM 1974.04, lot no. 4).
Description. — Carapace slightly wider than long, ratio of CW/CL = 1.03-1.05, rectangular in outline;
surface mostly flat, convexity greater in anterior-posterior direction than laterally, smooth, except for deeply
marked grooves (see below). Carapace surface densely covered with setae of only one kind: very short, soft setae,
clothing entire surface. Percopods covered with short setae as well as a few longer filiform setae (5 x length of
short setae and 0.10 x CW) which fringe limbs, locally concentrated on dactyli. Density of short setae completely
obscures body surface and on carapace they present a symmetrical undulating aspect, reflecting gentle undulations
in carapace surface: one oblique trough lies behind supraorbital margin, with a short median longitudinal trough
extending posteriorly, then a trough curving anterolaterally which marks cervical groove, followed by a trough
running across midline, just in front of cardiac area, which splits into two lateral troughs, and finally a short
trough crossing cardiac area. Microscopic details of setae not investigated.
A shallow frontal carapace groove separates a pair of low rounded protuberances. Just in front of cardiac region
two laterally-directed grooves originate: first groove (cervical) arises separately (but very close together) from
small gastric pits curving (slightly sinuously) anterolaterally on to branchial region towards a faint notch mid-way
along anterolateral margin, while second groove extends across mid-line and initially runs almost directly towards
lateral margin but then splits into an anterior portion which follows posterior to the first groove but does not
reach lateral carapace margin, while second portion curves posterolaterally, bordering anterior cardiac region, to
become branchial groove, meeting margin at posterolateral tooth. Posterior cardiac area marked by a shallow
groove crossing mid-line. Anterolateral carapace margin begins at level of postorbital corner, evenly convex,
without teeth, and interrupted by a shallow notch where cervical groove meets the edge, midway towards branchial
notch which marks beginning of posterolateral border. Reduced last leg lies alongside posterolateral border which
angles obliquely towards rear of carapace. Posterior carapace margin recessed in order to accommodate distal section
of first segment of abdomen which is visible dorsally.
Frontal margin continuous, V-shaped, ventrally-directed, joined to epistome (which separates orbits).
Supraorbital margin not projecting, continuous above orbits, not interrupted by a notch, and without granules.
Suborbital margin, convex, without teeth, projecting, shelf-like and visible dorsally. Orbits oblique and clearly
exposed dorsally.
First article of antennule large, filling a large part of ventral orbital region, distal margin obliquely angled and
not continuous with distal margin of second antennal article. Remainder of antennule folded into orbit. First article
of antenna moveable, wider than long, medially beaked; inferior tooth well developed, blunt; superior tooth, above
the opening of antennal gland, smaller but more acute and directed ventrally. Second article wider than long, distal
margin widest, to which is fixed the exopod curving over base of eyestalk, becoming broader and terminating
bluntly. Third antennal article is longer than wide, and attached to remaining distal border of second article,
slotting in behind exopod, and just matching length of exopod. Fourth antennal article smaller, as long as wide;
remainder of the antennal articles directed laterally, extending well beyond postorbital corner, and can be partially
folded under the supra-orbital margin. Ratio of length of antennal flagella to CW = 0.33. Eyestalk can be
completely folded into orbit; cornea well developed, occupying all of tip. Epistome broadly triangular, surface
slightly concave; dorsal arm, joined to tip of carapace, narrow; lateral arms longer and thicker. Joint between
epistome and carapace is marked by a narrow suture.
Subhepatic area smooth, very convex. A groove begins near base of antenna, curving round under branchial
region and meets branchial groove at posterolateral notch. A short continuation of cervical groove bends anteriorly
Source : MNHN. Paris -
FAMILY DYNOMENIDAE
519
Fig. 26. — Metadynomene devaneyi (Takeda, 1977): a-c, 9 15.6 x 15.2 mm, allotype, S. E. Oahu, Hawaii, "Star //".
367 m (BPBM-S 8510): a, dorsal view of right half of carapace; b, outer face of right cheliped; c, dorsal view of
right cheliped.
to meet groove at margin of branchial area. Third maxillipcds operculiform, bases widely separated by tip of
sternum. Crista dentata has twelve teeth increasing in size distally. Female sternal grooves short, ending wide
apart in a V-shaped groove created by a low medial parallel ridge, just behind bases of second walking legs. Sternal
sutures 7/8 concealed by a dense layer of long soft setae from adjacent coxa of third walking leg.
Branchial formula same as M. tanensis , except than there is no podobranch on last pereopod. A cross section
of an arthrobranch shows lateral margin deeply notched, dividing gill into a hypobranchial plate (containing
efferent vessel) and an epibranchial lobe. Between these marginal lobes are a pair of shorter lobes. Thus the
cpibranchial surface shows four rows of blunt lobes, which are arranged above afferent blood vessel. Towards tip
of gill the length and number of lobes is gradually reduced. Hypobranchial setae in posterior region of branchial
chamber well developed.
Cheliped stout, much longer and stouter than first leg. Mcrus trigonal, inner face smooth and fitting closely
against pterygostomial region of carapace; borders granulate, superior border has a subterminal broad, restriction
which separates a thickened, smooth distal ridge, from a row of three small granules; inferior face has three sub-
distal, blunt tubercles. Outer face of carpus convex, smooth, no longitudinal channel, two prominent blunt
tubercles on distal margin; inner superior border with three blunt tubercles, distal one largest which abuts against
the proximal inner surface of propodus thereby restricting closure of cheliped against frontal area. In a similar way,
the inferior carpal margin is produced as a smooth obtuse flange fitting against mcrus when limb is withdrawn.
Outer face of propodus smooth, superior face with three small granules, inner and inferior faces smooth, except
that there is a small proximal tubercle on inner propodal face. Fixed finger almost straight with seven or eight
almost obsolete teeth increasing in size distally; moveable finger not strongly curved, with one small proximal
tooth and three teeth at tip, interlocking with opposing teeth. A narrowing band of setae extend on to outer face of
520
C. L. McLAY
moveable finger. Just below proximal teeth of fixed finger are two small pits in which several long setae are
inserted. Both fingers, thick, hollowed out internally, with a small proximal gape, touching for about three-
quarters their length. In hollowed out interior of each finger there are small tufts of setae which come together
when fingers arc closed.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of meri of first two
legs and anterior face third leg merus smooth and nacreous, inferior distal margin hollowed out. Superior border of
meri of these legs with several small granules, length of merus of second leg about 2 x width and equal to about
half of CL. Anterior and posterior dorsal margins of carpi without granules, and produced distally to overhang the
base of propodi. Surface of propodi smooth. Dactyli curved, inferior margin armed with 2-3 small distal spines,
tip pale brown and subacute.
Last pair of legs greatly reduced, lying along posterolateral border of carapace, reaching to about two-thirds of
meral article of preceding pereopod; borders of articles unarmed. Legs subchelate, sexually dimorphic: female with
well developed distal extension of propodus which opposes dactyl, male with only weakly developed propodal
extension. Female propodal extension bearing several, small, stout, spines. Female dactyl as long as propodal
extension, bearing several, small, stout, spines. Male propodal extension bearing five unequal curved spines. Male
dactyl longer than propodal extension and ending in a single acute claw. Microscopic details of propodus and dactyl
unknown.
All segments of abdomen freely moveable. Telson much wider than long, anterior margin essentially straight,
posterior margin broadly rounded. In both sexes uropod plates arc large, filling all of space between penultimate
abdominal segment and telson, excluding all of last abdominal segment from reaching lateral margin of abdomen.
No effective abdominal locking mechanism: abdomen only loosely held against sternum in both sexes. Sideways
movement restricted by small spines (can be bifid) on coxae of second and third pereopods. adjacent to the margins
of the telson and penultimate abdominal segments. In mature female abdomen occupies all of ventral surface,
covering coxae of all pereopods with telson covering proximal third of third maxillipeds. In male abdomen not
quite so broad and telson only extends as far as bases of third maxillipeds.
Five pairs of pleopods in female, first pair vestigial and uniramous. remainder biramous. Five pairs of
pleopods in male; first pleopod a semi-rolled tube ending in a curved apical plate surrounded by long setae; second
pleopod needle-like with an exopod on basis (as illustrated by Takeda. 1977, fig. 1A-C). Microscopic details of
second pleopod unavailable. Remaining pleopods rudimentary. Third pleopod comparatively well developed, fourth
and fifth pleopods smaller, all are biramous.
COLOUR. — Covered in a pale tan velvet tomentum.
Geographic Distribution. — Metadynomene devaneyi is known from Oahu Id. Hawaii (type locality)
where it was collected from precious coral ( Corallium sp.) beds, and the Marquesas Islands (new record).
Depth. — The depth off southeast Oahu was 367 m, and at the Marquesas Islands 283-448 m and 341-344 m.
SIZE. — Seven specimens of Metadynomene devaneyi (4 males and 3 females) are known: maximum size for
males is 29.0 x 27.3 mm and for females is 25.0 x 25.3 mm. An ovigerous female has been collected from the
Marquesas in September. It carried approximately 2200 eggs, diameter 0.7 mm, a smaller number than comparable
M. tanensis females.
DISCUSSION. — Some details can be added to the original description. Takeda (1977, fig. 1 A-C) figured the
first two male pleopods ot Metadynomene devaneyi but did not show any details, especially of the second
pleopods, which are only visible under high magnification. The male type specimen has three pairs
rudimentary biramous pleopods. The dactyli of the walking legs are armed with two or three small distal spines.
The major differences between the three species of this genus are discussed below (see Table 3) under M. crosnieri
sp. nov.
The branchial formula of Metadynomene devaneyi (19 gills + 7 epipods) is very similar to M. tanensis except
that there is no podobranch on the last pereopod. McLay (1991, fig. 4b, as Dynomene devaneyi) described the
structure of the gills of M. devaneyi showing several asymmetrical epibranchial lobes. The number of lobes is
Source : MNHN. Paris
FAMILY DYN0MEN1DAE
521
dependent upon where, along the gill, the cross section is made. The epibranchial lobes and hypobranchial plates
do not always correspond exactly. Whereas the plates are clearly arranged in sequence, the lobes tend to arise
independently so that in one section there may be as many as five lobes, and in another section there may be only
three. Furthermore, the number of lobes decreases towards the ends of the gill. In a scanning electron microscope
picture the lobes are sometimes clumped and it is difficult to see the exact relationship between the lobes and the
hypobranchial plates. However, the multi-lobed gill structure is similar to that found in the species of
Hirsutodynomene (see above). Hypobranchial setae at the back of the gill chamber are much better developed in
M. devaneyi.
McLay (1991) compared some features of Metadynomene devaneyi with those of the primitive dromiid
Sphaerodromia, contrasting the shape of the front of the carapace, the structure of the antenna, female sternal
sutures 7/8, and male pleopods - all of which are very similar. However, M. devaneyi has a large number of
pereopodal epipods and podobranchs, the gills are multi-lobed, trichobranchiate-like. rather than phyllobranchiatc.
In Sphaerodromia the abdomen is held in place by denticulate ridges on coxae of first two walking legs against the
lateral margins, while in M. devaneyi there is no effective abdominal locking mechanism.
The gut of a female, 18.4 x 18.3 mm, from the Marquesas Islands contained amorphous organic material and
sand grains.
Metadynomene tanensis (Yokoya, 1933)
Figs 4 d, 6 c, 7 f, 9 d-e, 11,13 c, e-f, 14 e, 25 b. 27, 28
Dynomene tanensis Yokoya, 1933: 96, text-fig. 38. — Sakai, 1936: 45; 1940: 54 (list). — Serene, 1968: 36 (list). —
Takeda, 1977: 35 (list). — Poupin, 1996b: 26, pi. 12f (colour photo).
Dynomene praedator - Sakai. 1976: 30, text-fig. 17. — Naga & Tsuchida, 1995: 108, pi. 1, fig. 2. Not Dynomene
praedator A. Milne Edwards, 1879.
MATERIAL EXAMINED. — Taiwan. Nan-Fan-Auo Fishing Harbour, from a boat, no locality, no depth, no date,
coll. J. F. Huang : 1 6 9.0 x 8.0 mm. (only colour photo seen) (NTOU).
Indonesia. Danish Expedition Kei Islands: stn 12, 5°3rS, 132°35’E. 325 m. coll. T. Mortensen, 9.04.1922: I 9
5.5 x 5.0 mm (ZMUC).
Karubar: stn DW 18. 5°18'S, 133°OrE, 205-212 m. 24.10.1991: I 6 9.0 x 8.2 mm; 1 9 10.9 x 10.0 mm.
New Caledonia. Dragage du " Vauban ": stn D 15, 22°49’S, 167°12'E, 390-395 m, 10.04.1978: 1 9 6.4 x 6.0 mm.
— Stn D 24, 22°48’S, 167°12’E, 355-360 m, 13.04.1978: 3 9 11.2 x 10.5 - 13.4 x 12.6 mm.
Biocal: stn DW 50. 23°06.50’S, 167°53.74'E, 240-260 m. 31.08.1985: 1 9 22.8 x 20.5 mm.
Musorstom 4: stn 193, 18°56.30’S, I63°23.20’E, 415 m. 19.09.1985: 1 9 15.9 x 14.6 mm. — Stn 212,
22°47.40'S, 1 67° 10.50'E, 375-380 m, 28.09.1985: 1 6 7.2 x 6.8 mm; 1 9 13.0 x 12.1 mm. — Stn 213, 22°51.30’S,
167°1 2.00’E, 405-430 m, 28.09.1985: 1 6 18.9 x 18.1 mm. — Stn 214, 22°53.80'S, 167°13.90'E, 425-440 m,
28.09.1985: 1 6 13.6 x 12.6 mm; 1 9 13.0 x 12.2 mm. — Sin 215, 22°57.70’S, 167°17.00'E, 485-520 m,
28.09.1985: 3 6 7.2 x 7.0 - 16.9 x 16.0 mm; 4 9 14.4 x 13.9 - 19.2 x 18.9 mm. — Stn 230. 22°52.50’S. 167°1 1.80'E.
390-420 m, 30.09.1985: 1 8 22.8 x 21.6 mm; 1 9 15.8 x 14.6 mm.
Smib 2: stn DW 1, 22°52.7'S, 167°12.6'E, 438-444 m, 17.09.1986: 2 8 13.5 x 12.5. 17.8 x 16.6 mm. —
Stn DW 3. 22°56.0‘S, 167°18.8'E. 412-428 m, 17.09.1986: 1 8 13.1 x 12.3 mm. — Stn DW 5, 22°56.3'S, I67°14.4'E.
398-410 m, 17.09.1986: 2 9 11.0 x 10.4, 13.1 x 12.6 mm. — Stn DW 6, 22°56.2,S. 167°15.9’E. 442-460 m.
17.09.1986: I 9 16.7 x 15.2 mm. — Stn DW 16, 22°51.2'S, 167°11.7'E, 390 m, 19.09.1986: 1 9 19.6 x 17.5 mm. —
Stn DW 17. 22°55.1'S. 167°14.5'E. 428-448 m, 19.09.1986: 2 8 12.9 x 12.0. 18.4 x 17.5 mm; 1 9 13.7 x 12.6 mm.
Smib 3: stn DW 25, 22°56.1’S, 167°16.2'E, 437 m, 24.05.1987: 1 8 16.5 x 15.8 mm. — Stn DW 27. 22°55.2'S.
167°16.2'E. 457 m, 24.05.1987: I 9 ovig. 17.6 x 16.3 mm.
SMIB 4: stn DW 65, 22°55.3’S, 167°14.5’E. 420 m, 10.03.1989: 1 9 15.6 x 14.4 mm.
Smib 6: stn DW 125, 18°57.4'S, 163°23.5’E, 335-350 m, 3.03.1990: 1 8 19.8 x 18.0 mm.
Beryx II: stn CP 46, 23°42'S. 168°0rE, 300-350 m, 20.10.1992: 2 9 11.2 x 10.0, 20.0 x 18.0 mm.
Smib 8: stn DW 163, 24°49'S, 168°09'E, 310-460 m. 28.01.1993: 1 8 11.9 x 11.0 mm. — Stn DW 185. 23°I6'S.
168°04'E, 311-355 m, 31.01.1993: 3 9 4.8 x 4.8 - 16.0 x 15.4 mm. — Stn DW 190. 23°18’S, 168°05'E. 305-310 m.
31.01.1993: 1 9 16.2 x 14.8 mm. — Stn DW 193, 22°59'S. 168°19'E. 500-508 m. 1.02.1993: I 8 7.4 x 6.8 mm. —
Stn DW 197, 22°51'S, 168°12'E. 414-436 m, 1.02.1993: 1 8 22.1 x 21.2 mm; 4 9 8.6. x 7.5 - 19.0 x 18.4 mm. —
Stn DW 198, 22°52'S, 168°12'E, 414-430 m, 1.02.1993: 2 9 5.0 x 4.8, 16.2 x 15.3 mm. — Stn DW 199. 22°52’S.
168°12‘E, 408-410 m, 1.02.1993: 2 9 11.3 x 10.6, 21.0 x 19.8 mm.
Bathus 2: stn CP 736, 23°03'S, 166°58’E, 452-464 m, 13.05.1993: 1 9 ovig. 16.3 x 15.4 mm.
Source
522
C. L. McLAY
Bathus 3: stn CP 805, 33°41'S, 168°01'E, 278-310 m, 27.11.1993: 1 6 23.7 x 21.4 mm. — Stn CP 811. 23°41’S.
168°15'E, 383-408 m, 28.11.1993: 1 9 24.8 x 22.4 mm. — Stn DW 829, 23°21'S, 168°02'E, 386-390 m, 29.1 1.1993:
I 6 17.4 x 16.1 mm. — Stn DW 830, 23°20’S, 168°0I’E, 361-365 m, 29.11.1993: I 6 8.7 x 8.2 mm, 7 9 4.6 x 4.5 -
17.3 x 16.3 mm.
Halical 1: stn DW 01, 18°56’S, 163°24'E. 380-400 m, 23.11.1994: 1 6 12.9 x 12.0 mm. — Stn DW 04, 18°55'S.
163°24'E, 350-365 m, 26.11.1994: 1 9 14.5 x 13.7 mm.
Loyalty Islands. Musorstom 6: stn DW 459, 21°01.39’S, 167°31.45’E, 420 m, 20.02.1989: 1 9 16.8 x
15.4 mm. — Stn 460, 21°01.72’S, 167°31.45’E, 420 m. 20.02.1989: 1 6 16.4 x 15.2 mm. — Stn 464, 21°02.30'S,
167°31.60'E, 430 m, 21.02.1989: 2 9 15.5 x 14.6, 16.9 x 15.8 mm.
Chesterfield Islands. Chalcal 1: stn CP 8, 19°43.80'S, 158°35.25'E. 348 m. 19.07.1984: 1 9 ovig. 16.0 x
14.3 mm.
Musorstom 5: stn 361, 19°52.50'S, 158°38.10'E, 400 m, 19.10.1986: 1 6 19.8 x 17.9 mm.
Vanuatu. Musorstom 8: stn DW 965, 20°20’S, 169°51'E, 361-377 m, 21.09.1994: I 9 9.0 x 8.4 mm. —
Stn CP 982. 19°22'S. 169°26'E, 408-410 m, 23.09.1994: 1 6 7.7 x 7.2 mm.
French Polynesia (coll. SMSRB and J. Poupin). Tuamotu Archipelago, Fangalafu: stn 487, 22°14.1'S,
138°47.2' W, 310 m, 25.04.1995: I d 29.1 x 27.6 mm.
TYPES. — Dynomene tcmensis Yokoya, 1933: holotype is an ovigerous female 23.5 x 22.2 mm. collected by
the S.S. " Soyo-Maru ", from 30°06’N, 130°50'E (approximate coordinates), east of Tanegasima Id, Japan. 219 m,
between 1923 and 1930. 1 have been unable to trace the specimen so I assume that the specimen no longer exists.
DESCRIPTION. — Carapace slightly wider than long, ratio of CW/CL 1.05-1.10, rectangular in outline, surface
smooth, quite convex, with a few minute granules in branchial area. Carapace surface densely covered with setae of
only one kind: very short, soft setae, which are minutely serrated, clothing entire surface. Pereopods covered with
short setae as well as a few longer Filiform setae (5 x length of short setae and 0.10 x CW) which fringe limbs.
Density of short setae completely obscures body surface and on carapace they present a symmetrical undulating
aspect reflecting gentle undulations in carapace surface: one oblique trough lies behind supraorbital margin, with a
short median longitudinal trough extending posteriorly, then a trough curving anlerolatcrally which marks cervical
groove, followed by a trough running across midline, just in front of cardiac area, which splits into two lateral
troughs, and finally a short trough crossing cardiac area. All setae have same structure: proximal 20% of shaft
smooth, followed by a region covered in long setules which increase in size distally, projecting almost at right
angles to shaft, and finally an unornamented region, slightly curved and narrowing to an acute tip.
A shallow frontal carapace groove separates a pair of low rounded protuberances, and then divides into two
separate, short, faint grooves on a flattened area. Just in front of cardiac region two laterally-directed grooves
originate: first groove (cervical) arises separately (but very close together) from small gastric pits curving (slightly
sinuously) anterolaterally on to branchial region towards gap between second and third anterolateral teeth, while
second, shallower groove extends across mid-line and initially runs almost directly towards lateral margin but then
splits into an anterior portion which follows first groove but does not reach lateral carapace margin, while second
portion curves posterolaterally, bordering anterior cardiac region, meeting a branchial groove running to base of
last anterolateral tooth. Posterior cardiac area marked by a distinct groove crossing mid-line. Anterolateral carapace
margin begins at level of postorbital corner, slightly convex and bears three distinct, broad-based blunt teeth. First
tooth smallest and close to postorbital corner followed closely by second tooth, both directed almost anteriorly.
Third tooth (largest) more distant and directed laterally. A posterolateral tooth marks beginning of convergent
posterolateral border alongside which lies reduced last leg. Posterior carapace margin recessed in order to
accommodate distal section of first segment of abdomen which is visible dorsally.
Frontal margin continuous, V-shaped, minutely granulated, ventrally-directed, joined to epistome (which
separates orbits). Supraorbital margin not projecting, continuous above orbits, not interrupted by a notch, and
without granules. Suborbital margin, convex, without teeth, projecting, shelf-like and easily visible dorsally.
Orbits oblique and clearly exposed dorsally.
First article of antennule large, filling a large part of ventral orbital region: distal margin obliquely angled and
not continuous with distal margin of second antennal article. Remainder of antennule folded into orbit. First article
of antenna moveable, wider than long, medially beaked; inferior tooth well developed, blunt; superior tooth, above
the opening of antennal gland, smaller but more acute and directed ventrally. Second article wider than long, distal
Source : MNHN. Paris ,
FAMILY DYNOMENIDAE
523
Fig. 27. — Metadynomene tanensis (Yokoya, 1933), 2 19.2 x 18.9 mm, New Caledonia, MUSORSTOM 4, stn 215, 485-
520 m: a, dorsal view of right half of carapace; b, ventral view of right orbital area; c, outer face of right cheliped;
d, dorsal view of right cheliped; e, posterior view of terminal articles of right fourth pereopod; f. posterior view of
terminal articles of right fifth pereopod; g. ventral view of telson and terminal segments of female abdomen.
Source : MNHN. Paris
524
C. L. McLAY
margin widest, to which is fixed the exopod curving over base of eyestalk and becoming broader and terminating
bluntly. Third antennal article longer than wide, and attached to remaining distal border of second article, slotting
in behind exopod, and just matching length of exopod. Fourth antennal article smaller, as long as wide; remainder
of antennal articles directed laterally, extending well beyond postorbilal corner, and partially folded under supra¬
orbital margin. Ratio of length of antennal flagella to CW = 0.33. Eyestalk can be completely folded into orbit,
and cornea is well developed, occupying all of tip. Epistome broadly triangular, surface slightly concave; dorsal
arm, joined to tip of carapace, very elongate and narrow; lateral amis shorter and thicker. Joint between epistome
and carapace is marked by a narrow suture.
