MEMOIRES

- - - s

Re suit at s des Campagnes MUSORSTOM

DU MUSEUM

NATIONAL

D’HISTOIRE

NATURELLE

Volume 14

Coordonne par

TOME 167

Philippe BOUCHET

ZOOLOGIE

1995

Source : MNHN, F

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Ce volume des Resultats des Campagnes MUSORSTOM est
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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.

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Volume 14 : Mem. Mus. natn. Hist, nat., 167 : 1-654, 987 fig., 3 pi. couleur (1995). ISBN : 2-85653-217-9.

Source MNHN. Paris

esultats des campagncs

Volume 14

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ISBN : 2-8565 3-217-9
ISSN : 1243-4442

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MEMOIRES DU MUSEUM NATIONAL D’HISTOIRE NATURELLE

TOME 167

ZOOLOG IE

Resultats des Campagnes Musorstom

Volume 14

Coordonne par

Philippe BOUCHET

Museum national d’Histoire naturelle
Laboratoire de Biologie des Invertebres marins et Malacologie
5 rue Buffon
F-75005 Paris

EDITIONS
DU MUSEUM
PARIS

1995

Source : MNHN , Paris

Couveriure : Cardiomya gouldiana

Source : MNHN , Paris

SOMMAIRE

Contents

Pages

1 . Bathyal Pectinoidea (Bivalvia: Propeamussiidae, Entoliidae, Pectinidae) from New Caledo¬
nia and adjacent areas . 9

Henk H. Dijkstra

2. Systematic revision of living species of Meiocardia , Glossidae and Glossocardia, Trape-

zidae (Bivalvia) . 75

Akihiko Matsukuma & Tadashige Habe

3. Carnivorous bivalve molluscs (Anomalodesmata) from the tropical western Pacific Ocean,

with a proposed classification and a catalogue of Recent species . 107

Jean-Maurice Poutiers & Frank R. Bernard

4. Scaphopoda of the tropical Pacific and Indian Oceans, with description of 3 new genera and

42 new species . 1 89

Victor Scarabino

5. Calliostomatidae (Gastropoda: Trochoidea) from New Caledonia, the Loyalty Islands, and

the northern Lord Howe Rise . 381

Bruce A. Marshall

6. The Trophoninae (Gastropoda: Muricidae) of the New Caledonia region . 459

Roland Houart

7. A review of the deep-water volute genus Calliotectum (Gastropoda: Volutidae) . 499

Philippe Bouchet & Guido T. Poppe

8. A revision of the drilliid genera Splendrillia and Plagio strop ha (Gastropoda: Conoidea) from

New Caledonia, with additional records from other areas . 527

Fred E. Wells

9. Deep-water Cones (Gastropoda: Conidae) from the New Caledonia region . 557

Dieter Rockel, Georges Richard & Robert G. Moolenbeek

10. Mathildidae from New Caledonia and the Loyalty Islands (Gastropoda: Heterobranchia). . 595

Rudiger Bieler

Index . 643

Source : MNHN, Paris

FATS DES CAMPAGNES MUSORSTOM. VOLUME 14 RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 14 RESULT/

Bathyal Pectinoidea

(Bivalvia: Propeamussiidae, Entoliidae, Pectinidae)
from New Caledonia and adjacent areas

Henk H. DIJKSTRA

Institute of Systematics and Population Biology
(Zoological Museum)

University of Amsterdam
P.O. Box 94766, 1090 GT Amsterdam
The Netherlands

ABSTRACT

The biological exploration of deep-sea benthos off New Caledonia during the years 1978-1989 has yielded a rich mollusc
fauna, including 30 species of Pectinoidea. The highest diversity, with 14 species, is observed in the 600-800 m depth interval,
and only three species have been collected below 1500 m. The fauna belongs to Propeamussiidae (21 species, all taken alive),
Entoliidae (1 species, alive), and Pectinidae (8 species, 6 taken alive). Nine species are new to science: Parvamussium
multiliratum, P. retiaculum, P. retiolum, P. squalidulum, P. undisonum. P. vesiculatum, Cyclopecten horridus, C. pellucidulus
(Propeamussiidae), and Hyalopecten mireilleae (Pectinidae). Most of the other species are new records for the region. Ten
lectotypes are designated, one new synonym and one new1 combination recognized. This pectinoid fauna shows a strong
similarity to that oT the wider Indo-Pacific, and marginally to that of northern New Zealand and southeastern Australia.

RESUME

Les Pectinoidea bathyaux (Bivalvia: Propeamussiidae, Entoliidae et Pectinidae) de Nouvelle-Caledonie et des regions
voisines.

Les campagnes oceanographiques realisees de 1979 a 1989 autour de la Nouvelle-Caledonie ont recolte une riche faune
de moliusques bathyaux et abyssaux, dont 30 especes de bivalves Pectinoidea. Le maximum de diversite est observe entre 600
et 800 m de profondeur, avec 14 especes. alors que trois especes seulement ont ete recoltees a plus de 1500 m. Cette faunule
se repartit en Propeamussiidae (21 especes, toutes recoltees vivantes). Entoliidae (1 espece, vivante), et Pectinidae (8 especes.
dont 6 recoltees vivantes). Neuf especes nouvelles sont decrites : Parvamussium multiliratum. P. retiaculum. P. retiolum. P.

Oijkstra, H.H., 1995. Bathyal Pectinoidea (Bivalvia; Propeamusiidae, Entoliidae, Pectinidae) from New Caledonia and adjacent areas. In: P. Bol'CHET (ed.).
Resultats des Campagnes Musorstom, Volume 14 Mem. Mus. natn. Hist. mu.. 167; 9-73. Paris ISBN 2-85653-217-9.

Published 29'h December 1995.

Source . MNHN. Paris

10

HENK DIJKSTRA

squalidulum, P. undisonum, P. vesiculatum, Cyclopeclen horridus, C. pellucidulus (Propeamussudae), et Hyalopecten miredleae
(Pectinidae). La plupart des autres especes sont signalees pour la premiere fois de ce secteur geographique. Des lectotypes soni
desienes pour dix taxons, et une nouvelle synonymie et une nouvelle combinaison sont etablies. Cette faune de Pectinoidea fait
incontestablement partie de ITndo-Pacifique avec, marginalement. quelques affinites avec le Nord de la Nouvelle-Zelande et
le Sud-Est de 1'Australie.

INTRODUCTION

Deep-sea pectinoid bivalves from the tropical South-West Pacific are poorly known, and only
a few species were described by previous authors (E.A. Smith, 1885; Hedley, 1902). During the years
1978-1989 several French expeditions have collected rich samples of marine biota from bathyal depths
around New Caledonia, including many pectinoidean bivalves. The species treated here are those
occurring at depths below 100 m, i.e. all the littoral and reef associated species are not considered.
With these limitations, the collection comprises 21 species of Propeamussiidae (all taken alive), one
species of Entoliidae (alive), and 8 species of Pectinidae (6 taken alive), a total of 30 pectinoids. To
make the survey of New Caledonia Propeamussiidae more complete, Parvamussium pauciliratum, a
shallow-water species, is also treated and illustrated.

The present paper completes the revision of the pectinoids of New Caledonia. In a series of
papers, Dijkstra (1983 to 1994) and Dijkstra et al. (1989, 1990) recorded the occurence of 33 species
of Pectinidae from the reefs and coral reef lagoons.

As for other papers in this volume, the material studied was collected during the cruises
reported on by Richer de Forges (1990, 1993). For descriptions of deep-sea bottom topography and
faunal zonation around New Caledonia, 1 refer to Roux (1991) and Roux et al. (1991). In species
descriptions, morphological terminology follows Waller (1978, 1984, 1991, 1993) and Waller &
Marincovich (1992).

Comparative material from the Indo-Pacific and type material was studied from various
museum collections, viz. ams, bmnh, kbin, mnhn, nmnz, nmp, nmw, rmnh, zma, zmc, zsi. The
present material is stored in MNHN; paratypes have been distributed to other museums, including
the reference collection of the author.

ABBREVIATIONS AND TEXT CONVENTIONS

Repositories

AIM

AMS

BMNH

HD

IOAS

KBIN

MCZ

MNHN

NMNZ

NMP

NMV

NMW

NSMT

RMNH

USNM

ZMA

ZMC

ZSI

: Auckland Institute and Museum, Auckland
: Australian Museum, Sydney
: The Natural History Museum, London
: H.H. Dijkstra collection

: Institute of Oceanology, Academia Sinica, Qingdao
: Koninklijk Belgisch Instituut voor Natuurwetenschappen, Brussels
: Museum of Comparative Zoology, Cambridge (U.S.A.)

: Museum national d’Histoire naturelle, Paris
: Museum of New Zealand Te Papa Tongarewa, Wellington
: Natal Museum, Pietermaritzburg
: National Museum of Victoria, Melbourne
: National Museum of Wales, Cardiff
: National Science Museum, Tokyo
: Nationaal Natuurhistorisch Museum, Leiden
: National Museum of Natural History, Washington, DC
: Zoologisch Museum, Amsterdam
: Zoologisk Museum, Copenhagen
: Zoological Survey of India, New Alipur, Calcutta

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

11

Station data

CC : Chalut a crevettes (Shrimp Trawl)

cp : Chalut a perche (Beam Trawl)

DC : Drague Charcot (Charcot Dredge)

de : Drague epibenthique (Epibenthic Sledge)

ds : Drague epibenthique Sanders (Sanders epibenthic Sledge)

dw : Drague Waren (Waren Dredge)

kg : Carottier Usnel grande surface (Usnel Box-Corer)

Other abbreviations

db : doublet (paired valves, dead collected)

Iv : left valve(s)

rv : right valve(s)

spm(s) : live-taken specimen(s)

v : valve(s)

od : Original designation

so : Subsequent designation.

SYSTEMATIC ACCOUNT

Class Bivalvia Linnaeus, 1758
Subclass Pteriomorphia Beurlen, 1944 [emend., Boss 1982]

Superorder Eupteriomorphia Boss, 1982
Order Ostreoida Waller, 1978
Suborder Pectinina Waller, 1978
Superfamily Pectinoidea Wilkes, 1810 [emend., Waller 1978]

Family Propeamussiidae Abbott, 1954

Propeamussiidae Abbott, 1954: 361. 369.

1 f Diagnostic characters. - Free or byssate Pectinoidea with outer foliated calcitic layer on
let t valve and prismatic calcitic layer on right valve present on the main part of the disc; inner layer
crossed-lamellar aragonite beyond pallial line, sometimes nearly to distal margins; byssal notch
without ctenolium.

Remarks. Hertlein (1969: N350) placed Propeamussium (with Parvamussium), as a
subgenus together with Amusium Roding, 1798 and Korobkovia Glibert & van de Poel, 1965 in the
Amusium-group, with the note that these genera may have been derived from different groups in
lectimdae. Abbott (1954), however, had introduced a new family Propeamussiidae [emended by
waller (1978: 353)], for Propeamussium. Waller (1984) also included several other related genera
m Propeamussiidae, viz. Parvamussium , Cyclopecten Verrill, 1897, Similipecten Winckworth, 1932,
and Latdlopecten Iredale, 1939. Hayami (1988a) mentioned also Polynemamussium Habe, 1951 as a
ecent genus of Propeamussiidae, and treated Parvamussium as a synonym of Propeamussium.
u sequently, Hayami & Kase (1993: 54) raised Parvamussium in rank to genus.

Hayami (1988b) and Waller (1991, 1993) mentioned, the suprageneric classification of the
i ectinoidea is still in disorder and under appraisal.

12

HENK DIJKSTRA

Genus Propeamvssium de Gregorio, 1884

Propeamussium de Gregorio, 1884: 119. [Proposed as a subgenus of Pecten}. Type species (OD): Peclen ( Propeamussium)
ceciliae de Gregorio, 1884; Miocene, Sicily, Italy.

Synonyms:

Paramusium Verrill 1897: 72. Type species (OD): A nuts sium dalli E. A. Smith, 1885; Recent, oil Bermuda. W Atlantic, 796 m.
Occultamussium Korobkov, 1937: 56. [Proposed as a subgenus of Amusium], Type species (OD): Peclen semiradiatus Mayer,
1861; Upper Eocene, Austria. ,

Pseudopalliorum Oyama, 1944: 244. [Proposed as a subgenus of Propeamussium]. Type species (OD): Pecten interradmtus Gabb.
1869; Eocene, California. USA.

Bathymussium Oyama. 1951: 79. [Proposed as a subgenus of Ctenamusium). Type species (OD): Amussium je/Jreysu E.A. Smith.
1885; Recent, N Sulu Sea. Philippines, 686 m.

Micramussium Oyama, 1951: 80. [Proposed as a subgenus of Ctenamusium]. Type species (OD): Ctenamusium ( Micramusstum)
siratama Oyama. 1951; Recent, Japan, 234-291 m.

Flavamussium Oyama. 1951: 81. [Proposed as a subgenus of Parvamussium). Type species (OD): Amussium caducum E.A. Smith,
1885; Recent. Philippines, 1280 m. .

Luteamussium Oyama. 1951: 82. Type species (OD): Amussium sibogai Dautzenberg & Bavay. 1904; Recent, Indonesia, 289 m.

Diagnosis. — Shell equivalve, fragile, usually rather small, mostly transparent, laterally
compressed, gaping along lateral margins; left valve smooth or sculptured with fine radial and/or
concentric riblets or striae, right valve with concentric lines or lirae; auricles nearly equal to equal;
byssal notch moderately slight; no ctenolium; internal riblets extend to submarginal region.

Distribution. — Jurassic-Recent. Worldwide; 275-2740 m (Waller, 1971).

Remarks. — Grau (1959) repeated the original diagnosis of Propeamussium , and of the type
species P. ceciliae, with a translation in English. The red colour of the fossil shell described by de
Gregorio, is probably the colour of iron oxyde, as commented by Gale (in Grant & Gale, 1931:
232). The type specimen has been subsequently figured by de Gregorio (1898: pi. 4, figs 10-12).
Figures 13 and 14, also referred to by Grau (1959; 9) do not belong to the type species. Hertlein
(1969: N350) has reproduced de Gregorio’s illustration of the right (sic) valve of the holotype.
North (1951b: 123) mentioned that he could not trace the holotype.

Grau (1959: 9) placed Paramussium, Pseudopalliorum, Flavamussium and Luteamussium in the
synonymy of Propeamussium, and Hertlein (1969: N350) subsequently added also Occultamussium,
and Actinopecten with a question mark. Hayami (1988a) enumerated also Parvamussium, Ctenamus-
sium (sic), Glyptamusium “Oyama, 1944“ (sic) [= Glyptamusium Iredale, 1939], Bathyamussium and
Micramussium as synonyms of Propeamussium. However, the type species of Ctenamusium and
Glyptamusium are morphologically similar to Parvamussium, whereas the type species of Bathymus¬
sium and Micramussium are more similar to Propeamussium.

Differences in shell characters of Propeamussium and Parvamussium are summarized in Table 1

Table 1

. — Characters of Propeamussiw

<n and Parvamussium.

Propeamussium

Parvamussium

dimension

10-85 mm

5 - 20 mm

shape

orbicular to oval

orbicular to oblique

lateral gape

present

absent

byssal notch

absent

well developed

auricles

equal or nearly equal

unequal

internal ribs

starling in early
ontogeny and developed
to central or sub-marginal area

starling later

and developed to marginal area

Source : MNHN. Paris

BATHYAL PF.CTINOIDEA FROM NEW CALEDONIA

13

Propeamussium alcocki (E.A. Smith, 1894)
Figs 1-4, 133-137

Amussium alcocki E.A. Smith, 1894: 172, pi. 5, figs 15-16.

Other references:

Amussium alcocki - At. cock & Anderson, 1897: pi. 2, figs 3-3a
Thiele & Jaeckel. 1931: 8. - Winckworth, 1940: 26.
Propeamussium alcocki - Abbott & Dance. 1982: 303.

Alcock, 1902: 282, fig. 79

E.A. Smith, 1906: 255.

.,,,/QT|YPEtM^TER,^,L- - Lect°tyPe (Figs 133-7, H 40.4, L 39.8, D 7.9 mm) here designated zs.
6154,9 live taken. Three paralectotypes: bmnh 1894.9.11.1, nmw 1955.158 785 zmc The^tvnes are
somewhat damaged near the margins, and the right valves of three of them (zsi, bmnh nmw) are

Type locality. — " Investigator ”, stn 105, 15H02'N, 72°34' E, Laccadive Sea, 1353 m.

Material examined. — The type material.

Chesterfield Islands, corail 2: stn DW 171, 18°24'S, 155°22'E 650 m 2 Iv I rv
New Caledonia, biocal: stn CP 75, 22° 18' S, 167°23' E, 825-860 m 93 'sDms'
biogeocal: stn CP 232, 21°33' S, 166°27' E, 760-790 m 3 spins

5(11 cp 421 ■ 2o°23'Si i66°2o'E' 800 m- 6 sPms- - stn cp «8

zu 15 8, 166 20 E, 780 m, 5 spms.

Distribution. Laccadive Sea and Bay of Bengal (Smith, 1894; Alcock, 1902) Gulf of
Aden (Thiele & Jaeckel, 1931) and New Caledonia, Chesterfield Islands and Loyalty Islands
Present material living at 760-860 m. *

Description. — Shell inequivalve, circular, up to ca.
M) mm high, inequilateral, fragile, creamy, translucent
Prodmoconch ca. 240 pm in height.

Left valve somewhat more convex than right, with some
concentric growth lines, and 1-3 fine radial lirae near
posterior margin. Anterior margin more convex than poste-
nor. Auricles equal, smooth, anterior margin somewhat
raised, bight to 10 internal riblets entering from near resilifer

to two-thirds length of adult disc, somewhat finer than on
right valve; one auricular lira on each side.

Righi valve covered with wide-set concentric costae,
commencing at ca. 3 mm shell length, with granulate
interstitial microsculpture. Auricles with small concentric
striations, prominent scales on posterior and anterior dorsal
margins. Resilifer triangular. No byssal notch, or ctenolium.

althmmbEH,ARKSi ThC present material of Propeamussium alcocki is similar to the type specimens,
gh the coloration is somewhat different (creamy instead of transparent white) All of the

Sntrd n?retC,I?fbnSJaCk Cr°"ce,n!.ric lamelIae near the ventral margin and minute radiations on the
central part of the disc of the left valve. Knudsen (1967: 281) considered P. alcocki a junior synonym

TTeted aS Variation the differences in shape (circular to nearly oval), concentric
of S n,8r Va V,e (yar-es in setting)’ and radiating sculpture of left valve (varies in development)

New V^l T‘a fr0m. thLC nd‘an °Cean‘ 1 did 1101 °bserve these variations in the material from around
Ldiedonia and the two species are well defined conchologically (see descriptions).

I7QR1 k ‘ a cock‘h'dS rbeen refered to Amussium Hermannsen, 1846 [emendation of Amusium Rodins,
m by a number of authors, but that genus belongs to Pectinidae.

Source MNHN.

14

HENK DIJKSTRA

Figs 1-8. — 1-4. Propeamussium alcocki, biocal: stn CP 75, 46.1 x 39.8 mm (db). — 1, left valve, exterior. — 2, left valve,
interior. — 3, right valve, exterior. — 4, right valve, interior. — 5-8. P. andamanicum , biocal: stn CP 74, 44.y x
36.8 mm (db)’. — 5, left valve, exterior. — 6, left valve, interior. - 7, right valve, exterior. — 8, right valve, interior.

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

15

Propeamussium andamanicum (E.A. Smith, 1894)
Figs 5-8, 138-142

Amussium andamanicum E.A. Smith, 1894: 172-173. pi. 5, figs 13-14.
Other references:

Amimmm andamanicum - E.A. Smith, 1895b: 265. Alcock & Anderson, 1897: pi 2 figs 1-la - E A Smith 19P4-

14. Winckworth. 1940: 26. — Knudsen, 1967: 273, pi. 1, fig. 22, text fig. 15. P ’ 8 t. A. Smith, 1904.

74.R Q1iYPEtTTERxALL' r Lectotype (F'8s 138'142’ H 30-2> L 22.4, D 6.4 mm) here designated zs.
7418/9, live taken. The type specimen is somewhat damaged on the posterior margin of the left valve
and ventral margin of the right valve. Therefore its current dimensions differ ''from those in the
original description. Smith (1895b, 1904) indicated material from different stations of “Investigator”
however he mentioned only one station in the original description. Material from “Investigator”
stn 122 and 192 do not belong to the type series.

Tyre locality. — “ Investigator ”, stn 1 13, 12°59' N, 93°23'10" E, Andaman Sea, 1249 m. Smith
1894) mentioned a depth of 688-922 fathoms [= 1258-1686 m], which should be 683 fathoms
[- 1249 m] (stn 1 13) and 922 fathoms [= 1686 m] (stn 1 14). On a printed label of the Indian Museum
is written in ink station number “113” and a depth of “683” fathoms (see Fig. 138).

Material examined. — The type material.

New Caledonia, biocal: stn CP 74, 22° 14' S, 167°29' E, 1300-1475 m 1 spm
biogeocal: stn CP 238, 21°27'S, 166°23'E, 1260-1300 m. 1 spm.

Distribution. — Andaman Sea, Laccadive Sea and Arabian Sea (Smith, 1895b 1904)
hvinaT 1260 1 475 'm °f (KNUDSEN’ 1967)' and now New Caledonia. Present material

')ES™™ — Shell fragile, hyaline, up to ca. 45 mm
high, elongate, umbonal angle ca. 90”. left valve transparent-
white, right valve creamy.

Prodissoconch ca. 250 pm in height (Knudsen. 1967).

Leji valve somewhat more convex than right, ornamented
with widespread concentric lirae that commence near the
central part of disc and extend to ventral margin. Auricles
equal and rather small, with delicate concentric striations.
Anterior and posterior margin of auricles somewhat raised.

Small scales on dorsal edge of auricles. Interior lirae
commencing directly below resilifer, 9-11, with one small
auricular lira on each side, all gradually enlarging.

Right valve covered with concentric lirae and interstitial
granulated microsculpture (prismatic calcite layer). Auricles
with delicate concentric striae. Dorsal margin with prominent
scales. No byssal notch, no clenolium. Resilifer rather
triangular elongate.

Remarks. — The present specimens resemble the type material, although the concentric lirae
or tne left valve are more delicate and start somewhat earlier. The internal riblets are variable in
length and number. P. andamanicum is closest to P. alcocki, and is a typical Propeamussium.

Propeamussium caducum (E.A. Smith, 1885)
Figs 9-10, 129-132

Amussium caducum E.A. Smith, 1885: 309. pi. 23, figs 1-lc.
Synonyms:

Amussium weberi Dautzenberg & Bavay, 1912: 32. pi. 28, figs 9-13.
lo/wawwar/wn \ ndkazawai Kuroda, 1932. 87, figs 101-102 ( nomen nudum). Synonymy established by Oyama (1951), and

subsequently followed by Habe (1958. 1977) and Knudsen (1967).

16

HENK DIJKSTRA

Figs 9-14. — 9-10. Propeamussium caducum, “Vauban" 1978-79- sin II 77 Q v is q „ , r.

Source : MNHN, Paris

BATHYAL PECT1NOIDEA FROM NEW CALEDONIA

17

Other references:

Amussium caducum - E.A. Smith. 1894: 173; 1895a: 18; 1904: 13; 1906: 255. — Melvill & Standen, 1907: 807. Thiele
& Jaeckel, 1931: 7. Winckworth, 1940: 26.

Parvamussium ( Flavamussium) caducum - Oyama, 1951: 81, pi. 13. figs 11-12. Kira, 1967: 138. pi. 49. fig. 15.
Flavamussium caducum — Habe, 1958: 267. pi. 11, fig. 26.

Propeamussium caducum - Hayami. 1988a: 476. Okutani. Tagawa & Horikawa. 1989: 58, figs. Dukstra, 1991: 6.
Propeamussium I Propeamussium ) caducum — Wang, 1984: 599. pi. 1, figs 3-4. textlig. 2. Dukstra. 1990: 9-10.

Type material. — A. caducum: lectotype (Figs 129-132, H 20.9, L 18.8, D 4.5 mm), here
designated bmnfi 1887.2.9.3310, 4 paralectotypes bmnh 1887.2.9.331 1/1-4. The anterior and posterior
margins of the left valve and the marginal apron of the right valve of the holotype are broken off;
in consequence the measurements differ slightly from the original ones. A. weberi : lectotype, here
designated zma 3.12.013, paralectotypes ZMA, RMNH, kbin, mcz. P. nakazawai: holotype not seen.

Type locality. A. caducum: “ Challenger ", stn 207, 12°2T N, 122°15' E, W ol Luzon.
Philippines, alive, 1280 m. - A. weberi: “Siboga”, stn 316, 7°19.4' S, 116°49.5'E, Bali Sea, alive.
538 m. — P. nakazawai: Suruga Bay, Japan, alive, 549-732 m.

Material examined. - New Caledonia. “Vauban’’ 1978-79: stn 31, 22"31'S. 166" 25' E.
450-550 m, 2 spms. — Stn 32, 22°32' S. 166°25' E, 430-500 m. 2 rv. - Stn 33, 22°33' S, 166°25' E,
290-350 m, 1 lv. - Stn 34, 22°32' S. 166°26' E, 350-420 m, 2 lv.

Distribution. — Japan (Oyama, 1951; Kira. 1967), Philippines (Smith 1885; Oyama,
1951; Knudsen, 1967). Indonesian Archipelago (Dautzenberg & Bavay. 1912; Thiele & Jaeckel,
1931; Dukstra, 1991), Arabian Sea and Bay of Bengal (Smith, 1894, 1895a, 1904. 1906), Gull of
Aden and Zanzibar area (Knudsen, 1967). Now also New Caledonia. Present material living at

450-550 m.

Description. Shell slightly inequivalve, fragile, nearly
equilateral, up to ca. 25 mm high, somewhat higher than
wide, rather opaque, glossy, creamy, umbonal angle about
90'’, left valve somewhat more convex than the right.

Prodissoconch ca. 215 pm in height (Knudsen. 1967).

Left valve smooth, ornamented with concentric growth
lines, no radial slriations. Auricles rather small, without
sculpture, somewhat raised near margins.

Right valve sculptured with wide-set concentric lirae.

commencing at 3 mm shell height and extending to submar¬
ginal area, microscopic radial scratches between them. Auri¬
cles with very fine concentric striae, strong scales produced
on the marginal areas of hinge. Hinge line straight near
umbo, then rising near anterior and posterior dorsal margins.
Internal lirae generally 10 in number, sometimes 9 or II.
slightly nodulose at distal ends. Lirae of right valve somewhat
more strongly developed. No byssal notch or ctenolium.
lateral gape present.

Remarks. Other specimens seen from the western Paciltc dilfer slightly from the present
material, mainly in coloration (somewhat paler) and in having usually I or 2 more internal lirae. The
specimens are similar to the type material, although the concentric growth lines are weaker and there
are no radial striae on the left valve. Amussium weberi is similar in all features, and is interpreted as
junior synonym of Propeamussium caducum. P. nakazawai treated by Oyama (1951). Habe (1958,
1977) and Knudsen (1967) as a synonym of P. caducum is a nomen nudum (no description and not
compared with any other species).

Descriptions of the soft parts, food and reproduction are given by Knudsen (1967; 275).
Hayami (1988a) described shell crystallography.

P. caducum is the type species of Parvamussium (Flavamussium) . Kuroda & Habe (1981:62)
treated Flavamussium as a subgenus of, and Hertlein (1969: N350) as a synonym ol Propeamussium.
P. caducum is a typical Propeamussium.

18

HENK DIJKSTRA

Propeamussium maorium (Dell. 1956)
Figs 11-14

Parvamussium maorium Dell, 1956: 20, figs 30-31.

Other references:

Parvamussium maorium - Powell. 1979: 381, figs 93.1-2. Rombouts 1991- 69
Parvamussium maorum (sic) - Dell, 1962: 75; 1963: 206.

Type material. Holotype dead-taken, nmnz M9171, 5 paratypes nmnz M9169

Type locality. — Portobello " Alert ”,
Zealand, 476-640 m.

stn 54-17. Canyon A, ENE of Taiaroa Head, New

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn CP 387, 20°53' S, 160°52'E, 650-660 m 1 snm.

S“2: St" PE l5’ 20°51' s' 160°56' E’ 580-590 m, 1 lv, 1 rv. — Stn DE 15b,' 20°5T S, 160°55'
580-590 m, 3 lv, 1 rv.

New Caledonia, musorstom 4: stn CP 169. 18°54' S, 1 63°1 1 ' E, 590 m 1 db
Loyalty Islands, biogeocal: stn CP 232, 21°33'S, 166°27'E, 760-790 m 1 lv 2 rv.

musorstom 6: stn DW 469, 21°03' S, 167°34' E, 630 m, 1 spm. Stn DW 483 21°19' S 167°47'
600 m, I spm.

E,

E,

Kerm,?,STMBU1ION' S°Uth Islands °f Ne* Zealand> Chatham Islands (nmnz),

'imU ndv ,lt S Rnd5 dl N°rf° k IS a,n.d <Dijkstra- unpubl. data); now also the Chesterfield Islands
and Loyalty Basin. Present material living at 600-660 m.

Description. Shell somewhat inequivalve, fragile,
suborbicular. up lo ca. 20 mm high, semi-translucent left
valve brighter brownish or creamy than right, umbonal angle
about 90°. b

Prodissoconch ca. 210 pm in height.

Left valve somewhat more convex than right, covered with
prominent, irregularly arranged radial costae, that extend to
marginal area; the costae covered with fine lamellae, that
become more closely spaced near ventral margin. Auricles
small, equal in size, smooth.

Right valve with widely spaced concentric lirae. Auricles
unequal sculptured with fine concentric striae. Small scales
produced on hinge line near dorsal margin. Internal lirae
generally 10-12. sometimes a few secondary riblets developed
between primary riblets near anterior and posterior margins
Resi lifer triangular, elongate. Byssal notch small. Lateral
gape present, no ctenolium.

Remarks. - The present specimens agree very well with the type material. The closest species

ad^cosmeTnH (EA' Smith’ l906) from the SE Arabian Sea. The irregularly spaced

radial costae and concentric striae commence earlier on the disc of the latter species whereas P

Smhh rdptSSy at lhC SamC StagC °f gr0Wth- Another allied species is P. jeffreysii (EA

th“ I extend Lfr^Ct 1PPnfPLW5iCh haS, fmer scu,Pture (somewhat cancellate) on the left valve
at' maturity P of the d,sc’ and Promment concentric lamellae near the ventral margin

Although Dell (1956) placed P. maorium in Parvamussium , it is a typical Propeamussium.

Source : MNHN , Paris

BATHYAL PECTI NOIDEA FROM NEW CALEDONIA

19

Propeamussium meridionale (E.A. Smith, 1885)
Figs 15-18, 143-146

Amussium meridionale E.A. Smith, 1885: 316, pi. 24. figs 1-la.

Other references:

Amussium meridionale - Knudsen, 1967: 277. pi. 1, fig. 16, textfig. 17,

Propeamussium meridionale — Grau, 1959: 12, pi. 1. Knudsen, 1970: 94, pi. 12. figs 5-9, textfig. 58. Dell, 1990: 37.
Propeamussium ( Propeamussium ) meridionale - Dijkstra, 1990: 2, 9. pi. 1, figs 1-2.

Varlamussium (sic) meridionale - Powell, 1960: 175.

Verlamussium (sic) meridionale — Clarke. 1962: 60.

Type material. — Lectotype (Figs 143-6, H 13.8, L 13.4, D 4.0 mm) here designated, live
taken, bmnh 1887.2.9.3337 (corresponding to the original description and measurements),
2 paralectotypes bmnh 1887.2.9.3335/1-2, undamaged left valve (corresponding to the figured shell
from " Challenger ”, stn 146). Posterior margin of left valve and marginal apron of right valve of the
lectotype is somewhat damaged. — A. meridionale var. bmnh 1887.2.9.3336/1-4, 3 lv and 1 rv (stn 302)
does not belong to the type series according to iczn art. 72b(i).

Type locality. — " Challenger ”, stn 158, 50°01' S, 123° E, southern Indian Ocean, 3292 m.

Material examined. — The type material.

Chesterfield Islands, corail 2: stn DW 172, 18°26' S, 155°12' E, 1100 m, 1 spm.

New Caledonia, biocal: stn DS 04, 21° 15' S, 166°39' E, 2340 m, 5 spms, 1 lv. — Stn DW 53, 23”09 S,
167°42' E, 975-1005 m, 3 lv. — Stn DW 56, 23°34' S, 1 67°1 1' E, 695-705 m, 1 rv. — Stn CP 61,
24° 1 1 ' S, 167° 31' E, 1070 m, 3 lv, 4 rv. — Stn CP 63, 24°28' S. 168°07' E, 2160 m, 1 spm. — Stn DW
66, 24°55' S, 168°21' E, 505-515 m, 1 rv. — Stn DW 70, 23°24' S, 167°53' E, 965 m, 4 lv, 1 rv. -
Stn DW 80, 20°31' S, 166°48' E, 900-980 m, 4 rv. — Stn DS 98, 21°24' S, 166°29' E, 2365-2470 m,
3 spms.

musorstom 4: stn DW 160, 18°42' S, 163° 13' E, 675 m, 4 lv, 2 rv. — Stn CP 169, 18°54' S, 163°H' E,
600 m, 6 lv, 2 rv. — Stn CP 178, 18°56' S, 163° 12' E, 520 m, 1 lv.

Loyaltv Islands, biogeocal: stn CP 250, 21°24' S, 166°28' E, 2350 m, 1 spm. — Stn CP 290, 20°36' S,
167°03' E, 920-760 m, 2 spms, 3 lv, 2 rv. — Stn DW 296, 20°38' S, 167° 10' E, 1230-1270 m, 3 lv,
2 rv. — Stn CP 297, 20°38' S, 167° 10' E, 1230-1240 m, 1 spm, 1 lv.

musorstom 6: stn DW 396, 20°48' S, 167°00' E, 1400 m, 3 lv, 1 rv. — Stn DW 397, 20°47' S,
1 67°05' E, 380 m, 1 lv. — Stn DW 488, 20°49' S, 167°06' E, 800 m, 4 spms, 37 lv, 29 rv.

Distribution. — Smith (1885) recorded this species from three widely scattered localities:
South of Western Australia, East of Marion Island (southern Indian Ocean), and West of Patagonia
(southeastern Pacific). Knudsen (1967) added several new records from the northern Arabian Sea,
Zanzibar area, Maidive Islands, the Gulf of Aden, and the Kermadec area (NE of New Zealand).

Dijkstra (1990) mentioned a new record from the
Islands, New Caledonia and the Loyalty Islands

Dlscription. — Shell fragile, somewhat inequivalve,
nearly circular, up to ca. 15 mm high, left valve slightly more
convex than right, auricles equal, pellucid white, umbonal
angle about 120°.

Prodissoconch ca. 200 pm in height (Knudsen, 1967).

Left valve sculptured with delicate radial lirae, commencing
at 3 mm shell length, extending almost to ventral margin.
Prominent concentric lamellae extend to ventral margin,
intersected by radial lirae to produce a somewhat cancellate
sculpture. Auricles equal, sculptured with radial striae.

Flores Sea (Indonesia). Now also the Chesterfield
Present material living at 760-2470 m.

Some delicate lamellae near anterior and posterior mar¬
gins.

Right valve covered by regular concentric lirae. which are
more close-set near margins. Auricles differently sculptured:
Anterior auricle with stronger radial lirae than posterior,
concentric lamellae near dorsal margin. Interior somewhat
iridescent, hyaline near margins. Internal riblets variable in
number, generally 12, without distinct terminal nodules.
Hinge line straight with delicate scales on dorsal margin.
Resilifer triangular. No byssal notch, or ctenolium.

Source MNHN . Paris

20

HENK DIJKSTRA

Eigs 15-22 15-18. Propeamussium meridionale, musorstom 6: stn DW 488, 18.0 x 18.5 mm (Iv) 14 5 x 14 2 mm (rv)

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

21

Remarks. — The present specimens correspond very well with the type material, although the
sculpture on the left valves is somewhat more pronounced, while a few (1-3) internal riblets are more

strongly developed. .

Powell (1960: 175) placed P. meridionals in Varlamussium , but all its conchological characters

indicate that it is Propeamussium.

Hicks & Marshall (1985: 227) describe the food of this species.

Propeamussium ruhrotinctum (Oyama, 1951)
Figs 23-26

Parvamussium (Parvamussium) ruhrotinctum Oyama, 1951: 81, pi. 13, figs 8-10.

Synonym:

Propeamussium (Propeamussium) Stella Wang, 1984: 600, 602, pi. 1, figs 11-14. (Syn. nov.)

Other reference:

Propeamussium ruhrotinctum — Hayami, 1988b: 80.

Type material. — P. ruhrotinctum: probably in the private collection of Dr K. Oyama at
Toba (not seen). P. Stella: holotype ioas m25778, paratype ioas m25779.

Type locality. P. ruhrotinctum: Shikoku, Gulf of Tosa, Japan, depth not mentioned,
alive?. — P. Stella: South China Sea, 19° N, 112.5° E, 290 m.

Material examined. — New Caledonia. “ Vauhan " 1978-79: stn 2, 22" 17 S, 167 14 E,
425-430 m, 4 lv. — Stn 3, 22°17'S, 167°12'E, 390 m, 2 lv. — Stn 33, 22°33' S, 166°25' E, 290-
350 m, 1 lv.

biocal: stn DW 104, 21°30' S. 166°21' E, 375-450 m, 1 lv.

musorstom 4: stn DC 235, 22°13' S, 167°12' E, 405-415 m, 5 lv, 2 rv. — Stn DW 244, 22 02 S,
167"08' E, 435-445 m, 2 lv. — Stn CC 246, 22"08' S, 167°11' E, 410-420 m. 4 lv, 4 rv.

Loyalty Islands, biogeocal: stn DW 253, 21°3TS, 166°28' E, 310-315 m, 1 lv.
musorstom 6: stn DW 391, 20°47' S, 167"05' E, 390m, 1 lv, 4 rv. — Stn DW 406, 20 40, S, 67 06 E,
373 m, 3 lv, 5 rv. — Stn DW 407, 20°40' S, 167°06' E, 360 m, 1 lv. Stn CP 408, 20"41 S, i67 ' 07 E,
380 m, 2 spms. - Stn DW 410, 20°38' S, 167°06' E, 490 m, 1 spin, 1 lv, 1 rv. — Stn DW 41 1, 20' 40 S,
167°03' E, 424 m, 1 1 spms, 22 lv, 13 rv. — Stn DW 412, 20°40' S, 167° 03' E, 437 m, > 50 spms
Stn DW 413, 20°40' S, 167° 03' E, 463 m, 1 spm. 2 rv. — Stn CP 415, 20040' S, ,167 03 E, 46i
m, 4 spms. Stn DW 416, 20°42' S, 166°59' E, 343 m, 3 rv. — Stn DW 426, 20°24 S, 166 22 E,
610 m, 1 lv, 1 rv. — Stn DW 428, 20°23' S, 166° 12' E. 420 m, 1 spm, 1 rv. — Stn DW 447, 20 54 S,
167° 19' E, 460 m, 1 lv. — Stn DW 458, 21°00' S, 167°29' E, 400 m, 1 rv. — Stn DW 459, -T 01 S,
167° 31' E. 425 m, I lv. — Stn CP 464, 21°02' S, 167°31' E, 430 m, 3 lv, 1 rv.

Distribution. — Oyama (1951) recorded this species from Shikoku and Kyushu, Japan.
Wang (1984) added a new record from South China Sea. Now also New Caledonia and the Loyalty

Islands. Present material living at 380-490 m.

Description. Shell fragile, suborbicular, inequivalve,
inequilateral, up to ca. 20 mm high, left valve somewhat more
convex than right. Valves gaping at lateral margins. Auricles
relatively small, unequal. Umbonal angle about 90".

Prodissoconch ca. 200 pm in height.

Left valve covered with minute concentric lamellae in late
ontogeny near periphery, otherwise smooth and glossy with a
few concentric plications, one or two delicate radial plicae
near posterior margin. Shell transparent, with light orange
patches and small white dots. Auricles smooth. Hinge line
straight.

Right valve covered with regular concentric lirae and
interstitial microscopic radial scratches. Auricles with concen¬
tric striae and somewhat serrated on dorsal margin. Shell
whitish, nearly opaque. Internal lirae generally 10. an
auricular lira on each side, commencing directly below the
resilifer, extending to submarginal area, slightly nodulous at
distal ends. Internal lirae of right valve somewhat broader.
Resilifer triangular, elongate. Outer ligament rather broad.
Adductor scar large, nearly circular, with whitish spots. Small
byssal notch, no ctenolium.

22

HENK DIJKSTRA

Figs 23-30 ,2r^'26: Propeamussium rubrotinctum , musorstom 6: stn DW 412, 21.8 x 20 5 mm (dh) - 23 lefi vilv,- ,,,Prillr

stnCfc'Wi TZ7 mm (db)ri8h,27al|^eXt,eri0r- ^ ^ SSSSli

30, right valve, interior ( ’' “ 'efl Va‘Ve’ eX,en°r- “ 28‘ left valve' inlerior- ~ 29- right valve, exterior. -

Source : MNHN , Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

23

Remarks. — Wang (1984: 603) compared Propeamussium Stella with P. caducum, and
observed that these species differ in shape, the number of internal lirae, and in having a glossy surface
and a colour pattern, but he overlooked P. steindachneri (Sturany, 1901) from the Red Sea and Gulf
of Oman, and P. rubrotinctum (Oyama, 1951) from Japan. Both of the latter are similar to P. Stella,
although P. steindachneri is smaller (ca. 14 mm) and has fewer internal lirae (8), but other shell
features are very similar, including the coloration.

Oyama (1951) placed P. rubrotinctum in Parvamussium s. str., but Hayami (1988b) indicated
that it should be refered to Propeamussium, despite its unequal auricles and weakly developed byssal
notch.

Propeamussium sibogai (Dautzenberg & Bavay, 1904)
Figs 19-22

Amussium sibogai Dautzenberg & Bavay, 1904: 207, figs 1-4.

Other references:

Amussium sibogai - Dautzenberg & Bavay, 1912: 31, pi. 28, figs 1-4.

Amussium cf. sibogai - Barnard, 1969: 655. pi. 1, figs a-d.

Luteamussium sibogai — Oyama. 1951: 82, fig. 1. — Kira, 1967: 138, pi. 49, fig. 14.

Luteamusium (sic) sibogae (sic) - Kosuge, 1985: 58, pi. 23, fig- 12.

Propeamussium sibogai - Knudsen, 1967: 272, pi. 1, figs 23-24. Abbott & Dance, 1982. 303, fig.
Whitehead, 1992: pi. 6, no. 34, fig. 34. rnn , , _ , . . ,r .

Propeamussium (Propeamussium) sibogai - Wang, 1984: 599, pi. 1, figs 1-2, textlig. I.

Lamprell &

Type material. — Holotype, live taken, zma 3.04.001.

Type locality. — “ Siboga ", stn 12, 7°15' S, 115°15' E. Bali Sea, Indonesia, 289 m.
Material examined. — Chesterfield Islands, musorstom 5: stn DC 381, 19 37 S, 158 46 E,

corail 2: stn DE 16, 20°48' S, 160°56' E, 500 m, 1 lv, 1 rv. — Stn DC 169, 18°21'S, 155°20'E,

575 m, 2 rv. , „ ,-n

New Caledonia, musorstom 4: stn CP 169, 18°54' S, 163°1 T E, 600 m, 6 spms, 1 rv. Stn CP
18°57' S. 163° 12' E, 485 m, 1 spm. 0

Loyalty Islands, biogeocal: stn DW 292, 20°28' S, 166°48' E, 465-470 m, 1 rv. - Stn DW 308,
20°40' S, 166°58' E, 510-590 m, 1 rv.

musorstom 6: stn DW 410, 20°38' S, 167°06' E, 490 m, 4 spms, 2 lv, 5 rv. — Stn DW 413, 20 40 S,
1 67°03' E, 463 m, 1 spm. Stn DW 415, 20°40' S, 167°03' E, 461 m, 2 spms. — Stn CP 464, 21 0_ S.
167°31' E, 430 m, I spm. — Stn CP 465, 21°03' S, 167°32' E. 480 m, 6 spms, 3 lv, 4 rv. — Stn CP
467, 21°05' S, 167°32' E, 575 m, 2 spms, 4 lv, 4 rv. — Stn DW 487, 21°23' S, 167 46 E, 500 m, 2 lv.
volsmar: stn DW 37, 22°23' S, 168°43' E, 500-550 m. 3 rv.

Distribution. — Dautzenberg & Bavay (1912) recorded this species from the Bali Sea
(Indonesia). Oyama (1951), Kira (1967) and Okutani et al. (1989) mentioned it from the southern
Japanese waters to Indonesia. Knudsen (1967) enumerated several records from Indonesia, the
Philippines, the Arafura Sea, and off Durban (S. Africa). Kosuge (1985) added a new record from
the Timor Sea (Northwestern Australia). Now this species is also known from New Caledonia and
the Loyalty Islands. Present material living at 430-575m. Knudsen (1967) indicated a broader vertica
range (183-710 m).

Description. Shell fragile, nearly orbicular, inequivalve,
somewhat inequilateral, oblique, up to ca. 55 mm high, left
valve slightly more convex than right, gaping rather strongly,
transparent, auricles rather small and equal in size, umbonal

angle about 125°; left valve creamy brown, the right cream
coloured.

Prodissoconch ca. 210 pm in height (Knudsen, 1967).

Left Vahe covered with a few delicate concentric growth

24

HENK DIJKSTRA

lines, a few minute concentric lamellae sometimes present
near ventral margin. Delicate radial lirae developed near
posterior margin. Auricles smooth and somewhat raised on
lateral margins. Hinge line straight. Internal lirae generally 8,
deep brown on left valve, whiter and broader on right.
Marginal lirae with 4 or 5 small nodules that are visible
through lateral gape.

Right valve covered with widely spaced concentric lirae that
are much weaker at periphery, and microscopic interstitial
granules. Marginal apron of right valve often missing due to
the very thin layer of prismatic calcile (Hayami, 1988a:
fig. 4). Auricles smooth. Resilifer triangular, elongate. No
byssal notch, or ctenolium.

Remarks. — The present material is very similar to the holotype from the Bali Sea, although
they have 1 or 2 additional internal riblets.

Dautzenberg & Bavay (1904) incorrectly indicated the type locality as “Celebes Sea” which
they subsequently corrected (1912: 31).

P. sibogai is the type species of Luteamussium, which was treated by Hertlein (1969: N350)
as a synonym of Propeamussium. Indeed, P. sibogai is a typical Propeamussium.

Descriptions of the soft parts, reproduction and food are given by Knudsen (1967: 272).

Propeamussium watsoni (E.A. Smith, 1885)

Figs 27-30, 123-124

Amussium watsoni E.A. Smith, 1885: 309. pi. 22, figs 8-8c.

Synonym:

Propeamussium watsoni hayonnaisense Okutani. 1962: 15-16, pi. 2. figs 1-2; 1966: 8.

Other references:

Amussium alcocki - Thiele & Jaeckel. 1931- 8

Amussium watsoni - Clarke. 1962: 61. Knudsen. 1967: 280, pi 1 fig 18

Propeamussium watsoni - Kiseleva, 1971: 221. Abbott & Dance,' 1982: 303, fig. - Hayami. 1988a: 476, figs 2-6.

Type material. — A. watsoni : lectotype (Figs 123-4, H 51.8, F 50.0, D 9.5 mm) here
designated, bmnh 1887.2.9.3307, 2 paralectotypes bmnh 1887.2.9.3308/1-2. — P. watsoni bavonnai-
sense: holotype, alive, nsmt Mo.62759 and 5 paratypes nsmt Mo.62760.

Type locality. — A. watsoni : “ Challenger ”, stn 218, 2°23' S, 144°04' E, NE of New Guinea,
alive, 195/ m — - P. watsoni bayonnaisense : 24 miles off Bayonnaise Rocks, 32°00.0' N, 140"21.4' E,
Japan, 2140-2160 m.

.... Material examined. — Chesterfield Islands, musorstom 5: stn DC 321, 21°20' S 158°02' E

1 57-sr’ f Q7(? rV' « Stn CP 32c 21 °l8< s' 157057 E- 970 m’ ca- 90 sPms- — Stn CP 32'4- 2 1 0 1 5' S,’
i 'S iSS'p irS' 7 Sm CCo365' 19°42'S- l58°48'E, 710 m, 3 spins. - Stn CC 366,

Stn^CC^383,8 19°40' s!^15£^46 E^OO-Ils'm ^"rv ^ ’ * '58°53'E' 830'855 25 -

“^AL: Stn CPC°- 23°08'S’ 166°40'E" 1140 ">• 1 SP“- - St" CP 31. 23°07' S,
66 50 E 850 m, 15 spms. — Stn CP 75, 22° 18' S, 167l)23' E, 825-860 m, 4 spms

K ?6ar">9%r^rOM,o6: Stn ? 427 20°23'S’ 166°20'E* 800m, 13 spms. - Stn CP 438,
-u IS 8, 166 20 E, 780 m, 42 spms, 2 rv.

Philippines, musorstom 2: stn 69, 14°06' N, 120°03' E, 1800-1950 m, 2 spms.

Distribution — Smith (1885) described this species from NE of New Guinea and
subsequently recorded it from the Bay of Bengal and Laccadive Sea. Thiele & Jaeckel (1931)
recorded it from the Gulf of Aden, and Okutan, (1962) from SE Japan. Knudsen S
records from the Arabian Sea, Zanzibar area and the Strait of Malacca. Kiseleva (1971) added a new

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

25

record from the Red Sea. This species is also found in the Chesterfield Islands, New Caledonia and
the Loyalty Islands. Present material living at 650-1140 m.

Description. - Shell inequivalve, nearly circular, ca. 60
mm high, slightly higher than wide, somewhat inequilateral,
umbo mil angle ca. 125°, opaque, left valve milky-white, right
valve creamy.

Prodissoconch ca. 240 pm in height.

Left valve more convex than right, with concentric lamellae
near the ventral margin and delicate radial lirae that
commence at ca. 3 mm shell length and extend to central part
of disc, lamellae and lirae variable. Auricles equal, concentric
lamellae prominent on anterior, finer and more closely-set on

posterior. Generally 10 interior lirae and a small auricular
lira on each side, commencing just below resilifer and
extending to pallial line.

Right valve with fine regularly concentric lirae and granu¬
late interstitial microsculpture (prismatic calcite layer). Auri¬
cles with concentric lirae, anterior auricle with a few radial
lines near suture, absent on posterior auricle. Prominent
scales on anterior and posterior dorsal margins of auricles.
Resilifer triangular, erect. No byssal notch, or ctenolium.

Remarks. — The present specimens are similar to the type material in shape, sculpture and
coloration, although the concentric lirae of the left valve commence closer to the ventral margin, and
the radial lirae are somewhat more delicate. Both of the latter two characters are variable.

I follow Knudsen (1967: 281) in interpreting P. watsoni bayonnaisense as a form of P. wat-

soni.

Genus Parvamvsswm Sacco, 1897

Parvamussium Sacco. 1897a: 102. Proposed as a subgenus of Amussium Herrmannsen 1846 (unjustified ^emendation of
Amusium Roding, 1798); no diagnosis given, but type species designated; Sacco. 1897b: 48 (diagnosis). T>pe species
(OD): Pecten (Pleuronectes) duodecimlamellatus Bronn, 1832; Upper Miocene, northern Italy.

Synonyms:

Variamussium Sacco, 1897a: 102. Proposed as a subgenus of Amussium: no diagnosis given but type species designated^ Sacco.
1897b: 49 (diagnosis). Type species (OD): Amussium cancellatum E.A.Smith, 1885; Recent, off Bermuda, W Atlantic,

Ctenamusium Iredale, 1929: 164. Type species: Amusium thetidis Hedley, 1902; Recent, ofl New South Wales, Australia, 115-

Glyptamusiurii Iredale, 1939: 370. Type species (OD): Amussium torresi E.A.Smith, 1885; Recent. E of Cape York, Queensland.

Squam amussium Oyama, 1944: 245. Proposed as a subgenus of Propeamussium. Type species (OD): Amussium squamigerum

E.A.Smith. 1885; Recent, off E Puerto Rico, West Indies, 713 m.

Polvnemamussium Habe. 1951: 72. Proposed as a subgenus of Parvamussium. Type species (OD): Pecten mtuscostalus
Yokoyama, 1920; Pleistocene, Miura City, Kanagawa Prefecture, Japan.

Diagnosis. — Shell inaequivalve, orbicular to oblique, usually small to ca. 20 mm, laterally
compressed, lateral gape absent; left valve usually strongly sculptured with radial and/or concentric
riblets or striae, right valve with concentric lamellae; auricles unequal; byssal notch well-developed,
no ctenolium; internal lirae extend to submarginal or marginal region.

Distribution. — Cretaceous to Recent. Worldwide, living in 18-2110 m.

Remarks. — The characters separating Parvamussium from Propeamussium are summarized
in Table 1.

Grau (1959; 15) and subsequently Hertlein (1969: N350) treated Parvamussium as a
subgenus of Propeamussium and enumerated Variamussium. Ctenamusium. Glyptamusium. Xenamus-
sium, Squamamussium, Polynemamussium. Bathymussium and Micramussium as synonyms ol
Parvamussium. Hertlein (1969: N350) also mentioned in the synonymy “Grapt amussium Oyama
1944”, which is an error for Glyptamusium. Bathymussium and Micramussium are here considered
synonyms of Propeamussium , ami Xenamussium a synonym of Cyclopecten.

26

HENK DIJKSTRA

Parvamussium multiliratum sp. nov.
Figs 31-34, 91-92

Type material. — Holotype mnhn. Paratypes: II mnhn, 1 ams C201711, 1 lid, 1 usnm.

Type locality. — Southern New Caledonia, biocal, stn CP 72 22°10'S 167°33'E
2100-2110 m.

Material examined. — New Caledonia, biocal: stn CP 72, 22° 10' S, 167°33' E, 2100-21 10 m
2 spm. 1 lv (holotype and paratypes).

Loyalty Islands, biogeocal: stn KG 240, 21°29' S, 166°27' E, 1520 m, 1 rv (paratype). Stn CP 243
21°28' S, 166°26' E, 1820 m, 1 lv (paratype). — Stn CP 260, 21°00' S, 167°58' E, 1820-1980 m 2 spins
(paratypes). — Stn CP 272, 2 POO'S, 166°57'E, 1615-1710 m, 1 spm (paratype). — Stn CP 273,
21°02' S, 166°57' E, 1920-2040 m, 5 spms (paratypes: 1 hd, 1 ams, 2 mnhn, 1 usnm). — Stn CP 317’
20°48' S, 166"53'E, 1620-1630 m, 1 lv, 1 rv (paratypes).

Distribution. New Caledonia, 1520-2110 m, living in 1615-2110 m.

Description. — Shell rather small, fragile, orbicular,
inequivalve, semi-transparent white, up to ca. 9 mm high, left
valve slightly more convex than right, auricles subequal,
umbonal angle about 1 10°.

Prodissoconch ca. 220 pm in height.

Left valve sculptured with widely spaced concentric lamel¬
lae that commence at 3 mm shell height, and extend to
ventral margin. Irregularly spaced, delicate radial riblets
arising near submarginal region, a few commencing earlier.
Umbonal region glossy with some white spots, otherwise
transparent and dull due to microscopic granules. Auricles
with concentrically lamellate riblets that are closer near
lateral margins.

Right valve with regularly spaced concentric lirae, some¬
what interrupted near submarginal region by a disturbance of
growth. Auricles with concentric lamellae that are somewhat
stronger on anterior, and more closely spaced on posterior.
Hinge line straight, with some scales near margins.that are
more prominent on anterior. Byssal fasciole small. Each valve
with 14 internal lirae plus 2 rudimentary interstitial lirae: one
auricular lira on both auricles. External sculpture clearly
visible from the interior. Resilifer triangular. Byssal notch
present, no ctenolium.

Dimensions of the holotype: H 7.9, L 8.9, D 2.4 mm.

Remarks. The most closely related species is Parvamussium permirum (Dautzenbera. 1925)
recorded from the Bay of Biscay (E Atlantic) and adjacent abyssal depths. P. permirum has more
regularly spaced radial riblets on the left valve, and fewer concentric lamellae. Internal lirae generally
12 with 2-4 additional rudimentary interstitial lirae and the hinge line has fewer scales. P. retiaculum
is somewhat similar to P. multiliratum in sculpture, although the latter has fewer radial riblets, and
a more strongly orbicular shape, with more numerous internal lirae. Dautzenberg (1925: 1 1) placed

P. permirum in Amussium (Variamussium), but all the conchological characters are similar to
Parvamussium.

Etymology. So named because the internal lirae are more numerous than usual for a
propeamussnd (Lat. liratus, adj. = with lira(e)).

Parvamussium pauciliratum (E.A. Smith, 1903)
Figs 107-110, 151-152

Amussium paucilirata E.A. Smith, 1903: 622, pi. 36, figs 23-24.

Other reference:

Parvamussium pauciliratum - Dukstra. 1991: 14, figs 23-24.

Source . MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

27

51-38. 31-34. Parvamussium mulliliraium , holotype, 7.9 x 8.9 mm (db). 31. left valve «tenor 32’ ’

interior. 33. rieht valve, exterior. 34. right valve, interior. - 35-38. P. reUaculum , holotype. 7.3 * : 7.2 mm (db).
35. left valve, exterior. — 36. left valve, interior. - 37. right valve, exterior. - 38. right valve, interior.

28

HENK DIJKSTRA

Type material. — Lectotype (Figs 151-2, H 7.5, L 7.4, D 2.0 nun), here designated, live taken,
bmnh 1903.9.17.17; 2 paralectotypes bmnh 1903.9.17.18/1-2. Although the original registration
number indicated 4 specimens, only three syntypes are now present.

Type locality. S Nilandu Atoll, Maidive Islands, 2-66 m.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DW 263, 25°21'S, 159°46'E, 150-225 m, 1 Iv, 1 rv.

New Caledonia, lagon: stn 348, 22°42' S, 166°55' E, 45 m, I lv. — Stn 622, 22°02' S, I66°53'E.
67 m, 1 rv. — Stn 698, 21°29'S, 166°09'E, 40-43 m, I spin. — Stn 729, 21°19'S, 1 65°54' E, 42-
45 m, 1 spm. — Stn 853, 20°41' S, 165°07' E. 27 m, 1 rv. — Stn 873, 20°39' S, 164°46' E, 27 m, 1 spm.
Stn 1191, 19°35'S, 163°38' E. 45 m, 1 spm.

Distribution. This is a shallow-water species, which is included here for a comprehensive
treatment of the Propeamussiidae of New Caledonia. Beside the type locality, the Maidive Islands,
it has been recorded from Indonesia by Dijkstra (1991). Present material from New Caledonia alive

in Z/-45 m. Also shells in the Chesterfield Islands

Description. — Shell rather small, semi-transparent, whi¬
tish. sub-orbicular, inequivalve, up to ca. 8 nun high, auricles
nearly equal in size, right valve more convex than left,
umbonal angle about 105°.

Prodissoconch ca. 200 pm in height.

Left v alve glossy, with a few minute growth lines and covered
with microscopic granules. Auricles smooth, sometimes [see
Remarks] with delicate concentric lirae near anterior margin.

m 150-250 m.

Right valve glossy, smooth, apart from microscopic granu¬
les. Anterior auricle with minute concentric lirae. Hinge line
straight.

Internal lirae rudimentary. I or 2. usually a small auricular
lira on each side. Outer ligament well developed. Left valve
creamy-white, stained with white dots, right valve
transparent-white. Resilifer triangular. Byssal notch very
small, no ctenolium.

Remarks. — The present specimens are similar in all aspects to the type material from the
Maldive Islands, although microsculpture is sometimes variable. The New Caledonian specimens,
which are from shallower depths, sometimes have microscopic divergent scratches near the
postero-dorsal margin on the left valve. Microscopic granules are generally present on both valves.
Juveniles often lack the internal rudimentary liration.

Parvamussium retiaculum sp. nov.

Figs 35-38

Type material. Holotype mnhn. Paratypes: 25 mnhn, 2 ams C201712, 2 hd, 2 nmnz
M268536, 2 nsmt, 2 usnm.

Type locality. - Southern New Caledonia, biocal, stn DW 51, 23°05' S, 167°45' E, 680-

700 m.

Material examined. — New Caledonia, biocal: stn DW 33, 23° 10' S, 167" 10' E 675-680 m
2 lv (paratypes). — Stn DW 36, 23°09' S, 167°11' E. 650-680 m, 2 lv (paratypes). — Stn DW 48,
23°00' S, 167°29' E, 775 m, 2 lv (paratypes). Stn DW 51. 23°05' S, 167°45' E, 680-700 m, 7 spms,
8 lv, 15 rv (holotype and paratypes: 2 ams, 2 hd, 19 mnhn, 2 nmnz, 2 nsmt, 2 usnm).

Distribution. Southern New Caledonia

Description. Shell small, fragile, suborbicular. inequi¬
valve, up lo ca. 7 mm high, left valve slightly more convex
than right, semi-transparent white, auricles of unequal size
umbonal angle about 100°.

Prodissoconch ca. 210 pm in height.

650-775 m, living 680-700 m.

Left valve sculptured with 14 prominent, widely spaced,
concentric lamellae, starting weakly at ca. 2 mm shell height,
extending to ventral margin. Delicate, regularly spaced radial
riblets, commence at about 3 mm shell height, and extend to

Source MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

29

ventral margin. Exterior of disc with semi-reticulated sculp¬
ture produced by concentric lamellae and intersecting radial
riblets. Anterior auricle stronger than posterior sculptured
with close-set. irregularly spaced concentric lamellae: poste¬
rior auricle with identical though weaker sculpture. Hinge

line straight. , ,

Right valve with regularly spaced concentric lirae. close-set
near umbonal area and becoming more widely spaced toward
ventral margin. Anterior auricle more strongly sculptured

near lateral margin with concentric lamellae. Hinge line
somewhat elevated due to strong concentric lamellae. Byssal
fasciole present.

Each valve with 12 internal lirae and 1 auricular lira that
commence rather late and extend to near the periphery: one
rudimentary interstitial lira on left valve near the ventral
margin. Outer ligament well developed and rather broad on
posterior auricle. Small byssal notch, no ctenolium.

Dimensions of the holotype: H 7.3. L 7.2, D 1.9 mm.

Remarks. — P. retiaculum is somewhat similar to P. multiliratum , although the latter is
slightly larger and more orbicular, while the auricles are more equal and smaller, fewer radial riblets
are developed on the left valve, there are more internal lirae, and the hinge line of the right valve is
straight. P. multiliratum lives at much great depths (lower bathyal to upper abyssal).

The right valve of P. torresi is somewhat similar to that in P. retiaculum, although the internal
lirae commence earlier, and are fewer in number (generally 10). Moreover, the left valve of P. torresi
is glossy and nearly smooth, whereas P. retiaculum is prominently sculptured and dull.

The left valve of P. thetidis is covered with prominent irregularly spaced, radial costae and
delicate close-set concentric lamellae, whereas P. retiaculum has prominent concentric lamellae, and
delicate radial riblets.

Etymology. — From the cancellate sculpture of the left valve (Lat. retiaculum , n. = net,
lattice-work, wattle-work).

Parvamussium retiolum sp. nov.
Figs 39-42, 97

Type material. — Holotype mnhn. Paratypes: 36 mnhn, 2 ams C201713, 2 hd, 2 nmnz
M268537, 2 nsmt, 2 usnm.

Type locality. — Chesterfield Islands, musorstom 5, stn CP 363, 19°47' S, 158°44'E,
685-700 m.

Material examined. — Chesterfield Islands, musorstom 5: stn DW 340, 19°48' S, 158°40' E,
675-680 m. 1 spm, 1 lv. 3 rv. - Stn DW 341. 19°45' S, 158”43' E. 620-630 m i - spins 3 lv 3 rv.

Stn DC 357, 19°37' S, 158°45' E, 630 m, 2 lv, 2 rv. — Stn DC 358, 19' 38 S, 158 47 E, 680-700 m
4 spins, 1 lv, 5 rv. - Stn CP 363, 19°47 S, 158"44'E, 685-700 m, 41 spm, 6 Mv (holotype and

paratypes). — Stn CP 364, 19°45' S, 158°46' E, 675 m, 1 spm. — Stn DC 380, 19 37 S, 158 43 E,

555-570 m, 1 lv. , , „

New Caledonia, biocal: stn DW 56, 23°34' S, 167° I T E, 695-705 m, lv, 7 rv. . ,.0]n, P

musorstom 4: stn CP 158, 18°49'S, 163°15'E, 620 m, 1 lv. — Stn DC 168, 18 48 S, 163 10 E,

720 m, 1 spm.

Distribution. New Caledonia and Chesterfield Islands, 555-720 m, living 620-720 m

Description. - Shell rather small, inequivalve. inequila¬
teral, up to ca. 16 mm high, left valve somewhat more convex
than right, translucent 'white, auricles unequal, umbonal
angle about 95".

Prodissoconch ca. 220 pm in height.

Left valve covered with delicate concentric lamellae cros¬
sing closely spaced radial riblets. somewhat coarser near
umbonal area, finer near ventral margin. Anterior auricle
well developed, with many fine radial riblets that are crossed

near margin by fine concentric lamellae: posterior auricle
similary sculptured. Hinge line straight.

Right valve with regular concentric lirae. closely spaced
near umbonal area, becoming more widely spaced towards
the ventral margin. Microscopic interstitial radial scratches
near margins. Auricles strongly developed, with concentric
lamellae that are more prominent anteriorly. Strongly deve¬
loped scales on antero-dorsal margin. . . ..

Internal lirae 10. plus 1 posterior auricular lira and 4 rudi-

30

HENK DIJKSTRA

Figs 39-46. — 39-42 Parvamussium retiolum, holotype, 16.0 x 14.8 mm (db). — 39. left valve, exterior. 40. left valve
intQn°nT 41' ,r',uU val<v,e-e*ten?r- 42- nght valve, interior. 43-46. P. scilulum , musorstom 6: stn DW 462
interior 4 mm ' 43‘ eft V3 VC’ exlenor- 44- left valve- interior. 45, right valve, exterior. 46. right valve.

Source :

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

31

mentary auricular lirae on left valve and 3 on right, ends nodose.
Resilifer triangular, elongate. Strongly developed outer

ligament near resilifer. Byssal notch small, no ctenolium
Dimensions of the holotype: H 16, L 14.8, D 3.5 mm.

Remarks. - Parvamussium retiolum is somewhat similar to P. thetidis, which also occurs in
the study area (see below). P . retiolum is larger, with a fine cancellate sculpture on left valve; the latter
species has irregularly spaced, strong radial costae crossed by delicate concentric lamellae. Auricles
and right valve of P. retiolum more sculptured with closely spaced, concentric lamellae. Internal lirae
variable, somewhat similar to P. thetidis.

Parvamussium retiolum differs from P. retiaculum in its more delicate cancellate sculpture on
left valve, and in having more closely spaced, concentric lamellae on right valve. Internal lirae of P.
retiolum start earlier.

In all aspects P. retiolum is representative of Parvamussium.

Etymology. — From the cancellate sculpture of the left valve (Latin retiolum , n. = small net).

Parvamussium scitulum (E.A. Smith, 1885)
Figs 43-46, 153-154

Amussium scitulum E.A. Smith, 1885: 312, pi. 23, figs 4-4b.

Synonym:

Amusium ( Propeamusium) (sic) scitulum var.? cmadoritinclum Kuroda, 1931: 77, figs 81-82.

Other references:

Ctenamusium (Ctenamusium) cmadoritinclum - Oyama, 1951: 80, pi. 13, figs 3-4.

Ctenammssium (sic) cmadoritinclum - Habe, 1964: 173. pi. 53, fig. 4.

Ctenamussium (sic) scitulum - Kuroda & Habe. 1981: 63.

Propeamussium cmadoritinclum - HaYami, 1988a: 479, figs 2-1,2.

Propeamussium (Parvamussium) scitulum - Dijkstra, 1990: 3.

Parvamussium scitulum — Dijkstra, 1991: 16, figs 35-43.

Type material. — A. scitulum-. lectotype (Figs 153-4, left valve: H 4.4, L 4.5 mm), here
designated bmnh 1887.2.9.3319/1; 6 paralectotypes bmnh 1887.2.9.3319/2-7. — A. scitulum var.
cmadoritinclum : Kikuchi Shell Museum, Kuroda coll, (not seen).

Type locality. — A. scitulum: “ Challenger ", stn 188, 9°59' S, 139°42' E, S of New Guinea,
dead. 51m. A. scitulum var. cmadoritinclum: Yakushima Island, Kagoshima Prefecture, Kyushu.
Japan.

Material examined, - Chesterfield Islands, musorstom 5: stn DW 258, 25°32' S, 159°46' E,
300 m, 1 spm. — Stn DW 270, 24°48' S, 159° 34' E, 223 m, 3 lv, 3 rv. — Stn DW 285, 24°09' S,
1 59°34' E, 245-255 m, 1 lv. — Stn DW 296, 23° 12' S, 159°36' E, 278 m, 1 lv.

New Caledonia. “Vauhan" 1978-79: stn 40, 22°30' S, 166°24' E, 250-350 m, 17 lv, 21 rv.
biocal: stn DW 64, 24°48' S, 168°09' E, 250 m, 3 spms, 3 lv.
smib 5: stn DW 96, 23°00' S, 168°19' E, 245 m, 1 lv.

Loyalty Islands, musorstom 6: stn DW 462, 21°05' S, 167°26' E, 200 m, 3 spms, 4 lv, 1 rv.

Distribution. — Smith (1885) described this species from the southern area of New Guinea.
Kuroda (1931) mentioned it from Japanese waters, whereas Kuroda & Habe (1981) recorded a
larger distribution from Honshu (Japan) to Indonesia. Dijkstra (1990, 1991) summarized several
records from the Indonesian Archipelago. Now it is also known from the New Caledonian region.
Present material living at 200-300 m.

Source :

32

HENK DIJKSTRA

Description. — Shell rather small, sub-orbicular,
compressed, inequivalve. inequilateral, right valve more
convex than left, up to ca. 10 mm high, right valve
semi-transparent white, left valve stained and opaque, auri¬
cles rather small and subequal, umbonal angle about 120°.

Prodissoconch ca. 210 pm in height.

Left valve flat, weakly sculptured with delicate, variable
radial striations, somewhat stronger near posterior margin.
Microscopic close-set concentric striae sometimes developed
on disc, somewhat more widely spaced near umbonal area.

Anterior auricle with prominent concentric lamellae close to
Hank of disc, decreasing in prominence or absent near
antero-dorsal margin; posterior auricle covered with fine
radial and concentric striations, sometimes nearly absent.

Right valve either weakly sculptured with a few growth
lines or completely smooth. Internal lirae extending to
submarginal area, and generally 9 or 10, sometimes with I or
2 rudimentary interstitial lirae. Resilifer triangular. Hinge line
straight. Byssal notch present, no ctenolium.

Remarks. — The present specimens are similar to the type material, although larger in size.
Smith (1885: 312) suggested that the syntypes of P. scitulum could be juveniles, which would seem
to be the case. Sculpture and coloration are rather variable. P. cmadoritinctum has slightly stronger
sculpture, but is otherwise similar to the present specimens. Parvamussium sinense Wang. 1980 (: 259),
from the East China Sea, is somewhat broader, with stronger, scaly radial costae.

Oyama (1951: 80) used the genus Ctenamusium for this species, but the shell characters are
typical of Parvamussium. as indicated by Hertlein (1969).

Parvamussium squalidulum sp. nov.

Figs 47-50

Type material. — Holotype mnhn. Paratypes: 2 ams C201714, 3 hd, 6 mnhn, 2 usnm.

Type locality. - Lord Howe Rise, Kelso Bank, musorstom 5, stn DW 277, 24°1TS,
159°35' E, 270 m.

Material examined. — Chesterfield Islands, musorstom 5: stn DW 255, 25°15' S, 159°55' E,
280-295 m. 1 lv, 2 rv. — Stn DW 258, 25° 33' S. 159°46' E, 300 m, 7 lv, 3 rv. — Stn DW 260, 25°29' S,
1 59"44' E, 285 m, 1 lv. — Stn DW 261, 25°27' S, 159°46' E, 300 m, 2 rv. — Stn CP 268, 24°45' S,
159°39' E, 280 m, 4 lv. — Stn CP 269. 24°47' S, 159°37' E, 250-270 m. I lv. — Stn DW 272, 24°41' S,
1 59°43' E, 500-540 m, 1 lv, 3 rv. — Stn DW 274, 24°45' S, 159°41' E, 285 m, 1 lv. — Stn DW 277,
24°1 1' S. 159°35' E, 270 m, 4 spms, 2 lv, 8 rv (holotype and paratypes: 2 ams, 3 hd, 6 mnhn, 2 usnm).

Stn CP 279, 24'W S, 159°38' E, 260-270 m, 6 lv, 1 rv. — Stn DW 299, 24°48' S. 159° 24' E,

360-390 m, 2 lv, 3 rv. — Stn DW 301, 22°07' S, 159°25' E, 487-610 m. 1 lv.

Loyalty Islands, musorstom 6: stn DW 480, 21°08'S. 167°56' E, 380 m, 1 lv, 1 rv.

New Hebrides Arc1, volsmar: stn DW 7, 22°26' S, 171°44'E, 325-400 m, 1 spm, 1 lv. - Stn

DW 17, 22°23' S. 171°41' E, 260-300 m, 1 lv.

Distribution. — Chesterfield Islands, New Caledonia to the New Hebrides Arc; shells in
260-610 m, alive in 260-400 m.

Description. — Shell rather small, suborbicular, inequi¬
valve, inequilateral, semi-transparent, up to ca. 1 1 mm high,
left valve slightly more convex than right; auricles rather
small, unequal; umbonal angle about 125°.

Prodissoconch ca. 210 pm in height.

Left valve with numerous irregularly spaced, scaly radial
costae, a few commencing weakly at about I mm shell height,
increasing in number towards the ventral margin; small
delicate concentric lamellae developing between them, more
closely spaced near ventral margin. Anterior auricle well

developed, posterior rather small. Three very weak radial
lirae on anterior auricle, absent on posterior. Both auricles
covered with concentric lamellae. Hinge line straight.

Right valve with minute concentric lirae near the lateral
margin, absent from central region of disc. Marginal apron
(outer prismatic calcite layer) mostly broken off. covered with
fine concentric lirae (marginal region below the internal
lirae). Anterior auricle well developed, provided with 5 radial
riblets that are covered with con-centric lamellae, stronger on
dorsal margin. Posterior auricle with delicate lamellae.

Although the condominium of the New Hebrides has become independant under the name of Vanuatu, the term New
Hebrides is retained here to designate the Hunter and Matthew area of the New Hebrides Arc. Hunter and Matthew are a
dependency of New Caledonia. Similarly, geographers and geophysicists have retained the expression New Hebrides Trench.

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

33

47-54. 47-50, Parvamussium squalidulum , holotype. 1 1.1 x 11.4 mm (db). 47, lefl valve, exterior. 48. left valve,

interior. 49, right valve, exterior. 50. right valve, interior. - 51-54. P. lorresi , musorstom 5: stn DW 329, 9.0 x
8.9 mm (Iv), 7.9 x 7.7 mm (rv). - 51. left valve, exterior. 52. left valve, interior. 53. right valve, exterior. -
54, right valve, interior.

34

HENK DIJKSTRA

Ten irregularly spaced internal lirae. plus 2 rudimentary
lirae. One auricular lira on posterior auricle, absent on
anterior. External sculpture and coloration slightly visible
internally. Resilifer triangular. Byssal notch present, no

ctenolium. Left valve creamy, with white and orange-brown
maculations, right valve semi-transparent white and glossy.
Dimensions of the holotype: H 1 1.1, L 1 1.4, D 2 mm.

Remarks. — The closest species is Parvamussium sinense, the left valve of which is more
strongly sculptured with closely spaced scaly radial costae than in P. squalidulum. The latter is slightly
more orbicular, and the right valve is covered with delicate concentric lirae, which are absent in P.
sinense. The internal lirae of P. sinense are more regularly spaced with more stronger developed
terminal nodules at the distal ends. Another related species is P. scitulum, the left valve of which is
more delicately sculptured with close-set radial striae (sometimes absent), while concentric lirae are
also absent on the right valve. P. squalidulum is slightly more convex than P. scitulum.

Etymology. — From numerous scaly radial costae of the exterior of the left valve (Latin
squalidulus, adj. = with small scales).

Parvamussium texturatum (Dautzenberg & Bavay, 1912)
Figs 103-106

Amussium texturalum Dautzenberg & Bavay, 1912: 37. pi. 27. figs 19-22.

Other reference:

Parvamussium cf. texturalum — Dijkstra. 1991: 17. figs 44-52.

Type material. — Holotype. live taken, zma 3.12.022.

Type locality. — Sulu Archipelago, “Siboga", stn 105, 6°08' N, 1 2 1 ° 1 9' E, 275 m.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DW 300, 22°48' S, 159°24' E, 450 m, 2 spms.

New Caledonia, biogeocal: stn DW 308, 20°40' S, 166°58' E, 510-590 m, 4 lv.

Loyalty Islands, musorstom 6: stn DW 417, 20°42' S. 167°04' E, 283 m, 1 lv. — Stn DW 456, 21°01' S,
167°26' E, 240 m, 1 lv, 2 rv. — Stn DW 461, 21°06' S, 167°26' E, 240 m, 1 spm.

Distribution. Beside the type locality in the Sulu archipelago (Philippines), somewhat
similar specimens were recorded by Dijkstra (1991) from Indonesia. Now it is also known from the
Chesterfield Islands, New Caledonia and the Loyalty Islands. Present material live collected in
240-450 m.

Description. Shell small, suborbicular, inequivalve,
inequilateral, nearly opaque, up to ca. 5 mm high, left valve
slightly more convex than right, auricles unequal in size,
umbonal angle about 95°. Left valve creamy white, someti¬
mes stained with few white and brown spots, right valve
whitish.

Prodissoconch ca. 200 pm in height.

Left valve sculptured with strongly developed, irregularly-
spaced radial costae, starting at 1-2 mm shell height and
enlarging towards ventral margin. Regularly spaced concen¬
tric lamellae cross radial costae, which are ornamented with
strongly developed spines or scales. Auricles sculptured with

concentric lamellae, more strongly developed on anterior.
Hinge line straight.

Right valve covered with regularly spaced, close-set concen¬
tric lirae that enlarge near ventral margin to form lamellae.
Anterior auricles with 4-7 weak radial riblets covered with
concentric lamellae. Posterior auricle with delicate concentric
lamellae, very strongly developed on dorsal margin.

Internal lirae 10-15, sometimes one or more rudimentary
interstitial lirae, plus 1 or 2 auricular lirae. Lirae commencing
in late ontogeny, enlarging into submarginal region. Resilifer
triangular. Byssal notch small, no ctenolium.

Remarks. — The present specimens are similar to the type material from the Sulu Sea,
although most have fewer internal lirae. The internal lirae, however, are variable in number, and it
is possible that the number in the type specimen (16) is exceptional.

The closest species is Parvamussium vesiculatum sp. nov., see below.

Source : MNHN. Paris

BATHVAL. PECTINOIDEA FROM NEW CALEDONIA

35

Parvamussium thetidis (Hedley, 1902)
Figs 99-102

Amusium thetidis Hedley, 1902: 304, fig. 49.

Synonym:

Ctenamusium salacon I redale, 1929: 164.

Other references:

Amusium thetidis - Hedley & Petterd, 1906: 223, pi. 38, figs 18-19.

Amussium thetidis - Pritchard & Gatliff. 1904: 266.

Ctenamusium thetidis - Iredale. 1929: 164, 188. Cotton & Godfrey, 1938: 98, fig. 84. Cotton, 1961: 102, fig. 88.
Parvamussium thetidis - Dijkstra, 1991: 14. Lamprell & Whitehead, 1992: no. 35. pi. 6, fig. 35.

Type material. A. thetidis-. holotype, rv, ams Cl 32 10, 5 paratypes, v, ams Cl 71 634. — C.
salacon. holotype, lv, ams C24345.

Type locality. — A. thetidis : 5-8 miles off Port Kembla, New South Wales, Australia,
“ Thetis ”, stn 49, 34°28' S, 151°06-03' E, 137-1 15 m. C. salacon. 27 miles E of Sydney Heads, New
South Wales, Australia, 549 m.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DW 272, 24°40' S, 159°43' E, 500-540 m, 1 spm. Stn DW
305, 22°09' S, 1 59°24' E, 430-440 m, 1 lv, 1 rv.

Norfolk Ridge, chalcal 2: stn DW 73, 24"40' S, 168°38' E, 573 m, 2 spms, 5 lv, 4 rv. Stn DW 74,
24°40' S, 168°38' E, 650 m, 2 spms, 4 lv, 5 rv. Stn DW 75, 24°39' S. 168°39' E, 600 m, 2 spms,
2 rv.

Loyalty Islands, musorstom 6: stn DW 479. 21°09' S, 167°55' E, 310 m, 1 lv. - Stn DW 485, 21°23' S,
167°59' E, 350 m, 1 lv.

New Hebrides Arc. volsmar: stn DW 30, 22°17' S, 1 7 1°1 8' E, 450-550 m, 5 lv, 10 rv.

Gemini: stn DW 51, 20°59' S, 170°03' E, 450 m, 4 lv, 6 rv.

Distribution. — Eastern Australia, Chesterfield Islands as far east as New Hebrides Arc.
Present material living at 500-650 m.

Description. Shell rather small, fragile, inequivalve,
inequilateral, up to ca. 10 mm high, left valve slightly more
convex than right, auricles unequal in size, umbonal angle
about 95°. Left valve creamy, sometimes with white spots,
opaque; right valve uniform whitish, semi-transparent.

Prodissoconch ca. 210 pm in height.

Left valve sculptured with irregularly spaced radial costae,
commencing at ca. 2 mm shell height, enlarging to ventral
margin. Concentric lamellae delicate, close-set transversing
radial costae. Auricles with delicate, closely-spaced concentric
lamellae. Hinge line straight.

Right valve with regularly spaced concentric lirae. Anterior
auricle with closely-spaced, concentric lamellae and 1-3
weakly developed radial riblets.

Internal lirae generally 10. with some rudimentary inters¬
titial lirae, plus I or sometimes 2 auricular lirae. Lirae fine at
first, becoming stronger distally near submarginal region.
Hinge line slightly raised at margins, with small scales on
dorsal margin. Resilifer triangular. Byssal notch small, no
ctenolium.

Remarks. — Closely related species are Parvamussium cristatellum (Dautzenberg & Bavay,
1912) from Indonesia and the Bay of Bengal, and P. siebenrocki (Sturany, 1901) from the
northwestern Indian Ocean. It is possible that both are synonyms, as 1 have already suggested
(Dijkstra, 1991: 14).

P. thetidis is the type species of Ctenamussium which Hertlein (1969: N350) synonymized with
Parvamussium. Indeed, it is a typical Parvamussium.

Source :

36

HENK DIJKSTRA

Parvamussium torresi (E.A. Smith, 1885)
Figs 51-54, 125-128

Amussium torresi E.A. Smith, 1885: 311, pi. 23, figs 3-3b.

Other references:

Glyptamusium torresi — Iredale, 1939: 370.

Propeamussium (Parvamussium) torresi — Dijkstra, 1990: 3.

Type material. Lectotype (Figs 125-8, H 7.8, L 7.9, D 2 mm) here designated, live taken,
bmnh 1887.2.9.3316, 5 paralectotypes bmnh 1887.2.9.3317/1-5. Size of the lectotype is somewhat
different from Smith's measurements, due to some damages near ventral margin of left valve, and
marginal apron of right valve broken off.

Type locality. — E of Cape York, Queensland, Australia, ‘■Challenger”, stn 185B. 1 1°38'15"
S, 143°59'38" E, 285 m.

Material examined. — The type material.

Chesterfield Islands, chalcal 1: stn D 13, 19°33'S, 158° 38' E, 390 m. 3 lv.

musorstom 5: stn DC 291, 23°08' S, 159°28' E, 300 m. 1 rv. — Stn DW 328, 20°23' S, 158°44' E,

340-355 m, 1 spm, 8 lv, 25 rv. — Stn DW 329, 20°23' S, 158°47' E, 320 m, 23 lv, 34 rv. — Stn DW

334, 20°06' S, 158°48' E, 315-320 m. 1 spm, 7 lv, 3 rv. — Stn DW 335, 20°03' S, 158°45' E, 315 m,

1 spm, 3 lv. — Stn DC 376, 19°51' S, 158°30' E, 280 m, 7 lv, 1 rv.

corail 2: stn DC 169, 18°21'S, 155°20'E, 575 m, 1 lv, 2 rv.

New Caledonia, musorstom 4: stn CC 174, 19°00' S. 163°18' E, 365 m. 2 lv.

Norfolk Ridge, chalcal 2: stn DW 71, 24°42' S, 168°09' E, 230 m, 1 lv.

Loyalty Islands, musorstom 6: stn DW 459, 21°0T S, 1 67°3 1 ' E, 425 m, 22 lv. 58 rv. — Stn DW 468
21°06'S, I67°33'E, 600 m, 1 lv, 1 rv.

New Hebrides Arc. volsmar: stn DW 17, 22°23' S, 1 7 1°4 1' E, 260-300 m, 1 spm, 1 lv, I rv.

Distribution. — Torres Strait and Sulu archipelago (Smith, 1885; Dijkstra. 1990). Now also
recorded from the Chesterfield Islands, New Caledonia and Loyalty Islands. Present specimens living
at 260-355 m. 6

Description. Shell rather small, fragile, suborbicular,
inequivalve, semi-transparent white, up to ca. 9 mm high, left
valve somewhat more convex than right, auricles unequal in
size, umbonal angle ca. 100".

Prodissoconch ca. 200 pm in height.

Left valve glossy, very weakly sculptured with irregularly
spaced concentric growth lines. Sometimes with minute,
close-set concentric lamellae near ventral margin. Auricles
smooth: anterior auricle considerable larger than posterior.
Hinge line straight.

Right valve covered with regularly spaced concentric lirae.
close-set near the umbonal area, becoming more widely

spaced towards ventral margin. Anterior auricle sculptured
with 1-5 weakly de-veloped radial riblets, absent on posterior.
Prominent concentric lamellae near anterior margin, more
weakly developed near posterior margin. Hinge line somew¬
hat elevated due to strong lamellation on antero-and postero-
dorsal margins.

Internal Tirae generally 10. plus 1 small auricular lira on
each side. Sometimes with the addition of 1 or 2 rudimentary
interstitial lirae. Left valve transparent white, sometimes with
white spots, right valve semi-transparent white. Resilifer
triangular. Outer ligament well developed. Small byssal
notch, no ctenolium.

Remarks. — The present specimens are very similar to the type material. Small specimens of
P. torresi and P. scitulum are easily confused, because of their variable sculpture. The left valve of
P. torresi is rather convex, that of P. scitulum strongly compressed to nearly flat, and more brightly
coloured. The right valve of P. torresi is covered with regularly concentric lirae, which are generally
lacking on P. scitulum.

A closely related species is Parvamussium formosum (Melvill & Standen, 1907) from the Gulf
of Oman, which is sculptured with weak radial striae near the posterior and anterior margins of the
left valve, while the auricles are more strongly sculptured than in P. torresi.

Source

BATHYAL PECTINOIDF.A FROM NEW CALEDONIA

37

P. torresi is the type species of Glyptamusium, which was synonymized with Parvamussium by
Hertlein (1969: N350). Oyama (1944: 242, 244) misspelled the name “ Graptamussium

Parvamussium undisonum sp. nov.

Figs 55-58

Type material. Holotype mnhn. Paratypes: 5 mnhn, 1 usnm, 2 hd.

Type locality. Loyalty Islands, musorstom 6, stn DW 489, 20°48' S, 167°05' E, 700 m.

Material examined. New Caledonia, biocal: stn KG 06, 20°34' S, 166°52' E, 735 m, 2 rv
(paratypes: mnhn, usnm).

Loyalty Islands, musorstom 6: stn DW 488, 20°49' S, 167°06' E, 800 m, 4 lv (paratypes: 3 mnhn,
1 hd). Stn DW 489, 20°48' S, 167°05' E, 700 m, 1 spin (holotype).

New Hebrides Arc. Gemini: stn DW 55, 20°59' S. 170°01' E, 710 m, 1 spm, I rv (paratypes: mnhn, hd).

Distribution. — New Caledonia and New Hebrides Arc, 700-800 m, living 700-710 m.

Description. Shell rather small, inequivalve, inequila¬
teral. up to ca. 14 mm high, somewhat higher than wide, left
valve slightly more convex than right, translucent white:
auricles nearly equal in size, relatively small, umbonal angle
ca. 90".

Prodissoconch ca. 210 pm in height.

Left valve covered with undulating concentric lamellae that
cross irregularly spaced radial costae, enlarging towards
ventral margin. Auricles with close-set concentric lamellae,
which is somewhat more prominent on the anterior auricle.
Hinge line straight.

Right valve ornamented with regularly spaced concentric
lirae. very weak near submarginal and marginal areas, which
are covered by a layer of delicate undulating concentric
lamellae (often worn off in dead specimens). Anterior auricle

well developed, sculptured with 5 radial riblets, covered with
concentric iamellae, which is strongly developed on anlero-
dorsal margin. Posterior auricle covered with concentric
lamellae, which are more prominent on postero-dorsal mar¬
gin. Marginal apron broken off in all specimens seen.

Internal lirae 10. 4 of them rudimentary, plus 1 small
posterior auricular lira on the left valve; 9 lirae plus 2
rudiriientary. and I posterior auricular lira on right valve.
Lirae commencing near adductor insertion scar and enlarging
towards to submarginal area. Exterior sculpture of left valve
slightly visible from the interior. Outer ligament small.
Resilifer triangular. Byssal notch small. Fasciolar pseudo-
ctenolium present, with one active tooth.

Dimensions of the holotype: FI 14.2, L 13.9, D 3.1 mm.

Remarks. — The most closely related species is P. sayanum (Dali. 1886) from the Gulf of
Mexico and West Indies, which has almost identical sculpture. It differs, however, in having more
prominent, radial and more widely spaced concentric lamellae, which are more delicate on the
auricles.

Waller (1984: 213) discussed the pseudo-ctenolium in P. sayanum , which is an unusual
feature in Propeamussiidae. P. undisonum is another propeamussiid with an active tooth, only visible
in adult specimens. One immature right valve (9 mm high) from biocal stn KG 06 lacks this tooth.

No other propeamussiid species known from the Indo-Pacific region approaches P. undisonum.

Etymology. — From the undulating sculpture (Latin undisonus, adi. = surrounded by wave).

Parvamussium vesiculatum sp. nov.

Figs 59-62, 93-96

Type material. Holotype mnhn. Paratypes: 2 ams C201715, 12 hd, 43 mnhn, 2 nmnz
M268538, 2 nsmt, 2 usnm.

Type locality. South-east New Caledonia, biocal. stn DW 44, 22°47' S, 167°14'E,
440-450 m.

38

HENK DIJKSTRA

Figs 55-62. — 55-58. Parvamussium undisonum, holotype. 14.2 x 13.9 mm (db). 55. left valve, exterior. - 56, left valve,

interior. - 57. right valve, exterior. — 58. right valve, interior. — 59-62. P. vesiculatum , holotype, 6.0 x 5.8 mm
(db). 59, left valve, exterior. — 60. left valve, interior. 61. right valve, exterior. — 62, right valve, interior.

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

39

Material examined. — New Caledonia. " Vauban " 1978-79: stn 15, 22°49' S. 167°12' E,
390-395 m. I lv.

biocal: stn DW 08, 20°34' S, 166°54' E, 435 m, 1 rv. — Stn DW 44, 22°47' S, 167°14' E, 440-450 m,
37 spms, 13 lv, 14 rv (holotype and paratypes). — Stn CP 45, 22°47' S, 167°15' E, 430-465 m, I lv.
- Stn DW 46. 22°53' S, 167° 17' E, 570-610 m, 1 rv. — Stn DW 49, 23°03' S, 167°32' E, 825-830 m,
1 rv. _ stn DW 51, 23°05' S, 167°45' E, 680-700 m, 2 lv. — Stn DW 106, 21°36' S, 166°29' E, 625-
650 m, 2 spms.

musorstom 4: stn DW 184, 19°04' S, 163°27' E, 260 m, 1 spm. — Stn DW 210, 22°44' S, 167°09' E,
340-345 m, 1 spm, 2 lv, 1 rv. — Stn DW 222, 22°58' S, 167°33' E, 410-440 m. 2 lv. — Stn DW 234.
22°15'S, 167°08'E, 350-365 m, 1 spm.

Norfolk Ridge, chalcal 2: stn DW 76, 23°41' S, 167°45' E, 470 m, 2 spms.
smib 3: stn DW 22, 23°03' S, 167°19' E, 503 m, 7 lv.

Loyalty Islands, musorstom 6: stn DW 479, 21°09' S. 167°55' E. 310 m, 1 lv.

Distribution. — New Caledonia and Loyalty Islands, shells in 260-830 m, living in 260-

650 m.

Dkscription. — Shell small, suborbicular. inequivalve.
inequilateral, semi-transparent, up to 6 mm high, left valve
slightly more convex than right, auricles unequal in size,
umbonal angle ca. 95°.

Prodissoconch ca. 200 pm in height.

Left valve widely spaced concentric lirae with strongly
developed nodules or hollow scales, broken off in all
specimens seen below umbonal region, commencing almost
immediately, close-set at first, becoming stronger and more
widely spaced towards ventral margin. Irregular radial riblets
between concentric lirae. Auricles covered with concentric
lamellae, stronger on anterior auricle, slightly closer on
posterior auricle. Hinge line straight.

Right valve with regularly spaced concentric lirae. very
weakly developed near umbonal region. Prodissoconch and

dissoconch pellucid. Anterior auricle with 4 delicate radial
riblets, which are covered by concentric lamellae that deve-
lope into scales on the dorsal margin. Posterior auricle with
concentric lamellae, strongly developed on dorsal margin.
Byssal fasciole well developed.

Internal lirae rudimentary. 5 near anterior and 4 near
posterior margin. I delicate auricular lira on anterior and a
stronger one on posterior of left valve; right valve with 3
rudimentary lirae on anterior, posterior margin with I
auricular lira on posterior auricle. External sculpture visible
from interior of both valves. Resilifer triangular. Byssal notch
small, no ctenolium.

Both valves semi-transparent white; right valve with a large
white spot at centre of disc.

Dimensions of the holotype: H 6, L 5.8, D 1.8 mm.

Remarks — A few paratypes lack external sculpture on the left valve (Figs 95-96) after the
shell attains 2 mm high, and are very fragile, smooth, glossy and transparent.

Parvamussium vesiculatum most closely resembles P. texturatum which is sympatric and is
slightly more orbicular with somewhat larger auricles. The radial costae on the left valve are more
prominent in the latter species, whereas concentric lirae are more strongly developed and more widely-
spaced in P. vesiculatum. P. vesiculatum is ornamented with nodules on the concentric lirae, whereas
P. texturatum bears more scales. There are also differences in the internal lirae, which are rudimentary
and fewer in number in P. vesiculatum. and well developed in P . texturatum.

Etymology. — From the nodular sculpture of the left valve (Latin vesiculatus. adj. — with
nodules).

Genus Cyclochlamys Finlay, 1926

Cvclochlamys Finlay, 1926: 452. Type species (OD): Pecten transenna (sic) Suter, 1913. Recent, near the Snares. New Zealand.
91 m.

Synonym:

Chlamydella Iredale, 1929: 164. 188. Type species (OD): Cyclopecten favus Hedley, 1902. Recent, off New South Wales,
Australia, 75-91 m.

Diagnosis. Shell inequivalve. minute, orbicular to strongly oblique; left valve sculptured
with radial and/or concentric riblets or striae, right valve with commarginally elongate hexagonal

40

HENK DIJKSTRA

(honeycomb-like) microstructure (simple calcitic prismatic outer layer); auricles unequal; byssal notch
well-developed; no ctenolium; no internal lirae.

Distribution. — Recent. Subantarctic, South Australia and West Pacific from New Zealand
to Japan (Hayami & Kase, 1993).

Remarks. — Cyclochlamys was erroneously mentioned as a “nom. null." by Hertlein (1969:
N353). It differs from Cyclopecten by a pteriform shell, a triangular prodissoconch and the
commarginally elongate hexagonal (honey-comb like) microstructure of the right valve.

Cyclochlamys favus (Hedley, 1902) comb. nov.

Figs 87-90

Cyclopecten favus Hedley, 1902: 305, fig. 50 (as C.Jlavus, sic).

Synonyms:

Cyclopecten obliquus Hedley. 1902: 306. fig. 51.

Cyclopecten nepeanensis Pritchard & Gatliff, 1904: 338. pi. 20. fig. 5.

Other references:

Chlamydella favus - Cotton & Godfrey, 1938: 98, fig. 85 (after Hedley). — Cotton. 1961: 103, fig. 89.

Type material. — C. favus: holotype (H 3.0, L 3.4, D 0.5 mm) live taken, ams C 1 323 1 .
20+ paratypes v, ams Cl 7 1633. — C. obliquus : holotype lv, ams Cl 3234, 70+ paratypes v, ams
C13232, Cl 3233. — C. nepeanensis: holotype lv, nmv F558.

Type locality. — C. favus: 5. 5-7. 5 miles NE of Cape Three Points, New South Wales,
Australia, "Tltelis", stn 13, 33°28' S, 1 5 1 °33' E, 75-91 m. — C. obliquus: 5-8 miles off Port Kembla,
New South Wales, " Thetis ", stn 49, 34°28' S, 151°06-03'E, 115-137 m. — C. nepeanensis: Back
Beach. Point Nepean. Victoria, Australia.

Material examined. — The type material.

Australia. Off Moreton Bay, Queensland. 27°3T S, 153°40' E, 75-80 m.l rv (ams). — E of Sydney,
New South Wales, "Gascoyne", stn G2-56-62, 150 m, several valves (ams). — NE of Woolongong,
New South Wales, “ Kapala ”, stn K75-05-06, 34°18-24' S, 151°26-21'E. 466-494 m, 13 v (ams). —
2.5 miles NE Beaching Bay, Maria Island, SE Tasmania, " Penghana ”, 42°27.5' S, 148° 12' E, 82.5 m,
several spms (ams). Between Eucla and Esperance. Western Australia, " Gascoyne ”, stn G2-96-62,
79-147 m, several v (ams).

New Caledonia, biocal: stn DW 44, 22°47' S, 167°14' E, 440-450 m, 1 spm (typical). — Stn DW 46,
22°53' S, 167° 17' E, 570-610 m, 6 spms (typical), 2 spms (var. obliqua).

— Southern Australia and New Caledonia. Present material living at 440-

Distribution.

610 m.

Description. — Shell small, up to ca. 3 mm high,
translucent. Valves dissimilar in shape, size and sculpture.
Prodissoconch ca. 240 pm in height (Figs 87-88).

Left valve somewhat larger, more convex, transparent,
smooth except for a few minute concentric growth lines
(typical) or with a few more prominent concentric lirae
(var. obliqua ); auricles nearly equal, with some concentric

lines. Outer ligament insertion with small holes near resilifer
(Fig. 89).

Right valve somewhat smaller, transparent, smooth, with
hexagonal microsculpture (Fig. 90). Anterior auricle with
close-set concentric lamellae. Byssal notch well developed; no
ctenolium.

Remarks. — The present material is similar in all respects to the type specimen from off New
South Wales. Cyclochlamys favus is rather variable in sculpture on the left valves with weak or

Source

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

41

pronounced concentric growth lines, and sometimes secondary radial striae. Completely smooth
specimens are also known.

Genus Cyclopectex Verrill, 1897

Cyclopecten Verrill. 1897: 70. Type species (SD, Sykes. Smith & Crick, 1898): Pecten pustulosus Verrill. 1873: 14: Recent,
off Newfoundland. Canada. 274 m.

Synonym:

Xenamussium Oyama, 1944: 244. [Proposed as a subgenus of Propeamussium). Type species (OD): Pecten lioskynsi Forbes,
1844; Recent, "Asia minor" [= off Turkey].

Diagnosis. — Shell equivalve, small, usually orbicular, laterally compressed, prodissoconch
flat curved; left valve usually sculptured with radial and/or concentric costae or striae, right valve with
concentric lamellae; auricles unequal; byssal notch well-developed; no ctenolium; no internal lirae.

Distribution. — Miocene to Recent. Worlwide, subtidal to hadal (Hayami & Kase, 1993:
59). In the Indo-Pacific Cyclopecten , however, only lives in deep-water (Dijkstra, unpubl. data).

Remarks. In the original diagnosis of Cyclopecten, the characters “few pectinidial teeth
[= ctenolium], byssus small and a few threads”, refer to “ Cyclopecten ” orbicularis (Sowerby)
[= Lissochlamis exotica (Dillwyn, 1817)], a pectinid species. These characters are not present in
Cyclopecten, as defined by the type species. There are many described species of Cyclopecten but they
are currently placed in different genera, for example Delectopecten (a pectinid) (Hertlein, 1969;
Knudsen, 1970; Habe, 1977; Bernard. 1978; Rombouts, 1991) or Palliolum Monterosato, 1884 (also
a pectinid) (Abbott, 1974, pro parte). Also, several “ Cyclopecten ” species described from the
subantarctic region, belong to Cyclochlamys (Dijkstra, unpubl. data). Cyclopecten is morphologi¬
cally closest to Parvamussium. As Hayami & Kase (1993: 59) suggested, Cyclopecten may have been
derived from Parvamussium through paedomorphic evolution. Shell characters of Xenamussium are
sufficiently similar to Cyclopecten to suggest synonymy.

Cyclopecten horridus sp. nov.
Figs 63-64, 98

Type material. — Holotype mnhn. Paratypes: 5 mnhn, 2 hd.

Type locality. — Loyalty Islands, musorstom 6, stn DW 420, 20°29' S, 166°43' E, 600 m.

Material examined. — New Caledonia, biocal: stn KG 06, 20°34' S, 166°52' E, 735 m, 1 lv
(paratype). — Stn DW 08, 20°34' S, 166° 53' E, 435 m, 3 lv, 2 rv (paratypes: 3 mnhn, 2 hd).
biogeocal: stn DW 307, 20°35' S, 166°55' E. 470-480 m, 1 lv (paratype).

Loyalty Islands, musorstom 6: stn DW 420, 20°29' S, 166°43' E, 600 m. 1 spm (holotype).

Distribution. — New Caledonia and Loyalty Islands, shells in 435-735 m. living in 600 m.

Description. Shell small, suborbicular, inequivalvc. up
to ca. 6 mm high, left valve more convex than right, marginal
apron of right valve pressed against margin of interior of left
valve, auricles of nearly equal size, umbonal angle ca. 125°,
whitish opaque.

Prodissoconch ca. 220 pm in height.

Left valve covered with regular concentric lamellae, closely-

set in early ontogeny, enlarging towards periphery. Erect
hollow spines developing on lamellae (Fig. 98), which are
stronger near the ventral margin (broken off near umbonal
area in all specimens seen). Crowded microscopic granules
between concentric lamellae, radial scratches added in late
ontogeny. Auricles covered with sculpture identical to that on
exterior of disc.

42

HENK DIJKSTRA

Figs 63-70. — 63-64. Cyclopecten horridus, holotype. 5.9 x 6.0 mm (db). — 63. left valve, exterior. 64. right valve, exte¬
rior. — 65-68. C. pellucidulus, holotype, 6.1 x 6.8 mm (db). 65. left valve, exterior. — 66. left valve, interior.

67, right valve, exterior. 68. right valve, interior. — 69-70. Delectopecten musorstomi, biocal: stn DW 64. 4.3 x
4.3 mm (db). — 69. left valve, exterior. 70. right valve, exterior.

Source : MNHN, Paris

BATHYAL PF.CTINOIDEA FROM NEW CALEDONIA

43

Right valve with regular concentric lirae, close-set to 2 mm
shell height, then transformed into more widely spaced,
undulating concentric lamellae with delicate spines near
ventral margin. Granulate microsculpture between the
concentric lirae and lamellae. Anterior auricle sculptured
with 3 radial riblets covered with concentric lamellae. Poste¬

rior auricle poorly differentiated from disc, covered with
concentric lirae. Hinge line straight. Antero-dorsal margin
with scales, squamae weakly developed on postero-dorsal
margin. Byssal notch strongly developed. No ctenolium.

Dimensions of the holotype: H 5.9. L 6, D 1.6 mm.

Remarks. — A somewhat similar species is Parvamussium araneum Dijkstra, 1991 from
the Indonesian Archipelago. This species has a few internal rudimentary lirae, and micro-sculpture
on the right valve has radial scratches. Another related species is P. vidalense (Barnard, 1964) from
off Zululand, South Africa. The radial sculpture of P. vidalense is somewhat weaker than in P.
araneum , with overriding concentric lamellae, while the erect spines are weaker. Moreover, a few
more internal rudimentary lirae are present in P. vidalense than in P. araneum , but they are absent
in C. horridus.

Some shell features of C. horridus resemble those characteristic of Delectopecten, such as
arrangement of erect spines on concentric lamellae, and strongly developed byssal notch. A ctenolium
however, is absent, and the sculpture of the right valve is more similar to that in Cyclopecten.

Etymology. — From the spiny sculpture (Latin horridus , adj. = bristly).

Cyclopecten pellucidulus sp. nov.

Figs 65-68

Type material. — Holotype mnhn. Paratypes: 1 hd, 2 mnhn.

Type locality. — South of New Caledonia, biocal, stn DW 51, 23°05' S, 167°44' E, 680-

700 m.

Material examined. — Only known from the type material.

Distribution. — Only known from the type locality, northern Norfolk Ridge, alive in 680-

700 m.

Description. — Shell small, fragile, pellucid, suborbicular.
inequivalve, up lo ca. 6 mm high, left valve slightly more
convex than right, auricles equal in size, umbonal angle ca.
125°.

Prodissoconch ca. 230 pm in height.

Left valve covered with curved concentric lamellae,
commencing at 2 mm shell height, enlarging towards ventral
margin. Umbonal area smooth and glossy. Auricles with
close-set concentric lamellae.

Right valve covered with delicate regularly spaced concen¬
tric lirae that gradually enlarge, interspaces with microscopic
radial scratches. Anterior auricle with concentric lamellae,
stronger than on posterior auricle, strongly developed scales
on antero-dorsal margin. Exterior sculpture visible from
interior of both valves. Interior with one weak liration on the
inner side of each auricle. Resilifer triangularly elongate.
Byssal notch present, no ctenolium. Outer ligament small.

Dimensions of the holotype: H 6.1, H 6.8, D 1.2 mm.

Remarks. — A somewhat similar species is Cyclopecten aupouria Powell, 1937 from northern
New Zealand, which attains larger size, is more convex, and is opaque white with more delicate
concentric lirae and finer radial lirae. Another related species is C. bavayi Dijkstra, 1990 from the
Ceram Sea (Indonesia), which is also slightly larger, opaque, and covered with numerous concentric
lamellae on the early disc, and small radial riblets, which are absent in C. pellucidulus.

Etymology. — From the translucence of the shell (Latin pellucidulus , adj. = translucent).

44

HENK DIJKSTRA

Family Entoliidae Teppner. 1922

Entoliidae Teppner. 1922: 89. 274.

Synonym:

Syncyclonemidae Waller. 1978: 353.

Diagnosis. Byssate or free Pectinoidea. lacking prismatic calcite; foliated calcite thin or
absent; crossed-lamellar aragonite dominant extending to margins; byssal notch without ctenolium.

Remarks. Waller (1978) described a new family Syncyclonemidae in Pectinoidea. and
treated Pectinella as a junior synonym of Syncyelonema Meek. 1864. Later, he reduced Syncyclone¬
midae in rank within the family Entoliidae, and reinstated Pectinella as a Recent genus (Waller.
1984: 219).

Genus Pectinella Verrill, 1897

Pectinella Verrill, 1897: 68. Type species (OD): Pecten ( Pseudamusium) sigsbeei Dali. 1886; Recenl. Cuba. 289 m.

Diagnosis. — Shell equivalve, small, transparent, nearly smooth, with convex and nearly
equal valves; auricles very unequal, oblique; byssal notch well-developed; no ctenolium.

Distribution. - Recent, Gulf of Mexico, Caribbean Sea, central and western Pacific; 240-

300 m.

Pectinella aequoris Dijkstra, 1991
Figs 71-74

Pectinella aequoris Dijkstra. 1991: 23. figs 78-86.

Type material. Holotype lv, rmnh 56567, 4 paratypes rv, rmnh 56568-56569.

Type locality. Indonesia, Bay of Sanagar, N of Sumbawa, “Snellius-II", stn 4.111
8°19.3'S, 118°15.6'E, 175-185 m.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DW 272, 24°41' S, 159°43' E, 500-540 m. I lv.

Loyalty Islands, biogeocal: stn DW 296, 20°38' S, 167°10' E, 1230-1270 m, 1 lv.

musorstom 6: stn DW 397, 20'’47' S, 167°05' E, 380 m, 1 lv. — Stn DW 474, 21°09' S, 167°55' E,

260 m, 1 spm.

volsmar: stn DW 38, 22°22' S, 168°44' E, 380-420 m, 1 lv.

Distribution. Indonesia, Hawaiian Islands and Fiji Islands (Dijkstra, 1991). Now also
found in the Chesterfield Islands and the Loyalty Islands. Present material living at 260 m.

Description. — Shell rather small, convex, elongate,
slightly equivalve, semi-transparent, up to ca. 20 mm high,
right valve somewhat more convex than left, auricles unequal
in size, umbonal angle ca. 90°.

Prodissoconch ca. 220 pm in height.

Left valve smooth, glossy, with microscopic radially diver¬
ging scratches. Delicate close-set concentric striae enlarging

towards ventral margin. A few concentric iirae near umbonal
region. Some specimens with a few delicate, irregularly
spaced radial grooves that commence near centre of disc and
enlarge ventral margin (visible from interior as very weak
lirae). Anterior auricle with concentric lirae, weaker on pos¬
terior. Delicate concentric striae near lateral margin. Hinge
line straight. Internal surface glossy, and microscopically

Source MNHN. Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

45

-78 71-74, Peclinella aequoris, musorstom 6: stn DW 474. 18.2 x 16.6 mm (db). - 71, left valve, exterior

72. left valve, interior. 73, right valve, exterior. 74. right valve, interior — 75-78. Pseudohmmies levu BIOGE°Cal.
itn CP 273, paratypes, 41.8 x 45.1 mm (lv), 28.9 x 37.2 mm (rv). - 75. left valve, exterior. — 76. left valve, mte-
-ior. 77. right valve, exterior. 78, right valve, interior.

Source

46

HENK DIJKSTRA

granulated. One lira! tooth (auricular crura) formed near
border of each auricle. Resilifer triangularly elongate. Car¬
dinal crura rather broad near resilial pit on antero-dorsal
region. A prominent tooth near pit on postero-dorsal region.

Right valve with similar microsculpture to that of left valve.
Auricles strongly produced on dorsal margin. Antero-dorsal

region of auricles with narrow straight groove, hinge line
strongly raised. Postero-dorsal side" of auricles a small
depression. Byssal notch well developed, no ctenolium.

Creamy transparent with orange-brown and white macula-
tions on left valve, right valve transparent and uniformly
cream.

Remarks. — The present specimens are similar to the type material, although larger in size.
For comparison with Pectinella sigsbeei and other species, see Dijkstra (1991: 25).

Family Pectinidae Wilkes, 1810
Pectinidae [as Pectinoidae] Wilkes, 1810: 32 [emend. Waller, 1978]

Diagnosis. — Byssate, cemented, or free living Pectinoidea with outer, prismatic calcited layer
on right valve; crossed-lamellar aragonite inside of pallial line or absent; byssal notch with ctenolium,
at least at early growth stages.

Remarks. — The family Pectinidae is usually attributed to Rafinesque (1815). Wilkes,
however, has priority. For synonymy see Hertlein (1969: N348).

Genus Pseudohinnites Dijkstra, 1989

Pseudohinnites Dijkslra. 1989: 29. Type species (OD): Pseudohinnites levii Dijkstra, 1989.

Diagnosis. — Shell inequivalve, medium-sized, fragile, very irregular, with a Hinnites-Uke
appearance; auricles unequal; left valve convex, right valve flat; exterior of left valve with irregular
radial costae and concentric lamellae overlying the radial costae and interstices; right valve smooth,
with irregular concentric waves or irregular depressions, giving a deformed appearance; byssal notch
with ctenolium.

Distribution. — Recent. Western to southwestern Pacific; 700-2000 m.

Remarks. — Pseudohinnites somewhat resembles Hyalopecten, although the former has
different shape (“Hinnites- like”), whereas the latter is regular undulated. Pseudohinnites is still under
study and will be discussed elsewhere (Knudsen & Dijkstra, in prep.).

Pseudohinnites levii Dijkstra, 1989
Figs 75-78

Pseudohinnites levii Dijkstra. 1989: 29, figs 1-3; 1990: 7. figs 14-15.

Type material. — Holotype and paratypes mnhn.

Type locality. — SE of New Caledonia, biocal, stn DW 70, 23°25' S, 167°53' E, 965 m.

Material examined. — New Caledonia, biocal: stn CP 26, 22°40' S, 166°27' E, 1618-1740 m
2 fragm. lv, 3 fragm. rv (paratypes). — Stn CP 30, 23°09' S. 166°41' E, 1 140 m, 4 fragm. lv, 1 fragm’
rv (paratypes). - Stn DW 70, 23°25' S, 167°53' E, 965 m, 1 lv (holotype). — Stn DW 80. 20°32' S,
166°48' E, 900-980 m, 1 rv (paratype).

Source MNHN. Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

47

biogeocal: stn CP 232, 21°34' S, 166°27' E, 760-790 m, 1 lv. - Stn CP 243, 21°27' S, 166°26' E, 1820
m, 3 lv. - Stn CP 260, 21°00' S, 166°58' E, 1820-1980 m, 3 lv, 2 rv (paratypes). — Stn CP 265,
21°04' S, 167°00' E, 1760-1870 m, 1 fragm. lv. — Stn CP 273, 21°02' S, 166"57' E, 1920-2040 m, 4 lv,
3 rv (paratypes).

calsub: dive 13, 21°26' S, 166°23' E, 1567-1807 m, 1 spm Guv.).

Loyalty Islands, musorstom 6: stn CP 438, 20°23' S, 166°20' E, 780 m, 2 rv. Stn DW 488, 20°49 S,
1 67°06' E, 800 m, 2 lv, 2 rv. — Stn DW 489, 20°48' S, 167°06' E, 700 m, 2 fragm. rv.

New Hebrides Arc. subpso: dive 7, sample 706, 15°50' S, 166°42' E, 2000 m, 2 spms.

Distribution. — Dijkstra (1989) described this species from New Caledonia, and recorded
material from the Philippines and Indonesia (Dijkstra 1989, 1990). Now also found from the Loyalty
Islands and the New Hebrides Arc. Present specimens living at 1567-2000 m.

Description. Shell medium-sized, fragile, semi¬

transparent white, interior nacreous, suborbicular. inequi-
valve, variable in shape, left valve Strongly convex, right valve
flat to slightly concave, up to ca. 35 mm high, auricles
unequal, umbonal angle ca. 110°.

Prodissoconch ca. 300 pm in height.

Left valve covered with rather regular, closely spaced,
radial costae, which multiply and enlarge towards ventral
margin. Irregular close-set concentric lamellae between and
across radial costae, commencing at ca. 10 mm shell height
and enlarging to form slightly scaly lamellae near periphery.

Auricles continuous with disc margins, sculpture similar to
that on disc. Hinge line straight.

