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REVISION OF THE
NORTHEAST ATLANTIC
BATHYAL AND ABYSSAL
NEOGASTROPODA
EXCLUDING TURRIDAE
(MOLLUSCA, GASTROPODA)

PHILIPPE BOUCHET

Museum National d’Histoire Naturelle, Paris
and

ANDERS WAREN
Zoologiska Institutionen, Goteborg

This supplement has been published with support from the Swedish Natural Science

Research Council

societA italiana di malacologia

Bollettino Malacologico, Supplemento 1
Milano novembre 1985

S.I.M. - societA italiana di malacologia

c/o Acquario Civico, viale Gadio 2, I - 20121 Milano

La SIM e una Societa senza fini di lucro che si propone di incoraggia-
re e diffondere lo studio dei molluschi continentali, marini e fossili con
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© 1985, The Authors & Societa Italiana di Malacologia

Typesetting by Angus Graham Associates Ltd., Reading
Printed in Italy by GRAFICHE ATA, Paderno Dugnano (Milano)
Published november 1985

REVISION OF THE
NORTHEAST ATLANTIC/
BATHYAL AND ABYSSAL
NEOGASTROPODA ~ ,
EXCLUDING TURRIDAE
(MOLLUSCA, GASTROPODA

PHILIPPE BOUCHET

Museum National d’Histoire Naturelle, Paris

and

ANDERS WAREN

Zoologiska Institutionen, Goteborg

This supplement has been published with support from the Swedish Natural Science

Research Council

This paper constitutes the second part of a monograph of the N.E.
Atlantic bathyal and abyssal Gastropoda. Page and figure numbers in this
volume follow numbers in the first volume.

Part 1, covering the family Turridae, was published as J. Moll. Stud, sup¬
plement 8 (1980).

Part. 3, covering the families Eulimidae, Aclididae and Epitoniidae, will
also be published by Societa Italiana di Malacologia towards the end of

1985.

societA italiana di malacologia

Bollettino Malacologico, Supplemento 1
Milano novembre 1985

NORTH-EAST ATLANTIC NEOGASTROPODA 123

CONTENTS

Contents . . . 123

Abstract . 124

Introduction and material . 125

Relocated species . . 125

Taxonomical part . 126

Muricidae . . 126

Coralliophilidae . 151

Nassariidae . . 158

Fasciolariidae . . 160

Columbellidae . 165

Buccinidae . . . 170

Olividae . . 249

Volutomitridae . 251

Turbinellidae . . 254

Cancellariidae . 257

Marginellidae . . 267

Acknowledgements . 276

References . . 277

Index . 286

Distribution of the species

294

124 P. BOUCHET & A. WAREN

ABSTRACT

A revision of the deep sea neogastropods (excluding Turridae) is presented. The area surveyed
covers the Atlantic N of 33° N, E of the Mid Atlantic Ridge, including the Mediterranean and the
Norwegian Sea. Only species normally occurring below 300 m are included. All previously published
material and unpublished material from the French, British and German deep sea programmes is
included. The revision includes species of Muricidae, Coralliophilidae, Nassariidae, Fasciolaridae, Co-
lumbellidae, Bucciniaae, Olividae, Volutomitridae, Turbinellidae, Cancellariidae and Marginellidae.
About 385 names of the species category are discussed and reduced to 80 species.

The new species Pterynotus atlantideus, Microvoluta superstes, Metzgeria gagei, Iphinopsis alba, I.
fuscoapicata, Admete azorica, Volvarina ingolfi and Marginella aronnax are described. Buccinum sulca¬
tum Friele is renamed B. kjennerudae, Turrisipho lachesis var. bicarinata (Friele) is renamed T. voerin-
gi and Colus ventricosus (Gray) is renamed C. terraenovae because of homonymy.

RIASSUNTO

La presente revisione dei Neogastropodi di grande profondita segue la revisione dei Turridae ba-
tiali e abissali dell'Atlantico nord-orientale, pubblicata dagli stessi autori (1980). La regione esamina-
ta comprende 1’Atlantico settentrionale a nord del 33° parallelo e a est della dorsale medio-oceanica,
ivi compresi il Mediterraneo e il mare di Norvegia. Sono prese in considerazione solo le specie che
vivono normalmente sotto i 300 m di profondita.

La revisione si basa su tutto il materiale storico e sul materiale proveniente dai programmi di
studio del benthos profondo francesi, britannici e tedeschi. In questo volume sono trattate le seguenti
famiglie: Muricidae, Coralliophilidae, Nassariidae, Fasciolariidae, Columbellidae, Buccinidae, Olivi¬
dae, Volutomitridae, Turbinellidae, Cancellariidae e Marginellidae. Sono state discusse 385 denomi-
nazioni di gruppo-specie con conseguente riduzione a 80 specie valide. Infine sono state istituite otto
specie nuove e sono state proposte tre nuove denominazioni.

Per quanto concerne la nomenclatura e la sistematica delle specie mediterranee si perviene alle
seguenti conclusioni:

1. Trophon echinatus (Kiener, 1840) e il nome valido per la specie nota generalmente come Pago-
dula vaginata e P. multilamellosa. Questa specie presenta un cline batimetrico: le popolazioni batiali
atlantiche e mediterranee (-250 / -600 m) appartengoono alia forma grimaldii Dautzenberg & Fischer,
caratterizzata da un maggior numero di lamelle assiali rispetto a quelle della piattaforma continen-
tale.

2. Coralliophila squamosa (Bivona, 1838) e il nome valido per la specie nota come C. alucoides e
C. squamulosa, entrambe pre-occupate.

3. Coralliophila lactuca Dali, 1889 e sinonimo di C. richardi (P. Fischer, 1882) che ha quindi prio¬
ri ta.

4. Nassarius frigens (v. Martens) ha distribuzione limitata all’Atlantico dell 'Africa tropicale e sub-
tropicale. La presenza di Nassarius wolffi (Knudsen, 1956) in Mediterraneo resta da confermare.

5. Fusinus gigliolii (Monterosato, 1890) e da considerarsi come una forma di profondita, priva di
valore tassonomico, di F. rostratus (Olivi, 1792).

6. Amphissa acutecostata (Philippi, 1844) e il nome valido della specie nota sotto il nome, pre-
occupato, Amphissa costulata.

7. Il genere Colus e rappresentato in Mediterraneo solamente da C. jeffreysianus (Fischer, 1868).

8. Granulina Jousseaume, 1888 e il nome del genere valido per le piccole Marginellidae general¬
mente note sotto il nome Gibberulina e Cypraeolina.

NORTH-EAST ATLANTIC NEOGASTROPODA 125

INTRODUCTION

This revision forms part 2 in a series treating the NE Atlantic deep sea gastropod fauna. For the first
part, see Bouchet & Waren (1980). The revision is based on all material we have been able to
examine, from numerous sources, which are listed in part 1. For part 2 we have to a higher degree
been able to base our work on material in Scandinavian museums, because much of the revision
involves northern members of the family Buccinidae. Detailed information about this material can
be found in Thorson (1941, 1944, 1951) and Friele & Grieg (1901).

We have also been able to use a large supply of material from the Rockall Trough, collected by Dr
John Gage from R.V. Challenger II. This material is now in the Royal Scottish Museum, Edinburgh.

A rich supply of NW Atlantic buccinids was collected by Dr J.C. Mahe, Institut Scientifique et
Technique des Peches Maritimes, St Pierre & Miquelon, and put at our disposal. The material is now
in MNHN.

Several members of the Bologna group of the Unione Malacologica Italiana have contributed
material of rare Mediterranean shelf species and fossil material from their private collections.

Material from the French Seabed and Walvis expeditions has also been used. The area investigated
by these expeditions is to the south of the area we have covered, but the material was useful for
comparisons. It is now in MNHN.

Abbreviations are listed in part 1 .

SPECIES REFERRED TO TAX A NOT TREATED IN THIS PART

Sipho leptaleus Verrill, 1884:175, pi. 31, fig. 14. We have examined the holotype (USNM 202879),
from off Martha’s Vineyard, 810 m (off the NE US), and it proved to be a turrid, probably a species
of Ta ranis.

Nassa nigrolabra Verrill, 1880:371, from off Martha’s Vineyard, 280 m. We have examined the
holotype and it is a larval shell, probably of a cymatiid.

Trophon maclaini (Dali, 1902), described from off Greenland, does not belong to Trophon, but was
based on a species of Oenopota (Turridae). We have examined the holotype, USNM 126 194, and it
must have been a slip of the pen when Dali placed it in Boreotrophon.

126 P. BOUCHET & A. WAREN

Family MURICIDAE

Among the muricids, it is the genus Trophon that constitutes the largest group below the continental
shelf in the NE Atlantic. This seems to be the case also in other seas. Other genera are represented by
scattered species of groups normally inhabiting shallow water or the continental shelf. Trophon on
the contrary extends from shallow to deep water in high northern or southern areas, while in
intermediate latitudes it is restricted to deep water.

Genus TROPHON Montfort, 1810

Type species: Murex magellanicus Gmelin, 1791 , by original designation. Several generic names have
been proposed for the species we include in Trophon viz.:

Trophonopsis Bucquoy, Dautzenberg & Dollfus, 1882. Type species: Murex muricatus Montagu,
1803, by original designation.

Boreotrophon P. Fischer, 1884. Type species: Murex clathratus Linne, 1758, by monotypy.
Pagodula Monterosato, 1884. Type species: Murex carinatus Bivona, 1832 (= Murex vaginatus De
Cristofori & Jan, 1832), by monotypy.

Pinon De Gregorio, 1885. Type species: Pinon vaginatus Philippi (sic!), by original designation.
Chalmon De Gregorio, 1885. Type species: Chalmon muricatus (Montagu) by original designation.

Pinon and Chalmon are thus objective synonyms of Pagodula and Trophonopsis respectively.
There are, however, some doubts whether the Recent or the fossil species known under the name
Trophon vaginatus should be considered the type species.

Recent workers on the genus (Radwin & D’Attilio, 1976; Vokes, 1976; Houart, 1981) have
advocated a separation of the southern species usually included in Trophon and the northern ones,
which they place in Boreotrophon and Pagodula. (Houart also used the genera Nodulotrophon
Habe & Ito, 1965 and Nipponotrophon Kuroda & Habe, 1971 for some NE Atlantic species.) We
have examined radulae and opercula of the type species, T. magellanicus, and of all the NE Atlantic
species and are not able to separate them into different groups. The species are so variable that they
will usually fit equally well into two or more “genera”. It is only T. fabricii that to some degree
stands out from the others, by its strongly-shouldered whorls and strong spirabsculpture. It also has
a radula that is proportionally longer in relation to the height of the shell. The significance of these
characters can, however, not be evaluated until they are known for many more species of the group.

The separation of the S American Trophon from the N Atlantic genera has been made only on the
basis of unspecified differences in shell morphology and on the grounds that SE American Trophon
should have a “purpuroid” operculum, while the NE Atlantic species should have a “muricoid”
operculum (Radwin & D’Attilio, 1976, Vokes, 1976). We have figured the operculum of Trophon
geversianus and some NE Atlantic species for comparison (Figs 319-327) and although there is a
difference in strength of the concentric ridges, this (or any other difference) is not sufficient to divide
the S American and NE Atlantic species into two groups. In a comparison of opercula, T. fabricii
again stands out, by having a much more distinct pattern. (It should be noticed that opercula of
young Trophon are paucispiral cf. Fig. 322.)

Comparison of the radulae did not reveal anything except details in the proportions of the cusps
(Figs 329-332), differences insufficient to warrant separation.

NORTH-EAST ATLANTIC NEOGASTROPODA 127

Neither was a separation based on shell characters possible. Strebel (1905) in his treatment of the S
American Trophon shows very well the variation in shell morphology of the genus, from the large,
stout forms like geversianus to species like laciniatus (Dillwyn, 1817) that in shell morphology agree
well with our NE Atlantic species like T. clathratus.

The only remaining difference between the S American and NE Atlantic species (pointed out by
Thorson (1946), and to us in a letter by E. Vokes) is the egg capsules. Thorson (1940, 1946) reported
that the capsules of T. truncatus and T. clathratus were planoconcavely lens-shaped, with a few
embryos, and attached by the flat side to some hard substratum. Jeffreys (1867) described similar
egg capsules in T. muricatus.

Strebel (1905 & 1920) and Penchaszadeh (1976) reported those of T. geversianus and two other S
American species to be lens-shaped and attached by a short broad stalk from the edge. It is
impossible to understand what this difference means taxonomically, especially as the spawn is
known for very few species.

Because of this absence of distinguishing characters we have decided to use Trophon, to include
all the NE Atlantic species.

The species of Trophon are unusually variable (easily seen from the long lists of synonyms) and it
may seem possible to arrange a clade connecting all the different forms if one uses specimens from
many different localities. We have, however, found it possible to distinguish nine species (possibly
only eight), by the characters listed below. These nine (eight?) species have been examined from all
or large parts of their geographic and bathymetric distribution. They typically exhibit a variation
from strongly-shouldered and spiny specimens, to more slender forms with weak sculpture (cf. Figs
286-287, 309-312). This variation is usually correlated with a depth or geographical cline, but one can
also find local populations with a unique appearance. Two exceptions from this are T. dabneyi and
T. droueti, which are fairly constant. A lower degree of variation is consistent with their biology. T.
dabneyi is the only NE Atlantic species with planktotrophic larvae, which possibly leads to less local
variation and T. droueti is restricted to the continental slopes around the Azores.

Distinctive characters of the species

Trophon fabricii is an arctic shallow-water species with very distinct spiral sculpture, strongly-
shouldered whorls, rather thick and solid axial ribs and usually a crenulated outer lip.

Trophon dabneyi has a yellowish, large larval shell of more than two whorls, slender, rounded
whorls and low, never spiny ribs. It is a decidedly bathyal species that is never found on the shelf.

Trophon droueti has a low spire, rather solid shell, and a thickened flaring outer lip with internal
crenulations. It is known from the bathyal area of the Azores.

Trophon truncatus is an arctic shallow-water species with rather slender and rounded whorls,
rounded larval shell, no spiral sculpture and low, crowded axial lamellae that never form spiny
projections.

Trophon clathratus is an arctic shallow-water species with a distinctly angulated larval whorl, high,
sparse axial ribs that often form spiny projections from the rather low and convex whorls. No spiral
sculpture.

Trophon clavatus is a boreal species occurring in intermediate depths. It resembles T. clathratus, but
has a rounded larval shell and is half the size of T. clathratus. It may have some indication of spiral
sculpture, but lacks it normally. Specimens from shallow water often are quite slender, compared
with T. clathratus.

Trophon barvicensis is a boreal species, possibly reaching the Mediterranean. The shell is greyish to
whitish, rather slender, with distinct spiral ribs that give the axial lamellae a fringed appearance.
Young specimens may have spiny projections from the lamellae, but we have not seen them in full-
grown ones.

128 P. BOUCHET & A. WAREN

Trophon muricatus is a rather southern species, reaching northwards to S Great Britain. It usually
has a fairly solid, brownish or reddish shell, strong spiral ribs, low and solid axial ribs and a quite
slender shape.

Trophon echinatus is an extremely variable, southern species that occurs from fairly shallow (100 m)
down to the lower bathyal. In shallow water the shell is semitransparent and of a glassy structure, in
deep water more chalky. The most characteristic feature is the long slender siphonal canal and
distinctly shouldered whorls. Shallow-water specimens lack spiral sculpture, while it is well
developed in specimens from deep water. Shallow-water specimens have very well-developed spiny
projections from the axial ribs, but they may be lacking in specimens from deep water.

Trophon truncatus (StrOm, 1768) Figs 282-283

Buccinum truncatum StrOm, 1768:369, fig. 26.

Trophon truncatus vars alba & scalaris Jeffreys, 1867:320.

Trophon truncatus var. abbreviata MOrch, 1869:195.

Trophon truncatus var. major BrPgger, 1901:654 (1900:50).

Trophon (Boreotrophon) truncatus var. intermedia Harmer, 1918:350, pi. 36, fig. 24.

Trophon (Boreotrophon) clathratus var. attenuata Harmer, 1918:351, pi. 36, figs 7-8.

Trophon (Boreotrophon) clathratus var. exilis Harmer, 1918:352, pi. 36, figs 9-1Q.

Type material: T. truncatus, lost.

Type locality: T. truncatus, N Norway.

Material examined : Several hundred shells from all parts of its distribution in the N Atlantic (SMNH, ZMC,
MNHN, USNM, coll. AW).

Distribution: N Atlantic coastal areas; Parts of Ireland and Scotland (Seaward, 1982), the Kattegat,
absent in the Skagerrak and W Norway, common in N Norway (G.O. Sars 1878), the Faroes,
Iceland and Greenland (Sparck & Thorson, 1933; Thorson, 1941, 1944, 1951), NE America S to
George’s Bank (off Massachusetts) (Abbott 1974). The depths are normally 3 — 150 m, although
Golikov (1964) has recorded it from 950 m. Probably panarctic.

Remarks: T. truncatus has often been considered a variety or form of T. clathratus, but we consider
it a good species. It can be distinguished by its more evenly convex whorls, more numerous axial
lamellae and rounded protoconch, from T. clathratus, which has a distinctly angulated initial whorl.
Neither does it have any tendencies to form spiny processes from the ribs as clathratus.

The specimens figured by Houart (1981, pi. 6, figs 9-10) as Boreotrophon sp. are large specimens
of T. truncatus.

NORTH-EAST ATLANTIC NEOGASTROPODA 129

Trophon clathratus (Linne, 1767) Figs 284, 285, 325-326

Murex clathratus Linne, 1767:1223.

Buccinum lyratum Gmelin, 1790:3494.

Buccinum lamellatum Gmelin, 1790:3498.

Murex bamffius Donovan, 1804: pi. 169.

Fusus costatus Hisinger, 1837:43 (not Philippi, 1836). {Fide Hagg, 1925).

Fusus lamellosus Gray, 1839:118, pi. 36, fig. 13 (not Borson, 1821).

Fusus scalariformis Gould, 1840:197.

Tritonium gunneri Loven, 1846:144.

Tritonium clathratum f. normalis, elatior, ventricosa Middendorff, 1849:455-457.

Tritonium rossi Leach (MS?) Morch, 1858:85.

Trophon richardsoni Gray (MS?) Morch, 1869:21.

Trophon clathratus var. grandis Morch, 1869:21 .

Trophon cepula Sowerby , 1880:61 , pi. 404 fig. 14, pi. 405 fig. 27 (replacement name for Fusus lamellosus Gray).
Trophon clathratus var. intermedius, maximus Verkriizen, 1881 : 84-85.

Trophon gunneri var. coddi Harmer, 1918:353.

Type material T. clathratus , in the Linnean Society, London.
Type locality T. clathratus , Icelandic Seas.

Material examined: Several hundred shells and specimens from the Swedish west coast (glacial fossils), N
Norway, Spitzbergen, Iceland and Greenland (ZMC, SMNH, coll. AW).

Distribution: Arctic circumpolar in coastal waters and on the continental shelf. It reaches south to
the New England area (Macpherson, 1971), E and W Greenland (Thorson, 1951), Iceland
(Oskarsson, 1962), the Faroes (Sparck & Thorson, 1933) and Lofoten (N Norway) (G.O. Sars,
1878). The bathymetric distribution is 4.5 m (Macpherson, 1971) to about 300 m.

Remarks: Thorson (1940) described the reproduction. The species has direct development and lens¬
shaped egg capsules. Some attention should, however, be paid to the fact that Oenopota elegans
(Turridae) has a very similar larval shell, and we are not aware how Thorson, who identified the
capsules from the shells of the embryos, could distinguish between these two species.

We have listed Fusus lamellosus Gray as a synonym of T. clathratus . This is not certain, but this
will not affect any other name, because the name is preoccupied. We have not examined any Pacific
material of T. clathratus , but the names based on such material are all younger than that of Linne,
so they will not change it, if they should prove to be based on the same species. The distribution
mentioned under that heading is much more arctic than is usually found in the literature. This is
because we have distinguished the southern deep-water populations as a distinct species, T. clavatus.
For a comparison with that species see p. 130.

Some Icelandic specimens seem to have a more rounded protoconch than the typical arctic form,
but the specimens are quite worn. Their shells, however, are typical for the arctic form. This point
should be examined when better material becomes available.

130 P. BOUCHET & A. WAREN

Trophon clavatus G.O. Sars, 1878 Figs 286-290, 322-324, 331, 339
Trophon clavatus G.O. Sars, 1878: 249, pi. 15, fig. 12; pi. 23, fig. 14.

Type material: Lectotype and 2 paralectotypes, ZMO D1396 a and b, selected by Houart (1981).
Type locality: Lofoten, N Norway, 200-360 m.

Material examined: Several thousand specimens from SW Norway and the Swedish west coast, 50 — 500 m, coll.
AW; THOR 1903, S of Iceland, probably 1000 m, 3 shs; THOR 1904, st 99, 61° 15 N,09°35 W, 900 m, 1
spm; INGOLF st 90, 64°45 N, 29°06 W, 1000 m, 2 shs; INGOLF st 78, 60°37 N, 27°52 W, 1505 m 1 sh.

Distribution: Uncertain, known with certainty only from the material examined, and some
specimens determined by G.O. Sars (1878).

Remarks: We have decided to keep this species or form separate from T clathratus, as did Houart
(1981), although other authors have united it with J. clathratus. The distribution is incompletely
known because of this 'Tumping”, but probably some specimens referred to T. clathratus by Sykes
(1911) and Massy (1930) from off NW Scotland and Ireland will prove to belong to clavatus.

T. clavatus is very common off the W coast of Norway and Sweden, in 50 — 500 m, often together
with T. barvicensis (cf. that species), but it seems to be very rare elsewhere.

The reasons why we have decided to distinguish between the two species are:

1 . T. clavatus has a distinctly rounded larval shell while that of typical T. clathratus has a distinctly
angulated periphery.

2. In Norway typical T. clathratus is restricted to Lofoten and northwards, in 20 — 90 m (G.O. Sars,
1878). This is the large form with angulated larval shell. Further to the south we find the smaller
form, with rounded larval shell, and then usually below 100 m with its main distribution between 200
and 500 m.

3. In Iceland typical T. clathratus is common, mainly off the N and NW coasts, in 11 — 260 m
(Thorson, 1941; Oskarsson, 1962). It has not been recorded deeper. We have, however, found some
specimens in ZMC (see above) from 900 — 1500 m. These belong to the same form as we find in deep
water in Scandinavia, although they have a tendency to be still more spiny (Fig. 286).

4. In the Faroes typical T. clathratus is known from 8 — 50 m (Sparck & Thorson, 1933). The
clavatus form is not known from there.

5. T. clathratus in its typical form is common along the coasts of Greenland and E Canada, S to

Maine, in shallow water (30 100 m, Verrill, 1882), while in deep water we find Trophon

abyssorum Verrill, 1885, with a more southern distribution. The clavatus form is not known here.
Therefore we suppose that two species, (or at least subspecies) have been confused; the more or less
panarctic T. clathratus » and T. clavatus with a rather restricted distribution in the NE Atlantic. G.O.
Sars (1878) was not aware of this when he described T. clavatus , because he used the name for some
extremely large specimens with a typical appearance, while he regarded the S Scandinavian
specimens from deep water as a small form of T. clathratus.

As our figures illustrate, T. clavatus is quite variable. Some specimens resemble T. barvicensis
closely, but we have noticed this only in young specimens, when sometimes T. barvicensis has a
poorly-developed sculpture and T. clavatus may show some undulations in the axial lamellae. We
have never had any difficulties in separating half-grown or larger specimens. Neither have we had
any problems separating T. clavatus from T. clathratus, but that may be because we have not been
able to examine more than a few specimens from areas where both species occur.

NORTH-EAST ATLANTIC NEOGASTROPODA 131

Trophon barvicensis (Johnston, 1825) Figs 291-296, 341

Fusus barvicensis Johnston, 1825:221.

Type material : Not known.

Type locality: Near Berwick, NE England.

Material examined: Several thousand specimens from S Norway and the Swedish W coast, coll. AW; a few

hundred shells and specimens from the British Isles, the French W coast and Iceland, MNHN, USNM, ZMC;
some doubtful specimens from the Mediterranean, private collections, MNHN, USNM; off Morocco, a few
shells, MNHN.

Distribution : From W Finmark, N Norway (G.O. Sars, 1878), Iceland (Thorson, 1941), southwards
on the shelf and in coastal waters to off Morocco (Locard, 1897), where it occurs in the upper
bathyal zone.

Remarks: T, barvicensis is often very common on the shelf off S Norway, where it occurs together
with T. clavatus in large numbers. Some young specimens may then be difficult to distinguish from
that species, but they are quite rare and can be separated by having a microsculpture of scattered
tubercles on the protoconch, while T. barvicensis has these tubercles arranged in lines.

We have not seen any unquestionable Mediterranean specimens. Some specimens reported on by
Di Geronimo & Panetta (1973) and Franchini & Frilli (1970), determined as T. richardi, resemble T.
barvicensis closely, but we have seen all interrmediates between these specimens and T. muricatus
from closely-situated localities. (Similar observations have been made by Chaster (1898) and Chaster
& Heathcote (1894) in the British Isles.) Therefore we are not certain if T, barvicensis lives in the
Mediterranean. The question arises that T. barvicensis may be only a deep-water form of T.
muricatus, an assumption that is supported by the doubts expressed by Chaster (1898). However, in
Scandinavia barvicensis occurs more shallowly, from 50 m, thus occupying depths where muricatus
should be expected, yet barvicensis does not change form. The problem can, however, not be
considered settled.

Some specimens from the Azores ( Trophon sp., p. 131, Fig. 297) resemble T. barvicensis closely,
but have more convex larval whorls, they are more spiny and occasionally have a coloured shell. This
may be a local form of barvicensis, but because of the differences mentioned above and the
geographical separation, we prefer to keep it separate, at least until we know if T. barvicensis occurs
on any of the intermediate sea-mounts.

Trophon sp. Fig. 297

Material examined: BIACORES st 84,38°58 N, 28°03 W, 190 — 220 m, 2 spms; st 90, 39°04 N, 28°07 W, 205 —
210 m, 1 sh; st 159, 37°26 N, 25°51 W, 525 — 600 m, 1 spm.

Remarks: We have only seen four specimens of this form of Trophon, all from the Azores. It differs
from T. droueti, which occurs in the same area and the same depths, by being more slender and
having stronger projecting spines on the shoulder of the whorls. We have not seen any intermediates
between the two. There is a possibility that this is a form of barvicensis, but as nothing is known
about the occurrence of barvicensis on the sea-mounts between the Azores and Europe, it cannot be
discussed.

132 P. BOUCHET & A. WAREN

Figs 282-290. Genus Trophon. 282, T. truncatus, TromsO area, 8 m, 10.2 mm. 283, T. truncatus,
Godhavn, Greenland, 50-100 m, apex 1.3 mm. 284, T. clathratus, TromsO area, 8 m, 6.8 mm. 285, same
sample, apex 1.1 mm. 286, T. clavatus, THOR 1904 st 99 (61° 15 N, 09°35 W, 900 m), 1 1.2 mm. 287, T.
clavatus, off Korsfjorden, Bergen area, 240-250 m, 7.8 mm. 288, T. clavatus, Sacken, Swedish west
coast, 13.9 mm. 289 and 290, T. clavatus, Bergen area, 285-290 m, apex 0.9 mm.

NORTH-EAST ATLANTIC' N HOG ASTRO POD A 133

Figs 291-297. Genus Trophon. 291, T. barvicensis, Sacken, Swedish west coast, 10.9 mm. 292, T.
barvicensis, THALASSA st Z428 (48°27 N, 10°50 W, 850 m), 17.1 mm. 293, T. barvicensis, THALASSA
st Y378 (41 °31 N, 09° 16 W, 1000 m), 9.05 mm. 294, T. barvicensis, Bergen area, 240-250 m, 4.7 mm. 295
and 296, T. barvicensis, Bergen area, 322-330 m, apex 0.6 mm. 297, T. sp., BIACORES st 84, 9.5 mm.

134 P. BO U CHET & A. WAREN

Trophon muricatus (Montagu, 1803)

Figs 298-301,342

Mur ex muricatus Montagu, 1803:262.

Fusus asperrimus Leach in Brown. 1827: pi. 47, fig. 2.

Fusus cancellatus Bivona, 1838:14 (not Sowerby, 1826 or Valenciennes, 1832).

Trophon muricatus var. lactea Jeffreys, 1867:317.

Fusus longurio Weinkauff, 1866:244, pi. 5, fig. 4.

Trophon breviatus Jeffreys, 1882:426.

Trophon muricatus var. aspera Monterosato MS, Bucquoy, Dautzenberg & Dollfus, 1882:40.
Trophonopsis curta Locard, 1892:109 (partim).

Trophonopsis muricatus var. minor Locard, 1897:349.

Trophon breviatus var. striata & var. lactea Milaschewitsch, 1916:105, pi. 3 figs 51-53.
Trophonopsis muricatus albinus Monterosato in Settepassi, 1977:7.

Trophonopsis muricatus major Monterosato in Settepassi, 1977:7.

Type material: T. muricatus , not examined; T. longurio, holotype USNM 192693; T. breviatus,
syntypes USNM 192695.

Type locality: T. muricatus, Salcombe Bay, G.B; T. longurio, Alger; T. breviatus, off the mouth of
the Danube, Black Sea.

Material examined: The types ennumerated above and a few hundred shells from all parts of the distribution
area USNM, BMNH, MNHN, coll. AW.

Distribution: European coastal areas, from Great Britain south to the Mediterranean and Morocco,
in 20 — 300 m.

Remarks: Several authors (Franchini & Frilli, 1970; Houart, 1981) have recently synonymized T.
barvicensis and T. muricatus. As we have discussed under T. barvicensis we do not find the evidence
for this convincing enough. We have seen samples with the two forms occurring sympatrically, from
the Bay of Biscay and N Morocco, from 120-200m. We have, however, also seen samples where it
seems impossible to distinguish the two (e.g. from the Adriatic, coll. G. Spada, Bologna). The
Mediterranean material we have seen has been insufficient to ascertain whether barvicensis really
occurs in the Mediterranean and the problems are caused by a higher variability of muricatus there,
or if they really integrade. The latter seems to be supported by Chaster’s (1898) observation that they
intergrade in the British Isles, but contradicted by our observations from other areas. Therefore we
prefer to leave the question open and treat them as two species.

Jeffreys (1867), Lebour (1936) and Fioroni (1966) have described the egg capsules, which are lens¬
shaped and attached to some hard substratum.

The Black Sea form, breviatus, is probably worth subspecific status. It has been figured by Houart
(1981) and Waren (1980).

NORTH-EAST ATLANTIC NTOdASTROPODA 135

Figs 298-301. Trophon muricatus. 298. Middle Adriatic. 100 m. 3.9 mm. 299 and 300 same sample,
apices 870 /xm and 970 /xm. 301 . Specimen figured by Jeffreys ;n British Conchology. '9.5 mm.

Trophon fabricii (Beck in Moller, 1842) Figs 302-303, 320-321, 332

Tritonium craticulatum Fabricius, 1780:401 (not Linne, 1767).
Trophon fabricii Beck in Moller, 1842:87.

Mu rex borealis Reeve, 1845: sp. 145.

Trophon heuglini Morch. 1876:368.

Trophon interstriatus Sowerby, 1880:61, pi. 1, fig. 6.

Type material: T. fabricii, lectotype, here selected, in ZMC (Fig. 302); T. borealis, holotype in
BMNH.

Type locality: S Greenland.

Material examined : The type material of fabricii and borealis and about 100 specimens and shells from all parts
of the distribution area.

Distribution: NE America from Maine and northwards (Abbott, 1974), E and W Greenland
(Thorson, 1944, 1951), Iceland (Thorson, 1944), Spitzbergen (Odhner, 1915) and possibly N Norway
(Sars, 1878). It has been found in 6 — 160 m.

Remarks: T. fabricii is not a deep-water species, but has been included to complete the genus. It is
the most characteristic of the species of Trophon discussed here and should not cause any problems
with determination (cf. introduction to Trophon ).

136 P. BOUCHET & A. WAREN

Trophon droueti Dautzenberg, 1889 Figs 306-307, 340

Trophon droueti Dautzenberg, 1889:37, pi. 2, fig. 1.

Trophon richardi Dautzenberg & Fischer, 1896:438, pi. 18, fig. 6.

Trophon droueti var. elongata Locard, 1897:351.

Type material : T. droueti, holotype, now partly destroyed by an acid glass tube in MOM; T.
richardi, holotype in MOM.

Type locality: T. droueti, MONACO st 1 12, 38°34 N, 28°06 W, 1287 m; T. richardi, MONACO st
234, 39°02 N, 27°55 W, 454 m.

Material examined: The type material and MONACO st 198, 38°26 N, 28°39 W, 800 m, 1 sh; st 203, 39°27 N,
30°55 W, 1557 m, 1 sh; st 233, 38°33 N, 28°09 W, 1300 m, 18 shs; st 553, 37°43 N, 25°05 W, 1385 m, 3 shs; st
703, 39°21 N, 31 °06 W, 1360 m, 2 shs; st 1349, 38°35 N, 28°06 W, 1250 m, 15 shs; st 2450 (probably erroneous
st-number), 1 sh; TALISMAN dr 127, 38°38 N, 28°21 W, 1257 m, 4 shs; BIACORES st 46, 37°34 N, 28°54 W,
784 m, 1 spm; st 148, 37°34 N, 25°34 W, 870 m, 1 spm; st 157, 37°33 N, 25°43 W, 787-826 m, 1 spm; st 240,
37°35 N. 25° 32 W. 810 m. 1 spm; st 241. 37°37 N. 25° 32 W, 395-463 m. 1 sh.

Distribution: Endemic to the bathyal area of the Azores; known only from the material examined.

Remarks: As already noticed by Houart (1981) the name richardi ( = droueti) has recently been used
by Mediterranean authors for shelf populations of T. muricatus or T. banicensis. T. droueti can be
distinguished from these populations by its broader outlines and its aperture, furnished with
denticles on the inside of the outer lip. The development of spines from the axial lamellae is variable
in the two forms and cannot be used for separation.

There is, however, some possibility that T. droueti should be considered only an isolated
population or subspecies of barvicensis, which certainly is its closest relative. At present nothing is
known about the occurrence of barvicensis or droueti on the sea-mounts between the Azores and
Europe, which prohibits further speculation on the relationship.

Trophon dabneyi Dautzenberg, 1889 Figs 304-305, 319, 334

Trophon dabneyi Dautzenberg, 1889:36, pi. 2, fig. 7.

Trophon deeoratus Locard, 1897:340, pi. 17, fig. 5-11, and var. minor Locard, 1897:341; var. lanceolata
Locard, 1897:345.

Type material: T. dabneyi, holotype in MOM; T. deeoratus, syntypes in MNHN.

Type locality: T. dabneyi, MONACO st 1 12, 38°34 N, 28°06 W, 1287 m; T. deeoratus, TALISMAN
dr 127, 38°38 N, 28°21 W, 1257 m.

Material examined: The type material and MONACO st 184, 40° 05 N, 27°28 W, 1850 m, 5 shs; st 213, 39°23 N,
31 °25 W, 1384 m, 2 shs; st 224, 39°44 N, 31°06 W, 1213 m, 2 shs; st 233, 38°33 N, 28°09 W, 1300 m, 3 shs; st
624, 38°57 N, 28°19 W, 2102 m, 2 shs; st 684, 38°20 N, 28°05 W, 1550 m, 1 sh; st 698,39°11 N, 30°45 W, 1846
m, 2 shs; st 702. 39°21 N, 31 °06 W. 1360 m, 1 sh; st 719, 39° 1 1 N,30°24W. 1600 m. 6 shs; st 738, 37°40N,
26°26 W, 1919 m, 1 spm, 1 sh; st 1190, 15° 14 N, 23°04 W, 628 m, 1 sh; st 1193, 15° 17 N, 23°02 W, 1311 m, 1 sh;
st 1209, 16°34 N, 23°03 W, 1447 m, 1 sh; st 1331, 38°40 N, 26°01 W, 1805 m, 4 shs; st 1344, 38°45 N, 28°08 W,
1095 m. 2 shs; st 1349. 38°35 N. 28°06 W, 1250 m, 2 shs; st 3150. 38°01 N. 25°21 W, 1740 m. 1 sh;
NORATLANTE st B19, 47°28 N, 08°25 W, 2149 m, 1 spm; THALASSA st Z 453, 48°34 N, 10°51 W,
1975-2070 m, 2 spms; BIACORES st 36, 38° 1 1 N, 29°39 W, 2670 m, 1 spm; st 66, 38°34 N, 28° 19 W, 1225 m, 1
spm; st 95, 39°03 N. 28°25 W. 2440 m, 3 spms; st 179, 38°05 N, 25°46 W, 1590 m. 12 spms; st 191 , 37°56 N.
24°49 W. 1650-1750 m, 2 spms; st 217. 37°05 N, 24°52 W, 1735 m, 2 spms; st 227, 37°10 N, 25°19 W, 2160 m. 1
spm; st 252, 47°35 N, 08°47 W, 2550-2700 m, 3 spms; BIOGAS st CV6, 47°31 N, 08° 16 W, 2200 m, 3 spms; st
CV9, 47°31 N, 08° 44 W, 2199 m, 1 spm; st CV 10, 47°31 N, 08°41W, 2108 m, 2 spms; st CV 39, 47°33 N, 08°45
W, 2350 m, 2 spms; st CP 1, 47°35 N, 08°38 W, 2245 m, 4 spms; st CP 33, 47°43 N, 08°48 W, 2115 m, 1 spm; st
CP 34, 47°32 N, 08°23 W, 1970 m, 1 spm; st DS 15, 47°35 N, 08°40 W, 2246 m, 1 spm; st DS 17, 47°32 N, 08°45
W, 2103 m, 2 shs; st DS 65, 47°36 N, 08°40 W, 2360 m, 2 spms; st DS 71, 47°34 N, 08°34 W, 2194 m, 2 shs;
THOR, 49°27 N, 13°33 W, 2100 m, 2 shs, ZMC.

Distribution: The bathyal parts of the Eastern Atlantic, from the northern part of the Bay of Biscay
to the Cape Verde Islands and Azores.

Remarks: T. dabneyi can be recognized from its multispiral, brownish larval shell which indicates
planktotrophic larval development. It is the only species, in the area studied, with such larvae. The
protoconch, however, is frequently corroded in adult specimens and the species becomes difficult to
distinguish from T. echinatus (the cossmanni “form”). Distinctive characters then, are the more
blunt spire, less distinct spiral sculpture and less shouldered axial lamellae in T. dabneyi.

NORTH-EAST ATLANTIC NEOGASTROPODA 137

Trophon echinatus (Kiener, 1840) Figs 308-318, 333, 335-338

Murex vaginatus of authors on Recent Mollusca, not de Cristofori & Jan, 1832: 1 1 (fossil).

Murex carinatus of authors on Recent Mollusca, not Bivona, 1832:23, pi. 3, fig. 12 (fossil) (non Murex carinatus
Pennant, 1777, or Fischer de Waldheim, 1807 or Laskey, 1811).

Fusus echinatus Kiener, 1840:19, pi. 2, fig. 27.

Murex multilamellosus of authors on Recent Mollusca, not Philippi, 1844:182, pi. 27, fig. 8 (fossil).

Trophon carinatus Jeffreys, 1883:395, pi. 44, fig. 4 (non Murex carinatus Bivona, 1832 which is a Trophon).
Trophon grimaldii Dautzenberg & Fischer, 1896:439, pi. 18, figs 1-2.

Pagodula carinata var. tenuis and var. cinara Monterosato, 1884:116.

Trophonopsis carinatus var . major, var. elongata, var. depressa, var. spinosa and var. mutica Locard, 1897:346.
Trophonopsis varicosissimus var. major Locard, 1897:350.

Trophon cossmani Locard, 1897:342, pi. 17, figs 12-14.

Trophon deversus Locard, 1897:343, pi. 17, figs 15-17.

Trophonopsis carinatus aculeatus Settepassi, 1977:4, pi. 3, figs 24-28.

Trophonopsis carinatus multiaculeatus Settepassi, 1977:4, pi. 3, fig. 41 .

Trophonopsis carinatus hirtus St ttepassi, 1977:4, pi. 2, figs 17-18.

Type material: F. echinatus, not found in MNHN; T. carinatus, holotype in USNM; T. grimaldii,
holotype in MOM; T. cossmanni and T. deversus, holotypes in MNHN.

Type locality: F. echinatus, coasts of Sicily; T. carinatus, TRITON st 13, 59°51 N, 08° 18 W, 1050
m; T. cossmanni, TALISMAN dr 99, 17° 12 N, 17°07 W, 1617 m; T. deversus, TALISMAN dr 130,
37°55 N, 27° 02 W, 2235 m.

Material examined: The type material mentioned and about one hundred specimens and shells from the
Mediterranean (inch 6 shs and spms from the Eastern Basins), from depths between 100 and 650 m. From the
Atlantic we have seen about 80 specimens and shells from the Southern Bay of Biscay, Iberian Coasts and
Morocco, from 175 — 1000 m. From greater depths we have examined:

TRAVAILLEUR 1880 dr 10, 43°39 N, 03°28 W, 1960 m, 1 sh; 1881 dr 31 , 36°27 N, 08°03 W, 1383 m, 1 sh; 1882
dr 16, 42°48 N, 08°37 W, 627 m. 1 sh; dr 19, 41 °32 N, 08°21 W, 1350 m, 2 spms; dr 40, 33°09N, 08°38 W, 1900
m, 4 spm + shs; TALISMAN dr 99, 1 spm; dr 130, 2 spms; NORATLANTE st Bl, 55°08 N, 15° 11 W,
2215-2233 m, 2 spms; st B12, 36°21 N, 08°43 W, 2875 m, 1 spm; st B16, 45°35 N, 03°51 W, 3950-4150 m, 2
spms; B1ACORES st 202, 37°26 N, 25°00 W, 2900 m, 10 spms; st 206, 37°21 N, 25°28 W, 2090 m, 6 spms; st
235, 37° 18 N, 25°32 W, 2085-2115 m, 5 spms; st 252, 47°35 N, 08°47 W, 2550-2700 m, 1 sh; BIOGAS st DS86,
44° 05 N, 04° 19 W, 1950 m, 1 sh; INCAL st DS01, 58°OON, 10°40 W, 2091 m, 1 sh; st DS06, 56°26 N, 11 MOW,
2494 m, 1 sh; st CP01, 57°57 N, 10°55 W, 2068 m, 1 spm; st CP02, 57°58 N, 10°43 W, 2091 m, 1 spm; st CP03,
56°38 N, 1 1 °06 W, 2466 m, 2 shs; st CP04, 56°33 N, 1 1 0 1 1 W, 2483 m, 2 spms; st CP07, 55°03 N, 12°46 W,
2895 m, 1 spm; THALASSA st X336, 44° 11 N, 05° 10 W, 1850-2050 m, 1 sh; st X339, 44° 10 N, 04°30 W,
1520-1600 m, 1 spm, 2 shs; st X362, 44°06 N, 04°51 W, 585-600 m, 1 sh; st Y374, 41 °31 N,09°20W, 1250 m, 2
shs; st Y401 , 40°37 N, 09°21 W, 1040 m, 3 shs; st Z409, 47°43 N, 08°04 W, 1035-1080 m, 1 sh; st Z428, 48°27 N,
10°50 W, 850 m, 1 sh; st Z435, 48°40 N, 09°53 W, 1050 m, 2 shs; SARSIA st 7612, 43°44 N, 03°41 W, 1540-1570
m, 5 spms; st 7614, 43°43 N, 03°38 W, 1 100 m, 1 sh; st 7622, 43°44 N, 03°43 W, 1600 m, 1 sh; st 7627, 43°47 N,
03°46 W, 1925-1990 m, 9 spm + shs; METEOR st 36/16-17-99, 21 °37 N, 18°41 W, 2768-2971 m, 1 spm.

Distribution: The continental shelf of the Mediterranean and the E Atlantic north to the Bay of
Biscay (typical echinatus ), and the continental slopes of the same area and north to the Rockall
Trough (Houart, 1981 as T. cossmanni and Nodulotrophon scolopax ) ( cossmanni “form”). Also
present in the bathyal zone of the Azores {cossmanni “form”).

138 P. BOUCHET & A. WAREN

Remarks on the nomenclature

The present species is usually known as Trophon vaginatus (De Cristofori & Jan, 1832), as far as
specimens from the shelf are concerned. We have examined the lectotype in Museo Civico di Storia
Naturale, Milano. It was described as a Pliocene fossil from Tabiano, N. Italy. It is certainly quite
similar to the Recent Mediterranean form (Fig. 314 and Pinna, 1971, pi. 77, fig. 6), but there is a
distinct difference in the larval shell, which in the Recent form consists of about 1 .5 whorls (Fig. 316)
while the Pliocene form has more than 2.5 whorls (Fig. 315), and possibly had planktotrophic larval
development. This observation was confirmed by examining numerous specimens from other N
Italian Pliocene deposits (in the rich collection of Della Bella). We therefore consider the Recent
form specifically distinct from the Pliocene fossil.

The two subsequent names Murex carinatus Bivona, 1832 and Murex calcar Scacchi, 1836 which
have been used for the Recent Mediterranean form, are both based on N Italian Pliocene specimens
and also preoccupied, and thus not available.

Fusus echinatus Kiener, 1840 was based on Recent specimens and well figured. There are no
doubts about its identity. The case is, however, complicated by Sowerby’s (1818) use of the name
Murex echinatus. Sowerby’s species was transferred to Fusus by Fleming (1828) and Philippi (1836)
and is in fact a synonym of Trophon muricatus (Montagu, 1803). In the description, however,
Sowerby writes “Syn, Murex echinatus Brocchi, 2, 423, t. 8, f. 3”. Rossi-Ronchetti (1955)
considered M. echinatus Brocchi to be a species of Philbertia (Turridae) which probably is correct.
Therefore Sowerby’s use of the name is based on a misidentification and has no taxonomical status
and does not make Trophon echinatus preoccupied.

Several later names reflect the high variability of T. echinatus.

Remarks on the variation

Trophon echinatus exhibits considerable variation, especially in relation to depth. The
Mediterranean and the Atlantic shelf form do not differ very much. They have a porcellaneous,
smooth, very slender and spiny shell (Figs 312-313), without spiral sculpture. Sometimes the
Mediterranean form is a little more slender and spiny, but not enough to cause any problems with
the identification. At the uppermost part of the continental slope there is a tendency to more
numerous but less spiny axial lamellae. Typical specimens of this form have been called grimaldii
Dautzenberg & Fischer in the Atlantic and multilamellosa Philippi in the Mediterranean. Still
deeper, in the Atlantic in 600-1000 m, the spiral sculpture starts to become conspicuous and the
spines are still less developed (Figs 308-309). This form is known as cossmanni Locard and has been
found down to 3000 m.

Franchini & Frilli (1970) considered grimaldii a subspecies of “ carinata de Cristofori & Jan” ( =
T. echinatus) (and considered “ multilamellosa Philippi” a distinct species). To discuss this one has
to keep in mind that the typical T. echinatus has a continuous distribution along the shelf, via the
Straits of Gibraltar, and there exists one Atlantic and one Mediterranean deep-water outcrop of the
species. The bathymetric variation in shell form of these two outcrops is identical, except that the
cossmanni form is very scarce and perhaps extinct in the Mediterranean. Only a few old shells are
known. Therefore we prefer to regard the variation not as a result of genetic isolation (subspecies),
but as more directly influenced by the depth. It would be more than pure coincidence if the Atlantic
and Mediterranean deep populations had diverged from the shallow-water form because of genetic
isolation in exactly the same way in the two outcrops. Therefore we prefer to call them simply
different “forms”, viz. typical echinatus (Figs 312-313), the grimaldii “form” (Figs 310-311) and the
cossmanni “form” (Figs 308-309).

