The lower Pliocene gastropods of Le Pigeon Blanc (Loire-Atlantique, northwest France). Part 3 – Muricidae

Abstract

In this paper we review the Muricidae of the Zanclean lower Pliocene assemblage of Le Pigeon Blanc, Loire-Atlantique department, which we consider the ‘type’ locality for Assemblage III of Van Dingenen et al. (2015). 14 species are recorded, of which 2 are new: Hexaplex (Trunculariopsis) ledoni spec. nov. and Ocinebrina lauriatrageae spec. nov. Murex lineatus Millet, 1865 is considered a junior homonym of Murex lineatus Gmelin, 1791 and replaced by Favartia milleti nom. nov. Ocinebra erinacea var. minor Harmer, 1914 is considered a junior subjective synonym of Murex subcontabulatus Millet, 1854.
The muricid assemblage has a far stronger thermophilic character than seen at the latitude of north western France today and we suggest that average Sea Surface Temperatures off the NW French coast in the Zanclean lower Pliocene may have been warmer than they are at these latitudes today, possibly similar to those found today off the southern Portuguese coasts.

Full text

VITA MALACOLOGICA 15: 35VITA MALACOLOGICA 15: 35
et al., 2014, 2016; Van Dingenen et al., 2014, 2015). Here the
Muricidae are revised, and the study is restricted to the locality
of Le Pigeon Blanc, which we consider to be the ‘type’ locality
for Assemblage III gastropods (of Van Dingenen et al., 2015).
In his unpublished thesis, Brébion (1964) of the Centre
National de la Recherche Scientique, Paris recorded seven
muricid species from Le Pigeon Blanc, one of which was
described as new. However, as the thesis was never published,
the names do not comply with article 13 of the ICZN code
(1999) and must be considered nomina nuda.
For geological setting and material and methods (see
Ceulemans et al., 2016).
ABBREVIATIONS
MNHN.F Muséum national d’Histoire naturelle (collection
de Paléontologie), Paris (France).
NHMW Naturhistorisches Museum Wien collection,
Vienna (Austria).
FVD Frank Van Dingenen private collection, Brecht
(Belgium).
LC Luc Ceulemans private collection (Rixensart,
Belgium).
The descriptions adopt the terminology suggested by Merle
(1999, 2001, 2005), see next page:
ABST RAC T
In this paper we review the Muricidae of the Zanclean lower
Pliocene assemblage of Le Pigeon Blanc, Loire-Atlantique
department, which we consider the ‘type’ locality for
Assemblage III of Van Dingenen et al. (2015). 14 species are
recorded, of which 2 are new: Hexaplex (Trunculariopsis) ledoni
spec. nov. and Ocinebrina lauriatrageae spe c. nov. Murex lin-
eatus Millet, 1865 is considered a junior homonym of Murex lin-
eatus Gmelin, 1791 and replaced by Favartia milleti nom. nov.
Ocinebra erinacea var. minor Harmer, 1914 is considered a jun-
ior subjective synonym of Murex subcontabulatus Mil let , 1854.
The muricid assemblage has a far stronger thermophilic char-
acter than seen at the latitude of north western France today and
we suggest that average Sea Surface Temperatures off the NW
French coast in the Zanclean lower Pliocene may have been
warmer than they are at these latitudes today, possibly similar
to those found today off the southern Portuguese coasts.
INTRODUCTION
In this paper we continue our studies on the Neogene gastro-
pod fossil assemblages of northwestern France (see Ceulemans
Vita Malacologica 15: 35-55 8 October 2016
The lower Pliocene gastropods of Le Pigeon Blanc
(Loire-Atlantique, northwest France). Part 31) – Muricidae
Luc CEULEMANS
Avenue Général Naessens de Loncin 1, B-1330 Rixensar t, Belgium
e-mail: luc.ceulem@skynet.be
Frank VAN DINGENEN
Cambeenboslaan A 11, B-2960 Brecht, Belgium
e-mail: fvd@telenet.be
Didier MERLE
Muséum national d’Histoire naturelle, (CR2P, Sorbonne Universités, MNHN, CNRS), 8 rue Buffon, F750 05 Paris CP 43, France
e-mail: dmerle@mnhn.com
Bernard M. LANDAU
Naturalis Biodiversit y Center, P.O. Box 9517, 2300 RA Leiden, Netherlands
Instituto Dom Luiz da Universidade de Lisboa, Campo Grande, 1749- 016 Lisboa, Portugal
International Health Centres, Av. Infante de Henrique 7, Areias São João, P- 8200 Albufeira, Portugal
e-mail: bernielandau@sapo.pt (corresponding author)
Key words: northwestern France, lower Pliocene, Muricidae, new taxa
1) For No. 1 in this series see: Ceulemans, L. Dingenen, F. Van & Landau, B.M.
2016. The lower Pliocene gastropods of Le Pigeon Blanc (Loire-Atlantique,
Northwe st France). Patellogast ropoda and Vetigas tropoda. Cai nozoic Research
16(1): 51-100. (not numbered)
For No. 2 in this series see: Ceulemans, L. Dingenen, F. Van & Landau, B.M.
(in press). The lower Pliocene gastropods of Le Pigeon Blanc (Loire-Atlanti-
que, Northwest France). 2. Caenogastropoda. Cainozoic Research 16(2).

VITA MALACOLOGICA 15: 36
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
VITA MALACOLOGICA 15: 36
Trunculariopsis Cossmann, 1921
Purpura Rö d i n g, 1798: 139. Typ e specie s (by sub s e quent de sig-
nation, Winckworth, 1945): Murex trunculus Linnaeus,
1758, present-day, Mediterranean. Invalid: junior homonym
of Purpura Bruguière, 1789.
Polyplex Perry, 1810: sgn. M7. Type species (by subsequent
designation, Opinion 911 (ICZN, 1970): Polyplex pur-
purescens Perry, 1810 (= Murex trunculus Linnaeus, 1758),
present-day, Mediterranean.
Truncularia Monterosato, 1917: 20. Type species (by mono-
ty py): Murex trunculus Linnaeus, 1758, present-day,
Mediterranean. Invalid: junior homonym of Truncularia
Wiegmann, 1832 [Bryozoa].
Trunculariopsis Cossmann, 1921: 79. Type species (by typi-
cation of replaced name): Murex trunculus Linnaeus, 1758,
present-day, Mediterranean, nom. nov. pro Truncularia
Monterosato, 1917, non Wiegmann, 1832.
Murithais Grant & Gale, 1931: 729. Type species (by origi-
nal designation): Murex trunculus Linnaeus, 1758, pres-
ent-day, Mediterranean. Invalid: junior objective synonym
of Trunculariopsis Cossmann, 1921, with the same type
species.
Hexaplex (Trunculariopsis) ledoni spec. nov.
Pl. 1 Figs 1-3, Textg. 1
Hexaplex bourgeoisi Tournouër, 1875 – Brébion, 1964: 362
(partim, Assemblage III specimens only).
Type material. — Holotype MNHN.F.A57390, height 25.3
mm, width 14.9 mm (Pl. 1 Fig. 1, Text-g. 1); paratype 1
MNHN.F.A57391, height 20.0 mm, width 12.2 mm (Pl. 1 Fig.
2); paratype 2 NHMW 2015/0133/0308, height 46.3 mm, width
29.5 mm (Pl. 1 Fig. 3); paratype 3 NHMW 2015/0133/0405,
height 28.6 mm, width 17.4 mm; paratype 4, height 25.4 mm,
width 14.9 mm.
Other material. — Maximum height 40.9 mm. NHMW
2015/0133/0103 (23); LC (4). Same locality.
Et y molog y. — Named after Daniel Ledon, excellent collec-
tor of French fossil shells and friend to the authors. Hexaplex
gender male.
Locus typicus. — Le Pigeon Blanc, Le Landreau, Nantes
area, Loire-Atlantique department, NW France.
Stratum typicum. — Zanclean, lower Pliocene.
Diagnosis. — A medium-sized Hexaplex (Trunculariopsis)
species with a paucispiral protoconch consisting of two smooth
whorls, spire moderately elevated, four varices per whorl, one
intervarical node, last whorl: P1-P6 subequal; only P1 forming
a short spine on apertural varix, secondary and tertiary spiral
sculpture present, aperture with six pairs of denticles within,
siphonal canal relatively long: ADP, MP, ABP well-developed,
forming small spines where they cross the varices.
Description. — Shell medium-sized, solid, fusiform, spire
P Primary cord
s secondary cord
t tertiary cord
Ad adapical (or adapertural)
Ab abapical (or abapertural)
SP Subsutural cord
IP Infrasutural primary cord (primary
cord on shoulder)
adis adapical infrasutural secondary cord
(shoulder)
abis abapical infrasutural secondary cord
(shoulder)
P1 Shoulder cord
P2-P6 Primary cords of the convex part of
the teleoconch whorl
s1-s6 secondary cords of the convex part
of the teleoconch whorl
example: s1 = secondary cord between P1 and P2; s2 =
secondary cord between P2 and P3, etc.
ADP adapertural primary cord on the
siphonal canal
MP median primary cord on the siphonal
canal
ABP abapertural primary cord on the
siphonal canal
ads : adapertural secondary cord on the
siphonal canal
ms : median secondary cord on the
siphonal canal
abs : abapertural secondary cord on the
siphonal canal
APERTURE
ID: Infrasutural denticle
D1 to D6: Abapical denticles
SYSTEMATICS
Remarks. — The muricids are a group of marine predatory
gastropods. The classication adopted here is according to
Houart (2001). According to Oliverio & Mariottini (2001)
and Barco et al. (2010), the Coralliophilinae are also arranged
within the Muricidae.
MURICOIDEA Ranesque, 1815
MURICIDAE Ranesque, 1815
Muricinae Ranesque, 1815
Hexaplex Perry, 1810
Hexaplex Perry, 1810: sgn. M7. Type species (by subsequent
designation, Iredale, 1915): Hexaplex foliacea Perry, 1811
a junior synonym of Hexaplex cichoreum (Gmelin, 1791),
present-day, Philippines.