Subhepatic area smooth, very convex. A groove begins near base of antenna, curving round under branchial
region and meeting lateral carapace margin just anterior to last tooth at beginning of posterolateral border. A short
cervical groove branches off and ascends towards gap between first and second anterolateral teeth. Third maxillipeds
opcrculiform, bases widely separated by tip of sternum. Crista dentata has twelve teeth increasing in size distally.
Female sternal sutures 7/8 short, ending wide apart in a V-shaped groove created by a low medial parallel ridge,
just behind bases of second walking legs. Sternal sutures 7/8 concealed by a dense layer of long soft setae from
adjacent coxa of third walking leg.
The branchial formula is 20 gills and 7 epipodites on each side:
A podobranch is definitely present on last pereopod but there is no epipod. A cross section of an arthrobranch
shows lateral margin deeply notched, dividing gill into a hypobranchial plate (containing efferent vessel) and an
epibranchial lobe. Between these marginal lobes are a pair of shorter lobes. Thus epibranchia! surface shows four
rows of blunt lobes, which are arranged above afferent blood vessel. Towards tip of the gill, length and number of
lobes is gradually reduced. Second maxilla has two long setae extending into gill chamber. Hypobranchial setae in
posterior region of branchial chamber are well developed. Hypobranchial margin of each podobranch armed with
long setae as found on epipods to which they are attached. Posterior margin of scaphognathite bears two long
setae. Hypobranchial margin of podobranchs bears same setae as on epipod.
Cheliped stout, much longer and stouter than first leg. Merus trigonal, inner face smooth and fitting closely
against pterygostomial region of carapace; borders granulate, superior border has a subterminal broad, restriction
which separates a thickened, smooth distal ridge, from a row of four to five small granules; inferior face has three
subdistal, blunt tubercles. Outer face of carpus convex with many small granules, separated by a smooth
longitudinal channel, two more prominent blunt tubercles on distal margin; inner superior border with three blunt
tubercles, distal one largest which abuts against proximal inner surface of propodus thereby restricting closure of
cheliped against frontal area. In a similar way, inferior carpal margin is produced as a smooth obtuse flange fitting
against merus when limb is withdrawn. Outer and superior faces of propodus with several small granules, inner
and inferior faces smooth, except that there is a small proximal tubercle on inner propodal face. Fixed finger
almost straight with seven or eight almost obsolete teeth increasing in size distally; moveable finger not strongly
curved, with one large proximal tooth and four teeth at tip, interlocking with opposing teeth. A narrowing band of
setae extend on to outer face of moveable finger. Both fingers, thick, hollowed out internally, gaping basally,
touching for about half their length. In hollowed out interior of each finger there are prominent small tufts of long
setae which come together when fingers are closed.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of meri of first two
legs and anterior face third leg merus smooth and nacreous, inferior distal margin hollowed out to accommodate
carpal article. Superior border of meri of these legs with several small granules, length of merus of second leg
Source : MNHN. Paris .
FAMILY DYNOMENIDAE
525
about 2.5 x width and equal to about half of CL. Anterior and posterior dorsal margins of carpi without granules,
and produced distally to overhang base of propodi. Surface of propodi smooth. Dactyli curved, inferior margin
armed with 3-4 small distal spines, tip dark brown and subacute.
Last pair of legs greatly reduced, lying along posterolateral border of carapace, reaching almost to end of meral
article of preceding limb; borders of articles unarmed. Legs subchelate, sexually dimorphic: female with well de¬
veloped distal extension of propodus which opposes dactyl, male with only weakly developed propodal extension.
Female propodal extension bearing eight, unequal, stout, hooked, spines with tiny flattened teeth along most of
concave inner surface, and distal area free of teeth. Female dactyl as long as propodal extension, bearing eleven
unequal, stout, hooked spines (arranged asymmetrically around perimeter of the dactyl) whose concave inner
surface is devoid of tiny teeth. Male propodal extension bearing five unequal curved spines without teeth. Male
dactyl longer than propodal extension and ending in a single acute claw which has a tiny acute spine on
its outer margin.
All segments of abdomen freely moveable. Tclson much wider than long, anterior margin essentially straight,
posterior margin broadly rounded. In both sexes uropod plates are large, filling all of space between penultimate
abdominal segment and tclson, excluding all of last abdominal segment from reaching lateral margin of abdomen.
Abdominal locking mechanism consists of a small ridge or spine on coxae of first and second walking legs
adjacent to uropods and penultimate abdominal segment respectively. Abdomen only loosely held against sternum
in both sexes. In mature female, coxal spines absent; abdomen occupies all of ventral surface, covering coxae of
all pcrcopods with tclson covering proximal half of third maxillipeds. In male, abdomen not quite so broad and
telson only extends as far as bases of third maxillipeds.
Five pairs of plcopods in female, first pair vestigial, remainder biramous. (Seventeen percent of females had
male first pleopods instead of the normal vestigial type. See discussion below). First male pleopod a semi-rolled
tube with a small apical plate surrounded by long setae. Second pleopod with an exopod on basis, needle-like
distally, armed with a series of twenty four tiny, curved, acute, inset spines and ending in two larger curved spines.
Subterminal spines evenly spaced and not following a sinuous path. Third to fifth male pleopods rudimentary and
biramous, both articles about same length and not separated from base by a joint.
COLOUR. — The colour picture in POUPIN (1996b) shows the whole body covered with a pale tan velvet
tomentum, which is a darker brown colour on the carpi of each pereopod. Cheliped fingers may have a pink
colouration.
Geographic Distribution. — The type locality given by Yokoya (1933) is "east of Tanegasima Id". The
distribution of Metadynomene tanensis includes Japan, Taiwan, Indonesia, New Caledonia (including Loyalty Ids,
and Chesterfield Ids), and Tuamotu, French Polynesia. The records for Taiwan, Indonesia, New’ Caledonia and
Tuamotu are new.
Depth. — The depth range of the material examined is 205-520 m, although the majority of specimens come
from 300-400 m. The low'er depth limit is a little uncertain but it is at least 500 m. The type specimen,
an ovigerous female approx. 23.5 x 22.2 mm, came from a depth of 219 m. Metadynomene tanensis is clearly
a deep water western Pacific species.
Size. — The maximum size for males is 29.1 x 27.6 mm, for females 24.8 x 22.4 mm, and the smallest
ovigerous female is 16.0 x 14.3 mm. Although sexually mature females have been recorded from almost every
month, only 4 ovigerous females are known. These females were collected in May, when the eggs were newJy laid
or showed little development, and July, when the eggs were eyed and ready to hatch. This suggests that the
breeding season of Metadynomene tanensis may be restricted to a short period and that females may only have
a single brood of eggs each year. This may be a consequence of living at a depth where water temperatures are
much lower. Only one female, 17.6 x 16.3 mm. gave a reliable egg count of 2800. The mean egg diameter is
0.62 mm, which is somewhat larger than for the shallow water dynomenids, but probably still consistent with the
assumption that this species has planktotrophic larvae. The largest female with an immature sized abdomen was
1 1.2 x 10.5 mm, and the smallest female with a mature sized abdomen was 1 1.3 x 10.6 mm. Therefore females
probably mature at a size within the range of 1 1.0-12.0 mm CW.
526
C. L McLAY
DISCUSSION. — This enigmatic species has been largely ignored and in the Japanese carcinological literature
its status has been uncertain (see below). The only report of Metadynomene tanensis is the original description by
Yokoya (1933). The description is brief and his figure 38 is not entirely in agreement with the text which is as
follows:
" Carapace and appendages of legs covered with short and dense hairs; and with a few long setae on the margins
of the legs. Carapace subcircular, regions marked by some transverse grooves; five anterolateral teeth very shallow
and obtuse. Front broad triangular, grooved in the medial line. Upper and lower margin of orbit smooth, outer
angle rounded. Meri and carpi of legs irregularly tuberculated. Chelipeds subequal, stouter and longer than
succeeding legs; fingers naked near the extremities, with deeply excavated extremities, inner edges dentate, gaping
at the base when closed; last leg slender and shorter than one half of the preceding leg, the tip minutely chelate."
YOKOYA (1933, fig. 38) shows an ovigerous female which still has all the setae and the last pair of legs is
shown as having the same orientation as the other walking legs. However this cannot be accurate because in all
dynomenids the last pair of legs are always straight and lie alongside the posterolateral margins of the carapace.
Most of the text could apply to any of the other species in this genus except for the details about the anterolateral
teeth. Thus the interpretation of these teeth is critical to establishing the use of this name. The text states that
there are five shallow and obtuse anterolateral teeth, but the figure shows only two teeth plus the posterolateral
tooth. In their natural state the specimens examined here closely resemble YOKOYA's figure and they agree in
having the stated number of teeth. If it is assumed that there were indeed five teeth on the type specimen, then
they must have been made up as follows: the first tooth close to the orbit (shown in his figure), the second and
smaller third teeth (not shown in the figure), the fourth tooth (shown), and the fifth tooth (shown) must be the
posterolateral tooth. Thus the features of the present material can be reconciled with both the text and the figure.
One additional feature of YOKOYA's figure, which is not mentioned in the text, is the shape of the chclipcd carpal
article. The outer face of this article appears deeply sculptured because of the arrangement of tubercles and apart
from the differences in the anterolateral margin, this characteristic is sufficient to separate it from M. devaneyi in
which the surface of the carpal article is much less sculptured. For these reasons it seems valid to identify the
present material as being Metadynomene tanensis (Yokoya, 1933). The major differences between the three species
of this genus are discussed below (see Table 3) under M. crosnieri sp. nov..
Although the two species are clearly different, there has been some confusion between Dynomene praedator and
Metadynomene tanensis from Japan. Sakai (1936) listed Dynomene tanensis but stated that he had not seen any
specimens. Without giving any reasons, Serene (1968) stated that these two species were synonyms. For
D. praedator from Japan, Sakai (1976) used a text-figure which showed M. tanensis and placed Dynomene
tanensis Yokoya, 1933 in the list of synonyms for Dynomene praedator. The habitat information which he gives
is almost certainly for D. praedator , i.e. "coral reefs, shallow waters", and the records of material examined (from
Yoron Id, Taketomi Id, and the Bonin Ids) are probably for this species. There is no record from Sagami Bay
(Sakai, 1965) and it is only in 1976 that there is any evidence he had seen a specimen of M. tanensis (from
Ishigaki Id. depth not given, 1 male approx. 15.0 x 13.5 mm) which he referred to D. praedator. In his list of the
Japanese fauna Miyake (1983) also assumed that D. tanensis is a synonym of D. praedator. Thus it is important
to clearly establish that these are two valid species and that both occur in Japanese waters. D. praedator is a
shallow water species (0-50 m) while M. tanensis occurs in much deeper water (205-520 m).
The setae of M. tanensis clothe almost the entire body surface and are for the most part densely covered in long
setules, giving the surface of the crab a soft velvet-like appearance. Although there are a few longer setae, all setae
have the same microscopic structure. In this respect M. tanensis is similar to Dynomene hispida and D. praedator
but the setules are longer and cover more of the shaft. Also the bare tip is relatively much shorter.
Metadynomene tanensis is the only dynomenid examined in which the branchial formula is 20 gills +
7 epipods. This species has a small podobranch on the last pereopod but this is absent in the other dynomenids.
Besides the seven epipods M. tanensis has two long setae on the posterior margin of the scaphognathite and a well
developed field of hypobranchial setae at the back of the gill chamber. These are all part of the mechanism used for
gill cleaning and they are aided by the presence of long cleaning setae on the hypobranchial margin of each
podobranch. These setae probably clean the bases of the adjacent gills. The structure of the gills is very similar to
that of M. devaneyi with four epibranchial lobes associated with each gill plate.
Source :
FAMILY DYNOMENIDAE
527
The propodus and dactyl of the last pair of legs in females show some similarity to those found in the species
of Dynomene : they have dentate propodal spines, and edentate dactylar spines. Similarly, the males have five
edentate propodal spines, as in D. filholi and D. pilumnoides. but the dactyl is most unusual in having an acute
spine on one side. This identical structure has only been observed in one other dynomenid species, D. filholi.
although a similar spine on the dorsal margin is present in Paradynomene tubercidata. The structure is reminiscent
of that found in some dromiids (see Discussion under D. filholi).
The first two pairs of male pleopods agree closely with those of Metadynomene devaneyi as figured by
Takeda (1977, fig. la-c). Examination of the M. tanensis second pleopod, using the scanning electron
microscope, shows that it is armed with a large number (24) of tiny curved spines. Similar large numbers of
spines arc found in Hirsutodynomene Ursula , H. spinosa . and Dynomene pilumnoides although in these species
the spines are not curved. As with most of the other dynomenids the last three pairs of pleopods in M. tanensis are
biramous but there is no sign of any joint between the separate articles and the basal article.
Examination of the stomach contents of a male, 19.8 x 17.9 mm, from New Caledonia showed mostly fine
sand grains, soft unidentifiable organic material as well as chopped dark brown chitinous fragments. Like most of
the other dynomenids, Metadynomene tanensis may obtain most of its food by sieving sand, using the setae on
the cheliped fingers and third maxillipeds, and perhaps some by scavenging.
Occurrence of gynandromorph Females
Metadynomene tanensis was one of the more common species in the collection studied. A total of
73 specimens were examined: 25 males and 48 females. The sex ratio was 1.92 females/male, but the most
interesting feature was the occurrence of females with male first pleopods. While the second to fifth biramous
pleopods were developed in the normal way, these gynandromorphic females had male pleopods in place of the
normally vestigial uniramous first pleopods. Seventeen percent of the females were gynandromorphs and most of
them were sexually mature: the smallest was CW = 13.0 mm and the largest was CW = 19.2 mm. One female,
CW = 13.7 mm, had one male pleopod and one normal vestigial pleopod. Although it was not possible to
examine the internal organs, these abnormal females seemed to be normal in every other respect, having well
developed genital apertures in the coxae of the third pereopods and an abdomen as well developed as in normal
females. They are probably reproductively successful, although none were carrying eggs. The female pleopods on
the second to fifth abdominal segments were of the normal size and structure, and there was no evidence of
parasitism. The male pleopods of the gynandromorphs were only about half the size that would be expected for a
male of the same size. None of the females of any of the other dynomenid species examined showed any evidence
of developing male first pleopods.
Relative growth of the abdomen and chelipeds in Metadynomene tanensis is shown in Fig. 28 a-b. The size
of normal females ranged from CW = 4.8 to CW = 24.8 mm and included three ovigerous females. Relative
width of the last abdominal segment for small crabs shows no difference between females and males, but there is
a sudden increase for females larger than about CW =11.0 mm, suggesting a pubertal moult. The abdomen
of males continues to grow at the same relative rate as for small crabs. The abnormal crabs do not differ in
their abdomen size from mature females. With cheliped propodus length (or depth for that matter), both males
and females show a similar pattern until they reach about CW = 15.0 mm when male chelipeds become relati¬
vely larger. This suggests that males may reach sexual maturity at a larger size than females. For abnormal crabs,
the relative growth of both the abdomen and the chelipeds shows a pattern which conforms to that of females.
The smallest of the abnormal specimens, CW = 6.4 mm, was unusual in that the male pleopods were the
longest while the remaining female pleopods (biramous) were very small. Since the genital apertures are not
developed at this size, and both sexes have five pairs of pleopods, the only criterion for deciding on gender is
whether the last four pairs of pleopods are biramous (female) or uniramous (male). Since the abdomen size of
males and females up to about CW =11.0 mm does not differ, it is unclear whether this specimen is a
masculinized female or a feminized male. In Fig. 28 a-b this specimen is shown as an abnormal female. Another
small female, CW = 8.6 mm, had uniramous vestigial first pleopods with the rest biramous but all the pleopods
were very small and not properly developed. Again, the original sex of this specimen is uncertain because it is the
same as the first case, except that its first pleopod did not show any male characteristics.
528
C. L. McLAY
Assuming that the mature abnormal crabs have passed through the same stages as the two small crabs
discussed above, then it is not clear whether they represent crabs which started out as males or as females. If we
assume that they were originally males, then this would help to explain why the overall sex ratio is biased in
favour of females. Perhaps there are some individuals in the population which have delayed sex determination,
retaining the potential to become either male or female. While the rate of occurrence of abnormal crabs would be
17% if we assume that they are masculinized females, it would be 8/40 = 20% if they were feminized males and
the overall sex ratio would be 1.21 females/male rather than 1.92. There is no evidence of sex change such as
would be found in a sequential hermaphrodite because females reach sexual maturity at about CW = I I mm and
males at about 15 mm CW. Another hypothesis is that these abnormal crabs represent "errors in development",
but 17% seems to be a very high rate of occurrence. An alternative explanation is that the development of these
two small abnormal crabs has been modified by a parasite, but there was no external evidence of parasitism.
Moreover, the parasite hypothesis would not account for the occurrence of the abnormal mature females.
The occurrence of gynandromorphic individuals is rare amongst Malacostraca and appears to be limited to the
decapods (Charniaux-Cotton, 1975). Bilateral gynandromorphs have been reported for Metapenaeus monoceros
(George, 1963), a species of Lucifer (see Manning & Holthuis, 1981), Nephrops norvegicus (Farmer.
1972), Homarus americanus (Chace & Moore, 1959) and H. gammarus (Gordon, 1957), Jasus frontalis
(reported as Palinurus frontalis by BURGEN, 1902) and J. edwardsii from New Zealand (pers. obs.), and Cambarus
propinquus (reported as Orconectes propinquus by Hay, 1905). Some cases involve the presence of supernumerary
genital openings e.g. Astacus astacus (reported as A. fluviatilis by Benham. 1891). Immature female
Procambarus clarkii which received androgenic gland implants developed male first pleopods and. when mature,
vitellogenesis was inhibited (Taketomi & Nishikawa, 1996).
Bilateral gynandromorphs have also been reported in brachyurans such as Chionoecetes opilio (Taylor.
1986), and Callinectes sapidus (Cargo, 1980; Johnson & Otto, 1981). More complex and peculiar
malformations have been found in several other brachyurans. Veillet (1945) described an unusual Carcinus
maenas (Portunidae) in which there were two male pleopods and a male genital aperture on the right side, while on
the left side there was a male first pleopod and four female pleopods, with a female genital aperture in the thorax.
Froglia and Manning (1978) reported a gynandromorph specimen of the grapsid Brachynotus gemmellari which
had male first pleopods but female second pleopods. Manning (1993) reported a similar case of a segmental
gynandromorph in the pinnotherid Nepinnotheres androgynus which had a wide abdomen with male pleopods and
female pleopods, but lacked female gonopores. Another peculiar case was reported by GORDON (1963) for a male
Pleistacantha moseleyi which had five pairs of pleopods: normal male first and second pleopods, except that the
second carried an exopodite, followed by three pairs of semi-biramous pleopods. MANNING and Holthuis (1981)
found one gynandromorph specimen of Ebalia tuberculata Miers (Lcucosiidae) which had one half of the abdomen
with female characteristics while the other half had male characteristics. Sternal female gonopores were present and
four normal female pleopods were found on the right hand side of the abdomen, but on the left hand side there were
male pleopods on the first two segments, followed by normal female pleopods on the third to fifth segments.
Evidently, female characters were dominant in this crab. ROPER (1979) found that around 1% of Leptomithrax
longipes (Majidae) had poorly developed female gonopores, abdomens of intermediate size and varying degrees of
development of male pleopods on the first abdominal segment while the third to fifth segments had normal female
pleopods. He concluded that these specimens were males showing various degrees of feminization which may have
been attributable to the effects of a bacterium. Hartnoll (1960) found a specimen of Hyas coarctatus Leach
(Majidae) which had both female and male genital openings while there was a pair of male pleopods followed by
three pairs of female pleopods on the abdomen. Several morphological features were intermediate in size between
typical males and females. This crab most closely approximates the hermaphroditic condition.
In the abnormal specimens ot Metadynomene tanensis only the pleopods of the first abdominal segment were
modified to the male form. Whereas in Ebalia tuberculata the first male pleopod was of normal size, in
M. tanensis the pleopods were only about half the size that would be expected in a male of the same size. In both
these species female characters were evidently dominant. The E. tuberculata specimen is a case of bilateral
gynandromorphism while B. gemmellari , N. androgynus and most of the M. tanensis specimens, were segmental
gynandromorphs, although one M. tanensis female had one male first pleopod as well as a typical female
Source : MNHN. Pans
FAMILY DYNOMENEDAE
529
first pleopod. In most of the decapods mentioned above, gynandromorphism is very rare, except in the case of
Nephrops norvegicus where up to 12% of males can be affected (RlDEWOOD, 1909). CHARNIAUX-COTTON
(1975) concludes that, at least in the case of bilateral gynandromorphs, the cause of feminization is genetic in
origin, resulting from the loss of the male chromosome from a blastomere at an early stage of cleavage, so that
Fig. 28. — Relative growth of Metadynomene tanensis (Yokoya, 1933): a. cheliped propodus length and depth plotted
against carapace width; b. width of penultimate abdominal segment plotted against carapace width (mm). Open circles
are for males, closed circles are for females, and triangles are for gynandromorphic individuals.
Source
530
C. L. McLAY
one half of the animal develops as a male while the other develops as female. It seems most likely that the cause
of the pleopod abnormalities in M. tanensis is also genetic in origin and may be environmentally induced. Given
that male decapods are usually the heterogametic sex (Lecher et al. , 1995), it may well be that the abnormal
animals are feminized males rather than masculinized females.
Metadynomene crosnieri sp. nov.
Figs 25 c, 29
Dynomene devaneyi - Guinot, 1993: 1227 [Not Takeda, 1977].
Material EXAMINED. — Glorieuses Islands. BENTHEDl: 11°32.00'S, 47°16.40'E, 330-440 m. 7.06.1977: I <3
23.2 x 22.7 mm (MNHN-B 22510, originally identified as D. devaneyi by C. L. McLay).
Types. — The holotype is a male 23.2 x 22.7 mm, collected during the BENTHEDl Expedition from
1 1°32.0'S, 47°16.4'E. off the Glorieuses Ids, Indian Ocean, 330-440 m, 7.06.1977, held at the Museum national
d'Histoirc naturelle, Paris, registration number MNHN-B 22510.
Description. — Carapace- about as wide as long, ratio of CW/CL = 1.02. rectangular in outline; surface
smooth, quite convex, no granules. Carapace surface densely covered with setae of only one kind: very short, soft
setae, which arc minutely serrated, clothing entire surface. Pereopods covered with short setae as well as a few
longer filiform setae (5 x length of short setae and 0.10 x CW) which occur sparsely on limbs with a tuft
associated with dactyli of second to fourth pereopods. Density of short setae completely obscures body surface and
on carapace they present a symmetrical undulating aspect reflecting gentle undulations in carapace surface: one
oblique trough lies behind supraorbital margin, with a short median longitudinal trough extending posteriorly,
then a trough curving anterolaterally which marks cervical groove, followed by a trough running across midline,
just in front of cardiac area, which splits into two lateral troughs, and finally a short trough crossing cardiac area.
Microscopic details of setae not investigated.
A shallow frontal carapace groove separates a pair of low rounded protuberances, and then divides into two
separate, short, faint grooves on a flattened area. Just in front of cardiac region two laterally-directed grooves
originate: first groove (cervical) arises separately (but very close together) from small gastric pits curving (slightly
sinuously) anterolaterally on to branchial region towards gap between second and third anterolateral teeth, while
the second, shallower groove extends across mid-line and initially runs almost directly towards lateral margin but
then splits into an anterior portion which follows the first groove for a short distance, while the second portion
curves posterolaterally, bordering anterior cardiac region, meeting a branchial groove running to base of last
anterolateral tooth. Posterior cardiac area marked by a distinct groove crossing mid-line. Anterolateral carapace
margin begins at level of postorbital corner, slightly convex and bears three similar, very small, subacute teeth.