Right valve irregularly indented, ornamented with delicate,
rather regularly spaced, undulating radial lirae. Anterior
auricle separated from disc by a distinct suture, posterior
continuous with disc margin, auricular sculpture similar to
that on disc. Byssal notch present. Inactive and active
ctenolium (4-7 teeth). Hinge line somewhat elevated. Two
indistinct resilial teeth are present; no dorsal teeth (crura).
Resilifer triangular, outer ligament very small.

Remarks. — The present specimens of P. levii will make it possible to supplement the original
description with information on the anatomy (Knudsen & Dijkstra, in prep.). For comparison with
Pseudohinnites adamsi (Dali, 1886) from the West Indies, see Dijkstra (1989. 32).

Genus Hyalopecten Verrill, 1897

Hyalopecten Verrill, 1897: 63, 71. pi. 18, fig. 5. Type species (OD): Peeler, rndatus Verrill & S. Smith in Verrill, 1885; Recent,
off Chesapeake Bay, Maryland, USA, 2603 m.

Remarks. — P. undatus Verrill & S. Smith is a homonym of Pecien undatus Defrance, 1825
and a senior synonym of Hvalopecten ddectus Verrill & Bush in Verrill, 1897 [Recent, off Martha s
Vineyard, Massachusetts, USA, 3316 m], which then becomes the valid name of the type species (see
North, 1951a: 234).

Diagnosis. — Shell inequivalve, compressed, hyaline; valves with concentric undulations or
corrugations and sculptured with fine radial lirae on one or both valves; auricles unequal, byssa
notch well-developed, ctenolium present.

Distribution. — Miocene to Recent, worldwide, 800-7000 m (Knudsen, 1970).

Remarks. — Grau (1959: 55) treated Hyalopecten as a subgenus of Cyclopecten (a
propeamussiid), and was followed by Knudsen (1970: 97) and Rombouts (1991: 79) Hertlein (1969:
N354) considered Hvalopecten a subgenus of Palliolum (a pectmid), placed in the Eburneopecten
group, and Vaught' (1989: 119) placed Hyalopecten in the tribe Eburneopectmim. Hyakrpecten is
considered a valid Recent genus of Pectinidae by other authors (Bernard. 1983; Sc.hein, 1988, 789
Dell, 1990; Dijkstra, 1991). Waller & Marincovich (1992: 219) consider Hyalopecten as related
to a radially sculptured ancestor, Praechlamys Allasinaz, 1972 from the Triassic, without placing it
in a higher taxon.

Source MNHN. Paris

48

HENK DIJKSTRA

Type material.
Type locality.

710 m.

Hyalopecten mireilleae sp. nov.

Figs 79-82

Holotype mnhn. Paratypes: 2 mnhn, 2 hd.

S New Hebrides Arc, volsmar, stn DW 55, 20°59'

S,

1 70°02' E,

Material examined. — New Caledonia, biocal: stn CP 72, 22°09' S, 167°33' E, 2100-21 10 m
1 juv. rv (paratype). — Stn KG 85, 20°59' S, 167°00' E, 1639 m, 1 lv (paratype).

Loyalty Islands, musorstom 6: stn CP 438, 20°23' S, 166°20' E, 780 m. 1 lv (paratype hd) — Stn
DW 489, 20°48' S, 167°06' E, 700 m, 1 rv (paratype hd).

New Hebrides Arc. gemini: stn DW 55, 20°59' S. 170°02' E, 710 m, 1 spm (holotype).

Distribution. — Eastern New Caledonia to the New Hebrides Arc, 700-2110 m living in

710 m.

Description. Shell small, up to ca. 14 mm high, fragile,
oblique, elongate, inequivalve, inequilateral, semi-transparent
white, right valve more convex than left, auricles unequal,
umbonal angle ca. 90”.

Prodissoconch ca. 200 pm in height.

Left valve externally sculptured over entire surface with
regular spaced concentric lamellae that enlarge towards
ventral margin. Lamellae near periphery more strongly
curved and more irregularly and closely spaced. Delicate
radial lirae between concentric lamellae, more prominent
directly beneath each concentric lira. Anterior and posterior
auricles with curved, closely spaced concentric lirae. somew¬
hat more prominent and widely arranged on anterior.
Anterior auricle larger than posterior auricle, sculpture

similar to that of disc margins. External sculpture visible
from interior. Interior surface nacreous.

Right valve with regular concentric lamellae and many
delicate interstitial radial lirae similar to those on left valve.
Anterior auricle separated from disc by distinct suture,
sculptured with irregularly spaced, concentric lamellae, deve¬
loped into spinous scales on dorsal margin. Posterior auricle
with close-set concentric lamellae and continuous w'ith the
disc margin. Resilifer triangular. No cardinal crura exposed.
Only an inactive ctenolium is developed (two teeth of active
ctenolium are developed in paratype from biocal: stn CP 72).
Byssal notch present.

Dimensions of the holotype: H 13.4, L 11, D 5.6 mm.

Remarks. — A somewhat similar species is Hyalopecten tydemani Dijkstra, 1990 from the
Ceram Sea (Indonesia), which is smaller, less oblique, and ornamented with very closely spaced
concentric lamellae. Moreover, the delicate interstitial radial lirae are absent. The western Atlantic
species H. slrigillatus (Dali, 1889) is more like H. tydemani, but is somewhat more strongly sculptured
with closely spaced concentric lamellae. The anterior auricle of the right valve in both species is
relative smaller in H. mireilleae. Hyalopecten pudicus (E.A. Smith, 1885) from the southern Indian
Ocean is more depressed, slighly more orbicular, with an undulating surface. H. pudicus is more
similar to H. dilectus from the Atlantic Ocean. Hyalopecten neoceanicus (Dali, 1908) from the
Ga apagos Islands is more orbicular and depressed with a slightly undulating surface, and reticulate
sculpture. Hyalopecten profundicola (Okutani, 1962) from abyssal depths of Japan is more depressed,
more oblique and orbicular, and the right valve has more closely spaced lamellae. Hyalopecten hadalis
(Knudsen. 1970) from the Kermadec Trench is somewhat larger (up to 20 mm in height), oblong,
strongly undulated, and sculptured with fine, close-set radial striae, anterior auricle of the right valve
very small and byssal notch well developed. Hyalopecten sp. (Knudsen, 1970: 102) from the Tasman
Sea is much larger (up to ca. 35 mm), also oblong and strongly undulated, and sculptured with radial

Etymology. Named after Mrs Mireille Moreau, technician at mnhn, who has skilfully been
processing the bivalves from the various expeditions around New Caledonia.

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

49

19-86. 79-80. Hyalopecten mireiileae, hololype. 1 3.4 x 1 10 mm (lv)_ — 79. left ^'1.7^^' exterior"' ^'s^'righTvalve.

81-82. musorstom 6: stn CP 438. paratype. 12.1 x 10 4 mm (rv). - SLnght vaKe. extern) • g

interior. — 83-86. Delectopecten fluctuate, calsub: dive 15, 4.5 x 5.1 mm (dO).

84. left valve, interior. 85. right valve, exterior. - 86. right valve, interior.

50

HENK DIJKSTRA

Subfamily Camptonectinae Habe. 1977

Diagnosis. — Pectinidae with antimarginal microsculpture, weak radial costae or striae
sometimes present; shell microstructure of outer lathic or foliated calcite, inner crossed-lamellar
aragomtic layer to margins; auricles unequal; byssal notch with ctenolium.

Genus Delectopecten Stewart, 1930

^7r,es (0D): Pecm m

c„ , t DlAGN(’SIS- - Shell inequivalve, small, fragile, nearly orbicular, valves nearly equal convex
sculptured with concentric rows of scales or vesicles, spinose radial ridges and/or fine diverged radial
striae, byssal notch well-developed; ctenolium present.

Distribution. — Early Cretaceous to Recent, worldwide, 27-4256 m (Clarke, 1962).

Remarks. — Grau (1959: N38) considered Delectopecten a subgenus of Cvclooecten and
synonymized Arctmula Thiele, 1934 and Catillopecten Iredale, 1939. Characters o/ Arc t inula
however, are similar to those of Similipecten Winckworth, 1932 (Schein, 1989; 96) now placed in

a»° i°nSidered a RCCent genUS ^ Propeamussiidae (Zv» \m.
89), and most closely related to Bathypecten Schein-Fatton, 1985.

(1969: N354) treated Delectopecten as a subgenus of Palliolum, placed in the

p < J ill grouP; Current|y’ however, Delectopecten is interpreted as a Recent genus of

199I: & M— c„, ,992: 2,9) an§d JSUd

Delectopecten alcocki (E.A. Smith, 1904)
Figs 111-114, 147-150

Pecien alcocki E.A. Smith, 1904: 13.

Other references:

“ " ALCOCK' Annandale & McGilchrlst. 1907: pi. 18, figs 4. 4a-b. - Knudsen. 1967: 282, pi. 2, figs 3-4.

PDecten (Pseudamussium) alcocki - Thiele & Jaeckel, 1931: 6.

rropeamusstum (Hyalopecien) alcocki - Winckworth, 1940- 26

Delectopecten alcocki - Poutiers. 1981: 331, pi. 1, fig. i. - Dukstra, 1991: 26, fig. 87.

Type material. — Lectotype live taken, (Figs 147-150' H 19 0 1 18 0 mn mrn\

designated, zsi M669/1, and 29 paralectotypes zsi M667-668, M670-698/1.

Type locality. Off southern India, “ Investigator ”, stn 232, 7° 17' N, 76°54' E, 786 m.
Material examined. - The type material

* lv- ■ rv.

jAECKB?a™nUSDS7Nmm(1^ H°n describ=d‘his sPeci« from off southern India. Thiele &
jaeckel (1931), Knudsen (1967) and Dukstra (1991) added records from the Philippines, the Banda

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

51

Sea and Sumatra (Indonesia), Gulf of Aden and eastern Africa. Now its range is extented to the
Chesterfield Islands. Present material living at 650-660 m.

Dfscription. — Shell up to ca. 15 mm high, orbicular,
semi-transparent white, auricles nearly equal in size, umbonal
angle ca. 125°.

Prodissoconch ca. 280 pm (Knudsen, 1967).

Left valve with delicate diverging radial striae over entire
disc Crenulated concentric growth lines with radially aligned
scales. Auricles identically sculptured and continuous with
the disc. Hinge line straight.

Right valve sculptured as on left valve, although somewhat
weaker. Anterior auricle separated from disc by a suture,
with 2-4 radiating ridges and radial lamellae. Byssal fasctole
and notch strongly developed. Inactive and active ctenolium
(3-5 teeth) present. Resilifer triangular. External sculpture
barely visible from interior.

Anatomy described by Knudsen (1967: 282).

Remarks. — The present material corresponds in all aspects with the type material. Further
notes on soft parts and ecology are given by Knudsen (1967: 283) and Dijkstra (1991: 26).

Delectopecten fluctuatus (Bavay, 1905)
Figs 83-86

Pecten (Clilamys) fluctuatus Bavay, 1905: 188. pi. 17, Figs 3a-b.

Type material. — Holotype, ?live taken, zsi M 3359/1.

Type locality. — Off Andaman Islands, depth unknown.

Material examined. — The type material.

Loyalty Islands, calsub: dive 15, 20°37' S, 166°58'E, 538 m, 1 spm (juv.).

Distribution. Andaman Islands and the Loyalty Islands. The present specimen living at

538 m.

Description. Shell up to ca. 10 mm high, fragile,
suborbicular, undulating, slightly inequilateral, semi¬
transparent white to opaque cream, auricles unequal, umbo¬
nal angle 90°-105°.

Prodissoconch ca. 200 pm in height.

Left valve undulating and complexly sculptured with line,
regularly spaced, scaly radial riblets, commencing at ca. 2
mm shell height and progressively enlarging and multiplying
by intercalation to ca. 30. Delicate, closely spaced lamellae
between radial riblets commencing early on disc, enlarging
towards ventral margin. Entire surface of disc covered with
microscopic diverging striae. Anterior auricle larger than
posterior, both auricles sculptured with delicate, close-set.

concentric lamellae, and very fine radial striae; posterior
auricle more nearly continuous with disc margin. Hinge line
straight. A few small scales on dorsal margin. External
sculpture sometimes visible from interior of valves.

Right valve undulating, sculptured as in left valve but with
smaller scaly radial riblets. Anterior auricle separated from
disc by distinct suture, posterior continuous with disc.
Anterior auricle with 4 very weak radial riblets. covered with
concentric lamellae. Strongly developed scales on antero-
dorsal margin. Byssal fasciole and notch well developed.
Inactive and active ctenolium (3 teeth) present. Resiliter
triangular. No cardinal crura.

Remarks. — The present immature specimen is similar to the type specimen, although
semi-transparent white instead of opaque cream as in the holotype. Moreover, the scaly sculpture is
somewhat more prominent in the holotype. Delectopecten alcocki is closely related but is more
orbicular and larger, with an almost smooth exterior surface, and without any undulations. D.
macrocheiricola (Habe, 1951), from Japan, is more weakly sculpture with diverging scratches and the
surface of its disc is more weakly undulating, especially on the early disc. Habe s species is a so muc
larger (up to 25 mm high). D. musorstomi is orbicular with a stronger sculpture, diverging radia
scratches and weaker concentric lirae. Dijkstra (1989: 32) suggested that D. fluctuatus could possibly
be a Pseudohinnites, but after examining the badly damaged holotype (zsi), I have come to tne
conclusion that it is a Delectopecten.

52

HENK DIJKSTRA

Figs 87-92. 87-90. Cyclochlamys favus, biocal: sin DW 46. 87. right valve, exterior, scale bar 200 urn. - 88. right valve,

dissoconch stage, scale bar 50 pm. 89, right valve, hollow sections internal ligament, scale bar 25 urn. 90. right
valve, hexagonal prismatic microstructure, scale bar 10 pm. 91-92. Parvamussium multiliratum, biocal: stn CP 75,
paratypes. 91. left valve, exterior, scale bar 350 pm. 92. right valve, interior, scale bar 350 pm.

Source : MNHN. Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

53

Delectopecten musorstomi Pouliers, 1981
Figs 69-70

Delectopecten musorstomi Poutiers, 1981: 331. pi. 1, figs 2-3

Other reference:

Delectopecten musorstomi — Dijkstra. 1991: 26.

Type material. — Holotype live taken, mnhn.

Type locality. — N of Lubang, Philippines, musorstom 1, stn 18, 13°57' N. 120° 17’ E.
150-159 m.

Material examined. — The type material.

New Caledonia, biocal: stn DW 64, 24”48' S, 168°09' E, 250 m, 1 spm.

Distribution. — Philippines (Poutiers, 1981), Indonesian Archipelago (Dijkstra, 1991), now

extended to New Caledonia. Present material liv

Description. Shell small, up to ca. 5 mm high,
orbicular, transparent, equivalve. equilateral, anterior auricle
somewhat larger than posterior, umbonal angle ca. 120 .

Prodissoconch ca. 200 pm in height.

Left valve covered with microscopic diverging radial striae
and 14-20 fine, irregularly spaced, scaly radial lirae. that
commence at ca. 1 mm shell height and progressively enlarge.
Fine regularly spaced, somewhat crenulated, concentric
growth fines. Scales often broken. Auricles continuous with
The disc and similarly sculptured. Hinge line straight.

I at 2j0 m.

Right valve sculptured similar to that of left valve, with
slightly stronger microscopic concentric lirae in early disc,
that commence at about 1.5 mm shell height. Anterior auricle
separated from the disc by distinct suture, with 4-5 radia
ridges and concentric lamellae, the latter strongly developed
at dorsal margin. Byssal fasciole and notch well developed.
Inactive and active ctenolium with 2-3 teeth. Resilifer trian¬
gular. Valves sometimes with a few white spots. External
sculpture visible from the interior.

Remarks. — The present specimen is very similar to the holotype of D. musorstomi from the

' Poutiers (1981: 332) compared Delectopecten musorstomi with D. polyleptus (Dali, 1908) trom
the Galapagos Islands and Chile. Another related species is D.fluctuatus which has similar sculpture,
though it is larger (up to 10 mm high) and the disc is undulating, and there are more numerous sea y
radial lirae.

Subfamily Chlamydinae Teppner, 1922
Tribe Chlamydini Teppner, 1922

Diagnosis. Chlamydinae with microsculpture of diverged striae (antimarginal); shagreen
(reticulated) microsculpture present at least in early growth stages.

Genus Laevichlamys Waller, 1993

Laevichtamys Waller. 1993: 204. Type species (OD): Pecten multisquamalus Dunker, 1864; Recent, Cuba.

Diagnosis. Byssate, non-cemented, free living Chlamydini; shagreen (reticulated) micros-
culpture absent; sculptured with closely spaced, weak radial costae on both valves, reguia
commarginal lirae absent; byssal notch with ctenolium.

54

HENK DIJKSTRA

600 am Id ll , , vesiculaturn, biocal: stn DW 44. paratypes. 93. left valve, exterior, scale bar

600 un ’ o 4-, rf i ’ fXten0^ antero-rnarginal detail, scale bar 100 pm. - 95. left valve, exterior, scale bar

n6' ff Va r, e en0r' <?°,rsal1regl0n detaiK scale bar 200 97 • p retiolum, musorstom 5: si

paratype, left valve, exterior, central detail, scale bar 100 pm.
exterior, postero-marginal detail, scale bar 50 pm.

no _ stn CP 363,

98. Cyclopecten horridus, biocai.: stn DW 08, left valve.

Source MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

55

•9-106. 99-102. Parvamussium iheiidis, Chalcal 2: sin DW 73, 10.0 x 9.8 mm (lv), 8_9 x 8.9 mnJ ^ ^vamusshm
exterior. 100. left valve, interior. 101 . right valve, extenor. 102 nght valve ntenor. ™ ££££“_

lexturalum , musorstom 5: stn DW 300, 6.5 x 6.6 mm (db). 103, left valve, ex . .

105. right valve, exterior. 106. right valve, interior.

Source .

56

HENK DIJKSTRA

f '/ 107-110. Parvamussium pauciliratum, musorstom 5: stn DW 263, 8.2 x 8.1 mm (Iv). 7.2 x 7.2 mm (rv). —
07, left valve, exterior. - 108. left valve, interior. 109, right valve, exterior. 110. right valve, interior. 111-
1 14, Delectopecten alcocki, corah. 2: stn DE 14. 10.4 x 10.1 mm (db). 111. left valve, exterior - 112 left valve

interior. 1 13, right valve, exterior. 1 14, right valve, interior.

Source :

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

57

Distribution. — Miocene to Recent. Indo-West Pacific and western Atlantic; littoral to
sublittoral.

Laevichlamys kauaiensis Dali, Bartsch & Rehder, 1938
Figs 121-122

Chlamys kauaiensis Dali, Bartsch & Rehder. 1938: 92, pi. 22, figs 9-10.

Other references:

Chlamys kauaiensis - Kay, 1979: 525. figs 168 F-G. Earle, 1985: 1, 4. fig.

Type material. — Holotype, rv, usnm 335674.

Type locality. Off Hanamaulu warehouse, Kauai. Hawaii Islands, Albatross , stn 4132,
470-571 m.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DW 329, 20°23' S, 158"47' E, 320 m, 1 rv.
20°45' S, 1 60°54' E, 500-510 m, 1 rv.

New Caledonia. “Vauban” 1978-79: stn 40, 22"30' S, 166"24' E, 250-350 m, 1 rv.

Stn DC 388,

Distribution. Hawaiian Islands (Dall, Bartsch & Rehder, 1938; Kay, 1979; Earle.
1985), now also Chesterfield Islands and New Caledonia. Present material taken dead at 250-510 m

Description. Shell small, up to ca. 6 mm high, Iragile.
suborbicular, equivalve, right valve slightly more convex than
left, anterior and posterior auricles very unequal in size,
umbonal angle 90-95°.

Prodissoconch ca. 240 pm in height.

Left valve sculptured with 24-28 fine regularly spaced
radial riblets, with delicate scales (often eroded). Microscopic
diverging scratches between radial riblets. Auricles sculptured
with 3 or 4 scaly axial riblets that are somewhat more
prominent on anterior. Hinge line straight.

Right valve sculptured like the left. Anterior auricle with
5 of 6 scaly radial scaly. Antero-dorsal margin strongly
denticulate. Posterior auricle narrow, with 3 or 4 spinous
riblets. Byssal fasciole and notch well developed. Inactive and
active ctenolium with 4-6 teeth. Resilifer triangular. Anterior
resilial tooth more prominent than posterior. Internal plicae
extending from central sector to ventral margin.

Left and right valves usually cream, white or bright yellow
or orange-red. sometimes with small white spots.

Remarks. - The present specimens are very similar to the holotype of L. kauaiensis, ^although
somewhat more brightly coloured, with slightly stronger sculpture on the posterior auricle ol the right
valve. L. kauaiensis is rather variable in radiation and coloration.

Also similar to L. kauaiensis is L. allorenti (Dijkstra, 1988) from Reunion and adjacent areas
(see Dijkstra, 1988: 21).

Genus Veprichlamys I redale, 1929

Veprichlamys Iredale, 1929: 164. 188 (Proposed as a subgenus of Mimaclilamys]. Type species (OD): Chlamys penllustns
Iredale, 1925; Recent, off Victoria. Australia, 274-457 m.

Diagnosis. — Non-cemented thin Chlamydini, obliquely oval; slender radial costae with
prickly lamellae, interstices broad with or without secondary radial riblets; microsculpture
longitudinal or diverged scratches; auricles very unequal; inner ribs undulated, lacking cannae, yssa
notch well-developed with ctenolium.

Distribution. — ?Miocene to Recent, Pacific Ocean, including New Zealand and the
Galapagos Islands; 120-1280 m (Dijkstra. unpubl. data).

58

HENK DIJKSTRA

Veprichlamys kiwaensis Powell, 1933
Figs 119-120

Chiamys kiwaensis Powell, 1933: 371, pi. 40. figs 1-5.

Other reference:

Clilamys kiwaensis - Powell, 1979: 378. pi. 72. fig. 14.

Type material. — Holotype live taken, aim AK70167.

Type locality. — 400 miles W of New Plymouth, New Zealand, 39‘W S, 168°50' E.
1097-1280 m.

Material examined. — New Zealand. Off Taiaroa Heads, 475-640 m, 2 v (nmnz M9138).
41°35'S, 174°53' E, Palliser Bay, 366-549 m, 5 spms (nmnz M9850). — 44°45.6' S, 171°05'E, off
Taiaroa Heads, 549 m, 2 v (nmnz M12827). — 41°30.7' S, 174°58.4'E, Turakirae Trench, Cook
Strait, 448-512 m, 7 spms (nmnz M29148). — 41°30.5' S. 174°54' E, Turakirae Trench, Cook Strait,
640-658 m, 3 spms (nmnz M29129). — 43°I4' S, 173°39' E, Pegasus Canyon, off Banks Peninsula,
512-1006 m. 18 spms + many valves (nmnz M52769). — 42°35' S, 173°41' E, off Kaikoura, 640 m.
5 spms (nmnz M53676). — 41°55.9' S, 174°43.2' E, off Cape Campbell, 424-454 m, 14 spms
(nmnz M59679). — 41°55.8'S, 174'’40.7' E, SE of Cape Campbell, 434-446 m, 10 spms + many
valves (nmnz M60407). — 37°22.0' S, 176°28.5' E, off Mayor I., 388-448 m, 2 spms (nmnz M60094).
— Conway Rise, Kaikoura, 400 m, 10 spms (nmnz M76106).

New Caledonia, musorstom 4: stn DW 225, 22°52' S, 167°23' E, 590-600 m, 1 rv.

New Zealand, living in 366- 1006 m; New Caledonia, taken dead at 590-600 m.

Distribution.

Description. — Shell equivalve, up to ca. 35 mm in height,
suborbicular to oblique, compressed, fragile, auricles strongly
unequal.

Valves sculptured with 20-24 scabrous radial costae, inters¬
paces with Camptonectes-\ike radial striae, sometimes absent
in late growth stages. Anterior auricle of left valve sculptured

with 8 radial riblets, 5 on right valve: posterior auricles nearly
smooth.

Byssal notch well-developed, with ctenolium. Colour out¬
side dull-white or creamy, sometimes with the radials
brownish-pink; inside nacreous.

Remarks. — The present material from off New Caledonia is very similar to the type
specimens of C. kiwaensis from New Zealand, although smaller (height 12 mm). The microscopic
diverging striae between the radial costae (“ Camptonectes " — like divarications of Powell’s
description) are strongest near the antero-lateral margin in the present material, and absent near the
ventral margin.

Related species are Veprichlamys jousseaumei Bavay, 1904 from southern Japan to Indonesia,
V. perillustris from SE and S Australia, and V. incantata (Hertlein, 1972) from the Galapagos Islands.
Waller (1993: 236) mentioned also V. onzola (Olsson, 1964), a Neogene species from northwestern
Ecuador. V. jousseaumei is slightly orbicular in shape, and has numerous radial costae (30-38). V.
perillustris is more oblique, and has fewer radial costae (20-26), which are more strongly imbricated.
V. incantata is larger (height up to ca. 55 mm), less obliquely oval, and sculptured with ca. 20 radial
costae and interstitial radial riblets.

Tribe Aequipectinini Nordsieck, 1969

Diagnosis. — Chlamydinae with complex sculpture of radiating costae, regularly lamellose
with series of hollow chambers or secondary interstitial concentric lamellae, equilateral and auricles
more equal in late growth stages.

Source :

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

59

Figs

115-122. 115-118, Cryplopecten bulla t us, MUSORSTOM 6: sin CP 464. 15.9 x 17.0 ram (db).

rior. - 116, left valve interior. 117. right valve, extenor. - 118. right valve, mter or.

kiwaensis, musorstom 4: stn DW 225. 12.2 x 10.8 mm (rv). 119 nght valve, exterior. 120. nght valve, in

rior. - 121-122. Laevichlamys kauaiensis. " Vauban " 1978-79: stn 40. 6.9 x 6.9 mm (rv). - 121. right valve,
rior. — 122. right valve, interior.

Source

60

HENK DIJKSTRA

Genus Cryptopecten Dali, Bartsch & Rehder, 1938

Bartschn& Rehder, 1938: 93. Type species (OD,: Cryptopecten alii Dali, Bartsch & Rehder. 1938; Recent,

Synonym:

Corymbtchjamys Iredale, 1939: 367.Type species (OD): Chlamys corymbiatus Hedley, 1909; Recent, off Queensland. Australia.

Diagnosis. — Shell small, orbicular, convexity variable, laterally compressed, auricles nearly
equal, valves sculptured with 12-25 radial costae, regularly lamellose, interspaces with fine imbricated

Shct^^P10^ WUh 3 ^ radiaI riblets; byssal notch well-developed and moderately deep.

Distribution. — Lower Miocene-Recent. Indo-Pacific, western Atlantic-
bathyal (Hayami 1984: 116).

upper shelf to upper

Cryptopecten buUatus (Dautzenberg & Bavay, 1912)

Figs 115-118

Pecten (Chlamys) bullatus Dautzenberg & Bavay, 1912: 17. pi. 27, figs 1-2.

Synonyms:

Cryptopecten alii Dali. Bartsch & Rehder, 1938: 93 pi ’3 figs 1-4 7
Cryptopecten complanus Wang, 1983: 402, 405. figs 1.1-7. ’

Other references:

Chlamys ( Aequipecten ) tissotii - Kuroda, 1932: app 95
Chlamys alii - Kay. 1979: 524. fig. 168A.

g It * 2- P'- P«- '«• % 3- PI- . I- fig. 3; WaoNER, ,989:

r- T/YPu ^AtTER1AI-; - 9 bullatus: holotype, live taken, zma 3.12.006, 2 paratypes zma 3.12.007.
paratype'iOAS °M lTo^r6 l?3194‘ _ Complanus: hol«type. live taken, ioas Ml 1072.

171°19'TFP?7s°mALITz; ~n Cc bu!!atus: Su>u Archipelago. Philippines. “Siboga”, stn 105, 6°08' N,
, -75 m C. al/i. South coast of Oahu. Hawaii Islands, “ Albatross ”, stn 381 1, 436-461 m
C. complanus : East China Sea, 31°05' N, 128°00' E, 147 m. ’

Material examined. — The type material.

Chesterfield Islands. CHALCAL 1: stn D 35, 19°45'S, 158°26'E 210 m 1 lv

musorstom 5: stn DW 255, 25*>15' S. 159°55' E, 280-295 m. 3 lv. - Stn DW 258, 25"3T S 159*46' E

300 m, I spm. - Stn DW 277, 24“ 1 1' S, 159"35' E, 270 m, 1 rv — Stn CP 279 24*09' s’ 159"38' F

360*300 iV- T Stn CP 289’ 24°°2' S- 1 59°38' E" 273 2 lv- - Stn DW 299, 22*48' S 59*24' E

20 23 S misl& E T 3°6- 22°08^ 159°21' E- 375-4.5 m. 2 lv, I 'rv. - Stn DW 3li

ZU Z3 S 158 44 E, 340-355 m, 1 rv. — Stn DC 388, 20°45' S, 160°54' E, 500-510 m I lv

corail : stn DW 114, 19°25' S, 158*38' E. 217 m, 1 lv, I rv. — Stn DW 129 19*28' S 158°34'E
215 m, 1 rv. - Stn CP 131, 19*25' S, 158°38' E, 215-217 m, 1 lv ’

l978;79: Stn 33- 22°33' S' I66°25' E- 29°-350 m- I lv. - stn 37, 22°32' S
166 26 E, 175-250 m, 1 rv. - Stn 40, 22*30' S, 166*24' E, 250-350 m 1 lv
biocal: stn CP 105, 21*31' S, 1 66°22' E, 330-335 m 1 lv
musorstom 4: stn DW 219, 23°03' S, 167°33' E, 760 m, 1 rv.

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

61

smib 2: stn DW 14, 22°53' S, 167°13' E, 405-444 m, 1 spm.

smib 5- stn DW 77, 23°41' S, 168°01' E, 270 m, 1 spm. — Stn DW 91, 22°18' S, 168°41' E, 340 m,

V rv. — Stn DW 92, 22°20' S, 168°41' E, 280 m, 1 rv. — Stn DW 98, 23°02' S, 168°16' E, 335 m,

I iv;_ stn DW 105, 23° 14' S. 168°05' E, 310 m, 1 lv.

Lovaltv Islands, musorstom 6: stn DW 391, 20°47' S, 167°06' E. 390 m, 1 spm, 2 rv. — Stn DW 392,
20°47' S 167”05' E 340 m, 1 rv. — Stn DW 398, 20°47' S, 167°06' E, 370 m, 1 lv. — Stn DW 406,

20°4 1 ' s' 1 67°07' E, 373 m, 1 spm. 1 lv. Stn DW 416, 20°42' S, 166°60' E, 343 m, 3 rv. — Stn DW

421 20°26' S 1 66°40' E, 245 m, 1 rv. Stn DW 428, 20°24' S, 166° 13' E, 420 m, 1 rv. Stn DW

457’ 2 1 "00' S, 167°29' E, 353 m. 2 lv, 2 rv. — Stn DW 459, 21°01' S, 167°31' E, 425 m. 1 spm, 1 lv,

] rv _ Stn DW 464, 21°02' S, 167°32' E, 430 m, 3 spms. 4 lv. — Stn DW 479, 21° 09' S, 167°55' E,

310 m 1 lv ? rv — Stn DW 480, 21°09' S. 167°56' E, 380 m, 1 rv. - Stn DW 481, 21°22' S, 167°

50' E, 300 m, 1 lv. — Stn DW 485, 21°23' S, 167°59' E, 350 m, 1 rv. — Stn DW 487, 21°23' S.

167°46' E, 500 m, 1 spm.

New Hebrides Arc. volsmar: stn DW 7, 22°26' S, 171°44' E, 325-400 m, 2 spms, 2 lv, 2 rv. — Stn DW
8. 22°25' S, 171 °43' E, 630 m, 1 fragm. lv. — Stn DW 9, 22l>23' S, 171°42' E, 275-300 m, 1 lv. - Stn
DW 16, 22°25' S, 171°4r E, 420-500 m, 1 lv, 1 rv. — Stn DW 17, 22°23' S, 171°42' E, 260-300 m,
1 spm. stn DW 38, 22"2T S, 168°43' E, 380-420 m, 1 lv.

Distribution. — Throughout the western, southwestern and central Pacific, western Indian
Ocean, Chesterfield Islands, New Caledonia, Loyalty Islands and New Hebrides Arc. Present material
living at 260-500 m.

Description. Shell up to ca. 25 mm high, usually
20 mm, moderately thin, depressed, somewhat oblique
towards posterior half, equivalve, inequilateral, right valve
slightly more convex than left, auricles unequal in size,
umbonal angle ca. 110°.

Prodissoconch ca. 230 pm in height.

Left valve with 17-20 radial costae (commonly 18-19).
commencing at 2 mm shell height and gradually enlarging,
strong on central ridge, with numerous cavities on their sides.
Interspaces commonly narrower than costae, finely lamellate.
Anterior auricle slightly larger than posterior, sculptured
with 2 prominent radial riblets and 2-4 interstitial radial
riblets. Posterior auricle with 1 strong radial riblet and 3-

5 smaller ones between disc margin. A few denticulated scales
on antero - and postero-dorsal margins. Hinge line straight.

Right valve similar to that of left valve, with finer
interstitial lamellae, and stronger denticulate scales on dorsal
marein. Hinee line somewhat raised. Byssal fasciole relative
small, byssafnotch rather deep. Inactive and active ctenolium
with 4 or 5 teeth. Antero — and postero-lateral margins of
disc scarcely gaping. Resililer elongate triangular. Inner
surface of both valves strongly plicate.

Coloration very variable, commonly reddish brown with
pale oblique or zigzag markings, sometimes uniform yellow
or purple.

Remarks. — The present material is very similar to the type specimens of C. bullatus from the
Sulu Archipelago, but the latter have more radial costae (22). The auricles also have a lower number
of radial riblets. The convexity and sculpture vary with the bathymetric range and geographic
distribution. For further discussion see Hayami (1984: 98), Wagner (1989: 61), and Dijkstra (1991:
36).

62

HENK DIJKSTRA

FlGS ‘ viw2«teriOT I241mT^^ WW0”'’ lectot,yPe- 51.8 x 50.0 mm (db), - 123. left valve, exterior. - 124 right
valve! imerior - 127 S. Cal “rior "Ttt ™ X 7’-9 mm [t1 ,25' lef* valve- exterior. 126 fef!

Se? iSr (db)' - 129’ 'S°Si

Source . MNHN. Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

63

Figs 133-142. — 133-137. Propeatnussium alcocki, lectotype. 40.4 x 39.8 mm (db). 133. original label ^’Jamussiwn

exterior. 135. left valve, interior. 136. right valve exterior. 137 ngh valve Sleft valve.
andamanicum, lectotype, 30.2 x 22.4 mm (db). — 138. original label. 1.9,
interior. — 141, right valve, exterior. — 142. right valve, interior.

Source .

64

HENK. D1JKSTRA

Figs 143-150. 143-146. Propeamussium meridionale , lectotype, 13.8 x 13.4 mm (db). 143. left valve, exterior. 144. left

valve, interior. - 145, right valve, exterior. — 146, right valve, interior. - 147-150, Delectopeclen alcocki, lectotype,
19.0 x 18.0 mm (db). — 147. left valve, exterior. 148. right valve, exterior. 149, left valve, interior. 150. right
valve, interior.

Source : MNHN. Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

65

Figs 151-154. 151-152. Parvamussium pauciliralum , leclotype. 7.5 x 7.4 mm (db). 151. left valve, exterior. 152 right

valve, exterior. 153-154, Parvamussium scitulum , lectotype, 4.4 x 4.5 mm (lv). 153, left valve, exterior. 154, lei t

valve, interior.

DISCUSSION

Thirty species of Pectinoidea are now known from the Chesterfield Islands. New Caledonia
and the Loyalty Islands, at depths below 100 m. Including Pseudohinnites levii, which had been
described earlier in a seperate paper. 10 species, precisely a third of the fauna, were undescribed
before this survey. The collection studied here comprised 236 lots, i.e. 8.7 lots per species. Only
4 species ( Cyclopecten pellucidulus, Delectopecten fluctuatus, D. musorstomi and Vepriehlamys
kiwaensis), or 13.3% of the total fauna, are known from a single sample. All these observations
indicate that the fauna is probably now rather adequately sampled, at least at depths shallower than
1500 m. The bathymetric distribution of the species is summarized in Table 2. The highest diversity
is in the 600-800 m depth interval, with 14 live-taken species; respectively 9 and 8 species have been
taken alive in the 200-400 m and 400-600 m depths intervals. Below 800 m, the diversity drops to
4 species. Only 3 species ( Propeamussium meridionale, Parvamussium multiliratum and Pseudohinnites
levii ) have been taken alive deeper than 1500 m, but this may reflect the fact that much fewer samples
have been taken in very deep water.

Up to 4 species have been collected from a single station: musorstom 6: stn CP 438, 780 m
(Propeamussium alcocki and P. watsoni, alive; Hyalopecten mireilleae and Pseudohinnites levii, dead).
A maximum of 2 species have been collected alive in a single station. However, ca. 70% ol the
stations with pectinoids contain a single species.

66

HENK DIJKSTRA

Table 2. Bathymetric range of deep-water Pectinoidea from the New Caledonia region and
Indonesia. Thick bar: confirmed living depth range; thin line: depth range indicated by empty
shells only. Solid black: data for New Caledonia; stippled: data for Indonesia.

0 400 800 1200 1600 2000 2400 m

Propeamussium alcocki

- ™

P. andanianicum

11 ““

P. caducum

P. maorium

-

P. lacteuin

P. meridionale

P. rubrotincium

P. sibogai

P. siratama

P. walsoni

P. zoniferum

Parvamussium araneum

P. car base uin

P. cassium

P. cristatellum

P. dautzenbergi

P. ina

P. margaritiferum

P. multiliraium

P. reiiaculum

“ —

P. retioluin

" 1

P. scitulum

~

P. squalidulum

P. lexuiratum

P. thetidis

P. lorresi

P. undisonum

" -

P. undosum

P. vesiculaium

P. virgaium

Cyclochtamys favus

—

Cyclopecten bavayi

C. cancellus

C. horridus

C . pellucidulus

"

Similipecten eous

Pectinella aequoris

Pseudohiniles levii

Caiillopecien micaceus

C. translucens

llyalopecien lydemani

11. mireilleae

Deleciopeclen alcocki

”

D.jluctuatus

"

D. musor stand

. “ .

Lacvichlamys kauaiensis

Veprichlamys kiwaensis

—

Crypiqpecten bullatus

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

67

Table 3. — Occurence and abundance of Pectinoidea in deep-water off Indonesia and New
Caledonia.

Stn: number of stations where the species has been recovered.

Live/dead: number of specimens taken alive and dead respectively.

Species Indonesia NC region

Sin

live

dead

Sin

live

dead

Propeamussium alc.oc.ki

-

-

-

5

37

3

P. andamanicum

-

-

-

2

2

P. caducum

11

20

11

4

2

5

P. maorium

-

6

3

9

P. lacteum

1

-

2

-

-

-

P. meridionale

1

1

20

18

46

P. rubrolincluin

-

-

-

24

70

52

P. sibogai

1

1

-

15

23

12

P. siratama

1

-

1

-

-

-

P. watsoni

-

-

12

226

4

P. zoniferum

1

1

5

-

-

-

Parvamussium araneum

3

2

1

-

-

-

P. carbaseum

14

-

73

-

-

-

P. cassium

1

1

-

-

-

-

P. cristatellum

6

1

43

-

-

-

P. dauizenbergi

2

-

3

-

-

-

P. ina

1

-

3

-

-

-

P. margaritiferum

1

1

-

-

-

-

P. muliiliratum

-

-

-

7

10

5

P. reiiaculum

-

-

-

4

7

6

/’. retiolum

-

-

-

10

53

14

P. scitulum

14

-

>100

8

7

47

P. squalidulum

-

-

-

15

5

42

P. texiuraium

3

9

1

5

3

8

P. thetidis

-

-

-

9

7

29

P. lorrcsi

1

i

-

13

4

156

P. undisonum

-

-

-

4

2

6

P. undosum

5

-

6

-

-

-

P. vesiculalum

-

-

-

15

44

17

P. virgaium

3

5

-

-

-

Cyclochlamys favus

-

-

-

2

9

-

Cyclopecien bavayi

1

i

-

-

-

-

C. canceltus

6

-

15

-

-

-

C. horridus

-

-

-

4

1

7

C. pelluc.idulus

-

-

1

4

-

Similipecten eons

3

-

7

-

-

-

Peciinclla aequoris

O

-

5

5

1

4

Pseudohinniies levii

1

i

-

14

3

36

Catillopecten micaceus

1

i

-

-

-

C. ir ansi uc.e ns

1

i

-

-

-

-

llyalopecicn lydemani

2

-

2

-

-

-

II. mireilleae

-

-

.

5

1

4

Delcc.iopcc.ten alcocki

I

i

2

1

2

D.fluctuatus

-

-

-

1

1

-

D. musorstomi

10

i

19

1

1

-

Laevic.hlamys kauaiensis

-

-

3

-

3

Veprichlamys kiwaensis

-

-

-

1

-

1

Crypiopecien hullaius

4

i

6

45

13

50

Source : MNHN, Paris

68

HENK D1JKSTRA

Cryptopecten bullatus is the species represented by the highest number of samples (45),
followed by Propeamussium rubrotinctum (24) and Propeamussium meridionale (20). The species new
to science are not particularly the rare ones: they are represented, as a mean, by 8.1 samples per
species, with a minimum of 1. and a maximum of 15 samples.

Between 200 and 1000 m, propeamussiids are not only a diverse group, they may also be
locally abundant as individuals, as the examples below indicate:

Propeamussium rubrotinctum : Loyalty Islands. MUSORSTOM 6: stn DW 412, 437 m (> 50 spins).

Propeamussium watsoni: Chesterfield Islands, musorstom 5: stn CP 323. 970 m (ca. 90 spms).

— Parvamussium retiolum : Chesterfield Islands, musorstom 5: stn CP 363, 685-700 m (40 spms).

Parvamussium vesiculatum: New Caledonia, biocal: stn DW 44. 440-450 m (37 spms).

The faunal assemblage from the Chesterfield Islands, New Caledonia and the Loyalty Islands
may be compared to that of the Indonesian Archipelago (Tables 2 and 3). The sampling efforts are
not directly comparable. In Indonesia, there are fewer than 150 deep-water stations (“ Siboga 91
stations, “ Snellius-JI 54 stations), as compared to ca. 700 stations for the New Caledonia region.
Conversely, the Indonesian Archipelago covers a larger geographical area, with more complex
topography, than the New Caledonia region. However, several papers (Dautzenberg & Bavay 1912;
Dijkstra, 1990, 1991) cover the deep-water pectinoid fauna of Indonesia, and a comparison between
the two faunal assemblages is not meaningless.

It is remarkable that, despite the difference in sampling intensity. Indonesia and New
Caledonia have exactly the same number of deep-water pectinoids, 30 species. It is not less
remarkable that only 12 species are shared: 60% of the pectinoids of New Caledonia are not recorded
from Indonesia and vice versa. Considering the lower number of stations in Indonesia, it is not
surprising that these 30 species are represented by fewer samples (Table 3): there is a total of 102
deep-water pectinoid samples, or 3.5 samples per species. As many as 14 species (46.7% of the fauna)
have been recorded from Indonesia based on a single sample. This indicates a more highly diversified
fauna, and it is very likely that additional species will be recorded when the sampling intensity
increases.

Table 4. — Summary of faunal affinities on the deep-water Pectinoid fauna from the New
Caledonian region.

Species shared with New Zealand

2

6.7%

Species shared with temperate Australia

2

6.7%

Species shared with tropical Indo-Paclfic
(of which: Species shared with Indonesia: 12)

17

56.6%

New species (actual distribution unknown,
some may be endemic)

9

30.0%

Notwithstanding the relatively low proportion of species shared with Indonesia, the New
Caledonia fauna consists mostly of species with larger Indo-West Pacific distribution (Table 4). Four
species have a more restricted South-West Pacific distribution:/,an'amw.v.v/wm thetidis and Cycloch-
tamys favus are shared with southern and eastern Australia, and Propeamussium maorium and
Veprichlamys kiwaensis are shared with New Zealand.

Source : MNHN, Paris

BATHYAL PECTINOIDEA FROM NEW CALEDONIA

69

ACKNOWLEDGEMENTS

I am much indebted to P. Bouchet and B. Richer de Forges for allowing me to study the
Pectinoidea from the New Caledonian region. 1 am also grateful to P. Bouchet and the two referees
for their valuable advise and critical comments on this paper. Also many thanks are due to B.
Metivier and the technical staff, notably M. Moreau. P. Maestrati and P. Lozouet, for their kind
assistance during my visits to the mnhn. Furthermore, I wish to express my gratitude to the following
persons, who provided access to collections, assisted me, and lent or donated study material: W.
Ponder and I. Loch (ams), B.A. Marshall (nmnz), A.B. Stephenson and B.W. Hayward (aim), F.E.
Wells and S.M. Slack-Smith (Western Australian Museum. Perth), Subba Rao and Surya Rao (ZSl),
T. Okutani (Tokyo University of Fisheries), I. Hayami (Geological Institute, University of Tokyo),
T. Habe (nsmt). A. Matsukuma (Department of Earth and Planetary Sciences, Kyushu University,
Fukuoka), Z. Wang (ioas), R.N. Kilburn (nmp), H.K.Mienis (Hebrew University. Jerusalem), E.
Gittenberger and J. Goud (rmnh). H.E. Coomans and R.G. Moolenbeek (zma). J. Van Goethem and
A. Lievrouw (kbin), C. Vaucher and Y. Finet (Museum d'Histoire Naturelle, Geneva), J. Knudsen
and T. Schiotte (zmc), S. Morris (formerly of the bmnh), and P.G. Oliver and A. Trew (National
Museum of Wales, Cardiff). For technical assistance I thank R.G. Moolenbeek (Figs 87-98) and L.
Van Der Laan (zma) (Figs 1-86, 99-122) for taking scanning electron micrographs and light
photographs.

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Vaught. K. C. 1989. A classification of the living Mollusca. Melbourne. USA. xii + 195 pp.

Verrill A. E„ 1885. - Third catalogue of Mollusca recently added to the fauna of the New England coast. Consisting mostly
and Science Sfr395-452h '10teS °n °lher SpeC'eS prevlously recordcd- Transactions of the Connecticut Academy of Arts

v errill A E 1897. - A study of the family Pectinidae. with a revision of the genera and subgenera. Transactions of the
Connecticut Academy oj Arts and Science, 10: 41-95.

WAGNEpeclinidae)l9sL^aTh5e3 0-3U): %^[opecten Dall‘ Burlsch & Rehder- l938- in the Indo-Pacific (Mollusca: Bivalvia:

^ KLLE%pon fof 970- ~5-7he 8'aSS SCalI°P Pr°Peamussium’ a livin8 relict of the past. American Malacological Union. Annual

WALLEp’,„T;RV| l9J8f ~ Morphology, morphoclines and a new classification of the Pteriomorphia (Mollusca: Bivalvia).
Philosophical Transactions of the Royal Society of London. (B). 284: 345-365.

WALLEfn .rh,Rp, '984„., 'Pl;c ctenohum of scallop shells: functional morphology and evolution of a key family-level character
in the Pectinacea (Mollusca: Bivalvia). Malacologta. 25 (1): 203-219.

Source : MNHN, Paris

BATHYAI. PECTINOIDEA FROM NEW CALEDONIA

73

Waller T. R.. 1991. Evolutionary relationship among commercial scallops (Mollusca: Bivalvia: Pectinidae). Pp. 1-73. In:
S.E. Shumway (ed.). Scallops: Biology, ecology and aquaculture. Amsterdam. xx+ 1095 pp.

Waller, T. R.. 1993. — The evolution of "Chlamys" (Mollusca: Bivalvia: Pectinidae) in the tropical western Atlantic and
eastern Pacific. American Malacological Bulletin, 10 (2): 195-249.

Waller, T. R. & Marincovich, L., 1992. — New species of Camptochlamys and Chlamys (Mollusca: Bivalvia: Pectinidae)
from near the Cretaceous/Tertiary boundary at Ocean Point, North Slope, Alaska. Journal of Paleontology, 66 (2):
215-227.

Wang, Z., 1980. — Study on Chinese species of the family Pectinidae (Mollusca, Bivalvia). 1. A new species of the subfamily
Propeamussiinae. Oceanologia et Limnologia Sinica, 11 (3): 259-262.

Wang. Z„ 1983. — Studies on Chinese species of the family Pectinidae. III. Chlamydinae (Genus Bractechlamys). Oceanologia
et Limnologia Sinica. 14 (6): 531-535.

Wang, Z.. 1984. — Studies on Chinese species of the family Pectinidae. VI. Subfamily Propeamussiinae. Oceanologia et
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Wilkes. J., 1810. Conchology. In: Encyclopaedia Londinensis; or. Universal Dictionary of Arts. Sciences, and Literature:
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Source : MNHN. Paris

DES CAMPAGNES MUSORSTOM. VOLUME 14

RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 14

RESULT/

Systematic revision of living
species of Meiocardia , Glossidae
and Glossocardia , Trapezidae (Bivalvia)

Akihiko MATSUKUMA

Department of Earth and Planetary Sciences
Faculty of Science, Kyushu University
Hakozaki, Fukuoka, Japan 812-81

&

Tadashige HABE

National Science Museum
3-23-1, Hyakunin-cho, Shinjuku-ku
Tokyo, Japan 169

ABSTRACT

Living species of Meiocardia, Glossidae, are reviewed on the basis of specimens stored in various museums and
institutions, including the MUSORSTOM collection of Museum national d’Histoire naturelle, Paris. Six species, one of them new,
are reported from the Indo-West Pacific. The type species, M. moltkiana (Gmelin, 1791), has been variously interpreted by
authors, so we redescribe it and give a new diagnosis of the genus. Other species of Meiocardia are: M. sanguineomaculala
(Dunker. 1882) (Philippines to Seychelles); M. vulgaris (Reeve. 1845) (China to Philippines); M. globosa sp. nov. (eastern
Indian Ocean to Taiwan and Philippines); M. samarangiae Bernard, Cai & Morton, 1 993 (Japan); and M. Iiawaiana Dali,
Bartsch & Rehder, 1938 (western Indian Ocean to Hawaii). Meiocardia lamarckii (Reeve. 1 845) is synonymised with M.
moltkiana. Meiocardia lamarckii of Japanese authors is not the same as M. lamarckii (Reeve), but is conspecific with M.
Iiawaiana. Meiocardia samarangiae Bernard. Cai & Morton, 1993 is a replacement name for Isocardia tetragona Adams &
Reeve, 1850 non Koch & Dunker, 1837.

The genus Glossocardia, Trapezidae, is redescribed on the basis of the type-species. Glossocardia obesa (Reeve. 1843)
(tropical West Pacific). It includes Glossocardia sloliczkana Prashad. 1932 (Philippines and New Caledonia) and the tropical
western Atlantic G. agassizii (Dali, 1886), which was originally assigned to Meiocardia. There are no records of living or fossil
species of Meiocardia from the western Atlantic or eastern Pacific.

Matsukuma. A. & Habe, T„ 1995.— Systematic revision of living species of Meiocardia. Glossidae and Glossocardia. Trapezidae (Bivalvia). In: P. Bouchet (cd.).
Resullats des Campagnes Musorstom, Volume 14. Mem. Mus. nam. Ilisl. not.. 167: 75-106. Paris isbn 2-85653-217-9.

Published 29" December 1995.

76

A. MATSUKUMA & T. HA BE

RESUME

Revision systematique des especes actuelles do Meiocardia (Glossidae) et Glossocardia (Trapezidae) (Mollusca: Bivalvia).

Les especes acluelles de Meiocardia (Glossidae) sont revisees sur la base du materiel conserve dans divers niusees et
tnstituts. y compris les collections constituees pendant les campagnes musorstom. Six especes, dont une nouvelle sont
recon n ues dans la region Indo-Ouest-Pacifique. L’espece-type, M. moltkiana (Gmelin. 1791 ). a ete diversement interpretee dans
la litterature; nous la redecrivons done, et donnons une diagnose revisee du genre. Les autres especes de Meiocardia sont ■ M
sangumeomacutaia (Dunker. 1882) (des Philippines aux Seychelles) ; M. vulgaris (Reeve. 1845) (de la Chine aux Philippines) •
M glohosa sp. nov (de I'Est de l’ocean Indien a Taiwan et aux Philippines) ; M . samarangiae Bernard, Cai & Morton 1997
(du Japon) : et M. hawaiana Dali. Bartsch & Rehder, 1938 (de l'ouest de l’ocean Indien a HawaT). Meiocardia lamarckii ( Reeve.
1845) est mis en synonymie de M moltkiana. mais M. lamarckii des auteurs japonais est un synonyme de M hawaiana
Meiocardia samarangiae Bernard, Cai & Morton, 1993 est un nom de remplaceme.it pour Isocardia tetraeona Adams & Reeve
1850. non Koch & Dunker. 1837.

Le genre Glossocardia (Trapezidae) est redecrit sur la base de son espece-type, G. ohesa (Reeve 1843) (du Pacifique
Ouest tropical). Ce genre comprend egalement Glossocardia stoliezkana Prashad. 1932 (des Philippines et de Nouvelle-
Caledome), et G agassizii (Dali, 1886), de l’Atlantique tropical americain, decrite a I’orieine coniine Meiocardia. On ne connait
aucun Meiocardia. actuel ou fossile, dans le Pacifique oriental ou I’Atlantique americain.

INTRODUCTION

The bivalve family Glossidae is characterized by trapezoidal to cordiform shells with a
cyprinoid hinge and conspicuous enrolled prosogyrous beaks. There are several living species
belonging in two genera, Glossus and Meiocardia. Almost all living species of Meiocardia are
restricted to the Indo-West Pacific. However, Dall (1886) described a living species from the tropical
western Atlantic under the name Meiocardia agassizii.

Several fossil species from the Miocene of the Netherlands (Janssen, 1984), Tertiary of
Piemonte, Italy (Sacco. 1900). and the Eocene of California (Squires & Advocate, 1986) and North
Carolina (Harris, 1919a, b) have been included in Meiocardia. The taxonomic position of these fossil
and living species from the Atlantic and the eastern Pacific is reassessed below.

Although several investigators, including Reeve (1845), Roemer ( 1 868), Buelow (1906), Lamy
(1920) and Prashad (1932), have reviewed the living species of Meiocardia. the taxonomy of
Glossidae is still confused. For example, some authors have considered Meiocardia moltkiana. the
type species of Meiocardia, to be strongly ribbed, with conspicuous nodules on the keel and a distinct
smuation just anterior to the keel (Reeve, 1845; Adams & Reeve, 1850; Buelow, 1906). whereas
Japanese authors have adopted the name for a species with an antero-posteriorly elongated, thick
shell with a sharp keel lacking conspicuous nodules (Okutani & Matsukuma, 1982), and other
authors have illustrated a shell with rounded postero-dorsal martiin as this species (Keen & Casey
1969; Abbott & Dance, 1982).

We review living species of Meiocardia, Glossidae, based on the musorstom collection in
Museum national d Histoire naturelle. Paris; National Science Museum. Tokyo; Institute of
Systematics and Population biology, Universiteit van Amsterdam; Natural History Museum
London; Museum of Comparative Zoology, Harvard University; Academy of Natural Sciences!

I hiladelphia; Los Angeles County Museum, and others.

ABBREVIATIONS AND TEXT CONVENTIONS

Repositories

ANSP

BMNH

IC

IMT

Academy of Natural Sciences. Philadelphia, Pennsylvania
The Natural History Museum, London
Mr Hiroshi Ito’s private collection, Tokyo
Institute of Malacology of Tokyo, Tanashi

Source . MNHN, Paris

REVISION OF MEIOCA RDIA AND GLOSSOCARDIA

77

kem : Kanagawa Prefectural Museum. Yokohama

lacm : Los Angeles County Museum of Natural History. Los Angeles. California

mcz : Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts

mnhn : Museum national d'Histoire naturelle, Paris

nsmt : Department of Zoology, National Science Museum, Tokyo

sbmnu : Santa Barbara Museum of Natural History. Santa Barbara, California

USNM : National Museum of Natural History, Washington, DC

yc : Mrs Yoshie Yamasaki's private collection, Nagoya

zma : Institute of Systematics and Population Biology (Zoologisch Museum), Universiteit van

Amsterdam, Amsterdam

Measurements (Fig. 1):

Cb

Convexity of both valves

Cs

Convexity of single valve

H

Shell height

L

Shell length

lv

left valve

rv

right valve

V

valve.

Fig. 1 . - Shell measurements of Meiocardia and Glosso-
cardia. Cb, convexity of both valves. Cs, convexity of
single valve H. shell height. L, shell length.

SYSTEMATIC ACCOUNT
Family Glossidae Gray, 1847

Diagnosis. Shell rounded to cordiform, strongly inequilateral, equivalve, with strongly
enrolled, prosogyrous beaks. Ventral margin not gaping, smooth. Ligament external, opisthodetic.
parivincular. Hinge with two lamellar cardinals and two laterals. Outer surface ornamented with
commarginal sculpture. Dimyarian, with subequal adductor muscles. Pallial line simple, without
sinus.

Remarks. The family Glossidae is apparently very close to the family Trapezidae, but
differs from the latter by having a glossy shell and by lacking radial sculpture and microscopic scales
on the outer surface.

Genus Meiocardia H. & A. Adams. 1857

Meiocardia H. & A. Adams. 1857: 461. Type species: Bucardia mohkiana, Chem[nitzJ [as listed by Ft. & A. Adams] -
Meiocardia ruolikiana Spengler [as cited by Stoliczka] = Chama mohkiana Gntelin, 1791, subsequently designated by
Stoliczka. 1870: 187.

78

A. MATSUKUMA & T. HABE

Diagnosis. — Shell small, usually less than 50 mm long, cordiform, strongly inflated, strongly
inequilateral, equivalve. Beaks prominent, prosogyrous, strongly incurved. Larval ligament anterior
to beaks. Outer surface glossy or polished, ornamented with commarginal sculpture. A strong
primary keel from beak to lower posterior end separating posterior slope; weak secondary keel from
beak to upper posterior end separating postero-dorsal margin. Cardinals (1, 3a, 3b) and laterals (lai,
lam, lpi, lpiii) in the right valve; cardinals (2a, 2b, 4b) and laterals (laii, lpii) in the left valve.
Posterior laterals (lpi, lpii, lpiii) finely striated or nodulated (Figs 1-2, 4-5).

Remarks. — All living species of the Glossidae are placed in either G/ossus Poli, 1795, or
Meiocardia. Meiocardia clearly differs from Glossus in the having a smaller, subquadrate shell with
very thin periostracum and a prominent primary keel separating the posterior slope. The genus
Glossus includes only one living species, G. humanus (Linnaeus, 1758) of the Mediterranean to
northeastern Atlantic, and is characterized by having a large, rounded shell with thick, reddish-brown
periostracum and in lacking a dorsal keel. Isocardia Lamarck, 1799 is an objective junior synonym
of Glossus Poli, 1795.

Meiocardia moltkiana (Gmelin, 1791)
Figs 3-4, 24-28

Chama Moltkiana Gmelin, 1791 [ex Chemnitz, non binominal]: 3303-3304.

Synonyms:

Isocardia lamarckii Reeve, 1845: Isocardia sp. 5. pi. I, fig, 5 [China].

Meiocardia nishimurai Kosuge & Kase, 1994: 28-29. pi. 11, figs 1-5 [Bonin (= Ogasawara) Islands, Japan], (Syn. nov.).
Other references:

■•Die Moltkische Chama" Spengler, 1783: 321-325. pi. 14, figs 1-4 [South Seas; invalid, not latinized]

Chama Moltkiana Chemnitz, 1784: 105-108. pi. 48. figs 484-487 [Ostindien; appeared in invalid work], - Bruguiere. 1797-
pl. 233, figs la-d [fig. only].

Isocardia moltkiana - Lamarck. 1819: 31 [India, China], — Borv nr St. Vincent 1827- 149

Isocardia lamarckii - Hanley, 1856: appendix, 370-371. app. pi. 18, f. 34 [China], - Buelow. 1906: 7-8, pi. 8, figs 6a-b

Lamy. 1911: 132 [Mauritius].

Isocardia (Meiocardia) moltkiana - Rof.mer, 1869: 8-9, pi. 1, figs 4-7 [Australia to Philippines]. - Prashad 1932- 149-150
[pars] [Indonesia]. '

Isocardia ( Meiocardia ) Moltkeana var. lamarcki - Lamy. 1920: 298-300 [Mauritius]

Istcardta (sic) (Meiocardia) moltkiana - Abrard. 1947: 39, pi. 3, fig. 11 [Miocene, New Hebrides]

Meiocardia moltkiana sanguinomaculata (sic) - Kira. 1959: 194. pi. 71, fig. 5 [Japan], - Shikama, 1964: 67, pi, 41 fig 13
[Mie Pref.. central Japan]. ’ y ’ 6

~ KUR?nDoA,el,4'I''?7l:, ™ (Japanese), 440 (English), pi. 97, figs 11-12 [Sagami Bay, central Japan],
r<-ic°KUT1 & Matsukuma, 1982: 175-176, pi. 10. fig. 5 [Izu Peninsula, central Japan], Matsukuma, 1986: 324-325 (f )
[Okinawa]. Kosuge & Kase. 1994: pi. 10. fig. 10 [Spengler's type material]

Meiocardia letragona - Drivas & Jay. 1988: 142, pi. 56, fig. 5 [Reunion & Mauritius],

Meiocardia lamarckii - Kosuge & Kase. 1994: pi. 10. fig. 12 [one of the syntypes].

Tvpe Material. Die Moltkische Chama": Spengler’s illustrated specimens are stored in
Zoologisk Museum, Copenhagen (not examined). — Meiocardia lamarckii: syntypes. Reeve
collection, bmnh. — Meiocardia nishimurai: holotype, a conjoined specimen imt-94-1.

Type Locality. — “Die Moltkische Chama": South seas [tropical Indo-West Pacific].
Meiocardia lamarckii: China. — Meiocardia nishimurai: Ogasawara Islands, Japan.

Material Examined. — Pleistocene: Japan. Ryukyu Limestone, Kikaijima Is., Kagoshima
Pref. (nsmt-M o69745).

Recent. Japan. Kameki-sho reef, Sagami Bay (nsmt-Mo43435). — 1.5 km SW of Jogashima, Sagami
?^yorJ°’-75 m (nsmt‘Mo69739)- — Futami Bay, Chichijima Island, Bonin Islands, 27°04.5' N,
142 07.1 E, 115 m (nsmt-Mo59790). — Bonin Islands, holotype of Meiocardia nishimurai (imt-

Source : MNHN, Paris

REVISION OF MEIOCA RDIA AND GLOSSOCARDIA

79

2-6. Dentition and gross anatomy of Meiocardia. — 2a-b. Meiocardia samarangiae. - 3a-b & 4a-b. Meiocardia
moltkiana. 5a-b. Meiocardia sanguineomacutata. — 6a-b. Meiocardia hawauma. 1, 3a-3b. cardinals in the nght valve.
2a-2b & 4b, cardinals in the left valve, lai & law: Anterior laterals in the nght valve, laii: Anterior lateral in the left
valve, lpi & lpiii: Posterior laterals in the right valve, lph: Posterior lateral in the left valve, aa: Anterior adductor,
al: Adult ligament, es: Exhalant siphon, f: Foot, id: Inner demibranch. is: Inhalant siphon, od. Outer demibrantn.
pa: Posterior adductor, s: Siphons.

Source

80

A. MATSUKUMA & T. HABE

94-1). — Aichi Pref. (nsmt-Mo69740, Kawamura Coll.). - Off Kirimezaki, Wakayama Pref.
(nsmt-Mo69741). — Wakayama Pref.. 54 m (ansp 274370). - Tokushima Pref. (nsmt-Mo69742,
Kawamura Coll.). — Okinoshima Island. Kochi Pref. (nsmt-Mo42307). - Kochi Pref. (nsmt-

Mo69743, Kawamura Coll.). — Nishinohama, Tomioka, Kumamoto Pref. (nsmt-Mo43455). _ 20-

40 m, Tomioka Bay. Kumamoto Pref. (lacm 82-23). — Tomioka Bay (nsmt-Mo43436). Okinawa
Pref. (nsmt-Mo63233). 60 m. I km WNW of Onna-son. Okinawa Pref. (lacm 79-76, 78-101).

China, (zma). — (ansp 54195, 189703, 217140). (bmnh reg. no. 1907.12.30.505; Reeve coll.;
Cuming coll.; Winckworth coll.; Trochman coll.).

Philippines. (nsmt-Mo69744. Kawamura Coll.). - Luzon Island (bmnh)
musorstom 1: stn 57. 13°53' N. 120° 13.5' E, N of Lubang Is., 96-107 m.

MUSORSTOM 2: stn CP 21. 1 4°00' N. 120° 17' E. N of Lubang Is.. 191-192 m (all mnhn).

Indonesia. Off Maroepi Is.. Ambai Group. Japen Is., Geelvink Bay, 36-45 m. mud and shells (ansp
279722). 1.6 km NE of Roemwakon. Aoeri Is.. Geelvink Bay, 36-45 m (ansp 205650. 275802). —

3.6 km N of Matas. Aoeri Is., Geelvink Bay, 32-36 m (ansp 208922).

Coral Sea. chalcal 1: stn CP 7, 19° 18' S. 158°36' E. Plateau Chesterfield-Bellona, 65-68 m (mnhn).

New Caledonia, lagon: stn 836. Secteur de Poindimie, 20°46' S, 165°16'E. 57 m. a conjoined
specimen. — Stn 928, Secteur de Koumac, 20°45' S. 164°23' E, 7-10 m. a conjoined specimen — Stn
983. Secteur de Poum. 20°23' S, 163°57' E, 38-68 m (mnhn).

"Vauhan" 1978-79: stn 40. 22°30' S, 166°24' E. 250-350 m (mniin)
smib 5: stn DW 81, 22°38' S, 167°35' E. 110 m (mnhn).

Thailand. Phuket (nsmt-Mo43437).

0.6

0.5

0.4

Cs/H

o

o

o

CO

o

o

o

o

oo
o o

o o o

QD

8o° &8o °

• • o°° cP

O ## 8

o

o

••

o

n9>

°0§o

o

o o

►o

o

o

L (mm)

o

10

20

30

FlG- 7 -ln(1 China |.Sr°Win^ ’,r*lationshiP between L. and Cs/H in Meiocardia moIlkUma. Dots: Specimens from Japan

and China. N - 25, r - 0.325: regression equation Cs/H = 0.00 129L + 0.451. Circles: Specimens from Reunion N
- 55. r - -0.258; regression equation Cs/H = -0.000818L + 0 500

Source : MNHN, Paris

REVISION OF MEIOCARDIA AND GLOSSOCARDIA

81

Cs/H

0.6

O

O

1.0 1.2 1.4

Fig. 8. Scatter diagram showing relationship
between Cs/H and L H in Meiocardia moltkiana.
Dots: Specimens from Japan and China. N = 25;
r = -0.210; regression equation Cs/H =

-0.0874L H + 0.587. Circles: Specimens from
Reunion. N = 55; r = 0.207; regression equation
Cs/H = 0.132L/H + 0.316.

Andaman Sea. Port Blair, Andaman Island (bmnh). 19.2 km NW of Port Blair. 1 1"49' N, 92 ’53' E,
90 m. shelly sand (ansp 292189). Maldives, NW of Maduwari Is., Fadiffolu Atoll, ca. 5°18'N,
73°29' E. 45-63 m (ansp 325451).

Seychelles. Saya de Mala Bank, 100 m (bmnh).

Malagasy. Tulear (mnhn); 7.2 km W of Nosy N'Tangan. SW Nossi Be (ansp 260078).

Reunion, md 32 Reunion; stn DC 41, 2 1 °2 1 ' S, 55°27' E, 75 in. — Stn CP 42, 21°21' S. 55' 27 E, 74-77
m. Stn DC 43, 21°2F S, 55°27' E, 73-77 m. — Stn DR 47, 21°23' S, 55°37' E, 205-215 m. Stn
DC 54, 21°06' S, 55°13' E, 80-83 m. — Stn CP 55, 21°05' S. 55°13' E. 97-1 10 m. - Stn DC 56,
21°05' S, 55°12' E, 170-225 m. — Stn DC 85, 21°00' S. 55°15' E. 58-70 m. Stn DC 124. 20°52' S.
55°37' E, 40 m. Stn DC 126. 20°52' S, 55°38' E, 1 10 m. Stn CP 129. 20°51' S. 55°36' E, 290-300
m. — Stn DC 176, 21°02' S. 55°11' E, 165-195 m (all mnhn).

Distribution. Miocene: New Hebrides [Vanuatu] (Abrard, 1947). Pleistocene: Japan.
Recent: Western Japan. China, Philippines, Coral Sea, New Caledonia, Thailand, Seychelles,
Malagasy, Reunion, Mauritius.

This species lives in coarse sediments in shallow water, approximately 7-70 m, but empty shells

have been collected from deeper than 300 m.

Description. Shell small, thick, subcircular, strongly
inequilateral, equivalve. Mean of L H in Japanese and
Chinese specimens 1.257 (N = 25, SD = 0.070). not signi¬
ficantly correlated with L (r = 0.158, a = 0.05); mean of L H
in Reunion 1.297 (N = 55. SD = 0.043). not significantly
correlated with L (r = -0.0418. a = 0.05). Mean of Cs H in
Japanese and Chinese specimens 0.478 (N = 25. SD =
0.029), not significantly correlated with L (r = 0.325. a =
0.05): mean of Cs/H in Reunion 0.487 (N = 55, SD =
0.027). not significantly correlated with L (r = -0.258. a =
0.05) (Fig. 7); L/H not significantly correlated with Cs/H in
both areas (r = -0.210 in Japan and China, r = 0.207 in
Reunion, a = 0.05) (Fig. 8).

Beak prosogyrate, spirally twisted. Strong keel from beak
to postero-ventral margin. Postero-ventral margin pointed,
protruding, with a sinuation just anterior to smooth keel.
Outer surface yellowish white, ornamented with regularly
spaced concentric ribs on anterior and central slopes. Poste¬
rior slope widely concave, smooth except fine growth striae.
Inner surface creamy or milky white, shining. Anterior
adductor scar semicircular: posterior adductor scar subcir¬
cular; both scars nearly equal in size.

Measurements. See Appendix. Table 1.

Remarks. - Spengler (1783) fully described the species, but did not give it a Latin name.
Although the name, "die Moltkische Chama" of Spengler (1783). was latinized by Chemnitz (1784).

82

A. MATSUKUMA & T. HABE

the work of Martini & Chemnitz (1769-1795) was placed by the Commission of Zoological

spedeT atUfe °n th£ St °f InValld W°rkS’ S° Gmel'n °791) mUSt be accePted as the author of the

Although Prashad (1932) cited figures given by Reeve (1845- nl 1 n AnA..c o D_

iltemS by Rffiwrmsf dT* (W°* 37'* P'' «• 7> » Meiocardia llMana, the shells

rStJ u u ^ ( 8f5) dnd 0thers rePresent Meiocardia sanguineomaculata, which has a smaller
AitKdedkSi? WItPh ^Uch coarser nbs and has conspicuous nodules on the primary keel
Although Keen & Casey (1969) assigned the specimen figured by Chenu 0862: 113 fig 533) to

haS a" antt;ro-P°steriorly elongated shell with a smooth and gently rounded
postero-dorsal area. The outer surface of shell is ornamented with fine concentric ribs and lacks

bS* LAMY (1920) COrreCt,y POinted 0Ut that this "Ot M. moltkiana.

Juveniles of M moltkiana, which have thin, quadrate shells, are similar to Meiocardia
awaian“' - but - they differ from the latter in having stronger ribs and a straight cardinal (2a).

Shells from Bonin Islands (nsmt-Mo59790 and imt-94-1) have a somewhat distinct sulcation
n front of a strong primary keel and reddish brown brotches. Although Kosuge & Kasf (1994)
proposed for ,t the name Meiocardia nishimurai, Holocene shells from KikaijVm Is and Kagoshima

Reunion shows continuous variation between the two forms (Figs 25, 26a-d).

Meiocardia sanguineomaculata (Dunker, 1882)

Figs 5, 22-23

Isocarctia moltkiana var. sanguineomaculata Dunker, 1882: 213 ("Korea"].

Other references:

8Pfig. t '' ^ ^ ~ Adams & Rbhvh. ,850: 77, pl. 22,

Isocardta ( Meiocardia ) moltkeana var. sanguineomaculata - Lamy, 1920: 298-299.

*0 ** hiustrateb by

“Coreaf’r™ iS"!; 7tvUh°,Ugh A°AMS REEVE <1850) ”oted the locality of their shell is

m, sand and coral (usnm 431 190) — Stn 5139 off Inin inin ic tr ' , J Is” 34

— Stn 5146 off Sulade Is «5„i„ Ar(.u , ’ Jol°' Jol° Is’ 36 m’ coarse sand (usnm 294513).

5148 off 9ir„n to, *’ , ArchlPelaS°- m, coarse sand with shells (USNM 292393) — Stn

Tnnnwf is ’ Ai;ch,pelaS0' 31 m, coarse sand (usnm 235929). - Stn 5149 W of Lanac
Tapul Is., 18 m, coarse sand with shells (usnm 235950). _ Stn 5151 off sin.n u- t. • i LaPac’

Archipelago, 43 m, coarse sand with shells (usnm 292027) — Stn 5253 Pakinntan h’ Su U

50 m (usnm 237240) ' ’ Mn ~>433, ‘ dkiputan Strait, Mindanao,

Andaman Sea. Port Blair, Andaman Is. (bmnh 1989178)

?6%C6 FeS55RmESa2; Stn 55°44' E> 43 m, a conjoined specimen. - Stn 16 05°36' S

55°25'F L /f J°in? specimen. - Stn 18, 05°45' S, 56°35' E, 50 m. - Stn 24' 05°09' S

MNHN) ’ JOin Specimen' ~ Stn 3°' °4°42' s’ 54'>24' E’ 47 a conjoined specimen (all

Source : MNHN. Paris

REVISION OF MEIOCARDIA AND GLOSSOCARDIA

83

C n,„r Ornirn 9°41'S 5 1°03' E, 16 Dec. 1964 (USNM 718999).

Saya de Malha Bank, Indian Ocean, 99 m, 2 conjoined specimens (bmnh 1910.8.31.688-689, J.S.
Gardiner Coll.)

Distribution. — Recent: Philippines, Andaman Islands, Seychelles. Meiocardia sanguineo-
maculata is a species of coarse sediments in 10 to 100 m.

Description. — Shell small, thick and solid, subquadrate
strongly inhaled, strongly inequilateral, equivalve. Mean ol
l II r i45 (N = 28, SD = 0.073), not significantly correlated
with L (r = 0.159, a = 0.05) Mean of Cs/H 0.515 (N - 28.
SD = 0.034), not significantly correlated with L (r U.iro,
a = 0 05) (Fig. 9). L/H significantly correlated with Cs/H
(r = 0 519, a = 0.05) (Fig. 10). Outer surface ornamented
with strong concentric ribs. The primary keel with prominent

tubercles, wide, rounded. Postero-ventral margin with pro¬
minent sulcation just in front of the roundly protruded
primary keel. Outer coloration whitish yellow with reddish
brown triangular blotches. Beaks spirally enrolled. Inner
surface creamy white, pale brown posteriorly. Anterior
adductor scar elongated oval to semicircular; posterior
adductor scar subcircular; both scars nearly equal in size.

Measurements. See Appendix, Table 2.

Remarks — M. sanguineomaculata has a thick, globular shell with Cs/H often exceeding 3.5
and L/H around 1.1. The shell form suggests that this species is most probably an inactive shallow
burrower. This species differs from Meiocardia moltkiana by having an antero-postenody shortened
rounded shell with stronger concentric ribs and a distinct sinuation just anterior to the keel which has
prominent tubercles.

Meiocardia vulgaris (Reeve, 1845)
Figs 12, 31-32

Isocardia vulgaris Reeve, 1845: Isocardia sp. 2, pi. 1, figs 2a-b [China).

Synonym:

Meiocardia delicata Kosuge & Kase, 1994: 29-30. pi. 11, figs 6-8, pi. 12. figs 1-3 [Okinawa, Japan). (Syn. nov.).

Other references: f

Isocardia moltkiana - Sowerby, 1852: 82. pi. 6, fig- 126. — 9^ oUj [China)3’— 8PraSHAd ^ 932M 50- 1 sTp ndonesia).
Isocardia ( Meiocardia ) vulgaris - Roemer, 1868: 9-10, pi 1, figs 9-10 [China). i-rashad. iyjz. i

Isocardia vulgaris - Buelow. 1906: 37-38, pi. 8, fig. 5 [China]. . p , r H inHial

Isocardia ( Meiocardia ) moltkeana var. vulgaris - Lamy, 1920: 297-298 [ i > • 1 994- p| 10 fig. II [one of

Meiocardia vulgaris - Sh.kama, 1964: 67, pi. 41, fig. 14 [East China Sea). - Kosuge & Kase, 1994. pi. ng 1

G$^uTlSeioSrdia) moltkiana - Keen & Casey. 1969: N657. fig. E134-7 [= Chenu, 1862, fig. 533).

Type Material. — Isocardia vulgaris : 5 syntypes in bmnh, without registration number.
Meiocardia delicata: holotype, a conjoined specimen imt-94-2.

Type locality. — Isocardia vulgaris : China. — Meiocardia delicata : Okinawa, Japan.

Material examined. - Recent: Japan. Holotype of Meiocardia delicata (MT»

China. 5 syntypes oUsocardia vulgaris (bmnh). - R. Winckworth Coll. (bmnh). — L. A. I Lee> /e Co ^
(BMNH). - (ZMA). - (ANSP 54063, 54064. 138495). - (usnm 17495, 32049 and 759 12). (u ac

13513, 48064, A1463, A5076). — Hong Kong (usnm 47939). — Taiwan (kpm 761-2357). Ch

(usnm 186123). , ... . „ , _ , q „ H

Philippines. “Philippines" (nsmt-Mo69747, Kawamura Coll.). — ‘™1PP'!ie? ’ 5Q'm (I acm

Cuming coll. (bmnh). — Nazasa Bay, Zambales Prov., W Luzon, 14 48.9 N, f ~ Turtle

60-40). — NE Villa Carmen, Bataan Prov., W Luzon, 2-9 m (lacm 89954). W of Boan .,

Is, SW Sulu Sea, 29-32 m (lacm 89904). ,, ,

*■ Albatross ” stn 5097, S of Corregidor Is., Manila Bay, 54 m, gray mud, sand and shells (usnm
283971). — Stn 5107, off Corregidor Is., Manila Bay, 50 m, gray mud (usnm 235242). Mn a ,

84

A. MATSUKUMA & T. HABE

Cs/H

0.5

oo

o

®

CO

CO

%

A

O

O

O

O

O

10

20

30

'J'J 40 50

D„B, _

regress, on equation Cs/H = 6.00 1 03 L - 0.505. Triangles N = 28; r = 0.155;

?L“Tnaw!,a»; f"' °°T Tt (USNM 302604)- - St" ^ 56, off

off Tinakta Is., Tawitewi, SW Si, ^ArcSaao « m fi 'n** (USNM 2921 l7>- ~ Stn 5,57.
Tawitawi. SW Sulu Archioehpn 2? m , 8 ' \2 m" bne sand <USNM 2361 10). - Stn 5164, off SE

Panay, 47 m. fine SE ofV t" '° ' NE

58 m, green sand (usnm 293088) - Stn 5?m M -,r rt , ' 1^“’ ? _ of Bantayan Is.. NW of Cebu,
248221). - Stn 5220. N of Mannduaue 9(i m f “ " 90 m- muddy “ad (usnm

Magabao Is., E. Mindanao, 79 m Xmud <US™,2483767 - Stn 5235, off

Palawan. 83 m, sandy mud (usnm 229~’~’9) - Stn 5335 I ' 8t~ 5335, , ltl;ipacan Str., N of

Palawan. 83 m, sandy mud Jusnm 297^ - s " Ills' pZTp ' N of

coarse sand and mud (usnm 237803 297338) Stn 5477 ff' r u aSS’ ofF Preservatory Is., 77 m.
(usnm 238563). - Stn 5478. off Talbac Pt I Ztf !n? V ^ Pt'’ Leyte’ 86 m- 8™Y mud
Pt., Leyte, 112 m, fine sand (usnm ">3858 1 ) E end of Cn ^ SNNI -dI13-2). Stn 5480, off Tacbac
247061). SW of San Nicholas Shoals Lmht M f„i? ^orrfg,dor £ Manila Bay, 11-18 m (ansp
SE Luzon. 54 m (lacm 89903) _ Ragav Gulf SF I * ^ ^f6662’ 246705,‘ ~ RaSay Gulf.
Hulagaan Is., Occidental Mindoro (aSp ?<>207n T (NSMT-M°69746’ Kawamura Coll.,. -
Maqueda Bay, Samar Is. (mcz) - N end of C>h I Rorongai1: Samar Prov.(LACM 89907).
60557). ' 1 N end of Cebu ,s- (ansp 245942). - Cebu (nsmt-Mo54938,

musorstom 3: stn DR 140 11°43'N m°34'c „ , «. ,,

Malaysia' * 7* « Panay is~^44 -mShn) ^ ^ 93‘" m (MNHN>' St" CP 14R

Indonesia, corindon: stnCH 205° LWS ?q m’ mud (USNM 297635).

(mnhn). ^ 1 08 S’ 1 17 19 E, Makassar Strait, 49 m, a conjoined specimen

Welkmost Bay, Bantam, Java (usnm 761 i 371 __ d

1 34°04' E. W of Babi Is., Wokam NE Aru 54 63 gdnler- Bantam- Java (usnm 261 138). 3°54' S.

, U-P.CU11, INC Aru, 54-63 m, mud (usnm 747420).

Source : MNHN, Paris

REVISION OF MEIOCA RDIA AND CLOSSOCARDIA

85

Australia, (usnm 95524, Jeffrey coll.); 14.4 km SE of Double Is., Tin Can Bay, Queensland (usnm

681655") _ Bundaberg, Queensland, 45 m (lacm 30008).

cpa 08°29' N 97"59' E. 40 km nnw of Phuket. Thailand. 42 m, sandy mud (ansp _917_9).

tTkm NE or Lighthouse Is., Phuket. 144 m (ANSP 286326). - 13=00' N, 97=41- E. 56 km W of
Tavov Is Andaman Is., 68 m, mud with shells (ansp 292935). - 13=28' N, 97=19' E, 91.2 km NW
If Tavoy'lk. 39 m. shelly sand (ansp 293036). - 12=03' N. 92=57' E, 40 km NNW of Port Blatr,
Diligent Strait, Andaman Is. 38 m, shelly sand (ansp 292612).

o ® f Bengal 1 5°08' N 94°04' E, 80 km SW of Irrawaddy River. Prepans North Channel, Burma,
53 m grav mud (ansp 293697). 19^32' N, 921'52' E, 27.2 km SSE of Akyab, Burma. 55 m. muddy

s ind (ansp 294080). - 17°35' N, 83°25' E, 28.8 km SW of Vizagapatnam, NE India, 79 m shelly
sand (ansp 294702). - 17°35' N, 83°25' E, 16 km SE of Vizagapatnam, NE India, 58 m, shelly sand
(ansp 294183).

Oman 25°50' N, 58°08' E, Oman Gulf (usnm 716879).

Zanzibar, 3.2 km W of Chango Is., 27 m, shelly sand (ansp 214511).

Malagasy. 96 km NE of Cape St. Andre (usnm 719126).

Distribution. - Recent; China, Taiwan. Philippines, Malaysia, Indonesia Queensland,
Andaman Is., Burma, NE India, Oman, Zanzibar; Malagasy. This species lives in mud in to 50 m,
and empty shells have been collected in approximately 110 m on a mud bottom with shells.

Description. Shell large for genus, occasionally attai¬
ning 45 mm in length, yellowish white. L/H significantly
correlated with L (N = 19, r = 0.900, a = 0.05). Cs/H
significantly correlated with L. (N = 19. r = -0.541, a
0 05) (Fig. 9). L/H significantly correlated with Cs/H (N -
19, r = -0.449, a = 0.05) (Fig. 15). Keel running from beak
to postero-ventral corner not prominent. Postero-dorsal area

smooth. Antero-ventral area ornamented with weak concen¬
tric ribs. Postero-dorsal margin rounded. Inner surface
creamy white, umbonally yellowish, shining. Anterior adduc¬
tor scar semicircular; posterior adductor scar subcircular;
both scars nearly equal in size.

Measurements. See Appendix. Table 3.

0.6 .

Cs/H

o

o

o

9-5 - ° oo CD

0.4

°8

o

• •

o

L/H

Fig 10 - Scatter diagram showing relationship between L/H
and Cs H in Meiocardia. Dots: Meiocardia hawaiana. N -
17- r = 0.557; regression equation Cs/H - 0.48/l/m +
0.113. Circles: Meiocardia sanguineomaculata. N - -8; r -
0.519; regression equation Cs/H - 0.245L/H + 0—34.

1.0

1.2

1.4

Remarks. This is the largest species in the genus and differs from M. moltkiana in ha g
a gently protruded postero-dorsal margin, weaker commarginal ribs, and a smooth primary iceei
Although Kosuge & Kase (1994) noted that Meiocardia dehcata ddters from M. vugans in
having weaker comarginal sculptures and less globular shells, a gently arched postero- orsa margi
with delicate ornamentations is a characteristic of M. vulgaris.

86

A. MATSUKUMA & T. HABE

" 1 4- Dentition of Meiocardia
1 la-b, Meiocardia globosa sp. nov. -
stoliczkana.

; S '

* • . •* v \ \

LAII I A

3 | \

( f jLAJ

: ^ A
■ 1 \ ■

y j

\

... AA'.A

Figs I

Source : MNHN, Paris

REVISION OF MEIOCARDIA AND GLOSSOCARDIA

87

0.6,-

Cs/H

0.5

0.-1

A O

O o
Cfa o

1.0

1.2

l m

1.6

Fig. 15. — Scatter diagram showing relationship between
L/H and Cs/H in Meiocardia. Dots: Meiocardia samaran-
giae. N = 8; r = 0.701; regression equation Cs/H =
0.00380L/H + 0.347. Circles: Meiocardia vulgaris.
N = 19; r = -0.541; regression equation Cs/H =
-0.000934L./H + 0.514. Triangles: Meiocardia globosa sp.
nov. N = 3.

Meiocardia globosa sp. nov.

Figs 11, 30

Type material. — Holotype: a conjoined specimen, nsmt-Mo69749. Paratypes: 1 conjoined
specimen nsmt-Mo69750, 1 left valve nsmt-Mo69751; usnm 236796, 283380a.

Type locality. — Pescadores, Taiwan, coll, by K. Nakayasu.

Material examined. — Holotype. — “ Albatross ”, stn 5181, E of Gigante Is., Philippines,
47 m, mud and fine sand. — “ Albatross ", stn 5164, SE of Tawitawi Is., Sulu Archipelago, Philippines,
32 m. — Makham Bay, Phuket, Thailand.

Distribution. — Recent: Taiwan, Philippines, Indian Ocean coast of Thailand. This species
lives on muddy sand bottoms in 30 to 50 m.

Description. — Shell rounded, more or less thin, strongly
inequilateral, equivalve with strongly enrolled beaks. Postero-
dorsal margin gently rounded. Outer surface ornamented
with regularly spaced, strong, concentric ribs. Outer colora¬
tion yellowish brown. Primary keel sharp, smooth, without

nodules, separating smooth postero-dorsal area. Postero-
ventral margin with prominent sulcation just in front ol the
pointed primary keel. Inner surface milky white, umbonally
yellowish. Anterior adductor scar semicircular; posterior
adductor scar subcircular; both scars nearly equal in size.

Measurements (mm):

V

L

nsmt-Mo69749 (holotype)

rv + lv

24.8

nsmt-Mo69750 (paratype)

rv + lv

12.1

nsmt-Mo69751 (paratype)

lv

25.1

H L/H Cb _ Cs _ Cs/H

2L8 1.14 22.1 - (0.507)*

11 7 1.03 114 - (0.487)

23.7 1.06 - 12-7 0.536

•Numerical in parentheses show Cb/2H, instead of Cs/H.

Remarks. — Meiocardia globosa is closest to M. vulgaris which is sympatric Meiocaidia
globosa has a more rounded, antero-posteriorly shorter shell, with stronger commarginal ribs an a
more prominent keel (Figs 30a-d).

A. MATSUKUMA & T. HABE

Meiocardia samarangiae Bernard. Cai & Morton, 1993
Figs 2, 16, 29

Meiocardia samarangiae Bernard, Cai & Morton. 1993; 67.

Synonym:

Isocardia tetragona Adams & Reeve, 1850: 76 pi fie I , ,

, P ’ s' 1 |Jdpan- Preoccupied by /. tetragona Koch & Dunker. 18371.

Other references:

MS && & isn -W* i"" «* 243'245 '»■

1874. “ Supp0sed Syntype: a “"J0™'1 speam® sucked on board, bmnh
Type locality. — Japan.

Pref. (nsmt-Mo59576),NI D R vuk vu' ° Limes'! o n i! " Ka'm i k ' 7^ Moeshima Is” Kagoshima

(nsmt-Mo54638). " ' ’ Kamikatetsu, Kikaijima Is., Kagoshima Pref.

Kan^awa^^ef°^(^SMT-^M!M44),en^SU‘‘s'aRarnia^RWa”^>I7^ 43442). - Hayama.

Kawamura Coll.). - Amadaiba Sagan" Bay (nsLm^T545 Off'' fhV9’ ^
(nsmt-Mo43450). — “Wakayama Pref” a a. * 1-.V \ ,r 7~ 0,f Issh,kl’ Aichi Pref.

Wakayama I^re?^(NSM^Mo43454^n— Wakayama ^*^'I^,0^^^an“dTatsuga^ma"

Fukui Pref. (nsmt-Mo43453 69755) "S’ u ’ •• ^ ’ Kochi I ref. - Kuriya, Echizen-cho,

et •, v 09/00). - Soyo-Maru stn 483 34°44'5()" N nn°47'in" n t u-

- 0 m, sand (nsmt-Mo43440). - ‘'Soyo-Maru” stn 46s’, 34‘>25'40” N P9"47'iS" F
otrait, 112 m. mud sand (nsmt-Mo4244Xi <*cv, u .. „ , ’ ,w'' ^ k, Isushima

Satamisaki, 181 m. 4 ’ 3£13'i0'N- B. off

130-26W E. off Kaimondake. 192 m, Sander Tfrw T V' 3n0’15' N-

- Miyakojiraa Island, Okinawa Pref (nsmt-M™ 1917) “ Chma Sea «™T-Mo43449).

190 m.

D,s™bu™n. - Japan, China. This species hves in mnddy sand in approximately

110 to

a Description. Shell antero-posteriorly elongate subaua

TST^^iT^: oofs?* »oT

correlated with L (r = 0 701 a = 0 05) I H n, t Slgn!pcan, y
correlated with cS/H ,N =’9°, r

Measurements (mm):

bw, prosogyrate. Primary keel running from beak to pointed

face wahTr^ imargm- Secondary keel obscure. Outer sur¬
face with irregular commarginal ribs except on nearly smooth
umbonal area. Hinge teeth thin: I long, parallel^ dorsal
^HH8'?- LPI" Smal!' Inner surlace creamy white Anterior

chrhnthCtr SemiC'rfular; Pos[erior adductor scar subcir¬
cular, both scars nearly equal in size.

V

L

H

L/H

Cb

nsmt-Mo 69752 (1)

Mo 69752 (2)

Mo 69752 (3)

Mo 69752 (4)
nsmt-Mo 69753

rv + |v

rv + Iv
rv + lv
rv + Iv
rv + lv

29.9

29.7

25.1

20.0

28.0

20.1

21.1

17.0

15.4

18. 1

1.49

1.41

1.48

1.30

1.55

18.2

20.3

15.0

(14.1)

16.1

Cs

Cs/H

(0.453)*

(0.481)

(0.441)

(0.445)

Source : MNHN, Paris

REVISION OF MEIOCA RDIA AND GLOSSOCARDIA

89

nsmt-Mo 69754 (I)
Mo 69754 (2)
nsmt-Mo 69755
nsmt (Tosa)

V

L

H

rv + Iv

31.1

21.7

rv + Iv

29.5

20.9

rv + lv

27.7

18.9

rv + Iv

23.0

16.4

./»

Cb

Cs

Cs/H

.43

20.4

-

(0.470)

.41

18.7

-

(0.447)

.47

17.0

-

(0.450)

.40

14.4

-

(0.439)

*Numericals in parentheses show Cb/2H, instead of Cs/H.

Remarks - Isocardia tetragona Adams & Reeve, 1850 is a primary junior homonym of
Isocardia tetragona Koch & Dunker, 1837. Meiocardia samarangiae Bernard. Cat & Morton, 1993 is
a replacement for Adams & Reeve’s name. The scatter diagram of L/H — Cs/H shows that M.
samarangiae clearly differs from any living species of Meiocardia (Figs 8, 10. 15).

Although Bernard et al. (1993) recorded M. samarangiae from the Philippines to Japan, we
think the species is restricted to seas around Japan. Prashad (1932) reported a Meiocardia species
from Sulu Sea and Indonesia under the name M. tetragona. These specimens are an antero-postenorly
elongated form of Meiocardia hawaiana , because they have cardinals (1 and 2a) that are oblique to

the dorsal margin (Fig. 35b). . . r , .

Although Habe (1977) considered Meiocardia hawaiana to be a junior synonym ol M.
tetragona. the Hawaiian species differs from M. tetragona in having thinner, waxy white shells with

oblique cardinals (1 and 2a). . .

The supposed life position of M. samarangiae which is suggested by attached animals,

including bryozoans and sponges with siliceous spicules, is shown in Fig. 16. A specimen Irom Tosa
Bay, Kochi Prefecture, has a muricid borehole at the ventral margin.

Fig. 16. Supposed living position of
Meiocardia samarangiae.

Meiocardia hawaiana Dali. Bartsch & Rehder, 1938
Figs 6, 33-37

Meiocardia hawaiana Dali, Bartsch & Rehder. 1938: 121-122, pi. 34, figs 17-18 [Hawaii].

Synonym'?:

Bucardia ( Meiocardia ) cumingii Adams, 1864: 142.

Ollier references:

Isocardia (Meiocardia) tetragona - Prashad, 1932: 151-152.pl. 5. figs 3-4 (Sulu Archipelago]. ,, ,

Meiocardia tamarckii - Habe, 1951: 1 17 [western Japan]; 1977: 236 [western Japan], Kira. 1959: 131. pi. . — tig. pap 1-

Matsukuma, 1986: 324-325(f.) [Kochi Pref.. western Japan], _ .... . „ m

Meiocardia hawaiana - Weaver, 1963: 2, figs 1-2 [Oahu. Hawaii]. Kay, 1979: 568, figs 184 A-B [Hawaii].

90

A. MATSUKUMA & T. HABE

Type material. Meiocardia hawaiana : holotype, a right valve, usnm 173048 (Boss
Rosewater & Ruhoff, 1968). Paratypes, usnm 338242, 338243, 346246, 428546.

Type locality. — “ Albatross ”, stn 4133, near Kauai, Hawaii, 74-562 m.

Hawaii. 21°02.TN, 156°47.25'W, off Oahu, 220

m, a

Material examined. — Recent:
conjoined specimen (usnm 807654).

Japan. Off Isshiki, Aichi Pref. (nsmt-Mo43438). — Kumanonada Sea (yc). — Off Wakayama (ic)
— Kochi Pref. (nsmt-Mo63224, Kawamura Coll.). - Tosa Bay off Kochi Pref. (nsmt-Mo43439!
tvawamura Coll.). Off Amitori, Iriomote Is., Okinawa Pref. (nsmt-Mo69757, 69758).

ol|,l’!PPI?rec- Albatross" stn 5173’ off Jol°’ Jol° Is- 335 m, shelly coarse sand (usnm 236529). — Stn
^-12, off Sibugay Is., east of Masbate, 194 m, gray sand and mud (usnm 292687) - Stn 5268 off

Matocot Point, west of Luzon, 306 m (usnm 295937)

— OM 1: stn 19> 13°56'N, 120° 19' E, north of Lubang Is., 167-187 m, a conjoined specimen

nUn^TpM 2: r',7’ l4°00T'N> 1 2°° 1 7' E, north of Lubang Is., 174-193 m. — Stn 19, 14°0TN
20 17 E, north of Lubang Is., 189-192 m. - Stn DR 33, 13°32' N, 12P07' E, north of Mindoro Is
uu-lj/ m, a conjoined specimen and many empty shells, (all mnhn)

™™3;-CP f’ 1f4:01'N’ 1207J E. north of Lubang Is'., 183-187 m. - Stn CP 98,
400 N’ i2° I8 E’ north of Lubang Is., 194-205 m. — Stn DR 102, 14°0T N, 120°18' E, north of

12? II'WN9 W Ftn Cf ^ 14"0'' N' ;20°18' E’ north of Lubang Is., 188-195 m. — Stn DR
P6’. f..N> 121 22 E, west of Panay Is., 266 m. — Stn CP 131, 11°37'N, 121°43' E, west of Panay

Is., 120-1_2 m. Stn CP 143, 1 1°29' N, 124°1 T E, northwest of Leyte Is., 205-214 m, a conjoined
specimen and several empty shells, (all mnhn). J

Siboga stn 95, Sulu Archipelago, 2 conjoined specimens (zma). — Stn 105, Sulu Archipelago
3 conjoined specimens (zma). h 6 ’

Indonesia. 6°0 T S, 133°57’ E, 12.8 km southwest of Tg Ratoe, Matkoor, Aru, Moluccas 45 m a

tITkJ rn (UT 755534!i - 5:32' S- 1 «* of Mitduau reef, west coast of Nu’hu

ljut, Rat s., 54-56 m, 1 conjoined specimen (usnm 747158). - "Siboga"'. stn 66, south of Salavar
Sulawesi, 1 conjoined specimen (zma). ydr’

Chesterfield Islands, chalcal 1: stn D 63, Banc Nova, 22°11'S, I59°15'E, 305 m (mnhn).
musorstom 5: stn 270, 24°49' S, 159°34' E, Banc Capel, 223 m. — Stn 277, 24° 1 V S 159°35' E Banc
Ke'so 270 m - Stn 285 24-09' S, 1 59^34' E. Banc Kelso, 245-255 m. - Stn 309 22« 0 S 59'>23 E

StJ S’ 1c58"47' E- 320 m- - Stn 350’ l9°34' S’ 158"33' H 280 m!

ToS4V5Joilot15&)80 m- - S,n 378- I9”54' s- I58°38' E- 355 m- - s“ 388’ 2°”«' s.

2rir s- i67°i2'E- 390 -

“o-M5 “S?1' S' 166°21' E' 330 m- - St” DW 253' 21°32' s' 166°29' E' ~

CHALCAL 2: stn DW 71, 24°42' S, 168°10' E, 230 m, 1 conjoined specimen (mnhn).

smIr 4 Stfn D^24:42;,S’ I68°08, E’ 265 m’ 1 conJ°ined specimen (mnhn).

smib 4. stn DW 44, S New Caledonia, 24°46' S, 168°08' E, 270-300 m. — Stn DW 47 S New

Caledonia, 24 46 S, 168°08' E, 250-280 m (all mnhn)

musorstom 6: stn DW 391, 20"47' S. 167°06' E, 390 m. - Stn DW 406 20°41' S

20°42' S 167W ET343n ”^nh) '' ^ * 36° m’ 1 C°nj°ined ~ Stn DW ^

DW RMe dS LOyaU“' 350 m’ 2 ^toens. - Stn

i v* U’. A 168 42 E- Rlde des Loyaute, 340 m. — Stn DW 97, 23°0T S I68°18' E 300 m
1 conjomed specimen. - Stn DW 98, 23°02’ S, 168=16’ E, 335 m. (all mnhn) '

dw n "4) srCi7r4,T:7i„n E? 7- F2T s' ‘a7'044' E’ Ridc des Loyamd' 325-40» - St”

w i / , zz zj n, i / 1 41 b, 260-300 m, 1 conjoined specimen, 6 rv, 6 Iv. — Stn DW 38, 22°22' S.

Source : MNHN, Paris

REVISION OF MEIOCARDIA AND GLOSSOCARD1A

91

, 68°44' E 380-420 m. - Stn DW 42, 22° 17' S, 168*42' E, 340-400 m. (all mnhn).

Reunion md32 Reunion: stn CP 129, 20*51' S, 55*36' E, 290-300 m (mnhn).

Distribution. — Hawaii, Japan, Philippines, Indonesia, Coral Sea, New Caledonia; Indian
Ocean side of Thailand, Reunion. This species lives on sand and mud bottoms in 45 to -20 m, and
empty shells are occasionally recorded from 500 m or deeper.

Description. - Shell small to medium for genus thim
subquadrate, glossy white. Mean of. L/H 1.279 (N 17, SD
= 0.038). not significantly correlated with L (r 0342 a
0.05). Mean of Cs/H .0.480 (N = 17, SD = 0 019) not
significantly correlated with L (r -0311, a _ 0.05). L/H
significantly correlated with Cs/H (N - 17, r 0.557, a
0 05) ( Fic 10) Outer surface around keel and postero-dorsal
area occasionally yellowish or reddish brown. Sharp, angular,

primary keel separating very smooth postero-dorsa area
from antero-ventral area with irregular commarginal ribs.
Anterior cardinal (1) oblique to dorsal margin; 2a dorsally
deeply concave, oblique to dorsal margin. Inner surface
creamy white, occassionally posteriorly pale brown, shining.
Anterior adductor scar semicircular; posterior adductor scar
subcircular; both scars nearly equal in size.

Measurements. See Appendix, Table 4.

Rfmarks — Adams (1864) described Bucardia (. Meiocardia ) cumingii on the basis of material
from CWna“ 1, hough the species was said to have a waxy white shell, he only gave a bnef descnphon
without anv measurements The type material was not found in bmnh during a visit by one of us (, )
" %9 ZS of ado pting a'doubtful and obscure name for this species, we prefer to use

Meiocardia hawaiana.

Family Trapezidae Lamy, 1920

Diagnosis — Shell trapezoidal, strongly inequilateral, equivalve, with prosogyrous beaks near
anteriofend Wntral margin" not gaping, smooth. Ligament external, ^J?***™^
Hinge with two lamellar cardinals and two laterals. Outer surface : ornamen ed w^^
radial sculpture. Dimyarian, with subequal adductor muscles. Foot small, grooved, often byssc .

Pallial line simple, without sinus.

Genus Glossocardia Stoliczka, 1870

Type species: Cypricardia obesa Reeve, 1843.

Diagnosis. - Shell subquadrate, thin in young, thick and solid in ^^^^'^t^osteriSr
Beaks not prominent. Larval ligament anterior to beaks^ Primary keel grating P but

slope, sharp. Two additional keels on posterior slope, the secondary and ^ tertiary _ , ‘orner

distinct; secondary keel, the weakest of the three, running from the ea o p , ’

separating postero-dorsal margin. The stronger, tertiary keel located on the posterior slop
primary keel. Outer surface nearly smooth except for weak, irregular commarginal ribs and ad.al
rows of microscopic scales. Hinge cyprinoid; teeth parallel to hinge margins an e ,, ,

right valve (1) lamellate in juvenile, nodulous in adult; posterior lateral m left valve (lpii) small, fn y

nodulated.

Remarks. - Only three living species, Glossocardia agassizii, G. obesajnd
belong to this genus. The Eocene Meiocardia carolinae Harris, 1919a, has a mo c P , •

tertiary keel on the posterior slope near the primary keel (Harris, 1 191 9b) and is safely placed n tins
genus. Meiocardia suzukii Squires & Advocate, 1986, from the Eocene anio Although M.
California, has shells that are not as inflated and lack strongly spirogyra e um • .

suzukii apparently lacks the tertiary keel on the posterior slope, it possib y e ongs

92

A. MATSUKUMA & T. HABE

Glossocardia obesa (Reeve, 1843)
Figs 18-19

Cypricardia obesa Reeve. 1843: Cvpricardia sp. 10: pi. 2, fig. 10 [locality unknown].
Other references:

Glossocardia obesa - Hare, 1951: 118-119, figs 247-248 [Okinawa. Japan],

Trapezium ( Glossocardia ) obesa (pars) - Solem, 1955: 73-74, pi. 6. figs I 1-12 (holotype).

Type material. - Holotype, a conjoined specimen, b.mnh 1952. 1 0. 1 0.7.8.

Type locality. — Not given.

■ .... , ,MateR'al examined. - Recent: Indonesia. Silale, Ambon (nsmt-Mo69759)

Ph.hpp.nes^ Batangas Is.. N of Cebu, coll. Poppe, 1 conjoined specimen (mnhn).

E- 55 m- 1 co"joi"ed ^ - st"
New Caledonia, lagon: stn 612, 22''09’ S. 167°0I'E. 46-48 m. I rv (mnhn).

Chesterfiel™™' " ReCe"t: °ki”aWa' Japan: Philipp,r,es; Indonesia; New Caledonia;

Description. Shell trapezoidal, thin in juveniles but
thick and solid in full-grown specimens. Beaks prosogyrous.
'ow- no* strongly enrolled. Larval ligament small, anterior to
beaks I rimary keel low, separating slightly concave postero-
aorsal area. Outer surface ornamented' with irregularly

spaced commarginal lamellae with numerous microscopic
scales. Postero- ventral corner bluntly pointed. Inner surface
creamy white. Anterior adductor scar semicircular, thickened
ventrally; posterior adductor scar subcircular: both scars
nearly equal in size.

Measurements (mm):

nsmt-Mo69759-1

M 069759-2 _

*Numericals in parentheses

V L H

rv + Iv 31.5 20.5

rv 61.6 43.3

show Cb/2H, instead of Cs/H,

L.H

1.53

1.42

Cb

Cs

Cs/H

16.4

-

(0.400)

-

21.2

0.490

Remarks. \ oung specimens of Glossocardia obesa are apparently very close to Meiocardia
species m having a more or less translucent shell with the larval ligament anterio to Ae bSto a^d

the^outer Sc^ Cardma'S’ bUt ^ cteari* differ in havin^S^

Glossocardia stoliczkana (Prashad. 1932)

Figs 17, 20, 39

m,lakam 1931 «>■ s. figs 15-18 [Sulu Archipelago, Philippine*

Type material. — Holotype, a conjoined specimen, ZMA no registration number.

564 m TVPE LOCAL,ty- - 'Sihoga", stn 97, 5"48.7'N, 119"49.6'E, Sulu Archipelago. Philippines,

Recent^hmMtaefsidir?’ w C" (?): Niuc' a Gonj°ined specimen, G. Paulay coll, (usnm)

(NSMT M?57S6 n P 8° (h0l°type- “*>• M«an Island. Cebu, a conjoined specimen

Source MNHN. Paris

REVISION OF ME IOC A RDIA AND GLOSSOCARDIA

93

Figs 17-20. Shells of Glossocardia. 17a-c. Glossocardia sioliczkana. 1 sheli^'enetli1' 7 1°^ !m" 19a-b!

length 19.8 mm. 18a-d, Glossocardia obesa. Ambon Indonesia (wm-MoWM). She! I'engtn icz m
Glossocardia obesa. Ambon. Indonesia (nsmt-Mo69759) Shell length 61.6 mm. 20a b. Gtossocaraia
Loyalty Ridge, New Caledonia (mnhn). Shell length 19.3 mm.

MUSORSTOM 2: stn DR 33, I3"32' N, 121°08' E, north of Mindoro ^ 130-137 m 1 rv (mnhn).
musorstom 3: stn DR 137. 12°03' N, 122°06' E, north of Panay Is., 56 m, 1 lv (mnhn).

Loyalty Islands, musorstom 6: stn DW 391, 20°47 S, 167 06 E, 390 m, 1 rv - SJn ™ ^
20'’47' S, 1 67-06' E, 370 m. 1 rv. Stn DW 418. 20°42' S. 167*03'. E a 283 m, 1 rv - Stn DW 439,
20"46' S. 167° 17' E, 288 m, 1 rv. — Stn DW 440, 20"49 S, 167 17 E, -88 m, 1 lv. (c

Distribution. - Pleistocene: Niue. Recent: Cebu, Mindoro, Panay, Sulu Archipelago,
Philippines; New Caledonia. This species is known from 56 to 564 m.

Description. Shell small, thin, inflated, high, quadran¬
gular. strongly inequilateral, equivalve. Mean of L FI 1.399
(N = 8, SD = 0.132). not significantly correlated with L
(r = -0.486. a = 0.05). Cs/H significantly correlated with L
(r = 0.885, a = 0.05) (Fig. 21). L/H not significantly
correlated with Cs/H (N = 8, r = -0.660, a - 0.05).
Umbones prosogyrous, low, not prominent, translucent.
Outer surface white, ornamented with weak commarginal
ribs and radial rows of fine scales: posterior area orange,
smooth, with two narrow radial riblets (Fig. 39b, s and t) and
fine growth striae, concave, separated from middle area by a

harp keel. Postero-ventral margin pointed without a sinua-
lon like M. sanguineomaculala. Inner color white, oran^
iosteriorly. Hinge- teeth thin; 1 thin and lamellar in young
pecimens^ and solid, and rounded-lamellar in older speci¬
mens 3a long. thin; 3b weakly bifid; la, small; la,., small
iut distinct; i an long, strong; lphi small, long and narrow 2a
mall, thin; 2b rather long, thin; 4b long and narrow. _th n;
aii very small, indistinct; semicircular, thickened ventrally.
iosterior adductor scar subcircular or ovale. Pallial scar
■ntire. wide, integripalliate.

94

A. MATSUKUMA & T. HABE

0.6

0.5

Cs/H

FlG' ? Scatter diagram showing relationship between L
and Cs/H in Glossocardia. Dots: Glossocardia stoliczkana.
N - 8: r = 0.885; regression equation Cs/H = 0.0103L
+ 0.258. Circles: Glossocardia agassizii. N = 14; r =
0.676; regression equation Cs/H = 0.00470L + 0.345.

0.4

o

* o

o

o °

L (mm)

10 20

Measurements (mm):

30

V

L

H

L/H

Cb

Cs

Cs/H

nsmt-Mo57861
musorstom 6: stn DW391
stn DW398
stn DW418
stn DW439
stn DW440

musorstom 2: stn DR33
musorstom 3: stn DR 137

rv +

rv +

rv

rv

rv

rv

lv

rv

lv

lv 19.8

lv 24.7

18.5

27.8

30.7

12.1

19.3

9.9

15.2

12.3

18.2

14.9

22.3

23.5

8.3

13.1

6.9

10.0

1.61

1.36

1.24

1.25

1.31

1.46

1.47

1.43

1.52

9.6

17.5

7.0

11.9

14.5

3.3

6.3

2.6

4.2

(0.390)*

(0.481)

0.470

0.534

0.617

0.398

0.481

0.377

0.420

‘Numericals in parentheses show Cb/2H, instead of Cs/H.

39,. S,nRDW 39S

stnae- Addit,onal spec'imms are -eeded

vou„s SK? T 'iCZkrana suPerficially resembles the tropical western Atlantic G. agassizii and
young shells of G. obesa from the tropical West Pacific. These three species have radial rows of
microscopic scales on the outer surface and a larval ligament at the anterior margin of beaks

um boT ‘"d Shorter Tntero domal ^ a^assizii. haVing the higher and more slronS'y enrolled
postero-dorsa of? 3 poster°-dorsaI Commarginal striae in the

postero dorsal area of G. agassizii are finer and more regularly spaced than those of G stnlir-kmm

Glossocardia stoliczkana clearly differs from G. obesa in having a distinct tertiary ^ee! and fine

7e la,e °n thC postenor sI°Pe- The cardinal (1) in the right valve of G stoliczkana in
young and adult specimens, is lamellar and parallel to ventral margin whereas the cardinal 7 h
of G. obesa ,s thin and lamellar in young shells, and is strong and triangXr !n adults 0>

them .^!OSSOCardl“ St<>^zkana is very similar to M. hawaiana and M. samara, u-iae but differs from
them in having a higher, thinner, shell with lower umbones, radial rows of microscopic scales^ «n t e
outer surface, and a distinct tertiary keel on the yellow posterior slope (F™ 21) P

Source . MNHN. Paris

REVISION OF MEIOCARD1A AND GLOSSOCARDIA

95

Glossocardia agassizii (Dali, 1886)
Fig. 38

Meiocardia agassizii Dali, 1886: 271 [Trinidad].

Other refe ^ [Trinidad] . — Abbott, 1974: 5847, fig. 5848 (- Dall, 1889: pi. 40,

£T[wtt Fafonda the CaribbelS Bemuda]. - NaIlchi, 1976: 205-210. figs 1-3 [Rio Doce. Espirito Santo, Braz.1], - Rtos.
I985 259. pi. 91, fig- 1286 (= Dall, 1889: pi. 40. fig. 7) [Espirito Santo, Brazil],

Type material. — Depository unknown.

Type locality. Off Trinidad.

Material examined. — Recent: Bermuda. “Bermuda” (mcz 152851); 1.2 km S of Castle
Rock, 144-180 m (MCZ 161960).

Tnha 22°09'30" N 81°11'W, Bahia de Cochinos, 315-405 m (mcz 16504).

British Guiana. 08°10.5' N-08°10.0' N, 57°48' W, 153.6 km N of Georgetown, 96-106 m (mcz 27.570).

Distribution. Bermuda; Cuba; Trinidad; British Guinea; Brazil (19°35' S i, 39"20' W, off the
mouth of the Rio Doce, Espirito State, 73 m, mud, January 1972, coll, by R.S. Wladmir Besnard ,
Oceanographic Institute of the University of Sao Paulo) (Narchi pers. comm.). This species is known
from muddy bottoms in 73 to 405 m.

Description. — Shell cordiform, thin, inflated, strongly
inequilateral, equivalve, with prosogyrous beaks. Mean oi
L H 1 377 (N = 14, SD = 0.045), not significantly correlated
with L (r = -0.146, a = 0.05). Cs/H significantly correlated
with L (r = 0.676, a = 0.05) (Fig. 21). L/H not significantly
correlated with Cs/H (N = 14, r = -0.000, a = 0.05). Larval
ligament anterior to beak. Outer surface ornamented with

weak commarginal lamellae and microscopic scales. Primary
keel sharp, separating slightly concave postero-dorsal area
which has an obscure secondary keel and a narrow but
conspicuous tertiary keel. Postero-dorsal area with regularly
spaced commarginal lamellae. Inner surface glossy white.
Anterior adductor scar semicircular; posterior adductor scar
ovate; both scars nearly equal in size.

Measurements (mm):

mcz 16504
mcz 152851
mcz 161960
MCZ 273570-1
273570-2
273570-3
273570-4
273570-5
273570-6
273570-7
273570-8
273570-9
273570-10
273570-11
273570-12

V

L

_

13.9

rv + lv

13.7

rl

23.6

rv

27.0

rv

22.6

rv

17.9

rv

15.8

rv

16.5

lv

22.6

lv

19.0

lv

17.7

lv

18.6

lv

18.7

lv

15.2

lv

14.8

H

L/H

10.0

1.39

9.3

1.47

17.7

1.33

18.9

1.43

16.7

1.35

13.1

1.37

12

1.32

12.1

1.36

16.9

1.34

14.1

1.35

13.3

1.33

13.0

1.43

13.2

1.42

11.4

1.33

10.7

1.38

*Numericals in parentheses show Cb/2H. instead of Cs/H.