VITA MALACOLOGICA 15: 37VITA MALACOLOGICA 15: 37
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
Distribution. — Lower Pliocene: Atlantic, NW France
(Brébion, 1964).
Re mark s. — At rst gl ance these shel ls rese mble sma ll spec -
imens of Hexaplex (Trunculariopsis) bourgeoisi (Tournouër,
1875) from the middle Miocene Langhian of NW France.
However, H. (T.) bourgeoisi has a tall dome-shaped multispi-
ral protoconch, with a small nucleus (Pl. 1 Fig. 4), whereas
the species from Le Pigeon Blanc have a paucispiral proto-
conch, with a large, bulbous nucleus. The teleoconch is similar
to that of smaller specimens of H. (T.) bourgeoisi such as that
illustrated by Merle et al. (2011: 40, g. 6). The spire in the
NW French Pliocene species is possibly slightly lower and less
scalate, but we can nd no objective character by which to sep-
arate the teleoconch shell. Nevertheless, the differences in the
protoconch are sufcient to separate the middle Miocene and
Pliocene shells from NW France. Therefore, the protoconch
type is the only reliable distinguishing character between the
lower Pliocene Hexaplex (Trunculariopsis) from Le Pigeon
Blanc and the middle Miocene H. (T.) bourgeoisi, also from
NW France. The species with the paucispiral protoconch is
here described as new: Hexaplex (Trunculariopsis) ledoni
sp e c. n ov.
Millet (1854: 163) erected the name Murex asper (nomen
nudum), later made available by a description in Millet (1865:
593). This referred to the Hexaplex-like muricid from the
“Redonian”. Unfortunately this name is invalid as it is a jun-
ior homonym of M. asper Linnaeus, 1758. Millet’s taxon was
based on material from Assemblage I localities of Sceaux-
d’Anjou, Thorigné and Reneauleau. Until we review the pro-
scalate, moderately high, ornament spinose only at shoulder.
Protoconch paucispiral, consisting of two smooth, convex
whorls with a large nucleus. Teleoconch consisting of ve
whorls. Suture impressed, irregular. Spire of moderate height,
with moderately broad sutural ramp, roundly angulated at
shoulder, convex below, with periphery just below mid-whorl.
Axial sculpture on rst visible whorl (third teleoconch whorl)
consisting of four raised rounded varices, with two prominent
intervarical ridges, axial sculpture weak on sutural ramp, high
and rounded below. Spiral sculpture consisting of narrow,
rounded cords, overlain by numerous nely scabrous spiral
threads. P1 forming a short, blunt spine at shoulder, subse-
quent cords producing a subobsolete spine where they cross
the varices. Last whorl with a relatively narrow sutural ramp,
roundly angulate at shoulder, convex below, strongly con-
stricted at base, with ve rounded varices and one intervari-
cal ridge. Spiral sculpture; P1-P6 subequal; only P1 forming
a short spine on apertural varix, secondary cords s3-s5 most
strongly developed in interspaces, whole surface covered in
close-set tertiary threads, interrupted by the growth lines,
giving them a nely beaded appearance. Aperture small,
roundly ovate; outer lip erect, with a crenulated edge, dentic-
ulate within, denticles split, arranged in six pairs. Anal notch
marked by a small adapical notch, bordered medially by a
ridge of parietal callus. Columellar lip concave, smooth, nar-
row, adpressed. Siphonal canal relatively broad, long, open,
weakly recurved. Siphonal fasciole low, rounded; pseudo-um-
bilicus absent; ADP, MP, ABP well-developed, forming small
spines where they cross the varices.
Fig. 1. Hexaplex (Trunculariopsis) ledoni spec. nov., holotype MNHN.F.A57390, height 25.3 mm, width 14.9 mm, Le Pigeon Blanc, Le
Landreau, Nantes area, Loire-Atlantique department, NW France, Zanclean, lower Pliocene. Sculptural detail following terminology of Merle
(1999, 2001, 2005).