NORTH-EAST ATLANTIC NEOGASTROPODA 139

Comparison with other Atlantic species of Trophon

The sculpture of T. echinatus is variable and this does not facilitate identification. A few features
are, however, quite constant: paucispiral larval shell, long slender siphonal canal and strongly-
shouldered whorls. T. dabneyi has a larval shell of 2.5 brownish whorls while that of echinatus has
1.5 whorls and is colourless. It also has more rounded whorls and more solid, rib-like axial sculpture,
not thin lamellae as echinatus. T. droueti has a shorter siphonal canal and more pronounced spiral
sculpture, approximately as strong as the axial one. T. barvicensis has a shorter siphonal canal,
stronger spiral sculpture and a higher shoulder of the whorls.

The Caribbean T. aculeatus Watson, 1883 (= lacunella Dali, 1889) resembles T. echinatus,
especially specimens from intermediate depths, but is higher, more slender and has a deeper suture
than echinatus. We have not seen specimens of aculeatus with spiral sculpture.

T. abyssorum Verrill, 1885 (? = limicola Dali, 1902) resembles the cossmanni “form”, in the
shape of the shell and the sculpture. It is known from off the NE U.S. from 1500 to 3500 m. The
available material of that species, however, is scanty and there is a large gap in the distribution in the
N Atlantic, from Rockall to U.S., where no Trophon species related to these species are known.
Therefore we find it better to keep them separate.

More problems arise when going further south. Trophon guineensis Thiele was described from the
Gulf of Guinea (03° 10 N, 05°28 E, 2278 m). The larval shell is not known, but the postlarval shell
can hardly be distinguished from the cossmanni “form”. We have examined two specimens from off
Angola (WALDA st CY15, 10°30 S, 11°55 E, 1756 m). The shell is identical, but the larval shell is
destroyed. One of the specimens was a female and had a couple of egg clusters, 1100-1250
diameter and containing a few hundred eggs each, of a diameter of 120 fim. These egg clusters were
kept in the pallial cavity. The diameter of the eggs indicates planktonic larval development, because
the diameter of the eggs of species of Trophon with direct development is 230 — 500 \xm (5 species
known). This female also differed from T. echinatus in having unpigmented eyes, while T. echinatus
has black eyes.

Further to the south we have examined many specimens from the sides of the Walvis Ridge
(WALVIS st CP13, 32°18 S, 13°16 E, 3550 m and CP 14, 32°29 S, 13°25 E, 3675 m). They were all
heavily encrusted by zoantharians and the larval shells were badly corroded. They had normal eyes,
but no eggs were observed. They resemble T. acceptans Barnard, 1959 (see also 1963), described
from 33°34 S, 1160 — 3275 m. Barnard (1963) doubted the validity of this species and said “I fully
expect that acceptans will not be accepted when further material is obtained from the Atlantic
trough along the west coast of Africa”. It is still impossible to say anything about the validity of
these two names, but the differences in the development of the eyes indicate that they may not be
synonyms.

140 P. BOUCHET & A. WAREN

Figs 302-307. Genus Trophon. 302, T, fabricii, lectotype, 44 mm. 303, T. fabricii, Berufjord, Iceland,
40 mm. 304, T. dabneyi, BIOGAS st CV10, 37.2 mm. 305, T. dabneyi, larval shell, 850 /mi. 306, T.
droueti, BIACORES st 157, 14.0 mm. 307, T. droueti, holotype of T. richardi, 15.0 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 141

Figs 308-313. Trophon echinatus . 308, INCAL st CP4 (56°33 N, 1 1 ° 1 1 W, 2483 m), 20.0 mm. 309,
syntype of T. cossmanni , 26.0 mm. 310, WASHINGTON expedition, 370-420 m, 13.1 mm. 311,
NORATLANTE st B1 (55°08 N, 15° 1 1 W, 2215 m), 19.1 mm. 312, Gulf of Napoli, 19.0 mm. 313,
TALISMAN dr 20 (33°43 N, 09°02 W, 1 105 m), 15.8 mm.

142 P. BOUCHET & A. WAREN

Figs 314-318. Genus Trophon. 314, T. vaginatus, upper Pliocene of Carignano, Imola, Bologna, 19.5
mm. 315, apex of a specimen from the same locality, 1.13 mm. 316, T. echinatus, apex of a Recent
Mediterranean specimen, 1.07 mm. 317, T. sp., WALVIS st CP14, 33.3 mm. 318, T. sp., WALDA st
CY15, 25.5 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 143

Figs 319-327. Trophon opercula. 319, T. dabneyi, Bay of Biscay, 10.9 mm. 320 and 321, T. fabricii,
Berufjord, Iceland, 2.5 and 9.5 mm. 322, 323 and 324, T. clavatus, Koster area, Swedish west coast,
0.8 mm, 2.8 mm and 3.4 mm. 325 and 326, T. clathratus, Godhavn, Greenland, 1.4 mm and 4.6 mm.
327, T. geversianus, Magellan Strait, 11.2 mm.

144 P. BOUCHET & A. WAREN

Figs 328-332. Genera Tromina and Trophon. 328, Tromina unicarinata, type species of Tromina Dali,
1918, Magellan Strait, 35 m, 7.4 mm. 329, Trophon geversianus, breadth of central tooth 167 jum. 330,
Tromina unicarinata, breadth of central tooth 27 /jm. 331, Trophon clavatus, 41 f^m. 332, Trophon
fabricii, 110 /urn.

NORTH-EAST ATLANTIC NEOGASTROPODA 145

341

Figs 333-342. Radulae of Trophon. 333, T. echinatus , INCAL st CP2, breadth of central tooth 56 pro.
334, T. dabneyi, 94 pm. 335, T. echinatus, BIACORES st 202, 68 pm. 336, T. echinatus, spiny form, N.
Spain 500-540 m, 51 pm. 337, T. sp., WALVIS stCP14, 60 pm. 338, T. sp.. WALDA siCYIS, 77 pm.
339, T. clavatus, 40 pm. 340, T. droueti, 47 pm. 341, T. barvicensis, 33 pm. 342, T. muricatus, 36 pm.

146 P. BOUCHET & A. WAREN

Figs 343- 347 Murick! radulae. 343, Pterynotus ieucas, breadth of central tooth 56 .urn, 344, Orania grayi,
98 (jm. 345, Muricopsis blainvillei, 41 fum. 346, Ocinebrina aciculata 43 jk m. 347, Orania fusulus , 50 pm.

NORTH-EAST ATLANTIC NEOGASTROPODA 147

Genus PTERYNOTUS Swainson, 1833

Type species: Murex pinnatus Swainson, 1822, by original designation (pi. 122).

Remarks: The genus Pterynotus and its subgenera have been discussed by Ponder (1972) and the W
Atlantic species were monographed by Harasewych & Jensen (1979). There are two species of the
genus in the Central E and NE Atlantic, P. leucas Locard and P. atiantideus which is described here.
They can easily be separated by the presence of intervarical nodes or bulges in P. atiantideus, easily
seen in an apical view (cf. Figs 350, 353). These bulges are absent in P. leucas .

Figs 348-350

Pterynotus atiantideus n.sp.

Type material: Hoiotype in MNHN.

Type locality : BIACORES st 186, 37°51 N, 25°40 W, 370-455 m.

Material examined: The type material and BIACORES st 38, 37°59 N, 29° 16 W, 350-400 m, 1 spm; st 146, 37° 39

1 sh; st 236, 37°21 N, 25°45 W, 470-500 m, 1 spm.

Distribution: In addition to the material from the Azores enumerated above, P. atiantideus is known
from the Great Meteor Bank, in 295-343 m (Fechter, 1977 as P. leucas ).

Description: Shell solid, fusiform, consisting of six postlarval whorls, larval shell white, consisting
of one smooth whorl. The first postlarval whorl has 7 fragile foliaceous varices that adhere to the
protoconch; there is no intervarical node between these early varices. The regular adult sculpture
starts on the second whorl and there are 3 varices and 3 intervarical nodes per whorl. The spiral
sculpture consists of spiral cords, of which a few stronger ones extend on to the varices where they
tend to produce spines; there are also extremely fine spiral threads which, together with the fine axial
sculpture present on some specimens (including the hoiotype), may give a reticulate appearance
under high magnification. Aperture oval, outer lip with 7 denticles; siphonal canal of moderate
length for the genus, open; colour light tan with a few brownish blotches on the varices or the nodes;
aperture white.

Dimensions : Height of the shell 21 mm, breadth 11 mm.

Remarks : P. atiantideus differs from P, leucas in its smaller larval shell, by having intervarical nodes
and by its coloured shell. In P. leucas » there are several postlarval whorls with numerous varices
(Fechter, 1977: fig. 11), while there are only 1 to 1.5 such whorls in P. atiantideus. P. leucas also
reaches a larger size.

Pterynotus leucas (P. Fischer in Locard, 1897)

Figs 343, 351-353

Murex leucas P. Fischer in Locard, 1897:306, pi. 15, figs 10-13.

Type material: Lectotype (selected here) in MNHN.

Type locality: TALISMAN st 71, 25°39 N, 15°58 W, 640 m.

Material examined: The type material and 6 additional spms + shs from the type locality (paralectotypes).

Remarks: We have included P. leucas for comparison with atiantideus although it lives outside the
area covered by this series. It has not been found since the original description. The record in Fechter
(1977) is based on P. atiantideus.

148 P. BOUCHET & A. WAREN

Figs 348-353. Genus Pterynotus. 348 and 349, P. atlantideus, BIACORES st 186, 21.0 mm. 350, P.
atlantideus, BIACORES st 41, 27.0 mm. 351, 352 and 353, P. leucas, lectotype, 40.5 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 149

Figs 354-356. Genus Orania. 354, O. fusulus, SE Bay of Biscay, 160 m, 23.0 mm. 355, O. fusulus, apex
1.0 mm. 356, O. grayi, MONACO st 1242, 18.7 mm.

Genus ORANIA Pallary, 1900

Type species: Murex spadae Libassi, 1859, by original designation.

Synonym: Nemofusus Cossmann, 1903. Type species: Murex fusulus Brocchi, 1814, by original
designation.

Remarks: We have kept the generic name Orania for Murex fusulus Brocchi. This species has usually
been referred to Pseudomurex or Urosalpinx. It also resembles Ocinebrina and Muricopsis.
Pseudomurex belongs to Coralliophilidae and has no radula, while fusulus has one. Urosalpinx
differs considerably in radular characters and shell characters. Ocinebrina and Muricopsis have quite
similar shells, compared with fusulus , but have a very distinct radula (Figs 345-346). (It should here
be pointed out that the radula of Ocinebrina aciculata Lamarck (type species of Ocinebrina) was
figured by Radwin & D‘Attilio 1976, but their figure differs considerably from what we have seen.)
We know of no other genus in common use where fusulus fits, and have therefore kept Pallary’s
genus, which was not included by Radwin & D’Attilio (1976). Lack of any well-fitting genus has also
induced us to place grayi here, although it also resembles Cytharomorula Kuroda, 1953.

Pallary (1900) designated Murex spadae Libassi as type species, when describing the new genus
Orania. There exists some doubt about the identity of spadae which was described as a Pliocene
fossil (Kobelt 1887), but evidently Pallary introduced the new generic name for the Recent form,
which should be considered the type species if more than one species proves to be involved.

150 P. BOUCHET & A. WAREN

Orania fusulus (Brocchi, 1814)

Figs 347, 354-355

Murex fusulus Brocchi, 1814:409, pi. 8, fig. 9.

Murex spadae Libassi, 1859:43, pi. 1, fig. 29.

Murex spadae var. tiberianus Brusina, 1871, fide Settepassi, 1977.

Murex pyrrhias Watson, 1883:603.

Rojlia fusulus var. major Locard, 1897:325, pi. 16, figs 14-16.

Pseudomurex spadae var. obesa & var. minor Pallary, 1900:287.

Pseudomurex spadae robustus Settepassi, 1977:3, pi. 1, fig. 4.

Pseudomurex spadae elongatus Pallary in Settepassi, 1977:2, pi. 1, fig. 6.

Type material : M. fusulus, holotype in Museo Civico di Storia Naturale, Milano, figured by Pinna
(1978, pi. 33, fig. 2); M pyrrhias, BMNH.

Type locality: M. fusulus, Pliocene of Val d’Ancona, N. Italy; M. pyrrhias, CHALLENGER st. 75,
38°38 N, 28°28 W, 830-920 m.

Material examined: The types listed and about 75 specimens and shells from the Bay of Biscay (northernmost
record 45°43 N, 02° 14 W, 160 m, 2 spms) to the Cape Verde Peninsula (95 — - 200 m) and Tristao (Guinea) in the
south (MNHN), the Azores and the western Mediterranean, where it seems to be rather common off the N
African coast.

Distribution: The western parts of the Mediterranean, from Sicily (Monterosato, 1890), Corsica
(Locard & Caziot, 1901), Oran, Algeria (Pallary, 1900), westwards to the Atlantic, where the
extremes are given in “Material examined”. It is known from 35-440 m.

Remarks: Murex fusulus has often been placed in Pseudomurex (Coralliophilidae), but it has a
radula so it fits better in the Muricidae. This was already noticed by Kobelt (1887), but has not been
mentioned by most later authors.

Orania fusulus has a smooth, multispiral larval shell, which indicates planktotrophic larval
development.

When this paper was in press, the work by Sabelli & Tommasini (1983) was published, with
anatomical datas.

Orania grayi (Dali, 1889) Figs 344, 356

Nassarina grayi Dali, 1889:183, pi. 32, fig. 12a.

Trophon lowei Watson, 1897:244, pi. 19, fig. 12.

Cantharus (Tritonidea) laevis Smith, 1890:261, pi. 21, fig. 11.

Type material: N. grayi , holotype USNM: C. laevis, 5 syntypes BMNH 89.10.1.2362-7.
Type locality: N. grayi, off St Kitts, W Indies: T. lowei, Madeira; C. laevis, St Helena.

Material examined: The types of N. grayi and C. laevis; J. CHARCOT MADERE st 49, 32°27 N, 16°32 W,
450-500 m, 1 sh; MONACO st 1242, Seine Bank, 240 m, 1 spm, 1 sh; TRAVAILLEUR 1882, dr 52, 32°30 N,

1 6 ° 3 1 W, 100 m, 4 shs; no st number, 6 spms + shs; TALISMAN 1883, dr 53, 28°35 N, 13°19 W, 905 m, 2
shs; La Palma, Dacia Bank, Tenerife (Canaries), from fishermen, MNHN and private collections.

Distribution: Known from the Caribbean area, 100 — 200 m (Abbott, 1974) and some sea mounts
and islands in the E Atlantic. Not known from European or African coasts.

Remarks: O. grayi was transferred to Muricidae by Radwin & D’Attilio (1976) who placed it in
Evokesia Radwin & D’Attilio, 1972. We have, however, preferred to place it in the same genus as O.
fusulus.

The larval shell is multispiral and indicates a planktotrophic larval development, which possibly
contributes to the scattered distribution.

NORTH-EAST ATLANTIC NEOGASTROPODA 151

Family CORALLIOPHILIDAE

The taxonomy of the family Coralliophilidae is at present in a state of confusion, presumably
because attention has been focused on the many extreme-looking species that represent the most far-
reaching sprouts of the evolutionary tree, and give the least information about relations within the
family. We have therefore not made any attempts to divide the species treated here into different
genera, but use the generic name Coralliophila for all. We shall, however, give some nomenclatorial
remarks on the generic names that have been in use for the NE Atlantic species.

Pseudomurex Monterosato, 1872 is the most often used name. D’Attilio (1978) and other authors
have considered Murex bracteatus Brocchi, 1814, to be the type species, but Monterosato only
included two species in his genus, viz. Murex lamellosus Jan (sic!) and Murex spadae Libassi. Later
he restricted the genus to include only M. lamellosus and transferred M. spadae to Pollia. M. spadae
is a muricid and is discussed on p. 150. Therefore Dautzenberg (1927) was correct when he designated
Fusus lamellosus Philippi as type species of Pseudomurex.

Aradomurex Coen, 1943. Type species, Murex so fiae Aradas & Benoit, 1876.

Babelomurex Coen, 1922. Type species, Fusus babelis Requien, 1849.

Fusomurex Coen, 1922. Type species, Murex alucoides de Blainville, 1829 (not Olivi, 1792). A
synonym of Pseudomurex.

Hirtomurex Coen, 1922. Type species, Fusus lamellosus Philippi, 1836.

Latimurex Coen, 1922. Type species, Murex meyendorffi Calcara, 1845.

Lepadomurex Coen, 1922. Type species, Purpura brevis de Blainville, 1832.

The species of Coralliophilidae live as parasites or predators on anthozoans. Coralliophila brevis
has been found on at least three species of gorgonian ( Paramuricea chamaeleon, Eunicella stricta
and Lophogorgia sarmentosa) and C. meyendorffi has been found on Anemonia sulcata and
Cladocora cespitosa (G. Spada, pers. comm.), so evidently they are not very host specific.

The species are quite rare in deep water, below the continental shelf, which makes a treatment of
these species more difficult. The Mediterranean shallow-water species were discussed by Sabelli &
Spada (1980) in a very useful paper, and by Settepassi (1977) who lists 14 names of specific rank and
a multitude of lower ranking names.

We have examined the larval shells of all species discussed here except C. sentix. They proved to
have a multispiral larval shell, of very uniform shape and sculpture (Figs 370-371, also Richter &
Thorson 1975, figs 70-71). Because of the uniformity and also because it is normally corroded, even
in young specimens, the larval shell is of almost no use for taxonomical purposes.

Pleurotomoides obliquispira Nordsieck, 1977, is based on a juvenile of a species of
Coralliophilidae, but we are not able to say which species.

152 P. BOUCHET & A. WAREN

Genus CORA LLIOPHILA H. & A. Adams, 1853
Type species: Fusus neritoideus Lamarck, 1816, by subsequent designation (Iredale, 1912).

Corailiophiia richardi (P. Fischer, 1882) Fig. 368

Mur ex richardi P. Fischer, 1882:49.

Corailiophiia lactuca Dali, 1889:220, ph 16, fig. 6.

Pseudomurex richardi P. Fischer, Locard 1897:316, pi. 16, figs 3-8.

Type material: M. richardi, holotype in MNHN; C. lactuca , syntypes in USNM.

Type locality: M. richardi, TRAVAILLEUR 1881 dr 42, 44°01 N, Q7°05 W, 896 m; C. lactuca,
USFC st 2669, off Fernandina, Florida, 635 m.

Material examined: The type material and TRAVAILLEUR 1881 dr 39,44°05 N, 07°06 W, 953-1226 m, 1 spm;
off N. Spain, 496 m, 1 spm; TALISMAN dr 10,35°26 N, 06°49 W, 717 m, 1 spm; SARSIA st 7614, 43°43 N,
03°38 W, 1 100 m, 1 sh; MONACO st 553, 37°43 N, 25°05 W, 1385 m, 1 sh; st 584, 38°30 N, 26°50 W, 845 m, 1
spm; 43°58 N, 02°06 W, 320 m, Lagardere coll., 1 sh; W of Tampa, Florida, 400 m, 1 sh; EOLIS st 128, off
Fowey Light, Florida, 1 10 m, 1 spm; st 166, off Ragged Key, 120 m, 1 spm, USNM; and the material reported
by Taviani & Colantoni (1979) and Sabelli & Spada (1980).

Distribution: Both sides of the subtropical North Atlantic, in the upper bathyal zone. The species
has been reported in the Mediterranean by Monterosato (1890:23), Taviani & Colantoni (1979:141),
and Sabelli & Spada (1980:2) from empty shells. The species probably lives associated with Cnidaria
of the Lophelia- reefs and is possibly now extinct in the Mediterranean.

Remarks: C. richardi is the most characteristic of the NE Atlantic species of Corailiophiia. The axial
sculpture consists of lamellose varices instead of solid ribs as in the other species. These lamellae are
given a slightly fringed appearance by the spiral sculpture. It lacks the fine, squamulose sculpture
characteristic of many species of the family.

Corailiophiia panormitana (Monterosato, 1 869)

Fig. 369

Pyrula panormitana Monterosato, 1869: 17, fig. 9.

Type material: Not seen.

Type locality: Palermo, Sicily.

Material examined: About 20 spms from the Western Mediterranean and TALISMAN dr 63-64, 26° 17 N,
14°52 W, 355-640 m, 1 spm.

Remarks: We have seen only one specimen from the Atlantic, but that one is very large in
comparison with W Mediterranean specimens. It closely resembles, however, an unusually large
specimen from the continental shelf off Mdiq (S Alboran.Sea, coll.S.Gofas).

C. panormitana can be distinguished from C squamosa and C. basileus by having a more inflated
body whorl.

NORTH-EAST ATLANTIC NEOGASTROPODA 153

Coralliophila squamosa (Bivona, 1838) Figs 357-361

Murex alucoides de Blainville, 1830:128, pi. 5B, fig. 1 (not Murex alucoides Olivi, 1792: 153).

Fusus lamellosus de Cristofori & Jan in Philippi, 1836:204, pi. 11, fig. 30 (not Fusus iamellosus Borson,
1821:317).

Fusus squamulosus Philippi, 1836:204, pi. 11, fig. 31 (not Fusus squamulosus Deshayes, 1835:540).

Fusus squamosus Bivona, 1838:14, fig. 22.

Pseudomurex perfectus P. Fischer, 1882:274 (nomen dubium).

Pseudomurex ruderatus Monterosato in Sturany, 1896:25, pi. 2, figs 42-43.

Pseudomurex monterosatoi Locard, 1897:315, pi. 15, figs 21-23.

In addition, the names (Coralliophila) inflata, elongata, turns, longicauda, depressa, curta, squamulata ,
modesta and ( Ocinebrina ) pianosana have been introduced by Settepassi (1977).

Type material: F. squamosus, neotype in MNHN; P. ruderatus, syntypes in NHMW; P.
monterosatoi, syntypes in MNHN.

Type locality: F. squamosus , Palermo and Messina, Sicily; P. ruderatus, Corsica; P. perfectus,
TRAVAILLEUR 1882, Atlantic (probably N. Spain), 400 m; P. monterosatoi, TRAVAILLEUR
1882, dr 1 & 2, 44° 05— 44° 07 N, 05°35 W, 564-608 m.

Material examined: the type material listed; c. 250 spms and shs from shallow water (60-200m) in the
Mediterranean; and TALISMAN dr 63-64, 26° 18 N, 14°52 W, 640 m, 6 spms; MONACO st 503, 47° 10 N, 05°48
W, 748-1262 m, 2 spms; st 684, 38°20 N, 28°05 W, 1550 m, 1 spm; st 2720, 36°42 N, 08°40 W, 310-749 m, 1 sh;
THALASSA st W399, 43°48 N, 05° 11 W, 165-210 m, 1 sh; st W424, 44° 02 N, 07°09 W, 300-700 m, 1 sh; st
Till , 41 °32 N, 09° 14 W, 320 m, 1 sh; st Y383, 41°26 N, 09°12 W, 420 m, 2 shs; 45°43 N, 02°14 W, 78 m, 2 shs,
Lagardere coll.; METEOR st 36-98, 25°31 N, 16°02 W, 700-900 m, 2 spms.

Distribution: From the bay of Biscay, along the Iberian peninsula, 'the Mediterranean, NW Africa to
the Canaries and the Azores. It occurs mainly on the continental shelf and in coastal water, but has
occasionally been found on the upper part of the continental slope. Possibly also in the Caribbean
area (Bayer 1971).

Remarks: The present species has usually been known under the names alucoides de Blainville, 1830
and lamellosus de Cristofori & Jan in Philippi, 1836. Both these names are, however, preoccupied,
as is also the third name suggested by Philippi (1836). The name Fusus squamosus Bivona, 1838 is
not preoccupied, but the description and figure are not very good. To avoid this problem we have
selected a neotype from Corsica that agrees fairly well with Bivona’s figure. Another name that has
been in use for the present species is bracteatus Brocchi, 1814. The name was based on a Pliocene
fossil. The type has been figured by Rossi-Ronchetti (1955), Pinna & Spezia (1978) and we have
personally examined it in Museo Civico di Storia Naturale, Milano; we find it more like the Recent
W African C. angolensis Knudsen. We think that Knudsen’s species is the Recent descendant of C.
bracteata, as is the case for many other Italian Pliocene species, the descendants of which now live in
W Africa.

The names introduced by Settepassi (1977) need no comments.

C. squamosa occurs in coastal waters and on the shelf in the Mediterranean (but seems to be
absent from the continental slopes). Such specimens are characterized by having a strongly
squamose sculpture, well-defined siphonal canal and a rather high spire. Atlantic specimens have a
broader and more open siphonal canal and reach a larger size (50 mm instead of 40 mm maximum
size). In these respects the Atlantic form approaches fossil shells from old coral associations in the
Mediterranean, reported by Taviani & Colantoni (1979).

For a discussion of basileus, see that species, and for separation from the Mediterranean shallow-
water species, see Sabelli & Spada (1980). Bayer (1971) reported C. lamellosa ( = squamosa) from
the Caribbean area and synonymized it with Trophon lintoni Verrill & Smith, 1882. We have
examined this and additional material in USNM. We agree that there are specimens that closely
resemble squamosa, although the specimen figured by Bayer (fig. 51) more closely resembles
basileus. Other facts also, like the occurrence at the Azores and the presence of planktotrophic
larvae, suggest an amphiatlantic distribution, but the chaotic state of the taxonomy of the Caribbean
Coraliiophilidae makes it impossible to coordinate the names of the species without a complete
revision of the W Atlantic species.

154 P. BOUCHET & A. WAREN

Coralliophila basileus (Dautzenberg & H. Fischer, 1896) Figs 362-366

Pseudomurex basileus Dautzenberg & H. Fischer, 1896: 440, pi. 18, fig. 3.

Type material: Holotype in MOM.

Type locality: MONACO st 233, 38°33 N, 28°08 W, 1300 m.

Material examined: The type material and MONACO st 105, 38°23 N, 28°31 W, 927 m, 1 spm; st 247, 38°24 N,
28°01 W, 318 m, 1 spm; st 680, 38° 19 N, 28°08 W, 1550 m, 1 sh; TALISMAN dr 53, 28°35 N, 13°09W,905 m, 1
sh; dr 63-64, 26° 18 N, 14°52 W, 640 m, 3 spms + shs; dr 122, 37°35 N, 29°26 W, 1440 m, 1 spm; BIACORES st
4, 38° 1 1 N, 28°38 W, 1080-1200 m, 1 spm; st 148, 37°34 N, 25°34 W, 847-870 m, 5 spms; st 151, 37°37 N, 25°39
W, 780-788 m, 1 spm; st 180, 37°57 N, 25°33 W, 1069-1235 m, 2 spms; st 230, 36°54 N, 25°09 W, 665-742
m, 1 spm.

Distribution: The upper part of the continental slope around the Azores and the Canaries.

Remarks: We are not convinced that basileus is truly distinct from squamosus, but as we have no
definite proof we have preferred to keep them distinct. There is a difference in shape, basileus having
a broader siphonal canal that is less set off from the aperture, and in size, basileus barely exceeding
half the size of squamosus. There is also a difference in distribution. C. basileus occurs only around
the Azores and the Canaries, while it is not known from the E Atlantic coasts. C. squamosus also
occurs around these islands, but its main distribution is Mediterranean and along the S European
and N African coasts. The two forms thus occur sympatrically in the Atlantic islands, but we have
not seen specimens of both forms in a single dredge haul (except a TALISMAN sample, from several
stations), which indicates some degree of separation. The two forms are very variable and we have
not examined very large numbers, so we prefer to leave the question of conspecificity unsettled.

A second possibility, although we find it less probable, is that C. basileus is a local form of C.
brevis (de Blainville, 1826). That species, however, has a thinner shell and the aperture, at least in
adult specimens, is deformed by its sessile life on gorgonians, which we never have seen in basileus.

Coralliophila sentix (Bayer, 1971)

Fig. 367

Coralliophila sentix Bayer, 1971: 189, fig. 49.

Type material: Holotype USNM 701155.

Type locality: PILLSBURY st P876, 13° 14 N, 61 °05 W, 231-258 m (Lesser Antilles).

Material examined: The type material and MONACO st 1 12, 38°34 N, 28°06 W, 1287 m, 1 sh; BIACORES st
41, 37°43N,29°04W, 450-475 m, 1 spm; st 236, 37°21 N, 25 °45 W, 470-500 m, 1 spm; Alboran Sea? (taken
by fishermen), 2 shs, coll. G. Spada.

Distribution: The upper bathyal of the subtropical North Atlantic.

Remarks: Specimens from the W Atlantic have longer and more developed shoulder projections that
join each other more extensively than those from the E Atlantic. We have, however, considered
them conspecific. C. fax Bayer, 1971, from off Florida, has the same type of non-squamose
sculpture as C. sentix, but has a broader shape, lower spire and a smoother shell. In the Atlantic only
C. juliae Clench & Aguayo, 1939 shares the rather high spire and non-squamose sculpture, but it has
more or less open coiling. No other Atlantic coralliophilid can be confused with them.

NORTH-EAST ATLANTIC NEOGASTROPODA 155

Figs 357-361 . Coralliophila squamosa. 357, neotype, Calvi, 30.0 mm. 358, MONACO st 503, 49.3 mm.
359, MONACO st 54, 27.5 mm. 360, THALASSA st Y383, 40 mm. 361, syntype of Pseudomurex
rude rat us, 23 mm.

156 P. BOUCHET & A. WAREN

Figs 362-366. Coralliophila basileus. 362, BIACORES st 4, 27.3 mm. 363, B1ACORES st 180, 24 mm.
364, BIACORES st 148, 18.9 mm. 365, TALISMAN dr 53, 21.3 mm. 366, MONACO st 105, 18.7 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 157

Figs 367-371. Genus Coralliophila. 367, C. sentix, BIACORES st 236, 26.2 mm. 368, C. richardi ,
TRAVAILLEUR off NW Spain, 16.5 mm. 369, C. panormitana , TALISMAN dr 63-64, 24.7 mm. 370
and 37 1 , Coralliophila larvae from plankton of Curacao, 1 .0 mm.

158 P. BOUCHET & A. WAREN

Family NASSARIIDAE

Genus NASSARIUS Dumeril, 1806

Type species: Buccinum arcularia Linne, 1758.

Remarks: Only two nassariid species can be considered to belong to the deep sea fauna, and they
occur in the uppermost bathyal zone and on the shelf. A few additional species are known from the
NW African shelf and were treated by Adam & Glibert (1974, 1976).

Nassarius wolffi (Knudsen, 1956) Fig. 373

Nassa wolffi Knudsen, 1956:58, pi. 1, fig. 7.

Nassa sadurnii Bot, 1972:15.

Hinia frigens malacitanae Ghisotti & Spada in Maldonado, 1973:233, pi. 3.

Type material: N. wolffi, holotype in ZMC.

Figs 372-373. Genus Nassarius. 372, N. frigens, TALISMAN dr 82, 16.9 mm. 373, N. wolffi. Dakar,
200-300 m, 34.6 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 159

Type locality: N. wolffi, ATLANTIDE st 120, 02°09 N, 09°27 E, 260-650 m; N. sadurnii , off
Huelva, Spain, 1500-2000 m; H. frigens malacitanae , triangle between Huelva, Casablanca and
Gibraltar, 1400 m and the Alboran Sea, 500 m.

Material examined: The type material mentioned and Cabo Frio, SW Africa, 1 sh; WALDA st CM01, 20° 24 S,
1 2°3 1 E, 256 m, 8 spms; 05° 00 S, 1 1 °2 1 E, 296-308 m, 14 spms, 4 shs, Crosnier coll.; off Abidjan and Grand
Bassam, Ivory Coast, 13 shs; W. of Jacqueville, Q5°30 W, 200-400 m, 5 spms, Le Loeuff coll.; off Dakar,

200-300 m, 5 spms, Marche-Marchad coll.; PILLSBURY 1966, st 254, 03° 50 N, 07°08 E, 148-174 m, 7 spms; st
255, 03° 49 W, 07°38 E, 269-264 m, 9 spms. See also “Remarks”.

Distribution : The upper bathyal of West Africa, from Namibia to Senegal at least; doubtful records
further to the north (see below).

Remarks : Although the type material of N. sadurnii has never been figured and we have not
examined it, there is no doubt that the name is a synonym of wolffi . In a letter of November 1975 Mr
J. Bot informed us that his species is very close to Nassa wolffi Knudsen. He said he had seen
material from Angola with all the characters of sadurnii. The same species was again described by
Maldonado (1973) from specimens taken by shrimp trawlers. In the same paper, Ghisotti & Spada
argued that the description of Nassa sadurnii Bot did not meet the requirements of ICZN because it
was not accompanied by an illustration. At the same time (p. 234) they “provisionally name”
Maldonado’s material Hinia frigens malacitanae n.ssp. “until further material can be studied for a
definitive identification”. In fact Bot’s description is perfectly valid, while Ghisotti & Spada do not
fulfil the requirements of ICZN §15 which states that “after 1960, a new name proposed
conditionally (...) is not available”.

N. wolffi has been recorded a few times from the W Mediterranean, but all records are based on
specimens with unreliable locality information, taken by fishing boats and said to come from the
Alboran Sea and the fishing grounds off Catalonia. It is possible that the species occurs in the W
Mediterranean, but the records need confirmation.

N. wolffi has a smooth larval shell of 2.5 whorls, which indicates planktotrophic larval
development.

Nassarius frigens (von Martens, 1878) Figs 372, 387

Nassa frigens von Martens, 1878:134; 1881 : 14, pi. 22, figs 17-18.

Nassa brychia Watson, 1882:365; 1886:189, pi. 11, fig. 15.

Nassa brychia vars. decorata, major and minor Locard, 1 897:274.

Type material: N. frigens , holotype in the ZMHU; N. brychia , holotype in BMNH (figured by
Cernohorsky 1975:157, fig. 77).

Type locality: N. frigens , GAZELLE 10° 13 N, 17°25 W, 650 m; N. brychia , CHALLENGER st 8,
28°03 N, 17°27 W, 1135 m.

Material examined: The type of N. brychia and TALISMAN dr 82, 23°00 N, 17° 30 W, 932 m, 2 shs; dr 83
and 85 (mixed samples), 22°52-22°57 N, 17°23-17°30 W, 830 m, 9 spms + shs; dr 87, 22°03 N, 17° 23 W,
1013-1113 m, 9 spms + shs; dr?, 820-860 m, 32 spms + shs; off Dakar, 150-250 m, 4 shs, Marche-Marchad
coll.; off Abidjan, 1 sh, Le Loeuff coll.; PILLSBURY 1966 st P44, 05°05 N, 04°00 W, 586-403 m 15
spms + shs; ATLANTIDE st 135, 07°55 S, 12°38 E, 235-460 m, 1 spm.

Distribution: Beside the material cited above, the species is known from Angola, in 225-750 m
(Adam & Knudsen 1955:10).

Remarks: All the shells have corroded apices, so the type of larval development is not known. We
have not examined the type of N. frigens , but our material fits with the specimens figured and
described by Adam & Knudsen (1955) who had examined the type.

N. frigens bears some resemblance to Profundinassa Thiele, 1929 (type species Nassa babylonica
Watson, from the bathyal zone of the Philippines). Typical species of Profundinassa are present in
the Pliocene of Sicily (Moroni & Torre 1966:32, pi. 1, fig. 7).

160 P. BOUCHET & A. WAREN

Family FASCIOLARIIDAE

We follow Ponder (1973) and include the “fusinid” genera in Fasciolariidae. Several genera have
been proposed for the shallow water species of S Europe:

Aptyxis Troschel, 1868. Type species Fusus syracusanus Linne.

Gracilipurpura Jousseaume, 1880. Type species Fusus strigosus Lamarck (= rostratus Olivi, 1792,
not Solander in Brander, 1766).

Pseudofusus Monterosato, 1884. Type species Murex rostratus Olivi, 1792, here designated.
Sinistralia H. & A. Adams, 1853. Type species Murex maroccensis Gmelin.

We figure the radulae for the type species of these genera (figs 380, 381, 385). Of these it is only
Aptyxis that shows any difference in the central tooth, which has longer and#more slender cusps.
This genus also differs in its shell characters and could probably be kept as a subgenus. We consider
the other genera synonyms of Fusinus Rafinesque, 1815.

The family has very few species in deep water in the NE Atlantic. We recognize three species, but
the names below need some comments.

Clathurella rugosissima Locard, 1897. The holotype in MNHN is an extremely worn old shell,
probably a Lathyrus, but too poor to be identified. It is probably a shallow-water species.

Pseudofusus gigliolii Monterosato, 1890. The species was described from specimens taken by the
WASHINGTON expedition, without any information about the locality, but the expedition dredged
around Sardinia and Sicily. Nothing is mentioned about this name by D’Amico (1912), who
prepared a report on the Mollusca. The type material could not be found in ZMR, although it was
figured by Franchini & Zanca (1977). We have however examined specimens taken by the
MONACO expeditions and the geological cruises of BANNOCK (in Istituto di Geologia, Bologna),
that correspond with Monterosato’ s description and the figures of Franchini & Zanca. From this we
conclude that gigliolii was based on the same white, slightly-keeled, deep-water form of rostratus as
bengasiensis.

Fusus bengasiensis Sturany, 1896. We have examined the holotype in Naturhistorisches Museum,
Wien. It is an old, broken specimen of a form of Fusinus rostratus , known as var. carinata
Monterosato in Bucquoy, Dautzenberg & Dollfus, 1882.

Pseudofusus locardi Pallary, 1904, is based on Locard ’s figures (1897, pi. 16, figs 26-30) of
‘Fusus rusticulus Monterosato”. The specimen came from 530 m, off Morocco. Examination of
the specimen in question showed that it is a specimen of F. rostratus .

It may be mentioned here that the name Murex rostratus Olivi 1792 actually is preoccupied by
Solander in Brander’s use of the name (1766). The next name available seems to be Murex
sanctaeluciae von Salis, 1793, but a change of name should not be done until the shallow-water
species are carefully revised, to avoid unnecessary name changes.

The genus Troschelia, which usually has been placed in Fasciolariidae, is here regarded to belong
to Buccinidae.

Genus FUSINUS Rafinesque, 1815
New name for Fusus Lamarck, 1799, not Helbling, 1779.

Type species: Murex colus Linne, 1758, subsequent designation by Schumacher, 1817.
Remarks. See remarks on the family.

Fusinus bocagei (P. Fischer, 1882) Figs 374-375, 383

Trophon rugosus Jeffreys, 1880:318 (nom. nud.).

Fusus bocagei P. Fischer, 1882:49.

Fusus azoricus Dautzenberg, 1889:32, pi. 2, fig. 3.

Fusus bocagei vars. major, minor, ventricosa, longicaudata, ecaudata Locard, 1897:329.

NORTH-EAST ATLANTIC NEOGASTROPODA 161

Type material: F. bocagei: lectotype in MNHN; F. azoricus: holotype in MOM.

Type locality: F. bocagei: TRAVAILLEUR 1881, dr 1 or dr 2, 43°01 N, 09°37 W, 2018 m or
41°43 N, 09° 19 W, 1068 m (mixed labels); F. azoricus: MONACO st 112, 38°34 N, 28°06 W,

1287 m.

Material examined: The type material and many samples from MONACO, BIACORES and TALISMAN
around the Azores; the shallowest record is from BIACORES st 168, 225-260 m, 12 spms; 18 samples with 173
spsm + shs in 250-500 m; 15 samples with 232 spms + shs in 500-750 m; 1 1 samples with 96 spms + shs in 750-1000
m; 3 samples with 7 spms + shs in 1000-1500 m; the deepest samples are BIACORES st 235, 2085-21 15 m, 1 spm,
1 sh; BIACORES st 171, 3215 m, 1 spm. Material outside the Azores: TRAVAILLEUR 1882, dr 2, 44° 05 N;
05°36 W, 608 m, 5 spms + shs; dr 3, 44° 04 N, 05° 34 W, 512 m, 2 spms; TALISMAN 1883, dr 90, 2 1 0 5 1 N, 17°28
W, 175 m, 1 sh; drill, 16°52 N, 25°10W, 590 m, 32 spms; MONACO st 66, 43°12 N, 09°32 W, 363 m, 1 sh; st
475, 37°52 N, 09° 16 W, 552 m, 2 shs; st 1190, 15°14 N, 23°04 W, 628 m, 2 spms; THALASSA st W401, 43°57
N, 05° 29 W, 630-920 m, 1 sh; st W407, 43° 54 N, 06°06 W, 960-1200 m, 1 sh; st W414, 43°51 N, 06° 10 W,
520-625 m, 1 sh; st W415, 43°55 N, 06° 1 1 W, 860-1150 m, 2 spms; st W423, 44°04 N, 07°07 W, 710-1070 m,

1 sh; st Y372, 41 °34 N, 09° 17 W, 800 m, 1 spm; st Y385, 41 ° 19 N, 09° 14 W, 810 m, 2 shs; st Y400, 40°46 N,
09°19 W, 800 m, 1 sh; st Z421, 48°22 N, 09°33 W, 950 m, 1 sh; CALYPSO, Gorringe Bank, 510 m, 3 shs;
METEOR, st 36-98, 25°31 N, 16°02 W, 700-900 m, 1 sh.

Distribution: The upper part of the continental slope, from the Bay of Biscay, south to the Cape
Verde Islands and the Azores. Unkown in the Mediterranean.

Remarks: Fusinus sectus (Locard, 1897) from 882 m, off Mauretania is probably a distinct species.
The whorls increase their diameter more rapidly, there are more numerous axial fibs and the
siphonal canal is distinctly curved in that species. It is known from the holotype only. Its larval shell
is too corroded to allow examination.

The animal of F. bocagei has eyes. The larval shell consists of three whorls, the last with indistinct
axial ribs, and indicates planktotrophic larval development.

The species here called Fusinus sp. and also F. rostratus can be distinguished from F. bocagei by
having a paucispiral larval shell.

Fusinus amiantus (Dali, 1889)

Figs 377-378, 386

Fusus amianta Dali, 1889: 169, pi. 15, fig. 11.

Fusus grimaldii Dautzenberg & Fischer, 1896:434, pi. 18, figs 10-1 1 .

Fusus grimaldii var. major Locard, 1897:330
Meyeria decorata Locard, 1897:337, pi. 16, figs 31-34.

Meyeria decorata var. ecaudata Locard, 1897:338.

Type material: F. amiantus , holotype USNM 508726; F. grimaldii, holotype in MOM; M. decorata,
3 syntypes in MNHN.

Type localities: F. amiantus, off Havanna (Cuba), 1480 m; F. grimaldii, MONACO st 213, 39°23 N,
31°25 W, 1384 m; M. decorata , TALISMAN 1883 dr 127, 38°38 N, 28°21 W, 1257 m.

Material examined: The type material and MONACO st 703, 39°21 N, 31 °06 W, 1360 m, 4 shs; st 719, 39° 1 1 N,
30° 24 W, 1600 m, 6 shs; st 743, 37°36 N, 25° 17 W, 1494 m, 1 sh; st 1338, 38°41 N, 28°45 W, 950 m, 1 sh; st
1349, 38°35 N, 28°06 W, 1250 m, 1 sh; st 3293, 38°47 N, 30° 16 W, 133 1 m, 2 shs; TALISMAN dr 122, 37°35
N, 29°26 W, 1440 m, 4 spms; dr 127, 38°38 N, 28°21 W, 1257 m, 15 spms + shs; BIACORES st 66, 38°34 N,
28°19 W, 1225-1260 m, 1 spm; st 179, 38°05 N, 25°46 W, 1590-1665 m, 1 spm.

Distribution: The continental slopes of the Azores and the N Caribbean.

Remarks: F. amiantus can be distinguished from F. bocagei, by having much finer spiral sculpture
and by having more pointed knobs on the axial ribs. Also the larval shells differ: F. amiantus has
more distinct axial ribs and 2 or 3 spiral cords just above the suture, instead of a single cord as in F.

bocagei.

162 P. BOUCHET & A. WAREN

Figs 374-379. Genus Fusinus . 374, F. bocagei, B1ACORES st 2 12 (37° 1 8 N, 24°45 W, 6 10 m), 26.0 mm.
375, F. bocagei, larval shell, 900 /urn. 376, F. sp., TRAVAILLEUR 1882 dr 39, 16.5 mm. 377, F.
amiantus, holotype, 15.6 mm. 378, F. amiantus, BIACORES st 66, 24.5 mm. 379, F. amiantus, larval
shell. 1 100 /urn.

NORTH-EAST ATLANTIC NEOGASTROPODA 163

Figs 380-386. Fusinid radulae. 380, Sinistralia elegcms (Reeve), height of central tooth 29 /um. 381,
Aptyxis syracusanus, 28 /urn. 382, Granulifusus nipponicus, 39 /urn. 383, Fusinus bocagei, 35 /urn. 384,
Fusinus pulchellus, 19 /urn. 385, F. rostratus, 32 /urn. 386, F. amiantus, 25 /im.

164 P. BOUCHET & A. WAREN

Figs 387-390. Radulae. 387, Nassarius frigens, breadth of central tooth 1 10 jjm. 388, Mitrella nitidulina,
30 pm. 389, Metzgeria gagei, 47 ^m. 390, Latiromitra barthelom, 80 pm.

Fusinus sp. Fig. 376

Fusinus gigliolii, Locard 1897:332, pi. 16, figs 21-25.

Material examined: TRAVAILLEUR 1882, dr 38, 34° 13 N, 07°43 W, 636 m, 1 sh; dr 39, 34° 1 1 N, 07°39 W,
530 m, 2 shs.

Remarks: Locard considered the specimens listed above to be Fusinus gigliolii (Monterosato). We
consider F. gigliolii a deep-water form of F. rostratus (Olivi), but find Locard’s specimens to differ
from any F. rostratus we have examined, in having a larger larval shell and more regularly convex
whorls, a more slender spire and a more curved siphonal canal. The specific distinction is supported
by the fact that F. rostratus was found at st 39, (listed by Locard under the name Fusus rusticulus
Monterosato and later named Pseudojusus locardi Pallary, 1904) together with Fusinus sp. The
diameter of the larval shell of Fusinus sp. is 1.3 — 1.4 mm, while F. rostratus varies from normally
700 pm, up to 1.2 mm (in deep-water specimens).

We prefer to leave this form undescribed until more material is known.

NORTH-EAST ATLANTIC NEOGASTROPODA 165

Family COLUMBELLIDAE

The genera of W Atlantic Columbellidae are treated by Radwin (1977-1978), who however was
concerned mainly with shallow-water species. We have transferred Nassarina grayi Dali from the
Columbellidae to the muricid genus Orania.

Genus AMPHISSA H. & A. Adams, 1853
Type species: Buccinum corrugatum Reeve, 1847.

Remark: The type species lives on the continental shelf of the American North Pacific.

Amphissa acutecostata (Philippi, 1844) Figs 392, 395-398

Fusus costulatus Cantraine, 1835:393 (non Fusus costulatus Lamarck, 1822).

Buccinum acutecostatum Philippi, 1844:192, pi. 27, fig. 14.

Buccinum testae Aradas, 1847:84 (fide Monterosato 1878:104).

Columbella haliaeeti Jeffreys, 1867:356, pi. 6, fig. 5.

Columbella haliaeeti var. albula Jeffreys, 1869:219, pi. 88, fig. 3.

Bela grimaldii Dautzenberg, 1889:26, pi. 2, fig. 2.

Bela lima tula Locard, 1896:141, pi. 5, fig. 3.

Anachis costulata vars. major, minor, elongata, costulata (all nomina nuda) Locard, 1897:145.

Anachis haliaeeti vars. major, curta, costulatissima, attenuata LoCard, 1897:149.

Anachis nicolayi Nordsieck, 1974:12, fig. 14.

Type material: B. acutecostatum, not located; C. haliaeeti, lectotype USNM 191601; B. grimaldii,
holotype in MOM; B. limatula, lost; A. nicolayi, holotype in coll. Giannini.