Cb

7.3

Cs

4.5

7.7

9.6

7.6

5.7

4.8

7.3
6.0

5.8

5.4

5.7

4.8
4.6

Cs/H

0.450

(0.392)*

0.435

0.508

0.455

0.435

0.397

0.432

0.426

0.436

0.415

0.432

0.421

0.430

Source . MNHN, Paris

96

A. MATSUKUMA & T. HABE

Remarks. Dall (1886. 1889) described this species as a member of the genus Meiocardia,
and American authors have apparently followed this assignment (Abbott, 1974; Narchi, 1976; Rios.'
1985). It has a thin, inflated, cordiform shell, with a sharp primary keel, a prominent tertiary keel,
and a larval ligament anterior to the beaks. The hinge teeth of this species are also superficially close
to Meiocardia species of the Indo-West Pacific. The outer surface of Dali’s species is ornamented with
weak commarginal lamellae and numerous microscopic scales, which is similar to the ornament on
Glossocardia obesa of the Trapezidae but which is never found on shells of Meiocardia. This should
be therefore transfered from the genus Meiocardia, Glossidae. to the genus Glossocardia, Trapezidae.

ACKNOWLEDGEMENTS

We thank the following persons and institutions for allowing us access to the material stored
in their collections: Professor Walter Narchi. Universidade de Sao Paulo. Brazil; Dr Gustav Paulay,
formerly at University of Niue; Dr Henry E. Coomans and Mr Robert Moolenbeek. Zoologisch
Museum, Amsterdam; Dr Philippe Bouchet and B. Metivier, Museum National d'Histoire Naturelle,
Paris; Dr Bertrand Richer de Forges, orstom, Noumea. Dr Solene Morris, formerly of The Natural
History Museum, London; Professor Kenneth Boss, Museum of Comparative Zoology, Harvard
University; the late Dr Richard Houbrick, National Museum of Natural History, Smithsonian
Institute; Dr George Davis. Academy of Natural Sciences, Philadelphia; Dr James McLean and the
late Mr Clifton Coney. Los Angeles County Museum; Dr Paul Scott, Santa Barbara Museum of
Natural History.

Dr Eugene Coan, San Francisco, and B. Metivier kindly read our draft and gave us fruitful
comments and suggestions for improving this paper.

Some specimens stored in National Science Museum, Tokyo, were collected by members of the
overseas research project ••Diversity of marine organisms from Indonesian shallow waters”
financially supported by Ministry of Education, Culture and Science, Japanese Government (project
no. 04041103). This paper constitutes Contribution no. VII of Studies on the Kawamura Collection
(Mollusca) stored in the National Science Museum, Tokyo.

Figs 22-29. 22 a-b. Meiocardia sanguineomaculata. REVES 2: stn 10. Seychelles (mnhn). Shell length 1 1.9 mm. - a. left side

view. b. postero-dorsal view. 23 a-f. Meiocardia sanguineomaculata. reves 2: stn 30, Seychelles (mnhn). Shell length

15.0 mm. a anterior view. — b, posterodordal view. c, left side view. d. inside view of right valve. e. right
w i/’ rnS,‘de V‘?W of ’5rt val»e T 24' Meiocardia moltkiana. Pleistocene Ryukyu Limestone. Kikaijima Is.,

Kagoshima Prefecture, Japan (nsmt-Mo69745). Shell length 33.4 mm. 25, Meiocardia moltkiana. Chichi jima
Ogasawara Islands. Japan (nsmt-Mo59790). Shell length 28.2 mm. - 26 a-d. Meiocardia moltkiana . China (zma). Shell
length 33 4 nun. a. inside view of right valve. b. left side view. - c. anterior view. d. postero-dorsal view -
27 a-b Meiocardia moltkiana MD 32: stn CP 43. Reunion (mnhn). Shell length 25.3 mm. a. right side view
Ll,?,hd™TW ° tHe n8h* va|ve. 28 a-b. Meiocardia moltkiana. smib 5: stn DW 81, New Caledonia (mnhn). Shell
H?,, w V d Vgh Sld.e v'evv' b- ms,de Vlew °r the right valve. 29 a-e. Meiocardia samarangiae. Tanabe
Bay, Wakayama Prefecture. Japan (nsmt-Mo69752). Shell length 29.7 mm. a. left side view I, right side
view. c. inside view of the right valve. d. anterior view. — e, postero-dorsal view.

Source : MNHN , Paris

REVISION OF MEIOCA RDIA AND GLOSSOCA RD1A

97

Source : MNHN, Paris

98

A. MATSUKUMA & T. HABE

F,°S t ,‘4 - '“o'yP" “7-M069749,. Shell length 24.8

valve. f. inside view of the right valve 11 ah uliZ i d'- f>*?1 ,Slde vlew‘ "" e- lnside VICW of the left
29 mm. - a. anterior view - b left side view 17 Menard, a vulgar, s. Philippines (Habe Coll.). Shell length appr.
Coll.). Shell length 46.1 mm - a left tide view f’ h' *!°c“rd,a.vul8a™ Philippines (nsmt-Mo69747, Kawamura
d, postero-dorsaf view. 'el'righ, s^de^view^f, inside view' oftlnf leffvalve. - «■ -

Source : MNHN, Paris

REVISION OF MEIOCARDIA AND GLOSSOCARDIA

99

Source : MNHN, Paris

100

A. MATSUKUMA & T. HABE

Figs 33-39. 33. Meiocardia hawaiana. " Siboga stn 105. Sulu Archipelago (zma). Shell lcnalh 17.1 mm

34 a-b. Meiocardia hawaiana. smib 5: stn DW 90. New Caledonia (mnhn). Shell length 17.1 mm. a. right side
view. b. inside view ol the right valve. 35 a-e. Meiocardia hawaiana. "Siboga”'. stn 105. Sulu Archipelago (zma).
Shell length 27.2 mm. a. left side view. b. inside view of the right valve, c. postero-dorsal view.

36 a-e. Meiocardia hawaiana. Oahu. Hawaii (usnm 807654). Shell length 14.4 mm. a. left side view. b. inside view
of the left valve. c. inside view of the right valve. d. dorsal view. e. postero-dorsal view. 37 a-b. Meiocardia
hawaiana. Kochi Prefecture. Japan (nsmt-Mo43439-1 ). Shell length 16.9 mm. a. left side view. b. inside view of
the right valve. - 38 a-b. GlossoCardia agassizii. Castle Rock. Bermuda (mcz 161960). Shell leneth 23.6 mm. a, rieht
side vew. p: Primary keel, t: Tertiary keel. b. inside view of the right valve. 39 a-f. Glossocardia sloliczkana. Cebu.
Philippines (nsmt-Mo57861 ). Shell length 24.7 mm. p: Primary keel, s: Secondary keel, t: Tertiary keel. a. anterior
view. Arrows indicate larval ligaments. b. postero-dorsal view. - c. left side view'. d. inside view of the left
valve. - e. right side view. — f, inside view of the right valve.

Source : MNHN , Paris

REVISION OF MEIOCA RDIA AND GLOSSOCARDIA

101

Source

102

A. MATSUKUMA & T. HABE

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Abrard. R.. 1947. — Fossiles neogenes et Quaternaires des Nouvelles-Hebrides. Annates de Paleonto\ogie, 32: 1-112, pis 1-5.

Adams, A.. 1864. On some new genera and species of mollusks from the seas of China and Japan. Annates and Magazine
of Natural History, ser. 3, 13: 140-144.

Adams, A. & Reeve, L. A.. 1850. — The Zoology of the voyage of H.M.S. Samarang. Mollusca. 87 pp.. 24 pis.

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Bernard, F. R., Cai. Y.-Y. & Morton, B., 1993. — Catalogue of the living marine bivalve molluscs of China. Hong Kong
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Boss. K. J., Rosewater, J. & Ruhoff, F.. 1968. — The zoological taxa of William Healey Dali. Bulletin of the United Slates
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Bruguiere, J. G., 1797. Tableau encyclopedique et methodique des trois regnes de la nature. Vingt-troisieme partie,
mollusques et polypes divers, par M. Lamarck. Pis 190-286.

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Gesellscliaft, (1906[1]): 33-38. pis l(7)-2(8).

Chemnitz, J. F., 1784. — Neues Systematisches Conchylien-Cabinet, vol. 7. 356 pp., vignette 8-11, pis 37-69.

Chf.nu, J. C., 1862. — Manuel de conchyliologie et de paleontologie conchyliologique, vol. 2. Masson, Paris. 327 pp.

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Source : MNHN, Paris

REVISION OF MEIOCA RDIA AND GLOSSOCARDIA

103

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104

A. MATSUKUMA & T. HABE

Appendix: Tables of measurements.

Table 1. Meiocardia moltkiana. Measurements (mm) of selected material.

V

L

H

L/H

Cb

Cs

Cs/H

nsmt-Mo 42307

TV + Iv

18.9

15.1

1.25

13.6

(0.450)

nsmt-Mo 43435

TV + IV

18.4

14.9

1.23

13.0

.

(0.436)

nsmt-Mo 43436 (1)

Iv

24.9

19.6

1.27

_

10.3

(0.526)

Mo 43436 (2)

Iv

13.0

10.6

1.23

.

4.8

0.453

Mo 43436 (3)

Iv

11.6

9.7

1.20

.

5.1

0.526

Mo 43436 (4)

TV

10.1

7.8

1.29

.

3.7

0.474

nsmt-Mo 43455 (1)

Iv

21.1

15.4

1.37

.

7.7

0.500

Mo 43455 (2)

Iv

17.4

14.1

1.23

.

6.7

0.475

Mo 43455 (3)

rv

13.9

11.0

1.26

.

5.0

0.455

Mo 43455 (4)

TV

13.7

11. 1

1.23

5.4

0.486

Mo 43455 (5)

rv

12.2

9.3

1.31

.

4.3

0.462

Mo 43455 (6)

TV'

11.4

9.3

1.23

.

4.3

0.462

nsmt-Mo 59790

rv

28.2

25.5

l.ll

.

14.1

0.553

nsmt-Mo 63233
nsmt-Mo 69739

rv + Iv
rv + Iv

17.1

22.9

13.5

17.7

1.27

1.29

1 1.4

16.2

(0.422)

(0.458)

nsmt-Mo 69740

rv + Iv

17.5

14.8

1.18

13.3

(0.449)

nsmt-Mo 69741

rv + lv

28.1

21.7

1.29

21.0

(0.484)

nsmt-Mo 69742

rv + lv

29.7

22.9

1.30

22.3

(0.487)

nsmt-Mo 69743

rv + Iv

22.5

18.8

1.20

18.0

(0.479)

nsmt-Mo 69745 (1)

Iv

33.4

30.1

111

14.9

0.495

Mo 69745 (2)

rv

22.1

20.3

1.09

Mo 69745 (3)

Iv

19.1

14.9

1.28

.

7.4

0.497

zma China- 1

rv + Iv

33.4

24.3

1.37

23.4

(0.48 1 )

China-2
md32 DC41 (1)

rv + lv

rv

33.8

17.3

24.2

13.7

1.40

1.26

23.3

6.4

(0.484)

0.467

DC41 (2)

rv

17.4

13.3

1.31

.

5.9

0.444

DC41 (3)

rv

15.6

12.7

1.23

6.1

0.480

DC4I (4)

rv

14.0

10.9

1.28

.

5.1

0.468

DC4 1 (5)

rv

11.3

8.7

1.30

4.1

0.471

DC41 (6)

rv

5.4

4.3

1.26

2.2

0.512

DC4I (7)

lv

33.2

25.0

1.33

.

12.3

0.492

DC4I (8)

lv

26.0

20.4

1.27

.

10.0

0.490

DC41 (9)

lv

25.4

20.0

1.27

.

9.6

0.480

DC41 (10)

Iv

17.4

13.6

1.28

6.6

0.485

DC41 (11)

Iv

14.5

10.8

1.34

5.2

0.48 1

DC4I (12)

lv

13.6

10.6

1.28

5.1

0.481

DC41 (13)

lv

8.5

6.9

1.23

3.7

0.536

DC41 (14)
md32 DC43 (1)

lv

rv

7.8

31.4

6.2

24.0

1.26

1.30

-

3.2

11.9

0.516

0.496

DC43 (2)

DC43 (3)

rv

rv

31.0

29.9

23.1

23.1

1.34

1.29

-

10.9

1 1.4

0.472

0.494

DC43 (4)

rv

26.6

20.4

1.30

9.8

0.480

DC43 (5)

DC43 (6)

DC43 (7)

DC43 (8)

DC43 (9)

DC43 (10)

DC43 (11)

DC43 (12)
md32 DC 1 26 (1)

DC 126 (2)

DC 126 (3)

DC 126 (4)

rv

rv

lv

Iv

lv

lv

Iv

Iv

rv

rv

rv

rv

26.0

22.3

32.2

28.7

28.2

27.3

26.5

26.2

23.2

19.6

17.4

13.6

20.3

16.8

23.8

22.6

22.1

21.2

20.9

20.4

18.6

16.1

13.4

10.8

1.28

1.33

1.35

1.27

1.28

1.29

1.27

1.28

1.25

1.22

1.30

1.26

-

9.7

7.9

11.8

10.9

10.3

10.1

10.0

9.9

8.5

7.6

6.0

5.0

0.478

0.470

0.496

0.482

0.466

0.476

0.478

0.485

0.457

0.472

0.448

0.463

Source : MNHN, Paris

REVISION OF MEIOCA RDIA AND GLOSSOCARDIA

105

V L

DC 126 (5)

rv

11.4

DC 126 (6)

rv

10.8

DC 126 (7)

rv

9.8

DC 126 (8)

rv

9.7

DC 126 (9)

rv

8.4

DC 126 (10)

rv

8.5

DC 1 26 (11)

rv

7.7

DC 126 (12)

rv

7.5

DC 126 (13)

rv

7.3

DC 126 (14)

rv

6.7

DC 126 (15)

rv

5.2

DC 126 (16)

rv

3.8

DC 126 (17)

lv

16.9

DC 126 (18)

lv

14.5

DC 126 (19)

lv

14.4

DC 126 (20)

lv

13.3

DC126 (21)

lv

12.1

DC 126 (22)

lv

9.8

DC 126 (23)

lv

8.7

DC 126 (24)

lv

6.9

DC 126 (25)

lv

7.1

DC 126 (26)

lv

6.8

DC 126 (27)

lv

5.9

md32 DC 1 76 (1)

rv

10.4

DC 176 (2)

lv

7.9

DC 176 (3)

lv

6.7

H

L/H

Cb

Cs

Cs/H

8.7

1.31

.

4.2

0.483

8.1

1.33

-

3.9

0.481

7.5

1.31

-

3.6

0.480

7.2

1 .35

-

3.6

0.500

6.7

1.25

-

3.4

0.507

6.2

1.37

-

3.2

0.516

5.6

1.38

-

2.8

0.500

5.7

1.32

-

2.8

0.491

5.5

1.33

-

2.7

0.491

5.3

1.26

-

2.7

0.509

4.0

1.30

-

1.9

0.475

3.1

1.23

-

1.3

0.419

13.0

1.30

-

6.0

0.462

11.6

1.25

-

5.4

0.466

10.7

1.35

-

5.1

0.477

9.8

1.36

-

4.7

0.480

9.0

1.34

-

4.5

0.500

7.4

1.32

-

3.6

0.486

7.0

1.24

-

3.1

0.443

5.6

1.23

-

3.1

0.554

5.1

1.39

-

2.7

0.529

5.2

1.31

-

2.4

0.462

4.7

1.26

-

-

-

7.5

1.39

-

4.0

0.533

6.1

1.30

-

3.5

0.574

5.1

1.31

-

2.7

0.529

*Numericals in parentheses show Cb/2H, instead of Cs/H.

Table 2 — Meiocardia sanguineomaeulata. Measurements (mm) of selected material.

V

L

liMNit 1910.8.31.688

rv

+

lv

13.6

1910.8.31.689

rv

+

lv

7.8

bmnii 1989178

rv

+

lv

17.4

1989179 (1)

rv

+

lv

19.3

1989179 (2)

rv

+

lv

19.7

1989179 (3)

rv

+

lv

6.0

usnm 235929

lv

10.3

usnm 235950

rv

10.5

usnm 237240

rv

(18)

usnm 294513 (1)

lv

9.0

294513 (2)

lv

5.0

294513 (3)

lv

3.0

294513 (4)

lv

2.2

294513 (5)

rv

4.6

294513 (6)

rv

4.0

usnm 294545 (1)

rv

4.5

294545 (2)

lv

9.0

USNM 431190 (1)

rv

8.0

431190 (2)

rv

8.1

431190 (3)

rv

4.6

431190 (4)

lv

12.0

431190 (5)

lv

8.5

431190 (6)

rv

4.0

rhves 2 Stn 30

rv

+

lv

15.0

Stn 24

rv

+

lv

13.2

H

L/H

Cb

Cs

Cs/H

11.3

1.20

13.0

-

(0575)*

6.7

1.16

7.3

-

(0.545)

14.9

1.17

14.9

-

(0.500)

15.7

1.23

17.7

-

(0.564)

17.2

1.15

18.0

-

(0.523)

5.0

1.20

4.6

-

(0.460)

10.1

1.02

-

5.1

0.505

9.8

1.07

-

4.9

0.500

15.4

_

_

7.0

0.455

7.9

1.14

-

3.8

0.481

4.5

1.11

-

2.2

0.489

2.5

1.20

-

1.5

0.600

2.0

1.10

-

1

0.500

3.9

1.18

-

2.0

0.513

3.4

1.18

-

1.8

0.529

3.4

1.18

-

1.8

0.529

7.3

1.23

-

4.1

0.562

7.4

1.08

-

3.6

0.486

6.6

1.23

-

3.6

0.545

4.4

1.05

-

2.2

0.500

10.8

l.ll

-

5.4

0.500

8.2

1.04

-

4.1

0.500

3.4

1.18

-

1.18

0.529

12.4

1.21

13.6

-

(0.548)

11.9

1.11

11.8

-

(0.496)

Source

106

A. MATSUKL’MA & T. HABE

V

L

H

L/H

Cb

Cs

Cs/H

Stn 18

rv

+

Iv

19.3

16.6

1.16

17.8

-

(0.536)

Stn 16

rv

+

Iv

10.9

9.7

1.12

9.0

-

(0.464)

Stn 10

rv

+

Iv

11.9

10.0

1.19

9.5

-

(0.475)

*Numericals in parentheses show Cb/2H, instead of Cs H.

Table 3. Meiocardia vulgaris. Measurements (mm) of selected material.

V

L

H

L/H

Cb

Cs

Cs/H

bmnh syntypes 1

rv

+

Iv

48.6

38.6

1.26

36,0

_

(0.466)*

2

rv

+

Iv

37.6

31.9

1.18

31.4

-

(0.492)

3

rv

+

Iv

38.2

30.5

1.25

30.7

-

(0.503)

4

rv

+

Iv

36.4

29.3

1.24

28.6

-

(0.488)

MUSORSTOM 3 CPI41

rv

+

lv

25.0

21.1

1.18

19.7

-

(0.467)

CORINDON Stn 205 (1)

rv

+

Iv

14.6

13.8

1.06

13.4

-

(0.486)

Stn 205 (2)

rv

+

Iv

14.1

13.2

1.07

13.3

-

(0.504)

Stn 205 (3)

Iv

14.2

12.7

1.12

-

6.7

0.528

zma China (1)

rv

+

lv

34.3

27.8

1.23

26.7

-

(0.480)

China (2)

rv

+

lv

33.3

27.3

1.22

26.5

-

(0.485)

China (3)

rv

+

lv

23.9

21.5

1.11

21.2

-

(0.493)

China (4)

rv

+

lv

23.5

20.0

1.18

20.0

-

(0.500)

nsmt-Mo 54938

rv

24.3

20.3

1.20

-

9.9

0.488

nsmt-Mo 60557

rv

+

lv

31.5

26.2

1.20

24.9

-

(0.475)

nsmt-Mo69746

rv

+

lv

29.6

25.2

1.17

23.7

-

(0.470)

nsmt-Mo 69747 (1)

rv

+

lv

39.3

30.5

1.29

30.3

(0.497)

Mo 69747 (2)

rv

+

lv

36.4

29.3

1.24

27.7

-

(0.473)

nsmt-Mo 69748 (1)

rv

+

lv

46.1

35.0

1.32

32.4

-

(0.463)

Mo 69748 (2)

rv

+

Iv

35.5

28.7

1.24

26.3

-

(0.458)

*NumericaIs in parentheses show Cb/2H, instead of Cs H.

Table 4. — Meiocardia hawaiana. Measurements (mm) of selected material.

V

L

H

L/H

Cb

Cs

Cs/H

nsmt-Mo 43439 (1)

rv + Iv

16.9

13.0

1.30

12.0

_

(0.462)

Mo 43439 (2)

rv + lv

19.1

14.8

1.29

14.5

.

(0.490)

MUSORSTOM 3 CP 143 (1)

rv + lv

27.4

20.5

1.34

19.2

_

(0.468)

CP 143 (2)

rv + Iv

22.4

17.6

1.27

16.3

-

(0.463)

CP 143 (3)

rv

28.9

21.9

1.32

-

10.9

0.498

CP 143 (4)

rv

24.4

19.7

1.24

_

9.5

0.482

CP 143 (5)

rv

23.7

18.4

1.29

-

8.8

0.478

CPI43 (6)

rv

22.3

18.1

1.23

.

8.3

0.459

CP 143 (7)

rv

20.9

16.0

1.31

_

7.6

0.475

CP 143 (8)

rv

17.9

14.3

1.25

.

6.9

0.483

CP 143 (9)

Iv

27.7

22.1

1.25

_

10.4

0.471

CP 1 43 (10)

Iv

26.6

21.7

1.23

-

10.5

0.484

CPI43 (11)

Iv

26.4

20.1

1.31

_

10.3

0.512

CP143 (12)

Iv

25.7

19.2

1.34

_

10.0

0.521

CP143 (13)

Iv

25.4

19.5

1.30

-

9.7

0.497

CP 143 (14)

Iv

22.2

17.2

1.29

_

8.0

0.465

CP 143 (15)

Iv

21.9

17.8

1.23

_

7.9

0.444

CP 143 (16)

lv

20.6

16.1

1.28

-

7.9

0.491

MUSORSTOM 1 Stn 19

rv + lv

12.3

9.8

1.26

9.3

-

(0.474)

*Numericals in parentheses show Cb/2H, instead of Cs H.

Source

ILTATS DES CAMPAGNES MUSORSTOM. VOLUME 14 RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 14 RESUl

3

Carnivorous bivalve molluscs (Anomalodesmata)
from the tropical western Pacific Ocean,
with a proposed classification
and a catalogue of Recent species

Jean- Maurice POUTIERS

Museum National d'Histoire Naturelie
55 rue de Buffon, 75005 Paris
France

&

Frank Reinhold BERNARD f

Department of Fisheries & Oceans
Pacific Biological Station, Nanaimo, B.C.

Canada V9R 5K6

CONTENTS

Abstract . 108

Introduction . 108

Species accounts . 110

Classification . 138

General remarks . 138

Synopsis of classification . 139

Diagnoses of supraspecific taxa . 141

Catalogue of recent species .

Appendix 1: Station data . 169

Appendix 2: Translation of Scarlato & Starobogatov 1983 . '

References . ' 7 6

rciuiieio. j.-ivi. i.in.. ^ - . . “ . . . . . - i - - ----- n

classification and a catalogue of Recent species. In: P. Bouchet (ed ), Resultats des Campagnes MUSORSTOM, Volume 14. Mem. Mm. nam. His!, nal.. 167: 107-18/. Paris
isb\ 2-85653-217-9.

Published 29th December 1995.

Source : MNHN, Paris

108

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

ABSTRACT

This paper deals with 37 species of carnivorous anomalodesmatan bivalves usually assigned to the septibranchs,
originating from the Philippines. Indonesia and New Caledonia. Eight species are described as new: Cetoeoncha boucheti , C.
exigua, Cuspidaria morrisae, C. (Soyomya) claihraia , Halicardia houbricki, Haliris leporis, Myonera rostra and Policordia
olivacea. New names are also provided here for two species with preoccupied names: Policordia ivanovae , for P. japonica
Ivanova, 1977, and Myonera alleni for Cuspidaria (Myonera) atlantica Allen & Morgan, 1981. A new classification of the
carnivorous Anomalodesmata, based on F. R. Bernard's latest works, is proposed together with diagnoses of the supraspecific
taxa and a catalogue of the living known species.

RESUME

MoIIusques Bivalves carnivores (Anomalodesmata) des regions tropieales du Pacifique occidental, avec une nouvelle
classification et un catalogue des especes actuelles.

Cet article decrit 37 especes de bivalves carnivores Anomalodesmata traditionnellement qualifies de septibranches
provenant des lies Philippines, d'lndonesieet de Nouvelle-Caledonie. Huit especes sont nouvelles pour la science : Cetoconclia
boucheti, C. exigua, Cuspidaria morrisae , C. ( Soyomya ) clathrata, Halicardia houbricki, Haliris teporis, Myonera rostra et
Policordia olivacea. De nouveaux noms ont aussi ete attribues a deux especes aux noms preoccupes: Policordia ivanovae, pour
P. japonica Ivanova, 1977, et Myonera alleni pour Cuspidaria (Myonera) atlantica Allen & Morgan, 1981. Une nouvelle
classification des Anomalodesmata carnivores, fondee sur les derniers travaux de F. R. Bernard, est presentee ainsi que des
diagnoses des differents taxa supraspecifiques et un catalogue des especes vivantes acluellement connues.

INTRODUCTION

FOREWORD

The present paper is based on a manuscript of the second author, the late Dr Frank R.
Bernard. At the end of 1988, Dr P. Bouchet showed me a first draft of a manuscript on West Pacific
septibranch bivalves for which Dr Bernard wanted to have my comments before its completion,
because I knew of the material he dealt with, having had the opportunity to study it partly and to
publish a few papers about it. I quickly gave him an answer, but Dr Bernard could not work further
on this paper and died shortly after, 29 March 1989. Later, the material on loan to him was returned
to Paris, through the courtesy of Drs Glen S. Jamieson of the Pacific Biological Station, Nanaimo,
and James A. Cosgrove of the Royal British Columbia Museum, Victoria, British Columbia
(Canada). I was then asked by P. Bouchet to revise Bernard's draft for publication.

As it was already apparent to me in 1988, the manuscript was far from complete and could
not be published without substantial revision. But it seemed worthwhile to do it, especially because
of its classification of carnivorous Anomalodesmata, which can be considered as the outcome of Dr
Bernard's ideas on the systematics of the septibranch group (for a review of F. R. Bernard's
contributions, see Matsukuma, 1989).

Although I have greatly amended and completed the original manuscript, or even written
many points in it, I tried to respect F. R. Bernard's personal opinions. However, in a few instances,
when I could not agree with his interpretation of the material reported in this paper, I have decided
against it. One of the new species described here (Policordia olivacea) is in this case.

J.-M. Poutiers

MATERIAL AND METHODS

The bivalve mollusc material reviewed in this paper, kindly provided by Dr P. Bouchet,
comprises carnivorous anomalodesmatan species usually assigned to the septibranchs. It originates
from several French expeditions organized in recent years in the Pacific Ocean, and comprises:

Source :

ANOMALODESMATA FROM THE TROPICAL PACIFIC

109

Material collected during two expeditions by mnhn and orstom to the central Philippines under
the direction of Prof. J. Forest (mnhn): musorstom 1 in 1976 aboard N.O. “Vauban" (Forest, 1981),
and musorstom 2 in 1980 aboard N.O. “ Coriolis " (Forest, 1986).

Material collected during the corindon 2 cruise organized in 1980 by orstom to the Straits of
Makassar between Borneo and Sulawesi (Indonesia) aboard N.O. “ Coriolis ”, with J. Forest
responsible for biological data.

Material collected by P. Bouchet and A. Waren off southern New Caledonia in 1978-79, aboard
N.O. " Vauban " (Richer de Forges, 1990).

Most of the finer residues from these expeditions was sorted by the Centre National de Tri
d’Oceanographie Biologique (centob, ifremer, Brest).

Unless otherwise stated, all material including holotypes is deposited in mnhn. Parts of the
material collected in New Caledonia and in the Philippines during musorstom 1 expedition have been
already investigated (Morton, 1987: Poutiers, 1981, 1982a, b, 1985).

A species from the USNM collections, dredged in 1909 by the '' Albatross ” during its
Philippines expedition (Bowers, 1910), has been added through the courtesy of the late Dr R. S.
Houbrick.

The material comprises 37 species taken at 69 stations at depths from 70 to 1628 m. A list of
stations, with main geographical and bathymetrical data and collected species, is presented in
Appendix 1. Illustrations, bathymetric and geographic distributional data and a brief systematic
description are given for each species taken. The known synonyms are listed, but no attempt has been
made to cite all subsequent usages.

The descriptive section of this paper is followed by a brief summary of the classification,
reflecting mainly F.R. Bernard’s personal opinion. This synopsis has been improved and updated
with as few modifications as possible, notably through the insertion of genus-group taxa introduced
in 1983 by Scarlato & Starobogatov. Short diagnoses are provided for the supraspecific Recent
taxa, with a listing of all the species belonging in each genus (or subgenus). A general list of the
binomina has also been included at the end of this section. The authors have attempted to make this
catalogue as comprehensive as possible, but it is likely to be incomplete. A free translation of the key
paper by Scarlato & Starobogatov (1983) referred above is provided in Appendix 2.

Since 1984. when the material reported in this paper was sent to F. R. Bernard for study, a number
of additional deep-sea expeditions have been led by the Paris Museum and orstom in the Philippines
(musorstom 3, 1985), in Indonesia (karubar, 1991), and mainly in the New Caledonian economic
zone (for a review of cruises in this area, see Richer de Forges, 1990, 1993). They have resulted in
many samples of carnivorous anomalodesmatan Bivalves which will form the subject of forthcoming
papers in which the first author will provide more personal views on septibranch classification.

ABBREVIATIONS AND TEXT CONVENTIONS

Repositories

AMS

MNHN

NMNZ

NSMT

USNM

: Australian Museum, Sydney

: Museum national d’Histoire naturelle. Paris
: Museum of New Zealand. Wellington
: National Science Museum, Tokyo
: National Museum of Natural History, Washington, dc

Other abbreviations

v

OD

db

spm(s)

paired valves, dead collected
live-taken specimen(s)
valve(s)

original designation.

110

JEAN-MAUR1CE POUTIERS & FRANK R. BERNARD

SPECIES ACCOUNTS

Family Verticordiidae

Spinosipella costeminens (Poutiers, 1981)
Figs 1-2

Verticordia (Spinosipella) costeminens Poutiers, 1981: 351, pi. 4, figs 1-4, text-lig.5.

Material examined. — Philippines. musorstom 1: stn 49, 13°49'N, 120°00.5' E, 750-925 m,
3 spms (holotype and paratypes).

musorstom 2: stn 55, 13°53.4' N, 19°57.8' E, 865-866 m, 3 spms, 3 db, 5 v.

Distribution. — Only known from the Central Philippines, in 750-925 m.

Description. — Shell inflated, rather large for the genus,
subquadrate, with spirally twisted umbones. With 16-17 thin,
produced, blade-like radial ribs, one of which is much more
prominent than the others and runs from umbo to the
posteroventral angle of shell. Exterior surface covered with
radial row's of small spines. Internal ligament short, arcuate.

reinforced by a large Iithodesma. Right valve with a strong,
conical, posteriorly recurved cardinal tooth. Left valve with a
small corresponding denticle. Interior pearly white, with the
external ribbing showing through. Margins deeply indented.
Length: 23.5 mm.

Remarks. — This elegant large species is distinguished by the blade-like shape of its radial ribs and
the prominent umbono-posterior keel.

Spinosipella deshuyesiana (P. Fischer, 1862)
Figs 7-9

Verticordia deshuyesiana P. Fischer, 1862: 35, pi. 5, figs 10-11 (Published 7 January 1862).

Synonym:

Verticordia japonica A. Adams, 1862: 224 (Published in March 1862).

Material examined. — Philippines, musorstom 2: stn 33, 13°32' N, 121°07.5' E, 130-137 m.
3 v. — Stn 51, 13°59.8' N, 120°17' E, 170-187 m, 1 v.

New Caledonia. “Vauban”\9T&-19\ stn 2, 22°17' S, 167° 14' E, 425-430 m, 2 v. — Stn 4, 22°17'S,
167° 13' E, 400 m. 3 v. — Stn 15, 22°49' S, 167° 12' E, 390-395 m, 3 v. — Stn 16, 22°46' S, 167° 12' E,
390-400 m, 2 v. — Stn 42, 22°08' S, 167°04' E, 230-260 m, 2 v.

Distribution. — Indo-Pacific, East Africa, Indonesia, Thailand, South and East China Sea,
Honshu, Japan to Hawaii and New Caledonia, in 40-693 m (Higo & Goto, 1993).

Description. — Shell solid, globose, rounded to ovate.
Umbones prominent, spirally twisted, overhanging a small,
deeply impressed lunule. Radial ribs strong and sharp,
covered on crests with small prickly granules; interspaces
with numerous, closely set, fine pustules more or less radially

aligned. Internal ligament reinforced by a strong Iithodesma.
Hinge of right valve with a large, conical, cardinal tooth. Left
valve with a smaller corresponding denticle. Interior na¬
creous. Inner margins deeply crenulated. Length: 16.1 mm.

Remarks. — The shell of this species is somewhat variable in rib number and general shape,
larger specimens tending to be proportionately higher. Its relationships to the Atlantic S. acuticostata
(Philippi, 1844) and the Indo-Pacific S. ericia (Hedley, 1911) have not yet been clearly elucidated. The
latter is said to have a wide distribution (Crozier, 1966) and to be smaller, rounder, with less

Source :

ANOMALODESMATA FROM THE TROPICAL PACIFIC

111

Figs 1-10. — Verticordiidae — 1-2, Spinosipella costeminens , musorstom 2: stn 55, L = 19.2 mm, exterior of left valve (1),
interior of right valve (2). — 3-6, Halicardia Iwubricki sp. nov., “Albatross"', stn 5582, holotype, L = 34.6 mm, interior
of left valve (3), exterior of right valve (4), exterior of left valve (5), interior of right valve (6). 7-9, Spinosipella

deshayesiana: 7-8. "Vauban" 1978-79: stn 42. L = 9.1 mm. exterior of left valve (7), L = 11.2 mm, interior of right
valve (8); 9. "Vauban" 1978-79: stn 4, L = 16.1 mm, exterior of right valve (9). 10, Haliris multicostata, musorstom

2: stn 33, L = 6.9 mm, exterior of right valve.

Source : MNHN, Paris

112

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

prominent umbones. However, these features seem rather mutable and may prove to fall within the
range of individual or ontogenetic variations.

Haliris multicost at a (A. Adams, 1862)
Fig. 10

Verlicordia multicostata A. Adams. 1862: 224.

Synonym:

Verlicordia (Haliris) moeshimaensis Habe. 1953: 133.

Material examined. — Philippines, musorstom 2: stn 33, 13°32' N, 121°07.5' E, 130-137 m.
5 v.

New Caledonia. “Vauban” 1978-79: stn 2, 22°17' S, 167°14' E. 425-430 m, 7 v. — Stn 40. 22°30' S.
1 66°24' E, 250-350 m. 5 v.

Distribution. — Western Pacific. New Caledonia, the Philippines. South and East China Sea
to Honshu, Japan, in 50-450 m.

Dkscription. Shell small, inflated, cordiform. Inequila¬
teral. with anterior, prosogyrate umbones and strongly
developed, shallow lunule. Sculpture of rounded, imbricated
or nodulose radial riblets and numerous fine granules.

Lithodesma thin, lamelliform. Right valve with a stout conical
cardinal tooth and a lateral ridge under posterodorsal
margin. Interior of valves subnacreous. Inner shell margin
regularly crenulale. Length: 6.9 mm.

Haliris teporis sp. nov.
(Figs 11-14)

Type material. — Holotype live taken, mnhn.

Type locality. — New Caledonia. " Vauban " 1978-79, stn 3, 22°17'S, 167° 12' E, 390 m.

Material examined. — Only known from the type material.
Distribution. — Only known from the type locality.

Description. Shell minute, oval subquadrate, rather
compressed, inequilateral. Umbones prosogyrate, not promi¬
nent. situated at approximately anterior one third of shell
length. Slightly inequivalve, left valve somewhat deeper and
larger, overlapping a little the opposite valve along margins.
Anterodorsal margin short, a little depressed along lunule,
somewhat curled up to meet the rounded anterior and ventral
margins. Posterodorsal margin elongate, oblique and feebly
convex, forming a blunt angle with the broadly arcuate
posterior margin. Sculpture of about 25 delicate radial riblets;
riblets and interspaces covered with tiny prickly granules.
Lunule small, relatively deep. Periostracum adherent, te¬
nuous. pale straw in colour, extending somewhat over the

inner shell margin. Internal ligament reinforced by a litho¬
desma. subquadrate in outline with slightly produced poste¬
rior angles. Right valve with a conical cardinal tooth and an
obscure lateral ridge under the posterodorsal margin. Hinge
of left valve edentate, with a small subumbonal indentation to
fit the opposite tooth. Interior polished, subnacreous, with
radial furrows and inner margin crenulations corresponding
to outer radial sculpture.

Measurements: Length 3.0 mm, height 2.8 mm, inflation
2.0 mm. The single specimen consists of a set of valves with
its lithodesma. The soft parts were sent to F. R. Bernard, but
are now missing.

Remarks. — This small species is distinct in outline from H. multicostata , which also occurs
in New Caledonia, and the delicate radial riblets of its outer surface are characteristic.

Etymology. — The specific name alludes to the habitat of the species. It is derived from the
Latin tepor, meaning lukewarm.

Source : MNHN , Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

113

Figs 11-22. — 11-14. Verticordiidae: Haliris teporis sp. nov., "Vauban” 1978-79: stn 3, holotype, L = 3.0 mm, exterior of left
valve (11), interior of right valve (12), interior of left valve (13), exterior of right valve (14). — 15-18, Lyonsiellidae:
Policordia olivacea sp. nov., corindon: stn 231, holotype, L = 15.3 mm, exterior of left valve (15), interior of right valve
(16), interior of left valve (17), exterior of right valve (18). — 19-22. Euciroidae 19-20. Euciroa crassa, musorstom
2: stn 68, L = 12.3 mm, exterior of right valve (19), L = 14.1 mm, interior of left valve (20). — 21-22, Euciroa
millegemmala, musorstom 2: stn 19, L = 15.3 mm, exterior of left vaive (21), interior of right valve (22).

Source : MNHN, Paris

114

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Halicardia houhricki sp. nov.

Figs 3-6

Type material. — Holotype empty shell, usnm 239020.

Type locality. — Northeastern Borneo, “ Albatross ”, stn 5582, 4°19.9'N, 118°58.6'E,
1628 m.

Material examined. Only known from the type material.
Distribution. — Only known from the type locality.

Description. — Shell large, solid, inflated, nearly as long
as high, irregularly trigonal in outline. Inequilateral, with the
anterior half of disc ventrally expanded and an alate,
somewhat compressed laterally, posterodorsal slope. Lunule
small, deeply impressed, overhung posteriorly by the umbo-
nes, more developed in right valve. Umbones prosogyrate,
inflated and moderately prominent, in front of midlength of
shell. Right valve tending to overlap the opposite valve, a
little along posterodorsal margin, more evidently on lunuiar
area, making the shell slightly inequivalve. External sculpture
of six strong radial undulations that scallop ventral margin,
and numerous, narrow, unequal radiating grooves. Concen¬
tric growth marks well impressed, giving a finely decussate
effect with the radial elements. Outer shell surface covered
with densely set small granules. Periostracum thin, adherent,
light beige in colour, slightly thicker on periphery of shell and
somewhat reinforcing the radiating grooves. External liga¬
ment a long, narrow brown band stretching along postero¬
dorsal and lunuiar margins. Internal ligament opisthodetic.

leaving in each valve a trigonal, elongate groove pointing
obliquely under umbo. Lithodesma large, asymmetrical.
Hinge feeble, nearly edentate: left valve slightly protruding on
middle of posterodorsal margin; cardinal tubercle of right
valve obsolete, reduced to a faint ridge of lunuiar margin
under umbo. Inner side of shell pearly white, smooth, largely
scalloped by the exterior radial undulations. Anterior adduc¬
tor scar distinct, semilunar, bordered internally by a slight
thickening of shell. Posterior adductor scar a little larger, less
impressed, roughly subquadrate, with a short dorsal expan¬
sion corresponding to posterior pedal retractor scar. Anterior
retractor scar deeply impressed, globular, hooked ventrally,
situated beneath lunuiar margin. Umbonal cavity with a very
small, ovate muscle scar under resilifer. Pallial line narrow,
not indented by a sinus, remote from ventral margin.

Measurements: Length 34.6 mm, height 35.3 mm, inflation
28.8 mm. The single specimen consists of a set of valves in
fresh condition. The lithodesma is unfortunately now mis¬
sing, but was observed by F. R. Bernard.

Remarks. — The new species closely resembles the North Eastern Pacific H. perplicata (Dali,
1890) but is readily distinguished by the straighter posterodorsal margin, the proportionately larger
lithodesma, the smaller dental tubercle of the right valve and the much finer surface granulations.

It is very distinct from the three other Recent Halicardia species previously known from the
North Pacific area: H. nipponensis Okutani, 1957, from Japan, and H. gouldi Dali. Bartsch & Rehder,
1938, from Hawaii, are characterized by radial flexures rather than distinct ribs; H. philippinensis
Poutiers, 1981, from the central Philippines, has a slender, much thinner shell with delicate external
sculpture.

Etymology. — The species is named in honour of the late Dr R. S. Houbrick of the
Smithsonian Institution, Washington, DC for his numerous and valuable contributions to malaco¬
logy.

Halicardia philippinensis Poutiers, 1981
Figs 81-82

Halicardia philippinensis Poutiers, 1981: 353, textfig. 6, pi. 4. figs 7-8.

Material examined. — Philippines, musorstom 1: stn 44, 13°46.5' N, 120°29.5' E, 592-610 m,
1 spm (holotype), 2 spms, 1 db (paratypes) (mnhn).

Distribution. — Only known from the Central Philippines, in 592-610 m.

Source : MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

115

Description. — Shell inflated, brittle, higher than long,
subequivalve, roughly trigonal in outline. Inequilateral, ven¬
tral margin anteriorly expanded, posterodorsal margin so¬
mewhat alate. Sculpture reduced, with a shallow depression
radiating to the slight sinuosity of posteroventral margin, and
with numerous radial grooves and densely set small granules.
Internal ligament opisthodetic, leaving in each valve a long

and narrow oblique groove. Lithodesma large, horseshoe-
shaped, with pointed posterior lobes. Right valve with a
posteriorly recurved cardinal tooth. Left valve edentate.
Interior nacreous, with the external sculpture showing
through posteriorly. Inner shell margins sharp and smooth.
Length: 21.3 mm.

Remarks. — Halicardia philippinensis is closely related to the Hawaiian species H. gouldi Dali,
Bartsch & Rehder, 1938. However, it differs from the latter by the higher shape with more rounded
outline, a posterodorsal expansion more weakly demarcated from disk, and an evenly convex anterior
margin, not flattened or depressed in the middle.

Family Lyonsiellidae

Policordia olivacea Poutiers, sp. nov.

Figs 15-18

Type material. — Holotype live taken, mnhn.

Type locality. — Indonesia, corindon, stn 231, 0°04.9' N, 119°47.8'E, 980-1080 m.

Material examined. Only known from the type material.
Distribution. Only known from the type locality.

Description. — Shell brittle, trapezoidal, rather compres¬
sed. Inequivalve, with a flexuous commissural plane and a
slightly larger but less inflated right valve. Inequilateral,
umbones anterior, prosogyrate, not very prominent. Antero-
dorsal margin rather short, abruptly curved; posterodorsal
margin long, slightly convex. Ventral margin forming a
narrowly rounded expansion. Posterior slope set off by a
slightly depressed radial undulation. Sculpture of 34-39 fine
radial striae with raised periostracal fringes, and many
concentric lines forming fine lirae on lateral slopes. Shell
translucent, whitish, with an oily iridescence, covered with a
thin adherent periostracum, pale olive-brown in colour.
Hinge margin thin, edentate, only slightly protruding under
and behind umbo of left valve. Anterodorsal margin covered
in both valves with a thickened outgrow of periostracum.
Ligament opisthodetic, with a strong, bifid, recurved litho¬
desma, obliquely inserted under posterodorsal margin of
valves. Interior nacreous, with the extemak sculpture showing
through. Attachment scars obscure.

Anatomy : Adductor muscles ovate, subequal, the posterior
one slightly larger. Mantle rather solid, marginally thickened.

extensively fused, with a small pedal opening. Entire free edge
of each mantle lobe with a series of squat sensory tubercles.
Exhalant opening with lateral tissue pads and five unbran¬
ched conical tentacles, three dorsally and two apart ventrally.
Inhalant aperture larger, with an extensive raptorial hood,
surrounded on either side by ten prominent, partially fused
tentacles. These are branched into seven lobes and densely
covered with microscopic pointed papillae. Two additional
pairs of small, simple, conical tentacles inserted obliquely on
outer edge of the ring of inhalant tentacles, one pair in the
interstices between the fourth and fifth tentacles, and another
pair between the sixth and seventh ones. Septum thin and
delicate; gills with inner and outer demibranch, the ascending
lamella of the outer demibranch absent. Gill filaments short.
Foot small, rather tongue-shaped, with a byssal groove but
no trace of a functional byssus. Mouth large, with overhan¬
ging lips; labial palps small.

Measurements: Length 15.3 mm, height 15.8 mm, inflation
9.2 mm. The single specimen consists of a set of valves with
preserved soft parts. The lithodesma is now missing, but was
observed by F. R. Bernard.

Remarks. — This new species shows marked similarities with Policordia densicostata (Locard,
1898) of the subtropical Atlantic, a rather variable form, difficult to separate from P. gemma (Verrill,
1880) and P. atlantica Allen & Turner, 1974 on shell characters alone. Considering there were some
doubt about the relationships between these three species, F. R. Bernard postulated that a variable
complex is represented in the Atlantic, and tentatively assigned the corindon specimen to P.
densicostata. However, after an examination of Locard's type specimen in mnhn, and the study of
material assigned to P. densicostata by Allen & Turner (1974: 478), I cannot agree with the opinion
of Bernard and consider the corindon specimen a distinct species [J.-M. P.].

1 16

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Policordia olivacea differs from P. densicosiata by its compressed shell (length/inflation ratio
1.66 instead of 1.30). a more protruding ventral margin giving a somewhat trigonal outline, less
prominent umbones. and a more conspicuous, light olive-brown periostracum (tenuous and pale erey
to beige in P. densicosiata). On anatomical grounds, both species are easily distinguishable by^th'e
mantle free margins (devoid of tubercles in P. densicosiata) and the tentacles surrounding the inhalant
aperture (P. densicosiata has two more branched tentacles but no additional simple tentacles on their
outer edge).

Policordia olivacea has a general resemblance to the West Pacific P. pilula (see Prashad. 1932 for
description and illustration of the shell), a much smaller, differently shaped species. It is likely that
P. media Okutani, 1962. is identical to P. pilula.

Etymology. The specific name is derived from the Latin oliva, olive, with reference to the
colour and oily iridescence of the new species.

Family Euciroidae

Euciroa crassa Thiele & Jaeckel, 1931
Figs 19-20

Euciroa crassa Thiele & Jaeckel. 1931: 248. pi. 10. fig. 130.
Synonym:

Euciroa cistagemma Kuroda, 1952: 14, pi. 1. figs 16-18.

Material examined. — Philippines, musorstom 2: stn 2. 14°00.5' N. I20°17.3' E. 184-186 m.

1 v. — Stn 6. 13°56.5'N, 120°21.5'E, 136-152 m, 1 v. — Stn 10. I4°00.7'N, 1 20° 18.2' E. 188-195 m.

1 v. Stn 17. 14"00.5' N. 120°17.8' E, 174-193 m, 1 v. Stn 26. 13°49' N. 120°50.3' E. 299-320 m.

5 v. — Stn 51. 13°59.8' N, 120° 17' E, 170-187 m, 5 v. — Stn 64. 14°00.8'N. 1 20” 18.6' E. 191-195 m,

1 v. Stn 68, 14°01.2'N, 1 20° 1 8.2' E. 195-199 m, 4 v. — Stn 71. 14°00.6' N, 120° 18 5' E 189-
197 m, 1 v. Stn 72, 14°00.4' N. I20°18.6'E. 182-197 m, 1 v.

Distribution. Indo-Pacific. off East Africa. Maldives area, the Philippines, South and East
China Sea to Honshu. Japan, in 136-1463 m.

Description. Shell thick, ralher small; umbones promi¬
nent. Outline variable but generally rounded-triangular.

Inequilateral, post-umbonal part of shell short, truncate.

Sculpture of numerous, nodulated or dorsally imbricated
radial riblets. set closer on the posterior slope, which is set off
by two shallow radial sulci. Primary riblets rather strong,
with weaker intercalated riblets. Ligament opisthodetic, in a

deep resilifer. Lithodesma strong, rectangular, with two short
posierior lobes. Right valve with a slurdy cardinal tooth. Left
valve with an anterior tubercle and a weak, remote posterior
lateral tooth. Interior nacreous, with a radial ridge running
from the umbonal cavity to the posteroventral marain. Inner
shell margins crenulate. Length 19.2 mm.

Euciroa ehurnea (Wood-Mason & Alcock, 1891)
Figs 23-24

Verticorclia (Euciroa) eburnea Wood-Mason & Alcock. 1891: 447, figs 14a-d.

Synonyms:

Verticordia optima G.B. Sowerby III, 1894a: 39. pi. S fig 3- 1894b- 82
Euciroa tlalli Pilsbry, 1911: 523.

Material examined. Philippines, musorstom I: stn 44, 13°46.5' N. 120°29 5' E 599-610 m
1 db. Stn 47, 13°41.5' N. 120°30' E. 685-757 m, 5 spins.

Source : MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

117

Figs 23-29. Euciroidae — 23-24. Euciroa eburnea, musorstom 2: stn 79, L = 38.6 mm, interior of right valve (23), exterior
of left valve (24). 25-29, Acreuciroa rostrata, musorstom 2: stn 75: 25-26. L = 32.8 mm (juvenile specimen): dorsal

view of shell (25), exterior of left valve (26); 27-28, L = 61.4 mm (mature specimen): exterior of left valve (27), interior
of right valve (28); 29, L = 63.5 mm (gerontic form): exterior of left valve.

Source :

118

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

MUSORSTOM 2: stn 50, 13°37.4' N, 120°33' E, 810-820 m, 1 v. — Stn 79, 13°44' N, 120°31 7' E 682-
770 m, 3 spms, 1 db, 10 v. — Stn 82, 13°47' N, 120°28.8' E, 550 m, 1 spm.

New Caledonia. “Vauban” 1978-79: stn 15, 22°49' S, 167°12'E, 390-395 m, 1 v.

Distribution. — Widely distributed in the Indo-Pacific region, from South Africa and the
Gulf of Aden to the Philippines, the South China Sea and New Caledonia, in 340-1473 m.

Description. — Shell rounded-ovate, subequivalve, with
umbones somewhat in front of midlength. Posterior slope set
off by shallow radial sulcus. Sculpture of weak radial cords
with rows of small spines and numerous microscopic nodules.
Ligament opisthodetic. in a deep resilifer. Lithodesma long,
slightly bifurcated. Right valve with a strong conical cardinal
tooth, articulating in a pit of the opposite valve, and a
shallow lateral ridge behind the resilifer. Left valve with
anterior and posterior dorsal margins each produced in a
tooth-like ridge, and with a small cardinal tooth edging the
resilifer. Interior of shell brilliantly nacreous, with minute
radial striae. Inner ventral margin finely crenulated. Length
38.6 mm.

Anatomy. Adductor muscles subequal. Mantle with large
pedal opening, three-lobed, but lobes narrow. Ventral half of
mantle much thickened. Taenioid muscles absent. Margins of
pedal opening with minute erect spicules. Pallial apertures
not developed into siphons, placed in a muscular plate,
consisting of a small rounded exhalant aperture and a narrow

fissure for the inhalant aperture. The apertures are surroun¬
ded by a row of simple conical tentacles. Foot laterally
compressed, pointed, without a byssal groove. Gills appres-
sed to the septum. Outer demibranch appearantly with
ascending lamella only. Mouth broad, overhung by the lips
that are produced into lateral bulbs, the “lateral sacs” of
Dale (1895b). Ventrally is a tongue-like process, the “central
posterior lappet" of Dall. A similar structure has been
reported in Lyonsiella formosa by Allen & Turner (1974)
and is probably connected with adaptations to macrophagy.
A similar appendage is also present in Pholadomya Candida
(Morton, 1980). Labial palps long and prominent, similar to
those of Pholadomya (Morton, 1980). Oesophagus develo¬
ped. Stomach large and globular. Two ducts to the digestive
diverticula. Mid-gut and style-sac conjoined. Rectum proba¬
bly penetrating the heart and passing over the posterior
adductor muscle. Anus not projecting. Hermaphroditic, with
superficial, arborescent testis.

Remarks. — This species is closely related to Euciroa pacifica (Dali, 1895b) from Hawaii. The
presence of some preserved specimens (musorstom 1: Stn 47) allowed the elucidation of the gross
anatomy and comparison with Dall's (1895b) description of the Hawaiian species.

Euciroa eburnea shows a number of interesting similarities to the Verticordiidae and the
Poromyidae. The presence of large labial palps separates it at family level. It shows a number of
adaptations to macrophagy, but lacks any vestige of the “valvular membrane” associated with the
inhalant siphon which can be extruded as a raptorial hood (Morton, 1981b, 1985a, c ). It is difficult
to visualize how the minute inhalant fissure could be readily expanded, but the thickening of the
mantle margins containing haemocoels and numerous muscle fibres may play a role. Mechanism of
predation in Euciroa could be similar to the method supposedly used by verticordiids to capture
bottom living preys (Morton, 1987).

Euciroa millegemmata Kuroda & Habe, 1952
Figs 21-22

Euciroa millegemmata Kuroda & Habe in Kuroda, 1952: 14-15 (footnote 1 a), pi. 1, figs 12-15.

Material examined. — Philippines, musorstom 2: stn 19, 14°00.6' N 120° 17 4' E 189-192 m
1 spm. — Stn 32, 13°40.5' N, 120°54' E, 192-220 m, 1 spm.

IS™JION- — Limited to Western Pacific area, the Philippines and China Sea to Honshu
Japan, in 100-365 m.

Description. — Shell small, ovate, inflated. Inequivalve,
nght valve deeper. Anterior margin rounded, posterior
margin slightly produced, obtusely angulate. Sculpture of
numerous, fine narrow radial striae and rows of pustules or
spines. Lunule deep, small. Ligament opisthodetic, in a deep
resilifer. Lithodesma robust, with two pointed posterior

lobes. Right valve with a small, conical cardinal tooth and a
shallow posterior lateral ridge merged into shell margin;
anterodorsal margin somewhat thickened internally. Dorsal
margins of left valve slightly produced anteriorly and poste¬
riorly. Interior of shell nacreous, with fine radial striae.
Length 17.5 mm.

Source . MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

119

Remarks. — This species has been placed without explanation in Acreuciroa by Bernard et
al. (1993).

Euciroa trapeza Poutiers, 1982
Figs 79-80

Euciroa trapeza Poutiers. 1982: 332-333, figs 1-2.

Material examined. — New Caledonia. " Vauban ” 1978-79: stn 14. 22°16'S. 167°17' E,
465-495 m, 6 v, fragments. — Stn 33, 22°33' S, 166°25' E, 290-350 m, 2 db (holotype). Stn 34,
22°32' S, 166°26' E, 350-420 m, 4 v. — Stn 39, 22°29' S, 166°23' E, 375-550 m, ldb, 6 v. — Stn 40,
22°30' S, 166°24' E, 250-350 m, 7 v. All paratypes: ams, mnhn, nmnz, nsmt.

Distribution. — West Pacific, off southern New Caledonia, in 250-550m.

Description. — Shell large, inflated, subequivalve, trans-
versally elongated. Inequilateral, anterior margin broadly
rounded, posterior margin rostrate, obtusely angulated and
strongly sloping dorsally. Sculpture of thin radial threads
fading out ventrally and posteriorly, and numerous irregular
rows of small prickly granules. Ligament opisthodetic, in a
deep resilifer. Lithodesma not known. Right valve with a

strong conical cardinal tooth, articulating in a recess of the
opposite valve, and a shallow lateral ridge behind the
resilifer. Left valve with anterior and posterior dorsal margins
each produced in a lateral ridge, and a weak tubercle in front
of the resilifer. Interior of shell brilliantly nacreous, finely
striated on margins. Length 58.0 mm.

Acreuciroa rostrata (Thiele & Jaeckel, 1931)

Figs 25-29

Euciroa (Acreuciroa) rostrata Thiele & Jaeckel, 1931: 249, pi. 10, figs 1 32- 1 32a.

Synonym:

Euciroa (Acreuciroa) teramachii Kuroda. 1952: 15, figs 19-20.

Material examined. Philippines, musorstom 1: stn 42, 13°54.5' N. 120°29' E, 379-407 m,
1 db. — Stn 43, 13°50' N, 120°28' E. 448-484 m, 1 db.

musorstom 2: stn 15, 13°55'N, 120°28.9' E, 326-330 m, 1 v. — Stn 75, 13°51.9'N, 120°30.1'E,
300-330 m, 4 spms, 4 db, 18 v. — Stn 78, 13°49.5'N, 120°28.5' E, 441-550 m, 2 v. — Stn 82, 13°47' N,
120°28.8' E, 550 in, 2 v. — Stn 83, 13°55.9' N, 120°30.5' E, 318-320 m, 4 v.

Distribution. — From Indonesia to the Philippines, South and East China Sea to Honshu,
Japan, in 200-550 m.

Description. — Shell large, rounded-ovate, with a well-
developed posterior rostrum. Sculpture of numerous, spinose,
intercalating radial riblets. tending to be more prominent
towards the anterior end of rostrum. Ligament opisthodetic.

broad, strong, heart-shaped. Right valve with a strong
cardinal tooth and a posterior lateral lamina. Left valve
edentate. Interior richly iridescent, finely striate on margins.
Length 66.8 mm.

Remarks. — The juveniles tend to be more elongate (Poutiers, 1981; Tsuchida, 1986).
Following Habe (1977, 1981) and Abbott & Dance (1982), the posteriorly produced Acreuciroa is
here considered a separate genus, distinct from the ovate and inflated species of Euciroa s. s.

Source .

120

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Family Cuspidariidae

Cuspidaria convexa Pelseneer, 1911
Figs 30-3 1

Cuspidaria convexa Pelseneer. 1911: 80. pi. 26. fig. 10.

Other reference:

Cuspidaria (Cuspidaria) convexa - Prashad. 1932: 329. pi. 7. tigs 36-37

Material examined. Philippines, musorstom I: stn 5, 14°01.5'N, 120°22' E. 200-215 m,

1 spm.

musorstom 2: stn 49, 13°38.8' N. 121°43.2' E. 416-425 m, 1 db.

Western Pacific area. East of Java, the Central Philippines and South China

Distribution.

Sea, in 100-694 m.

Description. Shell globular, thin and white, slightly
inequivalve, with a deeper left valve. Umbones prominent,
anteriorly situated: postumbonal part equals about 60% or
shell length. Anterior side of valves rounded, posterior drawn
out in a short, wide rostrum, poorly marked off from disc by
shallow radial depression. Posterodorsal area of rostrum with
a few indistinct radial threads. Sculpture of irregular concen¬

tric striae; surface with mucus-bound mud layer. Resilifer a
narrow, opisthogyrate groove almost parallel to dorsal
margin and not protruding under the hinge. Right valve with
a trigonal, elongate, prominent posterior lateral tooth. Left
valve edentate. Interior of shell milky white, glossy. Lengih
12.3 mm.

Remarks. - This species is very similar to the Indian Ocean C. approximate i E. A. Smith,
1896. but Knudsen (1967: 309) gave anatomical reasons to distinguish them. In C. approximate!, the
concentric sculpture is said lo be thinner, the rostrum seems to be more ventrally placed and has a
distinct lidge radiating from umbo to posteroventral end of shell. However, some of these characters
may be mutable, and nothing is known about the individual variation of shell in the two species.

Cuspidaria convexa is compared by Prashad to his C. mills, a less globular and more elongate
species with less protruding umbones.

Cuspidaria corrugata Prashad, 1932
Figs 32-33

Cuspidaria (Cuspidaria) cprrugata Prashad. 1932: 329, pi. 7. fig. 38.

Material examined. Philippines, musorstom 2: stn 26. 13"49' N. 120°50.3' E. 299-320 m.
1 db. Stn 72. 14°00.4' N, 120° 18.6' E, 182-197 m. 1 v. Stn 80. 13°45.2'N P0°37 5' E 178-
205 m, 2 v. ’

Distribution. Western Pacific, Indonesia and the Philippines, in 38-320 m

Description. Shell thin, ovate, shortly rostrate poste¬
riorly. Umbones not elevated, a little behind midlength of
shell. Dorsal margin strongly sloping on either side of
umbones. Ventral margin arcuate, curving up sharply in its
posterior half. Rostrum short, subcentral, tapered, with a
truncate end. set off by a broad radial depression that also
produces a sinuous ventral margin. Sculpture of strong
concentric riblets with wide interspaces, more regularly
placed anteriorly, somewhat sinuous and lamellose poste-
riorly. Concentric riblets ending on rostrum along a strong

oblique ridge radiating from umbones to the posteroventral
margin. Posterodorsal area of rostrum crowded with very
fine concentric threads, with a second, lower radial ridge
bordering dorsal margin of shell. Internal ligament opis-
thogyrate. in a narrow, oblique resilifer that is not produced.
Left valve edentate. Posterior denticle of right valve obsolete.
Posterior adductor scar well impressed, roughly trigonal,
bordered on umbonal margin by an internal reinforcement of
shell. Anterior adductor scar indistinct. Inner surface with
concentric undulations. Lengih 17.3 mm.

Source .

Figs 30-42. Cuspidariidae 30-31. Cuspidaria convexa, musorstom 2: sin 49. L = 12.3 mm. interior of right valve (30),
exterior of left valve (31). 32-33. Cuspidaria corrugaia, musorstom 2: stn 80. L = 16.5 mm. interior of left valve (32),

L = 17.3 mm, exterior of right valve (33). — 34. Cuspidaria gigantea, musorstom 2: stn 10. L = 33.7 mm, exterior
of right valve. 35-36, Cuspidaria hindsiana, musorstom 1: stn 71, L = 22.0 mm, interior of left valve (35). exterior
of right valve (36). 37, Cuspidaria japonica, musorstom 2: stn 11, L = 27.7 mm. exterior of right valve.

38-39, Cuspidaria kyushuensis, musorstom 2: stn 25. L = 12.7 mm, interior of right valve (38). exterior of left valve

(39) . 40-42. Cuspidaria morrisae sp. nov., "Vauban" 1978-79: stn 42, holotype, L = 20.0 mm, interior of left valve

(40) . exterior of left valve (41), interior of right valve (42).

Source . MNHN, Paris

122

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Remarks. The present material agrees well with Prashad's original description, although
his figure of the outside of shell shows a somewhat more centrally placed umbo and more irregular
concentric sculpture. However, the shape of musorstom 2 valves is rather variable, and theses
differences may represent individual variation. Prashad did not describe nor figure the hinge, stating
only it is “as in typical Cuspidaria ”, but in the right valves described here, the posterior tooth is
indistinct.

Cuspidaria gigantea Prashad, 1 932
Fig. 34

Cuspidaria (Cuspidaria) giganlea Prashad, 1932: 329, pi. 7, fig. 38.
Synonym:

Cuspidaria kawamurai Kuroda. 1948: 11, pi. 1, fig. 4.

Material examined. — Philippines, musorstom 1: stn 71, 14°09.5' N, 120°26.5' E, 174-204 m

1 spm.

musorstom 2: stn 10, 14°00.7' N, 120°18.2' E, 188-195 m, 1 v. — Stn 26, 13°49'N 120°50 3'E
299-320 m, 1 v. — Stn 51, 13°59.8' N, 120° 17' E, 170-187 m, 2 v.

Distribution. — Indo-Pacific, from Southeast Africa to the Philippines, South and East
China Sea to Honshu, Japan, in 100-1030 m.

Description. — Shell solid, subequivalve, oblique ovate,
strongly rostrate, very inequilateral. Umbones not promi¬
nent, feebly opisthogyrate. Rostrum long, narrow, usually
recurved, set off from disc by a broad radial depression that
deeply sinuates the ventral shell margin. Rostrum with an
inflated ridge radiating from umbones to the posterodorsal
margin, giving the appearance of an elongate escutcheon.

Sculpture of a concentric striation, becoming coarse and
irregular posteriorly in the depressed area of the posteroven-
tral sinuation. Periostracum thick, dehiscent. Internal liga¬
ment reinforced by a lithodesma, in vertical trigonal resilifer
that is strongly protruded under hinge margin. Right valve
with a strong posterior lateral tooth. Left valve edentate.
Interior of shell with fine radial striae. Length 33.7 mm.

Remarks. — Knudsen (1967: 314) synonymized Cuspidaria kawamurai with C. giganlea , but
used Kuroda s name on the basis that the specific name gigantea was preoccupied by Verrill (1884)
Barnard (1964: 579) also mentioned this homonymy, but did not give any replacement name for C
gigantea Prashad. However, Verrill's species was described under Neaera, and actually belongs to
Myonera (Verrill & Bush, 1898). Thus, the secondary homonymy detected by Barnard and
Knudsen doesn’t exist any more, and the specific name C. gigantea Prashad, 1932 remains.

Cuspidaria hindsiana (A. Adams, 1864)
Figs 35-36

Neaera hindsiana A. Adams, 1864: 207.

Material examined. —

1 spm.

Philippines, musorstom 1: stn 71, 14°09.5' N, 120°26.5' E, 174-204 m.

Distribution. - Northwestern Pacific, the Philippines, East China Sea to Taiwan and
Honshu, Japan, in 50-200 m.

Description. — Shell thin and white, inflated, elongate-
ovate, with a narrow central recurved rostrum. Ventral
margin broadly convex, with a weak posterior constriction at
base of rostrum. Sculpture of thin, irregular sharp concentric

ridges, and many very fine growth lines. Rostrum with
numerous transverse grooves and a few indistinct radial
threads. Periostracum thin, adherent, translucent. Internal
ligament with broad lithodesma, attached in each valve to a

Source .

ANOMALODESMATA FROM THE TROPICAL PACIFIC 123

trigonal resililer that is inclined posteriorly. Hinge of right Left valve edentate. Interior of shell with outer concentric
valve with a strong, elongate, trigonal posterior lateral tooth. ribbing showing through. Length 22.0 mm.

Cuspidaria japonica Kuroda, 1948
Fig. 37

Cuspidaria japonica Kuroda, 1948: 14. pi. 1, fig. 2.

Material examined. — Philippines, musorstom 2: stn 11, 14°00.3' N, 120° 19.3' E, 194-196 m
1 v. — Stn 68, 14°01.2'N, 120°18.2'E, 195-199 m. 1 db.

Distribution. — Northwest Pacific, from

Description. - Shell rather large, globose-ovate, with a
moderately developed, attenuate dorsal rostrum. Umbones
submedian, inflated but not very prominent, slightly opis-
thogyrate. Anterodorsal margin broadly convex, oblique.
Anterior and ventral margins rounded, the latter somewhat
sinuate posteriorly. Posterodorsal margin straightish to a
little recurved, gently sloping. Base of rostrum relatively
broad, with weak radial depression ending in the postero-

the Philippines to Honshu, Japan, in 100-200 m.

ventral sinuosity of margin. Sculpture of irregular concentric
striae, becoming coarser on rostrum. Periostracum thin,
adherent, beige in colour, somewhat darker and fibrous on
ventral margin and rostrum. Internal ligament reinforced by
a short lithodesma, in a small pit under and behind the umbo.
Hinge of right valve with a short posterior lateral tooth and
a small internal thickening of margin in front of umbo. Left
valve edentate. Interior of shell milky white. Length 27.7 mm.

Remarks. — This species is similar to C. lubangensis Poutiers, 1981, from the Central
Philippines, but the latter has a proportionately shorter rostrum, and more delicate hinge in the right
valve.

Cuspidaria kyushuensis Okutani, 1962
Figs 38-39

Cuspidaria kyushuensis Okutani, 1962: 35, pi. 3, fig. 12.

Material examined. — Philippines, musorstom 2: stn 25, 13H40'N, 120°43' E, 520-550 m.
1 db, 1 v.

Distribution. — Northwest Pacific, from Kyushu, Japan, to the central Philippines, in
520-760 m.

Description. — Shell solid, globose, quadrangular-ovate,
with a ventral enlargement and a rather short, compressed,
slightly upturned rostrum. Inequi valve, left valve somewhat
deeper and larger ventrally. Umbones small, inflated, anterior
to midlength of shell. Sculpture of rather regular concentric
lirae, disappearing on rostrum. Shell chalky white, with a
feebly glowing olive-yellow periostracum. Internal ligament
in a small, deep, oblique pit under umbo. Right valve with a

stout, trigonal posterior lateral tooth and a thickened
anterior hinge margin. Hinge of left valve edentate. Interior
of shell whitish, porcelaneous. Posterior adductor scar trigo¬
nal, deeply impressed. Anterior adductor scar elongate-ovate,
with a small pedal retractor scar near its posterodorsal end.
Pallial line hardly distinct, with a large sinus at base of
rostrum. Length 12.7 mm.

Remarks. — The present material agrees well with the short diagnosis and figure of C.
kyushuensis given by Okutani (1962). Although he compared it with C. circinata (Jeffreys, 1876), it
seems more closely related to the Atlantic C. ventricosa Verrill & Bush, 1898. In the latter species,
however, the sculpture is coarser and more closely spaced, and the posterior lateral tooth is closer to
the ligament pit.

Source :

124

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Cuspidaria lubangensis Poutiers. 1981
Fig. 83

Cuspidaria lubangensis Poutiers. 1981: 348. pi. 3. figs 4-5

Material examined. — Philippines, musorstom 1: stn 63, 14°00.5' N. I20°16'E, 191-195 m
(holotype db, mnhn).

Distribution. Only known from the central Philippines, in 191-195 m

Description. — Shell brittle, globose-ovate, with a mode¬
rately developed and rather broad dorsal rostrum. Umbones
inflated, on midlength of valves, nearly orthogyrate. Anterior
and ventral margins rounded, the former subangulate dor-
sally, the latter somewhat sinuate posteriorly. Posterodorsal
margin concave, dorsally recurved towards” posterior end.
Base of rostrum with a shallow radial depression ending in
the posteroventral sinuosity of margin. Sculpture of irregular.

concentric grooves and narrow wrinkles, the former more
developed posteriorly. Periostracum beige to brown, adhe¬
rent. fibrous, darker and coarser towards margins. Internal
ligament fitting in an oblique resilifer that is slightly protru¬
ding under hinge margin. Hinge of right valve with a shallow
posterior lateral tooth. Left valve edentate. Interior of shell
bright white in colour. Length 19.5 mm.

Remarks. — For comparison with Cuspidaria japonica Kuroda, 1948, see under that species

Cuspidaria macrorhynchus E. A. Smith. 1895
Figs 43-44

Cuspidaria macrorhynchus E. A. Smith. 1895: 12, pi. 2, figs 5-5a.

Material examined. Philippines, musorstom 2: stn 39, 13°08' N. 122°36 3' E 1030-1190 m
8 v. - Stn 44. 13°23.5' N, 122°20.6' E. 760-820 m, 2 spms, 3 db.

• aoo nooR,BUTI°N' Indo-Pacific, from East Africa to Indonesia, the Philippines and China Sea,

Description. — Shell thin, strongly inequilateral, ovate to
slightly oblique with a very long and narrow, tube-like
rostrum. Umbones prominent, in front of midlength of shell
Anterodorsal margin straight, horizontal, close to umbones
slightly convex and strongly sloping anteriorly. Anterior and
ventral margins rounded, the latter deeply sinuate at base of
rostrum. Posterodorsal margin straight, horizontal, someti¬
mes slightly recurved distally. Sculpture of a fine irregular
concentric striation, becoming coarser and transverse on
posterodorsal area of rostrum. An obsolete diagonal ridge
radiating from umbo to posteroventral extremity of rostrum.

Periostracum thin, pale ochre in colour, becoming thicker
and darker on periphery of disc and particularly on rosirum
Internal ligament with a short lithodesma, attached in each
valve to a minute, oblique resilifer. rounded ventrally and
pointing under umbo. Right valve with a prominent posterior
lateral tooth and an anterior lateral groove for the opposite
valve margin. Left valve edentate. Interior of shell glossy
white, with a very fine radial striation and one or two shallow
longitudinal ridges in dorsal half of rostrum Length
36.2 mm.

Remarks. I he shell of this species is somewhat variable, and the rostrum tends to be

from V I inkin' r^"ghtly retUrVnd l'P 011 °'der sPecimens- Cuspidaria sugamtmai Nomura. 1940,
rom Japan in 106-220 m. is usually synonymized with C. macrorhynchus, but has a shorter

rftfUm and Stronger dentition in the right valve. It is considered here as a
valid species limited to Japan.

Source :

Figs 43-54. Cuspidariidae - - 43-44. Cuspidaria macrorhynchus, musorstom 2: sin 44. L = 23.7 mm. exterior of left valve
(43). interior of right valve (44). - 45-47. Cuspidaria prolaiissima, musorstom 2: stn 80, L = 23.9 mm, interior of right
valve (45), exterior of left valve (46); L = 21.0 mm, dorsal view of shell (47). — 48-49. Cuspidaria steindachneri ,
musorstom 1: stn 63, L = 18.6 mm. exterior of left valve (48), interior of right valve (49). — 50. Myonera (Rengea)
caduca, “Vauban 1978-79: stn 42, L = 30.5 mm. exterior of left valve. 51-52. Myonera roslra sp. nov., musorstom
2: stn 40. holotype, L = 13.4 mm, interior of left valve (51), exterior of left valve (52). — 53-54. Myonera dautzenbergi,
CORindon: stn 280. L = 18.4 mm, exterior of left valve (53), interior of right valve (54).

Source : MNHN, Paris

126

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Cuspidaria morrisae sp. nov.
Figs 40-42

Type material. — Holotype db. mnhn.

Type locality. — New Caledonia .“Vauban” 1978-79, stn 42, 22°08' S, 167°04' E, 230-260 m.
Material examined. — Known only from the type material.

Distribution. — Known only from the type locality.

Description. — Shell solid, inflated, oblique ovate, ine¬
quilateral. Anterior side rounded, posterior side with a short
central rostrum, relatively narrow, abruptly truncate distally.
Inequivalve, left valve slightly deeper and commissural plane
somewhat distorted ventrally, a little drawn out to the left
anteriorly and to the right posteriorly. Umbones prominent,
opisthogyrate, well behind midlength of shell, the postumbo-
nal part forming about 42% of the total length. Anterodorsal
margin elongate, oblique, broadly arcuate, meeting rounded
anterior margin at an obtuse angle. Posterodorsal margin
concave, gently sloping. Ventral margin rounded, becoming
sinuous and oblique posteriorly. Rosfrum laterally compres^
sed, set off from disc by a radial depression, with a rounded

diagonal ridge running from umbo to the postero-ventral
margin. Sculpture of numerous irregular concentric lines and
grooves. Periostracum thin, translucent, yellow-ochre, stron¬
ger and folded ventrally. Internal ligament with broad,
arched lithodesma, attached to a shallow oblique resilifer
protruding a little beyond hinge margin. Hinge feeble. Right
valve with an obscure posterior lateral tooth. Left valve
edentate. Posterior adductor scar well impressed, kidney¬
shaped; anterior adductor scar indistinct. Interior of shell
porcelaneous white, with fine radial striae.

Measurements: Length 20.0 mm, height 12.3 mm, inflation
9.8 mm. The single specimen consists of a set of valves in
fresh condition.

Remarks. — This new species can be compared to C. capensis (E. A. Smith, 1885) from the
Southeast Atlantic, but the latter is rather more compressed, the umbones are less prominent, more
anteriorly placed and not strongly opisthogyrate.

Etymology. — The specific name is for Dr S. Morris formerly of the British Museum
(Natural History) in recognition of abundant and cheerful assistance.

Cuspidaria nobilis (A. Adams, 1864)
Fig. 61

Neaera nobilis A. Adams, 1864: 207.

Synomym:

Cuspidaria nobilis consimilis Habe, 1961: 146 & App. 42, pi. 65, fig. 21.

Material examined. — Philippines, musorstom 2: stn 19, 14°00.6' N, 120° 17.4' E, 189-192 m
1 db. — Stn 21. 14°01.2' N. 120°17.6' E, 191-192 m, 1 v. — Stn 51, 13°59.8' N, 120°17' E, 170-187
m, 2 v. — Stn 59, 14°00.4' N, 120°17' E, 186-190 m, 1 v. — Stn 68, 14°01.2' N, 120°18.2' E, 195-199
m, 1 spm, 7 v. — Stn 71, 14°00.6' N. 120°18.5' E, 189-197 m, 1 v. — Stn 80, 13°45.2' N, 120°37.5' E,
178-205 m, 2 v.

Indonesia, corindon: stn 267, 1°56.6' S, 119°16.7'E, 134-186 m, 1 spm.

Distribution. — Western Pacific, Indonesia, the Philippines, East China Sea to Honshu and
Shikoku, Japan, in 50-300 m.

Description. — Shell large, inflated, inequilateral,
elongate-ovate, with an attenuate, rostrate posterior end.
Umbones inclined posteriorly, situated a little in front of
midlength of shell. Anterodorsal margin convex, meeting

without discontinuity the rounded anterior margin. Rostrum
large, ventral, truncate, with a weak angle radiating to
posteroventral corner. Posterodorsal margin strongly recur¬
ved. Ventral margin arcuate, a little sinuate posteriorly by a

Source :

ANOMALODESMATA FROM THE TROPICAL PACIFIC

127

faint radial groove at base of rostrum. Sculpture of many
strong concentric ribs. Surface white, periostracum thin and
yellowish, often becoming stronger and darker on rostrum.
Resilifer broad, subvertical. Right valve with a low, triangular

posterior lateral tooth. Left valve edentate. Inner face of
shell undulated by the outer concentric sculpture. Length
51.7 mm.

Cuspidaria prolatissima Poutiers, 1981
Figs 45-47

Cuspidaria prolatissima Poutiers, 1981: 348, pi. 3, figs 2-3.

Material examined. — Philippines, musorstom 1: stn 25, 14°02.5' N, 120°22' E, 191-200 m.

1 spm. — Stn 31, 14°00'N, 120°17.5'E, 187-195 m, 1 spm. — Stn 42, 13°54.5'N, 120°29' E, 379-
407 m, 2 spms (holotype and paratype).

musorstom 2: stn 17, 14°00.5' N, 120°17.8' E, 174-193 m, 1 spm. — Stn 21, 14°01 2' N 120°17 6' E
191-192 m, 2 spms. — Stn 26, 13°49' N, 120°50.3' E, 299-320 m, 1 spm. — Stn 51, 13°59.8'N
120° 17' E, 170-187 m, 1 v. — Stn 63, 14H07.3' N, 120°15.5'E, 215-230 m, 1 spm. — Stn 64,
14°00.8'N, 120°18.6'E, 191-195 m, 1 spm, 3 v. — Stn 68, 14°01.2'N, 120°18.2' E, 195-199 m’

2 spms. — Stn 80, 13°45.2' N, 120°37.5' E, 178-205 m, 2 spms, 1 v. — Stn 83, 13°55.9' N, 120°30.5' E,
318-320 m, 1 spm, 2 v.

Distribution. — Fairly common in the central Philippines, in 170-407 m.

Description. — Shell equivalve, moderately inflated,
elongate-ovate, with a long, narrow, straight rostrum, some¬
times a little upturned posteriorly. Anterodorsal slope stron¬
gly depressed, set off by an oblique radial fold forming an
angle at anterior end. Sculpture of numerous irregular
concentric striae, becoming coarser and transverse on ros¬
trum, and overlain by shallow concentric ribs on umbonal

and anterior parts of the disc. Rostrum demarcated by a
radial sulcus, with a smoothish dorsal margin and an
umbonoventral ridge becoming indistinct posteriorly. Inter¬
nal ligament in a small trigonal socket beneath and behind
the umbo. Hinge plate narrow. Right valve with an elongate
posterior lateral tooth. Left valve edentate. Length 26.7 mm.

Remarks. — This elongate species is distinguished by the infolded area anterior to the umbo,
rather like a large lunule, and the consequently acute anterior shell margin.

Cuspidaria steindachneri Sturany, 1901
Figs 48-49

Cuspidaria steindachneri Sturany, 1901: 261, pi. 1, figs 5-9.

Synonym:

Cuspidaria hirasei Kuroda, 1948: 10, pi. I, fig. 3.

Material examined. — Philippines, musorstom 1: stn 63, 14°00.5' N, 120°16' E, 191-195 m,

1 db.

Distribution. — Indo-Pacific, Red Sea, Indian Ocean, the Philippines, South China Sea to
Honshu, Japan, in 106-1308 m.

Description. — Shell solid, elongate-ovate, with a long,
straight central rostrum. Umbones prominent, well anterior
to midlength of shell. Anterodorsal margin oblique, slightly
convex; posterodorsal margin concave. Ventral margin
broadly rounded, with a posterior sinuation. Rostrum de¬
marcated from disc by a shallow radial furrow. A fine ridge
running from umbo to posteroventral extremity of rostrum,
and becoming indistinct distally. Sculpture of irregular

concentric striae, coarser and recurved at a right angle on
posterodorsal area of rostrum. Internal ligament in a narrow,
oblique resilifer. Right valve with a strong posterior lateral
tooth and a distinct internal thickening of the anterodorsal
margin. Left valve edentate, hinge margin somewhat promi¬
nent anterior to umbo. Interior of shell with fine radial lines.
Length 18.6 mm.

128

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Cuspidaria (Soyomya) clathrata sp. nov.

Figs 62-65

Type material. — Holotype paired valves with dried soft parts, mnhn.

Type locality. — New Caledonia. “Vauban" 1978-79, stn 42. 22°08' S. 167°04' E, 230-260 m.

Material examined. — Only known from the type material.
Distribution. — Only known from the type locality.

Description. Shell solid, ovate, laterally compressed,
slightly inequivalve; posterior end with a short rostrum.
Umbones not very prominent, slightly prosogyrate and
submedian, the postumbonal part forming about 51% of
shell length. Anterodorsal margin horizontal just anterior to
umbones. gently convex and sloping towards rounded ante¬
rior margin. Posterodorsal margin oblique, straight, subtrun¬
cate posteriorly. Posterior truncation a little stronger on right
valve. Ventral margin broadly rounded, somewhat flattened
posteriorly. Rostrum broad, laterally compressed, largely
merged with disc, set off only by a shallow radial depression
that becomes less prominent ventrally. Surface with irregular
feeble concentric lirae and oblique rows of shallow pustules
in a divaricate pattern. Pustules developed on main part of

the disc, disappearing towards anterodorsal margin and on
umbonal and rostral regions. Periostracum thin, adherent,
translucent. Internal ligament light tan in colour, reinforced
ventrally by a broad, subquadrate lithodesma, attached in
each valve to a trigonal, posteriorly directed resilifer. Hinge
of right valve with a strong, oblique posterior lateral tooth
and a very small lamina just in front of resilifer. Left valve
edentate. Inner side of shell shiny, milky white, with a
shallow thickening in front of the posterior adductor scar and
a number of thin unequal lines radiating from the umbonal
cavity.

Measurements: Length 17.4 mm. height 12.5 mm. inflation
8.7 mm.

Remarks. — The external oblique sculpture of the new' species, recalling somewhat that of a
Strigilla Turton, 1822. is highly distinctive and warrants separation of the taxon at the subgeneric
level. It is here tentatively assigned to Soyoniva Okutani, 1985, and is similar to C. kitrohijii Okutani,
1972, from Japan, which is a more inflated species (inflation/length ratio 0.60-0.63, instead of 0.50
in C. clathrata), with more protruding umbones and a continuous sculpture resulting in a divaricate
appearance.

Etymology. The specific name is derived from the Latin clathratus, latticed, with reference
to the highly distinct sculpture.

Halonympha leiomyoides (Poutiers. 1981)
Figs 84-87

Cuspidaria leiomyoides Poutiers. 1981: 349-350. pi. 3. figs 6-7.

Material examined. — Philippines, musorstom I: stn 50. 13°49' N. 120°02' E. 415-510 m, 1
db (holotype). 1 v (paratype) (mnhn).

Distribution. — Only known from the type locality.

Description. - Shell globular, thin and translucent,
whitish, equivalve. Inequilateral, rounded anteriorly and
drawn out posteriorly in a short, trigonal blunt rostrum.
Sculpture of low concentric ridges that are more densely set
ventrally and fading out on posterior half of valves. Perios¬
tracum translucent and iridescent. Internal ligament fitting in
each valve in an oblique, ventrally protruding resilifer. Right

valve with a posteriorly recurved anterior cardinal tooth,
bordered on anterior and posterior dorsal margins by a faint
lateral ridge. Left valve with a small pointed posterior lateral
tooth. A raised oblique ridge under posterodorsal margin of
each valve. Interior of shell shiny, with an irregular tiny
granulation. Length 9.0 mm.

Source : MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

129

Figs 55-65. — Cuspidariidae 55-56, Cardiomya (Kurodamya) fortisculpta, Tosa Bay. Japan (mnhn, Coll. Staadt 1969),
L = 13.1 mm, dorsal view of right valve (55). exterior of left valve (56). — 57-58. Cardiomya alcocki, mijsorstom 2:
stn 25, L = 18.2 mm. dorsal view of right valve (57), exterior of left valve (58). 59-60, Cardiomya gouldiana.

musorstom 1: stn 58, L = 14.0 mm, interior of right valve (59), exterior of left valve (60). — 61. Cuspidaria nobilis,
MUSORSTOM 2: stn 71, L = 51.7 mm. exterior of left valve. 62-65, Cuspidaria (Soyomya) clathrata sp. nov., " Vauban "
1978-79: stn 42. holotype. L = 17.4 mm, interior of right valve (62), exterior of right valve (63). exterior of left valve
(64). interior of left valve (65).

Source : MNHN, Paris

130

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Cardiomya alcocki (E. A. Smith, 1894)
Figs 57-58

Cuspidaria alcocki E. A. Smith. 1894: 170. pi. 5. fig. 8.

Synonyms:

Cuspidaria /Cardiomya) polii Sturany, 1901: 264. pi. 1. figs 10-16.
Cuspidaria (Cardiomya) persculpta Prashad. 1932: 332. pi. 7, fig. 44.
Cuspidaria (Cardiomya) mullicarinata Prashad. 1932: 332. pi. 7~ figs 45-46.

Material examined. - Philippines, musorstom 2: stn 25. 13°39.5' N. 120°42.9' E. 520-550 m

1 db.

Indonesia, corindon: stn 281. 1°59'S. 119°09.9'1

Distribution. — Indo-Pacific, Red Sea.
60-1150 m.

Description. Shell thin, fragile, ovate, inflated, sube-
quivalve; left valve slightly larger, more inflated than right.
Rostrum short, recurved, with obscure radial riblets. Sculp¬
ture of disc with radial riblets increasing in number towards
ventral margin, and a few strong radial ribs on posterior

715-800 m, 1 db.

northern Indian Ocean and Southeast Asia, in

slope. Concentric ornementation of numerous fine lirae.
Resilifer narrow, subvertical. Right valve with a strong
elongate posterior lateral tooth. Interior with main external
ribbing showing through. Length 22.5 mm.

Remarks. — Cardiomya alcocki is usually considered a rather variable species and exhibits a
somewhat discrepant radial sculpture on the disc. Knudsen (1967). following Melvill & Standen’s
(1907) suggestion, synonymized the Red Sea species C. potli under Smith’s name. Habf. (1964. 1977,
1981) merged with C. alcocki the Japanese C. fordsculpta (Kuroda, 1948) and the Southeast" Asian
C. persculpta . C. mullicarinata and C. semicostata (all of Prashad, 1932). However, C. fortisculpta,
erected as the type of subgenus Kurodamya by Okutani & Sakurai (1964: 25), can be considered
a valid species restricted to Japan. It is easily distinguishable from the above-mentioned forms by its
sculpture being completely devoid of radial elements on the anterior slope, and by its especially short,
strongly protruding posterior lateral tooth in the right valve. A specimen from Tosa Bay (Shikoku)
is figured here for comparison (Figs 55-56). Following Okutani & Sakurai (1964: 26), C.
semicostata is a distinct species also referable to Kurodamya ; it is characterized by its very short
rostrum and radial sculpture restricted to the posterior third of shell.

Cardiomya gouldiana (Hinds, 1843)

Figs 59-60

Neaera gouldiana Hinds. 1843: 77.

Synonyms:

Cuspidaria (Cardiomya) gouldiana septentrionalis Kuroda, 1948: 18 pi 2 fie 12
Cardiomya lindbergi Scarlato, 1972: 125, figs 14-16.

Cardiomya lindbergi batialis Scarlato. 1972: 126, figs 17-19.

Material examined. — Philippines, musorstom I: stn 58, 13°58.5'N, 120°14' E, 143-178 m,

1 db.

,, , D|STRIBUTI0N- — Indonesia, the Philippines, South China Sea. Yellow Sea and Japan Sea, in
13-1030 m.

Source :

ANOMALODESMATA FROM THE TROPICAL PACIFIC

131

Description. — Shell small, obliquely ovate, inequivalve;
left valve somewhat larger, more inflated. Rostrum rather
short. central, slightly recurved, with concentric lirae. Sculp¬
ture of a dozen or more strong, sharp radial ribs, slightly

more developed in the right valve. Interspaces of ribs with
regular concentric lirae. Resilifer small. Right valve with a
strong, triangular posterior lateral tooth. Length 14.0 mm.

Remarks. — This species is similar to C. singaporensis (Hinds, 1843) but is proportionately
thinner, with less tumid umbones and more delicate radial ridges.

My oner a dautzenbergi Prashad. 1932
Figs 53-54

Myonera dautzenbergi Prashad, 1932: 334. pi. 7, fig. 51.

Material examined. — Philippines, musorstom 2: stn 81, 13°35.3'N, 121° 01.3' E, 856-
884 m, 4 spins.

Indonesia, corindon: stn 280, 1059' S, 1 19°09.9' E, 715-800 m, 4 spins, 3 db, 3 v.

Distribution. — Indo-Pacific, from southern Indonesia to the Philippines and Japan
(Okutani, 1968, 1976), in 715-959 m.

Description. Shell relatively large, thin, inflated ovate,
inequilateral. Left valve slightly larger, deeper, overlapping
right along ventral margin. Umbones prominent, incurved,
pointing behind midlength of shell. Anterior margin rounded.
Rostrum broad, short, slightly recurved dorsally, compres¬
sed, set off from disc by a well-marked radial fold. Sculpture
of numerous fine concentric striae, some raised in low

undulations anteriorly. Surface of rostrum quite smooth.
Periostracum adherent, straw-coloured, somewhat thickened
towards margins, wrinkled on posterior end. Internal liga¬
ment oblique, with a thin lithodesma. Resilifer narrow,
posteriorly directed. Interior of valves milky while, with the
external sculpture showing through. Length 20.8 mm.

Remarks. — This species has a somewhat variable outline and shows a slight allometry of
growth, the height/length ratio tending to increase in large specimens.

Contrary to Habe’s opinion (1981: 193, 195), Myonera dautzenbergi is not identical with M.
dispar (Dali, Bartsch & Rehder, 1938), which has two radiating keels between the rostrum and the
disc, and strong concentric undulations extending over the whole disc. Rather curiously, Bernard el
al. (1993: 1 18-119) repeated Habe’s error and recorded M. dautzenbergi under two different generic
names (namely Cuspidaria and Myonera). They also gave different synonymic and biogeographical
data for these two denominations of the same species.

Myonera rostra sp. nov.

Figs 51-52

Type material. — Holotype left valve, mnhn.

Type locality. — Philippines, musorstom 2, stn 40. 13°08' N. 122°40.2' E, 280-440 m.

Material examined. — Only known from the type material.

Distribution. Only known from the type locality.

Description. Shell extremely thin, inflated, triangular,
inequilateral, the postumbonal part forming about 60% of
total length. Umbo large, rounded, prominent. Anterodorsal
margin slightly convex, abruptly sloping towards the rounded
anterior margin. Posterodorsal margin oblique, somewhat

flexuous. Ventral margin broadly convex, with a slight
sinuosity on its rostral side. Rostrum broad, compressed,
wedge-shaped, rounded posteriorly. Rostrum set off from
disc by a strong, narrow radial ridge. Sculpture of the disc of
strong, distant concentric ribs, fading out towards the

132

JEAN-MAURICE POUTIF.RS & PRANK R. BERNARD

anterodorsa! margin and extending posteriorly to the umbo-
noventral ridge. Rostrum with a wide sulcus bordering the
radial ridge, well marked towards the umbo, shallower near
ventral margin. Surface of shell polished, with many fine
concentric lines, translucent white under an adherent, straw-
coloured periostracum. Hinge edentate, thin shelled, slightly
upturned under the umbo, with a well-developed ligament in
a long, narrow, oblique furrow. Interior of shell glossy, with
the external sculpture showing through. Muscle scars indis¬

tinct. Margins smooth, apart from the shallow flexure on
ventral side of rostrum.

Measurements: Length 13.4 mm: height 10.9 mm; inflation
(one valve) 4.8 mm. The single specimen consisted of a set of
valves with remains of soft parts. The right valve was
fragmented according to F. R. Bernard and has been subse¬
quently lost, so that the description and measurements are
based only on the remaining left valve.

Remarks. — This new species is very similar to M. garret ti Dali, 1908, and M. mexicana
Knudsen, 1970. two closely related eastern Pacific species. However, both American species are
devoid of the radial furrow bordering the umbonoventral ridge of rostrum and are less inequilateral.
Myonera mexicana is also more elongate, with a height/length ratio of 0.70 (fide Knudsen, 1970; 135)
instead of 0.81 in the present species. In M. garret ti, the concentric ribs do not reach the
umbonoventral ridge, are indistinct on the central part of the disc (as shown in Bernard, 1974: pi. 19
fig- 1 ) and less numerous for a same shell length, whereas in M. rostra they are continuous and more
numerous (about 24 in number instead of 15 in M. garretti).

Myonera rostra bears a general resemblance to Cuspidaria undata (Verrill, 1884) of the Atlantic
and Indian oceans. However, the latter is easily distinguishable by a more rounded outline, hinge
teeth on right valve (Knudsen, 1970: 137; Poutiers, 1984: 288) and the absence of radial ridge
between rostrum and disc.

The shell erroneously figured by Habe (1981: pi. 8, fig. 4) as M. daulzenbergi is in fact another
species identical or very similar to M. rostra.

Etymology. — The specific name alludes to the well-developed rostral side of the shell.

Myonera (Rengea) caduca (E. A. Smith, 1894)
Fig. 50

Cuspidaria (Myonera) caduca E. A. Smith. 1894: 170. pi. 5. figs 9-10.

Synonym:

Myonera fluctuosa Kuroda, 1948: 25. pi. 2, fig. 20.

Material examined. — New Caledonia. " Vauban " 1978-79: stn 42, 22°08' S 1 67°04' E
230-260 m, 1 v.

Distribution. - Indo-Pacific, from Southeast Africa to New Caledonia and South China Sea
to Honshu, Japan, in 50-1134 m.

Description. Shell thin, white, inequilateral, elongate-
ovate, with a long, broad, ventral, laterally compressed
rostrum. Umbones not prominent, posteriorly inclined, in
front of midlength of shell. Anterodorsal margin rounded,
posterodorsal margin recurved. Ventral margin broadly ar¬
cuated. slightly sinuate posteriorly. Rostrum' with two fine

ridges diverging from umbo to the truncate posterior end of
shell. Sculpture of irregularly concentric ridges and lirae.
Resilifer subvertical, protruding under umbo.'Hinge feeble,
edentate. Outer concentric wavy sculpture visible from the
inner side of shell. Length 30.5 mm.

Remarks. - Bernard et al. (1993: 118-119) erroneously recorded Myonera caduca twice in
their catalogue of the living marine bivalves of China; once under Cuspidaria and once under Rengea.
Under these two denominations of the same species, they provided different bathymetric, substrate,
and geographic distributional data.

Source : MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

133

Family Poromyidae

Poromya (Cetomya) butoni (Prashad, 1932)
Figs 77-78

Poromya (Cetoconcha) butoni Prashad. 1932: 327, pi. 7. figs 33-34.

Material examined. — Philippines, musorstom 1: stn 26, 14°00'N 120°17'E 189 m
1 spm. — Stn 31, 14‘W N, 120°17.5' E, 187-195 m, 1 db. - Stn 34, 13°59.5' N, 120° 17 5' E 188-
191 m, 1 db. — Stn 61, 14°01' N, 120°17.5' E, 124-129 m, 1 db. — Stn 72 14°12 5' N 120°29' E 12?-
127 m, 1 spm. — Stn 73, 14°16' NT 120°31.5' E, 70-76 m, 1 db

musorstom 2: stn 19, 14°00.6' N, 120° 17.4' E, 189-192 m, 1 spm, 1 db. — Stn 26 13°49'N
120°50.3'E, 299-320 m, 1 db. — Stn 59, 14°00.4'N, 120° 17' E, 186-190 m, 1 spm — Stn 72
14°00.4'N, 120° 18.6' E, 182- 197 m, 1 v. P ’

Distribution. — Western Pacific, Indonesia and the Philippines, in 70-535 m

Description. Shell rather small, whitish to hyaline,
inequilateral, slightly rostrate, subtruncate posteriorly. Ine-
quivalve, right valve somewhat overlapping left at margins.
Umbones prominent, situated in front of midlength of shell.
Surface iridescent, sculpture of many rows of minute granu¬

les, mostly retained on shell periphery. Ligament mainly
external, stretching along posterodorsal margin. Hinge feeble,
a little strengthened under umbones, completely edentate in
both valves. Interior glossy, with faint concentric growth
undulations. Length 13.0 mm.

Remarks. — Poromya butoni is completely edentate and has no trace of a cardinal tooth in
right valve. The observation [by J.-M. P.] of the soft parts of a preserved specimen in mnhn
(musorstom 1: stn 26) has shown that, in this species, the septum has two pairs of ostial openings.
Then. P. butoni is not a Cetoconcha and instead belongs in the subgenus Cetomya. This doesn't brina
P. (Cetomya) butoni in secondary homonymy with P. (Dermatomva) buttoni (Dali, 1916b) under
Article 58 of the Code of Nomenclature (dealing with single or double consonants), because the two
names are evidently not of the same origin and meaning: the epithet butoni is derived from Buton
Strait, Banda Sea (the type locality of Prashad’s species); the eastern Pacific buttoni , is named in
honour of Mr Fred. L. Button, as indicated by Dall (1900a: 321; 1916a: 5, 22). Thus, Article 57f
is applicable, and the two names are not homonyms.

Poromya ( Cetomya) exirnia (Pelseneer, 1911)
Figs 74-75

Poromya exirnia Pelseneer. 1911: 78. pi. 26, figs 3-4.

Others references:

Poromya ( Cetoconcha ) exirnia - Prashad, 1932: 327, pi. 7, figs 31-32.
Cetoconcha (exirnia var?) intermedia - Habe, 1952b: 158, pi. 21. figs 18-19.

Material examined. — Philippines, musorstom 1: stn 50, 13°49'N, 1 20°02' E, 415-510 m,
1 v.

musorstom 2: stn 25, 13°39.5' N, 120°42.9' E, 520-550 m, 3 spms. — Stn 39, 13°08'N, 122°36.3' E,
1030-1190 m, 1 spm. — Stn 44. 13°23.5' N, 122°20.6' E, 760-820 m, 1 db, 2 v. — Stn 46, 13°26.2' N,
1 22° 17.3' E, 445-520 m, 1 db. — Stn 75, 13°51.9'N, 120°30.1'E, 300-330 m, 1 v. — Stn 78,
13°49.5' N, 120°28.5' E, 441-550 m, 2 spms. — Stn 82, 13°47' N, 120°28.8' E, 550 m, 1 v.

134

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Figs 66-78. — Poromyidae 66-69 Cetoconcha boucheti sp. nov., musorstom 2: stn 75. holotype. L = 17.7 mm. exterior ol'
left valve (66), interior of right valve (67), interior of left valve (68). exterior of right valve (69). - 70-71 Cetoconcha
gloriosa, corindon: stn 208. L = 21.1 mm, exterior of left valve (70), interior of right valve (71). 72-73. Cetoconcha

ex‘gx‘1 sp. nov musorstom 2: stn 56. holotype. L = 6.0 mm, exterior of left valve (72), interior of right valve
r, r, , ,4 J<,ron'ya (Cetomya) eximia, musorstom 2: stn 25. L = 15.4 mm. interior of right valve (74): exterior

ol leit valve (75). 76. Cetoconcha japonica, " Vauban ” 1978-79: stn 9, L = 7.9 mm. exterior of right valve. - 77-78

Poromya ( Cetomya) hutom, musorstom 1: stn 26. L = 1 1.0 mm, exterior of left valve (77). interior of right valve (78)'

Source . MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

135

~ Distribution. Indo-Pacific, off East Africa, Flores Sea, the Philippines and South China
Sea to Honshu Japan, in 300-1190 m. Poromya intermedia (Habe, 1952) is said to be a local form
of this species, living in Japan in 50-200 m.

Description. — Shell inequilateral, highly inflated,
rhomboidal-ovate, with a short, broad posterior rostration.
Right valve slightly overlapping left along ventral and
posterodorsal margins. Umbones much inflated, prominent
strongly incurved, prosogyrate. Posterior side laterally
compressed, with a shallow, indistinct depression radiating
from umbo to posteroventral margin, where it appears as a
faint sinus. A small ridge, more distinct on the right valve,
extending along posterodorsal margin. Sculpture of concen¬

tric growth lines and striae, and easily eroded radial rows of
granules, most developed posteriorly. Shell whitish under a
pale dull brown periostracum. becoming irregularly wrinkled
postero vent rally. Ligament short, visible from the outside
Hinge margin narrow, slightly upturned, thickened under
umbones. Left valve edentate. Cardinal tubercle of right valve
reduced to absent. Interior of shell pearly white, smooth, with
a rounded ridge corresponding to the eternal umbonoventral
depression. Length 15.7 mm.

Remarks. — This species is well characterized by its plump shape and broad posterior
rostration. It is somewhat variable in outline and hinge features, with a more or less expanded
posterior rostration and a sometimes completely edentate right valve.

u C^trary to the opinion of Bernard et al. ( 1 993), which followed that of Prashad ( 1 932) and
Habe (1952b 1977, 1981), P. exuma cannot belong to Cetoconcha, as it has only two pairs of septal
openings with crossed filaments and interfilaments. Those features of the septum, investigated bv
Pelseneer (1911: 78) and Knudsen (1967: 305), have been also observed on a dried specimen of the
present material (musorstom 2: stn 25). Considering its ligament and hinge characters, P eximia is
best refered to the subgenus Cetomya.

Cetoconcha houcheti sp. nov.
Figs 66-69

Type material. Holotype db and 1 paratype v, mnhn.

Type locality. — Philippines, musorstom 2, stn 75, 13°51.9'N, 120°30.1'E, 300-330 m
Material examined. — Only known from the type material.

Distribution. — Only known from the type locality.

Description. - Shell thin, hyaline, inflated, trigonal-
ovate, subequivalve. right valve slightly deeper. Strongly
inequilateral, rounded anteriorly, produced and rostrate
posteriorly. Umbones slightly prosogyrate. about midlenuth
of shell, not very prominent. Anterodorsal margin oblique,
regularly rounded; posterodorsal margin sloping, nearly
straight, abruptly truncate at posterior end. Ventral margin
broadly convex, a little more arcuate near the posterior
truncation. Posterior slope of shell laterally compressed,
giving a gently concave outline in dorsal view. Sculpture of
concentric lines and striae, and of densely set pustules in
irregular concentric and radial rows, most abundant near
lateral and ventral margins. In addition, there is a fine

umbonoventral radial cord, reinforced by periostracal exten¬
sions near posteroventral shell margin. Outer surface oily
iridescent. Periostracum adherent, light ochre in colour,
becoming darker on periphery of valves. Ligament mainly
external, long and narrow, supported by a small thickening of
hinge margin under the umbones. Hinge feeble. Left valve
edentate, with slightly protruding anterior and posterior
margins. Right valve with a small, erect, cardinal denticle.
Interior polished, iridescent, with minute radial striae and
low concentric growth undulations. Muscle scars obscure.

Measurements: Length 17.7 mm. height 14.5 mm. inflation
1 1.3 mm.

Remarks. In the left valve of the holotype, a small transverse ridge can be seen on inner
side of the anterodorsal margin, at about one-third the distance from umbo to the anterior end. This
does not appear on the paratype, and must be considered merely as an individual variation.

From the other species of the genus, C. boucheti is distinguished by its characteristic outline
recalling somewhat that of a Macoma, with rounded anterior half and asymmetrically attenuated
posterior. It differs from C. exigua by its much larger size, a different posterior shape and more
crowded rows of pustules.

Etymology. — The specific epithet is named for Dr P. Bouchet.

Source .

136

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Cetoconcha exigua sp. nov.
Figs 72-73

Type material. — Holotype paired valves with traces of dried soft parts, mnhn.
Type locality. — Philippines, musorstom 2, stn 56, 13°54.1'N, 119°56.7'E, 970 m
Material examined. — Only known from the type material.

Distribution. — Only known from the type locality.

Description. — Shell small, thin, hyaline, ovate, subtrun¬
cate posteriorly. Umbones prosogyrate. anterior to midlength
of shell, prominent. Anterodorsal margin steeply sloping,
evenly curved. Anterior and ventral margins rounded, the
latter becoming somewhat flattened posteriorly. Posterodor-
sal margin oblique, shallowly curved, ending in a deep
posterior truncation. Outer surface moderately inflated, with
an obscure rounded keel extending from umbo to postero-
ventral end and delimiting a slightly depressed, triangular

posterodorsal slope. Sculpture of radial rows of fine pustules,
more prominent posteriorly. Periostracum tenuous but adhe¬
rent. translucent, shiny pale yellow in colour. Ligament
mainly external, long and narrow, supported by a small
thickening of hinge margin under and behind the umbones.
Hinge feeble, edentate in left valve, with a small cardinal
tubercle in right valve. Interior smooth, faintly iridescent.

Measurements: Length 6.0 mm. height 5.4'mm. inflation
4.1 mm.

Remarks. By its relatively high and rostrate outline, C. exigua somewhat recalls Poromya
eximia , but it has a much less inflated shape and sparser pustules on the outer surface. Its cardinal
tubercle is also more prominent than that of P. eximia. It resembles the Atlantic C. braziliensis Allen
& Morgan, 1981, which has a similar shape but no umbonoposterior ridge, rays of granules restricted
to the posterior region, a less sharply truncate posterior margin, and a completely edentulous hinge
in the right valve.

Etymology. The specific name is derived from the Latin exiguus, meaning small and

scanty.

Cetoconcha gloriosa (Prashad, 1932)
Figs 70-71

Poromya (Cetoconcha) gloriosa Prashad, 1932: 326. pi. 7. figs 29-30,

Material examined. — Indonesia, corindon: stn 208, 0°14.6' S, 117°52' E, 150 m, 3 spms

Distribution. — Indonesia, East of Flores and Makassar Strait, New Guinea and South
China Sea (Bernard el a/., 1993), in 150-400 m.

Description. — Shell thin, rather elongate, elliptical-
ovate, with a widely rostrate, somewhat compressed posterior
end. Subequivalve, right valve slightly larger than left.
Umbones subcentral, prosogyrate, inflated, prominent.
Sculpture of concentric striae and numerous rows of minute

granules, denser on rostrum. Periostracum thin, adherent,
light straw in colour. Ligament opisthodetic, mainly external!
Hinge feeble, edentate in left valve, with a small cardinal
denticle in front of ligament in right valve. Interior glossy,
somewhat pearly. Length 21.1 mm.

Remarks. — The present material corresponds rather well with the original diagnosis of C.
gloriosa except for the shape of the rostral region, which appears less attenuate than on Prashad’s
ngures. It is thus with some misgiving that these specimens are identified with Prashad’s species.

A complete but dead shell of C. gloriosa has also been reported by Kilburn (1973: 577) from
‘ atal waters, in 35 m depth. However, description of its hinge leaves doubt about the identity of this
torm and suggests it belongs to a different species.

Source .

ANOMALODESMATA FROM THE TROPICAL PACIFIC

137

Figs 79-87. 79-80. Euciroidae: Euciroa irapeza. ” Vauban " 1978-79: stn 40. paratype. L = 42.0 mm. exterior of left valve

(79), interior of right valve (80). 81-82. Verticordiidae: Halicardia philippinensis, musorstom 1: stn 44, L = 16.2 mm.

interior of right valve (81), exterior of left valve (82). 83. Cuspidariidae: Cuspidaria lubangensis, musorstom 1: stn

63. holotype. L = 19.5 mm. exterior of right valve. — 84-87. Cuspidariidae: Halonympha leiomyoides, musorstom I:
stn 50. holotype. L = 9.0 mm. exterior of left valve (84), interior of left valve (85), interior of right valve (86), exterior
of right valve (87).

138

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Cetoconcha japonica Ha be, 1952
Fig. 76

Cetoconcha japonica Habe. 1952b: 159. pi. 22, figs 2-4.

Material examined. — New Caledonia. “Vauban” 1978-79: stn 9, 22°20' S, 167° 10' E.
175-200 m, 1 v.

Distribution. — Japan, Hokkaido to Shikoku, in 200-650 m (Habe, 1977); also in Izu
Peninsula, Honshu, in 60-120 m (Okutani & Matsukuma, 1982). New Caledonia in 175-200 m
(dead).

Description. — Shell ovate to elongate. Umbones slightly
in front of midlength of valves. Translucent, thin, fraizile at
juvenile stages; becoming thicker, covered with a velvety,
brownish periostracum and attaining a rather large size for

the genus (up to 37.5 mm length, fide Habe. 1964). Outer
surface with numerous minute pustules. A small cardinal
tooth present in front of ligament in right valve.

Remarks. — It is with some misgiving that this small damaged right valve (length: 7.9 mm)
is identified with C. japonica. Cetoconcha japonica was first noted as Cetoconcha sp. (Habe, 1952a:
274) and described as a new species later in the same year. Anatomy is not known.

CLASSIFICATION

GENERAL REMARKS

Jhe S0',(fa!Led sePt*branchiate bivalves, currently assigned to the Anomalodesmata (Keen.
1969; Habe, 1977; Vokes, 1980; Boss, 1982) or to several orders (Nevesskaia et al. 1971-
Starobogatov, 1977, 1992; Scarlato & Starobogatov, 1978, 1979, 1983, 1985; this paper) are. as
ar as known exclusively carnivorous with unique features of their alimentary canal and feeding style
(Yonge, 1928; Purchon, 1956, 1987, 1990; Morton, 1981a, b, 1985c, 1987), and free proteolytic
enzymes are present in the stomach (Reid, 1977). Although a few benthic Foraminifera, spicules and
detrita material may be found in the stomach, the muco-ciliary system necessary for suspension and
detntal Reding 1S absent or vestigial, and septibranchs are not likely to be partially carnivorous as
suggested by Nakazima (1967).

The analysis of dietary records (Knudsen, 1967, 1970; Bernard, 1974; Krylova 1989) and
study of functional anatomy strongly suggest that cuspidariids feed mainly on swimming prey while
poromyids and verticordiids catch bottom-dwelling prey (Morton, 1987). Prey capture has been
observed in Cuspidaria by Reid & Reid (1974), and in Cardiomya by Reid & Crosby (1980) It is
not simply ascomphshed by aspiration, but utilizes the protrusion of the inhalant siphon, effected by
a complex muscular and haemocoel system. Poromya is thought to use a large raptorial hood formed
y the eversible, inhalant siphon (Morton, 1981a). In verticordiids, prey capture has been incorrectlv
ascribed to sticky tentacles (Allen & Turner, 1974), but these have been revealed to be only
sensory and not glandular. The mechanism of capture seems rather to depend on the eversion of a
snort conical siphon or of a raptorial hood (Morton, 1985a, 1987).

Recent progress, particularly in the functional anatomy, makes necessary a re-evaluation of
the systematics of the group. Early workers associated verticordiids. poromyids and cuspidariids in
a continuous series ol modifications from typically eulamellibranch to entirely gill-less forms but the
new data rather suggest that similarities are the consequence of adaptation to macrophagy

ANOMALODESMATA FROM THE TROPICAL PACIFIC

139

(Salvini-Plawen, 1980; Bernard, 1983; Morton, 1985c). However, the origin of the septum is still
uncertain the gill origin school (Pelseneer, 1888a, b. 1891) and the pallial origin school (Dall,
1886a, b. 1888) still have adherents — , and this issue may be finally resolved only by embryologicai
studies (Yonge, 1947). The discovery of cuspidariids with vestigial gills (Allen & Morgan, 1981)
does not alter the case. It suggests a progressive reduction of the gills and concurrent formation of
the septum. But, while innervation of the septum of Cardiomya is distinctly of mantle origin (Plate,
1897; Bernard. 1974), it is likely, as suggested by Morton (1981b), that the development of the
septum involved a significant pallial element and may be derived from the taenioid muscles present
in Parilimyia (Morton, 1981a, 1982). The family Parilimyidae of the superfamily Pholadomyoidea,
while not septibranch, has so many parallel adaptations and so prefigures the superfamilies
Verticordioidea and Cuspidarioidea, that it should be included as a potential root lineage.

The following synopsis of classification, based on an original frame elaborated by F.R.
Bernard, considers septibranchs as representative of two orders of the subclass Anomalodesmata.

As proposed by Runnegar (1974), cuspidariids are not thought to be related to the
poromyids, and the discovery of Protocuspidaria by Allen & Morgan (1981) shows them to be
unrelated to the protobranchs (Purchon 1956, 1960, 1963; Bernard 1974). The poromyids have a
significantly different statocyst structure (Morton, 1985b) and are placed in the order Poromyoida.

In a short paper overlooked by F. R. Bernard. Scarlato & Starobogatov (1983) proposed
a new classification of septibranchiate bivalves, mainly based on the structure of septum. A free
translation of this paper is given in Appendix 2. Its major difference with other classifications, is a
proliferation of family-group or higher-rank taxa. many of which are new. Although the usage of
several ordinal taxa can reflect the probable polyphyletic nature of septibranchs, and while some of
the divisions may prove to be useful, many of them seem presently unwarranted. This agrees with the
conclusions of Maxwell (1988), who discussed a classification of the Protobranchia established by
the same authors. Their system of Septibranchia, which incorporates some of the oldest and most
questionable bivalve-like fossils currently known as the Rostroconcha (Pojeta & Runnegar, 1976),
so greatly differs from the classification expressed in the present paper that it has not been attempted
to reconcile the two. Only new genera proposed by Scarlato & Starobogatov are tentatively
included here, because it did not alter the original frame of classification.

SYNOPSIS OF CLASSIFICATION

Subclass Anomalodesmata Dali, 1889 [nom. trails/, et correct. Keen, 1963]

Order Pholadomyoida Newell, 1965

Superfamily Pholadomyoidea Gray, 1847 [nom. transl. Newell, 1965]

Family Parilimyidae Morton, 1982

Genus Parilimya Melvill & Standen, 1899
Genus Panacea Dall, 1905

= Aporema Dall, 1903, non Scudder, 1890 (Insecta; Capsidae)

Genus Nipponopanacca Habe, 1977

Superfamily Verticordioidea Stoliczka, 1871 [nom. transl. Bernard, 1974]

Family Verticordiidae Stoliczka, 1871

Genus Verticordia J. de C. Sowerby, 1844

= Hippagus Philippi 1844, non Lea, 1833 (Mytilidae); Hippella Dall, 1903, non Morch, 1861
(Condylocardiidae); Iphigenia Costa, 1850, non Schumacher, 1817 (Donacidae)

Genus Pecchiolia Savi & Meneghini, 1851 (fossil: Caenozoic)

Genus Vertambitus Iredale, 1930

Genus Simplicicordia Kuroda & Habe in Kuroda, Habe & Oyama, 1971
Genus Trigonulina d'Orbigny, 1846

140

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Genus Spinosipella Iredale, 1930
Genus Haliris Dali, 1886b

Subgenus Haliris s. s.