VITA MALACOLOGICA 15: 38VITA MALACOLOGICA 15: 38
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
Dermomurex (Dermomurex) scalaroides (Blainville, 1829) –
Landau et al., 2007: 22, pl. 6, g. 1 (with synonymy) ; Merle
et al., 2011: 214, text-g. 72A, B, G, H, pl. 166, g. 9, pl. 168,
gs 1-5 ; Goret et al., 2013: 12, text-g. 9, pl. 3, g. 3.
Material and dimensions. — Height 16.9 mm. NHMW
2015/0133/0310 (1, Pl. 1 Fig. 5). Le Pigeon Blanc, Le Landreau,
Nantes area, Loire-Atlantique department, NW France.
Distribution. — Middle Miocene: Proto-Mediterranean
Sea, Italy (Bellardi, 1873). Middle Miocene: Atlantic, Loire
Basin (Glibert, 1952). Upper Miocene: Atlantic, NW France
(Brébion, 1964); Proto-Mediterranean Sea, Italy (Bellardi,
1873). Lower Pliocene: Atlantic, NW France (Brébion, 1964);
western Mediterranean, France (Goret et al., 2013). Upper
Pliocene: western Mediterranean, Estepona (Landau et al.,
2007); central Mediterranean (Sacco, 1904; Ruggieri et al.,
1959; Andreoli & Marsigli, 1992). Present-day: Atlantic coast
of West Africa, Senegal, southern Mediterranean, 0.5-100 m,
on dead coral, algae and rocks (Houart, 2001).
Discussion. — Dermomurex (Dermomurex) scalaroides
(Blai nville, 1829) is uncommon in the Le Pigeon Blanc assem-
blage. The single specimen at hand is more slender than usual
for the species, although slender specimens also occur in the
Mediterranean today (Merle et al. 2011, pl. 168, g. 2). The
protoconch is paucispiral (Pl. 1 Fig. 4c), similar to that of pres-
ent-day specimens illustrated by Houart (2001: 32, g. 26).
Brébion (1964: 376) recorded this species from Assemblage
II localit y (Apigné) and Assembla ge III (Le Girondor), to which
we add Le Pigeon Blanc. Like many other gastropods present in
Assemblage III, D. (D.) scalaroides is a relatively thermophilic
species, with its present-day distribution bounded to the north
by the southern Mediterranean. Its presence in the Zanclean of
NW France suggests warmer Sea Surface Temperatures (SST)
than those that prevail at this latitude today.
Dermomurex (Dermomurex) tenellus (Mayer, 1869)
Pl. 1 Fig. 6
Murex tenellus Mayer, 1869: 83, pl. 3, g. 5; – Benoist, 1880:
165, pl. 9, gs 5-6; Cossmann, 1903: 47, pl. 2, g. 14.
Murex (Hexachorda) tenellus Mayer – Cossmann & Peyrot,
toconch of the Assemblage I specimens we are unsure if they
belong to H. (T.) bourgeoisi or H. (T.) ledoni. We hope to return
to this question in our revision of the Assemblage I muricids.
Brébion (1964: 362) recorded Hexaplex bourgeoisi from
Assemblage I localities (Reneauleau, Thorigné, Sceaux-
d’Anjou, St-Clément-de-la-Place, St-Michel), Assemblage III
(Le Girondor, Le Pigeon Blanc, Palluau, La Dixmérie) and
Assemblage IV (St-Jean-la-Poterie). However, each of these
records will have to be reassessed individually and protoconch
type assessed. We provisionally restrict the distribution of the
new species to lower Pliocene Assemblage III.
Dermomurex Monterosato, 1890
Dermomurex s.s. Monterosato, 1890
Poweria Monterosato, 1884: 113. Type species (by original
designation): Murex scalarinus Bivona, 1832, present-day,
Mediterranean. Invalid: Junior homonym of Power ia
Bonaparte, 1840 [Pisces].
Dermomurex Monterosato, 1890: 181. Type species (by typ-
ication of replaced name): Murex scalarinus Bivona,
1832, present-day, Mediterranean. Nom. nov. pro Poweria
Monterosato, 1884, non Bonaparte, 1840 [Pisces].
Hexachorda Cossmann, 1903: 10, 47. Type species (by orig-
inal designation): Murex tenellus Mayer, 1869, Miocene-
Pliocene, France. Junior subjective synonym.
Exachorda Sacco, 1904: 20, 203. Incorrect subsequent spell-
ing.
Remarks. — The placement of Dermomurex Monterosato,
1890 in the subfamily Muricinae is doubtful and provisional,
as molecular data presented by Barco et al. (2010) showed the
subfamily to be polyphyletic. See Merle et al. (2011) for further
discussion.
Dermomurex (Dermomurex) scalaroides (Blainville, 1829)
Pl. 1 Fig. 5
Murex scalaroides Blainville, 1829: 131, pl. 5, gs 5-6.
Aspella scalarioides Blainville, 1826 [sic] – Brébion, 1964: 375.
PL AT E 1
Figs 1-3. Hexaplex (Trunculariopsis) ledoni nov. sp. 1a-b. holotype MNHN.F.A57390, height 25.3 mm, width 14.9 mm. 1a. ventral view. 1b.
dorsal view. 2. paratype 1 MNH N.F.A57391, height 20.0 mm, width 12.2 mm (view showing the paucispiral protoconch). 3a-b. paratype 2
NHMW 2015/0133/0308 height 46.3 mm, width 29.5 mm. 3a. ventral view. 3b. dorsal view. Fig. 4. Hexaplex (Trunculariopsis) bourgeoisi
(Tournouër, 1875), MNHN.F.A57363 Pontlevoy, Loir-et-Cher, France, Langhian, middle Miocene (view showing the multispiral protoconch)
(photo Didier Merle). Figs 5a-c. Dermomurex scalaroides (Blainville, 1829), NHMW 2015/0133/0310, height 16.9 mm. 5a. ventral view. 5b.
dorsal view. 5c. view showing the paucispiral protoconch. Figs 6a-b. Dermomurex tenellus (Mayer, 1869), NHMW 2015/0133/0311, height 16.7
mm. 6a. ventral view. 6b. dorsal view. Figs 7a-c. Favartia milleti nom. nov., NHMW 2015/0133/0312, height 13.4 mm. 7a. ventral view. 7b.
dorsal view. 7c. view showing the paucispiral protoconch. Figs 8-9. Favartia suboblonga (d’Orbigny, 1852). 8a-b. NH MW 2015/0133/0314,
height 11.3 mm. 8a. ventral view. 8b. dorsal view. 9a-b. FVD coll., height 15.5 mm. 9a. ventral view. 9b. dorsal view. All: Le Landreau, Le
Pigeon Blanc, Loire-Atlantique department, France, Zanclean, lower Pliocene (except g. 4).

VITA MALACOLOGICA 15: 39VITA MALACOLOGICA 15: 39
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VITA MALACOLOGICA 15: 40
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
VITA MALACOLOGICA 15: 40
0313 (8); LC (23). Le Pigeon Blanc, Le Landreau, Nantes area,
Loire-Atlantique department, NW France.
Distribution. — Upper Miocene (Tortonian): Atlantic, NW
France (Brébion, 1964). Lower Pliocene: Atlantic, NW France
(Brébion, 1964). Upper Pliocene-Pleistocene: Atlantic, NW
France (Brébion, 1964).
Discussion. — Favartia milleti nom. nov. is characterised
by its strongly scalate spire. The spire whorls are sculptured
by two narrow, elevated spiral cords crossed by about six to
eight axial lamellae. The last whorl is short, with a shallow,
at to weakly concave subsutural platform, sharply angled at
the shoulder, convex below and constricted at the base. P1-P5
are narrow, strongly developed, P1, P2 and P3 are hooked at
the labial varix. P6 is weaker, forming a short erect spine at the
aperture. ADP and MP are also strongly developed, forming
an open spine at their extremities. Over the entire surface the
lamellae are spinose where they cross the spiral cords. The
aperture is ovate and the siphonal canal is open and moder-
ately long. Favartia lineata has a paucispiral protoconch (Pl. 1
Fig. 7c).
Landau et a l. (2007: 45) recognised fou r species of Favartia in
the European Miocene: Favartia absona (De Cristofori & Jan,
1832), F. suboblonga (d’Orbigny, 1852), F. exc isa (Grateloup,
1833) and F. czjzeki (Hörnes, 1856), the rst two of which sur-
vived into the Pliocene, but not the Pleistocene. The authors
omitted a fth species: Favartia collega (Boettger, 1906)
described from the Romanian middle Miocene of “Kostej”
(= Coșteiu de Sus) and “Lapugy” (= Lăpugiu de Sus). This is a
small, slender species, with the spiral cords elevated and well
developed across the entire whorl width. A juvenile specimen
at hand (NHMW coll.) has a multispiral, dome-shaped proto-
conch. Favartia milleti represents a sixth species, with an upper
Miocene to lower Pliocene distribution, restricted geographi-
cally to NW France. Favartia excisa, F. suboblonga and F. col-
lega can be separated from F. m ille ti by having a planktotroph-
ic-type protoconch consisting of about 3.5 whorls. Favar tia
absona has a more globose shell than F. m ille t i, the spire is less
scalate, the siphonal canal less slender and the sculpture more
spinose, P1-P6 all forming spines where they cross the axial
ribs. We have not seen any specimens of F. czjzeki, but this
seems to be a more slender species than F. millet i.
Brébion (1964: 379) recorded this species from Assemblage
I localities (Thorigné, Sceaux-d’Anjou), Assemblage III
(Le Girondor, Le Pigeon Blanc, Palluau, La Dixmérie) and
Assemblage IV (Gourbesville).
Favartia suboblonga (d’Orbigny, 1852)
Pl. 1 Figs 8-9
Murex oblongus Grateloup, 1833: pl. 100 (non M. oblongus
Brocchi, 1814).
Murex suboblongus d’Orbigny, 1852: 73 (nom. nov. pro Murex
oblongus Grateloup, 1833, non Brocchi, 1814).
Aspella (Favartia) incisa var. excisa Grateloup, 1833 –
Brébion, 1964: 377.
1924: 137, pl. 12, gs 48-49.
Aspella tenella Mayer, 1869 – Glibert, 1952: 297, pl. 6, g. 9;
Mayer, 1869 – Brébion, 1964: 376.
Dermomurex (s.s.) tenellus (Mayer, 1869) – Merle et al., 2011:
215, text-g. 73A, pl. 165, gs 7-9.
Material and dimensions. — Height 16.7 mm. NHMW
2015/ 0133/0311 (1, Pl. 1 Fig. 6); LC (2). Le Pigeon Blanc, Le
Landreau, Nantes area, Loire-Atlantique department, NW
France.
Distribution. — Lower Miocene (Burdigalian): Aquitaine
Basin, France (Cossmann & Peyrot, 1924). Middle Miocene:
Loire Basin, France (Benoist, 1880; Cossmann, 1903; Glibert,
1952; Merle et al., 2011). Upper Miocene (Tortonian): Atlantic,
NW France ( Brébion, 1964; Merle et al., 2011). Lower Pliocene:
Atlantic, NW France (this paper).
Discussion. — Dermomurex (Dermomurex) tenellus
(Mayer, 1869) is characterised by its beaded spiral cords.
Uncommon at Le Pigeon Blanc, the shells are all abraded,
but the beaded cords can be seen in the specimen illustrated.
Brébion (1964: 376) recorded this species from Assemblage
I localities (Sceaux-d’Anjou, Thorigné, St-Clément-de-la-
Place), to which we add the Assemblage III locality of Le
Pigeon Blanc.
Muricopsinae Radwin & D’Atillio, 1971
Favartia Jousseaume, 1880
Favartia Jousseaume, 1880: 335. Type species (by original
designation): Murex breviculus G.B. Sowerby 2nd, 1834,
present-day, Philippines.
Murexiella Clench & Pérez Farfante, 1945: 49. Type species
(by original designation): Murex hidalgoi Crosse, 1869.
Recent, Gulf of Mexico to southern Caribbean. Junior sub-
jective synonym.
Minnimurex Woolacott, 1957: 115. Type species (by original
designation): Minnimurex phantom Woolacott, 1957. Recent,
New South Wales, Australia. Junior subjective synonym.
Caribiella Perrilliat, 1972: 82. Type species (by original desig-
nation): Favartia (Caribiella) alveata Kiener, 1842. Recent,
Florida to North Brasil. Junior subjective synonym.
Favartia milleti nom. nov.
Pl. 1 Fig. 7
Murex lineatus Millet, 1854: 163 (nomen nudum); Millet,
1865: 593 (non Murex lineatus Gmelin, 1791; junior syno-
nym of Buccinum linea Martyn, 1784: valid name, ICZN,
Opinion 479).
Aspella (Favartia) absona lineata Millet, 1854 – Brébion,
1964: 377, pl. 9, gs 5, 6.
Material and dimensions. — Maximum height 14.0 mm.
NHMW 2015/0133/0312 (1, Pl. 1 Fig. 7), NHMW 2015/0133/