Type locality: F. costulatus, Pleistocene, Pelore, S. Italy; B. acutecostatum, fossil “in Calabria inter
Stilo et Monasterace’’; C. haliaeeti, 25 miles NNW of Unst, Shetland, 155-175 m; B. grimaldii,
MONACO st 112, 38°34 N, 28°06 W, 1287 m; B. limatula, CAUDAN st 14, 44°05 N, 02°25 W, 960
m; A. nicolayi, Cape Comino, Corsica.

Material examined: The types of haliaeeti, grimaldii and nicolayi; over 1500 spms from all parts of the
distribution area, USNM, MNHN, BMNH, ZMC.

Distribution: From the Lofoten Is. southwards on the outer banks off the Norwegian coast in
150-700 m; along the continental slope of the British Isles, the Bay of Biscay and south to 31°43 N
(MONACO st 116); the bathyal of the Mediterranean and the Azores; S of Iceland (Thorson
1941:75, and INGOLF material in ZMC); also off northern N America south to North Carolina.

Remarks: The present species has usually been known under the name Amphissa costulata or A.
haliaeeti . The name of Cantraine (1835) is however preoccupied by Fusus costulatus Lamarck, 1822.
The second available name, B. acutecostatum was used by Locard (1897) without any comments,
although Watson (1886) who had received specimens supposed to be acutecostatum from Seguenza,
wrote that he had “no doubts that the Buccinum acutecostatum is not the Columbella haliaeeti
Jeffreys”. Dautzenberg (1927) on the contrary, said he could see no differences between Recent
specimens and fossil acutecostata that he had received from Monterosato. He apparently used the
name costulata because it escaped his notice that it was preoccupied. None of these authors figured
any fossil specimens to support their conclusions.

Through the kindness of S. Palazzi, we have been able to examine fossil specimens from S Italy
(Boretto) that agree well with Philippi’s description and occur in deposits having a similar fauna as
Philippi’s specimens. Fig. 398 shows the larval shell of such a specimen. We have not been able to
find any difference from Recent specimens and conclude that acutecostatum is the correct name for
the Recent species too.

166 P. BOUCHET & A. WAREN

Figs 391-394. Radulae. 391, Volutomitra groenlandica, breadth of (central) tooth 57 /urn. 392, Amphissa
acutecostata, 55 /urn. 393, Metzgeria alba, 45 /urn. 394, Gibberula abyssicola, 20 /um.

A. acutecostata is a rather variable species. Mediterranean specimens tend to be uniformly rather
solid, in shallow water with blotches of light tan. Atlantic specimens are white or greyish and the
solidity of the shell is variable. The development of the axial ribs is variable, even in a single
population.

The larval shell is multispiral and has an elaborate sculpture (Fig. 397). It has been figured by
Bouchet (1977) and Bouchet & Waren (1979) as A haliaeeti. Simroth (1895) and Richter & Thorson
(1975) obtained the larva in plankton (described as C. haliaeeti and species 4 respectively). Bouchet
& Fontes (1981) examined the isotope composition of the larval shell and concluded that the larval
shell had been formed in more shallow water than the postlarval shell.

NORTH-EAST ATLANTIC NEOGASTROPODA 167

Figs 395-398. Amphissa acutecostata. 395 and 396, THALASSA st Z442 (48°55 N, 1 1°02 W, 975 m), 7.0
and 6.1 mm. 397, larval shell of a recent specimen, 1025 pm. 398, larval shell of a Quaternary fossil from
Boretto, 1050 pm.

168 P. BOUCHET & A. WAREN

Genus MITRELLA Risso, 1826

Type species: Mitrella flaminea Risso, 1826.

Mitrella nitidulina (Locard, 1897)

Figs 388, 399-400

Columbella nitidulina Locard, 1897:143, pi. 14, fig. 10-13.

Type material: Lectotype (here selected) in MNHN.

Type locality: TALISMAN dr 76, 25°01 N, 16°55 W, 2638 m.

Material examined: The type material and TALISMAN dr 40, 30°03 N, 1 1°42 W, 2212 m, 1 sh; dr 131, 38°38 N,
25° 06 W, 2995 m, 4 spms + shs; MONACO st 536, 37°54 N, 24°43 W, 2178 m, 1 spm; st 698, 39° 1 1 N, 20°45 W,
1846 m, 26 shs; st 738, 37°40 N, 26°26 W, 1919 m, 7 spms + shs; st 1209, 16°34N, 23°03 W, 1477 m, 1 spm, 1 sh;
st 1334, 39° 30 N, 29°02 W, 1900 m, 1 sh; st 1349, 38°35 N, 28°06 W, 1250 m, 1 sh; BIACORES st 126, 39°19N,
33°47 W, 3360 m, 1 sh; st 173, 37°57 N, 26°08 W, 3225 m, 1 sh; BIOGAS st CV38, 47°31 N, 08°59 W, 2695 m, 1
spm; J. CHARCOT MADERE st 17, 32°58 N, 16°26 W, 1390-1630 m, 1 sh.

Distribution: The deep bathyal and upper abyssal parts of the NE Atlantic, from the Bay of Biscay
and the Azores to Morocco. Only known from the material examined.

Remarks: M. nitidulina is closely related to Astyris profundi Dali, 1889 from the western Atlantic
basin, but is broader, has smaller apertural teeth and coarser incremental lines. M. nitidulina also
has a more inflated larval shell and lacks the characteristic axial and spiral microsculpture of
profundi.

The protoconch is multispiral and has an elaborate sculpture, indicating planktotrophic larvae.

Mitrella sp.

Fig. 402

A single worn shell of another Mitrella species was taken at BIACORES st 101, 39°23 N, 31°05 W,
843-900 m.

It is a broken shell in poor condition (Fig. 402) but shows some resemblance to Astyris profundi
Dali, 1889 (Fig. 401). The specimens, however, are too few to allow certain identification because of
the often very minute differences between columbellids.

Genus ANACHIS H. & A. Adams, 1853

Type species: Columbella scalarina Sowerby, 1832.

Anachis sp

'Tg. 403

A single shell of a small Anachis was taken by INGOLF at 60° 47 N, 04°00 E, in 350 m. It probably
represents an undescribed species.

NORTH-EAST ATLANTIC NEOGASTROPODA 169

Figs 399-403. Columbellidae. 399, Mitrella nitidulina, TALISMAN dr 131, 1 1.6 mm. 400, M. nitidulina,
MADERE st 17, larval shell, 820 ^m. 401, M. profundi, holotype, 8.1 mm. 402, M. cf. profundi,
BIACORES st 101, 8.5 mm. 403, Anachis sp., INGOLF, 2.4 mm.

170 P. BOUCHET & A. WAREN

Family BUCCINIDAE

Most previous workers on the family have divided it into several subfamilies or families (Cossmann,
1901 ; Wenz 1941; Golikov & Starobogatov, 1975; Habe & Sato, 1973; Kantor, 1981), while Ponder
(1973) united Buccinidae, Fasciolariidae, Nassariidae, and Melongenidae to a single family,
Buccinidae, and considered the former families subfamilies. Our experience from genera like
Kryptos, Troschelia and Belomitra supports Ponder’s view. We have, however, kept the old families
for some genera of typical Fasciolariidae and Nassariidae, but placed the doubtful genera in
Buccinidae.

The taxonomy of the N Atlantic buccinids has been found confused and in order to reduce the
confusion we have included a number of shallow-water species and W Atlantic species from N
Carolina and northwards.

Below we discuss some of the taxonomic criteria we have used and give a provisional key to the N
Atlantic genera.

Larval shell. The larval development of Buccinum undatum and Colus stimpsoni has been
described in detail (Portmann, 1925, 1930; West, 1973). Several hundred to several thousand eggs
are laid in each capsule. A few of these develop to shell-less veligers. These ingest further eggs and
store them in the digestive tract, which thus acts as a yolk sac. In B. undatum (Portmann, 1925) and
Colus gracilis (own observations) the shell is formed rather late in the development and takes a shape
according to the shape of the visceral sac, which varies depending on how many nurse eggs the larva
has eaten. In some specimens of C. gracilis we noticed that the formation of the shell started with the
lower parts and the apical part was the last to be formed. After the shell has been formed, normal
“postlarval” growth starts, while the larva still lives in the egg capsule, feeding on less developed
embryos. Fig. 588 shows larvae from capsules where these two stages can be seen.

In C. stimpsoni a larval shell is said to develop rather early, but the apical part is later thrown off,
leaving a scar where it has been attached (West, 1973). We have not noticed this scar in perfect
protoconchs of C. stimpsoni.

In other species the larva starts to form a shell at a very small size, presumably before it starts
feeding by adelphophagy. We have observed this in Mohnia mohni and M. danielsseni. The two
species hatch with a shell of three whorls, starting with a protoconch of a diameter of about 0.5 mm.
In M. abyssorum we have observed veligers with a shell of 0.6 mm, consisting of slightly less than
one whorl. (This species, however, seems not to have nurse eggs, because our preserved capsules are
filled with a solid yolk mass.) Also M. abyssorum hatches at a size of three whorls (about 4 mm).

When the shells of embryos ready to hatch of these species of Mohnia are compared to those of
embryos of Buccinum, the difference is striking (Figs 547, 590). The same applies when apices of the
different groups are compared. We assume, however, that the difference does not mean as much
systematically as one might believe, but that the development of Buccinum (and other genera with
similar larval shell) represents a more advanced type, where the formation of the shell is delayed until
shortly before the young leave the shelter of the capsule. Such an evolution, with delay of shell
formation may have taken place several times and we do not assume that similar types of larval shell
necessarily indicate relationship.

Operculum. Opercula of adult buccinids have often been divided into three groups: the Buccinum
type with a central or subcentral nucleus, the normal type with an apical nucleus and the Mohnia
type with a paucispiral operculum (Figs 442, 413, 431). As in most neogastropod species, the apical
portion of the operculum gets eroded with increasing age and it is usually impossible to study the
nucleus in adult specimens of Buccinidae. We have examined opercula of both young and adult
specimens of almost all species treated in the revision and found opercula of recently hatched young
to be much more informative than those of adult specimens.

Thus we have been able to distinguish the following types of opercula:

NORTH-EAST ATLANTIC NEOGASTROPODA 171

A. Mohnia type (Fig. 431). Paucispirdl. Often this can be seen only in young specimens because the
lower part of the operculum is lacking. Occurs in Mohnia.

B. Buccinum type (Fig. 442). Concentric growth. Occurs in Buccinum.

C. Turrisipho type (Fig. 407). The lower part of the larval operculum is very pointed. The axis of this
point forms a small angle with later growth. Occurs in Turrisipho, possibly also in Beringius.

D. Colus type (Fig. 413). The operculum of the embryo is regularly ovate and it grows in an open
spiral, so the axis of the adult operculum forms a more or less right angle with the length axis of the
larval operculum. Occurs in Colus.

E. Neptunea type (Fig. 404). Similar to that of Colus, but the growth takes place in the same
direction throughout the life of the snail, so the axis of the larval operculum is the same as that of the
adult one. Occurs in Neptunea.

F. Some other types of opercula, that do not fit into any of these groups, occur in Liomesus,
Volutopsius and Kryptos.

Radula. The radulae of N Atlantic buccinids have not been used very much for taxonomic
purposes, presumably because of the large intraspecific variation reported by Friele (1879) and
verified by later authors. We have, however, found the radula to be a useful accessory tool in the
classification of the N Atlantic Buccinidae, and give figures of most of them. Two main types can be
distinguished:

A. Buccinum type (Fig. 469) with a short, broad central tooth with 4-9 cusps with individual bases.
Occurs in Buccinum, Neptunea, Volutopsius.

B. Mohnia type (Fig. 480) with a more or less square, often thin or reduced central plate with 0-3
cusps with fused bases, often appearing as processes from the anterior edge of the plate. Occurs in
Mohnia, Colus, Turrisipho. In addition to these, there are a number of species that have more
modified radulae: Belomitra, Liomesus, Kryptos, Troschelia, Beringius . The phylogenetic
interpretation of these radulae is discussed under each genus, but our general impression is that they
represent groups of low taxonomic level.

Among the genera here considered to be buccinids, Belomitra stands out by having a central tooth
resembling that of Fasciolariidae. We do not know if this is convergence or indication of
relationship: Similarities to Fasciolariidae in the radula are also found in Troschelia. Here, however,
it is the lateral teeth that are similar (cf. Figs 484, 486), while the central teeth are of normal Mohnia
type.

Concluding remarks. The N Atlantic species of Buccinidae may be distributed rather naturally
into ten genera, with the exception of Mohnia glypta, M. caelata and M. carolinensis which do not
fit very well into any described genus, when radula, operculum and shell shape are used in
combination. The two last mentioned of these species show some resemblances to tropical deep¬
water buccinids of the genus Manaria E.A. Smith, 1906, which are very poorly known, the first
mentioned species resembles no known buccinid. We have preferred to place these species
provisionally in Mohnia until the tropical species are better known.

To test our generic concept, we have tried to classify the N Pacific species of which operculum and
radula are known, which worked very well.

172 P. BOUCHET & A. WAREN

Key to the genera of N Atlantic Buccinidae

A. Operculum with central or subcentral nucleus (Figs 442, 445)

A. Operculum without central or subcentral nucleus

B. Operculum, at least in very young specimens, paucispiral

B. Operculum not paucispiral in young specimens

C. Whorls distinctly shouldered with angular axial ribs

C. Whorls rounded

D. Central teeth of the radula without cusps

D. Central teeth of the radula with cusps

E. Shell short, ovate, almost smooth

E. Shell tall, fusiform, with distinct spiral sculpture

F. Lateral teeth with 5-10 cusps of equal size

F. Lateral teeth with 2-5 cusps of distinctly unequal size

G. Aperture very large, siphonal canal short, broad and open, only 2 large cusps
on the lateral teeth

G. Aperture constricted at the siphonal canal which is distinctly set off

H. Periostracum hardly visible

H. Periostracum usually well developed, always easily visible in specimens taken
alive

I. Larval operculum pointed, periostracum with rows of bristles along the spiral
ridges

I. Larval operculum oval, periostracum smooth or with bristles along the
incremental lines

Buccinum

B

Mohnia

C

Kryptos

D

E

F

Liomesus

Beringius

Troschelia

G

Volutopsius

H

Neptunea

I

Turrisipho

Coins

NORTH-EAST ATLANTIC NEOGASTROPODA 173

Figs 404-412. Buccinid opercula. 404, Neptunea brevicauda, E. Canada, 6.6 mm. 405, Neptunea antiqua,
off Korsfjorden, Bergen area, 300 m, 4.2 mm. 406, Turrisipho fenestratus, W of Korsfjorden, 280 m, 2.5
mm. 407, Turrisipho moebii, same locality, 3.9 mm. 408, Turrisipho moebii, off Reykjanes, Iceland, 19.2
mm. 409, Turrisipho lachesis, INGOLF st 28, 5.3 mm. 410 and 41 1 , Turrisipho voeringi, INGOLF st 105,
6.5 mm and 14.4 mm. 412, Troschelia berniciensis, southern Bay of Biscay, 400-690 m, 15.7 mm.

174 P, BOUCHET & A. WAREN

Figs 413-421. Buccinid opercula. 413, Coins terraenovae, Newfoundland Banks, 7.3 mm. 414 and 415,
Coins gracilis, W Norway, 2.6 and 16.3 mm. 416, Coins gracilis , E Greenland, 10.0 mm. 417, Coins
tnrgidnlns, Norwegian Sea, 11.7 mm. 418, Coins sabini, Baffin Bay, 8.0 mm. 419 and 420, Coins
holboelli, Greenland, 3.2 mm and 7.7 mm. 421, Coins ho/boe/li, Baffin Bay, 12.1 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 175

424

,

W

• >

-

427

T< -';T-: ■''

Figs 422-430. Buccinid opercula. 422, Colus turgidulus, Norwegian Sea, 1 1.7 mm. 423, Colus laterieeus,
Norway, 2.3 mm. 424, Belomitra quadruplex, INCAL st CP20, 7.9 mm. 425, Mohnia krampi, syntype,
7.5 mm. 426 and 427, Mohnia abyssorum, Bay of Biscay, 1.9 mm and 9.5 mm. 428, Colus jeffreysianus,
Northumberland, UK, 7.3 mm. 429, Colus jeffreysianus, Bay of Biscay, 13.3 mm. 430, Mohnia glyptus,
INGOLF st 32, 1.8 mm.

176 P. BOUCHET & A. WAREN

Figs 431-439. Buccinid opercula. 431 and 432, Mohnia danielsseni, Norwegian Sea 3.0 mm and 8.2 mm.
433, Mohnia caelata, off NE United States, 3.6 mm. 434, Mohnia carolinensis, off NE United States, 1 .6
mm. 435, Mohnia mohni, N of Iceland, 7.1 mm. 436, Mohnia parva, off NE United States, 2.8 mm. 437
and 438, Mohnia simplex, INGOLF st 35, 3.6 mm and 3.4 mm. 439, Mohnia blakei, off NE United
States, 3.7 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 177

Figs 440-448. Buccinid opercula. 440, Liomesus ovum, W Norway, 5.1 mm. 441, Kryptos koeh/eri, off
Portugal, 2.0 mm. 442, Buccinum kjennerudae, INGOLF st 28, 2.3 mm. 443, Volutopsius norwegicus,
off E Canada, 3.0 mm. 444, Volutopsius norwegicus, TromsO, N Norway, 20.8 mm. 445, Buccinum
abyssorum, off NE United States, 7.6 mm. 446, Beringius turtoni, North Sea, 3.5 mm 447, Beringius
turtoni, W Norway, 23.7 mm. 448, Buccinum abyssorum, off NE United States, 8.7 mm.

178 P. BOUCHET & A. WAREN

Figs 449-452. Radulae of Mohnia. 449, M. krampi, breadth of central tooth 112 /urn. 450, M. blakei, 41
/u m. 451, M. parva, 36 /urn. 452, M. simplex, 50 /um.

NORTH-EAST ATLANTIC NEOGASTROPODA 179

Figs 453-460. Buccinid radulae. 453, Mohnia glyptus, breadth of central tooth 30 /urn. 454, M. caelata, 58
fum. 455, Turrisipho moebii, 38 /urn. 456, Neptunea brevicauda, 64 /urn. 457, Mohnia carolinensis, 19/um.
458, Turrisipho lachesis, 19 /urn. 459, Buccinum abyssorum, 223 (jm. 460, B. kjennerudae, 81 /urn.

180 P. BOUCHET & A. WAREN

A

Figs 461-465. Buccinid radulae. 461 , Liomesus ovum from egg capsule, breadth of central tooth, 46 pm.
462, Colus terraenovae, 67 pm. 463, C. stimpsoni, 69 pm. 464, C. pygmaeus, 49 pm. 465, C. pubescens,
94 pm.

NORTH-EAST ATLANTIC NEOGASTROPODA 181

Figs 466-47 1 . Buccinid radulae. 466, Beringius turtoni, breadth of central tooth 190 /xm. 467, Volutopsius
norwegicus, 1 12 /xm. 468, Buccinum humphreysianum, 290 /xm. 469, Buccinum undatum, 318 /xm. 470,
Colus turgidulus, 258 /xm. 471, Colus holboelli, 155 /xm.

182 P. BOUCHET & A. WAREN

Figs 472-477. Buccinid radulae. 472, Colus sabini, breadth of central tooth 169 )jm. 473, Colus gracilis,
164 (jm. 474, C. jeffreysianus, Bergen, S. Norway, 106 /urn. 475, C. jeffreysianus, N Bay of Biscay, older
part of the ribbon, 131 /urh. 476, Neptunea antiqua, 3 12 /urn. 477, C. jeffreysianus, same specimen as Fig.
475, younger part of the ribbon, 114 /urn.

NORTH-EAST ATLANTIC NEOGASTROPODA 183

Figs 478-483. Buccinid radulae. 478, Coins latericeus, breadth of central tooth 32 pan. 479, Coins
islandicus , 238 ^m. 480, Mohnia danielsseni, 81 /jm. 481, Mohnia abyssorum, 97 ^m. 482, Mohnia
mohni, 68 /^m. 483, Turrisipho fenestratus, 48 ptm.

184 P. BOUCHET & A. WAREN

Figs 484-489. Buccinid and fusinid radulae. 484, Troschelia berniciensis, Bergen, S Norway, breadth of
central tooth 1Q2 /urn. 485, Troschelia berniciensis, N Bay of Biscay, 68 /am. 486, Fusinus bocagei, 19 /am.
487, Kryptos koehleri, 17 /am. 488 and 489, Belomitra quadrup'ex, 23 /am.

NORTH-EAST ATLANTIC NEOGASTROPODA 185

Genus LIOMESUS Stimpson, 1865

Type species: Buccinum dalei Sowerby, 1825, by monotypy (Fig. 490).

Synonym: Buccinopsis Jeffreys, 1867 is an absolute junior synonym. Type species Buccinum dalei
Sowerby. (Not Buccinopsis Conrad, 1856).

Remarks: There is some doubt about the identity of the type species. We have examined fossil
specimens agreeing perfectly with Sowerby’s original description, both from Belgian and British
localities and found them to differ from the Recent form by having a larger, more solid shell, blunter
and more rounded spire, and coarser striation. Nowhere in the present distribution can populations
be found that agree in these respects with the Crag fossil. We have, therefore, preferred to use the
specific name ovum Turton, 1825, which is a few months younger than Sowerby’s name and based
on the Recent form. The same opinion was expressed by Dali (1916).

The genus Liomesus has been separated from Buccinum because of its radula (with a single large
lateral cusp and a cusp-less rounded central plate (Sars, 1878)) and an operculum without central or
subcentral nucleus. The North Pacific species previously referred to Liomesus have proved to have a
more normal buccinid radula and were referred to Pseudoliomesus by Habe & Sato (1973). Liomesus

Figs490-491. Genus Liomesus. 490, L. dalei , upper Pliocene of Orford, 40 mm. 491, L. ovum, off Ireland,
36 mm.

186 P. BOUCHET & A. WAREN

stimpsoni Dali, 1889 has a bicuspid lateral tooth of normal buccinid shape and a monocuspidate
central (Weber, 1959). Therefore L. ovum is the single remaining species of the genus. Young
specimens of L. ovum have a lateral tooth with two large and two smaller cusps and a central plate
with several cusps (Fig. 461) and we question if the peculiar radula of L. ovum should not be
considered a specific adaptation and the classification of these species should be reconsidered.

Liomesus ovum (Turton, 1825)

Figs 440, 461,490-491

Buccinum ovum Turton, 1825:366
Halia flemingiana Macgillivray, 1843:189.

Tritonium eburneum M. Sars, 1851:192 (not Lowe 1846).

Buccinopsis dalei of authors on Recent molluscs, not Sowerby, 1825.

Type material: B. ovum , holotype USNM 192257; T. eburneum: holotype ZMO D. 25954.

Type locality: B. ovum, off Plymouth, Great Britain; T. eburneum, Vestfjorden, N. Norway, 75-90 m.
75-90 m.

Material examined: About 50 shells and specimens from all parts of the distributional area, BMNH, MNHN,
ZMO, ZMC, USNM, SMNH.

Distribution: From Lofoten (NW Norway) and W and NW of Iceland (Oskarsson, 1966) along the
European shelf south to the extreme north of the Bay of Biscay (THALASSA st Z422, 48°21 N,
09°39 W, 1 175 m, 1 fresh shell). Normally it is a shelf species, occurring in 100-300 m but sometimes
it is found on the upper part of the continental slope (several records from the Porcupine Expedition
1869 from off Ireland and the Thalassa station above).

Remarks: Jeffreys (1867) suspected the type locality given by Turton to be wrong and that it actually
came from Co. Cork, Ireland. Fie also figured and described the animal and the egg capsules. M.
Sars (1851) described the animal.

Except for the synonymy with B. dalei there are no taxonomical problems with L. ovum. It is
easily recognized by the almost perfectly smooth, ovate shell of a Buccinum- like shape, but without
a central or subcentral nucleus in the operculum.

Genus BUCCINUM Linne, 1758

Type species: Buccinum undatum Linne, 1758, subsequent designation De Montfort (1810).
Synonyms: Tritonellium Valenciennes, 1858. Type species T. barthae Valenciennes, 1858, by
monotypy.

Mada Jeffreys, 1867 (not Mulsant, 1850). Type species B. Humphreys ianum Bennett, 1824, by
monotypy.

Mala Cossmann, 1901 (not Distant, 1854). Replacement name for Mada Jeffreys.

Madiella Wenz, 1943. Replacement name for Mada Jeffreys.

Remarks: We can see no reason for separating Madiella as a distinct genus. The only character ever
given for this is the fine, regular, incised sculpture of B. Humphrey sianum, but this occurs in other
species (e.g. B. meridionale Harmer, from Newfoundland), often in combination with a coarser
sculpture.

NORTH-EAST ATLANTIC NEOGASTROPODA 187

There are some problems with the type designation of Buccinum. Lamarck (1799) cited B.
undatum as an example of the genus “pour se faire mieux entendre”, but this can hardly be
considered a type designation (R.V. Melville, pers. comm.), although the intention was the same. It
will, however, in this case make no difference, because De Montfort designated the same species as
Lamarck as type species.

The northern species of Buccinum, both of the Atlantic and the Pacific are highly variable and
hundreds of different species and forms have been described. We do not intend to make a revision of
all these forms, and we have included only the species restricted to deep water, S of the Iceland-
Faroe Ridge. N of this ridge, there are some species in bathyal depths, but they normally occur more
shallowly and there are no species of Buccinum in the arctic, abyssal fauna (Bouchet & Waren, 1979;
Friele & Grieg, 1901).

The more southern species of Buccinum are less variable and may be separated by the following

characters:

A. Axial sculpture dominating B. undatum

A. Axial sculpture absent or weak B

B. Only a very fine and regular sculpture of equal sized, incised spiral B. humphreysianum
lines present

B. Spiral sculpture consisting of larger and smaller raised ribs C

C. Some spiral ridges with knobs, apertural height greater than spire height B. oblitum

C. No spiral ribs with knobs, apertural height less than spire height D

D. Two or three sharp spiral keels much larger than the others, animal

without eyes B. abyssorum

D. Several equal sized, rounded spiral keels present, animal with eyes B. kjennerudae
A revision of Buccinum was recently published by Golikov (1980), who recognized about 90
species, but synonymized B. oblitum and kjennerudae with arctic shallow-water species.

Buccinum undatum Linne, 1758 Figs 469, 492-493, 590

Buccinum undatum Linne. 1758:740.

Type locality: “O. Europaeo”.

Type material: Four syntypes in the Linnean Society, London.

Distribution: Along the North Atlantic shores, from New Jersey, U.S.A. (Abbott, 1974), north to
Greenland (Thorson 1951) (70° N), Spitzbergen (Odhner, 1915), N Norway (Sars, 1878) and S to N
Spain (Bay of Biscay, Ortea, 1977), Portugal (Hidalgo, 1917) and possibly the W Mediterranean
(Altimira, 1975, Ghisotti, 1974, Jeffreys, 1867, et ai). Normally it occurs in shallow water, 0-100 m,
but it has been recorded from bathyal depths (Friele & Grieg, 1901, 1500 m; Sykes 1911, 1450 m,
etc.). We have, however, not seen specimens taken alive from depths greater than 500 m
(Korsfjorden, W. Norway, coll. AW, cf. Fig. 492).

Remarks: B. undatum has been included here only to exclude it from the bathyal fauna. It was not
represented by a single shell in the collections made by Travailleur, Talisman, Thalassa, Biacores,
Monaco Expeditions, etc., from deep water in the Bay of Biscay, where it is scarce along the S coast.
Scattered empty shells occur off Ireland and Scotland (Sykes, 1911) where it is common at the coast
and the shelf (Massy, 1930). Therefore we suspect shells to have been carried out by fish, many
species of which are known to eat Buccinum.

188 P. BOUCHET & A. WAREN

Figs 492-493. Buccinum undatum. 492, Korsfjord, Bergen area, 150-500 m, 71.3 mm. 493,
Lieholmsrannan, Bergen area, 70-73 m, 34 mm.

Buccinum humphreysianum Bennett, 1824 Figs 468, 494-496

Buccinum humphreysianum Bennett, 1824:398.

Buccinum fusiforme Kiener, 1834:5 (not Borson, 1820).

Buccinum ventricosum Kiener, 1834:4 (not Grateloup, 1827).

Buccinum striatum Philippi, 1844:193 (not Pennant, 1777).

Buccinum puxleianum Leach, 1847:270 (nom. mid.).

Buccinum humphreysianum var. lactea Jeffreys, 1867:294 (not Kiener, 1834).

Buccinum inf latum Aradas & Benoit, 1876:287 (not Shaw, 1811).

Buccinum kieneri Monterosato, 1872:32 (new name for B. fusiforme Kiener, 1834, not Broderip, 1830).
Buccinum atractodeum Locard, 1886:107.

Buccinum monterosatoi Locard, 1886:109.

Buccinum monterosatoi var. flammulata Locard, 1886:110.

Buccinum monterosatoi var. minor Locard, 1897:280.

Buccinum humphreysianum var. azonata Locard, 1886:106.

Buccinum liocephalum Pallary, 1931:9.

Buccinum lusitanicum Pallary, 1931:7.

Buccinum humphreysianum var. euthriaeformis Paulus & Mars, 1942:73.

Buccinum inf latum panormitanum Settepassi, 1977, app.:6.

Buccinum inf latum partenopaeum Settepassi, 1977, app.:6.

Buccinum gracile Monterosato in Settepassi, 1977, app.:6 (not Reeve, 1846).

NORTH-EAST ATLANTIC NEOGASTROPODA 189

Type material: B. Humphrey sianum: not known.

Type locality: B. Humphrey sianum: Cork, Ireland.

Material examined: About 150 specimens from all parts of the distribution area, USNM, MNHN, ZMC,
SMNH, ZMB, coll. AW.

Distribution: Along the European shelf and the uppermost continental slope, from Finmark (N
Norway) (Sars, 1878) and E of Iceland (Oskarsson, 1977) to W Morocco (Locard, 1897) and the
western part of the Mediterranean (Settepassi, 1977; Casamor & Ghisotti, 1968). The depth
distribution is 15 m (Kattegat, ZMC) — 360 m (W Norway, coll. AW) in the northern part and 70 m
(Casamor & Ghisotti, 1968) — 1190 m (Locard, 1897) in the southern part of the range.

Remarks: The rich synonymy of B. Humphrey sianum is not the result of a high intraspecific
variation, but more the result of vain splitting by “la nouvelle ecole” and its recent representatives.

We have examined Pleistocene specimens from Sicily (B. striatum Philippi) and found them to fall
well inside the variation of Recent B. Humphrey sianum.

B. Humphrey sianum has often (the last occasion by Abbott, 1974) been recorded from the W
Atlantic and the Arctic, but we have seen no such specimens, despite searching many collections.
The egg capsules are hemispherical and are laid singly (Jeffreys, 1867).

Buccinum abyssorum Verrill & Smith, 1884 Figs 445, 448, 459, 497-499
Buccinum abyssorum Verrill & Smith in Verrill, 1884:167, pi. 31, fig. 11.

Type material: 2 syntypes USNM 35644.

Type locality: USFC st. 2111, off North Carolina, 1700 m.

Material examined: 24 samples with about 80 specimens from the W Atlantic; INGOLF st 65, 61°33 N, 19°00
W, 2051 m, 2 spms; st 95, 65°14 N, 30°39 W, 1416 m, 2 shs; INCAL CP 05, 55°00 N, 12°30 W, 2884 m, 5 spms;
CHALLENGER II, the Rockall Trough, c. 2000 m, several specimens.

Distribution: On the lower part of the continental slope, from N Carolina to SW Greenland
(Thorson, 1951), SW of Iceland and W of Ireland. Depth range 1400—2900 m.

Remarks: There is some degree of variation in the proportions of the shell, but the absence of eyes is
a good character distinguishing B. abyssorum from B. kjennerudae. The sculpture is also different in
the two species. B. kjennerudae lacks the strong, sharp spiral keels characteristic of B. abyssorum
and has evenly rounded ridges instead.

Buccinum oblitum Sykes, 1911 Figs 500-501

Buccinopsis striata Jeffreys in Wyville-Thomson, 1873: fig. 76 (not Buccinum striatum Muller, 1774 or any of
the other ten B. striatum that have been described).

Buccinum oblitum Sykes, 1911:342, Fig. unnumbered
Buccinum perexiguum Dautzenberg, 1925:5, fig. 1.

Type material: B. oblitum, lectotype (Waren 1980) USNM 191839; B. perexiguum, holotype in
MOM.

Type localities: B. oblitum, PORCUPINE 1869, st 55, 60°04 N, 06° 19 W, 1100 m; B. perexiguum
MONACO st 1096, 36°07 N, 08°03 W, 1440 m.

Material examined: The types and 5 shs from the type locality of B. oblitum, USNM and BMNH; 1 sh, off
Korsfjorden, W Norway, coll. AW; THOR st 164, S of Iceland, 1900 m, 1 sh; INGOLF st 90, 64°45 N, 29°06
W, 1070 m, 2 shs.

Distribution: Only known from the material examined, from the shelf and upper continental slope,
from Iceland to S of Portugal.

Remarks: B. oblitum can always be recognized by its small size combined with a thickened outer lip,
large aperture and spiral ribs with small knobs. The animal is not known, but the shell agrees in all
details with that of Buccinum while the sculpture is quite different from the smooth shell of
Liomesus. We have therefore preferred to place oblitum in Buccinum, as did Sykes (1911) and
Dautzenberg (1925).

190 P. BOUCHET & A. WAREN

Buccinum kjennerudae nom. nov. Figs 442, 460, 502-504

Buccinum sulcatum Friele, 1882:32, pi. 3, fig. 18 (not Born, 1778, Meuschen, 1787, Bruguiere, 1792, Sowerby,
1822, Kiener, 1834).

Type material: Holotype ZMB 22917.

Type locality: Norwegian North Atlantic Expedition st 192, N of Lofoten, 1150 m.

Material examined: The holotype and INGOLF st 28, 65° 14 N, 55°42 W, 791 m, 2 spms; st 35, 65° 16 N, 55°05
W, -682 m, 2 shs; TJALFE st 367, 66° 22 N, 57° 16 W, 686 m, 1 spm.

Distribution: Only known from the material examined, from N Norway, to Davis Strait, in
600—1100 m.

Remarks: The present species has never been recognized as a good species, probably because of its
rarity. Golikov (1980) synonymized it with B. polare, a high arctic shallow-water species with distinct
axial sculpture.

The differences from B. abyssorum are listed under that species. An additional difference is the
shape of the spire, which is more convex in B. abyssorum.

The name B. sulcatum which was suggested by Friele is preoccupied several times, so we found it
appropriate to name it after Miss Johanne Kjennerud, curator at ZMB, who has been of great help
in our work.

NORTH-EAST ATLANTIC NEOGASTROPODA 191

Figs 494-499. Genus Buccinum. 494, B. humphreysianum, syntype of B. monterosatoi, off St Henri, SE
France, 41 mm. 495, B. humphreysianum, 35°31 N, 06°35 W, 535 m, 68.9 mm. 496, B.
humphreysianum, Bergen area 55-65 m, apex 1.3 mm. 497, B. ahyssorum, 1NCAL st CP5, 30.1 mm. 498
and 499, B. abyssorum, off Nova Scotia, ca. 40 mm and apex 1.3 mm.

192 P. BOUCHET & A. WAREN

Figs 500-504. Genus Buccinum. 500, B. oblitum, N. Marsteinen, Bergen area, 300-330 m, 14.7 mm. 501,
INGOLF st 90, 16.6 mm. 502, B. kjennerudae, holotype of B. sulcatum, 26 mm. 503 and 504, INGOLF st
28, 1 1 .0 mm and apex 1 .9 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 193

Genus TROSCHELIA MOrch, 1876

Type species: Fusus berniciensis King, 1846, by monotypy.

Remarks: Boreofusus G.O. Sars, 1878 has the same type species and is an objective junior synonym.
Buccinofusus Conrad, 1868 was synonymized with Troschelia by Tryon (1881) and several later
authors, but we find that genus more similar to typical species of Fusinus, as did Dali (1909).

We do not know any additional species of Troschelia. Troschelia (Thalassoplanes) moerchi Dali,
1908 (figured by Kosuge, 1975) has almost no siphonal canal and quite a different operculum, which
is very slender. Fusus perminutus Dali, 1927 from off Florida, resembles the young of Troschelia
berniciensis , but was described from immature specimens with no animal, so its systematic position
remains uncertain.

The radula of Troschelia berniciensis made G.O. Sars (1878) place the genus in Fasciolariidae.
Thalassoplanes Dali, 1908 has a similar radula, but the central tooth of Thalassoplanes moerchi
(which possibly is a synonym of Brevisiphonia circumreta Lus, 1973) is quite reduced and only a
small plate remains. Costaria borealis Golikov, 1977 has lateral teeth with 7 cusps of equal size and a
typical buccinid central tooth. Calliloconcha solida Lus, 1978 has the same kind of central tooth as
C. borealis but lateral teeth with only three cusps of equal size. We therefore suppose that these
groups represent a trend in the Buccinidae, of reduction of the central tooth and a corresponding
increase in the number of cusps on the lateral teeth and we prefer to place Troschelia in the
Buccinidae.

Troschelia berniciensis (King, 1846) Figs 412, 484=485, 505-510.

Fusus berniciensis King, 1846:246.

Fusus berniciensis var. elegans Jeffreys, 1867:342.

Fusus berniciensis var. inf lata Jeffreys, 1877:327.

Boreofusus berniciensis var. solida Jeffreys in G.O. Sars, 1878: 279.

Fusus (Siphonorbis) amblyterus Watson, 1886:205.

Neptunia aquitanica Locard, 1897:356.

Neptunia berniciensis vars carinata, major , minor , ventricosa, elongata Locard, 1897:354.

Type material: F. berniciensis, not known.

Type locality: F. berniciensis , deep water off Northumberland (North Sea).

Material examined: About 250 shells and specimens from all parts of the distributional area (MNHN, USNM,
ZMC, coll. AW).

Distribution: Along the NE Atlantic continental shelves and upper slopes, from 56°01 N, 32°42 W,
1200 m (Jeffreys, 1877) (between Iceland and Greenland) to NW Norway (Sars, 1878) and
southwards to 25 °N (Locard, 1897). It is not known from the Mediterranean. In Norway it has been
found from 90 m (Sars, 1878) down to 600 m on the continental slope (Friele & Grieg, 1901), but
occurs deeper (1000 m) in the fjords (AW). Off North-West Africa it occurs in depths between 600
and 2000 m (Locard, 1897). The deepest occurrence is BIACORES stn 252, 47°35 N, 08°47 W, 2700
m, where empty but fresh shells were found.

Remarks: T. berniciensis is a rather variable species. Northern specimens are rather broad (Fig. 505),
southern specimens more slender (Locard’s N. aquitanica, Fig. 507), and specimens from the
deepest part of the distribution are smaller and slender (Watson’s F. amblyterus, Fig. 509).

There is also a variation in the number of cusps of the lateral teeth, from 8-10 in northern
specimens from shallow water, to about 5 in southern specimens from deep waters (Figs 484-485).
Specimens from intermediate localities have intermediate numbers of cusps.

194 P. BOUCHET & A. WAREN

Figs 505-510. Troschelia berniciensis. 505, Akrafjord, Norway, 620 m, 97.4 mm. 506, THALASSA st
W405 (43°56 N, 05°44 W, 400-690 m), 64.4 mm. 507, THALASSA st X322 (44°00 N, 04°45 W, 980-1080
m), 90.4 mm. 508, BIOGAS st CV39 (47°33 N, 08°45 W, 2350 m), 61.4 mm. 509, Bay of Biscay, deep
water form, 61.2 mm. 510, off Korsfjorden, Bergen area, apex 1.8 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 195
Genus KRYPTOS Jeffreys in Dautzenberg & Fischer, 1896

Type species: Kryptos koehleri (Locard, 1896) (= Kryptos e/egans Jeffreys, 1896), by monotypy.

Remarks: This genus was placed in Fasciolariidae by Dautzenberg & Fischer (1896), Thiele (1931)
and Wenz (1943) while Locard (1897) considered it to belong to Turridae. For reasons given in the
introduction to Buccinidae, we prefer to place it in Buccinidae.

We do not know of any additional species from the Atlantic which should be classified in Kryptos.

Kryptos koehleri (Locard, 1896) Figs 441, 487, 51 1-513

P/eurotomella koehleri Locard, 1896 (June): 208.

Pleurotomella koehleri Locard, 1896:137, pi. 5, fig. 1.

Kryptos elegans Jeffreys in Dautzenberg & Fischer, 1896:435, pi. 15, fig. 20.
Pleurotomella atlantica Locard, 1897:240, pi. 12, figs 1-8.

Pleurotomella demulcata Locard, 1897:243, pi. 12, figs 9-13.

Fig 511-513. Kryptos koehleri. 511, BIOGAS st CP25, 21.8 mm. 512 and 513, off Portugal, 1 1 .4 mm and
apex 1.6 mm.

196 P. BOUCHET & A. WAREN

Type material: P. koehleri ; lost; K. elegans, holotype in MNHN; P. atlantica and P. demulcata ,
syntypes in MNHN.

Type localities: P. koehleri , N of Spain, 44°02 N, 07°42 W, 1300 m; K. elegans, bathyal, W of Spain
(here designated).

Material examined: The type material and PORCUPINE st 16, 17, 17a, 24, 30, off Spain and Portugal,
500—2000 m, several shs; TRAVAILLEUR 1881, drag 1, 43°01 N, 09°37 W, 2018 m, 1 spm; drag 34, 38°18 N,
09°24 W, 1226 m, 1 spm; 1882, drag 19, 41 °32 N, 09°21 W, 1350 m, 2 shs; drag 40, 33°09 N, 09°38 W, 1340m, 2
shs; drag 70, 43°59 N, 05°35 W, 1000 m, 1 sh; TALISMAN drag 20, 33°43 N, 09°02 W, 1 105 m, 1 sh; drag 22,
33°47 N, 09°02 W, 1635 m, 1 spm; drag 3 1 , 32°40 N, 09°50 W, 1917 m, 5 spms; drag 37, 31 °34 N, 10°21 W, 900
m, 1 spm; THALASSA W415, 43°53 N, 06°11 W, 860-1150 m, 1 spm; X339, 44°10 N, 04°30 W, 1560 m, 3
spms; Y373, 41 °35 N, 09° 19 W, 1200 m, 3 shs; Y374,41°31 N,09°20W, 1250 m, 1 sh; Y395, 41 0 19 N, 09° 14 W,
810 m, 1 sh; Y401, 40°37 N, 09°22 W, 1040 m, 1 sh; POLYGAS CV16, 44° 07 N, 04° 17 W, 1909 m, 1 spm;
BIOGAS DS40, 47°36 N, 09°04 W, 3345 m, 1 spm; DS86, 44°09 N, 04° 19 W, 1950 m, 1 sh; CP07, 44° 10 N,
04° 16 W, 2170 m, 1 sh; CP23, 44°05 N, 04°21 W, 1980 m, 1 spm; CP25, 44°05 N, 04°17 W, 1894 m, 1 sh.

Distribution: The continental slope from the N parts of the Bay of Biscay to S Morocco, 800—2000
m normally.

Remarks: K. koehleri is not very variable and no species from the Atlantic can be confused with it
except young specimens of Mohnia abyssorum, which resemble the young of K. koehleri.

We have not been able to find a precise date of publication of Dautzenberg & Fischer’s (1896)
description of K, elegans, but in the same paper (page 434), they refer to Locard (1896), which
therefore must be older.

The rectum of a specimen soaked for radula preparation was filled with detritus and numerous
polychaete bristles. The animal has very small tentacles and lacks eyes.

NORTH-EAST ATLANTIC NEOGASTROPODA 197

Genus BERINGIUS Dali, 1887

Type species: Ghrysodomus crebricostaius Dali, 1877, by original designation.

Synonyms: Jumaia Friele, 1882:6. Type species Fusus turtoni Bean, 1834, by original designation.
Ukko Friele in Norman, 1893:352. New name for Jumaia Friele, 1882.

Remarks: The genus is characterized by its short, indistinct siphonal canal, small aperture and the
radula with a cuspless central and claw-like, bicuspidate lateral teeth. There is a single N Atlantic
and a few N Pacific species. B. crebricostaius is quite similar to B. turtoni in the shell characters, but
the sculpture of crebricostaius consists of very strong spiral ribs.

Friele’s name was rejected by 1CZN (Opinion 469, 1957) because it was supposed to be
blasphemous. Jumaia was a lappish name for the Christian god. At this decision the older and
available name Brongus De Gregorio, 1885 was overlooked, but we have selected a type species for
Brongus so it becomes a junior synonym of Coins to keep stability in the nomenclature.

Beringius brychius (Verrill & Smith, 1885) is here referred to Beiomitra (see that genus).

The radula of Beringius was the reason why Habe & Sato (1973) placed the genus in the subfamily
Liomesusinae (sic!). Japelion Dali, 1916 and the questionable genus Noeberingius Habe & Ito, 1965
have a. radula very similar to Beringius and are probably closely related, as the shells also indicate.
Liomesus on the contrary has quite a different shell, and the similarities in the radula are caused by
enlargement of the base of a normal bu canid lateral tooth in Beringius while in Liomesus there is
reduction of the inner cusps of the same tooth. When drawings of the teeth are compared, this
difference is not evident, but in SEM pictures it is.

Beringius turtoni (Bean, 1834) Figs 446-447, 466, 51 4-5 16, 519

Fusus turtoni Bean, 1834:493, fig. 62.

Tritonium schantaricum Middendorff, 1849:475.

Neptunea ossiania Friele, 1879:279.

Fusus turtoni var. brevispira Norman, 1893:352.

Fusus turtoni var. tumida Norman, 1893:352.

Fusus turtoni var. attenuata Simpson in Marshall, 1902:41. (See also Simpson, 1903:83.)

Type locality: Off Scarborough, North Sea.

Type material : Not known.

Material examined: About 75 specimens from all parts of the distributional area (MNHN, USNM, ZMC,
SMNH, ZMB, coll AW).

Distribution: From 80° N, between Spitzbergen and Greenland and N of Franz Joseph’s Land
(Friele, 1882, Gorbunov, 1946), on the continental shelves southwards to NE Norway (Verkruzen,
1875), along the Norwegian coast (Sars, 1878), the North Sea (numerous records), Shetland
(numerous records), the Faroes (unpublished in ZMC), Iceland (Thor son, 1941; Oskarsson, 1962),
E, S, W Greenland (Thorson, 1944; Clarke, 1974; unpublished in ZMC), Arctic Canada from 95° W
(McPherson, 1971), S to 150 miles S of Cape Race, Newfoundland (unpublished, in USNM). In
Greenland it is known from 25 m, in other places the normal upper limit is about 100 m and it is
known down to 1447 m (Golikov, 1964) although the lower limit usually is 600 m.

Remarks: Several attempts have been made to distinguish two species, B. turtoni and B. ossiania
(Friele, 1882; McPherson, 1971; Thorson, 1935). We have examined the types of B. ossiania and the
differences from the typical form in the North Sea were noticed, but most specimens we have seen
are intermediate and there is no difference in the distribution of the forms, so the names should not
even be used for subspecific separation.

198 P. BOUCHET & A. WAREN

Figs 514-516. Beringius turtoni. 514, NW of Disco, W Greenland, 202 m, 121 mm. 515 and 516, off
Korsfjorden, Bergen area, 240-250 m, 62.8 mm.

The differences in the egg capsules mentioned by Friele (1882) are probably the result of variation
because his description of the egg capsules of B. ossiania agrees very well with Jeffreys’ description
(1867) which was based on typical B. turtoni. Jeffreys described the colour of the capsules as “pale
orange’’, Friele (1882) said that the capsules of B. ossiania were “citron-yellow’’ while Thorson
(1935) said that the capsules of B. turtoni were more yellow than those of B. ossiania. Howse (1847)
figured in the capsules and described them as “pale yellow’’.