Subgenus Setaliris Iredale, 1930
Subgenus Vertisphaera Iredale, 1930
Genus Halicardia Dali, 1895b

= Halicardissa Dali, 1913; Haloconcha Dali, 1900b nom. null.

Genus Kurinuia Marwick. 1942 (fossil: Paleogene)

Family Lyonsiellidae Dali. 1895a

= Policordiidae Scarlato, 1981

Genus Policordia Dali, Bartsch & Rehder. 1938
Subgenus Policordia s. s.

= Latebranchia Ivanova in Scarlato & Starobogatov, 1983
Subgenus Angustebranchia Ivanova in Scarlato & Starobogatov, 1983
Subgenus Dallicordia Scarlato & Starobogatov, 1983
Genus Laevicordia Seguenza, 1876a
Genus Lyonsiella G. 6. Sars, 1872

= Proagorina Iredale, 1930; Rectilyonsiella Scarlato & Starobogatov, 1983; Allenicordia

Scarlato & Starobogatov, 1983; ISpinolyonsiel/a Scarlato & Starobogatov, 1983

Family Euciroidae Dali. 1895a
Genus Euciroa Dali, 1881
Genus Acreuciroa Thiele & Jaeckel, 1931

Superfamily Cuspidarioidea Dali, 1886b [nom. transl. Scarlato & Starobogatov in Nevesskaia

et al., 1971]

Family Cuspidariidae Dali, 1886b

Genus Protocuspidaria Allen & Morgan, 1981
Subgenus Protocuspidaria s. s.

Subgenus Bident aria Allen & Morgan, 1981
Subgenus Edentaria Allen & Morgan, 1981
Genus Pseudoneaera Sturany, 1901

= Bendoneaera Cossmann, 1904, nom. null.; Jeffreysomya Nordsieck, 1969
Genus Cuspidaria Nardo, 1840

- Neaera Gray in Griffith & Pidgeon, 1834, non Robineau-Desvoidy, 1830 (Insecta-

Diptera); Aulacophora Jeffreys, 1882, non Chevrolat in Dejean, 1835 (Insecta: ColeopteraV

AUenineaera Scarlato & Starobogatov, 1983
Subgenus Cuspidaria s. s.

Subgenus Rhinoclama Dali & Smith in Dali, 1886b

= Rhinomya A. Adams, 1864, non Robineau-Desvoidy, 1830 (Insecta: Diptera)-

Austroneaera Powell, 1937

Subgenus Luzonia Dali & Smith in Dali, 1890

Subgenus Leiomya A. Adams, 1864

Subgenus Soyomya Okutani, 1985

Subgenus Tergulina Nosky, 1939 (fossil: Paleogene)

Subgenus Tropidomya Dali & Smith in Dali, 1886b

= Tropidophora Jeffreys, 1882, non Troschel, 1847 (Mollusca: Gastropoda); Gonio-
phora Jeffreys, 1883, non Phillips, 1848 (Modiomorphidae)

Subgenus Nordoneaera Okutani, 1985
Subgenus Vulcanomya Dali, 1886b
Genus Plectodon Carpenter, 1864
Genus Halonympha Dali & Smith in Dali, 1886b

Source .

ANOMALODESMATA FROM THE TROPICAL PACIFIC

141

Genus Cardiomya A. Adams, 1864

= Spathophora Jeffreys, 1882, non Amyot & Serville, 1843 (Insecta: Hemiptera)
Subgenus Cardiomya s. s.

Subgenus Bowdenia Dali, 1903 (fossil: Neogene)

Subgenus Kurodamya Okutani & Sakurai, 1964
Genus Bathyneaera Scarlato & Starobogatov, 1983

= Semicardiomya Scarlato & Starobogatov, 1983; Labromysa Bernard, 1989
Genus Boriesia Doncieux, 1911 (fossil: Paleogene)

Genus Octoporia Scarlato & Starobogatov, 1983
Genus Myonera Dali, 1886a

Subgenus Myonera s. s.

Subgenus Rengea Kuroda & Habe in Kuroda, Habe & Oyama, 1971
Genus Fabagella Cossmann, 1886 (fossil: Paleogene)

Order Poromyoida Pelseneer, 1906 [nom. correct. Newell, 1965]

Superfamily Poromyoidea Dali, 1886b [nom. transl. Dall, 1895a]

Family Poromyidae Dali, 1886b
Genus Poromya Forbes, 1844

= Embla Loven, 1846; Thetis H. Adams & A. Adams, 1856, non J. de C. Sowerbv, 1826
(Mactromyidae); Ectorisma Tate, 1892; Questimya Iredale, 1930
Subgenus Poromya s. s.

Subgenus Mioporomya Sacco, 1901 (fossil: Neogene)

Subgenus Dermatomya Dall, 1889
Subgenus Cetomya Dall, 1889
Genus Perlaporomya Scarlato & Starobogatov, 1983
Genus Neaeroporomya Cossmann, 1886 (fossil: Paleogene)

Genus Pseudocuspidaria Eames, 1951 (fossil: Paleogene)

Genus Cymella Meek, 1864 (fossil: Cretaceous)

Genus Cetoconcha Dall, 1886b

= Silenia E.A. Smith, 1885, non Mulsant & Rey, 1874 (Insecta Coleoptera); Cribrosoconcha
Krylova, 1991

Genus Liopistha Meek, 1864 (fossil: Cretaceous)

= Psilomya Meek, 1876

DIAGNOSES OF SUPRASPECIFIC TAXA

In this section, diagnoses are only for taxa with Recent representatives. They include also the
pholadomyoid family Parilimyidae.

Order Pholadomyoida Newell, 1965
Superfamily Pholadomyoidea Gray, 1847
Family Parilimyidae Morton. 1982

Shell thin, equivalve, gaping at both ends. Sculpture radial, either feeble or strong.
Periostracum thin, usually with adherent sand grains. Ligament external, opisthodetic. Hinge plate
feeble, edentate or with an anterior tubercle. Pallial line with a moderately deep sinus.

Mantle lobes fused, with an anteroventral pedal gape and a fourth pallial aperture. Mantle
margins with arenophilic radial glands. Inhalant siphon very large, eversible; exhalant siphon small.
Taenioid muscles present. Foot rounded, elongate. Gills eulamellibranch, with reduced outer

142

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

demibranch. Labial palps short, ridged. Stomach features transitional between types II and IV
(Purchon, 1990). Midgut and style-sac conjoined. Intestine passing through the ventricle of heart.
Statocyst with a multicellular capsule of ciliated cells and a large free statolith within, either single
or associated with a number of small statoconia. Hermaphroditic.

Remarks: This family is included here because it has many of the adaptations shown by the
carnivorous groups. It is probably at least partly raptorial, and may be ancestral to the verticordiids
(Morton, 1981a, 1982, 1987).

Superfamily Verticordioidea Stoliczka, 1871

Shell inflated, inequilateral, ovate to quadrate or trapezoidal, subequivalve. Outer surface
usually with radial sculpture, granulate to spinose, sometimes smooth. Ligament opisthodetic, with
resilium supported by a lithodesma. Hinge variable, feeble and edentate to more or less thickened,
with cardinal and lateral elements. Interior of valves nacreous. Microstructure of shell with prismatic
outer, lenticular nacreous middle, and sheet nacre inner layers.

Mantle lobes fused, with a variable pedal opening and arenophilic radial glands on margins.
Inhalant and exhalant apertures usually with short siphonal apparatus, surrounded by large sensory
tentacles. Foot digitiform or laterally compressed, often weakly byssate in adult. Septum diaphanous,
with reduced eulamellibranch gills. Labial palps mostly small; lips enlarged as an oral funnel.
Stomach very muscular, of type II. Two ducts to the digestive diverticula. Midgut and style-sac
conjoined. Intestine passing through the ventricle of the heart. Statocyst with a large multicellular
capsule of ciliated cells and a single free statolith within. Hermaphroditic.

Family Verticordiidae Stoliczka, 1871

Shell ovate to trapezoidal, generally with a conspicuous radial sculpture and a well-demarcated
lunule. Outer surface frequently granulate or spinose. Hinge with at least a cardinal tubercle in right
valve.

Gills reduced. Foot digitiform, often byssate. Labial palps reduced.

Genus Verticordia J. de C. Sowerby, 1844

Type species (by monotypy): Hippagus? cardiiformis J. de C. Sowerby, 1844. Pliocene, Northern
Europe.

Shell inflated, rather solid, usually granulate, pustulate or spinose. Umbones prosogyrate,
prominent. Sculpture often with radial riblets or plications. Lunule large, deeply impressed. Hinge
plate with a hooked cardinal tubercle in right valve, occasionally with posterior lateral tooth.

Included species; australiensis E. A. Smith, bordaensis, expansa, granulifera, guineensis,
inornata , monosteira, perversa , quadrat a, seguenzae, tenerrima, woodi.

Genus Vertambitus Iredale, 1930

Type species (OD); Verticordia vadosa Hedley, 1907a. Recent, Australia.

Shell rather compressed, with prominent umbones. Sculpture of feeble wide radial riblets and
rows of pustules on entire surface. Hinge heavy, with at least a strong cardinal tooth in right valve.

Included species: affinis, cuneatus, excoriatus, torridus, triangularis , vadosus.

ANOMALODESMATA FROM THE TROPICAL PACIFIC

143

Genus Simpucicordia Kuroda & Habe in Kuroda. Habe & Oyama, 1971
Type species (OD): Thyasira trigonaia Yokoyama, 1922. Pleistocene to Recent, Japan.

Shell small, thin, inflated, subtrigonal. Umbones prosogyrate, not prominent. Anterodorsal
margin concave; posterodorsal margin convex. Surface minutely granulated. Sculpture of feeble
concentric irregular riblets, corrugating the interior of the shell. No radial sculpture. Hinge plate
feeble, with an obscure cardinal denticle in right valve.

Included species: trigonaia.

Genus Trigonulina d'Orbigny, 1846

Type species (by monotypy): Trigonulina ornata d’Orbigny. 1846. Recent, Caribbean.

Shell rather compressed, solid. Radial ribs prominent, irregularly spaced, crenulating the
ventral shell margin. Lunule deeply impressed. Right valve with a stout, posteriorly recurved cardinal
tooth, and a long socket to accomodate the posterior lateral tooth of the opposite valve. Interior
brilliantly nacreous.

Included species: hancocki, ornata.

Genus Spinosipella I redale, 1930

Type species (OD): Verticordia ericia Hedley, 1911. Recent, Indo-Pacific.

Shell inflated, solid, with strongly enrolled, prosogyrate umbones overhanging the lunule.
Sculpture of prominent, radial ribs crenulating the ventral shell margin. Lunule small, deeply
invaginated. Outer surface densely granulate to spinose. Hinge of right valve with a strong cardinal
tooth. Left valve usually with a smaller corresponding denticle.

Remarks: Usually treated as a subgenus in Verticordia. but the near absence of lunule and hinge
features support its separation.

Included species: acuticostata, costeminens, deshayesiana, ericia.

Genus Haliris Dali, 1886b

Type species (OD): Verticordia fischeriana Dali, 1881. Recent, Caribbean.

Shell inflated, finely granulate, with small and regular radial ribs, or with obsolete sculpture.
Hinge usually with cardinal tubercles and posterior lateral teeth in both valves, more developed in
the right.

Subgenus Haliris s. s.

Shell solid, sculpture with wide radial riblets. Lunule shallow. Hinge plate strong; right valve
with a stout cardinal tooth and a lateral ridge under posterodorsal margin; left valve with a minute

144

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

ephemeral cardinal tubercle and a small posterior lateral tooth. Inner ventral margin of valves
crenulated.

Included species: berenicensis, crebrilirata, fischeriana , granulata, jaffaensis, lamothei, multi-
costata, spinosa, teporis.

Subgenus Setaliris Iredale. 1930

Type species (OD): Verticordia setosa Hedley, 1907a. Recent, New Zealand.

Shell small, solid, quadrate, with numerous radial riblets and intercalated concentric striae.
Lunule shallow. Hinge with cardinal and lateral teeth in both valves. Ventral shell margin crenulate,
somewhat flexuous.

Included species: accessa , pygmaea, setosa.

Subgenus Vertisphaera Iredale, 1930

Type species (OD): Vertisphaera cambrica Iredale, 1930. Recent. Australia.

Shell thin, sculpture obsolete. Lunule depressed. Hinge plate feeble, right valve with an
obscure cardinal tooth. Inner ventral margin of shell smooth.

Remarks: Frequently placed as a subgenus in Verticordia , the hinge shows it to be referable to Hali-
ris.

Included species: cambrica.

Genus Haucardia Dali, 1895b

Type species (OD): Mytilimeria flexuosa Verrill & Smith in Verrill, 1881. Recent, North Atlantic.

Shell large for the group, inflated. Radial sculpture developed either as ribs or strong to feeble
plications and furrows, scalloping the ventral shell margin. Outer surface with small granules. Lunule
small, deeply impressed. Hinge plate feeble, right valve with a small to obsolete cardinal tooth.
Visceral mass with an opisthopodium posterior to the foot.

Remarks: The separation of Halicardissa Dali, 1913 (type species: Verticordia perplicata Dali, 1890)
does not seem to be warranted, neither on conchological nor on anatomical grounds.

Included species: angulata , carinifera, ferruginea, fischeri. flexuosa , gouldi. houbricki, maoria,
nipponensis, perplicata, philippinensis, saharica.

Family Lyonsiellidae Dali, 1895a

Shell thin, inequilateral, suborbicular to quadrate, with a reduced sculpture. Lunule obscure.
Outer surface with or without granulations. Hinge edentate.

Gills reduced to absent. Foot digitiform, usually with a byssal groove. Labial palps small.

Source . MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

145

Genus Policordia Dali, Bartsch & Rehder, 1938
Type species (OD): Policordia diomedea Dali, Bartsch & Rehder, 1938. Recent, Hawaii.

Shell suborbicular to trapezoidal, usually with radiating lirae and concentric marks, often
somewhat reinforced by periostracum. Outer surface completely devoid of spines or granulations.

Gills variably reduced to absent.

Remarks: Ivanova (1977) divided Policordia into two new subgenera ( Angustebranchia and
Latebranchia), thereby extinguishing Policordia s. s. However, Ivanova originally failed to designate
type species for her two subgenera which are then unavailable, under Article 13b of the Code of
Nomenclature, whereas the species described as new in her work are available under Article 11, h,
iii, 1.

Subsequently, Ivanova made Angustebranchia and Latebranchia available (in Scarlato &
Starobogatov, 1983) as full genera, by the designation of type species and the explicit reference to
the diagnoses published in 1977. We agree with Scarlato & Starobogatov to consider that
Latebranchia is equivalent to Policordia s. s.

Subgenus Policordia s. s.

Inhalant aperture with one row of tentacles. Gills wide, fused posterior to the foot. Mouth
broad. Bathyal to abyssal.

Included species: atlantica, cordata, densicostata , diomedea, gemma , grandis, insolita, ivanovae ,
jeffreysi (Friele), lisbethae, murrayi, obliqueovata, olivacea, ovata, papyracea , pilula, radiata, subro-
tundala.

Subgenus Angustebranchia Ivanova in Scarlato & Starobogatov, 1983

Type species (OD): Policordia (Angustebranchia) rectangulata Ivanova, 1977. Recent, Kurile-
Kamchatka Trench.

Inhalant aperture with two rows of tentacles. Gills narrow, free posterior to the foot. Mouth
narrow. Hadal.

Included species: extenta, laevigata, maculata, rectangulata.

Subgenus Dallicordia Scarlato & Starobogatov, 1983

Type species (OD): Lyonsiella alaskana Dali, 1895b. Recent, East Pacific.

Inhalant aperture with one row of tentacles. Gills absent. Septum thin, with a few pairs of
small pores. Mouth large. Bathyal to abyssal.

Included species: alaskana, ochotica, uschakovi.

146

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Genus Laevicordia Seguenza, 1876a

Type species (subsequent designation by Soot-Ryen, 1966): Verticordia ( Laevicordia ) orbiculata
Seguenza, 1876a. Pliocene to Recent, Mediterranean.

Shell suborbicular, with fine granules and obscure radial striae. Hinge margin may be
somewhat thickened anteriorly. Gills moderately reduced.

Included species: abscissa, axinoides , frieli, galatheae , horrida, insculpta, orbiculata, pacifica,
sinuosa, smidti.

Genus Lyonsiella G. O. Sars, 1872

Type species (by monotypy): Pecchiolia abyssicola G. O. Sars, 1872. Recent, Arctic, North Atlantic.

Shell small, oval to subquadrate, generally with fine pustules or spines and radial striae. Hinge
plate feeble, edentate, but anterior of the left valve may be thickened. Inhalant siphon with
an eversible raptorial valvule. Taenioid muscles sometimes well developed. Gills moderately redu¬
ced.

Included species: abyssicola, agulhasensis, compressa, curta, formosa, fragilis, magnifica, parva,
perplexa, quadrata , quaylei, subquadrata.

Family Euciroidae Dali, 1895a

Shell usually thick, solid, with numerous radial riblets or striae. Outer surface finely pustulose
or spinose. Lunule weakly demarcated. Hinge plate robust, with variably developed lateral laminae
and one or two cardinal teeth. Interior of valves highly nacreous, radially striated.

Mantle margins strongly muscular. Gills with reduced outer demibranch. Foot late¬
rally compressed, without a byssal groove. Labial palps large, striated. Lips produced into lateral
bulbs.

Genus Euciroa Dali, 1881

Type species (by monotypy; genus name Euciroa cited in synonymy): Verticordia elegantissima Dali,
1881. Recent, Caribbean.

Shell ovate, inflated. Hinge of right valve with a strong, curved cardinal tooth, and usually a
posterior ridge. Left valve with a small cardinal tooth edging the resilifer, and often shallow lateral
laminae.

Included species: aethiopica, crassa, eburnea, elegantissima, galatheae, granifera, mediopacifi-
cal, millegemmata , pacifica, spinosa, trapeza.

Source . MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

147

Genus Acreuciroa Thiele & Jaeckel, 1931

Type species (by monotypy): Euciroa (Acreuciroa) rostrata Thiele & Jaeckel, 1931. Recent,
Indo-Pacific.

Shell posteriorly produced, somewhat compressed. Right valve with a conical cardinal tooth.
Left valve nearly edentate.

Included species: rostrata.

Superfamily Cuspidarioidea Dali, 1886b

Shell inflated, strongly inequilateral, rounded to ovate anteriorly, rostrate posteriorly. Most of
the time slightly inequivalve, with left valve larger. Sculpture variable, often reduced to absent,
sometimes with radial or concentric elements. Outer surface with thin periostracum, generally smooth
(rarely granulate). Ligament sunken into a small resilifer, supported by a lithodesma. Hinge feeble,
edentate or with small cardinal tubercles and lateral ridges. Interior of valves porcelaneous!
Microstructure of shell with homogenous outer and inner layers.

Mantle lobes fused, with a small pedal opening. Inhalant and exhalant apertures with well
developed siphons united at their base and with seven prominent sensory tentacles. Exhalant siphon
short and eversible; inhalant siphon relatively large and forcefully extensible to capture swimming
prey. Foot digitiform, with a byssal groove. Adult not byssate. Gills generally absent. Septum usually
muscular, with a number of paired pores (usually four or five pairs of pores). Labial palps small,
flap-shaped (posterior palps cup-shaped, when developed). Stomach very muscular, of type II. Two
ducts to the digestive diverticula. Midgut and style-sac separated. Intestine passing through the
ventricle of the heart. Statocyst with a small capsule comprising a few swollen and microvillose cells,
and a large, ovate, not freely mobile statolith within. Dioecious.

Family Cuspidariidae Dali, 1886b
Characters same as for the superfamily.

Genus Protocuspidaria Allen & Morgan, 1981
Type species (OD): Protocuspidaria verityi Allen & Morgan, 1981. Recent, Atlantic.

Shell equivalve, rounded, laterally compressed; rostrum short, truncate. Hinge edentate, or
with anterior lateral tooth in one or both valves. Septum thin, membranous, with two longitudinal
rows of gill filaments but no muscle attachments to the shell. Posterior labial palps large, cup-shaped.

Remarks: This very interesting genus may represent a transitional stage in the development of the
gill-less cuspidariids (Allen & Morgan 1981). The division of the representatives into three
subgenera may be premature as the hinge dentition may be mutable.

Subgenus Protocuspidaria s. s.

Hinge of right valve with an anterior lateral tooth. Left valve edentate.
Included species: verityi.

Source .

148

JEAN-MAUR1CE POUTIERS & FRANK R. BERNARD

Subgenus Bident aria Allen & Morgan, 1981

Type species (OD): Protocuspidaria ( Bidentaria ) atlantica Allen & Morgan, 1981. Recent, North
Atlantic.

Hinge of both valves with an anterior lateral tooth.

Included species: aequatorialisl, atlantica , colpodesl

Subgenus Edentaria Allen & Morgan, 1981

Type species (OD): Protocuspidaria (Edentaria) simplis Allen & Morgan, 1981. Recent, North
Atlantic.

Hinge of both valves edentate.

Included species: ruginosal, simplis, thomassini.

Genus Psevdoneaera Sturany, 1901

Type species (by monotypy): Pseudoneaera thaumasia Sturany, 1901. Recent, Red Sea.

Rostrum short, trigonal. Hinge of right valve with anterior and posterior cardinal teeth. Left
valve with an anterior cardinal tooth.

Included species: minor, periplomoidesl , semipellucida, tasmanical, thaumasia, trigonalisl,
truncata, wellmani.

Genus Cvspioaria Nardo, 1840

Type species (by monotypy): Cuspidaria typus Nardo, 1840 = Tellina cuspidata Olivi, 1792. Recent,
Northeast Atlantic, Mediterranean.

Outline globular to ovate; rostrum variably developed, often prominent. Left valve slightly
more convex. Hinge teeth present, at least in the right valve. Septum muscular, usually with four or
five pairs of pores, with both anterior and posterior muscle attachments to the shell. Labial palps
small.

Remarks: The general arrangement of the genus depends on the type and presence of hinge teeth,
which seem rather mutable.

Subgenus Cuspidaria s. s.

Outer surface smooth or with concentric lirae. Rostrum often well developed. Hinge of right
valve with a posterior lateral tooth. Left valve edentate.

Included species: abyssopacifica, angasi, annandaleV., apodema, approximata, arctica, arcuata,
ascoldica, atlantica, azorical, barnardi, bicar inata, brachyrhynchus, braziliensisl, buccina, capensis,

Source .

ANOMALODESMATA FROM THE TROPICAL PACIFIC

149

chinensis, circinata , cochinensisl , concentrica, consociata, contracta, convexa Pelseneer, corrugata,
cowani , cuspidata, delli, dissociata, elegans, elliptical , erma, exarata , exigua, fairchildi, formosa,
fraterna , giganlea Prashad, glacialis, gracilis, guineensisl, haasi, halei, hawaiensis, hindsiana, hyalinal,
imbricata , infelix,japonica, Jeffrey si, jugosa Wood?, kerguetensis, kurodai, kyushuensis, lamellosa G. O.
Sars, latesulcata, lubangensis, macrorhynchus, maxima, media, meridionalis, microrhina , mitis,
morelandi, moriorial, morrisae, nasuta A. Adams, natalensis, nobilis, obesa, obtusirostris, occidua,
okezoko, optima, panamensis , papyrial, parapodema, parkeri, parva, palagonica, pellucida, platensis,
prolatissima, rosea, rostrata, sadoensis, semirostrata, solidula, sleindachneri, subglacialis, subtorta,
suganumai, sulcifera, tene/la, teramachii, tomlini, trailli, trigonal, truncatal, tuhua, turgida, undata,
variola, ventricosa, willetti, wollastonii.

Subgenus Rhinocluma Dali & Smith in Dali, 1886b

Type species (iczn Opinion 1376): Cuspidaria (Rhinoclama) adamsi Morgan & Heppell in Allen &
Morgan, 1981. Recent, Philippines.

Rostrum moderately extended, with two radial ridges. Outer surface with fine concentric
striations. Hinge of right valve with triangular anterior and posterior lateral teeth. Left valve
edentate.

Remarks: The group was established as Rhinomya A. Adams, 1864, as a subgenus of Neaera, but was
preoccupied. Rhinoclama Dali & Smith (in Dali, 1886b) was proposed as a replacement name, but a
nomenclatural problem existed for a number of years, because the type species was a nomen nudum.
The result of an inadequate nomenclatural action by Stoliczka (1871) merged Luzonia Dali & Smith
(in Dali, 1890) as a synonym, although the differences between the two had already been quoted by
E. A. Smith (1885). The situation was finally regularized by Heppell & Morgan (1983) and Opinion
1376 of the iczn (1986).

Included species: abrupta, adamsi Heppell & Morgan, alta, aupouria, benthedii, brevirostris
Powell, dorsirecta, dubia Pelseneer, filatovae, ftnlayi, halimera , nitens, notabilis, raoulensis, rugata A.
Adams, similis, semistrigosa, simulans , teres, testai, valdiviae.

Subgenus Luzonia Dali & Smith in Dali, 1890
Type species (iczn Opinion 1376): Neaera philippinensis Hinds, 1843. Recent, Philippines.

Rostrum short, tapered, largely confluent with the disc. Hinge of right valve with an anterior
cardinal tooth, frequently twisted. Left valve edentate.

Remarks: Stoliczka (1871) designated Neaera phillipinensis (a lapsus for philippinensis) Hinds as the
type of Rhinomya, thereby merging Luzonia as an objective synonym. Thiele (1934), following the
brief diagnosis of E. A. Smith (1885), proposed Cuspidaria adamsi as a substitute name for Neaera
philippinensis “A. Adams” non Hinds, 1843. As this was a nomen nudum, its substitute has no
nomenclatural status. Heppell & Morgan (1983) reviewed the situation and showed that N.
philippinensis Hinds, 1843, is the type of Luzonia. This genus name has been validated by Opinion
1376 of the iczn (1986).

Included species: chilensis, philippinensis Hinds, simplex, walleri.

150

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

Subgenus Leiomya A. Adams, 1864

Type species (by monotypy): Neaera adunca Gould, 1861. Recent, Japan.

Outer surface smooth. Rostrum short, trigonal. Hinge of both valves with an anterior cardinal
tooth, that of the right valve usually strongly bifid. Right valve also with an anterior and a posterior
lateral tooth.

Included species: adunca Gould, injlata.

Subgenus Soyomya Okutani, 1985

Type species (by monotypy): Cuspidaria kurohjii Okutani, 1972. Recent, Japan.

Rostrum short, wide. Outer surface with a complex oblique sculpture. Right valve with a
posterior lateral tooth. Left valve edentate.

Included species: clathrata, kurohijii.

Subgenus Tropidomya Dali & Smith in Dali, 1886b

Type species (by monotypy): Neaera abbreviata Forbes, 1843. Recent, Northeast Atlantic, Mediter¬
ranean.

Rostrum short, trigonal. A small anterior cardinal tooth in both valves.

Included species: abbreviata , diagonalis.

Subgenus Nordoneaera Okutani, 1985

Type species (by monotypy): Cuspidaria trosaetes Dali, 1925. Recent, Japan.

Shell stout, rostrum very short, with obscure boundary. Hinge of right valve with a posterior
lateral tooth. Left valve edentate.

Included species: trosaetes.

Subgenus Vulcanomya Dali, 1886b

Type species (by monotypy): Vulcanomya smithii Dali, 1886b = Neaera adunca “Gould” E. A. Smith,
1885 (non Gould, 1861).

Rostrum short, trigonal. Hinge of right valve with a short trigonal lateral tooth on either side
of the resilifer. Left valve with a tiny double notch in the anterior dorsal margin.

Included species: smithii.

Source :

ANOMALODESMATA FROM THE TROPICAL PACIFIC

151

Genus Plectodon Carpenter, 1864

Type species (OD): Plectodon scaber Carpenter, 1864, Recent, East Pacific.

Outer surface dull, usually granulate. Anterodorsal margin of valves spirally incurved. Hinge
of right valve with elongate, reflected anterior and posterior lateral teeth. Left valve edentate. Septum
relatively thin, with generally five pairs of pores, and both anterior and posterior muscle shell
attachments. Labial palps small.

Remarks: This genus has been frequently but erroneously listed as a subgenus of Leiomya. It is
treated as a subgenus of Cuspidaria by Allen & Morgan (1981).

Included species: brazieri, granulatus, ligula, scaber.

Genus Halonympha Dali & Smith in Dali, 1886b

Type species (OD): Neaera claviculata Dali, 1881. Recent, West Central Atlantic.

Shell ovate; rostrum merged with disc. Hinge of right valve with a small knob-like cardinal
tooth. A raised oblique buttress or ridge along the posterodorsal margin of either valve. Septum with
more than five pairs of pores, without posterior dorsal attachments to the shell. Posterior labial palps
large, cup-shaped.

Remarks: Allen & Morgan (1981) separated the group at the genus level on the basis of significant
anatomical differences.

Included species: aethiopica , asiatica, at lant a. claviculata, congenita , depressa, ledaeformisl,
leiomyoides , ros, salamensis , striatella.

Genus Cardiomya A. Adams, 1864

Type species (by monotypy): Neaera gouldiana Hinds, 1843. Recent, Western Pacific.

Radial sculpture developed, at least on posterior half of disc. Rostrum frequently elongate.
Right valve with a posterior lateral tooth. Septum muscular, with four pairs of pores; a small lateral
septal muscle generally present, besides the anterior and posterior muscle shell attachments.

Remarks: Anatomically, the genus does not differ much from Cuspidaria , and Allen & Morgan
(1981) merged them. The consistent differences in sculpture and general more shallow distribution of
Cardiomya suggests its separation.

Subgenus Cardiomya s .s.

Radial sculpture developed over entire disc. Right valve with a posterior lateral tooth. Left
valve edentate.

Included species: abyssicola , alcocki, alternata, alveata, andamanica , angusticauda, balboae,
behringensis, bruuni, californica, casta , chuni, cleryana, concinna , cos tat a, costellata, curt a Jeffreys,
didyma, ecuadoriana, filatovae, forticostata, fragilissima, fujitai, gilchristi, gouldiana , greenii, iturupica,
kashimana, knudseni, lanieri, nipponica , obliqua , ochotensis , orientals, ornatissima, pectinata, perros-
trata, pinna, planetica, rectimarginata, reticulata, saba, sibogai, singaporensis, sinica, striata, striolata,
surinamensis, tosaensis.

Source .

152

JEAN-MAURICE P0UT1ERS & FRANK R. BERNARD

Subgenus Kurodamya Okutani & Sakurai, 1964

Type species (by monotypy): Cuspidaria (Cardiomya) fortisculpta Kuroda, 1948. Recent, Japan.

Radial sculpture absent from the anterior slope of the disc. Right valve with a prominent,
often hook-shaped, posterior lateral tooth. Left valve edentate.

Remarks: A useful division, merged by Allen & Morgan (1981) into Cardiomya.

Included species: fallaxl, fortisculpta, levifrons, semicostata.

Genus Bathyneaera Scarlato & Starobogatov, 1983

Type species (by monotypy): Cuspidaria hadalis Knudsen, 1970. Recent, Banda Trench.

Sculpture a complex of radial riblets and concentric lirae, more developed on posterior slope.
Rostrum compressed, truncate, merged with disc. Hinge of both valves edentate. Septum muscular,
with four pairs of pores; an extra lateral septal muscle attached on posterior part of shell, besides the
other posterior muscles.

Remarks: Bernard (1989) created Labromysa (type species: Cardiomya (Labromysa) disa Bernard,
1989) as a subgenus of Cardiomya, to accomodate species with external shell features similar to
Myonera Dali, 1886a, but anatomically assignable to Cardiomya, and with a feeble edentate hinge
plate. He tentatively included in it Myonera laticella Dali, 1888b and Cuspidaria hadalis (the type
species of Bathyneaera). Consequently, Labromysa is considered here to be a synonym of
Bathyneaera. Krylova (1993) revised the genus, merging in it Semicardiomya Scarlato &
Starobogatov, 1983.

Included species: bernardi, disa, globulosa, hadalis, laticella, paleifera, quadrostrata, tillamoo-

kensis.

Genus Octoporia Scarlato & Starobogatov, 1983

Type species (OD): Cuspidaria (Myonera) octaporosa Allen & Morgan, 1981. Recent, North Atlantic.

Shell ovate, with a well developed, broad rostrum. Sculpture of low concentric ribs. Hinge of
both valves edentate. Septum with eight to twenty pairs of pores, with feeble posterodorsal
attachments to the shell. Posterior labial palps large, cup-shaped.

Remarks: Krylova (1994b) revised Octoporia and assigned to it a number of new species in addition
to the type and only known species, showing then it is worth of generic recognition. The genus
accomodates species with shell features recalling Myonera Dali, 1886a, but anatomically similar to
Halonympha Dali & Smith in Dall, 1886b.

Included species: octaporosa, podobeda, rugosa, sinuosa.

Genus Myonera Dali, 1886a

Type species (subsequent designation by Dall & Smith in Dall, 1886b): Myonera paucistriata Dali
1886b. Recent, North Atlantic.

Source .

ANOMALODESMATA FROM THE TROPICAL PACIFIC

153

Sculpture with concentric, and sometimes radial elements. Hinge plate feeble, edentate in both
valves. Septum muscular, with four pairs of pores. Labial palps small.

Remarks: In a short note to Nature published 10 June 1886, Dall introduced Mvonera, as a subgenus
of Neaera (an older but invalid name for Cuspidaria). The anatomical description he gave, based on
a new (but unnamed) species of Myonera from the Gulf of Mexico, made it available as a genus-group
name without named species. Later (September 1886), Dall & Smith (in Dall 1886b- 302) defined
Myonera as a full genus, providing a list of species to be included in it. and designating Myonera
paucistriata Dall as the type species.

Subgenus Myonera s. s.

Sculpture complex of radial and concentric elements. Resilifer posteriorly directed, or nearly
vertical. ’ J

Included species: acutecarinata, alleni, angularis, bicarinata, canariensis, centobi, dautzenbergi,
dispar, garretti, gigantea, lamellifera , limatula, lischkei, mexicana , pailoloana , paucistriata, pretiosal,
rostra, tasmanica.

Subgenus Rengea Kuroda & Habe in Kuroda, Habe & Oyama, 1971

Type species (OD): Myonera fluctuosa Kuroda, 1948 = Cuspidaria (Myonera) caduca E. A. Smith
1894. Recent, Indo-Pacific.

Sculpture of the disc with strong concentric plications. Umbones depressed, opisthogyrate.
Resilifer projecting, subvertical.

Included species: caduca, murrayi.

Order Poromyoida Pelseneer, 1906
Superfamily Poromyoidea Dall. 1886b

Shell inflated, ovate to rhomboidal or trigonal, subequivalve. Posterior rostrum absent. Outer
surface with adherent periostracum, smooth or granulate to spiculate. Ligament external, opistho-
detic, without a lithodesma. Dorsal margins of valves united by fused periostracum. Hinge edentate,
or with a variable cardinal tooth in right valve. Interior of valves nacreous. Microstructure of shell
with prismatic outer and nacreous middle and inner layers, or with a two-layered homogenous
structure.

Mantle lobes fused, with a large pedal opening. Inhalant and exhalant apertures with siphons,
surrounded with up to 15 stout sensory tentacles. Exhalant siphon short; inhalant siphon eversible
in a large raptorial hood. Foot digitiform, with a byssal groove. Adult usually not byssate. Gills
absent. Septum muscular, with two or three (rarely one) pairs of ostial openings. Labial palps
cup-shaped, the anterior ones large. Stomach very muscular, of type II. Two ducts to the digestive
diverticula. Midgut and style-sac conjoined. Intestine passing through the ventricle of the heart.
Statocyst with a large multicellular capsule of ciliated cells and a single free statolith within.
Hermaphroditic.

Source .

154

JF.AN-MAUR1CE POUTIERS & FRANK R. BERNARD

Family Poromyidae Dali, 1886b
Characters same as for the superfamily.

Genus Poromva Forbes, 1844

Type species (by monotypy): Poromya anatinoides Forbes, 1844 = Corbula granulata Nyst &
Westendorp, 1839. Pliocene to Recent, Arctic, North Atlantic and Mediterranean.

Shell surface smooth or granulate. Hinge with or without a cardinal tooth in right valve.
Septum with two pairs of ostial openings.

Subgenus Poromya 5. s.

Shell surface with small granules, frequently set in rows. Ligament external, but deeply sunken.
Right valve with an anterior cardinal tooth.

Included species: adelaidis, australis, curta, cymata, flexuosa, gilchristi, granosissima, granulata
Nyst & Westendorp, granuloderma, hayashii, houhricki, laevis, neaeroides, neozelanica, rostrata,
sansibarica, spinulosa, striata, sumatrana, tornata, transversa, umbonata, undosa.

Subgenus Dermatomya Dali, 1889

Type species (by monotypy): Poromya (Dermatomya) mactroides Dali, 1889. Recent, Eastern Pacific.

Shell surface smooth, without granulation. Ligament external, but deeply sunken. Hinge
rather strong, cardinal tooth of right valve large, frequently knob-like.

Included species: buttoni , castanea, chilensis, hyalina, mactroides, tenuiconcha, trosti.

Subgenus Cetomya Dali, 1889

Type species (subsequent designation by Glibert, 1936): Poromya elongata Dali, 1886b. Recent, West
Central Atlantic.

Shell surface granulate. Ligament external, not deeply sunken. Hinge plate feeble, cardinal
tooth of right valve vestigial or absent.

Included species: albida, butoni, elongata, eximia Pelseneer, malespinae Ridewood, niasensisl,
orientalis, scapha.

Genus Perlaporomya Scarlato & Starobogatov, 1983
Type species (OD): Poromya perla Dali, 1908. Recent, East Pacific.

Shell surface granulate, with very high, bulbous umbones. Ligament external, but deeply

Source .

ANOMALODESMATA FROM THE TROPICAL PACIFIC

155

sunken. Hinge with a strong cardinal tooth in right valve, and a corresponding socket in left valve.
Septum with only one pair of ostial openings.

Included species: per la.

Genus Cetoconcha Dali, 1886b

Type species (OD): Lyonsia bulla Dali, 1881. Recent, tropical West Atlantic.

Shell surface granulate or spiculated. Ligament external, slightly sunken. Hinge feeble; anterior
cardinal tooth of right valve reduced or absent. Septum with three pairs of ostial openings.

Included species: alephtinae, angolensis, atypha , boucheti, braziliensis, bulla, ceylonensis, elegans,
e.xigua, galatheae, gloriosa, indica, japonica, margarita , pelseneeri, sarsii, smithii , tenuissima, transversa.

CATALOGUE OF RECENT SPECIES

In the following alphabetical list, the generic attributions and synonymies are only tentative
because, for many species, information is scarce, too scattered or obsolete. Some genera are mainly
distinguishable on anatomical grounds, but these data are not available for many species, hence many
confusions in generic or even familial allocations (see, for example. Poutiers, 1984: 285; Dell, 1990
61).

In two publications dealing with the anatomy of bivalves, Ridewood (1903) and Pelseneer
(1911) described the soft parts of some new septibranch species. The shells of these species were later
named and described as new, respectively by Dall (1916a, b) and Prashad (1932). Although a few
authors, among which F. R. Bernard, rejected the names introduced by Ridewood and Pelseneer,
these are tentatively included herein. However their somewhat uncertain nomenclatural status has to
be reconsidered on an individual basis in a thorough revision of the group.

Names of valid species are in bold. Current generic allocation is in square brackets.

abbreviala, Neaera Forbes, 1843: 75. Northeast Atlantic. Mediterranean. 55-1350 m. | Cuspidaria
(Tropidomya)\

abrupta, Cuspidaria ( Rhinoclama ) Allen & Morgan, 1981: 479. Southeast Atlantic. 619-1014 m.
[ Cuspidaria ( Rhinoclama ) |

abscissa , Lyonsiella Pelseneer, 1911: 76. Indo-Pacific. 700-850 m. [Laevicordia]
abyssicola, Cardiomya Verrill & Bush, 1898: 806. Northwest & West Central Atlantic. 2840-3316 m.
\Cardiomya\

abyssicola, Lyonsiella M. Sars, 1869: 257, nom. nud. = Lyonsiella abyssicola (G.O. Sars, 1872)
abyssicola, Pecchiolia G.O. Sars, (ex M. Sars MS) 1872: 25. Arctic, North Atlantic. 38-3909 m.

| Lyonsiella]

abyssopacifica, Cuspidaria Okutani, 1975b: 74. Northwest Pacific. 3420-5620 m. \Cuspidaria]
accessa, Setaliris Iredale, 1930: 388. Southwest Pacific. 457-550 m. [Haliris (Setaliris)\
actoni, Neaera Tiberi, 1855: pi. I. = Cardiomya costellata (Deshayes, 1833)
acutecarinata. Cuspidaria Dautzenberg & Fischer, 1906: 95. East Central Atlantic. 628 m. \Myonera\
acuticostatus, Hippagus Philippi, 1844: 42. Central Atlantic, Mediterranean. 99-4255 m. \Spinosipella]
adamsi, Cuspidaria Thiele, 1934 (nom. nov. pro Cuspidaria philippinensis “Hinds”, A. Adams, 1864):
948, nom. nud. = Cuspidaria (Rhinoclama) adamsi Heppell & Morgan, 1981

156

JEAN-MAUR1CE POUTIERS & FRANK R. BERNARD

adamsi. Cuspidaria (Rhinoclama) Heppell & Morgan, 1981: 546. West Pacific. 38-46 m. [Cuspidaria
( Rhinoclama )\

adelaidis, Pholadomya Hedley, 1916: 29. Antarctic. 110-2154 m. \Poromya\
adunca , Neaera Gould, 1861: 24. Northwest Pacific. 10-600 m. | Cuspidaria (Leiomya)\
adunca, Neaera “Gould” E.A. Smith 1885: 37, non Gould 1861. = Cuspidaria (Vulcanomya) smithii
Dali, 1886b

aequacostata, Verticordia Howard, 1950: 109. = Haliris Jischeriana (Dali, 1881)
aequatorialis, Cuspidaria Thiele & Jaeckel, 1931: 253. Indian Ocean. 693-750 m. [ 1 Protocuspidaria
( Bident aria )\

aethiopica, Cuspidaria Thiele & Jaeckel, 1931: 254. West Indian Ocean. 693 m. \Ha!onympha\
aethiopica , Euciroa Thiele & Jaeckel, 1931: 248. West Indian Ocean. 818-977 m. \Euciroa\
affinis, Verticordia Thiele & Jaeckel, 1931: 247. West Indian Ocean. 693-1134 m. | Vert ambitus]
agulhasensis, Cuspidaria Thiele & Jaeckel, 1931: 253. = Cuspidaria optima Sowerby, 1904
agulhasensis. Lyonsiella Thiele & Jaeckel, 1931: 250. South Africa. 50-250 m. [? Lyonsiella]
alaskana. Lyonsiella Dali, 1895b: 703. East Pacific. 800-3570 m. | Policordia ( Dallicordia)]
alhida, Poromya ( Cetoconcha ) Dali. 1886b: 282. Northwest & West Central Atlantic. 175-1337 m.
[Porotnya ( Cetomya ) \

alcocki. Cuspidaria E.A. Smith, 1894: 170. Indo-Pacific. 60-1150 m. \Cardiomya]
alephtinae, Cribrosoconcha Krylova, 1991: 133. Southeast Pacific, 1014-1058 m. [Cetoconcha]
alleni, Myonera Poutiers, nom. nov. pro Cuspidaria (My oner a) atlantica Allen & Morgan, 1981: 470.

North Atlantic. 1427-4680 m. [Myonera] (see note 1 below).
alta. Cuspidaria Verco, 1908: 198. Southern Australia. 165-275 m. [Cuspidaria ( Rhinoclama )[
alternata, Sphena d’Orbigny in de la Sagra, 1846: 286. West Central Atlantic. 24-340 m. [Cardiomya]
alveata. Cuspidaria Hedley, 1907b: 362. Southwest Pacific. 1500 m. [Cardiomya]
anatinoides , Poromya Forbes, 1844: 191. = Poromya granulata (Nyst & Westendorp, 1839)
andamanica, Cardiomya Preston, 1916b: 99. Indian Ocean. 3-10 m. [Cardiomya]
angasi, Neaera E.A. Smith, 1885: 47. Southern Australia. 238-750 m. [Cuspidaria]
angolensis , Cetoconcha Allen & Morgan, 1981: 528. Southeast Atlantic. 5124 m. | Cetoconcha [
angularis, Neaera Jeffreys, 1876: 498. North Atlantic. 530-3265 m. [ Myonera |
angulata, Pecchiolia Jeffreys, 1882: 933. Northeast Atlantic, Azores. 454-1429m. [Halicardia]
angusticauda, Cardiomya Scarlato, 1972: 124. Northwest Pacific. 135-664m. [Cardiomya]
annandalei. Cuspidaria Preston, 1915: 308. Indian Ocean. Shallow water. [Taxonomic status
uncertain]

antarctica, Pholadomya Hedley, 1916: 28. =? Poromya adelaidis (Hedley, 1916), fide Dell, 1990.
apodema, Cuspidaria Dali, 1916a: 23, nom. nud.\ 1916b: 407. Northeast Pacific. 1098-2900 m.

[Cuspidaria]

approximata. Cuspidaria E.A. Smith, 1896: 373. Indian Ocean. (46?), 400-786 m. [Cuspidaria]
arctica. Neaera M. Sars, 1859: 62. Arctic, North Atlantic. 35-1190 m. [Cuspidaria]
arcuata , Neaera Dali, 1881: 113. West Central Atlantic. 1170 m. [Cuspidaria]
ascoldica, Cuspidaria Scarlato, 1972: 121. Northwest Pacific. 100-800 m. | Cuspidaria |
asiatica , Halonympha Hayami & Kase, 1993: 103. Northwest Pacific. 7-20 m. [ Halonympha |
atlanta, Halonympha Allen & Morgan, 1981: 494. Northwest & East Central Atlantic.’ 2644-3128 m.
| Halonympha\

atlantica. Cuspidaria Allen & Morgan, 1981: 455. Atlantic. 530-2154 m. [Cuspidaria]
atlantica , Cuspidaria ( Myonera ) Allen & Morgan, 1981: 470, non Cuspidaria atlantica Allen &
Morgan, 1981. = Myonera alleni Poutiers, nom. nov. (see note 1 below).
atlantica , Policordia Allen & Turner, 1974: 484. North Atlantic. 458-2186 m. [Policordia]
atlantica , Protocuspidaria ( Bidentaria ) Allen & Morgan, 1981: 499. North Atlantic, Canaries.

1150-4706 m. [Protocuspidaria (Bidentaria)]
attenuata, Neaera Forbes, 1843: 75. = Cuspidaria rostrata (Spengler, 1793)
atypha. Cetoconcha Verrill & Bush, 1898: 814. Northwest Atlantic. 2602 m. [Cetoconcha]
aupouria. Cuspidaria Dell, 1950: 21. Southwest Pacific. 137-805 m. [Cuspidaria ( Rhinoclama )[

ANOMALODESMATA FROM THE TROPICAL PACIFIC

157

australiensis, Verticorciia Hedley, 1907a: 303, non E. A. Smith, 1885. = Verticordia bordaensis Cotton
& Godfrey, 1938.

australiensis, Verticordia E. A. Smith, 1885: 167. Southwest Pacific. 285 m. [Verticordia]
australis , Poromya E. A. Smith, 1885: 54. West Pacific. 83-283 m. \Poromya\
axinoides, Verticordia (Laevicordia) Seguenza, 1876a: 111. Mediterranean. 250-400 m. I Laevicor-
dia 1

azorica, Neaera E.A. Smith, 1885: 41. Mid Atlantic. 1829 m. | ICuspidaria [

halboae, Cardiomya Dali, 1916b: 407. East Central Pacific. 45-170 m. \Cardiomya\

barnardi , Cuspidaria Knudsen, 1970: 139. Atlantic, Indian Ocean. 2178-3828 m. \Cuspidaria\

batialis, Cardiomya lindbergi Scarlato, 1972: 124. = Cardiomya gouldiana (Hinds, 1843)

hehringensis, Neaera Leche, 1883: 438. North Pacific. 18-2900 m. [Cardiomya]

benthedii , Cuspidaria Poutiers, 1984: 289. Indian Ocean. 3700-3716 m. | Cuspidaria ( Rhinoclama)]

berenicensis , Pecchiolia Sturany, 1896: 15. Mediterranean. 700 m. [Haliris]

beringiana, Dermatomya Dali, 1916a: 22, nom. nud.; 1916b: 406. = Poromya (Dermatomva)
tenuiconcha Dali, 1913

bernardi, Bathyneaera Krylova, 1993: 58. Tropical West Pacific. 7800-7870 m. [Bat hy neaera]

hicarinata, Myonera E. A. Smith, 1896: 374. Indian Ocean. 660-1163 m. [My oner a] '

bicarinata , Neaera Jeffreys, 1880: 316, nom. nud.; 1882: 939. Northeast Atlantic. 1262-2004 m.

[Cuspidaria]

bordaensis , Verticordia Cotton & Godfrey, 1938: 149, nom. nov. pro Verticordia australiensis Hedley.

1907, non E.A. Smith, 1885. Southwest Pacific. 16-549 m. | Verticordia]
boucheti , Cetoconcha Poutiers & Bernard sp. nov. West Central Pacific. 300-330 m. [Cetoconcha]
brachyrhynchus , Cuspidaria Sturany, 1901: 263. Red Sea. 375-2160 m. [Cuspidaria]
brazieri, Neaera E.A. Smith, 1885: 51. Southwest Pacific. 4-200 m. | Plectodon]
braziliensis, Cetoconcha Allen & Morgan, 1981: 521. Atlantic. 2250-3730 m. [Cetoconcha]
hraziliensis, Cuspidaria E.A. Smith, 1915: 104. Southwest Atlantic. 72-100 m. [ICuspidaria]
brevirostris , Anatina Brown, 1829: 11. = Cuspidaria cuspidata (Olivi, 1792)
brevirostris, Austroneaera Powell, 1937: 174. Southwest Pacific. 260 m. [Cuspidaria ( Rhinoclama )[
brucei, Cuspidaria Melvill & Standen, 1907: 122. = Cuspidaria undata (Verrill, 1884)
hr uuni. Cardiomya Dell, 1956a: 34. South Pacific. 610 m. [Cardiomya]
buccina. Cuspidaria ( Cuspidaria ) Bernard, 1989: 62. Northeast Pacific. 3585 m. | Cuspidaria [
bulla , Lyonsia Dali, 1878: 61, nom. nud.; 1881: 107. Northwest Atlantic. 3506-5860 m. [Cetoconcha]
butoni, Poromya ( Cetoconcha ) Prashad, 1932: 327. West Pacific. 70-535 m. [Poromya ( Cetomya )\
buttoni, Dermatomya Dali, 1916a: 22, nom. nud.; 1916b: 406. Northeast Pacific. 1 19-1063 m. [Poromya
( Dermatomya )[

caduca, Cuspidaria (Myonera) E. A. Smith, 1894: 170. Indo-Pacific. 50-1134 m. [Myonera ( Rengea )[
caelata, Verticordia Verrill, 1882: 566. = Trigonulina ornata d'Orbigny, 1846
californica, Cuspidaria ( Cardiomya ) Dali, 1886b: 296. East Pacific. 15-640 m. [Cardiomva]
cambrica, Vertisphaera Iredale, 1930: 387-388. Southwest Pacific. 146 m. | Haliris ( Ver'tisphaera )[
canadensis, Poromya (Dermatomya) Bernard, 1969: 2232. = Poromya (Dermatomva) tenuiconcha
Dali, 1913

canariensis, Cuspidaria (Myonera) De Boer, 1985: 102. North Atlantic. 500-2300 m. [Myonera]
capensis, Neaera E.A. Smith, 1885: 45. Southeast Atlantic. 91-564 m. | Cuspidaria |
carinifera, Verticordia Locard, 1898: 208. East Central Atlantic. 2083 m. [Halicardia]
casta, Neaera Hinds, 1843: 77. West Central Pacific. 15 m. [ Cardiomya |
castanea, Poromya Habe, 1952b: 156. Northwest Pacific. 30-880 m.[Poromya\
centohi. Cuspidaria Bouchet & Waren, 1979: 218. Northeast Atlantic, Arctic Ocean (fide Knudsen
1985). 2330-3700 m. [Myonera]

ceylonensis, Cetoconcha Knudsen, 1970: 119. Indian Ocean. 3310 m. [Cetoconcha]
chilensis, Cuspidaria (Luzonia) Dali, 1890: 282. Southeast Pacific. 1238-1895 m. | Cuspidaria ( Luzonia )|
chilensis, Poromya (Dermatomya) Dali, 1908: 430. Southeast Pacific. 821 m. [Poromya ( Dermato¬
mya )\

158

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

chinensis, Neaera Gray in Griffith & Pidgeon, 1834: 12 & 598. West Pacific. 100-250 m. [Cuspidaria]
chuni, Cuspidaria ( Cardiomya ) Thiele & Jaeckel, 1931: 257. Indian Ocean. 400-1134 m. | Cardiomya]
cinerea, Neaera cuspidata var. Jeffreys, 1865: 54. = Cuspidaria cuspidata (Olivi, 1792)
circinata, Neaera Jeffreys, 1876: 497. North and Central Atlantic. 564-4382 m. [Cuspidaria]
cistagemma, Euciroa Kuroda, 1952: 14. = Euciroa crassa Thiele & Jaeckel, 1931
clathrata, Cuspidaria ( Soyomya ) Poutiers & Bernard sp. nov. West Pacific. 230 m. [Cuspidaria
( Soyomya) \

claviculata , Neaera Dali, 1881: 112. West Central Atlantic. 183-985 m. | Halonympha \
cleryana, Sphaenia d'Orbigny, 1842: 708, pi. 83, figs 16-18. Southwest Atlantic. 50-225 m. [ Cardio¬
mya 1

cochinensis, Cuspidaria Preston, 1916a: 39. Indian Ocean. Shallow water. [Taxonomic status
uncertain]

cochlearis, Neaera Hinds, 1844: 98. = Leptomya cochlearis (Hinds, 1844) [Semelidae]
colpodes , Cuspidaria Dautzenberg & Fischer, 1897b: 223. Azores, West Indian Ocean. 693-1644 m.
|? Protocuspidaria ( Bidentaria)\

compressa, Lyonsiella Allen & Turner, 1974: 458. Northeast Atlantic. 119 m. [Lyonsiella]
compressa, Mytilimeria Locard, 1898: 211. Not a Halicardia (see note 2 below).
concentrica, Cuspidaria Thiele, 1912: 233. Antarctic. 351 m. [ Cuspidaria |
concinna. Neaera Hinds, 1843: 77. Habitat unknown. [Cardiomya]
congenita, Neaera E. A. Smith, 1885: 52. West Central Atlantic. 800 m. | Halonympha ]
consimilis, Cuspidaria nobilis Habe, 1961: 146 & App.42. = Cuspidaria nobilis (A. Adams, 1864)
consociata, Neaera E. A. Smith, 1885: 41. West Central Atlantic. 340-850 m. [ Cuspidaria [
contractu, Neaera Jeffreys, 1882: 941. Northeast Atlantic. 1354-2967 m. [Cuspidaria]
convexa, Cuspidaria Pelseneer, 1911: 80. Tropical West Pacific. 100-694 m. [Cuspidaria]
convexa , Cuspidaria ( Cuspidaria ) Prashad, 1932: 329. = Cuspidaria convexa Pelseneer, 1911
cordata, Lyonsiella Verrill & Bush, 1898: 818. Northwest Atlantic. 2602-3338 m. [Policordia]
corpulent a, Neaera costellata var. Dali, 1881: 111. = Cardiomya costellata (Deshayes, 1833)
corrugata , Cuspidaria ( Cuspidaria ) Prashad, 1932: 329. Tropical West Pacific. 38-320 m. [Cuspidaria]
costata, Anatina Sowerby, 1834: 87. Tropical East Pacific, 4-84 m. [Cardiomya]
costata, Neaera Bush, 1885: 472, non Sowerby, 1834. = Cardiomya ornatissima (d’Orbigny, 1846)
costellata , Corbula Deshayes, 1833: 86. North Atlantic, Mediterranean. 5-2000 m. [Cardiomya]
costeminens, Verticordia ( Spinosipella ) Poutiers, 1981: 351. Tropical West Pacific. 750-925 m.
[Spinosipella]

cowani. Cuspidaria ( Cuspidaria ) Bernard, 1967: 2629. Northeast Pacific. 1318 m. [ Cuspidaria [

crassa, Euciroa Thiele & Jaeckel, 1931: 248. Indo-Pacific. 136-1463 m. [Euciroa]

crebrilirata, Verticordia ( Verticordia ) Prashad, 1932: 324. Tropical West Pacific. 564 m. [Haliris]

cuneata , Verticordia ( Vertambitus ) Kuroda, 1952: 8. Northwest Pacific. 200 m. [Vertamhitus]

curta, Lyonsiella Poutiers, 1984: 298. West Indian Ocean. 3700-3716 m. [ Lyonsiella |

curta. Neaera Jeffreys, 1876: 495, nom. nud.; 1882: 943. Arctic, North Atlantic. 32-2338 m.

[ Cardiomya \

curta , Neaera cuspidata var. Jeffreys, 1865: 54. = Cuspidaria cuspidata (Olivi, 1792)
curta, Neaera multicostata var. Verrill, 1882: 560, non Neaera curta Jeffreys, 1882. = Cardiomya
striata (Jeffreys, 1876)

curta, Poromya Sowerby, 1904: 17. Southwest Indian Ocean. 805 m. [Poromya]
cuspidata, Neaera Hinds, 1843: 76, non Olivi, 1792. Northwest Pacific. 154 m. Nom. dub.
cuspidata, Tellina Olivi, 1792: 101. Northeast & East Central Atlantic, Mediterranean. 20-1850 m.
[Cuspidaria]

cymata, Poromya Dali. 1890: 289. Southwest Atlantic. 72-108 m. [Poromya]

dalli, Euciroa Pilsbry, 1911: 523. = Euciroa eburnea (Wood-Mason & Alcock, 1891)

dautzenbergi, Myonera Prashad, 1932: 334. Northwest Pacific to Indonesia. 715-959 m. [Myonera]

delectabile, Euciroa Dell, 1956b: 42. = Euciroa galatheae (Dell, 1956)

delli, Cuspidaria Knudsen, 1970: 141. Southwest' Pacific. 4400 m. [ Cuspidaria [

ANOMALODESMATA FROM THE TROPICAL PACIFIC

159

demistriata, Cuspidaria (Myonera) Allen & Morgan, 1981: 469. = Bathyneaera hadalis (Knudsen
1970)

densicostata, Verticordia Locard, 1898: 202. Central Atlantic. 1002-2325 m. \Policordia\
depressa, Neaera Jeffreys, 1882: 940. East Atlantic, Mediterranean. 164-2351 m. \Halonympha]
deshayesiana, Verticordia Fischer, 1862: 35. Indo-Pacific. 40-693 m. \Spinosipella\
diagonalis, Cuspidaria (Tropidomya) Allen & Morgan, 1981: 487. Southeast Atlantic. 527-542 m.
| Cuspidaria ( Tropidomya ) |

didyma. Neaera Hinds, 1843: 78. Tropical East Pacific. 18-48 m. \Cardiomya\
diomedea, Policordia Dali, Bartsch & Rehder, 1938: 217. Mid-North Pacific. 44-530 m. \Policordia\
disa, Cardiomya (Labromvsa) Bernard, 1989: 64. Northeast and Northwest Pacific, tropical West
Indian Ocean, tropical West Atlantic. 3700-6850 m. \Bathyneaera\
dispar, Cuspidaria ( Myonera ) Dali, Bartsch & Rehder, 1938: 225. Northwest & Mid-North Pacific
400-914 m. [ Myonera \

dissociata, Cuspidaria Sturany, 1901: 262. Red Sea. 805 m. [Cuspidaria]

dorsirecta, Cuspidaria Verco, 1908: 198. Southeast Indian Ocean, Southern Australia. 73-1150 m.

| Cuspidaria ( Rhinoclama ) ]

dubia, Cuspidaria ( Myonera ) Pelseneer, 1911: 80. Indo-Pacific. 200-2798 m. | Cuspidaria ( Rhinoclama)\
dubia, Cuspidaria (Rhinoclama) Prashad, 1932: 333. = Cuspidaria ( Rhinoclama ) dubia (Pelseneer,
1911)

dulcis, Cuspidaria (Cardiomya) Pilsbry & Lowe, 1932: 10. = Cardiomya coslata (Sowerby, 1834)
eburnea , Verticordia (Euciroa) Wood-Mason & Alcock, 1891: 447. Indo-Pacific. 340-1500 m. \Euciroa\
ecuadoriana, Cuspidaria (Cardiomya) Olsson, 1961: 465. Tropical East Pacific. 55-146 m. | Cardiomya]
elegans, Cribrosoconcha Krylova, 1991: 135. Southeast Pacific, 218-575 m. \Cetoconcha\
elegans, Neaera Hinds, 1843: 76. Indo-Pacific. 13-200 m. \Cuspidaria\
elegant issima, Euciroa Dali, 1878: 61, nom. nud. = Euciroa elegantissima (Dali, 1881)
elegantissima, Verticordia Dali, 1881: 106. West Central Atlantic. 300-1500 m. \Euciroa\
elliptica , Cuspidaria (Cuspidaria) Di Geronimo, 1974: 157. Mediterranean. 2300 m. [ICuspidaria]
elongata, Poromya (Cetoconcha) Dali, 1886b: 283. West Central Atlantic. 183-364 m. | Poromya
( Cetomya )\

equatorialis , Poromya (Dermatomya) Dali, 1908: 429. = Poromya ( Dermatomya) mactroides Dali,
1889

ericia, Verticordia Hedley, 1911: 96. South and Central Indo-Pacific. 146-805 m. \Spinosipella]
erma , Cuspidaria Cotton, 1931: 347. Southern Australia. 148-550 m. [Cuspidaria]
exarata, Cuspidaria Verco, 1908: 199. Southern Australia. 190 m. | Cuspidaria \
excoriata, Verticordia Poutiers, 1984: 297. West Indian Ocean. 3700 m. [Vertambitus]
exigua. Cetoconcha Poutiers & Bernard, sp. nov. West Pacific. 970 m. | Cetoconcha |
exigua , Neaera Jeffreys, 1876: 496. Arctic. North Atlantic. 640-1836 m. [ Cuspidaria )
eximia, Poromya Pelseneer, 1911: 78. Indo-Pacific. 50-1190 m. | Poromya (Cetomya)\
eximia, Poromya (Cetoconcha) Prashad, 1932: 327. = Poromya (Cetomya) eximia Pelseneer, 1911
expansa, Verticordia ? Prashad, 1932: 324. West Pacific. 835 m. [ Verticordia J
extenta , Policordia (Angus tebranchia) Ivanova, 1977: 182. Northwest Pacific. 8220-8430 m. | Policordia
( August ebranchia)]

fairchildi, Cuspidaria Suter, 1908: 372. Southwest Pacific. 137-640 m. | Cuspidaria]
fallax, Neaera E. A. Smith, 1885: 49. West Pacific. 283 m. [? Cardiomya ( Kurodamya )]
ferruginea. Halicardia Di Geronimo, 1974: 158. Mediterranean. 2300-2400 m. | Halicardia ]
filatovae, Cardiomya Scarlato, 1972: 127. Northwest Pacific. 3350 m. | Cardiomya ]
filatovae, Cuspidaria Bernard, 1979: 14. Northeast Pacific. 3500-4882 m. [ Cuspidaria (Rhinoclama)]
filocarinata, Neaera E. A. Smith, 1885: 44. = Cuspidaria (Rhinoclama) notabilis (Jeffreys, 1876)
finlayi , Austroneaera Powell, 1937: 175. Southwest Pacific. 110 m. | Cuspidaria ( Rhinoclama )]
fischeri, Mytilimeria Jeffreys, 1880: 316; 1881: 384. Nom. nud. = Halicardia fischeri (Locard, 1898)
fischeri, Mytilimeria Locard, 1898: 212 (ex Jeffreys MS). Northeast Atlantic, 1140-2650 m.
[Halicardia]

160

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

fischeriana, Verticordia Dali, 1881: 106. West Atlantic, East Pacific. 80-510 m. \Haliris\
flexuosa, Mytilimeria Verrill & Smith in Verrill, 1881: 302. North Atlantic. 137-2330 m. \Halicardia [
flexuosa , Poromya Yokoyama, 1922: 173. Northwest Pacific. 30-350 m. \Poromya\
fluctuosa, Myonera Kuroda, 1948: 25.= Myonera caduca (E. A. Smith, 1894)
formosa, Cuspidaria Verrill & Bush, 1898: 803. Northwest Atlantic. 2173 m. \Cuspidaria\
formosa, Lyonsia Jeffreys, 1874: 112, nom. nud.\ 1880: 316, nom. nud.\ 1882: 930. Atlantic,
Mediterranean; Indian Ocean & Pacific? (Morton, 1985a) 366-3783 m. [? Lyonsiella [ (see
note 3 below).

forticostata , Cuspidaria ( Cardiomya ) Sowerby, 1904: 18. Southwest Indian Ocean. 805 m. \Cardiomya\
fortisculpta, Cuspidaria ( Cardiomya! ) Kuroda, 1948: 20. Northwest Pacific. 100-200 m. \Cardiomya
( Kurodamya)\

fragilis, Lyonsiella Allen & Turner, 1974: 456. North Atlantic. 1102-1470 m. [ Lyonsiella]
fragilis, Neaera A. Adams, 1856: 226.= Theora fragilis (A. Adams, 1856) [Semelidae]
fragilissima. Neaera E. A. Smith. 1885: 53. Southwest Indian Ocean. 349 m. \Cardiomya\
fraterna, Cuspidaria Verrill & Bush, 1898: 803. Northwest Atlantic. 552-1800 m. [Cuspidaria]
frieli, Lyonsiella Allen & Turner, 1974: 440. North Atlantic. 3301-4429 m. \Laevicordia\
fujitai, Cuspidaria ( Cardiomya ) Kuroda, 1948: 18. Northwest Pacific. 100-300 m. \Cardiomya\
galatheae, Cetoconcha Knudsen. 1970: 120. Southwest Pacific. 4400 m. [Cetoconcha]
galatheae , Laevicordia Knudsen, 1970: 128. West Indian Ocean. 4820 m. [Laevicordia]
galatheae , Questimya Dell, 1956a: 33. Southwest Pacific. 475-622 m. \Euciroa\
garretti , Myonera Dali, 1908: 434. Tropical East Pacific. 1644 m. \Myonera\
gemma, Cardiomya Verrill & Bush, 1898: 809. = Cardiomya costellata (Deshayes, 1833)
gemma. Lyonsiella Verrill, 1880: 396. North Atlantic, Mediterranean. 73-3917 m. \Policordia\
gigantea. Cuspidaria (Cuspidaria) Prashad, 1932: 329. Indo-Pacific. 100-1030 m. [Cuspidaria]
gigantea , Neaera Verrill, 1884: 223. Northwest Atlantic. 2286-3506 m. [Myonera]
gilchristi, Poromya Sowerby, 1904: 15. Southwest Indian Ocean. 86-99 m. [ Poromya |
gilchristi , Cuspidaria ( Cardiomya ) Sowerby, 1904: 18. Southwest Indian Ocean. 73-229 m. | Cardio¬
mya |

glacialis. Neaera G. O. Sars, 1878: 88. Arctic, North Atlantic. 71-2500 m. [ Cuspidaria \
globulosa, Bathyneaera Krylova, 1993: 55. North Atlantic. 4440-4480 m. [ Bathyneaera ]
gloriosa, Poromya ( Cetoconcha ) Prashad, 1932: 326. Tropical West Pacific. 150-400 m. [Cetoconcha]
glypta, Cardiomya Bush in Verrill & Bush, 1898: 810, nom. nov. pro Neaera costata Bush, 1885 non
Sowerby, 1834. = Cardiomya ornatissima (d’Orbigny, 1846)
gouldi, Halicardia Dali, Bartsch & Rehder, 1938: 218. Mid-North Pacific. Depth unknown.
[Halicardia]

gouldiana, Neaera Hinds, 1843: 77. West Pacific. 13-1030 m. [Cardiomya]
gracilis, Cuspidaria “(G. O. Sars)” Nordsieck, 1969: 176. = Cuspidaria gracilis (Jeffreys, 1882)
gracilis, Neaera Jeffreys, 1882: 938. Arctic, Northeast Atlantic. 70-2032 m. [Cuspidaria]
granatina , Poromya (?) Dali, 1881: 109. = Basterotia quadrata (Hinds, 1843) [Sportellidae]
grandis , Lyonsiella E. A. Smith, 1885: 74. South Atlantic. 3470 m. [Policordia]
granifera, Questimya Cotton, 1931: 345. Southwest Pacific. 20-250 m. [Euciroa]
granosissima, Poromya Sowerby, 1904: 16. Southwest Indian Ocean. 49-165 m. [Poromya]
granulata, Corbula? Nyst & Westendorp, 1839: 6. North Atlantic, Mediterranean. 30-1262 m.
[Poromya]

granulata, Ectorisma Tate, 1892: 127. = Poromya laevis E. A. Smith, 1885.

granulata, Neaera Dali, 1881: 111. West Central Atlantic. 37-274 m. [ Plectodon \

granulata, Verticordia Seguenza, 1858: 356. North & Central Atlantic, Mediterranean. 55-1245 m.

| Haliris /

granulifera, Pecchiolia Verrill, 1885: 434. Northwest Atlantic. 2450-3400 m. [Verticordia]
granuloderma, Poromya granuloderma Scarlato, 1981: 428. Northwest Pacific. 300-664 m. [Poromya]
greenii , Cuspidaria ( Cardiomya ) E. A. Smith, 1889: 423. Northeast Atlantic. 1829 m. [Cardiomya]
guineensis, Cuspidaria Knudsen, 1970: 143. East Central Atlantic. 2550 m. ^.Cuspidaria ]

ANOMALODESMATA FROM THE TROPICAL PACIFIC

161

guineensis, Verticordia Thiele & Jaeckel, 1931: 246. East Central Atlantic. 2278 m. ( Verticordia]
haasi. Cuspidaria Knudsen, 1970: 145. Tropical East Pacific. 3570 m. \Cuspidaria\
hadalis. Cuspidaria Knudsen, 1970: 146. West Atlantic, tropical West Indian Ocean, West Pacific.
1135-8430 m. \Bathyneaera\

halei. Cuspidaria Cotton & Godfrey, 1938: 158. Southern Australia. 238-550 m. | Cuspidaria ]
halimera, Cuspidaria ( Leiomya ) (( Rhinoclama )) Dali, 1886b: 300. North Atlantic. 1337-2934 m.

| Cuspidaria ( Rhinoclama ) ]

hancocki, Verticordia ( Trigonulina ) Bernard, 1969: 2233. Tropical East Pacific. 73-1 10 m. [Trigonulina]
hawaiensis , Cuspidaria ( Cuspidaria ) Dali, Bartsch & Rehder, 1938: 226. Mid-North Pacific. 468-
874 m. \Cuspidaria\

hawaiensis, Euciroa “ Dali. Bartsch & Redher ” Habe, 1964: 212, nom. null. Error for Euciroa gouldi
Dali, Bartsch & Rehder, 1938.

hayashii , Poromya Habe, 1958: 175, 180. Northwest Pacific. 50-200 m. \Poromya]
hindsiana , Neaera A. Adams, 1864: 207. Northwest Pacific. 50-200 m. \Cuspidaria\
hirasei, Cuspidaria Kuroda, 1948: 10. = Cuspidaria steindachneri Sturany, 1901
horrida, Laevicordia Allen & Turner, 1974: 507. Northeast Atlantic. 2862-2886 nr. \Laevicordia ]
houhricki , Halicardia Poutiers & Bernard sp. nov. Borneo. 1629 m. | Halicardia]
houbricki, Poromya ( Poromya ) Bernard, 1989: 65. Northeast Pacific. 95-100 m. \Poromya\
hyalina. Neaera Hinds [ex Sowerby MS), 1843: 76. Northwest Pacific. Depth unknown. \?Cuspidaria\
hyalina. Poromya ( Dermatomya ) Bernard, 1989: 66. Northeast Pacific. 4882 m. | Poromya (Derma-
tomya) ]

illevis, Poromya Hedley, 1913: 265, nom. nov. pro Ectorisma granulaia Tate, 1892 non Poromya
granulata (Nyst & Westendorp, 1839). = Poromya laevis E.A. Smith, 1885
imhricata , Neaera Jeffreys, 1880: 316; 1881: 383; 1882: 942. Nom. nud. = Cuspidaria imbricata
Locard, 1898

imhricata. Cuspidaria Locard [ex Jeffreys MS), 1898: 187. Northeast Atlantic. 1107-1960 m.

[Cuspidaria]

indica. Cetoconcha Ray, 1951: 187. Indian Ocean. Depth unknown. \Cetoconcha\
infelix. Cuspidaria Thiele, 1912: 233. Antarctic. 91-752 m. [Cuspidaria]
inf lata , Neaera Jeffreys, 1882: 942. North Atlantic. 1000-2000 m. | Cuspidaria (Leiomya)]
inornata. Verticordia Thiele & Jaeckel. 1931: 245. Southwest Indian Ocean. 49-229 m. | Verticordia]
insculpta, Pecchiolia Jeffreys, 1874: 112, nom. nud.-, 1882: 932. = Policordia gemma (Verrill, 1880)
insculpta. Verticordia (Laevicordia) Seguenza, 1876a: 112. Mediterranean. Depth unknown. | Laevi¬
cordia ]

insolita. Policordia Allen & Turner, 1974: 502. North Atlantic. 1546-2178 m. | Policordia |
intermedia, Cetoconcha [eximia var?) Habe, 1952b: 158. = Poromya ( Cetomya ) eximia Pelseneer, 1911
iridella, Cuspidaria ( Pseudoneaera ) Kuroda, 1948: 25. = Pseudoneaera semipellucida (Kuroda, 1948)
iridescens, Neaera Hinds, 1843: 78. = Theora iridescens (Hinds, 1843) [Semelidae]
isocar dioides, Poromya ( Cetoconcha ) Dautzenberg & Fischer, 1897a: 30. = Poromya tornata (Jeffreys,
1876)

isolirata, Cardiomya Bernard. 1969: 2231. = Cardiomya pectinata (Carpenter, 1865)
iturupica, Cardiomya Scarlato, 1972: 127. Northwest Pacific. 414 m. [ Cardiomya ]
ivanovae, Policordia Poutiers & Bernard, nom. nov. pro Policordia japonica Ivanova, 1977. non Habe.
1961. Northwest Pacific. 3042 m. | Policordia 1

jajfaensis, Verticordia Cotton & Godfrey, 1938: 151. Southwest Pacific. 183-549 m. [Haliris]

japonica. Cetoconcha Habe, 1952b: 159. West Pacific. 175-650 m. [Cetoconcha]

japonica. Cuspidaria Kuroda, 1948: 14. Northwest Pacific. 100-300 m. [ Cuspidaria ]

japonica , Lyonsiella Habe, 1952a: 269, nom. nud.; 1961: 145 & App.41. = Policordia pilula (Pelseneer.

191 i)

japonica , Policordia ( Latebranchia ) Ivanova, 1977: 193, non Habe, 1961. = Policordia ivanovae
Poutiers & Bernard

japonica, Verticordia A. Adams, 1862: 224. = Spinosipella deshayesiana (Fischer, 1862)

162

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

jeffreysi , Lyonsiella Friele, 1879: 269. Northeast Atlantic, Arctic Ocean {fide Knudsen, 1985).
1198-4429 m. [Policordia]

jeffreysi, Lyonsiella E. A. Smith. 1885: 73, non Friele. 1879. = Laevicordia smidti (Friele, 1886)
jeffreysi , Neaera Dali, 1881: 111. Northwest & West Central Atlantic. 849-2022 m. \Cuspidaria\
jugosa, Neaera G. O. Sars. 1878: 88, non Wood. 1850. = Cuspidaria lamellosa (G. O. Sars, 1878)
jugosa, Neaera Wood, 1850: 272. Northeast Atlantic, Mediterranean. 200-1100 m. [1 Cuspidaria]
kashimana, Cardiomya Okutani & Sakurai, 1964: 20. Northwest Pacific. 200-1030 m. \Cardiomya]
kawamurai, Cuspidaria Kuroda, 1948: 11. = Cuspidaria {Cuspidaria) gigantea Prashad, 1932
kerguelensis, Neaera E. A. Smith, 1885: 46. Antarctic. 60-574 m. [Cuspidaria]
knudseni. Cuspidaria {Cardiomya) Allen & Morgan, 1981: 466. Southwest & North Atlantic.
1661-3806 m. [Cardiomya]

korenii, Embla Loven, 1846: 200. = Poromya granulata (Nyst & Westendorp, 1839)
kurilensis, Dermatomya Scarlato, 1981: 427. = Poromya (Dermatomya) tenuiconcha Dali, 1913
kurodai, Cuspidaria Okutani, 1975a: 196. Northwest Pacific. 158-177 m. [Cuspidaria]
kurohijii , Cuspidaria Okutani, 1972: 126. Northwest Pacific. 130-190 m. | Cuspidaria (Soyomya)]
kyushuensis. Cuspidaria Okutani, 1962: 35. Northwest Pacific. 520-760 m. [Cuspidaria]
lactea, Neaera costellata var. Jeffreys, 1865: 50. = Cardiomya costellata (Deshayes, 1833)
laevigata, Policordia { Angustebranchia ) Ivanova, 1977: 184. West Pacific. 8160-8900 m. | Policordia
( A ngustebranchia ) ]

laevis, Poromya E. A. Smith, 1885: 55. South Pacific. 15-805 m. | Poromya]
lamellifera, Neaera Dali, 1881: 113. Northwest Atlantic. 153-457 m. [My oner a]
lamellosa , Neaera M. Sars, 1859: 62; 1869: 257. Nom. nud. = Cuspidaria lamellosa (G. O. Sars, 1878)
lamellosa , Neaera G. O. Sars, 1878 {ex M. Sars MS): 88. North Atlantic. 91-1015 m. [Cuspidaria]
lamothei, Verticordia Dautzenberg & Fischer, 1897a: 30. East Atlantic, West Indian Ocean. 126-
608 m. [Haliris]

lanieri, Cuspidaria Strong & Hertlein, 1937: 163. East Central Pacific. 37-402 m. [Cardiomya]
lata, Neaera Hinds, 1843: 79. = Theora lata (Hinds, 1843) [Semelidae]
latesulcata, Neaera Tenison-Woods, 1878: 123. West Pacific. 30 m. [ Cuspidaria ]
laticella, Myonera Dali, 1886b: 305. West Atlantic. 3111 m. [ Bathyneaera ]

ledaeformis. Cuspidaria Dautzenberg & Fischer, 1897a: 29. Central Atlantic. 1300-1600 m.

[IHalonympha]

leiomyoides, Cuspidaria Poutiers, 1981: 340. Tropical West Pacific. 415-510 m. [Halonympha]
leonina, Dermatomya Dali, 1916a: 22, nom. nud:, 1916b: 406. = Poromya {Dermatomya) tenuiconcha
Dali, 1913

levifrons, Cuspidaria Cotton, 1930: 235. Southern Australia. 550 m. [Cardiomya ( Kurodamya)]
ligula, Cuspidaria Yokoyama, 1922: 169. Northwest Pacific. 10-300 m. [Plectodon]
limatula. Neaera Dali, 1881: 112. Northwest & West Central Atlantic. 230-1000 m. | Myonera ]
lindbergi, Cardiomya Scarlato, 1972: 125. = Cardiomya gouldiana (Hinds, 1843)
lisbethae. Policordia Knudsen, 1970: 132. East Central Atlantic. 2690 m. [Policordia]
lischkei, Cuspidaria {Myonera) E. A. Smith, 1891: 438. Northwest Pacific. 3429 m. [ Myonera [
longirostris, Anatina Lamarck, 1818: 463. = Cuspidaria rostrata (Spengler, 1793)
lubangensis, Cuspidaria Poutiers, 1981: 348. Tropical West Pacific. 191-195 m. [Cuspidaria]
lucifuga, Cuspidaria P. Fischer. 1887: 1155, nom. nud:, Locard, 1898: 184. = Cuspidaria undata
(Verrill, 1884)

Ivrata, Neaera Hinds, 1844: 97. = Raeta lyrala (Hinds, 1844) [Mactridae]
macrorhynchus, Cuspidaria E. A. Smith, 1895: 12. Indo-Pacific. 400-1190 m. | Cuspidaria [
mactroides, Poromya (Dermatomya) Dali, 1889: 448. East Pacific. 637-3060 m. [Poromya ( Derma¬
tomya )[

maculata, Policordia (Angustebranchia) Ivanova, 1977: 184. West Pacific. 9000-9050 m. | Policordia
( A ngustebranchia ) [

magnifica. Lyonsiella Dali, 1913: 595. Tropical East Pacific. 115-300 m. [Lyonsiella]

Source : MNHN. Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

163

major, Cuspidaria rostrata var. Dautzenberg & Fischer, 1897b: 220. = Cuspidaria rostrata (Spengler,
1793)

makiyamai, Verlicordia (Hal iris) Habe in Kuroda, 1952: 10. = Haliris ( Setaliris ) pygmaea (Kuroda,
1952)

malespinae, Cetoconcha Dali, 1916a: 22, nom. nud.; 1916b: 407. = Poromya (Cetomya) malespinae
Ridewood, 1903

malespinae, Poromya Ridewood, 1903: 272. Northeast Pacific. 2104-2871 m. [Poromya (Cetomya)]
maoria, Halicardia Dell, 1978: 162. Southwest Pacific. 676-713 m. | Halicardia \
margarita, Poromya ( Cetoconcha ) Dali, 1886b: 284. West Central Atlantic. 715-1864 m. [Cetoconcha]
marmorea, Cuspidaria (Rhinoclama) Bernard, 1989: 64. = Cuspidaria (Rhinoclama) fdatovae (Bernard,
1979)

maxima, Cuspidaria Dautzenberg & Fischer, 1897a: 28. East Atlantic. 1850-3200 m. [ Cuspidaria ]
media, Cuspidaria Verrill & Bush, 1898: 800. Northwest Atlantic. 115-283 m. [Cuspidaria]
media, Lyonsiella Okutani, 1962: 30. = Policordia pilula (Pelseneer, 1911)
mediopacifica, Euciroa Kosuge, 1979: 35. Central Pacific. 170-370 m. [? Euciroa]
meridionalis , Neaera E. A. Smith, 1885: 43. South Indian Ocean. 3566 m. | Cuspidaria \
mexicana, Myonera Knudsen, 1970: 134. East Pacific. 2110-3557 m. [My oner a]
microdonta. Poromya ( Cetomya ) Dali, 1890: 290. = Poromya tornata (Jeffreys, 1876)
microrhina, Cuspidaria rostrata var. Dali, 1886b: 295. Northwest Atlantic. 183-931 m. [Cuspidaria]
millegemmata, Euciroa Kuroda & Habe in Kuroda, 1952: 14. West Pacific. 100-365 m. | Euciroa |
minor, Pseudoneaera Thiele & Jaeckel, 1931: 258. West Indian Ocean. 50 m. [ Pseudoneaera [
mitis , Cuspidaria (Cuspidaria) Prashad, 1932: 328. Indo-Pacific. 500-1620 m. [Cuspidaria]
moeshimaensis, Verlicordia (Haliris) Habe, 1953: 133. = Haliris multicostata (A. Adams, 1862)
moluccana, Neaera Adams & Reeve, 1850: 84. = Cuspidaria elegans (Hinds, 1843)
monosteira, Cuspidaria (?) Dali, 1890: 281. West Central Atlantic. 850-1864 m. [Verticordia]
morelandi, Cuspidaria Dell, 1956b: 39. South Pacific. 238-549 m. [Cuspidaria]
morioria, Cuspidaria Dell, 1956b: 40. South Pacific. 238 m. [1 Cuspidaria]
morrisae, Cuspidaria Poutiers & Bernard sp. nov. West Pacific. 230 m. | Cuspidaria ]
multicarinata, Cuspidaria (Cardiomya) Prashad, 1932: 322. = Cardiomya alcocki (E. A. Smith, 1894)
multicostata, Neaera Verrill & Smith in Verrill, 1880: 398. = Cardiomya striata (Jeffreys, 1876)
multicostata, Verticordia A. Adams, 1862: 224. West Pacific. 50-450 m. [Haliris]
murrayi, Lyonsiella Knudsen, 1967: 297. West Indian Ocean. 1207-1463 m. [Policordia]
murrayi, Neaera E. A. Smith, 1885: 319. Mid-North Pacific. 5304 m. [Myonera ( Rengea)]
nasuta, Cuspidaria Sowerby, 1904: 16, non A. Adams, 1864. = Cuspidaria capensis (E. A. Smith, 1885)
nasuta, Neaera A. Adams, 1864: 207. Northwest Pacific. 100-150 m. | Cuspidaria ]
natalensis, Cuspidaria Knudsen, 1970: 148. Southwest Indian Ocean. 2640 m. [Cuspidaria]
neaeroides , Poromya Seguenza, 1876b: 270. North and Central Atlantic, Mediterranean. 208-1350 m.
[Poromya]

neozelanica, Eetorisma Dell, 1956b: 43. South Pacific. 549 m. [Poromya]

niasensis, Poromya Thiele & Jaeckel, 1931: 252. East Indian Ocean. 660 m. I? Poromya (Cetomya)]
nipponensis, Halicardia Okutani, 1957: 30. Northwest Pacific. 400-1500 m. [Halicardia]
nipponica, Cuspidaria (Cardiomya) abyssicola Okutani, 1962: 35. Northwest Pacific. 360-1480 m.
| Cardiomya \

nitens, Cuspidaria Locard, 1898: 181. Northeast Atlantic. 905-2200 m. [Cuspidaria ( Rhinoclama )]
nitida, Poromya Adams & Reeve, 1850: 43. = Leptomya nitida (Adams & Reeve, 1850) [Semelidae]
nitida, Thracia Verrill, 1884: 221. = Cetoconcha bulla (Dali, 1881)
nobilis, Neaera A. Adams, 1864: 207. West Pacific. 50-300 m. | Cuspidaria ]

notabilis , Neaera Jeffreys, 1876: 497. Northeast & East Central Atlantic. 1100-4734 m. [Cuspidaria
( Rhinoclama )]

novemcostatus, Hippagus Adams & Reeve, 1850: 76. =? Trigonulina ornata d'Orbigny, 1846
nybelini, Cuspidaria Odhner, 1960: 381. = Cuspidaria undata (Verrill, 1884)

164

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

oahuensis, Policordia "Dali, 1895” Higo & Goto, 1993: 679. = Allogramma oahuensis (Dali, 1913)
[?Lyonsiidae]

obesa, Neaera Loven, 1846: 202. Arctic, North Atlantic. 73-4453 m. \Cuspidaria\
obliqua , Cuspidaria ( Cardiomya ) De Boer, 1985: 101. North Atlantic. 60-245 m. \Cardiomya\
obliqueovata, Policordia ( Latebranchia ) Ivanova, 1977: 189. West Pacific. 1100-1640 m. | Policordia/
obtusirostris, Cuspidaria Okutani, 1962: 34. Northwest Pacific. 620-1640 m. [ Cuspidaria ]
occidua, Cuspidaria Cotton, 1931: 347. Southern Australia. 132-148 m. [Cuspidaria]
ochotensis , Cardiomya Scarlato, 1972: 125. Northwest Pacific. 144-207 m. \Cardiomya]
ochotensis, Poromya granuloderma Scarlato, 1981: 429. = Poromya granuloderma Scarlato, 1981
ochotica , Policordia Scarlato, 1981: 419. West Pacific. 660 m. | Policordia ( Dallicordia )]
octaporosa , Cuspidaria ( Myonera ) Allen & Morgan, 1981: 476. North Atlantic. 3459-5000 m.
\Octoporia\

okezoko , Cuspidaria Okutani, 1985: 146. Northwest Pacific. 400 m. \Cuspidaria\
okutanii, Cardiomya behringensis Scarlato, 1972: 122. = Cardiomya behringensis (Leche, 1883)
olivacea , Policordia Poutiers sp. nov. Tropical West Pacific. 980-1080 m. | Policordia \
oldroydi, Cuspidaria ( Cardiomya ) Dali in Oldroyd, 1924: 33. = Cardiomya pectinata (Carpenter, 1864)
opalina, Neaera Hinds, 1843: 78. = Theora opalina (Hinds. 1843) [Semelidae]
optima, Cuspidaria Sowerby, 1904: 17. Southwest Indian Ocean. 44-564 m. [Cuspidaria]
optima, Verticordia Sowerby, 1894a: 39. = Euciroa eburnea (Wood-Mason & Alcock, 1891)
orbiculata, Verticordia ( Laevicordia ) Seguenza, 1876a: 112. Mediterranean. Depth unknown. | Laevi-
cordia |

oregonensis , Poromya Ridewood, 1903: 274. Norn. dub.

orientalis , Cuspidaria ( Cardiomya ) Thiele & Jaeckel, 1931: 257. Indian Ocean. 50 m. [Cardiomya]
orientalis , Poromya Thiele & Jaeckel, 1931: 251. West Indian Ocean. 693-1644 m. [ Poromya
( Cetomya )\

ornata. Trigonulina d'Orbigny, 1846: 292. West Atlantic, East Pacific. 18-850 m. [Trigonulina]
ornatissima, Sphena d'Orbigny in de la Sagra, 1846: 286. West Atlantic. 34-620 m. [Cardiomya]
ovata, Policordia ( Latebranchia ) Ivanova, 1977: 191. West Pacific. 5740-6040 m. [Policordia]
pacifica, Euciroa Dali, 1895b: 688. Mid-North Pacific. 435-910 m. [Euciroa]
pacijica, Lyonsiella Dali, 1908: 428. Mid Pacific. 2090 m. [Laevicordia]
paleifera, Bathyneaera Krylova, 1993: 57. Tropical West Atlantic. 6800-8330 m. [Bathyneaera]
pailoloana, Cuspidaria ( Myonera ) Dali. Bartsch & Rehder, 1938: 225. Mid-North Pacific. 506-604 m.
[Myonera]

panamensis, Cuspidaria Dali, 1908: 432. Tropical East Pacific. 915-1281 m. [Cuspidaria]
papyracea, Lyonsiella E. A. Smith, 1885: 73. Southeast Indian Ocean. 3562 m. [Policordia]
papyria, Neaera Jeffreys, 1876: 498. Northeast Atlantic. 2652 m. [ICuspidaria]
parapodema, Cuspidaria Bernard, 1969: 2232. East Pacific. 53-320 m. | Cuspidaria \
parkeri. Cuspidaria Knudsen, 1970: 150. Tropical East Pacific. 2790-2817 m. [Cuspidaria]
parthenopaea, Cumingia Tiberi, 1855: 10. = Poromya granulata (Nyst & Westendorp, 1839)
parva, Cuspidaria Verrill & Bush, 1898: 801. North & Central Atlantic. 257-4659 m. [Cuspidaria]
parva , Lyonsiella Okutani, 1962: 29. Northwest Pacific. 1230-1350 m. [Lyonsiella]
patagonica, Neaera E. A. Smith, 1885: 39. Southeast Pacific. 302 m. [Cuspidaria]
paucistriata , Myonera Dali, 1886b: 302. Atlantic. 600-3806 m. [Myonera]

paucistriata, Neaera “Dali” Bush, 1885: 473, nom. nud. = Cardiomya gemma Verril & Bush, 1898
pectinata, Neaera Carpenter, 1864: 602, 637, nom. nude, 1865: 54. East Pacific. 45-210 m. [Cardiomya]
pellucida, Neaera Stimpson, 1853: 21. Northwest Atlantic. 95-944 m. [Cuspidaria]
pelseneeri, Cetoconcha Pelseneer, 1911: 79. Habitat unknown. [Cetoconcha]
pergranosa, Poromya Pelseneer, 1911: 78. = Poromya australis E. A. Smith, 1885
pergranosa, Poromya ( Poromya ) Prashad, 1932: 326. = Poromya australis E. A. Smith, 1885
periplomoides. Cuspidaria Sakurai & Habe in Habe, 1961: 146 & App. 41. Northwest Pacific. 200 m.
[TPseudoneaera]

perla. Poromya Dali. 1908: 428. Northeast Pacific. 2071-3518 m. [Perlaporomya]

Source

ANOMALODESMATA FROM THE TROPICAL PACIFIC

165

perplexa, Lyonsiella Allen & Turner, 1974: 437. West Atlantic. 2041-4429 m. \Lyonsiella\
perplicata , Verticordia Dali, 1890: 278. East Pacific. 1000-1500 m. [Halicardia]
perrostrata, Neaera ornatissima var. Dali, 1881: 110. West Atlantic. 35-761 m. \Cardiomya\
persculpta, Cuspidaria ( Cardiomya ) Prashad, 1932: 332. = Cardiomya alcocki (E. A. Smith, 1894)
perversa, Verticordia Dali, 1886b: 289. Northwest Atlantic. 1337 m' \ Verticordia\
philippinensis. Halicardia Poutiers, 1981: 353. Tropical West Pacific. 592-610 m. \Halicardia\
philippinensis, Neaera ( Rliinomya ) A. Adams, 1864: 207, nom. nud.; non Neaera philippinensis Hinds,
1843. = Cuspidaria ( Rhinoclama ) adamsi Heppell & Morgan, 1981
philippinensis. Neaera Hinds, 1843: 78. West Pacific. 36-55 m. [ Cuspidaria (Luzonia)\
pilula, Lyonsiella Pelseneer, 1911: 76. West Pacific, Gulf of Alaska (Ivanova, 1977). 100-2980 m.
[ Policordia \

pilula, Lyonsiella Prashad, 1932: 325. = Policordia pilula (Pelseneer, 1911)
pinna. Cuspidaria ( Cardiomya ) Verco, 1908: 200. Southern Australia. 220-549 m. | Cardiomya]
planetica, Cuspidaria ( Cardiomya ) Dali, 1908: 433. East Pacific. 25-3000 m. [Cardiomya]
planulata, Lysonsiella Thiele, 1912: 232. = Lyons ia arcaeformis Martens, 1885 [Lyonsiidael. See Dell
(1990: 63)

platensis, Neaera E. A. Smith, 1885: 45. Southwest Atlantic. 100-1100 m. \Cuspidaria\
plicata, Cuspidaria Thiele, 1912: 233. = Cuspidaria tenella E. A. Smith, 1907
podobeda, Octoporia Krylova, 1994b: 42. Southeast Pacific, 2140 m. [Octoporia]
polpodes, Cuseidaria (sic) Dautzenberg & Fischer, 1897a: 28, nom. null. Error for Cuspidaria colpodes
Dautzenberg & Fischer, 1897

potli, Cuspidaria ( Cardiomya ) Sturany, 1901: 264. = Cardiomya alcocki (E. A. Smith, 1894)
pretiosa. Myonera (?) Verrill & Bush, 1898: 812. Northwest Atlantic. 618 m. |? My oner a\
prolatissima, Cuspidaria Poutiers, 1981: 348. Tropical West Pacific. 170-407 m. \Cuspidaria]
pseustes, Cuspidaria (Cardiomya) Dali, 1908: 432. = Cardiomya planetica (Dali, 1908), fide E. Coan
(in litt. 1992)

pulchella, Neaera (Cardiomya) H. Adams, 1871: 789. = Cardiomya singaporensis (Hinds, 1843)
pulchella, Poromya Adams & Reeve, 1850: 83. = Raeta (Raetellops) pulchella (Adams & Reeve,
1850) [Mactridae]

pygmaea, Verticordia (Haliris) Kuroda, 1952: 10. Northwest Pacific. 30-200 m. \ Ha Hr is (Setaliris) \
quadrata, Lyonsiella Hedley, 1907a: 303. Southwest Pacific. 146 m. [ Lyonsiella \
quadrata. Verticordia Dali, 1886b: 290. Atlantic. 630-4980 m. | Verticordia)

quadrostrata. Myonera angularis Poutiers, 1984: 292. West Indian Ocean. 3700-3716 m. \Bathyneaera\
quaylei, Lyonsiella Bernard, 1969: 2232. East Pacific. 350-1800 m. \ Lyonsiella]
radiata, Anatina Calcara, 1840: 40. = Cardiomya costellata (Deshayes, 1833)
radiata, Lyonsiella Dali, 1889: 442, nom. nud:, 1 890: 276. Southeast Pacific. 675-821 m. | Policordia |
raoulensis, Austroneaera Powell, 1958: 78. Southwest Pacific. 75-85 m. | Cuspidaria ( Rhinoclama )]
rara, Cuspidaria Thiele & Jaeckel, 1931: 253. = Cuspidaria optima Sowerby, 1904
rectangulata, Policordia (Angustebranchia) Ivanova, 1977: 180. West Pacific. 8175-9380 m. | Policordia
( Angustebranchia )\

rectimarginata. Cardiomya Dell, 1962: 69. South Pacific. 550 m. \Cardiomya\
renovata, Neaera Tiberi, 1855: 9. = Cuspidaria rostrata (Spengler, 1793)

reticulata, Cuspidaria (Cardiomya) Kuroda, 1948: 19. Northwest Pacific. 100-800 m. \Cardiomya\
rhomboidea, Verticordia Hedley, 1906b: 72, non Tate, 1887. = Haliris (Setaliris) setosa (Hedley, 1907)
robiginosa, Cardiomya Okutani & Sakurai, 1964: 23. = Cardiomya behringensis (Leche, 1883)
romanchensis, Poromya Odhner, 1960: 374. = Poromya tornata (Jeffreys, 1876)
ros, Cuspidaria (Halonympha) Verco, 1908: 201. Southern Australia. 235-550 m. \Halonympha\
rosea, Neaera Hinds, 1843: 78. West Pacific. 9-55 m. [Cuspidaria]
rostra. Myonera Poutiers & Bernard sp. nov. West Central Pacific. 280-440 m. | Myonera \
rostrata, Euciroa (Acreuciroa) Thiele & Jaeckel, 1931: 249. West Pacific and East Indian Ocean.
200-550 m. [Acreuciroa]

rostrata, Mya Spengler, 1793: 42. Arctic, North and Central Atlantic. 18-2997 m. | Cuspidaria |

166

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

rostrata, Poromya Rehder, 1943: 189. Northwest & West Central Atlantic. 110-183 m. [Poromya]
rostratocostellata , Corbula Acton, 1855: 3. = Cardiomya costellata (Deshayes, 1833)
rotundata, Poromya Jeffreys, 1876: 494. = Poromya granulata (Nyst & Westendorp, 1839)
rugata, Neaera (Rhinomya) Angas. 1867: 914, non A. Adams, 1864. = Plectodon brazieri (E.A.
Smith, 1885)

rugata , Neaera ( Rhinomya ) A. Adams, 1864: 207. Northwest Pacific. 46 m. \Cuspidaria ( Rhinoclama)\
ruginosa , Neaera Jeffreys, 1882: 942. East Atlantic. 250-3400 m. [1 Protocuspidaria ( Edentaria )\
rugosa, Octoporia Krylova, 1994b: 44. Southern Australia. 5020 m. | Octoporia]
saha, Cardiomya Knudsen, 1982: 133. West Central Atlantic. 235 m. | Cardiomya]
sadoensis , Cuspidaria Okutani & Ito, 1983: 167. Northwest Pacific. 200-275 m. \Cuspidaria]
sagamiana, Cardiomya Okutani & Sakurai, 1964: 21. = Cardiomya tosaensis (Kuroda, 1948)
sagamiensis, Dermatomya tenuiconcha Okutani, 1962: 32. = Poromya ( Dermatomya ) tenuiconcha
Dali, 1913

saharica, Mytilimeria Locard, 1898: 213. East Central Atlantic. 1495 m. \Halicardia\
salamensis, Cuspidaria Thiele & Jaeckel, 1931: 254. West Indian Ocean. 404 m. \Halonympha\
sansibarica, Poromya Thiele & Jaeckel, 1931: 251. West Indian Ocean. 404-463 m. | Poromya |
sarsii , Silenia E.A. Smith, 1885: 75. Southwest Atlantic, Southeast Indian Ocean. 3566-4846 m.
\Cetoconcha\

scaber, Plectodon Carpenter, 1866: 207. East Pacific. 20-250 m. | Plectodon \
scapha, Cetoconcha Dali, 1902: 561. Tropical East Pacific. 183 m. \Poromya ( Cetomya )\
seguenzae, Verticordia Dali, 1886b: 290. West Atlantic. 227-1 170 m. \ Verticordia \
semicostata, Cuspidaria ( Cardiomya ) Prashad, 1932: 333. Tropical West Pacific. 411 m. | Cardiomya
( Kurodamya )\

semipellucida, Cuspidaria (1 Plectodon) Kuroda, 1948: 24. Northwest Pacific. 100-250 m. | Pseudoneae-
ra\

semirostrata , Cuspidaria Locard, 1898: 177. Atlantic. 2030-3175 m. \Cuspidana\
semistrigosa , Neaera Jeffreys, 1882: 941. Northeast and tropical Atlantic. 330-1220 m. [ Cuspidaria
( Rhinoclama) \

septentrionalis, Cuspidaria (Cardiomya) gouldiana Kuroda, 1948: 18. = Cardiomya gouldiana (Hinds,
1843)

setosa , Verticordia Hedley, 1907a: 303, nom. nov. pro Verticordia rhomboidea Hedley, 1906, non Tate,
1887. Southwest Pacific. 146-293 m. [ Haliris ( Setaliris )\
sibogai, Cuspidaria ( Cardiomya ) Prashad, 1932: 331. Indonesia. 2060 m. \Cardiomya )
simillima, Cuspidaria ( Cardiomya ) E.A. Smith, 1915: 104. = Cardiomya cleryana (d’Orbigny, 1845)
similis , Rhinoclama (Austroneaera) Krylova, 1994a: 61. Southeast Pacific. 380-570 m. | Cuspidaria
( Rhinoclama )\

simplex, Cuspidaria {Luzonia) Allen & Morgan, 1981: 485. East Central Atlantic. 619-2357 m.

\Cuspidaria ( Luzonia )]

simplis, Protocuspidaria ( Edentaria ) Allen & Morgan, 1981: 498. North Atlantic. 1624-4825 m.
| Protocuspidaria ( Edentaria ) |

simulans, Cuspidaria Tate, 1897: 44. Southern Australia. 75-275 m. | Cuspidaria ( Rhinoclama )\
singaporensis, Neaera Hinds, 1843: 77. Indo-Pacific. 10-200 m. [Cardiomya]
sinica, Cardiomya Xu, 1980: 338. Northwest Pacific. 104-220 m. | Cardiomya \
sinuosa, Octoporia Krylova, 1994b: 40. Northwest, Southern Australia. 4440-5540 m. | Octoporia \
sinuosa, Pecchiolia Jeffreys, 1882: 932. North Atlantic, Mediterranean. 1200-2500 m. \Laevicordia\
smidti, Lyonsiella Friele, 1886: 38, nom. nov. pro Lyonsiella jeffreysi E.A. Smith, 1885, non Friele,
1879. Atlantic. 3300-4400 m. \Laevicordia]

smithi, Lyonsiella Dautzenberg, 1927: 348, nom. nov. pro Lyonsiella jeffreysi E. A. Smith, 1885, non
Friele, 1879. = Laevicordia smidti (Friele, 1886)

smithi , Lyonsiella Prashad, 1932: 325, non Dautzenberg, 1927. = Laevicordia abscissa (Pelseneer,
1911)

smithii , Cetoconcha Dali, 1908: 431. East Central & Southeast Pacific. 302-3436 m. [ Cetoconcha \

Source : MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

167

smithii, Vulcanomya Dali, 1886b: 299, nom. nov. pro Neaera adunca E.A. Smith, 1885 non Gould,
1861. Habitat unknown. \Cuspidaria ( Vulcanomya )[
solidula, Cuspidaria ( Cuspidaria ) Prashad, 1932: 330. Tropical West Pacific. 522 m. \Cuspidaria\
soyoae, Poromya (Dermatomya) Habe, 1952a: 274, nom. nud. = Dermatomya tenuiconcha var. soyoae
Habe, 1952b

soyoae, Dermatomya tenuiconcha var . Habe, 1952b: 158. = Poromya (Dermatomya) tenuiconcha Dali,
1913

spinosa. Euciroa Thiele & Jaeckel, 1931: 249. West Indian Ocean. 404-463 m. | Euciroa]
spinosa , Verticordia ( Haliris ) Bernard, 1969: 2233. Tropical East Pacific. 275 m. \HaUris)
spinulosa, Poromya Thiele, 1912: 232. Antarctic. 300-400 m. [Poromya]
steindachneri, Cuspidaria Sturany, 1901: 261. Indo-Pacific. 106-1308 m. [Cuspidaria]
striata. Neaera Jeffreys, 1876: 495. Atlantic. 150-2650 m. [Cardiomya]
striata. Poromya Sowerby, 1904: 16. South Africa. 97-384 m. [ Poromya [
striatella , Halonympha Verrill & Bush, 1898: 810. West Atlantic. 618-760 m. [Halonympha]
striolata. Cuspidaria Locard, 1898: 195. Northeast Atlantic, Mediterranean. 322-2100 m. [Cardiomya]
subglacialis, Cuspidaria Dali, 1913: 593. Northeast Pacific. 2000 m. [ Cuspidaria [
sublevis, Poromya Verrill, 1884: 221. = Poromya tornata (Jeffreys, 1876)
subrotundata, Policordia (Latebranchia) Ivanova, 1977: 192. North Pacific. 1050 m. [Policordia]
subquadrata. Pecchiolia Jeffreys, 1882: 932. North Atlantic. 298-3340 m. [Lyonsiella]
subtorta. Neaera G. O. Sars, 1878: 87. Arctic, North Atlantic. 0-990 m. [ Cuspidaria |
suganumai, Cuspidaria Nomura, 1940: 101. Northwest Pacific. 106-220 m. | Cuspidaria [
sulcata, Corbula Wood, 1840: 243, nom. nud. = Cuspidaria jugosa (Wood, 1850)
sulcata. Neaera Loven, 1846: 202. = Cardiomya costellata (Deshayes, 1833)
sulcifera, Neaera Jeffreys, 1880: 316, nom. nud.; 1882: 937. Northeast Atlantic. 80-1250 m.
[Cuspidaria]

sumatrana, Poromya Thiele & Jaeckel, 1931: 252. East Indian Ocean. Depth unknown. [Poromya]
surinamensis, Cardiomya Altena, 1971: 78. West Central Atlantic. 6-95 m. [ Cardiomya [
tanabensis, Plectodon ( Pseudoneaera ) Habe, 1960: 288. = Cuspidaria (Leiomya) adunca (Gould,
1861)

tasmanica, Cuspidaria Knudsen, 1970: 152. Southwest Pacific. 4400 m. [Myonera]
tasmanica. Neaera Tenison-Woods, 1876: 27. Southern Australia, (fide Tate, 1897: 44). [IPseudo-
neaera]

tenella, Cuspidaria E. A. Smith, 1907: 1. Antarctic. 183-1674 m. [Cuspidaria]

tenerrima , Verticordia Thiele & Jaeckel, 1931: 246. West Indian Ocean. 463 m. | Verticordia [

tenuiconcha. Poromya ( Dermatomya ) Dali, 1913: 596. Northeast & Northwest Pacific. 100-2200 m.

| Poromya ( Dermatomya)]

tenuis, Neaera Hinds, 1844: 97. = Raeta (Raetella) tenuis (Hinds, 1844) [Mactridae]
tenuissima. Cetoconcha Okutani, 1966: 9. Northwest Pacific, 500-1000 m. [Cetoconcha]
teporis, Haliris Poutiers & Bernard, sp. nov. West Pacific. 390 m. [Haliris]
teramachii. Cuspidaria Kuroda, 1948: 14. Northwest Pacific. 100 m. | Cuspidaria [
teramachii, Euciroa (Acreuciroa) Kuroda, 1952: 15. = Acreuciroa rostrata (Thiele & Jaeckel, 1931)
teres, Neaera Jeffreys, 1882: 939. North Atlantic. 252-3000 m. [Cuspidaria ( Rhinoclama)]
testai, Cuspidaria Knudsen, 1970: 154. Northwest Atlantic. 4380 m. [Cuspidaria ( Rhinoclama )[
thaumasia, Pseudoneaera Sturany, 1901: 11. Red Sea, Indian Ocean. 247-1082 m. | Pseudoneaera \
thomassini. Protocuspidaria ( Edentaria ) Poutiers, 1984: 295. West Indian Ocean. 3716 m. [Protocus-
pidaria (Edentaria)]

tillamookensis, Myonera Dali, 1916a: 23, nom. nud.; 1916 b: 407. East and Southwest Pacific, tropical
East Atlantic. 436-2850 m. | Bathyne aera]

tomlini, Cuspidaria (Cuspidaria) Prashad, 1932: 330. West Central Pacific. 275 m. [ Cuspidaria [
tornata, Pecchiolia Jeffreys, 1876: 494. North and Central Atlantic, West & Central Indian Ocean.
2085-5300 nt. | Poromya [

torrida, Verticordia Hedley, 1906a: 473. Southwest Pacific. 31-37 m. | Vert ambitus]

168

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

tosaensis, Cuspidaria ( Cardiomyc ) Kuroda, 1948: 18. Northwest Pacific. 28-300 m. \Cardiomya\
trailli , Neaera Hutton, 1873: 62. Southwest Pacific. 9-202 in. [Cuspidaria]

transversa, Poromya Dali, Bartsch & Rehder, 1938: 224. Mid-North Pacific. 474-487 m. | Poro-
mya\

transversa, Verticordia Locard, 1898: 201. Northeast Atlantic, West Indian Ocean. 3700-4165 m.
[Cetoconcha]

trapeza, Euciroa Poutiers, 1982: 331. West Pacific. 250-550 m. [Euciroa]
trapezoidea, Verticordia Seguenza, 1876a: 110. = Haliris granulata (Seguenza, 1858)
triangularis , Verticordia Locard, 1898: 207. Atlantic. 200-3862 m. [ Vert ambitus]
trigona, Neaera Hinds, 1843: 78. Habitat unknown. |? Cuspidaria]
trigonalis , Cuspidaria Tate, 1897: 45. Southern Australia. 27-64 m. |? Pseudoneaera]
trigonata. Thyasira Yokoyama, 1922: 158. Northwest Pacific. 50-150 m. | Simplicicordia ]
trosaetes , Cuspidaria Dali, 1925: 16. Northwest Pacific. 50-650 m. | Cuspidaria (Nordoneaera)]
trosti, Poromya Strong & Hertlein, 1937: 163. Northeast Pacific. 37-398 m. | Poromya ( Dermatomya )]
truncata, Cuspidaria Hedley, 1905: 47. Southwest Pacific. 203 m. \l Cuspidaria]
truncata. Neaera Jeffreys, 1880: 316, nom. nude, 1882: 936. North Atlantic. 710-1340 m. [Pseudoneae¬
ra]

tuberata, Poromya Jeffreys, 1882: 936. = Poromya neaeroides Seguenza, 1876
tuhua, Cuspidaria Dell, 1962: 67. South Pacific. 494 m. [Cuspidaria]
turgida, Cuspidaria Verrill & Bush. 1898: 799. Northwest Atlantic. 3338 m. ] Cuspidaria \
typus, Cuspidaria Nardo, 1840: 50. = Cuspidaria cuspidata (Olivi, 1792)
umbonata , Poromya Knudsen, 1982: 129. West Central Atlantic. 850 m. [Poromya]
undata, Neaera Verrill, 1884: 223. Atlantic, Indian Ocean. 4320-5300 m. [Cuspidaria]
undosa, Poromya Hedley & Petterd, 1906: 224. Southwest Pacific. 457-549 m. | Poromya]
uschakovi, Lyonsiella Gorbunov, 1946: 321. Arctic. 1475-2209 m. | Policordia ( Dallicordia )]
vadosa, Verticordia Hedley, 1907a: 303. Southwest Pacific. 146 m. | Vert ambitus]
valdiviae, Cuspidaria Thiele & Jaeckel. 1931: 225. Indian Ocean. 693-1644 m. [Cuspidaria (Rhinocla-
ma)\

variola , Cuspidaria Bernard, 1979: 16. Northeast Pacific. 2520-2884 m. | Cuspidaria ]
velvetina, Leiomya (Plectodon) granulata var. Dali, 1886b: 300. = Plectodon granular us (Dali, 1881)
ventricosa, Cuspidaria Verrill & Bush, 1898: 802. Atlantic. 349-3235 m. [Cuspidaria]
verityi, Protocuspidaria (Protocuspidaria) Allen & Morgan, 1981: 496. Atlantic. 943-4706 m.
| Protocuspidaria]

verticordia, Verticordia Nordsieck, 1969: 170, non S.Wood, 1840 (nom. nud. = Verticordia
cardiiformis (Sowerby, 1844), a fossil species of Europe). = Spinosipella acuticostata (Philippi,
1844)

vitrea, Neaera Loven, 1846: 202. = Cuspidaria abbreviata (Forbes, 1843)

walleri, Cuspidaria (Luzonia) Bernard, 1989: 64. Northeast Pacific. 100-450 m. | Cuspidaria (Luzo-
nia) |

wellmani. Austroneaera Fleming, 1948: 82. Southwest Pacific. 7-55 m. | Pseudoneaera ]
willetti, Cuspidaria Fleming, 1948: 81. South Pacific. 26-64 m. | Cuspidaria \
wollastonii. Neaera E. A. Smith, 1885: 40. Atlantic. 103-3175 m. | Cuspidaria ]
woodi, Verticordia E. A. Smith, 1885: 168. West Atlantic. 180-1850 m. [Verticordia]

Note 1: Though Myonera is considered here as a full genus, distinct from Cuspidaria, the name
Cuspidaria ( Myonera ) atlantica Allen & Morgan, 1981, is a primary homonym of Cuspidaria
(Cuspidaria) atlantica Allen & Morgan, 1981, and consequently invalid, according to Article 57d of
the Code dealing with irrelevance of subgeneric names on homonymy between species-group names.
Then, the species has to be renamed, and I am pleased to give it herein the new name of Mvonera
alleni, in honour of my talented friend. Dr John A. Allen of the University Marine Biological
Station, Millport.

Source : MNHN, Paris

ANOMALODESMATA FROM THE TROPICAL PACIFIC

169

Note 2: 1 have examined the type of Mytilimeria compressa Locard in mnhn. It consists of a
single left valve showing lucinoid but not verticordioid affinities: the shell is lenticular, without an
external granulation; the interior is not nacreous, the pallial line without a sinus, and the anterior
adductor muscle scar elongate, with a ventral expansion parallel to the pallial line.

Note 3: The systematic placement of Lyonsia formosa Jeffreys has been much disputed, and
is still controversial. First considered a member of family Lyonsiidae, this species has been made a
Lyonsiella by Allen & Turner (1974) on anatomical grounds. Later, on the basis of Allen &
Turner’s study, Scarlato & Starobogatov (1983) created for it the monogeneric family
Spinolyonsiellidae and new genus Spinolyonsiella. However, as Dall (1903) already made L. formosa
the type species of Allogramma , Spinolyonsiella cannot stand and is an objective junior synonym of
Allogramma.

I refrain from adopting these nomenclatural changes, as it appears that the identity of L.
formosa ( sensu Allen & Turner) is problematical. Actually, it is possible that two species (perhaps not
closely related) have been mixed, one mainly bathyal, the other abyssal (Poutiers, 1984). Then, the
form Dall had in view when erecting Allogramma might be different from that studied by Allen &
Turner.