First tooth close to postorbital corner followed closely by second tooth, both directed almost anteriorly. Third and
fourth teeth, directed laterally, more distant and separated from first two by a marginal swelling above which is a
small tubercle. There is also a small tubercle just above base of posterolateral tooth. On righthand side (illustrated)
lourth anterolateral tooth is absent. A posterolateral tooth, behind branchial groove, marks beginning
of convergent posterolateral border alongside which lies the reduced last leg. The posterior carapace margin
recessed in order to accommodate distal section of first segment of abdomen which is visible dorsally.
Frontal margin continuous, V-shaped, minutely granulated, ventrally-directed, joined to epistome (which
separates orbits). Supraorbital margin not projecting, continuous above orbits, not interrupted by a notch, and
without granules around postorbital corner. Suborbital margin, convex, without teeth, not projecting, scarcely
visible dorsally. Orbits oblique and clearly exposed dorsally. First article of antennule large, filling a large part of
ventral orbital region, distal margin obliquely angled and not continuous with distal margin of second antennal
article. Remainder of antennule folded into orbit.
Fiist article ol antenna moveable, wider than long, medially beaked; inferior tooth well developed, subacute;
superior tooth, above opening of antennal gland, blunt, smaller and directed ventrally. Second article wider than
Source : MNHN, Pans -
FAMILY DYN0MEN1DAE
531
Fig. 29. — Metadynomene crosnieri sp. nov.. 6 23.2 x 22.7 mm, holotype, Glorieuses Ids, BENTHEDI, 330-440 m:
a. dorsal view of right half of carapace; b. ventral view of right orbital area; c. outer face of left cheliped: d. dorsal
view of left cheliped; e. posterior view of terminal articles of right fourth pereopod; f. posterior view of terminal
articles of right fifth pereopod; g, ventral view of telson and terminal segments of male abdomen.
Source : MNHN, Paris
532
C. L. McLAY
long, distal margin widest, to which is fixed the exopod curving over base of eyestalk and becoming broader and
terminating bluntly. Third antennal article longer than wide, and attached to remaining distal border of second
article, slotting in behind exopod, and exceeding length of exopod. Fourth antennal article smaller, as long as
wide, remainder of antennal articles directed laterally, extending well beyond postorbital corner, and can be partially
folded under supra-orbital margin. Ratio of length of antennal flagella to CW = 0.35. Eyestalk can be completely
folded into orbit; cornea well developed, occupying all of tip. Epistome broadly triangular, surface slightly
concave; dorsal arm joined to tip of carapace, very elongate and narrow; lateral arms shorter and thicker. Joint
between epistome and carapace marked by a narrow suture.
Subhepatic area smooth, except for three or four minute granules, very convex. A groove begins near base of
antenna, curving round under branchial region and meeting lateral carapace margin just anterior to tooth at
beginning of posterolateral border. A short cervical groove branches off and ascends towards base of marginal
swelling between first and second pairs of anterolateral teeth. Third maxillipeds operculiform, bases widely
separated by tip of sternum. Crista dentata has thirteen well developed teeth increasing in size distally. Female
sternal sutures 7/8 unknown.
Since there is only one specimen, branchial formula could not be determined but examination of an
arthrobranch from second pereopod shows that gills differ slightly from preceding species: a cross section shows
lateral margin deeply notched, dividing gill into a hypobranchial plate (containing efferent vessel) and an
epibranchial lobe. Between these marginal lobes are two pairs of shorter lobes. Thus the epibranchial surface
shows six rows of blunt lobes, which are arranged above afferent blood vessel. Towards lip of gill, length and
number of lobes is gradually reduced. There are well separated afferent and efferent channels in basal plate but with
a series of six elongate lobes developed on outer margin. These lobes are arranged into a longer pair on each side
of gill with another, smaller pair medially. At base of outer lobes there is a small notch on each side, but this is
not produced as a lobe.
Chelipcd stout, much longer and stouter than first leg. Merus trigonal; inner face smooth and fitting closely
against pterygostomial region of carapace; superior border has a sublerminal broad, restriction which separates a
thickened, smooth distal ridge, from a row of five to six small granules; inferior face has three blunt subdistal
tubercles. Outer face of carpus convex with many small granules, which tend to be arranged in parallel rows, and
divided by a smooth longitudinal channel, two more prominent blunt tubercles on distal margin; inner superior
border with three blunt tubercles, all of similar size and most distal one abuts against proximal inner surface of
propodus thereby restricting closure of cheliped against frontal area. In a similar way. inferior carpal margin
produced as a smooth obtuse flange fitting against merus when limb is withdrawn. Outer and superior faces of
propodus with scattered small granules; inner and inferior faces smooth, except that there is a small proximal
tubercle on inner propodal face. Fixed finger almost straight with seven or eight small teeth increasing in size
distally; moveable finger not strongly curved, with one large proximal tooth and four teeth at tip, interlocking
with opposing teeth. A narrowing band of setae extend on to outer face of moveable finger. Both fingers, thick,
hollowed out internally, gaping basally, touching for about half their length. In hollowed out interior of each
finger there are small tufts of long setae which come together when fingers are closed.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of meri of first two
legs and anterior face third leg merus smooth and nacreous, inferior distal margin hollowed out to accommodate
carpal article. Superior border of meri of these legs with several small granules, length of merus of second leg
about 2.6 x width and equal to about half of CL. Anterior and posterior dorsal margins of carpi without granules,
and produced distally to overhang the base of propodi. Surface of propodi smooth. Dactyli curved, inferior margin
armed with four small distal spines, tip brown and subacute.
Last pair of legs greatly reduced, lying along posterolateral border of carapace, reaching about halfway to end of
meral article of preceding pereopod; borders of articles unarmed. Male propodal extension bearing Five unequal
curved spines. Male dactyl longer than propodal extension and ending in a single acute claw. Microscopic details
of dactyl and propodal spines unknown. Female unknown.
All segments of abdomen freely moveable. Telson much wider than long, anterior margin essentially straight;
posterior margin broadly rounded. Uropod plates large, filling all of space between penultimate abdominal segment
and telson, excluding all of last abdominal segment from reaching lateral margin of abdomen. No effective
Source : MNHN, Paris .
FAMILY DYNOMENIDAE
533
abdominal locking mechanism: small bifid tubercle on coxa of first walking leg beside uropods restricts sideways
movement. Abdomen only loosely held against sternum. Tclson only extends as far as bases of third maxillipeds.
Five pairs of pleopods in male; first pleopod a semi-rolled tube ending in a curved apical plate surrounded by
long setae; second pleopod needle-like with an exopod on basis, remaining pleopods rudimentary and biramous.
Microscopic details unavailable. Female unknown.
ETYMOLOGY. — This new species is named after Alain CROSNIER in recognition of his enormous
contribution to the study of decapod crustaceans.
COLOUR. — Whole body covered with a pale tan velvet tomentum.
Geographic Distribution. — Metadynomene crosnieri is known only from the type locality near the
Glorieuses Ids, south of the Seychelle Ids. Indian Ocean.
Depth. — The depth of the type locality is between 330 and 440 m.
Size. — Only the male type specimen is known, 23.2 x 22.7 mm.
DISCUSSION. — At a casual glance the three species in this genus are extraordinarily similar: they are relatively
large dynomenids and all have a short, soft, undulating tomentum. Indeed the specimen of M. crosnieri was
originally mis-identified as M. devaneyi. Of the external differences between the species (see Table 3) the most
important are those of the anterolateral carapace margin and the prominence of the suborbital margin. M. devaneyi
is distinctive in not having any anterolateral teeth while M. crosnieri has four small subacute teeth.
The anterolateral teeth in M. tanensis are somewhat variable but there are usually at least three well developed
teeth. In all cases it is necessary to carefully remove the tomentum to expose the anterolateral margin.
Table 3. — Comparison of the Metadynomene species.
There are some minor differences between the gills of the three species of Metadynomene. M. crosnieri has the
largest number of epibranchial lobes (six) while the other two species have four. Thus in all species the
epibranchial portion of the gills is trichobranchiate-Iike, while the hypobranchiai portion is more
Source
534
C. L. McLAY
phyllobranchiate-like. In M. devaneyi and M. tanensis the hypobranchial cleaning setae are well developed. In
M. tanensis there is a podobranch on the last pereopod, but this gill is absent in M. devaneyi.
GuiNOT (1993, as Dynomene devaneyi) described the abdominal locking structures of Me tadynomene crosnieri
and the rudimentary third male pleopod. The male abdomen is only loosely retained by small bifid teeth on the
coxae of the second pereopods. The rudimentary third to fifth male pleopods are biramous as in the other two
species of this genus.
The species of Metadynomene show a remarkable resemblance to the dromiid Dromia wilsoni (Fulton &
Grant, 1902). The short undulating tomentum gives a superficial similarity, but inspection of the last two pairs of
legs reveals that these dynomenids are very different. Material sorted by a non-specialist often has these species
mixed together. Since all these species live in deeper water, these features may be convergent.
None of the species of Metadynomene have overlapping distributions: M. crosnieri is known from the Indian
Ocean, M. tanensis from the western Pacific and M. devaneyi from much further east at Hawaii. Given their high
degree of similarity it may be that these species are of quite recent origin.
Genus ACANTHODROMIA A. Milne Edwards, 1880
Acanthodromia A. Milne Edwards, 1880: 31. — BouviER, 1896: 56. — ALCOCK, 1899: 134; 1901: 36. — Ortmann.
1899: 1 155. — A. Milne Edwards & Bouvier. 1902: 22. — Rathbun, 1937: 55.
Diagnosis. — Carapace longer than wide, convex, ovoid; surface bristling with long spines. Lateral carapace
margin poorly defined, without distinct teeth but bearing numerous spines. Carapace grooves not well marked, but
lateral cardiac and branchial grooves faintly evident. Frontal carapace margin broadly triangular, spinous; eyestalks
short; eyes protected by well defined orbits. Sternal sutures 7/8 of female end well apart on low tubercles behind
bases of second walking legs.
Antennule can be concealed inside orbit at base of eyestalk. Antennal flagella shorter than carapace width.
All articles of antenna moveable, first article (urinal) not beaked medially and second article has an exopod firmly
fixed. Third maxillipeds opercular, completely covering buccal cavern, separated at their bases by a plate at same
level as sternum; basis and ischium of endopod fused but joint always marked by a shallow groove. Crista dentata
absent. Chclipeds equal, stouter than walking legs; last pair of legs very reduced; dactyl rudimentary, forming
an obsolete subchelate mechanism with an extension of propodus. Gills usually 19 (including 6 podobranchs) +
7 epipods. Gill structure basically phyllobranchiate.
Abdomen of six segments and telson folded loosely under thorax, uropods large. Abdominal locking
mechanism present, using well developed coxal projections. Surface of abdominal segments spinous, except for
fourth segment which bears one or more large, pearl-like, medial tubercles. Females have five pairs of pleopods,
first pair vestigial, remainder biramous and of normal size. Male pleopods unknown.
Type Species. —Acanthodromia erinacea A. Milne Edwards. 1880, by monotypy. Gender is feminine.
Other Species. — Dynomene margarita Alcock, 1899.
DISCUSSION. — The original definition of Acanthodromia by A. Milne Edwards (1880) included reference to
the shape of the carapace (narrow and ovoid) and its fronto-orbital region, third maxillipeds, walking legs, and the
rudimentary and cheliform last legs. A. Milne Edwards placed this genus in his family "Dromiens", between
Dromia and Dynomene , along with other dromiids and the homolodromiid genus Dicranodromia A. Milne
Edwards, 1880. Later A. Milne Edwards and Bouvier (1902) separated these into three sub-families, retaining
Dynomene and Acanthodromia in the Dynominae. They noted that Acanthodromia represented a curious mixture of
primitive (e.g. carapace longer than wide) and "secondary" (e.g. non filamentous gills) characters which
distinguished it from Dynomene. To these can be added the absence of a beaked first antennal article and a crista
dentata. The main reason for including Acanthodromia in the Dynomenidae is the structure of the reduced last pair
of legs. To include it any where else would require the assumption that these limbs had evolved more than once.
Source : MNHN. Paris -
FAMILY DYNOMENIDAE
535
Wright and Collins (1972: 24) have suggested that the genus Acanthodromici should be included in the
fossil family Prosopidae Von Meyer, 1860, sub-family Pithonothinae Glaessner. 1933, thus making it the only
genus of this family to survive to recent times. They considered that " Acanthodromici appears to be no more than a
spinose Plagiophthalmus Bell, 1863 in which the main furrows are obsolescent." However this seems ill-advised
because, although the carapace of Acanthodromici is longer than wide, it lacks a distinct carapace margin which is a
feature of Plagiophthalmus. Thus the indication by BRIGGS et al (1993) that the Prosopidae have a fossil record
extending from the mid Jurassic to the present day is incorrect.
Acanthodromia erinacea A. Milne Edwards. 1880
Figs 6 d, 9 f, 10 a, 30
Acanthodromia erinacea A. Milne Edwards, 1880: 31. — Bouvier, 1896: 56, figs 18-21. — Young. 1900: 336. —
Alcock, 1901: 75 (list). — A. MILNE Edwards & Bouvier, 1902: 23, text-figs 7-8, pi. 3, figs 5-15. pi. 4, figs 1-4.
— Ihle. 1913: 92 (list). — Rathbun, 1937: 55, pi. 12. figs 5-6. — Rice, 1981: 174.
Material EXAMINED. — Guadeloupe. "Blake": stn 166, 275 m. coll. A. Agassiz. 21.01.1878: 1 9 ovig. 16.7 x
17.7 mm, holotype (MCZ 6509).
Mexico. Yucatan. ", Albatross ": stn 2354, Arrowsmith Bank. 238 m, 22.01.1885: 1 9 9.5 x 11.5 mm. (USNM
9547).
Types. — The holotype is an ovigerous female 16.7 x 17.7 mm, collected by A. AGASSIZ from "Blake"
stn 166, 15°55.50'N, 61°37.05'W, Leeward Ids, off Guadeloupe, 275 m, 21.01.1878, held at the Museum of
Comparative Zoology, Harvard University, registration number MCZ 6509. A paratype specimen, consisting of
only a carapace, was collected from "Blake" stn 232, 13°6.45'N. 61°6.55’W. Windward Ids, off St. Vincent,
158 m, 21.02.1879, registration number MCZ 2641.
Description. — Carapace longer than wide, ratio of CW/CL = 0.90-0.94, ovoid in outline, surface evenly
convex, with a dense cover of acute spines and spinules with occasional long setae. Microscopic details of setae
not investigated. Density of spines completely obscures body surface. Frontal, cervical, and post-cervical grooves
not evident, crescentic lateral cardiac grooves and branchial groove faint, posterior cardiac area not defined.
Anterolateral carapace margin poorly defined, begins below level of postorbital corner, subparallel and adorned
with longer spines but these are not arranged in a well-defined row. Posterolateral border convergent alongside
which lies the reduced last leg. Posterior carapace margin recessed in order to accommodate first segment of
abdomen which is visible dorsally.
Frontal margin V-shaped, spinous, ventrally-directed. joined to epistome (which separates orbits). Supraorbital
margin projecting, continuous above orbits, eave-like, not interrupted by a notch, adorned with long spines which
become smaller near postorbital corner and curved posteriorly. Suborbital margin concave adorned by long spines
with a larger spine at inner corner. Cornea of eye and suborbital margin clearly exposed dorsally.
First article of antennule large, filling a large part of ventral orbital region; prominent spine mid-way along
length; distal margin spinous obliquely angled and not continuous with distal margin of second antennal article.
Remainder of antennule folded into orbit. First article of antenna moveable, wider than long, bearing a row of
three or four spines, not medially beaked, opening of antennal gland is on medial margin and concealed against
base of antennule. Second article spinous, longer than wide; medial margin longest produced as a spine, to which
third article is fixed. Exopod short, spinous, blunt extending behind third antennal article and curving over base of
eyestalk. Third and fourth antennal articles as wide as long, lying over distal end of exopod and continued as
a flagellum which is 0.6 x CW. Eyestalk can be completely folded into orbit (where it is well protected by
marginal spines), outer surface spinous; cornea light brown, well developed, occupying al! of tip. Epistome
triangular, surface slightly concave; dorsal arm joined to tip of carapace, margins bearing spines; lateral arms
shorter. Joint between epistome and carapace marked by a narrow suture.
Subhepatic area inflated, covered with tubercles. A groove (pleural suture or linea dromica) begins near base of
the antenna, curving round under branchial region and towards anterolateral carapace margin about mid-way along
Source :
536
C.L. McLAY
its length. A short groove branches off, ascending and curving towards postorbital corner marking posterior
margin of inflated subhepatic area. Third maxillipeds operculiform, bases separated by tip of sternum, without
crista dentata; coxae armed with prominent medial projections under which telson fits when at rest. Female sternal
sutures 7/8 short, ending wide apart under an overhanging lip immediately below female gonopores.
Branchial formula according to A. MILNE Edwards and Bouvier (1902: 24) is 19 gills + 7 cpipods
(see Discussion below). In cross section gills consist of pairs plates, one on each side of gill axis, with
epibranchial tips of each plate ending in a blunt thickened lobe. Lateral margins of each plate faintly notched about
mid-way along their length.
Chcliped stouter and longer than first leg. Merus trigonal, inner face roughened with rounded tubercles and
fitting closely against pterygostomial region of carapace; borders spinous, superior border has a faint subterminal
restriction which separates a thickened distal ridge, on which there are several spines, from a row of five or six
similar spines on superior border. Outer face of carpus convex with small blunt tubercles interspersed among
longer, sharper spines; inner superior border lacks a flattened, distomedially directed, spur restricting closure of
cheliped. Instead, both superior and inferior inner margins of carpus are spinous like outer face. Entire surface of
propodus covered with spines which are longer on superior and outer faces. Outer surface of Fingers covered with
tubercles and small spines. Dactyl strongly downcurved, margin sinuous but not interrupted by teeth except at lip
where there are two blunt teeth, roof of finger strongly concave. Fixed finger almost straight with three evenly
spaced, blunt teeth on outer margin, three further teeth on tip (interlocking with pair of teeth on dactyl), inner
margin without teeth and floor strongly concave. Small groups of long stiff setae, inserted near base of dactyl and
fixed finger, are directed across space between the two fingers. Spoon-shaped fingers gape proximally on internal
face but there is only a small gape externally.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of meri of First two
legs and anterior face third leg merus covered with low rounded tubercles; inferior distal margin hollowed out to
accommodate carpal article. Superior border of meri of these legs with numerous long spines, length of merus of
second leg about 1.5 x width and equal to about one third of CL. Dorsal margins of carpi bearing several long
spines, and produced distally to overhang base of propodi. Propodi bearing numerous long spines. Daclyli curved,
bearing numerous shorter spines; inferior margin armed with five small spines similar to lip which is dark brown
and subacute.
Last pair of legs greatly reduced, lying along posterolateral border of carapace above bases of walking legs,
reaching to about half-way along meral article of preceding limb; borders of articles spinous; basis-ischium and
merus fused. In female last pair of legs subchelate with well developed distal extension of propodus which opposes
dactyl. Propodal extension bearing seven, unequal, stout, hooked, spines facing laterally, with marginal rows
of 6-10 tiny flattened teeth proximally on concave inner surface, and distal area free of teeth. These marginal teeth
are curved inwards, without meeting the opposite row, and directed distomedially. Female dactyl as long as
propodal extension, bearing eight unequal, stout, hooked spines (arranged asymmetrically around perimeter of
dactyl) whose inner surface is edentate. Male unknown.
All segments of abdomen freely moveable, length and breadth of all segments similar, surface spinous, with
occasional setae, margins unarmed. First segment partially concealed under posterior border of carapace, visible
portion fits into recess and articulates with carapace margin; anterior margin of second segment sinuous; medial
region convex and inserted under margin of preceding segment; lateral margins produced as a flange which Fits over
posterior margin of first segment thereby preventing forward slippage of abdomen. Subsequent segments
articulated in a similar manner. Anterior half of fourth segment with two small pearl-like medial tubercles almost
totally united except for a narrow proximal fissure. Rest of medial region behind these two rounded tubercles bears
several spines. Fifth abdominal segment has a pair of similar but smaller tubercles, separated by a Fissure which
extends posteriorly on to sixth segment, between the tubercles. Telson spinous, much wider than long; anterior
margin angled to accommodate uropod; posterior margin broadly rounded. In female uropod plates are large, Filling
about half of the space between last abdominal segment and telson, excluding much of last abdominal segment
from reaching lateral margin of abdomen. Male characters unknown. Abdominal locking mechanism well
developed: when at rest, abdomen of mature female lies between bordering flanges on first three pereopods with
telson beneath coxal projections of third maxillipeds.
Source : MNHN, Paris .
FAMILY DYNOMENIDAE
537
1 cm a-b
1 cm c-e
1 mm f
Fig. 30. — Acanthodromia erinacea A. Milne Edwards, 1880. 9 ovig. 16.7 x 17.7 mm, holotype, Guadeloupe, "Blake",
stn 166, 275 m (MCZ 6509): a, dorsal view of right half of carapace; b. ventral view of right orbital area; c, outer
face of right cheliped; d. dorsal view of right cheliped; e. posterior view of terminal articles of right fourth pereopod;
f. posterior view of terminal articles of right fifth pereopod; g. ventral view of telson and terminal segments of
female abdomen (uropods concealed by strong spinulation).
Source :
538
C.L. McLAY
Five pairs of pleopods in female, first pair vestigial and not carrying eggs, remainder biramous. Male pleopod
characters unknown.
COLOUR. — Pale cream when preserved.
Geographic Distribution. — Along the chain of Caribbean islands from the Arrowsmith Bank (20°N.
86°W), off the Yucatan coast of Mexico, to St. Vincent (I3°N, 61°W) in the Windward Ids. Acanthodromia
erinacea seems to be an insular species like the other Atlantic dynomenid, Dynomene filholi.
Depth. — The depth range for this species is 158-540 m. Acanthodromia erinacea lives at a greater depth than
most of the species of Dynomene. The only information about its habitat comes from the specimen collected at
158 m off St. Vincent, Windward Ids where the substrate is described as "coral bottom".
Size. — Only three females of Acanthodromia erinacea are known as well as a carapace of unknown sex.
These collection records have been summarized by Rathbun (1937). The largest female (the holotype), which
was ovigerous, measured 16.7 x 17.7 mm and carried about 100 eggs of 0.5 mm diameter. This female was
collected in January, 1878 and the eggs were newly laid. The egg size is similar to that of the other dynomenid
species. RICH ( 1 98 1 ) examined eggs from a female collected in February 1933 which were very close
to hatching.
DISCUSSION. — RICE (1981) described the pre-zoca of Acanthodromia erinacea dissected from the eggs of a crab
collected by the Johnson-Smithsonian Expedition off Mona Id, West Indies. Although the material was rather
unsatisfactory, he concluded that the larva of this species is most similar to those of the Dromiidae and that
anomuran characteristics are strongly evident in both families. This pre-zoea remains the only known larval
material of any of the dynomenids.
The most accurate and detailed description of the reduced last pereopods of any dynomenid is given for
Acanthodromia erinacea by A. MlLNE Edwards and BOUVIER (1896, pi. 3, fig. 12) where they describe for the
female "...la pince parfaite qui lermine ces appendices est munie de soies spiniformes sur le bord des doigts, qui
sont deux fois plus courts que la region palmaire". The presence of spines on the propodus and dactyl has largely
escaped the attention of other dynomenid researchers. The number and arrangement of these spines in
Acanthodromia erinacea is similar to that found in the other dynomenids, but the teeth on the propodal spines are
unusual in being curved inwards. The number of these teeth is very reduced compared to Dynomene hispida , for
e.g.. but similar to the situation found in Paradynomene tuberculata and Hirsutodynomene Ursula females. In these
species the teeth are only found on the margins and are restricted to the proximal region of the spines. The last leg
of the A. erinacea male is unknown. In both P. tuberculata and A. erinacea the basis-ischium and merus articles of
the fifth pereopods are fused to make a single bent or curved article.