NORTH-EAST ATLANTIC NEOGASTROPODA 199

Figs 517-519. Egg capsules. 517, Unknown (buccinid?) egg capsule, WALDA st DS7 (19°57 S, 1 1°02 E,
1227 m), length 7.0 mm. Side and top views. 518, Volutopsius norwegicus, off NE United States,
diameter 32 mm. Top and side views. 519, Beringius turtoni, Dogger Bank, 75 m, diameter 28 mm. Side
views.

200 P. BOUCHET & A. WAREN

Genus VOLUTOPSIUS Morch, 1857

Type species: Fusus largillierti Petit de la Saussaye, 1851, by original designation.

Synonym: Strombella Gray, 1857:13 (not Schliiter, 1838). Type species Strombus norwegicus
Gmelin, 1790 by monotypy.

Remarks: The genus Volutopsius contains a few arctic species. Tiba & Kosuge (1979) listed the N
Pacific species. In the N Atlantic there are two species, V. norwegicus and V. (Pyrulojusus) deformis
(Reeve, 1847). V. deformis is sinistral and has lateral teeth of normal Buccinum type. V. harpa
(Morch, 1857) has often been considered a subspecies of deformis, but has the same kind of lateral
teeth as norwegicus (Habe & Sato 1973) and should be kept distinct. The variation in the lateral teeth
in this otherwise uniform group of species clearly shows that it is of little or not value above the
specific level.

Volutopsius norwegicus (Gmelin, 1790) Figs 443-444, 467, 518, 520-524

Strombus norwegicus... Chemnitz, 1788:218 (not binominal).

Strombus norwegicus Gmelin, 1790:3250.

Fusus largillierti Petit de la Saussaye, 1851:255.

Type material : S. norwegicus, lost? Not found in ZMC; F. largillierti, lost? Not in MNHN.

Type locality: S. norwegicus, Norway; F. largillierti, Newfoundland.

Material examined: About 150 shells and specimens from all parts of the range.

Distribution: From Spitzbergen, E and W Greenland southwards on the continental shelves, to the
North Sea, Shetland, The Faroes, and Iceland (Odhner, 1915; Thorson, 1944; Schlesh, 1926; Sars,
1878; Jeffreys, 1867), and from Arctic Canada, c. 95° W, to George’s Bank, 42° N (Macpherson,
1971 and unpublished material in USNM). It is restricted to the continental shelf, 25 m (E
Greenland, Thorson, 1941) to about 600 m (numerous records) except S of Iceland where it occurs
down to 2000 m (unpublished material in ZMC).

Remarks: There is a distinct cline in the size of the larval shell in V. norwegicus, W Atlantic
specimens having a larger one, but examination of a large collection in ZMC from Iceland and
Greenland proved that it is hardly possible to separate V. largillierti more than possibly as a western
subspecies, not sharply limited from the nominal species. Examination of further Siberian material
may show that V. middendorffi (Dali, 1891) belongs to the present species and it thus has a more or
less circumpolar distribution.

NORTH-EAST ATLANTIC NEOGASTROPODA 201

Figs 520-524. Volutopsius norwegicus. 520, norwegicus form, Balsfjord, Tromso area, 100 m, 85 mm.
521, largillierti form, NW Atlantic, 500 m, 92.5 mm. 522, embryonic shell from egg capsule, Shetland,
8.8 mm. 523, Shetland, 15.7 mm. 524, INGOLF st 81 (61°44 N, 27°00 W, 913 m), 30.6 mm.

202 P. BOUCHET & A. WAREN

Genus NEPTUNEA Roding, 1798

Type species: Fusus antiquus Linne, 1758, subsequent designation Sandberger (1861) and Nelson
(1976).

Synonym: Aulacofusus Dali, 1918. Type species Fusus spitzbergensis Reeve, 1855.

Remarks: Aulacofusus spitzbergensis is quite similar to Neptunea antiqua, in shell, radula and
opercular characters and differs mainly by its smaller size. We can therefore see no reason for
separating Aulacofusus, even as a subgenus.

It is not our intention to discuss the despecta - antiqua complex or the numerous arctic forms, but
we have limited our discussion to the two species that occur on the upper parts of the continental
slope in Europe and N. brevicauda which has a similar distribution in the NW Atlantic.

Neptunea brevicauda (Deshayes, 1832) Figs 404,456,528=529

Fusus brevicauda Deshayes, 1832:159.

Fusus spitzbergensis Reeve, 1855:395, pi. 32, figs 6a-b.

Tritonium schantaricum Middendorff, 1849:475.

Neptunea terebralis Gould, 1860:326 (not Lamarck, 1803).

Fusus lividus Morch, 1862:36.

Type material: F. brevicauda, holotype in MNHN; F. spitzbergensis, holotype BMNH 197654.
Type localities: F. brevicauda, not known; F. spitzbergenis , said to come from Spitzbergen.
Material examined: 20 samples with about 40 shells and specimens, all from the NW Atlantic.

Distribution: From Maine and northwards along the N American Arctic coast, to the state of
Washington in 2 — 548 m (Macpherson, 1971), Bering Sea and the Sea of Okhotsk (Middendorff,
1849). Not known from the Siberian coast, the NE Atlantic or Greenland.

Remarks: Fusus spitzbergensis was said to originate from Spitzbergen in the original description, but
no later writers on molluscs from Spitzbergen have reported it, nor is it present in any collections we
have examined from the NE Atlantic. Pain (1978) figured a specimen from “west of Spitzbergen,
400 m”, but his figure shows a specimen of Colus sabini. Therefore we suppose the species to be
absent from the N Atlantic, except the American coast. We.have therefore preferred to use the older
name of Deshayes, which does not imply an erroneous distribution.

No locality was given for Fusus brevicauda, but on the label one can read “Kamchatka” (NW
Pacific), which probably has been added later.

Probably some specific names described in various papers by Dali have to be added to the
synonymy.

N. brevicauda looks like a slender, miniature N. antiqua, but has more regular sculpture.

Neptunea antiqua (Linne, 1758) Figs 405, 476, 525-526

Murex antiquus Linne, 1758:754
Type locality: “In oras Europaeo”.

Type material: Two syntypes in the Linnean Society, London.

Material examined: THALASSA st W406, 43°55 N, 05°44 W, 700-400 m, 1 sh; st Y418, 41° 19 N, 09° 15 W, 585
m, 1 sh; st Z435, 48°39 N, 09°54 W, 720 m, 1 sh; BIOGAS CV22, 47°42 N, 08° 19 W, 1331 m, 1 sh; several
hundred specimens from the Scandinavian and British coasts, 15 — 600 m (coll. AW, USNM, ZMC, SMNH).

NORTH-EAST ATLANTIC NEOGASTROPODA 203

Distribution: Uncertain to the north and west. It is common in S Scandinavia and around the British
Isles, 15-1000 m (Seaward, 1982; Pearce & Thorson, 1967).

Remarks : In our material from deep water, there are a few empty shells from the Bay of Biscay and a
single one from off N Portugal, which agree well with the form common off the British Isles and
Scandinavia in deep water. They are all old and rather worn and probably fossil. This form can
neither be attributed to typical N. antiqua which is the form common in shallow water in the
Skagerrak, the Kattegat and the southern Baltic, nor can it be identified with typical N. despecta ,
which is a more heavily sculptured northern form. Intermediates between all these forms occur, but
each of these populations can be recognized from the shell with certainty. The relations between
these populations are not known.

Locard ( 1 886) published several records from the Channel and the Bay of Biscay, in shallow water.
We have not seen any specimens in his collection, which now is in MNHN, and doubt the validity. ■

The biology of N. antiqua is fairly well known. Pearce & Thorson (1967) summarized what is
known, and Fange (I960) reported on its poisonous salivary secretion.

Neptunea contraria (Linne, 1771)

Fig. 527

Murex contrarius Linne, 1771:551.

Fusus sinistrorsus Deshayes, 1832:160.

Fusus perversus Kiener, 1840: pi. 20, fig. 1 (name used only on the plate).

Type material: Holotype in the Linnean Society, London.

Type locality: Vigo Bay, N Spain (here designated). See also Rolan (1983:240).

Material examined: TALISMAN 1883 dr 11, 35°21 N, 07°05 W, 1084 m, 1 spm; dr 18, 33°33 N, 08°59 W, 550
m, 2 shs; THALASSA W400, 43°48 N, 05° 10 W, 350-200 m, 3 spms; W405, 43°56 N, 05°44 W, 690-400 m, 1
sh; W426, 44°08 N, 08°24 W, 337-215 m, 1 spm; ¥377, 41°32 N, 09° 14 W, 320 m, 1 sh; Fosse de Cap Breton,
Bay of Biscay, 1 sh; a few dozen shells and specimens with less precise localities: Vigo Bay, Bay of Biscay, Spain
etc. (MNHN, USNM, ZMC).

Distribution: Along the shelf and upper continental slope from the southern parts of the Bay of
Biscay to Morocco (Locard, 1897) in 100—1000 m. Mediterranean records seem to be based on fossil
specimens (Barsotti & Frill i, 1970).

Remarks: There have been numerous discussions in the literature whether M. contrarius was based
on the normally sinistral S European species or on a sinistral specimen of N, antiqua. Examination
of the holotype proved that the name was based on the S European species, as Hanley (1855)
concluded.

Dautzenberg (1910) recorded N. contraria from Cap Blanc, Mauretania, without any comments
about the depth. We have examined this specimen in IRSN, and although the shell looks very fresh,
the label says that it was collected at 32 m above sea level in a quaternary deposit.

Neptunea contraria resembles sinistral specimens of N. antiqua , which occasionally occur, but can
be separated by having a sculpture of coarser but more equal sized spiral ribs, that are rounded,
never sharp.

Some details on the anatomy of N . contraria were given by Fischer & Bouvier (1890).

Golikov (1963) regarded N. contraria as a subspecies of N. antiqua but we cannot agree. The fossil
history of N. contraria goes back to the Pliocene, as does that of N. antiqua , and there has never
been any mixing of them (cf. Harmer, 1914 and Strauch, 1972).

204 P. BOUCHET & A. WAREN

Figs 525-529. Genus Neptunea. 525, N. antiqua , Skagerrack, 155 mm. 526, Great Britain (Jeffreys coll.),
sinistral specimen, 88.3 mm. 527, N. contraria, off Vigo, NW Spain, 92.5 mm. 528, N. brevicauda, Egg
Harbor, Labrador, 13 m, 27.7 mm. 529, Newfoundland Banks (45°38 N, 55°00 W), 52.0 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 205

Genus MOHNIA Friele in Kobelt, 1878

Type species: Fusus mohni Friele, 1877, by monotypy.

Remarks: In Mohnia we have included a number of bathyal and abyssal buccinids that have a more
or less distinctly spirally-coiled operculum and a small initial whorl in the larval shell that increases
its diameter regularly (Figs 534, 537, 540, 543). The radula has a more or less square central tooth
with one or three cusps and lateral teeth with two major and 0-2 minor cusps between these.

All species we know of Mohnia are true deep-water species. They are known from the N Atlantic,
the N Pacific, and the Arctic basins.

In some species the spiral coiling of the operculum can be observed only in young specimens
because its older parts get worn rather early in life.

We have divided Mohnia in two subgenera, Mohnia s.str. and Tacita. The species of Tacita have a
more or less distinct axial sculpture in addition to spiral lines and lose the spirally-coiled part of the
operculum early in life. The species of Mohnia laek axial sculpture and usually the operculum keeps
the apical part throughout life.

In Mohnia we have also included some species for which we could not find a better genus,
although we are not satisfied with their position here, viz. M. carolinensis, M. caelatus and M.
glyptus.

The species referred to Mohnia can be determined by the following key:

A.

Only spiral sculpture present, suture not very deep

B (Mohnia s.str.7

A.

Axial sculpture usually present

D

B.

Three or four major spiral ribs on the upper whorls

M. mohni

B.

Several to many spiral ribs or lines present

C

C.

Spiral ribs of unequal size, separated by broader interstices

M. parvus

C.

Numerous fine spiral ribs of equal size, separated by narrow

interstices

M. simplex

D.

Whorls with a distinct shoulder

M. (T.) blakei

D.

Whorls evenly rounded

E

E.

Broad, rounded axial ribs present on all whorls

F

E.

Axial sculpture usually present only on the upper whorls

H

F.

Spiral sculpture forms knobs at the intersections with the axial ribs

M. carolinensis

F.

Spiral sculpture disappears on the axial ribs

G

G.

Columella distinctly curved

M. caelatus

G.

Columella almost straight

M. glyptus

H.

Axial sculpture of strong short ribs on the upper whorls

M. (T.) abyssorum

H.

Axial sculpture of rather indistinct, broad growth lines

I

I.

Spiral lines sharp

M (T.) krampi

I.

Spiral lines low and rounded

M (T.) danielsseni

Mohnia (Mohnia) mohni (Friele, 1877)

Figs 435,482,530-531

Fusus mohni Friele, 1877:6.

Fusus concinnus Jeffreys, 1883:396, pi. 44, fig. 8.

Type material: F. mohni in ZMB, not seen; F. concinnus , holotype BMNH 85.11.5.4576.

Type localities: F. mohni N. of the Faroes, 1000-2000 m; F. concinnus , 60°05 N, 06°21 W, S of the
Faroes, 1 100 m.

206 P. BOUCHET & A. WAREN

Material examined: 3000 shs and spms, listed by Bouchet & Waren (1979) and from INGOLF st 104, 105, 107,
110, 111, 112, 113, 116, 117, 118, 119, 120, 124, 125 and 139, W and NW of Iceland, 650-2300 m; 81° 14 N,
22°50 E, 150 m, 1 sh, SMNH.

Distribution: The Norwegian and Arctic abyssal basins (Bouchet & Waren, 1979; Lus, 1981), in
650-3800 m.

Remarks: M. mohni is only likely to get confused with M. danielsseni, with which it occurs together,

but it can easily be separated by having the apical whorls of half the diameter of M. danielsseni (Figs
531, 540).

Some details on the reproduction were given by Bouchet & Waren (1979). It can be added that the
capsules are not attached to some firm substratum as are those of M. danielsseni.

The specific names Mohnia alba and Fusus tener which have been mentioned in the literature, are
manuscript names of Friele and Jeffreys. Kobelt (1878) used the name Mohnia alba on plate 9 (fig.
5), but in the text he said that this name should not be used.

Mohnia (Mohnia) parva (Verrill & Smith, 1882) Figs 436, 451, 532-534

Sip ho parvus Verrill & Smith in Verrill, 1882:504, pi. 57, fig. 20.

Type material: Holotype USNM 38013.

Type locality: LJSFC st 997, off Delaware, 600 m.

Material examined: Numerous shells and specimens, USFC stns 925, 937, 947, 1030, 1093, 1154, 2048, 2076,
2183, 2203, 2486, 2547, 2586, 2687 and George’s Bank, 200 m (USNM); THOR 1902, st 164, SW of Iceland ca.
1900 m, 2 shs; INGOLF st 10, 64°24 N, 28°50 W, 1484 m, 1 sh; 74° 17 N, 15°20 W, 200 m, 4 shs.

Distribution: NE America from 39° N, to SW Iceland and NE Greenland, 200 — 2000 m.

Remarks: This species is characterized by its uneven sculpture, which in worn specimens has an

additional incised line on each spiral rib.

The specimens from NE Greenland were determined dalli by Posselt (1895) and formed the base
for the only record from Greenland. Thus dalli can be cancelled from the list of gastropods from
Greenland.

Mohnia (Mohnia) simplex (Verrill, 1884) Figs 437-438, 452, 535-537

Sipho (Mohnia) simplex Verrill, 1884:174.

Type material: Holotype USNM 35573.

Type locality: USFC st 2115, off N Carolina, 1500 m.

Material examined: The holotype and USFC st 2470, 44°47 N, 56°34 W, 440 m, 1 spm; USFC st 54B, 42°49 N,
68°48 W, 200 m, 2 shs; Lindenows Fjord, SE Greenland, 400—600 m, 1 sh, ZMC; INGOLF st 35, 65° 16 N,
55°05 W, 682 m, 5 spms.

Distribution: On the upper continental slope, from N Carolina to S Greenland, only known from the
material examined.

Remarks: The opercula from the specimens from USFC st 2470 and INGOLF st 35 show only traces
of being paucispiral. Verrill described it as “distinctly but less than M. mohni paucispiral”. No
original material with soft parts was available to check this, but a specimen in the Locard collection,
without precise locality data, had a more distinctly coiled operculum, corresponding to the
description. This may indicate that more than one species is involved, but we find it more likely that
it depends on intraspecific variation.

One of the specimens from the INGOLF expedition had a large parasitic copepod in the oviduct.

NORTH-EAST ATLANTIC NEOGASTROPODA 207

Figs 530-534. Genus Mohnia. 530 and 53 1, M. mohni, NORBI st DS18 (73°36N, 13°35 W, 2470 m), 23.6
mm and apex 1 .7 mm. 532, M. parva, 74° 17 N, 15°20 W, 234 m, 14.2 mm. 533, M. parva off Martha’s
Vineyard, 12.8 mm. 534, M. parva, S of Iceland, apex 1 .5 mm.

208 P. BOUCHET & A. WAREN

Figs 535-540. Genus Mohnia. 535, M. simplex, INGOLF st 35, 20.1 mm. 536, NE America, 202 m, 9.9
mm. 537, off Nova Scotia, 412 m, apex 1.4 mm. 538, M. krampi, syntype, 34.5 mm. 539 and 540, M.
danielsseni, NORBI st CP17 (73°31 N, 13°40 W, 2500 m), 42.7 mm and apex 2.1 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 209

Figs 541-545. Genus Mohnia. 541, M. abyssorum, NORATLANTE st B17 (45°13 N, 05°31 W, 4700 m),
43.5 mm. 542 and 543, INGOLF st 36 (61 °50 N, 56°21 W, 2702 m), 5.4 mm. 544 and 545, M. blakei, off
Maryland, 3360 m, 14.9 mm and apex 2.3 mm.

210 P. BOUCHET & A. WAREN

Subgenus TACITA Lus, 1971

Type species: T. holoserica Lus, 1971, by original designation.

Remarks: The type species was described from the Kuril Trench, in 6100 m. Its anatomy was
described in the original description. The shell, operculum and radula resemble rather closely those
of M. abyssorum and M. danielsseni. Lus introduced the genus after comparison with Neptunea and
Sipho and did not give any distinctive characters in relation to Mohnia. From shell morphology,
radular characters and operculum we find this genus very close to Mohnia, but keep it as a subgenus
for a number of species that have a less distinctly spirally-coiled operculum, usually are of a larger
size and often have some axial sculpture.

Mohnia (Tacita) danielsseni (Friele, 1879) Figs 431-432, 480, 539-540

Type material: M. danielsseni, holotype in ZMB.

Type locality: M. danielsseni, The abyssal area between Norway, Spitzbergen and Greenland; C.
hunkinsi, the abyssal of the North Canadian Basin.

Neptunea (Sipho) danielsseni Friele, 1879:282.

Co/us hunkinsi Clarke, 1960:3, pi. 1, fig. 9.

Material examined: The material enumerated by Bouchet & Waren (1979) and numerous shells and specimens,
INGOLF st 112, 113, NW of Iceland, 2400 m; 77°52 N, 03°05 W, 2750 m, 15 shs, SMNH; Recherche Bay,
Spitzbergen, 90 m(7), 3 shs, SMNH.

Distribution: The abyssal parts of the Norwegian and Arctic Basins 1400-3700 m.

Remarks: Some information about the biology was given by Bouchet & Waren (1979). For
differences from M. mohni, see that species. M. krampi resembles M. danielsseni, but has sharper
spiral ridges, flatter whorls and a more narrow aperture.

Lus (1981) placed M. danielsseni in Colicryptus, which she regarded as a subgenus of Coins .
Colicryptus is here considered a synonym of Turrisipho and is characterized by an apical, pointed
nucleus of the operculum (Fig. 407). Therefore we prefer to keep danielsseni in Mohnia.

Mohnia (Tacita) blakei (Verrill, 1885)

Figs 439,450, 544-545.

Bela blakei Verrill, 1885:417, pi. 44, fig. 8.

Type material: Holotype USNM 44655.

Type locality: USFC st 2226, 37°00 N, 71°54 W, 3850 m.

Material examined: The type material and USFC stn 2714, off Maryland, 4 spms, 3330 m.

Distribution: Only known from the material examined.

Remarks: This species has always been regarded as a turrid but the radula (Fig. 450) clearly shows
that it is a buccinid. For differences from M. abyssorum, see under that species.

NORTH-EAST ATLANTIC NEOGASTROPODA 211

Mohnia (Tacita) abyssorum (Fischer, 1883) Fig 426-427, 481, 541-543, 546-547.
Fusus abyssorum Fischer, 1883:391.

Sip ho hispid ulus Verrill, 1884:239.

Sipho profundicola Verrill & Smith, 1884: 170, pi. 31, fig. 13.

Sipho profundicola var. dispar Verrill, 1884: 171.

Type material: Lectotype of F. abyssorum (here selected) in MNHN; S. profundicola holotype
USNM 37999; 5. profundicola var. dispar holotype USNM 37955; S. hispidulus holotype USNM
34840.

Type localities: F. abyssorum, TALISMAN st 131, 38°38 N, 25°06 W, 2995 m; S. profundicola,
USFC st 2037, 38°53 N, 69°23 W, 3300 m; 5. profundicola var. dispar, USFC st 2042, 39°33 N,
68°26 W, 2800 m; 5. hispidulus, USFC st 2038, 38°30 N, 69°08 W, 3650 m.

Material examined: 25 samples, mainly USFC material, with about 50 specimens from off Virginia, to George’s
Bank, S of Newfoundland, material in USNM; INGOLF st 20, 58°20 N, 40°48 W, 3100 m, 1 sh; st 37, 60° 17 N,
54°05 W, 3100 m, 1 spm; 95 samples with about 640 shells and specimens from TALISMAN 1883,
NORATLANTE, BIACORES, THALASSA, BIOGAS, INCAL, and MONACO Expeditions, mainly in
MNHN, a few in MOM and IRSN. The extreme S distribution is TALISMAN st 106 (not 100 as cited by
Locard), 15°48 N, 20°23 W, 3655 m; the SW extreme is TALISMAN st 1 17, 32° 19 N, 35°44 W, 3432 m and 118,
34°46 N, 33°51 W, 3175 m. The deepest material is from MONACO expeditions st 2994, 44°08 N, 10°44 W,
5000 m. Four samples come from depths between 1900 and 2100 m; the remaining samples are deeper.

Distribution: The lowermost part of the continental slope and the upper part of the abyssal area of
the Atlantic, S to about 15° N.

Remarks: This is one of the most characteristic species of the upper abyssal area of the N Atlantic. It
is represented by numerous samples from 2500 m— 4800 m but only four more shallow and a single
below 4850 m. The S distribution is uncertain because of lack of sampling stations, but it is not
present in material from the Verna fracture zone, about 11° N, 5100-5900 m.

Young specimens of M. abyssorum resemble M. blakei slightly but M. blakei has more narrow
axial ribs and a larger larval shell. Young specimens also resemble Kryptos koehleri of the same size
but are broader and increase their diameter more rapidly (Figs 513, 542).

Several egg capsules were present in our material. The capsule is ovate, 5 mm long and about 3
mm in diameter. It is attached to the substratum by a stalk, Y-shaped in cross section and 1.5 mm
high, originating centrally from the longer side of the capsule. The capsule is filled by a
homogeneous yolk. In two capsules there was a very young embryo in the veliger stage, in a third
capsule (Fig. 546) solid yolk; in two capsules there were young almost ready to hatch.

Figs 546-547. Mohnia abyssorum. 546, BIOGAS st DS87 (44°05 N, 04° 19 W, 1950 m), egg capsule, 5.1
mm. 547, INCAL st DS 10 (50°13 N, 13°17 W, 2719 m), embryo, 5.1 mm.

212 P. BOUCHET & A. WAREN

Mohnia (Tacita) krampi (Thorson, 1951) Figs 425, 449, 538

Sipho krampi Thorson, 1951: 54, fig. 16.

Type material: Holotype and 7 paratypes in ZMC.

Type locality: 69°50 N, 61°36 W, 1880 m, Baffin Bay.

Material examined: The type material.

Distribution: Clarke (1974) recorded A/, krampi from 570-2300 m, (4 stations) in Baffin Bay. It has
not been taken outside the deep parts of Baffin Bay.

Remarks: This species resembles M. danielsseni rather closely but is more slender and its whorls are
higher.

All material we have examined is badly corroded and the larval shell is not known.

Mohnia carolinensis (Verrill, 1884) Figs 434, 457, 555-556

Urosalpinx carolinensis Verrill, 1884:237.

Mohnia carolinensis , Radwin 1972:339.

Type material: Numerous syntypes USNM 35735.

Type locality: USFC st 2109, off Cape Hatteras, North Carolina, 250 m.

Material examined: The material examined by Verrill (1884).

Distribution: Only known from off North Carolina, 250-1700 m.

Remarks: A7. carolinensis was described as as muricid, but was referred to Mohnia by Radwin
(1972), who examined the radula. We have examined the operculum of a young specimen, 4 mm
high, and it is slightly paucispiral. The shell, however, differs widely from Mohnia , in being solid
with strong axial sculpture and it will probably prove to be more closely related to certain tropical
buccinids from deep water that now are considered to be fusinids.

The only species that M. carolinensis can be confused with is M. caelatus, which differs from M.
carolinensis in having a weaker spiral sculpture, where the spiral lines disappear on the axial ribs.

Mohnia caelata (Verrill & Smith, 1880) Figs 433, 454, 548-551

Neptunea caelata Verrill & Smith in Verrill, 1880:369.

Sipho caelatus, Verrill 1882:506, pi. 57, fig. 19.

Sipho obesus Verrill, 1884:168.

Sipho caelatus var. hebes Verrill, 1884:172.

Sipho caelatulus Verrill, 1884: 172.

Type material: N. caelata, syntypes USNM 38026; 5. obesus, holotype and several syntypes USNM
35600; var. hebes, lectotype (here selected) USNM 35424; S. coelatulus, lectotype (here selected)
USNM 35226.

Type localities: N. caelata USFC st 894, off Pennsylvania, 700 m; S. obesus USFC st 2115, off
North Carolina, 1500 m; var. hebes USFC st 2103 off Delaware 1800 m; S. coelatulus USFC st
2076, off New York, 1700 m.

Material examined: The types and numerous specimens from 79 USFC stations and INGOLF st 25, 60°30 N,
54°25 W, 1065 m, 6 spms; 62°57 N, 19°58 W, 957 m, 12 shs, ZMC.

Distribution: along the upper part of the continental slope, from North Carolina to S of Iceland,
1000-2500 m.

Remarks: Comparison of the rich material of the species proved that all intermediate forms exist,
although there is a tendency to form uniform populations. The name obesus was based on a juvenile
specimen, var. hebes on an unusually broad specimen.

For differences from M. carolinensis, see under that species.

The operculum is slightly paucispiral.

M. caelata is quite similar to M. frielei (Dali, 1891)andM. robusta ( Dali, 1913) from deep water in
the N Pacific.

NORTH-EAST ATLANTIC NEOGASTROPODA 213

Mohnia glyptus (Verrill, 1882) Figs 430, 453, 552-554

Sipho glyptus Verrill, 1882:505, pi. 52, fig. 22, pi. 53, fig. 1.

Sip ho (Siphonorbis) iindahii Posselt, 1898:180, ph 1, fig. 5.

Type material: S. glyptus holotype USNM 38005; S. Iindahii holotype SMNH 1566.

Type localities: S. glyptus USFC st 895, off New Jersey, 450 m; S. Iindahii Umanak, W. Greenland,
270m.

Material examined: The type material and USFC st 1154, off Martha’s Vineyard, 350 m, 4 shs; 62°57 N, 19°58
W, 957 m, 4 shs, ZMC; 63°05 N, 20°07 W, 957 m, 1 sh, ZMC; INGOLF st 25, 60° 30 N, 54°25 W, 1096 m, 1 sh;
st 27, 64° 54 N, 55°10 W, 740 m, 1 sh; st 28, 65°14 N, 55°42 W, 791 m, 5 spms; st 32, 65°35 N, 56°38 W, 599 m,
14 spms; Umanak, W Greenland, 740 m, 1 spm, SMNH.

Distribution: From off New Jersey, along the upper continental slope, northwards to S of Iceland,
c. 300-1000 m. Only known from the material examined.

Remarks: The shell of M. glyptus does not resemble Mohnia, especially in the characters of the
larval shell, but we have not found any genus that resembles this species and we have not wanted to
describe a new genus for it.

The shell of M. glyptus resembles that of Metzgeria, but species of Metzgeria have distinct
columellar folds.

The larval shell of M. glyptus is remarkable (Fig. 554) because of its irregular shape and it does not
resemble that of any buccinid known to us. It is, however, a good character for recognizing this
species.

214 P. BOUCHET & A. WAREN

Figs 548-554. Genus Mohnia. 548, M. caelata, SE Delaware bay, 1492 m, 17.6 mm. 549, THOR st
166 (62°57 N, 19°58 W, 957 m), 9.5 mm. 550, INGOLF st 25, 11.4 mm. 551, INGOLF st 10 (64°24
N, 28°50 W, 1484 m), apex 1.2 mm. 552, M. glyptus, off Martha’s Vineyard, 355 m, 15.8 mm. 553
and 554, INGOLF, st 32, 17.4 mm and apex 1.4 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 215

Genus TURRISIPHO Dautzenberg & Fischer, 1912
Type species : Fusus lachesis Morch, 1869, by original designation.

Synonyms: Colicryptus Iredale, 1918. Type species Fusus fenestratus Turton, 1834, by original
designation.

Undacolus Nordsieck, 1968. Type species Neptunea undulata Friele, 1882, by original designation.
Remarks: The species of Turrisipho are characterized by having the nucleus of the operculum drawn
out into a small apical point, a squarish central tooth of the radula with 1-3 cusps, a distinct, hairy
periostracum and a high slender shell with numerous whorls.

The species here included in Turrisipho have previously been placed in a number of different
genera, and Lus (1981) referred Mohnia danielsseni to Colicryptus. We find, however, the group
here put together to be a very uniform group, especially when young specimens are compared. The
pointed apical part of the operculum is especially characteristic and we have not seen anything

similar in other buccinids.

The species of Turrisipho can be separated by the following key:

A. Aperture constitutes 0.4 of the total height of the shell or less B

A. Aperture constitutes more than 0.4 of the total height C

B. Diameter of third whorl c. 4 mm T. voeringi

B. Diameter of third whorl c. 2.5 mm T. lachesis

C. Siphonal canal well developed, occupying 1/3 of the height of the aperture T. dalli

C. Siphonal canal short and broad D

D. Distinct, broad axial ribs present T. fenestratus

D. No axial ribs j. moebii

Turrisipho dalli (Friele, 1881) Figs. 557-558

Sipho dalli Friele in Tryon, 1881:133 (Not Fusus ( Neptunea ) dalli Watson, 1882:379, renamed Fusus (Sipho)
liberatus Watson, 1886:206).

Type material: Holotype, ZMB 20644.

Type locality: Between Bear Island and Norway, 350 m.

Material examined: PORCUPINE 1869, 61 °21 N,03°44W, 1160 m, 1 sh; MONACO st 922, 58° 16 N, 05°48
W, 343 m, 1 spm; Varangerfjord, N Norway (G.O. Sars leg., ZMO), 1 sh; off Korsfjorden, W Norway,
250-325 m, 1 spm, 10 shs, coll. A.W.; the material listed by Friele (1882).

Distribution: From Barents Sea, along the Scandinavian shelves, S to the Shetland — Faroe Channel
and E of Iceland (Oskarsson, 1977).

Remarks: Live specimens of T. dalli are easily recognized by the greyish shell with rows of
periostracal bristles along the spiral keels. When dead, it is well characterized by the rounded,
slightly-shouldered whorls, polygyrate larval shell, its slender siphonal canal and the rather sharp
axial sculpture of incremental lines and alternating spiral keels and smaller ribs. On the shoulder, the
spiral ribs are of equal strength.

T. dalli was recorded from NE Greenland by Posselt (1895), but examination of the material
proved that the record was based on Mohnia parva, and the species thus is unknown from
Greenland.

We have not seen opercula of very young specimens, but shells of young specimens are very
similar to those of other species of Turrisipho.

Turrisipho lachesis (Morch, 1 869) Figs 409, 458, 560-566

Fusus (Siphonorbis) lachesis Morch, 1869:397.

Sipho undulatus Friele in Tryon, 1881:133.

Sipho (Tritono fusus) costiferus Posselt, 1898:183, pi. 1, fig. 6.

Type material: F. lachesis, holotype in ZMC; S. undulatus holotype ZMB 20656; S. costiferus,
holotype SMNH 1567.

Type locality: F. lachesis, Ikersak, W Greenland, 144 m; 5. undulatus, NW of N Cape, N Norway,
390 m; S. costiferus, off Julianehaab, W Greenland, 310 m.

216 P. BOUCHET & A. WAREN

Material examined: The types and MONACO st 161, 46°05 N, 46°42 W, 1267 m, 7 spms; 73°03 N, 18°30 E, 410
m, 1 sh, SMNH: Lindenowsfjord, SW Greenland, 125-150 m, 1 spm, ZMC; Lindenowsfjord, SW Greenland,
400-600 m, 1 sh, ZMC; 66° 18 N, 18°36 W, 360 m, 1 spm, ZMC; TJALFE st 366, 66°22 N, 57° 16 W, 686 m, 1
spm; INGOLF st 10, 64°24 N, 28°50 W, 1484 m, 1 spm; st 25, 63° 30 N, 54°25 W, 1096 m, 2 spms; st 28, 65° 14
N, 55°42 W, 791 m, 25 spms; st 32, 66°35 N, 56°38 W, 599 m, 1 spm; st 35, 65° 16 N, 55°05 W, 682 m, lsh; st
138, 66°26 N, 07°56 W, 887 m, 1 spm; the material listed by Frieie (1882).

Distribution: N and W of Norway, 200-1100 m (Frieie, 1882, as lachesis, typical form and undulata),
E and NW of the Faroes (Sykes, 1911 and ZMC) 500-900 m, N and NW of Iceland 200-1500 m
(Thorson, 1941 and ZMC), SE Greenland, 400-600 m(Thorson, 1944 as lachesis smooth variety),
SW Greenland and Davis Strait, 600-1100 m (Thorson, 1940 as undulatus, INGOLF & TJALFE,
ZMC) and off Newfoundland, 1267 m (Dautzenberg & Fischer, 1912 as undulatus ).

Remarks: Morch’s original description of lachesis was based on a very poor specimen (Fig. 565) and
this led to much confusion. Frieie (1882) and G.O. Sars (1878) interpreted it correctly, but after
Frieie’ s description and illustration of var. bicarinata (see voeringi) later authors used the name
lachesis for the “variety”.

Thorson (1940) described the egg capsules and larval development of what he considered to be
Sipho undulatus but we doubt his identification. To give an impression of a higher certainty in his
identification of the capsules on which his description was based, he mentioned that S. undulatus
was taken at two adjacent INGOLF stations, nos 25 and 28, but he did not mention that they
originated from depths of 1096 and 791 m, while the egg capsules came from 2260 m, 700 m deeper
than any of the about 20 records we know of. There is also a difference in the sculpture of his
drawing of the larva and actual specimens of T. lachesis. His drawing does not show the faint, but
distinct shoulder of the body whorl of the larva which is well shown in his drawing of the apical
whorls of an adult specimen. We have not been able to find his specimens in ZMC and are not able
to say on which species his description was based.

Some specimens of T. voeringi resemble lachesis considerably, but can always be separated by
their proportionally shorter siphonal canal, more convex whorls of more slowly increasing diameter,
and the diameter of the third whorl is about 4 mm in T. voeringi , while it is 2.5 mm in lachesis. T.
fenestratus can be separated by its broader and more solid shell of a larger apical angle and shorter
siphonal canal. T. dalli has a much stronger spiral sculpture and T. moebii has more rapidly growing
diameter of the apical whorls and a much broader shape.

Turrisipho voeringi nom.nov. Figs 410-41 1 , 567-570

Neptunea (Siphonorbis) lachesis var. bicarinata Frieie, 1879:282.

(Not Neptunea despecta var. bicarinata Kobelt, 1878.)

Type material: Holotype, ZMB 26054.

Type locality: Norwegian N Atlantic Expedition st 312, S of Spitzbergen, 1170 m.

Material examined: The types and INGOLF st 101, 66°23 N, 12°05 W, 1011 m, 1 spm; st 105, 65°34 N, 07°31
W, 1435 m, 4 spms; st 116, 70°05 N, 08°26 W, 699 m, 6 spms; st 126, 67° 19 N, 15°52 W, 552 m, 2 spms;
Dusenfjord, E Greenland, 240 m, 8 spms, ZMC; Franz Joseph Fjord, E Greenland, 320 m, 1 sh, ZMC; Hurry
Inlet, E Greenland, 90 m, 3 spms, ZMC; Forsblads Fjord, E Greenland, 90-160 m, 1 spm, ZMC; Lindenows
Fjord, E Greenland, 200-350 m, 2 spms, ZMC; Ella Island, E Greenland, 48-51 m, 2 spms, ZMC; Kap Hooker,
Jameson Land, E Greenland, 150 m, 2 egg capsules, ZMC; off Cap Powlet, W Greenland, 1000-600 m, 2 spms,
ZMC; 75°54 N, 81 °00 W, 600 m, 1 spm, ZMC; 75°26 N, 62°26 W, 820 m, 1 spm, ZMC; 63°36 N, 55° 15 W,
1200 m, 1 spm, ZMC; 73°20N,21°21 W,70m, 1 spm, ZMC;66°02N, 1 1 °05 W, 900-1040 m, 1 spm, ZMC;
M. SARS 1902, st 35 & 96, Faroes, 540-1000 m, 5 spms, ZMC; 72° 10 N, 20°37 E, 550-600 m, 1 sh, SMNH.

Distribution: S of Spitzbergen 1170 m (Frieie, 1882), the Kara Sea (Thorson, 1941), the Shetland-
Faroe Channel 700-1000 m (Jeffreys, 1883; Sykes, 1911), N and E of the Faroes 550-1100 m
(material in ZMC), Jan Mayen (material in MNHN), E, N and NW of Iceland 500-1500 m (Thorson,
1941 and material in ZMC), E and W Greenland 25-1200 m (Thorson, 1944, 1951, material in ZMC)
(all listed as lachesis)..

Remarks: The present species was well described and figured by Frieie (1882) under the name
lachesis var. bicarinata , a name that cannot be kept because of Kobelt’s earlier use of it. Later
authors have used the name lachesis in that sense. We consider lachesis and var. bicarinata two

NORTH-EAST ATLANTIC NEOGASTROPODA 217

distinct species, because we have never seen any transitional specimens and the two forms can always
be separated by the characters mentioned under lachesis ; it therefore has to be given a new name;

Turrisipho fenestratus (Turton, 1834) Figs 406, 483, 571-574, 591

Buccinum fusiforme Broderip, 1830:45 (not Borson, 1822).

Fusus fenestratus Turton, 1834:351.

Neptunia peregra Locard, 1897:371, pi. 18, figs. 8-1 1 .

Neptunia ecaudis Locard, 1897:368, pi. 18, figs. 5-7.

Neptunia fusiformis vars gracilis & minor Locard, 1897:370.

Type material: F. fenestratus, holotype USNM 192347; Locard’s species, syntypes in MNHN.
Type localities: F. fenestratus, Cork, Ireland; N. ecaudis & peregra, TRAVAILLEUR 1882, dr 2,
44° 05 N, 05°36 W, 608 m.

Material examined: The types and about 100 specimens from all parts of the distribution area.

Distribution: From Angmagsalik (SE Greenland), SW and S of Iceland (Thorson, 1944, 1941;
Clarke, 1974; material in ZMC), to Lofoten (N Norway) and southwards to W Norway (G.O. Sars,
1878), the Faroes (material in ZMC), W of the British Isles (Seaward, 1982; Massy, 1930), Bay of
Biscay (Dautzenberg, 1927; THALASSA, NORATLANTE, BIOGAS), W of Spain (Sykes, 1911)
and off Morocco (35° N) (Locard, 1897). It does not occur in the Mediterranean. The depth range is
from 50 m in the N and 400 m in the S, down to 1200 m.

Remarks: T. fenestratus is well characterized by its solid shell and short siphonal canal. For
differences from T. lachesis, see that species.

Two specimens from the Faroes (432m, in ZMC) have been found in stomachs of Anarrhichas
latrifrons (Pisces).

A single egg capsule of T. fenestratus was found 26 Jan 1971 in the S part of Hjeltfjord, W
Norway in 90-120 m. It has a diameter of 5.5 mm, a tough, rather thick, leathery wall, it is yellowish
brown and spherical with a well-defined attachment area covered with sand grains. It contained a
single embryo of 3 whorls and several groups of cells, that evidently were used as nourishment.

Turrisipho moebii (Dunker & Metzger, 1874) Figs 407-408, 455, 575-578

? Fusus sarsi Jeffreys, 1869:168 (nom. nud.) (not Trophon sarsi Jeffreys in Wood, 1872:25).

Tritonofusus moebii Dunker & Metzger, 1874:8.

Fusus moebii Dunker & Metzger, 1875:260, pi. 6, fig. 1.

Fusus ebur, Friele 1877:7 and most later authors, not Fusus ebur Morch, 1869.

Sipho sarsi Jeffreys in G.O. Sars, 1878:275, pi. 15, fig. 2.

Type material: F. moebii, not found, not in ZMHU.

Type locality: F. moebii, between Lindesnes and Listerfjord, S Norway, 195 m.

Material examined: About 40 shells and specimens from all parts of the distribution area.

Distribution: From between Bear Island and Norway, to NE Norway and along the Norwegian coast
into the Skagerrak (Friele, 1882; G.O. Sars, 1878; material in ZMC) and from about 27° W, off
Iceland, along S Iceland (Thorson, 1941; material in ZMC), the Faroes (material in ZMC and Sykes,
1911) and off Ireland (Massy, 1930). The depth range is 200 — 957 m.

Remarks: The nomenclatorial history of the present species is very confused. The main source of this
confusion was Morch’s description of Fusus ebur from W Greenland (= sabini Gray). That species
often has quite convex whorls and other malacologists did not recognize it from the description but
believed the description to be based on T. moebii. This mistake can also be partly blamed on the
poor condition of Morch’s specimen. The manuscript name of Jeffreys (sarsi) that was validated by
G.O. Sars (1878) has been in common use, but is preoccupied by Wood’s use of the name for a
British Pleistocene fossil (secondary homonym).

One of the specimens from the Faroes (in ZMC), from 400 m, had been taken from the stomach of
Anarrhichas lupus (Pisces).

T. moebii can be recognized from its convex whorls, deep suture, hairy periostracum (greenish
brown in fresh specimens), regular sculpture and multispiral larval shell.

218 P. BOUCHET & A. WAREN

Figs 555-559. Genera Mohnia and Turrisipho. 555 and 556, M. carolinensis, holotype, 1 1.0 mm and apex
1.2 mm. 557 and 558, T. dalli, off Korsfjorden, Bergen area, 300-330 m, 25.0 mm and 20.3 mm. 559,
PORCUPINE 1869 st 65 (61° 10 N, 02°21 W, 635 m), apex 1.8 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 219

Figs 560-565. Turrisipho lachesis. 560, INGOLF st 25, 59.5 mm. 561, no loc. data, 20.4 mm. 562,
INGOLF st 28, 9.5 mm. 563 and 564-holotype of Sipho undulatus, VORINGEN st 290, 18.2 mm and
apex 3.0 mm. 565, holotype of Fusus lachesis, W Greenland, 144 m, 40.3 mm.

220 P. BOUCHET & A. WAREN

Figs 566-570. Genus Turrisipho. 566, T. lachesis, INGOLF st 28, 46.8 mm. 567, T. voeringi, INGOLF st
105, 77. 1 mm. 568, T. voeringi, VORINGEN st 3 12 (74°54 N, 14°53 E, 1203 m), 53.3 mm. 569 and 570,
T. voeringi, 65°34 N, 07°31 W, 1435 m, apex 6.0 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 221

Figs 571-574. Turrisipho fenestratus. 571, Korsfjorden, Bergen area, 130-150 m, 41 mm. 572, same
locality, 75-100 m, 23.0 mm. 573, same locality, 55 m, 11.2 mm. 574, same locality 440-500 m, apex 2
mm.

222 P. BOUCHET & A. WAREN

Figs 575-578. Turrisipho moebii. 575, Skagerrak, 48.6 mm. 576, SSE of Reykjanes, Iceland, 216-326 m
80 mm. 577 and 578, outer pant of Korsfjorden, Bergen area, 340-420 m, 14.0 mm and apex 2 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 223

Genus BELOMITRA Fischer, 1882

Type species: B. paradoxa Fischer, 1882, by monotypy (= B. quadruplex (Watson)).

Synonyms: Pleurobela Monterosato in Locard, 1897. Type species P. speita Monterosato in Locard,
1897, by monotypy.

Bathyclionella Kobelt, 1905. Type species Pleurotoma quadruplex Watson, 1881, by monotypy.
Cryptomitra Dali, 1924. Type species by original designation Pleurotomella climacella Dali, 1895.
Morrisonella Bartsch, 1945. Type species by original designation Leucosyrinx pacifica Dali, 1908.
Dellina Beu, 1970. Type species by original designation Waipaoa munida Ponder, 1962.

Remarks: Belomitra has always been considered a turrid genus (Powell, 1966), although Vayssiere
(1930) showed that it has a buccinid radula (Figs 488-489). The radula of Morrisonella is known and
agrees well with our figures. Dellina is known only from empty shells, and was considered to belong
to Cancellariidae, but we find the shell so similar to Belomitra that we have synonymized it.

Our material of B. quadruplex shows great variation and there exist several areas where the species
occurs in more shallow water and forms morphologically separable populations. There is, however,
a continuous transition to the more uniform deep-water form. Such shallow-water populations exist
in the Bay of Biscay and the Azores. These shallow-water animals lack eyes, like the deep-water form;
we therefore find it more likely that they are shallow-water outcrops of a deep-water species, than the
opposite, because it is most unusual for shallow-water neogastropods to have no eyes, while many
deep-water species do lack them.

Belomitra richardi (Dautzenberg & Fischer, 1906) is known only from a few specimens and we
have not been able to decide whether it is a valid species or only another shallow-water offshoot of
B. quadruplex.

The species of Belomitra resemble mitrids, but lack columellar folds and are easily distinguished
by the radula. The genus is characterized by the distinctly shouldered whorls and the peculiarly
shaped lateral teeth.

The central tooth of Belomitra resembles that of Fasciolariidae more than that of Buccinidae, but
the lateral teeth are more like those of Buccinidae. We have preferred to place Belomitra in
Buccinidae simply because there is more variation in the buccinid radula than in the fasciolariid one,
as far as one can judge from the literature.

Belomitra quadruplex (Watson, 1882) Figs 424, 488-489, 579-586

Pleurotoma (Clionella) quadruplex Watson, 1882:253.

Belomitra paradoxa Fischer, 1882:275.

Pleurotoma climakis Watson, 1886:315, pi. 26, fig. 7.

Clionella quadruplex Watson, 1886:370, pi. 19, fig. 7.

Jumala brychia Verrill & Smith in Verrill, 1885:422, pi. 44, fig. 10.

Bela guernei Dautzenberg, 1891:614.

Belomitra fischeri Locard, 1897:261, pi. 13, figs 7-11.

Clionella delicatulina Locard, 1897:222, pi. 10, figs 9-16.

Clionella delicatulina var. costulata Locard, 1897:224.

Clionella conspicienda Locard, 1897: 225, pi. 10, figs 22-27 .

Clionella quadruplex var. minor Locard, 1897:224.

Belomitra speita Monterosato in Locard, 1897:263, pi. 13, figs 12-16, pi. 21, figs 15-16.

Belomitra speita var. minor Locard, 1897:264.

Belomitra speita var. major Locard, 1897:264.

Belomitra lyrata Monterosato in Locard, 1897:264, pi. 13, figs 23-28.