APPENDIX 1

STATION DATA

musorstom 1 (Philippines)

Stn 5, 19. III. 76, North of Lubang Island, 14°01.5' N, 120°22' E, 200-215 m; gear: 4 m beam trawl:
Cuspidaria convexa.

Stn 25, 22.111.76, North of Lubang Island, 14°02.5' N, 120°22' E, 191-200 m, sand and mud; gear: 4
m beam trawl: Cuspidaria prolatissima.

Stn 26, 22. III. 76, North of Lubang Island, 14°00' N, 120° 17' E, 189 m; gear: 4 m beam trawl:
Poromya butoni.

Stn 31, 22. III. 76, North of Lubang Island, 14°00' N, 120°17.5' E, 187-195 m, mud; gear: 4 m beam
trawl: Cuspidaria prolatissima , Poromya butoni.

Stn 34, 23. III. 76, North of Lubang Island, 13°59.5' N, 120° 17.5' E, 188-191 m; gear: 5 m beam trawl:
Poromya butoni.

Stn 42, 24.111.76, channel between Lubang and Luzon islands, 13°54.5' N, 120°29' E, 379-407 m, hard
mud; gear: 5 m beam trawl: Acreuciroa rostrata, Cuspidaria prolatissima.

Stn 43, 24. III. 76, channel between Lubang and Luzon islands, 13°50' N, 120°28' E, 448-484 m, mud
with plant remains and stones; gear: 5 m beam trawl: Acreuciroa rostrata.

Stn 44, 24.111.76, South of channel between Lubang and Luzon islands. 13°46.5'N, 120"29.5' E,
592-610 m, mud with plant remains; gear: 5 m beam trawl: Halicardia philippinensis , Euciroa
eburnea.

Stn 47, 25.111.76, Southeast of Lubang Island, 13°41.5'N, 120°30'E, 685-757 m, mud with plant
remains; gear: 5m beam trawl: Euciroa eburnea.

Stn 49, 25. III. 76, West of Lubang Island, 13°49' N, 120°00.5' E, 750-925 m, mud; gear: 4 m beam
trawl: Spinosipella costeminens.

Stn 50, 25. III. 76, West of Lubang Island, 13°49' N, 120°02' E, 415-510 m. mud; gear: 4 m beam trawl:
Halonympha leiomyoides. Poromya eximia.

Stn 58, 26.Ili.76. North of Lubang Island, 13°58.5' N, 120°14' E, 143-178 m, sand and mud; gear: 4
m beam trawl: Cardiomya gouldiana.

170

JEAN-MAUR1CE POUTIERS & FRANK R. BERNARD

Stn 61, 27.111.76, North of Lubang Island, 14°01' N, 120°17.5' E, 124-129 in, mud; gear: 4 m beam
trawl: Poromya butoni.

Stn 63, 27.III.76, North of Lubang Island, 14°00.5' N, 120°16' E, 191-195 m, mud; gear: 4 m beam
trawl: Cuspidaria lubangensis, Cuspidaria steindachneri.

Stn 71, 28. III. 76, Northeast of Lubang Island, 14°09.5' N, 120°26.5' E, 174-204 m, sand and mud;
gear: 4 m beam trawl: Cuspidaria gigantea, Cuspidaria hindsiana.

Stn 72, 28. III. 76, Southwest of Corregidor Island, 14°12.5' N, 120°29' E, 122-127 m, hard mud; gear:
4 m beam trawl: Poromya butoni.

Stn 73, 28.111.76, Southwest of Corregidor Island, 14°16'N, 120°31.5'E, 70-76 m, sand and mud;
gear: 4 m beam trawl: Poromya butoni.

musorstom 2 (Philippines)

Stn 2, 20. XI. 80, North of Lubang Island. 14°00.5' N, 120°17.3' E, 184-186 m; gear: 4 m beam trawl:
Euciroa crassa.

Stn 6, 20. XI. 80, North of Lubang Island, 13°56.5' N, 120°21.5' E, 136-152 m; gear: 4 m beam trawl:
Euciroa crassa.

Stn 10, 21. XI. 80, North of Lubang Island, 14°00.7' N, 120°18.2' E, 188-195 m; gear: 4 m beam trawl:
Cuspidaria gigantea , Euciroa crassa.

Stn 11, 21.XI.80, North of Lubang Island, 14°00.3' N, 120°19.3' E, 194-196 m; gear: 4 m beam trawl:
Cuspidaria japonica.

Stn 15, 21. XI. 80, channel between Lubang and Luzon islands, 13°55' N, 120°28.9' E, 326-330 m; gear:
4 m beam trawl: Acr euciroa rostrata.

Stn 17, 22. XI. 80, North of Lubang Island, 14°00.5' N, 120° 17.8' E, 174-193 m; gear: 4 m beam trawl:
Euciroacrassa, Cuspidaria prolatissima.

Stn 19, 22.XI.80, North of Lubang Island, 14°00.6' N, 120°17.4' E, 189-192 m; gear: 4 m beam trawl:

Euciroamillegemmata, Cuspidaria nobilis, Poromya butoni.

Stn 21, 22.XI.80, North of Lubang Island, 14°01.2' N, 120°17.6' E, 191-192 m; gear: 4 m beam trawl:
Cuspidaria nobilis , Cuspidaria prolatissima.

Stn 25, 23. XI. 80, Verde Island Passage, 13°39.5'N, 120°42.9' E, 520-550 m; gear: 4 m beam trawl:

Cuspidaria kyushuensis, Cardiomya alcocki , Poromya eximia.

Stn 26, 23. XI. 80, Verde Island Passage, 13°49'N, 120°50.3'E, 299-320 m; gear: 4 m beam trawl:
Euciroa crassa , Cuspidaria corrugata, Cuspidaria gigantea, Cuspidaria prolatissima, Poromya
butoni.

Stn 32, 24.XI.80, Verde Island Passage, 13°40.5'N, 120°54' E, 192-220 m; gear: geological dredge:
Euciroa millegemmata.

Stn 33, 24.XI.80, Verde Island Passage, 13°32'N, 121”07.5' E, 130-137 m; gear: 1.20 x 0.50 m
rectangular dredge: Spinosipella deshayesiana, Haliris multicostata.

Stn 39, 25. XI. 80, South of Mompog Passage, 13°08' N, 122°36.3' E, 1030-1190 m; gear: 4m beam
trawl: Cuspidaria macrorhynchus , Poromya eximia.

Stn 40, 25.XI.80, South of Mompog Passage, 13°08' N, 122°40.2' E, 280-440 m; gear: 4 m beam trawl:
Myonera rostra.

Stn 44, 26.XI.80, Mompog Passage, 13°23.5'N, 122°20.6' E, 760-820 m; gear: 4 m beam trawl:
Cuspidaria macrorhynchus, Poromya eximia.

Stn 46, 26.XI.80, Mompog Passage, 13°26.2'N, 122°17.3' E, 445-520 m; gear: 4 m beam trawl:
Poromya eximia.

Stn 49, 26.XI.80, Northwest of Marinduque Island, 13°38.8' N, 121°43.2' E, 416-425 m; gear: 4 m
beam trawl: Cuspidaria convexa.

Stn 50, 27.XI.80, East of Golo Island, 13°37.4'N, 120°33' E. 810-820 m; gear: 4 in beam trawl:
Euciroa eburnea.

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ANOMALODESMATA FROM THE TROPICAL PACIFIC

171

Stn 51, 27. XI. 80, North of Lubang Island, 13°59.8' N, 120°17' E, 170-187 m; gear: 4 m beam trawl:
Spinosipella deshayesiana, Euciroa crassa, Cuspidaria gigantea, Cuspidaria nobilis, Cuspidaria
prolatissima.

Stn 55, 27. XI. 80, West of Cabra Island, 13°53.4' N, 119°57.8' E, 865-866 m; gear: 4 m beam trawl:
Spinosipella costeminens.

Stn 56, 28. XI. 80, West of Cabra Island, 13°54.1'N, 119°56.7'E, 970 m; gear: 4 m beam trawl:
Cetoconcha exigua.

Stn 59, 28. XI. 80, North of Lubang Island, 14°00.4' N, 120°17' E, 186-190 m; gear: 4 m beam trawl:
Cuspidaria nobilis , Poromya butoni.

Stn 63, 29. XI. 80, North of Lubang Island, 14°07.3' N, 120°15.5' E, 215-230 m; gear: 4 m beam trawl:
C uspidaria pro la l issima .

Stn 64, 29.XI.80, North of Lubang Island, 14°00.8' N, 120°18.6' E, 191-195 m; gear: 4 m beam trawl:
Euciroa crassa, Cuspidaria prolatissima.

Stn 68, 29. XI. 80, North of Lubang Island, 14°01.2' N, 120°18.2' E, 195-199 m; gear: 4 m beam trawl:

Euciroa crassa, Cuspidaria japonica , Cuspidaria nobilis, Cuspidaria prolatissima.

Stn 71, 30. XI. 80, North of Lubang Island, 14°00.6' N, 120°18.5' E, 189-197 m; gear: 4 m beam trawl:
Euciroa crassa, Cuspidaria nobilis.

Stn 72, 30.XI.80, North of Lubang Island, 14°00.4' N, 120°18.6' E, 182-197 m; gear: 4 m beam trawl:

Euciroa crassa, Cuspidaria corrugata, Poromya butoni.

Stn 75, 01. XII. 80, channel between Lubang and Luzon islands, 1 3°5 1 .9' N, 120°30.1' E, 300-330 m;

gear: 4 m beam trawl: Acreuciroa rostrata, Cetoconcha boucheti, Poromya eximia.

Stn 78, 01. XII. 80, channel between Lubang and Luzon islands, 13°49.5' N, 120°28.5' E, 441-550 m;

gear: 4 m beam trawl: Acreuciroa rostrata, Poromya eximia.

Stn 79, 01. XII. 80, channel between Lubang and Luzon islands, 13°44'N, 120°31.7' E, 682-770 m;
gear: 4 m beam trawl; material: Euciroa eburnea.

Stn 80, 01. XII. 80, South of Cape Santiago, Luzon Island, 13°45.2'N, 120°37.5' E, 178-205 m; gear:

4 m beam trawl: Cuspidaria corrugata, Cuspidaria nobilis, Cuspidaria prolatissima.

Stn 81, 01.X1I.80, Verde Island Passage, 13°35.3' N, 121°01.3' E, 856-884 m; gear: 4 m beam trawl:
Myonera dautzenbergi.

Stn 82, 02. XII. 80, channel between Lubang and Luzon islands, 13°47' N, 120°28.8' E, 550 m; gear:

4 m beam trawl: Euciroa eburnea, Acreuciroa rostrata, Poromya eximia.

Stn 83, 02. XII. 80, channel between Lubang and Luzon islands, 13°55.9' N, 120°30.5' E, 318-320 m;
gear: 4 m beam trawl: Acreuciroa rostrata, Cuspidaria prolatissima.

corindon 2 (Makassar Strait, Indonesia)

Stn 208, 3 LX. 80, East of Borneo, 0°14.6' S, 117°52' E, 150 m; gear: 4 m beam trawl: Cetoconcha

gloriosa. , , .

Stn 231, 04.XI.80, Northwest of Sulawesi, 0°04.9' N, 1 19°47.8' E, 980-1080 m; gear: 4 m beam trawl:

Policordia olivacea.

Stn 267, 07.XI.80, West of Sulawesi, 1°56.6' S, 119°16.7'E, 134-186 m; gear: 4 m beam trawl:
Cuspidaria nobilis.

Stn 280, 08.XI.80, West of Sulawesi, 1°59' S, 1 19°09.9' E, 715-800 m; gear: 4 m beam trawl: Myonera
dautzenbergi.

Stn 281, 08.XI.80, West of Sulawesi, 1°59' S, 119"09.9'E, 715-800 m; gear: 4 m beam trawl:
Cardiomya alcocki.

172

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

" Albatross ” (Northeastern Borneo)

Stn 5582, 26.IX.09, Si Amil Island, off Darvel Bay, 4°19.9' N, 1 18H58.6' E, 1628 m, gray mud and fine
sand; gear: 12-foot beam trawl: Halicardia lioubricki.

“Vauban" 1978-1979 (Southern New Caledonia)

Stn 2, 23.V.78, 22°17' S, 167°14' E, 425-430 m: Spinosipella deshayesiana, Haliris multicostata.

Stn 3! 23.V.78, 22° 17' S, 167° 12' E, 390 m: Haliris teporis.

Stn 4, 23.V.78, 22°17' S, 167° 13' E, 400 m: Spinosipella deshayesiana.

Stn 9, 24.V.78, 22°20' S, 167°10' E, 175-200 m: Celoconcha japonica.

Stn 14, 28.V.78, 22° 16' S, 167° 17' E, 465-495 m: Euciroa trapeza.

Stn 15, 10.IV.78, 22°49' S, 167° 12' E, 390-395 m: Spinosipella deshayesiana , Euciroa eburnea.

Stn 16, 19. IV. 78, 22°46' S, 167° 12' E, 390-400 m: Spinosipella deshayesiana.

Stn 33, 06.V1.79, 22°33' S, 166°25' E, 290-350 m: Euciroa trapeza.

Stn 34, 06.VI.79, 22°32' S. 166°26' E, 350-420 m: Euciroa trapeza.

Stn 39, 07.VI.79, 22°29' S, 166°23' E, 375-550 m: Euciroa trapeza.

Stn 40, 07. VI. 79, 22°30' S, 166°24' E, 250-350 m: Haliris multicostata. Euciroa trapeza.

Stn 42, 26. IX. 79, 22°08' S, 167°04'E, 230-260 m: Spinosipella deshayesiana, Cuspidaria clathrata,
Cuspidaria morrisae , Myonera caduca.

APPENDIX 2

System of the bivalve molluscs of the superorder Septibranchia.

O. A. Scarlato & Ya. I. Starobogatov (1983)

[Translation by J.-M. Poutiers & J. P. Rocroi ]

Septibranch bivalve molluscs are distinguished by an extreme uniformity of their conchological
characters. Each of the 3 Recent orders of this superorder (Scarlato & Starobogatov, 1979) is
characterized on the whole by 1-2 types of shells. Hinges generally differ the one from the other only
in their degree of reduction (i.e. by a negative feature); moreover, even the most complete hinge is
so reduced that it is difficult to conclude whether it results from an heterodont or a preheterodont
hinge (the second proposal seems to us the most likely). Ligaments are also completely uniform
that is, either marginal and visible from the outside (in Fordillidae and in Recent poromyoid forms),
or with a developed resilium and provided with a lithodesma (in verticordioid and cuspidarioid
forms). Simultaneously, the accumulation of anatomical data (among the most recent ones: Knudsen
1970; Allen & Turner, 1974; Bernard, 1974; Ivanova, 1977; Allen & Morgan, 1981; Morton,
1982) instances the rather important taxonomic diversity of the superorder’s components. One is
entitled to conclude that the arguments of conchological order generally used for the generic
classification of bivalves are inapplicable to septibranchs, and that one is confronted with a basic
problem, the elaboration of new criteria suitable for the superorder in question.

To classify families and high level taxa, we propose to use anatomical characters, and first of
all the branchial apparatus-septum structure — the organ that best characterizes the representatives
of the superorder. It appears especially appropriate since study of the septibranchs anatomy is by far
much more complete than the one of other bivalve molluscs.

Nowadays, it is very well established (Allen & Morgan, 1981) that septum developed mainly
at the expense of the inner demibranchs and by reduction of the reflexion of their filaments. The

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ANOMALODESMATA FROM THE TROPICAL PACIFIC

173

following can be added to that is exposed in the above mentioned work. The hypothesis of Allen
& Morgan, attributing the origin of posterodorsal septal muscles in muscular fibres of the outer
demibranch’s filaments, perfectly suit for the poromyoid forms, especially since it is easy to relate the
posterior group of the septal openings of Cetoconchidae to the interfilamental grooves of the outer
demibranch. On the contrary, posterior septal musculature of the cuspidarioid forms seems to have
another origin. Judging on their disposition, posterolateral muscles (but in primitive cuspidarioids,
only them and those situated in posterior part of septum) derive from musculature of the
intersiphonal wall. Then, to increase efficiency of the septum’s function, posterodorsal muscles
originate, probably completely at the expense of the gathering of posterior fibres of the lateral
muscles; posterodorsal muscles are no more necessary, and this leads to their gradual reduction
(Allen & Morgan, 1981). The lateral septal muscles, initially diffuse, even after separation of the
posterodorsal muscles, and adhering generally along a line, concentrate into 2 pairs of fibres. A series
of basic differences can be noticed about the septal openings disposition (reticulate windows, groups
of openings, variable number of the series of openings).

By means of basic features of the branchial apparatus, verticordioid forms can be grouped in
two suborders, according to the degree of development of the outer demibranchs and to the stomach's
structure (Ivanova, 1977). A small group, representatives of which have a complete septum, can be
even related to cuspidarioid forms, and placed in a particular suborder. In the group of cuspidarioid
forms, differences in the septum structure allow to split these forms into several families which are
grouped together in two superfamilies; a third superfamily comprises the Eocene cuspidarioids with
a developed nacreous layer (for that, they are often placed among poromyoids, despite their shell
shape). For similar reasons, poromyoid forms must also be grouped in 2 superfamilies and 4 families.

We are not concerned with systematics of genera, since it needs a thorough study. However,
grouping of the main known genera in families compels us to establish series of genera, when
representatives of a traditional conchological genus fall in different families. The extinct order of
conocardioids has been already considered (Starobogatov, 1977). It is more varied when shell
structure is taken into account, and since a conchological classification is easy to do. About the
septum structure of its representatives, one may notice that two types of distribution can be observed
there for the lateral muscles, and that the same exists in cuspidarioids: along a line or in a few
bundles.

In the above exposed system, to designate the geological age of each genus, we use the latin
abbreviation of the epoch (with R for Recent). Besides, to avoid coincidence between denominations
of orders and suborders and those of genera, we use standardized suffixes, which are not of common
usage in malacology, but are accepted in the systematics of other groups of animals: -oidei for
suborders, -iformes for orders, -iformii for superorders; and we recommend that, in future, their use
becomes widespread in malacology, because they are more convenient.

Superorder Septibranchia Pelseneer, 1889 ( = Conocardiiformii Neumayr, 1891).

Order Verticordiiformes Scarlato & Starobogatov, 1971.

Suborder Fordilloidei Pojeta, 1975.

Gills composed of two demibranchs; a dorsal projection in stomach.

Superfamily Fordilloidea. Fordillidae Pojeta, 1975: Fordilla Barrande, 1881 — Cm, Neofordilla
Krasilova, 1977 — O.

Superfamily Parilimyoidea. Parilimyidae Morton, 1981: Procardia Meek, 1871 — J-Cr, Bucardiomya
Rollier in Cossmann, 1912 — J-Pg, (?) Triplicosta Cooper, 1897 — Pg, Parilimya Melvill &
Standen, 1899 — R, Panacea Dali, 1905 — R, Nipponipanacca Habe, 1977 — R.
Superfamily Euciroidea. Euciroidae Dali, 1894: Euciroa Dali, 1881 R, Acreuciroa Thiele & Jaeckel,
1931 — R, Kurinuia Marwick, 1942 — Pg; Lyonsiellidae Scarlato & Starobogatov, 1971:
Lyonsiella G. Sars, 1872 — R. Proagorina Iredale, 1930 — R. Rectilyonsiella gen. nov. (type

174

JEAN-MAUR1CE POUTIERS & FRANK R. BERNARD

species Lyonsiella compressa Allen & Turner, 1974. Shell rectangular, elongate, compressed;
sculpture of raised and distinct small radial lines; hinge without teeth) — R; Policordiidae
Scarlato, 1980: Halicardia Dali, 1895 — R, Halicardissa Dali, 1913 — R, Vertisphaera Iredale,
1930 — R, Policordia Dali, Bartsch & Rehder, 1939 — R (in this genus we provisionally
include genus Latebranchia Ivanova gen. nov.; for its diagnosis, see: Ivanova, 1977: 189, type
species Policardia obliquaeovata Ivanova, 1977 here designated by Ivanova).

Suborder Verticordioidei Scarlato & Starobogatov, 1971.

Gills are only composed of the inner demibranchs and sometimes by remnants of posterior part of
the outer demibranchs; no dorsal projection in stomach.

Spinolyonsiellidae fam. nov. Shell rectangular, as in lyonsiellids, with prosogyrous and sharply
prominent umbones. Surface with small spines arranged in radial rows; radial sculpture absent
or formed by a keeled flexure posteriorly and a few nearby ribs; hinge without teeth:
Spinolyonsiella gen. nov. (type species Lyonsia formosa Jeffreys, 1881. Shell with slightly
anterior umbones; dorsal margin feebly curved; radial ribs well marked) — R, Allenicordia
gen. nov. (type species Pecchiolia subquadrata Jeffreys, 1881. Shell with strongly prosogyrous
and decidedly anterior umbones; dorsal margin strongly curved, especialy in front of umbones;
radial ribs absent) — R; Verticordiidae Stoliczka, 1871: Verticordia Sowerby, 1844 (with
subgenus Spinosipella Iredale, 1930) — Pg-R, Vertambitus Iredale, 1930 - R, Haliris Dali,
1886 (with subgenus Setaliris Iredale, 1930) — R, Pecchiolia Savi & Meneghini in Murchison,
1850 — Pg-Ng, Trigonulina d’Orbigny, 1846 — R, Simplicicordia Kuroda & Habe, 1971 — R,
Laevicordia Seguenza, 1876 — Ng-R, Angustebranchia Ivanova gen. nov. (for its diagnosis, see:
Ivanova, 1977: 177-180, type species Policardia rectangulata Ivanova, 1977, here designated by
Ivanova) — R.

Order Conocardiiformes Neumayr, 1891.

Suborder Eopterioidei subordo nov.

Valves open under the action of a ligament and are closed by 1 or 2 adductors; septal muscles
attached to valves by 3 pairs of bundles.

Superfamily Eopterioidea. Two adductors. Eopteriidae Miller, 1889: Eopteria Billings, 1865 O;
Stolidotidae Starobogatov, 1977: (?) Myona Kobayashi, 1935 — Cm, Eoischyrina Kobayashi,
1933 — O, Maminka Barrande, 1881 — O-S, Siolidotus Hede, 1915 — S.

Superfamily Pseudotechnophoroidea. One adductor. Pseudotechnophoridae Starobogatov, 1977:
Pseudotechnophorus Kobayashi, 1933 — O.

Suborder Conocardioidei Neumayr, 1891.

Valves heavy; in living animal, they remain closed and are only pulled apart as growth goes on;
adductors reduced; septal muscles not gathering in bundles and attached along a line.

Superfamily Euchasmatoidea. Wanwaniidae fam. nov. Keel flexure feeble. Wanwania Kobayashi,
1933 — O, Apoplopegma Pojeta, G.-T., Sherg., 1977 — O, Euchasmatidae Starobogatov, 1977:
Euchasma Billings, 1865 — O, Euchasmella Kobayashi, 1933 — O, Pseudeuchasma Kobayashi,
1933 — O, Tenka Barrande, 1881 — O-S, Telinka Barrande, 1881 — S-D, Conocardiopsis
Beushausen, 1895 D.

Superfamily Conocardioidea. Bransoniidae Pojeta & Runnegar, 1976: Bransonia Pojeta & Runnegar,
1976 — O-P, Mulseodens Pojeta & Runnegar, 1976 — S-D, Pseudoconocardiwn Zavodowsky,
1960 — C-T; Hippocardiidae Pojeta & Runnegar, 1976: Hippocardia Brown, 1843 — O-C,
Rhipidocardium Fischer, 1887 — S, Bigalea Pojeta & Runnegar, 1976 — S-D; Conocardiidae
Miller, 1889: Conocardium Brown, 1835 — D-C, Arceodomus Pojeta & Runnegar, 1976
C-P.

Source :

ANOMALODESMATA FROM THE TROPICAL PACIFIC

175

Suborder Ribeirioidei Kobayashi, 1933.

Superfamily Ribeirioidea. Shell oval or elongate, with a rounded anterior margin, without keel or
radial sculpture. Ozomiidae Starobogatov, 1977: Ozomia Walcott, 1934 O; Ribeiriidae
Kobayashi, 1933: Ribeiria Sharpe, 1853 — Cm-O, Pinnocaris Etheridge, 1878 — Cm-O,
Ribeirina Billings, 1865 — O.

Superfamily Technophoroidea. Shell quadrate, elongate, with a sharp keel and often a radial
sculpture on carinal area. Technophoridae Miller, 1889: Kimopegma Pojeta, Gilbert-
Tomlinson & Shergold, 1977 — Cm, Pleuropegma Pojeta, Gilbert-Tomlinson & Shergold,
1977 — Cm, Opikella Runnegar & Pojeta, 1974 — Cm, Anisotechnophorus Pojeta & Runnegar,
1976 — O, Technophorus Miller, 1889 — O, Mvocaris Salter, 1864 — O.

Superfamily Ischyrinioidea. Tomalchoviidae Starobogatov, 1977: Cymatopegma Pojeta, Gilbert-
Tomlinson & Shergold, 1977 — Cm, Tolmachovia Howell & Kobayashi, 1936 — O,
Ptychopegma Pojeta, Gilbert-Tomlinson & Shergold, 1977 — O, Pauropegma Pojeta,
Gilbert-Tomlinson & Shergold, 1977 — O; Ischyriniidae Kobayashi, 1933: Ischirinia Billings,
1866 — O.

Order Poromyiformes Pelseneer, 1906.

Superfamily Dermatomyoidea. Septum with pairs of reticulated areas. Dermatomyidae fam. nov.
Two pairs of reticulated areas: Dermatomya Dali, 1889 — R, Ectorisma Tate, 1892 — We
include here most species of genus Poromya, but those grouped with P. granulata (Nyst &
Vestendorp) — R, Cetomya Dali, 1889 — R, Liopislha Meek, 1864 — Cr, Psilomya White,
1874 — Cr, Cymella Meek, 1864 — Cr; Perlaporomyidae fam. nov. One (medial) pair of
reticulated areas: Perlaporomya gen. nov. (type species Poromya perla Dali, 1908. Shell
quadrate, quite as long as high; very small tubercles covering the outer surface; posterior part
demarcated by a feeble groove; one subumbonal tooth on right valve, with a corresponding
socket on left valve; reticulated areas with 18 to 20 filaments) — R.

Superfamily Poromyoidea. Septum with groups of paired pores. Cetoconchidae Ridewood, 1903:
Cetoconcha Dali, 1886 — R; Poromyidae Dali, 1886: Poromya Forbes, 1884 — R,
Mioporomya Sacco, 1901 — Ng.

Order Cuspidariiformes Scarlato & Starobogatov, 1971.

Suborder Dallicordioidei subordo nov. Shell verticordioid; a very large incoming siphonal opening,
with many tentacles around; septum with 2 groups of openings on each side (in front of and
behind the foot); septal musculature developed towards left; bundles obvious, corresponding
with the existing filaments of inner demibranch.

Dallicordiidae fam. nov. Shell interiorly nacreous, septum with 4 pairs of openings: 2 pairs near the
oral funnel and 2 pairs behind the foot; posterior part of oral lobes largely united with septum,
Dallicordia gen. nov. (type species Lyonsiella alaskana Dali, 1895. Shell rounded, irregularly
pentagonal, truncated on anterior margin and rounded towards posterior margin; sculpture
composed of narrow radial threads; hinge without teeth. L. ushakovi Gorbunov and Policordia
ochotica Scarlato can also be placed in this genus) — R.

Suborder Cuspidarioidei Scarlato & Starobogatov, 1971.

Shell cuspidariid, with reduced siphonal openings, thus located both inside the rostrum at siphon s
extremity; septum with only one group of openings (4 to 20) on each side, strongly
muscularized; besides are developed, firstly 2 anterior pairs of muscles, and also posterior (1
or 2 pairs) and lateral (in a line or in 2 pairs) muscles.

Superfamily Neaeroporomyoidea. Neaeroporomyidae fam. nov. Shell interiorly nacreous, its
posterior part somewhat drawn out into a rostrum; hinge with a subumbonal tooth on each
valve: Neaeroporomya Cossmann, 1887 — Pg, Pseudocuspidaria Eames, 1951 — Pg.

Superfamily Protocuspidarioidea. Protocuspidariidae fam. nov. Shell not nacreous, rostrum short so
that the anterior part of dorsal shell margin is straight and horizontal; hinge without teeth or

176

JEAN-MAURICE POUTIERS & FRANK R. BERNARD

with an anterior tooth on only one valve or on both valves; septum comprising more than 10
pairs of short filaments among which septal openings are distributed: Bidentaria Allen &
Morgan, 1981 — R, Protocuspidaria Allen & Morgan, 1981 — R, Edentaria Allen & Morgan,
1981 — R.

Superfamily Cuspidarioidea. Halonymphidae fam. nov. 8-20 pairs of septal openings; among the
posterior septal muscles, posterolateral ones are only present; lateral septal muscles attached
along a line: Halonympha Dali & Smith, 1886 — R. Allenineaera gen. nov. (type species Neaera
circinnata Jeffreys, 1881. Shell oval quadrate, with a well-marked, straight, short rostrum and
with a straight dorsal margin; sculpture composed of fine concentric ribs) — R, Octoporia gen.
nov. (type species Myonera octoporosa Allen & Morgan. 1981. Shell drawn out in its posterior
part in a long, rounded at the end, rostrum; dorsal and ventral margins of rostrum slightly
concave; sculpture composed of concentric ridges, more strongly impressed on anterior part;
hinge without teeth) — R; Cardiomyidae fam. nov. 4-5 pairs of septal openings; posterodorsal
and posterolateral muscles simultaneously present; lateral septal muscles attached along a line:
Cardiomya A. Adams, 1864 — Cr-R, Kurodamya Okutani & Sakurai, 1964 R.
Semicardiomya gen. nov. (type species Myonera demistriata Allen & Morgan, 1981. Shell with
radial ribs on posterior half or on central and posterior parts, but with concentric ribs on
anterior part; hinge without teeth) — R. Bathyneaera gen. nov. (type species Cuspidaria hadalis
Knudsen, 1970. Shell compressed, with a feebly demarcated, rather short rostrum; strong
radial ribs on posterior half of valves; hinge without teeth) — R; Myoneridae fam. nov. 4-5
pairs of septal openings; posterodorsal muscles only present; lateral septal muscles attached
along a line: Jeffreysiomya Nordsieck, 1969 — Cr-R. Vulcanomya Dali, 1886 — R, Leiomya
A. Adams, 1864 — R. Pseudoneaera Sturany, 1902 — R, Rhinoclama Dali & Smith, 1886
R, Luzonia Dali & Smith, 1886 — R, Tergulina Doncieux. 1911 - Pg. Bowdenia Dali, 1963
Ng, Rengea Kuroda & Habe. 1971 — R, Myonera Dali & Smith, 1886 — R. Plectodon
Carpenter, 1864 — Ng-R; Cuspidariidae Dali, 1886. 4-5 pairs of septal openings; postero¬
dorsal muscles only present; lateral septal muscles united in 2 pairs of fibres, attached at
extremity to valves: Cuspidaria Nardo, 1840 — R — Here, we only place C. cuspidata (Olivi),
C ■ Jeffrey si (Dali), C. ventricosa Verrill & Bush, C. parkeri Knudsen, C. barnardi Knudsen — ,
Austroneaera Powell, 1937 — R — Here, because of the shell form, we include also Rhinoclama
abrupta Allen & Morgan, 1981 and we determine the systematic placement of genus because
of the anatomical features of this species.

ACKNOWLEDGEM ENTS

The first author wishes to thank Drs P. Bouchet and the late R.S. Houbrick for
communication of material on which this paper is based, as well as Drs E. Coan and K.J. Boss for
fruitful suggestions to improve this paper, and Dr A. Matsukuma for kind help about Japanese
literature. He is also grateful to Mr J.P. Rocroi and Ms A. Sasaki for translating papers written
respectively in Russian and in Japanese; and to Mr P. Lozouet for the careful photography.

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Verrill, A. E., 1880. - Notice of recent additions to the marine Invertebrata of the Northeastern coast of America, with
descriptions of new genera and species and critical remarks on others. Part II. — Mollusca, with notes on Annelida,
Echinodermata, etc., collected by the United States Fish Commission. Proceedings of the United States National
Museum. 3: 356-409.

Verrill, A. E., 1881. — Notice of the remarkable marine fauna occupying the outer banks off the Southern coast of New
England, No. 2. American Journal of Science, (3) 22: 292-303.

Verrill, A. E., 1882. — Catalogue of marine Mollusca added to the fauna of the New England region, during the past ten
years. Transactions of the Connecticut Academy of Arts and Science, 5: 447-587, pis 42-44 & 57-58.

Verrill, A. E., 1884. — Second catalogue of Mollusca recently added to the fauna of the New England coast and the adjacent
parts of the Atlantic, consisting mostly of deep-sea species, with notes on others previously recorded. Transactions of
the Connecticut Academy of Arts and Science, 6 (6): 139-294, pis 27-32.

Verrill, A. E., 1885. — Third catalogue of Mollusca recently added to the fauna of the New England coast and the adjacent
parts of the Atlantic, consisting mostly of deep-sea species, with notes on others previously recorded. Transactions of
the Connecticut Academy of Arts and Science, 6 (10): 395-452, pis 42-44.

Verrill, A. E. & Bush, K. J., 1898. — Revision of the deep-water Mollusca of the Atlantic coast of North America, with
descriptions of new genera and species. Part I. — Bivalvia. Proceedings of the United States National Museum. 20 (1 139):
775-901, pis 71-97.

Vokes, H. E., 1980. — Genera of the Bivalvia: a systematic and bibliographic catalogue (revised and updated). Paleontological
Reserch Institution, Ithaca, xxvii + 307 pp.

Wood, S. V., 1840. — A catalogue of shells from the Crag. Annals and Magazine of Natural History, 6: 241-253.

Wood, S. V., 1850. — A monograph of the Crag Mollusca. or, descriptions of shells from the Middle and Upper Tertiaries of
the East of England. Part II. Bivalves. Palaeontographical Society, London. 341 pp., pis 1-31.

ANOMALODESMATA FROM THE TROPICAL PACIFIC

187

Wood-Mason, J. & Alcock, A., 1891, — Natural history notes from H.M. Indian Marine Survey Steamer 'Investigator' .
Series II., No. I . On the results of deep-sea dredging during the season 1 890-91 . Annals and Magazine of Natural History ,
ser. 6, 8: 427-452.

Xu, Fengshan. 1980. — Two new species of Bivalvia (Mollusca) from the East China Sea. Oceanologia et Limnologia Sinica,
1 1 (4): 337-340.

Yokoyama, M., 1922. — Fossils from Upper Musashino of Kazusa and Shimosa. Journal of the College of Science, Imperial
University of Tokyo, 44 (1): 1-200, pis 1-17.

Yonge, C. M., 1928. — Structure and function of the organs of feeding and digestion in the septibranchs, Cuspidaria and
Poromya. Philosophical Transactions of the Royal Society of London, (B) 216 (434): 221-263, pis 12-14.

Yonge, C. M., 1947. — The pallia! organs in the aspidobranch Gastropoda and their evolution throughout the Mollusca.
Philosophical Transactions of the Royal Society of London, (B) 232: 443-517.

Note added in press:

The present paper was in press when the following paper by Krylova was published, containing the description of one new
genus, one new subgenus, and nine new species.

Krylova, E. M., 1995. — [Bivalve molluscs of the family Protocuspidariidae (Septibranchia, Cuspidarioidea): composition
and distribution.] Zoologicheskyi Zhurnal, 74 (9): 20-38. [in Russian]

New taxa:

Dentaria Krylova, 1995. Type species (OD): Multitentacula ( Dentaria) parvula Krylova, 1995.
Multitentacula Krylova, 1995. Type species (OD): Multitentacula admirabilis Krylova, 1995.

amoena, Multitentacula (Dentaria) Krylova, 1995. Kuriles-Kamchatka Trench, 5060 m.
composita, Multitentacula (M.) Krylova, 1995. Caribbean, 2970-3080 m.
fragilis, Protocuspidaria (P.) Krylova, 1995. Norhteast Atlantic, 3714 m.
parilis , Multitentacula (Dentaria) Krylova, 1995. Peru Trench, 2330 m.
parvula, Multitentacula (Dentaria) Krylova, 1995. Western Atlantic, 4205 m.
paulula, Multitentacula (Dentaria) Krylova, 1995. Southeast Atlantic, 4725 m.
pusilla, Protocuspidaria (Edentaria) Krylova, 1995. Northeast Pacific, 5840 m.
speciosa, Protocuspidaria (Edentaria) Krylova, 1995. Eastern Atlantic, 4278 m.
venusta, Multitentacula (M.) Krylova, 1995. Great Australian Bight, 3880 m.

Source : MNHN, Paris

ULTATS DES CAMPAGNES MUSORSTOM, VOLUME 14 RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 14

RESU

Scaphopoda of the tropical Pacific and Indian
with description of 3 new genera and 42 new

Oceans,

species

Victor SC A RABIN O

Museum national d'Histoire naturelle
55 rue de Buffon
75005 Paris, France

ABSTRACT

New data on the scaphopod fauna of the Indo-West Pacific are presented, based on new material from recent
oceanographic expeditions, mostly in the SW Indian Ocean, SE Asia and the New Caledonia region. Over 780 stations yielded
a total of 139 species. Of 81 species of Dentaliida and 58 Gadilida, 42 species (16 Dentaliida and 26 Gadilida), as well as
3 gadilid genera, are described as new. Many range extensions are documented, and new synonymies are established. With
73 recorded species, New Caledonia is currently the geographic area with the highest documented scaphopod diversity. Their
bathymetric distribution shows a peak in species numbers in deep water around 800 m, with a second, minor peak for Gadilida
at around 2,000 m. Including genera not represented in the Indo-Pacific, 44 Recent scaphopod genera are recognized. The
radula of 42 of these is described, and an update of the general classification of the class Scaphopoda is proposed.

RESUME

Scaphopodes des regions tropicales de I’ocean Indien et du Pacifique, avec la description de 3 nouveaux genres et 42 especes
nouvelles.

La faune de Scaphopodes de I'Indo-Pacifique est etudiee sur la base du materiel recolte par diverses campagnes
oceanographiques recentes, en particulier dans le Sud-Ouest de l'ocean Indien, l'Asie du Sud-Est et la region neo-caledonienne.
Au total, plus de 780 stations ont livre 139 especes. Sur 81 especes de Dentaliida et 58 Gadilida, 42 especes (16 Dentaliida et
26 Gadilida), de raeme que 3 genres de Gadilida, sont decrits comme nouveaux. L'aire de distribution connue de nombreuses
especes se trouve etendue, et de nouvelles synonymies sont etablies. Soixante-treize especes sont recensees en Nouvelle-
Caledonie, ce qui en fait le secteur de I’lndo-Pacifiique avec la faune de Scaphopodes la plus diversifiee. La distribution
bathymetrique de cette faune montre un maximum de diversite autour de 800 m, et un deuxieme maximum, plus attenue, pour
les Gadilida autour de 2000 m. En comprenant les genres non representes dans I'Indo-Pacifique, 44 genres actuels de
Scaphopodes sont reconnus. La radula de 42 d'entre eux est decrite, et la classification de la classe est mise a jour.

Scarabino, V.. 1995. — Scaphopoda of the tropical Pacific and Indian Oceans, with description of 3 new genera and 42 new species. In: P Bouchet (ed.),
Resultats des Campagnes MUSORSTOM, Volume 14. Mem. Mas. naln. Hist. rial.. 167: 189-379. Paris isbn 2-85653-217-9.

Published 29'1' December 1995.

190

VICTOR SCARABINO

INTRODUCTION

The present paper is based essentially on the collections made over the last 20 years in the
Indo-Pacific by mnhn staff and other French scientists, notably from orstom. It covers parts of the
Western Pacific, with emphasis on the Philippines, Indonesia and New Caledonia, and parts of the
Indian Ocean, with emphasis on its southwestern portion around Madagascar and the Mascarenes.
Much of the material was collected aboard R.V. “Vauban", “Coriolis” , and “ Alls' ' during the six
musorstom expeditions to the Philippines (1976-1985; for narratives and station lists, see Forest,
1981, 1986) and New Caledonia (1985-1989; for narratives of these, and other, cruises in the New
Caledonia area, together with station lists, see Richer de Forges, 1990 and 1991). There is also a
substantial amount of material from a dozen other expeditions on French research vessels, among
others Cruise 32 of R.V. “ Marion- Dufresne" (Reunion island; see Guille, 1982), CORINDON (R.V.
“ Coriolis ”, Makassar Channel, 1980) and benthedi (R.V. “Suroit”, Mozambique Channel, 1977).

Beside the material in mnhn, I have also studied the Scaphopoda taken during the “Snellius"
1 (1929-1930; see Boschma, 1936) and “Snellius” II Expeditions (1984; see Anonymous) in Indonesia
(rmnh), the “Galathea” Expedition (1950-52; see Bruun, 1959) (zmc) and the “Meiring Naude”
cruises (1976 and 1979; see Louw, 1977, 1980) off South Africa (sam). Whenever possible and
necessary, I have examined relevant type material and other reference material, as listed under each
species.

There has been no recent review of the scaphopod fauna of the Indo-Pacific. Many species
descriptions are scattered in the reports of major expeditions such as those of the “ Challenger ”
(Watson, 1879), “Investigator” (Smith, 1894 to 1906), “Siboga” (Boissevain, 1906), “Valdivia”
(Plate, 1908a, Jaeckel, 1932), and “John Murray" (Ludbrook, 1954). Part of the extensive Japanese
literature concerns directly the tropical Indo-Pacific, and Chistikov (1979 to 1983) studied the
material from several Russian expeditions in the Indian and Pacific Oceans. In parallel with the
present report, the scaphopod fauna of Australia is being revised by Dr J. Healy and Mr K.
Lamprell, and it is anticipated that the two papers combined will hopefully lay new foundations for
the taxonomy and biogeography of Indo-Pacific Scaphopoda. As a help for further revisionary work,
I have given under each genus a list of valid Indo-Pacific taxa not critically examined by me for this
paper, complete with type locality and literature references.

ABBREVIATIONS AND TEXT CONVENTIONS

For shells, usage of terms such as small, medium, long refers to the approximate mean length
for specimens considered adults. Known lengths range from 9 to 180 mm in Dentaliida, and from 2
to 50 mm in Gadilida. All measurements are in mm. L = length; W = maximum diameter; m =
diameter at aperture (this is not always the maximum diameter); w = minimum diameter (apex); arc
= the distance of maximum concavity of the dorsal side from a line between apex and mouth. Shell
terminology follows Figs 1-2.

Illustrations: in drawings of apical and oral sections, the specimens are placed dorsal side up.

Charts: distribution maps are given for all species. The star notes the type locality; in the case
of questionable type locality, a question mark has been added, see text for comments. Filled circles
represent locality records checked for the present paper, open circles represent records taken from the
literature. Circles may represent several neighbouring localities. In the case of records in Japanese
waters, I have not tried to map the distribution there in detail, since the present monograph only
marginally covers that country.

Radula illustrated teeth have been taken on the middle part of the radular ribbon, avoiding
both juvenile rows and rows showing wear. Radula terminology follows Figs 3-4.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

191

Fig. 1. — Illustrated glossary of shell structures and descriptive terms used in this publication: general shape and sculpture.

Repositories

amnh : American Museum of Natural History, New York
ams : Australian Musem, Sydney

ansp : Academy of Natural Sciences of Philadelphia, Philadelphia

bmnh : The Natural History Museum, London

cnrs : Centre National de la Recherche Scientifique

ioas : Institute of Oceanology, Academia Sinica

mhng : Museum d'Histoire Naturelle, Geneve

MNHN : Museum national d'Histoire naturelle, Paris

nmp : Natal Museum, Pietermaritzburg

rmnh : Nationaal Natuurhistorisches Museum, Leiden

nmnz : Museum of New Zealand, Wellington

nsmt : National Science Museum, Tokyo

orstom : Institut Frangais de Recherche Scientifique pour le Developpement en Cooperation

sam : South African Museum, Cape Town

usnm : National Museum of Natural History, Washington, DC

zin : Zoological Institute, Russian Academy of Sciences, St Petersburg

zmb : Zoologisches Museum der Humboldt-Universitat, Berlin

zmc : Zoologisk Museum. Copenhagen

Station data

CC : Shrimp trawl

CH : Otter trawl

CP : Beam trawl

DC : Charcot dredge

DG : Boillot geological dredge

DR : Rectangular dredge (1.20 x 0.50 m)

DS : Sanders epibenthic sledge

DW : Waren dredge

Source :

192

VICTOR SCARABINO

section

apical structures

o

triangular

subtriangular \\

quadrangular

pentagonal

in

© hexagonal

heptagonal

octogonal

n

apical callus

preapical callus

slit

fissure

bilobed

four-lobed

simple

five-lobed

Solenoxiphus type ~ six-lobed dorsal lobe ventral lobe

Fig. 2. — Illustrated glossary of shell structures and descriptive terms used in this publication: sections and apical structures.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

193

lateral rachidian

granules

F : Faubert [a group of old cotton nets towed on the bottom, especially on steep hard

bottoms]

KG : Usnel Box-Corer
B : Okean dredge

S : scuba operated air lift

stn : Station

T : Troika dredge

Other abbreviations

Coll.

: collection of

coll.

: collected by

Iv

: live-taken specimens

dd

: empty shells

OD

: Original designation

SD

: Subsequent designation

Source :

194

VICTOR SCARABINO

lateral rachidian

Fig. 4. — Radular nomenclature used in this publication: Gadilida.

SYSTEMATIC ACCOUNT

This paper follows, with some modifications based on unpublished information, the
classification proposed by Steiner (1992b), which is currently the best approach to a biologically
based ordination for the class (Table 1). Modifications proposed here are supported by radular and
shell characters. For historical reviews of former classifications see Emerson (1978) and Steiner
(1992b). This latter paper as well as those of Steiner (1991 and 1992a) are recent and provide
descriptions of the anatomy of representatives of genera, therefore I will here discuss only shell and
radular characters, based on new, unpublished observations. (Exceptions are Sagamicadulus and
Eudentalium, for which I have not had access to live-taken material). For general information about
scaphopod radula, I refer to Ivanov & Chistikov (1990).

Table 1. — Synopsis of the classification of Indo-Pacific Scaphopoda used in the present paper.

Class Scaphopoda Bronn, 1862
Order Dentaliida da Costa, 1776
Family Dentaliidae Gray, 1847

Genus Dentalium Linne, 1758 = Lentigodentalium Habe, 1963
Genus Paradentaliutn Cotton & Godfrey, 1933
Genus Tesseracme Pilsbry & Sharp, 1897
Genus Eudentalium Cotton & Godfrey, 1933
Genus Antalis H. & A. Adams, 1854

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

195

Genus Plagioglypta Pilsbry in Pilsbry & Sharp, 1897

Genus Striodentalium Habe, 1964

Genus Graptacme Pilsbry & Sharp, 1897

Genus Fissidentalium Fischer, 1885

Genus Schizodentalium Sowerby, 1894

Genus Compressidentalium Habe, 1963

Genus Coccodentalium Sacco, 1896

Genus Pictodentalium Palmer, 1974

Family Calliodentaliidae Chistikov, 1975
Genus Calliodentalium Habe, 1964

Family Fustiariidae Steiner, 1991

Genus Fustiaria Stoliczka, 1868

Family Gadilinidae Chistikov, 1975

Subfamily Gadilininae Chistikov, 1975
Genus Gadilina Foresti, 1895

Subfamily Episiphoninae Chistikov, 1975

Genus Episiphon Pilsbry & Sharp, 1897

Subfamily Anuudentaliinae Chistikov, 1975
Genus Anulidentalium Chistikov, 1975

Family Laevidentaliidae Palmer, 1974

Genus Laevidentalium Cossmann, 1888

Family Omniglyptidae Chistikov, 1975

Genus Omniglypta Kuroda & Habe in Habe, 1953

Family Rhabdidae Chistikov, 1975

Genus Rhabdus Pilbry & Sharp, 1897

Order Gadilida Starobogatov, 1974

Suborder Entalimorpha Steiner, 1992

Family Entalinidae Chistikov, 1979

Subfamily Entalininae Chistikov, 1979
Genus Entalina Monterosato, 1872

Subfamily Heteroschismoidinae Chistikov, 1982
Genus Heteroschismoides Ludbrook, 1960
Genus Costentalinci Chistikov, 1982
Genus Entalinopsis Habe, 1957
Genus Spadentalina Habe, 1963
Genus Pertusiconcha Chistikov, 1982

Subfamily Bathoxiphinae Chistikov, 1983

Genus Bathoxiphus Pilsbry & Sharp, 1897
Genus Rhomboxiphus Chistikov, 1983
Genus Solenoxiphus Chistikov, 1983

Suborder Gadilimorpha Steiner, 1992

Family Pulsellidae Scarabino in Boss, 1982
Genus Pulsellum Stoliczka, 1868
Genus Annulipulsellum Scarabino, 1986
Genus Striopulsellum gen. nov.

Source : MNHN. Paris

196

VICTOR SCARABINO

Family Wemersoniellidae Scarabino, 1986
Genus Wemersoniella Scarabino, 1986
Genus Chistikovia gen. nov.

Family Gadilidae Stoliczka, 1868
Subfamily Siphonodentaliinae Simroth, 1894
Genus Siphonodentalium M. Sars, 1859
Genus Sagamicadulus Sakurai & Shimazu, 1963
Genus Slriocadulus Emerson, 1962
Genus Polyschides Pilsbry & Sharp, 1898
Genus Dischides Jeffreys, 1867

Subfamily Gadilinae Stoliczka, 1868
Genus Cadulus Philippi, 1844
Genus Bathycadulus gen. nov.

Genus Gadila Gray, 1847 = Platyschides Flenderson, 1920

Families incertae sedis

Genus Megaentalina Habe, 1963

Genus Compressidens Pilsbry & Sharp, 1897

In the present study, 400 species-level taxa, including synonyms, are listed or discussed: they
represent 237 valid species (141 Dentaliida and 96 Gadilida), of which 139 (81 Dentaliida and
58 Gadilida) have been taken during the different expeditions here reported on. Forty-two are
described as new (16 Dentaliida and 26 Gadilida), and three new genera are erected in the Order
Gadilida. Of the other 97 species (65 Dentaliida and 32 Gadilida) cited from the Indo-Pacific in the
literature, I have examined the type material of many in their respective repositories.

Order Dentaliida Da Costa, 1776
Family Dentaliidae Gray, 1847

Genus Dentalium Linne, 1758

Type species (SD by Montfort. 1810): D. elephantinum Linne, 1758.

Synonym: Lentigodentalium Habe, 1963. Type species (OD): Dentalium variabile Deshayes, 1825.

Diagnosis. — Shell medium to large, generally well curved and strong, usually polished. White
to yellow, orange, red, or green. Longitudinally sculptured by 8 to 12 primary ribs of varying
strength, simple or channeled. Secondary ribs variable in number, smooth or sculptured. Intercostal
spaces concave or convex, smooth or sculptured by longitudinal or transversal striae, or both. Apex
generally with a flat V-shaped notch on the ventral side. Apical callous, lumen circular and pipe are
common. Transversal section polygonal at the apex, usually circular at the mouth.

Radula rachidian very regular, well curved in section, anterior margin smooth or granulous;
lateral solid with primary cusp sharp and secundary shorts; marginal short and slightly sinusoi¬
dal.

Distribution. — Cretaceous-Recent, worldwide; temperate to tropical; sublittoral-shelf-bathyal.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

197

Dentalium elephantinum Linne, 1758
Figs 5, 16 a

Dentalium elephantinum Linne, 1758: 785.

Synonyms:

Dentalium arcuatum Gmelin, 1791: 3738 (after Gualtieri, 1757: pi. 10, figs G, I).

Dentalium viridis Perry, 1811: pi. 52, fig. 3.

Other references:

Dentalium elephantinum - Martini, 1769: 31, pi. 1, fig. 5a (first illustration of a live specimen). — Lister, 1770: pi. 547,
fig. 1 {pars). Gmelin, 1791: 3736. — Lamarck, 1801: 326. — Deshayes. 1825: 347, pi. 3, fig. 7. — Sowerby, 1860: 102.
pi. 223 ( Dentalium 1), fig. 4; 1873: pi. 1. fig. 5. — Pilsbry & Sharp. 1897: 1, pi. 1, figs 1-7. — Boissevain, 1906: 7, pi. 1, fig.
1. - Oostingh. 1925: 228. — Habe, 1962: 106. pi. 47. fig. 16; 1963: 253, pi. 37, fig. 3; 1964a: 6. pi. 1, fig. 3. - Emerson,
1962: 468, pi. 77, figs la-c. Habe & Kosuge, 1964: I. — Springsteen & Leobrera, 1985: 286. pi. 82, fig. 1. Higo &
Goto, 1993: 685.

Type material. Syntypes, Uppsala University Zoological Museum (fide Wallin, 1992; not
seen), and Linnean Society of London, no. 612 (fide Wheeler, 1993, not seen).

Type locality. — D. elephantinum'. Amboina [Ambon], Indonesia. — D. arcuatum: unknown.
— D. viridis: “South Seas”.

Material examined. — Papua New Guinea. Manokwari, Coll. Staadt, 2 dd. — Port Dorey,
Raffray coll., 4 dd (both mnhn).

Indonesia. Moluccas, Coll. Jousseaume, 3 dd (mnhn).

Philippines. Davao, Coll. Staadt, 2 dd. — Philippines, 1 dd (both mnhn).

Northern Indian Ocean. India, Coll. Denis, 1 dd. — Malabar, 2 dd (both mnhn).

Fig. 5. — Distribution of Dentalium elephantinum.

Distribution. — The Philippines, Indonesia, New Guinea, Northern Indian Ocean (present
paper), 5-40 m (Habe, 1964a).

198

VICTOR SCARABINO

Dentalium aprinum Linne, 1767
Figs 6, 16 b

Dentalium aprinum Linne, 1767: 1263.

Synonyms:

Dentalium striatulum Gmelin, 1791: 3738 (after Lister. 1770: pi. 574, fig. 1) (pars).

Dentalium interstriatum Sowerby, 1860: 102, pi. 223 (Dentalium 1), fig. 7.

Dentalium aprinum incolor Boissevain. 1906: 9, pi. 4, figs 3-6.

Dentalium aprinum taiwanum Kuroda, 1941: 149.

Other references:

Dentalium aprinum - Lister, 1770: pi. 547, fig. 1 (pars). — Sowerby, 1860: 102, pi. 223 (Dentalium 1), figs 5-6; 1873: pi. 1,
figs 2a-b. — Martens, 1880: 311. — Clessin. 1896: 12, pi. 3, figs 1-2. — Boissevain, 1906: 9, pi. I, fig. 3, pi. 4, fig. 2.
Dautzenberg, 1929: 553. — Dawydoff, 1952: 144. — Habe, 1963: 253, pi. 37, fig. 6; 1977: 330. Habe & Kosuge, 1964:
1; 1966: 117, figs 24-25. — Springsteen & Leobrera, 1985: 286, pi. 82, fig. 2. — Dharma, 1992: 79, fig. 14. — Higo & Goto,
1993: 685.

Dentalium aparinum (sic) — Cheriyan, 1968: 126. Saytamurti, 1956: 4, pi. I figs 2 a-b.

Dentalium apricum (sic) - Paetel, 1873: 78.

Dentalium aprinum taiwanum — Habe, 1962: 106. pi. 47, fig. 15.

Dentalium interstriatum - Clessin, 1896: 13, pi. 3, fig. 8. — Pilsbry & Sharp, 1897: 4, pi. 1, fig. 15.

Fig. 6. — Distribution of Dentalium aprinum.

Type material. — D. aprinum : syntypes in the Linnean Society of London, no. 61 1 (not seen).
Dentalium aprinum incolor, lectotype (here designated) zma 3.06.055 and paralectotypes zma
3.06.066-007. — D. interstriatum : syntypes (3 dd) bmnh 1950.11.28.42-44.

Type locality. — D. aprinum ; Indian Ocean. — D. aprinum incolor : Indonesia, anchorage off
Lirung, Silababu Island, “Siboga”, stn 133. — D. aprinum taiwanum : Taiwan, Takao. — D.
interstriatum-. Philippines, Bohol Island. — D. striatulum ; unknown.

Material examined. — Type material of D. aprinum incolor and D. interstriatum.

New Caledonia, musorstom 4: stn DW 150, 19°07' S, 163°22' E, 110 m, 1 lv, 20 dd.

Papua New Guinea. New Britain, 3 dd (mnhn).

Source : MNHN Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

199

Indonesia, corindon: stn CH 206, 01°06' S, 117°45'E, 85 m, 1 dd. — Stn DR 216, 00°40' N
1 17°51' E, 94 m, 26 dd. — Stn B 256, 01°56' S, 1 19° 174' E, 24 m, 3 dd.

“ Snellius " 1: Timor, 123 m, 1 dd (rmnii).

Philippines. Philippines, no further data, 24 dd (mnhn).

Distribution. — From Southern Japan, 5-40 m (Habe & Kosuge, 1964) through Taiwan,
the Philippines and Sulu Sea, to Indonesia and Papua New Guinea. Now extended to New Caledonia,
alive in 110 m.

Dentaliuni octangulatum Donovan, 1804
Figs 7, 16 c

Dentalium octangulatum Donovan. 1804: pi. 162.

Synonyms:

Dentalium octogonum Lamarck, 1818: 344.

Dentalium yokohamense Watson, 1879: 517; 1886: 11, pi. 2, fig. 1.

Dentalium japonicum Dunker, 1882: 153, pi. 5, fig. 2.

Other references:

Dentalium octangulatum - Pilsbry & Sharp. 1898: 16, pi. 2. figs 16-18, 22. Boissevain, 1906: 17, pi. 1, fig. 8, pi. 4.
fig. 8-9. Dautzenberc & Fischer, 1906: 209. Tokunaga. 1907: 10. — Winckworth, 1927: 167, pi. 14, fig. 4. — Kuroda
& Kikuchi, 1933: 7. Nomura, 1938: 155. — Dawydoff, 1952: 144. Saytamurti, 1956: 126. — Ahmed, 1975: 29.
fig. 32. — Habf., 1977: 330. Springsteen & Leobrera. 1985: 286, pi. 82. fig. 3. — Qi & Ma, 1989: 112, fig. 2. — Ieyama,
1993: 245, figs 1-3.

Dentalium ( Paradentalium ) octangulatum - Habe. 1963: 254, pi. 37, figs 1-2; 1971: 486 (Japanese text), 305 (English text),
pi. 65, figs 10-11. — Habe & Kosuge, 1964: 1. — Kira, 1955: 80. pi. 40, fig. 8. — Higo & Goto, 1993: 685.

Dentalium ( Dentalium ) octangulatum - Hirase, 1931: 133, figs 1-4. — Ludbrook. 1954: 96. fig. 1. Kira, 1962: 116, pi. 41.
fig. 8.

Dentalium octogonum - Deshayes. 1825: 352. pi. 16, figs 5-6. — Delessert. 1841: pi. 1, figs 1-la-b. — Chenu, 1843: 4,
pi. 1, figs 21-23. — Reeve, 1842b: 130. pi. 130, fig. 8. Sowerby, 1860: 102, pi. 223, fig. 9; 1873: pi. 2, fig. 12. — Martens.
1874: 102. — Lischke, 1874: 75, pi. 5, figs 1-3. — Smith. 1875: 25. — Brazier. 1877: 55. Angas, 1878: 868. — Dunker,
1882: 153. Pilsbry. 1895: 116. Clessin, 1896: 10. pi. 1, fig. 6. — Hall & Standen, 1907: 65.

Dentalium octogonum - Stearns. 1891: 13.

Dentalium ( Paradentalium ) octangulum - Fukuda, 1992: 81, fig. 421.

Fig. 7. Distribution of Dentalium octangulatum.

200

VICTOR SCARABINO

Type material. — D. octangulatum: neotype, designated by Ludbrook (1954), bmnh
1952.2.23.1. — D. octogonum : syntypes (2 dd) mnhn (one of which figured by Delessert, 1841). —
D. yokohamense : syntypes (3 dd) bmnh 1887.2.9.45-47. — D. japonicum : holotype zmb 101995 ( fide
Kilias, 1995).

Type locality. D. octangulatum: Ludbrook (1954) designated Japan as type locality, while
Donovan’s original was China Seas. — D. octogonum : "Mers de la Chine”. — D. yokohamense :
Japan, Yokohama, “ Challenger ”, stn 233, 34°39' N, 135°14' E, 8 fms [15 m], — D. japonicum: Japan.

Material examined. — The type material of D. octangulatum, D. octogonum and D.
yokohamense.

Japan. No further data, Coll. Jousseaume, 2 dd (mnhn).

China. No further data, 6 dd (mnhn).

Northern Indian Ocean. Karikal, Bay of Bengal, 4 dd (mnhn). — Persian Gulf, Dubay’ah, S. Pras
coll., 20 dd (mnhn). — Jebel Dana and Abu Dhabi, 10 dd (bmnh). — Oman, Salalah, shore, 32 dd
(bmnh).

Distribution. — Japan, China Seas, Philippines, Northern Indian Ocean, Persian Gulf, shells
in 0-100 m (Habe & Kosuge, 1964).

Dentalium variabile Deshayes, 1825
Figs 8, 16 d-e

Dentalium variabile Deshayes, 1825: 352, pi. 2, fig. 30.

Synonyms:

Dentalium multistriatum Deshayes, 1825: 358, pi. 4, fig. 11 (Syn. nov.).

Dentalium belcheri Sowerby, 1860: 101, pi. 24 ( Dentalium 2), figs 28-29 (Syn. nov.).

Other references:

Dentalium variabile - Sowerby, 1860: 101, pi. 224 ( Dentalium 2), fig. 30; 1873: pi. 4, fig. 26. Martens 1887: 200. -

Boissevain, 1906: 36. pi. 1, fig. 17. — Clessin, 1896: 14. pi. 4, fig. 1. — Pilsbry & Sharp, 1897: 60, pi. 14, figs 26-28.
Lentigodentalium variabile - Habe, 1977: 332. — Habe el a!.. 1986: 24. — Chistikov, 1979b: 112. Higo & Goto, 1993:
686.

Dentalium ( Lentigodentalium ) variabile - Habe, 1963: 258, figs 53-54. — Habe & Kosuge, 1964: 2.

Dentalium belcheri - Sowerby, 1873: pi. 1, figs la-b. Pilsbry & Sharp, 1897: 60, pi. 14, figs 29-30. — Boissevain, 1906:
35, pi. 1, fig. 18. — Smith, 1906b: 58.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PAC1FIC

201

Type material. — D. variabile : lectotype (here designated) (L 22.5, W 2.4, w 1.4) from
164 syntypes (mnhn). — D. multistriatum : lectotype (here designated) (L 18.8, W 2, w 1) from 14 syn-
types (mnhn). — D. belcheri : lectotype (here designated) the largest (25 mm) of the 3 syntypes bmnh.

Type locality. D. variabile-. “could be from India”. — D. multistriatum : “presumably from
India” (Desha yes 1825). — D. belcheri : East Indian Archipelago [= Indonesia],

Material examined. — The type material.

Chesterfield Islands, chalcal 1: stn DC 40, 20°32' S, 158°5T E, 65 m, 1 dd.

New Caledonia, lagon: stn 7, 22°24' S, 166°20' E, 14 m, 7 dd. — Stn 8, 22°23' S, 166°18' E, 15 m,
1 dd. — Stn 21, 22°23' S, 166°23' E, 10 m, 1 dd. — Stn 50, 22°17' S, 166°12' E. 12 m, 1 lv, 2 dd. —
Stn 64, 22°28' S, 166°25' E, 15 m, 2 lv. — Stn 98, 22°36' S, 166°32' E, 15 m, 1 lv. — Stn 162, 22°13' S,
1 66°09' E, 10 m, 4 lv. — Stn 212, 21°56' S, 165°53' E, 10 m, 2 lv, 1 dd. — Stn 214, 21°55' S, 165°48'E,
12 m, 4 dd. — Stn 293, 22°42' S, 166°41' E, 20 m, 1 lv. — Stn 867, 20°39' S, 165°01' E, 25 m, 1 dd.
Philippines. No further data, 11 dd. — ibid.. Coll. Jousseaume, 73 dd, (all mnhn).

West Indian Ocean, md 32 Reunion: stn DC 85, 21°00' S, 55°15' E, 58-70 m, 60 dd. — Stn DC 86,
20°59' S, 55° 15' E, 75-90 m, 7 dd.

Distribution. — Japan, China Seas, the Philippines, Indonesia and India (?), now extended
to New Caledonia and Reunion Island, alive in 10-75 m (present paper).

Dentalium strigatum Gould, 1859
Figs 9, 16 f

Dentalium strigatum Gould. 1859: 166.

Synonym:

Dentalium agulhasense Plate. 1908a: 349, pi. 30, figs 21-23 (Syn. nov.).

Other references:

Dentalium strigatum - Smith, 1903: 393. Bartsch, 1915: 180, pi. 44, fig. 5. — Tomlin, 1931: 337. — Jaeckel, 1932: 303.
— Barnard. 1963b: 345; 1974: 742. — Johnson. 1964: 153.

Source :

202

VICTOR SCARABINO

Type material. — D. strigatum : lectotype (designated by Johnson, 1964) usnm 159,
paralectotypes mcz 169382. — D. agulhasense : lectotype, designated by Kilias (1995), zmb 61087a.

Type locality. — D. strigatum: False Bay, Cape of Good Hope, 20-80 m. — D. agulhasense:
South Africa, off Cape Agulhas, “ Valdivia ”, stn 95, 34°51' S, 19°38' E, 80 m.

Material examined. — The type material of D. strigatum.

West Indian Ocean, md 32 Reunion: stn CP 172, 20°52' S, 55°38' E, 105-120 m, 1 dd.

“Meiring Naude": stn SM 180, 33°29' S, 27°21' E, 80 m, 2 dd (sam).

Distribution. — South Africa, Mozambique Channel (Barnard, 1963b); now extended to
Reunion Island; shells in 80-120 m.

Denta'.ium bisexangulatum Sowerby, 1860
Figs 10, 16 j

Detualium bisexangulatum Sowerby, I860: 102, pi. 223 (Dentalium 1), fig. 8.

Other references:

Dentalium bisexangulatum - Sowerby, 1873: pi. 3, fig. 15. — Brazier, 1877: 57. — Cooke, 1885: 273. — Pilsbry & Sharp,
1897: 15, pi. 2, fig. 25. — Boissevain, 1906: 22, pi. 1, fig. 7.

Dentalium (Dentalium) bisexangulatum - Ludbrook, 1954: 92. — Higo & Goto, 1993: 685.

D. ( P .) bisexangulatum - Habe & Kosuge, 1964: 2.

Fig. 10. — Distribution of Dentalium bisexangulatum.

Type material. — 3 syntypes dd, bmnh no reg. no.

Type locality. — Java.

Material examined. — The type material.
Indonesia. Java, Coll. Jousseaume, 1 dd (mnhn).

Philippines. Philippines, Coll. Jousseaume, 1 dd (mnhn).
Japan. Booshu, Coll. Denis, 1945, 2 dd (mnhn).

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

203

Northwestern Indian Ocean. Persian Gulf, shore of Iran, 26°35' N, 54°00' E, 35 m, 2 dd, “ Calypso "
1954 (mnhn).

West Indian Ocean. Madagascar, Tulear, N of jetty, shore, 10 dd. — Tulear, Grand Recif, 1 m, 4 dd,
R. v. Cosel coll. 1986. — Zanzibar, Coll. Cloue, 1850, 5 dd (mnhn).

Distribution. — Japan, Singapore, Java, Torres Strait, Gulf of Suez, Mozambique (14-55 m)
(Ludbrook, 1954); Philippines, Persian Gulf, Zanzibar (present paper). No data about living depth
range.

Dentalium javanum Sowerby, 1860
Figs 11, 16 i

Dentalium javanum Sowerby, 1860: 102, pi. 223 ( Dentalium 1), fig. 12.

Other references:

Dentalium javanum - Sowerby, 1873: pi. 3, fig. 14. — Smith, 1884: 77. — Pilsbry & Sharp, 1897: 4, pi. 4, fig. 49. — Melvill
& Standen, 1899: 181. — Boissevain, 1906: 18, pi. 1, fig. 6; pi. 4, fig. 7, textfig. 13. — Habe, 1977: 331."

Dentalium ( Paradentalium ) javanum - Habe, 1963: 254, pi, 37, figs 7-8; 1964a: 9, pi. 1, figs 7-8.

Dentalium (Paradentalium) javanicum — Higo & Goto, 1993: 685.

Type material. — Not found in bmnh.

Type locality. — Java, Malacca.

Material examined. — Indonesia. “Java, Malacca”, Coll. Wright 1869, 2 dd. — Java, Coll.
Jousseaume, 1 dd (both mnhn).

Distribution. — Taiwan, Philippines, Indonesia, New Guinea, North Australia, India, 5-
50 m (Habe, 1964b).

204

VICTOR SCARABINO

Dentalium reevei Fischer, 1871
Figs 12, 16 g-h

Dentalium reevei Fischer (ex Deshayes, MS), 1871: 212.

Synonyms:

Dentalium aratorum Cooke. 1885: 273.

Dentalium lineolatum Cooke. 1885: 274.

Dentalium clavus Cooke, 1885: 275.

Dentalium laugieri Jousseaume, 1894: 103.

Dentalium macandrewi Boissevain, 1906: 25, pi. 5, figs 25-34.

Other references:

Dentalium reevei - Boissevain, 1906: 27. — Moazzo, 1939: 221.

Dentalium (Dentalium) reevei - Ludbrook. 1954: 98.

Dentalium aratorum - Pilsbry & Sharp, 1897: 10. - Boissevain, 1906: 27, pi. 5. figs 31-34. — Franc, 1956: 49.
Dentalium lineolatum - Pilsbry & Sharp, 1897: 11. — Boissevain, 1906: 26, pi. 5, figs 25-27. — Lamy, 1910: 334; 1938: 88.
— Dautzenberg, 1929: 554.

Dentalium laugieri - Pilsbry & Sharp, 1897: 12. Boissevain, 1906: 25.

Dentalium clavus - Boissevain. 1906: 27. pi. 5, figs 28-30.

Fig. 12. Distribution of Dentalium reevei.

Type material. — D. reevei : lectotype (here designated), the only remaining syntype, mnhn.

— D. laugieri ; 10 syntypes dd mnhn. — D. macandrewi : lectotype (here designated), the specimen
figured in Boissevain, 1906, pi. 5, fig. 29 (zma). — D. aratorum , D. lineolatum and D. clavus ; not
located.

Type locality. D. reevei. Suez. — D. aratorum , D. lineolatum and D. clavus : Gulf of Suez.

— D. laugieri. Aden, Suez. — D. macandrewi Gulf of Suez.

Material examined. — The type material listed.

Northwestern Indian Ocean. Red Sea: Saudi Arabia, 24°12' N, 37°55' E, Marioni coll., 2 dd. Port
Sudan, harbour, 1 m, Reid coll., 9 Iv, 1 dd (both bmnh). — Souakin, Sudan, 1 dd. — Aden, 2 dd
(both Coll. Jousseaume, mnhn).

West Indian Ocean, benthedi: stn S 18, 12°45' S, 45° 1 6' E, 15 m, 1 lv, 6 dd. Stn S 23, 12°46' S,

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

205

45° 15' E, 6 m, 1 lv, 1 dd. — Stn S 32, 12°45' S, 45° 18' E, 15-20 m, 1 dd. — Stn S 36, 12°52'S,
45° 16' E, 30 m, 2 dd.

Shimoni, Kenya, 2 lv, 5 dd. — 1 mile E Nyango, W Zanzibar, 18 fms [33 m], 1 dd (both bmnh).
Kiloa, Zanzibar, 3 dd. — Tulear, Madagascar, 2 dd. — Tulear, S beach, Thomassin coll., 1 dd (all
mnhn). — Nosy Be Island, NW Madagascar, 5 m, 2 lv, Plante coll. (bmnh).

Distribution. — Red Sea and the Gulf of Aden; now extended to East Africa and
Madagascar, alive in 6-32 m.

Remarks. — Although Boissevain (1906) introduced the name D. macandrewi as a nomen
novum for D. aratorum, D. lineolatum and D. clavus, none of the Cooke’s names is preoccupied. D.
macandrewi is to be treated as a nominally different species, with its own type material. D. reevei is
extremely variable in shape (W/w ratio 3. 5-1. 5) and colour (white to green). The ribs can be simple
to deeply channeled longitudinally, smooth to slightly sculptured transversally. The apex is also
variable in width, probably due to reabsorption.

Dentalium oryx Boissevain, 1906
Figs 13, 16 k, 71 d

Demalium oryx Boissevain, 1906: 20, pi. 6, fig. 24.

Synonym:

Dentalium paucicontortum Boissevain, 1906: 28, pi. 6, figs 25-28 (Syn. now).

Other references:

Entalinopsis ( Entalinospsis ) oryx — Habe & Kosuge, 1964: 9.

Dentalium ( Paradentalium ) paucicontortum — Habe, 1963: 255.

Type material. — D. oryx: lectotype (here designated) zma 3.06.014, paralectotypes zma
3.06.015. — D. paucicontortum: lectotype (here designated) zma 3.06.022, paralectotypes zma
3.06.023.024.

Type locality. — D. oryx: "Siboga”, stn 302, 1 0°28' S. 123°29' E, 216 m, South of Sunda
Island. — D. paucicontortum: Indonesia, Sulu Archipelago, " Siboga ”, stn 95, 05°44' N, 119°40'E,
522 m.

Material examined. — Chesterfield Islands, musorstom 5: stn DW 306. 22°08' S, 1 59°2 1 ' E,
375-415 m, 1 lv, 1 dd. — Stn DW 340, 19°49' S, 158°41' E, 675-680 m, 1 dd. — Stn DW 341, 19°46' S,
158°43' E, 620-630 m, 1 lv, 1 dd. — Stn DW 357, 19°37' S, 158°46' E, 630 m, 6 dd. — Stn DW 358,
19°39' S, 1 58°47' E, 680-700 m, 1 dd. — Stn DC 361, 19°53' S, 158°38' E, 400 m, 1 dd. — Stn DC
362, 19°53' S, 158°40' E, 410 m, 1 lv. Stn DC 381, 19°38' S, 158°47' E, 620 m, 1 dd. — Stn DC
382, 19°37'S. 158°43'E, 580 m, 1 dd.

New Caledonia. “Vauban” 1978-79: stn 40, 22°30' S, 166°24' E, 250-350 m, 28 dd.
biocal: stn DW 36, 23°09' S, 167°1 1' E, 650-680 m, 1 dd. — Stn DW 44, 22°47' S, 167° 14' E, 440-450
m, 4 dd. — Stn DW 66, 24°55' S, 168°22' E, 505-515 m, 3 lv, 4 dd. — Stn DW 70, 23°25' S, 167°53' E,
965 m, 1 lv.

musorstom 4: stn DW 226, 22°47' S, 167°22' E, 390 m. 1 dd.
chalcal 2: stn DW 72, 24°55' S, 168°22' E, 527 m, 2 lv.
lagon: stn 993. 20° 15' S, 163°53' E, 375-400 m, 2 dd.

Passe de Boulari. 400 m, B. Richer/ORSTOM coll., 6 dd.

Loyalty Islands, musorstom 6: stn DW 397, 20°47' S, 167°05'E, 380 m, 1 dd. — Stn DW 398,
20°47' S, 167°06' E, 370 m. 1 dd. — Stn DW 416, 20°42' S, 167°00' E, 343 m, 1 dd. — Stn DW 418,

206

VICTOR SCARABINO

20°42' S, 167°03' E, 283 m, 1 dd. — Stn DW 425, 20°24' S, 166°25' E, 594 m, 3 dd. — Stn DW 439,

20°46' S, 167° 17' E, 288 m, 3 dd. — Stn DW 440, 20°49' S, 167°17' E, 288 m, 2 dd. — Stn DW 451,

20°59' S, 167°25' E, 330 m, 3 dd. — Stn DW 452, 21°00' S, 167°25' E, 300 m, 1 dd. — Stn DW 481,

21°22' S, 167°50' E, 300 m, 3 dd. — Stn DW 485, 21°23' S, 167°59' E, 350 m, 2 dd. — Stn DW 487,

21°23' S, 1 67°46' E, 500 m, 2 dd. — Stn DW 488, 20°49' S, 167°06' E, 800 m, 1 lv, 1 dd.

Indonesia, corindon: stn B 248, 00°54' S, 119°29' E, 170 m, 3 dd.

“Snell ius" II: stn 4.166, 06°26' S, 120°27' E, 300 m, 5 dd.

Philippines, musorstom 3: stn DR 126, 11°49'N, 121°22' E, 266 m, 1 dd.

West Indian Ocean, benthedi: stn DR 08, 11°29'S, 47°18' E, 250 m, 3 lv, 2 dd. — Stn DS 10,
1 1°29' S, 47° 18' E, 440 m, 1 dd. — Stn F 49, 12°55' S, 44°59' E, 300-450 m, 1 lv. — Stn DR 34,
12°54' S, 45° 16' E, 500 m, 1 lv. — Stn 37, 12°54’S, 45°16' E, 520-830 m, 1 dd. — Stn DS 64, 12°41' S,
44°57' E, 770-860 m, 3 lv. — Stn DR 104, 1 1°26' S, 47°22' E, 330-530 m, 2 lv, 1 dd. — Stn DS 120,
1 1°30' S, 47°25' E, 335-390 m, 2 dd.

md 32 Reunion: stn DR 62, 21°09' S, 55°12' E, 630-710 m, 46 dd.

Fig. 13. — Distribution of Denialium oryx.

Distribution. — Indonesia; now extended to the Philippines, New Caledonia, and SW Indian
Ocean; alive in 216-965 m.

Dentalium leucoryx Boissevain, 1906
Figs 14, 16 1

Denialium leucoryx Boissevain, 1906: 20, pi. 6, fig. 23.

Synonym:

Denialium sinuosum Boissevain, 1906: 28, pi. 6, fig. 22 (Syn. nov.).

Other references:

Denialium leucoryx - Habe & Kosuge, 1964: 2.

Denialium sinuosum - Habe & Kosuge, 1964: 2. — Qi & Ma, 1989: 112, fig. 1.

Paradenlalium sinuosum - Chistikov, 1979b: 109.

Type material. — D. leucoryx : lectotype (here designated) zma 3.06.016, paralectotypes zma
3.06.017. — D. sinuosum : lectotype (here designated) zma 3.06.021, paralectotypes zma 3.06.018-020.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

207

Type locality. — D. leucoryx : Indonesia, off Sunda Island, “ Siboga ”, stn 285, 08°39' S,
127°04'E, 34 m. — D. sinuosum : Indonesia, Timor Sea, “ Siboga ”, stn 294, 10°12'S, 124°27' E,
73 m.

Material examined. — The type material.

Indonesia, corindon: stn B 207, 00°15' S, 1 17°52' E, 150 m, 8 dd. — Stn CH 208, 00°15' S, 1 17°52' E,
150 m, 1 lv. — Stn CH 214, 00°31'N, 117°50 E, 595 m, 2 dd. — Stn B 251, 00°54' S, 119°30'E,
65 m, 2 dd.

“Snellius” I: stn 123, 10°29' S, 126°44' E, 250 m, 1 dd.

“ Snellius ” II: stn 4.045, 05°57' S, 123°49' E, 250-300 m, 3 dd. — Stn 4.131, 08°18' S, 118°18'E,
600-800 m, 1 dd.

Distribution. South China Seas to Indonesia, alive in 150-157 m. shells from 34 m
(Boissevain, 1906) to 800 m (present paper; possibly washed down).

Dentalium pluricostatum Boissevain, 1906
Figs 15, 16 m, 28 k

Dentalium pluricostatum Boissevain, 1906: 30, pi. 6, figs 6-7.

Other reference:

D. pleuricostatum (sic) - Habe & Kosuge. 1964: 2.

Type material. Lectotype (here designated) zma 3.06.027, paralectotypes zma 3.06.025-

026.

Type locality. — Indonesia, Buton Strait, “Siboga”, stn 204, 04l,20' S, 122°58' E, 75-94 m.
Material examined. — The type material.

New Caledonia, lagon: stn 69, 22°23' S, 166°32' E, 13 m, 4 lv, 8 dd. — Stn 131, 22°28' S, 166°50' E,
38 m, 1 lv, 2 dd. — Stn 133, 22°24' S, 166°52' E, 59-62 m, 1 dd. — Stn 146, 22°24' S, 166°55' E. 40-

Source :

208

VICTOR SCARABINO

52 m, 2 lv, 3 dd. — Stn 147, 22°26' S, 166°54' E, 50-60 m, 5 dd. — Stn 149, 22°29' S, 166°51' E,
48 m, 2 lv, 4 dd. — Stn 234, 22°33' S, 166°51' E, 56 m, 6 lv, 12 dd. — Stn 234bis, 22°33' S, 166°51' E,
60 m, 2 dd. — Stn 235, 22°31' S, 166°52' E, 70 m, 1 lv, 3 dd. — Stn 236, 22°29' S, 166°54' E, 67 m,
2 lv, 3 dd. — Stn 237, 22°27' S, 166°55' E, 62 m, 1 dd. — Stn 238, 22°26' S, 1 66°56' E, 50 m,

1 lv. — Stn 240, 22°23' S, 166°59' E, 42 m, 2 dd. — Stn 245, 22°27' S, 166°58' E, 62 m, 3 lv, 5 dd.

— Stn 257, 22°22' S, 166°20' E, 9 m, 1 lv. — Stn 301, 22°35' S, 166°52' E, 46 m, 1 lv. — Stn 317,
22°33' S, 166°53' E, 66 m, 1 lv, 2 dd. — Stn 318, 22°34' S, 166°55' E, 71 m, 1 lv, 3 dd. — Stn 319,
22°32' S, 166°57' E, 75 m, 3 lv, 3 dd. — Stn 320, 22°32' S, 166°54' E, 70 m, 1 lv, 2 dd. — Stn 321,
22°30' S, 1 66°56' E, 70 m, 1 lv. — Stn 322, 22°30' S, 166°58' E, 71 m, 1 lv, 20 dd. — Stn 323, 22°29' S,
166°59'E, 80 m, 1 dd. — Stn 325, 22°27' S, 167°01' E, 75 m, 4 lv, 2 dd. — Stn 326, 22°26' S,

1 67°02' E, 67 m, 4 lv, 18 dd. — Stn 328, 22°27' S, 167°03' E, 72 m, 3 lv, 2 dd. — Stn 329, 22°29' S,
167°02' E, 80 m, 7 lv, 4 dd. — Stn 330, 22°31' S, 167°00' E, 82 m, 1 lv, 3 dd. - Stn 331, 22°33' S,

1 66°59' E, 79 m, 3 lv, 3 dd. — Stn 332, 22°34' S, 166°57' E, 80 m, 3 lv, 1 dd. — Stn 333, 22°37' S,
166°56' E. 71 m, 7 lv. — Stn 350, 22°39' S, 166°57' E, 67 m, 4 lv, 6 dd. — Stn 352, 22°35' S, 166°60' E,

82 m, 3 lv, 7 dd. — Stn 354, 22°32' S, 167°02' E, 78 m, 7 lv, 3 dd. — Stn 355, 22°30' S, 167°04' E,

82 m, 5 lv, 1 dd. — Stn 356, 22°29' S, 167°05' E, 78 m, 4 lv, 3 dd. — Stn 357, 22°30' S, 167°07' E,

77 m, 3 lv, 5 dd. — Stn 359, 22°33' S, 167°04' E, 74 m, 3 lv. — Stn 361, 22°36' S, 167°02' E, 78 m,

I lv. — Stn 362, 22°38'S, 167°00' E, 83 m, 1 dd. — Stn 376, 22°34' S, 167°06'E, 75-76 m, 1 lv,

II dd. — Stn 386, 22°37' S, 167°09' E, 128 m, 1 lv. — Stn 429, 22°40' S, 167° 15' E, 95 m, 6 lv. -
Stn 537, 19°07' S, 163°22' E, 200 m, 3 dd. — Stn 538, 19°07' S, 163°21' E, 195 m, 1 lv, 7 dd. — Stn
539, 19°05' S, 163°17' E, 240 m, 2 lv, 2 dd. — Stn 603, 22°16' S, 167°05' E, 78-80 m, 1 lv, 6 dd.

Stn 604, 22° 14' S, 167°04' E, 80 m, 10 lv, 10 dd. — Stn 605, 22°15' S, 167°02' E. 65-70 m, 4 lv. -
Stn 606, 22° 13' S, 167°01' E, 46-48 m, 1 lv. — Stn 61 1, 22°09' S, 166°59' E, 56-57 m, 1 dd. — Stn 615,
22°07' S, 166°57' E, 56-60 m, 3 lv. — Stn 618, 22°05' S, 166°56' E, 53-58 m, 1 dd. — Stn 619, 22°03' S,
166°54' E, 27-42 m, 1 lv. — Stn 622, 22°02' S, 166°53' E, 67 m, 3 lv, 5 dd. — Stn 628, 22°00' S,

1 66°49' E, 55-56 m, 1 lv, 1 dd. — Stn 630, 21°59' S, 166°46' E, 60-68 m, 2 lv, 1 dd. — Stn 638,

21°56' S, 1 66°40' E, 56-58 m, 2 lv. — Stn 643, 21°53' S, 166°40' E, 56-66 m, 3 lv, 8 dd. — Stn 646,
21°52' S. 166°38' E, 66-70 m, 5 lv, 4 dd. — Stn 647, 21°54' S, 166°37' E, 50-52 m. 3 lv, 3 dd. — Stn

648, 21°53' S, 166°35' E, 22-25 m, 1 dd. — Stn 649, 21°51' S, 166°37' E, 64-65 m, 3 lv, 7 dd. — Stn

652, 21°50' S, 166°35' E, 55-62 m. 6 dd. — Stn 655, 2P48' S, 166°31' E, 35-40 m, 2 lv. — Stn 656,
21°49' S, 166°33' E, 30-40 m, 1 lv. — Stn 660, 21°47' S, 166°33' E, 48-52 m, 1 dd. — Stn 665, 21°45' S,
166°28' E, 40-42 m, 1 lv, 1 dd. — Stn 674, 21°38' S, 166°23' E, 48 m, 1 dd. — Stn 692, 21°32' S,

166° 12' E, 44-48 m, 1 dd. — Stn 694, 21°32' S, 166° 10' E, 45-47 m, 2 lv. — Stn 695, 21°31'S,

166°1 1' E, 54-55 m, 1 lv, 1 dd. — Stn 699, 21°31' S, 166°08' E, 50-52 m, 1 dd. — Stn 712, 21°25' S,

1 66°00' E, 47-49 m, 2 lv. — Stn 728, 21°2E S, 165°52' E, 43-47 m, 2 lv, 2 dd. — Stn 737, 22°08' S,

166°59'E, 49-50 m, 1 lv. — Stn 738, 22°10'S, 167°00' E, 59-61 m, 1 lv. — Stn 749, 21°18' S,
165° 18' E, 49 m, 1 dd. — Stn 768, 21°14' S, 165°39' E, 28 m, 1 dd. — Stn 785, 21°08' S, 165°33' E,
37 m, 1 dd. — Stn 791, 21°07' S, 165°31' E, 33 m, 1 dd. — Stn 809, 20°56' S, 165°28' E, 34 m, 1 dd.

— Stn 813, 22°51' S, 165°25' E, 47 m, 1 lv. — Stn 814, 21°56' S, 165°26' E, 38-50 m, 1 lv, 1 dd. -
Stn 815, 21°54' S, 165°27' E, 32 m, 2 lv. — Stn 821, 20°52' S, 165°23' E, 32 m, 1 dd. — Stn 827,
20°52' S, 165°18' E, 53 m, 1 dd. — Stn 831, 20°50' S, 165°18' E, 73 m, 1 dd. — Stn 832, 20°51' S,
165°13' E, 32 m, 1 lv. — Stn 833, 20°50' S, 165°18' E. 52-70 m, 2 lv, 2 dd. — Stn 834, 20°48' S,
165° 16' E, 58 m, 1 dd. — Stn 878, 20°32' S, 164°48' E, 54 m, 2 dd. — Stn 901, 20°13' S, 164°22' E,
22-40 m, 1 lv.

musorstom 4: stn DW 149, 19°08' S, 163°23' E, 155 m, 3 lv, 70 dd. — Stn DW 150, 19°07' S,
163°23' E, 110 m, 6 lv, 18 dd. — Stn DW 151, 19°07' S, 163°22' E, 200 m, 1 lv, 10 dd. — Stn DW
186, 19°07' S, 163°30' E, 190 m, 1 dd. — Stn DW 203, 22°36' S, 167°05' E, 105-1 10 m, 1 lv. — Stn
DW 232, 22°29' S, 167°05' E, 77 m, 1 lv.

Loyalty Islands, musorstom 6: stn DW 441, 20°54' S, 167°17' E, 80 m, 1 dd. — Stn DW 442, 20°54' S,
167° 17' E, 200 m, 1 lv, 10 dd.

Philippines, musorstom 2: stn DG 32, 13°40'N, 1 20°54' E, 192-220 m, 1 dd.
musorstom 3: stn DR 140, 11°43' N, 122°34' E, 93-99 m, 14 dd.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

209

West Indian Ocean, benthedi: stn DR 38, 12°55' S. 45°16' E, 200-500 m, 1 dd. - Stn F 49, 12°55' S,
44°57' E, 300-450 m, 1 dd.

Distribution. — The Philippines and Indonesia, now extended to New Caledonia and NW
Madagascar; living from 9 to 240 m, shells down to 500 m (present paper).

Dentalium caledonicum sp. nov.

Figs 17, 28 a, 71 c

Type material. — Holotype and 2 paratypes. mnhn.

Type locality. New Caledonia, Northern Norfolk Ridge, chalcal 2, stn DW 74,
24°40'S, 168°38'E, 650 m.

Material examined. — New Caledonia, chalcal 2: stn DW 73, 24°40' S, 168°38' E, 573 m.
1 lv (paratype). — Stn DW 74, 24°40' S, 168°38' E, 650 m, 1 lv (holotype). — Stn DW 75, 24°39' S,
168°40' E, 600 m, 1 lv (paratype).

Distribution. — SE New Caledonia, alive in 573-650 m.

Description. — Shell to 30 mm long, nearly straight, shiny
white, solid, 15 primary ribs with rounded edges. Secondary
ribs present on the dorsal side or with only vestigal undula¬
tions reaching the fragile, circular oral aperture. Apex simple,
truncate, with a shallow V-shaped notch in the ventral side.

Lumen circular, suboval at the end of a short projecting pipe
fissured on dorsal side.

Measurements: holotype L 26.4, W 3.5, w 2.1, arc 0.8;
paratypes L 30.1, W 3.4, w 2.2, arc 0.5; L 27.5. W 3.5, w 2.1,
arc 0.6. W/w ratio 1.54-1.67.

Etymology. — From New Caledonia.

Fig. 16. - a, Dentalium elephantinum, shell (72 mm) and oral section, India (mnhn). b. Dentalium aprinum , shell (75 mm),
apex, apical and oral sections, detail of the sculpture. Papua New Guinea (mnhn). - c, Dentalium octangulatum, shell
(40 mm), apex, apical and oral sections. — d, Dentalium variabile, lectotype (22.5 mm), apex and apical section, detail
of the sculpture. — e. Dentalium variabile , apex and apical section (1.7 mm), detail of rib and sculpture. New Caledonia,
lagon: stn 7. f, Dentalium strigatum. shell (18 mm), apical section, detail of the sculpture, md 32 Reunion: stn
CP 172. - g, Dentalium reevei, lectotype (29.8 mm), shell and apical section. — h, Dentalium reevei , shell (23 mm), apex,
apical section, detail of the sculpture, Tulear (mnhn). i, Dentalium javanum , shell (55 mm) and oral section, Java,
Malacca (mnhn). — j, Dentalium bisexangulatum , shell (72 mm), apex and apical section, detail of the sculpture and
oral section, Tulear (mnhn). — k, Dentalium oryx, shell (32 mm), apex, apical and oral sections, detail of sculpture.
benthedi: stn DR 08. — I, Dentalium leucoryx, shell (47 mm), apex, apical and oral sections, corindon: stn CH 208.

m. Dentalium pluricostatum, shell (49 mm), apex, apical section and detail of the sculpture, musorstom 4: stn
DW 149.

Source : MNHN. Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

211

Fig. 17. — Distribution of Dentalium caledonicum.

Dentalium crosrtieri sp. nov.

Figs 18, 28 b

Type material. — Holotype and 7 paratypes, mnhn.

Type locality. — New Caledonia, Loyalty Islands, musorstom 6, stn DW 428, 20°24' S,
166° 13' E, 420 m.

Material examined. — New Caledonia, biogeocal: stn DW 292, 20°28' S. 166”48' E. 465-
470 m, 1 dd (paratype).

Loyalty Islands, musorstom 6: stn DW 428, 20°24' S, 166°13' E, 420 m, 2 lv, 1 dd (holotype lv and
paratypes). — Stn DW 459, 21°0T S, 167°31' E. 425 m, 1 1 dd (2 paratypes). — Stn DW 487, 21°23' S,
167°46' E, 500 m, 1 lv, 1 dd (paratypes).

Distribution. — New Caledonia and Loyalty Islands, alive in 420-500 m.

Source :

212

VICTOR SCARABINO

Description. — Shell to 32 mm long, needle-like, almost
straight, solid, white, shiny, with 15-17 primary ribs with
rounded edges. Secondary ribs less prominent, but more so
towards oral aperture. Intercostal spaces with coarse growth
lines only. Oral aperture circular. Apex simple, truncate,

circular to slightly flattened dorso-ventrally, lumen circular.

Measurements: holotype L 30.7, W 1.5, w 0.7, arc 0.9;
paratypes L 31.5, W 1.6, w 0.9, arc 0.9; L 30.5, W 1.7, w 1.3,
arc 0.8; L 23.5, W 1.4, w 0.6, arc 0.8. W/w ratio 1.31-2.34.

Remarks. — The slight curvature and W/w ratio are the main distinguishing characters of this
species. D. oryx, its closest relative, differs in apical section and W/w ratio (range 3.6-5. 8 in
10 specimens).

Etymology. — Named for Alain Crosnier, orstom, without whom there would have been no
musorstom expeditions. In addition to making possible and participating himself in seven cruises at
sea between 1976 and 1992, A. Crosnier has edited most of the report volumes in the series where
the present paper is published.

Dentalium flavum sp. nov.

Figs 19, 28 c, e

Type material. — Holotype mnhn. Paratypes; 10 mnhn, 1 ams C201722, 1 nmnz M268960.

Type locality. — Chesterfield Islands, musorstom 5, stn DW 301, 22°07' S, 159°25'E,
487-610 m.

; i —

Fig. 19. — Distribution of Dentalium flavum.

Material examined. — Chesterfield Islands, musorstom 5: stn DW 301, 22°07' S, I59°25' E,
487-610 m, 2 Iv (holotype and paratype). — Stn DW 306, 22°08' S, 159°21'E, 375-415 m, 1 lv
(paratype).

New Caledonia, lagon: stn 444, 18° 15' S, 162°59' E, 300-350 m, 1 lv (paratype).
ciialcal 2: stn DW 72, 24°54' S, 168°22' E, 527 m, 2 dd (paratypes).
biocal: stn DW 38, 23°00' S, 167°15' E, 360 m, 1 dd (paratype).

smib 3; stn DW 01, 24°56' S, 168°22' E, 500 m, 1 dd (paratype). — Stn DW 05, 24°55' S, 168°22' E,
502-512 m, 1 dd (paratype).

Source : MNHN , Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

213

Loyalty Islands, musorstom 6: stn DW 478, 21°09' S, 167°54' E, 400 m, 1 dd (paratype).
479, 21°09' S, 167°55' E, 310 m, 3 dd ( paratypes: mnhn, ams, nmnz).

Stn DW

Distribution. Coral Sea, New Caledonia and Loyalty Islands, alive in 300-610 m

Description. Shell to 42 mm long, solid, shiny, slightly
curved, white with alternating yellow bands. Sculpture
consisting of 13-14 rounded primary ribs, secondary ribs
appearing below apex, fading by the anterior quarter.
Intercostal spaces smooth, except for growth lines and
occasional vestigial longitudinal lines. Oral aperture thin.

subcircular, slightly depressed on ventral side. Apex trunca¬
ted, round and wide in cross section, with a projecting pipe
fissured at dorsal side.

Measurements: holotype L 36.4, W 3.6. w 1.2, arc 1 5-
paratype: L 23.4, VV 3.1. w 1.4, arc 1. W/w ratio 2.2-3.0

Etymology. — From the Latin flavus (yellow)

Dentalium deforgesi sp. nov.

Figs 20, 28 d, 71 a-b

Type material. — Holotype and 6 paratypes, mnhn.

Type locality. — New Caledonia, Northern Norfolk Ridge, chalcal 2, stn DW 73
24°40' S, 168°38' E, 573 m.

Material examined. — New Caledonia, chalcal 2: stn DW 73, 24°40' S, 168°38' E, 573 m,
3 lv (holotype and 2 paratypes ).

lagon: stn 1146, 19°08' S, 163°31'E, 185 m, 4 lv (paratypes).

Fig. 20. — Distribution of Dentalium deforgesi.

Distribution. New Caledonia, alive in

Description. - Shell to 42 mm long, solid, regularly
curved, shiny, cream-white. Sculpture of 11-13 primary ribs,
secondary ribs beginning at apex. Ribs decreasing in strength
shortly below apex on ventral side, later on dorsal side. Shell
smooth for first quarter of length except for growth lines,
circular in section, aperture thin, straight. Apex truncate with
terminal callus and a deep V-shaped notch on ventral side.

185-575 m.

Lumen protected by pseudo-pipe fissured on the ventral side;
lumen oval, laterally compressed.

Measurements: holotype L 36.3. W 4, w 1 .4, arc 2:
paratypes L 41.5. W 3.8, w 2.1. arc 1.8; L 37.8. W 3.8. w I.
arc 2; L 35.7, W 2.7, w 1.4, arc 2.2; L 27.1. W 2.8, w I. arc
1.6. W/w ratio 1.81-3.8.

Source .

214

VICTOR SCARABINO

Remarks. — This species is similar to D. obtusum Qi & Ma, 1989, an intertidal species from
China Seas, which differs in having 0-12 primary ribs, and lacking secondary ribs. In D. deforgesi the
ribs encircle the oral aperture, thus producing an irregular outline.

Etymology. Named for Dr Bertrand Richer de Forges (orstom, Noumea), organizer of
many dredging expeditions in New Caledonia and collector of numerous novelties in every group of
marine animals.

Fig. 21. — Dentalium tessellation sp. nov., holotype. — a. apex area. — c. lateral view (9.6 mm),
sculpture. Scale bars: 100 pm (a), 10 pm (b. d. e).

b. d-e, details of the

Dentalium tessellatum sp. nov.

Figs 21 a-e, 22

Type material. — Holotype and 3 paratypes dd, mniin.

Type locality. — Philippines, musorstom 3, stn DR 140, H°43' N, 122°34' E, 93-99 m.
Material examined. — Only known from the type material.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

215

The Philippines. Only known from the type locality, in 93-99 m, not

Distribution.
recorded alive.

Description. - Shell to 16 mm long, solid, opaque,
cream, slightly curved, with 8 high, rounded primary ribs,
lacking secondary ribs. At SEM magnification the entire
surface has a mosaic sculpture which appears finely reticula¬
ted under an optical microscope. Apex simple or with apical

callus, lumen circular.

Measurements: holotype L 9.6, W 1.5, w 0.5, a 0.6;
paratypes L 15.1, W 1.6. w 0.6, a 0.6: L 10.1, W 1.6, w 0.5.
a 0.4. W/w ratio 2.67-3.2.

Remarks. The absence of secondary ribs and the fine sculpture characterize this new

species.

Etymology. — From the Latin tessellatus (mosaic).

Other Indo-Pacific species of Dentalium cited in the literature

Dentalium aciculum Gould, 1859: 165. Hong Kong. Holotype usnm 24149.

Dentalium adenense Ludbrook, 1954: 97, fig. 2. Gulf of Aden, “John Murray ”, stn 28, 12°00' N,
50°38' E, 201 m. Holotype bmnh 1952.3.25.124.

Dentalium huccinulum Gould. 1859: 166. Kagoshima, Japan. Holotype usnm 24160.

Dentalium cancellatum Sowerby, 1860: 101, pi. 224 (Dentalium 2), fig. 36. China. Type material
apparently missing.

Dentalium cheverti Pilsbry & Sharp. 1897: 9. Evans Bay, Cape York, N. Australia, 11 m. ansp.

Dentalium eookei Pilsbry & Sharp, 1897: 29. Gulf of Suez. ansp.

Dentalium decemcostatum Brazier, 1877: 55. Katow, New Guinea. 2 syntypes dd ams A90.

Dentalium duodecemcostatum Brazier, 1877: 56. New Guinea. Holotype ams A91.

Dentalium laseroni Colman, 1958: 143, fig. 7. Broken Bay. New South Wales. Holotype ams C62221.

Dentalium lessoni Deshayes, 1825: 357, pi. 2, fig. 13. New Guinea. Lectotype (here designated)
20.3 mm mni-in.

Dentalium letsonae Pilsbry & Sharp. 1897: 4, pi. 1. fig. 13; pi. 5, figs 66-68. Island of Bohol,
Philippines, ansp.

Dentalium nannarense Winckworth, 1927: 167. fig. 4. Nannar Island, Sri Lanka, 6 m. 7 syntypes lv
BMNH 1952.3.21.6-12, and nmw ( fide Oliver, 1984).

Source : MNHN. Paris

216

VICTOR SCARABINO

Dentalium obtusum Qi & Ma, 1989: 69. Zhejiang Province, China.

Dentalium regulare Smith, 1903: 393, pi. 15, fig. 2. Port Shepstone, South Africa. 6 syntypes dd bmnh
1903. 9.9. 24-24a and 1904.7.26.29-32.

Dentalium robustum Brazier, 1877: 56. Katow, New Guinea. Syntype, labelled “lectotype”, but
designation apparently never published, ams A95.

Dentalium tignum Colman, 1958: 141, fig. 1. 35 miles East of Sydney, New South Wales, 1463 m.
Holotype ams C24485.

Dentalium tomlini Melvill, 1918: 155, pi. 5, fig. 31. Karachi, Pakistan. 5 syntypes dd bmnh
1921.1.28.36-40.

Dentalium woalacottae Colman, 1958: 142, fig. 4. Middle Harbour, Sydney, New South Wales.
Holotype ams C21230.

Genus Paradentalium Cotton & Godfrey, 1933
Type species (OD): D. intercalatum Gould, 1859.

Diagnosis. — Shell medium to large, moderately curved, solid, shiny, polished; translucent,
white or yellow. Longitudinally sculptured with 6 primary ribs, the most prominent dorsal, 1 ventral
and 4 latero-ventral. Secondary ribs present, variable number. Ribs round, flat or angled in cross
section, intercostal spaces generally smooth and straigth on dorsal side and convex on ventral side.
Apex simple, truncate, lumen circular, frequently with short terminal pipe. Section hexagonal at apex,
subcircular at mouth. Oral aperture generally thin and translucent in fresh specimens.

Radula similar to Dentalium.

Distribution. — Recent, worldwide, temperate and tropical regions. Sublittoral to bathyal.

Paradentalium pseudosexagonum (Deshayes. 1825)

Figs 23, 28 f

Dentalium pseudosexagonum Deshayes, 1825: 358, pi. 2, figs 14-16.

Olher references:

Dentalium pseudosexagonum - Sowerby, I860: 103. pi. 224 ( Dentalium 2), fig. 34; 1873: pi. 4, fig. 23. - Brazier, 1877: 56.

Pilsbry & Sharp. 1897: 23. pi. 4, figs 47-48. — Melvill & Standen, 1899: 181. — Boissevain, 1906: 14 pi 1 fig 10
— Melvill. 1909: 120. — Ludbrook, 1954: 97, fig. 3.

Dentalium (Paradentalium) pseudohexagonum (sic) - Habe & Kosuge, 1964: 1.

Type material. — 6 syntypes dd, mnhn.

Type locality. — “Unknown” (Deshayes 1825).

Material examined. — The type material.

Indonesia, corindon: stn B 210, 00°13'S, 117°53'E, 338 m, 1 dd.

“Snellius” II: stn 4.181, 06°21' S, 120°26' E, 34 m, 1 dd (rmnh).

Philippines. Philippines, Coll. Jousseaume, 6 dd (mnhn).

China. Hong Kong, Coll. Jousseaume, 1 dd (mnhn).

Distribution. — China Sea, the Philippines to Indonesia, Northern Australia, Seychelles and
the Gulf of Aden. Probably a shallow-water species, shells down to 499 m (Ludbrook, 1954).

Source : MNHN , Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

217

Fig. 23. — Distribution of Paradenialium pseudosexagonum.

Paradentalium intercalation (Gould, 1859)
Figs 24, 28 g

Denialium intercalation Gould. 1859: 106.

Other references:

Denialium intercalalum - Sowerby, 1873: pi. 7, fig. 45. — Boissevain, 1906: 22, pi. I, fig. 9. — Pilsbry & Sharp. 1897: 23,
pi. 11, figs 88-89. — Habe, 1977: 330. — Johnson, 1964: 93.

Denialium (Paradentalium) intercalalum - Habe & Kosuge. 1964: 1.

Type material. — Holotype usnm 24183.

Type locality. — China Seas.

Material examined. — The type material.

Philippines, musorstom 3: stn DR 137, 12°03'N, 122°06' E, 56 m, 1 dd.

Distribution. — China Sea, now extended to the Philippines; no living records.

Source

218

VICTOR SCARABINO

Paradentalium hexagonum (Gould. 1859)
Figs 25. 28 h

Demalium hexagonum Gould, 1859: 166.

Synonyms:

Demalium sexcostalum Sowerby, I860: 103, pi. 223 (Demalium 1), fig. 44.

Demalium minus Boissevain, 1906: 14. pi. 6, fig. 3 (Syn. nov.).

Other references:

Demalium hexagonum - Sowerby, 1860: 103, pi. 223 (Demalium 1), fig. 10; 1873: pi. 2. fig. 6. Clessin. 1896: 14 pi 24
fig. 2. Pilsbry & Sharp, 1897: 18. pi. 2, figs 20-21, 23-24. — Boissevain, 1906: 12. pi. 1. fie. 14. pi. 6. fie. I. Hirase,
1931: 133. pi. 3, fig. 2. Johnson. 1964: 88. pi. 22, fig. 4.

Demalium hexagonum var. sexcostalum - Pilsbry & Sharp, 1897: 19. pi. 2, figs 27-28. Boissevain, 1906: 13. pi. 6,
fig- 2.

Paradentalium hexagonum sexcostalum — Chistikov. 1979b: 109.

Demalium ( Paradentalium ) hexagonum - Kira, 1955: 80. pi. 40, fig. 7.

Dent ahum sexcostalum — Smith, 1875: 25.

Paradentalium minus - Habe & Kosuge, 1964: 2. Chistikov, 1979b: 109.

Type material. — D. hexagonum: holotype usnm 2053. — D. sexcostalum: not located. — D.
minus: lectotype (here designated) zma 3.06.008, paralectotypes zma 3.06.009.

Type locality. — D. hexagonum: Hong Kong, China. — D. sexcostalum: China. - D. minus:
Indonesia, Java Sea, " Siboga ”, stn 319, 06° 17- S, 1 14°37' E, 82 m.

Material examined. — The type material of D. hexagonum and D. minus.

Indonesia, corindon: stn B 202, 01°10' S, 1 1 7°06' E. 21 m, I lv. Stn CH 203. 0TW S, 1 17°08' E
25 m. 1 dd. — Stn CH 204, 01°09' S. 1 17° 1 9' E, 49 m. 2 dd.

Fig. 25. Distribution of Paradentalium hexagonum.

Distribution. — China Sea to Indonesia; alive in 15-29 m (Chistikov, 1979b and present

paper).

Remarks. Nomura (1938) synonymized P. hexagonum with Demalium octangulatum based
on rib count in 1500 specimens of D. octangulatum , of which 33 shells were found to have 6 ribs. The

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

219

material identifed by us as P. hexagonum presents 6 ribs, no secondary rib, and intercostal spaces are
smooth or with traces of longitudinal threads. D. octangulatum has secondary ribs that, although they
never reach the size of the primary ones, obscure the octangular section at the mouth. Variation in
the number ol ribs is observed in several species and I consider the presence of six ribs in occasional
specimens of D. octangulatum insufficient to support the synonymy established by Nomura. In
specimens of P. hexagonum 1 have studied, characters were constant.

Paradentalium natalensis (Barnard, 1963)

Figs 26, 28 i

Dentalium natalensis Barnard, 1963b: 350, fig. 30 e; 1974: 742.

Type material. 49 syntypes ( fide Barnard) sam A9364. 6 syntypes dd bmnh 1964.2.57.

Type locality. South Africa, off Cape Natal, Durban. 85 fms [156 m].

Material examined. - The type material in bmnh.

West Indian Ocean. Madagascar, Nosy Be Island, 50 m, Plante coll., 2 lv, 4 dd (bmnh).

South Africa. "Meiring Naude stn SM 94, 28° 16' S, 32°29' E, 670 m, 1 dd (sam).

Fig. 26. — Distribution of Paradentalium natalensis.

Distribution. South Africa, from off East London to Madagascar. Alive in 50 m. shells
from 70 to 670 m.

Paradentalium rudoi sp. nov.

Figs 27, 28 j, 1

Type material. Holotype mnhn. Paratypes: 9 mniin. 1 nmp.

Type locality. Western Indian Ocean. NE Madagascar, 12°39' S. 48°17'E, 240 m.

Source :

220

VICTOR SCARABINO

Material examined. — West Indian Ocean. NW Madagascar, “ Vauban 12°39' S,
48° 17' E. 240 rn, A. Crosnier coll. 1974, 13 dd (holotype and 6 paratypes mnhn, I paratype nmp).
benthedi: stn DS 72, 12°3 1 ' S, 45°02' E, 300-350 m, 2 lv, 1 dd ( paratypes).

Fig. 27. Distribution of Paradentalium rudoi.

Distribution. Off North Madagascar, alive in 300-350 m.

Description. Shell to 60 tnm long, solid, polished,
yellow, regularly curved. Six primary angle-edged ribs distri¬
buted one on the dorsal side, one on each side and 3 on the
ventral side. Secondary ribs appear at the posterior third of
the shell, fading at the aperture. Intercostal spaces with fine
longitudinal and transverse lines, which give the apical area

a reticulate appearance. Apex simple, lumen slightly irregu¬
larly compressed dorsoventrally.

Measurements: holotype L 52,8, W 3.8, w 1.2, arc 2.8-
paratypes L 60.2, W 3.9. w 1.5, arc 4.2; L 51. W 3.5, w 1. arc
3.5; L 47.5, W 3.4, w 1. arc 2.1; L 45.2. W 3, w 0.9, arc 2; L
50.9. W 3.5, w 1.1, arc 2.1. W/w ratio 2.6-3. 5.

Remarks. — P. pseudosexagonum is a related species from which P. rudoi differs in having
sculptured intercostal spaces, a longer and more slender shell with differently placed ribs and different
brightness and color.

Etymology. — Named after Rudo von Cosel, mnhn, who has contributed to our knowledge
of the Madagascar fauna by collecting remarkable and spectacular mollusc species during a survey
of shrimp resources conducted in 1989.

FlG- 28,n DuTn,Z sp; noV" ,hol°type> shell (26.4 mm), apex and apical section. - b. Den, ahum crosnieri

h°lolype- shell (30 7 mm), apical section and detail of the sculpture. c, Denialiwn flavum sp. nov.. holotype,
shell (36.4 mm), apex, apical and oral sections, detail of the sculpture. d, Dentalium deforgesi sp. nov., holotype, shell
L6J mm) apex and apical section. — e Dentalium flavum sp, nov., extremely damaged and repaired shell (33 mm).

t, Paradentalium pseudosexagonum, she 11 (38 mm), apex and apical section, corindon: stn B 210. g. Paradentalium
mtercalatum, shell (20 mm), apex, apical and oral sections, detail of the sculpture, musorstom 3: stn DR 137 h
Paradentalium hexagonum, shell (35 mm), apical and oral sections, corindon: stn CH 203. i. Paradentalium
Shell — ,mr)’ilpex1and, apical sf tl®n- de,ai1 of the sculpture. j. Paradentalium rudoi sp. nov., holotype.
IP. rS srnov ) &ale lines! lOOpm.' " '*** ^ ^ '• Paradentalium type radula

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

221

Source . MNHN, Paris

222

VICTOR SCARABINO

Other Indo-Pacific species of Paradentalium cited in the literature

Paradentalium angustistriatum Chistikov, 1979b: 110, fig. 2. Tonking Bay, Viet Nam, 47 m. zin.
Paradentaiium gradile Chistikov, 1979b: 109, fig. 1. Tonking Bay, Viet Nam, 60 m. zin.
Paradentalium katowense (Brazier, 1877): 56. Katow, New Guinea, 15 m. Holotype ams A92.
Paradentaiium pistis (Winckworth, 1940b): 43. fig. 6. Madras, India. Holotype bmnh 1940.7.227.

Genus Tesseracme Pilsbry & Sharp, 1897
Type species (SD by Woodring, 1925): Dentalium quadrapicale Sowerby, 1860.

Diagnosis. Shell medium to large, moderately curved, solid, generally polished; white,
cream to orange-yellow at apical area. Longitudinally sculptured with 4 primary ribs, prominent at
the apex, simple or bifurcated, one ventral, one dorsal and two lateral. Secondary ribs present,
variable in number. Intercostal spaces straight or concave, smooth or with longitudinal striae. Apex
simple, truncate, lumen circular; frequently with short terminal pipe. Transverse section quadrangular
at apex, subquadrangular to subcircular at mouth; oral aperture generally thin and translucent in
fresh specimens. Radula similar to Dentalium.

Distribution. — Eocene-Recent, Pacific and Indian Oceans, absent in the Atlantic Ocean;
sublittoral to shelf in temperate and warm waters.

Tesseracme quadrapicalis (Hanley, 1860)
Figs 29, 33 a

Dentalium quadrapicale Hanley in Sowerby, 1860: 103. pi. 225 (Dentalium 3), fig. 61.

Synonyms:

Dentalium conspicuum Melvill, 1897: 21, pi. 7, fig. 28 (Syn. nov.).

Dentalium dipsycha Pilsbry & Sharp, 1897: 33. pi. 4, figs 57-60 (Syn. nov ).

Other references:

Dentalium quadrapicale - Sowerby, 1873: pi. 7. fig. 46. - Smith, 1896: 371. Pilsbry & Sharp, 1897: 34. pi. 4. fig. 50.

Boissevain, 1906: 42, pi. 1, fig. 13. Winckworth, 1940b: 25. Ahmed, 1975: 29, fig. 33.

Tesseracme quadrapicale — Habe & Kosuge, 1964: 5.

Dentalium ( Tesseracme ) quadrapicale - Plate, 1908a: 350, pi. 30, fig. 53. Ludbrook. 1954: 102, tig. 5.

Dentalium quadripicale (sic) - Clessin, 1896: 13, pi. 3, fig. 6.

Dentalium conspicuum - Pilsbry & Sharp. 1898: 248. pi. 33. fig. 60. Boissevain. 1906: 42, pi. 2. fig. 26. Dinamani,
1964: 1.

Dentalium dipsyclia - Boissevain, 1906: 42, pi. 2, figs 24-25.

Type material. — D. quadrapicale : holotype bmnh 1907.10.28.147, paratypes bmnh
1907.10.28.148-149. — D. conspicuum : 2 syntypes dd BMHN 1897.30.80-81. D. dipsycha : syntype
ansp.

Type locality. — D. quadrapicale : Cochin, Malabar, India. — D. conspicuum: Karachi.

D. dipsycha: unknown.

Material examined. — The type material.