Bouvier (1896) compared the chelate nature of the last legs o f Acanthodromia erinacea with that found in
Dicranodromia ovata A. Milne Edwards, 1880 and Homolodromia paradoxa A. Milne Edwards, 1880. He noted
that in the sequence D. ovata - H. paradoxa - A. erinacea the propodus becomes more developed so that a
subchelate limb becomes a "..pince parfaitement caracterisee". He argued that A. erinacea was probably derived
from a very primitive form in which the last pair of legs were similar to that found in H. paradoxa. This idea
about the origin of dynomenids may well be correct but Bouvier could not have known that there are considerable
differences in the fine structure of the limbs between dynomenids and homolodromiids. His intuitive idea ignores
the fact that the propodal and dactyl spines of A. erinacea are constructed on a different plan, and the dactyl itself is
also very different from that found in any brachyuran crab: at least in females, it is more like a blunt, flattened,
articulated plate, with the spines arranged around the margins, rather than an articulated claw with a single row of
medial spines. Considerable evolutionary change is required to transform a camouflage-carrying homolodromiid
limb into what may have been some kind of cleaning appendage.
The branchial formula of Acanthodromia erinacea is summarized in the table at the top of the next page.
The branchial formula given by A. Milne Edwards and Bouvier (1902) for this species is 19 gills +
7 epipods. However there seem to some errors in this interpretation. Firstly, they omitted the podobranch of the
second maxilliped and secondly, they indicated that the third maxilliped had two arthrobranchs. It seems likely that
Source : MNHN, Paris -
FAMILY DYNOMENIDAE
539
the correct number is only one arthrobranch, as found in all other dynomenids which have been examined. Thus
the total number of gills is still 19. but for different reasons. The gill structure of A. erinacea is very different
from other dynomenid species: there are no epibranchial lobes and the gills consist of paired plates surrounding the
gill axis. A. MlLNE Edwards and Bouvier (1902, pi. 4, fig 4) show a small epibranchial projection between
these plates, but I could not find this. The anterior and posterior margins of each plate are slightly notched. The
gill structure of 4. erinacea most closely resembles that found in Dynomene hispida and D. praedator. The gill
structure of Acanthodromia margarita is virtually identical. A. MlLNE Edwards and Bouvier (1902, pi. 3, fig 7)
show three long setae on the posterior margin of the scaphognathite. This number of setae is only found in
Dynomene fdholi and D. pilumnoides. All the other dynomenid species only have two such setae.
The mature female Acanthodromia erinacea has spinous coxal projections on the third maxillipeds and the first
three pereopods which appear to help hold the abdomen in place. In order to release the abdomen, it would be
necessary for these pereopods to move posteriorly. In other dynomenids, neither males nor females, have any
really effective abdominal locking mechanism, and the abdomen is only held loosely against the sternum. In
immature dromiids both sexes have an abdominal locking mechanism, usually involving pereopodal coxae and the
uropods, which only persists in mature males. Therefore A. erinacea is unusual in having a locking mechanism in
mature females. Unfortunately the male of this species is unknown but presumably it also has a similar
mechanism.
For two species inhabiting widely separated regions (Indo-West Pacific and Caribbean Sea), Acanthodromia
margarita and A. erinacea are extraordinarily similar: the only differences are in the shape of the supraorbital
spines, and the pearl-like lobes on the fourth and fifth abdominal segments. A likely scenario for the origins of
these two species might be as follows: species of Acanthodromia originated in the Tethys Sea and spread to seas
that eventually became the Atlantic and Caribbean, perhaps as early as the Upper Jurassic (145 mybp). This
dispersal could have been as late as the Middle Miocene (25 mybp), when connections between these two oceans
was severed, but an earlier date seems more likely because even by Palaeocene times (65 mybp) the Atlantic was
already well formed and the Caribbean isolated. Therefore these two species could have been separated at least since
the Palaeocene (or at latest the Middle Miocene), but there has scarcely been any divergence in their morphology.
Both are deeper water species although A. erinacea extends to a greater depth (540 m). They grow to a similar
maximum size and inhabit muddy coral bottoms. A. erinacea appears to be a relict Tethyan species.
Acanthodromia margarita (Alcock, 1899)
Fig. 31
Dynomene margarita Alcock, 1899: 19, pi. 2, fig 3.
Acanthodromia margarita - ALCOCK, 1900: 134; 1901: 36, pi. 1, fig. 3, 3a. — lHLE, 1913: 92 (list). — Sakai, 1965b:
43; 1976: 31. pi. 7, fig 2. — Serene, 1968: 37 (list). — Miyake, 1983: 196 (list). — Nagai. 1989: 43.
MATERIAL EXAMINED. — Japan. Wakayama, 120 m, 1989. S. Nagai coll.: 1 9 12.0 X 13.0 mm (see Nagai,
1989).
Types. — Dynomene margarita Alcock, 1899: holotypc is a male 4.5 x 5.0 mm, collected by the
" Investigator ”, from 13°16.00'N, 93°08.00'E, Andaman Sea, 135 m, held by the Indian Museum. Calcutta,
registration number 2690/10.
Source :
540
C. L. McLAY
Description. — Carapace longer than wide, ratio of CW/CL = 0.92, ovoid in outline; surface evenly convex,
with a dense cover of acute spines and spinules with occasional long setae. Microscopic details of setae not
investigated. Density of spines completely obscures body surface. Frontal, cervical, and post-cervical grooves not
evident, crescentic lateral cardiac grooves and branchial groove faint, posterior cardiac area not defined.
Anterolateral carapace margin poorly defined, begins below level of postorbital corner, subparallel and adorned
with longer spines but these are not arranged in a well-defined row. Posterolateral border convergent alongside
which lies the reduced last leg. Posterior carapace margin recessed in order to accommodate first segment of
abdomen which is visible dorsally.
Frontal margin V-shaped, spinous, ventrally-directed. joined to epistomc. Supraorbital margin projecting,
continuous above orbits, eave-like, not interrupted by a notch, adorned with long spines which become smaller
and blunter near postorbital corner and are not curved posteriorly. Suborbital margin concave, adorned by small
spines with a larger spine at inner comer. Cornea of eye and suborbital margin clearly exposed dorsally.
First article of antennule large, filling a large part of ventral orbital region, prominent spine mid-way along
length; distal margin spinous, obliquely angled and not continuous with distal margin of second antennal article.
Remainder of antennule folded into orbit. First article of antenna moveable, wider than long, bearing a row of
spines, not medially beaked, opening of antennal gland is on medial margin and concealed against base of
antennule. Second article spinous, longer than wide, medial margin longest produced as a spine, to which is fixed
the third article. Exopod short, spinous, blunt extending behind third antennal article and curving over base of
eyestalk. Remaining antennal articles absent from specimen. Eyestalk can be completely folded into orbit, outer
surface spinous; cornea light brown, well developed, occupying all of tip. Epistome triangular, surface slightly
concave; dorsal arm joined to tip of carapace margins bearing spines; lateral arms shorter. Joint between epistome
and carapace marked by a narrow suture.
Subhepatic area inflated, covered with tubercles. A groove (pleural suture or linea dromica) begins near base of
antenna, curving round under branchial region and on to carapace to meet faint branchial groove. A short groove
branches off, ascending and curving towards postorbital corner marking posterior margin of inflated subhepatic
area. Third maxillipcds operculiform, bases widely separated by tip of sternum. Female sternal sutures 7/8 short,
ending wide apart under an overhanging lip immediately below female gonopores.
Branchial formula unknown. In cross section gills consist of pairs plates, one on each side of the gill axis,
with epibranchial tips ol each plate ending in a blunt thickened lobe. Lateral margins of each plate are faintly
notched about mid-way along their length.
Cheliped stouter and longer than first leg. Merus trigonal, inner face roughened with rounded tubercles and
fitting closely against pterygostomial region of carapace; borders spinous, superior border has a faint subterminal
restriction which separates a thickened distal ridge, on which there are several spines, from a row of five or six
similar spines on the superior border. Outer face of carpus convex with small blunt tubercles interspersed among
longer, sharper spines; inner superior border lacks a flattened, distomedially directed, spur restricting closure of
cheliped. Instead, both superior and inferior inner margins are spinous like the outer face. Entire surface of
propodus covered with spines which are longer on superior and outer faces. Outer surface of fingers covered with
small spines. Dactyl strongly downcurved, margin sinuous but not interrupted by teeth except at tip where there
are two blunt teeth, roof of finger strongly concave. Fixed finger almost straight with three evenly spaced blunt
teeth on the outer margin, three further teeth on tip (interlocking with pair of teeth on dactyl), inner margin
without teeth and floor strongly concave. Small groups of long stiff setae, inserted near base of dactyl and fixed
finger, are directed across the space between the two fingers. Spoon-shaped fingers gape proximally on internal
face but there is no gape externally.
First three pairs of walking legs decreasing in length posteriorly. Meri elongate, both faces of ineri of first two
legs and anterior face third leg merus covered with low rounded tubercles, inferior distal margin hollowed out to
accommodate carpal article. Superior border of meri of second to fourth pereopods with numerous long spines,
length of merus of third pereopod about 2.0 x width and equal to about one quarter of CL. Dorsal margins of carpi
bearing several long spines, and produced distally to overhang base of propodi. Propodi bearing numerous long
spines. Dactyli curved, bearing numerous shorter spines; inferior margin armed with four small spines similar to
tip which is dark brown and subacute.
Source : MNHN, Pahs
FAMILY DYNOMENIDAE
541
Fig. 31. — Accmthodromia margarita (Alcock, 1899): a-f. 9 12.0 x 13.0 mm, Wakayama. Japan. 120 m: a. dorsal view
of right half of carapace; b. ventral view of left orbital area (note that most of antenna is missing) ; c, outer face of
right cheliped; d. dorsal view of right cheliped; e, posterior view of terminal articles of right fourth pereopod;
f, ventral view of telson and terminal segments of female abdomen (note that uropod plates are missing).
Source : MNHN. Paris
542
C. L. McLAY
Last pair of legs greatly reduced; articles spinous, lying along posterolateral carapace border above bases of
posterior walking legs. Structural details of subchelate mechanism not available.
All segments of abdomen freely moveable, length and breadth of all segments similar, surface spinous,
margins unarmed. First segment partially concealed under posterior border of carapace, visible portion fits into a
recess and articulates with carapace margin, anterior margin of second segment sinuous, medial region convex and
inserted under margin of preceding segment, lateral margins produced as a flange which fils over posterior margin
of first segment thereby preventing forward slippage of abdomen. Subsequent segments articulated in a similar
manner. Fourth segment with two large pearl-like medial tubercles separated by a narrow fissure filled with short
setae. Telson spinous, much wider than long; anterior margin angled to accommodate uropod; posterior margin
broadly rounded. In female uropod plates are large, filling about two-thirds of space between last abdominal
segment and telson, excluding last abdominal segment from reaching lateral margin of abdomen. Male characters
unknown. Abdomen of mature female occupies all of ventral surface, covering coxae of all pereopods with telson
covering proximal half of the third maxillipeds. Abdominal locking mechanism well developed: when at rest
abdomen of mature female lies between bordering flanges on first three pereopods with telson beneath coxal
projections of third maxillipeds.
Five pairs of pleopods in the female, first pair vestigial, remainder biramous. Male characters unknown.
Colour. — Pale cream. Eyes deeply pigmented when preserved.
Geographic Distribution. — Andaman Sea, Indian Ocean, Tosa Bay, and Kii Peninsula, Japan. This is a
rare species: there are only three records of Acanthodromia margarita (1 6.2 9 ).
Depth. — The depth range for Acanthodromia margarita is 120-200 m. The depth at the type locality, in the
Andaman Sea, was 135 m. Sakai (1976) describes the bottom from which his specimen came as being muddy.
Size. — The only male specimen known is the type which measured approximately 4.5 x 5.0 mm.
The largest female has been reported by Sakai (1976) but the ratio of the dimensions given in the text do not
agree with his figure: assuming that the CL is correct, then the dimensions must be 15.5 x 17.0 mm.
No ovigerous females of Acanthodromia margarita are known.
DISCUSSION. — The description given above is based on the specimen of Acanthodromia margarita reported by
Nagai (1989) and is largely in agreement with the original description except that the branchial groove is not
clearly evident as claimed by Alcock (1899). Admittedly the branchial groove is difficult to discern amongst
the forest of spines. There are no features of the Japanese specimen which are different from those of the type
from the Indian Ocean.
In the text ALCOCK (1899) placed this species in Dynomene , but in the Corrigenda, at the beginning of his
paper, he indicated that it should be placed in Acanthodromia. There was a delay in publication of the "Account of
the Deep-Sea Brachyura..."and meanwhile the generic designation had been corrected in his "Materials for
a carcinological fauna of India" (Alcock, 1900) where reference is made to A. Milne Edwards (1880).
Essentially the same description is repeated by ALCOCK (1899, 1900, and 1901) but no comparison of
A margarita with A. erinacea is made.
The gill structure of Acanthodromia margarita is essentially the same as that of A. erinacea. Both species have
gills which lack epibranchial lobes, making them almost phyllobranchiate, except for the marginal notch which is
characteristic of dynomcnid gills. The relationships of these two species is discussed under A. erinacea.
Genus PARADYNOMENE Sakai, 1963
Paradynomene Sakai, 1963: 230; 1965a: 13. — Guinot, 1993: 1226.
DIAGNOSIS. — Carapace subquadrangular in shape, slightly longer than wide; surface convex, granulate, well
areolated; individual areolae each having one or two low conical tubercles. Lateral carapace margins well defined.
Source : MNHN. Paris
FAMILY DYNOMENIDAE
543
subparallcl and armed with irregular teeth. Narrow frontal groove split in two posteriorly; cervical, postcervical
grooves evident. Frontal carapace margin well produced anteriorly, cut into three teeth; median tooth small; lateral
teeth broad and cristate; eyestalks short; eyes protected by well defined orbits. Sternal sutures 7/8 of female end
well apart on low tubercles behind bases of second walking legs.
Antennule can be concealed inside orbit at base of eyestalk. Antennal flagella shorter than half of carapace
width. All articles of antenna moveable; first article (urinal) beaked medially and second article has an exopod
firmly fixed. Third maxillipeds opercular, completely covering buccal cavern, separated at their bases by a plate at
the same level as the sternum; basis and ischium of endopod fused but joint always marked by a shallow groove.
Crista dentata present. Chelipeds robust, equal, stouter than walking legs. Last pair of legs very reduced; dactyl
rudimentary, forming an obsolete chelate mechanism with an extension of propodus only in female. Branchial
formula 19 gills + 7 epipods.
Abdomen of six segments and telson folded loosely under thorax; uropods comparatively small; effective
abdominal locking mechanism absent. Both sexes have five pairs of pleopods, first pair vestigial in female, last
three pairs rudimentary in male. First pair of male pleopods consist of a stout, setose semi-rolled tube with an
apical plate; second pair needle-like bearing a linear row of tiny inset spines along anterior surface (modified from
Sakai, 1963).
Type Species. — Paradynomene tuberculata Sakai, 1963 by original designation and monotypy.
DISCUSSION. — Paradynomene Sakai, 1963, collected from Sagami Bay, Japan, was the third extant genus of
dynomenids to be discovered. As with Acanthodromia, this genus is radically different from the type genus of the
family ( Dynomene ). While both have a carapace which is longer than wide, in Paradynomene the carapace is
subquadrangular with tubcrculate areolae and in Acanthodromia it is ovoid and densely covered with spines.
However, all three genera share the distinctive characters associated with the last pereopods. The characters of the
second male pleopod are also similar to those found in Metadynomene and Dynomene. It is intriguing to make the
same kind of comparison between members of the Dromiidac: the genus Epigodromia McLay, 1993 is to Dromia
or Cryptodromia , what Paradynomene is to Dynomene. Epigodromia has a highly areolate carapace and obsolete
last two pairs of legs, while Dromia and Cryptodromia have a smooth setose carapace and functional last two pairs
of legs. Perhaps this tendency to evolve a thickened, ornamented carapace has occurred independently in the two
families (see Discussion below). Also Sakai (1963) noted that the general features of the front, thoracic
appendages and the external maxillipeds of Paradynomene resemble those of the family Dromiidae. rather than
those of Dynomene or Acanthodromia.
GUINOT (1993) suggested that there are certain resemblances between Paradynomene and the fossil genera
Rathbunopon Stenzel, 1945 and Mithracites Gould, 1859. The similarities relate to the ornamentation of the
carapace.
Paradynomene tuberculata Sakai, 1963
Figs 4 e-f, 6 e-f, 7 c, e, 10 b-d, 14 a, 25 d. 32
Paradynomene tuberculata Sakai, 1963: 231, fig. 8; 1965a: 13, pi. 6, fig. 1; 1976: 31. pi. 7, fig. 1. — SERENE, 1968: 37
(list). — Miyake, 1983: 196 (list). — Nagai. 1989: 43. — Guinot, 1993: 1227, figs 1-2.
Material EXAMINED. — Gulf of Aden. " Meteor" : stn Me 5/230-KD2. I2°43.5’N. 43°14.8'E, 214-277 m.
5.03.1987: 1 8 7.8 x 8.6 mm (SMF).
Indonesia. Karubar: stn DW 18. Kai Ids, 5°18'S, 133°0rE, 205-212 m. 24.10.1991: 1 9 ovig. 13.8 x 14.5 mm.
— Stn DW 49, Tanimbar Ids. 8°00'S. 132°59,E, 210-206 m, 29.10.1991: 1 8 11.2 x 12.5 mm.
Chesterfield Islands. Corail 2: stn DW 159, 19°46.04'S, 158°19.98’E, 52 m. 1.09.1988: 1 9 18.5 x 18.3 mm.
New Caledonia. Lagon: stn 444. 18°15.3’S, 162°58.8’E. 300-350 m. 28.02.1985: I 8 17.0 x 17.2 mm.
Smib 3: stn DW 14, 23°40.rS. 167°59.7’E, 246 m, 22.05.1987: 1 8 22.0 x 22.8 mm.
Smib 4: stn DW 44, 24°46.0'S, 168°8.2’E, 300 m, 8.03.1989: 1 8 17.0 x 17.8 mm.
Smib 8: stn DW 184, 23°18'S, 168°05’E, 305-320 m, 31.01.1993: 1 8 19.2 x 19.0 mm. — Stn DW 189. 23°18,S,
168°05'E, 400-402 m, 31.01.1993: 2 8 12.2 x 13.1, 21.0 x 21.9 mm.
Bathus 3: stn DW 830, 23°19’S, 168°0rE, 361-365 m, 23.11.1993: I 8 14.7 x 15.8 mm.
Source
544
C. L. McLAY
BATHUS 4: sin DW 931, 18°55'S. 163024’E, 360-377 m, 7.08.1994: 1 6 23.7 x 23.0 mm.
Halical I: stn DW 02. 18°54’S. 163°24'E, 352-397 m. 23.11.1994: 1 9 23.2 x 24.0 mm.
Loyalty Islands. MUSORSTOM 6: DW 406. 20°40.65'S, 167°06.80'E, 373 m. 15.02.1989: 1 6 21.5 x 22.3 mm;
1 9 21.5 x 21.2 mm; 1 9 ovig. 20.6 x 21.4 mm.
Guam (H. T. Conley coll.). Piti Lagoon. 13°27'N, 144°47'E, 1.2-7. 5 m deep in dead coral, 26.05.1994: I <3 21.7 x
20.0 mm (UGM). — Ibidem , 4-8 m, among dead coral, 5.06.1994: Id 17.2 x 16.4 mm (UGM). — Ibidem . 1.5-5 m, in
coral rubble, 12.05.1997: 1 9 ovig. 20.5 x 19.4 mm (UGM).
Types. — The holotype is a male 9.5 x 10.5 mm, collected by His Majesty the Emperor of Japan, from
35°08.00'N, I39°37.00'E, west of Jogashima Misaki, Sagami Bay, Japan, 85 m, deposited in His Majesty's
Museum at the Imperial Palace.
Description. — Carapace sparsely setose, longer than wide, ratio of CW/CL approx. 0.95, oblong in outline;
posterior margin truncate; surface convex, areolate and granulate. There are about eighteen to nineteen swellings
each marked by a subacute tubercle carrying three or four long (0.1 1 x CW), stiff serrate setae, and whole surface
is covered in evenly distributed, rounded granules. Largest tubercles on inner branchial area. Pereopods also cany a
few stiff setae, and abdomen margins and bases of third maxillipeds are densely covered with long, soft serrate
setae. General body surface thickly covered with short setae, although these are only evident under high
magnification. Structure of short and long setae are different. In short setae the proximal 40% of shaft is erect and
lacks ornamentation, then the setae bend at about 45° and bear two opposite rows of fine setules decreasing in size
distal ly . These setae are feather-like. lack an acute smooth tip, and are especially common in areas between
carapace swellings. In long setae the proximal 50% of shaft is covered with small setules, then next 45% bears
longer setules, projecting at right angles, increasing distally in size and finally last 5% is smooth, narrowing to an
acute tip. Distal 50% of each setae curved, forming a U-shape, so that tip of setae is directed towards body.
A narrow frontal carapace groove separates a pair of small tubercles behind frontal margin, and then divides
into separate grooves which terminate beside a similar median tubercle. Just in front of cardiac region two
laterally-directed grooves originate: first groove (cervical) arises separately (but close together) from small gastric
pits and runs directly anterolateral just in front of the largest tubercles on to branchial region. Second, shallower
groove extends across mid-line and joins two deep, longitudinal pits bordering anterior cardiac region. Cardiac area
is defined by grooves and adorned by two large tubercles. Branchial groove not evident. A row of four large
tubercles curve across posterior region of carapace. Anterolateral carapace margin begins below level of postorbital
corner, is slightly convex and bears six rather irregular, laterally directed, subacute teeth or tubercles. First, fourth
and sixth teeth largest, and about equidistant, while second, third and fifth are mere tubercles in between.
Posterolateral margin bears two teeth which are much larger than any of preceding anterolateral teeth. Posterior
carapace margin is recessed in order to accommodate first segment of abdomen part of which is visible dorsally.
Frontal margin projecting, tridentate, ventrally-directed, joined to epistome (which separates orbits). Median
tooth on a lower level, lateral teeth lie above on beginning of the orbital margin. Supraorbital margin has two
small tubercles followed by a small notch closer to postorbital corner, which is granulated; suborbital margin with
similar granules followed by a subacute tooth (visible dorsally) and then notched before inner corner. Orbits
obliquely arranged, clearly exposed dorsally.
First article of antennulc large, granulated, filling a large part of ventral orbital region; distal margin obliquely
angled and not continuous with distal margin of second antennal article. Remainder of antennulc folded into orbit.
Antennal articles granulated; first article moveable, wider than long, medially beaked; inferior tooth well
developed, blunt; superior tooth above opening of antennal gland is much smaller. Second article wider than long;
distal margin widest, to which exopod is fixed, curving over base of eyestalk and becoming broader and
terminating bluntly. Third antennal article longer than wide, and attached to remaining distal border of second
article, slotting in behind exopod, and along with fourth article just surpassing length of exopod. Fourth antennal
article smaller, as long as wide; remaining antennal articles directed laterally, extending well beyond postorbital
corner, and can be partially folded under supra-orbital margin. Ratio of length of antennal flagella to CW = 0.35.