Type material: P. quadruplex holotype BMNH 1887.2.9.1127; B. paradoxa neotype = holotype of
B. guernei in MOM; P. climakis holotype BMNH 1887.2.9. 1007; J. brychia holotype USNM 44647;
other types in MNHN.

224 P. BOUCHET & A. WAREN

Type localities: P. quadruplex CHALLENGER st 73, W of the Azores, 1800 m; J. brychia USFC st
2224, off North Carolina, 1800 m; B. paradoxa TRAVAILLEUR 1882, dr 16, off N Portugal, 627
m.

Material examined: The types and TALISMAN 1883 st 44, 30°03 N, 09°22 W, 2212 m, 1 sh; st 45, 30°01 N,
09°25 W, 2115 m, 3 shs; st 47, 29°52 N, 09° 24 W, 2075 m, 3 shs; st 48-49, 29°52 N, 09° 24-27 W, 1235-2083 m, 3
shs; st 86, 22°57 N, 15°01 W, 930 m, 2 spms; st 131, 27°35 N, 27°06 W, 1440 m, 7 shs; st 137, 38°37 N, 26°01 W,
1258 m, 10 spms; st 145, 40°35 N, 21°14 W, 4415 m, 2 spms; TRAVAILLEUR 1881 dr 42, 44°01 N, 04°45 W,
900 m, 2 shs; NORATLANTE st B019, 47°29 N, 08°24 W, 2149 m, 3 spms; BIACORES st 39, 37°44 N, 27°03
W, 420 m, 3 shs; st 66, 38°35 N, 28°20 W, 1245 m, 4 spms; st 120, 39°04 N, 32°44 W, 2100 m, 1 sh; st 150, 37°37
N, 25°35 W, 575 m, 2 spms; st 157, 37°34 N, 25°43 W, 800 m, 1 sh; st 165, 37°33 N, 25°58 W, 2060 m, 1 sh; st
202, 27°36 N, 24°59 W, 2850 m, 3 shs; st 216, 37°05 N, 24°53 W, 1735 m, 2 shs; st 232, 36°55 N, 25°11 W,
390-620 m, 1 spm; st 240,' 37°35 N, 25°33 W, 815 m, 1 spm; st 245, 40°57 N, 22°16 W, 4270 m, 3 spms;
THALASSA Y 374, 41°31 N, 09°20 W, 1250 m, 1 sh; BIOGAS CV16, 44°07 N, 04° 17 W, 1909 m, 1 sh; CV23,
47°33 N, 08°34 W, 2034 m, 1 spm; CP01, 47°35 N, 08°41 W, 2245 m, 1 sh; CP02, 47°33 N, 08°41 W, 2177 m, 2
spms; CP07, 44°10 N, 04°16 W, 2170 m, 1 sh; CP17,46°31 N, 10°20 W, 4706 m, 1 spm; CP24, 44°08 N, 04° 16
W, 1995 m, 1 sh; DS52, 44°06 N, 04°22 W, 2006 m, 1 spm; DS57, 47°31 N, 09°08 W, 2906 m, 2 shs; DS86,
44° 05 N, 04° 19 W, 1950 m, I spm; DS87, 44°05 N, 04° 19 W, 1913 m, 1 spm; st 2, 47°30 N, 09°07 W, 3050 m, 1
sh; st 1, 44°33 N, 08°38 W, 2170 m, 4 shs; INCAL CP01, 57°58 N, 10°55 W, 2068 m, 1 spm; CP02, 57°58 N,
10°43 W, 2091 m, 2 spms; CP03, 56°38 N, 1 1 °06 W, 2466 m, 4 shs; CP13, 46°02 N, 10° 14 W, 4800 m, 0SO5,
47°31 N, 09°34 W, 4296 m, 1 sh; 0SO6, 46°27 N, 09°36 W, 4316 m, 1 sh; WS 07, 47°30 N, 09°37 W, 4281 m, 2
spms; WS 08, 47°31 N, 09°34 W, 4300 m, 1 sh; WS 09, 47°29 N, 09°34 W, 4277 m, 3 spms; WS 10, 47°27 N,
09° 40 W, 4354 m, 2 spms; INGOLF st 10, 64°24 N, 28°50 W, 1484 m, 1 sh; SEABED II, st CP19, 19° 15 N,
29° 49 W, 4958 m, 1 spm; CP20, 19° 13 N, 29° 48 W, 4940 m, 2 spms.

Distribution: The lower parts of the European continental slopes, from S of Iceland, south to 19° N.
In the Azores and the Bay of Biscay it also occurs more shallowly, up to about 300 m. It also occurs
all over the N Atlantic abyssal area, but does not reach the N American continental slope. The
deepest records are from about 5300 m (Fechter, 1979).

Remarks: The animal lacks eyes and has direct development. The variability is discussed under the
introduction to the genus. Abyssal specimens tend to be larger and have a thinner shell (“brychia”
Verrill), bathyal specimens are more solid and smaller ( “ paradoxa ’ ’ Fischer).

NORTH-EAST ATLANTIC NEOGASTROPODA 225

Figs 579-586. Belomitra quadruplex. 579? NORATLANTE st B19, 30.4 mm. 580, syntype of Clionella
delicatulina, 35 mm. 581, BIACORES st 240, 11.4 mm. 582, BIACORES st 232, 15.5 mm. 583,
holotype of Pleurotoma richardi (see text), 27.6 mm. 584, BIACORES st 39, 14.7 mm. 585, Azores,
young specimen. 586, TALISMAN dr 145, apex 1 mm.

226 P. BOUCHET & A. WAREN

Genus COLUS Roding, 1798

Type species: Murex islandicus Mohr, 1786, subsequent designation Dali (1906).

Synonyms: Sipho Klein, 1753 and many subsequent authors. Prelinnean.

? Colus Humphreys, 1797. Rejected work.

? Rhombus Humphreys, 1797. Rejected work.

Atractus Agassiz, 1839. Type species M. islandicus Mohr, 1786, subsequent designation Cossmann
(1901).

Tritonofusus Morch, 1857. Type species M. islandicus Mohr, 1786 subsequent designation
Cossmann (1901).

Brongus De Gregorio, 1885. Type species M. islandicus Mohr, 1786, here designated.

Remarks: The history of the name Sipho, which has been much used for the species here placed in
Colus, needs to be discussed because the name Sipho has been used until recently and the case has
never been thoroughly treated. Klein (1753) used the name for a number of slender neogastropods
and this was repeated by Bruguiere (1792:532). This use by Bruguiere (1792) does not validate the
name, because Bruguiere only repeated the classification used by Klein, to give the nomenclatorial
history of these gastropods, and he did not use the name himself.

Fabricius (1823:83) is cited by Sherborn, and Fabricius (dated 1822) is also cited by Herrmannsen
as the first validation of Sipho. Fabricius’s work is a list of his collection and no descriptions are
given. He cited four names under Sipho viz. infundibulum, turritus, polygonus and striatus. These
names are nomina nuda and the work has been rejected by ICZN (Opinion 521). Therefore the name
cannot be ascribed to Fabricius.

The next time Sipho was used was by Brown (1833), who described Sipho striatus. This is a junior
synonym of Puncturella noachina (Linne, 1758). No other species were mentioned in Sipho by
Brown, and therefore Sipho becomes a junior synonym of Puncturella Lowe, 1827.

We have therefore followed Dali (1906, 1918), Iredale (1915) and Winckworth (1932) in their use
of the name Colus, with Murex islandicus as type species. They had, however, overlooked the
validation of this name by Mohr (1786), and ascribed the name to Gmelin (1790). This does not
change anything, because these two authors both refer to Chemnitz’s non-binominal use of the
name.

We have used Colus for a rather heterogeneous assortment of more or less slender buccinids with
an operculum with apical nucleus, a radula with a mono- or tricuspidate, square central tooth and
laterals with 2 major and 0-3 smaller cusps.

There are several names available for a subgeneric division of Colus. The following names are
based on N Atlantic species:

Siphonorbis Morch, 1869. Type species Fusus (Siphonorbis) ebur Morch, 1869, subsequent
designation by Cossmann (1901). (Friele’s (1882) designation is not valid because the species
designated as type was not originally included in Siphonorbis.)

Siphonellona Wenz, 1941 . New name for Neptunella Verrill, 1873 (not Meek, 1864) and Siphonella
Verrill, 1879 (not Macquart, 1835). Type species Fusus pygmaeus Gould, 1841, by original
designation.

Plicijusus Dali, 1902. Type species Fusus kroeyeri Moller, 1842 by original designation. Synonym:
Parasipho Dautzenberg & Fischer, 1912, with the same type species.

Anomalosipho Dautzenberg & Fischer, 1912. Type species Sipho verkruezeni Kobelt, 1876, by
original designation.

None of these names, however, has a type species that is a member of any significantly different
group of species, except perhaps Siphonorbis. Colus sabini ( = ebur), the type of Siphonorbis, has a
perfectly smooth larval shell of about three whorls, almost identical to that of C. holboelli and C.
terraenovae. We are not able to take any decision about this until the N Pacific species of the group
are better known and prefer to leave the N Atlantic Colus species without a subgeneric classification.

NORTH-EAST ATLANTIC NEOGASTROPODA 227

In addition to the species treated here, we have seen two forms from the N American east coast,
that we were unable to classify (Figs 644-646). They may prove to be forms of C. pygmaeus, but
more material from other localities is needed to show this.

Because of the great variation of several of the species, it is close to impossible to give a key to the
species, but this attempt may give some help in their determination.

A.

A.

B.

B.

C.

C.

D.

D.

E.
E.

F.

F.

G.

G.

H.

H.

I.

I.

Siphonal canal very short and indistinct, broad
Siphonal canal long and slender

All the shell covered by spiral sculpture, sometimes also axial ribs
Only the apical whorls with spiral sculpture, no axial ribs

No axial sculpture, shell usually whitish or greyish under the periostracum
Axial sculpture present, shell usually brownish or reddish under the
periostracum

Outer lip flaring in adult specimens, spiral sculpture indistinct
Outer lip regularly arched, spiral sculpture distinct

Larval shell consisting of 2-3 distinctly set off, smooth, inflated whorls
Larval shell consisting of about one distinctly set off whorl or several, not
distinctly set off, whorls

B

C

C. verkruezeni
C. turgidulus

E

D

C. kroeyeri
C. later iceus

F

H

Shell very broad and inflated
Shell slender

Outer lip forms a distinct angle with the siphonal canal
Outer lip and siphonal canal form an even arc

Larval shell inflated, usually broader than the subsequent whorl (Fig.
616)

Larval shell more slender than subsequent whorls
Larval shell with 1-1.5 distinctly set off whorls

Larval shell with a very small initial whorl, not distinctly set off

C. terrae novae
G

C. holboelli
C. sabini

C. islandicus

I

C. gracilis
C. stimpsoni
C. pubescens
C. jeffreysianus
C. pygmaeus

Colus gracilis (Da Costa, 1778) Figs 414-416, 473, 587-589, 592-605

Buccinum gracile Da Costa, 1778:124, pi. 6, fig. 5.

Fusus listen Jonas, 1846:106, pi. 10, fig. 13.

Tritonium (Fusus) islandicum var. striata Middendorff, 1849:472.

Fusus gracilis var. convoluta Jeffreys, 1867:336.

Sipho glaber Verkriizen in Kobelt, 1876:174, pi. 3, fig. 3.

Fusus gracilis vars coulsoni and belliana Jordan, 1890:232.

Neptunia nicolloni Locard, 1891:34.

Neptunia gracilis var. major Locard, 1896:150.

Sipho togatus var. frielei Harmer, 1914:182, pi. 22, figs 3-4.

Sipho turgidula var. minor Thorson, 1944:81, fig. 10.

Type material: B. gracile not known; F. listen not known; S. glaber lectotype, here selected, SMF
190099/1.

Type localities: B. gracile, British Isles; F. listen, the Kattegat; S. glaber, Porsangerfjorden, N
Norway.

228 P. BOUCHET & A. WAREN

Material examined: More than 1000 specimens from all parts of the distributional area, MNHN, USNM, ZMC,
SMNH, coll. AW, ZMB.

Distribution: From the Murmansk area (Middendorff, 1849; Herzenstein, 1885) and southwards
along the Scandinavian coasts, to S Kattegat (Thorson, 1946), the British Isles (Seaward, 1982),
the Bay of Biscay and south to N Portugal (several THALASSA stations). It occurs in the Faroes
(Sparck & Thorson, 1933), S, W and N Iceland (Thorson, 1941) and SE Greenland (Thorson, 1944
as turgidula var. minor and unpublished material in ZMC). C. gracilis lives on the continental shelf
and the upper slope in 50 — 1500 m.

Remarks: C. gracilis is a highly variable species that occurs in several morphologically different
geographical and bathymetrical forms. These forms may look identical over large areas and easily
give an impression that several species are involved.

For Norwegian specimens the name glaber has been used. This is a smooth form with flatter
whorls and proportionally greater breadth than typical gracilis. The glaber form reaches the Faroes,
Shetland and Iceland. In the Faroes and Shetland there is a transition to typical gracilis , while in
Iceland there is a tendency to get still smoother, broader and with flatter whorls, which misled
Thorson (1944) to refer specimens to C. turgidulus (Fig. 598). The transition to typical gracilis can
also be seen in the North Sea, where the specimens are quite intermediate. To the W and NW of the
British Isles and along the Iceland - Faroe Ridge the glaber form enters deeper water and at the
same time the periostracum becomes less distinct, the shell smoother and the siphonal canal more
curved (Fig. 599). To the south, in shallow water, there is little variation in the gracilis form,
although there exist local populations with a more solid shell and convex whorls, from which Locard
described nicolloni (Fig. 596). In deeper water there is an increase in size, a reduction of the thickness
of the shell and an increase in the convexity of the whorls, from the Faroe-Rockall area southwards.
The typical form from the Bay of Biscay in 700 — 1300 m looks very distinct from the shallow-water
form of the same area (Figs 600, 603). These three different forms, gracilis (Fig. 597), glaber (Fig.
594) and the S deep-water form (Fig. 600) may possibly be worth subspecific distinction.

The egg capsules were described by Jeffreys (1867) but have never been figured so we give figures
of specimens from W Norway and deep water in the Bay of Biscay, to show the differences between
the populations and to emphasize that differences in the egg capsules should not be used uncritically
to prove specific separation.

C. gracilis has been reported from the Mediterranean several times (Crosse, 1875; Spada &
Maldonado, 1974, e.g.) but these records have been based on C. jeffreysianus.

Colus holboelli (Moller, 1842) Figs 419-421, 471 , 606-61 1

Fusus holbolli Moller, 1842:88.

Fusus tortuosus Reeve, 1855:394, pi. 32, fig. 5.

Sipho tortuosus var. attenuata G.O. Sars, 1878:273, pi. 15, fig. 5.

Fusus delicatus Jeffreys, 1883:396, pi. 44, fig. 6.

Sipho ( Siphonorbis ) turritus var. distincta Posselt, 1895:179.

Sipho togatus var. crassa Harmer, 1914:182, pi. 22, fig. 6.

Type material: F. holboelli , holotype in ZMC; F. tortuosus, neotype, here designated BMNF1
1968377.

Type localities: F. holboelli, W Greenland; F. tortuosus, N of Beechey Island, NE Canada.

Material examined: About 300 shells and specimens from all parts of the distribution area, ZMC, SMNH,
USNM, MNHN, BMNH.

Distribution: Arctic Canada from 90° W (Macpherson, 1971 as tortuosus), W Greenland (Thorson,
1951 as turritus), E Greenland (Thorson, 1944 as tortuosus ), W, N and E coasts of Iceland
(Thorson, 1941 as tortuosus), between Shetland and the Faroes (Jeffreys, 1883, as delicatus), deep
water off Norway and the Norwegian coast to Spitzbergen (Sars, 1878; Friele, 1882). We have seen
no material from Spitzbergen. Normally it occurs in 400-1500 m, although it occurs more shallowly
in Greenland and Iceland, 10-500 m.

NORTH-EAST ATLANTIC NEOGASTROPODA 229

Remarks: As was pointed out by Friele (1882), Jeffreys and G.O. Sars were using the name attenuata
in different ways. Jeffreys used it for a deep-water form of C. jeffreysianus.

The specific name holboelli could possibly be considered a nomen oblitum, but we have preferred
to use it because of the confusion in the use of tortuosus.

We have selected a specimen of C. holboelli in the collection of H. Cuming, BMNH, from “Arctic
Seas” as neotype of C. tortuosus , to stabilize the nomenclature.

Harmer (1914) described Sipho exiguus from the British Crag formation and mentioned that he
had seen specimens in SMNH from Iceland. We cannot judge the validity of the fossil species, but
suppose the Recent record to be based on C. holboelli of which certain Icelandic specimens resemble
Harmer’s figures closely.

C. holboelli is a rather variable species. The type and other specimens from W Greenland are
almost as solid as C. gracilis while specimens from deep water have a thin, fragile shell. Norwegian
specimens are smaller and more slender.

C. holboelli resembles C. sab ini, but has a smooth periostracum, a more curved siphonal canal
and the outer lip is not evenly arched where it goes over into the siphonal canal as in C. sabini. Very
young specimens may resemble C. terraenovae in having a similar shape and larval shell, but that

species has a proportionally higher aperture.

When the larval shell is too corroded to be examined C. holboelli can be distinguished from other
species by its more slender and twisted siphonal canal.

Colus islandicus (Mohr, 1786)

Figs 479,612-616

Fusus islandicus... Chemnitz, 1780:159, figs 1312, 1313 (not binominal).

Murex islandicus Mohr, 1786:136.

Tritonium islandicum var. sulcata Middendorff, 1849:471.

Fusus obesus Sowerby, 1880:sp. 129.

Neptunia islandica var. gallica Locard, 1896:149.

Type material: F. islandicus, lectotype, here selected, in ZMC (Fig. 612).

Type locality: F. islandicus, Iceland.

Material examined: About 200 specimens from all parts of the distributional area.

Distribution: Along the shelf and upper part of the continental slope from aVctic Canada (90° W,
Macpherson, 1971), to 150 miles S of Cape Race, Newfoundland (unpublished, USNM), W and E
Greenland (Thorson, 1944, 1951), Iceland (Thorson, 1941), the Faroes (Sparck & Thorson, 1933),
Spitzbergen (Odhner, 1915), Norway (G.O. Sars, 1878), the outer parts of the Skagerrak and the
North Sea (unpublished material in ZMC), Shetland and W of the British Isles (Seaward, 1982;
Massy, 1930), the Bay of Biscay, off Spain and Morocco (Locard, 1897; Dautzenberg, 1927). In the
arctic area it occurs in 5-500 m, in the southern parts in 400-2000 m.

Remarks: We have not seen E Siberian or N Pacific material of the species and do not exclude that
the records from these areas can be ascribed to misidentifications (cf. Odhner, 1915).

We have selected a specimen from the collection of Chemnitz as lectotype. It does not agree very
well with the figure in Chemnitz (1780), but Chemnitz had already criticized the figures for not
resembling his specimens. Mohr’s (1786) validation of the name was based on Chemnitz’s figures
which therefore have to be considered types.

The size of C. islandicus is highly variable. It reaches a maximum size of about 200 mm, but we

have seen deep-water specimens of 90 mm that evidently are full-grown. Specimens from the Bay of
Biscay and southwards are more slender and solid than arctic specimens.

The development was described by Thorson (1935), who gave figures of egg capsules and young
specimens. C. islandicus hatches at a size of 5-8 mm.

C. islandicus can be recognized from its large size, unusually inflated larval whorl, the oblique
suture and the long, straight, siphonal canal.

230 P. BOUCHET & A. WAREN

Colus jejfreysianus (Fischer, 1868) Figs 428-429, 474-475, 477, 617-630

Fusus propinquus Alder, 1848: 157 (not Goldfuss, 1843).

Tritonium turritum M. Sars, 1859:39 (nom. nud.).

Fusus propinquus var. turrita Jeffreys, 1867:339 (not F. turritus Schafhautl, 1863).

Fusus jeffreysianus Fischer, 1868:37.

Fusus attenuatus Jeffreys, 1877:326 (not Philippi, 1847).

Fusus (Sipho) pupula Fischer, 1882:274.

Neptunia jeffreysiana vars major, minor, ventricosa, elongata & bicolor Locard, 1896:151-152.

Neptunia torra Locard, 1897: 361, pi. 17, figs 21-25.

Neptunia pupoidea Locard, 1897:363, pi. 17, figs 26-28.

Fusus consimilis Marshall, 1902:49.

Fusus propinquus var. howsei Marshall, 1902:46; 1911:226.

Fusus propinquus var. laevis Marshall, 1902:46; 1911:225.

Fusus marshalli Iredale, 1918:33.

Type material: F. jeffreysianus, lectotype , here selected, in MNHN.

Type locality: F. jeffreysianus, Arcachon, Gironde, SW France.

Material examined: About 600 specimens from all parts of the distribution area.

Distribution: From the Kattegat and SW Skagerrak (20-200 m, Petersen, 1888, ZMC), the British
Isles, Shetland (Seaward, 1982; Massy, 1930), the Faroes (unpublished material ZMC) and S Iceland
in deep water (400-2100 m, unpublished material in ZMC). From the British Isles southwards on the
continental shelf and slope S to Casablanca (Morocco) and into the Mediterranean, at least to the
Alboran Sea (Locard, 1897; material in MNHN), possibly also S France (Monterosato, 1877).

Remarks: C. jeffreysianus is an extremely variable species which has given rise to a prolific
synonymy. In S Scandinavia and N Great Britain including the Shetlands, the predominant form is
small, solid, with convex whorls and a rather coarse periostracum. The name howsei was based on
this form. Off W Great Britain and Ireland this form goes over into typical jeffreysianus, which is
larger, higher, has flatter whorls and a thinner periostracum. This transition has already been
pointed out by Marshall (191 1) and Massy (1930), but has never been accepted in the literature. The
jeffreysianus form occurs mainly in 100-200 m and is distributed southwards to the Spanish N coast,
where it goes deeper and becomes smaller. In the area off N Great Britain the howsei form becomes
more slender, more cylindrical and gets a thinner periostracum when it is found in deeper water, the
attenuata form (Fig. 622). This is a continuous change and we have observed a similar change in the
Bay of Biscay from typical jeffreysianus, via the torra and pupoidea forms to typical attenuatus (cf.
Figs 624, 626-628). There cannot be put any sharp upper limit for the attenuata form, but usually it
ocdurs below 1000 m, while specimens from 200-1000 m are usually transitional between
jeffreysianus and attenuatus.

Thorson (1944) excluded C. jeffreysianus from the Greenlandic fauna, which we find correct.
There are a few specimens in ZMC labelled “Greenland” by Moller, but they belong to the Kattegat
form of howsei and there is no doubt that the labels are erroneous. Locard (several papers) reported
C. jeffreysianus from “Provence”, but we have not seen any material in his collection.

Monterosato (1877) discussed the occurrence of C. jeffreysianus in the Mediterranean and wrote
that Martin’s (Jeffreys, 1867) record of Fusus buccinatus from the stomach of Trigla gurnardus
(Pisces), from Golfe du Lion, S France was based on C. jeffreysianus. He also mentioned that he
had often found broken specimens in that fish, and referred Crosse’s record (1875) of two specimens
of Fusus gracilis from Golfe du Lion to C. jeffreysianus. It seems therefore that the species occurs
further into the Mediterranean than our material indicates.

Thorson (1946) described the larval development under the name Sipho propinquus.

Colus jejfreysianus and C. gracilis are often quite difficult to separate. When the larval shell is
present they can easily be distinguished {cf. figs 604-605, 629-630), but this is often corroded. Then
one can use the general shape of the shell, which is more slender, with flatter whorls in C. gracilis,
but often it is necessary to wait until one gets some young specimens with preserved larval shell
from the population in question, especially if the specimens come from deep water.

NORTH-EAST ATLANTIC NEOGASTROPODA 231

Colus kroeyeri ( Moller, 1842)

Figs 631-633

Fusus kroyeri Moller, 1842:88.

Fusus arcticus Philippi, 1850:119, pi. 5, fig. 5.

Fusus (Tritonofusus) kroeyeri vars major & pumila Morch, 1869:19.

Plicifus johanseni Dali, 191 9:21 A.

Type material: F. kroeyeri, lectotype, here selected, in ZMC.

Type locality: F. kroeyeri, W Greenland.

Material examined: The type and the material listed by Thorson (1944, 1951) and Odhner (1915) and about 25
samples scattered over the N Atlantic part of the range (MNHN, SMNH, USNM, ZMC).

Distribution: Circumpolar in 0-200 m (Macpherson, 1971). Not present in Norway (Sars, 1878) or
Iceland (Thorson, 1941).

Remarks: We have included this species to make the ennumeration of N Atlantic species of Colus
complete. There are no species in the N Atlantic that are easily confused with C. kroeyeri. The
distinct axial ribs in combination with the brownish shell make it easily recognizable.

Fusus cretaceus Reeve, 1847 has usually been synonymized with C. kroeyeri but Cernohorsky
(1977: Rec. Auckland Inst.Mus. , 14:97) has shown that it is a synonym of the New Zealand muricid
Xymene ambiguus (Philippi, 1844).

Colus latericeus (Moller, 1842)

Figs 423,478,634-636

Fusus latericeus Moller, 1842:88.

Fusus pellucidus Hancock, 1846:330, pi. 5, fig. 3.

Fusus pullus Reeve, 1848:sp. 89.

Fusus (Tritonofusus) latericeus var. laevis Morch, 1869:19.

Sipho geminostriatus Pfeffer, 1886:4, fig. 1.

Neptunea (Sipho) pertenuis Sykes, 1911:339, fig. page 340.

Type material: F. latericeus, lectotype, here selected, in ZMC.

Type locality: W Greenland.

Material examined: The material enumerated by Thorson (1941, 1951) and Odhner (1915) and a few additional
localities from the known distributional area.

Distribution: Along the W Greenland coast, Jan Mayen (absent from E Greenland) (Thorson 1944,
1951), NW and N Iceland (Thorson, 1941), N Norway (G.O. Sars, 1878), Spitzbergen (Odhner,
1915) and Barents Sea, in 20-200 m.

Remarks: This is a coast and shelf species, but we have included it to make this a complete treatment
of the N Atlantic species of Colus. It has been recorded from 1 187 m (Friele, 1882) but this record is
based on an empty shell. Fusus pertenuis (holotype BMNH 1885.11.5.2931) was based on an
unusually slender and thin specimen, but the sculpture and apex are normal.

C. latericeus may be recognized by the reddish brown shell (occasionally colourless), the sculpture
of strong spiral lines and axial ribs and its small size.

Colus pubescens (Verrill, 1 882)

Figs 465,639-640

Sipho pubescens Verrill, 1882:501, pi. 43, fig. 6, pi. 57, fig. 25.

Type material: Three syntypes, USNM 37767.

Type locality: USFC st 793, off Newport, Rhode Island, 35 m.

Material examined: Many hundred specimens, from off Daytona, Florida, to Newfoundland, 20-800 m
(USNM).

Distribution: As above.

232 P. BOUCHET & A. WAREN

Remarks: C. pubescens has often been confused with C. stimpsoni, but differs by having a smaller
larval shell and a hairy periostracum; it is more slender and has more convex whorls than stimpsoni.
It also resembles C. gracilis very much, but that species has a smooth periostracum.

Macpherson (1971) recorded C. pubescens from arctic Canada, but her figure (pi. 5, fig. 1) clearly
shows a specimen of C. sabini, so we consider her record uncertain.

Colus pygmaeus (Gould, 1841) Figs 464, 641-643

Fusus islandicus var. pygmaeus Gould, 1841:284, fig. 199.

Fusus trumbulli Linsley, 1845:285.

Sipho pygmaeus var. planulus Verrill, 1882:505.

Type material: All lost.

Type locality: F. pygmaeus, “In fish, our harbour” (Boston, Massachusetts).

Material examined: A few hundred shells and specimens from all parts of the distribution area.

Distribution: From Gulf of St. Lawrence to North Carolina, NE America, in 1 — 1100 m (Abbott,
1974).

Remarks: There seem to be no problems with the identity of this species, but the young looks rather
different from the adult specimens (Figs 641, 642). We have, however, seen two more forms, one
from Florida (Fig. 644) and one from off N. Carolina (Fig. 645), which differ from pygmaeus by
their larger size and more straight columella. These two forms also resemble C. jeffreysianus, but we
consider them too imperfectly known to allow any further speculations.

Colus sabini (Gray, 1 824)

Figs 41 8, 472, 647-652

Buccinum sabini Gray, 1824:240.

Fusus reeveanus Petit de la Saussaye, 1851:365, pi. 10, fig. 7.

Fusus (Siphonorbis) togatus Morch, 1869:398.

Fusus (Siphonorbis) ebur Morch, 1869:398.

Fusus (Siphonorbis) pfaffi Morch, 1876:369.

Neptunea (Sipho) hanseni Friele, 1879:281.

Fusus hirsutus Jeffreys, 1883:396, pi. 44, fig. 7.

Sipho brevispira Brogger, 1900:49, pi. 3, fig. 4.

Sipho togatus vars vallensis & sinuosa Brogger, 1900:47, pi. 2, fig. 2 and pi. 3, fig. 5.

Type material: B. sabini, one neotype in BMNH, here selected, no. 1968376; F. reeveanus, holotype
in MNHN; F. togatus, ebur, pfaffi, syntypes in ZMC; TV. hanseni, holotype ZMB 20621.

Type localities: B. sabini, NW Arctic Atlantic; F. reeveanus, presumably the Newfoundland Banks.
Material examined: About 400 specimens from all parts of the N Atlantic range, USNM, ZMC, SMNH, ZMB,
MNHN.

Distribution: From arctic Canada (Macpherson, 1971 as pubescens and togatus) south to 150 miles S
of Cape Race, Newfoundland (unpublished in USNM), W Greenland (Thorson, 1951, as togatus ), E
Greenland (Thorson, 1944, as curtus ), N and E Iceland (Thorson, 1941, as togatus ), between
Shetland and the Faroes (Jeffreys, 1883, as hirsutus ), Spitzbergen (Odhner, 1915, as hanseni,
togatus and hirsutus), Novaja Zemlya (Leche, 1878, as sabini) and the Laptev Sea (Aurivillius,
(1885). Absent from Norway (Friele, 1882). It occurs in 5-1200 m, normally 100-600 m, in high
arctic areas in more shallow water.

Remarks: Some developmental stages were described by Thorson (1935) under the name S. curtus.
The young live on nurse eggs and hatch at a size of 5 mm.

C. sabini is quite variable, both in size (max. 83 mm, the type of reeveanus) the solidity of the
shell, and the sculpture. A good help in recognising it is the sculpture, which consists of alternating
strong and weak spiral ribs. Usually the apical whorls are completely corroded. The parietal callus is

NORTH-EAST ATLANTIC NEOGASTROPODA 233

often very solid in adult specimens and sometimes there is an umbilical chink.

We are not aware why Macpherson (1971) considered Buccinum sabini a nomen dubium. We have
selected a neotype from an old sample labelled sabini and collected by Sabine. The description is not
less precise than that of most other species described at the same time.

Colus stimpsoni (Morch, 1867) Figs 463, 653-657

Fusus striatus Reeve, 1847:sp. 42 (not Sowerb^ 1828).

Fusus curtus Jeffreys, 1867:336 (not Fleming, t828).

Fusus stimpsoni Morch, 1868:83.

Fusus americanus Bell, 1870:214.

Fusus solidulus Sowerby, 1880:sp. 131.

Neptunea (Sipho) arata Verrill, 1880:370 (/z^ Verrill 1882:500).

Sipho stimpsoni vars brevis & liratulus Verrill, 1882:500.

Neptunea tenuistriata Harmer, 1918:369, pi. 37, figs 1-2.

Type material: F. stimpsoni, neotype, here selected, in ZMC.

Type locality: F. stimpsoni, “Shores of New Jersey”, NE US.

Material examined: Several hundred specimens from all parts of the distributional area, mainly in USNM.
Distribution: Labrador to off N Carolina, 2 — 850 m (Abbott, 1974).

Remarks: C. stimpsoni is a highly variable species. The short broad form (Fig. 653) has often been
confused with C. terraenovae and has a more N and shallow distribution, from the New England
area northwards. All intermediates between this form and the more slender, more southern and
deeper form (Fig. 655) exist.

C. stimpsoni may be recognized from its smooth, shiny, naked periostracum and usually very
solid shell. These features are shared with C. terraenovae but that species has a smooth, multigyrate
larval shell and is still shorter and broader even than the form from the Banks.

To stabilize the nomenclature, we have selected a neotype from the Morch collection in
Copenhagen.

The development and reproductive anatomy were described by West (1973, 1978).

Colus terraenovae nom.nov.

Figs 413, 462, 658-660

Fusus ventricosus Gray, 1839:117 (not Anton, 1838).

Type material: Two syntypes in BMNH, no reg. no.

Type locality: Not mentioned, here restricted to Newfoundland Banks.

Material examined: About 40 specimens from the Newfoundland Banks.

Distribution: Uncertain because of confusion with the short forms of C. stimpsoni. We have only
seen reliable records from the Banks around Newfoundland, in 40 — 150 m. Verrill (1882) considered
the records from New England doubtful.

Remarks: Colus terraenovae has very often been confused with the short, stout forms of C.
stimpsoni. This is probably the reason why it has been recorded from the NE U.S. (Abbott, 1974)
and as a fossil from Great Britain (Pain, 1981). They can, however, easily be distinguished by the
outlines of the spire, which are convex in terraenovae and concave in stimpsoni. Also the larval shells
are widely different: that of terraenovae consists of 2-3 smooth, distinctly set off whorls while that of
stimpsoni is not distinctly set off, starts with a large initial whorl and has a rough surface (Figs 657,
660).

The name ventricosus under which it has been usually known is preoccupied by Anton’s (1838) use
of the name and it has to be given a new name. The date of publication of Anton’s catalogue was
discussed by Cernohorsky ( Veliger , 1978, page 299), who showed that it had already been published
in 1838 instead of 1839 as generally supposed.

234 P. BOUCHET & A. WAREN

Coins turgidulus (Jeffreys, 1877) Figs 417, 422, 470, 637-638

(no generic name) turgidulus Jeffreys in Wood, 1872:25, nomen nudum.

Fusus turgidula Jeffreys in Friele, 1877:8.

Not Sipho turgidula var. minor Thorson, 1944 (see C. gracilis).

Type material: Holotype ZMB 20619.

Type locality: Shetland-Faroe Channel.

Material examined: The material listed by Friele (1882) and TRITON st 9, 60°05 N, 06°21 W, 1130 m, 4 spms;
PORCUPINE 1869 st 78, 60° 15 N, 04°45 W, 550 m, 2 shs; st 65 + 84, Shetland-Faroe Channel, 400-800 m, 2 shs;
INGOLF st 59, 65°0ON, 1 1 ° 16 W, 584 m, 1 sh; st 101, 66°23 N, 12°05 W, 1011 m, 6 shs; st 103, 66°23 N,08°52
W, 1090 m, 2 shs; st 116, 70°05 N, 08°26 W, 699 m, 6 shs; 64°58 N, 11° 12 W, 564 m, 8 shs, ZMC.
Distribution: Along the slopes of the Norwegian Basin, from between Norway and Spitzbergen, to
the Shetland-Faroe Channel and E Iceland, in 400—1100 m.

Remarks: Colus turgidulus is a characteristic species but resembles Buccinum more than Coins at the
first glance. The operculum is, however, not of Buccinum type but normal for a species of Colus, as
is also the radula. The fine spiral sculpture is present only on the first postlarval whorls.

Thorson’s var. minor of C. turgidulus was based on specimens of C. gracilis and the species is thus
not known from Greenland.

Colus verkruezeni (Kobelt, 1876)

Figs 661-664

Sipho verkriizeni Kobelt, 1876:70, pi. 2, fig. 2.

Neptunea (Sipho) virgata Friele, 1879:261.

Buccinum brucei Melvill & Standen, 1900:9, fig. page 9.

Sipho verkriizeni var. plicifera Brdgger, 1900:50, pi. 1, fig. 1.

Anomalosipho dautzenbergi Dali, 1916:8.

Helicofusus paraelatior Kantor, 1981 : 1 147.

Type material: S. verkriizeni, lectotype SMF 253576/1, selected by Tiba & Kosuge, 1981; N. virgata,
holotype ZMB 20622; A. dautzenbergi, holotype in MOM.

Type localities: S. verkriizeni, Porsangerfjord, N Norway; S. virgata, VORINGEN st 124, SW of
Lofoten, N Norway, 640 m; A. dautzenbergi, off North Cape, N Norway 394 m.

Material examined: The type material listed and the material reported by Friele (1882), Thorson (1941, 1944),
Odhner (1915) and Dautzenberg & Fischer (1912). Also a few additional specimens from different parts of the
known distributional area (SMNH, ZMC).

Distribution: From E Greenland (Thorson, 1944), N Iceland (Thorson, 1941), W and N Norway
(Friele, 1882), Spitzbergen (Odhner, 1915) and the Kara Sea (Thorson, 1941), in 30—640 m.

Remarks: Macpherson (1971: pi. 5, fig. 6) figured a specimen that she had reluctantly determined as
C. verkruezeni, from NE Canada. We agree that it probably does not represent that species.

Golikov & Scarlato (1977) identified B. brucei with C. latericeus, but the very short siphonal canal
and the size (41 mm) indicate that it must be a synonym of C. verkruezeni.

C. altus (S. Wood, 1842) from the British Crag Formation, resembles C. verkruezeni closely, but
we hesitate to consider them conspecific.

NORTH-EAST ATLANTIC NEOGASTROPODA 235

Figs 587-591. Buccinid egg capsules and embryos. 587 -Colus gracilis, Hjeltefjord, Norway, egg-capsule in
side view showing opened hatching hole, diameter 6 mm. 588-Larval shell from the same egg-capsule,
diameter 3.0 mm. 589 -Colus gracilis, THALASSA st Z407 (47°44 N, 08°07 W, 1085-1115 m), egg-
capsule, diameter 17 mm. Top and side views showing the pre-formed hatching hole closed by a thin
membrane. 590 -Buccinum undatum, embryo from egg-capsule, diameter 2.5 mm. 59\-Turrisipho
fenestratus, Bergen, apical view, x 13.

236 P. BOUCHET & A. WAREN

1 1P?-^

Figs 592-597. Colus gracilis. 597-lectotype of Sipho glaber, 62 mm. 593-East of Faeroes 385-420 m, 34
mm. 594-Hjeltefjord, Bergen area, 100-180 m, 57.5 mm. 595-Moray Firth, Scotland, 50.9 mm. 596-
syntype of Neptunia nicolloni, 69 mm. 597-N of Jutland, Skagerrack 185-370 m, 99.5 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 237

Figs 598-605. Colus gracilis. 598-syntype of Sipho turgidula minor, 34.7 mm. 599-Hebridean Terrace,
750-1000 m, 62 mm. 600-THALASSA st W423 (44°04 N, 07°07 W, 710-1070 m), 44.3 mm.
601-THALASSA st Z414 (48°05 N, 08°30 W, 650 m), 47.4 mm. 602-BIOGAS st CV2 (47°42 N, 08°03
W, 994 m), 83.8 mm. 603-THALASSA st W346 (43°51 N, 02° 11 W, 460-520 m), 93.4 mm. 604-Bergen
area, apex 1.6 mm. 605-SE bay of Biscay, apex 2.3 mm.

238 P. BOUCHET & A. WAREN

Figs 606-611. Colus ho/boel/i. 606-Finmark, Norway, 39.1 mm. 607-paralectotype of Fusus delicatus,
TRITON st 8 (60°18 N, 06°15 W, 1178 m), 29 mm. 608-lectotype of Tritonium turritum, Oksfjord,
Tromsd, 44.3 mm. 609 and 610-INGOLF st 33 (67°57 N, 55°30 W, 66 m), 69 mm and apex 1.7 mm.
611-Finmark, Norway, apex 2.1 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 239

Figs 612-616. Coins islandicus. 612-lectotype, 129.5 mm. 613-Tromso, 110 m. 614 and 615-THALASSA
st W413 (43° 50 N, 06° 09 W, 500-540 m), 131 mm and 73.8 mm. 616-INGOLF st 75 (61°28 N, 26°25 W,
1469 m), apex 8.7 mm.

240 P. BOUCHET & A. WAREN

Figs 617-622. Colus jeffreysianus. 617-lectotype, 70 mm. 618 and 619-Great Britain, Jeffreys ex Jordan,
50 mm and 58.3 mm. 620-INGOLF st 10 (64°24 N, 28°50 W, 1484 m), 39 mm. 621-Shetland, 43 mm.
622-INCAL st CP2 (57°58 N, 10°43 W, 2091 m), 50.4 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 241

Figs 623-630. Colus jeffreysianus. 623-POLYMEDE st CM 15 (36°28 N, 04°09 W, 609 m), 49.5 mm.
624-THALASSA st W412 (43°50 N, 06°06 W, 272-310 m), 52.7 mm. 625-syntype of Neptunia torra, 38
mm. 626-THALASSA st W338, N Spain, 19.8 mm. 627-THALASSA st W409 (43°55 N, 06°03 W,
176-185 m), 37.6 mm. 628-THALASSA st X375 (44°07 N, 04°43 W, 625-650 m), 43.7 mm. 629-Great
Britain, Jeffreys ex Jordan, apex 2.7 mm. 630-apex 2.5 mm.

242 P. BOUCHET & A. WAREN

Figs 631-636. Genus Colus. 631-C. kroeyeri, lectotype, 68.0 mm. 632-Bellsund, Spitzbergen, 89.4 mm.
633-Spitzbergen, apex 2.3 mm. 634-C. latericeus, MONACO st 162 (46°50 N, 47°51 W, 155 m), 27.6
mm. 635 and 636-Lofoten, 19.5 mm and apex 1.7 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 243

Figs 637-640. Genus Colus. 631-C. turgidulus, VORINGEN st 286 (72°57 N, 14°32 E, 817 m) 52.7 mm.
638-VORINGEN st 323 (72°53 N, 21°51 E, 408 m), apex 2.8 mm. 639 and 640-C. pubescens, off
Martha’s Vineyard 616 m, 60.3 mm and apex 1.7 mm.

|

244 P. BOUCHET & A. WAREN

Figs 641-646. Genus Colus. 641-C. pygmaeus, off Martha’s Vineyard 185 m, 21.6 mm. 642 and
643-Georges Bank (41°30 N, 66°55 W), 11.8 mm and apex 1.2 mm. 644 -Colus sp. aff. pygmaeus, off
Fowey Light, Florida, 185 m, 13.3 mm. 645 and 646 -Colus sp. aff. pygmaeus, SSE of Cape Fear, North
Carolina, 977 m, 39 mm and apex 1.9 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 245

Figs 647-652. Colus sabini. 647-Hebron Fjord, 175 m, 59.9 mm. 648 and 649-74° 12 N, 77°05 W, 680 m,
69.2 mm and 24.1 mm. 650 and 651-KNIGHT ERRANT st 8 (60°03 N, 05°51 W, 994 m) 42 6 mm and
apex 3 mm. 652-INGOLF st 28 (65° 14 N, 55°42 W, 791 m), 8.0 mm.

246 P. BOUCHET & A. WAREN

Figs 653-657. Colus stimpsoni. 653, Coast of Maine, 45.3 mm. 654, off Martha's Vineyard 381 m, 74. 1
mm. 655, “E of Bunker’s Ledge”, NE America, 77.7 mm. 656, off Martha’s Vineyard 185 m, 19.8 mm.
657, Newfoundland Banks, apex 1.9 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 247

Figs 658-660. Coins terraenovae. 658, Newfoundland Banks, 57.8 mm. 659 and 660, NE part of Grand
Bank 74 m, 26.7 mm and apex 1.7 mm.

248 P. BOUCHET & A. WAREN

Figs 661-664. Colus verkruezeni. 661, lectotype, 49.7 mm. 662, N Norway, 45.5 mm. 633, Goteborg,
glacial deposit, 38.3 mm. 664, holotype of Neptunea virgata, apex 2.9 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 249

Family OLIVIDAE

Genus BENTHOBIA Dali, 1889.

Type species : Benthobia tryoni Dali, 1889 (by monotypy).

Synonym: Nux Barnard, 1960. Type species N. alabaster Barnard, 1960 by original designation.

Remarks: Dali originally suggested a cancellariid affinity for Benthobia and the genus remained
placed in this family without any particular reason. We have examined the radula of a specimen of
the type species and found it to be a rachiglossan type. The presence of the spiral sulcus on the body
whorl recalls that of several Olividae grouped as Pseudolivinae, both Recent, such as Fulmentum
sepimentum (Rang), and fossil, such as Pseudoliva elisae Briart & Cornet figured in Glibert (1973,
pi. 9), and several species figured by Cossmann & Pissarro (1907-1913, pi. 36, figs 177-1 - 3). We also
figure the radula of the Recent Fulmentum sepimentum from West Africa, for comparison with
Benthobia (Figs 669-670).

In the original description of Nux, Barnard noted a resemblance to Benthobia but assumed they
belonged to different superfamilies because Benthobia was then believed to be a cancellariid and
Nux had a rachiglossate radula. Barnard did not locate Nux in a family.

Benthobia tryoni Dali, 1889

Figs 665-668

Benthobia Tryonii Dali, 1889:132, pi. 35, fig. 6.

Lacuna cossmanni Locard, 1897:493, pi. 22, figs 4-9.

Nux alabaster Barnard, 1960:439, fig. 2.

Type material: B. tryoni, holotype in USNM; L. cossmanni, 2 syntypes in MNHN; N. alabaster,
holotype in the South African Museum, Cape Town.

Type locality: B. tryoni, USFC st 2678, off Cape Fear, N. Carolina, 1338 m; L. cossmanni,
TALISMAN dr 76, 25°01 N, 16°55 W, 2638 m; N. alabaster, 33°26 S, 16°33 E, 2392 m.

Material examined: the type material and BIACORES st 165, 37°33 N, 25°58 W, 2050-2085 m, 1 sh;
DISCOVERY st 8524-6, 20°46 N, 22°44 W, 4415 m, 1 spm; Gulf of Mexico, 28°13 N, 87°04 W, 800-1200 m, 1
sh; 27°01 N, 94°43 W, 1399 m, 1 sh; 25°39 N, 95°49 W, 960 m, 1 spm, MNHN; 33°52 S, 16°51 E, 2540-2797 m,
2 shs; ABYPLAINE st DS2, 37°20 N, 15°37 W, 4220-4380 m, 2 spms.

Distribution: Only known from the material examined.

Remarks: The animal has no eyes. The foot is large, with a distinct propodium. The operculum has
an apical nucleus (Fig. 668). There is a short proboscis with two large salivary glands. The large
protoconch, preserved on a single shell, is multispiral, which indicates planktotrophic larval
development.

250 P. BOUCHET & A. WAREN

Figs 665-670. Pseudolivinae. 665 and 666, Benthobia tryoni, syntypes of Lacuna cossmanni, 9.4 mm and
6.5 mm. 667, B. tryoni, radula, breadth of central tooth 48 /um. 668, Operculum, 4.1 mm. 669,
Fulmentum sepimentum (Rang), radula, breadth of central tooth 42 /^m. 670, Operculum, 4.5 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 251

Family VOLUTOMITRIDAE

The family was revised by Cernohorsky (1970) and two additional Atlantic species were described by
Bayer (1971). We have moved the genus Latiromitra to Turbinellidae after examination of the
radula, and we describe a new species of Microvoluta from the Josephine sea-mount.

Most Volutomitridae are known from very few specimens.

Genus Volutomitra H. & A. Adams, 1853

Type species: Mitra groenlandica Beck in Moller, 1842, subsequent designation Fischer, 1884.

Remarks: In our material Volutomitra is represented by two species, of which we leave one
undescribed. The genus is mainly known from Arctic, Antarctic and Subantarctic areas.

Volutomitra groenlandica (Beck in Moller, 1842) Figs 391, 671-672

Mitra groenlandica Beck in Moller, 1842:88.

Type material: Numerous syntypes in ZMC.

Type locality: W Greenland.