West Indian Ocean. Malabar, 28 dd. “Cotes de Malabar”, 60 dd. Karachi, Coll. Jousseaume,
1 lv. - Karachi, Coll. Denis, 2 dd (all mnhn).

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

223

Fig. 29. Distribution of Tesseracme quadrapicalis.

Distribution. - Malaya (Habe & Kosuge, 1964), Indonesia (Boissevain, 1906), S of India
in 742 m (Smith. 1896), Gulf of Oman (Ludbrook, 1954). Living depth range unknown, apparently
a shallow water species, shells cited from 9 m to 742 m .

Remarks. — The syntypes of Dentulium conspicuum are gerontic specimens with the apex wide
by reabsorption, 8-ribbed in section but the 4 primary ribs clearly noticed indicate its generic position.
Tesseracme are variable in shell shape, curvature, width of the apex and starting point of the
secondary ribs. I have observed this in specimens of T. quadrapicalis and specially in a large number
of specimens of T. tetrapleura , and which allows me to ascertain the present synonymy.

Tesseracme dispar (Sowerby, 1860)

Figs 30, 33 b

Denialium dispar Sowerby. I860: 103, pi. 224 (Dentulium 2). fig. 37.

Other references:

Denialium dispar - SoWerby. 1873: pi. 4. fig. 25. Brazihr, 1877: 58. Clessin:, 1896 1 1. pi. 2, fig- 4. - Pilsbry & Sharp.
1897: 32, pi. 4, figs 52-56. — Boissevain. 1906: 39, pi. 2, figs 22-23.

Tesseracme dispar - Habe & Kosuge, 1964: 5.

Type material. - Four specimens from the Island of Samar. 4 fms [7 m] in bmnh, could
belong to the type series, but this is far from certain.

Type locality. Singapore and Samar, the Philippines.

Material examined. Indonesia, corindon: stn B 253. 00°54' S. 119°30'E, 17 m, 2 dd.
China. Hong Kong, Coll. Jousseaume, 1 dd (mnhn).

Distribution. Philippines and China to Indonesia and North Australia, 0-54 m (Habe &
Kosuge, 1964).

224

VICTOR SCARABINO

Tesseracme tetrapleura (Boissevain, 1906)
Figs 31, 33 c-d

Dentalium telrapleurum Boissevain, 1906: 41, pi. 6, fig. 37.

Other reference:

Dentalium telrapleurum - Habe & K.OSUGE, 1964: 5 (as a synonym of Tesseracme quadrapicale ) .

Type material. — Lectotype (here designated) zma 3.06.49, paralectotype zma 3.06.50.
Type locality. — “Siboga”, stn 4, 07°42' S, 114°13' E, 9 m, anchorage off Djankar, Java.
Material examined. The type material.

New Caledonia, lagon: stn 41, 22°19' S, 166°16' E, 28-46 m, 1 dd. — Stn 58, 22°09' S, 166°13' E,
22 m, 1 dd. — Stn 63, 22°26' S, 166°26' E, 20 m, 1 lv. — Stn 65, 22°29' S, 166°26' E, 24 m, 1 lv. -
Stn 80, 22°3T S. 166°28' E, 33 m, 5 lv, 12 dd. — Stn 83, 22°32' S, 166°30' E, 22 m. 2 lv, 1 dd. — Stn
170, 22°09' S, 1 66°07' E, 22 m, 2 dd. — Stn 171.22°11'S, 166°06' E, 32 m, 1 dd. — Stn 192, 22°0T S,
166°00' E, 18 m, 2 lv. — Stn 201, 22°00' S, 165°59' E, 17 m, 1 lv. — Stn 202, 21°59' S, 165°57' E,
13 m, 1 lv. — Stn 211, 21°55' S. 165°52'E, 12 m, 1 dd. — Stn 214, 21°55' S, 165°48' E, 12 m, 1 lv,
1 dd. — Stn 216, 21°53' S, 165°49' E, 14 m, 1 dd. — Stn 226, 22°38' S, 166°39' E, 28 m, 1 lv, 1 dd.

— Stn 239, 22°24' S, 166°58' E. 43 m, 2 dd. — Stn 244, 22°25' S, 167‘W E, 37 m, 1 lv, 1 dd.

Stn 290, 22°16' S, 166°32' E, 1 1 m, 1 dd. — Stn 31 1, 22°44' S, 166°47' E, 36 m, 2 lv, 2 dd. — Stn 348,
22°42' S, 166°55' E, 45 m, 1 dd. — Stn 353, 22°34' S, 167°01' E, 70 m, 1 dd. — Stn 360, 22°35' S,
167°03'E, 60 m, 1 lv, 1 dd. — Stn 470, 18°28'S, 163°09'E, 41 m, 1 dd. — Stn 473, 18°24' S,
163°03' E, 50 m. 2 dd. — Stn 517, 19°09' S, 163°35' E, 42 m, 2 dd. — Stn 541, 19°06' S, 163°13' E,
45 m, 2 lv. 3 dd. — Stn 601, 22° 18' S, 167°03' E, 47-48 m, 1 lv, 2 dd. — Stn 602, 22° 16' S, 167°03' E,
43-48 m, 4 dd. — Stn 607, 22° 12' S. 167°03' E, 48-54 m, 2 lv. — Stn 619, 22°03' S, 166°54' E, 27-42
m, 1 dd. — Stn 632, 21°57' S, 166°50' E, 44-45 m, 2 lv, 4 dd. Stn 633, 2I°56' S, 166°48' E, 50 m,
1 dd. — Stn 667, 21°42' S, 166°28' E, 33-37 m, 2 lv, 2 dd. — Stn 676, 21°35' S, 166°23' E, 41 m.
1 dd. — Stn 682. 21°34' S. 166°19' E. 36-37 m. 2 lv. — Stn 687, 21°33' S, 166°17' E, 37-40 m, 1 lv.

— Stn 688, 21°3T S, 166°15' E, 36-40 m, 1 lv. — Stn 696, 21°29' S, 166°12' E, 41-57 m, 1 lv, 4 dd.

— Stn 697, 21°28' S, 166°10' E. 35-36 m, 1 dd. — Stn 701, 21°28' S, 166°07' E, 36-39 m, 1 lv, 1 dd.

— Stn 702, 21°27' S. 166°08' E, 37 m, .1 lv. — Stn 703, 21°25' S, 166°07' E, 38-40 m, 5 lv, 1 dd. -

Stn 704, 21°27' S, 166°06' E, 46-58 m, 1 lv. — Stn 707, 21°25' S, 166°04' E, 24-38 m, 1 lv, 1 dd.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

225

Stn 716, 2 1 °22' S, 165°59' E, 30 m, 1 lv. — Stn 726, 21°20' S, 165°55' E, 50-51 m, 1 dd. — Stn 729
21°19'S, 165°54'E, 42-45 m, 4 lv, 4 dd. — Stn 730, 21°07' S, 165°55' E, 40-43 m, 1 lv 3 dd —
Stn 747, 21°15' S, 165°5I' E, 31-34 m, 1 lv. — Stn 749, 21°18' S, 165°18' E, 49 m, 1 dd — Stn 754
21°13' S, 1 65°49' E, 36 m, 1 lv, 1 dd. — Stn 762, 21°12' S, 165°46' E, 43 m, 1 dd. - Stn 772, 21°08' s’
165°41' E, 30 m, 1 dd. — Stn 781, 21°05' S, 165°38' E, 36 m, 3 dd. — Stn 787, 21°04' S, 165°36' E
39 m, 1 lv. — Stn 801, 21°02' S, 165°29' E, 29 m, 2 lv, 1 dd. — Stn 822, 20°51' S. I65°21' E, 33 m
1 dd. — Stn 856, 20°37' S, 165°1 1' E, 30 m, 2 lv. — Stn 865, 20°39' S. 165°04' E, 24 m 1 lv —
Stn 866. 20°38' S, 165°03' E, 26 m, 1 dd. — Stn 867, 20°39' S, 165°01' E, 25 m. 1 dd — Stn 898
20° 14' S, 1 67°27' E, 22 m, 5 dd. - Stn 913, 20°58' S, 164°32' E, 10-13 m, 1 dd. — Stn 916 20°56' s’
1 64°28' E, 13 m, 1 lv, 2 dd. — Stn 931, 20°45' S, 164° 17' E, 28-29 m, 1 lv, 1 dd. — Stn 932 20°46' s'
164° 17' E, 23 m, 3 lv, l dd. — Stn 972, 20°25' S, 163°58'E, 27 m, 1 dd. — Stn 995 20°15'S
163°55' E, 35-36 m, 1 lv, 2 dd. — Stn 1007, 20°12' S, 163°52' E, 23-24 m, 1 lv. — Stn 1015 90°10' S
163°52' E, 25 m, 1 lv. — Stn 1025, 20°07' S, 163°49' E, 25-28 m, 9 lv, 9 dd. — Stn 1063, 20°03' s’
1 63°47' E. 31 m, 1 dd. — Stn 1126, 19°33' S, 163°46' E, 41 m, 1 dd. — Stn 1168, 19°16' S, 163°09' E,
50 m, 2 dd. — Stn 1182, 19°27' S, 163° 16' E, 48 m, 1 lv. — Stn 1205, 19°42'S, 163°26'E, 38 m,
1 lv.

Loyalty Islands, musorstom 6: stn DW 435, 20°21' S. 166°08' E, 32 m, 1 dd.

Indonesia, corindon: stn B 251, 00°54' S, 1 19°30' E, 95 in, 1 dd.

“Snellius" II: stn 4.025, 05°57' S, 123°49' E, 250-300 m, 2 dd (rmnh).

Fig. 31. — Distribution of Tesseracme letrapleura.

Distribution. — Indonesia, now extended to New Caledonia. Alive in 10-70 m (present

paper).

Genus Eudehtauvm Cotton & Godfrey, 1933

Type species (OD): Denralium quadricostatum Brazier, 1877.

Diagnosis. — Shell small to medium with four ribs, intercostal spaces smooth and straight.
Apical callus prominent, lumen circular, pipe present. Ribs simple, high, with rounded section,
irregular due to growth lines.

Radula unknown, assumed to be similar to Tesseracme.

Distribution. — Recent. Eastern Australian waters. New Guinea, sublittoral-shelf.

226

VICTOR SCARABINO

Eudentalium quadric out at um (Brazier, 1877)
Figs 32, 33 e

Denlalium quadricostatum Brazier, 1877: 58.

Other references:

Denlalium quadricostatum — Pilsbry & Shari1, 1897: 33. — Cotton & Godfrey, 1933: 140.

Type material. — 5 syntypes dd, ams A94.

Type locality. — Northeastern Australia, Princess Charlotte Bay, 13 fms [24 m].
Material examined. — The syntypes.

Northeastern Australia. Off Pioneer Bay, Orpheus Island, 26 m, Reid coll., 1 dd (bmnh).

Distribution. — East Australia, Papua New Guinea (Brazier, 1877), shells in 15-24 m.

Genus Antalis H. & A. Adams, 1854

Type species (SD by Pilsbry & Sharp, 1897): Denlalium entalis Linne, 1758. Recent, Europe.

Diagnosis. — Shell medium to large, moderately to well curved; general aspect variable, solid
to thin and fragile, generally polished; white, cream, and/or orange-yellow at the apex. Longitudinally
sculptured, primary ribs variable in number, and strength; secondary ribs present, also variable in
number and can equal primary ribs in size at the oral area. Ribs generally round in section,
occasionally flat or angled; smooth or sculptured. Intercostal spaces concave or convex, smooth or
sculptured by longitudinal or transverse striae. Apex simple, truncate, with a V-shaped notch or
irregular slit on ventral side; lumen circular, commonly with short terminal pipe. Section “polygonal”
at apex, circular, subcircular dorsoventrally compressed, or subpolygonal at mouth; generally thin
and translucent in fresh specimens.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC 227

Fig. 33. - a. Tesseracme quadrapicalis, shell (42 mm), apical section, detail of the sculpture at the mouth, Karachi
mnhn. — b, Tesseracme dispar , shell (30 mm), apex, apical section, section of a rib, sculpture at the mouth, corindon:
stn B 253. c, Tesseracme leirapleura , shell (35 mm), apical sections, apex, detail of the sculpture. New Caledonia
lagon: stn 83. d, Tesseracme type radula (T. leirapleura). Scale line: 100 pm. — e, Eudentalium quadricostatum , shell
(22 mm), apex, apical and oral sections. Orpheus Isl. (bmnh).

Radula rachidian similar to Dentalium , anterior margin with granulae; laterals well armed, head area
granulose.

Distribution. — Triassic- Recent, worldwide. Sublittoral to abyssal.

Antalis longitrorsum (Reeve, 1842)
Figs 34, 45 a

Dentalium longitrorsum Reeve, 1842a: 197; 1842b: 6, pi. 130, fig. 6.

Synonym:

Dentalium lamarcki Chenu. 1843: 2, pi. 6, figs 15- 15a.

Other references:

Dentalium longitrorsum - Reeve. 1842b: 6, pi. 130. fig. 6. - Sowerby, 1860: 98. pi. 225 ( Dentalium 3). figs 59-60; 1873:
pi. 2, figs 9a-b. — Brazier. 1877: 59. — Watson, 1879: 515; 1886: 9. Cooke, 1885: 273. - Melville & Abercrombie. 1893:
41. Ci.essin, 1896: 23, pi. 1, fig, 2. - Pilsbry & Sharp, 1897: 111, pi. 20, figs 35-36. — Smith. 1903: 393; 1906b: 58. —
Boissevain, 1906: 52, pi. 2, figs 33-33a. Lamy, 1938: 88. — Moazzo, 1939: 222. — Dawidoff. 1952: 114. — Kuroda &
Habe, 1952: 36. — Dharma. 1992: 78, fig. 14.

Dentalium ( Laevidentalium) longitrorsum — Ludbrook, 1954: 104.

Laevidentalium longitrorsum - Matsukuma et al„ 1991: 183. Habe & Kosuge, 1964: 7.

Dentalium longirostrum - Paetel, 1873: 79. - Mastaller, 1978: 136.

Antalis longitrorsum - Kilburn & Rippey. 1982: 148, pi. 34, fig. 5.

Laevidentalium longitorsum (sic) - Higo & Goto, 1993: 687.

Source . MNHN. Paris

228

VICTOR SCARABINO

Type material. — D. longitrorsum : holotype bmnh 1887.3.4.21. D. lamarcki : probably in

MHNG.

Type locality. — D. longitrorsum: Zanzibar. — D. lamarcki: China Seas.

Material examined. — The holotype of D. longitrorsum.

West Indian Ocean. Madagascar, Tulear. Harbour, 0-3 m, 1 lv, 86 dd. — N harbour, 2-4 m, 5 dd.

Pointe Anosy, 0.5-2 m, 1 dd. — N of jetty, tidal flat, fine sand at low tide, 28 dd. — NW of jetty,
tidal flat, fine sand at low tide, 33 dd. — S of jetty, low tide, 16 dd (all R. von Cosel coll., mnhn).

Lagoon, 0-13 m, 6 dd. — Grand Recif, shore, 15 dd. — Foly, 7 m, 1 lv, 2 dd. — Nosy Be Island,
NW Madagascar, coll. Plante, 5 dd (all bmnh).

“ Meiring Naude stn SM 114, 29°1 1' S. 31°43' E, 40 m, 1 dd (sam).

Distribution. — China, the Philippines, New Guinea, Northern Australia, India, Red Sea,
Zanzibar and Natal, 65-68 m (Ludbrook, 1954); Madagascar, alive in 0-3 m (present paper).

Antalis porcatum (Gould, 1859)
Figs 35, 45 b

Deni alium porcatum Gould, 1859: 166.

Other references:

Demalium porcatum - Sowerby, 1873: pi. 7, fig. 47. Clessin, 1896: 20, pi. 6, fig. 3. — Pu.sbry & Sharp. 1897: 15,
pi. 6, fig. 80. — Boissevain, 1906: 15, pi. 1, fig. 15. — Habe & Kosuge, 1964: 2. — Johnson, 1964: 130, pi. 18, fig. 5.

Type material. — Holotype mcz 169304 ( fide Johnson, 1964), paratype usnm 24142.

Type locality. — Hong Kong Harbour, China.

Material examined. — The paratype.

West Indian Ocean, md 32 Reunion: stn DC 86, 20°59' S, 55° 15' E, 75-90 m, 2 lv, 8 dd.

Source : MNHN , Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

229

Distribution. — China, Karachi (Boissevain, 1906). Now extended to Reunion Island, alive
in 75-90 m.

Antalis weinkauffi (Dunker, 1877)
Figs 36, 45 c

Dentalium weinkauffi Dunker, 1877: 68; 1882: 153, pi. 5, fig. 1.

Synonym:

Antalis septentrionalis Kuroda & Habe in Habe. 1963' 262. pi. 38, fig. 34. textfigs 15-17 (Syn. nov.).

Other references:

Dentalium weinkauffi - Pilsbry, 1895: 116. — Pilsbry & Sharp, 1897: 40, pi. 2. fig. 26. Hirase. 1931: 135. pi. 3 fig 4
— Habe, 1957: 128, fig. 7. - Habe et a!.. 1986: 24.

Dentalium (Dentate) weinkauffi - Kira. 1955: 80. pi. 40, fig. 6.

Antalis weinkauffi - Habe, 1963: 261, pi. 38, fig. 30. textfig. 27; 1971: 488 (Japanese text): 307, (English), pi. 65, figs 12-13;
1977: 333, pi. 70. figs 1-4. — Habe & Kosuge. 1964: 5. — Springsteen & Leobrera. 1985: 287, pi. 82, fig. 7. — Qi & Ma.
1989: 118. — Hick) & Goto, 1993: 686.

Antalis weinkauffi weinkauffi — Habe & Kosuge, 1964: 5.

Type material. — D. weinkauffi. holotype zmb 101996 (fide Kilias. 1995). — A. septen¬
trionalis-. nsmt (fide Habe, 1963).

Type locality. — D. weinkauffi. Japan. — A. septentrionalis-. Japan, off Hachinoe, Honshu,

30 m.

Material examined. — Japan. Coll. Jousseaume, 2 dd. — Kadena, Okinawa, Coll. Staadt,
1 dd. — Kii, Japan, Coll. Staadt, 1 dd (all mnhn).

New Caledonia, musorstom 4: stn CP 189, 19°07' S, 163°29' E, 210 m, 1 dd.
lagon: stn 267, 22° 16' S, 166° 17' E, 26 m, 1 dd.

West Indian Ocean, md 32 Reunion: stn CP 41, 21°21' S, 55°27' E, 75 m, 4 dd. — Stn DC 43, 21°21' S,
55°27' E, 73-77 m. 5 dd. — Stn DC 86, 20°59' S, 55° 15' E, 75-90 m, 2 lv, 3 dd. — Stn DC 126,
20°52'S, 55°38'E, 110 m, 1 dd.

Source

230

VICTOR SCARABINO

Fig. 36. — Distribution of Antalis weinkauffi.

Distribution. — Japan, 30-500 m (Habe & Kosuge, 1964); East and South China Seas,
67-195 m (Qi & Ma, 1989); the Philippines (Springsteen & Leobrera, 1985). Now extended to New
Caledonia and Reunion Island, alive in 75-90 m.

Antalis usitatum (Smith, 1894)
Figs 37, 45 d

Denialium usitatum Smith, 1894: 168, pi. 4, fig. 16.

Other references:

Denialium usitatum - Smith, 1906a: 250. — Pilsbry & Sharp, 1897: 29, pi. 10, figs 68-69. — Winckworth, 1940a: 25.
Graptacme usitatum - Habe & Kosuge, 1964: 5.

Dentalium (Antalis) usitatum — Ludbrook, 1954: 99, fig. 4.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

231

Type material. — Holotype presumably in the Zoological
paratype bmnh 1894.9.1 1.8, Gulf of Bengal, 1094 m.

Survey of India (not seen);

Type locality. — Off Colombo, “ Investigator ”, 06°32' N, 79°37' E, 675 fms [1542 m].

Material examined. — Paratype in bmnh.

New Caledonia, biocal: stn CP 57, 23°44' S, 166H58'E, 1490-1620 m, 1 dd.

Indonesia, corindon: stn B 244, 00°57' S, 119°22'E, 970 m. 1 dd.

Philippines, musorstom 2: stn CP 55, 13°54'N, 119°58'E, 865 m, 1 lv.

1O0 Distribution. ~ Indonesia to the Bay of Bengal, the Maldives and the Gulf of Aden in
183-1542 m (Ludbrook, 1954), now extended to the Philippines and New Caledonia; living in 865 m
(present paper).

Antalis tibanum (Nomura, 1940)

Figs 38, 45 e

Dentalium ( Antalis ) tibanum Nomura, 1940: 101, pi. 1. figs 11-1 la.

Synonyms:

Dentalium entalis var. indicum Boissevain, 1906: 44, pi. 6, fig. 15 ( non Dentalium indicum Chenu, 1843).

Antalis hotssevainae Palmer, 1974b: 124. Norn. nov. pro Dentalium entalis var. indicum Boissevain. 1906 non Chenu, 1843 (Syn.

Other references:

Dentalium pretiosum - Hirase, 1931: 136, pi. 3, fig. 6.

Dentalium lubricatum - Hirase. 1931: 138, pi. 3 fig 9

Antalis tibanum - Habe ,1963: 262, pi. 38. fig. 14: 1971: 488 (Japanese text), 307 (English text), pi. 65. figs 6-7
Antalis indicum - Habe & Kosuge, 1964: 4.

Antalis indicum tibanum - Habe & Kosuge, 1964: 4.

& Gmob\m~6^b^ l9?7: 331 ~ SpRrNGSTEEN & Leobrera:. 1985: 287, pi. 82, fig. 10. - Habe et a!., 1986: 24. - Higo

Type material. — D. entalis var. indicum : lectotype (here designated) zma.

Type locality. — D. entalis var. indicum : Indonesia, “Siboga”, stn 159, 00°59’1 S, 129°48.8'
E, 41 1 m. — D. tibanum: Japan, off Boso Peninsula.

232

VICTOR SCARABINO

Material examined. — The type material of D. entalis var. indicum.

New Hebrides Arc. volsmar: stn DW30, 22° 17' S, 171°18' E, 450-550 m, 8 dd.

Indonesia. “ Snellius " II: stn 4.128, 08°18'S, 1 1 8° 1 6' E, 700-835 m, 1 lv, 2 dd (rmnh).

Distribution. — From Japan. 0-200 m (Habe & Kosuge, 1964) to Indonesia. Now extended
to New Hebrides Arc, alive in 700-835 m (present paper).

Antalis gardineri (Melvill, 1909)
Figs 39, 45 f

Dentalium gardineri Melvill, 1909: 120, pi. 5, fig. 9.

Other reference:

Dentalium (Antalis) gardineri — Ludbrook, 1954: 93.

Type material. — Holotype bmnh 1910.3.17.12.

Type locality. — Indian Ocean, Amirantes Is., 293-383 m.

Material examined. — The type material.

New Caledonia. “ Vauban ” 1978-79: stn 22. 22°59' S. 167°17' E, 540-545 m, 1 dd. — Stn 33, 22°33' S,
166°25' E, 290-350 m, 4 dd.

smib 2: stn DW 21, 22°40' S, 167°41' E, 460-500 m, 1 dd. — Stn DW 23, 22°31' S, 167°37' E, 410-
420 m, 1 dd.

2 dd. — Stn DW 75, 24°39' S, 168°40' E, 600 m, 1 dd.
biogeocal: stn DW 308, 20°40' S, 166°58' E, 510-590 m, 1 lv, 3 dd.

Loyalty Islands, musorstom 6: stn DW 410, 20°38' S, 167°07' E, 490 m, 1 lv, 2 dd.
New Hebrides Arc. gemini: stn DW 51, 20°59' S, 170°03' E, 450 m. 4 lv, 14 dd.
West Indian Ocean, benthedi: stn DR 28, 12°42' S, 45°20' E, 705 m, 1 dd.

Fig. 39. — Distribution of Antalis gardineri.

Distribution. — Seychelles. Now extended to NW Madagascar and New Caledonia, live
records in 450-590 m.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

233

Antalis perinvolutum (Ludbrook, 1954)

Figs 40, 45 g

Dentalium (Fissidentalium) perinvolutum Ludbrook, 1954: 101, fig. 7.

Other references:

Dentalium (Graptacme) usitatum - Boissevain, 1906: 44, pi. 4, figs 6-8.

Type material. — Holotype bmnh 1952.3.25.65.

Type locality. — ‘‘John Murray”, stn 185, 13°48' N, 49°16' E, 2000 m, Gulf of Aden.

Material examined. — The type material. Specimens identified by Boissevain (1906) as D.
usitatum in zma.

Indonesia, corindon: stn DR 231, 00'’05' N, 119°48' E, 1080 m, 1 lv, 1 dd.

Distribution. — Indonesia, Andaman Sea and Gulf of Aden, 918-2000 m. Alive in 918-
1080 m.

Antalis phaneum (Dali, 1895)

Figs 41, 45 i

Dentalium phaneum Dali, 1895: 686, pi. 26, fig. 1.

Other references:

Dentalium phaneum - Boss et al., 1968: 253. — Kay, 1979: 586, fig. 193F.

Type material. — Holotype usnm 107025, paratypes usnm 107026.

Type locality. — ‘‘Albatross”, stn 3476, near Sandwich [Hawaii] Is., 298 fms [545 m].
Material examined. — The type material.

234

VICTOR SCARABINO

New Caledonia, biogeocal: stn CP 232, 21°34' S, 166°27' E, 760-790 m, 23 lv.

biocal: stn DW 56, 23°35' S, 167° 12' E, 695-705 m, 1 lv. — Stn CP 75, 22° 19' S, 167°23' E, 825-860

m, 8 lv. — Stn DW 106, 21°36' S, 166°29' E, 625-650 m, 1 dd.

smib 2: stn DW 21, 22°40' S, 167°41' E, 460-500 m, 7 dd.

Loyalty Islands, musorstom 6: stn DW 488, 20°49' S. 167°06' E. 800 m, 1 dd.

French Polynesia. 1 1°22' S, 139°43' W, 614 m, J. Poupin-SMCB coll., 3 lv (mnhn).

Distribution. — Central and South Western Pacific from Hawaii to French Polynesia and
New Caledonia, live records in 614-860 m.

Antalis boucheti sp. nov.

Figs 42, 45 h, j, 73 g

Type material. — Holotype mnhn. Paratypes: 12 mnhn, 1 ams C201723, 1 nmnz M268959,
1 USNM.

Type locality. — Loyalty Islands, musorstom 6, stn 428, 20°24' S, 166° 13' E, 420 m.

Material examined. — New Caledonia, musorstom 4: stn CC 175, 18°59'S, 163°17'E,
355 m, 1 dd (paratype).

Loyalty Islands, musorstom 6: stn DW 397, 20°47' S, 167°05' E, 380 m, 4 dd (2 paratypes: 1 mnhn,
I usnm). — Stn DW 398, 20°47' S, 167°06' E, 370 m, 1 dd. — Stn DW 406, 20°41' S, 167°07' E,
373 m, 1 dd (paratype). — Stn DW 41 1, 20°40' S, 167°03' E, 424 m, 1 lv, 1 dd (paratypes). — Stn
DW 413, 20°40' S, 167°03' E, 463 m, 1 dd. — Stn DW 428, 20°24' S, 166° 13' E, 420 m, 3 lv (holotype
and paratypes), 3 dd (paratypes: 1 mnhn, 1 ams, 1 nmnz). — Stn DW 444, 20°54' S, 167°18'E,
300 m, 2 dd. — Stn DW 446, 20°54' S, 167° 19' E, 360 m, 1 lv, 2 dd (1 paratype lv, 1 paratype dd).
- Stn DW 451, 20°59' S, 167°25' E, 330 m, 5 dd (1 paratype). — Stn DW 481, 21°22' S, 167°50' E,
300 m, 1 lv (paratype), 4 dd.

Philippines, musorstom 3: stn DR 94, 13°47'N, 120°03' E, 842 m, 1 dd.

Distribution. — New Caledonia and the Philippines. Alive from 360 to 424 m, shells down
to 842 m.

Source . MNHN , Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

235

Fig. 42. — Distribution of Antalis boucheti.

Description. — Shell to 50 mm long, white to yellow,
solid, polished, very slightly curved, rapidly tapering to apex,
16 to 18 rounded primary angled ribs. Secondary ribs begin
near the apex, 58 in number at mouth. Transversal sculpture
of fine and dense subequal striae, round in section. Under
magnification, the longitudinal and transverse sculpture give
this species a reticulate appearance. Apex fine with a deep

fissure in fresh specimens, but this is usually missing due to
breakage or wear.

Measurements: holotype L 42, W 4.4, w 0.7, a 1; mean of
8 paratypes L 32.2. W 3.71, w 0.67, arc 0.66. W/w ratio
3. 9-6. 3.

Etymology. — Named after Dr Philippe Bouchet, who contributed to this monograph all the
way from collecting the material in the field to reviewing and editing my manuscript.

Antalis guillei sp. nov.

Figs 43, 45 k

Type material. — Holotype mnhn. Paratypes: 1 1 mnhn, 1 nmp.

Type locality. — West Indian Ocean, md 32 Reunion, stn DC 136, 20°46' S, 55"36' E,
915-922 m.

Material examined. — West Indian Ocean, md 32 Reunion: stn FA 40, 21°21' S, 55°27' E,
150 m, 4 dd. — Stn DR 47, 21°23' S, 55°37' E, 205-215 m, 7 dd. — Stn DC 56, 21°05' S, 55°12' E,
170-225 m, 8 dd (7 paratypes mnhn, 1 paratype nmp). — Stn DC 136, 20°46' S, 55°36' E, 915-922 m,
5 dd (holotype and 4 paratypes).

Distribution. — Reunion Island. Shells only in 150-915 m.

Description. — Shell to 9 mm long, well arched, translu¬
cent. Sculpture of 10 primary ribs. Secondary ones begin
equal in strength to. but less prominent than the primary ribs
near the anterior aperture where both are present. Conspi¬
cuous growth lines also present. Apex simple, straight.

lacking callus. Mouth thin, slightly depressed ventrally.

Measurements: holotype L 8.7. W 0.9, w 0.3, arc 0.8;
paratypes L 7.5, W 0.7, w 0.3, arc 0.7; L 8.6. W 0.9, w 0.4,
arc 0.8; L 9, W 0.9, w 0.4. arc 0.9; L 8.7, W 0.8, w 0.3, arc
0.6; L 8.5. W 0.8, w 0.3, arc 0.8. W/w ratio 2.3-3.

Remarks. — Despite the small size, specimens appear to be mature.

Etymology. — Named for Dr Alain Guille, now at Laboratoire Arago, Banyuls, formerly
curator of Echinoderms at mnhn and senior scientist of Marion-Dufresne cruise md 32.

Source : MNHN, Paris

236

VICTOR SCARABINO

Fig. 43. — Distribution of Antalis guillei.

Other Indo-Pacific species of Antalis cited in the literature

Antalis diarrhox (Watson, 1879): 511. " Challenger ", stn 169, 37°34' S, 178°22' E, 700 fms [1280 m],
New Zealand. Holotype bmnh 1887.2.9.65.

Antalis glaucarena (Dell, 1953): 48. Chatham Rise, New Zealand, 200-300 fms [366-548 m], nmnz.
Antalis inflexion (Sowerby, 1903): 224, pi. 5, fig. II. South Africa, Natal, Tugela River mouth, 14 fms
[26 m], Paratype sam A9364.

Antalis marukawai (Otuka, 1933): 159, textfig. la-f. Off Koshikizima, Satuma, Japan, 200 m. Central
Fisheries Experimental Station, Japan.

Antalis suteri (Emerson, 1954): 185, nom. nov. pro D. arenarium Suter, 1907. Stewart Island, New
Zealand, 18 fms [33 m]. New Zealand Geological Survey, Wellington (fide Dance, 1986);
paratype South Australian Museum D 16001 (fide Zeidler & Macphail, 1978).

Antalis tosaensis (Habe, 1963): 23, pi. 2, fig. 2. Tosa Bay, Shikoku, 200 m. nsmt.

Genus Plagioglypta Pilsbry in Pilsbry & Sharp, 1897

Type species (OD): Dentatium undulalum Munster, 1844. Carboniferous.

Diagnosis. — Shell medium to large, almost straight, solid, polished, shiny, white. Sculptured
at apex by close-set, fine, encircled wrinkles; prominent, oblique growth lines over the entire shell.
Apex simple or with a fiat V-shaped notch. Subcircular in section, slightly compressed dorsoventrally.
Radula rachidian similar to Dentalium, but with smooth anterior margin; lateral with large primary
cusp; marginal short, sinusoidal.

Distribution. — Ordovician (fide Emerson, 1962)-Recent. SW Pacific and Indian Ocean in
temperate and cold water, absent in the Atlantic Ocean; shelf-bathyal.

Plagioglypta pertracheata (Plate, 1908)
Figs 44, 45 1-m

Dentalium (Plagioglypta) pertracheatum Plate. 1908a: 357, pi. 30, figs 45-46.

Source .

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

237

Type material. — Lectotype, designated by Kilias (1995), zmb 61099a (not seen).

Type locality. — “Valdivia”, stn 185, 03°41' S, 100°59' E, 614 m, SW Sumatra.

Material examined. — New Caledonia, biocal: stn CP 26, 22°40' S, I66°27' E, 1618-1740 m.
1 lv, 6 dd.

biogeocal: stn CP 214, 22°43' S, 166°28' E, 1590-1665 m, 4 dd.

West Indian Ocean, benthedi: stn DS 42, 13°05' S, 45°08' E, 400-520 m, 1 lv. — Stn F 49, 12°55' S,
44°57' E, 300-450 m, 2 dd. — Stn F 61, 12°46' S, 44°58' E, 475-510 m, 2 lv, 5 dd. — Stn DS 64,
12°4T S, 44°57’ E, 770-860 m, 1 dd. — Stn F 68, 12°30' S, 45°02' E, 400-460 m, 1 dd. — Stn DS 71,
12°30' S, 45°02' E, 450 m, 1 lv, 4 dd.

md 32 Reunion: stn CP 60, 21°03' S, 55°10' E, 460-490 m, 1 lv. — Stn DC 58, 21°03' S, 55°10' E,
450 m, 4 lv, 6 dd. — Stn DS 178, 21°04' S, 55°10' E, 412-460 m, 3 lv, 7 dd.

Fig. 44. — Distribution of Plagioglypta periracheala.

Distribution. — W Indonesia, now extended to New Caledonia, Reunion Island and NW
Madagascar, live records in 460-1740 m.

Genus Striodentauum Habe, 1964a
Type species (OD): Dentalium rhabdolum Pilsbry, 1905. Recent.

Diagnosis. Shell medium to large, slightly curved to almost straight, solid, opaque, light
brown. Longitudinal sculpture of 7-8 primary ribs; secondary ribs present, variable in number. Rib
section angled, smooth or sculptured. Intercostal spaces concave, smooth or sculptured with
longitudinal striae. Apex simple, section starlike at apex, starlike to polygonal at mouth.

Radula rachidian with the granulose anterior border; lateral with prominent primary cusp, head
granulose; marginal sinusoidal.

Distribution. — Recent, Pacific and Indian Oceans, temperate and cold waters. Absent in the
Atlantic Ocean. Shelf to bathyal.

Source . MNHN, Paris

238

VICTOR SCARABINO

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

239

Striodentalium thetidis (Hedley, 1903)
Figs 46, 52 b

Dentalium ihelidis Hedley, 1903: 327, fig. 61.

Other references:

Dentalium ihelidis — Hedley, 1918: 112.

Entalinopsis (Entalinopsis) thetidis - Habe & Kosuge, 1964: 8.

Type material. — 2 syntypes dd ams Cl 62 12.

Type locality. — Off Port Kembla, New South Wales, Australia, 115-137 m.

Material examined. — The type material.

Philippines, musorstom 3: stn CP 143, 11°29'N, 1 24° 1 1 ' E, 205-214 m, 2 dd.

Distribution. — Southeastern Australia, now extended to the Philippines; shells in 115-

214 m.

Fig. 45. a, Antalis longitrorsum , shell (80 mm), apex and apical section, Tulear, (mnhn). — b, Antalis porcatum, shell
(18 mm), apex and apical section, md 32 Reunion: stn DC 86. — c, Antalis weinkauffi, shell (68 mm), apex, apical
section, detail of the sculpture, New Caledonia lagon: stn 267. — d. Antalis usitatum, shell (45.5 mm), apex, detail of
the sculpture, corindon: stn B 244. — e, Antalis tibanum, shell (21 mm), apex and apical section, volsmar: stn
DW 30. — f, Antalis gardineri, shell (63 mm), apex, apical and oral sections, Gemini: stn DW 51. — g, Antalis
perinvolutum, shell (68 mm), apex, and apical section, corindon: stn DR 231. h, Antalis type radula (A. boucheti sp.
nov.). — i, Antalis phaneum, shell (37 mm), apex, apical and oral sections, biogeocal: stn CP 232. — j, Antalis boucheti
sp. nov., holotype, shell (42 mm), apex and detail of the sculpture. k, Antalis guillei sp. nov., holotype, shell (8.7 mm),
apex, apical and oral sections, section of the ribs and detail of sculpture. — 1, Plagioglypta pertracheata , shell (60 mm),
apex, apical and oral sections, detail of the sculpture, benthedi: stn F 61. — m, Plagioglypta type radula (P.
pertracheata ). Scale lines: 100 pm (h, m).

240

VICTOR SCARABINO

Striodentalium rhabdotum (Pilsbry, 1905)

Figs 47, 52 a, d

DenlaUum rhabdotum Pilsbry, 1905: 116, pi. 5, figs 45-47.

Other references:

Dentalium (Antalis) rhabdotum — Kuroda & Kikuchi, 1933: 8, pi. 1, figs 1-2.

Dentalium (Dentate) rhabdotum — Habe, 1953: 294; 1957: 129, fig. 11.

Antalis rhabdotum - Habe, 1962: 106, pi. 47, fig. 12; 1963: 263, pi. 38, figs 17-18. — Habe & Kosuge, 1964: 5. — Okutani,
1964: 73.

Striodentalium rhabdotum - Okutani. 1966: 12. — Habe, 1964a: 22, pi. 2, figs 17-18; 1971: 489 (Japanese text). 307 (English
text), pi. 65. figs 4-5. — Habe, 1977: 334, pi. 72, fig. 5. — Habe et a!.. 1986: 24, pi. 1, figs 11-12. — Qi & Ma, 1989: 118,
figs lOa-b. - Higo & Goto. 1993: 687.

Type material. Holotype ansp 88319.

Type locality. — Heda, Izu, Japan, 306 m.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DW 341, 19°46' S, 158°43' E, 620-630 m, 3 dd.

Indonesia. ", Snellius ” II: stn 4.135, 06°29' S, 121 °09' E, 495 m, 2 lv, 3 dd. — Stn 4.267, 08°18'S,
1 18°21' E, 650 m, 1 dd.

Philippines, musorstom 1: stn CP 44, 13°47' N, 120°30' E, 592-610 m, 8 dd.
musorstom 2: stn CP 82, 13°46' N, 120°28' E, 550 m, 5 lv.

musorstom 3: stn CP 106, 13°47'N, 120°30' E, 640-668 m, 6 lv, 60 dd. — Stn CP 118, 11°58' N,
121°06' E, 448-466 m, 1 lv, 3 dd. — Stn CP 122, I2°20' N, 121°42' E, 219-220 m, 2 dd.

Distribution. — Japan, from 61 m to 1350 m (Habe, 1957 and Okutani, 1964); East and
South China Seas, 300-550 m (Qi & Ma, 1989). Now extended to the Philippines and New Caledonia.
Live records in 448-668 m.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

241

Striodentalium kanakorum sp. nov.

Figs 48, 52 c

Type material. - Holotypc and 3 paratypes mnhn.

Type locality. New Caledonia, Coral Sea, chalcal 2, stn DW74,24°40' S. 168°38' E, 650 m.

Material examined. New Caledonia, chalcal 2: stn DW 72, 24°55' S, 168°22' E, 527 m,
1 lv (paratype). — Stn DW 73, 24°40' S. 168°38' E, 573 m, 1 dd (paratype). Stn DW 74. 24°40' S,
168°38' E, 650 m, 1 lv (holotype).

MUSORSTOM 4: stn DW 197, 1 8°5 1 ' S. 163°2T E, 550 m. 1 dd (paratype).

Fig. 48. Distribulion of Striodentalium kanakorum.

Distribution. New Caledonia, alive in 527-650 m.

Description. Shell to 32 mm long, solid, regularly
curved, white, shiny. Seven primary high, angulate and
irregular ribs and one secondary rib at each intercostal
concave space. Apex truncate with important callus, wide on
dorsal side, lumen drop-like shape. Cross section laterally
compressed throughout, mouth thin.

Measurements: holotype L 21, W 2.8. w 1.4, arc 1.3;
paratvpes L 18.5. W 2.7. w 1.4, arc 1: L 27.6. W 3.1. w 1.4,
arc 1.4; L 32, W 3.1, w 1.5, arc 1.5. W/w ratio 2.2-2.

Remarks. — S. kanakorum is shorter, more curved and shiny than the other species of the
genus, except S. thetidis, but in the latter species the intercostal spaces are finely striated.

Etymology. - After Kanak, name of the people of New Caledonia and Vanuatu Islands.

Other Indo-Pacific species of Striodentalium cited in the literature
Striodentalium chinensis Qi & Ma, 1986; 69, fig. 1. East China Sea.

Striodentalium concretum (Colman, 1958); 141, fig. 2. E of Sydney. NSW, Australia, 548 in. AMS.
Striodentalium hosoi Habe, 1963: 263, fig. 46. Off Tosa Bay, Shikoku, Japan, nsmt.
Striodentalium polycostatum Qi & Ma, 1986: 69. East China Sea, 184 m.

Source . MNHN , Paris

242

VICTOR SCARABINO

Genus Graptacme Pilsbry & Sharp, 1897

Type species (SD by Woodring, 1925): Denialium eboreum Conrad, 1846. Recent, Tampa Bay, Florida, USA.

Diagnosis. — Shell medium to large, slightly to well curved, generally fragile, polished to
shiny, except for apical portion; translucent white and salmon near the apex in some species.
Sculptured by close, fine, longitudinal striae, prominent near apex; anterior half of shell usually
smooth. Apex simple, truncate with apical callus and lumen variable in shape, or with deep irregular
slit on dorsum or side in some species. Circular in section, oral aperture generally thin and
translucent.

Radula similar to Denialium.

Distribution. Paleocene- Recent, worldwide; sublittoral of temperate and tropical regions
to abyssal.

Graptacme lactea (Deshayes, 1825)

Figs 49, 52 e-f, 71 f, 73 i

Denialium lacteum Deshayes, 1825: 362, pi. 2, fig. 27.

Other references:

Denialium lacteum - Sowerby, I860: 98, pi. 225 (Denialium 3), fig. 48: 1873: pi. 6, fig. 37. — Pilsbry & Sharp. 1897: 99,
pi. 19, fig. 1. — Boissevain, 1906: 54, pi. 1, fig. 21, pi. 6, fig. 35.

Eboreidens lacteum - Chistikov, 1975: 19; 1979: 113, fig. 5.

Fig. 49. — Distribution of Graptacme lactea.

Type material. — Lectotype (here designated; L 29, W 2.8, w 0.9) and paralectotype, mnhn.
Type locality. — India.

Material examined. — The type material.

Chesterfield Islands, corail 2: stn DW 72, 19°15' S, 158°21' E, 32 m, 2 dd. — Stn DW 100, 19°06' S,
158°27' E, 40 m, 1 lv. — Stn DW 122, 19°28' S, 158°17' E, 32 m, 2 dd. — Stn DW 155, 19°49' S,

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

243

1 58°25' E, 42 m, 1 dd. — Stn DW 158, 19°46' S, 158°17' E, 28 m, 1 dd. — Stn DW 160, 19°46' S.
1 58°23' E, 35-41 m, 1 Iv.

Philippines. Coll. Jousseaume, 3 dd (mnhn).

West Indian Ocean. Seychelles, 2 dd (mnhn).

Distribution. — Philippines and China Seas to Indonesia and the Indian Ocean, now
extended to the Coral Sea, alive in 32 to 42 m.

Remarks. — The type specimens are very worn but show typical Graptacme sculpture, as
noted by Boissevain (1906: 54). Chistikov (1975) proposed the genus Eboreidens and the family
Eboreidentidae using G. lactea as type species, but giving a description not corresponding to the
characteristics of G. lactea. I have not seen Chistikov’s specimens, but they do not appear to be the
present species. Examination of the figure of the radula (Chistikov, 1979b: fig. 5) indicates that
Chistikov’s species is Calliodentalium crocinum. If this is correct, Eboreidens and Eboreidentidae are
synonyms of Calliodentalium and Calliodentaliidae respectively.

Graptacme acutissima (Watson, 1 879)
Figs 50, 52 g

Denlalium acutissimum Watson, 1879: 514; 1886: 8, pi. 1, fig. 8.

Other references:

Denlalium acutissimum — Pilsbry & Sharp, 1897: 94, pi. 20, fig. 26. — Boissevain, 1906: 45, pi. 2, fig. 39, pi. 5, figs 9-12.
D. (Graptacme) acutissimum - Plate, 1908a: 351.

Graptacme acutissimum - Habf. & Kosuge, 1964: 5. — Okutani, 1974: 34.

Graptacme acutissima - Higo & Goto, 1993: 687.

Type material. — Three syntypes bmnii 1887.2.9.31-33. The specimen bmnh 1887.2.9.31 is the
figured syntype and will be designated lectotype by Lamprell & Healy (in press). The paralectotype
bmnh 1887.2.9.33 belongs to a different species.

Type locality. — Putative lectotype and paralectotype from “ Challenger ” stn 218, 02°33' S.
144°04' E, 1070 fms [1956 m], N off Papua New Guinea. Non-conspecific paralectotype from stn 246.
36° 10' N, 178°00' E, East of Japan, 3758 m.

Material examined. — The type material.

New Caledonia, biocal: stn KG 03, 21°15'S, 166°39' E, 2340 m, 1 lv. — Stn DS 04, 21°16' S,
1 66°40' E, 2340 m, 2 lv, 6 dd. — Stn CP 05, 21°16'S, 166°44' E, 2340 m, 11 dd. — Stn CP 23,
22°46' S, 1 66°20' E, 2040 m, 3 dd. — Stn CP 72, 22°10' S, 167°33' E, 2100-21 10 m, 3 lv, 13 dd. —
Stn CP 74, 22° 14' S, 167°29' E, 1300-1475 m, 1 dd. — Stn DW 79, 20°40' S, 166°52' E, 1320-1380 m,
1 dd. Stn KG 89, 21°03'S, 166°56' E, 2070 m, 1 lv, 2 dd. — Stn KG 95, 21°22'S, 166°33'E,
2365 m, 1 dd.

biogeocal: stn CP 238, 21°28'S, 166°23'E. 1260-1300 m, 2 lv, 1 dd. — Stn CP 250, 21°25' S,
166°28' E, 2350 m. 2 lv, 3 dd. — Stn CP 260, 21°00' S, 166°58' E, 1820-1980 m. 1 lv, 8 dd. — Stn
CP 266, 21°05' S, 166°57' E, 1990-2100 m, 2 dd. — Stn KG 269, 21°02' S. 166°58' E, 1810 m, 1 lv,
1 dd. - Stn CP 273, 21°02' S, 166°57'E, 1920-2040 m, 1 dd. — Stn CP 321, 21°12'S, 167°00'E,
2190-2205 m, 1 dd. Stn CP 341, 21°30' S, 166°47' E, 2334 m, 1 dd.

Indonesia, corindon: stn CH 231, 00°05' N, 119° 48' E, 980-1080 m, 1 dd.

Philippines, musorstom 2: stn CP 55, 13°54'N, 119°58'E, 865 m, 1 lv, 1 dd.

West Indian Ocean, md 32 Reunion: stn DS 109, 20°52' S, 55°06' E, 1050-1240 m, 1 lv, 1 dd.

244

VICTOR SCARAB1NO

Distribution. — The Philippines, Indonesia, N of Papua New Guinea and East Africa
(Plate, 1908a). Now extended to New Caledonia and Reunion Island. Living records from 1050 to
2350 m, mostly below 1800 m, shells known from 865 m (present paper).

Graptacme africana (Sowerby, 1903)

Figs 51, 52 h

Dentalium africanum Sowerby, 1903: 224, pi. 5, fig. 10.

Other references:

Dentalium africanum - Smith, 1906b: 58. Barnard, 1963b: 351: 1974: 742.

Type material. — Holotype bmnh 1903.7.27.54, paratype sam A5491.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

245

Type locality. North of Untwalumi river, Natal, South Africa, 25 fms [45 mj.

Material examined. — The type material.

West Indian Ocean. Tulear, Madagascar, Thomassin coll., 1 dd (bmnli).

South Africa. “Meiring Naude": stn SM 16, 27°33' S, 32°35' E, 384 m, 1 dd.

Distribution. South Africa, now extended to Madagascar, shells from 45 to 384 m.

Fig. 52. a. Striodentalium rhabdotum, shell (91 mm), apex, apical and oral sections, musorstom 3: stn CP 106.

b. Striodentalium tethidis, shell (22 mm), apex and apical section, detail of the sculpture, mnhn. musorstom 3: stn
CP 143. c, Striodentalium kanakorum sp. nov.. holotype, shell (21 mm), apex, apical and oral sections. -
d, Striodentalium type radula (S. rhabdotum). e. Graptacme lactea. shell (23 mm), apex and apical section, corail
2: stn CP 160. f. Graptacme type radula (O', lactea). g, Graptacme acutissima. shell (47 mm), apex and apical
section, riocal: stn DW 79. h. Graptacme africana , shell (44 mm), detail of the sculpture. Tulear (bmnh). Scale lines:
100 pm (d. fi.

246

VICTOR SCARABINO

Other Indo-Pacific species of Graptacme cited in the literature

Graptacme acuticostata (Plate, 1908): 352, pi. 30, fig. 37. Off Dar-es-Salaam, Tanzania, "Valdivia",
stn 247, 05°56' S, 39'T E, 50 m. Holotype zmb 61082 ( fide Kilias, 1995).

Graptacme elpis (Winckworth, 1927): 168, pi. 14, figs 6-7. West side of Nannar, Ceylon, 5 m.
2 syntypes bmnh 1952.3.21.13-14.

Genus Fissidentalium Fischer, 1885

Type species (by monotypy): D. ergasticum Fischer, 1885 (= D. capillosum Jeffreys, 1877). Recent, Atlantic Ocean.
1900 m.

Diagnosis. — Shell generally very large, moderately curved, solid, opaque or polished, white,
cream or light brown, usually with dark-brown markings. Longitudinal sculpture faint with primary
ribs variable in number; secondary ribs present, originating near the apex, variable in number and in
some species reaching the strength of the primary ribs at the oral area. Rib section round, fiat or
angled, striated, cancellated or serrated. Intercostal spaces concave or convex, smooth or with
longitudinal or transverse striae, generally less prominent than ribs. Apex with a deep irregular slit
on ventral side, frequently broken or worn. Section polygonal at apex, circular or subcircular slightly
dorsoventrally compressed at oral aperture.

Radula : rachidian strong, well curved with barely granulose anterior margin; lateral strong with short
but strong cusps or irregular grooves, anterior part of head usually granulose; marginal large and
sinusoidal.

Distribution. — Cretaceous-Recent, worldwide, shelf to abyssal.

Fissidentalium profundorum (Smith. 1894)

Figs 53, 62 f, 1, 70 a

Dentalium profundorum Smith, 1894: 167, pi. 4, fig. 18.

Other references:

Dentalium profundorum - Pilsbry & Sharp, 1897: 79, pi. 6, fig. 82. — Smith, 1906a: 249. — BoissEVArN, 1906: 37, pi. 4.
figs 14-16. - Winckworth, 1940a: 25.

Fissidentalium (Fissidentalium ) profundorum - Habe, 1964a: 13, pi. 1, fig. 15, pi. 5, fig. 59.

Type material. — Holotype presumably in the Zoological Survey of India; paratype bmnh
1894.9.11.11.

Type locality. — Off Sri Lanka, "Investigator", 06°32' N, 79°37' E, 675 fms [1235 mj.
Material examined. — The paratype.

New Caledonia, biocal: stn CP 57, 23°44' S, 166°58' E, 1490-1620 m, 1 dd. — Stn CP 62, 24°19' S,
167°49' E, 1395-1410 m, 1 dd.

biogeocal: stn CP 238, 21°28' S, 166°23'E, 1260-1300 m, 10 dd.

Indonesia, corindon: stn DR 229, 00°02' N, 119°50' E, 41 1-445 m, 1 lv, 1 dd.

West indian Ocean " Galathea ”: stn 217, 14°20' S, 45°09' E, 3560 m, 2 dd (zmc).

Distribution. — Indian Ocean, Indonesia, Japan, now extended to New Caledonia. Live
records in 441-445 m and shells down to 3560 m (present paper).

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

247

Fig. 53. — Distribution of Fissidentalium profundorum.

Fissidentalium magnificum (Smith, 1896)
Figs 54, 62 b, i, 78 a

Denialium magnificum Smith, 1896: 371.

Other references:

Denialium magnificum - Pilsbry & Sharp, 1897: 78, 251. — Alcock & Anderson, 1898: pi. 7, figs 5-5a. — Smith, 1904:
7; 1906a: 248; 1906b: pi. 7, figs 5-5a. — Boissevain, 1906: 37, pi. 2, figs 32-32a. — WiNCKWORTH,”l940a: 25.
Fissidentalium (Fissidentalium ) magnificum - Habe & Kosuge, 1964: 3.

Type material. — Holotype presumably in the Zoological Survey of India; paratype bmnh
1895.12.13.1.

Type locality. — Off Trincomalee, East coast of Sri Lanka, “Investigator", 08°40' N,
81°27' E, 637-800 fms [1165-1465 m].

Material examined. — Paratype in bmnh.

Chesterfield Islands, musorstom 5: stn CP 323, 21°19' S. 157°58' E, 970 m, 1 Iv, 3 dd. Stn CP 324,
21°15' S, 157°51' E. 970 m, 1 dd.

Loyalty Islands, musorstom 6: stn DW 426, 20°25' S, 166°23' E, 610 m, 2 lv. — Stn DW 489, 20°48' S,
167°06'E, 700 m, 1 Iv.

Indonesia, corindon: stn CH 214, 00°31'N, 117°50'E, 595 m, 1 dd. — Stn CH 240, 00°38' S,
1 19°34' E, 675 m, 2 lv.

" Snellius ” II: stn 4.127, 08°19'S, 118°18'E, 500-550 m, 1 dd. — Stn 4.128, 08°18'S, 118°16'E,

700-835 m, 2 lv, 6 dd. — Stn stn 4.130, 08°18' S, 118°18' E, 700-730 m, 1 lv, 5 dd. — Stn 4.267,

08° 18' S, 1 18°2T E, 650 m, 10 lv, 21 dd. (rmnh).

Philippines, musorstom 1: stn CP 47, 13°41'N, 120°30' E, 685-757 m, 2 lv, 7 dd.

musorstom 2: stn CP 24, 13°37' N, 120°42' E, 640-647 m, 1 dd. — Stn CP 25, 13”39' N, 120°43' E.

520-550 m, 1 lv. — Stn CP 55, 13°54' N, 119°58' E, 865 m, 8 lv, 8 dd.

musorstom 3: stn DR 95, 13°56' N, 119°59' E, 865 m, 13 dd. — Stn DR 102, 14°01' N, 120°18' E,

192 m, 1 dd. — Stn CP 106, 13°47'N, 120°30'E, 640-668 m, 2 dd. — Stn CP 133, 11°58'N,

121°52' E, 334-390 m, I dd.

West Indian Ocean. “Galathea": stn 314, 15°54' N, 90°17' E, 2600 m, 2 lv (zmc).
md 32 Reunion: stn CP103, 20°42' S, 54°57' E, 2950-2970 m, 2 dd.

248

VICTOR SCARABINO

iwr

I'Ki. 54. Distribution of Fissidentalium magnficum.

Distribution. — Indian Ocean, now extended to the
Caledonia, live records in 520-2600 m (present paper).

Philippines,

Indonesia and

New

Fissidentalium shoplandi (Jousseaume, 1894)

nr,, Fiss 55> 73 a-d

uentatium sliuplandi Jousseaume. 1894: 102.

Synonyms:

Dentalium transversostriatum Boissevain. 1906: 32. pi. 4 fig 23 (Svn nov i
Oenndnm, (, Ftss, dentalium ) chuni Plate, 1908a: 341. pi. 30. figs 1 -9 (Syn. nov.).

Other references:

- —

Dentalium (F.) shoplandi - Ludbrook, 1954: 99.

Dentalium (F.) chuni - Jaeckel, 1932: 305.

s ~ zma

MUSORSTOM 3: stn CP 123 12°H' N I21°4S' F 7no 7m , m’ 1 lv-

464-475 ns, 1 dd. - Stn 'CP ,28 Nn'50'N, ~ ^,7 dT' ‘ E-

C«led<S,Hr^11,^,19^r^,^r^,d the Ph4ippines, now extended to New

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

249

)

4

Jm

f 'l

O'

•

i»r

Fig. 55. Distribution of Fissidenlalium shoplandi.

Remarks. The type material of D. chuni was not examined but the detailed original
description allow's us to synonymize it.

Fissidenlalium cornubovis (Smith. 1906)
Figs 56. 62 h

Denlalium cornubovis Smith. 1906a: 249.

Other references:

Denlalium cornubovis — Annandalk & Stewart, 1909: pi. 23. figs 2-2a. Winckworth, 1940a: 25.

Denlalium ceras - Smith. 1906a: 248.

Type material. Lectotype (here desianated) the largest (63 mm) of the 3 syntypes bmnh
1906.10.12.142-144.

Type locality. Indian Ocean. 1154 fms [2109 m], "Investigator".

Material examined. — New Caledonia, biocal: stn CP 13, 20°20' S. 167° 19' E. 3690-3740 m,
1 Iv. 1 dd. Stn CP 17, 20°35' S, 167°25' E. 3680 m, 1 lv, 2 dd.

West Indian Ocean, benthedi: stn CH 87. 1 1°44' S. 47°35' E, 3716 m, 2 lv.
md 32 Reunion: stn DS 151, 20°51' S. 56°03' E. 3240-3300 m, 2 lv.

“Galathea": stn 234, 05°25' S. 47"09' E, 4830 m, 3 dd. — Stn 238, 03°23' S. 44°04' E, 3960 m, 2 lv.
1 dd (ZMC).

East Indian Ocean, safari 2: stn CP 10, 01°43'N, 87°08' E, 4350 m. 1 lv. Stn CP 12, 02°54' S.
89°43' E, 3344 m, I lv.

"Galathea": Stn 280, 01°66' N. 77°09' E. 4350 m, 1 lv, 8 dd. Stn 314, 15°54' N. 90°17' E, 2600 m,

I lv (zmc).

Distribution. — Widely distributed throughout the Indian Ocean, now' extended to New
Caledonia, recorded alive from 2600 to 4350 m.

Remarks. — The three syntypes of F. curnubovis are gerontic individuals, and their ribs fade
near the half of the shell. The specimens identified as Denlalium ceras by Smith, 1906 (S of Sri-Lanka,
bmnh 1906.10.12.1-2) are adults but not gerontic, and are ribbed throughout.

250

VICTOR SCARABINO

Fissidentalium ceras (Watson, 1879), from “Challenger” stations 246, 36° 10' N, 178°00' E, Mid
Pacific Ocean E of Japan, 2050 fms [3690 m] and 299, 33°31' S, 74°43' W, W of Valparaiso, 2160 fms
[3948 m], is here considered a senior synonym of Fissidentalium megathyris Dali, 1889, from
" Albatross ”, stn 2807, 00°24' S, 89°06' E, 812 fms [1485 m], near Galapagos Islands (holotype usnm
594262). This abyssal species is widely distributed throughout the East Pacific, from Chile to
Northern California and the Central Pacific.

Fissidert '•lium serrulatum (Smith, 1906)
Figs 57, 70 b

Denlalium serrulatum Smith, 1906a: 249.

Other reference:

Denlalium serrulatum — Winckworth, 1940a: 25.

Fig. 57. — Distribution of Fissidentalium serrulatum.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

251

Type material. — Holotype presumably in the Zoological Survey of India; paratype bmnh
1906.10.12.4.

Type locality. — Andaman Is., Gulf of Bengal, 60 fms [1 10 m].

Material examined. — Paratype in bmnh.

Indonesia, corindon: stn DR 216, 00°40' N, 117°5T E, 96 m, 2 dd.

Philippines, musorstom 2: stn CP 72, 14°00'N, 120°18'E, 182-197 m, 1 dd.

Distribution. — Gulf of Bengal, now extended to Indonesia and the Philippines. No records
of living material, shells from 96 to 182 m (present paper).

Fissidentalium malayanum (Boissevain, 1906)
Figs 58, 62 d, j

Denlalium malayanum Boissevain, 1906: 32, pi. 5, fig. 45.

Other reference:

Fissidenialium (F.) malayanum — Habe & Kosuge, 1964: 3.

Type material. — Lectotype (here designated) zma 3.06.033, paralectotypes zma 3.06.030-032.
Type locality. — “Siboga”, stn 300, 10°49' S, 123°23' E, 918 m, Timor Sea.

Material examined. — The type material.

New Caledonia, biocal: stn KG 03, 21°15'S, 166°39'E, 2340 m, 1 dd. — Stn CP 05, 21°16' S,
166°44' E, 2340 m, 2 lv, 30 dd. — Stn CP 72, 22°10' S, 167°33' E, 2100-21 10 m, 42 lv, 4 dd. — Stn
DS 98, 2 1° 1 4' S, 166°30' E, 2365-2470 m, 3 dd.

biogeocal: stn CP 250, 21°25' S, 166°28' E, 2350 m, 42 dd. — Stn CP 266, 21°05' S, 166°57T4' E,
1990-2100 m, 2 dd. — Stn CP 283, 21°22' S, 166°3T E, 2370-2375 m, 1 1 dd. — Stn DW 313, 20°59' S,
166°59' E, 1600-1640 m, 1 dd. — Stn CP 321, 21°12' S, 167°00' E, 2190-2205 m, 37 dd. — Stn CP

Source : MNHN, Paris

252

VICTOR SCARABINO

329. 21°09'S, 1 66°40' E, 2315-2310 m, 19 dd. — Stn CP 336, 21°12'S. 166°22' E. 2370-2380 m,
1 1 dd. Stn CP 341. 21°30' S. 166°47' E. 2334 m. 25 dd.

West Indian Ocean. "Galathea”: stn 190. 29°42' S. 39°19'E, 2760 m, 1 lv. 1 dd (zmc).

Distribution. Indonesia, now extended to southern Madagascar and New Caledonia.
Living from 2100 to 2760 m; shells from 918 m.

Fissidentalium viedani Kosuge, 1981
Figs 59, 62 c

Fissidentalium viedani Kosuge, 1981: 114, pi. 39, figs 5-7.

Other reference:

Fissidentalium (F.) viedani - Springsteen & Leobrera, 1985: 286. pi. 82, fig. 4.

Type material. Holotype, Institute of Malacology, Tokyo imt 81-36 (not seen).

Type locality. Bohol. Philippines.

Material examined. — Philippines, musorstom 3: stn CP 118. II°58'N, 121 "06' E, 448-
466 m, 1 dd.

Fig. 59. Distribution of Fissidentalium viedani.

Distribution. — Only known from the Philippines, shell in 448-466 m.

Fissidentalium levii sp. nov.

Figs 60, 70 c

Type material. Holotype mnhn. Paratypes: 8 mnhn, 1 ams C201724. I nmnz M268957.
Type locality. — North of New Caledonia, musorstom 4. stn DW 160, I8°42' S, 163°13' E.

668 m.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACII IC

253

Material examined. — Chesterfield Islands. MUSORSTOM 5: stn DW 341. I9°46' S. 158°43' E.
620-630 m. 1 dd (paratype). Stn DC 358, 19°39'S, I58°47' E, 680-700 m, I dd (paratype).

New Caledonia, biocal: stn DW 33, 23° 10' S, 1 67° 10' E, 675-680 m, 1 lv (paratype). Stn DW 39,
22°55' S, 167°23' E, 650 m, I dd (paratype ams). Stn CP 40. 22°55' S, 167°24' E. 650 m. 3 dd.

Stn DW 46, 22°53' S, 167"17' E, 570-610 m, 1 dd. - Stn DW 48, 23°00' S, 167°29' E. 775 m, 2 dd.
MUSORSTOM 4: stn DW 160, 18°42' S, 163° 13' E. 668 m, 3 lv (holotvpe and 2 paratypes).

SMIB 2: stn DW 20, 22°44' S. 167°42' E, 415-470 m, 2 dd. — Stn DW 21, 22°40' S,' 167°41' E, 460-
500 m, 3 dd.

smib 3: stn DW 21, 22l>59' S, 167°19'E, 525 m, 5 dd (1 paratype).

“ Vauban " 1976: Pointe du Grand Recif Sud, 200 m, 1 dd (1 paratype).

" Vauban ” 1978-79: stn 14, 22°16'S, 167°17'E, 465-495 m, 1 dd. Stn 22. 22°59' S. 167°17'E.
540-545 m, 1 dd.

Loyalty Islands, musorstom 6: stn DW 396, 20°48' S, 167°01' E, 1400 m, 1 dd (paratype nmnz).
New Hebrides Arc. gemini: stn DW 55, 20°59' S. 170°02' E. 710 m, 1 dd (paratype).

Fig. 60. Distribution of Fissidenlalium levii.

Distribution. — New Caledonia, alive in 668-680 m, shells from 200 to 1400 m.

DrscRii'TiON. Shell to 70 mm long, strong, white, shiny. Measurements: holotype L 51.7. W 5.1. w 1.6. arc 1.5:
slightly curved, regularly tapering. Sculptured by numerous paratypes L 61.5, W 6.5, w 1. arc 2; L 50.7, W 5. w 1.2, arc

close riblets often fading near the oral aperture. Apex with a 1.5; L 44.9, W 5, w 1. arc 1.5., W/w ratio 3.18-6.5.
narrow, irregular slit in ventral side in well preserved
specimens, slightly compressed dorsally. Mouth fragile, trans¬
lucent. subcircular in section.

Remarks. — This species can be compared with some growth stages of Antalis gar diner i, but
the latter species has better defined and regularly spaced ribs and transverse sculpture.

Etymology. Named for Prof. Claude Levi, mnhn, senior scientist of the biocal Expedition,
and former head of Marine Invertebrates in mnhn.

254

VICTOR SCARABINO

Fissidentalium metivieri sp. nov.

Figs 61, 62 a, 70 d-e

Type material. — Holotype mnhn. Paratypes 10 mnhn, 1 nmp, 1 usnm, 2 bmnh 1994.4041.

Type locality. — West Indian Ocean, SW Madagascar, "Mascareignes III", stn 106,
22°24' S, 43°03' E, 600 in.

Material examined. — West Indian Ocean. SW Madagascar, " Mascareignes III"', stn 65,
22°26' S, 43°05' E, 520 m, 1 lv (paratype). — Stn 67, 22°26' S, 43°06' E, 530 m, 1 dd. — Stn 74,
22°26' S, 43°03' E, 540 m, 1 lv, 1 dd. — Stn 76, 22°22' S, 43°04' E, 530 m, 2 lv (1 paratype). — Stn
82, 22° 1 1' S, 43°03' E, 520 m, 1 lv. — Stn 84, 22°21' S, 43°04' E, 535 m, 4 lv (1 paratype). — Stn 88,
22° 13' S, 43°04' E, 525 m, 2 dd (2 paratypes). — Stn 91, 22°25' S, 43°04' E, 535 m, 1 lv. — Stn 95,
22° 12' S, 43°03' E, 590 m. 4 lv (1 paratype mnhn, 1 paratype nmp). — Stn 97, 22°24' S, 43"04' E.
600 m, 3 lv. — Stn 100, 22°23' S, 43°03' E, 600 m, 2 lv. — Stn 104, 22°22' S, 43°03' E, 610 in, 1 lv
(paratype usnm). — Stn 106, 22°24' S, 43°03' E, 600 m, 3 lv (holotype and 1 paratype). — Stn 111,
22°17' S, 43°02' E, 610 m, 2 lv. — Stn 113, 22°11' S, 43°02' E, 650 m, 1 dd. — Stn 122, 22°17' S,

43°03' E. 600 m, 1 lv, 4 dd. — Stn 126, 22°18' S, 43°02' E, 590 m, 10 lv (2 paratypes), 4 dd. — Stn

127, 22°21' S, 43°02' E, 610 m, 8 lv, 2 dd.

Madagasgar. “Vauban" Crosnier coll. 1973: stn CH 46, 15°19'S, 46°12' E, 400 m, 3 dd. — Stn
CH 48, 1>18'S, 46° 12' E, 480-510 m, 1 dd. — Stn CH 49, 15°18' S, 46° 10' E, 500-550 m, 2 lv,
10 dd (1 paratype). — Stn CH 50, 15°19' S, 46°12' E, 405 in, 1 lv, 1 dd. — Stn CH 88, 18°54' S,

43°55'E, 280-310 m, 1 dd. — Stn CH 91, 21°26' S, 43°15' E, 425-550 m, 1 dd. — Stn CH 102,

22°30' S, 42°59' E, 995-1020 m, 2 dd (paratypes bmni-i).

SW Madagascar, Mozambique Channel, alive in 310-650 m, shells to 1020 m.

Distribution.

Description. — Shell to 180 mm long, solid, opaque, light
brown. Anterior half of shell straight and regularly curved to
posterior end, rapidly tapering to apex. Eight primary ribs, of
which two latero-dorsal are larger, prominent to mouth.
Secondary ribs begin earlier on dorsal than on ventral side,
never reaching the strength of the primary ribs. Transverse

sculpture cancellate over entire length of shell. Apex with
long irregular slit. Mouth straight, section oval dorsoven-
trally.

Measurements: holotype L 130.9, W 19.6, w 2.4, arc 0.5.
W/w ratio 7.1.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

255

Remarks. — Fissidentalium metivieri can be compared to F. magnificum , which is more
regularly curved, with circular to slightly flattened section, with ribs equal in size and different W/w
ratio 5.3/1. Fissidentalium metivieri also resembles the fossil F. rectum (Linne) in shape, but that
species has fine longitudinal striae over the entire surface. F. exasperatum is less curved and more
sculptured transversally and also has longitudinal striae throughout. F. metivieri is the largest known
living scaphopod species.

Etymology. Named for Bernard Metivier, mnhn, who participated in several of the
expeditions on which the present report is based, and who assisted in many ways during my visits to
Paris.

Other Indo-Pacific species of Fissidentalium cited in the literature

Fissidentalium ceras (Watson, 1879): 510. “ Challenger " stn 246, 36°10' N, 178°00' E, 2050 fms
[3747 m]. Mid Pacific E of Japan. Syntypes bmnh 1887.2.9.10-11.

Fissidentalium complexum (Dali, 1895): 687, pi. 26, fig. 3. " Albatross ”, stn 3472, 256-280 fms [468-
512 m], Near Hawaii Is. Holotype usnm 107023.

Fissidentalium eualdes (Barnard, 1963a): 444. Off South Africa, 33°36' S, 16°15'E, 2778-2870 m.
Syntypes sam (fide Barnard, 1963b, not seen) and bmnh (1 dd, no registration number).

Fissidentalium exasperatum (Sowerby, 1903): 225, pi. 5, fig. 12. South Africa, Umhloti river mouth,
49 m. 3 syntypes dd bmnh 1903.7.27.57-59.

Fissidentalium lima Kuroda & Habe in Habe, 1963: 260, pi. 37, fig. 15. Japan, Okinoshima, Shikoku,
40 m. nsmt.

Fissidentalium horikoshii Okutani, 1982: 1, figs 1-5. E of Japan, 38°22' N, 143°26' E, 2930-3020 m.
NSMT.

Fissidentalium kawamurai Kuroda & Habe in Habe. 1962: 106, pi. 47, fig. 14. Ashizurimisaki, Kochi
Prefecture, Japan, nsmt.

Fissidentalium laterischismum Shikama & Habe, 1963: 249, textfig. 1-2. Okhotsk Sea, off Monbetsu,
Hokkaido, Japan.

Fissidentalium platypleurum (Tomlin, 1931): 339. South Africa, off Itongazi River, Natal, 46 m. sam.

Fissidentalium salpinx (Tomlin, 1931): 338, fig. 1. Off South Africa, “Cape Point NE 3/4°E 40 miles,
1280-1462 m’\ sam.

Fissidentalium tenuicostatum Qi & Ma, 1986: 69. South China Seas.

Fissidentalium yokoyamai (Makiyama, 1931): 44, pi. 1, fig. 1. Yokosuka City, Honshu, Japan.

Fissidentalium zelandicum (Sowerby, 1860): 101, pi. 223 ( Dentalium 1), fig. 13. New Zealand, bmnh
(not seen).

Genus Schizodentalium Sowerby, 1894

Type species (by monotypy): Schizodentalium plurifissuratum Sowerby, 1894.

Diagnosis. — Shell long, moderately curved, solid, opaque, cream to light brown.
Longitudinally hardly sculptured with primary ribs; secondary ribs present, starting near the apex and
reaching almost the size of the primary ones at the oral area. Rib section angled; cancellated.
Intercostal spaces concave, sculptured. Apex a fissure furnished with a series of 3-5 elongated holes
on the ventral side. Section polygonal.

Radula as in Fissidentalium.

Distribution. — Cretaceous-Recent, worldwide, shelf to abyssal.

256

VICTOR SCARABINO

HG' 62A-f' Fissidentahum metivieri , holotype (130.9 mm). — b. Fissidentalium magnificum (115 mm), musorstom 2: stn
CP 25. — c. Fissidentahum vicdani (114 mm), musorstom 3: stn CP 118. — d, 'Fissidentalium malayanum (55 mm).
biogeocal: stn C P 250. - e, Compressidentalium suhcurvatum (78 mm), corindon: stn DR 231. f, Fissidentalium
projundorum (71 mm), corindon: stn DR 229. — g. Pictodentalium festivum (50 mm). New Caledonia i.agon: stn 424
h. Fissidentalium cornubovis (56 mm), benthedi: stn CI1 87. i. F. magnificum. detail of the sculpture. j. F.
malayanum, detail of the sculpture. — k, C. suhcurvatum , detail of the sculpture. I, F. profundorum. detail of the
sculpture. Scale lines: 100 pm (i, j, k), 1 mm (b).

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIF1C

257

Schizodentalium plurifissuratum Sowerby, 1894
Figs 63, 70 f

Schizodentalium plurifissuratum Sowerby, 1894: 158, pi. 12, fig. 24.

Other references:

Dentalium plurifissuratum - Pilsbry & Sharp, 1897: 82, pi. 6, figs 87-89. — Sowfrby, 1903: 231. Smith, 1906b: 57. -
Plate. 1908a: 344, pi. 30, figs 12-16. - Bartsch, 1915: 181. Barnard, 1963b: 347.

Dentalium multistriatum - Plate, 1908: 437. Jaeckel, 1932: 304.

Type material. — Holotype bmnh 95.4.29.179, paratypes bmnh 95.4.29.80-81.

Type locality. — Said to be from Hong Kong (erroneous).

Material examined. — South Africa. "Meiring Naude stn SM 184, 33°39' S, 27°12' E,
86 m, 1 dd. — Stn SM 185, 33°39' S, 27° 1 1' E, 90 m, 1 dd (sam).

Fig. 63. — Distribution of Schizodentalium plurifissuratum.

Distribution. — Apparently endemic to South Africa and Agulhas Bank (Sowerby, 1903).
No information on living depth range; shells from 86 m (present paper) to 564 m (Barnard, 1963b).

Genus Compressidentauum Habe, 1963

Type species (OD): Dentalium hungerfordi Pilsbry & Sharp, 1897.

Diagnosis. Shell medium to large, slightly curved to almost straight, usually curved only
at apical portion. Solid, polished, white to red. Longitudinal sculpture of more than 14 strong
primary ribs; secondary ribs present, reaching the oral area. Rib section flat at top, round, smooth.
Intercostal spaces concave, smooth or with longitudinal striae. Apex w'ith flat, V-shaped notch on
ventral side, apical callus with terminal pipe or irregular slit. Section strongly compressed
dorsoventrally. Oral aperture oval or subtriangular.

Radula similar to that of Antalis.

258

VICTOR SCARABINO

Distribution. — Pacific and Indian Oceans. Absent in the Atlantic Ocean. Temperate-
tropical, shelf to abyssal.

Compressidentalium hungerfordi (Pilsbry & Sharp, 1897)
Figs 64, 70 g

Dentalium hungerfordi Pilsbry & Sharp. 1897: 84, pi. 6, fig. 86 (nom. nov. pro Denialium compression Sowerby. 1888: 569.
pi. 28, fig. 18, non Watson, 1879).

Other references:

Denialium (Fissidenlalium) hungerfordi - Boissevain, 1906: 38. pi. 2. fig. 30. — Hirase, 1931: 293, figs 749-750. — Kira. 1955'
80. pi. 40, fig. 10.

Fissidenlalium ( Compressidentalium ) hungerfordi - Habe, 1963: 260, pi. 37, fig. 14, textfigs 25-26; 1964a: 17, pi. I. fig. 14.
pi. 4, figs 25-26; 1971: 487 (Japanese text), 306 (English text), pi. 65, figs 2-3. Habe, 1977: 332. — Higo & Goto. 1993:
686.

Type material. — Holotype bmnh 1881.2.1.1.

Type locality. — Hong Kong.

Material examined. — Japan. Tosa Bay, Shikoku, 100 fms [183 m], Coll. Staadt, 1 dd
(mnhn).

Distribution. — Indonesia, China, Japan, 50-1570 m.

Compressidentalium compressiusculum (Boissevain, 1906)
Figs 65, 70 h, 73 h

Denialium compressiusculum Boissevain, 1906: 33, pi. 6, fig. 12.

Other reference:

Fissidenlalium ( Compressidentalium ) compressiusculum — Habe & Kosuge, 1964: 4.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL 1NDO-PACIFIC

259

Type material. — Holotype zma 3.06.044.

Type locality. — " Siboga ”, stn 241, 04°24' S, 129°49' E, 1570 m, Banda Sea.

Material examined. — The type material.

Chesterfield Islands, corail 2: stn DW 172, 18°26'S, 155° 12' E, 1100 m, 5 dd.

New Caledonia, biocal: stn KG 95, 21°22' S, 166°33' E, 2365 m, 1 dd.

biogeocal: stn KG 248, 21°15'S, 166°29' E, 2340 m, 1 dd. — Stn DW 296, 20°38' S, 167° 10' E,
1230-1270 m, 4 dd.

Loyalty Islands, musorstom 6: stn DW 468, 21°06' S, 167°33' E, 600 m, 1 dd.

Indonesia, corindon: stn DR 231, 00°05' N, 119°48'E, 1080 m, 1 Iv.

" Galathea stn 490, 05°25' S, 117°03' E, 585 m, 2 dd (zmc).

Philippines, musorstom 2: stn CP 50, 13°37' N, 120°34' E, 810-820 m, 2 lv. — Stn CP 55, 13°54' N,
1 19°58' E, 865 m, 12 lv, 28 dd. — Stn CP 79, 13°44' N, 120°32' E, 682-770 m, 1 dd. — Stn CP 81,
13°34' N, 1 20°3 1 ' E, 856-884 m, 2 lv.

musorstom 3: stn DR 93, 13°49' N, 120°02' E, 540 m, 8 lv, 12 dd. — Stn CP 106, 13°47' N, 120°30' E,
640-668 m, 5 lv, 25 dd.

estase 2: stn CP 6, 04°38' N, 1 19°49' E, 2570 m, 4 lv, 1 dd.

West Indian Ocean, benthedi: stn DS 64, 12°4T S, 44°57' E, 860-770 m, 1 dd.

md 32 Reunion: stn DC 18, 21°19' S, 55°16' E, 3150-3225 m, 1 dd. — Stn DR 104, 20°49' S, 55°01' E,

1875-1920 m, 4 dd. — Stn DR 106, 20°48' S, 55°05' E, 1710-1730 m, 2 dd.

Fig. 65. — Distribution of Compressidentalium compressiusculum.

Distribution. — Indonesia, now extended to the Philippines, New Caledonia and the Western
Indian Ocean; live records from 540 to 2570 m, mostly between 540 and 700 m (present paper).

Compressidentalium sedecimcostatum (Boissevain, 1906)
Figs 66, 70 j

Denialium sedecimcostatum Boissevain, 1906: 33, pi. 6, figs 8-11.

Other references:

Denialium sedecimcostatum - Plate, 1908: 347.

Fissidentalium ( Compressidentalium? ) sedecimcostatum — Habe & Kosuge, 1964: 4.
Striodentalium sedecimcostatum — Ql & Ma, 1989: 118, figs lOa-b.

260

VICTOR SCARABINO

Fig. 66. — Distribution of Compressidenialium sedecimcostatum.

Type material. — Lectotype (here designated) zma 3.06.035, paralectotypes zma 3.06.036-043.
Type locality. — “Siboga", stn 52, 09°03' S, 1 19°57' E, 959 m, Banda Sea.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DC 321, 21°20' S, 158°02' E, 1000 m, 2 dd. - Stn CP 324
21°15'S, 157°51'E, 970 m, 1 lv.

New Caledonia. " Vauban ” 1978-79: stn 32, 22°32' S, 166°25' E, 430-500 m, 1 lv. — Stn 33, 22°33' S,
166°25' E, 290-350 m, 2 lv, 7 dd. — Stn 34, 22°32' S, 166°26' E, 350-420 m, 2 lv, 6 dd. — Stn 35,
22°32'S, 166°26'E, 250-375 m, 1 dd. — Stn 39, 22°29' S, 166°23'E, 375-550 m, 1 dd. — Stn 40,
22°30' S, 1 66°24' E, 250-350 m, 8 lv, 47 dd.

biocal: stn CP 29, 23°08' S, 166°40' E, 1 100 m, 1 dd. — Stn CP 30, 23°09' S, 166°41' E, 1140 m,
1 dd. — Stn CP 55, 23°20' S, 167°30' E, 1 160-1 175 m, 1 lv, 3 dd. — Stn DW 66, 24°55' S, 168°22' E.
505-515 m, 2 lv, 4 dd. — Stn CP 69, 23°52' S, 167°58' E, 1220-1225 m, 1 dd. — Stn DW 104, 21°31' S,
166°21'E, 375-450 m, 3 dd.
lagon: stn 872, 20°37' S, 165°58'E, 105 m, 1 dd.

Passe de Boulari, B. Richer/ORSTOM coll., 400 m, 2 lv, 7 dd.

biogeocal: stn KG 219, 22°39' S, 166°34' E, 570 m, 1 dd. — Stn CP 232, 21°34' S, 166°27' E, 760-
790 m, 2 dd. — Stn CP 238, 21°28' S, 166°23'E, 1260-1300 m, 4 dd. — Stn CP 273, 21°02' S,
166°57' E, 1920-2040 m, 2 lv, 1 dd. — Stn DW 313, 20°59' S, 166°59' E, 1600-1640 m, 2 dd.
musorstom 4: stn DC 168, 18°48' S, 1 63° 1 1 ' E, 720 m, 1 dd. — Stn DW 220, 22°58' S, 167°38' E,
505-550 m, 1 lv. — Stn CP 242, 22°06' S, 167°10' E, 500-550 m, 1 lv.

Loyalty Islands, musorstom 6: stn DW 394, 20°49' S, 167°09' E, 570 m, 3 lv, 6 dd. - Stn DW 424,
20°24' S, 166°25' E, 599 m, 1 lv, 1 dd. — Stn DW 425, 20°24' S, 166°25' E, 594 m, 4 dd. — Stn DW
468, 21°06' S, 167°33' E, 600 m, 1 dd. — Stn DW 483, 21°20' S, 167°48' E, 600 m, 1 lv, 1 dd.
Philippines, estase 2: stn CP 2, 14°05' N, 120°02' E, 2050 m, 1 lv.
musorstom 3: stn CP 106, 13°47' N, 120°30' E, 640-668 m, 3 lv, 1 dd.

West Indian Ocean, benthedi: stn DR 33, 12°54' S, 45°16' E, 275-400 m, 3 lv, 14 dd. — Stn DR 38,
12°55' S, 45° 16' E, 200-500 m, 7 lv, 6 dd. — Stn DS 42, 13°05' S, 45°08' E, 400-520 m, 3 lv. — Stn
F 61, 1 2°46' S, 44°58' E, 475-510 m, 2 lv, 7 dd. — Stn DS 62, 12°46' S, 44°58' E, 530-535 m, 1 dd.
md 32 Reunion: stn DC 10, 21°13' S, 55°52' E, 930-980 m, 2 dd. — Stn DC 58, 21°03' S, 55°10' E,
450 m, 5 lv, 14 dd. — Stn DC 64, 21°12' S, 55°04' E, 1150-1 180 m. 1 lv, 3 dd. - Stn DC 112, 20°53'S
55°09' E, 740-780 m, 1 dd. — Stn DS 178, 21°04' S, 55'W E, 412-460 m, 4 lv, 2 dd.

" Mascareignes III”: R. von Cosel coll. 1986: stn 97, 22°24’ S, 43°04' E, 600 m, 1 dd.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL 1NDO-PACIFIC

261

Distribution. — Indonesia, East China Seas, 550 m (Qi & Ma, 1989) to East Africa, 1 143 m
(Plate, 1908). Now extended to New Caledonia; live records from 250 to 2050 m (present paper).

Compressidentalium subcurvatum (Smith, 1906)

Figs 62 e, k, 67, 71 g

Dentalium subcurvatum Smith, 1906a [October]: 251.

Synonym:

Dentalium marlensi Boissevain, 1906 [December]: 34. pi. 4, fig. 19 (Syn. nov.).

Other references:

Dentalium subcurvatum - Annandale & Stewart, 1909: pi. 23. fig. 3. — Winckworth, 1940a: 25.
Dentalium marlensi - Ludbrook, 1954: 93.

Fissidentalium marlensi - Kosuge, 19S5: 59, pi. 23, fig. 4.

Type material. D. subcurvatum : holotype bmnh. — D. martensi: lectotype (here designated)

zma 3.06.045, paralectotypes zma 3.06.046.047.

Type locality. — D. subcurvatum: Indian Ocean, SW Cape Comorin, “ Investigator " stn 275,
08°27' N. 75°35' E, 731-771 fms [1336-1409 m], — D. martensi: Indonesia, “Siboga", stn 88, 00°35' N,
119°09'E, 1301 m.

Material examined. — The type material.

New Caledonia, musorstom 4: stn DC 168, 18°48' S, 163°11'E, 720 m, 1 lv.

Indonesia, corindon: stn DR 231, 00°05' N, 119°48' E, 980-1080 m, 1 lv.

“ Snellius ” II: stn 4.164, 06°24' S, 120°21' E (no depth data), 5 dd (rmnh).

Philippines, musorstom 1: stn CP 47, 13°41'N, 120°30'E, 685-757 m, 1 dd.
musorstom 2: stn CP 50, 13°37' N, 120°33'E, 810-820 m, 91 lv, 181 dd. — Stn CP 55, 13°54' N,
1 19°59' E, 865-866 m. 10 lv, 12 dd. — Stn CP 81, 13°34' N, 121°31' E, 856-884 m, 4 lv, 3 dd.
musorstom 3: stn DR 94, 13°47'N, 120°03' E, 842 m, 1 dd. — Stn DR 95, 13°56'N, 119°59'E,
865 m, 2 lv, 11 dd. — Stn CP 114, 13°34' N, 120°29' E, 1000-1040 m, 1 lv.

West Indian Ocean. NW Madagascar, Crosnier coll. 1975: stn CH 142, 13°46’S, 47"34' E, 1250-
1300 m, 5 dd.

Fig. 67. — Distribution of Compressidentalium subcurvatum.

262

VICTOR SCARABINO

Distribution. — Indian Ocean and Indonesia, now extended to the Philippines and New
Caledonia; live records from 685 to 1090 m (present paper) and shells from 142 to 1411 m.

Remarks. — Fissidentalium clathratum Martens, 1881, from Northeastern Australia, could be
a senior synonym of C. subcurvatum, but until type material of that species is examined, it appears
best to consider C. subcurvatum the valid name of the present species.

Compressidentalium zanzibarense (Plate, 1908)
Figs 68, 70 i

Dentalium zanzibarense Plate, 1908a: 384, pi. 3, figs 35-36.

Other reference:

Dentalium zanzibarense — Ludbrook, 1954: 93,

Type material. — Holotype zmb 61092 (fide Kilias, 1995).

Type locality. — " Valdivia ”, stn 245, 05°28' S, 39°18' E, 463 m, Zanzibar Channel.

Material examined. — West Indian Ocean, benthedi: stn DR 40, 12°56'S, 45°18' E,
1300-1480 m, 1 lv, 1 dd.

MD32 Reunion: stn CP 105, 20°47' S, 55°04' E, 1740-1850 m, 9 lv, 14 dd. Stn DS 106, 20°48' S
55°05' E, 1710-1730 m, 1 lv. — Stn DC 138, 24°4T S, 55°36' E, 1500-1590 m, 4 dd.

“ Meiring Naude stn SM 53, 26°51' S, 33°13' E, 720 m, 1 dd. — Stn SM 78, 27°32' S, 32°50' E, 750
m, 1 lv, 3 dd. — Stn SM 86, 28°00' S, 32°41' E, 550 m, 1 dd. — Stn SM 94, 28°16' S, 32°39' E, 670
m, 1 dd. - Stn SM 117, 30°18' S, 31°10' E, 820 m, 3 dd. — Stn SM 121, 30°32' S, 30°53' E. 625-900
m, 1 dd. — Stn SM 123, 30°33' S, 30°49' E, 690 m, 1 lv. — Stn SM 129, 30°53' S, 30°31' E, 850 m,
1 dd. — Stn SM 131, 30°43' S, 30°42' E, 780 m, 8 dd. — Stn SM 226, 32°29' S, 28°59' E, 710-775 m,
36 dd. — Stn SM 232, 32°15' S, 29°10' E, 260-620 m, 1 dd.

Fig. 68. Distribution of Compressidentalium zanzibarense.

Distribution. — Off Zanzibar, now extended to Madagascar, Reunion Island and South
Africa, recorded alive from 690 to 1850 m.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

263

Compressidentalium ceciliae sp. nov.

Figs 69, 70 k, 71 i

Type material. Holotype mnhn. Paratypes: 11 mnhn, 1 ams C201725, 1 nmnz M268958,

1 USNM.

Type locality. — Southern New Caledonia, musorstom 4, stn DW 220, 22°58' S, I67°38' S,
505-550 m.

Material examined. — New Caledonia, musorstom 4: stn DW 220, 22°58' S, 167°38'S,
505-550 m, 9 iv (holotype and paratypes: 7 mnhn, 1 nmnz).

biocal: stn DW 08, 20°34' S. 166°54' E, 435 m, 4 dd. — Stn DW 46, 22°53' S, 167°17' E, 570-610 m,

2 lv, 6 dd. Stn DW 51, 23°05' S, 167°45' E, 680-700 m, 13 lv (paratypes: 4 mnhn, 1 ams), 21 dd
(1 paratype usnm).

Fig. 69. Distribution of Compressidentalium ceciliae.

Distribution. — E and SE New Caledonia, living in 505-700 m.

Description. — Shell to 36 mm long, solid, polished,
white, straight for the anterior three fourths, then curved at
posterior end. Section subcircular at apex, strongly compres¬
sed dorsoventrally otherwise. Sculpture consists of 15 smooth
prominent, primary ribs, more polished than the interspaces.
Two lateral ribs noticeably more prominent are present on
some specimens. Secondary ribs originate on the posterior 1/3
of the shell, becoming as prominent as the primary ribs at the

anterior end. Mouth oval, more compressed on ventral side.
Apex narrow with V-shaped notch on ventral side.

Measurements: holotype L 34.7, W 3.6-3, w 0.5. arc 1 (in
the curved part); paratypes L 34. W 3.6-3. w 0.8. arc 0.8; L
31.9, W 3. 4-2. 9, w 0.7, arc 0.7; L 35.5. W 3.7-3. 2, w 0.7. arc
1; L 29. W 3.6-3. 1, w 0.9, arc 1.1; L 27.1. W 3.3-3. w 0.7. arc
0.9; L 15.7, W 2.3-2, w 0.7, arc 0.5. W/w ratio 7. 2-3. 2.

Etymology. Named for the author" s wife, Cecilia Valdes.

Other Indo-Pacific species of Compressidentalium cited in the literature

Compressidentalium clathratum (Martens, 1881): 66. Off Moreton Bay, NE Australia, 'Gazelle”,
1005 m. Holotype zmb 33122 ( fide Kilias, 1995).

264

VICTOR SCARABINO

Compressiden talium lardum (Barnard. 1963a): 445. Off South Africa, 33°26' S, 16°33' E, 2267-2376 m
sam and paratype bmnh 1964.2.42.

Compressidentalium sibogcie (Boissevain. 1906): 39, pi. 4, figs 17-18, textfig. 22. Indonesia " Siboea' ’
stn 159, 00°59'S. 129°49'E, 411 m. zma. * ’

Compressidentalium sumatrense (Jaeckel, 1932): 304, textfig. 3. Indonesia, Sumatra, " Valdivia ” stn
191, 00"39' S, 98-52' E, 750 m. Holotype zmb 61081.

Genus Coccodentalium Sacco, 1896

Type species (OD): Dentatium radula Schroter, 1784. Miocene. Italy.

Diagnosis. - Shell medium in length, nearly straight, solid, polished, white, usually with dark
brown markings. Longitudinal sculpture of 6 primary ribs, secondary ribs reaching oral area Rib
section round, irregular due to nodules, cancellate throughout. Intercostal spaces concave
transversally sculptured. Apex with fiat V-shaped notch on ventral side, lumen circular Section
polygonal at apex, subpolygonal at aperture.

Radula rachidian with anterior border irregular and granulose; lateral similar to Dentalium smooth-
marginal sigmoidal.

Distribution. — Eocene-Recent, Pacific and Indian Oceans, shelf to bathyal.

Coccodentalium gemmiparum (Melvill, 1909)
Figs 70 l-m, 72, 73 e-f
Dentalium gemmiparum Melvill, 1909: 120, pi. 5, fig. 10.

Type material. — Holotype bmnh 1910.3.17.5.

Type locality. — Indian Ocean, Chagos Archipelago, Diego Garc°a Lagoon.

Material examined. — The type material.

New Caledonia, biogeocal: stn CP 278, 22°48' S, 166°20' E, 2250 m 1 dd
Philippines, musorstom 3: stn DR 126, 11°49' N, 121°22' E, 266 m 2 dd —
122°14' E, 240-267 m, 153 lv, 96 dd.

Stn CP 139, 11°53' N,

alive inD740R?A7TI0^' Chagos Archipelago, now extended to the Philippines and New Caledonia,
alive in 240-267 m (present paper), shell (probably washed down) in 2250 m.

he s ulpiure -A dei ngZude~- stn m TcnZ ^cll (32 mm), apical and oral sections, detail of

section -Vauban- 1978-79 74 L >• J' ComfreSS'den'al,T ^deamcostatum, shell (52 mm), apex and apical

dorsal Views ap ex. apica mediS'and n0V” h°^ shuC" <347 m™>' lateral and

sections. S s» CP StSSil g«S^j=5S5 SSM JF “P“

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

265

Source : MNHN, Paris

266

VICTOR SCARABINO

Rc;. 71. — Radulae. — a. Dentalium deforgesi, general view, note the tooth of the right column in different positions. — b,
same specimen, detail of laterals. — c. Dentalium caledonicum, general view. d, Dentalium oryx, view of the heads
of laterals. e, Fustiaria langfordi, detail of laterals. — f. Graptacme lactea , external face of a lateral tooth. — g,
Compressidentalium subcurvatum, general view. h. Fustiaria vagina, general view. — i, Compressidentalium ceciliae,
general view. Scale bars: 200 pm (a, c, e), 100 pm (b. d, f, h, i), 500 pm (g).

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL 1NDO-PACIFIC

267

Fig. 72. Distribution of Coccodentalium gemmiparum.

Genus Pictodentalium Palmer, 1974
Type species (OD): Denlalium hirasei Kira, 1959.

Diagnosis. — Shell medium to very large, slightly to well curved, solid, polished, light yellow
to brightly coloured with red, violet and green patches. Longitudinal sculpture of wide primary ribs
and secondary ribs reaching the oral area. Rib section flat-topped to rounded. Intercostal spaces,
straight or concave, smooth or longitudinally sculptured. Apex with terminal callus and flat V-shaped
notch on ventral side; fissured terminal pipe. Section circular to slightly compressed dorsoventrally.
Oral aperture usually thin and coloured.

Radula rachidian flat, curved, anterior margin very slightly granulose; lateral with prominent primary
cusp and sculptured head; marginal slightly curved.

Distribution. Recent, Pacific and Indian Oceans, absent in the Atlantic Ocean, sublittoral
to bathyal.

Remarks. Pictodentalium was invalidly introduced by Kira (1959), who is generally cited
as author of this name. However, I refer to the discussion by Bieler & Petit (1990), who credit
authorship of Pictodentalium to Palmer (1974).

Pictodentalium formosum (Adams & Reeve, 1848)

Figs 74, 77 b

Denlalium formosum Adams & Reeve, 1848: 71. pi. 5, figs la-b.

Synonyms:

Pictodentalium formosum liirasei Kira. 1959: 117, pi. 41, fig. 11.

Fissidentalium formosum harrisoni Habe, 1970: 95.

Other references:

Denlalium formosum - Sowerby, I860: 102. pi. 223 ( Denlalium 1), fig. 2; 1873: pi. 2, fig. 7. — Clessin 1896: 21, pi. 6,
fig. 7. Pilsbry & Sharp, 1897: 2, pi. 1. figs 9-1 1. — Boissevain, 1906: 8, pi. 1, fig. 2. — Kira. 1955: 80, pi. 40, fig. II.
Denlalium (Denlalium) formosum - Hirasf.. 1931: 135, pi. 3, fig. 3.

Pictodentalium formosum — Habe, 1977: 332.

268

VICTOR SCARABINO

Fig. 73. — a-d Fissidentalium shoplandi (153 mm), musorstom 3: stn CP 128. — b-c, details of the sculpture. -
d. Fissidentalium shoplandi. holotype (71 mm). e-f, Coccodentalium gemmiparum, details of sculpture. g, Antalis
boucheti, detail of the sculpture. h. Compressidentalium compressiusculum, detail of the sculpture. i, Graptacme
lactea. apex and sculpture. Scale bars: iOO pm (b, e, g. h. i), 10 pm (c, f).

Source : MNHN. Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

269

Piciodentalium formosum hirasei — Okutani, 1983: 12, pi. 43, fig. 12.

Fissidenialium '( Piciodentalium) formosum - Habe, 1963: 255, pi. 37, figs 4, 12, textfig. 22. Habe& Kosuge, 1964: 4; 1977:
332. Habe & Kosuge, 1966: 117, figs 22-23. — Springsteen & Leobrera. 1985: 286, pi. 82, fig. 5. — Higo & Goto, 1993:
686.

Fissidenialium formosum — Qi & Ma, 1989: 116, figs 6a-b.

Fissidenialium formosum harrisoni - Higo & Goto, 1993: 686.

Type material. — D. formosunr. 3 syntypes dd bmnh 1951.2.14.1-3. — P. f. hirasei : not
checked. — F. f harrisoni : nsmt 37303 (fide Habe, 1970) [not seen].

Type locality. — D. formosum-. Sulu Archipelago, 29-36 m. P.f hirasei : Southern Japan,
20 fms [37 m]. — F. f. harrisoni : South China Sea.

Material examined. The type material of D. formosum.

China. South China Sea, Coll. Staadt, 1 dd (mnhn).

Distribution. — Japan, 0-50 m (Habe & Kosuge, 1964), South China Sea, 77-145 m (Qi &
Ma, 1989), the Philippines (Springsteen & Leobrera, 1985).

Pictodentalium festivum (Sowerby, 1914)
Figs 62 g, 75, 77 c, 78 b

Denialium feslivum Sowerby. 1914: 8, textfig.

Other reference:

Denialium feslivum - Habe, 1963: 255 (as synonym of P. formosum).

Type material. — Holotype bmnh 1914.4.21.

Type locality. — New Caledonia.

Material examined. — The holotype.

Chesterfield Islands, chalcal 1: stn DC 6, 20°57' S, 161°43' E, 45 m, 1 dd. — Stn DC 26, 19°1 1' S,
1 58°35' E, 48 m, 2 dd. — Stn DC 37, 19°54' S, 158°46' E, 50 m, 2 dd. — Stn DC 45, 20°50' S,

270

VICTOR SCARABINO

158°30' E, 50 m, 1 dd. — Stn DC 50, 21°04'S, 158°41'E, 70 m, 1 dd. — Stn DC 56, 21°24' S,
159°09' E, 60 m, 1 Iv. — Stn DC 59, 21°40' S, 159°21' E, 56 m, 1 dd.
musorstom 5: stn DW 264, 25°20' S, 159°44' E, 56 m, 1 lv, 2 dd.

corail 2: stn DW 71, 19°15'S, 158°24'E, 55 m, 1 lv. — Stn DW 91, 19°03'S, 158°55' E, 43 m,
1 lv. — Stn DW 103, 19°01' S. 158°32' E, 58 m, 1 dd. — Stn DW 117, 19°25' S, 158°32' E, 52 m, 1
dd. — Stn DW 136, 19°31' S, 158°16' E, 37 m, 2 dd.

New Caledonia, lagon: stn 120, 22°28' S, 166°44' E, 46 m, 1 dd. — Stn 129, 22°31' S, 166°47' E, 44-
46 m, 2 lv, 3 dd. — Stn 151, 22°32' S, 166°48' E, 31-33 m, 1 lv. — Stn 229, 22°39' S, 166°40' E,
41 m, 1 dd. — Stn 230, 22°38' S, 166°41' E, 35 m, 1 lv. — Stn 234bis, 22°32' S, 166°51' E, 60 m,
1 dd. — Stn 241, 22°21' S, 167‘W E, 35 m, 2 lv. — Stn 313, 22°40' S, 166°50' E, 30 m, 1 lv. — Stn
326, 22°26' S, 167°02' E, 67 m, 1 dd. — Stn 345, 22°46' S, 166°50' E, 39 m, 1 dd. — Stn 358, 22°31' S,
167°05' E, 50 m, 1 dd. — Stn 362, 22°38' S, 167W E, 83 m, 1 dd. — Stn 373, 22°28' S, 167°11' E,
52-57 m, 2 dd. — Stn 376, 22°34' S, 167°06' E, 75-76 m, 1 dd. — Stn 377, 22°35' S, 167°08' E, 56 m.

1 dd. — Stn 382, 22°30' S, 167°14' E, 57 m, 1 dd. — Stn 383, 22°32' S, 167° 13' E, 62 m, 1 lv, 2 dd.

Stn 403, 22°35' S, 167°18' E, 45 m, 2 dd. — Stn 414, 22°37' S, 167°16' E, 60 m, 1 dd. — Stn 416,
22°38' S, 167° 14' E. 40-50 m, 1 lv. — Stn 474, 18°02' S, 163°02' E, 52 m, 1 lv. — Stn 542, 19°06' S,
163° 10' E, 50 m, 2 dd. — Stn 603. 22° 16' S, 167°05' E, 78-80 m, 2 dd. — Stn 607, 22° 12' S, 167°03' E,
48-54 m, 1 dd. — Stn 619, 22°03' S, 166°54' E, 27-42 m, 1 dd. — Stn 729, 21°19' S, 165°54' E, 42-
45 m, 2 dd. — Stn 858, 20°37' S, 165°07' E, 220 m, 2 dd. — Stn 888, 20°22' S, 164°38' E, 20 m,

2 lv.

musorstom 4: stn DW 231, 22°34' S, 167° 10' E, 75 m, 1 dd.

“Vauban” 1978-79: stn 10, 22°17'S, 167°05'E, 80 m, 1 dd.

Hot Maitre, 20 m. Estival coll. 1980, 2 dd (mnhn).

Distribution. — New Caledonia, alive in 20-62 m (present paper).

Remarks. — Habe (1964) considered Pictodentalium festivum a junior synonym of P.
formosum but examination of the type material and other specimens of both species allows me to
reestablish Sowerby’s species. P. festivum has more ribs (17-19), is angled in section, and the material
examined always presented secondary ribs. The ribs of P. formosum are wide and round in section,
fewer in number (13-17) and have no secondary ribs. Also the color pattern is different, P. formosum
is basically dark purple with rose and green patches, while P. festivum is light purple, rose and cream.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

271

Pictodentalium vernedei (Sowerby, 1860)

Figs 76, 77 a

Dentalium vernedei Sowerby, 1860: 101, pi. 223 (Dentalium 1), fig. 3.

Other references:

Dentalium vernedei - Pilsbry & Sharp, 1897: 80, pi. 3, figs 35. 43. — Sowerby, 1873: pi. 1, fig. 3.

Dentalium ( Fissidentalium ) vernedei - Hirase, 1931: 137, pi. 3, fig. 8. Habe, 1953: 293, figs 741-743; 1963: 258, pi. 37,
fig. 9, textfigs 30-31.

Fissidentalium vernedei — Kira, 1962: 117, pi. 41, fig. 13.

Fissidentalium (Pictodentalium) vernedei - Habe, 1964a: 16, pi. 1, fig. 9, pi. 4, figs 30-31. - Habe & Kosuge, 1964: 3. Habe
1977: 332. — Ieyama 1993: 246, figs 4-5. — Higo & Goto, 1993: 686.

Type material. — bmnh (not seen).

Type locality. — Japan.

Material examined. — China. No further data, Coll. Jousseaume, 1 dd. Coll. Denis, 3 dd
(both mnhn).

Japan. No other data. Coll. Staadt, 3 dd (mnhn).

Fig. 76. Distribution of Pictodentalium vernedei.

Distribution. — China and Japan, 20-100 m.

Family Calliodentaliidae Chistikov, 1975

Genus Calliodentalwm Flabe, 1964

Type species (OD): Dentalium crocinum Dali, 1907.

Diagnosis. Shell medium to large, well curved, thin but not fragile, polished, shiny, white,
yellow or orange. Smooth or sculptured only at apical portion by longitudinal striae or close, fine,
encircled wrinkles. Apex simple or with fiat V-shaped notch on ventral side. Section subcircular,
slightly compressed dorsoventrally, more noticeable on ventral side.

272

VICTOR SCARAB I NO

Fig. 77. a. Pictodentahwn vernedei, shell (119 mm), apex. Japan (mnhn). b. Piciodenialium formoswn, shell (53 mm),
apical and oral sections, detail of the sculpture. South China Sea (mnhn). c. Piciodenialium type radula ( I’, festivum)-
see also Fig. 78 b.

Fig. 78. Radulae. — a, Fissidenlahum magmficum, rachidian, anterior border. b, Piciodenialium festivum , rachidian detail
ot the external tace. c. Fissidentalium candidum (North Atlantic Ocean), rachidian, external face. d Fustiaria
caesura, rachidian, anterior border. Scale bars: 100 pm (a). 200 pm (c). 20 pm (b d)

Source : MNHN, Paris

SCAPHOPODA OF THF TROPICAL INDO-PACIFIC

273

Radula rachidian slightly curved, anterior margin irregular, smooth; subrachidians (see below)
subpyramidal; laterals with the head slightly differenciated from body, one primary cusp almost
central and one secondary cusp sharp, another one with low cusp; marginals almost straight.

Distribution. — Recent, worldwide, temperate and warm waters, shelf to bathyal.

Remarks. - - This genus presently contains four species: C. callipeplum (Dali, 1889) from the
Western Atlantic Ocean and Caribbean Sea, C. crocinum, C. semitracheatum, and C. balanoides from
the Indo-Pacific. The unique radular formula ( 1 - 1 - 1 r- 1 - 1 r- 1 - 1 ) is characteristic, with an accessory
structure which I name subrachidian. Though considered by Scarabino (1981: 30) aberrant in C.
callipeplum (Dali, 1881) (Fig. 88 d), recent radular study confirmed this formula and tooth
morphology for three additional species; I now consider it a generic character. Shells of all four
species have similar characteristics, large, with pronounced curvature, rapidly increasing from apex
to oral aperture, with high W/w ratio, slight dorsoventral depression, smooth and shiny.

For remarks on the nomenclature of the names Eboreidens and Eboreidentidae, see under Graptacme
laclea.

Calliodentalium semitracheatum (Boissevain, 1906)
Figs 79, 82 a-d, 88 a

Dentalium semitracheatum Boissevain, 1906: 56, pi. 4, figs 20-21.

Synonym:

Dentalium ( Plagioglypta) curvotracheatum Plate, 1908a: 358, pi. 30. fig. 47 (Syn. nov.).

Type material. — D. semitracheatum : lectotype (here designated) zma 3.06.072, paralectoty-
pes zma 3.06.071. D. curvotracheatum-. holotype zmb 61086 ( fide Kilias, 1995).

Type locality. — D. semitracheatum-. “ Siboga ”, stn 284. 08°43' S. 127"17' E. 828 m, Timor
Sea. — D. curvotracheatum'. “Valdivia", stn 245, 05°28' S. 39"18' E. 463 m, Zanzibar Channel.

Material examined. — The type material.

Philippines. MUSORSTOM 1: stn CP 44, 13°47' N. 120°30' E, 592-610 m. 2 dd.

MUSORSTOM 2: stn CP 50, 13°37' N, 120°33' E, 810-820 m, 4 dd. - Stn CP 55. 13°54' N. 1 19°58' E.
865 m, 2 lv, 2 dd. Stn CP 82, 13°46' N. 120°28' E. 550 m, 2 lv.

274

VICTOR SCARABINO

musorstom 3: stn DR 93, 13°49' N, 120°02' E, 540 m, 2 lv, 2 dd. — Stn CP 106, 13°47' N 120°30' E
640-668 in, 3 dd.

Indonesia. “ Snellius " II: stn 4.113, 08° 19' S, 11 8° 16' E (no depth data), (dd). — Stn 4 131 08° 1 8' S
11 8° 18' E, 680-800 m, (dd) (rmnh).

" Galathea stn 490, 05°25' S, 117°03' E, 585 m, 1 dd (zmc).

West Indian Ocean, benthedi: stn DR 38, 12°55' S, 45° 15' E, 200-500 m, 4 dd. — Stn DS 53 13°00' S
44°54' E, 755-770 m, 1 dd.

SW Madagascar. “Mascareignes III”, R. von Cosel coll. 1986: stn 69, 22°22' S. 43°05' E, 350-420 m
1 dd. — Stn 78, 22°2 1 ' S, 43°03' E, 530 m, 1 lv. — Stn 81, 22°23' S, 43°03' E, 525 m, I dd. — Stn
127, 22°21' S, 43°02' E, 610 m, 1 lv. — Baie de Fanemotra, 450 m, 1 dd.

Madagascar. " Vauban ”, A. Crosnier coll. 1973: stn 49, 15°18' S, 46°10' E, 500-550 m, 2 dd Stn
50, 15° 19' S, 46° 12' E, 405 m, 3 dd. — Stn 112, 22°18'S, 43°02' E, 640-660 in, 1 lv. — Stn 114
22° 15' S, 43°05' E, 470-475 m, 2 dd.

Distribution. — The Philippines and Indonesia, now extended to Madagascar, live records
from 450 to 865 m (present paper).

Calliodentalium crocinum (Dali, 1907)
Figs 80, 88 b

Denialium crocinum Dali, 1907: 169, pi. 27, fig. 6.
Other references:

Denialium ( Laevidenialium ) crocinum - Kira, 1955: 80, pi. 40, fig. 9 — Habe 1955- 10
Laevidenialium crocinum - Habe, 1963: 268, pi. 38, figs 29, 35, textfigs 28-29.’- Habe & Kosuge, 1964- 7

;-,HrA’V964o 31» pL }’ ,35> P>- 4> figs 28-29: 1977: 336, pi. 70. figs 7-8. - Springsteen &

Leobrera, 1985. 287, pi. 82, fig. 9. — Habe et al.. 1986: 24. — Higo & Goto, 1993: 688.

Type material. — Holotype usnm 110508

Type locality. — Japan, Gulf of Tokyo, Albatross ”, stn 5094, in 188 fms [161 m].
Material examined. — The type material.

Philippines, musorstom 3: stn CP 99, 14°01' N, 120°19' E, 196-204 m, 1 lv. — Stn CP 101, 14°00' N
120° 19' E, 194-196 m, 1 lv. — Stn DR 126, 11°49'N, 121°22' E, 266 m, 1 lv.

Fig. 80. — Distribution of Calliodentalium crocinum.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

275

Distribution. Japan Sea (Habe, 1957) to the Philippines (Springsteen & Leobrera, 1985).
Recorded alive from 194 to 266 m (present paper).

Calliodentalium balanoides (Plate, 1908)
Figs 81, 82 e-f, 88 c

Denialium ( Laevidentalium ) balanoides Plate, 1908a: 357, pi. 30, figs 42-44.

Type material. — Holotype zmb 61109 (fide Kilias, 1995).

Type locality. — "Valdivia”, stn 186, 03°22' S, 1 0 1 ° 1 1 ' E, 903 m, West Sumatra.

Material examined. — New Caledonia. “ Vauban " 1978-79: stn 40, 22°30' S, 166°24' E,
250-350 m, 1 dd.

biocal: stn CP 108, 22°03' S, 167°06' E, 335 m, 1 dd.
musorstom 4: stn CC 247, 22°09' S, 167° 13' E, 435-460 m, 1 lv.
biogeocal: stn DW 253, 21°32' S, 166°29' E, 310-315 m, 1 dd.

Passe de Boulari, B. Richer/ORSTOM coll., 400 m, 1 dd.

Loyalty Islands, musorstom 6: stn DW 426, 20°25' S, 166°23' E, 610 m, 1 dd.

Philippines, musorstom 2: stn CP 50, 13°37' N, 120°33' E, 810-820 m, 3 dd. — Stn CP 55, 13°54' N,
119°58'E, 865-866 m, 4 dd. — Stn CP 71, 1 4°00' N, 120° 18' E, 189-197 m, 1 lv. — Stn CP 75,
13°51'N, 120°30'E, 300-330 m, 1 lv.

musorstom 3: stn DR 95, 13°56' N, 1 19°59' E, 865 m, 1 lv, ldd. — Stn CP 1 12, 14°00' N, 120°18' E.
187-199 m, 1 lv. — Stn DR 126, 1 1°49' N, 121°22' E, 266 m, 4 dd.

Distribution. — Sumatra, now extended to the Philippines and New Caledonia, living from
187 to 865 m.

276

VICTOR SCARABINO

Fig. 82. a-d, Calliodenialium semitracheatum, internal view of the radula, note the secondary teeth at left. — b, external face
of the rachidians and secondary teeth (arrows). — c, anterior view of the radula, note the position of the secondary
teeth. — d, secondary denticle. — e, Calliodenialium balanoides, lateral teeth. — f, internal face of the rachidians and
secondary teeth. Scale lines: 100 pm (a-c, e, 0. 10 pm (d).

Family Fustiariidae Steiner, 1991
Genus Fustiaria Stoliczka, 1868

Type species (SD by Pilsbry & Sharp, 1897): Dentalium circinatum Sowerby, 1823. Eocene, Paris Basin. France.

Diagnosis. — Shell medium, well curved, fragile but not thin, shiny, translucent white, yellow,
orange, red, lacking sculpture. Apex with long, regular slit or short v-shape notch; section circular
throughout.

Radula rachidian well curved in section, granulose; lateral with strong, armed granulose head;
marginal slightly sinusoidal.

SCAPHOPODA OF THE TROPICAL IN DO-PACIFIC

277

Distribution. Triassic-Recent, worldwide, temperate and tropical waters, sublittoral to
shelf and bathyal.