VITA MALACOLOGICA 15: 41
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
VITA MALACOLOGICA 15: 41
conch is not preserved in this French Zanclean material.
Favartia suboblonga differs from Favartia absona (De
Cristofori & Jan, 1832) in having a higher spire, broader,
rounded, rather than spinose varices and in having the siph-
onal canal much shorter and strongly dorsally recurved,
whereas that in F. absona is almost straight.
Brébion (1964: 379) recorded this species from Assemblage
I localities (Thorigné, Sceaux-d’Anjou), Assemblage III
(Palluau), we add Le Pigeon Blanc, and Assemblage IV
(Gourbesville).
Muricopsis Bucquoy & Dautzenberg, 1882
Muricopsis Bucquoy & Da utzenbe rg in Buc quoy, Dautzenber g
& Dollfus, 1882: 19. Type species (by original designation):
Murex blainvillei Payraudeau, 1826 (= Murex cristatus
Brocchi, 1814), present-day, Mediterranean.
Risomurex Olsson & McGinty, 1958: 40. Type species:
Ricinula deformis Reeve, 1846 (ICZN Opinion 1623), pres-
ent-day, Cuba. Junior subjective synonym.
Muricopsis cristata (Brocchi, 1814)
Pl. 2 Fig. 1
Murex cristatus Brocchi, 1814: 394, pl. 7, g. 15.
Muricopsis cristatus Brocchi, 1814, var. inermis Philippi, 1836
– Brébion, 1964: 380, pl. 9, g. 7.
Muricopsis (Muricopsis) cristata (Brocchi, 1814) – Landau et
al., 2007: 42, pl. 11, gs 10, 11 (with synonymy).
Muricopsis (Muricopsis) cristata (Brocchi, 1814) – Landau et
al., 2013: 160, pl. 24, g. 2 (with synonymy).
Muricopsis cristata (Brocchi, 1814) – Goret et al., 2013: 15,
pl. 4, gs 2, 3.
Material and dimensions. — Maximum height 21.8 mm.
NHMW 2015/0133/0316 (1, Pl. 2 Fig. 1), NHMW 2015/0133/
0317 (9); LC (17). Le Pigeon Blanc, Le Landreau, Nantes area,
Loire-Atlantique department, NW France.
Distribution. — Middle Miocene: Paratethys (Langhian-
Serravallian): Austria (Hörnes, 1856; Hoernes & Auinger,
1885; Schultz, 1998), Czech Republic (Hörnes, 1856;
Hoernes & Auinger, 1885), Hungary (Kókay, 1966; Strausz,
1966), Poland (Friedberg, 1912; Krach, 1981; Bałuk, 1995),
Romania (Hörnes, 1856; Hoernes & Auinger, 1885); Proto-
Mediterranean Sea (Serravallian): Karaman Basin, Turkey
(Landau et al., 2013). Upper Miocene (Tortonian and
Messinian): Atlantic, NW France (Brébion, 1964); Proto-
Mediterranean Sea: Italy (Montanaro, 1935; Venzo & Pelosio,
1963). Lower Pliocene: Atlantic, NW France (Brébion, 1964);
western Mediterranean, Estepona Basin (Landau et al., 2007),
France (Goret et al., 2013); central Mediterranean, Italy (Chirli,
2000), Tunisia (Fekih, 1975). Upper Pliocene: Atlantic, NW
France (Brébion, 1964), Mondego Basin (Landau et al., 2013);
central Mediterranean (Sacco, 1904; Ruggieri & Greco, 1965;
Favartia suboblonga (d’Orbigny, 1852) – Landau et al., 2007:
46, text-g. 12, pl. 12, gs 5-8 (with synonymy); Landau et
al., 2013: 161, pl. 24, g. 5; Goret et al., 2013: 17, text-g. 13
a, b, pl. 4, g. 6.
Material and dimensions. — Maximum height 12.2 mm.
NHMW 2015/0133/0314 (1, Pl. 1 Fig. 8), NHMW 2015/ 0133/
0315 (1); LC (4); FVD (1, Pl. 1 Fig. 9). Le Pigeon Blanc, Le
Landreau, Nantes area, Loire-Atlantique department, NW
France.
Distribution. — Middle Miocene (Langhian and
Serravallian): Atlantic, Loire Basin, France (Glibert, 1952),
Aquitaine Basin (Cossmann & Peyrot, 1924); Paratethys,
Austria (Hörnes, 1853), Poland (Bałuk, 1995), Romania
(Boettger, 1902; Zilch, 1934); Proto-Mediterranean Sea
(Serravallian), Karaman Basin (Landau et al., 2013). Upper
Miocene (Tortonian): Atlantic, NW France (Brébion, 1964);
Proto-Mediterranean Sea: Italy (Montanaro, 1935). Lower
Pliocene: Atlantic, NW France (Brébion, 1964); west-
ern Mediterranean, France (Goret et al., 2013); central
Mediterranean, Italy (Chirli, 2000). Upper Pliocene: western
Mediterranean, Estepona Basin (Landau et al., 2007); central
Mediterranean, Italy (Sacco, 1904; Cavallo & Repetto, 1992;
Andreoli & Marsigli, 1992). Upper Pliocene-Pleistocene:
Atlantic NW France (Brébion, 1964).
Discussion. — The complicated taxonomic history of this
species was discussed by Landau et al. (2007: 47). Two closely
similar species occur: F. suboblonga (d’Orbigny, 1852) and F.
excisa (Grateloup, 1833). The type locality for Murex excisus
is St-Paul-lès-Dax, Atlantic Lower Miocene, Aquitanian, of
France (Merle, 1999: 289). Both Favartia excisa and F. s ub-
oblonga have a planktotrophic-type protoconch consisting of
about 3.5 whorls. The main difference between F. exc isa and
F. suboblonga is the low growth of the secondary spiral cords
s2 and s3 in F. excis a (see Merle, 1999: 304). The relative fre-
quency of the presence of secondary cords in the two species
is as follows: for F. suboblonga, s1 (0%), s2 (99%), s3 (33%),
s4 (0%), s5 (86%, s6 (88%); for F. exci sa, s1 (0%), s2 (1%),
s3 (1%), s4 (1%), s5 (1%), s6 (11%). Therefore more than 80%
of specimens of F. suboblonga have s2-s5 developed, whereas
less than 1% of F. e xc isa do (Merle, 1999). Favartia excisa
occurs in the French Atlantic Late Oligocene, Chattian to late
Lower Miocene, Burdigalian. Favartia suboblonga coex-
isted with F. exc isa in the French Atlantic Lower Miocene,
Aquitanian, and persisted into the Middle Miocene, Langhian
(Merle, 1999). All the Late Miocene and Pliocene European
shells identied by authors as F. exci sa and F. in cisa are, in our
opinion also, F. suboblonga.
The French specimens from the upper Miocene Tortonian
Assemblage I locality of Sceaux-d’Anjou are indeed smaller
than the t ypical Favartia suboblonga from the Mediterranean
Pliocene, and the aperture is smaller and rounder, however
Merle (1999: 302) noted some degree of intraspecic varia-
bility in the size of the shells within other French Miocene
populations. The Assemblage III specimens from Le Pigeon
Blanc are also relatively small. Unfortunately, the proto-