Material examined: PORCUPINE 1869 st 23a, 56° 13 N, 14° 18 W, 773 m, 3 shs; st 65, 61° 10 N, 02°21 W, 635 m,
3 shs; st 74, 60°39 N, 03°09 W, 373 m, 3 shs; st 75, 60°45 N, 03°06 W, 460 m, 3 shs; st 78, 60° 14 N, 04°30 W,
534 m, 3 shs; USFC st 2486, between St Peters and Baguerau, Nova Scotia, 350 m, I sh; Wellington Channel, 2
spms, USNM; Fiskenaes, Greenland, ex Moller, 7 shs, ZMC; Godthaab, Greenland, 16 shs, ZMC; 76°08 N,
13°26 W, 250-300 m, 1 spm, ZMC; 61°00 N, 08°49 W, 105-100 m, 1 spm, ZMC; Iceland and the Faroes, 4
samples with 5 spms, ZMC (listed by Thorson, 1941 and Sparck & Thorson, 1933); 20 samples with 50
specimens, Greenland, ZMC; 18 samples with 65 specimens, Greenland, Iceland, N Norway, Spitzbergen,
SMNH.

Distribution: The N North Atlantic upper bathyal zone, in arctic areas also coastal waters; Canada
(Macpherson, 1971), E and W Greenland (Thorson, 1944, 1951), Iceland (Thorson, 1941), off
Ireland (Massy, 1916), the Faroes (Sparck & Thorson, 1933), Norway N of Lofoten (G.O. Sars,
1878), Spitzbergen (Odhner, 1915), Novaya Zemlya (Golikov, 1964). Depths 7 — 773 m, usually
20—100 m.

Remarks: V. groenlandica is quite variable in the shape of the shell and the colour. Arctic specimens
tend to be more light brown while southern specimens are more greyish. We have examined the
material found off Ireland by Massy (1916) and one specimen looked recently dead; other findings
from southern localities may be subfossil.

Volutomitra sp. Fig. 673

We figure a shell from THALASSA st Z424, 48°28 N, 09°44 W, 475 m, that does not agree with any
form of V. groenlandica we have seen. It is a rather fresh-looking shell and it differs from V.
groenlandica by having flatter whorls, being more slender and by having a grey shell. It may be an
undescribed species, but it may also be a more southern form of groenlandica. It is, however,
unusual for species that occur in arctic shallow water to go so far to the south.

252 P. BOUCHET & A. WAREN

Genus MICRO VOLUTA Angas, 1877
Type species: M. australis Angas, 1877, by monotypy.

Remarks: The related genus Conomitra Conrad, 1865 is represented by numerous fossil species in
the lower Tertiary beds of Europe. The species described below is the first of this group known
from the NE Atlantic as Recent.

Microvoluta superstes sp.n.

Fig. 674

Type material: Holotype and 18 paratypes in MNHN.

Type locality: NORATLANTE st E9, 36°41 N, 14° 15 W, 208-230 m.

Material examined: Only known from the type material.

Description: (Holotype) Shell white, glossy, consisting of 5.5 slightly convex whorls, with a well-
marked suture; protoconch of one globular smooth whorl; postlarval sculpture of strong axial ribs
of which 12 and 1 1 can be counted on the body- and penultimate whorl respectively; postlarval spiral
sculpture of rather deep grooves of which there are two below the suture and nine at the base of the
body whorl; at the base of the shell these spiral grooves are more closely-spaced and form granules
where they cross the base of the axial ribs; there is a narrow concave columellar zone; three
columellar folds of which the lowermost one is very oblique; outer lip thin, siphonal canal short.

Dimensions of the shell height: 5.55 mm, breadth 2.45 mm; height of the aperture 2.5 mm,
breadth 1.1 mm.

Some paratypes lack the spiral threads below the suture, while in others the spiral sculpture is
present even at the middle of the whorls.

Remarks: All shells are empty, and most of them were bored by naticids.

Because no animal is available, the generic placement is only tentative.

NORTH-EAST ATLANTIC NEOGASTROPODA 253

Figs 671-676. Genera Volutomitra, Microvoluta and Latiromitra. 671 and 672, V. groenlandica,
INGOLF st 32 (66°35 N, 56°38 W, 599 m), 19.7 mm and 26.8 mm. 673, Volutomitra sp., THALASSA st
Z424, 19.0 mm. 674, M. superstes, paratype, 5.5 mm. 675 and 676, L. cryptodon, syntype, 30.3 mm.

254 P. BOUCHET & A. WAREN

Family TURBINELLIDAE

The family Turbinellidae contains two genera present in the deep water of the NE Atlantic. A third
genus, Ptychatractus Stimpson, 1865 is represented by a single species in the NW Atlantic, in shallow
water. Several tropical species are known from the deep shelf or upper continental slope.

Genus METZGERIA Norman, 1879.

Meyeria Dunker & Metzger, 1874:150 (non Meyeria McCoy, 1849).

Type species: Lathyrus albellus Dunker & Metzger, 1874, by monotypy.

Metzgeria Norman, 1879:56, new name for Meyeria Dunker & Metzger, non McCoy.

Remarks: Metzgeria includes three NE Atlantic species, all living in deep water, of which M. gagei is
here described as new.

M. apodema Bouchet & Talavera, 1981 has a solid broad teleoconch and a multispiral larval shell.
M. alba (Jeffreys, 1873) and M. gagei n.sp. have a paucispiral larval shell and more hidden
columellar folds. M. gagei is smaller, more slender and fragile than alba.

Metzgeria decorata (Locard, 1897) is a synonym of Fusinus amiantus (Fasciolariidae).

Metzgeria alba (Jeffreys in Wyville-Thomson, 1873) Figs 393, 677-678

Tritonium pusillum M. Sars, 1859:39 (nom. nud.).

Latirus albus Jeffreys in Wyville-Thomson, 1873:464, fig. 77.

Lathyrus albellus Dunker & Metzger, 1874:150.

Lathyrus albellus Dunker & Metzger, 1875:257, pi. 6, fig. 4.

Meyeria pusilla M. Sars in G.O. Sars, 1878:245, pi. 13, fig. 8.

Type material: L. albus, lectotype, here selected, USNM 188881; L. albellus, not found, not in
ZMHU.

Type localities: L. albus, PORCUPINE 1869 st 74, 60°39 N, 03°09 W, 350 m; L. albellus, 2 miles
WSW of Haugesund, SW Norway, 195 — 220 m.

Material examined: INGOLF st 32, 66°35 N, 56°38 W, 599 m, 5 shs; st 64, 62°06 N, 19°00 W, 1960 m, 1 sh; st
98, 65°38 N, 26°27 W, 260 m, 5 shs; E of the Faroes, 433 m, from stomachs of Anarrhichas latifrons, 2 spms,
ZMC; E of the Faroes, 300 m, 1 sh, ZMC; between Norway and Shetland, 1 sh, ZMC; 64°45 N, 29°06 W, 1070
m, 1 sh, ZMC; 71°05 N, 20°00 E, 225 m, 1 sh; SMNH; 72° 10N, 20°37 E, 350-400 m, 3 shs; SMNH; Korsfjorden
and off Korsfjorden, W Norway, 200-400 m, 50 shs and spms, coll. AW.

Distribution: On the shelf and upper continental slope, from Davis Strait (see above), Iceland
(Oskarsson, 1962, 1967; Thorson, 1941), the Faroe Channel (Thomson, 1873) and along the
Norwegian coast, N to Finmarken (G.O. Sars, 1878), in 100-1960 m.

Remarks: Metzgeria alba was recorded from the Azores (Locard, 1897), but the specimen in
question proved to be an undeterminable fusinid, possibly Fusinus amiantus.

M. alba is quite a rare species, except possibly along the Norwegian west coast.

Metzgeria gagei sp.n. Figs 389, 679-68 1

Type material: Holotype RSM 1981089-13002.

Type locality: Challenger II 1981 st 30, 58°18 N, 14°31 W, 670 m.

Material examined: The type material and INGOLF st 10, 64°24 N, 28°50 W, 1484 m, 3 shs; st 81, 61°44
N, 27°00 W, 913 m, 1 sh; st 18, 61°44 N, 30°29 W, 2137 m, 2 spms.

Description: Shell whitish, fragile and slender. The larval shell consists of a little more than one,
rather inflated whorl, sculptured with some indistinct traces of spiral lines and its height is 1.1 mm.
The holotype has 7.3 postlarval whorls which are slightly shouldered and not very convex. The lower
whorl lacks this shoulder and is evenly convex. The sculpture consists of low spiral ribs absent on
the shoulder, and on the upper whorls rather strong, sharp axial ribs distinctly angled at the edge of
the shoulder. On the body whorl the axial sculpture becomes much fainter and looks more like
incremental lines. The columella is long and slender and sculptured by incremental lines. There are
two distinct columellar folds, which are not visible when looking straight at the aperture, but only
when the shell is rotated about 1/6 of a whorl clockwise. The outer lip is straight and thin.

Dimensions: Height of the shell 28.0 mm, diameter 8.1 mm, height of the aperture 13 mm,
breadth 3.2 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 255

Remarks: The operculum is almost straight, rather thin and yellowish. This new species differs from
M. alba in having a less solid shell, sharper apical axial ribs, a proportionally larger larval shell
(compared with the subsequent whorl), and by having a more slender aperture. Comparison with
specimens of M. alba from similar depths and adjacent localities from the Ingolf expedition, show
the same differences, so it can be concluded that M. gagei is not a geographical or deep variety.of M.
alba.

Metzgeria costata (Dali, 1890), from the Lesser Antilles, 1220 m, has stronger columellar folds,
easily visible in the apertue and a proportionally higher aperture.

Metzgeria apodema Bouchet & Talavera, 1981 Fig. 682

Metzgeria apodema Bouchet & Talavera, 1981:177.

Type material: holotype in MNHN.

Type locality: METEOR st 36-98, 25°31 N, 16°02 W, 700-900 m.

Distribution: Only known from the holotype.

Remarks: M. apodema differs from the two other species of Metzgeria from the area by its
multispiral larval shell, and its broader teleoconch with shouldered whorls.

Genus LA TIR OMITRA Locard, 1897

Type species: L. specialis Locard, 1897, by monotypy.

Synonym: Cyomesus Quinn, 1981. Type species Mesorhytis meekiana Dali, 1889.

Remarks: The systematic position of Latiromitra has been changed several times. Locard placed it in
Pisaniidae (i.e. Buccinidae!). Thiele (1929) transferred it to Costellariidae. Cernohorsky (1970)
considered it a subgenus of Volutomitra, hence a volutomitrid.

The radulae of two conchologically very similar species, Teramachia chaunax Bayer, 1971 and
Mesorhytis meekiana Dali, 1889 have been described by Bayer (1971). They are of turbinellid type
and Bayer concluded that the species belonged to the Turbinellidae. He therefore moved Teramachia
to Turbinellidae. The type species of Teramachia, however, is T. tibiaeformis Kuroda, 1931, of
which the radula has been figured by Habe (1952). There seems to be no doubt that the type species
is a volutid.

Rehder (1972) and Cernohorsky (1973) suggested that T. chaunax, Mesorhytis meekiana and two
other species, Mesorhytis costatus Dali, 1890 and Teramachia barthelowi (Bartsch, 1942) should be
placed in Benthovoluta Kuroda & Habe. Finally Quinn (1981) considered these species distinct
enough to separate them from Benthovoluta and suggested a new genus, Cyomesus, with meekiana
Dali as type species.

We have examined the radula of T. barthelowi (Fig. 390) from the Philippines and can confirm its
affinity to chaunax and meekiana. We cannot, however, see any conchological differences between
these four species and Latiromitra that can be used to maintain two genera, and consider Cyomesus
a synonym of Latiromitra. Presumably no one considered Latiromitra as a genus for these species
because it was considered a volutomitrid.

Mitra bairdi Dali, 1889, from the bathyal area off N Carolina and the Gulf of Mexico is a typical
mitracean, as we found from examination of the radula.

Latiromitra cryptodon (P. Fischer, 1882) Figs 675-676

Mitra cryptodon P. Fischer, 1882:273.

Latiromitra specialis Locard, 1897:321, pi. 14, fig. 30-34.

Type material: The two names are based on the same two syntypes in MNHN.

Type locality: TRAVAILLEUR 1882, dr 40, 33°09 N, 09°38 W, 1900 m.

Materia! examined: The type material and MONACO st 683, 38°20 N, 28°05 W, 1550 m, 1 sh; st 116, 31 °43 N,
10°47 W, 2165 m, 1 sh.

Distribution: Only known from the material examined.

Remarks: The larval shell is multispiral and probably indicates planktotrophic larvae.

256 P. BOUCHET & A. WAREN

Figs 677-682. Genus Metzgeria. 677 and 678, M. alba, INGOLF st 32, 19.8 mm and 2E4 mm. 679, M.
gagei, holotype. 680 and 681, M. gagei, INGOLF st 18, 14.1 mm and apex 1.6 mm. 682, M. apodema,
holotype.

NORTH-EAST ATLANTIC NEOGASTROPODA 257

Family CANCELLARIIDAE

Our collection of cancellarids is rather small. The family is not common in deep water, judging from
the literature. Clarke (1962) cites five species from below 1830 m (of which Admete aquilarum
(Watson) is a turrid). A few species have been described by Barnard (1960), Okutani (1964), Dali
(1908), Verrill (1885) and Knudsen (1964).

Several cancellarids were originally described as trichotropids.

The genus Benthobia which previously has been placed in Cancellaridae is here transferred to
Olividae (p.249).

Genus ADMETE Kroyer in Moller, 1 842

Type species: A. crispa Moller, 1842 ( = Tritonium viridulum Fabricius, 1780) by monotypy.

Remarks: There has been a controversy about the type species of Admete , since Dali (1887) wrote
that Tritonium viridulum Fabricius and Defrancia viridula Moller, 1842 were based on the same
specimen and suggested that T. viridulum was a turrid. European authors have continued to use the
name of Fabricius, while American authors have used the name Cancellaria couthouyi Jay, 1839
(replacement name for Cancellaria buccinoides Couthouy, 1838, not Sowerby, 1832).

Dali (1887) pointed out that this had been mentioned by Morch and that he had examined the
specimen in Copenhagen. Morch never mentioned this in any paper and even the last time he used
the name viridula (1877) shortly before his death, he used it in the accustomed sense. Posselt (1895)
mentioned Dali’s doubts on the validity of the name viridula and that it was caused by a catalogue
note by Morch, but evidently Posselt considered it of no importance, because he continued to use
the name Admete viridula. At this time there were no types of Admete crispa Moller or Tritonium
viridulum Fabricius in the museum, because they were not mentioned by Posselt and he always
listed the type material when it had been found. (There is, however, still alcohol material
determined “ Admete viridula Fabricius” by Moller.) Therefore we suppose that Moller (1842) had
misidentified Fabricius’s Tritonium viridulum and later, when he realized this, began to use this
name for his former Admete crispa.

The original description (Fabricius, 1780) also supports our view that T. viridulum was not based
on a turrid, especially the comparison with Buccinum undatum and the mention of a very short
siphonal canal.

To stabilize the nomenclature, we have therefore selected one of Moller’s specimens as neotype of
T. viridulum (Fig. 683). This solution will make it possible to keep the name, which has been used in
a single sense in Europe for 200 years.

The genus Admete contains a few species from high northern and southern latitudes. Many of the
species which have been placed here, actually belong to other groups.

We have included three species in Admete, the type species and A. nodosa and A. azorica. The
two last-mentioned species probably do not belong to Admete, but we have not been able to find a
genus for them among the numerous cancellariid genera that have been described and we hesitate to
introduce a new name before the other names have been thoroughly revised. Therefore we have kept
them here.

258 P. BOUCHET & A. WAREN

Admete viriduia (Fabricius, 1780)

Figs 683-689

Tritonium viridulum Fabricius, 1780:402.

Murex cos tel lifer Sower by, 1818:225.

Cancellaria huccinoides Couthouy, 1838:105 (not Sowerby, 183*2).

Cancellaria couthouyi Jay, 1839:77.

Admete crispa Moller, 1842:88.

Admete borealis A. Adams, 1855:122.

Admete viriduia var. grandis Morch, 1869:22.

Admete abnormis Gray in H. & A. Adams, 1858:278, fide Morch 1869:22.

Admete undatocostata Verkriizen in Kobelt, 1876:372.

Admete viriduia var. product a G.O. Sars, 1878:217, pi. 13, fig. 2.

Admete viriduia vars. undata , laevior » elongaia and distincta Leche, 1878:47-48.

Admete contabulata Friele, 1879:276.

Admete sadko Gorbunov, 1946:310, pi. 1, fig. 5.

Type material: A. viriduia , neotype in ZMC.

Type locality : A. viriduia , W Greenland.

Material examined: INGOLF st 32, 66°35 N, 56°38 W, 599 m, 10 spms; st 95, 65°14 N, 30°39 W, 1416 m, 1 sh;
st 98, 65°38 N, 26°27 W, 260 m, 1 sh; st 124, 67°40N, 15°40 W, 932 m, 1 sh; st 126, 67° 19 N, 15°52W, 552 m, 1
sh, ZMC; several hundred shells and specimens from the Arctic Atlantic, USNM, SMNH, ZMC; several
thousand specimens from S Scandinavia, 75—1000 m, coll. AW.

Distribution : From Massachusetts to Arctic Canada (Macpherson, 1971), E and W Greenland
(Thorson, 1944, 1951), Iceland (Thorson, 1941), the Faroes (Spate k & Thorson, 1933), deep water
off the British Isles (Jeffreys, 1885; Jordan, 1895; Sykes, 1911; Massy, 1930), along the Scandinavian
coasts (G.O. Sars, 1878), Spitzbergen (Friele, 1886; Odhner, 1915), Kara Sea and Novaya Zemlya
(Leche, 1878). In arctic areas A viriduia occurs in shallow water, from intertidaily, down to 1100 m
(Friele, 1886), in the southern parts of its distribution considerably deeper, in S Scandinavia in
75-1200 m. Probably A. viriduia has a panarctic circumpolar distribution.

Remarks : For discussion about the specific name, see under the genus.

Admete viriduia is extremely variable, which is reflected in the large number of varieties that have
been described. Probably also several names from the N Pacific should be added but that is beyond
the intention of our work.

Young specimens of A. viriduia may easily be confused with young Oenopota spp., but Oenopota
has a more slender aperture, distinct siphonal canal, and an operculum. Nordsieck (1968) considered
A. inf lata (see Iphinopsis ) as a subspecies of A. viriduia , but that is not because they are similar, but
because he had never seen A. inf lata.

Thorson (1944) referred the egg capsules he had previously (1935) assigned to Velutina undata » to
Admete viriduia. We can confirm this observation and figure egg capsules and young from W
Norway.

Admete nodosa Yerrill, 1885

Fig. 691

Admete nodosa Verrill, 1885:419, pi. 44, fig. 9.

Type material: Holotype USNM 44646.

Type locality: USFC st 2234, 39°09 N, 72°03 W (off New Jersey), 1475 m.

Material examined: The holotype and THALASSA st X 333, 44° 10 N, 04°32 W, 1925 m, 1 spm; st X 336, 44° 1 1
N, 05° 11 W, 1900 m, 1 spm; INCAL st DS 5, 56°28 N, 1 1°12 W, 2503 m, 1 spm; BIOGAS st CP 07, 44° 10N,
04° 16 W, 2170 m, 2 shs; st CP24, 44°08 N, 04° 16 W, 1995 m, 1 sh; POLYGAS st CV16, 44°06 N, 04° 17 W, 1909
m, 2 spms.

Distribution: Only known from the material examined, from both sides of the N Atlantic, 1400-
2500 m.

NORTH-EAST ATLANTIC NEOGASTROPODA 259

Figs 683-686. Admete viridula. 683, neotype, 13.7 mm. 684, holotype of Admete contabulata ,
VORINGEN st 192, 8.1 mm. 685, Spitzbergen, 26.0 mm. 686, Sacken, Swedish west coast, 10.6 mm.

260 P. BOUCHET & A. WAREN

Figs 687=689, Admete viriduia, 687, Lieholmsrannan, Bergen area, 55-65 m, egg-capsule. Length 5 mm,
688, shell of an embryo from the same capsule, 870 pm. 689, Korsfjorden, Bergen area, 306 m. Very
young specimen with postlarval growth, 1.0 mm.

Remarks: The larval shell is multispiral and indicates planktotrophic larvae.

A, nodosa does not resemble any N Atlantic gastropod known to us, except A. azorica, but that
species is smaller, more slender and has a proportionally smaller aperture.

A. nodosa has no radula. The rectum and intestine of the specimen which was examined for a
radula, were filled with droplets of some intensely red substance.

Admete azorica sp.n.

Fig. 690

Type material: Holotype in MNHN.

Type locality: BIACORES st 53, 38°07 N, 28°18 W, 1820 m, SE of Pico, Azores.

Material examined: MONACO st 112, 38°34 N, 28°06 W, 1287 m, 1 sh; st 1349, 38°35 N, 28°06 W, 1250 m, 1 sh;
TALISMAN dr 129, 38°00 N, 27°03 W, 2220 m, 1 sh.

Distribution: Only known from the material examined, from the Azores.

Description: Shell short, conical, rather broad, very solid, white. The larval shell consists of 2.25
whorls, sculptured by fine spiral lines and its height is about 850 pm. The holotype has 2.8 postlarval
whorls which are rather flat and sculptured by strong axial ribs; 1 1 ribs occur on the body whorl. There
are also numerous irregular growth lines. The spiral sculpture consists of three often interrupted spiral
lines on the upper whorls and two additional ones on the body whorl. The aperture is rather small with
two large columellar folds and a strongly oblique outer lip.

NORTH-EAST ATLANTIC NEOGASTROPODA 261

Dimensions: Height of the shell 7.6 mm, diameter 4.6 mm, height of the aperture 4.0 mm, breadth.
2.1 mm.

Remarks: A azorica differs from A. nodosa by its more slender shell and possibly its larval shell has
about half a whorl less, but this could not be positively ascertained because of the poor preservation of
all larval shells of A. nodosa. Furthermore, it reaches only half the size of A. nodosa.

Genus IPHINOPSIS Dali, 1924

Type species: Iphinoe kelseyi Dali, 1908, by original designation (Fig. 694).

Synonym: Iphinoella Habe, 1958. Type species I. choshiensis Habe, 1958, by original designation.

Remarks: Iphinopsis has previously, from shell characters, been placed in Trichotropidae, but on
evidence given below we transfer it to Cancellariidae.

There have been described several species resembling I. kelseyi in various families and genera:
Trichotropis inf lata Friele, 1879, from deep water N of Norway.

Trichotropis nuda Dali, 1927, from deep water off Florida (Fig. 704).

Iphinoella choshiensis Habe, 1958, off Japan, 500-1500 m.

Tromina traversensis Clarke, 1959, from the abyssal S Atlantic.

Cancellaria euthymei Barnard, 1960, from off S Africa.

We have examined specimens of kelseyi, inflata, nuda, traversensis and two species that are
described as new here, and found them to be very similar conchologically. We have also examined
the anatomy of four of these, inflata, traversensis and the two new species. They were identical,
except in the development of the eyes and sexual characters. The examination was made on dried
specimens reconstituted by soaking in a water detergent solution, and we noticed that after soaking,
all were extremely slimy and that the oviducts had swelled, producing a lot of mucus.

The foot is large and muscular and probably very large when the animal is crawling. The
propodium is distinct and there is no operculum. The tentacles are short and rounded with eyes
situated laterally at their base. The osphradium is large, bipectinate, of at least the same size as the
gill. The animal has no snout, a long pleurembolic proboscis with two pairs of salivary glands, but
no radula. No identifiable contents were found in the rectum or intestine. The diameter of the eyes
of I. traversensis was 40 pm, of alba 100 pm, of inflata 130 pm and of fuscoapicata 150 pm, in
specimens of the same size. The specimens of inflata, traversensis and fuscoapicata were females
with a well-developed pallial oviduct, while two specimens of I. alba had both penis and pallial
oviduct.

From these anatomical details and the presence of columellar folds, we conclude that the species
belong to Cancellariidae. Iphinopsis choshiensis Habe is the type species of Iphinoella Habe and I.
kelseyi Dali, is the type of Iphinopsis. Habe (1962) synonymized the two genera and considered them
to belong to Trichotropidae, which we find untenable on the basis of absence of radula, absence of
operculum, absence of snout, presence of pleurembolic proboscis and presence of columellar folds
in the species we have examined. We admit that we have not examined the animal of either of the
two type species, but conchologically they are very close to those which we have examined.

The species of Iphinopsis are easy to confuse with other groups of gastropods, as is evident from
the known species being described in three different families. Clarke (1961) pointed out that T.
traversensis could be confused with the young of Oocorys, but described it as a buccinid in the genus
Tromina Dali, 1918 (Fig. 328), probably misled by Powell (1951) who used Tromina for certain
antarctic buccinids. Iphinopsis can also be confused with the young of some cassids, some
buccinids and trichotropids. The species of Iphinopsis can, however, be distinguished by having
columellar folds, which may be very indistinct and visible only rather far back in the aperture. Some
turrids of the genus Lusitanops also resemble Iphinopsis, but they have a less well developed
umbilicus and a larval shell with cancellate sculpture.

262 P. BOUCHET & A. WAREN

Iphinopsis inf lata (Friele, 1879)

Figs 698-699

Trichotropis inflata Friele, 1879:275.

Admete inflata Friele, 1886:25, pi. 8, fig. 33.

Type material : Two syntypes, ZMB 20719.

Type locality: VORINGEN st 192, 69°46 N, 16°15 E, 1187 m.

Material examined : The type material and 20 shs from the type locality ZMB 20720; VORINGEN st 323, 72°53
N, 21 °51 E, 408 m, 2 spms; st 312, 74°54 N, 14°53 E, 1203 m, 6 spms, ZMB 20721; INGOLF st 90, 64°45 N,
29°06 W, 1070 m, 1 sh; st 102, 66°32 N, 10°26 W, 1412 m, 3 shs; st 103, 66°23 N, 08°52 W, 1090 m, 2 spms, 1
sh; st 116, 70°05 N, 08°26 W, 699 m, 6 spms; st 124, 67°40 N, 15°40 W, 932 m, 1 sh, 2 spms; st 139, 63°36 N,
07° 30 W, 1322 m, 16 spms, ZMC; 73°03 N, 18°30 E, 410 m, 1 spm, SMNH.

Distribution: Only known from between Norway and Spitzbergen and around Iceland, from the
material examined. VerrilFs (1884) record from off New England in 3200 m needs confirmation and
we were not able to locate his material in USNM.

Remarks: I. inflata differs from /. alba by having less developed, almost invisible columellar folds,
while those of /. alba are very distinct, especially the lowermost one. It differs from I. fuscoapicata
by having a white larval shell of about one whorl, instead of a brownish, multispiral larval shell.
To see the columellar folds of I. inflata it is necessary to look rather obliquely into the aperture.

Iphinopsis fuscoapicata sp.n.

Figs 700-703

Type material: Holotype in MNHN.

Type locality: INCAL st WS01, 50° 19 N, 13°07 W, 2550 m (SW of Ireland).

Material examined: INCAL DS06, 56°26 N, 1 1 ° 10 W, 2494 m, 5 spms; CP06, 55°02 N, 12°41 W, 2890 m, 1
spm; DS09, 55°08 N, 12°53 W, 2897 m, 1 spm; CP08, 50°15 N, 13°14 W, 2644 m, 1 spm; WS02, 50°19 N, 12°56
W, 2498 m, 1 spm; CHALLENGER 2 st 4, 56°52 N, 10°01 W, 1993 m, 1 spm; st ?, Rockall Trough, 2851 m, 1
spm.

Distribution: Only known from the material examined, off the British Isles, ca. 2500 m.

Description: Shell of medium size, fragile, inflated, dirty white. The larval shell is dark brownish and
consists of about two whorls, sculptured by faint spiral striae, and its height is about 650 pm. The
holotype has about three postlarval whorls of rather rapidly increasing diameter. The sculpture
consists of equidistant spiral cords, weaker on the lower part of the body whorl and almost
disappearing at the columella. The axial sculpture consists of a few stronger and numerous small,
sharp incremental ribs. The stronger ribs appear at irregular intervals, the smaller ones are close set
and regularly spaced. The outer lip, seen from the side, is slightly sigmoid. The inner lip forms a far-
protruding, evenly-curved parietal callus, continuous with the reflected part of the columella and
forming a deep umbilical chink behind it. There is no distinct siphonal canal.

Dimensions: Height of the shell 9.5 mm, diameter 6.0 mm, height of the aperture 6.0 mm,
breadth 2.9 mm.

Remarks: The present species can be distinguished from the two other species known from the NE
Atlantic by the larval shell, which is brownish in I. fuscoapicata and white in the others.

/. traversensis probably has a brownish larval shell, but the apex of the holotype is badly
corroded, so we could see only traces of this, in the old sutures. It differs from /. fuscoapicata by
having a finer and less distinct axial sculpture with more distant ribs and by having more convex and
strongly shouldered whorls.

There is hardly any columellar fold visible in the specimens we have examined, but this may be
because our specimens are not full grown.

NORTH-EAST ATLANTIC NEOGASTROPODA 263

Ip h inops is alba sp. n.

Figs 695-697

Type material: Holotype in MNHN.

Type locality: BIOGAS st CP01, 47°35 N, 08°39 W, 2245 m, NW part of the Bay of Biscay.

Material examined: BIOGAS st DS26, 47°33 N, 08°36 W, 2147 m, 2 shs; st DS52, 44°06 N, 04°22 W, 2006 m, 3
spms; st DS62, 47°33 N, 08°40 W, 2175 m, 1 sh; st DS63, 47°33 N, 08°35 W, 2126 m, 1 sh; st CP08, 47°33 N,
08°38 W, 2177 m, 1 spm; st CP12, 47°32 N, 09° 12 W, 2925 m, 1 spm; st CP25, 44°05 N, 04° 17 W, 1894 m, 1
spm; st DS71, 47°34 N, 08°34 W, 2194 m, 1 spm, 2 shs; st DS64, 47°29 N, 08°31 W, 2156 m, 1 spm, 1 sh;
THALASSA st X333, 44° 10 N, 04°32 W, 1925 m, 1 sh; st X334, 44° 10 N, 04°52 W, 1900 m, 1 sh; st X336, 44° 1 1
N, 05° 11 W, 1900 m, 1 sh; CHALLENGER 2, st 4, 56°52 N, 10°01 W, 1993 m, 1 spm.

Distribution: Only known from the continental slopes of the bay of Biscay and the Rockall Trough,
1900-2900 m.

Description: Shell of medium size, rather inflated, fragile, whitish, or dirty whitish. Larval shell
white, sculptured by a few rather coarse spiral lines, consisting of about 1 .2 whorls and of a height of
600 ^m. The holotype has about 4 postlarval whorls, but this figure is uncertain because of corrosion
of the spire. The sculpture is identical to that of fuscoapicata, possibly some specimens have coarser
axial sculpture, but this is difficult to ascertain from our small sample. The shape of the aperture and
the columella are also identical with fuscoapicata.

Dimensions: Height of the shell 14.5 mm, diameter 8.2 mm, height of the aperture 9.5 mm,
breadth 3.8 mm.

Remarks: Iphinopsis alba can be separated from /. fuscoapicata only by the larval shell (Figs 697,
702), which in /. alba has coarser sculpture, about 3/4 of a whorl less, and is white instead of brown.
Another difference, more difficult to see in juvenile specimens, is that /. alba has distinct columellar
folds while /. fuscoapicata has no trace of them. This is also the only difference from /. inf lata where
only traces of one fold can be seen.

Genus OLSSONELLA Petit, 1970
Type species: Cancellaria smithi Dali, 1883, by original designation.

Remarks: The type species was described from shallow water (35-80 m) off N Carolina. Dali (1889)
compared this species with Cancellaria minima Reeve, because of their similarity. We have used the
genus Olssonella on the advice of Mr R.E. Petit, with whom we have had valuable discussions about
the cancellariids.

Olssonella minima (Reeve, 1856)

Figs 692-693

Cancellaria minima Reeve, 1856: sp. 77.

Type material: Syntypes in BMNH.

Type locality: No locality mentioned.

Material examined: The types and: off Madeira, 410-580 m, 1 sh, MNHN; PORCUPINE 1870, off Cadiz, SW
Spain, 700 m, 7 shs, BMNH.

Distribution: The area W of Gibraltar, kriown from 600-700 m; Madeira, in shallow water (Watson,
1897).

Remarks: Cancellaria minima was described by Reeve without giving the locality but Jeffreys (1885)
said that the specimens originated from the dredgings of McAndrew and recorded it from Gibraltar
and Madeira on the authority of McAndrew. McAndrew (1852, 1857) also listed undescribed
cancellariids from Madeira and the Canaries. As Watson’s (1897) records from Madeira indicate,
this will probably prove to be a shallow-water species.

264 P. BOUCHET & A. WAREN

Figs 690-693. Genera Admete and Olssonella. 690, A. azorica, holotype, 7.8 mm. 691, A. nodosa,
BIOGAS st CP24, 13.8 mm. 692, O. minima, syntype, 6.5 mm. 693, O. minima, off Madeira, apex 707

pm.

NORTH-EAST ATLANTIC NEOGASTROPODA 265

Figs 694-699. Genus Iphinopsis. 694, I. kelseyi, type species of Iphinopsis, holotype 6.4 mm. 695, /.
alba, INCAL st CP8, 9.3 mm. 696, I. alba, BIOGAS st CPI, 14.9 mm. 697, /. alba, apex 0.5 mm.
698 and 699, /. inflata, INGOLF st 103, 7.5 mm and 7.9 mm.

266 P. BOUCHET & A. WAREN

Figs 700-704. Genus Iphinopsis. 700, I. fuscoapicata , INCAL st DS7, 3.9 mm. 701, INCAL st WS1
holotype, 9.5 mm. 702, Young postlarvae, INCAL st DS6, 870 /xm. 703, Embryonic shell of the same
specimen, diameter 390 /xm. 704, I nuda, syntype of Trichotropis nuda Dali, 2.9 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 267

Family MARGINELLIDAE

Examination of literature suggests that marginellids seem to be scarce in deep water (Clarke, 1962
lists four species). Our experience, however, is that they are not rare on the upper part of the
continental slopes, especially in tropical areas.

Several species of marginellids were described by Locard (1897) from just south of the area we
are treating. No further material has since been found so we refer to his treatment of these species.
A key to the species likely to be encountered in the area covered is presented below.

A.

A.

B.

B.

C.

C.

D.

D.

E.

E.

F.

F.

G.

G.

H.

H.

I.
I.

Spire visible C

Spire hidden (Granulina) B

Shell slender, breadth = 2/3 height G. parvulina

Shell broad, breadth = 4/5 height G. occulta

High spire, four or fewer equal columellar folds E

Short spire, more than four, unequal, columellar folds
(Gibberula) D

Shell small (2-2.5 mm), 5-6 columellar folds G. vignali

Shell bigger (3-5 mm), 4-5 columellar folds G. abyssicola

Three columellar folds (Volvarina) V. ingolfi

Four strong columellar folds (Marginella) F

Outer lip thickened into a broad varix, shell white to
yellowish white G

Outer lip not thickened into a varix, shell yellowish white to
orange I

Shell small (less than 10 mm), with very convex whorls M. hesperia

Shell larger, with almost flat whorls H

Suture polished, indistinct M. marocana

Suture distinct M. arronax

Hire spire with very convex whorls M subturrita

Shell broad, with almost flat whorls M. impudica

Genus GIBBERULA Swainson, 1840
Type species: Gibberula zonata Swainson, 1840, by monotypy.

The genus Gibberula comprises small marginellids, 3-10 mm high, that live mainly on the continental
shelf. Two species occur on the Atlantic slope of the Iberian peninsula, but do not enter the
Mediterranean where only shallow-water species are found.

Gibberula abyssicola Monterosato in Locard, 1897 Figs 394, 705-708, 714
Gibberula abyssicola Monterosato in Locard, 1897:130, pi. 4, figs 22-25.

Gibberula monterosatoi Locard, 1897:131, pi. 4, figs 26-28.

Gibberula retusa Monterosato in Locard, 1897:128, pi. 4, figs 19-21.

Type material: G. abyssicola , lectotype in MNHN; G. monterosatoi, 4 syntypes in MNHN; G.
retusa, 37 syntypes in MNHN.

Type localities: G. abyssicola, TRAVAILLEUR 1881 dr 40, 40°05 N, 07°05 W, 392 m; G.
monterosatoi, TRAVAILLEUR 1881 dr 1, 43°01 N, 09°38 W, 2018 m; G. retusa, TRAVAILLEUR
1881 dr 30, 35°25 N, 08° 00 W, 1205 m.

Material examined: The type material and TRAVAILLEUR 1881, dr 41, 44°02 N, 07°07 W, 1094 m, 2 shs; dr
42, 44°01 N, 07°05 W, 896 m, 8 spms -t- shs; THALASSA st W392, 44°07 N, 04°49 W, 600-1130 m, 1 spm; st
X305, 44° 05 N, 05°00 W, 463 m, 18 spms + shs; st X340, 44°07 N, 04°30 W, 860-910 m, 10 spms; st X341,
44° 07 N, 04° 31 W, 800-840 m, 2 spms; st X342, 44°08 N, 04°37 W, 700 m, 14 spms + shs; st X343,, 44°08 N,
04° 39 W, 600-655 m, 16 spms + shs; st X345, 44°06 N, 04°41 W, 525-550 m, 31 spms + shs; st X347, 44°07 N,
04° 44 W, 640-910 m, 36 spms + shs; st X348, 44°07 N, 04°44 W, 600-900 m, 4 shs; st X350, 44°04 N, 04°46 W,
505 m, 1 spm; st X352, 44°06 N, 04°45 W, 545-580 m, 5 spms; st X353, 44°07 N, 04°45 W, 635-655 m, 6 spms +

268 P. BOUCHET & A. WAREN

shs; st X360, 44°05 N, 04°50 W, 580-592 m, 1 sh; st X361 , 44°06 N, 04°50 W, 582-595 m, 7 spins; st X362, 44°07
N, 04° 51 W, 585-600 m, 18 spms + shs; st X363, 44°06 N, 04°54 W, 545-630 m, 8 spms; st Y377, 41°32 N,
09° 14 W, 320 m, 3 shs; st T482, 44°02 N, 08°44 W, 490 m, 4 spms; MONACO st 2717, 36°42 N, 08°40 W, 750
m, 53 spms -I- shs.

Distribution: Only known from the material examined, NW and S of the Iberian Peninsula, on the
upper part of the continental slope.

Remarks: G. abyssicola was reported from the Mediterranean (on the authority of Monterosato) by
Locard (1897). It was however, not mentioned by Monterosato or D’Amico in any paper. G. Spada,
who has examined the marginellids in the Monterosato collection has informed us that it is not
present there, except two shells from the TRAVAILLEUR dredgings, nor are any specimens of retusa
or monterosatoi; we conclude that there was some mistake behind Locard’s statement. The material
cited from shallow water in Madeira proved to belong to Granulina.

The name G. monterosatoi was based on young specimens without thickened outer lip.

The specimens examined from Le Danois Bank (THALASSA X stns) show a reduction in size
with increasing depth. The average size from 450 m is 4.3 mm, in 880 m, 3.3 mm. The name retusa
was based on more southern specimens, which have a more thickened outer lip and solid shell.

The egg capsule (Fig. 714) is a simple rounded capsule, slightly larger than the larval shell. It lies
free on the bottom. The size of the young corresponds with our smallest specimens, and indicates
direct development.

Fig. 714. Gibberula abyssicola. THALASSA st X315, egg capsule, length 1.9 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 269

Gibberula vignali (Dautzenberg & H. Fischer, 1896) Figs 709-710

Marginella vignali Dautzenberg & H. Fischer, 1896: 433, pi. 15, fig. 17.

Type material: lectotype in MOM.

Type locality: MONACO st 553, 37°43 N, 25°05 W, 1385 m.

Material examined: The type material and 7 shs from the type locality; MONACO st 1 12, 38°34 N, 28°06 W,
1287 m, 11 shs; st 1311, 37°37 N, 25°21 W, 1187 m, 2 shs; st 1349, 38°35 N, 28°06 W, 1250 m, 13 shs;
TRAVAILLEUR 1882, dr 34, 35°42 N, 06°20 W, 1 12 m, 2 shs; THALASSA st Y400, 40°46 N, 09° 19 W, 800 m,
5 spms; VANNEAU 1923, st 10, 29°54 N, 09°58 W, 110 m, 18 shs; st 35, 34°05 N, 07°40 W, 160 m, 9 shs.

Distribution: Only known from the material examined, off Portugal and Morocco, and the Azores,
on the continental shelf and upper parts of the slope.

Remarks: Our experience from shallow-water species of Gibberula shows that there exists very little
difference in shell characters between the species, and that living animals may be necessary to
distinguish the species. We will therefore not exclude the possibility that we may have confused two
species in C. vignali.

Gibberula vignali differs from abyssicola by being much smaller, about 2.5—2 mm instead of
3—4.5 mm, and by having five instead of four clearly visible columellar folds.

Genus GRANULINA Jousseaume, 1888

Synonym: Cypraeolina Cerulli-Irelli, 1911
Type species: Voluta clandestina Brocchi, 1814.

The group of marginellids treated here has consistently been called Gibberulina in the European
literature. However Gibberulina Monterosato, 1884 is a new name (“nom.sost.”) for Bui lata
Jousseaume, 1875 and therefore keeps the same type species, viz. Voluta hullata Born, 1778.
Bullata is a valid name and Gibberulina is an objective synonym of it.

Granulina was introduced for Marginella pygmaea Issel, 1869 (type species by monotypy), non
Sowerby 1846 (= Marginella isseli Nevill & Nevill, 1875) and can be used for the species formerly
included in Gibberulina in Europe.

Granulina parvulina (Locard, 1897)

Fig. 711

Volutella parvulina Locard, 1897:126, pi. 21, figs 3-5.

Volutella parvulina var. curta Locard, 1897:126.

Type material: Two syntypes in MNHN.

Type locality: TRAVAILLEUR 1881, dr 41, 44°02 N, 07°07 W, 1094 m.

Material examined: The type material and MONACO st 1713, 28°04 N, 16°49 W, 1340-1530 m, lsh;
VANNEAU 1923, st 10, 29°54 N, 09°58 W, 110 m, 2 shs.

Distribution: Only known from the material examined, off N Spain and Morocco.

Remarks: G. parvulina differs from the shallow-water species of the genus by being much more
slender. G. occulta (Monterosato, 1869) is a deep-shelf species occasionally occurring on the upper
slope in the Mediterranean and nearby Atlantic; it is figured here (Fig. 712) for comparison.

270 P. BOUCHET & A. WAREN

Genus MARGINELLA Lamarck, 1799
Type species: Marginella glabella (Linne) by monotypy.

The type species.of Marginella as well as four of the species discussed here lack a radula. The shape
of the shell of M. glabella resembles the four species, but the colour patterns are quite different. The
fifth species included in our revision, Marginella hesperia differs in shell characters; no animal was
available for examination for a radula, and it seems to be more closely related to certain deep shelf
species from West Africa, e.g. M. bojadorensis Thiele, 1925.

Marginella hesperia Sykes, 1905 Fig. 716

Marginella hesperia Sykes, 1905: 316, pi. 17, fig. 7.

Type material: 10 syntypes in BMNH.

Type locality: PORCUPINE st 24, 37° 19 N, 09° 13 W, 531 m.

Material examined : The type material and some more shells from the type locality (USNM);
JEAN-CHARCOT Madere st 29, 33°01 N, 16°15 W, 300-340 m, 2 shs; st 49, 32°27 N, 16°32 W, 450-500
m, 1 sh; st 58, 32°42 N, 16°41 W, 410-580 m, 1 sh.

Distribution: On the upper part of the continental slope, from off S Portugal and Madeira. Only
known from the material examined.

Remarks: Marginella hesperia can be recognized by the thickenings of the outer lip, which are
lacking in all other species known from the area treated. Attention must however, be drawn to M.
bojadorensis Thiele, 1925 from West Africa, which has similar thickenings on the outer lip but is
more slender.

Marginella subturritaP . Fischer, 1883 Fig. 717

Marginella impudica var. subturrita P. Fischer, 1883:392.

Marginella jousseaumei Locard, 1897: 11, pi. 3, figs 25-28.

Type material: M. impudica var. subturrita and M. jousseaumei are based on the same material. A
single lectotype in MNHN is designated for both.

Type locality: Not known precisely, but between Morocco and the Canaries.

Material examined: The type material and TALISMAN dr 45, 29°08 N, 12°26 W, 1235 m, 5 spm + shs; dr
49,52,53 (mixed), 28°33-28°37 N, 13°02-13°19 W, 865-946 m, 6 spms + shs; MONACO st 1118, 29°06 N, 13°03
W, 1098 m, 1 spm.

Distribution: The continental slope of western Sahara. Only known from the material examined.

Remarks: The animals lack a radula. It has eye swellings at the base of the tentacles and the
unpigmented eyes are 130 jum in diameter, while they are only 50 jum in M. impudica. M. subturrita
differs from M. impudica by being more slender, by its much more convex whorls and white shell,
while M. impudica is pinkish-orange when fresh.

Marginella marocana Locard, 1897 Figs 722-723

Marginella marocana Locard, 1897: 114, pi. 3, figs 29-31 (and var. curta, var. elongata).

Type material: Lectotype here selected in MNHN.

Type locality: TALISMAN st 49-52-53 (mixed lot), 28°33-28°37 N, 13°02-13°19 W, 865-946 m.
Material examined: The type material and 1 spm from the type locality; TALISMAN dr 44, 29°52 N, 1 1°47
W, 2083 m, 1 spm, 1 sh; dr 45, 29°08 N, 12°26 W, 1235 m, 3 shs.

Distribution: Only known from the material examined.

Remarks : See under aronnax.

NORTH-EAST ATLANTIC NEOGASTROPODA 271

Marginella impudica P . Fisher, 1 883 Figs 718-719

Marginella impudica P. Fischer, 1883:392.

Marginella impudica var. major, var. elongata Locard, 1897:109, pi. 3, figs 19-24.

Type material: Lectotype, here selected, in MNHN.

Type locality: “Cotes du Sahara, 800-1139 m”. Lectotype from TALISMAN dr 81, 23° 50 N, 17° 17
W, 1139 m.

Material examined: The type material and 4 spms from the type locality; TALISMAN dr 84, 22°54 N, 17°26 W,
860 m, 7 spms + shs; dr 85, 22°52 N, 17°23 W, 830 m, 1 sh, 5 spms; dr 86, 22°49 N, 17°21 W, 800 m, 5 spms, 1
sh; METEOR st 36-98, 25°31 N, 16°02 W, 700-900 m, 6 spms + shs.

Distribution: The continental slope of Mauretania and former Spanish Sahara. Only known from
the material examined.

Remarks: The animal lacks a radula. There are eye swellings at the base of the tentacles and the
unpigmented eyes measure 50 pm in diameter.

Neither M. impudica nor M. subturrita has a broad thickened labial varix when seen from the
side. For differences see remarks under subturrita.

Marginella aronnax n.sp. Figs 720-721

Marginella impudica var. minor, var. curta and var. marginata Locard, 1897:110 (not: Marginella curta
Sowerby, 1832, Marginella marginata Wood, 1828, Marginella minor C.B. Adams, 1852).

Type material: Holotype in MNHN.

Type locality: TALISMAN dr 84, 22°54 N, 17°26 W, 860 m.

Material examined: The type material and 1 sh from the type locality; TALISMAN dr 86, 22°49 N, 17°21
W, 800 m, 7 spms, 2 shs.

Description: Shell white, very solid, smooth, with a short blunt spire. The shell consists of 4.5
whorls which are very slightly convex, with a hardly visible suture. The body whorl occupies more
than 3/4 of the total height. Aperture elongate, narrow with a broad thickened labial varix when
seen from the side. There are four strong columellar folds of which the lowermost one is very
oblique. No columellar callus. Dimensions of the shell: height 28.7 mm, breadth 14.8 mm; height
of the aperture 21.2 mm, maximum breadth 3.3 mm.