Eyestalk can be completely folded into orbit, and cornea is well developed, occupying all of tip. Epistome broadly
triangular, surface granulate and concave; dorsal arm. joined to tip of carapace, is very elongate and narrow; lateral
arms shorter and thicker. Joint between epistome and carapace is marked by a faint suture.
Source : MNHN. Paris
FAMILY DYNOMENIDAE
545
v-**y
Fig. 32. — Paradynomene tuberculatci Sakai, 1963, 9 ovig. 13.8 x 14.5 mm, Kai Ids, Indonesia, KaRUBAR, stn DW 18,
205-212 m: a, dorsal view of right half of carapace; b, ventral view of right orbital area; c, outer face of right
cheliped; d. dorsal view of right cheliped; e, posterior view of terminal articles of right fourth pereopod; f, posterior
view of terminal articles of right fifth pereopod; g, ventral view of telson and terminal segments of female abdomen.
Source : MNHN. Paris
546
C. L. McLAY
Subhepatic area granulate, very inflated to an acute angle which is adorned with two or three larger granules
running diagonally down from first anterolateral tooth. A groove begins near base of antenna, curving round under
branchial region and meeting lateral carapace margin just behind fifth tooth. A short cervical groove branches off
and ascends to gap between third and fourth anterolateral teeth and also gives off a branch which curves around
under and behind fourth tooth. Third maxillipeds operculiform. sharply angled, distal articles granular, proximal
articles densely setose, bases widely separated by tip of sternum. Crista dentata has twelve or thirteen small, blunt
teeth on each side which tend to increase in size distally. Female sternal sutures 7/8 short, ending wide apart just
below female gonopore. almost completely covered by the coxa and its setae of third walking legs.
Branchial formula is 19 gills and 7 epipodites on each side:
Podobranchs lack setae on their hypobranchial margin. In cross section, gill structure shows four elongate
epibranchial lobes radiating from the afferent vessel and a deep notch separating off hypobranchial plates which are
produced at corners. Hypobranchial setae in posterior region of branchial chamber very well developed. Posterior
margin of scaphognathite bears two long setae.
Cheliped stout, slightly longer than first leg; anterior border of basis-ischium densely setose. Merus trigonal,
inner face mostly smooth and fitting closely against pterygostomial region of carapace; borders and other faces
granulate; outer face has a subterminal broad, restriction which separates a thickened distal ridge on which there are
three larger granules from a row of several smaller granules on superior border. Inner inferior margin of merus not
bearing an enlarged granule. Outer face of carpus convex with many small, unequal granules, three more
prominent granules on distal margin; inner superior border with a distomedially directed, granulated spur which
abuts against the proximal inner surface of propodus thereby restricting closure of cheliped against frontal area. In
a similar way, inferior carpal margin produced as an obtuse, flange fitting against merus when limb is withdrawn.
Outer and superior faces of propodus and base of dactylus densely granulated. There are three or four prominent
granules on superior margin, inner face smooth and densely setose in male (setae absent in female), except that
there is a small proximal granule on inner propodal face. Fixed finger almost straight with seven blunt teeth
increasing in size distally; moveable finger not strongly curved, two small apical teeth, both fingers thick,
hollowed out internally, touching for about half their length. In female there are small tufts of long stiff setae
inserted proximally on each finger and extending across space between fingers. In male long soft propodal setae
extend on to fingers, covering about half length of both fixed finger and dactyl, and embedding the tufts of stiff
setae seen in the female. Externally these setae fill angle between fingers. Proximal half of third maxillipcd
concave, allowing chelipeds to fold tightly away beneath them.
First three pairs of walking legs decreasing in length posteriorly, articles granular. Meri elongate, plane of
movement dorsoventral, faces of meri of first three legs granulate, inferior distal margin hollowed out to
accommodate carpal article. Superior border of meri of these legs with four to six prominent, blunt spines, in a
row, separated by a gap from two similar distal spines, and three or four spines on posterior margin. Length of
merus of second leg about 1.6 x its width and equal to about 0.4 x CL. Dorsal surface of carpi bearing three or
four acute spines, and produced distally to overhang base of propodi. Posterior margins of carpi have an acute
spine. Surface of propodi with several similar spines. Dactyli curved, inferior margin armed with 4-5 small spines,
tip brown and subacute.
Last pair of legs greatly reduced; surface granular but not spinous; basis-ischium and merus fused, lying along
posterolateral border of carapace, reaching only as far as half-way along meral article of preceding limb. Sub-
chelate, sexually dimorphic: female with well developed distal extension of propodus which opposes dactyl, male
Source :
FAMILY DYNOMENIDAE
547
without a propodal extension. Female propodal extension bearing 14-16, unequal, stout, hooked, spines with a
ridged, concave inner surface and marginal rows of 7-12 tiny flattened teeth proximally. These teeth are directed
laterally. Female dactyl as long as propodal extension, bearing 15-16 unequal, stout, hooked spines (arranged
asymmetrically around perimeter of dactyl) whose concave inner surface is devoid of tiny teeth. Distal margin of
male propodus not produced (as illustrated by GUINOT, 1993, fig. 1), but bearing five unequal curved spines
without teeth. Male dactyl largely withdrawn into propodus and ending in a single acute claw which has a tiny
subterminal spine on its dorsal margin.
All segments of abdomen freely moveable; surface granular except for central area of telson which is smooth:
margins unarmed but fringed with short setae. Medial area of abdomen convex and clearly marked. First segment
divided into two parts: first part hidden, narrower and longer inserted under posterior margin of carapace; second
part, bearing four tubercles, wider and shorter with its anterior margin abutting against posterior carapace margin
preventing forward slippage of abdomen. Second segment narrowest, anterior margin sinuous, medial region
convex, lateral margins produced as a flange which fits over posterior margin of first segment; third segment
wider; both segments have a lateral tubercle on each side. Subsequent segments gradually increasing in length and
breadth distally, not overlapping with preceding segments. Telson much wider than long, anterior margin slightly
angled to accommodate uropod, posterior margin broadly rounded. A row of granules follows entire border of
telson, surrounding central smooth area. In both male and female uropod plates comparatively small, filling about
half of space between last abdominal segment and telson, so that part of last abdominal segment reaches lateral
margin of abdomen. No effective abdominal locking mechanism: abdomen only loosely held against sternum in
both sexes. Margins of the abdomen are neatly surrounded by many coxal granules which restrict sideways
movement. In mature female abdomen occupies all of ventral surface, covering coxae of all pereopods w ith telson
reaching to base of third maxillipeds. In male, abdomen not quite so broad and again, telson only extends as far as
base of third maxillipeds.
Five pairs of pleopods in female, first pair vestigial, remainder biramous. First male pleopod a semi-rolled
tube with a small apical plate surrounded by long setae. Second male pleopod with an exopod on basis, needle-like
distally, armed with a series of fourteen tiny, straight, acute, inset spines and ending in two very short straight
spines. Subterminal spines evenly spaced. Third to fifth male pleopods rudimentary and biramous, exopod
articulated.
Colour. — Pale brown. Tips of dactyl i black.
Geographic Distribution. — Paradynomene tuberculata was previously known from Japan and
New Caledonia (including the Chesterfield Ids, Loyalty Ids, Ride dc Norfolk). The material examined includes
specimens from Guam, Mariana Ids, Indonesia and the Gulf of Aden and these provide a large extension of the
range of this species.
Depth. — The depth range of the material examined in this study was 1.5-402 m with most of the specimens
coming from greater depths than previously reported. The depth at the type locality was 85 m and Sakai (1976)
gives a depth range of 35-85 m for his Japanese material. Their habitat includes dead coral and rocky bottoms. The
specimens of Paradynomene tuberculata from Guam were collected from very shallow depths: only 1.5-8 nt.
Size. — The maximum size for males is 22.0 x 22.8 mm and for females 20.6 x 21.4 mm. Only three
ovigerous females have been collected: a female 13.8 x 14.5 mm carried 400 eggs, another female 20.5 x
19.4 mm carried 840 eggs, while the female 20.6 x 21.4 mm was damaged with most of the abdomen and brood
missing. The mean egg diameter was 0.5 mm which is similar to the other dynomenids and indicates the existence
of a planktotrophic stage. Two females, carrying newly laid eggs, were collected during October and February and
one female was collected in May carrying eggs almost ready to hatch.
DISCUSSION. — The description of Paradynomene tuberculata given here differs somewhat from that of Sakai
(1963). In the original description Sakai stated that the lateral carapace borders are armed with six principal teeth,
of which the third is somewhat dorsal in position and the last located on the posterolateral border, with several
additional small teeth between the principal teeth. It is debatable whether the third tooth mentioned above should
548
C. L. McLAY
be regarded as being marginal or not. In my description I have not treated it as being part of the anterolateral
margin and I have treated the last two teeth as being on the posterolateral border because the carapace margin
begins to curve inwards posteriorly. Exact enumeration of the anterolateral teeth is made difficult by the fact that
there arc a variable number of small teeth/tubercles following the first lacinated tooth. Sakai also stated that the
dactyl i of second to fourth pereopods are unarmed, but 4-5 very small spines are present.
As found in all the other genera of the Dy nomen idae, there are two sizes of setae and as in most species (except
for Dynomene hispida and D. praedator) there are morphological differences between the two types of setae in
Paradynomene tuberculata . While the long setae are similar to those found in other species, the short setae are very
different. They are feather-like, lack an acute tip. and because they are bent, form a dense mat over the body
surface. The only other dynomenid with feather-like setae is Dynomene pugnatrix as described by DE Man (1889).
The rarity of this species prevents a comparative microscopic study to confirm this similarity. The unusual short
setae of D. filholi have a subterminal brush of fine setules, but in other respects they are quite different from the
setae of P. tuberculata.
The gills of Paradynomene tuberculata have the typical dynomenid structure: there arc four epibranchial lobes
separated from the hypobranchial plate by a notch. This similar to that found in Dynomene pilumnoides and the
species of Metadynomene. The posterior margin of the scaphognathite bears two long setae and there are no
cleaning setae on the hypobranchial margin of the podobranchs. P. tuberculata is the only dynomenid in which
these podobranch setae arc absent. Particularly noteworthy is the extensive development of hypobranchial setae on
the wall at the back of the gill chamber. These tufts of setae have also been reported from some dromiids.
In his original description, Sakai (1963) did not include a description of the male pleopods. Although
Paradynomene tuberculata is, in many respects, very different from the other dynomenids, its pleopods are clearly
built on the same plan. The first pleopod has an apical oval lobe surrounded by a dense fringe of long setae and the
second pleopod has an exopod at its base and is armed with a row of small inset distal spines. The terminal spines
are reduced making the tip almost blunt as is found in dromiids such as Stimdromia lameUata (Ortmann, 1894)
and Epigodromia gilesii (Alcock, 1899) (unpublished photos). The second male pleopods in these species lack
spines of any description.
Several authors have commented upon the similarities between Paradynomene and certain dromiids. JAMIESON
et ai (1993) described the structure of Paradynomene sperm and listed four or five synapomorphies with the sperm
of some dromiids. With respect to the general features of the carapace front, thoracic appendages, and external
maxillipeds, Sakai (1963) noted the resemblance of Paradynomene to some dromiids rather than to Dynomene or
Acanthodromia. The front or "face” of Paradynomene is different to the other dynomenids and remarkably similar
to some dromiids like Epigodromia spp. In both these genera the rostrum is clearly tridentatc, a feature seen only
in Paradynomene and no other dynomenids. Also the "face" in these two genera is much flatter because the
rostrum, inflated pterygostomial areas, and third maxillipeds all extend further forward compared to Dynomene
spp. The longer coxa of the third maxilliped results in the whole of the endopod extending more anteriorly. In
Paradynomene the merus of the third maxilliped forms almost a right angle with the preceding basis-ischium
article whereas it forms a much greater angle in Dynomene with all of the articles lying more or less in the same
plane. In both Paradynomene and Epigodromia the third maxilliped is much more strongly operculiform and fits
more closely against the epistome so that there is a much narrower gap. Furthermore the chelipeds of these two
genera are modified so as to fit compactly against the body and the bases of the third maxillipeds. In Epigodromia
the anterior surface of the cheliped basis-ischium and merus is flattened so that the remainder of the cheliped can be
folded tightly away. In Paradynomene the modification goes even further because the basis-ischium is sculptured
so that the inferior surface of the cheliped propodus fits closely against a raised ridge (see also GuiNOT, 1993).
These two genera are also similar in having strongly calcified, areolate and tuberculate exteriors.
In both Paradynomene tuberculata and Acanthodromia erinacea the basis-ischium and merus articles of the fifth
pereopods are fused to make a single bent or curved article. GUINOT (1993) examined closely the last pair of legs
in P . tuberculata , noting the coxal extension which conducts sperm to the base of the male insemination organs,
a feature unique to all dynomenids, and the extreme reduction of the dactyl, which is almost totally withdrawn
inside the end of the propodus. However this is only true of the male. In the female the propodal extension, and
opposing dactyl, are better developed (see figs lOb-d, 32f) and equipped with 14-16 hooked spines, amongst the
Source : MNHN. Paris
FAMILY DYNOMENIDAE
549
largest number of such spines found in any dynomenid. In the male the propodal extension is absent but still bears
five spines (as found in most other dynomenid males) while the very small dactylus has a spine on its dorsal
margin. This kind of dactyl spine is only found in males and besides P. tuberculata , only found in Dynomene
filholi and Metadynomene tanensis (see these species for further discussion). However in the latter two species the
spine, although similar in shape, is not on the dorsal margin but on the lateral margin of the dactylus. The
dactylar spines are reminiscent of those found in both males and females of dromiids such as Dromidiopsis
Borradaile, 1900. Tunedromia McLay, 1993, and Lauridromia McLay, 1993 where they are used, along with other
spines, to assist in securing the sponge carried by the last two pairs of legs over the crab. However, in the
dynomenids the spines are closely flattened against the surface of the dactyl so that they could not presently
function in the same way as in the dromiids. These spines indicate a common ancestral relationship.
All these characters seem to indicate evolutionary convergence in Paradynomene and Epigodromia. It may be
that the species of both these genera normally live partially buried in the surface coral fragments and that by
tightly folding away their pereopods they resemble their surroundings because the rugose subquadrangular carapace
resembles a piece of coral. It should be noted that P. tuberculata and E. areolata (Ihlc, 1913), for example, have
a similar depth range and specimens of both species have been collected from similar depths during the
New Caledonia Lagoon survey, 1985 and the MUSORSTOM 6, 1989, expedition to the Loyalty Ids (see McLay,
1993). Epigodromia has the last two pairs of pereopods very reduced, carried subdorsal and they are not used for
carrying Camouflage as seen in many other dromiids. Paradynomene , like all the other dynomenids, has only the
last pair of pereopods reduced but they are held horizontally. Thus while the reduced limbs may be an adaptation to
a cryptic way of life in Epigodromia, the reduced limb in Paradynomene is an ancestral character. Observation of
living specimens in natural surroundings would allow the hypothesis of cryptic convergence to be tested.
Examination of the stomach contents of a large male 21.0 x 21.9 mm (Smib, 8 stn DW 189, 400-402 m)
showed sand grains, soft unidentifiable organic material and chopped fragments of a hydroid coenosarc. One
specimen of Paradynomene tuberculata was associated with the stylasterine hydrocoral Stylaster (New Caledonia,
Lagon, stn 444, 300-350 m).
DISCUSSION
Habitat. — In most collections, dynomenid crabs are usually comparatively rare (Richer DE Forges, pers.
comm.). The habitat of the shallow water dynomenids (maximum depth < 100 m) , i.e. Dynomene hispida.
D. praedator, Hirsutodynomene spinosa and H. Ursula, seems to be rocky substrates and corals such as Acropora
spp., Pocillopora damicornis , P. elegans , Seriatopora hystrix , Goniastrea retiformis , Favia stelligera , Oulophyllia
crispa , Porites sp. and crustose alga such as Amphiora foliacea (EDMONDSON, 1946; Peyrot-Clausade. 1977,
1981; Ribes, 1978; Naim, 1980; Odinetz, 1983, and HlGHSMlTH, 1981). They seem to occur more often in dead
than in live coral. Dynomenids living in deeper water (maximum depth > 100 m), such as D. filholi.
D. pilumnoides , Metadynomene spp., Acanthodromia spp., and Paradynomene tuberculata, seem to live on
lithothamnion algae, red coral, precious coral (e.g. Corallium sp.) as well as rock and sand. The deepest living
dynomenids are M. tanensis (520 m) and A. erinacea (540 m). Thus most dynomenids show some association
with corals but this link does not appear to be obligatory.
DIET. — Guts of nine dynomenid species, Dynomene hispida, D. praedator , D. filholi, D. pilumnoides ,
Hirsutodynomene spinosa, H. Ursula, Metadynomene tanensis , M. devaneyi, and Paradynomene tuberculata, were
examined (usually only one or two specimens for each species). The most common material found was sand grains
along with soft unidentifiable particulate organic material and. in some species, chopped chitinous fragments
which could have come from hydroids or perhaps other crustaceans. It seems likely that most of their food was
obtained by sieving organic fragments from the substrate, or in the case of coral inhabiting crabs, perhaps from
coral mucous. This is consistent with the presence of a well developed screen of stiff setae in the hollowed out
interior of the cheliped fingers and the setose palps of the third maxillipeds. The chopped chitinous fragments
suggest that some food might be also obtained by direct grazing. Based on the shape of the chelae, BALSS (1938)
550
C.L. McLAY
suggested that dynomenids consume coral polyps in the same way as, for example the xanthids, Chlorodiella and
Chlorodopsis , but this is not supported by the stomach contents. Feeding behaviour has only been observed for
D. praedator which seems to obtain most of its food by deposit feeding in sand. The gut contents of the other
dynomenids would be consistent with feeding behaviour similar to that of D. praedator.
Reproductive Strategy. — The concept of reproductive strategy includes size at sexual maturity,
maximum size, relationship of egg numbers to female size, egg size, and extent of the reproductive season. The
extremes of this strategy are brooding or broadcasting progeny. Some information about these characteristics are
available for all the dynomenid species except Metadynomene crosnieri , and Acanthodromia marganta. Egg size is
a useful indicator of whether the species has direct or indirect development. In terms of maximum body size the
dynomenids fall into two groups: small species whose maximum CW is less than 20 mm {D. hispida,
D. praedator , D. filholi , D. pugnatrix , Acanthodromia erinacea and A . margarita ), and larger species whose
maximum CW is greater than 20 mm (D. pilumnoides , Hirsutodynomene spinosa , H. Ursula , M. devan eyi,
M. tanensis , M. crosnieri, and Paradynomene tuberculata). For all species males have on average a CW 19%
larger. Size at sexual maturity is roughly related to maximum size: for the small species sexual maturity occurs at
5.8 - 8.0 mm CW while for the large species it is at 9.5 - 13.0 mm CW. Egg numbers increase logarithmically
with CW with a brood size of 30 to 900 eggs for small species and 120 to 3800 for large species. Mean egg
diameter for the small species is 0.46 mm, for large species 0.51 mm and for all species together it is 0.49 mm.
These egg sizes suggest that all dynomenids have indirect development with planktonic larvae, i.e. a broadcast
strategy. The only larvae reported arc from Acanthodromia erinacea but these were dissected from eggs (RlCE,
1981). No dynomenid larvae have ever been reported from plankton collections. Only a limited amount of
information about the timing of reproduction is available. Despite the fact that dynomenids are tropical crabs all of
the species for which there are adequate sample sizes have their reproduction confined to only part of the year.
D. hispida and D. praedator (from the Indo-Pacific) are ovigerous from January to July, D. filholi (from the
Atlantic) is ovigerous from May until December, D. pilumnoides from February to September, H. Ursula from
April to December, and P. tuberculata from October to February (all Indo-Pacific species). Therefore larvae could
be expected in the plankton in any month, but the average reproductive period for all species is only 6.7 months.
Amongst the dynomenids we do not find species which have unusually large eggs, such as are seen in
homolodromiids (see GuiNOT, 1995), and a small number of dromiids (see McLay, 1993). The modal size class
for dromiid eggs is within 0.7-0.8 mm diameter, and direct development (brooding) has been found in
Dromidiopsis globosa (Lamarck, 1818), Austrodromidia octodentata (Haswell, 1882), and Stimdromia lateralis
(Gray, 1931) all of which have eggs >1.0 mm diameter. Homolodromiids have large eggs which are 2. 0-2. 5 mm
diameter and brooding is known in Dicranodromia nagaii Guinot, 1995. When we examine these features in
relation to the hypothesized sister group relationships of these families (see Fig. 15), it becomes obvious that the
ancestral condition must have been females carrying small numbers of very large eggs (probably brooding the
young), as in the homolodromiids, and the derived condition must be females carrying much larger numbers of
smaller eggs (broadcasting the young). Viewed in this way, the dynomenids are the most derived group, having the
smallest eggs with indirect development, and the dromiids are intermediate. Judging by their egg size, most
dromiids must have indirect development (the development of more than ten species is known) with only a few
retaining the ancestral condition of large eggs. One Australian dromiid, Haledromia bicavernosa (Zietz, 1887) has
the largest eggs known for any brachyuran i.e. 2.8 mm diameter. It is interesting to note that all dromiids known
to have direct development live in Australian waters. Also the endemic dromiid genera, Dromidia Stimpson, 1858,
Exodromidia Stebbing, 1905, Pseudodromia Stimpson, 1858, and Speodromia Barnard, 1947, from South Africa
all have large eggs, implying direct development, although their mode of development is unknown (McLay,
1993: 159). Therefore we could hypothesize that the common ancestor shared by the dynomenids and dromiids also
had a reproductive strategy of a small number of large eggs. If this is true then the broadcast strategy, seen in
dynomenids and most dromiids, must have evolved independently. This interpretation is no doubt contrary to
accepted wisdom, but it is the most parsimonious conclusion.
Depth Distribution. — The greatest diversity of dynomenids occurs in the 0-50 m depth range. Seven
species from three of the genera, Dynomene (4), Hirsutodynomene (2) and Paradynomene (1) arc shallow water
Source : MNHN, Paris
FAMILY DYNOMENIDAE
551
inhabitants (see Fig. 33). Four species (D. hispida , D. praedator , //. spinosa , and //. Ursula) have been collected
from the lower intertidal range, and between 25 and 50 m, six species are known. Between 50 and 400 m all five
genera are represented with no more than five species at any one depth interval. Below 400 m the number of
species begins to decline with two species at 500 m and none at 550 m. The species of Dynomene and
Parody nomene occur from 0-400 m, those of Hirsutodynomene are shallow water inhabitants (<100 m), and
species of Metadynomene and Acanthodromia all inhabit deeper waters down to a maximum depth of 540 m.
D. pugnatrix , M. devaneyi and M. crosnieri are only known from a few specimens and so their depth range is
uncertain. The average depth range (maximum - minimum) for all the more common species (n = 10) is 196 m.
The bathymetric distribution of dynomenids is very similar to that of the dromiids (see McLay, 1993) where
the majority of species are found in shallow waters from 0 to 150 m, with a maximum of around 500 m. By
contrast most of the homolodromiids are found in depths of 300 to 900 m, with a maximum of 1330 m (see
GuiNOT, 1995), and the homolids are mostly found in depths from 200 to 1500 m, with a maximum of 2200 m
(see GuiNOT & Richer de Forges, 1995). So we have a depth "zonation" of these three families with
dynomenids + dromiids in shallow water, followed by homolodromiids and lastly the homolids at the greatest
depths. Thus the hypothesized sister group relationship of dynomenids and dromiids is supported by the depth
distribution data which shows that these two families share the same "habitat”. Furthermore, if these two families
shared a common ancestor with the homolodromiids, as is hypothesized above for the Dromiacea. then it would
seem that dynomenids and dromiids evolved from ancestors which must have lived in deep waters (approx. 300-
900 m). Similarly, if the Dromiacea and Archaeobrachyura shared a common ancestor then it must also have lived
in deep water, perhaps at even greater depths than the dynomcnid - dromiid ancestor, with descendants radiating
into very deep water. Thus, the dynomenids and dromiids are families of the continental shelves while the
homolodromiids and archacobrachyurans are inhabitants of the continental slopes.