VITA MALACOLOGICA 15: 42
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
VITA MALACOLOGICA 15: 42

VITA MALACOLOGICA 15: 43
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
VITA MALACOLOGICA 15: 43
Pterynopsis subcontabulata (Millet, 1854)
Pl. 2 Figs 2-4
Murex subcontabulatus Millet, 1854: 163. – Millet, 1865: 592;
Couffon, 1907: 185.
Murex torquis Dollfus, 1905: 367; Murex torquis Dollfus,
1906: 311.
Murex contabulatus Lmk. var. – Couffon, 1908: 56 (non
Lamarck, 1803).
Ocinebra erinacea var. minor Harmer, 1914: 125, pl. 12, gs
9-11.
Murex torquis G. Dollfus – Couffon, 1915: 47.
Ocinebra erinacea var. minor F.W. Harmer – Harmer, 1918:
343, pl. 35, g. 9.
Pterynotus (Pterochelus) canhami (?) var. minor Harmer, 1914
– Brébion, 1964: 367, pl. 8, gs 30-31.
Material and dimensions. — Maximum height 27.2 mm.
MNHN.F.A57350 (Pl. 2 Fig. 2), Saint-Jean-la-Poterie, Mor-
bihan department, Bretagne, France, upper Piacenzian-
Gelasian, upper Pliocene-lower Pleistocene. NHMW 2015/
0133/0406 (1, Pl. 2 Fig. 3), NHMW 2015/0133/0318 (1, Pl. 2
Fig. 4), NHMW 2015/0133/0319 (2); LC (10). Le Pigeon Blanc,
Le Landreau, Nantes area, Loire-Atlantique department, NW
France.
Distribution. — Upper Miocene (Tortonian): Atlantic, NW
France (Brébion, 1964). Lower Pliocene: Atlantic, NW France
(Brébion, 1964); NSB, Coralline Crag, England (Harmer,
1914). Upper Pliocene: NSB, Red Crag, England (Harmer,
1914). Upper Pliocene-Pleistocene: Atlantic, NW France
(Brébion, 1964). Pleistocene: Isle of Man, England (Harmer,
1914).
Discussion. — The original description is rather vague:
“Cette espèce a beaucoup d’analogie avec le M. contabula-
tus, Lamk., quoique moins eflée et plus petite dans toutes ses
dimensions, et que l’on trouve fossile dans l’étage Parisien
(Millet, 1854: 163)”. ?Pterochelus contabulatus (Lamarck,
1803) is a very distinctive Timbellus-like muricid and Millet
can only have been referring to this species. However, Millet’s
discussion goes on “Il se rapproche aussi du M. turoniensis
[sic], Duj.; mais les grosses épines arquées, qui garnissent
le dernier tour de spire, sufront pour l’en distinguer aus-
sitôt”. This sentence compares the species with Hexaplex
(Trunculariopsis) turonensis (Dujardin, 1837), which is quite
a different shaped species. Luckily Millet (1865) provided a
Palla, 1967; Malatesta, 1974; Caprotti, 1976; Chirli, 1988;
Cavallo & Repetto, 1992; Andreoli & Marsigli, 1992a).
Upper Pliocene-Pleistocene: Atlantic, NW France (Brébion,
1964). Pleistocene: western Mediterranean, Balearic Islands
(Cuerda Barceló, 1987); central Mediterranean, Italy (Cerulli-
Irelli, 1911; Bevilaqua, 1928; Malatesta, 1960). Present-day:
Atlantic, southern Portugal and the Canaries, Mediterranean,
intertidal to 120 m, on rocks and dead coral (Houart, 2001).
Discussion. — Muricopsis cristata (Brocchi, 1814) is a
highly polymorphic species, with many synonyms in the fossil
and Recent literature (see Houart, 2001: 92). All the specimens
from Le Pigeon Blanc are of the smooth inermis Philippi, 1836
morphotype (non Murex inermis Dujardin, 1837 = Muricopsis
dujardini Peyrot, 1938). The spinose M. cristata morphotype
cristata has not so far been found in the Pliocene NW French
assemblages. Muricopsis crassicosta (Benoist, 1873) from
the Atlantic upper Oligocene, Chattian and lower Miocene,
Aquitanian of France differs from M. cristata in having a pro-
toconch consisting of three whorls, whereas that of M. cristata
is paucispiral, consisting of only 1.5 whorls, and in developing
pointed spines between the fth and seventh whorls that are
not present in M. crassicosta (Merle, 1999). For further dis-
cussion see Landau et al. (2007: 43). Merle (1999) considered
the middle Miocene Atlantic Loire Basin specimens identied
by Glibert (1952: 299, pl. 6, g. 11) as Muricidea cristata iner-
mis Philippi, 1836 to be a distinct species, Muricopsis dujar-
dini Peyrot, 1938, distinguished from M. cristata by the late
appearance of the secondary spiral sculpture s1, 4-6, which do
not appear until 10 mm height. The specimens from Le Pigeon
Blanc t within the variability of M. cristata.
Brébion (1964: 382) recorded this species from Assemblage I
localities (Reneauleau, Thorigné, Sceaux-d’Anjou, St-Michel,
Beaulieu), Assemblage II (Apigné, Le Temple du Cerisier),
Assemblage III (Le Girondor, Le Pigeon Blanc, Palluau, La
Dixmérie, La Gauvinière) and Assemblage IV (Gourbesville).
Ocenebrinae Cossmann, 1903
Pterynopsis Vokes, 1972
Pterynopsis Vokes, 1972: 3. Type species (by original desig-
nation): Pter ynopsis prosopeion Vokes, 1972 (nom. nov. pro
Murex nysti Koenen, 1867; non Rouault, 1850), Miocene,
Germany.
PL AT E 2
Figs 1a-b. Muricopsis cristata (Brocchi, 1814), NHMW 2015/0133/0316, height 19.4 mm. 1a. ventral view. 1b. dorsal view. Figs 2-4. Pterynopsis
subcontabulata (Millet, 1854) 2. MNHN.F.A57350, height 27.2 mm. 2a. ventral view. 2b. dorsal view. Saint-Jean-la-Poterie, Morbihan
department, Bretagne, France, upper Piacenzian-Gelasian, upper Pliocene-lower Pleistocene. Specimen illustrated by Brébion (1964, pl. 8, g.
31). 3. NHMW 2015/0133/0406, height 22.2 m m. 3a. ventral view. 3b. dorsal view. 4. NHMW 2015/0133/0318, height 27.2 mm. 4 a. ventral view.
4b. dorsal view. Figs 5-7. Jaton dufrenoyi (Grateloup, 1845) 5. NHMW 2015/0133/0320, height 36.4 mm. 5a. ventral view. 5b. dorsal view. 6.
FVD coll., height 21.7 mm. 6a. ventral view. 6b. dorsal view. 7. NHMW 2015/0133/0407, height 26.2 mm. 7a. ventral view. 7b. dorsal view. All:
(except Fig. 2): Le Landreau, Le Pigeon Blanc, Loire-Atlantique department, France, Zanclean, lower Pliocene.