Fustiaria nipponica (Yokoyama. 1922)
Figs 83, 88 e

Denialium (Fustiaria) nipponicum Yokoyama. 1922: 119. pi. 6, fig. 7.

Synonyms:

Denialium nagoense Dali. 1927: I.

Denialium t Fustiaria) numatai Hirase. 1931: 139. pi. 3. fig. II.

Ollier references:

Denialium I Fustiaria) nipponicum — HiraSE, 1931: 139. pi. 3. fig. II. Habf., 1957: 131.

Fustiaria nipponica - Habe, 1964a: 27, pi. 2, figs 6-8; pi. 3, figs 8, 11; pi. 4, figs 1-2; 1971: 490 (Japanese text), 308 (English
text), pi. 116. figs 15-16; 1977: 336, pi. 69. lies 1-2. Habe & Kosuge, 1964: 6. Springsteen & Leobrera. 1985: 287.
pi. 82. fig. 8. Higo & Goto. 1993: 687.

Fustiaria ( Fustiaria ) nipponica — Habe, 1963: 266, pi. 38, figs 6-8, 26, textfigs 1-2.

Denialium polilum - Boissevain, 1906: 58. pi. I. fig. 20.

Denialium stenoscliizum - Boissevain, 1906: 59. pi. 6, figs 16-17.

Denialium nagoense — Habe, 1964b: 140, pi. 9, figs 1-2.

Denialium (Fustiaria) numatai - Habe, 1953: 295, figs 755-756.

Type material. — D. nipponicum and D. numatai. depository not checked. — D. nagoense :
holotype usnm 333718.

Type locality. D. nipponicum : Shito, Honshu, Japan (Pleistocene). — D. nagoense : Japan,
Nago, Okinawa, 15 fins [27 mj. — D. numatai Japan, Osumi and Oshima.

Material examined. — The holotype of D. nagoense. Material identified as D. stenoscliizum
and D. politum by Boissevain (1906).

Chesterfield Islands, chalcal I: stn DC 17, 19°12'S, 158°56'E, 44 m, 2 Iv, 1 dd. Stn D 34,
19°52'S, 158°20'E, 33-37 m. 1 dd.

corail 2: stn DW 72. 19°15'S, 1 58°21 ' E, 32 in. 1 Iv. — Stn DW 77. 19°12'S, 158°36' E, 60 in,
1 Iv. Stn DW 126, 19°28'S, 158°27'E, 46 m, 1 Iv, I dd. — Stn DW 147. 19°37' S. 158"14'E,
25 in, 1 Iv.

New Caledonia. lagon: stn 4, 22°13'S. 166°21'E. 9 m. 1 lv, 3 dd. - Stn 8, 22°23' S. 166°18'E,
15 in. 2 lv. 4 dd. Stn 21, 22°23' S. 166°23' E. 10 m. I dd. - Stn 50. 22°17' S. 166°12' E. 12 m,
1 dd. Stn 51. 22" 1 5' S, 166°11' E. 10 in, 1 lv. — Stn 63, 22°26' S, 166°26' E, 20 in, 1 lv. — Stn 64.
22°28' S, 1 66°25' E, 15 in, 1 lv. - Stn 65, 22°29' S. 166°26' E. 24 in. 2 lv. 2 dd. — Stn 66. 22°28' S.
166°27'E. 15 m, 2 lv, 1 dd. — Stn 68. 22°24' S, 1 66°30' E, 22-40 m, 1 lv. Stn 80, 22°3TS.
1 66°28' E. 33 m. I lv. Stn 83, 22°32' S, 166°30' E, 22 m, 1 dd. Stn 84, 22°30' S. 166"3T E,
17 m. 2 lv. Stn 98, 22"36' S. 166°32' E, 15 m. 3 dd. Stn 161, 22°34' S, 166°38' E, 20 m. 1 lv,

I dd. Stn 163, 22°12' S. 166"! 8' E, 15 in. I lv. 1 dd. Stn 170, 22"09' S. 166°07' E. 22 in. 3 lv.

Stn 185, 22°05' S. 166°02' E, 15 in, 1 lv. — Stn 211, 21°55' S. 165°52' E, 12 in, 1 lv. — Stn 212,
21°56' S, 165°53'E, 10 m, 1 lv. Stn 214. 2I"55'S, 165°48' E, 12 m. 1 dd. Stn 217, 21°53' S,
165°47'E, 16 in, 1 dd. Stn 226, 22°38' S, 166°39'E. 28 m, 4 lv, 3 dd. — Stn 233, 22°35' S,
1 66°46' E, 30 m, 1 Iv. - Stn 268, 22°20' S, 1 66° 17' E. 24 m, 1 Iv. Stn 281, 22"24' S. 166"24' E.
10 in. 1 lv. — Stn 284, 22°26' S, I66"25' E, 6 m, 3 lv, 1 dd. Stn 293. 22"42' S. I66°4T E, 20 in,
6 lv, 4 dd. Stn 296. 22°41' S. 166°44' E, 26 m, 1 Iv. — Stn 304, 22°40' S, 166°48' E, 27 m, 1 lv,

1 dd. Stn 31 1, 22°44' S. I66°47' E. 36 in, 1 lv, 1 dd. — Stn 322, 22°30' S, 166"58' E, 71 in, 2 lv.

Stn 340, 22"48' S. 166°47' E. 27 in, 1 Iv. Stn 438, 18° 10' S, 162°5T E. 37 in, 1 lv, 1 dd. Stn
441. 18°04'S. 162°56'E, 37 m. 3 dd. — Stn 442bis, 18°02'S. 162°56' E. 39 m, I dd. Stn 446.
18" 1 9' S, 163"04' E, 36 in. 7 lv, 3 dd. Stn 448, 18"22' S, 163"07' E, 30 in, 1 lv. Stn 449, 18°22' S,
163"09' E. 21 in. I lv. Stn 450. 18"24' S. 163"24' E, 29 in. 1 lv. 1 dd. Stn 453. 18"29' S, 163"12' E.

Source : MNHN, Paris

278

VICTOR SCARABINO

26 m, 1 lv, 4 dd. — Stn 465, 18°22' S. 163°05' E, 45 m, 2 Iv, 5 dd. — Stn 466, 18°24' S, 163°07' E,

42 m, 5 lv, 2 dd. — Stn 467, 18°25' S, 163°08' E, 41 m, 3 lv, 1 dd. — Stn 468, 18°27' S, 163°10' E,

40 m. 1 lv. — Stn 469, 18°29' S, 163° 10' E, 39 m, 1 lv, 1 dd. — Stn 470, 18°28' S, 163°09' E, 41 m,

1 dd. — Stn 471, 18°28' S, 163°07' E, 42 m, 2 lv, 1 dd. — Stn 473, 18°24' S, 163°03' E, 50 m, 1 lv,

3 dd. — Stn 481, 18°57' S, 163°32' E, 33 m, 3 lv, 7 dd. — Stn 482, 18°49' S, 163°31' E, 33 m, 1 lv,

8 dd. — Stn 512, 19°24' S, 163°35' E, 59 m, 1 lv, 3 dd. — Stn 519, 19°02' S, 163°34' E. 39 m, 1 lv,

2 dd. — Stn 536, 19°09' S, 163°23' E, 61 m, 14 lv. — Stn 554, 22°50' S, 166°54' E. 27 m, 1 lv. — Stn

866, 20°38' S, 165°03' E, 26 m, 1 lv. — Stn 871, 20°37' S, 165°00' E. 27 m, 1 lv. — Stn 916, 20°56' S,
1 64°28' E, 13 m, 2 lv. — Stn 932, 20°46' S, 164° 17' E, 23 m, 3 dd. — Stn 937, 20°40' S, 164° 15' E,
50-55 m, 1 lv. — Stn 943, 20°37' S, 164°11' E, 15 m, 1 dd. — Stn 984, 20°21' S, 163°56' E, 21-23 m,
1 dd. — Stn 1015, 20°10' S, 163°52' E, 25 m, 1 dd. — Stn 1025, 20°07' S, 163°49' E, 25-28 m, 1 lv.

Stn 1 105, 1 9°40' S, 163°57' E, 25 m, 4 dd. Stn 1118, 19°35' S, 163°52' E, 30 m, 2 lv, 2 dd.

Stn 1145, 19°21'S, 163°45' E, 38 m, 1 dd. — Stn 1146, 19°08' S, 163°31' E, 185 m, 1 dd. — Stn 1154,
19°09'S. 163°19'E, 40 m, 1 lv, 1 dd.

Loyalty Islands, musorstom 6: stn DW 430, 20°21' S, 166°07' E, 30 m, 1 lv, Idd. — Stn DW 431,
20°22' S, 166° 10' E, 21 m, 1 lv. — Stn DW 435, 20°21' S, 166°08' E, 32 m, 2 lv, 1 dd.

Indonesia. ", Snellius ” II: stn 4.020, 05°57' S, 123°46' E, 255-275 m, 1 dd. — Stn 4.153, 06°22, S,
120°26' E, 130-155 m, 1 lv. — Stn 4.155, 06°22' S, 120°26' E, 233-274 m, 1 dd (RMNH).
Philippines, musorstom 3: stn CP 143, 11°29'N, 124°11' E, 205-214 m, 1 dd.

Distribution. — Japan, the Philippines, China, 0-200 m (Habe & Kosuge, 1964), now
extended to New Caledonia, living from 6 to 71 m (present paper).

Fustiaria engischista (Barnard, 1963)

Figs 84, 88 f

Dentalium engischistum Barnard, 1963b: 351, fig. 30 f.

Type material. — Holotype sam A5463, paratypes bmnh 1964.256.

Type locality. — South Africa, off Cape Natal (Durban), 62 fms [113 m]

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

279

Material examined. — The type material.

West Indian Ocean, benthedi: stn DR 08, 1 P29' S, 47° 18' E, 250 m, 1 dd. — Stn F 77, 12°34' S.
44°54' E, 480-530 m, 1 dd. — Stn DR 104, 11°26' S, 47°22' E, 330-530 m, 1 dd.

MD32 Reunion: stn DS 178, 21°24' S, 55°10' E, 412-460 m, 1 lv.

Nosy Be Island, NW Madagascar, Plante coll., 6 dd (bmnh).

Distribution. — Southwestern Indian Ocean from Natal to NW Madagascar and Reunion
Island, alive in 412-460 m.

Fustiaria caesura (Colman, 1958)
Figs 78 d, 85, 88 g, j

Dentalium ( Pseudoantalis) caesura Colman, 1958: 145, fig. 11.

Fig. 85. - Distribution of Fustiaria caesura.

280

VICTOR SCARABINO

Type material. Holotype ams C62230. paratypes C62229.

Type locality. — Australia, off Willongong, New South Wales, 183 m.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DC 266, 25°20' S. 159°46'E, 240 m, 1 lv. — Stn DC 357,
19°37' S. 1 58°46' E, 630 m. 1 dd. — Stn DC 380, 19°38' S, 158°44' E, 555-570 m, 1 dd.

New Caledonia, musorstom 4: stn DW 159, 18°46' S, 163° 16' E, 585 m, 3 dd.

Philippines, musorstom 2: stn DR 34, 13H28'N, 1 2 1 °1 2' E, 155-167 m, 2 lv.
musorstom 3: stn CP 143, 11°29'N, 124°11'E, 205-214 m, 1 dd.

Distribution. E Australia, now extended to New Caledonia and the Philippines. Live
records from 155 to 240 m.

Fust i aria langfordi (Habe, 1963)
Figs 71 e, 86, 88 h

Laevidenialium langfordi Habe, 1963: 268, pi. 38. fig. 22.

Other references:

Laevidenialium langfordi - Habe, 1964a: 35. pi. 2, fig. 22; 1971: 491 (Japanese text), 309 (English text), pi. 65, figs 30-31.
Habe & Kosuge, 1964: 7. Habe el a!.. 1986: 24. Higo & Goto, 1993: 687 (as synonym of Laevidenialium longiiror-
sum).

Type material. Holotype nsmt.

Type locality. Itoman, Okinawa Island, Ryukyu Islands.

Material examined. — Chesterfield Islands, chalcal 1: stn DC 20, 19" 12' S. 158°42' E, 67
m, 6 dd. — Stn DC 31, 19°33'S. 158°30'E, 230 m, 1 dd. — Stn DC 33, 19°45'S, 158°26'E,
205 m, 2 lv, 10 dd. — Stn DC 35, 19°45' S, 158°26' E, 210 m, 2 dd. Stn DC 38, 20°00' S, 158°46' E,
250 m, 12 dd. — Stn DC 63, 22°11' S, 159°15' E. 305 m, 6 dd. — Stn DC 64. 22°12' S, 159°15' E,
305 m, 1 lv, 2 dd.

corail 2: stn DW 114, 19°25'S, 158°38'E. 217 m, ! dd. Stn DW 129, 19°28' S. 158°34'E.
215 m. 2 lv, 3 dd. — Stn DW 167, 19°46' S, 158°29' E, 270 m, 2 dd.

musorstom 5: stn DW 263, 25°2I' S, 159°46' E, 150-225 m, 9 dd. — Stn DW 265, 25°21' S, 159"45' E,
190-260 m, 3 lv, 5 dd. — Stn DW 266, 25"20' S, 159°46' E, 240 m, 1 lv, 6 dd. — Stn DW 270, 24°49' S
159°34'E, 223 m, 12 lv. — Stn DW 285, 24°09' S, 159°34'E, 245-255 m, 2 dd. — Stn CP 289
24"02' S. 159°38' E, 273 m, 1 dd. Stn DW 298, 22°44' S, 159°22' E, 320 m, 1 lv, 3 dd. — Stn DW

299, 22°48' S, 159°24' E, 360-390 m, 2 dd. Stn DW 302, 22°10' S. 159°23' E, 345-360 m, 3 dd.

Stn DW 303, 22° 12' S. 159"23' E, 332 m, I lv, 1 dd. — Stn CP 315, 22°25' S, 159°27' E, 330-335 m,
1 dd. Stn DW 328, 20°23' S, 158°44' E, 340-355 m, 1 lv, 4 dd. — Stn DW 329, 20°23' S, 158°47' E,
320 m, 1 lv, 9 dd. — Stn DW 334, 20°06' S. 158"48' E, 315-320 m, 1 lv, 1 dd. - Stn DW 344 19°39' S
158-34' E, 310 m, 2 dd. - Stn DW 346, I9"40'S, 158°27'E, 245-252 m, 6 dd. — Stn DW 349.
19°34' S, 158-34' E, 275 m, 1 lv. — Stn DC 376, 19°51' S, 158°30' E, 280 m, 6 dd.

Loyalty Islands, musorstom 6: stn DW 392, 20°47' S, 167°05' E, 340 m, 1 dd. — Stn DW 399
20-42' S, 167-00' E, 282 m, 2 dd. — Stn DW 417, 20°42' S, 167°04' E. 283 m, 3 dd. - Stn DW 418,’

20o42' S, 167-03' E, 283 m, 1 lv, 1 dd. — Stn DW 456, 2 POP S, 167°26' E, 240 m, 2 dd. — Stn DW

480, 21-08' S, 167-56' E, 380 m, 1 lv.

New Hebrides Arc. volsmar: stn DW 17, 22°23' S, 1 7 1 °4 1' E, 260-300 m, 4 dd.

Indonesia. “Snellius” II: stn 4.047, 09°53' S. 120°43' E, 100 m, 2 dd (rmnh).

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

281

Distribution. — Japan. Malaya, 50-200 in (Habe, 1964a), now extended to Indonesia and
New Caledonia, live records from 200 to 380 m.

Fustiaria vagina sp. nov.
Figs 71 h, 87, 88 i

Type material. Holotype mnhn. Paratypes: 8 mnhn, I ams C201726. 1 nmnz M268953,

1 USNM.

Type locality. - - New Caledonia, Chesterfield Islands, musorstom 5, stn DW 340, 1 9°49' S,
158°41' E, 675-680 m.

Fig. 87. — Distribution of Fustiaria vagina.

282

VICTOR SCARABINO

Material examined. - Chesterfield Islands, musorstom 5: stn DW 340, 19°49' S, 158°41' E,
675-680 m, 3 lv (holotype, 1 paratype ams). — Stn DW 341, 19°46'S, 158°43' E, 630-620 m, 1 lv
(paratype). — Stn DW 353, 19°27' S, 158°40' E, 290 m, 1 dd (paratype). Stn DW 357, 19°37' S,
158°46' E, 630 m, 1 lv (paratype).

New Caledonia, biocal: stn DW 46, 22°53' S. 167° 17' E, 570-610 m, 2 dd (paratypes). — Stn DW 49,
23°03' S, 167°32' E, 825-830 m, 3 dd (paratypes: 2 mnhn, 1 nmnz).
biogeocal: stn CP 290, 20°37' S, 167°03' E, 760-920 m, 1 dd (paratype).

Loyalty Islands, musorstom 6: stn DW 468, 21°06'S, 167°33' E, 600 m, 1 dd (paratype usnm).

Distribution. — New Caledonia, recorded alive in 630-680 m, shells from 290 to 820 m.

Description. — Shell to 30 mm, solid, white, glossy,
slightly curved, sculptured with growth lines only, conspi¬
cuous at posterior end. Apex truncate, oblique on ventral
side, with strong callus. Lumen longitudinally elongate, with
projecting irregular walls on ventral side. In section, walls are

wider on dorsal than on ventral side. Peristome oblique,
fragile. Section circular throughout.

Measurements: holotype L 29, W 2.8, w 1, arc 1; paratypes
range L 19-29, W 2-3.2, w 0.6-1. 5, arc 0.6-1. 5. W/w ratio 2.6.

Remarks. — Emerson (1962) explicity noted that the genus has varied apical structures. The
present species is provisionally placed in Fustiaria pending anatomical study.

Etymology. — Named for the resemblance of apical structure to the human female vagina.

Other species of Fustiaria cited in the literature

Fustiaria rubescens (Deshayes, 1825): 363, pi. 16, figs 23-24. Mediterranean Sea. mnhn. Cited
by Ludbrook (1954) as present off Zanzibar, " John Murray”, stn 103, 05°39' S, 39°11' E, 101 m. It
is doubtful that this Mediterranean species occurs in the Indo-Pacific and Ludbrook’s record should
be checked.

Family Gadilinidae Chistikov, 1975
Subfamily Gadilininae Chistikov, 1975

Genus Gadilina Foresti, 1895

Type species (by monotypy): Dentalium triquetrum Brocchi, 1814. Miocene, Italy.

Diagnosis. — Shell medium to large, well curved, solid, polished, shiny, white. Longitudinally
sculptured with two dorso-lateral ribs, more conspicuous at apex, slightly convex dorsally, strongly
concave ventrally. Apex truncate, with terminal callus, lumen circular. Section compressed dorsally,
circular ventrally.

Railula rachidian slightly curved in section, anterior margin irregular, three wide cusps ventrally;
lateral wide, with three primary cusps, the central quite pointed; marginal sinusoidal, with cusp in
anterior angle connecting with laterals.

Distribution. — Miocene-Recent, Pacific and Indian Oceans, absent in the Atlantic Ocean;
temperate-tropical belt, shelf to abyssal.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

283

Fig. 88. — a, Calliodenlalium semitracheatum, shell (68 mm) and apex, “ Mascareignes III": stn 78. — b, Calliodentalium
crocinum , shell (39 mm), apex, apical and medial sections, musorstom 3: stn CP 99. — c, Calliodenlalium balanoides ,
shell (65 mm) and apex, " Vauban " 1978-79: stn 40. — d, Calliodenlalium type radula ( Calliodenlalium callipeplum,
Caribbean Sea), left accessory denticle, relationship with the rachidian (see also Fig. 82 e). - e, Fustiaria nipponica, shell
(51 mm), apex and apical section. New Caledonia lagon: stn 214. — f, Fustiaria engischista, shell (41 mm), and apex,
Nosy-Be (bmnh). — g. Fustiaria caesura, shell (44 mm), apical views and apical section, musorstom 2: stn DR 34. —
h, Fustiaria langfordi, shell (49 mm), apex and apical section, musorstom 6: stn DW 417. — i, Fustiaria vagina sp. nov.,
paratype (26.4 mm), views of the apex. — j, Fustiaria type radula (F. caesura ); see also Fig. 78 d.

Source : MNHN, Paris

284

VICTOR SCARABINO

Gadilina insolita (Smith, 1894)
Figs 89, 90 a-f, 95 a-b

Dentalium insolitum Smith, 1894: 168, pi. 4. figs 1 7- 1 7a.

Synonym:

Dentalium stapes Boissevain. 1906: 50. pi. 5. figs 16-20, pi. 6, figs 79. 81. 83.

Other references:

Dentalium insolitum - Pilsbry & Sharp. 1897: 109, pi. 22, figs 56-57. — Smith. 1906a: 250. Boissevain. 1906: 49. pi. 5
fig. 15, pi. 6, figs 80-82, 84. — Winck worth, 1940a: 25.

Dentalium (Gadilina) insolitum - Plate, 1908a: 353. pi. 30, figs 50-51. — Jaeckel, 1932: 306. Ludbrook, 1954: 108.
Gadilina insoluta (sic) - Habe, 1964a: 32, pi. 2 fig. 23; pi. 4, figs 13-14. Habe & Kosuge, 1964: 7. Higo & Goto 1993-
688.

Dentalium (Gadilina) stapes — Ludbrook. 1954: 109.

Gadilina stapes - Habe, 1962: 105, pi. 47. fig. 6; 1963: 270. pi. 38. fig. 23. textfigs 13-14; 1964a: 32. pi. 2, fie. 23, pi 4 fies
13-14. — Okutani, 1964: 76. — Habe & Kosuge. 1964: 7.

O

O

•- ft
mr-p •

v«r

Fig. 89. Distribution of Gadilina insolita.

Type material. — D. insolitum: 2 syntypes dd bmnh 1952.3.25.83-93. D. stapes: svntypes
zma 3.06.061, 3.06.062.

Type locality. — D. insolita: Bay of Bengal, " Investigator ”, in 597 fms [1091 m], — D. stapes:
Indonesia, Banda Sea, " Siboga ”, stn 212, 05°55' S, 120°10' E, 462 m.

Material examined. — The type material.

New Caledonia, biocal: stn DW 56. 23°35' S, 167° 12' E, 695-705 m. 1 lv. — Stn DW 106 21°36' S
166°29' E, 625-650 m, 2 lv.

biogeocal: stn CP 232. 21°34' S. 166°27' E, 760-790 m, 2 lv, I dd.
lagon: stn 993, 20°15'S. 163°53' E, 375-400 m, 1 dd.

Indonesia, corindon: stn B 210, 00° 13' S. 1 17°53' E, 338 m, 1 dd. - Stn B 213. 00°3 1 ' N 117° 50' E
488 m, 1 dd.

' Snell ius II: stn 4.1 13, 08°19' S. 118°16' E (no depth data). I dd. - Stn 4.127, 08°19' S, 1 18°18' E
700-835 m. 2 lv, 1 dd. — Stn 4.128, 08°18' S, 1 18°16' E, 700-835 m, 2 lv, 1 dd. Stn 4.130, 08° 18' S,
1 1 8°1 8' E. 700-730 m, 2 lv. 1 dd. Stn 4.267. 08°18' S, 1 1 8°2 1 ' E. 650 m. 1 lv (rmnh).

Philippines, musorstom 1: stn CP 49, 13°49'N, I20°00' E, 750-925 m, I dd.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

285

musorstom 2: stn CP 25, 13°39' N, 120°43' E, 520-550 m, 1 dd.— Stn CP 50, 13°37' N, 120°33' E,
810-820 m, 13 dd, 9 lv. — Stn CP 51, 13°59' N, 120°16' E, 170-187 m, 1 lv. — Stn CP 55, 13°54' N,
1 19°58' E, 865 m, 6 lv. — Stn CP 78, 13°49' N, 120°28' E, 441-550 m, 2 lv, 6 dd. - Stn CP 81,
1 3°34' N, 120°31' E, 856-884 m, 3 lv. — Stn CP 82, 13°46' N, 120°28' E, 550 m, 1 lv, 2 dd.
musorstom 3: stn DR 94, 13°47'N, 120°03' E, 842 m, 1 dd. - Stn CP 106, 13°47'N, 120°30' E,
640-668 m, 1 lv, 2 dd. — Stn CP 118, 11°58' N, 121°06' E, 448-466 m, 3 lv, 6 dd. — Stn CP 122,
1 2°20' N, 1 2 1 °42' E, 219-220 m, 2 lv, 1 dd. - Stn CP 123, 12°10' N, 121°45' E, 700-702 m, 2 dd.

Stn CP 128, 1 1°50' N, 121° 42' E, 815-821 m, 5 dd.

Distribution. — Indonesia, Japan, Indian Ocean, now extended to New Caledonia, living
from 625 to 950 m (present paper), shells in 200-1134 in (Habe & Kosuge, 1964).

Remarks. — I concur with Habe (1964a) in considering this a single highly variable species
as examination of a large series shows clear gradation between forms.

Fig. 90. Radulae. a, Gadilina insolita. rachidian, view from the anterior border and internal face. — b, lateral internal
face. — c, lateral, head. — d, lateral, internal face. e, marginal, contact with lateral at left side. f, Gadilina insoliia
form stapes, general view. Scale bars: 100 (im (b-f), 10|am (a).

Other Indo-Pacific species of Gadilina cited in the literature

Gadilina pachypleura (Boissevain, 1906): 51. pi. 5, figs 21-22. "Siboga”, stn 208, 05°39' S, 122°12' E,
Banda Sea, 1886 m. zma.

286

VICTOR SCARABINO

Subfamily Episiphoninae Chistikov, 1975
Genus Episiphon Pilsbry & Sharp, 1897

Type species (SD by Suter, 1913): Dentalium sowerbyi Guilding, 1834. Recent, Caribbean Sea.

Diagnosis. — Shell small to medium, slightly curved, fragile but not thin, polished, shiny,
white, cream, orange or red. Sculptured apically with close, fine, encircled wrinkles; rarely
longitudinal striae near the apex or throughout. Section subcircular or subtriangular, slightly
compressed dorsoventrally, more on ventral side. Apex simple or truncate, with terminal callus,
lumen circular, small with short pipe.

Radula rachidian slightly curved in section, anterior margin irregular, nearly straight, three cusps on
internal face; lateral with few very sharp, pointed cusps; marginal sinusoidal, short.

Distribution. — Worldwide. Temperate-tropical waters, sublittoral-shelf.

Episiphon suhtorquatum (Fischer, 1871)

Figs 91, 95 c, f

Dentalium suhtorquatum Fischer, 1871: 212, pi. 11, figs 1-la.

Synonyms:

Dentalium annulosum Brazier, 1877: 58.

Dentalium tornatum Watson. 1879: 518; 1886: 13, pi. 2, fig. 3.

Dentalium (Episiphon) sewelli Ludbrook, 1954: 107, fig. 10 (Syn. nov.).

Other references:

mTlll1 suh'orq,,a!um ~ Pilsbry & Sharp, 1897: 101. — Boissevain, 1906: 57, pi. 2, fig. 27, pi. 6, fig. 36. Moazzo.

Dentalium ( Laevidentalium) suhtorquatum — Dell, 1964: 129.

Episiphon suhtorquatum - Habe & KosUGE, 1964: 8.

Plagioglypta suhtorquatum - Habe & Kosuge, 1964: 8.

Dentalium annulosum - Hedley, 1901: 129, pi. 17, fig. 36.

Episiphon tornatus (sic) - Habe & Kosuge, 1964: 6.'

Plagioglypta annulosum — Habe & Kosuge, 1964: 8.

Type material. — D. suhtorquatum-. lectotype (here designated: L 11.2, W 1.3, w 0.4) mnhn.
D. annulosum: syntype ams 189. — D. tornatum: 5 syntypes dd bmnh 87.2.9.57-60. — D. sewelli-
holotype bmnh 1952.3.25.125.

Type locality. — D. suhtorquatum: Suez. — D. anulosum: NE Australia, Princess Charlotte
Bay. — D. tornatum: Levuka, Fiji, 12 fms [22 m]. — D. sewelli: Gulf of Oman, " John Murray ”,
stn 75, 25° 10' N, 56°47' E, 201 m.

Material examined. — The type material and the material examined by Boissevain.

New Caledonia, lagon: stn 729, 21° 19' S, 165°54' E, 42-45 m, 2 lv.

West Indian Ocean, benthedi: stn S 18, 12°45' S, 45° 16' E, 15 m, 4 dd. — Stn S 23, 12°46' S, 45° 16' E,
6 m, 1 dd. — Stn S 32, 12°45' S, 47°18' E, 15-20 m, 1 dd. — Stn S 36, 12°52' S, 45° 16' E, 30 m]

4 lv. — Stn S 50, 12°55' S, 44°59' E, 32 m, 1 lv, 1 dd. — Stn DS 101, 1 1°26' S, 47°20' E, 26 m, 1 dd.
Shimont, Kenya, 1 lv (bmnh). — Kiloa, Zanzibar, 20 m, 1 lv, 2 dd. — Same locality, shore, 13 dd
(both mnhn). — Nosy Be Island, NW Madagascar, Plante coll., 2 lv, 28 dd. — Tulear. Madagascar

5 dd (both bmnh).

Red Sea. Suez, 107 dd. Gulf of Aden, Obock, Republic of Djibouti, 3 dd. — Hodeida, Yemen,
2 dd. Djibouti, I dd. — Aden, 5 dd (all Coll. Jousseaumf., mnhn).

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

287

Fig. 91. — Distribution of Episiphon subtorquaium.

Distribution. — Red Sea, Persian Gulf, and Zanzibar, 21-90 m (Ludbrook, 1954, as D.
sewelli ), Indonesia (Boissevain, 1906), North Australia and Fiji, now extended to Madagascar and
New Caledonia, alive in 20-45 m, shells down to 201 m.

Episiphon subrectum (Jeffreys, 1883)
Figs 92, 95 d

Denialium subrectum Jeffreys, 1883: 661.

Synonyms:

Denialium virgula Hedley, 1903: 328, fig. 62 (Svn. nov.).

Denialium carneum Boissevain, 1906: 48, pi. 6, figs 44-45 (Syn. nov.).

Denialium makiyamai Kuroda & Kikuchi, 1933: 11, pi. 5, fig. 8.

Other references:

Denialium subrectum - Pilsbry & Sharp, 1897: 120, pi. 8, fig. 5. — Boissevain, 1906: 47, pi. 6, figs 46-50. — Ludbrook,
1954: 108. — Habe & Kosuge, 1964: 6. — Habe, 1964a: 28, pi. 2, fig. 1.

Episiphon carneum — Habe & Kosuge, 1964: 6.

Episiphon makiyamai — Habe, 1963: 267, pi. 38, fig. 1. — Tsuchida el at., 1991: 14, pi. 3, fig. 16.

Type material. — D. subrectunr. syntypes usnm {fide Ludbrook, 1954) [not located]. — D.
virgula : holotype bmnh 1913.4.30.10-14, paratypes ams C162217. — D. carneum : lectotype (here
designated) zma 3.06.052, paralectotypes zma 3.06.053, 073, 055.

Type locality. — D. subrectum: Philippines. — D. virgula: off Port Kembla, NSW, Australia,
41-50 fms [80-91 m], D. carneum: Indonesia, Flores Sea, "Siboga”, stn 45, 07°24' S, H8°15'E,
794 m. - D. makiyamai: Toyama Bay, Japan.

Material examined. — The type material of D. virgula and D. carneum. Material identified
by Boissevain, 1906 (zma) and by Ludbrook, 1954 (bmnh 1952.3.25.78-82).

Philippines, musorstom 2: stn DR 34, 13°28'N, 1 2 1 ° 1 2' E, 155-167 m, 14 dd.

musorstom 3: stn DR 140, 1 1°43' N, 122°34' E, 93-99 m, 3 dd. — Stn CP 143, 1 1°29' N, 124°1 1 E,

205-214 m, 1 dd.

288

VICTOR SCARABINO

Distribution. — NW Indian Ocean, Indonesia, Philippines, Japan, SE Australia. Shells in
54-900 m.

Epi siphon virginieae sp. nov.

Figs 93, 95 e

Type material. — Holotype lv, mnhn. Paratypes dd: 9 mnhn, 1 ams C201727, 1 nmnz
M268954.

Type locality. — New Caledonia, Loyalty Islands, musorstom 6, stn DW 399, 20°42' S,
1 67°00' E, 282 m.

Material examined. — Only known from the type material.

Distribution. — Only known from the type locality.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

289

Description. — Shell to 9 mm long, translucent, fragile,
slender, slightly curved. Fine longitudinal sculpture of close,
minute lines throughout, noticeable under magnification.
Apex truncate, subcircular in section, lumen subcircular with
short pipe. Mouth suboval, straight.

Measurements: holotype L 8.6, W 0.9, w 0.7, arc 0.3;
paratypes L 8, W 0.8. w 0.4, arc 0.6; L 7.7, W 0.9, w 0.7, arc
0.3; L 8.4. W 0.8, w 0.5, arc 0.5; L 8.3. W 0.8, w 0.4, arc 0.3;
L 9.4, W 0.8, w 0.4, arc 0.5; L 7.7, W 0.9. w 0.6, arc 0.3. W/w
ratio 1.3-2.

Remarks. — The longitudinal sculpture throughout the shell is unique in the genus.

Etymology. — Named for Virginie Heros, mnhn, who processed much of the musorstom
material, and who has assisted in many ways with this report.

Other Indo-Pacific species of Episiphon cited in the literature

Episiphon candelatum (Kira, 1959): 105, pi. 40, fig. 5. Japan. Tosa Bay, Shikoku, 200 m.

Episiphon gazellae (Plate, 1908): 356, pi. 30, figs 40-41. Northwest Australia, 16 m. Holotype zmb
33195. Generic allocation uncertain.

Episiphon minutissimum Ludbrook, 1954: 108, fig. 11. Maidive area, “John Murray", stn 147,
04°53' N, 72°54' E, 27 m. Holotype and 9 paratypes bmnh 1952. 3.25.113-123. Generic
allocation uncertain.

Episiphon truncation (Boissevain. 1906): 51, pi. 6, fig. 33, textfig. Indonesia. Banda Sea, “Sihoga",
stn 90, 01°17'N, 118°53' E, 281 m. zma.

Fig. 94. — Distribution of Anutidenialium bambusa.

Subfamily Anulidentaliinae Chistikov, 1975

Genus Anulidentalium Chistikov, 1975

Type species (by monotypy): A. bambusa Chistikov, 1975.

Diagnosis. — Shell large, slightly curved to almost straight, thin, narrow, fragile, polished,
translucent white. Sculpture of encircling swellings at regular intervals. Apex simple. Section
subcircular, slightly compressed laterally.

290

VICTOR SCARABINO

Radula rachidian slighttly curved in section, with three central cusps on inner face; lateral with
prominent sharp primary cusp and two secondary cusps placed laterally to each other; marginal
short, sinusoidal.

Distribution. — Recent, western Pacific Ocean. Shelf-bathyal.

Anulidentalium bambusa Chistikov, 1975
Figs 94, 95 g

Anulidentalium bambusa Chistikov, 1975: 21.

Other reference:

Anulidentalium bambusa — Chistikov, 1979b: 112, fig. 4.

Type material. — zin (fide Chistikov, 1979) [not seen].

Type locality. — Gulf of Tonking, Viet Nam, 72 m.

Fig. 95. — a, Gadilina insolita, shell (49 mm), apex, apical, medial and oral sections, musorstom 2: stn CP 50. — b, Gadilina
insolita form stapes , shell (61 mm) apex, apical and oral sections, musorstom 2: stn CP 78. — c, Episiphon sublorqualum ,
shell (25 mm), apex and apical section, benthedi: stn S 36. — d, Episiphon subrectum, shell (14 mm), apex and apical
section, musorstom 2: stn DR 34. — e, Episiphon virginieae sp. nov., holotype, shell (8.6 mm), detail of sculpture and
apical section. — f, Episiphon type radula (E. subtorquatum). — g, Anulidentalium bambusa, shell (52 mm), apical
section, musorstom 3: stn CP 139; Anulidentalium type radula (A. bambusum ) (after Chistikov, 1979b).

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

291

Material examined. — Indonesia. “Snellius” II: stn 4.155, 06°22' S, 120°26' E, 233-274 m,

1 lv.

Philippines, musorstom 3: stn CP 139, 1 1°53' N, 122° 14' E, 240-267 m, 1 lv, 6 dd.

Distribution. — Gulf of Tonking, now extended to the Philippines and Indonesia. Alive from
72 m (Chistikov, 1979b) to ca. 270 m (present paper).

Family Laevidentaliidae Palmer, 1974

Genus Laevidentalium Cossmann, 1888
Type species (OD): Dentalium incertum Deshayes, 1825. Eocene. France.

Diagnosis. — Shell medium to large, slightly to well curved, thin but not fragile, shiny,
translucent white to cream. Sculpture usually fine longitudinal undulations and conspicuous growth
lines; in some species, slight swellings at regular intervals. Apex simple or with terminal callus and
projectig pipe cylindrical or fissured, lumen distinct. Section circular to subcircular.

Radula rachidian very slightly curved at central portion in transverse section; an irregular zone is
present in the center; in frontal view, the anterior margin is irregular with a high central projection.
Lateral head not well differentiated, no cusps, anterior margin finely undulated or granulose;
marginal long, slightly sinusoidal.

Distribution. — Triassic- Recent, worldwide, sublittoral-bathyal.

Laevidentalium eburneum (Linne, 1767)
Figs 96, 102 a

Dentalium eburneum Linne, 1767: 1264.

Synonyms:

Dentalium indicum Chenu, 1843: 1, pi. 3, fig. 11.

Dentalium philippinarum Sowerby, I860: 98, pi 225 ( Dentalium 3), fig. 54.

Other references:

Dentalium eburneum - Gmelin, 1791: 3737. — Sowf.rby. 1860: 98. pi. 225 ( Dentalium 3), fig. 53; 1873: pi. 3. fig. 16. —
Martens, 1887: 200. — Pilsbry & Share, 1897: 115, pi. 20, figs 33-34. — Smith. 1906a: 250. - Boissevain, 1906: 52,
pi. 2, fig. 31, pi. 4, figs 10-11. — Hedley, 1916: 223. — Winckworth, 1940a: 25. Dawidoff, 1952: 144. — Roberts,
Soemodih ardjo & Kastoro, 1982: 99.

Calliodentalium eburneum - Chistikov, 1979: 112.

Laevidentalium eburneum - Habf. & K.OSUGE, 1964: 7.

Dentalium philippinarum - Sowerby, 1872: pi 3, fig. 18. — Pilsbry & Sharp, 1897: 116. pi. 20, figs 31-32. — Boissevain,
1906: 53. - Habe, 1963: 269, pi. 38, fig. 33; 1964a: 34, pi. 2, fig. 33.

Laevidentalium philippinarum — Higo & Goto, 1993: 687.

Type material. — D. eburneum-. depository not located. — D. indicum: depository not
located. — D. philippinarum: depository not located.

Type locality. — D. eburneum: India. — D. indicum: Indian Ocean. — D. philippinarum:
Island of Samar, Philippines.

Material examined. — Indonesia, corindon: stn B 202, 01°1 1' S. 117°06' E, 27 m, 7 dd. —
Stn CH 205, 01°08' S, 1I7°19'E, 49 m, 1 dd. — Stn B 207, 00°15' S. 117°52'E, 150 m, 1 lv, 1 dd.
— Stn DR 216, 00°04' N, 1 1 7°5 1 ' E, 96 m, 1 dd. — Stn B 251, 00°54' S, 1 19°30' E, 65 m, 2 dd. -

Source . MNHN, Paris

292

VICTOR SCARABINO

Stn B 253, 00°54' S. 1 19°30' E, 17 m, 15 dd. — Stn DG 254, 00°58' S, 1 19°29' E, 53-62 m, 1 lv. -
Stn DG 258, 01°57' S, 119°17' E. 30 m, 1 dd. — Stn B 268, 01°57' S, 119°16' E, 200 m, 1 lv, 1 dd.
Philippines, musorstom 2: stn DR 33, I3°32' N, 121°07' E, 130-137 m, 2 dd.
musorstom 3: stn DR 140, 11°43' N, 122°34' E, 93-99 m, 18 dd. — Stn CP 141, 11°45' N, 122"44 E,
44 m, 1 lv.

Distribution. — Indian Ocean, China Seas, 75 m (Dawidoff, 1952), Malaya, Indonesia,
Thailand, Philippines, 10-100 m (Habe & Kosuge, 1964). Live records from 44 to 200 m (present
paper).

Laevidentalium leptosceles (Watson, 1879)
Figs 97, 102 b

Dentalium leptosceles Watson, 1879: 513.

Synonym:

Dentalium banale Boissevain, 1906: 55. pi. 6, fig. 30 (Syn. nov.).

Other references:

Dentalium leptosceles - Pilsbry & Sharp, 1897: 110, pi. 3, figs 44-46.

Dentalium leptosceles (?) (sic) - LuDBROOK, 1954: 104. fig. 6.

Dentalium leptoskeles (injustified emendation) - Watson, 1886: 7, pi. 1, fig. 6.

Laevidentalium banale — Habe & Kosuge, 1964: 7.

Type material. — D. leptosceles : holotype bmnii 1887.2.9.21. — D. banale : holotype zma
3.06.068.69.

Type locality. — D. leptosceles: S. of Australia, “ Challenger ", stn 160, 42°42' S, 134°10' E,
2600 fms [4758 m], — D. banale : Indonesia, Timor Sea, “ Siboga ”, stn 300, 10°49' S, 123°23' E,
918 m.

Material examined. — The type material.

New Caledonia, biocal: stn DS 59, 23°56' S. 166°4T E, 2650 m, 1 lv.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

293

biogeocal: stn CP 250, 21°25' S, 166°26' E, 2350 m, 1 dd. — Stn KG 276, 21°13'S, 167°00'E.
2200 m, 1 lv.

Tasman Sea. “ Galathea stn 574, 39°45' S, 159°39' E, 4680-4730 m, 1 dd.

Indian Ocean, safari 1: stn DS 05, 30°37' S, 48°30' E, 4500-4612 m, 2 lv. — Stn CP 06, 30°40' S,
48° 14' E, 4020-4035 m, 1 lv, 1 dd. — Stn CP 09, 30°49' S, 49°08' E, 4589-4730 m, 3 lv, 3 dd. — Stn
CP 17, 24°26' S, 58° 19' E, 4987-5025 m, 1 lv. — Stn SIPAN 7909, 30°4T S, 48°28' E. 4462 m, 1 dd.
safari 2: stn CP 13, 04°30' S, 86°55' E, 4950 m, 1 dd. — Stn CP 18, 06°02' S, 79°32' E, 5175 m,
1 lv. - Stn CP 29, 12°57' S, 79°37' E, 4928-4950 m, 1 dd. — Stn CP 31, 13°45' S, 76°56' E, 5300 m,
1 lv, 2 dd.

“Galathea": stn 194, 34°09' S, 30°45' E, 4360 m, 1 dd. — Stn 234, 05°25' S, 47°09' E, 4830 m, 1 dd.

Distribution. — Widely distributed at bathyal and abyssal depths in the Indian Ocean. Timor
Sea, Tasman Sea and New Caledonia. Recorded alive between 2200 and 5300 m (present paper).
Shells from 918 m (Boissevain, 1906. under D. hanale).

Laevidentalium coruscum (Pilsbry, 1905)

Figs 98, 102 c, f

Dentalium ( Laevidentalium ) coruscum Pilsbry, 1905:117, pi. 5. figs 42-43.

Other references:

Dentalium ( Laevidentalium ) coruscum - Kuroda & Kikuchi. 1933: 10. pi. 1, fig. 7.

Laevidentalium coruscum — Habe, 1963: 268, pi. 38. figs 15, 29.

Gadilina coruscum - Habe, 1964a: 32, pi. 2, figs 15, 26, 29.

Type material. — Syntype ansp.

Type locality. — Heda, Izu, Japan, 306 m.

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DC 376, I9°5TS, 158°30'E, 280 m, 1 dd.

Indonesia. “Snellius ” II: stn 4.127, 08°19'S, 118°18'E, 500-550 m, 1 dd. — Stn 4.267, 08°18'S,
1 1 8°2 1 ' E, 650 m, 2 dd.

294

VICTOR SCARABINO

Philippines, musorstom 2: stn CP 66, 1 4°0 1 ' N, 120°20' E, 192-209 m, 1 lv, 1 dd. — Stn CP 75,
13°50'N, 120°30'E, 300-330 m, 3 lv, 3 dd. — Stn CP 78, 13°49'N, 120°28' E, 441-550 m, 1 lv,
19 dd.

musorstom 3: stn CP 99, 14°01' N, 120°19' E, 196-204 m, 1 dd. — Stn CP 1 19, 12°00' N, 121°13' E,
320-337 m, 4 dd. — Stn CP 139, 1 1°53' N, 122°15' E, 240-267 m, 5 dd.

Fig. 98. Distribution of Laevidentalium coruscum.

Distribution. — Japan and Indonesia (Habe, 1964) now extended to the Philippines and New
Caledonia. Alive in 192-550 m.

Remarks. — Laevidentalium coruscum resembles L. eburneum, but it is white, shiny, more
slender and possesses apical callus.

Laevidentalium gofasi sp. nov.

Figs 99, 102 e

Type material. — Holotype mnhn. Paratypes: 8 mnhn, 1 ams C201728, 1 usnm.

Type locality. — Philippines, musorstom 2, stn CP 70, 14°01' N, 120°17'E, 191 m.

Material examined. — Loyalty Islands, musorstom 6: stn DW 481, 21°22' S, 167°50' E,
300 m, 1 lv.

Philippines, musorstom 2: stn CP 20, 14°00' N, 120°18'E, 185-192 m, 1 dd (paratype usnm). —
Stn CP 68, 14°01'N, 120° 18' E, 195-199 m, 2 lv (paratypes). — Stn CP 70, 14°01' N, 120° 17' E,
191 m, 2 dd (holotype and paratype). — Stn CP 83, 13°55'N, 120°30' E, 318-320 m, 1 lv (para-
type).

musorstom 3: stn CP 96, 14°00'N, 120°18'E, 190-194 m, 1 dd (paratype ams). — Stn CP 97,
14°00' N, 120° 18' E, 189-194 m, 1 dd (paratype). — Stn CP 102, 14°01' N, 120° 18' E, 192 m, 3 dd
(paratypes).

Distribution. — The Philippines and New Caledonia, live records from 195 to 320 m.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

295

Description. — Shell to 80 mm long, slender, chalky-
white. slightly translucent, polished, moderately to well
curved (2 to 6 mm for 27-56 mm length). Surface and sides
irregular due to encircling wrinkles of varying size and
position, prominent growth lines with associated white bands
and longitudinal threads, apparent under magnification,
more conspicuous on dorsal side. Section oval, slightly
laterally compressed. Apex truncate, simple, strong.

Measurements: holotype L 50, W 1.9-1. 7, w 0.53-0.50, arc
6; paratypes L 51.3. W 1.9-1. 8, w 0.52-0.50. arc 4.5; L 38.6.
W 1.7-1. 6, w 0.4-0. 3. arc 4; L 56.5, W 1.8- 1.7, w 0.6-0.5, arc
6; L 73.5. W 2.7-2.6, w 0.7-0.6. arc 4; L 44. W 1.7-1. 6, w
0.6-0.52, arc 2.5. W/w ratio 2.8-4.2.

Remarks. — Compared with Laexidentalium houbricki , L. gofasi is stronger, has slightly oval
section and longitudinal sculpture.

Etymology. Named for Dr Serge Gofas, mnhn.

Laevidentalium houbricki sp. nov.

Figs 100, 102 d

Type material. — Holotype mnhn. Paratypes: 7 mnhn, 1 ams C201729, 1 nmnz M268955,

1 USNM.

Type locality. New Caledonia, Poindimie area, lagon, stn 835, 20°47' S, 165°17' E,
135-150 m.

Material examined. — New Caledonia, lagon: stn 835, 20°47' S, 165°17' E, 135-150 m. 1 lv
(holotype), 2 dd (paratypes). — Stn 858, 20°37' S, 165°07' E, 220 m. 10 lv (8 paratypes: 5 mnhn,
1 AMS, 1 NMNZ, 1 USNM).

Passe de Boulari, B. Richer/ORSTOM coll., 400 m, 7 dd.

Indonesia, corindon: stn CH 208, 00°15' S, 117°52' E, 150 m, 1 dd.

Distribution. — Makassar Strait, Indonesia, and New Caledonia, live records in 135-220 m.

Description. — Shell to 67 mm, slender, solid, white,
polished, very slightly curved. Transversal sculpture consis¬
ting of close lines probably corresponding to grow th periods.
Longitudinal sculpture of undulating threads throughout,
observable under magnification. Section slightly compressed

laterally. Apex truncate, without notch, strongly walled,
lumen circular. Mouth straight.

Measurements: holotype L 60, W 2.3, w 0.8, arc 0.8;
paratypes L 66.6. W 2.4, w 0.7, arc 1.2; L 34.6. W 1.8, w 0.6,
arc. 1. W/w ratio 2. 9-3. 4.

296

VICTOR SCARABINO

Etymology. — Named for the late Dr Richard S. Houbrick, usnm, Smithsonian Institution.

Other Indo-Pacific species of Laevidentalium cited in the literature

Laevidentalium bisinuatum (Andre, 1896): 397, pi. 17, fig. 9a-c. Amboyna [= Ambon]. Indonesia.

Holotype mhng 1155/40. Generic allocation uncertain.

Laevidentalium pluteum Colman, 1958: 143, fig. 8. Off Willongong, New South Wales, Australia,
183 m. Holotype ams Cl 8217. Generic allocation uncertain.

Laevidentalium sominium Okutani, 1964: 75, fig. 3. Sagami Bay, Japan, 1320-1400 m.
Laevidentalium toyamense (Kuroda & Kikuchi, 1933): 11, pi. 1, figs 5-6. Toyama Bay, Honshu, Japan,
200 m.

Family Omniglyptidae Chistikov, 1975

Genus Ommglypta Kuroda & Habe in Habe, 1953

Type species (OD): Dentalium cerinum Pilsbry, 1905.

Diagnosis. — Shell large, slightly curved, narrow, fragile, translucent, light orange in fresh
specimens, cream when eroded. Sculptured with close, fine, regularly spaced encircling wrinkles
throughout. Apex simple, section circular.

Radula rachidian slightly curved in section, anterior margin and internal face with irregular
projections; lateral irregular, head poorly defined, cusp fiat, anterior margin finely undulated;
marginal long; nearly straight.

Distribution. — Recent, West Pacific and Indian Oceans, absent in the Atlantic Ocean.
Shelf-abyssal.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

297

Omniglypta cerina (Pilsbry, 1905)
Figs 101, 102 g-h

Dentalium (Rhabdus) cerinum Pilsbry, 1905: 117, pi. 5, figs 40-41.

Synonym:

Dentalium iracheatum Boissevain, 1906: 56, pi. 4, fig. 22 (Syn. nov.).

Other references:

Omniglypta cerina - Habe, 1953: 296, figs 753-754; 1955: 24, figs 1-2; 1962: 106, pi. 47, fig. 13; 1963: 270, textfigs 23-24; 1964a:
37, pi. 4. figs 23-24; 1971: 492 (Japanese text): 310 (English text), pi. 65, figs 24-25; 1977: 337, pi. 70, figs 5-6, pi. 72, fig. 9.

Habe & Kosuge, 1964: 7. — Okutani, 1983: 12, pi. 43, fig. 14. — Habe el al. , 1986: 24. — Higo & Goto, 1993:
688.

Dentalium ( Rhabdus ) cerinum - Clench & Turner, 1962: 29.

Dentalium ( Plagioglypta ) iracheatum — Plate, 1908a: 357.

Type material. — D. cerinum: holotype ansp 88305. — D. iracheatum : lectotype (here
designated) the specimen illustrated by Boissevain (1906), zma 3.06.076, paralectotypes zma 3.06.075,

077-080.

Type locality. — D. cerinum: Shimizu, Suruga Bay, Japan. — D. Iracheatum: Indonesia,
Banda Sea, " Siboga ”, stn 208, 05°39' S, 122°12'E, 1886 m.

Material examined. — The type material.

Indonesia. " Snellius ” II: stn 4.130, 08°18' S, 118°18'E, 700-730 m, (dd). — Stn 4.131, 08°18'S,
11 8° 18' E, 680-800 m, 6 dd.

Philippines, estase 2: stn CP 6, 04°38' N, 1 19°49' E, 2570 m, 2 dd. — Stn DR 5, 04°58' N, 125°19' E,
3925 m, 1 lv.

musorstom 3: stn DR 115, 12°32'N, 120°44'E, 794 m, 1 lv.

Fig. 101. — Distribution of Omniglypta cerina.

Distribution. — Japan and Indonesia, now the Philippines. Habe & Kosuge (1964) give the
depth range as 0-1886 m, but this species appears clearly to be a deep-water species, with live records
in 800-3925 m (present paper).

298

VICTOR SCARABINO

Family Rhabdidae Chistikov, 1975

Genus Rhabdus Pilsbry & Sharp, 1897
Figs 102 i-j

Type species (OD): Denialium reclius Carpenter, 1864. Recent, Northeastern Pacific.

Diagnosis. — Shell medium to large, almost sraight, fragile, translucent. Sculpture lacking,
cross section circular, apex and mouth simple. The pavilion secretes a secondary tube that cannot be
distinguished from the primary shell. The annular ciliary organ of the anterior mantle margin is
lacking. Instead, a pair of dorsolateral slits with ciliated walls is present. Bundles of longitudinal pedal
muscles keep the pedal ganglia in position. The ligament of the buccal septum is missing (Steiner,
1992a).

Radula rachidian slightly curved in section, anterior border irregular, with a prominent central
projection and two projections at each side that not reach the border; laterals with wide granulose
head, prominent cusp and wrinckles instead of denticles; marginals almost straight.

Distribution. — Worldwide, temperate to cold waters, shelf-bathyal.

The genus Rhabdus is not represented in the material reported on here.

Order Gadilida Starobogatov, 1974
Suborder Entalimorpha Steiner, 1992
Family Entalinidae Chistikov, 1979
Subfamily Entalininae Chistikov, 1979

Genus Entauna Monterosato, 1872
Type species (OD): Denialium letragonum Brocchi, 1814. Miocene, Italy.

Diagnosis. — Shell medium to large, arched, solid, usually polished, translucent white when
fresh, chalky when dead. Sculptured by 4-5 primary ribs, secondary riblets present. Rib section flat
to rounded, simple or bifurcated. Intercostal spaces straight to convex, smooth or longitudinally
sculptured. Apex simple. Section usually pentagonal at the apex; pentagonal or quadrate at mouth.
Radula rachidian high, narrow, anterior margin usually rounded, but occasionally irregular; lateral
high, with two lateral primary cusps, lateral denticles present, including 5-6 subequal secundary
denticles; marginal sinusoidal.

Distribution. — Miocene-Recent. Worldwide. Sublittoral-bathyal.

Fig. 102. — a, Laevidentalium eburneum, shell (58 mm), apex and apical section, corindon: stn DG 258. — b, Laevidentalium
leplosceles, shell (33 mm), apex and apical section, detail of sculpture, safari: stn CP 09. — c, Laevidentalium coruscum,
shell (43 mm), apex and apical section, musorstom 3: stn CP 139 . — d, Laevidentalium houbricki sp. nov., holotype,
shell (60 mm), apical, posterior 1/4 and oral sections. — e, Laevidentalium gofasi sp. nov., holotype, shell (50 mm), apex,
apical and oral sections. - f, Laevidentalium type radula (L. coruscum). — g, Omniglypta cerina, shell (62 mm), detail
of sculpture and oral section, estase 2: stn CP 6. — h, Omniglypta type radula (O. cerina). — i, Rhabdus perceptum,
(Southern Argentina; Montevideo Museum), shell (60 mm), lateral and dorsal views. j. Rhabdus type radula (R.
perceptum).

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

299

Source : MNHN . Paris

300

VICTOR SCARABINO

Entalina mirifica (Smith, 1895)
Figs 103, 107 a, c

Dentalium mirificum Smith, 1895: 9, pi. 2, fig. 1.

Synonyms:

Entalina quadrangularis Boissevain, 1906: 62, pi. 6, figs 73-75, 85-86 (Syn. nov.).

Entalina majestica Kira, 1959: 105, pi. 40, fig. 3.

Other references:

Dentalium mirificum - Winckworth, 1940a: 25.

Entalina mirifica — Pilsbry & Sharp, 1897: 134. pi. 20, fig. 29. — Boissevain, 1906: 62, pi. 2, fig. 37. — Ludbrook, 1954:
111, fig. 15. — Chistikov, 1982c: 1496, pi. 2, figs 2-3.

Entalina (E.) mirifica - Habe & Kosuge, 1964: 8.

Entalina quadrangularis — Plate, 1908a: 358. — Habe, 1964a: 39. pi. 3, fig. 6, pi. 4, figs 20-21. — Okutani, 1966: 13. — Habe.
1977: 339, pi. 71. figs 1-2. — Chistikov, 1982c: 1496. — Habe et ai. 1986: 24. “Higo & Goto, 1993: 689.

Entalina (E.j quadrangularis — Habe, 1964b: 8.

Entalina platamodes - Boissevain, 1906: 62, pi. 2, fig. 38, pi. 6. figs 76-78.

Entalina quadriangularis (sic) - Habe, 1963: 271, textfigs 20-21. — Okutani, 1964: 78, pi. 6, fig. 5; 1966: 13.

Type material. — D. mirificum : 4 syntypes dd bmnh 1895.7.2.26. — E. quadrangularis :
lectotype (here designated) zma 3.06.081 and paralectotypes z.ma 3.06.082.083.

Type locality. — D. mirificum: Indian Ocean, off Trincomalee, “Investigator”, stn 172,
200-350 fms [357-640 m], — E. quadrangularis : Indonesia, Celebes Sea, “ Siboga ”, stn 88, 00°35' N,
119°09' E, 1301 m. — E. majestica : Japan, 30-50 fms [55-91 m].

Material examined. — The type material listed above.

New Caledonia, biogeocal: stn DW 253, 21°32' S, 166°29' E, 310-315 m, 1 dd.

“ Vauban ” 1978-79: stn 40, 22°30' S, 166°24' E, 250-350 m, 7 lv, 10 dd.

Tasman Sea. “Galathea”: stn 554, 37°28' S, 151°51' E, 1330 m, 5 lv, 4 dd.

Indonesia, corindon: stn B 236, 00°07' N, 119°45' E, 173 m, 2 dd.

“ Snellius ” II: stn 4.112, 08°19' S, 118°16'E, 365 m, 5 dd. — Stn 4.128, 08°18'S, 118°16'E, 700-
835 m, 1 dd. — Stn 4.130, 08°18'S, 118°18'E, 700-730 m, 1 dd. — Stn 4.131, 08°18' S, 118°18'E,
680-800 m, 4 dd.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PAC1FIC

301

Philippines, estase 2: stn CP 2, 14°05' N, 120°02' E, 2050 m, 8 lv, 4 dd.

MUSORSTOM 2: stn CP 18, 14°00' N, 120°18'E, 188-195 m, 1 lv, 1 dd. - Stn CP 25, 13°39'N,
1 20°43' E, 520-550 m, 1 lv. — Stn CP 50, 13°37' N, 120°33' E, 810-820 m. 2 lv, 2 dd. — Stn CP 55,
13°54' N, 1 19°58' E, 865 m, 2 lv, 5 dd.

musorstom 3: stn CP 106, 13°47' N, 120°30' E, 640-668 m, 2 dd. Stn CP 1 18, 1 1°58' N, 121°06' E,
448-466 m, 1 dd.

Distribution. — Indonesia (Boissevain, 1906), Northern Indian Ocean, 55-1300 m (Habe &
Kosuge, 1964 as E. quadrangular is) as far West as Zanzibar and the Gulf of Aden (Ludbrook, 1954).
Now extended to the Philippines and the South Pacific, live records in 195 — 2050 m.

Remarks. — The shape of the oral section was the considered the main difference between
Entalina mirifica and E. quadrangular is. When a large series of individuals is examined, the variability
of this character indicates the weakness of the former specific distinction. I agree with Japanese
authors in considering E. majestica also a junior synonym of E. quadrangularis.

Entalina subterlineata (Tomlin, 1931)

Figs 104, 107 b

Denialium subterlineatum Tomlin, 1931: 337.

Other references:

Denialium subterlineatum — Barnard, 1963b: 348, fig. 30a.

Entalina subterlineata - Chistikov, 1982c: 1497.

Type material. Holotype sam A6192, paratype nmw (fide Oliver, 1984).

Type locality. Off South Africa, “Cape Point 17°85' E, 43 mi, 900 fms” [1645 m].

Material examined. — The holotype.

South Africa. “Meiring Naude": stn SM 94, 28°16' S, 32°29' E, 670 m, 2 dd. — Stn SM 123, 30°33' S,
30°49' E, 690 m, 2 lv. — Stn SM 129, 30°53' S, 32°31' E, 850 m, 16 lv, 20 dd. — Stn SM 131, 30°43' S,
30°40' E, 780 m, 1 dd. — Stn SM 184, 33°39' S, 27°11' E, 86 m, 6 lv, 10 dd.

Fig. 104. Distribution of Entalina subterlineata.

302

VICTOR SCARABINO

Distribution. Endemic to South Africa, recorded alive from 690-860 m, shells down
1645 m.

Other Indo-Pacific species of Entalina cited in the literature

Entalina adenensis Ludbrook, 1954: 112, fig. 17. Gulf of Aden, " John Murray", stn 185, 13°48' N,
16"48' E, 2000 m. Supposedly in bmnh, not located.

Entalina inaequisculpta Ludbrook, 1954: 111, fig. 16. Gulf of Aden, “John Murray”, stn 185,
13°48' N, 1 6°48' E, 2000 m. Supposedly in bmnh, not located.

Subfamily Heteroschismoidinae Chistikov, 1982

Genus Heteroschismoides Ludbrook, 1960
Figs 107 d-e

Type species (OD): Dentalium subterfissum Jeffreys, 1877. Recent, North Atlantic Ocean.

Diagnosis. Shell medium, slightly curved, regularly tapering, translucent when fresh,
milky-white when dead. Longitudinal sculpture of 10-12 primary prominent ribs, intercostal spaces
convex. Apex with a deep irregular fissure on dorsal side. Section polygonal, slightly compressed
laterally, oral aperture thin.

Radula rachidian with anterior margin rounded and lateral half folded to the ventral side; lateral with
sharp pointed primary cusps and 4 important denticles; marginal slightly curved with conspicuous
lateral processus.

Distribution. — Recent, North Atlantic Ocean, bathyal-abyssal.

Remarks. As in Spadentalina, the fissure is observed since the embryonic stage of the shell
(Fig. Ill g).

The genus Heteroschismoides has not been found in the Indo-Pacific region.

Genus Costentauna Chistikov, 1982

Type species (OD): C. elegans Chistikov, 1982. Recent, Indian Ocean.

Diagnosis. — Shell small to medium, slightly to well curved, fragile, translucent white when
fresh, opaque to polished when dead. Longitudinal sculpture of 10-12 prominent primary ribs;
intercostal spaces convex. Apex simple, truncate, irregular, preapical callus prominent, lumen
circular. Section polygonal, oral aperture thin.

Radula rachidian high, polygonal with rounded anterior margin and lateral half folded to ventral side;
lateral well armed, with sharp, pointed primary cusps; marginal slightly curved (Fig. 107 h: C. vemae
Scarabino, 1986).

Distribution. — Recent, worldwide, bathyal-abyssal.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

303

Costentalina tuscarorae Chistikov, 1982
Figs 105, 107 f, 111 h, k

Costentalina tuscarorae Chistikov, 1982b: 1316, pi. 1, figs 7-1 1, pi. 4, figs 1-2.

Type material. — zin ( fide Chistikov, 1982).

Type locality. — NW Pacific, “ Vitiaz ”, stn 3575, 38°02’1 N, 146°33’1 E, 5475 m.

Material examined. — Philippines, estase 2: stn CP 2, 14°05' N, 120°02' E, 2050 m, 3 lv,
2 dd. — Stn DW 1, 14°05' N, 120°0T E, 2200 m, 2 lv, 2 dd.

Distribution. — East of Japan and the Philippines. Live records from 2050 m (present paper)
to 5475 m (Chistikov, 1982).

Costentalina indica Chistikov, 1982
Figs 106, 107 g

Costentalina indica Chistikov, 1982b: 1318, pi. I, figs 12-14, pi. 4, figs 4-7.

Type material. — zin (fide Chistikov, 1982).

Type locality. — Indian Ocean, "Vitiaz", stn 4922, 06°54’2 S, 83°00'7 E, 3980 m.

Material examined. — West Indian Ocean, benthedi: stn CH 87, 1 1°44' S, 47°35' E, 3716 m,
8 lv, 5 dd. — Stn CH 90, 1 1°44' S, 47°30' E, 3700 m, 2 lv.

md 32 Reunion: stn DS 149, 20°26' S, 55°40' E, 3500-3510 m, 2 lv, 2 dd. — Stn DS 151, 20°51' S,
56°03' E, 3240-3300 m, 4 lv.

304

VICTOR SCARABINO

SAFARI 1: stn CP 06, 30°40' S, 48° 14' E, 4020-4035 m, 1 Iv. — Stn DS 05, 30°37' S, 48°30' E, 4500-
4612 m, 2 lv. — Stn DS 08, 24°22' S, 58°19' E, 5025-5825 m, 1 lv. — Stn SIPAN 79 09, 30°41' S,
48°28' E, 4462 m, 1 lv. — Stn CP 17, 24°26' S, 58° 19' E, 4987-5025 m, 1 lv.

Fig. 106. — Distribution of Costentalina indica.

Distribution. - Central Indian Ocean, SE of Sri Lanka, Madagascar and Reunion Island.
Live records from 3240 to 5285 m (present paper).

Other Indo-Pacific species of Costentalina cited in the literature

Costentalina elegans Chistikov, 1982. Indian Ocean, “Vitiaz”, stn 6744-5 T, 12°47' S, 88°54' E,
5100-5200 m, zin.

Genus E\taui\opsis Habe, 1957

Type species (OD): Denialium nivosum Kuroda & Kikuchi, 1933 [= Dentalium intercostalum Boissevain, 1906].

Diagnosis. — Shell medium to large, slightly curved, to nearly straight, solid, opaque or
polished, chalky white. Longitudinal sculpture of 7 angled primary ribs, one ventral; intercostal
spaces convex. Apex simple, with crownlike appearance in fresh specimens due to projection of the
primary ribs. Preapical callus prominent, lumen circular. Starlike in section.

Radula rachidian subpyramidal, anterior border rounded, base wide; lateral with one prominent
primary cusp and secondary denticles, base wide; marginal long, slightly sigmoidal.

Distribution. — Recent, West Pacific and Indian Oceans, absent in the Atlantic. Shelf,
bathyal.

Source : MNHNJ Paris

SCAPHOPODA OF THE TROPICAL 1NDO-PACIFIC

305

Fig. 107. - a. Entalina mirijlca , shell (21 mm), ventral view of apex (note double ribs), apical section, variation of oral section,
detail of sculpture, musorstom: stn CP 25 and oral section (right fig.), musorstom 2: stn CP 18 b. Entalina
subterlinieata, shell (17 mm) " Meiring Naude", stn SM 123. — c, Entalina type radula (E. mirifica and an isolated
rachidian of E. quinquangulare [Northeastern Atlantic Ocean] bottom right). - d. Heteroschismoides subterfissum
(Northeast Atlantic Ocean, Bay of Biscay, mnhn), shell (13 mm), lateral and ventral faces, apex, apical and oral sections.

e, Heteroschismoides type radula ( //. subterfissum). — f. Costentalina tuscarorae, shell (II mm), lateral and dorsal
faces, apical and oral sections, estase 2: stn CP 2. g. Costentalina indica, shell (10 mm), apical and oral sections,
safari 1: stn DS 5. h. Costentalina type radula (C. vemae, Argentine basin. South Atlantic Ocean). Scale lines:
100 pm.

Entalinopsis inter cost at a (Boissevain, 1906)
Figs 108, 115 a, d

Dentalium inlercostatum Boissevain. 1906: 14, pi. 6, fig. 14. textfig. II.

Synonyms:

? Dentalium siberutense Plate. 1908a: 348. pi. 30, figs 17-20 (Syn. nov.).

Dentalium nivosum Kuroda & Kikuchi, 1933: 7, pi. 1. figs 9-10, textfigs 1-2.
Dentalium tugaruense Nomura & Hatai, 1940: 73, pi. 3, fig. 4 (fide Habe, 1964a).

306

VICTOR SCARABINO

Other references:

Enialina intercostata - Habe. 1977: 339, pi. 71. figs 3-5.

Entalina ( Enlalinopsis ) intercostata - Habe. 1963: 272, pi. 38, figs 24-25.

Entalina (Enlalinopsis) nivosa - Habe, 1957: 135, fig 9.

Enlalinopsis intercostatus - Habe, 1964a: 40, pi. 2, figs 24-25, pi. 5, figs 63-65; 1971: 492 (Japanese text), 310 (English text),
pi. 65, figs 20-21. — Habe et a!., 1986: 24.

Dentalium (Entalina) intercostatum - Shikama, 1964: 35, fig. I.

Enlalinopsis intercostata - Higo & Goto, 1993: 689.

Type material. — D. intercostatum-. holotype zma 3.06.010.

Type locality. — D. intercostatum. Indonesia, Ceram Sea, “Siboga", stn 178, 02°40' S,
128°37' E, 835 m. — D. siberutense: Indonesia, Siberut Island, 00°39' S, 98°52' E, 750 m. — D.
nivosum: Toyama Bay, Japan, 80-150 m. — D. tugaruense: Japan.

Material examined. — The type material listed.

Indonesia. " Snellius ” II: stn 4.130, 08°18'S, 1 1 8°1 8' E, 700-730 m, 4 dd. — Stn 4.131, 08°18'S,
1 1 8° 1 8' E, 680-800 m, 1 dd. — Stn 4.135, 06°29' S, 121°09' E, 495 m, 1 lv. Stn 4.267, 08°18' S,
1 18°21' E, 650 m, 1 dd (rmnh).

Philippines, musorstom 2: stn CP 24, 13°37'N, 120°42' E, 640-647 m, 2 lv, 5 dd. — Stn CP 25,
13°39' N, 120°43' E, 520-550 m, 82 lv, 282 dd. — Stn CP 55, 13°54' N, 1 19°58' E, 865 m, 4 dd. -
Stn CP 78, 13°49'N, 120°28' E, 441-550 m, 1 dd.

musorstom 3: stn DR 93, 13°49'N, 120°02'E, 540 m, 1 dd. — Stn DR 94, 13°47'N, 120°03' E,
842 m, 1 lv, 5 dd. — Stn CP 106, 13°47' N. 120°30' E, 640-668 m, 30 lv, 43 dd.

Distribution. — Indonesia, East China Sea, Japan and the Philippines, living from 495 to
842 m (present paper), shells from 50 m (Habe & Kosuge, 1964).

Entalinopsis rnicra sp. nov.

Figs 109, 111 c-d, 115 c

Type material. — Holotype and 6 paratypes dd mnhn.

Type locality. — S New Caledonia, “ Vauban ” 1978-79, stn 40, 22°30' S, 1 66°24’ E, 250-

350 m.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

307

Material examined. — Only known from the type material.

Distribution. — New Caledonia. Shells

Description. — Shell to 8 mm long, arched, fragile,
translucent, cream in color. Sculpture of seven primary ribs
and one to three secondary ribs, originating near the apex,
less prominent at the mouth than the primary ribs. Apex fine
with protuberant ribs, star-like cross section. Subapical callus

only in 250-350 m.

fine, lumen circular. Mouth fragile, polygonal in section.

Measurements: holotype L 6.8, W 0.7. w 0.4, arc 0.5;
paratypes L 7.4, W 0.6, w 0.4, arc 0.6; L 6.3, W 0.5, w 0.3,
arc 0.5; L 6, W 0.5, w 0.3, arc 0.4. W/w ratio 1.8-1. 6.

Etymology. — From the Latin micro, very small.

Entalinopsis stellata sp. nov.

Figs 110, 111 a-b, 115 b

Type material. — Holotype mnhn. Paratypes: 6 mnhn, 1 ams C20I730, 1 usnm.

Type locality. — Philippines, musorstom 2, stn CP 18, 14°00' N, 120°18' E, 188-195 m.

Material examined. — New Caledonia, biocal: stn DW 80, 20’32' S, 166°48' E, 900-980 m,
1 lv, 1 dd.

Philippines, musorstom 2: stn CP 18, 14°00' N, 120°18' E, 188-195 m, 1 lv (holotype). — Stn CP 55,
13°54' N, 1 19°58' E, 865 m, 4 lv (paratypes: 3 mnhn, 1 ams), 3 dd (paratypes: 2 mnhn, 1 usnm).
musorstom 3: stn DR 94, 13°47'N, 120°03'E, 842 m, 1 dd (paratype).

Distribution. — The Philippines and New Caledonia, living depth range 195-900 m.

Description. — Shell to 26.2 mm long, regularly curved,
opaque, white. Sculpture of seven prominent, raised ribs, one
median on ventral side, two on dorsal one. Intercostal spaces
concave with growth lines. Ribs extending over apex, which
is star-like in cross section. No secondary ribs. Lumen

subcircular, prominent subapical callus. Mouth fragile, poly¬
gonal in section, slightly laterally compressed.

Measurements: holotype L 22.7. W 2.3, w 0.85, arc 2;
paratypes L 26.2, W 2.9-2. 6, w 1.15, arc 2.5: L 20.1, W 2.45,
w 1.5, arc 1.5. W/w ratio 1.6-2. 7.

308

VICTOR SCARABINO

Remarks. — One specimen is sculptured with nine ribs (Fig. Ill a).

Etymology. - - Named for the star-shaped transverse section.

Other Indo-Pacific species of Entalinopsis cited in the literature

Entalinopsis habutae (Kuroda & Kikuchi, 1933): 8, pi. 1, figs 3-4. 12-13. Japan, Toyama Bay. 1 50-
350 m. Geological Institute of Kyoto University. Japan.

Genus Spadentalina Habe. 1963
Type species (OD): Deitlalium lubiforme Boissevain. 1906.

Diagnosis. — Shell medium to large, slightly to well curved, solid, translucent when fresh,
opaque to polished when dead, white to light brown. Longitudinally sculptured by 8 angled or
rounded primary ribs, two of which are ventral, two dorsal and two pairs lateral. Secondary ribs
present, intercostal spaces convex to straight; the entire surface usually strongly cancellate. Apex with
the ventral wall wider and a ventral lobe, often with a regularly spaced series of holes creating the
lobe by reabsorption. Preapical callus thin. Section starlike to polygonal, oral aperture thin.

Radula rachidian high, subtriangular, similar to that of Pertusiconcha; lateral well armed, with two
primary cusps and denticles; marginal slightly curved.

Distribution. Recent, worldwide, bathyal-abyssal.

Spadentalina tubiformis (Boissevain, 1906)

Figs 111 i-j, 112, 115 e, 172 a-b

Dentalium lubiforme Boissevain. 1906: 19, pi. 6, fig. 5.

Other references:

Spadentalina tubiformis - Habe, 1964a: 11, pi. 4, figs 9-12; 1977: 333. Habe & Kosuge; 1964: 5. Chistikov, 1982a: 677.
pi. 3. figs 8-14. pi. 5, fig. 3. — Higo & Goto, 1993: 686.

Source : MNHN. Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

309

Fig. 111. a, Enialinopsis siellata, paratype, apex of a nine-ribbed specimen. — b, same specimen, detail of surface in
intercostal space. c, Enialinopsis micra , paratype, apex. d, same specimen, detail of surface in intercostal space.

e, Spaderlalina ingrata, paratype, detail of surface in intercostal space. — f, same specimen, apex. — g, Spadentalma
ingrata. specimen with the embryonic shell. — h, Costentalina tuscarorae , apex. — i. Spadentalina tubijormis, apex,
j, ' same specimen, detail of surface. k. Costentalina tuscarorae. detail of surface in intercostal space. —
I, Striopulsellum minimum, detail of surface. m, Solenoxiphus striatulus. detail of surface of a rib and intercostal
spaces. n, Dischides minutus , apex. — o, same specimen, detail of surface. Scale lines: 100 pm (a, c, t, g, h. 1, n), to pm

(1), 2 pm (b, d, e, j, k, m, o).

Source : MNHN, Paris

310

VICTOR SCARABINO

Type material. — Lectotype (here designated) zma 3.06.013, paralectotypes zma 3.06.012.
Type locality. — Indonesia, “ Siboga ”, stn 212, 05°56' S, 120°19' E, 462 m.

Material examined. — The type material.

Indonesia, corindon: stn B 210, 00°13' S, 1 17°53' E, 338 m, 2 dd. — Stn B 207, 00°15' S, 1 17°52' E,
150 m, 3 dd. — Stn B 248, 00°54' S, 119°29' E, 170 m, 14 dd.

“Snellius” II: stn 4.1 12, 08°19' S, 1 18°16' E, 365 m, 6 dd. — Stn 4.1 13, 08°18' S, 1 18°16' E (no depth
data), 2 dd. — Stn 4.135, 06°29' S, 121°09' E, 495 m, 1 lv (rmnei).

Philippines, musorstom 2: stn CP 82, 13°46'N, 120°28'E, 550 m, 3 dd. — Stn DR 83, 13H55'N,
1 20°30' E, 318-320 m, 3 lv.

musorstom 3: stn DR 94. 13°47'N, 120°03' E, 842 in, 1 dd. — Stn CP 106, 13°47'N, 120°30' E,
640-668 m, 3 dd. — Stn CP 122, 12°20' N, 121°42' E, 673-675 m, 2 dd. — Stn CP 123, 12°11' N,
121°45'E, 700-702 m, 1 dd.

Fig. 112. — Distribution of Spadentalina tubiformis.

Distribution. — Indonesia and Japan, now recorded from the Philippines, alive in 318-
495 m, shells down to 702 m (present paper).

Spadentalina ingrata sp. nov.

Figs 1 1 1 e-g, 113, 115 f

Type material. — Holotype mnhn. Paratypes: 12 mnhn, 1 ams C201731, 1 nmnz M268951,
1 USNM.

Type locality. — Loyalty Islands, musorstom 6, stn DW 444, 20°54' S, 167°18' E, 300 m.

Material examined. — Chesterfield Islands, chalcal 1: stn DC 38, 20°00' S, 158°46'E,
250 m, 2 dd.

musorstom 5: stn DW 280, 24°10' S, 159°36'E, 270 m, 1 dd (paratype usnm). — Stn DW 282,
24° 12' S, 159°32' E, 226-230 m, 3 dd ( paratypes: 2 mnhn, 1 ams). — Stn DW 285, 24°09' S, 159°34' E,

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

311

245-255 m, 3 dd. — Stn DW 334, 20°06' S, 158°48' E, 315-320 m, 1 dd (paratype). — Stn DW 335,

20°03' S, 1 58°45' E, 315 m, 1 dd. — Stn DW 344, 19°39' S, 158°34' E, 310 m, 1 dd (paratype).

New Caledonia. “Vauban" 1978-79: stn 33, 22°33' S, 166°25' E, 290-335 m, 3 dd. — Stn 34, 22°32' S.
1 66°26' E, 350-420 m, 1 dd (paratype). — Stn 35, 22°32' S, 166°26' E, 250-375 m, 1 dd. — Stn 37^
22°32' S, 166°26'E, 175-250 m, 2 dd. — Stn 40, 22°30' S, 1 66°24' E, 250-350 m, 15 lv, 166 dd
(paratypes: 3 mnhn, 1 nmnz).

biocal: stn DW 43, 22°46' S, 167°15' E, 400 m, 1 dd. — Stn DW 77, 22°15' S, 167°15' E, 440 m,
1 lv. — Stn DW 104, 21°31' S, 166°21' E, 375-450 m, 4 dd.

biogeocal: stn KG 252, 21°31'S, 166°21'E, 330 m, 1 dd. — Stn DW 253, 21°32' S, 166°29'E,
310-315 m, 17 dd (1 paratype).

Passe de Boulari, B. Richer/ORSTOM coll., 400 m, 5 lv.

Loyalty Islands, musorstom 6: stn DW 406, 20°41'S, 167°07'E, 373 m, 1 dd. — Stn DW 411,
20°40'S, 167°03'E, 424 m, 1 lv. — Stn DW 444, 20°54' S, 167°18' E, 300 m, 1 lv (holotype). —

Stn DW 446, 20°54' S, 167°19' E, 360 m, 3 dd (paratypes). — Stn DW 449, 20°54' S, 167°18' E, 300

m, 1 dd. — Stn DW 485, 2P23' S, 167°59' E, 350 m, 1 dd.

Indonesia. " Snellius " II: stn 4.031, 05°54' S, 123°58'E, 390 m, 1 dd.

Philippines, musorstom 1: stn CP 62, 14°00'N, 120°16'E, 179-194 m, 1 dd.

MUSORSTOM 2: stn CP 20, 14°00' N, 120° 18' E, 185-192 m, 1 lv, 1 dd. — Stn CP 72, 14°00'N,
120° 18' E, 182-197 m, 1 lv.

musorstom 3: stn CP 102, 14°01' N, 120°18' E, 192 m, 1 lv, 6 dd. — Stn DR 126, 1 1°49' N, 121°22' E,
266 m, 4 dd.

Fig. 113. — Distribution of Spadetualina ingrata.

Distribution. — The Philippines, Indonesia, New Caledonia, alive from 182 to 440 m.

Description. — Shell to 37 mm long, slender, solid.
Young specimens regularly curved, adults vary from almost
straight to regularly curved or curved only at the posterior
end. Sculpture of 8 primary ribs, prominent throughout.
Intercostal spaces with 6 to 7 riblets crossed by transversal
lines, giving a slightly cancellate appearance. Apex with long
lobe-like structure on the ventral side in fresh specimens,
showing only the base when broken. Mouth straight, octo-
gonal in section, slightly compressed laterally.

Measurements: holotype L 24.5, VV 2.65, w 0.82, arc 2.5;
paratypes L 36.4, W 2.2, w 1, arc 0.5; L 26.2, W 1.85, w 0.65,
arc 1; L 30.6, W 2, w 1.5, arc 2.5; L 30.5, W 2.2, w 1.2, arc
2; L 25.4; W 2.3, w 0.8, arc 1.5; L 27.6, W 2.5, w 0.85. arc 1.2;
L 34.7, W 2.6, w 1.1, arc I; L 19.2, W 1.2, w 0.4, arc 0.5. W/w
ratio 1. 3-3.2.

312

VICTOR SCARABINO

Remarks. - This species is variable in curvature, but constant in sculpture. It differs from
Spadentalina tubiformis from which it differs in the less prominent cancellate sculpture. The dorsal
lobe is olten regurlarly perforated on both sides (Fig. 1 15f ) in fresh specimens. This character is also
noted in Pertusiconcha and may be caused by the reabsorption process that forms the apical structure
This species may have been confused with Entalinopsis habutae but the later has 6 ribbs and a better
defined sculpture.

Etymology. — From the Latin ingrata , indicating the difficulties noted in defining the species.

Genus Pertusiconcha Chistikov, 1982

Type species (OD): Dentalium callithrix Dali, 1889. Recent, Yucatan strail, 640 fms [1170 m].

Diagnosis. — Shell medium to large, slightly curved, solid, opaque or polished, chalky white.
Longitudinal sculpture of 8 primary ribs and convex intercostal spaces. Apex truncate, irregular by
presence of regularly or irregularly spaced circular holes. Preapical callus prominent, lumen circular.
Section oval, laterally compressed.

RacJula rachidian high, subtriangular with rounded anterior margin, lateral half folded in, especially
at base; lateral head with one sharp pointed primary cusp and several subequal denticles; marginal
slightly curved (Fig. 1 15 g). 6

Distribution. Recent, worldwide, bathyal-abyssal.

Pertusiconcha tridentata Chistikov, 1982
Figs 114, 115 h

Pertusiconcha tridentata Chistikov, 1982a: 678, pi. 4, figs 2-13.

Type material. — zin (fide Chistikov).

,,An TYPE locality- — "Dimitri Mendeleyev", stn 1244T, Tasman Sea, 31°43'S 159°00'E
1640 m.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

313

Fig. 115. a, Entalinnpsis iniercostaia, shell (55 mm), apex, apical and oral sections, musokstom 2, stn CP 24.

b. Entalinopsis slellaia sp. nov., holotype, shell (22.7 mm), apex, apical and oral sections. — e. Entalinopsis micro sp.
nov., holotype. shell (6.8 mm), apex and sagital section of apex, apical section, detail of sculpture. d. Entalinopsis
type radula (E. iniercostaia). e. Spadentalina tubiformis, shell (37 mm), apex, apical and oral sections, musorstom
2: stn CP 82. f, Spadentalina ingrata sp. nov., holotype, shell (24.5 mm), apex, apical and oral sections, section near
the apex. — g, Pertusiconcha type radula (P. callithrix. Western Atlantic Ocean, Puerto Rico Trench),
h. Pertusiconcha tridentata, shell (29 mm), apex, apical, medial and oral section, biocal: stn DS 04.

Material examined. New Zealand. “ Galathea stn 665, 36"38' S, 1 78°2 1 ' E. 2470 m, 1 dd.
New Caledonia, biocal: stn DS 04, 21°16'S, 166"40' E, 2340 m, I dd. - Stn CP 26, 22°40' S,
166°27'E. 1618-1740 m, 1 dd. Stn KG 71, 22°10' S. I67°33' E, 2099 m. 1 dd. Stn CP 72,
22° 10' S, 167°33' E, 2100-2110 m, 1 Iv, I dd. Stn CP 75, 22° 19' S, 167°23' E, 825-860 m, 1 lv. -
Stn DW 79, 20°40'S. 166°52'E, 1320-1380 m, 3 dd. - Stn KG 86, 21°01'S. 166°58' E. 1860 in,

Source . MNHN. Paris

314

VICTOR SCARABINO

1 dd. — Stn KG 89, 21°03'S, 166°56'E, 2070 m. 1 Iv. — Stn DS 98, 21°24'S, 166°30' E 2365-
2470 m, 2 dd. — Stn KG 102, 21°28' S, 166°26' E, 1810 m, 1 lv.

biogeocal: stn CP 260, 21°00' S, 166°58' E, 1820-1980 m, 1 dd. — Stn KG 262 21°02' S 167°02' E
1380 m, 1 lv. — Stn CP 266, 21°05' S, 1 66°57' E, 1990-2100 m, 2 dd. — Stn CP 273 21°02' s’
166°57' E, 1920-2040 m, 1 dd. — Stn CP 321, 21°12' S, 167°00' E, 2190-2205 m 1 dd
Philippines, estase 2: stn CP 2, 14°05' N, 120°02' E, 2050 m, 1 dd.

West Indian Ocean, benthedi: stn DR 40, 12°56'S, 45° 18' E, 1300-1480 m, 1 lv.

Distribution. — Tasman Sea, now extended to New Zealand, New Caledonia, the Philippines
and Madagascar. Recorded alive from 825 to 2070 m, shells down to 2470 m.

Subfamily Bathoxiphinae Chistikov, 1983

Genus Bathoxiphus Pilsbry & Sharp, 1897

Type species (SD by Boissevain, 1906): Dentalium ensiculum Jeffreys, 1877. Recent, North Atlantic Ocean.

Diagnosis. — Shell small to medium in size, well arched, solid, polished, white. Almost
unsculptured, angled at dorsal and lateral sides. Apex truncate, lumen circular, located submedially
due to dorsal wall width in apical area. Section oval, strongly laterally compressed.

Radula rachidian high, anterior margin simple, rounded with latter half folded inward; lateral with
two pointed primary cusps and 5 to 6 subequal secondary cusps; marginal slightly curved.

Distribution. — Recent, worldwide, bathyal-abyssal.

Bathoxiphus soyomaruae Okutani, 1964
Figs 116, 121 a-b

Bathoxiphus soyomaruae Okutani, 1964: 77, fig. 4.
Other references:

Vo7v' 77%,4a: H p1' 5' nfn61-62/ 1977:,337’ P‘- 70- fi8s l7->8- - Habe & Kosuge, 1964: 6.
figs 6-9 L- h?go2 & Goto.' 1993 688 Syn0"ym """ ~ Ch,ST,kov’ 1983: 183- P1- 3- r>gs 6-22, pi. 4.

Fig. 116. — Distribution of Bathoxiphus soyomaruae.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

315

Type material. — nsmt, not checked.

Type locality. — Japan, off Aogashima Island, 3150-3350 m.

Material examined. — Chesterfield Islands, musorstom 5: stn DW 322, 21°19' S, 158°00' E,
975 m, 1 lv, 1 dd.

New Caledonia, biogeocal: stn KG 233, 21°31' S, 166°25' E, 1040 m, 2 lv, 2 dd.

West Indian Ocean, benthedi: stn CH 87, 1 1°44' S, 47°35' E, 3716 m, 1 dd. — Stn CH 90, 1 1°44' S,
47°30' E, 3700 m, 1 lv.

md 32 Reunion: stn DS 151, 21°5T S, 56°03' E, 3240-3300 m, 1 lv. — Stn DS 149, 20°26' S, 55°40' E,
3500-3510 m, 1 dd.

East Indian Ocean, safari 2: stn CP 10, 01°43' N, 87°08' E, 4350 m, 1 lv.

Distribution. — Japan, now extended to New Caledonia and the Indian Ocean, living from
975 m (present paper) to 5750 m (Chistikov, 1983).

Bathoxiphus inexpectatus sp. nov.

Figs 117, 121 c

Type material. — Holotype mnhn. Paratypes: 11 mnhn, 1 ams C201732, 1 nmnz M268952,
1 USNM.

Type locality. — New Caledonia, N Norfolk Ridge, biocal, stn DW 46, 22°53' S, 167° 17' E,
570-610 m.

Material examined. — Chesterfield Islands, musorstom: 5 stn DW 362, 19°53' S, 158°40' E,
410 m, 1 lv.

New Caledonia, biocal: stn DW 44, 22°47' S, 167° 14' E, 440-450 m, 6 lv, 4 dd. — Stn DW 46,
22°53' S, 167° 17' E, 570-610 m, 20 lv (holotype and paratypes: 10 mnhn, 1 ams, 1 nmnz, 1 usnm),
17 dd. — Stn DW 66, 24°55' S, 168°22' E, 505-515 m, 1 dd.
smib 3: stn DW 22, 23°03' S, 167°19' E, 503 m, 1 dd.

316

VICTOR SCARABINO

lagon: stn 830. 20°49' S. 165° 19' E, 105-110 m, 1 dd (paratype).

Loyalty Islands, musorstom 6: stn DW 459. 21°01'S, I67°31' E, 425 m. 2 Iv, 11 dd.

Distribution. - New Caledonia, live records between 410 and 570 m.

Description. Shell lo 13 mm long, lacking sculpture, V-shaped notches, preapical callus prominent, lumen cir-
polished. arched, translucent white. Section slightly compres- cular, mouth straight.

sed laterally throughout. Apex strong with two lateral flat Measurements: holotype L 11.9. W 1. 2-1.1, w 0.5-0.45, arc

1.6. W/w ratio 2.4.

Etymology. From the Latin, meaning unexpected.

Genus Rhomboxiphus Chistikov. 1983

Type species (OD): Denialium tricarinaium Boissevain, 1906.

Diagnosis. Shell small to medium, well arched, solid, polished, white, translucent.
Sculpture of 4 flat primary ribs, located ventrally, dorsally and laterally; secondary ribs may be
present. Apex truncate, lumen circular located submedially due to the dorsal side width at apical area.
Section rhomboidal, strongly laterally compressed.

Radula similar to that of Bathoxiphus.

Distribution. Recent, worldwide, bathyal-abyssal.

Rhomboxiphus tricarinatus (Boissevain, 1906)

Figs 1 18, 121 d, f

Denialium tricarinaium Boissevain, 1906: 48, pi. 6. figs 40-41.

Synonym:

Denialium (Compressidens ) capense Tomlin. 1931: 340 (Syn. nov.).

Other references:

Denialium (Bathoxiphus) tricarinatus — Plate, 1908a: 354.

Bathoxiphus tricarinatus - Hare, 1964a: 33, pi. 5, figs 69-70: 1977: 337. pi. 70. figs 15-16. Haul & Kosuge, 1964: 6.
Higo & Goto, 1993: 688.

Bathoxiphus tricarinaium Okutani. 1964: 76, pi. 6, fig. 10.

Denialium capense — Barnard, 1963a: 446: 1963b: 349, fig. 30d: 1974: 742.

Type material. — D. tricarinaium : lectotype (here designated) zma 3.06.060, paralectotypes
zma 3.06.056-059. D. capense: holotype sam.

Type locality. — D. tricarinatus: Indonesia, Ceram Sea, “ Siboga ”, stn 178, 02°40' S,
128°37' E, 835 m. - D. capense: South Africa, “Cape Point, N 86° E. 43 miles, 900 fins" [1645 m].

Material examined. - The type material.

Tasman Sea. " Galathea stn 574. 39°45' S. 159°39' E, 4680-4730 m, I dd.

New Caledonia, biocal: stn CP 26, 22°40' S. 166"27' E, 1618-1740 m, 9 dd.

biogeocal: stn CP 260, 2 TOO' S. 166°58' E. 1820-1980 m, 1 dd. Stn DW 313, 20°59' S, 166°59' E,

1600-1640 m, I dd.

calsub: dive 13. 21°26' S, 166°23' E, 1600 m. 1 dd.

Indonesia, corindon: stn B 244, 00°56' N. 119°22' E, 970 m, 4 dd.

“ Snellius ” II: stn 4.135, 06°29' S. 121°09' E. 495 m, 1 dd.

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SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

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Philippines, estase 2: stn DW 1. 14°05' N, 120"01' E. 2200 m, I dd. - Stn CP 2. 14°05' N. 120°02' E,
2050 m, 13 lv, 10 dd.

musorstom 2: stn CP 55, 13°54'N, 119°58'E, 865 m, 1 dd.
musorstom 3: stn DR 94, 13°47' N, 120°03' E, 842 m, I dd.

West Indian Ocean, benthedi: stn DS 02, 12°35' S, 47°40' E, 1750 m, 3 lv. — Stn DS 03, 1 2°36' S.
47°38' E, 1 100-1150 m, I dd. — Stn DR 11, 12°16' S, 46°42' E, 2300-2450 m, 1 dd. Stn DR 27,
12°38'S. 47° 1 2' E, 675 m, I dd. — Stn DR 28, 12°42'S, 45°20' E, 705 m, 1 dd. — Stn DS 64,
1 2°4 1 ' S. 44°57' E, 770-860 m, 1 dd.

md 32 Reunion: stn DC 64. 21°12' S, 55°04' E, 1150-1180 m, 3 dd. — Stn DR 67, 21°13' S, 55°01' E,
1390-1425 m, 1 dd. Stn DS 78, 21°l 3' S. 55°04' E, 1 175-1200 m, 2 lv, 2 dd. — Stn DS 100, 21°27' S,
55°47' E, 4180-4220 m, 1 lv. Stn DR 104, 20°49' S, 55"0r E, 1875-1920 m, 2 dd. - Stn DS 106,
20°48' S, 55°05' E, 1710-1730 m, I dd. Stn DS 109, 20‘’52' S. 55°06' E. 1050-1240 m, 7 lv. 6 dd.
“Galathea”: stn 234. 05°25' S. 47‘W E. 4830 m, 1 lv.

South Africa. “ Meiring Naude”: stn 94. 28°16' S. 32°29' E, 670 m, 1 lv, 1 dd.

Fig. 118. Distribution of Rhomboxiphus tricarinatus.

Distribution. From Japan to the East coast of Africa, 639-1900 m (Habe & Kosuge.
1964), now recorded from the Philippines and the SW Pacific. Live records from 495 to
4830 m.

Rhomboxiphus colmani (Palmer, 1974)
Figs 119, 121 e

DentaHum ( Ballioxiphus) applanatum Colman. 1958: 145, fig. 12 (non D. applanation Torley, 1908).
Dentalium (Ballioxiphus) colmani Palmer. 1974b: 124, nom. nov. pro D. applanatum Colman.

Other references:

Ballioxiphus colamni (sic) - Okutani, 1975: 77, pi. 3, figs 6-7 (erroneous subsequent spelling).
Rhomboxiphus colmani Chistikov, 1983: 183.

Type material. — Holotype and 3 paratypes dd ams 26652.

Type locality. Australia, New South Wales, 35 miles E of Sydney, 1463 m.

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VICTOR SCARABINO

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DC 379, 19°53' S, 158°40' E, 370-400 m, 1 lv.
New Caledonia, biogeocal: stn CP 232, 21°34' S, 166°27' E, 760-790 m, 2 lv, 4 dd.

Distribution. — E Australia now extended to New Caledonia, alive in 370-790 m, shells down
to 1463 m.

Remarks. — Chistikov (1983) synonymized this species with Bathoxiphus soyomaruae , but
study of type specimens confirms the validity of Rhomboxiphus colmani. It is similar to Rhomboxiphus
tricarinatus, but less sculptured.

Genus Solenoxiphus Chistikov, 1983
Type species (by monotypy): S', striatulus Chistikov, 1983.

Diagnosis. — Shell medium sized, slightly curved, solid, translucent when fresh, white opaque
to polished when dead. Longitudinally striated throughout. Apex oblique, the dorsal angle higher
than the ventral side, preapical callus wide, lumen circular. Section strongly laterally compressed,
dorsal and ventral sides rounded, laterals straight parallel or slightly concave.

Radula rachidian with lateral sides almost parallel, anterior margin with central lobe; laterals with
two main cusps and four denticles between, with a third cusp located at the base of the external cusp;
marginals slightly sinusoidal.

Distribution. — Recent, Pacific Ocean, bathyal-abyssal.

Solenoxiphus striatulus Chistikov, 1983
Figs 111m, 120, 121 g-h, 172 g

Solenoxiphus striatulus Chistikov, 1983: 187-188, pi. 3, figs 2-13, pi. 5, figs 10-11.

Type material. — zin (fide Chistikov).

Type locality. — “Vitiaz”, stn 5944-2, North Fiji basin, 14°2T S, 179°38' E, 2380 m.

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SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

319

Material examined. — New Caledonia, biocal: stn KG 03, 21°15'S, 166°39'E, 2340 m.
1 lv, 1 dd. — Stn DS 04, 21°16' S, 166°40' E, 2340 m, 3 lv, 6 dd. — Stn CP 05, 21°16' S, 166°44' E,
2340 m, 1 lv, 3 dd. — Stn DS 14, 20°19' S, 167°18' E, 3680-3700 m, 2 lv. — Stn CP 26, 22°40' S,
166°27' E, 1618-1740 m, 1 lv, 3 dd. — Stn KG 71, 22°10' S, 167°33' E, 2099 m, 2 dd. — Stn CP 72,
22° 10' S, 167°33' E, 2100-21 10 m, 10 lv. — Stn KG 85, 20°59' S, 167°00' E, 1639 m, 1 lv. — Stn KG
89, 21°03' S, 166° 5 6' E, 2070 m, 2 lv, 7 dd. — Stn KG 90, 21°08' S, 166°48' E, 2236 m, 4 dd.

Sin DS 98, 21°24'S, 166°30' E, 2365-2470 m, 7 lv, 1 dd. — Stn KG 101, 21°27'S, 166°24'E,
1790 m, 1 dd. — Stn KG 102, 21°28' S, 166°26' E, 1810 m, 1 dd.

biogeocal: stn KG 207, 22°38' S, 166°29' E, 1350 m, 1 dd. — Stn KG 240, 21°29' S, 166°27' E,
1520 m, 1 lv. — Stn CP 243, 21°27' S, 166°26' E, 1820 m, 1 dd. — Stn KG 248, 21°15' S, 166°29' E,
2340 m, 1 lv, 2 dd. — Stn CP 260, 21°00' S, 166°58' E, 1820-1980 m, 1 lv, 1 dd. — Stn KG 261,
21°02' S, 167°02' E, 1508 in, 1 lv, 1 dd. — Stn KG 267, 21°02' S, 166°59' E, 1935 m, 2 dd. — Stn KG
268, 21°03' S, 166°57' E, 1990 m, 1 dd. — Stn KG 269, 21°02' S, 166°58' E, 1810 m, 2 dd. — Stn KG
275, 21°06' S, 166°53' E, 1959 m, 2 lv, 1 dd. — Stn KG 277, 21° 17' S, 166°56' E, 2240 m, 1 lv. —
Stn KG 287, 20°43' S, 166°53' E, 1560 m, 1 lv. — Stn CP 317, 20°48' S, 166°53' E, 1620-1630 m,
1 dd.

calsub: dive 13, 21°26'S, 166°23'E, 1600 m, 1 lv.

Distribution. — North Fiji Basin, now extended to New Caledonia, alive in 1500-3700 m.

Suborder Gadilimorpha Steiner, 1992
Family Pulsellidae Scarabino in Boss, 1982

Genus Pulsellum Stoliczka, 1868

Type species (SD by Cossmann, 1888): Siphonodentalium lofoiense Sars, 1865. Recent, Norwegian Sea, bathyal.

Diagnosis. — Shell small, slightly to markedly curved, fragile, translucent to white in dead
shells. Sculptured by growth lines, occasionally conspicuous. Apex simple, preapical callus thin,
lumen circular. Section circular.

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VICTOR SCARAB1NO

Fig. 121. a, Baihoxiphus soyomaruae, shell (22 mm), apex, apical and oral sections, biogeocal: stn KG 233.

b Baihoxiphus type radula {B. soyomaruae). - c, Baihoxiphus inexpectaius sp. nov., holotype, shell (1 1.9 mm), apical
section. d. Rhomboxiphus tricarinatus, shell (19 mm), lateral and dorsal views, apex, apical and ora sections
MD 32 Reunion- stn DS 109. - e, Rhomboxiphus colmani, shell (16 mm), lateral and dorsal views, apical and oral
sections, biogf.ocal: stn CP 232. f. Rhomboxiphus type radula (R. tricarinatus). - g, Solenoxiphus striatulus shell
(16 mm), apical and oral sections, detail of the sculpture, biocal: stn K.G 85. h, Solenoxiphus type radula (5.
striatulus).

Radula rachidian polygonal, sides almost parallel, generally with a single cusp on anterior margin;
laterals high, with two primary cusps on inner side, one on outer side, 2 to 3 denticles between cusps;
marginals slightly sigmoidal.

Distribution. — Paleocene- Recent, worldwide, shelf-abyssal.

Pulsellum fragile sp. nov.

Figs 122, 125 a

Type material. — Holotype lv, mnhn. Paratypes lv: 14 mnhn, 1 nmp.

Type locality. — West Indian Ocean, mo 32 Reunion, stn DS 151, 20"51'S, 56"03 E,
3240-3300 m.

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SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

321

Fig. 122. Distribution of Pulsellum fragile.

Material examined. — Only known from

Distribution. — Only know from the type

Description. — Shell to 3 mm long, fine, delicate, regularly
curved, translucent white, opaque. Apical area with fine
encircled wrinkles. Section circular, apex simple, mouth
simple. Subapical callus weak.

the type material.

locality.

Measurements: holotype L 3, W 0.5, w 0.2, arc 0.3;
paratype L 3, W 0.5, w 0.2. arc 0.3; L 3, W 0.5, w 0.2, arc 0.3;
L 2.3, W 0.4. w 0.1. arc 0.2; L 2, W 0.4, w 0.1, arc 0.2. W/w
ratio 2.5-4.

Remarks. — Pulsellum fragile differs from P. thomassini in general shape, by its smaller W/w
ratio and better defined transversal sculpture.

Etymology. — From the Latin fragilis, fragile.

Pulsellum thomassini sp. nov.

Figs 123, 125 b

Type material. — Holotype and 6 paratypes mnhn.

Type locality. — West Indian Ocean, benthedi, stn DR 11, 12°16' S, 46°42' E, 2300-
2450 m.

Material examined. — New Caledonia, biocal: stn KG 86, 21°01'S, 166°58' E, 1860 m,

1 dd.

West Indian Ocean, benthedi: stn DR 11, 12° 16' S, 46°42' E, 2300-2450 m, 1 Iv (holotype). — Stn 88.
1 1°46' S, 47°34' E, 3700 m, 2 lv (paratypes).

md 32 Reunion: stn DS 106, 20°28' S, 55°05' E, 1710-1730 m, 2 lv (paratypes). — Stn DS 151,
20°51'S, 56°03'E, 3240-3300, 2 lv (paratypes).

Distribution. — SW Indian Ocean and New Caledonia, live records from 1730 to 3716 m.

Description. — Shell up to 6 mm long, fine, delicate, Measurements: holotype L 6, W 0.9, w 0.6, arc 0.2;
translucent-white, opaque, regularly curved. Circular in sec- paratypes L 5.6, W 0.8, w 0.5, arc 0.2; L 6, W 0.9, w 0.5, arc
tion, apex simple, mouth simple. Subapical callus weak. 0.2; L 6, W 0.8, w 0.6, arc 0.2. W/w ratio 1.3-1. 8.

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VICTOR SCARABINO

Fig. 123. — Distribution of Pulsellum thomassini.

Etymology. — Named for Dr Bernard Thomassin, Centre Oceanologique de Marseille, cruise
leader of the benthedi Expedition (1977).

Other Indo-Pacific species of Pulsellum cited in the literature

Pulsellum eboracense (Watson, 1879): 523. “Challenger", Torres Strait, Cape York, NE Australia, 9-
20 m, 4 syntypes dd bmnh 1 887.2.9.69.

Pulsellum hige Habe, 1963: 273, textfigs 47-48. Zushi, Kanagawa, Honshu, Japan, 350 m. nsmt.
Pulsellum kurogenge Habe & Kosuge, 1964: 9. Off Choshi, Chiba Prefecture, Japan, 350 m. nsmt.

Genus Annulipulsellum Scarabino, 1986
Figs 125 d-e

Type species (by monotypy): Annulipulsellum euzkadii Scarabino, 1986. Recent, Atlantic Ocean, 12°34' N, 59°09' W, " Knorr ",
cruise 25, stn 307, 3835-3862 m.

Diagnosis. — Shell small to medium sized, fine, regularly curved, translucent white. Sculpture
of angulated rings over the entire surface. Apex oblique, preapical callus prominent, cross section
circular. Two large central pedal retractor muscles originate directly from dorsoventral muscles and
cross the intestinal sinus; pedal base and central portion enter at terminal disk. Center of disk covered
by a mucoid epithelium (Steiner 1992b).

Radula rachidian small, high with central denticle at anterior margin; laterals high with two very
sharp primary cusps; marginals sinusoidal.

Distribution. — Atlantic Ocean, bathyal-abyssal.

The genus Annulipulsellum has not been recorded in the Indo-Pacific.

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SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

323

Genus Striopulsellum gen. nov.

Striopulsellum Scarabino, 1979 (published in a thesis, not available).

Striopulsellum Scarabino in Boss 1982: 1166 nom. nud. (no diagnosis, no species included).

Type species: Siphonodentalium minimum Plate, 1908.

Diagnosis. — Shell small to medium sized, sculpture of fine, numerous, smooth longitudinal
riblets, with secondary riblets near apex; primary riblets throughout. Apex simple, straight or oblique,
preapical callus wide.

Radula rachidian polygonal, anterior margin with single cusp; lateral with 1 to 5 denticles between
cusps; marginals slightly curved.

Distribution. — Pacific and Atlantic Oceans, Southern Ocean, Recent. Bathyal-hadal.

Remarks. — This genus contains 7 deep-sea species (4 undescribed), mostly known from
Atlantic Ocean basins. The named species are S. strialinum, S. minimum and S. galatheae. With a live
record from the Sunda Trench in 6900-7000 m, S', galatheae is the deepest known scaphopod taken
alive.

Striopulsellum minimum (Plate, 1908)

Figs 111 1, 124, 125 f-g

Siphonodentalium minimum Plate, 1908b: 4, fig. 5.

Type material. — Lectotype, designated by Kilias (1995), zmb 59728a.

Type locality. — Antarctic, “Gauss" Winterstation (approximately 66°02' S, 89°38' W),
3423 m.

Fig. 124. — Distribution of Striopulsellum minimum.

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VICTOR SCARABINO

Material examined. — New Caledonia, biocal: stn KG 16, 20°34' S, 167°22' E, 3680 m,

1 lv.

Distribution. — Circumantarctic, now extended to the New Caledonia basin. Alive from
3423 m (Plate, 1908b) to 6200 m (Scarabino, 1979).

Other Indo-Pacific species of Striopulsellum cited in the literature

Striopulsellum galatheae (Knudsen, 1964): 125, figs 1-2. Sunda Trench, " Galathea ”, stn 465, 10°20' S,
109°55' E, 6900-7000 m. zmc.

Fig. 125. — a, Pulsellum fragile sp. nov., holotype, shell (3 mm), apical and oral sections. — b, Pulsellum thomassini sp. nov.,
holotype, shell (6 mm), oral and apical sections. — c, Pulsellum type radula (P. affine Sars, 1865, Northeastern Atlantic
Ocean). — d, Annulipulsellum euzkadii, holotype. shell (8.7 mm), apical and oral sections, detail of sculpture. -
e, Annulipulsellum type radula (A. euzkadii). — f, Striopulsellum minimum, shell (5.7 mm), lateral and dorsal views,
apical and oral sections, apex and detail of sculpture, biocal: stn KG 16. — g, Striopulsellum type radula (.S’, mini¬
mum).

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

325

Family Wemersoniellidae Scarabino, 1986

Genus Wemersoniella Scarabino, 1986
Type species (OD): W. turnerae Scarabino, 19S6.

Diagnosis. — Shell medium to large, straight, white, opaque or polished. Sculpture lacking or
with longitudinal undulations on dorsal side. Apex oblique or with two lateral lobes, preapical callus
wide, lumen circular. Section slightly ovate to strongly depressed dorsoventrally, oral aperture thin,
oblique.

Radula rachidian polygonal with anterior margin simple or with central cusp; laterals strong, with a
sharp pointed primary cusp with irregular wrinckle on the outer part of the granulose head; marginals
almost straight, pointed at inner margin.

Distribution. — Recent, worldwide, abyssal.

Remarks. — The genus was erected for two abyssal species from the Atlantic Ocean, W.
turnerae from the North Atlantic in 4125-5150 m, and W. duartei Scarabino, 1986a, from the
Argentine basin in 5332-5781 m. We add here two new species from the Indo-Pacific. W. turnerae is
solid while the other three species are fragile.

Wemersoniella indica sp. nov.

Figs 126, 129 a

Type material. — Holotype mnhn.

Type locality. — West Indian Ocean, benthedi, stn CH 87, 11°44' S, 47°35' E. 3716 m.

Material examined. — West Indian Ocean, benthedi: stn CH 87, 1 1°44' S, 47°35' E, 3716 m,
2 dd (holotype and one other shell that has been accidentally broken after it was photographed).

Distribution. — Only known from the type locality.

Fig. 126. — Distribution of Wemersoniella indica.

326

VICTOR SCARABINO

Description. Shell up to 1 1 mm long straight, fragile,
dorsoventrally depressed, rapidly increasing in size from apex
to oral aperture, opaque white. Apex simple, oblique from
ventral to dorsal side, preapical callus prominent, lumen

circular. Maximum diameter at oral aperture. Mouth and
prominent growth rings oblique.

Measurements: holotype L 10.2, W 2-1.7, w 0.6-0. 5, W/w
ratio 3.3.

Remarks. — Apical and radular characteristics are shared with Wemersoniella turnerae and
with W. duartei, but W. indica is much more tapering and compressed than W. duartei and more
fragile than W. turnerae.

Etymology. — From Indian Ocean, where this species occurs.

Wemersoniella knudseni sp. nov.

Figs 127, 129 b-c

Type material. — Holotype zmc. Paratypes: 2 zmc, 2 nmnz M268545, 1 mnhn.

Type locality. — " Galathea ”, stn 664, Kermadec Trench, 36°34' S, 178°57' E, 4510-4570 m.

Material examined. — New Zealand. “Galathea"-. stn 602, 43°58' S, 165°24' E, 4510 m, 1 lv
(paratype zmc). — Stn 662, 36°22' S, 178°23' W, 4630 m, 1 lv (paratype zmc). — Stn 664, 36°34' S,
178°57' W, 4150-4570 m, 2 lv (holotype zmc and paratype mnhn).

“ Tangaroa stn P 934, 41°31' S, 165°6' E, 4405-4411 m, 2 lv (paratypes nmnz).

Distribution. — Kermadec Trench and Tasman Sea, alive in 4105-4630 m.

Description. — Shell up to 12 mm long, straight, fragile,
slightly dorsoventrally depressed, porcellaneous white. Apex
with two lateral lobes separated by two wide, rounded basal
notches, the dorsal one deeper. Maximum diameter at oral
aperture. Mouth oblique; oblique growth rings numerous and

pronounced in some specimens.

Radula as for the genus.

Measurements: holotype L 11, W 2. 5-2.4, w 1.5-1. 4;
paratypes L 12, W 2.1-2, w 0.99-0.91; L 6, W 1. 4-1.1, w
0.65-0.57; L 11.2, W 2-1.5, w 0.99-0.76. W/w ratio 1.6-2.2.

Remarks. — This species differs from Wermersoniella indica in its apical structure and W/w
ratio. W. indica is more conical. W. knudseni is related to W. turnerae in the form of the lobes and
to W. duartei in its fragility and slight dorsoventral compression.

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SCAPHOPODA OF THE TROPICAL INDO-PAC1FIC

327

Etymology. — Named for Dr Jorgen Knudsen (zmc) in recognition for his work on the
deep-sea Mollusca.

Chistikovia gen. nov.

Type species: Chistikovia kermadecae sp. nov.

Diagnosis. — Shell medium, straight, fragile, polished, white. Dorsal side straight, ventral side
very slightly curved, more noticeably so on the posterior half; lateral sides similarly curved to the
ventral. Maximum diameter at anterior third. Unsculptured except for very oblique, conspicuous and
close-set growth lines. Apex wide, with subtriangular lobe, margin thin on dorsal side. Section slightly
dorsoventrally depressed. Mouth thin, oblique.

Radula similar to that of Wemersoniella.

Distribution. — Recent, Pacific and Atlantic Oceans, abyssal.

Remarks. — In addition to Chistikovia kermadecae, an undescribed abyssal species from the
North Atlantic Ocean belongs to this genus. The general shape of the shell resembles that of
Wemersoniella, but the apical structures differ. Chistikovia is placed in Wemersoniellidae, but the
original diagnosis of the family may require modification to include forms with dorsally lobed apex
and maximum diameter not exclusively at oral aperture. The last mentioned character can be
observed in gerontic stages of Wemersoniella.

Etymology. — Named for the late Sergei Chistikov (zin) for his contribution to the
knowledge of Scaphopoda.

Chistikovia kermadecae sp. nov.

Figs 128, 129 d-e

Type material. — Holotype zmc. Paratypes: 4 zmc, 1 mnhn.

Type locality. — "Galathea" stn 664, Kermadec Trench, 36°34' S, 178°57' W, 4150-4570 m.

Fig. 128. — Distribution of Chistikovia kermadecae.

328

VICTOR SCARABINO

Material examined. — New Zealand. “ Galathea ”, stn 664, 36°34' S, 178°57'W, 4150-
4570 m. 3 lv, 2 dd (holotype lv and paratypes zmc). — Stn 665, 36°38' S, 178°21' W, 2470 m, 1 dd
(paratype mnhn).

Distribution. — Kermadec Trench, alive in 4150-4570 m, shell in 2470 m.

Description. — As for the new genus, dimension up to Measurements: holotype L 11.5, W 2.1, m 1.87, w 1. 1-9.3;
H.5 mm. paratype L 10.2, W 1.9, m 1.7, w 1-0.84.

Etymology. — From Kermadec Trench.