Fig. 33. — Depth distribution of dynomenid species. The "?" indicates an uncertain depth range because of a very small
sample size.
The hypothesis that dynomenids and dromiids represent shallow-water evolutionary radiations from ancestors
living in deeper water is similar to the hypothesis put forward by GEORGE and Main (1967) to explain the
Cretaceous evolutionary radiation of the Palinuridae which is supported by an analysis of phyllosoma characters of
the whole Palinuroidea by BAISTRE (1994).
Source :
552
C. L. McLAY
Biogeography. — Only two dynomenid species, representing two genera, are known from the Atlantic
Ocean: Dynomene filholi and Acanthodromia erinacea. Both of these are insular species (Figs 37, 40). /\. erinaceci
is restricted to the Caribbean area and D. filholi to the South Atlantic. In overall appearance, their obvious Indo-
Pacific counterparts are A. margarita and D. pilumnoides respectively.
The species of Acanthodromia no doubt originated from a tethyan ancestor and spread to seas that eventually
became the Atlantic and Caribbean, perhaps as early as the Upper Jurassic (145 mybp). This dispersal could have
been as late as the Middle Miocene (25 mybp). when connections between these two oceans were severed, but an
earlier date seems more likely because even by Palaeocene times (65 mybp) the Atlantic was already well formed
and the Caribbean isolated (Howarth. 1981). Therefore these two species could have been separated at least since
the Palaeocene (or at latest the Middle Miocene). The remarkable thing about A. erinacea and A. margarita is how
similar they are because there has scarcely been any divergence in their morphology (see Discussion under
A. erinacea).
Similarly, it seems to be a reasonable assumption that the ancestor of Dynomene filholi was a Tethyan crab
which also gave rise to D. pilumnoides. A southern colonization route for these crabs could have been open as
early as the Upper Cretaceous (90-80 mybp) or sometime later. At present there does not seem to be a dispersal
route via the Cape because it is blocked by the local Agulhas oceanic circulation pattern. This self-contained
circulation pattern seems to have been in existence for a considerable time because there is a suite of endemic
South African dromiid genera and species (see McLay, 1993) which have been isolated perhaps since the Upper
Cretaceous or Palaeocene (65 mybp). This interpretation would imply that colonization of the Atlantic by
Dynomene must have been during the late Mesozoic or very Early Tertiary. Again there are many features in
common between D. filholi and D. pilumnoides (see Discussion under D. pilumnoides).
Eleven species, representing all five genera, are known from the Indo-Pacific region. These species vary in the
extent of their distribution, but all of them lie within the limits of 40°N and 40°S. Within the genus Dynomene
two species (D. hispida , and D. praedator) are very widespread occurring from the coast of Africa to French
Polynesia in the east (Figs 34 & 35). D. pilumnoides is distributed over almost the same region except that it
only extends as far east as Hawaii (Fig. 36). The other species, D. pugnatrix , has a very limited distribution
restricted to the vicinity of Madagascar (Fig. 37). In the genus Hirsutodynomene one species, H. spinosa , has a
similar distribution to D. hispida and D. praedator while the other species, H. Ursula, is restricted to the Pacific
side of Central America (Fig. 38). The distributions of these two sister species do not overlap. It may well be that
H. ursula is of quite recent origin, perhaps not colonizing the eastern Pacific until after the formation of the
isthmus of Panama, because otherwise it would be reasonable to expect it to occur in the Caribbean as well as the
Pacific. Two of the species of Metadynomene are very rare: M. crosnieri is only known from the type locality,
north of Madagascar and M. devaneyi is only known from Hawaii and the Marquesas Islands. The third species,
M. tanensis , is common in the eastern Pacific and also occurs in French Polynesia (Fig. 39). While one species
of Acanthodromia, A. erinacea. lives in the Caribbean region, the other, A. margarita. has been recorded from the
Andaman Sea and Japan (Fig. 40). The last genus, Paradynomene , has until now been known only from the
eastern Pacific but I report herein a record of a specimen of P. tuberculata from the Gulf of Aden. Thus
Paradynomene is an Indo West Pacific genus (Fig. 40). Given the reproductive strategies outlined above, the large
geographic ranges of dynomenids is not unexpected. All dynomenids probably have planktonic larval stages.
The dynomenid fauna of insular Indo-Pacific localities is drawn from a suite of species which includes (in
decreasing order of frequency of occurrence): Dynomene praedator , D. hispida , Hirsutodynomene spinosa ,
D. pilumnoides. Metadynomene tanensis , Paradynomene tuberculata. H. ursula. and M. devaneyi. Almost without
exception wherever D. praedator is found, so is D. hispida. The largest number of species is found in Japanese
waters (6), followed by New Caledonia, Madagascar and French Polynesia (5). and Hawaii (4), while Mauritius,
Taiwan, Mariana Ids, Cocos Keeling Ids. Marshall Ids (Eniwetak) have 3 species. All other localities have only 1
or 2 species, usually D. praedator and/or D. hispida.
Although there are two shared genera ( Dynomene and Acanthodromia) in the Atlantic, there are no dynomenid
species which occur in both the Atlantic and Indo-Pacific regions. It can be safely assumed that the two Atlantic
species are derived from Indo-Pacific stocks and that their origin could have been as late as about 25 mybp before
the destruction of the Tethyan connection between these two oceans (Adams, 1981), although they could have
Source : MNHN , Paris
FAMILY DYNOMENIDAE
553
Fig. 34. — Geographic distribution of Dynomene hispida Guerin-Meneville. 1832.
Fig. 35. — Geographic distribution of Dynomene praedcitor A. Milne Edwards, 1879.
FIG. 36. — Geographic distribution of Dynomene pilumnoides Alcock, 1900.
Source : MNHN. Pans
554
C. L. McLAY
Fig. 37. — Geographic distribution of Dynomene pugnatrix dc Man, 1889 {■), and D. filholi Bouvicr, 1894 <•(
30° 60° 90° 120° 150°E 180° 150°W 120° 90°
Fig. 38. — Geographic distribution of Hirsutodynomene Ursula (Stimpson, 1860) (■) and H. spinosa (Rathbun, 1911) (•).
Fig. 39. — Geographic distribution of Metadynomene devaneyi (Takeda, 1977) (A). M. lanensis (Yokoya, 1933) (•). and
M. crosnieri sp. nov. <■).
Source : MNHN, Paris
FAMILY DYNOMENIDAE
555
FlG. 40. — Geographic distribution of Acanthodromia erinacea A. Milne Edwards, 1880 (•), A. margarita (Alcock, 1899)
(■). and Paradynomene tuberculata Sakai, 1963 (A).
been separate species much earlier. There does not seem to be any obvious reason why H i rs utody nomen e ,
Metadynomene and Paradynomene should not have become established in the Atlantic, unless these genera are of
more recent origin. In the Indian and Pacific Oceans there are eight and nine species respectively, five of which are
shared. All five genera have representatives in both oceans. Dynomenids are undoubtedly an ancient group of
crabs.
Fig. 41. — Longitudinal distribution of the Dynomcnidae.
Source : MNHN. Paris
556
C. L. McLAY
A similar picture emerges from our limited knowledge of the distribution of the Homolodromiidae (GUINOT,
1995) and the Homolidae (Guinot & Richer de Forges, 1995). The Atlantic and Indo-Pacific share genera but
not species. However, the species of both Hornolodromia and Dicranodromia have quite localized distributions
which may be a consequence of their tendency to have direct development. Dynomenid distribution is more like
that of the Homolidae where some genera are restricted to the Indo-Pacific and the species are widely distributed.
A similar pattern is found amongst the Dromiidae, although this family is a mixture of widespread genera along
with groups of genera endemic to a relatively small area. e.g. South Africa, or Australia (see McLay, 1993).
The relationship between dynomenid diversity and longitude shows that the highest diversity is found in the
Indo-West Pacific (Fig. 41). Diversity is lowest (0-2 species) in the Eastern Pacific and the Atlantic, and highest
(6-7 species) in East African and Japan/Philippines/New Caledonian latitudes. To some extent, this pattern
probably reflects the amount of collecting that has been done.
ACKNOWLEDGEMENTS
Many people have made this project possible by kindly loaning material from their collections and providing
answers to my questions. I would to particularly thank Michael TURKAY, Forschungsinstitut Scnckcnberg,
Frankfurt; Charles FRANSEN, Nationaal Natuurhistorisch Museum, Leiden; Daniele GUINOT, NGUYEN NGOC Ho,
Michele DE Saint Laurent, and Sylvie Secretan, Museum national d'Histoire naturelle, Paris; Joseph
POUPIN. Laboratoire d'Oceanographie. Ecole Navale du Poulmic, Brest; Niel BRUCE, Zoologische Museum,
Copenhagen; Marco VANNINI, Museo Zoologico de "La Specola", Firenze; Oliver COLEMAN, Museum fur
Naturkunde, Institut fur Systematische Zoologie. Berlin; Paul Clark, The Natural History Museum, London;
Austin Williams and Ray MANNING. Smithsonian Institution, Washington; David WiEDNER, The Academy of
Natural Sciences, Philadelphia; Hans KUCK, Natural History Museum of Los Angeles County; Ardis JOHNSTON,
Museum of Comparative Zoology, Harvard; Tin-Yam Chan, National Taiwan Ocean University, Taiwan;
Masayuki Osawa, Tokyo University of Fisheries; Seiji Nagai, Wakayama-ken. Japan; Roy Kropp, Battelle
Ocean Sciences, U.S.A.; Gustav Paulay, Guam; Lu Eldredge, Bishop Museum, Honolulu; Yang Chang
Man, Zoological Reference Collection, University of Singapore; Diana Jones, Western Australian Museum,
Perth; Alexander BRUCE and Peter Davie, Queensland Museum, Brisbane; Penny BERENTS, Australian Museum,
Sydney, and Bertrand Richer DE Forges, Institut fran^ais de Recherche scientifique pour Ic Developpement en
Cooperation (ORSTOM), Noumea.
Peter Ng, University of Singapore, provided invaluable assistance with the literature from China, and
Li Dai Qin and Wu Rongmei, University of Canterbury, kindly translated these papers. Keiji Baba, Kumamotu
University, and Hiroshi SUZUKI, Kagoshima University helped me by translating the Japanese literature.
Volker Neuman, Forschungsinstitut Senckenberg, Frankfurt and Franz DiTZ, University of Canterbury willingly
translated German papers for me. The photos in this paper were made by Jacques Rebiere, Museum national
d'Histoire naturelle, Paris, and Terry Williams, University of Canterbury. I also acknowledge the help of
Fran^oise Theureau, Museum national d'Histoire naturelle, Paris, who expertly prepared the figures.
Jan McKenzie, University of Canterbury gave invaluable help with the scanning electron microscope.
Gerhard Scholtz, and Stefan RICHTER, Institut fur Biologie, Humboldt Universitat zu Berlin helped to clarify
my thinking about the phylogeny of podotreme crabs and provided valuable comments on the manuscript.
John Hoover and Ron Holcom kindly collected Hawaiian specimens of Dynomene praedator and provided me
with a video tape of their behaviour. The manuscript was greatly improved by the comments of Professors
D. Guinot, and L. B. Holthuis.
Finally, all the efforts of the above persons would have been in vain if it were not for the tireless efforts and
enthusiasm of Alain Crosnier, ORSTOM and Museum national d’Histoire naturelle. Paris, to ensure the
completion of this project.
Source : MNHN. Pans
FAMILY DYNOMENIDAE
557
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Source : MNHN. Paris
FAMILY DYNOMENIDAE
567
INDEX
Family and genus names are given in capital letters, species names in italics. Bold family, genus and species
names indicate that die taxa are treated in detail.
Bold numbers indicate die pages where the subject is treated in detail.
Abdomen 439; 455
locking mechanism 455; 462; 539
sexual dimorphism 456
uropods 439
Acanthodromia 432; 433; 438; 439; 441; 443;
447; 449; 456; 464; 466; 468; 469; 534
A. erinacea 431; 432; 433; 442; 443; 453;
469; 470; 535
A. margarita 432; 433; 470; 539
Acanlhodromies 449
Achelata 466
Albuneidae 469
Alpheus lottini 480
Anomola 454; 466
Antennae 438; 441
Antennules 438; 441
Anterolateral teeth 439
variation 483; 501; 530; 544
Arachnodromia 468
Archaeobrachyura 463; 551
Astacus astacus 528
Austrodromidia octodentata 550
Ax ius Ion gi spina 499
Betaeus 454
Blepharipoda 469
Bopyridae
Gigantione 488
Brachynotus gemmellari 528
Brachyurites rugosus 434
Branchial formula 443; 476; 493; 500: 524; 526;
538; 546
Callianassa 466
Callinectes sapidus 528
Cambarus propinqnns 528
Camouflage behaviour 463; 466
Carapace 439
grooves 435; 439
Care inns maenas 528
Chelipeds 438; 449
Chionoecetes opilio 459; 528
Conchoecetes 447
Corals 549
ACROPORA 478; 48 1 ; 488; 5 1 0
CORALLIUM 520
Favia 481
Goniastrea 481
OULOPHYLLIA 481
Pocillopora 480; 488; 514
Porites 481; 488; 514
red coral 492
Seriatopora 48 1
Crista dentata 438; 443
Cronins tnmidnlns 444
Cryptodromia 543
C. canaliculata 5 1 0
Cryptodromiopsis
C. antillensis 467; 488
C. larrabnrei 444
Cyamocarcinns angnstifrons 434
Cyclodorippidae 463; 466
Cyclothyrens strambergensis 434
Cymonomidae 463
Depth distribution 478; 486; 492; 499; 502; 509;
514; 525; 533; 538; 542; 547; 550
DlCRANODROMIA 460; 463; 464; 468; 534; 556
D. felderi 460; 462
D. mahieuxii 447; 493
D. nagaii 550
D. ovata 447; 538
Diet 480; 494; 500; 510; 515; 527; 549
DROMIA 460; 464; 468; 543
D. erythropns 444
D. personata 44 1
D. wilsoni 534
Dromiacea 463; 464; 551
ancestor 463
Dromidia 468; 550
D. larrabnrei 444
D. spongiosa 433
Dromidiopsis 454; 493; 549
D globosa 550
D. edwardsi 467
Dromiens 468
DROMIIDAE 439; 441; 443; 444; 447; 454; 455;
460; 463; 464; 466; 468; 488; 493; 494;
538; 543; 548; 550; 551; 556
ancestor 466
Dromiopsis 434
DYNOMENE 433; 434; 439; 441; 443; 444; 449;
453; 455; 463; 471; 543
D. actaeiformis 432; 433; 494
D. depressa 433
D. devaneyi 432; 433; 530
Source :
568
C. L. McLAY
D. filholi 432; 433; 441; 442; 443; 447;
454: 460; 471; 488; 549
D. granulobata 432; 433; 473; 479
D. hispida 43 1 ; 432; 433; 438; 441; 443;
447; 453; 471; 473; 499; 505
D. hispide 473 '
D. huangluensis 432; 433; 481; 486
D. latreillii 432; 433; 473; 479
D. margarita 432; 433; 539
D. pilumnoides 432; 433; 441; 442; 447;
460; 471; 494
D. platyarthrodes 432; 433; 469
D. praedator 432; 433; 438; 441: 442; 447;
453; 456; 460; 470; 479; 480; 481; 521;
526
D. pugnatrix 433; 441; 447; 449; 471; 500
D. pugnatrix brevimana 432; 433; 500
D. sinense 48 1
D. sinensis 432; 433; 486
D. spinosa 432; 433
D. tanensis 432; 433; 526
D. tenuilobata 432; 433; 481 ; 486
D. ursula 432; 433; 510
DYNOMENIDAE 434; 444; 449; 464; 467
Dynomeninae 467
Ebalia tuberculata 528
E las so pod its stellatus 452
EODROMIA 460; 464
Epigodromia 447; 543; 549
E. gilesii 460; 548
Epipods 443
Eubrachyura 466
Evolution 449; 494; 539; 543; 550; 551; 552
convergence 549
Exodromidia 460; 550
Feeding behaviour 488; 550
Fossils 434; 441; 535
Gemmellarocarcinus loerentheyi 434
Geographic distribution 478; 486; 492; 498; 502;
509; 514; 515; 520; 525; 533; 538; 542;
547; 552
Gill cleaning 443; 447; 463
Gills 438; 443; 444; 447; 449; 479; 515; 548
evolution 466
Glyptodynomene alsasuensis 434
Graptocarcinus texanus 434
Gynandromorphs 527
Habitat 549
Haledromia bicavemosa 550
Hirsutodynomene 433; 434; 439; 441; 443;
444; 447; 449; 453; 455; 504
H. spinosa 433; 441; 460; 470; 473
H. ursula 433; 441; 460; 467; 510
Homarus
H. americanus 528
H. ganimarus 528
H. vulgaris 443; 447
Homola andamanicus 499
HOMOLIDAE 443; 454: 463; 466; 467; 468; 556
HOMOLODROMIA 460; 464; 556
H. kai 467
H. paradoxa 447; 453; 538
HOMOLODROMIlDAE 441; 443; 447; 455; 460;
463; 464; 466; 550; 551
HOMOLOGENUS 454
Hyas coarc tat us 528
Hypobranchial setae 444; 526; 548
Hypoconcha 447
H. arcuata 467
H. sabulosa 488
H. spinosissima 488
Identification key 469
JASUS
J edwardsii 528
J. frontalis 528
Krangalangia 466
Larvae 432; 469; 538; 550
Latreillia 466
Latreillidae 463; 466
Lauridromia 454; 493; 549
Leptomithrax longipes 528
Lucifera 528
Lyreidus 466
Maxillothrix 471
M. actaeiformis 432; 433; 494
Measurements 435
abdomen 435; 439
carapace 435
chelipcd propodus 435
pereopods 435
setae 435
Metadynomene 433; 434; 439; 441; 443; 449;
453; 455; 516
M. crosnieri 433; 447; 470; 530
M . devaneyi 433; 444; 447; 470; 517
M. tanensis 433; 441; 444; 447; 454; 456; 460;
467; 470; 481; 487; 493; 521
Metapenaeus monoceros 528
Mithracites 543
Mouthparts 438; 441; 487; 493
maxillipeds 443
Source :
FAMILY DYNOMENIDAE
569
Nephrops norvegicus 528
Nepinnotheres androgynus 528
Orbits 441
Palaemon 454
Palaeodromites octodentatus 434
Palinuridae 454; 551
PARADYNOMENE 433; 439; 443; 449; 455; 469;
542
P. tubercalata 433; 439; 441; 443; 444;
447; 454; 460; 467; 470; 493; 543
Penis 460
Pereopods 438; 449; 467
article fusion 449; 538; 548
dactyl spine 493; 527; 549
sexual dimorphism 453
vestigial 439; 449; 453; 454; 467; 493; 500;
515; 527; 538; 548
Petalomem pulchra 488
Phylogeny 462
brachyuran monophyly 463
Dromiacea 466
dromiacean monophyly 463
homolodromiid monophyly 464
Podotremata 466
PlLUMNUS 514
Plagiophthalmus 535
Pleistcicantha moseleyi 528
Pleopods 439; 456; 515
female 460
gynandromorphs 456
male first 458
male function 459
male second 459
rudimentary 460; 499; 527
structure 460
tegumental gland 459
Podotremata 463
ancestor 466
monophyly 464
paraphyly 466
PORCELLANIDAE 454
POUPINIIDAE 463
Procambarus clarkii 528
Prosopidae 467; 535
PSEUDODROMIA 550
Ranina 466
R. ranina 459
Raninidae 463; 466
Raninoides 466
Rathbunopon 543
Relative growth 527
Reproductive strategy 478; 486; 492; 499; 502;
509; 514; 525; 538; 547; 550
Scaphognathite setae 444; 499; 539
SCYLLARIDAE 454
SCYLLARUS 466
Setae 435; 441; 443; 449; 480; 493; 515; 526;
548
setules 441 ; 444
Sexual dimorphism
abdomen 456; 527
cheliped 449; 527
fifth pereopod 453; 480; 487; 493; 500; 515;
527; 538; 549
pleopods 456
rudimentary pleopods 460
sternal sutures 7/8 455
uropods 462
Speodromia 469; 550
S. platyarlhrodes 433
Sperm duct 460
SPHAERODROMIA 460; 463; 464; 521
Stephanometopon granulatum 434
Sternal Sutures 454
Sternal sutures 7/8 455
Stimdromia
S. lamellata 460; 548
S. lateralis 467; 550
Synalpheus charon 480
Tegumental gland 480
Thaumastochelidae 454
Trachynotocarcinus sulcatus 434
Tunedromia 454; 493; 549
Tymolus 466
Uropods 462
sexual dimorphism 462
Xanthidae 467; 550
Xeinostoma 466
Source : MNHN. Paris
IULTATS DES CAMPAGNES MUSOR.STOM. VOLUME 20 — RESULTATS DES CAMPAGNES MUSORSTOM, VOLUME 20 — RESULTATS DE
Pisces Anguilliformes: Deepwater snake eels
(Ophichthidae) from the New Caledonia region,
Southwest Pacific Ocean
John E. McCOSKER
California Academy of Sciences
Golden Gate Park, San Francisco. California 941 18
USA
ABSTRACT
This paper reports upon the snake eels collected by trawl during the 1985 Musorstom 4 New Caledonia Expedition.
The 14 specimens comprised five new ophichthid taxa which are described herein: Ophichthus exounis sp. nov. from 400-
520 m (also from Fiji); O. genie sp. nov. from 430-500 m (also from Maldives); O. mystacinus sp. nov. from 450-580 m;
Yirrkala insolitus sp. nov. from 59 m; and Rhinophichthus penicillatus gen. et sp. nov. from 435 m. The dorsal fin
location of the new species of Yirrkala provides an expanded character state within the genus. Rhinophichthus differs
from other generalized ophichthins in its very conical snout, and includes Ophichthys unicolor Regan from South Africa.
The affinities of the new taxa are with Indo-west Pacific ophichthids, however the collections are too few to allow
significant conclusions about the bathyal ophichthid fauna of the region.
RESUME
Pisces Anguilliformes: Poissons-serpents d'eau profonde (Ophichthidae) de la region neo-
caledonienne (Pacifique sud-ouest).
Les 14 specimens de poissons-serpents (famille des Ophichthidae) recoltes en 1985, au cours de la campagne de
chalutage prolond Musortom 4, au large de la Nouvelle-Cal£donie, sont etudies et cinq espfeces nouvelles sont decritcs :
Ophichthus exourus sp. nov. recoltee entre 400 et 520 m de profondeur (et aussi aux Fidji); O. genie sp. nov. recoltee
entre 430 et 500 m (et aussi aux Maldives); O. mystacinus sp. nov. recoltee a 450-580 m; Yirrkala insolitus sp. nov.
recoltee & 59 m et Rhinophichthus penicillatus gen. el sp. nov. recoltee a 435 m. La position de la nageoire dorsale de la
nouvelle especc de Yirrkala c onstitue un etat evolue de ce caractere. De mcme le genre Rhinophichthus . par son muscau
tres conique, semble particulierement 6volue; il inclut Ophichthys unicolor Regan decrite d'Afrique du Sud. Ces nouveaux
taxa ont des allinites indo-pacifiques, cependant la collection est trop reduite pour permettre des conclusions
significatives sur la faune ophichthienne bathyale de la region.