VITA MALACOLOGICA 15: 44
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
VITA MALACOLOGICA 15: 44
Jaton Pusch, 1837
Jaton Pusch, 1837: 135. Type species (by original designa-
tion): Murex decussatus Gmelin, 1791, present-day, West
Africa.
Jaton dufrenoyi (Grateloup, 1845)
Plate 2 Figs 5-7
Murex Dufrenoyi Grateloup, 1845: pl. 30, g. 9.
Murex (Inermicosta) dufrenoyi Grat. – Cossmann, 1903: 26,
pl. 1, g. 5; Cossmann & Peyrot, no. 738, p. 112, pl. 13, gs
29-30
Tritonalia (Tritonalia) dufrenoyi Grateloup, 1840 – Glibert,
1952: 308, pl. 7, g. 6.
Jaton dufrenoyi (Grateloup, 1847) – Lozouet et al., 2001: 55,
pl. 24, g. 6; Goret et al., 2013: 62, text-g. 8, pl. 4, gs 4-6.
Materia l and dimensions . — Maximum height 36.4 m m. NHM W
2015/0133/0320 (1, Pl. 2 Fig. 5), NHMW 2015/0133/0407 (1, Pl.
2 Fig. 6), NHMW 2015/0133/0321 (8 juveniles); LC (7 incom-
plete and juveniles); FVD (1, Pl. 2 Fig. 7). Le Pigeon Blanc,
Le Landreau, Nantes area, Loire-Atlantique department, NW
France.
Distribution. — Lower Miocene (Aquitanian and
Burdigalian): Aquitaine Basin, France (Cossmann & Peyrot,
1924; Lozouet et al., 2001). Lower-middle Miocene: Proto-
Mediterranean, France (Goret & Pons, 2013). Middle Miocene:
Atlantic, Loire Basin, France (Glibert, 1952). Upper Pliocene:
Atlantic, north western France (this paper).
Discussion. — Jaton dufrenoyi (Grateloup, 1845) is simi-
lar to J. sowerbyi (Michelotti, 1841), but differs by its slightly
more fusiform shell shape, and by its smaller, more elongate,
and less rounded aperture. Moreover, all the specimens we
have examined from Saucats (Le Peloua), France (NHMW
coll.) have denticles within the outer lip, whereas all speci-
mens we have examined of J. sowerbyi have a smooth outer
lip. The primary cords P1, P2, P3, P5 and P6 are all relatively
well developed, with P4 slighty weaker, s1 is also more prom-
inent than the rest of the secondary cords. Although not in a
perfect state of preservation, the specimen gured from Le
Pigeon Blanc illustrated well all the distinguishing characters
discussed above. Jaton dufrenoyi is typical of the Aquitanian
and Burdigalian, lower Miocene of the Aquitaine Basin. This
record in the Zanclean lower Pliocene of NW France is the
youngest record for the species, and put in doubt the sugges-
tion made by Landau et al. (2007) that J. dufrenoyi was proba-
bly the predecessor to J. sowerbyi as we have them together in
the Le Pigeon Blanc assemblage.
Lozouet et al. (2001: 55) placed the Middle Miocene
Paratethyan forms reported by Bałuk, (1995) within the syn-
onymy of J. dufrenoyi, however, they do not have denticles
within the outer lip either, and are closer to J. sowerbyi.
fuller description of his species: “Coq. comme fusiforme,
aiguë au sommet, composée de 8 tours de spire: les premiers
sont marqués de petites côtes verticales, et les deux derniers
portent chacun trois expansions lamellaires irrégulières, plus
ou moins épineuses et disposées triangulairement. Tous ces
tours sont anguleux et recouverts de grosses stries transver-
sales. L’ouverture, qui est ovale avec ses bords rééchis en
dehors, est terminée par un canal légèrement arqué; et le bord
droit, qui en se recourbant fait un pli tranchant à l’intérieur,
présente à sa partie supérieure une espèce de sinus très-pro-
noncé. Longueur: 20 millimètres; diamètre: 9-10 millimètres
(1865: 592)”, which clearly describes the species in question.
Vokes (1972) erected the genus Pterynopsis to include
muricid gastropods characterized by having a calcareous
shell structure, tending to exfoliate, a dilated, attened, sub-
circular aperture and an irregular number of varices, usually
about twelve on early whorls diminishing to about three on
later whorls. This seems to be a northern European genus that
rst appears in the Oligocene of Germany and continues to the
upper Pliocene of the North Sea Basin. There are no known
living members of the genus.
Several species have been described from the Pliocene
North Sea Basin. Pterynotus canhami (Wood, 1872) is imme-
diately separated by its relatively broad last whorl and having
the multiple varices typical of the early whorls persisting onto
the last whorl, which usually has six varices. In the remaining
Pliocene North Sea Basin species the varices become reduced
to three on the last whorl. Pterynopsis binominata (Sta adt
in Cossmann, 1909), P. boytonensis (Harmer, 1914) and P.
minor (Harmer, 1914) were all described based on material
from the Coralline Crag (lower Pliocene) and Red Crag (upper
Pliocene) of England. Pterynopsis boytonensis was considered
a large, well-developed specimen of P. binominata by Marquet
(1997), a position with which we agree. Pterynopsis minor is
a smaller, thinner-shelled, more delicate form. The specimens
from Le Pigeon Blanc agree with this form and are similar to
specimens at hand from the lower Pliocene North Sea Basin
of Belgium. Therefore P. minor is a junior subjective syno-
nym of P. subcontabulata. The main difference between P.
subcontabulata and P. binominata is the size and possibly the
slightly higher, more scalate spire in P. subcontabulata, dif-
ferences which seem consistent in the material at hand. We
therefore maintain P. binominata as a distinct species.
The shells from Le Pigeon Blanc are slightly larger than the
specimens from the Assemblage I localities. They have a tall
paucispiral protoconch composed of just under two whorls,
with a large nucleus. The protoconch is better preserved in
specimens from Assemblage I localities and will be illustrated
in the respective paper.
Brébion (1964: 369) recorded this species from Assemblage
I localities (Thorigné, Sceaux-d’Anjou, St-Clément-de-la-
Place, St-Michel), Assemblage III (Le Girondor, Le Pigeon
Blanc, Palluau) and Assemblage IV (St-Jean-la-Poterie,
Gourbesville).