Remarks : The animal is blind, without eye swellings on the tentacles. It lacks a radula.

Marginella aronnax is readily separated from M. impudica and M. subturrita which lack the broad
thickening of the labial varix. M. marocana is smaller with a much more obtuse spire, nearly flat
whorls and indistinct suture.

272 P. BOUCHET & A. WAREN

Genus VOLVARINA Hinds, 1844

Type species: Marginella (Volvarina) nitida Hinds, 1844.

Volvarina is here used in its conchological sense to include a new species that differs from
Marginella s.s. by its more slender shell with 3 instead of 4 columellar plaits. Volvarina species
have a radula while at least some of the Marginella spp. do not.

Volvarina ingolf i n.sp. Fig. 715

Type material : Holotype in ZMC.

Type locality: INGOLF st 92, 64°44 N, 32°52 W, 1776 m.

Material examined: Only known from the holotype.

Description: Shell solid, white, glossy consisting of 3.75 whorls which increase very rapidly in size;
apex blunt; spire low, the body whorl forming four fifths of the height of the shell; the penultimate
whorl is convex, but forms a concave slope below the suture; there are three strong columellar folds;
columellar callus very thin; outer lip regularly thickened, smooth, slightly constricted in the middle.
Dimensions: height 14.5 mm, breadth 7.6 mm; height of the aperture 11.7 mm, breadth 2.2 mm.
Remarks: There are additional Volvarina species in deep water south of the Canaries; all of them
however have a lower spire with less convex whorls, and hardly reach 10 mm. They also differ in
colour patterns, the non-banded species being uniformly orange or yellowish. Volvarina ingolfi is
the northernmost Atlantic marginellid.

NORTH-EAST ATLANTIC NEOGASTROPODA 273

Figs 705-713. Genera Gibberula and Granulina. 705, Gibberula abyssicola, MONACO st 2717, 2.9 mm.
706, THALASSA st X347, 3.3 mm. 707, THALASSA st T482, 4.3 mm. 708, MONACO st 2717, 2.9 mm.
709 and 710, Gibberula vignali, THALASSA st Y400, 2.3 mm and 2.2 mm. 711, Granulina parvulina,
MONACO st 1713, 2.1 mm. 712, Granulina occulta, Porto Empedocle, Sicily, 2.5 mm. 713, MONACO
st 2717, 2.1 mm.

274 P. BOUCHET & A. WAREN

Figs 715-719. Genera Volvarina and Marginella. 715, Volvarina ingolfi, holotype, 14.5 mm. 716,
Marginella hesperia, PORCUPINE st 24, 7.2 mm. 717, M. subturrita, lectotype, 34.0 mm. 718 and
719, M. impudica, TALISMAN dr 86, 31.6 mm.

NORTH-EAST ATLANTIC NEOGASTROPODA 275

Figs 720-723. Genus Marginella. 720 and 721, M. aronnax, holotype, 28.7 mm. 722 and 723, M
marocana, lectotype, 19.3 mm.

276 P. BOUCHET & A. WAREN

ACKNOWLEDGMENTS

We want to direct our thanks to the following persons who have contributed to this revision by
placing material at our disposal: R. Aldred (Institute of Oceanographical Sciences, Surrey), P.
Arnaud (Station marine d’Endoume, Marseille), KJ. Boss (MCZ), L. Cabioch (Roscoff), F.
Carrozza (Soiana), G. Della Bella (Bologna), J. Gage (Oban), F. Giannini (Empoli), S. Gofas
(Paris), T. Gosliner (South African Museum, Cape Town), J. van Goethem (IRSN), D. Heppell
(RSM), R. Houbrick (USNM), R. Janssen (SMF), R. Kilias (ZMHU), J. Kjennerud (ZMB), J.
Knudsen (ZMC), M. Lagardere (La Rochelle), J.C. Mahe (ISTPM, St. Pierre), C. Monniot
(MNHN), O. Paget (NHMW), S. Palazzi (Modena), L. Pequegnat (Texas A & M Univ.), R. Petit
(North Myrtle Beach, USA), P. Piani (Bologna), G. Pinna (Milano), R. Olerod (SMNH), J.
Rosewater (USNM), M. Sibuet (COB), S. Smith (RSM), G. Spada (Bologna), C. Tabanelli
(Cotignola), F. Talavera (Sta Cruz de Tenerife), M. Taviani (Bologna), G. Testa (MOM), K. Way
(BMNH), R. Work (Miami).

A large part of our material has been sorted by Centre de Tri d’Oceanographie Biologique, Brest.
The SEM photography was done at Centre de Microscopic du CNRS, Paris by Mme Guillaumin.
The work has been supported by the Swedish Natural Science Research Council and Centre
National de la Recherche Scientifique (France).

NORTH-EAST ATLANTIC NEOGASTROPODA 277

REFERENCES

ABBOTT, R.T. 1974. American Seashells . 2nd edition. New York, Van Nostrand Reinhold Co. 663 pp.
ADAM, W. & GLIBERT, M. 1974. Contribution a la connaissance de Nassarius semistriatus (Brocchi, 1814).

Bulletin de T Inst i tut Royal des Sciences Naturelles de Belgique, Biologic, 50(3): 1-78.

ADAM, W. & GLIBERT, M. 1976. Observations sur le “groupe” de Nassarius clathratus (Born, 1778). Bulletin
de Tlnstitut Royal des Sciences Naturelles de Belgique, Biologie, 5 1(4): 1-69.

ADAM, W. & KNUDSEN, J. 1955. Note sur quelques especes de Mollusques nouveaux ou peu connus de
PAfrique occidentale. Bulletin de Tlnstitut Royal des Sciences Naturelles de Belgique, Biologie, 31(61): 1-25.
ADAMS, A. 1855. Descriptions of two new genera and several species of Mollusca from the collection of Hugh
Cuming, Esq. Proceedings of the Zoological Society of London, 23:119-24.

ADAMS, H. & ADAMS, A. 1853-58. The Genera of Recent Mollusca, 1. London, van Voorst, 484 pp.
ALDER, J. 1848. Catalogue of the Mollusca of Northumberland and Durham. Transactions of the Tyneside
Naturalists Field Club (1848):97-209.

ALTIMIRA, C. 1975. Moluscos testaceos recolectados en el litoral de la parte norte de la provincia de Gerona.
Investigacion Pesquera, 39(l):63-78.

ARADAS, A. 1847. Descrizione delle conchiglie fossili di Gravitelli presso Messina. Atti dell’ Accademia
Gioenia di Scienze Naturali di Catania, (2)4:57-88.

ARADAS, A. & BENOIT, L. 1876. Conchigliologia vivente marina della Sicilia e delle isole che la circondano.

Atti dell’ Accademia Gioenia di Scienze Naturali di Catania, (3)6:227-324.

D‘ATTILIO, A. 1978. A catalogue of Coralliophilidae. Festivus (San Diego Shell Club), 10(10):69-96.
AURIVILLIUS, C.W.S. 1885. Ofversigt dfver de af Vega Expeditionen insamlade arktiske Hafsmollusker.

Vega Expeditionens Vetenskapliga Iakttagelser 313-383.

BARNARD, K.H. 1959. Contributions to the knowledge of South African marine Mollusca, 2: Gastropoda
Prosobranchiata Rachiglossa. Annals of the South African Museum, 45:1-255.

BARNARD, K.H. 1960. New species of South African marine gastropods. Journal of Conchology, London,
24:438-42.

BARNARD, K.H. 1963. Contributions to the knowledge of South African Marine Mollusca, III. Annals of the
South African Museum, 47:1-199.

BARSOTTI, G. & FRILLI, G. 1970. Sulla presenza di Neptunea sinistrorsa (Deshayes) nel Mediterraneo.
Conchiglie, 6(5-6): 52-58.

BARTSCH, P. 1945. Morrisonella, a new genus of East Pacific deep sea Mollusks. Nautilus, 59(1):23.
BAYER, F.M. 1971. New and unusual mollusks collected by R/V John Elliott Pillsbury and R/V Gerda in the
Tropical Western Atlantic. Bulletin of Marine Science, 21:111-236.

BEAN, W. 1834. Fusus turtoni Bean and Limnea lineata Bean, two new and hitherto undescribed species of
shells. Magazine of Natural History, 7:493-94.

BELL, A. 1870. On some new or little-known shells & c. of the Crag. Annals and Magazine of Natural History,
(4)6:213-17.

BELLARDI, L. 1873. I Molluschi dei terreni terziari del Piemonte e della Liguria, parte 1. Memorie della Reale
Accademia delle Scienze di Torino, (2)27:1-264.

BENNETT, E.T. 1824. Description of an hitherto unpublished species of Buccinum, recently discovered at
Cork. Zoological Journal, 1:398-99.

BEU, A. 1970. Bathyal upper Miocene Mollusca from Wairarapa district, New Zealand. Transactions of the
Royal Society of New Zealand (Earth Sciences), 7:209-40.

BIVONA, A. 1832. Caraterri di alcune nuove specie di conchiglie. Effemeridi scientifiche e letterarie per la
Sicilia, 1(2): 16-24.

BIVONA, A. 1838. Generi e Specie di Molluschi. Palermo, Giornale Letterario, xiv + 16 pp., 1 pi.
BLAINVILLE, H.M.D. de 1830. Faune Francaise. Malacozoaires. Paris, Levrault, 320 pp.

BOT, J. 1972. Nueva especie del genero Nassa. Miscellanea Zoologica, 3(2): 15.

BOUCHET, P. 1977. Mise en evidence de stades larvaires planctoniques chez des Gasteropodes Prosobranches
des etages bathyal et abyssal. Bulletin du Museum national d’Histoire naturelle, (3)400:947-72.
BOUCHET, P. & FONTES, J.C. 1981. Migrations verticales des larves de Gasteropodes abyssaux: arguments
nouveaux dus a l’analyse isotopique de la coquille larvaire et postlarvaire. Comptes Rendus de TAcademie
des Sciences, Paris, (3)292:1005-1008.

BOUCHET P. & TALAVERA, F.G. 1981. A new Metzgeria from North-West Africa Bollettino

Ma nrnlnpirn 17*177-80

BOUCHET, P. & WAREN, A. 1979. Planktotrophic larval development in deep-water gastropods. Sarsia,
64:37-40.

BOUCHET, P. & WAREN, A. 1979. The abyssal molluscan fauna of the Norwegian sea and its relation to
other faunas. Sarsia, 64:211-243.

BOUCHET, P. & WAREN, A. 1980. Revision of the North-East Atlantic bathyal and abyssal Turridae.

Journal of Molluscan Studies, Supplement 8. 120 pp.

BRANDER, G. 1766. Fossilia Hantoniensia collecta et in Musaeo Britannico deposita. Londini. 43 pp.
BROCCHI, G. 1814. Conchiologia fossile subapennina, 2 volumes. Milano, Stamperia reale. 712 pp.
BRODERIP, W.J. 1830. Descriptions of two new species of Buccinum from the English and Irish seas.
Zoological Journal, 5:44-46.

BROGGER, W.C. 1900-1901. Om de senglaciale og postglaciale nivaforandringer i Kristianiafeltet. Norges
Geologiske Undersogelse, 31:1-205 (1900), 206-731 (1901).

BROWN, T. 1827. Illustrations of the Recent Conchology of Great Britain and Ireland. London, Highley
5 pp., 52 pi.

278 P. BOUCHET & A. WAREN

BROWN, T. 1833. The Conchologist’s Textbook. Glasgow, Fullarton & Co. 180 pp.

BRUGUIERE, M. 1792. Encyclopedic Methodique. Histoire Naturelle des Vers, 1. Paris, Panckoucke, 757 pp.

BUCQUOY, E., DAUTZENBERG, P. & DOLLFUS, G. 1882. Les Mollusques marins du Roussillon, 1.
Gasteropodes. Fascicule 1. Paris, Bailliere. pp. 1-40.

CANTRAINE, F. 1835. Diagnoses ou descriptions succinctes de quelques especes nouvelles de Mollusques.
Bulletin de TAcademie royale de Bruxelles, 2:376-406.

CASAMOR. L. & GHISOTTI, F. 1968. Buccinum ( Madiella ) Humphrey sianum Bennett, 1825, in: Schede
Malacologiche del Mediterraneo (F. Ghisotti, ed.). Como, Societa Malacologica Italiana.

CERNOHORSKY, W.O. 1970. Systematics of the families Mitridae and Volutomitridae. Bulletin of the
Auckland Institute and Museum, 8:1-190.

CERNOHORSKY, W.O. 1973. The taxonomy of Benthovoluta hilgendorfi (von Martens) and allied turbinellid
genera. Records of the Auckland Institute and Museum, 10:123-31.

CERNOHORSKY, W.O. 1975. The taxonomy of some West American and Atlantic Nassariidae based on their
type-specimens. Records of the Auckland Institute and Museum, 12:121-73.

CHASTER, G.W. 1898. A report upon the Mollusca obtained by the Royal Irish Academy cruises of 1885, 1886
and 1888. Proceedings of the Royal Irish Academy, (3)5:1-33.

CHASTER, G.W. & HEATHCOTE, W.H. 1894. A contribution towards a list of the marine Mollusca and
Brachiopoda of the neighbourhood of Oban. Journal of Conchology, London, 7:289-312.

CHEMNITZ, J.H. 1780. Neues Systematisches Conchylien Cabinet, 4. Nurnberg, G.N. Raspe, 344 pp.

CLARKE, A.H. 1960. Arctic archibenthal and abyssal molluscs from drifting station Alpha. Breviora,
119:1-17.

CLARKE, A.H. 1961. Abyssal mollusks from the South Atlantic Ocean. Bulletin of the Museum of
Comparative Zoology, 125:345-387.

CLARKE, A.H. 1962. Annotated list and bibliography of the abyssal marine molluscs of the world. National
Museum of Canada, Bulletin, 181:1-114.

CLARKE, A.H. 1974. Molluscs from Baffin Bay and the northern North Atlantic Ocean. Publications in
biological Oceanography, National Museum of Natural Sciences of Canada, 7:1-23.

COEN, G. 1922. Del genere Pseudomurex, (Monterosato, 1872). Atti della Societa italiana di scienze naturali,
61:68-71.

COEN, G. 1943. Appunti di Malacologia mediterranea. Acta Pontificiae Academiae scientiarum, ll(7):79-94.

COSSMAN, M. 1901-1903. Essais de Paleonconchologie comparee. 4(1901) & 5(1903). Paris, Societe
d’Editions scientifiques, 293+215 pp.

COSSMANN, M. & PISSARRO, G. 1907-1913. Iconographie complete des coquilles fossiles de V Eocene des
environs de Paris, 2: Gasteropodes. Paris. 65 pis.

COUTHOUY, J.P. 1838. Descriptions of new species of Mollusca and shells and remarks on several polypi
found in Massachusettes Bay. Boston Journal of Natural History, 2:53-1 1 1.

CRISTOFORI, G. de & JAN, G. 1832. Catalogus in IV Sectiones divisus rerum naturalium in Museo exstantium
...Sectio II. Pars I. Conchylia fossilia ex formatione telluris tertiaria in collectione nostra exstantia. Parma,
Typ. Carmignani. 16 pp.

CROSSE, H. 1875. (Note on Mediterranean Fusus). Journalde Conchyliologie , Paris, 23:272.

DA COSTA, E.M. 1778. Historia Naturalis Testaceorum Britanniae or The British Conchology. 254 pp.
London, Millan, White, Elmsley & Robson.

DALL, W.G. 1887. Supplementary notes on some species of mollusks of the Bering sea and vicinity.
Proceedings of the United States National Museum, 9:297-309.

DALL, W.H. 1889. Reports on the results of dredging. ..in the gulf of Mexico and in the Caribbean sea. ..XXIX.
Report on the Mollusca, part 2. Gastropoda and Scaphopoda.i?«//^/>? of the Museum of Comparative
Zoology, 18:1-492.

DALL, W.H. 1895. Scientific results of explorations by the (...) “Albatross”: 34, Report on Mollusca and
Brachiopoda dredged in deep water, chiefly near the Hawaiian Islands (...). Proceedings of the United States
National Museum, 17:675-733.

DALL, W.H. 1902. Illustrations and descriptions of new, unfigured, or imperfectly known shells, chiefly
American, in the U.S. National Museum. Proceedings of the United States National Museum, 24:499-566.

DALL, W.H. 1906. Early history of the generic name Fusus. Journal of Conchology, London, 11:289-297.

DALL, W.H. 1908a. Descriptions of new species of mollusks from the Pacific coast of the United States, with
notes on other mollusks from the same region. Proceedings of the United States National Museum,
34:245-57.

DALL, W.H. 1908b. Reports on the dredging operations off the west coast of Central America to the
Galapagos, the west coast of Mexico, and in the gulf of California, 14: the Mollusca and Brachiopoda.
Bulletin of the Museum of Comparative Zoology, 43(6):205-487.

DALL, W.H. 1909 Contributions to the Tertiary paleontology of the Pacific coast, 1. The Miocene of
Astoria and Coos Bay, Oregon. United States Geological Survey Professional Papers, 59:1-278.

DALL, W.H. 1916. Prodrome of a revision of the chrysodomoid whelks of the boreal and Arctic regions.
Proceedings of the Biological Society of Washington, 29:7-8.

DALL, W.H. 1918a. Changes in and additions to molluscan nomenclature. Proceedings of the Biological
Society of Washington, 31:137-38.

DALL, W.H. 1918b. Notes on Chrysodomus and other mollusks from the North Pacific Ocean. Proceedings
of the United States National Museum, 54:207-234.

DALL, W.H. 1919. The Mollusca of the Arctic coast of America collected by the Canadian Arctic expedition
west from Bathurst Inlet with an appended report on a collection of Pleistocene fossil Mollusca. Report of
the Canadian Arctic Expedition, 8(A): 1 A-25A.

NORTH-EAST ATLANTIC NEOGASTROPODA 279

DALL, W.H. 1924. Notes on molluscan nomenclature. Proceedings of the biological society of Washington,
37:87-90.

DALL, W.H. 1927. Small shells from dredgings off the southeast coast of the United States... Proceedings of
the United States National Museum, 70(18): 1-134.

D’AMICO, A. 1912. I Molluschi raccolti nel Mediterraneo dalla R.N. “Washington” durante le campagne
talassografiche. Archivo Zoologico, 5:233-79.

DAUTZENBERG, P. 1889. Contribution a la faune malacologique des iles Agores. Resultats des campagnes
scientifiques. . . Albert Ier, 1:1-112.

DAUTZENBERG, P. 1891. Contribution a la faune malacologique du golfe de Gascogne. Memoires de la
societe zoologique de France, 4:604-19.

DAUTZENBERG, P. 1910. Contribution a la faune malacologique de l’Afrique occidentale. Actes de la
Societe linneenne de Bordeaux, 64:1-174.

DAUTZENBERG, P. 1925. Mollusques nouveaux provenant des croisieres du Prince Albert Ier de Monaco.
Bulletin de ITnstitut Oceanographique de Monaco, 457:1-12.

DAUTZENBERG, P. 1927. Mollusques provenant des campagnes scientifiques du Prince Albert Ier de Monaco
dans l’Ocean Atlantique et dans le golfe de Gascogne. Resultats des campagnes scientifiques... Albert Ier,
72:1-401.

DAUTZENBERG, P. & FISCHER, H. 1896. Dragages effectues par V Hirondelle et par la Princesse-AliceA .
Mollusques Gasteropodes. Memoires de la Societe zoologique de France, 9:395-498.

DAUTZENBERG, P. & FISCHER, H. 1906. Mollusques provenant des dragages effectues a l’ouest de
l’Afrique. Resultats des campagnes scientifiques. . .Albert Ier, 32:1-125.

DAUTZENBERG, P. & FISCHER, H. 1912. Mollusques provenant des campagnes de V Hirondelle et de la
Princesse- Alice dans les Mers du Nord. Resultats des campagnes scientifiques... Albert Ier, 37:1-629.

DESHAYES, G.P. 1832. Encyclopedie methodique: Histoire naturelle des Vers, 2, 145-594. Paris, Agasse.

DONOVAN, E. 1804. The natural history of British Shells, 2. London, C. Rivington.

DUNKER, W. & METZGER, A. 1874. Drei neue Meeres-Conchyhen der norwegischen Fauna. Nachrichtsblatt
der Deutschen Malakozoologischen Gesellschaft, 6:7-8.

DUNKER, W. & METZGER, A. 1874. Drei neue Meeres-Conchylien der norwegischen Fauna. Jahrbucher der
Deutschen Malakozoologischen Gesellschaft, 1:146-51.

DUNKER, W. & METZGER, A. 1875. Nordsee-Expedition 1872. 8: Mollusca. Jahresbericht der Commission
fur Untersuchung der Deutschen Meere, 3:229-64.

FABR1CIUS, O. 1780. Fauna Groenlandica. Hafniae et Lipsiae, J.G. Rothe. 468 pp.

FABRICIUS, O. 1823. Fortegnelse. Copenhagen. 114 pp.

FANGE, R. 1960. The salivary gland of Neptunea antiqua. Annals of the New York Academy of Sciences,
90:689-94.

FECHTER, R. 1977. Zwei bemerkenswerte Muriciden aus dem mittleren Ostatlantik. Spixiana, 1(2): 15 1-64.

FECHTER, R. 1979. Gastropoden aus der Iberischen Tiefsee. Meteor Forschungs-Ergebnisse, (D) 30:23-40.

FIORONI, P. 1966. Zur Morphologie und Embryogenese des Darmtraktes und der transitorischen Organe
bei Prosobranchiern. Revue suisse de Zoologie, 73(44): 62 1-876.

FISCHER, P. 1868. Note sur les especes du genre Fusus qui habitent les cotes oceaniques de la France. Journal
de Conchyliologie, Paris, 18:35-38.

FISCHER, P. 1882. Diagnoses d’especes nouvelles de Mollusques recueillis dans le cours des expeditions
scientifiques de l’aviso le Travailleur (1880 et 1881). Journal de Conchyliologie, Paris, 30:49-53, 273-77 .

FISCHER, P. 1883. Diagnoses d’especes nouvelles de Mollusques recueillis dans le cours de l’expedition
scientifique du Talisman (1883). Journal de Conchyliologie, Paris, 31:391-94.

FISCHER, P. 1884. Manuel de Conchyliologie, fascicule 7 (pp. 609-88). Paris, F. Savy.

FISCHER, P. & BOUVIER, E.L. 1890. Sur l’organisation des gastropodes prosobranches senestres (Neptunea
contraria). Comptes Rendus de TAcademie des Sciences, Paris, 110: 412-14.

FRANCHINI, D. & FRILLI, G. 1970. II genere Trophonopsis B.D.D. 1882, nel mar Mediterraneo. Atti delta
Unione Malacologica Italiana, l(l):30-55.

FRANCHINI, D. & ZANCA, M. 1977. II genere Fusinus nel Mediterraneo. La Conchiglia, 99-100:16-19.

FRIELE, H. 1877. Preliminary report on Mollusca from the Norwegian North Atlantic Expedition in 1876.
Nyt Magazin for Naturvidenskaberne, 23:1-10.

FRIELE, H. 1879. Uber die Variationen der Zahnstruktur bei dem Genus Buccinum. Jahrbucher der Deutschen
Malakozoologischen Gesellschaft, 1:256-63.

FRIELE, H. 1879. Catalog der auf der norwegischen Nordmeer-Expedition bei Spitzbergen gefundenen
Mollusken. Jahrbucher der Deutschen Malakozoologischen Gesellschaft, 1:264-86.

FRIELE, H. 1882. Mollusca I. Buccinidae. The Norwegian North- Atlantic Expedition 1876-78. 38 pp.

FRIELE, H. 1886. Mollusca II. The Norwegian North- Atlantic Expedition 1876-78. 3. 44 pp.

FRIELE, H. & GRIEG, J.A. 1901. Mollusca III. The Norwegian North- Atlantic Expedition, Zoology,
7.131 pp.

GERONIMO, I. DI & PANETTA, P. 1973. La malacofauna batiale del golfo di Taranto. Conchiglie, 9:69-122.

GHISOTTI, F. 1974. Rinvenimenti malacologici nel Mediterraneo. Conchiglie, 10:127-31.

GLIBERT, M. 1973. Revision des Gastropoda du Danien et du Montien de la Belgique. Memoires de ITnstitut
royal des sciences naturelles de Belgique, 173:1-1 16.

GMELIN, J.F. 1790. Systema Naturae, ed. 13, 1. Lipsiae.

GOLIKOV, A. 1964. Gastropods and Scaphopods from North Greenland sea, north of Spitzbergen, north of
Franz Joseph land and Novaja Zemlya. Trudy Arkticheskogo i antarkticheskogo nauchno-issledovateVs
skogo instituta, 259:340-53 [In Russian.]

GOLIKOV, A. 1977. Investigation of prosobranchs of the family Fasciolariidae in temperate waters.
Explorations of the fauna of the seas, 21(29): 102-104. Lin Russian.]

280 P. BOUCHET & A. WAREN

GOLIKOV, A.N. 1980. Buccininae mollusks of the world oceans. Fauna USSR, 121:465pp. [In Russian.].
GOLIKOV, A.N. & SCARLATO, O.A. 1977. Composition, distribution and ecology of gastropods and
bivalves off Franz Joseph Land. In Scarlato, O.A., Biocoenoses of the shelf of Franz Josef Land and the
fauna of adjacent waters. Explorations of the fauna of the Seas, 22:313-89. [In Russian.]

GOLIKOV, A.N. & STAROBOGATOV, Y.I. 1975. Systematics of prosobranch gastropods. Malacologia,
15:185-232.

GORBUNOV, G. 1946. Bottomlife of the Novosiberian shoalwaters and the central part of the Arctic Ocean.

Dreifuischchia ekspeditia Glavseomorputina nom Parokhodi G. Sedov 1937-40, 3:30-138. [In Russian.]
GORBUNOV, G. 1946. New and interesting species of Mollusca. Dreifuischchia ekspeditia Glavseomorputina
nom Parokhodi G. Sedov 1937-40, 3:308-22. [In Russian.]

GOULD, A. A. 1840. Descriptions of thirteen new species of shells. American Journal of Science, 38:196-97.
GOULD, A. A. 1841. Report on the Invertebrata of Massachusetts. Cambridge, Wright & Potter, 373 pp.
GOULD, A. A. 1860. Descriptions of new shells collected by the United States North Pacific Exploring
Expedition. Proceedings of the Boston Society of Natural History, 7:323-40.

GRAY, J.E. 1824. A supplement to the Appendix of Captain Perry's Voyage. Shells, pp. 240-246. London.
GRAY, J.E. 1839. Molluscous animals, and their shells. In: The Zoology of Captain Beechey’s Voyage,
pp. 102-155.

GRAY, J.E. 1857. Guide to the systematic distribution of Mollusca in the British Museum, part 1. London.
230 pp.

GREGORIO, A.DE. 1885. Appunti intorno al genere Trophon; Intorno a talune specie di Fasciolaria, Fusus,
Tudicla, Pyrula. Bullettino della Societa malacologica italiana, 11:26-32, 46-53.

HABE, T. 1952. Pholadomyidae, Clavagellidae, Pandoridae, Juliidae and Condylocardiidae of Japan.

Illustrated Catalogue of Japanese Shells, 1(18): 121-32.

HABE, T. 1958. Descriptions of ten new gastropod species. Venus, 20:32-42.

HABE, T. 1962. Trichotropidae in Japan. Bulletin of the National Science Museum. 6(21:67-77.

HABE, T. & SATO, J. 1973. A classification of the family Buccinidae from the North Pacific. Proceedings of
the Japanese Society of Systematic Zoology, 8:1-8.

HAGG, R. 1925. Svenska kvartara mollusker i Hisingers samling. Arkiv for Zoologi, 17A(19): 1-25.
HANCOCK, A. 1846. A list of shells dredged on the west coast of Davis’s Strait, with notes and descriptions
of eight new species. Annals and Magazine of Natural History, 18:323-38.

HANLEY, S. 1855. Ipsa Linnaei Conchylia. London, Williams & Norgate, 556 pp.

HARASEWYCH, M.G. & JENSEN, R.H. 1979. Review of the subgenus Pterynotus in the western Atlantic.
Nemouria, 22:1-16.

HARMER, F.W. 1914-1919. The Pliocene Mollusca of Great Britain, 1, 1-484. London, Paleontographical
Society.

HERZENSTEfN, S. 1885. Beitrage zur Kenntnis der Fauna der Murmankuste und des Weissen Meeres.

Materialy faune Murmanskago berega Belago morja, 1:635-814.

HIDALGO, J.G. 1917. Fauna Malacologica de Espana, Portugal y las Baleares. Trabajos del Museo Nacional
de Ciencias Naturales, Zoologia, 30:1-752.

HISINGER, W. 1837. Lethaea Svecica. Holmiae, Nordstedt & Son. 124 pp.

HOUART, R. 1981. Revision des Trophoninae d’Europe. Informations de la Societe beige de Malacologie,
(9)1-2:3-70.

HOWSE, R. 1847. Notes on a dredging excursion off the coast of Durham, with descriptions of the ova-capsules
of Fusus norwegicus and F. turtoni. Annals and Magazine of Natural History, 19:160-64.

IREDALE, T. 1915. Notes on the names of some British Marine Mollusca. Proceedings of the Malacological
Society of London, 11:329-43.

IREDALE, T. 1918. Molluscan nomenclatural problems and solutions, no. 1. Proceedings of the Malacological
Society of London, 13:28-40.

JAY, J.C. 1839. A catalogue of shells (...), 3rd edition. New York, Wiley & Putnam. 126 pp.

JEFFREYS, J.G. 1867. British Conchology, 4. London, van Voorst. 487 pp.

JEFFREYS, J.G. 1869. The deep-sea dredging expedition in H.M.S. Porcupine. Nature, 1:135-37, 166-68.
JEFFREYS, J.G. 1869. British Conchology, 5 . London, van Voorst. 259 pp.

JEFFREYS, J.G. 1877. New and peculiar Mollusca of the Eulimidae and other families of Gastropoda as well
as of the Pteropoda produced during the Valorous expedition. Annals and Magazine of Natural History,

(4) 19:317-39.

JEFFREYS, J.G. 1880. The deep-sea Mollusca of the bay of Biscay. Annals and Magazine of Natural History,

(5) 6:315-19.

JEFFREYS, J.G. 1882. Black Sea Mollusca. Annals and Magazine of Natural History, (5)10:425-27.
JEFFREYS, J.G. 1883. On the Mollusca procured during the cruise of H.M.S. Triton, between the Hebrides
and Faeroes in 1882. Proceedings of the Zoological Society of London (1883):389-399.

JEFFREYS, J.G. 1885. On the Mollusca procured during the Lightning and Porcupine expeditions, IX.

Proceedings of the Zoological Society of London (1885):27-63.

JOHNSTON, G. 1825. Contributions to the British fauna. Philosophical Journal, Edinburgh, 13:218-22.
JONAS, J.H. 1846. Molluskologische Beitrage. Abhandlungen von dem naturwissenschaftlichen Verein in
Hamburg, 1:99-130.

JORDAN, H.K. 1890. On the species and varieties of the genus Fusus. Journal of Conchology, London,
6:225-39.

KANTOR, Y. 1981. Species composition and distribution of Molluscs of the family Buccinidae in the East
Murman. Zoologicheskii Zhurnal, 60(8): 1 145-50. [In Russian.]

KIENER, L.C. 1834-1840. Species general et iconographie des coquilles vivantes: genre Buccin (1834, 108 pp.),
genre Fuseau (1840, 62 pp.). Paris, Rousseau.

NORTH-EAST ATLANTIC NEOGASTROPODA 281

KING, W. 1846. An account of some shells and other invertebrate forms found on the coast of Northumberland
and of Durham. Annals and Magazine of Natural History, 18:233-51.

KLEIN, J.T. 1753. Tentamen Methodi Ostracologicae. Lugduni Batavorum. 177 pp.

KNUDSEN, J. 1956. Marine prosobranchs of tropical West Africa. Stencglossa. Atlantide Report , 4:9-105.

KNUDSEN, J. 1964. Scaphopoda and Gastropoda from depths exceeding 6000 metres. Galathea Report,
7:125-36.

KOBELT, W. 1876. Beitrage zur arctischen Fauna. Jahrbucher der Deutschen Malakozoologischen
Gesellschaft, 3 : 6 1 -76 , 37 1 -73 .

KOBELT, W. 1878. Zur Kenntniss der nordischen Mollusken. Jahrbucher der Deutschen Malakozoologischen
Gesellschaft, 5:276-83.

KOBELT, W. 1887. Murex fusulus Brocchi. Jahrbucher dez Deutschen Malakozoologischen Gesellschaft,
14:120-24

KOBELT, W. 1905. Iconographie dez schalentragenden europaischen Meeresconchylien, 3. Wiesbaden,
Kreidel, 406 pp.

KOSUGE, S. 1975, Illustrations of type specimens of Molluscs described by William Healy Dali Privately
printed, 29 plates.

LAMARCK, J.B. de 1799. Nouvelle classification des coquilles. Memoires de la Societe d’Histoire Naturelle
de Paris 1799:63-91 .

LEACH, W.E. 1847. The classification of British Mollusca. Annals and Magazine of Natural History,
20:267-73.

LEBOUR, M.V. 1936. Notes on the eggs and larvae of some Plymouth Prosobranchs. Journal of the marine
biological Association of the U.K., 20:547-65.

LECHE, W. 1878. Hafsmollusker. Kongliga Svenska Vetenskaps Akademiens Hand/ingar, 16(2): 1-86.

LIBASSI, P.I. 1859. Memoria sopra alcune conchiglie fossili dei dintorni di Palermo. Atti dell' Accademia
di scienze, lettere ed arti, Palermo, (NS)3:l-47.

LINNE, C. 1758. Systema Naturae, ed. 10. Holmiae. 823 pp.

LINNE, C. 1771. Mantissa plantarum. Holmiae. 588 pp.

LINSLEY, J.H. 1845. Catalogue of the shells of Connecticut. American Journal of Science, 48:271-86.

LOCARD, A. 1886. Prodrome de malacologie frangaise. Catalogue general des Mollusques vivants de France.

Mollusques marins. Paris, Bailliere, 779 pp.

LOCARD, A. 1886. Contribution a la faune malacologique frangaise. X. Monographic des especes frangaises de
la famille des Buccinidae. Annales de la Societe Linneenne de Lyon, 33:17-127.

LOCARD, A. 1891. Sur une espece nouvelle du genre Neptunia. L'Echange, 77:34-35.

LOCARD, A. 1892. Les Coquilles marines des cotes de France. Paris, Bailliere, 384 pp.

LOCARD, A. 1896. Mollusques testaces et brachiopodes. In: Koehler, R.:Resultats scientifiques de la campagne
du Caudan dans le Golfe de Gascogne, fasc. 1. Annales de TUniversite de Lyon . 26:1-272 (Mollusca p.
129-242.) A shorter version was published 1897, in Annales de la Societe d’agriculture de Lyon (7)4:205-22.
Locard (1896) refers to this latter paper as being published 1895, but it was actually not published until 1897.

LOCARD, A. 1897. Mollusques Testaces, I. Expeditions scientifiques du Travailleur et du Talisman. Paris,
Masson, 516 pp.

LOCARD, A. & CAZIOT, E. 1901. Les coquilles marines des cotes de Corse. Annales de la Societe linneenne
de Lyon, (2)47:1-80, 159-291.

LOVEN, S. 1846. Index Molluscorum litora Scandinaviae occidentalia habitantium. Kongelige Vetenskaps-
Akademiens Fbrhandlingar, 3:135-204.

LUS, V.J. 1971. A new genus and species of gastropod moiluscs (Family Buccinidae) from the ultra-abyssal of
Kurile-Kamchatka Trench. Trudy Instituta okeanologii, 92:61-72 [In Russian.]

LUS, V.J. 1973. New abyssal Fasciolariidae. Transactions of the Shirshov Institute of Oceanology , 91:203-12
[In Russian.]

LUS, V.J. 1978. New genus and species of Buccinidae from the lower-abyssal zone of Idzu-Bonin Trench in
Pacific ocean. Transactions of the Shirshov Institute of Oceanology, 113:147-56. [In Russian.]

LUS, V.J. 1981. On abyssal species Mohnia mohni and Sipho danielsseni. Trudy Instituta okeanologii,
115:126-39. [In Russian.]

MACANDREW, R. 1852. Note of the Mollusca observed during a short visit to the Canary and Madeira
Islands, etc. in the months of April and May 1852. Annals and Magazine of Natural History, (2)10:100-108.

MACANDREW, R. 1857. Report of the marine testaceous Mollusca of the Northeast Atlantic and
neighbouring seas, and the physical conditions affecting their development. Report of the British Association
for the Advancement of Science (1856): 101-58.

MACGILLIVRAY, W. 1843. A history of the Molluscous animals of the counties of Aberdeen, Kincardine,
and Banff. London, Cunningham & Mortimer, 372 pp.

MACPHERSON, E. 1971. The marine Mollusca of arctic Canada. Publications in biological
oceanography, National Museum of Natural Sciences of Canada, 3:1-149.

MALDONADO QU1LES, A. 1973. Segnalazione di un Nassaridae nuovo per il Mediterraneo. Conchiglie,
9(1 1-1 2):23 1 -34.

MARSHALL, J.T. 1902. Notes on the British species of Buccinum, Fusus, etc. Journal of Malacology,
9:33-50.

MARSHALL, J.T. 1911. Additions to “British Conchology”. Journal of Conchology, London, 13:223-31.

MARTENS, E. von. 1878. Uber einige Crustaceen und Mollusken, welche das zoologische Museum in letzter
Zeit erhalten haben. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin ( 1 878): 131-35.

MARTENS, E. von. 1881. Conchologische Mittheilungen, 2. Heft 1 & 2:103-28. Cassel, T. Fischer.

MASSY, A. 1916. Mollusca and Brachiopoda of the Irish Atlantic slope. Journal of Conchology, London,
15:48-51.

282 P. BOUCHET & A. WAREN

MASSY, A. 1930. Mollusca of the Irish Atlantic slope, 50-1500 fathoms. Proceedings of the Royal Irish
Academy, 39B(13):232-342.

MELVILL, J.C. & STANDEN, R. 1900. Report on the Mollusca of the “Jackson-Harmsworth” Expedition to
Franz-Josef Land (1896-97), and the “Andrew Coats” Cruise (1898) to Kolguev, etc. Manchester Memoirs

44(4): 1-14.

MIDDENDORFF, A.T. 1849. Beitrage zu einer Malacozoologia Rossica, II. Aufzahlung und Beschreibung der
zur Meeresfauna Russlands gehOrigen Einschaler. Memoires de I’Academie imperiale des sciences de Saint-
Petersbourg, (6)8 (Sciences naturelles, 6):329-610.

MILASCHEWITSCH, K.O. 1916. Les Mollusgues de la mer Noire et de la mer d’Azov. Petrograd. 312 pp.

MOHR, N. 1786. Forsog til en islandsk naturhistorie. Kiobenhavn, C.F. Holm. 414 pp.

MOLLER, H. 1842. Index Molluscorum Groenlandiae. Naturhistorisk Tidsskrift, 4:7t>-97.

MONTAGU, G. 1803. Testacea Britannica, 1. London, J. White, 291 pp.

MONTEROSATO, A.T. di 1869. Testacei nuovi dei mari di Sicilia. Palermo, Mirto, 18 pp.

MONTEROSATO, A.T. di 1872. Notizie intorno alle Conchiglie fossili di Monte Pellegrino e Ficarazzi.
Palermo, M. Amenta, 44 pp.

MONTEROSATO, A.T. di 1877. Relativement au Fusus jeffreysianus Fischer. Journal de Conchy liologie,
Paris, 25:99-100.

MONTEROSATO, A.T. di 1878. Enumerazione e Sinonimia delle Conchiglie Mediterranee. Giornale di scienze
naturali ed economiche di Palermo, 13:61-115.

MONTEROSATO, A.T. di. 1884. Nomenclatura generica e specif ica di alcune conchiglie Mediterranee.
Palermo, Virzi, 152 pp.

MONTEROSATO, A.T. di. 1890. Conchiglie delle profondita del mare di Palermo. Naturalista Siciliano,
6-8:1-31.

MONTFORT, D. de. 1810. Conchy liologie systematique. Paris, F. Schoell, 676 pp.

MORCH, O. 1857. Fortegnelse over Grcfnlands Blctddyr. in: Rink, H.J. GrOnland geographisk og statistisk
beskrivet, pp. 75-100.

MORCH, O. 1862. Description d’une nouvelle espece de Fuseau. Journal de Conchy liologie, Paris, 10:36-37.

MORCH, O. 1868. Faunula molluscorum insularum Faeroensium. Videnskabelige Meddelelser fra den
naturhistoriske Forening i Kjobenhavn, (1867): 57-1 1 1 .

MORCH, O. 1869. Description de trois Fusus nouveaux du Greenland. Journal de Conchyliologie, 17:397-399.

MORCH, O. 1869. Catalogue des mollusques du Spitzberg recueillis par le Dr H. Kroyer pendant le voyage
de la corvette La Recherche en juin 1838. Memoires de la Societe malacologique de Belgique, 4:7-32.

MORCH, O. 1876. Description d‘especes nouvelles. Journal de Conchyliologie, Paris, 24:368-74.

MORCH, O. 1877. Mollusca. pp. 435-442, in: Rink, Danish Greenland, its people and its products. London,
S. King.

MORONI, M.A. & TORRE, G. 1966. Nuovi dati sul pliocene e il quaternario dei dintorni di Palermo, 4:
Macrofauna dei trubi (Pliocene inferiore) di Lascari. Rivista Mineraria Siciliana, 91-93:27-49.

NELSON, C.M. 1976. The type species of Neptunea Roding, 1798. Nautilus, 90:139-141.

NORDSIECK, F. 1968. Die europdischen Meeres- Gehauseschnecken . Stuttgart, G. Fischer, 274 pp.

NORDSIECK, F. 1974. Molluschi dei fondali della platea continentale fra la Corsica e la Sardegna. La
Conchiglia, 61:11-14.

NORDSIECK, F. 1977. The Turridae of the European Seas. La Conchiglia, Roma. 132 pp.

NORMAN, A.M. 1879. The Mollusca of the fjords near Bergen. Journal of Conchology, 2:8-77.

NORMAN, A.M. 1893. A month on the Trondheim Fiord. Annals and Magazine of Natural History, (6)12:
341-67.

ODHNER, N. 1915. Die Molluskenfauna des Eisfjordes. Kungliga Svenska Vetenskapsakademiens Handlingar,
54:1-274.

OPINION 469, 1957. Rejection (a) of the generic name “Jumala” Friele, 1882, as a name calculated to give
offence on religious ground, and (b) of the name “Beringius” Dali, 1879, as not having been duly
published. Opinions and declarations of ICZN, 16(8) :97- 128.

OPINION 521, 1958. Addition to the “Official Index of Rejected and invalid works in zoological
nomenclature” of the title of the paper by Otto Fabricius issued in Copenhagen in 1823 as “Fortegnelse over
afgang Biskop Fabriciusses efterladte Naturalier”. Opinions and Declarations of ICZN, 19(8):201 -208.

ORTEA, J.A. 1977. Moluscos marinos Gasteropodos y Bivalvos del litoral asturiano, 1. Thesis. Oviedo. 342 pp.

OSKARSSON, I. 1962. Skeldyrafdna Islands, 2. Reykjavik, Leiftur, 167 pp.

OSKARSSON, I. 1966. Nyjungar um islensk skeldyr. Ndtturufraedingurinn, 36:86-92.

OSKARSSON, I. 1967. Nyir fundarstadir skeldyra vid Island. Ndtturufraedingurinn, 37:58-63.

OSKARSSON, I. 1977. Fjogurra skeldyrategunda getid i fyrsta sinn fra strondum Islands,
Ndtturufraedingurinn, 47 : 1 80-83 .

PAIN, T. 1978. The genus Colus Roeding, 1798, in Western Europe. La Conchiglia, 114/115:3-7.

PAIN, T. 1979. The genus Buccinum Linne, 1758, in Western Europe La Conchiglia, 124/125:15-18.

PAIN, T. 1981. On the occurrence of Colus ventricosus (Gray, 1839), a prosobranch mollusc. Bulletin of the
Geological Society of Norfolk, 29:56-57.

PALLARY, P. 1900. Coquilles marines du littoral d’Oran. Journal de Conchyliologie, 48:211-420.

PALLARY, P. 1904. Addition a la faune malacologique du golfe de Gabes. Journal de Conchyliologie, 52:
212-48.

PALLARY, P. 1931. Les Buccins de la Mediterranee. Bulletin des Travaux publies par la Station d'Aquiculture
et de Peche de Castiglione “1930”(1):9-19.

PAULUS, M. & MARS, P. 1942. Guide malacologique des environs de Marseille. Bulletin du Museum
d’Histoire Naturelle de Marseille, 2:71-120.

NORTH-EAST ATLANTIC NEOGASTROPODA 283

PEARCE, J. & THORSON, G. 1967. The feeding and reproductive biology of the red whelk Neptunea antiaua
(L.). Ophelia, 4:277-314.

PENCHASZADEH, P. 1976. Reproduccion de Gastropodos Prosobranquios del Atlantico suroccidental. El
genero Trophon. Physis, (A)35(90):69-76.

PETERSEN, C. 1888. Om de skalbaerende molluskers udbredelseforhold i de danske have indenfor Skagen.
162 pp. Kjobenhavn, A.F. Host.

PETIT, R.E. 1970. Notes on Cancellariidae, II. Tulane Studies in Geology and Paleontology, 8(2):83-88.

PETIT DE LA SAUSSAYE, S. 185 1 . Description d’une nouvelle espece du genre Fuseau. Journal de
Conchyliologie , Pans, 2:254-255.

PETIT DE LA SAUSSAYE, S. 1851. Description de coquilles nouvelles. Journal de Conchyliologie, Paris,
2:365-368.

PFEFFER, G. 1886. Ubersicht der im Jahre 1881 vom Grafen Waldburg-Zeil im Karischen Meere gesammelten
Mollusken. Abhandlungen des Naturwissenschaftlichen Vereins zu Hamburg, 9(1): 1-14.

PHILIPPI, R.A. 1836-1844. Enumeratio Molluscorum Siciliae... Berlin & Halle. 1 (1836), 268 pp; 2 (1844),

. 303 pp.

PHILIPPI, R.A. 1850. Abbildungen und Beschreibungen neuer oder wenig gekannter Conchylien, 3. Cassel,
T. Fischer, 138 pp.

PINNA, G. 1971 . I tipi delle specie di Gasteropodi terziari istituite da Giuseppe De Cristofori e Giorgio Jan nel
1832 conservati nelle collezioni del Museo Civico di Storia Naturale di Milano. Atti della Societa Italiana de
Scienze naturale e del Museo Civico di Storia Naturale di Milano, 112:421-40.

PINNA, G. & SPEZIA, L. 1978. Catalogo dei tipi del Museo Civico di Storia Naturale di Milano. V, I tipi dei
Gasteropodi fossili. Atti della Societa Italiana de Scienze naturale e del Museo Civico di Storia Naturale di
Milano, 119:125-80.

PONDER, W. 1972. Notes on some Australian genera and species of the family Muricidae. Journal of the
Malacological Society of Australia, 2(3):2 1 5-48 .

PONDER, W. 1973. The origin and evolution of the Neogastropoda. Malacologia, 12:295-338.

PORTMANN, A. 1925. Der Einfluss der Nahreier auf die Larvenentwicklung von Buccinum und Purpura.
Zeitschrift fiir Morphologie und Okologie der Tiere, 3:526-41.

PORTMANN, A. 1930. Die Larvalnieren von Buccinum undatum L. Zeitschrift fiir Zellforschung, 10:401-10.

POSSELT, H.J. 1895. OstgrOnlandske Mollusker. Meddelelser om Gronland, 19(2):59-94.