McCOSKER, J.E., 1999. — Pisces Anguilliformes: Deepwater snake eels (Ophichthidae) from the New Caledonia
region, Southwest Pacific Ocean. In: A. Crosnier (ed.). Resultats des Campagnes Musorstom. Volume 20. Memoires du
Museum national d'Histoire naturelle : 180: 571-588. Paris ISBN 2-85653-520-8.
Source : MNHN, Paris
572
J.E. McCOSKER
INTRODUCTION
The 1985 Musorstom oceanographic survey of the bathyal fauna of New Caledonia resulted in many
interesting ichthyological discoveries. The specimens of ophichthid eels were captured from depths well below
those normally inhabited by most ophichthid species. They were forwarded to me by Bernard SERET for
identification and, although few in number - only 14 representing five taxa - the quality far exceeded the quantity,
in that those specimens comprise three genera (one undescribed) and five undescribed species of ophichthid eels.
The affinities of those new species seem to lie with Indo-west Pacific congeners. The new genus and its two
species cannot yet be identified with any known sister group.
TAXONOMY
The snake eels and worm eels of the anguilliform family Ophichthidae are the most diverse and speciose of true
eels (McCOSKER et al. , 1989). The more than 260 species distributed among 57 genera inhabit all tropical oceans
and seas. They occupy habitats ranging from the intertidal zone to depths of 750 m or more, living amongst coral
and rock reefs and sand and mud bottoms, some entering estuaries and rivers, and others having adapted to life in
midwater. Ophichthids are of limited direct economic importance, however all seem to be edible and several large
species are taken as a bycatch of trawl and trap fisheries and are consumed. In that they arc abundant in many
habitats, it is reasonable to assume that they are of indirect importance to commercial fisheries in their role as both
predator and prey.
A systematic revision ot the family Ophichthidae has yet to be achieved on a worldwide basis, although
McCOSKER (1977) provided a comprehensive treatment of the genera. Several species are so widespread in
distribution that they occupy both the Indian and Pacific oceans, however none are worldwide and others seem to be
limited to island archipelagos or to limited coastal zones. Faunal treatments of ophichthids that may be helpful for
the identification of genera and species have been prepared for: the western Atlantic (McCOSKER et al., 1989);
southeast Africa (McCOSKER & Castle, 1986); Indian Ocean and Red Sea (Smith. 1962); Hawaii (McCOSKER,
1979); eastern Pacific (McCOSKER & ROSENBLATT, 1995); and Australia (McCOSKER in preparation).
Deepwater trawling and trapping in recent years has resulted in the collection of many new species of
ophichthids. Because of their secretive, burrowing behavior, many species are able to avoid capture, particularly in
deep water. If the success of the MUSORSTOM project is any indication, it is likely that many species will be
discovered as a result of future deepwater surveys.
MATERIALS AND METHODS
All of the specimens collected from New Caledonia resulted from the 1985 MUSORSTOM 4 Expedition.
The first half of that expedition, aboard R. V. " Vauban ", investigated the "Grand Passage" between the northern
lagoon of the mainland and Surprise Atoll. A total of 57 stations were sampled with beam trawls (stations "CP"),
otter trawls (stations "CC"), Warren dredges, and Charcot dredges between 34-720 m depth (see Richer de
FORGES. 1990). The otter trawl had a 15 m headrope and is known as a "shrimp trawl"; the beam trawl was 4 m
wide and fitted with 20 mm mesh, lined with 5 mm mesh for the bag. The habitat, described by Richer de
Forges & Bargibant (1985) and Richer de Forges (1990), is that of a "narrow sill situated at about 600 m
depth. The bottoms encountered were mostly rocky, but with some soft areas carpeted with pumice stones. At the
far extremity of the northern lagoon, sand with Halimeda was found as deep as 550 m and more. To the south of
the Mainland, New Caledonia extends in a gentle sloping valley from 200 to 500 m. Here the bottoms are of
sandstone slabs, relatively flat, and trawling is possible." These efforts resulted in numerous congrid eels but only
14 specimens of ophichthids, captured during five beam trawls and three otter trawls.
Source :
OPHICHTHIDAE FROM THE NEW CALEDONIAN REGION
573
Specimen measurements are straight-line, made either with a 300 mm ruler with 0.5 mm gradations (for total
length, trunk length, and tail length) and recorded to the nearest 0.5 mm, or with dial calipers (all other
measurements) and recorded to the nearest 0.1 mm. Body length comprises head and trunk lengths. Head length is
measured from the snout tip to the posterodorsal margin of the gill opening; trunk length is taken from the end of
the head to mid-anus; maximum body depth does not include the median fins. Head pore terminology follows that
ol McCoSKER et al (1989: 257) such that the supraorbital pores are expressed as the ethmoidal pore + pores in
supraorbital canal, /.<?., 1 + 3, and the infraorbital pores are expressed as pores along the upper jaw + those in
vertical part of canal behind eye (the "postorbital pores"), 4 + 2, in that frequently the last pore included along
the upper jaw is part of the postorbital series. Radiographic techniques are described in Bohlke (1989). Gill arch
examination was accomplished after removal and clearing and counterstaining with alcian blue and alizarin red dyes
(DiNGERKUS & Uhler, 1977). The mean vertebral formula (MVF) is expressed as the average of predorsal,
preanal, and total vertebrae (Bohlke. 1982). Vertebral counts (which include the hypural) are taken from
radiographs. Institutional abbreviations follow the Standard Symbolic Codes for Institutional Research Collections
in Herpetology and Ichthyology (Leviton et al ., 1985). Other abbreviations are as follows: DFO = dorsal fin
origin; HL = head length; IO = interorbital width; TL = total length.
RESULTS
Key to the species of deepwater MUSORSTOM ophiehthid eels
from the New Caledonia region*
1 . Pectoral fins present; body moderately elongate, its depth < 45 times in total length .
. 2
— Pectoral fins absent; body elongate, its depth > 45 times in total length .
. Yirrkala insolitus
2. Snout conical, sharply pointed; jaw teeth uniserial ... Rhinophichthus penicillatus
— Snout rounded at tip, not conical and extended; maxillary teeth biserial . 3
3. Pectoral lin short, paddle-shaped, about 1.3- 1.4 in upper jaw; posterior margin of orbit
above rictus of jaw; teeth of mandible uniserial . Ophichthus exourus
— Pectoral fin elongate, nearly as long as jaw; posterior margin of orbit well in advance of
rictus or jaw; jaw teeth biserial . 4
4. Dorsal fin origin about mid-trunk; body depth 23-28 times in total length .
. Ophichthus my st acinus
— Dorsal fin origin above pectoral fins; body depth 31-42 times in total length .
. Ophichthus genie
*This key concerns only those ophichthids captured during the Musorstom collections. It is not meant to be
comprehensive in that the numerous shallow-water ophichthids from New Caledonia are not included.
FAMILY OPHICHTHIDAE
Diagnosis. — Body elongate, snake-like in large species, worm-like in some smaller species; cylindrical
anteriorly, generally laterally compressed posteriorly. Snout pointed to blunt, mouth either terminal or inferior.
Jaw teeth variable, from molariform to conical to large and fanglike; from single to multiple rows, a few species
without teeth on vomer. Nostrils widely separated, the posterior inside the mouth or penetrating through a valve or
hole in or above the upper lip, the anterior usually in a short tube. Gill openings midlateral to entirely ventral.
Source
574
J E. McCOSKER
a constricted opening in one subfamily (Myrophinae), variable in the other (Ophichthinae). Branchial region
supported by numerous overlapping branchiostegal rays. Gill arches uniquely developed with first epibranchial
connected by a continuous cartilaginous strap to the second infrapharyngobranchial; no more than first
basibranchial ossified; the third hypobranchial usually cartilaginous. Dorsal and anal fins, when present,
continuous around the caudal in one subfamily (Myrophinae) and absent, forming a hard finless point in the other
(Ophichthinae); pectoral fins present or absent; pelvic fins absent. Lateral line system extends onto head, the sides
connected through a frontal and a temporal canal. Coloration variable, from uniform (although usually darker
dorsally) to mottled, spotted, barred or striped.
REMARKS. — As currently recognized, the Ophichthidae comprise two subfamilies, the Myrophinae and the
Ophichthinae, and six tribes, the Myrophini, the Benthenchelyini, the Bascanichthyini, the Callechelyini, the
Sphagebranchini, and the Ophichthini (McCOSKER, 1977; McCOSKER et al., 1989). Ophichthid eels also differ
from related eel families in the following manner:
1) Congridae, which possess posterior nostrils at the level of the eye;
2) Muraenidae, which lack pectoral fins, possess small branchial openings, and have posterior nostrils at the
level of the eye;
3) Muraenesocidae, which possess posterior nostrils at the level of the eye, and have the vomer bearing a
median series of large teeth flanked by a row of small teeth;
4) Chlopsidae (previously the Xenocongridae), which usually have multiple rows of vomerine teeth and an
incomplete lateral line, limited to one or two pores in the branchial region.
Genus OPHICHTHUS Ahl, 1789
Innominado Parra, 1787: 96 (a junior synonym of Muraena ophis Linnaeus. 1758, non-binomial).
Ophichthus Ahl, 1789: 5 (type species Muraena ophis Linnaeus. 1758, by original designation. Improperly emended to
Ophichthys by other authors).
Ophis Turton, 1807: 87 (type species " O . niaculata ... Spotted Serpent. Shaw Zool., iv. p. 22 ... Bloch t. 154," = Muraena
ophis Linnaeus, 1758, by monotypy).
Cogrus Rafinesque, 1810: 62 (type species Cogrus maculatus Rafinesque, 1810, by monotypy).
Ophithorax McClelland. 1844: 212 (type species Ophisurus ophis Lacepede, 1800 = Muraena ophis Linnaeus. 1758 by
Jordan, 1919, as first reviser). ’
Centrurophis Kaup, 1856: 42 (type species Ophisurus spadiceus Richardson, 1844 = Ophichthys cephalazona Bleeker
1864, by Jordan, 1919, as first reviser).
Poeci locephalus Kaup, 1856: 43 (type species Poecilocephalus bonaparti Kaup, 1856, by monotypy).
Microdonophis Kaup, 1856: 43 (type species Microdonophis altipinnis Kaup, 1856. by monotypy).
Coecilophis Kaup. 1856: 44 (type species Ophisurus compar Richardson. 1844 = Ophisurus apica/is Bennett, 1830 by
monotypy). ' J
Scytalophis Kaup 1856: 46 (type species Scytalophis magnioculis Kaup, 1856 = Ophisurus gomesii Castelnau 1855 by
Jordan, 1919, as first reviser). ’ 3
Leptorhinophis Kaup, 1856: 46 (type species Ophisurus gomesii Castelnau, 1855, by Jordan 1919. as first reviser).
Cryptopterus Kaup, 1860: 1 1 (type species Cryptopterus puncticeps Kaup. 1860, by monotypy).
Uramchihys Poey, 1867: 256 (type species Muraena hauannensis Bloch & Schneider, 1801 = Muraena ophis Linnaeus
1758. by Jordan & Davis, 1891, as first revisers).
Parainyrus Gunther, 1870: 51 (type species Conger cylindroideus Ranzani, 1839. by Jordan & Davis 1891 as first
revisers).
Oxyodomichthys Poey, 1880: 254 (type species Ophichthys macrurus Poey, 1867 = Ophisurus gomesii Castelnau 1855
by original designation). *
Omochelys Fowler ,1918: 3 (type species Pisodonophis cruentifer Goode & Bean, 1896. by original designation*
described as a subgenus of Pisodonophis Kaup. 1856).
Syietor Jordan, 1919: 343 (type species Pisodonophis cruentifer Goode & Bean. 1895. by original designation).
Sty ie tor Jordan, 1919, in Jordan, Evermann & Clark, 1930: 86 (lapsus pro Syietor Jordan. 1919).
Acanthenchelys Norman 1922: 296 (type species Acanthenchelys spinicauda Norman. 1922, by original designation)
°0 fZX^ih^M^im]925'' 1 (SUbS"lU,e name for Cr>'P‘opterus Kaup, 1860, preoccupied; described as a subgenus
Source :
OPHICHTHIDAE FROM THE NEW CALEDONIAN REGION
575
Zonophichthus Whitley. 1930: 250 (type species Ophichthys cephalazona Bleeker, 1864, by original designation).
Gisenchelys Fowler, 1944: 188 (type species Ophicluhys zophochir Jordan & Gilbert, 1881, by original designation;
described as a subgenus of Ophichthus Ahl, 1789).
Syletophis Whitley, 1950: 44 (substitute name for Syletor Jordan, 1919, preoccupied).
Antobrantia Pinto, 1970: 13 (type species Antobrantia ribeiroi Pinto, 1970 = Muraena ophis Linnaeus, 1758. by
original designation).
DIAGNOSIS. — Moderately to very elongate ophichthid eels of the subfamily Ophichthinae. tribe Ophichthini,
with head and trunk shorter than tail; dorsal fin origin above or behind gill openings; pectoral fins present and
developed; snout and jaws moderately elongate; lips without numerous barbels or fringes; anterior nostrils opening
via a tube; posterior nostrils opening into mouth or along lower edge of lip; eye moderately developed; gill
openings lateral, elongate, nearly vertical and crescentic; teeth conical and numerous, but never caniniform; tail lip
a finless point; and coloration variable, often marked, but generally uniform and darker dorsally.
Remarks. — The genus Ophichthus (sensu lato ) is llte most speciose of ophichthids, with approximately
55 tropical and subtropical species worldwide. Several subgenera are recognizable within Ophichthus, though a
worldwide revision has yet to be attempted (McCoSKER, 1977).
Ophichthus exourus sp. nov.
Figs I -2a; Tabl. I
Material examined and Types. — 2 specimens.
New Caledonia. Musorstom 4: stn CP 178. I8°56.30'S, I63°12.90'E. 520 m, 18.09.1985: holotype, 590 mm TL
(MNHN 1995-425).
Fiji. Viti Levu. off Suva Barrier Reef, prawn irap set in 400 m, 28.08.1980: 1 paratype, 429 mm TL (CAS 89552).
OTHER MATERIAL EXAMINED. — Ophichthus brachynotoplerus: Madagascar, 1971: 1 paratype. 442 mm TL
(MNHN 1979-22); 1 paratype, 413 mm TL (MNHN 1979-23).
Ophichthys serpentinus: Cape of Good Hope, 1860: holotype, 495 mm TL (MCZ 9200).
Ophichthus mystacinus: holotype and paratypes. described below.
Fig. 1. — Ophichthus exourus sp. nov., holotype, 590 mm TL (MNHN 1995-425), New Caledonia. I8°56 30'S
163°12,90'E, Musorstom 4, stn CP 178. 520 m.
Source
576
J. E. McCOSKER
DIAGNOSIS. — A moderate length species of Ophichthus with the unique combination of characters: head and
trunk robust, tapering evenly to tail tip; tail 59-60% of total length; dorsal fin origin in anterior third of trunk;
pectoral fins small, paddle-shaped; jaw not elongate; eye large, above end of jaw; teeth conical, numerous and
small, biserial in maxillary, uniserial on mandible and vomer; coloration yellowish tan, brown dorsally, the fins
pale; and mean vertebral formula 20.5-60.5-176.5.
Description. — (Counts and measurements in mm of the holotype). Total length 590; head length 47.6;
trunk length 185.4; tail length 357; body depth at gill openings 20; body width at gill openings 17; body depth at
anus 15; body width at anus 14.5; tip of snout to dorsal fin origin 89; left pectoral fin length 12.7; base of left
pectoral fin 4.5; gill opening length 6; isthmus 6.7; snout 13.5; tip of snout to rictus of jaw 17.9; eye diameter
6.1; interorbital distance 6.7. Total vertebrae 176; predorsal vertebrae 20; preanal vertebrae 61.
Body moderately elongate, its depth behind gill openings 26-29 times and width behind gill openings
35-38 times in TL. Body cylindrical anteriorly, laterally compressed in tail region, tapering evenly from robust
head and trunk to much reduced tail tip. Head and trunk 2.5 times and head 12-12.5 times in TL. Head elongate,
conical when viewed from above, somewhat flattened dorsally. Snout pointed, conical from above. Lower jaw
reaches snout tip. Anterior nostrils in a short tube; posterior nostril in outer edge of upper lip and covered by a
Hap, appearing as a slit in the upper lip well before and beneath the orbit. A small but obvious protuberance in lip
between anterior and posterior nostrils. Eye large, its posterior margin slightly behind rictus of jaw.
Median tins moderately developed. Dorsal fin origin above anterior third of trunk and less than a head length
behind gill opening. Pectoral tins short and paddle-shaped. Median fins expanded before naked tail tip, lying within
a groove.
Cephalic pores (Fig. I) small and difficult to discern. Six mandibular, 2 preopercular, 1 ethmoidal +
4 supraorbital, 3 + 2 infraorbital, and single interorbilal and supratemporal pores. Nine lateral line pores above
branchial region. Lateral line pores of trunk and tail minute.
F,Gn2 ^ Schemal,c d,aSram of dentition of: A, Ophichthus exounis sp. nov., holotype, 590 mm TL (MNHN 1995-425)
^9552)€nie SP n°V ’ paralype' 337 mm TL (BPBM 34923 ); C. O. mystacinus sp. nov., paratype, 429 mm TL (CAS
Source :
OPHICHTHIDAE FROM THE NEW CALEDONIAN REGION
577
Teeth (Fig. 2a) small, nearly equal in size, conical, and separated at their bases. A small rosette of 7 teeth at
snout tip, followed by 15 neatly-aligned uniserial vomerine teeth. Maxillary teeth biserial, an outer row of
20-21 small teeth, flanked by 16 larger, more widely-spaced teeth. Lower jaw teeth uniserial, about 21-23 on each
side, with 4-5 inner teeth scattered on inner edge.
Body coloration in ethyl alcohol yellow, overlain on upper flanks with a dusting of microscopic brown spots
which are denser above the lateral midline. Median fins arise from pale longitudinal bands, the dorsal band
becoming darker in posterior half of body such that fin appears as a sharp white line along the dorsum. Fins
colorless except membrane of anal fin w'hich is darkened before the tail tip. Peritoneum black.
Size. — To 669 mm. The holotype, 590 mm, and the paratype are females with developing ova,
approximately 0.5 mm in diameter.
Etymology. — From the Greek, exourus , ending in a tapered point.
Distribution. — Known from the MUSORSTOM New Caledonia collections and from Fiji, between 450-
520 m depth.
Remarks and Comparisons. — Ophichthus exourus is within the subgenus Coecilophis Kaup
(cf McCosker, 1977). It appears most closely related to the deepwater species O. serpentinus Seale (1917)
from Madagascar, and O. mystacinus , described herein from deepwater off New' Caledonia. I have compared the
holotypes and paratypes of those species to the new' species, and find O. exourus to differ from
O. brcichynotopterus and O. mystacinus by having a shorter pectoral fin and uniserial (rather than biserial)
mandibular teeth. Both exourus and serpentinus have uniserial mandibular dentition, however they differ in their
vomerine dentition (strictly uniserial vs. biserial anteriorly, respectively) and in their vertebral numbers (176-177
vs. 165-167, respectively).
Ophichthus genie sp. nov.
Figs 2b-3; Tabl. 1
Material examined and Types. — 6 specimens.
New Caledonia. Musorstom 4: stn CP 170, I8°57.00'S. 163°12.60'E, 485 m. 17.09.1985: holotype, 230 mm TL
(MNHN 1998-43). — Stn CP 171, 18°57.80'S, 163°14,00’E, 435 m, 17.09.1985: 1 paratype, 217 mm TL (MNHN 1998-
44); 1 paratype 232 mm TL (CAS 89551); I paratype 206 mm TL (ANSP 174853). — Stn CC 201, 18°55.80’S.
I63°13,80'E, 500 m, 20.09.1985: 1 paratype. 196 mm TL (MNHN 1998-45).
Maidive Islands. Stn R3, 04°19'N, 72°55’E. 215 m, R. C. Anderson coll., 9.03.1991: 1 paratype 337 mm TL
(BPBM 34923).
OTHER Material EXAMINED. — Ophichthus kunaloa : Oahu, Hawaii. 1969: holotype. 440 mm TL (CAS 29136).
DIAGNOSIS. — A moderate length species of Ophichthus with the unique combination of characters: tail 55-
58% of total length; dorsal fin origin before pectoral tips; pectoral fins elongate, lanceolate; eye 2. 8-4. 2 times in
jaw length; teeth small, conical, biserial in jaws and uniserial on vomer; coloration pale, overlain on dorsal half
with fine brown punctations; and mean vertebral formula 14-57-142.
Description. — (Counts and measurements in mm of the holotype). Total length 230; head length 21.5;
trunk length 79.5; tail length 129; body depth at gill openings 7.3; body width at gill openings 4.8; body depth at
anus 5.9; body width at anus 4.5; tip of snout to dorsal fin origin 29; left pectoral fin length -10; base of left
pectoral fin 2; gill opening length 2.5; isthmus 3.9; snout 4.6; snout tip to tip of lower jaw -0.5; lip of snout to
rictus of jaw 10; eye diameter 3.1; interorbital distance 3.2. Total vertebrae 142; predorsal vertebrae 14; preanal
vertebrae 59.
Body moderately elongate, its depth behind gill openings 31-42 times and width behind gill openings
35-54 times in TL. Body cylindrical in anterior trunk region, laterally compressed posteriorly. Head and trunk
2. 2-2. 4 times and head 10.0-10.7 times in TL. Snout short, broad and swollen in appearance at anterior nostril
578
J. E. McCOSKER
bases. Lower jaw included, nearly reaches snout. Anterior nostrils in forward-directed tubes that reach snout tip;
posterior nostril in lip. covered by a flap with an anterior barbel that extends to or below the lip (see Fig. 1).
Eye large, its center above posterior 1/3 of upper jaw.
Dorsal fin low in anterior trunk region, elevated posteriorly. Its origin just in advance of end of pectoral rays.
Pectoral fins elongate, lanceolate, the middle rays longest.
Cephalic pores (Fig. 3) very reduced but discernible. Six or seven mandibular, 2 preopercular, 1 ethmoidal +
4 supraorbital, 4 + 2 intraorbital, and single interorbital and supralemporal pores. Eight lateral line pores above
branchial region. Lateral line pores of trunk and tail not possible to discern, minute and covered with a waxy
exudate.
Teeth (Fig. 2b) small and conical, nearly subequal. An intermaxillary rosette of 7 teeth, followed by a gap.
then a patch of 8 teeth followed by 15 uniserial vomerine teeth. Jaw teeth biserial, the rows irregular, each with
approximately 38-40 pairs.
Body coloration of small (195-232 mm) specimens in ethanol pale yellow, overlain on snout, nape, and dorsal
half with a line peppering of minute brown punctations which end. particularly in smaller specimens, abruptly
at the lateral line. All fins clear. Body coloration of largest specimen (337 mm) in isopropanol pale yellow, with
a pale brown band across shoulders beginning behind temporal pore band, becoming diffuse in region of lateral
line belore end of pectoral lins. Cheeks bright yellow. Body and tail covered throughout with a fine peppering of
minute brown spots. Peritoneum white with a fine gray/black speckling. Median fins and posterior half of pectoral
fins clear.
FIG sm cp'no' 4n8°5Vm h0l0lyPe’ ™ mm TL (MNHN l998'43)- NeW Caledo111^ 18°57,00'S, I63°12.60'E.
Size. — To 337 mm. The holotype, 230 mm TL, is a female with developing ova (~0.5 mm egg diameter)
The largest known specimen, BPBM 34923, is also a female with developing ova (slightly smaller in diameter
than those of the holotype).