VITA MALACOLOGICA 15: 45
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
Ocinebrina incincta Dollfus mss. – Brébion, 1964: 384, pl. 9,
gs 10-11 (nomen nudum).
Type material. — Holotype MNHN.F.A57351 (Pl. 3 Fig. 2),
height 21.4 mm; paratype 1, MNHN.F.A35352 (Pl. 3 Fig. 3),
height 19.3 mm; paratype 2 NHMW 2015/0133/0323, height
22.5 mm (Pl. 3 Fig. 4); paratype 3 NHMW 2015/0133/0324,
height 27.3 mm (Pl. 3 Fig. 5; Textg. 2).
Other material. — Maximum height 26.0 mm. NHMW
2015/0133/0325 (14); LC (23). Type locality.
Et y molog y. — Named after Agnès Lauriat-Rage of the
Muséum national d’Histoire naturelle, Paris, in recognition of
her work on Redonian bivalves. Ocinebrina gender feminine.
Locus typicus. — Le Landreau, Le Pigeon Blanc, Loire-
Atlantique department, NW France.
Stratum typicum. — Zanclean, lower Pliocene.
Diagnosis. — A medium-sized Ocinebrina species, with a
slender fusiform shell shape, a protoconch composed of two
whorls, an elevated spire with roundly angular whorls, sculp-
ture composed of low, rounded ribs and ne cords, a last whorl
bearing three weak varices, the base sharply delimited by cord
forming small tooth at outer lip, a small aperture bearing four
well-developed denticles and a long sealed siphonal canal.
Description. — Shell medium-sized, solid, fusiform.
Protoconch composed of two convex whorls. Teleoconch of
5.5 relatively elevated, shouldered whorls. Suture impressed,
weakly undulating. First teleoconch bearing two spiral cords
and 20-22 axial ribs of roughly equal strength, forming even
reticulated sculpture. Abapically, ribs decrease in number,
widen to become nine broad rounded ribs on penultimate
Jaton sowerbyi (Michelotti, 1841)
Plate 3 Fig. 1
Murex Sowerbyi Michelotti, 1841: 8, pl. 1, gs 14-15.
Jaton sowerbyi (Michelotti, 1841) – Landau et al., 2007: 38,
text-g. 9 (2), pl. 10, gs 3-4 (with synonymy).
Material and dimensions. — Maximum height 52.9 mm.
NHMW 2015/0133/0322 (1, Pl. 3 Fig. 1); LC (2 incomplete and
juveniles). Le Pigeon Blanc, Le Landreau, Nantes area, Loire-
Atlantique department, NW France.
Distribution. — Middle Miocene: Paratethys, Austria (Hörnes,
1856; Hoernes & Auinger, 1885), Hungary (Kókay, 1966), Poland
(Friedberg, 1912; Bałuk, 1995), Romania (Boettger, 1906; Zilch,
1934). Upper Miocene: Atlantic, Cacela, Portugal (Pereira da
Costa, 1867); Proto-Mediterranean Sea, Italy (Bellardi, 1873;
Montanaro, 1935). Lower Pliocene: Atlantic, NW France (this
paper); central Mediterranean, Italy (Chirli, 2000), Tunisia
(Fekih, 1975). Upper Pliocene: western Mediterranean, Estepona
(Landau et al., 2007); central Mediterranean (?Palla, 1967;
Malatesta, 1974; Cavallo & Repetto, 1992).
Discussion. — As discussed by Landau et al. (2007: 38),
Jaton sowerbyi (Michelotti, 1841) has often been misidenti-
ed as Ocenebra erinaceus (Linnaeus, 1758), or considered
a variety of the latter, however, it differs in having a rounder,
smaller aperture, usually pointed abapically in O. erinaceus,
in having always three varices, whereas O. erinaceus may
have three to seven varices, and in having a longer siphonal
canal. The scaly surface is worn in the Le Pigeon Blanc mate-
rial, but still apparent on the labial wing.
Ocinebrina Jousseaume, 1880
Ocinebrina Jousseaume, 1880: 332. Type species (by orig-
inal designation): Fusus corallinus Scacchi, 1836 [= O.
aciculata (Lamarck, 1822)], present-day, eastern Atlantic,
Mediterranean.
Corallinia Bucquoy & Dautzenberg in Bucquoy, Dautzenberg
& Dollfus, 1882: 24. Type species (by original designation):
Murex aciculatus Lamarck, 1822, present-day, eastern
Atlantic, Mediterranean.
Ocenebrina Cossmann, 1903: 38. Invalid: unjustied emenda-
tion of Ocinebrina Jousseaume, 1880.
Dentocenebra Monterosato, 1917: 21. Type species (by origi-
nal designation): Ocenebra corallinus Scacchi, 1836 [= O.
aciculata (Lamarck, 1822)], present-day, eastern Atlantic,
Mediterranean.
Ocinebrina lauriatrageae spec. nov.
Plate 3 Figs 2-5, Textg. 2
Ocinebra [sic] (Ocinebrina) funiculosa (Borson) – Harmer,
1918: 348, pl. 36, g. 1 (not g. 2 = Ocinebrina funiculosa)
[non Ocinebrina funiculosa (Borson, 1821)].
Fig. 2. Ocinebrina lauriatrageae spec. nov., paratype 3 NHMW
2015/0133/0324, height 27.3 mm, Le Pigeon Blanc, Le Landreau,
Nantes area, Loire-Atlantique department, NW France, Zanclean,
lower Pliocene. Sculptural detail following terminology of Merle
(1999, 2001, 2005).

VITA MALACOLOGICA 15: 46
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc

VITA MALACOLOGICA 15: 47
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
four denticles within the outer lip. Separating Ocenebra and
Ocinebrina species based on shell characters is not always
easy, but in our opinion the species from Le Pigeon Blanc is
closer to Ocinebrina.
Brébion (1964: 369) recorded this species from Assemblage
III localities (Le Pigeon Blanc, Palluau, La Gauvinière).
As noted by Brébion (1964: 385), the specimen from the
Pleistocene of St Erth, England illustrated by Harmer (1918:
pl. 36, g. 1) is likely to be this species, although we cannot
see the denticulation within the aperture. Harmer’s description
(1918: 348) mentions the presence of denticles, but not their
number.
Hadriania Bucquoy & Dautzenberg, 1882
Hadriania Buc quoy & Dautze nberg, 1882: 16, 33. Type species
(by original designation): Hadriania craticulata Bucquoy &
Dautzenberg, 1882, present-day, Mediterranean.
Hadriania brevituba (Millet, 1865)
Plate 3 Fig. 6
Fusus brevitubus Millet, 1854: 162 (nomen nudum); Millet,
1865: 590.
Hadriania brevituba Millet, 1854 – Brébion, 1964: 391, pl. 9,
g. 17.
Material and dimensions. — Maximum height 21.7 mm.
NHMW 2015/0133/0326 (1, Pl. 3 Fig. 6), NHMW 2015/0133/
0327 (1); LC (3). Le Pigeon Blanc, Le Landreau, Nantes area,
Loire-Atlantique department, NW France.
Distribution. — Upper Miocene-lower Pliocene (Tortonian):
Atlantic, NW France (Brébion, 1964). Lower Pliocene:
Atlantic, NW France (Brébion, 1964). Upper Pliocene-Pleisto-
cene: Atlantic, NW France (Brébion, 1964).
Discussion. — Couffon (1915) synonymised Fusus brevitu-
bus Millet with Trophon (Trophonopsis) muricatus (Montagu,
1803), but this is incorrect. Hadriania brevituba (Millet, 1865)
is characterised by its relatively small sized, elongate-fusiform
shell and its straight sealed siphonal canal. The protoconch is
whorl, equal in width to their interspaces. Ribs weaken
over subsutural ramp, obsolete just before adapical suture.
Secondary spiral sculpture develops on second teleoconch
whorl, with tertiary sculpture in the interspaces from the third
teleoconch whorl. Cords made very nely scabrose by close-
set axial growth lines. Last whorl with concave subsutural
ramp, rounded at shoulder, bearing three weakly-developed
varices, spiral cords P1-3, P5-6 roughly equal in strength,
sharply angled at base by stronger basal cord P4, which ter-
minates at lip in small tooth. Aperture small, ovate. Outer lip
thickened by labial varix, bearing stout denticles within; ID
weak, D1-D4 weakening consecutively. Siphonal canal rela-
tively long and broad, sealed.
Distribution. — Lower Pliocene: Atlantic, NW France
(Brébion, 1964). Pleistocene: St Erth, England (Harmer, 1918)
Discussion. — Ocinebrina lauriatrageae spec. nov.
bears features intermediate between those of Ocinebrina
Jousseaume, 1880 and Ocenebra Gray, 1847, to the latter, we
apply the more restricted generic concept of Vermeij & E. H.
Vokes (1997: 72) “a relatively small number of Miocene to
present-day species from western Europe, the Mediterranean
region and Tropical West Africa. These are characterized by
the tendency to form three varices on the last whorl, by the
presence of six to eight primary spiral cords on the last whorl,
a crenulated outer lip without a labral tooth, an adherent or
very lightly erect inner lip, and six weak to strong denticles
on the inner side of the outer lip. In species with varices, the
latter are separated from each other by a single intervarical
node”. The varices are weakly developed in Ocinebrina lau-
riatrageae spec. nov. when compared with some of its lower
Miocene congeners from the Aquitaine Basin (see Lozouet et
al., 2001: pl. 22, gs 6-7, pl. 23, gs 1-8) but nevertheless, they
are present. However, there is more than one intervarical node
and there are only four well-developed denticles within the
outer apertural lip. The presence of a well-delimited P4, mark-
ing an abrupt change in whorl prole from convex above the
basal cord to concave below, is common to several European
Ocinebrina species, such as the Pliocene Mediterranean and
adjacent Atlantic O. concerpta (Bellardi, 1873), but O. lau-
riatrageae can be separated from most of its European fos-
sil and present-day congeners by its rather slender fusiform
shape, with an elevated spire and by the presence of only
PL AT E 3
Figs 1a-b. Jaton sowerbyi (Michelotti, 1841), NHMW 2015/0133/0322, height 52.9 mm. 1a. ventral view. 1b. dorsal view. Figs 2-5. Ocinebrina
lauriatrageae nov. sp. 2a-b. holotype MNHN.F.A57351, height 21.4 mm. 2a. ventral view. 2b. dorsal view. Specimen illustrated by Brébion
(1964, pl. 9, g. 10). 3a-b. paratype 1 MNHN.F.A57352, height 19.3 mm. 3a. ventral view. 3b. dorsal view. Specimen illustrated by Brébion
(1964, pl. 9, g. 11). 4a-c. parat ype 2 NHMW 2015/0133/0323, height 22.5 mm. 4a. vent ral view. 4b. dorsal view. 4c. view showing t he paucispiral
protoconch. 5a-b. paratype 3 NHMW 2015/0133/0324, height 27.3 mm. 5a. ventral view. 5b. dorsal view. Figs 6a-b. Hadriania brevituba
(Millet, 1854), NHMW 2015/0133/0326, height 17.2 mm. 6a. ventral view. 6b. dorsal view. Figs 7a-b. Typhinellus labiatus (de Cristofori &
Jan, 1832), NHMW 2015/0133/0328, height 31.3 mm. 7a. ventral view. 7b. dorsal view. Figs 8a-b. Coralliophila cf. burdigalensis (Tournouer,
1874), NHMW 2015/0133/0331, height 21.5 mm. 8a. ventral view. 8b. dorsal view. Figs 9a-b. Hirtomurex aff. squamosus (Bivona Ant. in Bivona
And., 1838), NHMW 2015/0133/0330, height 14.0 mm. 9a. ventral view. 9b. dorsal view. All: Le Landreau, Le Pigeon Blanc, Loire-Atlantique
department, France, Zanclean, lower Pliocene.