FlG' 2u' 77 a‘ Wemerson'ella todica sp. nov., holotype, shell (10.2 mm), dorsal and lateral views, apical and oral sections. —
b, Wemersomella knudsem sp. nov., holotype, shell (11 mm), dorsal, lateral and ventral views, apex, apical and oral
sections; at right paratype, apex and apical section. — c, Wemersomella type radula (W. knudseni). — d, Chistikovia
type radula (C. kermadecae). — e, Chistikovia kermadecae sp. nov., holotype, shell (1 1.5 mm), dorsal, ventral and lateral
views; apex, apical and oral sections.

Family Gadilidae Stoliczka, 1868
Subfamily Siphonodentaliinae Simroth, 1894

Genus Siphonodentalium M. Sars, 1859

Type species^ (OD): Siphonodentalium vitreum M. Sars, 1859 [= S. lobatum Sowerby, I860). Recent, Norwegian Sea,

Source MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

329

Diagnosis. — Shell large, strong, polished, white, maximum diameter in anterior 1/3 or near
mouth. Sculpture lacking except for longitudinal threads in some species. Circular to subcircular in
section. Apex usually strong, with more than four notches and lobes (usually 6). Preapical callus wide,
prominent.

Radula rachidian variable in shape, usually with broad base and cusped anterior border; lateral strong
with wide head, well armed; marginal usually long [5. lobatum Sowerby, 1860, Fig. 135 f; S. dalli
(Pilsbry & Sharp, 1898), Fig. 135 g].

Distribution. — Eocene-Recent, worldwide, shelf-abyssal.

Siphonodentalium colubridens (Watson, 1879)
Figs 130, 135 a

Cadulus colubridens Watson, 1879: 523; 1886: 18, pi. 3, fig. 1.

Other references:

Cadulus colubridens - Pilsbry & Sharp, 1898: 184. pi. 26, fig. 71. — Boissevain, 1906: 71, pi. 3, fig. 41; pi. 6. fig. 66.
Cadulus (Gadila) colubridens — Plate, 1908a: 359, pi. 30, fig. 54.

Gadila colubridens — Okutani, 1966: 13.

Type material. — Hoiotype bmnh 1887.2.9.71.

Type locality. — “Challenger", stn 169, NE Point of New Zealand, 37°34' S, 179°22'E,
700 fms [1280 m].

Material examined. — The type material.

Chesterfield Islands, musorstom 5: stn DC 357, 19°37' S, 158°46' E, 630 m, 1 dd.

New Caledonia, biocal: stn CP 26, 22°40' S, 166°27' E, 1618-1740 m, 1 dd.

biogeocal: stn CP 238, 21°28' S, 166°23' E, 1260-1300 m, 1 dd. — Stn DW 313, 20°59' S, 166°59' E,

1600-1640 m, I dd.

Indonesia, corindon: stn DR 231, 00°05' N, 119°48' E, 980-1080 m, 1 dd.

Philippines. ESTASE 2: stn CP 2, 14°05' N, 120°02' E, 2050 m, 2 dd. — Stn DR 4, 06°08' N, 125°58' E,
2800 m, 1 dd. — Stn CP 6, 04°38' N, 119°49' E, 2570 m, 4 dd.

Fig. 130. — Distribution of Siphonodentalium colubridens.

330

VICTOR SCARABINO

Distribution. — North New Zealand, New Caledonia, Indonesia, the Philippines, Japan in
the Pacific Ocean; East Africa, 1280 m (Plate, 1908) in the Indian Ocean. Depth range 630 to
2800 m (dead).

Siphonodentalium magnum (Boissevain, 1906)

Figs 131, 135 c

Cadulus magnus Boissevain, 1906: 68, pi. 6, fig. 54, textfig. 33.

Other references:

Cadulus (Polyschides) magus (sic) — Habe 1964: 12.

Polyschides magnus - Habe, 1963: 278; 1977: 342; 1971: 496 (Japanese text), 313 (English text), pi. 65, figs 28-29. — Okutani,
1966: 14, fig. 7. — Habe, el al., 1986: 24. — Higo & Goto, 1993: 690.

Type material. Lectotype (here designated) zma 3.06.092, paralectotypes zma 3.06.091,
3.06.093.

Type locality. — “Siboga”, stn 88, Celebes Sea, 00°35' S, 119°08' E, 1301 m.

Material examined. — The type material.

New Caledonia, biogeocal: stn CP 250, 21°25' S, 166°28' E, 2350 m, 2 dd.

Distribution. — Southern Japan and Indonesia, now extended to New Caledonia. Shells fom
300 m (Habe, 1964a) to 2350 m (present paper).

Siphonodentalium hexaschistum (Boissevain, 1906)
Figs 132, 135 d

Cadulus hexaschistus Boissevain, 1906: 67, pi. 6, fig. 53, textfig. 33.

Type material. — Lectotype (here designated) zma 3.06.178.

Source :

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

331

Type locality. — “Siboga”, stn 178, Ceram Sea, 02°40' S, 128°37' E, 835 m.

Material examined. — The type material.

New Caledonia, biocal: stn CP 26, 22°40' S, 166°27' E, 1618-1740 m, 1 dd.

Indonesia. “Snellius" II: stn 4.128, 08°18'S, 118°16'E, 700-835 m, 1 dd.

Distribution. — Indonesia, now extended to New Caledonia. Shells in 700 to 1740 m (present

paper).

Siphonodentalium promontorii (Barnard, 1963)
Figs 133, 135 b

Cadulus promontorii Barnard, 1963b: 353, figs 30h-l; 1974: 743.

Fig. 133. — Distribution of Siphonodentalium promontorii.

332

VICTOR SCARABINO

Type material. — Syntypes sam A7460, and nmw (fide Oliver, 1984).

Type locality. — South Africa, 36 miles of Cape Point, 700 fms [1280 m].

Material examined. — The sam syntypes.

West Indian Ocean. “Meiring Naude stn SM 129, 30°53' S, 30°32' E, 850 m, 2 dd.

Distribution. — Endemic to South Africa, recorded alive in 1100 to 1280 m (Barnard
1963b), shells from 850 m.

Siphonodentalium jaeckeli sp. nov.

Figs 134, 135 e

Type material. Holotype and 1 paratype lv, mnhn.

Type locality. — West Indian Ocean. NW Madagascar, benthedi, stn DS 03, 12°35' S,
47°38'E, 1100-1150 m.

Material examined. — Only known from the type material.

Distribution. Only known from the type locality.

Description. — Shell to 1 1 mm long, strong, shiny, white.
Equator in anterior third, dorsoventrally compressed in
section. Apex subcircular, dorsoventrally compressed, with
five lobes, the largest ventral, two laterals and two dorsal.

Subapical callus prominent, lumen circular. Mouth simple,
subcircular, slightly laterally compressed in section.

Measurements: holotype L 11, W 2.72-2.51, m 1.6- 1.5, w
1.2-1. 1.

Remarks. — The apical area, short and wide, almost without neck, is the main distinguishing
feature of this species.

Etymology. — Named for S. Jaeckel, who studied the scaphopods of the Indian Ocean
collected by the German “ Valdivia ” Expedition, 1898-1899.

SCAPHOPODA OF THE TROPICAL INDO-PAC1FIC

333

Fig. 135. — a, Siphonodentalium colubridens , shell (15 mm), lateral and dorsal views, apex, apical and oral sections, musorstom
5: stn DC 357. — b, Siphonodentalium promontorii (after Barnard, 1963, fig. 30). c, Siphondentalium magnum, shell
(26 mm), apical and oral sections, biogeocal: stn CP 250. d, Siphonodentalium hexaschistum , shell (14 mm) and
apical section, “Snellius'l I, stn 4.128. — e, Siphonodentalium jaeckeli sp. nov., holotype. shell (11 mm), lateral and
dorsal views, apex, apical and oral sections. — f-g, Siphonodentalium type radula. — f, S. lobatum (North Atlantic
Ocean). g, 5. dalli (Antarctic).

Other Indo-Pacific species of Siphonodentalium cited in the literature

Siphonodentalium australasiae Boissevain, 1906: 64, pi. 6, fig. 68. “Siboga , stn 211, 05 41 S,
120°46' E, Banda Sea, 1158 m. Syntype zma.

Siphonodentalium booceras (Tomlin, 1926): 298, pi. 15, fig. 1 1. South Africa, Congella. sam and nmw
( fide Trew, 1990).

Siphonodentalium (?) delicatulum (Suter, 1913): 823, pi. 32, fig. 7. Southern New Zealand, Milford
Sound, 100-120 fms [183-220 m]. New Zealand Geological Survey, Wellington {fide Dance,
!986).

Siphonodentalium isaotakii Habe, 1953: 299. Tokyo Bay, Honshu, Japan, nsmt.

Siphonodentalium japonicum Habe, 1960: 294. Amakusa Island, Kyushu, Japan, nsmt.

Siphonodentalium longilobatum (Boissevain, 1906): 68. pi. 6, figs 55-56. Indonesia, Siboga , stn 133,
off Lirung, Salibabu Island, 36 m. 3 syntypes zma.

Siphonodentalium okudai Habe, 1953: 299, figs 759-760. Akkeshi Bay, Hokkaido. Japan, nsmt.

334

VICTOR SCARABINO

Genus Sagamicadulus Sakurai & Shimazu, 1963
Fig. 139 a

Type species (by monotypy): Striocadulus (Sagamicadulus) elegantissimus Sakurai & Shimazu, 1963.

Diagnosis. — Shell small to medium, thin, translucent white, glassy. Maximum diameter at
the anterior third. Surface smooth at apical portion, striated in median and anterior areas. Apex
round in section, with five lobes, two ventral and one flat, dorsal. Mouth ellipsoidal in section,
dorsoventrally compressed.

Radula unknown, assumed to be similar to that of Siphonodentalium.

Distribution. — Japan, shelf.

The genus Sagamicadulus is not represented in the material here reported on.

The type species is the only Indo-Pacific species: Sagamicadulus elegantissimus (Sakurai &
Shimazu, 1963): 250, textfig. 1. Sagami Bay, Japan, 55-128 m. nsmt.

Genus Striocadulus Emerson, 1962

Type species (OD): Cadulus albicomalus Dali, 1890. Recent, Ecuador, 401 fms [773 m].

Diagnosis. — Shell medium to large, strong, polished, white, maximum diameter in anterior
third. Sculptured with numerous close, fine, longitudinal striae throughout. Subcircular in section,
dorsoventrally compressed. Apex strong, with two weak, fiat lateral lobes, preapical callus prominent!
Radula similar to that of Siphonodentalium.

Distribution. — Recent, Pacific and Indian Ocean, absent in the Atlantic Ocean.
Shelf-bathyal.

Striocadulus pulchenimus (Boissevain, 1906)

Figs 136, 139 b

Cadulus pulcherrimus Boissevain, 1906: 74, pi. 6, figs 58-59.

Other reference:

Cadulus (Gadila) pulcherrimus - Habe & Kosuge, 1964: 11.

Type material. — Lectotype (here designated) zma 3.06.104, paralectotypes zma 3.06.105.

Type locality. — “Sihoga", stn 314, Flores Sea, 07°36' S, 1 1 7°3 1 ' E, 694 m.

Material examined. — The type material.

Indonesia, corindon: stn B 275, 01°54' S, 119°14'E, 530 m, 1 lv, 2 dd.

Distribution. — Indonesia, alive in 530-694 m.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

335

Striocadulus sagei sp. nov.

Figs 137, 139 d-e, 172 c-d

Type material. — Holotype and 4 paratypes mnhn.

Type locality. — Philippines, musorstom 2, stn CP 24, 13°37' N, 120°42' E, 640-647 m.

Material examined. — Indonesia. " Snellius ” II: stn 4.128, 08"18' S, 118°16'E, 700-835 m,
1 dd. — Stn 4.130, 08°18' S, 1 18°19' E, 700-730 m 3 lv, 9 dd. — Stn 4.131, 08°18' S, 118°18' E, 680-
800 m 1 lv, 2 dd (rmnh).

Philippines. MUSORSTOM 2: stn CP 24, 13°37' N, 120°42' E, 640-647 m, 5 lv (holotype and paratypes).
— Stn CP 25, 13°39' N, 120°43' E, 520-550 m, 3 dd.

Fig. 137. — Distribution of Striocadulus sagei.

Distribution. — The Philippines and Indonesia, alive in 640-800 m, shells from 550 m.

336

VICTOR SCARAB1NO

Description. — Shell to 43 mm long, strong, polished,
white, maximum diameter near the oral aperture. Longitudi¬
nal sculpture of numerous close, fine, flat round-section
riblets, crossed throughout by growth lines. Apex wide,
slightly dorsoventraily compressed, with two wide lateral

lobes. Oral aperture oblique, dorsoventraily compressed.

Measurements: holotype L 40. W 4.5. m 3.9, w 1.6, arc 3;
paratypes L 40, W 4.5, m 4, w 1.9, arc 3; L 43, W 4.6.1, m
4, w 1.8, arc 3; L 39.5, W 4.8. m 4.4, w 1.5, arc 3.5; L 40.2,
W 4.6, m 4, w 1 .7, arc 4.

Remarks. — This is the largest species in the order Gadilida. Siphonodentalium magnum
(Boissevain, 1906) has no sculpture and its apex is circular in section with traces of four flat, irregular
lobes. Other species of Gadilidae with longitudinal sculpture are S. striatulus (Pilsbry & Sharp, 1898)
and S. albicomatus (Dali, 1990) from Western North America, 5. pulcherrimus , S. lubrooki (here
described) and Sagamicadulus elegantissimus from Japan.

Etymology. — Named for Walter Sage (amnh), who kindly reviewed linguistically my
manuscript.

Striocadulus ludbrooki sp. nov.

Figs 138, 139 c

Cadulus (Polyschides) hexaschistus - Ludbrook, 1954: 118, fig. 14. [not C. hexaschisius Boissevain, 1906]

Type material. — Holotype and 3 paratypes bmnii 1952.3.25.342-345.

Type locality. — Gulf of Aden, "John Murray ”, stn 176, 12°05' N, 50°38' E, 655-732 m.

Material examined. — Northwestern Indian Ocean. "John Murray''-, stn 176, 12°05'N,
50°38' E, 655-732 m, 2 dd (holotype and paratype). — Stn 188, 13°46'N, 47°50' E, 528 m, 2 dd
(paratypes).

Distribution. — Northern Indian Ocean, shells in 528-732 m.

Description. — Shell to 20 mm, translucent, shiny, ventral observed in frontal view. Apex with two wide, flat lobes,
side regularly curved, dorsal with a swell at first fourth, also section oval, dorsoventraily compressed. Scultpure by longi-

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

337

tudinal striae, more prominent in first 3/4. Mouth oblique, Measurements: holotype L 19,5, W 3, m 2, w 1.4; paratype
subcircular in section. L 20, W 2.8, m 2.3, w 1.2.

Remarks. — This species is much less tapering than Striocadulus pulcherrimus and shorter and
less sculptured than S'. sagei, the other two Indo-Pacific species of the genus.

Etymology. — Named after Neils Ludbrook, who studied the scaphopods taken during the
“John Murray” expedition (1933-34) in the NW Indian Ocean.

Fig. 139. - a, Sagamicadulus elegantissiums (after Habe, 1964, pi. 5, figs 45-46), shell and apex. b, Striocadulus pulcherrimus,
shell (16 mm), lateral and dorsal views, apex, apical and oral sections, CORINDON: stn B 275. — c. Striocadulus ludbrooki,
holotype, shell (19.5 mm), lateral and dorsal views, apical and oral sections, apex. d, Striocadulus sagei sp. nov.,
holotype, shell (40 mm), apical section, apex, detail of sculpture, oral section. — e, Striocadulus type radula (S. sagei).

Genus Polysch/des Pilsbry & Sharp, 1898

Tvpe species (OD): Cadulus tetraschistus Watson, 1879. Recent, West Atlantic Ocean, " Challenger ", off Fernando de Noronha.
stn 113a. 03°47' S. 32°24' W, 7-25 fms [13-47 m].

Diagnosis. — Shell small to medium, strong, smooth, white, translucent when fresh, polished
when dead. Maximum diameter located in anterior third or in the mouth area. Apex wide, with four
deep notches and four lobes, two laterals, one dorsal, one ventral. Preapical callus weak.

Radula rachidian subpyramidal with wide base; lateral with two sharp cusps on inner surface, one on
outer face; marginal short.

Distribution. — Eocene- Recent, worldwide, littoral-bathyal.

338

VICTOR SCARABINO

Polyschides pelamide Chistikov, 1979
Figs 140, 147 a

Polyschides pelamide Chistikov, 1979b: 143, fig. 7.

Type material. — zin (fide Chistikov).

Type locality. — Gulf of Tonking, South China Sea, 56 m.

Material examined. — Indonesia, corindon: stn B 204, 0 1 ”09' S. 1 17" 18' E. 49 m, 1 lv, 8 dd.
Stn B 207, 00°15' S, 117°52' E, 150 m, 4 dd. — Stn B 251, 00°54' S, 119°30' E, 65 m, 4 dd.
Philippines. musorstom 3: stn DR 140. 11°43'N, 122°34' E, 93-99 m, 71 dd.

Fig. 140. Distribution of Polyschides pelamide.

Distribution. — Viet-Nam, now extended to the Philippines and Indonesia, alive in 49-56 m,
shells down to 150 m (present paper).

Polyschides arnaudi sp. nov.

Figs 141, 147 b, h

Type material. — Holotype mnhn. Paratypes: 13 mnhn, 1 nmp, 1 usnm.

Type locality. — West Indian Ocean, md 32 Reunion, stn DC 124, 20°52' S, 55°37' E,

40 m.

Material examined. — West Indian Ocean, md 32 Reunion: stn DC 10, 21" 13' S, 55"52' E,
930-980 m. 1 dd. — Stn DR 47, 21°23' S, 55°37' E, 205-215 m, 1 dd. — Stn DC 56, 21°05' S, 55°12' E,
170-225 m, 26 dd. — Stn DC 85, 2TW S, 55°15' E, 58-70 m, 7 dd. — Stn DC 86, 20°59' S, 55°15' E,
75-90 m, 7 lv, 38 dd. — Stn DR 90, I9°45' S, 54°09' E, 65 m, 3 dd. — Stn DC 124, 20°52' S, 55°37' E,
40 m, 13 lv, 129 dd (holotype dd, paratypes: 7 lv, 6 dd mnhn, 1 dd nmp, 1 dd usnm). — Stn DC 136,
20°46' S, 55°36' E, 915-922 m, 9 dd. — Stn DC 176, 21°02' S, 55°11' E, 165-195 m, 1 dd.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

339

Only known from Reunion Island. Alive in 40-90 m, shells (probably washed

Distribution.
down) to 980 m.

Description. Shell to 42 mm long, translucent, shiny,
smooth. Maximum diameter at anterior 1/8 of shell, swollen
at anterior aperture. Apex oval dorsoventrally, wide, with
four lobes created by four notches. The latero-dorsal notch is
deeper, forming a wide, flat dorsal lobe; two lobes are lateral;
the fourth is ventral, higher and posteriorly rounded. Prea-

pical callus prominent, lumen oval. Oral aperture circular,
slightly oblique.

Measurements: holotype L 7.5. W 1.2. w 0.9, apex 0.6;
paratypes L 6.4, W 1.2, w 0.7. apex 0.5: L 7.9, W 1.3, w 1.
apex 0.6; L 7.6, W 1.3, w 0.9, apex 0.5.

Remarks. The shell is characterized by alternating white and translucent rings throughout.
Polyschides arnaudi resembles the Western Atlantic P. tetrodon (Henderson, 1920), from which it
differs in having a less fusiform shell sape.

Etymology. Named for Dr Patrick M. Arnaud, Centre Oceanologique de Marseille.

Other Indo-Pacific species of Polyschides cited in the literature

Polyschides J'austus (Kuroda & Habe in Habe, 1971): 496 (Japanese text), 313 (English text), pi. 65.
figs 18-19. Sagami Bay. Japan.

Polyschides kaiyomaruae Okutani, 1975: pi. 3, tig. 5. N W Pacific, 29"49' N. 147' 09 E to 28 48 N,
147°09' E, 6200 m.

Polyschides sakuraii (Kuroda & Habe in Habe. 1962): 105, pi. 47, fig. 3. Off Choshi, Chiba Prefecture.
Honshu, Japan, 200 m. nsmt.

Polyschides vietnamicus Chistikov, 1979b: 113, fig. 6. Tonking Bay, Viet Nam. 8 m. zin.

Genus Dischides Jeffreys, 1867

Type species (by monotypy): Dentalium bifissum Wood. 1842. Recent. Mediterranean and East Atlantic; Pliocene of England
and Italy.

Diagnosis. - Shell small to medium sized, strong, smooth, white, translucent when fresh,
shiny when dead. Maximum diameter in anterior half of the shell or near the mouth. Apex wide, with

340

VICTOR SCARABINO

two deep lateral notches and two lobes, the ventral larger, usually with a central nodule in the inner
wall. Preapical callus weak.

Radula similar to that of Polyschides.

Distribution. — Eocene-Recent, worldwide, well represented in the Indo-Pacific region,
littoral-bathyal.

Dischides minutus (H. Adams, 1872)
Figs 1 1 1 n-o, 142, 147 c

Cadulus minutus H. Adams, 1872: 10, pi. 3, fig. 9.

Other references:

Dentalium minutus — Sowerby, 1873: pi. 7, fig. 48. — Cooke, 1885: 273.

Cadulus minutus - Pilsbry & Sharp, 1897: 188, pi. 26, fig. 78. - Boissevain, 1906: 67, pi. 3, fig. 49.

Type material. — Presumably in bmnh (not located).

Type locality. — Red Sea.

Material examined. — Red Sea. Suez, 145 dd. — Gulf of Suez, 15 dd. Souakin, Sudan,
3 dd. — Djeddah, 1 dd. — Aden, 65 dd (all Coll. Jousseaume, mnhn).

Fig. 142. — Distribution of Dischides minutus.

Distribution. — Red Sea and Gulf of Aden. Depth distribution unknown, shells probably
washed ashore.

Dischides dichelus (Watson, 1879)
Figs 143, 147 e

Siphonodentalium dichelum Watson, 1879: 521; 1886: 15, pi. 2, fig. 7.

Other references:

Cadulus dichelus - Pilsbry & Sharp, 1898: 145, pi. 26, fig. 73. Boissevain, 1906: 65, pi. 3, fig. 48, pi. 6, fig. 51.
C. ( Dischides ) dichelus - Habf. & Kosuge, 1964: 11.

Source : MNHN , Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

341

Type material. Holotype bmnh 1887.2.9.65.

Type locality. ‘‘Challenger", Levuka, Fiji, 12 fms [22 m].

Material examined. — The type material.

West Indian Ocean, md 32 Reunion: stn DC 56, 21°05' S, 55° 12' E, 170-225 m, 2 lv, 254 dd. — Stn
DC 85, 21°00' S, 55°15' E, 58-70 m, 2 dd. — Stn 90, 19°45' S, 54°09' E, 65 m, 1 dd. — Stn CP 129,
20°51' S, 55°36' E, 290-300 m, 1 dd.

NW Madagascar, Nosy Be Island, Plante coll., 1 lv, 2 dd (bmnh).

Distribution. — Fiji and Indonesia (Boissevain, 1906), now extended to Reunion Island and
Madagascar; alive in 170-225 m and in indeterminate condition up to 22 m.

Dischides prionotus (Watson, 1879)

Figs 144, 147 d, i

Siphonodentalium prionotum Watson. 1879: 522; 1886: 16, pi. 2, fig. 9.

Other references:

Cadulus prionotus — Pilsbry & Sharp. 1898: 146, pi. 26, fig. 74. — Boissevain, 1906: 66, pi. 3, fig. 47.

Type material. — 3 syntypes dd bmnh 1907.10.28.147-149.

Type locality. — “ Challenger ", stn 185b, 11°38'S, 143°59' E, 155 fms [283 m], off Cape
York, N Australia.

Material examined. — The type material.

New Caledonia. " Vauhan ” 1978-79: stn 33, 22°33' S, 166°25'E, 290-350 m. 1 dd.

West Indian Ocean, benthedi: stn DR 08, 11°29'S, 47°18'E, 250 m, 5 lv, 4 dd. — Stn DS 10,
11°29'S, 47° 18' E, 440 m, 1 lv, 1 dd. — Stn DS 64, 12°41'S, 44°57' E, 770-860 m, 1 lv. — Stn
DS 72, 1 2°3 1 ' S, 45°02' E, 300-350 m, 1 lv, 2 dd. — Stn DS 94, 1 1°32' S. 47°16' E, 450 m, 1 lv.

Stn DS 120, 1 1°30' S, 47°25' E, 335-390 m, 5 lv, 3 dd.

Source : MNHN, Paris

342

VICTOR SCARABINO

MD 32 Reunion: stn DC 85, 20°59' S. 55°15' E, 58-70 m. 2 dd. — Stn DC 128, 20°51' S, 55°36' E,
280-340 m, 1 dd.

Distribution. — N Australia, now extended to New Caledonia, Reunion Island and NW
Madagascar, alive in 250-450 m and shells from 70 m.

Dischides viperidens (Melvill & Standen, 1896)

Figs 145, 147 f

Cadulus viperidens Melvill & Standen, 1896: 314, pi. 11, fig. 79.

Type material. — Figured syntype Manchester Museum EE 3797; 6 other syntypes nmw
(Trf.w. 1987).

Fig. 145. - Distribution of Dischides viperidens.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

343

Type locality. — Lifu and Uvea, Loyalty Islands, New Caledonia.

Material examined. — Loyalty Islands. Niemak, Ouvea, beach, 27 July 1978, P. Bouchet
coll., 25 dd.

Distribution. — Loyalty Islands, shore.

Dischides yateensis sp. nov.

Figs 146, 147 g

Type material. — Holotype and paratype mnhn.

Type locality. — New Caledonia, lagon, stn 619, 22°03' S, 166°54' E, 27-42 m.

Material examined. — New Caledonia, lagon: stn 619, 22°03' S, 166°54' E, 27-42 m, 2 dd
(holotype and paratype).

Philippines. Coll. Jousseaume, 6 dd, mnhn.

Distribution. — New Caledonia and the Philippines. Shells from the shore to 42 m.

Description. — Shell to 13 mm long, strong, shiny,
translucent white. Maximum diameter in anterior fifth, where
a slight swelling originates near the mouth and rapidly curves
towards the peristome; growth lines pronounced. Apex
strong, wide, dorsoventrally compressed, with two lateral
notches forming two lobes, the dorsal with a rounded edge

and the ventral angled with a prominent central denticle.
Preapical callus prominent, lumen circular. Mouth straight,
oval dorsoventrally.

Measurements: holotype L 12.4. W 2-1.9, m 1.5-1. 4, apex
0.9-0. 8, arc 0.5: paratvpe L 12.1, W 2, m 1.8-1.65, apex
0.9-0. 8, arc 0.5.

Remarks. — This species resembles Dischides prionotus, which has similar sculpture but
different shape, and D. dichelus, which is fusiform in outline. The three species share the denticle
situated on the ventral lobe of the apex.

Etymology. — Named for Yate, the type locality.

344

VICTOR SCARABINO

Fig. 147. a, Polyschides pelamide, shell (11.5 mm), apical and oral sections, musorstom 3: stn DR 140. — b, Polyschides
arnaudi sp. nov., holotype, shell (7.5 mm) lateral and dorsal views, apex and apical section. — c, Dischides minutus, shell
(5 mm), apex, apical and oral sections, Aden (mnhn). - d, Dischides prionotus , shell (12 mm), lateral and dorsal views,
apex and apical sections, benthedi: stn DS 120. — e, Dischides dichelus, shell (9 mm), lateral and dorsal views, apex,
apical and oral sections, md 32 Reunion: stn DC 56. — f, Dischides viperidens, shell (6 mm), dorsal and lateral views,
apex, apical and oral sections, Loyalty Islands (mnhn). — g, Dischides yateensis sp. nov., holotype, shell (12.4 mm),
dorsal and lateral views, apex, apical and oral sections. — h. Polyschides type radula (/’. arnaudi). — i, Dischides type
radula (D. prionolus).

Source : MNHN , Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

345

Subfamily Gadilinae Stoliczka, 1868
Genus Cadulus Philippi, 1844

Type species (by monotypy): Cadulus ovulum Philippi, 1844. Recent, Mediterranean Sea.

Diagnosis. — Shell small to medium, swollen, solid, smooth, white, translucent when fresh,
shiny when dead. Maximum diameter in center of shell. Apex simple or coronate, preapical callus
usually prominent, lumen circular. Apical and oral sections variable, usually oval dorsoventrally or
laterally compressed.

Radula rachidian variable, polygonal to almost quadrangular, anterior margin generally with cusps;
lateral with two to three primary cusps with a series of denticles between cusps; marginal short,
slightly curved (Fig. 160 k, C. tumidosus Jeffreys, 1877).

Distribution. — Cretaceous- Recent, worldwide, shelf-abyssal.

Cadulus simillimus Watson, 1879
Figs 148, 160 a

Cadulus simillimus Watson, 1879: 526: 1886: 20, pi. 3, fig. 6.

Other references:

Cadulus simillimus - Boissevain, 1906: 69, pi. 3, fig. 46.

Cadulus (Gadila) simillima - Habe & Kosuge, 1964: 11.

Type material. — 2 syntypes dd, bmnh 1887.2.9.80-81.

Type locality. — Australia, off Cape York, Queensland, “ Challenger ”, stn 185b, 11°38'S,
143°59' E, 155 fms [283 m].

Material examined. — New Caledonia, biogeocal: stn KG 201, 22°40' S, 166°33' E. 595 m,
1 dd. — Stn KG 219, 22°39' S, 166°34' E, 570 m, 2 dd.

North Australia. Port Darwin, Coll. Denis, 2 dd (mnhn).

Fig. 148. — Distribution of Cadulus simillimus.

346

VICTOR SCARABINO

Distribution. Northern Australia, now extended to New Caledonia, shells from 6 fms
[11 m] (Watson, 1879) to 595 m (present paper).

Cadulus aratus Hedley, 1899
Figs 149, 160 b

Cadulus aratus Hedley, 1899b: 551, fig. 60.

Type material. — Syntypes ams C5635, 36, 38.

Type locality. Pacific Ocean, Tuvalu, Funafuti Atoll, 66 m.

Material examined. The type material.

New Caledonia, lagon: stn 729, 21°194 S. 165°54' E, 42-45 m. 1 dd.

Philippines, musorstom 2: stn DR 33, I3°32'N. 121°08' E, 130-137 m, 3 dd.
West Indian Ocean, benthedi: stn DS 120, 1 l°30' S, 47°25' E, 335-390 m, 10 lv.
md 32 Reunion: stn DS 178, 21°04' S. 55°10' E. 412-460 m, 2 lv.

Distribution. — Funafuti Atoll, now extended to New Caledonia, the Philippines, Reunion
Island and NW Madagascar, alive in 335-460 m. shells recorded as shallow as 45 m.

Cadulus aequatorialis Jaeckel, 1932
Figs 150, 160 c

Cadulus aequatorialis Jaeckel, 1932: 311, textfig. 10.

Other reference:

Cadulus (C.) aequatorialis - Habe & Kosuge, 1964: 9.

Type material. — Lectotype (Kilias, 1972) zmb 75372, paratype 75373.

Type locality. — Indonesia, West Sumatra, “Valdivia", stn 191, 00°39' S, 98°52' E, 750 m.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL 1NDO-PACIFIC

347

Material examined. — New Caledonia, biogeocal: stn CP 214, 22°43' S, 166°28' E,
1590-1665 m, 1 dd. — Stn KG 221, 22°42' S, 166°24' E, 1915 m, 1 lv.

Distribution. — Indonesia, now extended to New Caledonia, alive in 1915 m, shells from

750 m.

Cadulus cyathoides Jaeckel, 1932
Figs 151, 152, 160 d

Cadulus cyathoides Jaeckel, 1932: 308, textfig. 5.

Other reference:

Cadulus (C.) cyathoides - Ludbrook, 1954: 112, fig. 12.

Type material. — Lectotype, designated by Kilias (1995), zmb 75361a.

Type locality. — Indonesia, West Sumatra, “ Valdivia ”, stn 191, 00°39' S, 98°52' E, 750 m.

Fig. 151. — Cadulus cyathoides, detail of the coronate apex
common to all globose Cadulus. Scale line: 100 pm.

348

VICTOR SCARABINO

Material examined. — Philippines, estase 2: stn CP 2, 14°05' N, 120°02\ 2050 m, 2 dd.
musorstom 3: stn CP 143, 11°29'N, 124°11'E, 205-214 m, 1 lv.

West Indian Ocean, benthedi: stn 42, 13°05' S, 45°08' E, 400-520 m, 1 lv.

Fig. 152. — Distribution of Cadulus cyathoides.

Distribution. — Red Sea, Gulf of Aden (Ludbrook, 1954) and Indonesia, now extended to
the Philippines, alive in 205-520 m and shells down to 2050 m (present paper).

Cadulus chuni Jaeckel, 1932
Figs 153, 160 e

Cadulus chuni Jaeckel. 1932: 309, textfig. 6.

Other reference:

Cadulus (C.) chuni - Habe & Kosuge, 1964: 10.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

349

Type material. — Lectotype, designated by Kilias (1995), zmb 75374a.

Type locality. — East Africa, off Somalia, “ Valdivia ”, stn 256, 01°49' N, 45°30' E, 1134 m.
Material examined. — Indonesia. “ Snellius ” II: stn 4.112. 08° 1 9' S, 1 1 8°1 6' E, 365 m. 20 lv,

18 dd.

Philippines, musorstom 3: stn CP 139, 11°53'N, 122°14'E, 240-267 m, 1 lv, 1 dd. Stn CP 143,
11°29'N, 124°11'E, 205-214 m, 3 lv, 15 dd.

Distribution. — East Africa, now extended to Indonesia and the Philippines, alive in 205 to

365 m.

Cadulus martini sp. nov.

Figs 154, 160 f

Type material. Holotype and 6 paratypes mnhn.

Type locality. — New Caledonia, biogeocal, stn CP 232, 21°34' S, 166°27' E, 760-790 m.

Material examined. - New Caledonia, biogeocal: stn CP 232, 21°34' S, 1 66°27' E, 760-
790 m, 1 lv (paratype), 3 dd (holotype, 2 paratypes).

biocal: stn CP 75, 22°19' S, 167°23' E, 825-860 m, 1 lv (paratype). Stn DW 106, 21°36' S,
1 66°29' E, 625-650 m, 1 lv, 1 dd (paratypes).

West Indian Ocean, md 32 Reunion: stn DS 149, 20°26' S, 55°40' E, 3500-3510 m, 1 dd.

Fig. 154. — Distribution of Cadulus martini.

Distribution. — New Caledonia and Reunion Island, living in 625-860 m and shells
(probably washed down) down to 3500 m.

Description. — Shell to 5 mm long, globose, shiny, white.
Maximum diameter posterior to center of shell. Apical
constriction shorter than oral. In lateral view, the ventral side
shows a regular and pronounced curve to the neck. The
dorsal side shows alternating concave and convex lines. The

oral concave curve reaches maximum depth at 2/5 of the
shell; the apical curve at the posterior 1/5. In frontal view, the
posterior area from the maximum diameter is shorter and
more curved than the anterior area which has straight sides
tapering regularly to the mouth. Apex oval dorsoventrally.

350

VICTOR SCARABINO

with coronate structure. Preapical callus conspicuous, lumen Measurements: holotype L 5, W 1.9, m 1-0.8. apex 0.7-0. 6:
oval dorsoventrally. Mouth laterally compressed, curved on paratypes L 5.5. W 2-2, m 1-0.9, apex 0.7-0. 6; L 5, W 2-1.9,
ventral side, oblique. m 1-0.8, apex 0.7-0. 6; L 5.3, W 2-1.9, m 0.9-0. 8. apex 0.7-0. 6.

Remarks. — A coronate structure of the apex is common among the globose Cadulus, and can
be noticed in the apical part of the animal (Scarabino, 1979).

Etymology. Named for the author’s son Martin.

Cadulus glans sp. nov.

Figs 155, 160 g

Type material. - Holotype and 7 paratypes mnhn.

Type locality. - New Caledonia, Loyalty Basin, biogeocal, stn KG 227, 21°33'S,
166°24' E, 500 m.

Material examined. — New Caledonia, biogeocal: stn KG 219, 22°39' S, 166°34' E, 570 m,
2 dd (paratypes). — Stn KG 227, 21°33' S, 166°24' E. 500 m. 1 dd (holotype).

“Vauban” 1978-79: stn 40, 22°30' S, 166°24' E, 250-350 m, 5 dd (paratypes).

Distribution. Only known from New Caledonia, shells from 250-570 m.

Description. — Shell to 2 mm long, globose, glossy,
translucent white. Equator posterior to the center of the shell.
Apical constriction very short, formed by 11-12 lobes. In
lateral view, the ventral side is straight to the center of the
shell, then curves regularly and strongly to the apex; the
dorsal side is sinusoidal, with the concave sector short near
the mouth, with a long convex curve that reaches its
maximum diameter at the posterior quarter. In frontal view

the posterior area from the center is short and more curved
than the anterior area, which has straight sides regularly
tapering to the mouth. Apex oval laterally, larger than the
mouth, with prominent coronate structure. Preapical callus
prominent, lumen circular placed ventrally by the callus.
Mouth circular, oblique.

Measurements: holotype L 1.7, W 0.89, m 0.44, apex 0.47.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

351

Remarks. The coronate structure is formed by 11-12 wide lobes. The most similar species
is Cadulus martini , from which C. glans differs in being the only species with the apex, rather than
the lumen, larger than the mouth.

Etymology. From the latin glans : acorn-shaped.

Cadulus florenciae sp. nov.

Figs 156, 160 h

Type material. — Holotype and 3 paratypes mnhn.

Type locality. — West Indian Ocean, md 32 Reunion, stn DS 139, 20°47' S, 55°38' E,
1575-1600 m.

Material examined. — West Indian Ocean, md 32 Reunion: stn DR 104, 20°49' S, 55°01' E.
1875-1920 m, 3 dd (paratypes). — Stn DS 139. 20°47' S, 55°38' E. 1575-1600 m. 1 lv (holotype).

Distribution. — Only known from off Reunion Island, alive in 1575-1600 m.

Description. Shell to 2 mm long, delicate, fusiform,
shiny, white, translucent. Maximum diameter at center of
shell. The ventral side shows a regular curvature, less
prominent in oral area; the dorsal side shows a slight

Remarks. — The most similar species is
shorter oral area and neck.

swelling. Apex simple, oval dorsoventrally. Preapical callus
conspicuous, lumen oval. Mouth suboval laterally compres
sed.

Measurements: holotype L 2, W 0.6, mouth 0.2, apex 0.2

Cadulus aequatorialis , which differs in having .>

Etymology. — Named for the author's daughter, Florencia.

352

VICTOR SCARABINO

Cadulus sofiae sp. nov.

Figs 157, 160 i, 1

Type material. — Holotype mnhn. Paratypes: 7 mnhn, 1 nmp.

Type locality. — West Indian Ocean, md 32 Reunion, stn DS 151, 20°5T S, 56°03' E,
3240-3300 m.

Material examined. — New Caledonia, biocal: stn KG 03, 21°15'S, 166°39' E, 2340 m,
2 dd. — Stn KG 71, 22°10' S, 167°33' E, 2099 m, 1 dd. Stn KG 102, 21°28' S, 166°26' E, 1810 m,
1 dd.

biogeocal: stn CP 243, 21°27' S, 166°26'E, 1820 m, 3 dd. — Stn CP 260, 21°00' S, 166°58' E,
1820-1980 m, 4 dd. — Stn KG 276, 21° 13' S, 167°00' E, 2200 m, 1 dd. Stn CP 317, 20°48' S,
166°53' E, 1620-1630 m, 2 dd.
calsub: dive 13, 21°26' S, 166°23' E, 1600 m, 3 dd.

West Indian Ocean, md 32 Reunion: stn DS 149, 20°26' S, 55°40' E, 3500-3510 m, 1 lv (paratype nmp).
— Stn DS 151, 20°5 1 ' S, 56°03' E, 3240-3300 m, 4 lv (holotype and 3 paratypes).
benthedi: stn DS 11, 12°16' S, 46°42' E, 2300-2450 m, 1 dd (paratype). — Stn CH 87, 11°44'S,
47°35' E, 3716 m, 6 lv (2 paratypes). — Stn CH 90 11°44' S, 47°30' E, 3700 m. 1 dd (paratype).

Fig. 157. — Distribution of Cadulus sofiae.

Distribution. — Reunion Island, NW Madagascar and New Caledonia, alive in 3500 to
3716 m, shells from 1600 m.

Description. — Shell to 5 mm, globose, white, shiny,
translucent in fresh specimens. Maximum diameter at center
of shell. In lateral view the swell is prominent on ventral side;
in frontal view, both sides are similar. Rapidly tapering from
the maximum diameter to apex and mouth. Circular in

section. Apex simple, oval dorsoventrally. Preapical callus
conspicuous, lumen circular. Mouth laterally compressed,
oblique.

Mesurements: holotype L 4.5, w 2-2, d 2, m 1.9-1, 7, arc 1.

Remarks. — This species is similar in shape to Cadulus tumidosus Jeffreys, 1877, from the
Atlantic Ocean, but less globose.

Etymology. — Named for the author’s daughter Sofia.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

353

Cadulus labeyriei sp. nov.

Figs 158, 160 j

Type material. — Holotype mnhn. Paratypes: 9 mnhn, 1 usnm.

Type locality. — Philippines, estase 2, stn DW 1, 14°05' N, 120°01'E, 2200 m.

Material examined. — Philippines, estase 2: stn DW 1, 14°05' N, 120°01' E, 2200 m, 3 lv
(holotype and 2 paratypes). — Stn CP 2, 14°05' N, 120°02' E, 2050 m, 8 lv (paratypes: 7 mnhn,
1 usnm).

Fig. 158. — Distribution of Cadulus labeyriei.

Distribution. — Only known from the Philippines, living from 2050 to 2200 m.

Description. — Shell to 3 mm long, delicate, globose,
shiny, white, translucent to transparent in fresh specimens.
Maximum diameter near center of shell, oval in section,
prominent. In lateral view, the ventral side shows regular
curvature, while on the dorsal side the swelling is prominent
and shows oral and apical constrictions. In front view, the
maximum diameter is at the center of the shell, the constric¬

tion through mouth and the apex begins straight, near the
apex the slight curvature forms a short neck. Apex simple,
oval dorsoventrally. Preapical callus conspicuous, lumen
circular. Mouth suboval laterally compressed, oblique.

Measurements: holotype L 2.7, W 1.2, m 0.4. apex 0.4;
paratype L 3.1, W 1.3, m 0.4, apex 0.2.

Remarks. — The most similar species is Cadulus valdiviae , which differs in its shorter apical
and oral constrictions and is more swollen.

Etymology. Named for Dr Laurent Labeyrie (cnrs, Gif), cruise leader of the estase 2
Expedition aboard R.V. “ Jean-Charcot

Other Indo-Pacific species of Cadulus cited in the literature

Cadulus campylus Melvill, 1906: 80, pi. 8, Fig. 32. Gulf of Oman, 285 m. Syntype bmnh 1906.10.23.76,
and 3 syntypes nmw (fide Oliver, 1984) (not seen).

354

VICTOR SCARABINO

Cadulus euloides Melvill & Standen, 1901: 459, pi. 24, fig. 24. Gulf of Oman. Syntype bmnh

1901.12.9.12.

Cadulus oval is Boissevain, 1906: 66, pi. 6, fig. 52. Madura Bay. Indonesia, " Siboga ", stn 51, 69-91 m.

2 syntypes dd zma. _

Cadulus platei Jaeckel, 1932: 311, fig. 9. “ Valdivia ", stn 109, off South Africa, 35 19 S, 20 12 E,

126 m. Holotype zmb 75376. ,

Cadulus siberutensis Jaeckel, 1932: 309, fig. 7. Indonesia, Sumatra, '‘Valdivia", stn 191, 00' 39 S,
98°52' E, 750 m. Lectotype (Kilias. 1995) zmb 75368a.

Cadulus singaporensis Pilsbry & Sharp. 1898: 195, pi. 36, figs 30-32. Singapore, ansp. ,

Cadulus valdiviae Jaeckel, 1932: 312, fig. 11. “ Valdivia ", stn 251. off Somalia, 01°41' S, 41"47 E,

693 m. Lectotype (Kilias, 1995) zmb 75370a.

Genus Bathycadulus gen. nov.

Type species: Bathycadulus fabrizioi sp. nov.

Diagnosis. — Shell medium sized for the family, smooth, shiny. Maximum diameter at the 3/5
median zone, where sides are almost parallel, or in two swellings, one at 2/5 and the other at 4/5 of
the total length. Oral and apical constrictions pronounced, with short anterior and posterior areas.
Apex simple or with a series of narrow notches forming a coronate edge. Section at apex slightly
dorsoventrally compressed with a conspicuous preapical callus. Mouth oblique, laterally compressed.
Apical area of animal short with principal muscular ring simple or coronate, neck with conspicuous
dermic papillae.

Radula rachidian nearly as high as wide, anterior edge irregular, with a small cusp at the center and
two or three denticles at eaclfside. Lateral with well sculptured head, having three major cusps, one
located internally, the other two externally, and 4-5 denticles; sculpture of the head of laterals
accompanied by a chain of tooth like formations located at the outer side of primary cusps. Marginal
slightly sigmoidally depressed, with the union edge to the lateral large and a prominent lateral keel.

Remarks. The subfamily Gadilinae includes the genera Cadulus, Gadila (radula very
similar) and Sulcogadila Moroni & Ruggieri, 1981 (fossil from the Pleistocene of Sicily). Cadulus and
Sulcogadila have the maximum diameter centrally located in one swelling with concave sides, and
Gadila has the equator near the oral aperture, and usually concave dorsal and ventral sides.
Bathycadulus has unique shell characters for the class, with sides almost parallel and even slightly
concave at the maximum diameter.

Two additional new species of this genus, from bathyal and abyssal depths of the Atlantic
Ocean, will be described elsewhere.

Distribution. — Recent, Indian and Atlantic Oceans, abyssal.

Bathycadulus fabrizioi sp. nov.

Figs 159. 160 m-n

Type material. — Holotype sam A36258. Paratypes: 1 sam A36259, 1 usnm, 1 1 mniin, 1 nmp.
1 Museo Nacional de Historia Natural, Montevideo, Uruguay 14751.

Type locality. — South Africa, “ Meiring Naude”, stn SM 109. 28°4T S, 32°37' E, 1300 m.

Material examined. New Caledonia, biogeocal: stn KG 221, 22°42' S, 166"24' E, 1915 m.
1 dd. Stn KG 222, 22°45' S, 166°25' E, 1675 m, 1 dd.

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

355

West Indian Ocean, md 32 Reunion: stn DS 78, 21°13'S, 55°04' E, 1175-1200 m, 1 lv, 1 dd.

Stn DS 106, 20°48' S, 55°05' E, 1710-1730 m, 1 lv. — Stn DS 109, 20°52' S, 55°06' E, 1050-1240 m,
12 lv (paratypes: 11 mnhn, I usnm), 1 dd (paratype nmp). — Stn DS 139, 20°47' S, 55°38' E,
1575-1600 m, 8 lv, 2 dd.

“Meiring Naude stn SM 109, 28°41' S, 32°37' E, 1300 m, 2 lv (holotype and 1 paratype sam), 1 dd
(paratype Montevideo).

Fig. 159. — Distribution of Baihycadulus fabrizioi.

Distribution. — Southeast Africa, Reunion Island and New Caledonia, alive in 1200-1600 m,
shells down to 1915 m.

Description. — Shell to 7 mm long, shiny, translucent
to porcellaneous white. Maximum diameter at 3/5 of
shell length, sides parallel. Neck and mouth with a short
constriction. Apex simple with irregular coronate edge;
slightly dorsoventrally compressed in section, with a promi¬

nent preapical callus. Oral aperture simple, laterally compres¬
sed.

Riulula as for the genus.

Measurements: holotype L 5.2. W 1.05. w 0.8. apex 0.8. arc
(at the equator) 0.1.

Etymology. — Named for the author's son Fabrizio.

Genus Gadila Gray, 1847

Type species (OD): Dentalium gadus Montagu, 1803. Recent, type locality unknown.

Synonym: Platyschides Henderson, 1920. Type species (OD): Cadulus grandis Verrill, 1885. Recent, NW Atlantic Ocean.
906 fms [1703 m].

Diagnosis. — Shell small to medium, curved, smooth, white, translucent when fresh, shiny
when dead. Maximum diameter in anterior third of the shell; ventral side regularly curved, dorsal side
sigmoidal in section. Apex simple or with flat lobes, variable in number. Oral and apical section
different.

Radula similar to that of Cadulus (Fig. 169 h, Gadila sp.).

Distribution. — Cretaceous-Recent, worldwide, littoral-abyssal.

356

VICTOR SCARAB1NO

Gadila virginalis (Boissevain, 1906)
Figs 161, 169 a

Cadulus virginalis Boissevain, 1906: 72, pi. 6, figs 60-64.
Other references:

Type material. — Lectotype (here designated) zma 3.06.096, paralectotypes zma 3.06.097-

100.

Type locality. — C. virginalis: Indonesia, Savu Sea, "Siboga , stn 52, 09 03 S, 119 57 E,

959 m.

Material examined. — The type material.

New Caledonia, biocal: stn DW 49, 23°03' S, 167°33' E, 825-830 m, 1 dd.
Indonesia, corindon: stn B 236, 00°07' S, 119°45' E. 1730 m, 2 dd.
Philippines. musorstom 3: stn DR 93, 13°49'N, 120°02' E, 540 m, 1 dd.

Fig. 161. — Distribution of Gadila virginalis.

Distribution. — Southern Japan (?) to Indonesia, now extended to New Caledonia. Shells in
200-3010 m (Habe & Kosuge, 1964).

Fig. 160. a, Cadulus simillimus, shell (3 mm), lateral and dorsal views, apical and oral sections, biogeocal: stn KG 219
— b. Cadulus aratus. shell (2 mm), apical and oral sections, musorstom 2: stn DR 33. c. Cadulus aequatorialis shell
(4 mm), apical and oral sections, biogeocal: stn CP 214. d, Cadulus cyathoides, shell (2.5 mm), lateral and dorsal
views, apical and oral sections, estase 2: stn CP 2. e, Cadulus chuni, shell (2.5 mm), apical and oral sections,
musorstom 3: stn CP 143. - f, Cadulus martini sp. nov., holotype, shell (5 mm), apical and oral sections. g, Cadulus
glans sp. nov., holotype, shell (1.7 mm), lateral and dorsal views, apical and oral sections. h , Cadulus Jlorenciae sp.

nov., holotype, shell (2 mm), lateral and dorsal views, apical and oral sections. i, Cadulus sofiae sp. nov., holotype,
shell (4,5 mm), lateral and dorsal views, apical and oral sections. j, Cadulus labeyirei sp. nov., holotype, shell
(2.7 mm), lateral and dorsal views, apical and oral sections. — k-I, Cadulus type radula. — k, C. tumidosus Jeffreys,
1877 (North Atlantic Ocean). 1, C. sofiae. — m. Bathycadulus fabrizioi sp. nov., holotype, shell (5.2 mm), apical and
oral sections. — n, Bathycadulus type radula (B. fabrizioi).

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

357

Source : MNHN, Paris

358

VICTOR SCARABINO

Remarks. — Habe (1963, 1964b, 1977) includes Gadila novilunata Kira, 1959 in the synonymy
of virginalis, but I prefer to keep them separate pending further study. Consequently, it is uncertain
whether the records of Cadulus virginalis from Japan refer to the present species or to G. novilu¬
nata.

Gadila zonata (Boissevain, 1906)

Figs 162, 169 b

Cadulus zonatus Boissevain. 1906: 74. pi. 6, fig. 57, textfig. 37.

Type material. — Lectotype (here designated) zma 3.06.102, paralectotypes zma 3.06.103.
Type locality. — Indonesia, Banda Sea, “Siboga", stn 214, 06°30' S, 121°55' E, 2796 m.
Material examined. — The type material.

Philippines. musorstom 2: stn CP 18, 14°00' N, 120° 18' E, 188-195 m, 1 dd. — Stn CP 55, 13°54' N,
1 19°58' E, 865 m, 1 dd.

Distribution. — Indonesia and the Philippines, shells from 195 to 2796 m. no record of living
specimens.

Gadila boissevainae (Jaeckel, 1932)
Figs 163, 169 c

Cadulus boissevain i Jaeckel, 1932: 311, textfig. S.

Synonym:

Cadulus (Gadila) reesi Ludbrook, 1954: 115, fig. 20 (Syn. nov.).
Other reference:

Cadulus ( Cadulus ) boissevaini — Habe & Kosuge, 1964: 9.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

359

Type material. — C. boissevaini: lectotype, designated by Kilias (1995), zmb 75359a.

C. reesi: holotype bmnh 1952.3.25.150.

Type locality. C. bossevaini : East Africa, off Zanzibar, “ Valdivia ", stn 251, 01°41' S.
4 1 °47' E, 693 m. C. reesi : Zanzibar area, “John Murray’’, stn 105B, 05°34' S, 39°14' E, 238-293 m.

Material examined. — The type material of C. reesi.

Northwest Indian Ocean. Gulf of Aden, “John Murray stn 176, 12°04' N, 50°38' E, 655-732 m, 3 dd.

Stn 179b, 1 2°02' N, 50"40' E, 275 m, I dd. — Stn 191, 13°46' N, 47°49' E, 274 m, 7 dd (all bmnh)
West Indian Ocean. “Meiring Naude”: stn SM 53, 26°51' S, 33°13' E. 720 m, 1 lv, 5 dd. — Stn 59,
27° 10' S, 32°59' E, 820 m, 8 lv, 35 dd (1 lv, 1 dd mnhn). — Stn SM 69, 27°14' S, 33°12' E 680-
700 m, 1 lv, 1 dd. Stn SM 78, 27°32' S, 32°50' E, 750 m, 9 lv, 26 dd. — Stn SM 94, 28°50' S,
32°50' E, 750 m, 3 dd. — Stn SM 125, 30°32' S, 30°57' E, 1280 m, 1 lv, 6 dd. Stn 162, 32°55' S.
28°3 1 ' E, 630 m, 1 dd.

Distribution. — Western Indian Ocean from the Gulf of Aden to Natal, alive in 680-1280 m,
shells up to 274 m.

Remarks. — Because Maria Boissevain was a woman, the specific name is here emended to
boissevainae (iczn. Art. 3 1 (a)(i)). The specimens of Cadulus reesi are all fragments, without the neck
area. The rest of the shell corresponds to Gadila boissevainae and leads me to consider it a junior
synonym of Jaeckefs species.

Gadila elenae sp. nov.

Figs 164, 169 d

Type material. — Flolotype mnhn. Paratypes: 7 mnhn, 1 ams C201733, 1 nmnz M 268956.
Type locality. - New Caledonia, musorstom 4, stn DW 156, 18°54'S, 163°19' E. 525 m.

360

VICTOR SCARABINO

Material examined. — New Caledonia, biocal: stn DW 08, 20°34' S, 166°54' E, 435 m, 2 dd
(paratypes).

musorstom 4: stn DW 156, 18°54' S, 163° 19' E, 525 m, 1 lv (holotype), 2 dd (paratypes). — Stn DW
160. 18°42'S. 163°13'E, 668 m, 1 lv, 2 dd (paratypes: 1 mnhn, 1 ams, 1 nmnz).

Loyalty Islands, musorstom 6: stn DW 410, 20°38' S, 167°07' E, 490 m, 2 dd (paratypes).

Distribution. — New Caledonia, living from 525 to 668 m, shells from 435 m.

Description. — Shell to 12 mm long, smooth, shiny,
translucent white. Ventral side showing a regular curve,
dorsal side sinusoidal with convex curve at first quarter and
concave curve rising to the apex. Maximum diameter near the
anterior quarter of the shell, suboval in section. Apex oval,
dorsoventrally depressed with ventral and dorsal lobes, the

former larger and with a central denticle-like structure.
Preapical callus prominent, lumen circular. Mouth oblique,
subcircular in section.

Measurements: holotype L 11.5, W 2. 5-2. 3, m 1.35-1.20,
apex 1-1.8; paratype L 10.1, W 2.3-2. 1, m 1.3-1. 2, apex
0.9-0.85; L 8.5, W 2.3-2.2, m 1.6- 1.4, apex 1.50-1.51.

Remarks. — The apical structure is similar to that of Gadila monodonta, described below,
which is larger and fusiform.

Etymology. — Named for Elena Gofas-Salas (Paris, Malaga).

Gadila doumenci sp. nov.

Figs 165, 169 e

Type material. — Holotype and 3 paratypes mnhn.

Type locality. — West Indian Ocean, benthedi, stn DS 1 1, 12°16' S, 46°42' E, 2300-2450 m.

Material examined. — New Caledonia, biogeocal: stn CP 260, 21°00' S, 166°58' E,
1820-1960 m, 1 dd.

West Indian Ocean, benthedi: stn DS 03, 12°36' S, 47°38' E, 1100-1150, 1 lv (paratype). — Stn
DS 11, 12°16'S, 46°42'E. 2300-2450 m, 1 lv (holotype). — Stn DR 40, 12°56'S, 45° 18' E, 1300-
1480 m. 1 lv, 1 dd (paratypes).

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

361

Fig. 165. — Distribution of Gadila doumenci.

Distribution. — New Caledonia and NW Madagascar, living from 1100 to 2450 m.

Description. — Shell to 1 1 mm long, solid, shiny,
moderately curved, white. Maximum diameter at anterior
fifth of the shell. Apex almost as wide as mouth, simple.
Preapical callus wide, lumen circular. Mouth slightly laterally

compressed with the ventral side flattened, straight, simple.

Measurements: holotype L 10.5, W 2.2-2.15, m 1 .6-1.5, w
1.1-1.06; paratypes L 9.4. W 2.05. m 1.4, w 1; L 9, W 2, m
1.2, w 0.9.

Etymology. — Named for Prof. Dominique Doumenc, who made possible my work as
visiting curator in mnhn.

Gadila desaintlaurentae sp. nov.

Figs 166, 169 f

Type material. — Holotype mnhn. Paratypes: 8 mnhn, 1 ams C201734, 1 usnm.

Type locality. — Philippines, musorstom 2, stn DR 33, 13°32'N, 121°08'E, 130-137 m.

Material examined. — New Caledonia. “Vauban” 1978-79: stn 40, 22°30' S, 166°24' E,
250-350 m, 1 lv, 12 dd.

Indonesia, corindon: stn B 268, 01°57' S, 119°16' E, 200 m, 2 dd.

Philippines, musorstom 2: stn DR 33, 13°32' N, 121°08' E, 130-137 m, 3 lv (holotype and paratypes),

15 dd (paratypes: 6 mnhn, 1 ams, 1 usnm).

musorstom 3: stn CP 112, 14°00'N, 120°18' E, 187-199 m, 1 dd.

Distribution. — The Philippines, Indonesia and New Caledonia, living from 137 to

250 m.

Description. — Shell to 6 mm long, shiny, white, with
maximum diameter at anterior third. Ventral side regularly
curved, dorsal side describing a convexity followed by a
pronounced concavity that reaches the apex. Apex simple,
circular. Preapical callus faint, lumen circular. Mouth simple,
oblique, circular.

Measurements: holotype L 6, W 1.1, m 0.7, w 0.4;
paratypes L 5.8, W 1,2, m 0.7, w 0.4; L 5.5, W 1.1, m 0.7.
w 0.4; L 6.3. W 1.2, m 0.8. w 0.4; L 5.6, W 1. m 0.7,
w 0.4.

362

VICTOR SCARABINO

Etymology. — Named for Dr Michele de Saint-Laurent, formerly curator of Crustacea
(mnhn), who participated in the first three musorstom expeditions to the Philippines.

Gadila minutalis sp. nov.

Figs 167, 169 g

Type material. — Holotype and 8 paratypes lv, mnhn.

Type locality. — West Indian Ocean, md32 Reunion, stn DS 151, 20°5T S, 56°03' E,
3240-3300 m.

Material examined. — Only known from the type material.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

363

Distribution. Only known from Reunion Island, alive in 3240-3300 m.

Description. — Shell to 3 mm long, fragile, shiny, white, Measurements: holotype L 2.9. W 0.5, m 0.3, w 0.2.
moderately curved. Maximum diameter at anterior quarter of
the shell. Apex simple, circular. Preapical callus thin, lumen
circular. Mouth simple, straight, circular.

Etymology. — Very small (Latin).

Gadila monodonta sp. nov.

Figs 168, 169 i

Type material. — Holotype and 2 paratypes mnhn.

Type locality. — West Indian Ocean, benthedi, stn DS 10, 1 1°29' S, 47°18' E, 440 m.

Material examined. — New Caledonia, calsub: dive 20, 22°53' S, 167°23' E, 580 m, 1 dd.
West Indian Ocean, benthedi: stn DS 10, 1 1°29' S, 47° 18' E, 440 m, 1 lv (holotype). — Stn DS 120,
1 1°30' S, 47°25' E, 335-390 m, 1 lv (paratype). — Stn DS 122, 11°32' S. 47°23' E, 615-625 m, 1 dd
(paratype).

Distribution. — NW Madagascar and New Caledonia, alive in 390-440 m, shells down to

625 m.

Description. Shell to 15 mm long, shiny, white,
maximum diameter near the center of the shell, giving a
generally fusiform shape. Ventral side regularly curved; curve
of dorsal side interrupted at the maximum diameter. Apex
oval, with ventral lobe showing a central denticle at the edge.

Preapical callus thin, lumen suboval. Mouth simple, oblique,
slightly dorsoventrally compressed.

Measurements: holotype L 14.2, W 2.6-2.45, m 1.45-1.25,
w 0.75-0.7.

Remarks. The fusiform shape and distinctive apical feature easily characterize this new

species.

Etymology. From the Latin, meaning a single tooth.

364

VICTOR SCARABINO

Fig. 169. — a. Gadila virginalis, shell (16 mm), lateral and dorsal views, apical and oral sections, corindon: stn B 236. —
b. Gadila zonata, shell (14 mm), lateral and dorsal views, apical and oral sections, musorstom 2: stn CP 18. — c, Gadila
boissevainae, shell (7.5 mm), lateral and dorsal views, apical section, "Meiring Naude”, stn SM 59. d, Gadila elenae
sp. nov., holotype, shell (11.5 mm), lateral and dorsal views, apical and oral sections. e, Gadila doumenci sp. nov.,
holotype, shell (10.5 mm), lateral and dorsal views, apical and oral sections. - f. Gadila desaintlaurentae sp. nov.,
holotype, shell (6 mm), lateral and dorsal views, apical and oral sections. - g, Gadila minutalis sp. nov.. holotype, shell
(2.9 mm), dorsal and lateral views, apical and oral sections. — h, Gadila type radula (Gadila sp. nov., Caribbean Sea).
— i, Gadila monodonia sp. nov., holotype, shell (14.2 mm), lateral and dorsal views, apex, apical and oral sections.

Other Indo-Pacific species of Gadila cited in the literature

Gadila abruptoinflata Boissevain, 1906: 75, pi. 66, fig. 65, textfig. 39. Indonesia, Madura, “Siboga",
stn 5, 07°46 ‘S, 114°31' E, 330 m. Holotype zma.

Gadila anguidens (Melvill & Standen, 1898): 32, pi. 1, fig. 6. Off Madras, India. Holotype and
paratype, Manchester Museum (fide Trhw, 1987).

Gadila clavata (Gould, 1859): 166. Hong Kong, China, 11-36 m. Holotype usnm 24245.

Gadila honotuluensis (Watson, 1879): 89. " Challenger ", Reefs of Honolulu, Hawaii, 40 fms [73 m],
Holotype bmnh 1887.2.9.70.

Gadila opportuna (Kuroda & Habe, 1961): 105, pi. 47, fig. 2. Sagami Bay, Japan, 200 m. nsmt.

Gadila pseudolivae (Boissevain, 1906): 73, pi. 6, fig. 67, textfig. 36. “Siboga", stn 211, 05°41' S,
120°46'E, Banda Sea, 1158 m. Syntype zma.

Gadila subcolubridens Ludbrook, 1954: 115, fig. 21. Gulf of Aden, “John Murray ”, stn 185, 13°48' N,
49°16' E, 2000 m. Presumably in bmnh (not located).

Gadila subtilis (Plate, 1908a): 360, pi. 30, fig. 48. Off Dar-es-Salaam, Tanzania, “Valdivia", stn 244,
05°56' S, 39°0T E, 50 m. Lectotype (Kilias, 1995) zmb 61108a.

Gadila novilunata Kira, 1955: 80, pi. 40, fig. 2. Japan, Shikoku.

Source . MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

365

Family incertae sedis

The following two genera possess confusing characters that do not allow placement in the
present classification. Megaentalina is the only representative of the Order having a triangular section
and a Siphonodentalium type radula. Compressidens has dorsoventrally compressed and curved shell
as a diagnostic character in all species. Pending further data, I have decided to treat both genera
separate from the rest.

Genus Megaentalina Habe. 1963

Type species (OD): M. teramachii Kuroda & Habe in Habe, 1963 [= M. mediocarinata Boissevain, 1906].

Diagnosis. — Shell medium to large, well arched, solid, polished, white to cream. Sculpture
of 3 primary ribs, with one angle on dorsal side and a base at the ventral; secondary riblets present;
rib section flat-topped to rounded; smooth; intercostal spaces, almost straight. Apex simple or with
two flat notches, one dorsal and the other ventral. Section subtriangular at apex, less noticeable at
mouth.

Radula rachidian polygonal with wide base, anterior border irregular, with 5-6 prominent nodules in
the interior face that overpass the border; laterals similar to Siphonodentalium, strong, curved, with
short but strong cusps; marginals straight with internal side wider than the external side.

Distribution. — Recent, East Pacific and Indian Oceans, absent in the Atlantic Ocean.
Shelf-bathyal.

Remarks. — Steiner (1991) places Megaentalina in the Order Dentaliida, on my advice
(Scarabino, 1986, pers. com.). This suggestion was incorrect, since it was based on the radula of two
animals supposedly conspecific with two shells of Megaentalina mediocarinata found in the same vial.
Recent study of live animals has corrected this error the earlier specimens belong to a dentaliid
species. My erroneous placement can now be corrected and we await further data for proper
classification of this genus.

Fig. 170. — Distribution of Megaentalina cornucopiae.

366

VICTOR SCARABINO

Megaentalina cornucopiae (Boissevain, 1906)
Figs 170, 174 b

Entalina cornucopiae Boissevain, 1906: 63, pi. 6, fig. 89.

Other references:

Megaentalina cornucopiae - Habe, 1963: 212, pi. 38, figs 31-32; 1964a: 41, pi. 2, figs 31-32; 1977: 340. Hark & Kosuge,
1964: 9. Chistikov, 1982c: 1499. pi. 2. figs 4-6. Higo & Goto. 1993: 689.

Type material. — Holotype zma 3.06.086.

Type locality. Indonesia, Sumba Sea, " Siboga ”, stn 52, 09°03' S, 119°57' E, 959 m.

Material examined. — The type material.

Philippines. estase 2: stn DR 4, 05°02' N, 125°15' E, 3250 m, 1 dd.

Distribution. — Japan, the Philippines, Indonesia, shells from 959 to 3250 m.

Fig. 171. - Distribution of Megaentalina mediocarinala.

Megaentalina mediocarinala (Boissevain, 1906)
Figs 171, 172 e-f, 174 a, c

Entalina mediocarinala Boissevain, 1906: 63. pi. 6, figs 70-72, 87-88.

Synonym:

Megaentalina teramachii Kuroda & Habe in Habe. 1963: 273, pi. 38, figs 12-13.

Other references:

Megaentalina mediocarinala - Habe. 1964a: 42, pi. 2. figs 12-13; 1977: 340. — Habe & Kosuge, 1964: 9. Chistikov, 1982c:
1498. - Higo & Goto, 1993: 689.

Dentalium (Compressidens) comprimatum - Ludbrook, 1954: 106.

Type material. — E. mediocarinala: lectotype (here designated) zma 3.06.084, paralectotypes
zma 3.06.085. — M. teramachii : holotype, nsmt.

Type locality. E. mediocarinala ; Bali Sea, Indonesia, “ Siboga ”, stn 5, 07°46' S, 114°30' E,
330 m. — M. teramachii: Tosa Bay, Shikoku, Japan, 200 m.

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

367

Material examined. — The type material of E. mediocarinata.

Indonesia, corindon: stn B 213, 00°3T N, 117°50' E, 488 m, 1 dd.

“Snellius” II: stn 4.130, 08° 18' S, 1 18° 18' S, 700-730 m, 2 lv, 3 dd. — Stn 4.131, 08° 18' S, 1 18°18' E,
680-800 m, 1 dd (rmnh).

Philippines, musorstom 2: stn CP 24, 13°37' N, 120°42' E, 640-647 m, 1 dd. — Stn CP 25, 13°39' N,
120°43'E, 520-550 m, 4 lv, 11 dd. — Stn CP 26, 13°49' N, 120°50'E, 299-330 m, 3 dd. —
Stn CP 40, 13°08'N, 122°40’E, 280-440 m, 1 dd. — Stn CP 46, 13°26' N, 122°17'E, 445-520 m,
1 lv. — Stn CP 78, 1 3°49' N, 120°28' E, 441-550 m, 3 lv, 3 dd.

musorstom 3: stn CP 123, 12°10' N, 121°45' E, 700-702 m, 1 dd. — Stn CP 128, 11°50' N, 12P42' E,
815-821 m, 1 dd.

West Indian Ocean. Zanzibar area, "John Murray"-, st 105B, 05°24' S, 39°14' E, 238-293 m, 1 dd
[Dentalium (Compressidens) comprimatum ] (bmnh).

Northern Indian Ocean, safari 2: stn CP 06, 08° 1 1' N, 79°03' E, 1035 m, 1 lv.

Distribution. — Japan, the Philippines, now extended to Indonesia and off Sri Lanka and
Zanzibar, living depth range 441-1035 m, shells from 300 m.

Fig. 172. - Radulae. a. Spadenialina lubiformis, general view, note at upper left an internal face view of a lateral teeth,
to be compared with Entalinopsis and Pertusiconcha radulae (Figs 115 d and g respectively). — b, same specimen,
rachidians. c, Striocadulus sagei, margin of the head of a lateral tooth. - d, same specimen, general view,
e, Megaentalina mediocarinata, internal face of lateral tooth. — f, Megaentalina mediocarinata, external view of lateral
and rachidians. — g, Solenoxiphus striatulus, internal view of lateral tooth. Scale lines: 100 pm (a, d-0, 10 pm (b. c),
20 pm (0-

368

VICTOR SCARABINO

Genus Compressidens Pilsbry & Sharp, 1 897

Type species (OD): Dentalium pressum Pilsbry & Sharp, 1897. Recent, North of Culebra Island, West Indies, 390 fms
[714 m].

Diagnosis. — Shell small to medium, well curved, solid, translucent, opaque or polished.
Section oval, strongly compressed dorsoventrally. Sculpture variable, with longitudinal riblets,
undulations or fine, close encircling wrinkles; growth lines conspicuous. Apex simple or truncate,
preapical callus usually wide, lumen circular.

Radula rachidian high, with cusped anterior marging; lateral high, with very sharp-pointed cusps;
marginal short, sinusoidal (Fig. 174 e, C. ophiodon Dali, 1881).

Distribution. — Miocene- Recent, worldwide, shelf-bathyal.

Compressidens infimus sp. nov.

Figs 173, 174 d

Type material. — Holotype and 8 paratypes lv, mnhn.

Type locality. — West Indian Ocean, md 32 Reunion, stn DS 109, 20°52' S, 55°06' E,
1050-1240 m.

Material examined. — Only known from the type material.

Distribution. — Reunion Island, alive in 1050-1240 m.

Description. — Shell to 5 mm long, fragile, well curved.
Sculture of close, fine, encircling wrinkles throughout. Apex
subcircular, dorsoventally depressed. Preapical callus thin,

lumen circular. Mouth thin, slightly dorsoventrally depres¬
sed, especially on ventral side.

Measurements: holotype L 4, W 0.5-0.45, w 0.1, arc 0.22.

Source : MNHN , Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

369

Remarks. — Compared with Compressidens platyceras, a shallow water species from Eastern
Australia, C. infimus is smaller and less compressed. C. comprimatum (Plate, 1908a) from Zanzibar
Channel, has longitudinal striation.

Etymology. — From the Latin meaning very little.

Fig. 174. — a, Megaemalina mediocarinata, shell (34 mm), lateral and dorsal views, apex and apical sections (usnm). -
b, Megaemalina cornucopiae, shell (22 mm), lateral and dorsal views, apical and oral sections, section of sculpture,
estase 2: stn DR 4. — c, Megaemalina type radula ( M . mediocarinata ); see also Fig. 172 e. — d, Compressidens infimus
sp. nov., holotype, shell (4 mm), lateral and dorsal views, apical and oral sections, detail of sculpture. —
e, Compressidens type radula (C. ophiodon Dali, 1881, Northern Brazil).

370

VICTOR SCARABINO

Other Indo-Pacific species of Compressidens cited in the literature

Compressidens comprimatum (Plate, 1908a): 349, pi. 30, figs 26-34. Zanzibar Channel, Valdivia ,
stn 245. 05°28' S, 39°19' E, 463 m. Lectotype (Kilias, 1995) zmb 61101a.

Compressidens kikuchii (Kuroda & Habe, 1952): 9, pi. 1, figs 3-4. Toyama Bay. Honshu. Japan.

NSMT.

Compressidens platyceras (Sharp & Pilsbry in Pilsbry & Sharp, 1897): 126. pi. 22, figs 58-60. East
Australia, Salamander Bay, Port Stephens, New South Wales, 4-8 fms [7-15 m], Syntypes ansp
35565 and ams 55085, 11721.

GENERAL DISCUSSION

The total number of scaphopod species present in discrete geographical subregions of the
Indo-Pacific is presented in Table 2. Estimated figures are based on bibliographic sources cited in this
paper, as well as actual examination of material. It is likely that data in this Table will become
obsolete as new regional revisions, such as the scaphopod part of the ongoing Fauna of Australia by
Healy & Lamprell, are published. With 73 species currently recorded from there. New Caledonia
is now the area with the highest documented scaphopod diversity in the Indo-Pacific. Undoubtedly,
differences among different"regions reflect intensity in sampling effort, as well as genuine differences
in global faunal diversity. Thanks to the series of musorstom expeditions in the New Caledonia area,
the" collecting effort in that part of the SW Pacific has been considerably higher than in any other part
of the Indo-Pacific of similar size. The total of 73 New Caledonian species represents 53% of the total
number of scaphopod species recorded in the present paper, and 31% of the total Indo-West Pacific
fauna (237 species). This proportion is particularly remarkable in view of the small size (less than 2
million km2) of the New Caledonia Exclusive Economic Zone compared to the total extent of the
Indo-Pacific region (over 150 million km2).

Table 2. Estimated number of scaphopod species in selected regions

Region

Total

Dentaliida

Gadilida

Japan

50

30

20

Indonesia

65

41

24

Philippines

50

39

11

Indian Ocean

62

33

29

China Seas

26

22

4

N and E Australia

22

17

5

South Africa

15

10

5

New Zealand

12

5

7

Hawaii

5

3

2

New Caledonia

73

43

30

Source : MNHN, Paris

SCAPHOPODA OF THE TROPICAL INDO-PACIFIC

371

BATHYMETRIC DISTRIBUTION

Sampling effort in New Caledonia has been rather even between 0 and 2000 m. The fauna
deeper than 2000 m is less adequately surveyed, and there are no samples from deeper than 3750 m.
Figure 175 illustrates the number of species found at discrete depth intervals in New Caledonia:
0-100, 100-300, 300-500, 500-800, 800-1200, 1200-2000, 2000-3000 and 3000-4000 m. Because a
species may have different bathymetric ranges at different latitudes and/or in different hydrological
regimes, only depth data for New Caledonia have been considered. Of the 73 species recorded, 48
have been collected alive. For the remaining 25 species, bathymetric ranges are based on empty shells
since they apparently depict patterns that are both coherent and consistent with what is known of the
depth range of the same species elsewhere in the Indo-Pacific.

FIG. 175. - Bathymetric distribution of the scaphopod fauna of New Caledonia. Open square: Dentaliida; black square:
Gadilida.

The diversity of both Dentaliida and Gadilida increases from 0 to 500 m. is maximal in the
500-800 m depth interval, and decreases abruptly in the 800-1200 m interval. Deeper than 1200 m,
the number of Dentaliida continues to decrease, whereas Gadilida show a second, minor peak in
1200-2000 m. In Dentaliida, Fissidentalium is most diverse in the deep sea (mostly bathyal and
abyssal), other genera being equally represented at all depths. The second peak of the Gadilida is
mostly formed by representatives of Entalinomorpha and Siphonodentalium. Similar patterns have
been noted in the Atlantic Ocean (Scarabino, 1979, 1986a-b, and unpublished observations) and
point to distinct radiation patterns of Dentaliida and Gadilida in the deep-sea.

ACKNOWLEGDEMENTS

This paper has been done in the course of several periods where I was working as visiting
curator in the Laboratoire de Biologie des Invertebres Marins et Malacologie, mnhn, and I am
grateful to the director, staff and colleagues for inviting me and entrusting me with this very valuable
material: D. Doumenc, B. Metivier, A. Crosnier, S. Gofas, R. von Cosel, V. Heros and J.P. Rocroi.

372

VICTOR SCARABINO

Scaphopod material was collected during expeditions under the direction of J. Forest (mnhn), C. Levi
(mnhn), B. Richer de Forges (orstom, Noumea), A. Guille (then of mniin, now at Laboratoire Arago,
Banyuls), B. Thomassin (cnrs, Marseille) and L. Labeyrie (cnrs. Gif), centob (ifremer, Brest)
contributed to the sorting of residues. For access to material under their responsability, I thank R.
Moolenbeek (zma), A. Bogan, R. Robertson, G. Davis (ansp), the late R. Houbrick (usnm), B.
Marshall (nmnz), K. Way (bmnh), P. Colman (ams), E. Louw and M. van der Merwe (sam), J.
Knudsen, T. Schiotte (zmc), E. Gittenberger and J. Goud (rmnii). For technical assistance, I thank
D. Guillaumin (Centre Inter-Universitaire de Microscopie Electronique, Universite Paris 6) and the
staff of mnhn, P. Maestrati and M. Moreau. The manuscript owes especially to P. Bouchet (mnhn)
for his criticisms and highly valuable suggestions and to W. K. Emerson and Walter Sage (amnii) for
their suggestions and linguistic assistance.

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Source : MNHN, Paris

JLTATS DES CAMPAGNES MUSORSTOM. VOLUME 14 RESULTATS DES CAMPAGNES MUSORSTOM. VOLUME 14 RESUL"

Calliostomatidae (Gastropoda: Trochoidea)
from New Caledonia,

the Loyalty Islands, and the northern Lord Howe Rise

Bruce A. MARSHALL

Museum of New Zealand Te Papa Tongarewa
P.O. Box 467, Wellington
New Zealand

ABSTRACT

Thirty species (27 new) of Calliostomatidae are recorded from the study region, all but two of which are new records.
An additional new species is based on material from northern New Zealand. They are referred to Fautor Iredale. 1924,
Benthastelena Iredale. 1936, Ampullotrochus Monterosato, 1890 (as subgenera of Calliostoma Swainson, 1840), Baihyfautor
gen. nov., Daclylastele gen. nov., Laelifaulor Iredale, 1929, Selastele gen. nov., Fautrix gen. nov., and Thysanodonta Marshall,
1988. A new tribe, Fautricini, is introduced for species with a radula that is evidently the most primitive (plestomorphic) in
the family, and Fautricini either represents the common basal stock or an early offshoot from it. Calliostomatidae is treated
as a family within Trochoidea rather than a subfamily of Trochidae as has been traditional. Three calliostomatid genus group
taxa are newly synonymised: Tristichotrochus Ikebe. 1942 (= Benthastelena Iredale, 1936), Salsipotens Iredale, 1924 (= Astele
Swainson, 1840), Spicator Cotton & Godfrey, 1935 (= Laelifaulor Iredale. 1929). Criteria used for taxonomic discrimination,
evolutionary history, and some biogeographical observations are discussed. All calliostomatid genus group taxa and taxa
removed (some newly) from the family are listed in appendices. A lectotype is designated for Zizyphinus scobinatus A. Adams,
1863.

RESUME

Les Calliostomatidae (Gastropoda: Trochoidea) de Nouvelle-Caledonie, des ties Loyaute ct du Nord de la Ride de Lord

Howe.

Trente especes (dont 27 especes nouvelles) de Calliostomatidae sont recensees dans la region consideree. et toutes sauf
deux representent des signalisations geographiques nouvelles. Une autre espece nouvelle est decrite a partir de materiel du Nord
de la Nouvelle-Zelande. Les especes sont classees dans les genres Fautor Iredale, 1924. Benthastelena Iredale. 1936,
Ampullotrochus Monterosato, 1890 (traites comme sous-genres de Calliostoma Swainson. 1840), Baihyfautor gen. nov..

Marshall, B.A.. 1995. Calliostomatidae (Gastropoda: Trochoidea) from New Caledonia. Ihe Loyally Islands, and ihe northern Lord Howe Rise. In: P.
Bouchft (ed.), Resuitats des Campagnes Mlsoustom, Volume 14. Mem. Mus. natn. Hist. nat.. 167: 381-458. Paris ISBN 2-85653-217-9.

Published 29"1 December 1995.

382

BRUCE MARSHALL

Dactylastele gen. nov., Laetifautor Iredale, 1929, Selaslele gen. nov., Fautrix gen. nov., el Thysanodonta Marshall, 1988. La
nouvelle tribu Fautricini, creee pour les taxons dont la radula est apparemment la plus primitive (plesiomorphe), represente
le stock ancestral de la famille ou une branche ancienne de celui-ci. Calliostomatidae est traite comrae une famille de
Trochoidea, et non simplement corame une sous-famille de Trochidae comrae il est habituel de le faire. Trois noms du
groupe-genre sont mis en synonymie: Tristichotrochus Ikebe, 1942 (= Benthaslelena Iredale, 1936), Salsipotens Iredale, 1924
(= As tele Swainson, 1840) et Spicator Cotton & Godfrey, 1935 (= Laetifautor Iredale, 1929). Les caracteres utilises pour la
systematique et I'histoire evolutive de la famille sont discutes, de meme que di verses observations de biogeographie. Une liste
complete des taxons du groupe-genre appartenant a la famille Calliostomatidae, ainsi qu'une liste de ceux qui en ont ete exclus
(certains pour la premiere fois dans ce travail), figurent en appendice. Un lectotype de Zizyphinus scobinatus A. Adams, 1863
est choisi.

INTRODUCTION

During the last few decades French oceanographic expeditions have sampled benthic biota at
over 600 dredge and trawl stations off New Caledonia, the Loyalty Islands and on the Lord Howe
Seamount Chain (Richer de Forges, 1990). The resulting material (mnhn) is by far the richest and
most comprehensive collection of benthic tropical biota ever assembled from the Indo-Pacific,
particularly from depths greater than 100 metres on rugged substrata. The molluscan material
includes many hundreds of undescribed species and great numbers of new distributional records.
These collections dramatically reveal how little is actually known of the biotas occurring on
seamounts and off tropical islands in general, especially in the western Pacific, and they probably
afford a mere hint of the species richness in this huge region. Through the generosity of Dr Philippe
Bouchet (mnhn), I am privileged to be able to describe the rich calliostomatid material from these
collections.

Calliostomatids are trochiform gastropods that range from about 8-100 mm in shell height.
They are exclusively marine and occur in all oceans from the intertidal zone to about 3,000 metres
depth, typically on rocky substrata. Related to trochids they are characterised by a distinctive
protoconch sculpture and radular morphology. The group is richly speciated with more than 250
valid Recent species, including those named herein. About 300 fossil taxa have been named. While
many of the fossils are doubtfully or categorically not calliostomatids or are synonyms, I have
personally examined about 70 undescribed Tertiary species (mainly from Australia and New Zealand)
as well as a similar number of undescribed Recent species. Whereas some species are commonly found
associated with algae, gut contents and field observations reveal that all are carnivores when adult,
feeding mostly upon Cnidaria and, to a lesser extent, sponges or carion.

ABBREVIATIONS AND TEXT CONVENTIONS

Repositories

AMS

BMNH

LACM

MNHN

NMNZ

NMP

NSMT

NZGS

NZOI

USNM

: Australian Museum, Sydney
: The Natural History Museum, London
: County Museum of Natural History, Los Angeles
: Museum national d’Histoire naturelle, Paris
: Museum of New Zealand, Wellington
: Natal Museum, Pietermaritzburg
: National Science Museum, Tokyo

: Institute of Geological and Nuclear Sciences, Lower Hutt
: National Institute of Water and Atmospheric Sciences, Wellington
: National Museum of Natural History, Washington DC

CALLIOSTOMATIDAF. FROM NEW CALEDONIA

383

Other abbreviations

D

: Diameter

H

: Height

P

: Primary spiral (Fig. 1)

S

: Secondary spiral (Fig. 1)

TW

: Teleoconch whorls (number)

dd

: no live-taken specimens present

in sample

lv

: live-taken specimens present in

sample

spm

: specimen, condition unknown

OD

: Original designation

SD

: Subsequent designation.

Fig. I. — Key to standard calliostomatid spiral cord numbers on two whorls of Calliostoma (Maurea) osbornei Powell, 1926
(off Cape Maria van Diemen. New Zealand. 38-43 m. nmnz M74665, * 7).P = primary spiral cord, S = secondary
spiral cord.

Height precedes diameter in all given dimensions. All shell measurements were taken on the
longitudinal axis or at right angles to it. Unless specified, all material is in Museum National
d'Histoire Naturelle, Paris.

SYSTEMATIC ACCOUNT

Subclass prosobranchia Milne Edwards, 1848
Order vetigastropoda Salvini-Plawen, 1980
Superfamily trochoidea Rafinesque, 1815
Family calliostomatidae Thiele, 1924

Diagnosis. — Shell trochiform or turbiniform, 8-100 mm high, umbilicate or anomphalous,
internally nacreous. Protoconch sculptured with network of threads that enclose roughly hexagonal
spaces. First teleoconch whorl convex, subsequent whorls flat to convex or angulate; sculpture rarely
absent, usually consisting of nodular spiral cords that multiply by intercalation (rarely by fission) and

384

BRUCE MARSHALL

axial riblets, the axials generally restricted to earliest whorls, though in some species persistent
throughout. Operculum chitinous, thin, multispiral. Radula (adult) with the formula co + 1-17 +

1 + 17-1 + co, tooth morphology very diverse (see subfamily diagnoses); lateral teeth added
incrementally by intercalation in the central field in Calliostomatinae, and by morphological
transformation of marginal teeth in Fautricini. Jaw plates large, buccal musculature very strong.
Cephalic tentacles elongate, tapered, papillate, well developed eyes on swellings at their outer bases
in all species examined. Snout tip papillate, ventral lip split and rolled to form an incomplete tube
(pseudoproboscis), dorsal lip with a cuticularised lining that extends into buccal cavity. Epipodium
well developed with prominent neck lobes. Cephalic lappets large, small or absent. Up to five
prominent epipodial tentacles on each side, side of foot papillate. Sexes separate. (See Fretter &
Graham, 1962, 1977 for more detailed anatomical data).

Remarks. — Interpretation of the relationships between calliostomatid species and thus
resolution of actual or potential phylogenetic radiations within the family has proved to be extremely
and unusually difficult. This is due primarily to the fact that radular morphology and external
anatomy are in general highly conservative within the major groups. Many groups of species that
seem to represent independent phylogenetic radiations on the basis of distinctive combinations of
shell characters, cannot be neatly defined as genera or subgenera because, considered together with
other species, there is a bewildering mosaic of intermediate character states. Shell shape and sculpture,
though diverse, are generally variations on a very limited theme, and it is clear that many of the most
pronounced differences are merely the result of accelerated or retarded development, often in
apparently closely related species. In most calliostomatids (i.e. most Calliostomatinae), the first few
teleoconch whorls are convex and sculptured with four strong primary spiral cords (PI -P4, see Fig. 1 ), P4
being peripheral and more or less covered by succeeding whorls. On the first whorl P1-P4 or P2-P4
typically commence simultaneously, and PI is usually either weaker than P2 and P3 when
commencing simultaneously, or it appears later. With subsequent growth the whorls typically become
flatter-sided, P1-P3 remain similar, or PI becomes larger than P2 and P3, and spiral cords are
multiplied by intercalation of secondaries and sometimes tertiaries. In some species P2 and P3 become
relatively stronger after the first whorl and may remain relatively strong throughout, any secondary
spirals that intercalate between them typically remaining substantially weaker than the primaries
(compare Figs 49 and 55). While there are exceptions, particularly large spiral cords tend to occur
in species with convex whorls and in which PI either is relatively weak throughout, appears very late,
or entirely fails to develop. Adults of these species resemble their own juvenile stages to some extent,
and it seems that they are paedomorphic, with retarded somatic development. Polarity for these
character states, however, cannot be resolved without knowledge of the fossil record. Calliostomatids
that are considered to have “juvenilized" adult shell morphology include species of Laetifautor ,
Coralastele, Otukaia, Omphalotukaia, and some species of Benthastelena. Since phylogenetic groups
( e.g . genera) within the family naturally comprise species in which character states are accelerated or
retarded to varying degrees, conspicuous differences in shell morphology such as whorl convexity
alone may not necessarily be suitable for discrimination at generic or subgeneric level.

Major interspecific differences in radular morphology include the shape of the central tooth,
the number and shape of the lateral teeth, and features of the marginal teeth, especially the innermost
pair. As recently demonstrated by Waren (1990), calliostomatid radulae undergo pronounced
changes with increasing shell size/age, which include a progressive change in shape and increase in
number of lateral teeth (by intercalation) and a change in shape of the innermost pair of marginal
teeth. As with the shell, differences in the radula, particularly the number of lateral teeth, cannot be
utilised for genus group discrimination in isolation, since development of this organ structure is
obviously accelerated or retarded to varying degrees in related species.

When viewed on a worldwide basis, a number of more or less nebulous natural (phylogenetic)
groups are recognisable, though most of them have proved to be unusually difficult to quantify or

Source : MNHN, Paris

CALLIOSTOMATIDAE FROM NEW CALEDONIA

385

to define objectively with traditional methods, particularly with due consideration to heterochrony.
Molecular cladistic techniques would seem to be a promising source of data for resolution of the
phylogenetic structure of this family, and indeed for the whole of the Trochoidea.

Included genera and subgenera are listed in Appendix 1. For genus group taxa removed from
the family see Appendix 2.

Protoconch. The calliostomatid protoconch is highly distinctive in having a sculptural
network of crisp threads that enclose more or less hexagonal spaces, and in having a thickened
apertural rim. Thread thickness, enclosed space size, overall protoconch size, and degree of exsertion
(tilting) are interspecifically variable and intraspecifically highly stable. The shape of the tip of the
apical fold may be rounded or tightly pinched, an obese protoconch evidently reflecting a large yolk
supply. Bandel (1982) has shown that the surface sculpture is secreted after deformation (folding) of
the protoconch, presumably via solute transmission through the semipermeable outer shell layer at
an early stage of mineralisation. The fully mineralised shell is almost perfectly bilaterally symmetrical,
and so far no species have been noted in which the tip of the apical fold bulges abapically as in some
trochids (Hadfield & Stratiimann, 1990). Incidentally, these latter authors misinterpreted trochids
with abapically bulging protoconch tips as having "heterostrophic” shells. This is incorrect, because
the archaeogastropod protoconch is asymmetrically (in this case) or symmetrically folded about an
axis, and there is no accretionary shell growth.

Teleoconch /. All known calliostomatines exhibit a pronounced growth scar on the first
quarter teleoconch whorl, the position of which is interspecifically variable though intraspecifically
rather stable (Figs 2-3). Sculpture before the scar comprises 4-6 fine, crisp, similar spiral threads, and
usually 1, 2 or more varices. Sculpture immediately after the scar is usually discordant (Fig. 2),
though a few species exhibit a rather fluid transition to succeeding teleoconch sculpture, usually
accompanied by a sudden increase in the strength of the spiral sculpture (Fig. 3). From descriptions
of calliostomatids reared in aquaria (Lebour, 1936; Holyoak. 1988; Ramon, 1990; K. Bandel pers.
comm.), teleoconch accretion commences immediately or soon after formation of the terminal varix
of the protoconch (here interpreted as the point of settlement). During the period of initial teleoconch
formation the young evidently subsists on residual yolk, the resorbed velum, and perhaps dissolved
organic matter, probably increasingly supplemented by detritus ingested from the substratum. It
seems clear that the postlarval scar represents a growth pause or a crisis period and I suggest that
it may denote the transition to exclusive detritivory (later transitional to carnivory). It may actually
mark the transition to carnivory, at least in some species, though the radula may be insufficient to
deal with Cnidaria or sponges at such an early stage of development. This interpretation is at variance
with that of Hickman (1992; fig. 5G) who identified the terminal protoconch varix and the varix
following it as denoting the times of hatching and settlement respectively. Note that the postlarval
scar in the species illustrated by Hickman is situated immediately on the adapertural side of the varix
following that denoted as time of “settlement”.

Teleoconch II. — With the exception of a few more or less smooth species, the majority of
calliostomatids are sculptured with prominent (usually nodular) spiral cords and axial costae, the
latter generally confined to the early spire whorls. There are usually four primary spiral cords (P1-P4.
Fig. 1) on the early spire whorls, to which others are added by intercalation (rarely fission) of
secondary spirals (S1-S3) and sometimes one or more tertiary spirals that may enlarge to resemble
the primaries or secondaries. The number, spacing, relative size, rate of enlargement, position, point
of origin, and cross-sectional shape of spiral cords are primary criteria for discrimination of species
group taxa. Sculpture tends to be intraspecifically highly stable on the early spire whorls, and
increasingly variable on later whorls, due primarily to intercalation of varying numbers of tertiary
spirals at varying positions with varying rates of enlargement. Accordingly, for descriptions and
discrimination of taxa, careful attention must be paid to the origin and development of individual

386

BRUCE MARSHALL

elements of the spiral sculpture by tracing them from the beginning of the teleoconch mere
counting of the number of spiral cords on particular whorls is virtually useless. Unequivocal
objectivity can only be achieved through rigorous application of the excellent spiral designation
system initiated by Ikebe (1942), which is reproduced here for convenience (Fig. 1).

Sculptural features combine to produce an infinite variety of distinctive, intraspecifically stable
gross shell facies, particularly on the early teleoconch. Natural (phylogenetic) groups segregated on
the basis of highly distinctive radular morphology (e.g. Aside, Laetifautor , Fautrix gen. nov. and
Thysanodonta ) tend also to have distinctive early teleoconch facies. Nevertheless, even highly
distinctive shell facies are typically impossible to quantify and very difficult to describe objectively
(especially with verbal economy), so illustrations (SEM) are an integral component of my genus
group diagnoses.

Radula. — As recently shown by Waren (1990), the calliostomatine radula is distinctive
among trochoideans in that the innermost pair of marginals become greatly enlarged at an extremely
early stage of ontogenesis, after which the central and lateral teeth arise by intercalation in the central
field. In other trochoideans (and apparently in the new calliostomatid tribe Fautricini — see below)
the lateral teeth arise through progressive in-column morphological transformation of marginal teeth,
and any additional laterals are added pair by pair through transformation of successive columns of
marginals. For comparative purposes, therefore, it is clearly essential to compare radulae from
individuals at equivalent stages of development, ideally maturity since related species will naturally
differ in rates of somatic development.

Calliostomatine radular morphology is on the whole exceedingly conservative, and species
with highly divergent shell morphologies commonly have fundamentally similar radulae. Nevertheless
there is interspecific variation and intraspecific stability in the relative size, shape, and number of
teeth and cusps in adult radulae. The overall shape of the central and lateral teeth and the sizes of
their basal plates cannot be determined from standard SEM views of artificially protracted radulae
because these teeth are longer than broad, crowded, strongly curved and typically flexible. Tooth
outline can be accurately determined only by flattening excised individual teeth, while the relative
sizes of the basal plates are best determined by light microscopy of stained preparations using
substage lighting. While experimental use of these techniques on various species revealed many
taxonomically significant features, they are omitted from the present study not only because of time
constraints, but also because shell and gross radular characters are considered to be adequate for
discrimination of species and most supraspecific groups. Most natural (phylogenetic) groups that can
be discriminated on shell morphology tend also to have characteristic numbers of lateral teeth, for
example species of Laetifautor have only one pair of laterals, while the type species of Venustatrochus
has as many as 17 pairs per transverse row. The majority of calliostomatines have 5-9 pairs of laterals
at maturity and interspecific variation by 1 or 2 pairs is common within groups of closely related
species. The low number of lateral teeth in Laetifautor species and the exceptionally small size of the
lateral and central teeth with respect to the innermost marginals strongly suggests that development
of the radula is retarded (Figs 136-137). Unlike juvenile calliostomatids (Waren, 1990: fig. 6B), teeth
in the central field in Laetifautor are slender with serrated edges, which suggests that the condition
is the result of a paedomorphic process rather than a primary one. By contrast, the adult radula in
species of Selastele gen. nov. (Figs 138-139) have exceptionally high numbers of lateral teeth relative
to body size, yet few marginal teeth (as in juveniles), while all of the teeth are weakly solidified and
semigelatinous. The number of marginal teeth has evidently been reduced by a paedomorphic process
that has differentially failed to retard multiplication of the lateral teeth during ontogenesis. For
Selastele the large number of lateral teeth and dissimilarity between young and adult teleoconch
suggests that small size is the result of dwarfism rather than a paedomorphic process.

Diet and feeding mode. — Although shallow water calliostomatines are frequently observed in
association with algae, careful field observations and examination of gut contents reveals that all are

Source : MNHN, Paris

CALLIOSTOMATIDAE FROM NFAV CALEDONIA

387

obligate carnivores (Marshall, 1988; Ferro & Cretella, 1993). The majority of species feed on
Cnidaria (commonly hydroids), and a few feed exclusively on sponges. From examination of many
radulae from about 100 species of calliostomatids from throughout the world belonging to a variety
of genera and subgenera, the only teeth that exhibit any significant degree of wear are the enlarged
innermost pair of marginals. Evidently, therefore, all of the other teeth are involved primarily in food
retention and clasping for transportation into the mouth.

When an excised calliostomatine radula is stretched apart and bent as if it were being pulled
around the tip of the odontophore, the innermost pair of marginal teeth stand vertically and rigid
with their laterally compressed vertical cutting areas aligned on the longitudinal axis. Thus the
columns of innermost marginals function like a pair of chainsaws as the radula is worked back and
forth around the odontophore tip by the massively developed buccal muscles. The tips of the (flexible)
immediately adjacent laterals are commonly observed to be cleanly broken or entirely missing at the
anterior end of the radula. and this damage is clearly incurred through accidental emplacement
between the slashing innermost marginals and the prey/substratum. Several species are known in
which the "cutting" areas are naturally absent from the outermost 1-3 pairs of laterals throughout
the entire adult radula. It is possible that these "cutting" areas are progressively reduced during
ontogenesis or they may simply never develop (juvenile radulae not seen).

In the new tribe Fautricini (see below), all of the innermost 9 pairs of marginal teeth are
greatly enlarged and strengthened during ontogenesis, and clearly all of them are directly involved in
food preparation (Figs 144-145). Marshall (1988) hypothesised that the bizarre radula in
Thysanodontinae (Figs 148-149) may be somehow involved in suctorial feeding.

Subfamily Calliostomatinae Thiele, 1924

Diagnosis. — Shell as for Calliostomatidae. Radula with the formula co + 1-17 + 1 + 17-1
+ co. Central and lateral teeth typically thin or very thin in section, with angulate, finely serrate tips.
Innermost pair of marginal teeth typically shorter and stouter than other marginals, and modified to
some degree. Outer marginals slender with narrow, finely serrate tips; outermost lew marginals
usually with blunt, cuspless tips and often fused.

Included genera and subgenera: see appendix 1.

Genus Calliosto.ua Swainson, 1840

Calliostoma Swainson. 1840: 218, 351. Type species (SD by Herrmannsen. 1846: 154): Trochus conulus Linnaeus. 1758: Recent,
north eastern Atlantic and Mediterranean.

Synonyms:

Ziziphinus Gray, 1840: 151 = Nomen nudum. .

Conulus Nardo, 1841: 244. Type species (OD): Troclius conulus Linnaeus. 1758 (not Conulus Leske, 1778, not Fitzinger. 1833).
Ziziphinus Gray. 1842: 57. Type species (SD by Gray 1847: 145 and by tautonomy): " Trochus ziziphinus " - Trochus zizyphinus
Linnaeus. 1 758; Recent, north eastern Atlantic and Mediterranean.

Zizyphinus Gray. 1847: 237. Incorrect subsequent spelling of Ziziphinus Gray, 1842.

Callisioma C'ullisiomus Herrmannsen. 1846: 154. 1852: 22. Unnecessary emendations of Calliostoma Swainson.

Jacinthinus Monterosato, 1889: 79. Type species (SD by Clench & Turner. 1960: 11): Trochus conulus Linnaeus. 1758.

Remarks. — Swainson (1840: 218) first introduced Calliostoma as a subgenus of Trochus
Linnaeus, 1758 with the entry "... forming our subgenus Calliostoma ... Trochus zizyphinus of British
writers will give a very good idea of these shells ...”. This does not constitute original designation ot
a type species (ICZN Art. 67c). Contrary to the opinion of some authors (e.g. Clench & Turner.
1960; Abbott. 1974), T. zizyphinus is not type species by monotypy either (Art. 68d), since Swainson
(1840: 351) included eight taxa in the formal diagnosis of Calliostoma, including T. zizyphinus and 7.
conulus (as zizyphina and conula). Since no type species was designated, the type species is T. conulus
through subsequent designation by Herrmannsen (1846: 154) (Art. 69a).

388

BRUCE MARSHALL

At its earliest introduction by Gray (1840: 151), Ziziphinus appears in a list without associated
species, description or comment and is thus a nomen nudum (Iredale, 1913). At the second
introduction the name is again without associated species. Gray (1842: 57) stating (under Thalolia )
“in Ziziphinus, Cantharidus, and Thalolia the mouth is oblong and simple and the axis is covered by
the inner lip; the former is top-shaped, the Cantharidi are ovate and green within". The first use of
Zizyphinus (note spelling) in connection with nominate species was by Gray (1843: 237) who included
four New Zealand species and two mislocalised North American species. Gray (1847: 145)
subsequently designated the European species T. ziziphinus (i.e. Trochus zizyphinus Linnaeus, 1758)
as type species of Ziziphinus Gray, 1840, while Reiider (1937: 115) selected the North American
species Trochus canaliculatus “Martyn" Lightfoot as type species of Zizyphinus Gray. 1843. On the
face of it, therefore, Ziziphinus and Zizyphinus would seem to be distinct genus group taxa with
separate type species, and they were so interpreted by Keen (1960). It seems preferable, however, to
treat Ziziphinus in the spirit with which it was intended (tautonomy), and interpret T. zizyphinus as
type species of Ziziphinus Gray, 1842 by subsequent designation of Gray 1847. Gray's Zizyphinus is
eliminated as an incorrected subsequent spelling since it is neither a mandatory change nor a
demonstrably intentional emendation (ICZN Art. 33). Consequently Rehder's (1937: 1 15) selection of
T. canaliculatus as type species of Zizyphinus was unnecessary.

Subgenus Calliostoma (s. sir.) Swainson. 1840

Diagnosis. — Shell up to about 35 mm high, stout, narrowly to broadly conical, periphery
rounded or angulate, anomphalous. P1-P4 commencing immediately or PI later, strong to very
strong, persisting or becoming obsolete; axial sculpture very weak or absent, nodules absent or large
and bluntly rounded. Animal with large left and right neck lobes, and 3 or 4 small epipodial tentacles
on each side. Cephalic tentacles tapered, extending little beyond tip of snout. Central and lateral teeth
broad-based, thin in section and flexible, laterals with very slender tips. Jaw plates with short elements
at anterior margins.

Remarks. — The concept of Calliostoma (5. sir.) is here restricted to the following taxa: C.
conulus (Linnaeus, 1758), C. zizyphinum (Linnaeus, 1758), C. laugieri (Payraudeau, 1826), C.
occidentale (Mighels, 1842), C. gualtierianum (Philippi, 1848). C. hirondellei Dautzenberg & Fischer,

1896, C. grimaldii Dautzenberg & Fischer, 1896, C. cleopatra (Fischer, 1898), C. laqueatum (Locard.
1897) and C. oppansum (Locard, 1897). All of these occur in the north eastern Atlantic and/or
Mediterranean, with the exception of C. occidentale , which occurs in the north eastern and north
western Atlantic. The limits of morphological expression within the subgenus are uncertain, and 1
have only included species that seem most likely to represent a tight phylogenetic radiation. Inclusion
of C. occidentale , perhaps the most divergent in shell morphology, provides both morphological and
geographical links between eastern and western Atlantic species groups.

Three groups of western Pacific species with rather distinctive shell facies are here referred to
subgenera Ampullotrochus, Fautor , and Benthastelena. Despite the extreme conservatism of Callios¬
toma (5. lat.) world wide, these groups all seem to represent separate phylogenetic radiations. Indeed
one or more of them may eventually prove to be worth recognising as genera, but until the family
is better known I prefer a conservative approach. As stated above these groups are difficult to
objectively define in that the combinations of character states are difficult to quantify or verbalise.

Judging from illustrations, the following Recent European species may not belong in
Calliostoma (s. str.) and their precise relationships within Calliostomatidae are uncertain: Calliostoma
caroli Dautzenberg, 1927, C. leptophyma Dautzenberg & Fischer, 1896, Z. milneedwardsi Locard,

1897, C. normani Dautzenberg & Fischer. 1897, Z. triporcatus Fischer in Filhol, 1886.

Source : MNHN, Paris

CALLIOSTOMATIDAE FROM NEW CALEDONIA

389

Subgenus Factor Iredale, 1924

Fautor Iredale, 1924: 230. Type species (OD): Ziziphinus compius A. Adams. 1855: Recent, southern Australia.

Remarks. — Fautor is here utilised as a subgenus of Calliostoma for a group of predominantly
Indo-western Pacific taxa that differ from species of Calliostoma (x. str.) (Fig. 20) in having narrower,
more finely beaded spiral cords. 1 am unable to detect any constant differences between the two
groups in the radula. jaws or external anatomy. Although relative size of the spiral cords and nodules
admittedly seems rather trivial, viewed as a whole the two groups exhibit a distinctive mosaic
combination of character states that is difficult to objectively quantify or describe. Moreover, I am
strongly disinclined to treat them as synonyms because Fautor , as here limited, is almost certainly
polyphyletic.

In addition to the species described herein, the following species are considered to belong in
Fautor. Calliostoma admirandum Smith. 1906 (off Tranvancore), Zizyphinus allporti Tenison Woods,
1875 (southern Australia), Calliostoma funiculare Melvill, 1906 (Persian Gulf). Zizyphinus scobinatus
A. Adams, 1863 (Bombay), Calliostoma takujii Kosuge, 1986 (Bonin Islands), and perhaps the eastern
Atlantic species C. lithocplletum Dautzenberg, 1925.

The type species of Fautor (Z. comptus. Figs 4-6, lectotype Fig. 4) was based on specimens
reputedly from New Caledonia, although nothing strictly similar to it has ever been obtained in the
New Caledonia region. As correctly concluded by Brazier (1895), the type material is in fact
indistinguishable from the New South Wales form of a common indigenous southern Australian
species. C. (F.) comptum has at times been confused with Trochus poupineli Montrouzier, 1875 (e.g.
Fischer, 1879; Ikebe, 1942; Kosuge, 1984) although in fact Montrouzier's species differs widely and
is herein referred to Dactylastele gen. nov.

Calliostoma ( Fautor) boucheti sp. nov.

Colour Plate; Figs 3, 7-9, 117, 150, 155; Table 1

Type material. — Holotype (13.0 x 1 0.6 mm, 7.30 TW) mnhn. Paratypes; 90 mnhn, 1 ams
C201701, 1 bmnh 1995.013, 1 nmp, 1 usnm, 2 nmnz M262476.

Type locality. Off S. New Caledonia, chalcal 2, stn DW 75, 24°39' S, 168°40' E, 600 m.

Material examined. — All type material. New Caledonia, biocal: stn DW 66, 24°55' S.
168°22' E, 505-515 m, 5 dd (paratypes).

chalcal 2: stn DW 72, 24°55' S, 168°22' E. 527 m, 33 lv (31 mnhn, 2 nmnz). — Stn DW 73, 24°40' S,
168°38' E, 573 m, 8 lv (1 ams, 1 bmnh, 4 mnhn, 1 nmp, 1 usnm). — Stn DW 74, 24°40' S, 168°38' E.
650 m, 13 lv (paratypes). — Stn DW 75, 24°39' S, 168°40' E. 600 m, 4 lv (holotype and 3 paratypes).
smib 3: stn DW 1, 24°56' S, 168°22' E, 520 m. 2 lv (paratypes). - Stn DW 2, 24°53' S, 168°22' E.
530-537 m, 8 lv (paratypes). — Stn DW 3, 24°55' S, 168°22' E. 513 m, 7 lv paratypes). - Stn DW
5, 24°55' S, 168°22' E, 502-512 m. 4 lv (paratypes). — Stn DW 6. 24°56' S, 168°21' E. 505 m, 5 lv
(paratypes). Stn DW 7, 24°55' S, 168°21'E, 505 m. 5 lv (paratypes). — Stn DW 8, 24°45' S.
1 68°08' E, 233 m, 1 dd (paratype). — Stn DW 22, 23°03' S, 167° 19' E, 503 m, 2 lv (paratypes).

Distribution (Fig. 155). — South of lie des Pins, southern New Caledonia and northern
Norfolk Ridge, 233-650 m, living at 502-650 m.

390

BRUCE MARSHALL

Colour plate. — Holotypes of CaUiostoma species. Top row (left to right): Calliostoma ( Faulor ) boucheii, 13.0 x 10.6 mm;
C. (F.) houbricki, 12.8 x 10.8 mm; C. (F.) metivieri, 11.5 x 9.40 mm. Centre row (left to right): C. (F.)
Chester fieldense, 7.60 x 6.90 mm; C. ( Benthastelena) diadematum, 11.0 x 10.5 mm. Bottom row (left to right): C. (F.)
periglyptum, 10.2 x S.30 mm; C. (Amputloirochus) heros, 11.6 x 10.2 mm; C. (A.) xanthos, 8.00 x 6.90 mm.

Source : MNHN, Paris

CALLIOSTOMATIDAE FROM NEW CALEDONIA

391

Figs 2-3. Side views of tip of spire showing postlarva] grow th scar denoted by arrows. 2, Laetifaulor rubropunciaius (A.
Adams), off Swain's Reef. Queensland. 100 m, ams Cl 5321 5, x 147. 3. Calliostoma (Fautor) boucheti , paratype,

biocal: stn DW 66. x 123.

Description. — Shell up to 13 mm high, rather lightly
built, glossy, spire narrowly conical, very weakly cyrto-
conoid, 1.92-2.05x as high as aperture, mean spire angle
57-62°, anomphalous.

Colour of protoconch reddish brown. Teleoconch grading
from reddish to yellowish brown over 1st 2 whorls, subse¬
quent whorls bright orange to yellowish brown, pale green
nacreous layer showing through translucent outer layer in
spiral interspaces, inner and outer lip margins white.

Protoconch 400-430 u.m wide, sculptured with network of
fine threads that enclose roughly hexagonal spaces. Apical
fold tip rounded, terminal varix rounded.

Teleoconch of up to 7.30 whorls; 1st tenth whorl with 4
crisp spiral threads, delineated by growth scar. First 2 whorls
strongly convex, angulated at P3. 3rd whorl becoming flat,
subsequent whorls flat; periphery angulate. becoming tightly
rounded late on last adult whorl, S3 and P3 peripheral: base
weakly convex. Spire and basal spirals prominent, rounded,
multiplying by intercalation, their rounded conical nodules
stronger on spire; axial riblets strong on 1st 3 whorls,
becoming obsolete on 4th whorl. P2-P4 commencing imme¬
diately after post-larval growth scar: P2 and P3 strong,
similar, flange-like and weakly undulant on 1st whorl,
becoming nodular on 2nd whorl (P2) and 3rd whorl (P3). PI
a fine thread on 1st whorl, enlarging over 2nd whorl to
become as large as P2 and P3 on 3rd whorl, thereafter slightly
larger than others. P4 partly covered by succeeding whorls.

Secondary spirals enlarging to resemble primaries: SI
commencing late 3rd - mid 4th whorl, S2 commencing end
2nd - late 3rd whorl, S3 commencing mid - end 4th whorl.

Tertiary spirals numbering 1 or more, usually commencing
on last adult whorl, remain weaker than primaries and
secondaries. Base with 12-17 spiral cords, stronger and more
strongly nodular on inner half, nodular on outer half on last
adult whorl only. Microsculpture of fine crisp spiral threads
on 1st 2 whorls; fine collabral growth lines throughout,
crisply defined between P3 and P4 before appearance of S3.
Aperture subquadrate. Outer lip thin at rim, slightly thicke¬
ned within. Inner lip of moderate thickness. No parietal
inductura.

Animal creamy white, snout tip laterally expanded, set with
slender papillae. Cephalic tentacles long, tapered, finely
papillate, prominent eyes at outer bases. Neck lobes thin, left
considerably larger and more convoluted. 3 right and 4 left
epipodial tentacles. Operculum typical.

RaJula (Fig. 117). Central and lateral teeth thin in section
with broad, laterally expanded bases. Tip of central tooth
narrowly tapered, finely serrate. Lateral teeth similar. 6 pairs
in adults; tips very long, slender, finely serrate. Innermost
marginal teeth stout, broad-based, with 3 or 4 strong cusps.
Outer marginals numerous, tips finely serrate, outermost 5
marginals on each side fused, tips spathulate.

Jaw plate (Fig. 150) with broadly rounded anterior mar¬
gins, fringing elements short.

392

BRUCE MARSHALL

Table I . Calliostoma (Fautor) boucheti. Shell measurements (mm) and countings (chalcal 2: stns
DW 73, DW 74, DW 75).

Character

it

Range

Mean

SD

H

15

9.20-13.00

1 1.39

1.17

D

15

8.30-11.20

09.88

0.91

H/D

15

1.11-01.28

01.15

0.04

TW

15

6.25-07.30

06.85

0.29

Remarks. — Compared with the superficially similar Australian species C. (F.) comp turn, C.
(F.) boucheti differs in numerous details of shell, colour, shape and sculpture. These differences
include much finer protoconch sculpture, flange-like cords on the first teleoconch whorl, the later
development of PI, and the more evenly conical spire.

Etymology. Named after Philippe Bouchet (mnhn).

Calliostoma ( Fautor) richeri sp. nov.

Figs 10-12, 118, 153; Table 2

Type material. — Holotype (12.3 x 9.30 mm, 8.10 TW) mnhn. Paratypes: 34 mnhn, 1 ams
C201702, 1 bmnh 1995.014, 2 nmnz M262470. I nmp, 1 usnm.

Type locality. — S. New Caledonia, lagon, stn 400, 22°34' S, 167°14' E, 64 m.

Material examined. New Caledonia, lagon: stn 113, 22°23' S, 166°48' E, 32 m, 2 dd
(paratypes). — Stn 146, 22°24' S. 166°55' E, 40-52 m. 3 dd (paratypes). - Stn 234, 22°33' S, 1 66°5 1 ' E,
56 m, 1 dd (paratype). — Stn 240, 22°23' S, 166°59' E, 42 m, I dd (paratype). — Stn 297, 22°39' S,
166°46' E, 30 m, 1 dd (paratype). — Stn 301, 22°35' S, 166°52' E, 46 m, 1 dd (paratype). — Stn 303,
22°38' S, 166°49' E