POSSELT, H.J. & JENSEN, A.S. 1898. Gronlands Brachiopoder og Bloddyr. Meddelelser om Gronland,
23:1-298.

POWELL, A.W.B. 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda. Discovery
Reports, 26:49-196.

POWELL, A.W.B. 1966. The molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute
and Museum, 5:1-184.

QUINN, J. 1981. A new genus of Turbinellidae with the description of a new species from the Caribbean sea.
Nautilus, 95:72-77.

RADWIN, G.E. 1972. The systematic position of Urosalpinx carolinensis Verrill, 1884, with comments on
the genus Mohnia Friele, 1878. Transactions of the San Diego Society of Natural History, 16:339-42.

RADWIN, G.E. 1977-78. The family Columbellidae in the Western Atlantic. The Veliger, 19:403-17; 20: 1 19-33,
328-44.

RADWIN, G.E. & D’ATTILIO, A. 1976. Mur ex Shells of the World. Stanford, Stanford University Press,
284 pp.

REEVE, L. 1845-48. Conchologia Iconica, 3 (1845-47), 4 (1847-48). London, Reeve & Benham.

REEVE, L. 1855. Account of the shells collected by Captain Sir Edward Belcher, North of Beechey Island. 2,
392-399. In: Richardson et al., The last of the Arctic Voyages (...). London, L. Reeve.

REEVE, L. 1856. Conchologia Iconica, 10. London, Reeve & Benham.

REHDER, H. 1972. Some notes on the genus Teramachia. Veliger, 15:7-10.

RICHTER, G. & THORSON, G. 1975. Pelagische Prosobranchier-Larven des Golfes von Neapel. Ophelia,
13:109-85.

RIESER, N.W. 1969. Feeding behaviour of some New England marine gastropods. Nautilus, 82:112-113.

RODING, P.F. 1798. Museum Boltenianum . . .Pars secunda continens Conchylia, etc. ..Hamburg. 199 pp.

ROLAN, E. 1983. Moluscos de la ria de Vigo, 1. Gasteropodos. Privately published, Vigo. 383 pp.

ROSSI-RONCHETTI, C. 1955. I tipi della “Conchiologia fossile subapennine” di G. Brocchi, 7. Rivista
italiana di Paleontologia, 60(1): 187-202.

SABELLI, B. & SPADA, G. 1980. Guida illustrata all’identificazione delle conchiglie del Mediterraneo: Fam.
Coralliophilidae. Bollettino Malacologico, 16 (7-8, suppl.): 3 pp.

SABELLI, B. & TOMMASINI, S. 1983. Contributo alia conoscenza sistematica di “Murex” fusulus Brocchi,
1814. Bollettino Malacologico, 19:1-12.

SALIS MARSCHLINS, C.U. von. 1793. Reisen in verschiedene Provinzen des Konigreiches Neapel, 1. Zurich &
Leipzig, Ziegler & Sohne, 442 pp.

SANDBERGER, C. 1858-63. Die Conchylien des Mainzer Tertidrbeckens. Wiesbaden, C.W. Kreidels, 458 pp.
(pp. 73-232:1861.)

SARS, G.O. 1878. Mollusca Regionis Arcticae Norwegiae. Christiania, Brogger, 466 pp.

SARS, M. 1851. Beretning om en i sommeren 1849 foretagen zoologisk Reise. Nyt Magasin for Naturviden-
skapene, 6:121-211.

SARS, M. 1859. Bidrag till en skildring av den arktiske molluskfauna ved Norges nordlige Kyst. Forhandlinger i
Videnskabsselskabet i Kristiania (1858):34-87.

284 P. BOUCHET & A. WAREN

SCACCHI, A. 1836. Catalogus Conchyliorum Regni Neapolitani. Neapoli. 18 pp.

SCHLESH, H. 1926. Notes sur la distribution du Volutopsius norwegica Chemn. et du Beringius turtoni Bean.

Journal de Conchyliologie, Paris, 70:146-50.

SCHUMACHER, C.F. 1817. Essai d’un nouveau systeme des habitations des Vers testaces. Copenhague,

Sctjultz, 287 pp.

SEAWARD, D. 1982. Sea area atlas of the marine Molluscs of Britain and Ireland. Nature Conservancy

Council, Shrewsbury. 746 maps.

SETTEPASSI, F. 1977. / Molluschi marini viventi nel Mediterraneo, 2 (1977?, 1971 on title page is erroneous).

Roma.

SIMPSON, J. 1903. Marine conchology of “Dee”. Transactions of the Aberdeen working Men’s Natural

History and Scientific Society, 1:64-86.

SIMROTH, H. 1895. Die Gastropoden der Plankton-Expedition. Ergebnisse der Plankton Expedition, 2:1-208.
SMITH, E.A. 1890. Report on the marine Molluscan fauna of the Island of St. Helena. Proceedings of the
Zoological Society of London, (1890):247-317.

SMITH, E.A. 1906. On Mollusca from the Bay of Bengal and the Arabian sea. Annals and Magazine of Natural
History, (7)18:157-75.

SOWERBY, G.B. 1880. Thesaurus Conchyliorum, 4. Parts 35-36. London, Sowerby.

SOWERBY, J.C. 1818. Mineral Conchology of Great Britain, 2. London, Arding & Merrett. 251 pp.
SPADA, G. & MALDONADO, A. 1974. Nota preliminare sulle specie di molluschi a diffusione prevalente-
mente Atlantica e presente anche in Mediterraneo nel Mare di Alboran. Quaderni della Civica Stazione
Idrobiologica di Milano, 5:51 -69.

SPARCK, R. & THORSON, G. 1933. Marine Gastropoda Prosobranchiata. Zoology of the Faeroes, 52:1-56.
SPIGHT, T.M. 1976. Hatching size and the distribution of nurse eggs among prosobranch embryos. Biological
Bulletin, 150:491-99.

STIMPSON, W. 1865. Review of the northern Buccinums, and remarks on some other northern marine
moilusks, part 1. The Canadian Naturalist, (N.S.), 2:364-89.

STRAUCH, F. 1972. Phylogenese, Adaptation und Migration einiger nordischer mariner Molluskengenera
(Neptunea, Panomya, Cyrtodaria und Mya). Abhandlungen von der Senckenbergischen naturforschenden
Gesellschaft , 531 : 1-2 1 1 .

STREBEL, H. 1904. Beitrage zur Kenntniss der Molluskenfauna der Magalhaen Provinz. Zoologische
Jahrbucher, Abt. Systematik, 21:171-248.

STROM, H. 1768. Beskrivelse over norske insekter. Andet stykke. Konglige norske Videnskabers Selskab i
Trondheims Skrifter, 4:313-71.

STURANY, R. 1896. Mollusken I gesammelt von S.M. Schiff Pola 1890-1894. Denkschriften der
Mathematisch-Naturwissen-schaft lichen Classe der Kaiserlichen Akademie der Wissenschaften, 63:1-36.
SYKES, E.R. 1905. Descriptions of new forms of Marginellidae and Pleurotomidae. Proceedings of the
Malacological Society of London, 6:315-18.

SYKES, E.R. 1911. On the Mollusca procured during the Porcupine expeditions. Supplementary notes, part 4.

Proceedings of the Malacological Society of London, 9:331-48.

TAVIANI, M. & COLANTONI, P. 1979. Thanatocoenoses wurmiennes associees aux coraux blancs. Rapport
de la commission Internationale pour V exploration scientifique de la mer Mediterranee, 25/26(4): 141-42.
THIELE, J. 1929. Handbuch der systematischen Weichtierkunde, 1. Jena, G. Fischer, 376 pp.

THORSON, G. 1935. Studies on the egg-capsules and development of arctic marine Prosobranchs. Meddelelser
om Gronland, 100(5): 1-71.

THORSON, G. 1940. Notes on the egg-capsules of some North Atlantic Prosobranchs of the genus Troschelia,
Chrysodomus , Volutopsius, Sipho, and Trophon. Videnskabelige Meddelelser fra Dansk naturhistorisk
Forening i Kjobenhavn, 104:251-65.

THORSON, G. 1941. Marine Gastropoda Prosobranchiata. The Zoology of Iceland, 4(60): 1-150.
THORSON, G. 1944, The Zoology of East Greenland: Marine Gastropoda Prosobranchiata. Meddelelser om
Gronland, 121(13): 1-181.

THORSON, G. 1946. Reproduction and larval development of Danish marine bottom invertebrates.

Meddelelser fra Kommissionen for Havundersogelser, serie Plankton, 4:1-523.

THORSON, G. 1951. The Godthaab expedition 1928: Scaphopoda, Placophora, Solenogastres, Gastropoda
Prosobranchiata, Lamellibranchiata. Meddelelser om Gronland, 81(2): 1-1 17.

TIBA, R. & KOSUGE, S. 1979. North Pacific Shells: (1) Genus Volutopsius Moerch. 26 pp. (7) Genus Plicifusus
Dali. 29 pp. (1980).

TRYON, G. 1881. Manual of Conchology, 3. Philadelphia, Academy of Natural Sciences, 310 pp.
TURTON, W. 1825. Descriptions of some new British shells. Zoological Journal, 2:361-67.

TURTON, W. 1834. Description of some nondescript and rare British species of shells. Magazine of Natural
History, 7:350-53.

VAYSSIERE, A. 1930. Etude zoologique et anatomique sur quelques Gasteropodes Prosobranches provenant
des campagnes du Prince Albert ler de Monaco. Resultats scientifiques (...) Albert Ier, 80:1-26.
VERKRUZEN, T.A. 1875. Bericht liber einen Schabeausflug im Sommer 1874. Jahrbucher der Deutschen
Malakozoologischen Gesellschaft, 2:229-35.

VERKRUZEN, T.A. 1881. Bericht liber meinem Besuch der grossen Bank von Neufundland in Sommer 1880.

Jahrbucher der Deutschen Malakozoologischen Gesellschaft, 8:82-100.

VERRILL, A.E. 1873. Report upon the invertebrate animals of Vineyard sounds and the adjacent waters.

Report of the United States Commissioner of Fisheries, 1:295-778.

VERRILL, A.E. 1879. Preliminary checklist on the marine invertebrates. Newhaven. 32 pp.

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 2. Proceedings of
the United States National Museum, 3:356-405.

NORTH-EAST ATLANTIC NEOGASTROPODA 285

VERRILL, A.E. 1882. Catalogue of marine Molluscs added to the fauna of New England during the past ten
years. Transactions of the Connecticut Academy, 5(2):447-587.

VERRILL, A.E. 1884. Second catalogue of Mollusca..., consisting mostly of deep sea species ...Transactions
of the Connecticut Academy, 6(1): 139-294.

VERRILL, A.E. 1885. Third catalogue of Mollusca...., consisting mostly of deep sea species ...Transactions
of the Connecticut Academy, 6:395-452.

VOKES, E. 1976. Cenozoic Muricidae of the western Atlantic region, part 7. Calotrophon and Attiliosa. Tulane
Studies in Geology and Paleontology, 12(3): 101-32.

WAREN, A. 1980. Marine Mollusca described by John Gwyn Jeffreys, with the location of the type material.

Conchological Society of Great Britain and Ireland, Special publication, 1:1-60.

WATSON, R.B. 1882-83. Mollusca of H.M.S. ‘Challenger’ expedition, part 11 (1882). Journal of the Linnean
Society, 16:247-54; part 13 (1882), 358-72; part 15 (1883), 594-611.

WATSON, R.B. 1886. Report on the Scaphopoda and Gasteropoda collected by H.M.S. ‘Challenger’ during
the years 1873-76. Reports on the Scientific Results of the Challenger Expedition, Zoology, 42:1-756.
WATSON, R.B. 1897. Marine Mollusca of Madeira. Journal of the Linnean Society of London, 26:233-329.
WEBER, J. 1959. Radula of Liomesus stimpsoni Dali. The Nautilus, 72:99-100.

WEINKAUFF, H.C. 1866. Nouveau supplement au catalogue des coquilles marines recueillies sur les cotes de
1’Algerie. Journal de Conchy liologie, Paris, 14:227-48.

WENZ, W. 1941-43. Handbuch der Palaozoologie: Gastropoda 1, 5, (pp. 961-1200; 1941) 6, (pp. 1201-1506;
1943). Berlin, G. Borntraeger.

WEST, D.L. 1973. Notes on the development of Colus stimpsoni. Nautilus, 87:1-4.

WEST, D.L. 1978. Reproductive biology of Colus stimpsoni, 1. Male genital system. Veliger, 20:266-73; 2.

Spermatogenesis; 3. Female genital system. Veliger, 21:1-9, 432-38.

WINCKWORTH, R. 1932. The British marine Mollusca. Journal of Conchology, London, 19:211-52.
WOOD, S.V. 1872. Supplement to the Mollusca from the Crag. London, Paleontographical Society. 98 pp.
WYVILLE THOMSON, C. 1873. The Depths of the sea. London, Macmillan, 527 pp.

286 P. BOUCHET & A. WAREN

INDEX

Italic type indicates valid taxa as used here.

Names in Roman type are synonyms or superseded names.

Names of valid genera are in CAPITAL letters.

abbreviata MOrch, 1869 (Trophon clathratus) = truncatus . 128

abnormis Gray in H.&A. Adams, 1858 (Admete) = viridula . 258

abyssicola Monterosato in Locard, 1897 (Gibberula) . . 267

abyssorum Verrill & Smith, 1 884 (Buccinum) . 189

abyssorum P. Fischer, 1883 (Fusus): Mohnia 211

aculeatus Settepassi, 1977 (Trophonopsis carinatus) = echinatus . 137

acutecostata Philippi, 1844 (Buccinum): Amphissa . 165

ADMETE KrOyer in MOller, 1842 . 257

alabaster Barnard, 1960 (Nux) = tryoni . 249

alba Jeffreys, 1867 (Trophon truncatus) = truncatus . 128

alba Jeffreys in Wyville-Thomson, 1873 (Latirus): Metzgeria . 254

alba n.sp. (Iphinopsis) . .. . 263

alba Friele in Kobelt, 1878 (Mohnia) = mohni . 206

albellus Dunker & Metzger, 1874(Lathyrus) = Metzgeria alba . 254

albinus Monterosato in Settepassi, 1977 (Trophonopsis muricatus) = muricatus . 134

albula Jeffreys, 1869 (Columbella haliaeeti) - acutecostata . 165

alucoides de Blainville, 1830(Murex) = squamosus . 153

amblyterus Watson, 1886 (Fusus) = berniciensis . 193

americanus Bell, 1870 (Fusus) - stimpsoni . 233

amianta Dali, 1889 (Fusus): Fusinus . 161

AMPHISSA H.&A. Adams, 1853 . 165

Anomalosipho Dautzenberg & Fischer, 1912 = Colus . 226

antiqua Linne, 1758 (Murex): Neptunea . 202

apodema Bouchet & Talavera, 1981 (Metzgeria) . 254

AptyxisTrosehel, 1868 . 160

aquitanica Locard, 1897 (Neptunia) - berniciensis . 193

AradomurexCoen, 1943 = Coralliophila 151

arata Verrill, 1880 (Neptunea) = stimpsoni . 233

arcticus Philippi, 1850 (Fusus) = kroeyeri . 231

aronnaxn.s^. (Margineila) . 271

aspera Monterosato in B.D.D., 1882 (Trophon muricatus) = muricatus . 134

asperrimus Leach in Brown, 1827 (Fusus) = muricatus . 134

atlantica Locard, 1897 (Pleurotomella) = koehleri 195

atlantideus n.sp. (Pterynotus) . 147

atractodeum Locard, 1886 (Buccinum) = humphreysianum . 188

Atractus Agassiz, 1839 = Colus . 226

attenuataG.O. Sars, 1878 (Sipho tortuosus) = holboelli . 228

attenuatus Jeffreys, 1877 (Fusus) = jeffreysianus . 230

attenuata Simpson in Marshall, 1902 (Fusus turtoni) - turtoni . 197

attenuata Locard, 1897 (Anachis haliaeeti) = acutecostata . 165

attenuata Harmer, 1918 (Trophon clathratus) = truncatus . 128

Aulacofusus Dali, 1918 = Colus . 202

azonata Locard, 1886 (Buccinum humphreysianum) = humphreysianum . 188

azorica n.sp. (Admete) . 260

azoricus Dautzenberg, 1889 (Fusus) = bocagei . 160

Babelomurex Coen, 1922 = Coralliophila . 151

bamffius Donovan, 1804 (Murex) = clathratus . 129

barvicensis Johnston, 1825 (Fusus): Trophon . 131

basileus Dautzenberg & H . Fischer, 1896 (Pseudomurex): Coralliophila . 154

Bathyclionella Kobelt, 1905 = Belomitra . 223

belliana Jordan, 1890 (Fusus gracilis) = gracilis . 227

BELOMITRA P. Fischer, 1882 . 223

bengasiensis Sturany, 1896 (Fusus) = rostratus . 160

BENTHOBIA Dali, 1889 . 249

BERINGIUS Dali, 1887 . 197

berni ciensis King, 1846 (Fusus): Troschelia . 193

NORTH-EAST ATLANTIC NEOGASTROPODA 287

bicarinataFriele, 1879 (Neptunea lachesis) = voeringi . 216

bicolor Locard, 1896 (Neptunia jeffreysiana) = jeffreysianus 230

blakei Verrill, 1885 (Bela): Mohnia 210

bocagei P. Fischer, 1882 (Fusus): Fusinus 160

borealis Reeve, 1845 (Murex) = fabricii . 135

borealis Adams, 1855 (Admete) = viridula . 258

Boreofusus G.O. Sars, 1878 = Troschelia . 193

Boreotrophon P. Fischer, 1884 = Trophon . 126

bracteatus Brocchi, 1814 (Murex): see Pseudomurex and squamosus 151,153

breviatus Jeffreys, 1882 (Trophon) = muricatus . 134

brevicauda Deshayes, 1832 (Fusus): Neptunea 202

brevis Verrill, 1884 (Sipho stimpsoni) = stimpsoni . 233

brevispira Norman, 1893 (Fusus turtoni) = turtoni 197

brevispira Brogger, 1900 (Sipho) = sabini . 232

Brongus de Gregorio, 1885 = Colus . 226

brucei Melvill& Standen, 1900 (Buccinum) = verkruezeni . 234

brychia Watson, 1882 (Nassa) = frigens . 159

brychia Verrill, 1885(Jumala) - quadruplex . 223

buccinoides Couthouy, 1838 (Cancellaria) = viridula 258

Buccinopsis Jeffreys, 1867 = Liomesus . 185

BUCCINUM Linne, 1758 . 186

caelatulus Verrill, 1884 (Sipho) = caelatus . 212

caelatus Verrill & Smith, 1880 (Neptunea): Mohnia . 212

cancellatus Bivona, 1838 (Fusus) = muricatus . 134

carinata Locard, 1897 (Neptunia berni ciensis) = berni ciensis . . 193

carinatus Bivona, 1832: see echinatus . 137

carinatus Jeffreys, 1883 (Trophon) = echinatus . 137

carolinensis Verrill, 1884 (Urosalpinx): Mohnia . 212

cepula Sowerby, 1880 (Trophon) = clathratus 129

Chalmon de Gregorio, 1885 = Trophon 126

cinara Monterosato, 1 884 (Pagodula carinata) = echinatus . 137

clathratus Linne, 1767 (Murex): Trophon 129

clavatusG.O. Sars, 1878 (Trophon) . 130

climakis Watson, 1886 (Pleurotoma) = quadruplex . 223

coddi Harmer, 1918 (Trophon gunneri) = clathratus 129

Colicryptus Iredale, 1918 = Turrisipho 215

COLUS ROding, 1798 . 226

concinnus Jeffreys, 1883 (Fusus) = mohni . 205

consimilis Marshall, 1902 (Fusus) = jeffreysianus . 230

conspicienda Locard, 1897 (Clionella) = quadruplex . 223

contabulata Friele, 1879(Admete) = viridula . 258

contraria Linne, 1771 (Murex): Neptunea . 203

convoluta Jeffreys, 1867 (Fusus gracilis) = gracilis . 227

CORA LLIOPHILA H.&A. Adams, 1853 152

cossmanni Locard, 1897 (Lacuna) = tryoni . 249

cossmanni Locard, 1897 (Trophon) = echinatus . 137

costatus Hisinger, 1837 (Fusus) = clathratus . 129

costellifer Sowerby, 1818 (Murex) = viridula . 258

costiferus Posselt, 1898 (Sipho) = lachesis . 215

costulata Locard, 1897 (Clionella delicatulina) = quadruplex . 223

costulatissima Locard, 1897 (Anachis haliaeeti) = acutecostata 165

costulatus Cantraine, 1835 (Fusus) = acutecostata 165

coulsoni Jordan, 1890 (Fusus gracilis) = gracilis . 227

couthouyi Jay, 1839 (Cancellaria) = viridula 258

crassa Harmer, 1914 (Sipho togatus) = holboelli . 228

craticulatum Fabricius, 1780(Tritonium) = fabricii . 135

crispa MOller, 1842(Admete) = viridula . 258

cryptodon P. Fischer, 1882(Mitra): Latiromitra . 255

Cryptomitra Dali, 1924 = Belomitra . 223

curta Locard, 1892 (Trophonopsis) = muricatus p.p . 134

curta Locard, 1897 (Marginella marocana) = marocana . 271

curta Locard, 1897 (Marginella impudica) = aronnax . 271

288 P. BOUCHET & A. WAREN

curta Locard, 1897 (Volutella parvulina) = parvulina . 269

curta Locard, 1897 (Anachis haliaeeti) = acutecostata . 165

curta Settepassi, 1977 (Coralliophila aluccides) = squamosus . 153

curtus Jeffreys, 1867(Fusus) = stimpsoni . 233

Cyomesus Quinn, 1981 = Latirornitra . 255

Cypraeolina Cerulli-Irelli, 1911 = Granulina . 269

dabneyi Dautzenberg, 1889 (Trophon) . 136

daleiSowerby, 1825 (Buccinum): see under Liomesus . 185

dalli Friele in Try on, 1881 (Sipho): Turrisipho . 215

danielsseni Friele, 1879(Neptunea): Mohnia 210

dautzenbergi Dali, 1916 (Anomalosipho) = verkruezeni . 234

decorata Locard, 1897 (Meyeria) = amiantus . 161

decorata Locard, 1897 (Nassa brychia) = frigens . 159

decoratus Locard, 1897 (Trophon) = dabneyi . 136

delicatulina Locard, 1897 (Clionella) = quadruplex 223

delicatus Jeffreys, 1833(Fusus) = holboelli . 228

DellinaBeu, 1970 = Belomitra 223

demulcata Locard, 1897 (Pleurotomella) = koehleri 195

depressa Locard, 1897 (Trophonopsis carinatus) = echinatus 137

depressa Settepassi, 1977 (Coralliophiiaalucoides) = squamosus 153

despecta Linne, 1758 (Murex): see under Neptunea . 203

deversus Locard, 1897 (Trophon) = echinatus 137

dispar Verrill, 1884 (Sipho profundicola) = abyssorum (Mohnia) . 211

distincta Leche, 1878 (Admete viridula) = viridula . 258

distincta Posselt, 1895 (Sipho turritus) = holboelli . 228

droueti Dautzenberg, 1889 (Trophon) 136

eburMbrch, 1869(Fusus) = sabini . 232

eburneumM Sars, 1851 (Tritonium) = ovum 186

ecaudata Locard, 1897 (Fusus bocagei) = bocagei 160

ecaudata Locard, 1897 (Meyeria decorata) = amiantus . 161

ecaudis Locard, 1897 (Neptunia) = fenestratus . 217

echinatus Kiener, 1840 (Fusus): Trophon . 137

elatior Middendorf, 1849 (Tritonium clathiatum) = clathratus 129

elegans Jeffreys, 1867 (Fusus berniciensis) = berniciensis . 193

elegans Jeffreys in Dautzenberg & H. Fischer, 1896(Kryptos) = koehleri . 195

elongata Leche, 1878 (Admete viridula) = viridula . 258

elongata Locard, 1896 (Neptunia jeffreysiana) = jeffreysianus . 230

elongata Locard, 1897 (Marginella marocana) = marocana . 270

elongata Locard, 1897 (Trophon droueti) = droueti . 136

elongata Locard, 1897 (Trophonopsis carinatus) = echinatus . 137

elongata Locard, 1897 (Marginella impudica) = impudica . 271

elongata Locard, 1897 (Neptunia berniciensis) = berniciensis . 193

elongata Settepassi, 1977 (Coralliophila babelis) - squamosus . 153

elongatus Pallary in Settepassi, 1977 (Pseudomurex spadae) = fusulus . 150

euthriaeformis Paulus & Mars, 1942 (Buccinum humphreysianum) = humphreysianum 188

exilis Harmer, 1916 (Trophon clathratus) = truncatus . 128

fabricii Beck in MOller , 1 842 (T rophon) 135

fenestratus Turton, 1834 (Fusus): Turrisipho . 217

fischeri Locard, 1897 (Belomitra) = quadruplex . 223

flammulata Locard, 1 886 (Buccinum monterosatoi) = humphreysianum . 188

flemingiana Macgillivray, 1843(Halia) = ovum 186

frieleiKantor, 1981 (Anomalosipho) = Buccinum sp. 186

frielei Harmer, 1914 (Sipho togatus) = gracilis . 227

frigens von Martens, 1878 (Nassa): Nassarius . 159

fuscoapicaia n.sp. (Iphinopsis) . 262

fusiforme Broderip, 1830 (Buccinum) - fenestratus . 217

fusiforme Kiener, 1834 (Buccinum) = humphreysianum . 188

FUSINUS Rafinesque, 1815 160

Fusomurex Coen, 1922 = Coralliophila 151

fusulus Brocchi, 1814 (Murex): Orania . 150

Fusus Lamarck, 1 799: see under Fusinus . 160

NORTH-EAST ATLANTIC NEOGASTROPODA 289

gtfge/n.sp. (Metzgeria) . 254

gallica Locard, 1 896 (Neptunia islandica) = islandica . 229

geministriatus Pfeffer, 1886(Sipho)? = latericeus . 231

GIBBERULA Swainson, 1840 267

gigliolii Monterosato, 1 890 (Pseudofusus) = rostratus . 160

glaber Verkruzen in Kobelt, 1876(Sipho) = gracilis . 227

glyptusVe rrill, 1882 (Sipho): Mohnia . 213

gracile Locard, 1897 (Neptunia fusiformis) = fenestratus . 217

gracile Monterosato in Settepassi, 1977 (Buccinum) = humphreysianum . 188

gracilis DaCosta, 1778 (Buccinum): Colus . 227

Gracilipurpura Jousseaume, 1880 = Fusinus . 160

grandis Morch, 1869(Admete viridula) = viridula . 258

grandis Morch, 1869 (Murex clathratus) = clathratus . 129

GRANULINA Jousseaume, 1888 269

gray/ Dali, 1889 (Nassarina): Orania . 150

grimaldii Dautzenberg, 1889 (Bela) = acutecostata . 165

grimaldii Dautzenberg& H. Fischer, 1896 (Trophon) = echinatus 137

grimaldii Dautzenberg & H. Fischer, 1896 (Fusus) = amiantus . 161

groenlandica Beck in Mdller, 1842 (Mitra): Volutomitra . 251

guernei Dautzenberg, 1891 (Bela) = quadruplex . 223

gunneri Loven, 1846(Tritonium) = clathratus . . 129

haliaeeti Jeffreys, 1867 (Columbella) = acutecostata . 165

hanseni Friele, 1879 (Neptunea) = sabini . 232

hebes Verrill, 1884 (Sipho caelatus) = caelatus . 212

Hesperia Sykes, 1905 (Marginella) . 270

heuglini Mbrch, 1876 (Trophon) = fabricii . 135

hirsutus Jeffreys, 1883 (Fusus) = sabini . 232

Hirtomurex Coen, 1922 = Pseudomurex . 151

hirtus Settepassi, 1977 (Trophonopsis carinatus) = echinatus . 137

hispidulus Verrill, 1884 (Sipho) = abyssorum (Mohnia) . 211

holboelli Mbller, 1842 (Fusus): Colus . 228

howsei Marshall, 1902 (Fusus propinquus) = jeffreysianus . 230

humphreysianum Bennet, 1824 (Buccinum) . 188

hunkinsi Clarke, 1960 (Colus) = danielsseni . 210

impudica P. Fischer, 1883 (Marginella) 271

incarnatum M. Sars, 1851 (Tritonium) = latericeus 231

inflata Jeffreys, 1877 (Fusus berniciensis) = berniciensis 193

inf lata Friele, 1879 (Trichotropis): Iphinopsis 262

inflata Settepassi, 1977 (Coralliophila lamellosa) = squamosus 153

inflatum Aradas& Benoit, 1876 (Buccinum) = humphreysianum 188

ingolf in.sp. (Volvarina) . 272

intermedia Harmer, 1916 (Trophon truncatus) = truncatus 128

intermedius Verkruzen, 1881 (Trophon clathratus) = clathratus . 129

interstriatus Sowerby, 1880 (Trophon) = fabricii . 135

Iphinoella Habe, 1958 = Iphinopsis 261

IPHINOPSIS Dali, 1924 261

islandicus Mohr, 1786 (Murex): Colus 229

jeffreysianus P. Fischer, 1868 (Fusus): Colus . 230

johanseni Dali, 1919 (Plicifusus) = kroeyeri . 231

jousseaumei Locard, 1897 (Marginella) = subturrita . 270

Jumala Friele, 1882 -- Beringius . 197

kieneri Monterosato, 1872 (Buccinum) = humphreysianum 188

kjennerudae norn. nov. (Buccinum) . 190

koehleri Locard, 1896 (Pleurotomella): Kryptos . 195

krampi Thorson, 1951 (Sipho): Mohnia . 212

kroeyeri Moller, 1842 (Fusus): Colus . 231

KRYPTOS Jeffreys in Dautzenberg & Fischer, 1896 195

lachesis Morch, 1869 (Fusus): Turrisipho 215

lactea Jeffreys, 1867 (Buccinum humphreysianum) = humphreysianum 188

lactea Jeffreys, 1867 (Trophon muricatus) = muricatus . 134

lactea Milaschewitsch, 1916 (Trophon breviatus) = muricatus . 134

290 P. BOUCHET & A. WAREN

laetuca Dali, 1889 (Coralliophila) = richardi 152

laevior Leche, 1878 (Admete viridula) = viridula . 258

laevis MOrch, 1869 (Fusus latericeus) = latericeus . 231

laevis Smith, 1890 (Cantharus) = grayi . 150

laevis Marshall, 1902 (Fusus propinquus) = jeffreysianus . 230

lamellatus Gmelin, 1790 (Buccinum) = clathratus 129

lamellosus Philippi, 1836 (Fusus) = squamosus 153

lamellosus Gray, 1839 (Fusus) = clathratus 129

lanceolata Locard, 1897 (Trophon decoratus) = dabneyi 136

largillierti Petit de la Saussaye, 1851 (Fusus) = norwegicus 200

latericeus M Oiler, 1842 (Fusus): Colus 231

Latimurex Coen, 1922 = Coralliophila 151

LATIROMITRA Locard, 1897 . 255

LepadomurexCoen, 1922 = Coralliophila 151

leptaleus Verrill, 1 884 (Sipho): is a Taranis (Turridae) 125

leucas P. Fischer in Locard, 1897 (Murex): Pterynotus 147

limatula Locard, 1896 (Bela) = acutecostata 165

lindahli Posselt, 1898 (Sipho) = glyptus . 213

lintoni Verrill & Smith, 1882 (Trophon): see under squamosa 153

liocephalum Pallary, 1931 (Buccinum) = humphreysianum 188

LIOMESUS Stimpson, 1865 . 185

liratulus Verrill, 1884 (Sipho stimpsoni) = stimpsoni . 233

listen Jonas, 1846 (Fusus) = gracilis . 227

lividus Morch, 1862 (Fusus) = brevicauda 202

locardi Pallary, 1904(Pseudofusus) = rostratus 160

longicaudaSettepassi, 1977 (Coralliophila lamellosa) = squamosus 153

longicaudata Locard, 1897 (Fusus bocagei) = bocagei 160

longurio Weinkauff, 1866 (Fusus) = muricatus . 134

lowei Watson, 1897 (Trophon) = grayi 150

lusitanicum Pallary, 1931 (Buccinum) = humphreysianum 188

lyrata Monterosato in Locard, 1897 (Belomitra) = quadruplex . 223

lyratum Gmelin, 1790 (Buccinum) = clathratus . 129

maclaini Dali, 1902 (Boreotrophon): is an Oenopota (Turridae) . 125

Mada Jeffreys, 1867 = Buccinum . 186

Madiella Wenz, 1943 = Buccinum 186

major Morch, 1869 (Fusus kroeyeri) = kroeyeri . 231

major Brogger, 1901 (Trophon truncatus) = truncatus . 128

major Locard, 1896 (Neptunia jeffreysiana) = jeffreysianus 230

major Locard, 1896 (Neptunia gracilis) = gracilis . 227

major Locard, 1897 (Belomitra spelta) = quadruplex . 223

major Locard, 1897 (Marginella impudica) = impudica . 271

major Locard, 1897 (Anachis haliaeeti) = acutecostata . 165

major Locard, 1897 (Nassa brychia) = frigens . 159

major Locard, 1897 (Fusus bocagei) = bocagei . 160

major Locard, 1897 (Fusus grimaldii) = amianta . 161

major Locard, 1897 (Trophonopsis carinatus and T. varicosissimus) = echinatus . 137

major Locard, 1897 (Pollia fusulus) = fusulus . 150

major Monterosato in Settepassi, 1977 (Trophonopsis muricatus) = muricatus . 134

Mala Cossmann, 1905 = Buccinum . 186

malacitanaeGhisotti&Spada in Maldonado, 1973 (Flinia frigens) = wolffi . 158

Manaria Smith, 1906 . 171

marginata Locard, 1897 (Marginella impudica) = aronnax . 271

MARGINELLA Lamarck, 1799 270

marocana Locard, 1897 (Marginella) . 270

marshalli Ircdale, 1918 (Fusus) = jeffreysianus . 230

maximus Verkriizen, 1881 (Trophon clathratus) = clathratus . 129

METZGERIA Norman, 1879 . 254

Meyeria Dunker& Metzger, 1874 = Metzgeria . 254

MICROVOLUTA Angas, 1874 . 252

minima Reeve, 1856 (Cancellaria): Olssonella . 263

minor Locard, 1896 (Neptunia jeffreysiana) = jeffreysianus . 230

minor Locard, 1897 (Belomitra spelta) = quadruplex . 223

minor Locard, 1897 (Buccinum monterosatoi) = humphreysianum . 188

minor Locard, 1897 (Neptunia fusiformis) = fenestratus . 217

minor Locard, 1897 (Nassa brychia) = frigens . 159

minor Locard, 1897 (Trophon decoratus) = dabneyi . 136

minor Locard, 1897 (Trophonopsis muricatus) = muricatus . 134

minor Locard, 1897 (Fusus bocagei) = bocagei . 160

minor Locard, 1897 (Marginella impudica) = aronnax . 271

minor Pallary, 1900(Pseudomurex spadae) = fusulus . 150

NORTH-EAST ATLANTIC NEOGASTROPODA 291

minor Thorson, 1944 (Sipho turgidula) = gracilis . 227

modesta Settepassi, 1977 (Coralliophila alucoides) = squamosus . 153

moebii Dunker & Metzger, 1874 (Fusus): Turrisipho . 217

mohni Friele, 1877 (Fusus): Mohnia . 205

MOHNIA Friele, 1878 . 205

monterosatoi Locard, 1886(Buccinum) = humphreysianum . 188

monterosatoi Locard, 1897 (Pseudomurex) = squamosus . 153

monterosatoi Locard, 1897 (Gibberula) = abyssicola 267

Morrisonella Bartsch, 1945 = Belomitra . 223

multiaculeatus Settepassi, 1977 (Trophonopsis carinatus) = echinatus . 137

multilamellosus Philippi, 1844 (Murex): see under echinatus . 137

muricatus Montagu, 1803 (Murex): Trophon . 134

mutica Locard, 1897 (Trophonopsis carinatus) - echinatus 137

NASSARIUS Dumeril, 1806 . 158

Nemofusus Cossmann, 1903 = Orania . 149

NEPTUNEA Rbding, 1798 . 202

Neptunella Verrill, 1873 = Colus . 226

nicolayi Nordsieck, 1974 (Anachis) = acutecostata . 165

nicolloni Locard, 1891 (Neptunia) = gracilis . . 227

nigrolabra Verrill, 1880 (Nassa): is a larval Cymatiidae 125

nitidulina Locard, 1897 (Columbella): Mitrella . 168

nodosa Verrill, 1885 (Admete) . 258

normalis Middendorff, 1849 (Tritonium clathratum) = clathratus . 129

norwegicus Gmelin, 1790(Strombus): Volutopsius . 200

Nux Barnard, 1960 = Benthobia . 249

obesa Pallary, 1900 (Pseudomurex spadae) = fusulus . 150

obesus Sowerby, 1880 (Fusus) = islandicus . 229

obesus Verrill, 1884 (Sipho) = caelatus . 212

oblitum Sykes, 1911 (Buccinum) 189

obliquispira Nordsieck, 1977 (Pleurotomoides): larval shell of Coralliophilidae . 151

OLSSONELLA Petit, 1970 . 263

ORANIA Pallary, 1900 . 149

ossiania Friele, 1879 (Neptunea) = turtoni . 197

ovum Turton, 1825 (Buccinum): Liomesus . 186

Pagodula Monterosato, 1884 - Trophon .

panormitana Monterosato, 1869 (Pyrula): Coralliophila
panormitanum Settepassi, 1977 (Buccinum inflatum) = humphreysianum

paradoxaP. Fischer, 1882 (Belomitra) = quadruplex .

paraelatior Kantor, 1981 (Helicofusus) = verkruezeni
Parasipho Dautzenberg& Fischer, 1912 = Colus

partenopaeum Settepassi, 1977 (Buccinum inflatum) = humphreysianum

parvulina Locard, 1897 (Volutella): Granulina .

parva Verrill & Smith, 1 882 (Sipho): Mohnia .

pellucidus Hancock, 1846 (Fusus) = latericeus .

peregra Locard, 1897 (Neptunia) = fenestratus .

perexiguum Dautzenberg, 1925 (Buccinum) = oblitum .

perfectus P. Fischer, 1882 (Pseudomurex) = squamosus .

perminutus Dali, 1927 (Fusus): see under Troschelia .

pertenuis Sykes, 1911 (Neptunea) = latericeus .

perversus Kiener, 1840 (Fusus) = contraria .

pfaffiMbrch, 1876 (Fusus) = sabini .

pianosana Monterosato in Settepassi, 1977 (Ocinebrina) - squamosus .

Pinon de Gregorio, 1885 = Trophon .

planulus Verrill, 1882 (Sipho pygmaeus) = pygmaeus .

Pleurobela Monterosato in Locard, 1897 = Belomitra .

plicifera Brbgger, 1900 (Sipho verkruezeni) = verkruezeni

Plicifusus Dali, 1902 = Colus .

productaG.O. Sars, 1878 (Admete viridula) = viridula .

profundicola Verrill & Smith, 1884 (Sipho) = abyssorum (Mohnia) .

propinquus Alder, 1848 (Fusus) = jeffreysianus .

Pseudofusus Monterosato, 1884 = Fusinus .

Pseudomurex Monterosato, 1872 = Coralliophila .

PTER YNOTUS Swainson, 1833

pubescens Verrill, 1882 (Sipho): Colus .

pullus Reeve, 1848 (Fusus) = latericeus .

pumilaMOrch, 1869 (Fusus kroeyeri) = kroeyeri .

pupoidea Locard, 1897 (Neptunia) = jeffreysianus .

126

152
188
223
234
226
188
269
206
231
217
189

153
193

231
203

232
153
126
232
223
234
226
258
211

230
160
151
147

231
231
231
230

292 P. BOUCHET & A. WAREN

pupulaP. Fischer, 1882(Fusus) = jeffreysianus 230

pusillum M. Sars, 1859 (Tritonium) = alba (Metzgeria) 254

puxleianum Leach, 1847 (Buccinum) = humphreysianum 188

pygmaeus Gould, 1841 (Fusus islandicus): Colus . 232

pyrrhias Watson, 1883 (Murex) = fusulus 150

quadruplex Watson, 1882 (Pleurotoma): Belomitra 223

reeveanus Petit de la Saussaye, 1851 (Fusus) = sabini . 232

retusa Monterosato in Locard, 1897 (Gibberula) = abyssicola . 267

richardi P. Fischer, 1882(Murex): Coralliophila . 152

richardi Dautzenberg& H. Fischer, 1896 (Trophon) = droueti . 136

richardsoni Gray in Morch, 1869 (Trophon) = clathratus . 129

robustus Settepassi, 1977 (Pseudomurex spadae) = fusulus . 150

rossi Leach in Morch, 1857 (Tritonium) = clathratus 129

ruderatus Monterosato in Sturany, 1896 (Pseudomurex) = squamosus . 153

rugosissima Locard, 1897 (Clathurella): see under Fasciolaridae . 160

rugosus Jeffreys, 1880 (Trophon) = bocagei . 160

sabini Gray, 1824 (Buccinum): Colus . 232

sadko Gorbunov, 1946(Admete) = viridula . 258

sadurnii Bot, 1972(Nassa) = wolffi . 158

sarsi Jeffreys ms. (Fusus/Trophon): see under Turrisipho moebii . 217

sarsi Jeffreys in G.O. Sars, 1878 (Sipho) = moebii . 217

scalariformis Gould, 1840 (Fusus) = clathratus . 129

scalaris Jeffreys, 1867 (Trophon truncatus) = truncatus . 128

schantaricum Middendorff, 1849 (Tritonium) ? = turtoni, ? = brevicauda . 197,202

sentix Bayer, 1971 (Coralliophila) . 154

simplex \e rrill, 1884 (Sipho): Mohnia . 206

Sinistralia H.&A. Adams, 1853 = Fusinus . 160

sinistrorsus Deshayes, 1832 (Fusus) = contraria 203

sinuosa Brogger, 1900 (Sipho togatus) = sabini . 232

Sipho: see discussion under Colus . 226

Siphonella Verrill, 1879 = Colus . 226

Siphonellona Wenz, 1944 = Colus . 226

Siphonorbis Mdrch, 1869 = Colus . 226

solida Jeffreys in G.O. Sars, 1878 (Boreofusus berni ciensis) = berni ciensis . 193

solidulus Sowerby, 1880 (Fusus) = stimpsoni . 253

spadae Libassi, 1859(Murex) = fusulus . 150

specialis Locard, 1897 (Latiromitra) = cryptodon . 255

spelta Monterosato in Locard, 1897 (Belomitra) = quadruplex . 223

spinosa Locard, 1897 (Trophonopsis carinatus) = echinatus . 137

spitzbergensis Reeve, 1855 (Fusus) = brevicauda . 202

squamosus Bivona, 1838 (Fusus): Coralliophila . 153

squamulata Settepassi, 1977 (Coralliophila alucoides) = squamosus . 153

squamulosus Philippi, 1836 (Fusus) = squamosus . 153

stimpsoni Mdrch, 1868 (Fusus): Colus . 233

striatum Philippi, 1844 (Buccinum) = humphreysianum . 188

striatus Reeve, 1847 (Fusus) = stimpsoni . 233

striata Middendorff, 1849 (Tritonium islandicum) = gracilis . 227

striata Jeffreys in Wyville-Thomson, 1873 (Buccinopsis) = oblitum 189

striata Milaschewitsch, 1916 (Trophon breviatus) = muricatus . 134

StrombellaGray, 1857 = Volutopsius . 200

subturrita P. Fischer, 1883 (Marginella impudica) . 270

sulcata Middendorff, 1849 (Tritonium islandicum) = islandicus 229

sulcatum Friele, 1882 (Buccinum) = kjennerudae . 190

superstes n.sp. (Microvoluta) . 252

TACIT A Lus, 1971 . 210

tener Jeffreys ms. (Fusus) = mohni . 206

tenuis Monterosato, 1884 (Pagodulacarinata) = echinatus . 137

tenuistriata Harmer, 1918 (Neptunea) = stimpsoni . 233

terebralis Gould, 1860 (Neptunea) = brevicauda . 302

terraenovae nom.nov. (Colus) . 233

testae Aradas, 1847 (Buccinum) = acutecostata 165

tiberianus Brusina, 1871 (Murex spadae) = fusulus . 150

togatus M6rch, 1869 (Fusus) = sabini . 232

torra Locard, 1897 (Neptunia) = jeffreysianus . 230

tortuosus Reeve, 1855 (Fusus) = holboelli . 228

Tritonellium Valenciennes, 1858 = Buccinum . 186

NORTH-EAST ATLANTIC NEOGASTROPODA 293

Tritonofusus Morch, 1857 = Colus . 226

TROPHON Montfort, 1810 126

Trophonopsis B.D.D., 1882 = Trophon . 126

TROSCHELIA Morch, 1876 . 193

trumbulli Linsley, 1845 (Fusus) = pygmaeus . 232

tru neat us Strom, 1768 (Buccinum): Trophon . 128

tryoni Da\\, 1889 (Benthobia) . 249

tumida Norman, 1893 (Fusus turtoni) = turtoni 197

turgidulus Jeffreys in Friele, 1877 (Fusus): Colus . 234

turris Monterosato in Settepassi, 1977 (Coralliophila) = squamosus . 153

TURRISIPHO Dautzenberg & H. Fischer, 1912 . 215

turritum M. Sars, 1859 (Tritonium) = holboelli . 230

turrita Jeffreys, 1867 (Fusus propinquus) = jeffreysianus . 230

turtoni Bean, 1834 (Fusus): Beringius . 197

Ukko Friele in Norman, 1893 = Beringius . . 197

Undacolus Nordsieck, 1968 = Turrisipho . 215

undata Leche, 1878 (Admete viridula) = viridula 258

undatocostata Verkriizen in Kobelt , 1876 (Admete) = viridula . 258

undatum Linne, 1758 (Buccinum) . 187

undulatus Friele, 1881(Sipho) = lachesis . 215

vaginatus deCristofori & Jan,1832(Murex): see echinatus . 137

vallensis Brogger, 1900 (Sipho togatus) = sabini . 232

ventricosus Gray, 1839 (Fusus) = terraenovae . 233

ventricosum Kiener, 1834 (Buccinum) = humphreysianum . 188

ventricosa Middendorff, 1849 (Tritonium clathratum) = clathratus . 129

ventricosa Locard, 1897 (Neptunia jeffreysiana) = jeffreysianus . 230

ventricosa Locard, 1897 (Fusus bocagei) = bocagei . 160

ventricosa Locard, 1897 (Neptunia berniciensis) - berniciensis . 193

verkruezeni Kobelt, 1876 (Sipho): Colus . . 234

vignali Dautzenberg & H. Fischer, 1896 (Marginella):Gibberula . 269

virgata Friele, 1879 (Neptunea) = verkruezeni . 234

viridula Fabricius, 1780 (Tritonium): Admete . 258

voeringi nom.nov. (Turrisipho) . 216

VOLUTOMITRA Gray in H.&A. Adams, 1853 251

VOLUTOPSIUS Morch, 1857 . 200

VOL V A RINA Hinds, 1844 . 272

wolffi Knudsen, 1956 (Nassa): Nassarius

158

Table 3. Distribution of the species.

The bathymetrical distribution is indicated in two ways, either by figures or lines. The heading indicates 100 metres intervals, and the figures the number of samples from the
preceding interval. For some species of which numerous samples are known, a solid line indicates the normal distribution and a row of points scattered findings. In a few cases,
there are two such lines. In these cases the upper line indicates the distribution in the northern part of the area, the lower one in the southern part, n is the number of samples
examined. After the mean depth, the standard deviation is given. This gives some indication of the bathymetrical range.

294 P. BOUCHET & A. WAREN

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Mitrella sp. _ + _ 39 °N _ 1 _ 1 _ bathyal?

NORTH-EAST ATLANTIC NEOGASTROPODA 295

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296 P. BOUCHET & A. WAREN

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