Etymology. — Named in honor of Eugenia B. Bohlke, friend and contributor to knowledge of apodal
fishes.
from IhcTcddTv™’ ~ Kn°Wn fr°m Ser'eS’ C°lleCled bCIWeen 435‘500 m dcP,h in New Caledonia and
Source : MNHN , Paris
OPHICHTHIDAE FROM THE NEW CALEDONIAN REGION
579
Remarks and Comparisons. — The new species is similar to other species of Ophichthus , subgenus
Coecilophis Kaup (c/. McCosker, 1977), which share a preference for deep sand and mud substrates. All possess
small dentition, posterior nostrils along the lip (rather than opening into the mouth) and preceded by a Hap, two
rather than three prcopercular pores, and a plain coloration, often with a dark smudge along the anal fin near the tail
tip. Its closest relatives appear to be O. kunaloa McCosker, 1979. a deepwater Hawaiian species that has more
vertebrae (MVF 17.5-66.5-183 vs. 14-57-142 for O. genie) and appears to be more robust (depth 27.5-31.5 times
in total length vs. 31-42 times for O. genie).
Table 1. — Proportions (in thousandths) and counts of the holotypes and paratypes of Ophichthus genie , O. mystacinus
and O. exourus. Abbreviations are: TL = total length; HL = head length; DFO = dorsal fin origin; 10 = interorbital
width.
Ophichthus mystacinus sp. nov.
Figs 2c, 4; Tabl. 1
Material examined and Types. — 4 specimens.
New Caledonia. Musorstom 4: stn CC 202. 18°58,00’S. 163°10.50'E, 580 m, 20.09.1985: holotype. 426 mm TL
(MNHN 1998-46); I paratype. 429 mm TL (CAS 89552). — Stn CP 180, 18°56.0'S. I63°l 7,70’E, 450 m. 18.09.1985:
1 paratype, 336 mm TL (BPBM 37308). — Stn CC 201, I8°55,80’S, 163°13.80'E, 500 m, 20.09.1985: 1 paratype.
383 mm TL (MNHN 1998-47).
OTHER MATERIAL EXAMINED. — Ophichthus brachynotopterus : Madagascar, 1971: 1 paratype. 442 mm TL
(MNHN 1979-22); 1 paratype, 41 3 mm TL (MNHN 1979-23).
DIAGNOSIS. — A moderate length species of Ophichthus with the unique combination of characters: tail 59-
60% of total length; dorsal fin origin about mid-trunk; pectoral fins elongate, the central rays threadlike; jaw
elongate; eye large; teeth conical, numerous and small, biserial in jaws and anteriorly on vomer; coloration
yellowish tan, brown dorsally, the fins pale; and mean vertebral formula 33.8-61.8-172.8.
Description. — (Counts and measurements in mm of the holotype). Total length 426; head length 47.1;
trunk length 125; tail length 254; body depth at gill openings 16; body width at gill openings 14; body depth at
Source :
580
J. H. McCOSKER
anus 13; body width at anus 1 1; tip of snout to dorsal fin origin I 10: left pectoral fin length 20.3: base of left
pectoral fin 3.8; gill opening length 4.3; isthmus 8.7; snout II; snout tip to lip of lower jaw 2.5; tip of snout
to rictus of jaw 26; eye diameter 6; interorbital distance 5.6. Total vertebrae 174; predorsal vertebrae 35;
preanal vertebrae 62.
Body moderately elongate, its depth behind gill openings 23-28 times and width behind gill openings
29-35 times in IL. Body cylindrical anteriorly, laterally compressed in tail region. Head and trunk 2.4-2. 5 and head
8.6-9. 3 in TL. Head elongate, conical when viewed from above, somewhat flattened dorsally. Snout pointed,
sharply conical front above. Lower jaw nearly reaches snout tip. Anterior nostrils in a short tube; posterior nostril
in upper lip and covered by a flap, appearing as a slit in the upper lip before and beneath the orbit. Eye large,
its center above middle of upper jaw.
Median fins low and poorly developed, lying within a groove for most of their length. Dorsal fin origin just
anterior to mid-trunk. Pectoral tins elongate, the central rays threadlike and longest. Median fins expanded before
naked tail tip, lying within a groove.
Cephalic pores (Fig. 4) small and very difficult to discern. Six mandibular. 2 preopercular, 1 ethmoidal +
4 supraorbital, 4 + 2 infraorbital, and single intcrorbital and supratemporal pores. Nine lateral line pores above
branchial region. Lateral line pores of trunk and tail minute, not possible to discern.
leeth (Fig. 2c) small, nearly equal in size, conical and sharply pointed, and separated at their bases. A small
rosette of 5 teeth at snout lip, followed by 12 neatly-aligned pairs extending nearly to mid-vomer, followed by
about 14 umsena! teeth. Six uniserial, closely-spaced teeth in the outer row of the upper jaw. followed by two
rows ol about 45 teeth which form pairs for most of their length, the last few becoming more randomly placed.
Lower jaw with about 4-5 pairs anteriorly, followed by a row of about 15 teeth, then about 25-30 irregular pairs.
Body coloration in ethyl alcohol tan, overlain with a dusting of microscopic brown spots which are denser
above the lateral midlinc. giving the body and tail a brown dorsal cast. Area surrounding base of anterior nostril
tubes has a high density of dark punctations, particularly as seen from above, appearing like a faint mustache.
Chm. throat, belly, and lower edge of tail and tail tip yellow. Peritoneum white with a fine black spccklins.
Median fins clear. Pectoral fins yellow. The hindmost (ca. the length of the head) ventral surface of the tail Is
black, including a black smudge on the membrane of the anal fin.
Size. To 429 mm. The holotype, 426 mm, is a male with undeveloped gonads.
Etymology. — From the Greek, mystacinus, mustachioed.
Source :
OPHICHTHIDAE FROM THE NEW CALEDONIAN REGION
581
DISTRIBUTION. — Known only from the MUSORSTOM New Caledonia collections, between 450-580 m depth.
Remarks and Comparisons. — Ophichthus mystacinus is similar to other species of Ophichthus of the
subgenus Coecilophis Kaup (cf. McCoSKER. 1977). It appears most closely related to O. brachynotopterus Karrer,
1982, a deepwater (355-478 m) species from northwest Madagascar. I have compared the paralypes of that species
to the new species, and find O. brachynotopterus easily separable from O. mystacinus by the former's: vertebral
number (178 vs. 169-174); shorter pectoral fin (3. 6-4.0 times in head vs. 2. 2-2. 6); and shorter snout (4. 6-4. 7
times in head vs. 3. 6-4. 4). As well, O. brachynotopterus appears to have a somewhat shorter head and larger eye.
Genus RHINOPHICHTHUS nov.
Type Species. — Rhinophichthus penicillatus sp. nov.
DIAGNOSIS. — Elongate ophichthids, subfamily Ophichthinae, tribe Ophichthini (sensu McCoSKER, 1977),
with tail much longer than head and trunk; median fins low; dorsal fin arising behind pectoral fin; pectoral fin base
arising above and occupying less than half of gill opening; gill openings lateral, elongate, nearly vertical and
crescentic; eye moderately developed, above posterior third of jaw; jaws moderately developed, but not elongate;
snout conical, tapering evenly to a sharp point; underside of snout separated in advance of anterior nostril bases
exposing the teeth; teeth conical, numerous and small, uniserial or biserial; 2 preopercular pores; gill arches
developed, similar to those of Ophichthus , however fifth ceratobranchial present and ossified as a thin rod for
anterior 80 %, the remainder cartilaginous, and upper pharyngeal tooth plates fused. Other characters those of the
two species.
Etymology. — From the Greek, rhinos , snout, and Ophichthus , a genus of snake eel; gender masculine.
Remarks. — In general appearance, the species of Rhinophichthus appear similar to that of the elongate¬
snouted species of Ophisurus. They differ in the condition of the snout (clavate rather than conical) and in their
dentition (enlarged and fanglike, particularly those of the intermaxillary). With the exception of the condition of
the snout and its associated tissues, Rhinophichthus is very similar to the generalized species of Ophichthus. The
appearance of the snout of Rhinophichthus is more comparable to that of species within the tribe Sphagebranchini.
particularly those of the genera Apterichtus and Ichthyapus. Those eels generally occupy shallow water sand
habitats and are capable of burrowing quickly into the substrate with either end, an adaptation which I presume that
Rhinophichthus penicillatus enjoys as well. Other adaptations displayed by the new species, such as the nearly
uniform coloration, small and numerous teeth, fairly large eye, posterior nostril along upper lip and covered by a
Hap, and poorly developed median fins, are typical of other deep-dwelling ophichthids.
An additional taxon that I can assign to Rhinophichthus is Ophichthys unicolor Regan, 1908. It. and its
synonym Ophichthys algoensis Barnard, 1925 (a replacement name for Ophichthys triserial is Barnard. 1923,
preoccupied) were described from Algoa Bay, South Africa, from 40 and 50 fathoms, respectively (Smith. 1962;
McCoSKER & CASTLE, 1986). Differences between O. unicolor and the new species are described in the Remarks
below. A complete osteological analysis of Rhinophichthus may demonstrate that its ancestry is shared with that
of species of Ophichthus.
Rhinophichthus penicillatus sp. nov.
Figs 5-6a; Tabl. 2
Material examined. — 3 specimens.
New Caledonia. MUSORSTOM 4: stn CP 171, 18°57,80'S, 163°14,00’E, 435 m, 17.09.1985: holotype. 457 mm TL
(MNHN 1998-48); 1 paratype. 359 mm TL (MNHN 1998-49); 1 paratype, 427 mm TL, gill arches removed and cleared and
counterstained (CAS 89553).
Source :
582
J. E. McCOSKER
(BMNHIE1906.tn9 39) EXAMINED' ~ Ophichihys unicolor. Algoa Bay. Souih Africa, 1906: holotypc - 271 mm TL
Ophichihys algoensis: Algoa Bay. South Africa. 1904: holotypc. 300 mm TL (SAM 12776).
Diagnosis - An elongate ophichthine, genus Rhinophichthus. with the unique combination of characters-
tail 66-67 % of total length: dorsal fin origin behind pectoral lips; pectoral fins small and paddle-shaped, not
elongate; eye large; dentition small, conical and uniserial throughout; coloration pale, slightly darker dorsal Iv fins
pale; and mean vertebral formula 1 2-54- 173. "
Inn mESna,P,TI-??' ~ and measurcmenls i[> mm of the holotype). Total length 457; head length 35- trunk
0 5 h i eng °osb0dy dCP,h 31 gi" °PenmgS ,a2; b°d>' widlh a> openings 8.5; body depth a. Is
ln 2 3 dM,W' TUS . ;l‘P°f Sn°U1 10 d0rsal fin ori§in 47 ■ left Pecl°ral fin length 6.7; base of left pectoral
rintiK Vs ZTg kTh 5'2: ,SlhmUS 4,5; Sn0Ul leng,h 9 7 ' snoul l'P 10 liP of lower jaw 2 8- tip of snout to
veitebrae^54 ^ ^ ^ 3* ^ ™ «hae .73; predorsll vertebrTe 12 ”al
Body elongate its depth behind gill openings 45-53 times and widlh behind gill openings 39-59 times in TI
Body cylindrical laterally compressed only posteriorly. Head and trunk 3 times and hetfd 1 3- W limes i TL Snliut
inSd "sLrzt °nas rerz wiih
Source :
OPHICHTHIDAE FROM THE NEW CALEDONIAN REGION
583
/
Teeth (Fig. 6a) small, conical and nearly suhequal,
evenly spaced but not close-set. A narrow rosette of
5 teeth just behind snout tip, followed by a space,
3 teeth, and 12 uniserial vomerine teeth. Jaw teeth
neatly uniserial: 20-22 in upper jaw and 15-17 in
lower jaw.
Body coloration in ethanol uniformly pale yellow,
overlain on upper half of head, trunk, and tail with
numerous fine brown punctations. Chin, throat, belly
and underside of snout pale. Peritoneum while. All
fins clear.
SIZE. — The largest known specimen, the
hololype, is 457 mm long. It is a female with
developing ova (egg diameter -0.5 mm).
Etymology. — From the Latin penicillus , a
pencil, and - citus , having the nature of, in reference to
the sharpened-pcncil appearance of both ends of this
eel.
Fig. 6. — Schematic diagram of dentition of hoiotypes of
A. Rhinop/iic/uhus penicillatus sp. nov.. 457 mm TL
(MNHN 1998-48), and B, Yirrkala insolitus sp. nov.,
258 mm TL (MNHN 1998-50).
species is easily separable from any known ophichthid. Although quite generalized in most of its external
morphology, its conical snout and other ophichthin characters are shared only by the poorly-known Ophichthus
unicolor Regan, which I assign to the genus Rhinophichthus. I examined the dried and poorly-preserved holotype
of O. unicolor in 1982 and recently compared the holotype of its synonym. Ophichthys algoensis Barnard, to
specimens of R. penicillatus. R. unicolor differs in having fewer vertebrae (156 vs. 173) and biserial rather than
uniserial dentition.
Distribution. — Known only from the
MUSORSTOM New Caledonia collections, from 435 m
depth.
Remarks and Comparisons. — The new
Genus YIRRKALA Whitley, 1940
Yirrkala Whitley, 1940: 410 (type species Y. chaselingi Whitley, 1940, by original designation).
Pantonora Smith, 1964: 719 (type species Ophichthys tenuis Gunther, 1870. by original designation).
Diagnosis. — Ophichthin eels, tribe Sphagebranchini, with the following characteristics: body elongate,
cylindrical, shorter than tail: snout conical: dorsal fin arises above head, along trunk, or above anus; pectoral fins
absent; gill openings ventral, longer than isthmus; teeth conical, generally uniserial; coloration usually uniform or
darker dorsally, although striped in one species.
Remarks. — In earlier works (McCosker, 1977: 69; McCosker & Castle, 1984: 185), I considered the
type species, Y. chaselingi, to be a junior synonym of Sphagebranchus lumbricoides Bleeker. Subsequent
examination of additional material has demonstrated that Whitley's chaselingi is a valid species. Mv revision of
the genus, recognizing approximately 10 valid Indo- Pacific species, is currently in preparation.
The new species shares all of the external morphological characteristics of the genus Yirrkala . however its
dorsal fin origin is far posterior to that of its congeners. Such a condition is not significant enough to merit
generic recognition, and that character state within the genus should therefore be expanded to range from before the
gill openings to above the anus.
Source :
584
J. E. McCOSKER
Table 2. — Proportions (in thousandths) and counts of the holotype and paratypes of Rhinophichthus penicillatus and
the holotype of Yirrkala insolitus. Abbreviations are: TL = total length; HL = head length; DFO = dorsal fin origin;
10 = interorbital width.
Yirrkala insolitus sp. nov.
Figs 6b-7; Tabl. 2
Material examined. — i specimen.
New Caledonia. Musorstom 4: stn CP 148. 19°23.40'S. I63°31.90'E. 59 m. 14.09.1985: holotype. 258 mm TL
(MNHN 1998-50).
Other Material EXAMINED. — Yirrkala chaselingi : Queensland. Australia: holotype, 610 mm TL (AM IB. 481)
Sphagebranc/uis gjellerupi : New Guinea, 1910: holotype. 153 mm TL (ZMA 104.146).
Sphagebranchus lumbricoides : Timor: holotype. 230 mm TL (BMNH 1867.1 1.28.300).
Ophichthys misolensis : Misol Island: holotype, 283 mm TL (BMNH 1 870.8.3. 1 1 2).
Dal op his moluccensis : Ceram: holotype. 405 mm TL (BMNH 1867.11.28.289).
Ophichthys tenuis : locality unknown, possibly Mauritius: lectotype -530 mm TL (BMNH 1965:1.2.1)
Ophichthys timorensis : Timor: holotype -205 mm TL (BMNH 1867.11.28.322).
Diagnosis. — An elongate species of Yirrkala with the unique combination of characters: head 7% of total
length (TL); tail 47% of TL; dorsal fin origin above anus; teeth conical, moderately developed, uniserial on jaws
and vomer; coloration pale, overlain with fine brown punctations above midline; and vertebral formula 80-80-149.
Description. — (Counts and measurements in mm of the holotype). Total length 258; head length 1 8; trunk
length 149; tail length 121; body depth at gill openings 5.3; body depth at anus 4.6; body width at gill openings
3.3; body width at anus 3.3; origin of dorsal fin 165; left gill opening length 1.4; isthmus 1.2; snout length 2.3;
snout tip to tip of lower jaw - 1.8; tip of snout to rictus of jaw 5.0; eye diameter 1.0; intcrorbital distance 1.3.
lotal vertebrae 149; predorsal vertebrae 80; prcanal vertebrae 80.
Body very elongate, its depth at gill openings 49 times in TL. tapering posteriorly to an acute, finless point,
o y and tail nearly cylindrical throughout, becoming laterally compressed in posterior 10%. Head and trunk
times and head 14.3 times in TL. Snout acute at tip, conical from above, rounded on underside and split
Source : MNHN, Paris
OPHICHTHIDAE FROM THE NEW CALEDONIAN REGION
585
medially. Lower jaw included, falls far short of anterior nostril edge and barely exceeds anterior pupil edge. Anterior
nostril nearly Hush with snout; posterior nostril within upper lip, visible externally as a slit beneath center of eye.
Eye moderately developed, its center at middle of upper jaw.
Median fins low but obvious. Dorsal fin arises above level of anus and ends 1/2 HL from tail tip. Gill
openings low, their major axis nearly horizontal, without an anterior lateral membrane or duplication. Isthmus
narrow, slightly less than gill opening.
Cephalic pores minuscule, barely discernible (Fig. 7). Four mandibular, 2 preopercular, l ethmoidal +
4 supraorbital, 4 + 2 infraorbital, and single interorbital and supratemporal pores. Lateral line pores not possible
to discern.
Teeth (Fig. 6b) moderately developed, conical and uniserial. An anterior chevron of 3 intermaxillary teeth,
followed by a short gap and 9 vomerine teeth in a nearly uniscrial row, descending in size. Jaw teeth uniserial,
approximately 7 in upper jaw and 13 in lower.
Body coloration in ethanol pale, with fine brown punctations above the lateral midline, becoming more
numerous above dorsal midline. A few fine brown spots on throat and chin, forming fine dotted lines between eyes
and across nape. Median fins yellow, unspotted.
Size. — The only known specimen is 258 mm TL, a female with unripe gonads.
ETYMOLOGY. — From the Latin insolitus , meaning unusual or strange, in reference to the posterior location
of the dorsal fin origin, as well as the unusual depth of capture of an Yirrkala.
Distribution. — Known only from the holotypc, from 59 m off New Caledonia.
Remarks and COMPARISONS. — The new species differs from all its known congeners in the location ol its
dorsal fin origin, far posterior to that of all other species. Other body characteristics differ little from the condition
of other known species of Yirrkala , and I find no reason to not consider it congeneric. Its habitat is deeper than that
of most of its congeners, which are known from shallow water sand and mud bottoms, and some enter freshwater
streams.
1
Fig. 7. — Yirrkala insolitus sp. nov., holotype, 258 mm TL (MNHN 1998-50). New Caledonia. 19°23,40'S, 163°31.90E,
Musorstom 4, stn CP 148, 59 m.
Source :
586
J.E. McCOSKER
ACKNOWLEDGMENTS
I am most grateful to Bernard Seret for making the Musorstom eels and other specimens from the Museum
national d'Histoire naturelle available for study. R. Charles Anderson (Marine Research Section, Male, Maldives)
kindly provided the Maldives type specimen. J. Seeto kindly donated the Fijian type specimen. For permission to
examine specimens in their care, I also thank: D. F. HOESE, J. R. PAXTON, M. McGROUTHER and others at the
Australian Museum; E. BOHLKE and others at the Academy of Natural Sciences of Philadelphia; the curatorial staff
of The Natural History Museum (London); J. E. Randall and A. SUZUMOTO at the Bernice P. Bishop Museum;
W. N. Eschmeyer, T. Iwamoto, D. Catania, J. Fong and others at the California Academy of Sciences;
A. Graham and A. R. Williams at CSIRO; M. F. GOMON at the Museum of Victoria (Melbourne);
M. E. Anderson at the J.L.B. Smith Institute of Ichthyology (Grahamstown); the curatorial staff of the South
African Museum; R. Rosenblatt at the Scripps Institution of Oceanography; and the ichthyological staff of the
National Museum of Natural History (Washington). Tomio IWAMOTO (California Academy of Sciences) read
a draft of this manuscript and provided valuable advice. Molly Brown prepared the illustrations. Other individuals
assisted in many ways, including but not limited to Mysi HOANG, Dong LlN, Roy ElSENHARDT, and Carl
Ferraris of the California Academy of Sciences.
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Rapports scientifiques et techniques . Centre ORSTOM de Noumea . (37): 1-23.
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Whitley, G. P., 1950. — New fish names. Proceedings of the Royal Society of New South Wales for 1948-49: 44.
Source : MNHN. Paris
REMERCIEMENTS AUX RAPPORTEURS / ACKNOWLEDGEMENTS TO REFEREES
La Redaction tient & remercier les experts exterieurs au Museum national d'Histoire naturelle dont les noms suivent. d'avoir
bien voulu contribuer, avec les rapporteurs de rEtablissement, a revaluation des manuscrits (1995/1998) :
The Editorial Board acknowledges with thanks the following referees who, with Museum referees, have reviewed papers
submitted to the M^moires du Museum (1995/1998):
Source : MNHN, Paris
Acheve d imprimer en janvicr 1999 sur les presses de Plmprimerie de Monlligeon
La Chapelle Monlligeon - Depot legal I irimeslre 1999 - N Imp. 19185 - Distribute 10 fevrier 1999
Source : MNHN. Paris
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2 2 FEV. 1999
Source : MNHN. Paris
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Volume 4. 400 pp. (ISBN : 2-85653-505-04) 450 FF.
Tome 170 : Peter A. ZIEGLER & Frank HORVATH (eds), 1996. — Peri-Tethys
Memoir 2: Structure and Prospects of Alpine Basins and Forelands. 552 pp. +
atlas. (ISBN : 2-85653-507-0) 450 FF.
Informations sur les Publications Scientifiques du Museum national d'Histoire naturellc :
Informations about the Scientific Publications of the Museum national d'Histoire naturelle:
Internet http://www.mnhn.fr/
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Prices in French Francs, postage not included.
Resultats des Campagnes MUSORSTOM is an ongoing series on the deep-sea fauna of
the tropical Indo-Pacific. Initially focussed on the New Caledonia region, the series has now
expanded to cover other South Pacific islands and island groups, in particular Vanuatu,
Wallis and Futuna, Fiji and the Marquesas. MUSORSTOM, which rests on an international
network of collaborating taxonomists, is the most ambitious ongoing program of deep-sea
zoological exploration.
The present volume contains contributions by eleven authors from Japan, New Zealand,
USA and Europe, and deals with deep-sea corals (I paper), bryozoans (1), isopod and
decapod crustaceans (6), and snake eels (1). The results present new data on the
morphology, systematics and distribution of more than 250 species. One new subfamily,
five new genera and 60 new species are described. Of broader biological interest are the
descriptions of new biological associations, e.g., between hermit crabs and Zoanthidea, and
between acrothoracican cirripedes and corals. Carrier shells (Xenophora) were found to
incorporate into their shells species of deep-sea corals that have otherwise never been
collected by conventional means, which draws attention to the still inadequate sampling
intensity of the tropical deep-sea benthos. The bryozoan genus Pseudothyracella,
previously known as a Paleogene fossil in Europe and North America, is added to the list of
"living fossils", being represented by a new, living species.
The MUSORSTOM series is a joint program of the Museum national d'Histoire naturelle
and the Institut Fran?ais de Recherche Scientifique pour le Developpement en Cooperation
(ORSTOM).
PUBLICATIONS
SCIENTIFIQUES
DU MUSEUM
57, RUE CUVIER
75005 PARIS
ISBN 2-85653-520-8
ISNN 1243-4442
600 FF TTC
Source : MNHN, Pans