VITA MALACOLOGICA 15: 48
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
Mediterranean (Sacco, 1904; Ruggieri et al., 1959; Caprotti,
1976; Cavallo & Repetto, 1992; Andreoli & Marsigli, 1992).
Lower Pleistocene: central Mediterranean, Italy (Cerulli-
Irelli, 1911). Present-day: Mediterranean, adjacent Atlantic
coast, West Africa to São Tome and Principe (Houart et al.,
2011).
Discussion. — Houart (2015) described the Caribbean spec-
imens as a distinct species T. jacolombi. The Caribbean spe-
cies differs from T. labiatus in having a generally smaller shell,
which is narrower compared to its length, with a length/width
ratio of 1.68-1.77 compared to 1.50-1.72 in T. jacolombi and
in having a different spiral sculpture morphology. Typhinellus
labiatus has the primary cords IP to MP with s2 in all shells
and an occasional s4. Typhinellus jacolombi always has a
s2 spiral cord, often s3, less often s4 and rarely s5 and abs.
Lastly, T. labiatus has a less broad and less expanded apertural
wing abapically and a more constricted last teleoconch whorl
(Houart, 2015: 431). Therefore, the Caribbean Miocene to
present-day specimens identied in the literature as T. labia-
tus are transfered to T. jacolombi Houa rt, 2015.
Brébion (1964: 375) recorded this species from Assemblage I
localities (Thorigné, Sceaux-d’Anjou, St-Clément-de-la-Place,
Beaulieu), Assemblage II (Apigné, Le Temple du Cerisier) and
Assemblage III (Le Girondor, Le Pigeon Blanc, Palluau, La
Dixmérie).
Coralliophilinae Chenu, 1859
Coralliophila H. Adams & A. Adams, 1853
Coralliophila H. Adams & A. Adams, 1853: 135. Type spe-
cies (by subsequent designation, Cossmann, 1903): Murex
neritoideus Gmelin, 1791 [Coralliophila violacea (Kiener,
1836)], present-day, East Africa, Indo-West Pacic.
Coralliophila cf. burdigalensis (Tournouër, 1874)
Plate 3 Fig. 8
cf. Coralliophila burdigalensis (Tournouër, 1874) – Lozouet &
Renard, 1998: 173, g. 2.1-10.
Material and dimensions. — Maximum height 21.5 mm.
NHMW 2015/0133/0331 (1, Pl. 3 Fig. 8); NHMW 2015/0133/
0332 (1). Le Pigeon Blanc, Le Landreau, Nantes area, Loire-
Atlantique department, NW France.
Distribution. — Lower Pliocene: Atlantic, NW France
(Brébion, 1964).
Discussion. — The material from Le Pigeon Blanc at hand
is poorly preserved and incomplete. Nevertheless, the sculp-
ture of narrow, close-set, subequal spiral cords, in which nei-
ther the shoulder nor the base are strongly delimited, and the
broad subobsolete axial ribs, are evident.
This sculpture closely matches that of Coralliophila burdi-
galensis (Tournouër, 1874) (see Lozouet & Renard, 1998, g.
2.1-10). However, in view of the wide strat igraphic gap between
missing in the Le Pigeon Blanc material. The upper Miocene
to present-day Mediterranean and adjacent Atlantic H. crat-
iculata Bucquoy & Dautzenberg, 1882 differs in being larg-
er-shelled, having more shouldered whorls and a bent sipho-
nal fasciole, which may or may not be sealed. We note that
Bouchet et al. (2015) considered H. craticulatoides (Vo kes,
1964) an unnecessary replacement name, as Murex craticula-
tus Linnaeus, 1758 is accepted as a buccinid: Turrilatirus crat-
iculatus (Linna eus, 1758). Murex craticulatus Linnaeus, 1758
sensu Brocchi, 1814 is a muricid and was placed by Bucquoy
& Dautzenberg (1882) in Hadriania Bucquoy & Dautzenberg,
1882. These authors therefore validated the name and it is
therefore not a homonym.
Brébion (1964: 392) recorded this species from Assemblage
I localities (Thorigné, Sceaux-d’Anjou, St-Michel) and the
Assemblage IV locality of Gourbesville. We here add the
Assemblage III locality of Le Pigeon Blanc.
Typhinae Cossmann, 1903
Typhinellus Jousseaume, 1880
Typhinellus Jousseaume, 1880: 335. Type species (by original
designation): Murex sowerbyii Broderip, 1833 (= Murex
labiatus de Cristofori & Jan, 1832), Pliocene to present-day,
Mediterranean, eastern and western Atlantic.
Typhinellus labiatus (de Cristofori & Jan, 1832)
Plate 3 Fig. 7
Murex labiatus de Cristofori & Jan, 1832: 11.
Typhinellus labiatus (de Cristofori & Jan, 1832) – Houart et al.
(partim), 2011: 92, gs 1-24 (not gs 25-29 and Caribbean
records = T. jacolombi Houart, 2015); Landau et al., 2013:
162, pl. 24, gs 8, 9, pl. 64, g. 13 (with synonymy, exclud-
ing Caribbean records = T. jacolombi Houart, 2015); Goret
et al., 2013: text-g. 25, pl. 9, g. 1.
Material and dimensions. — Maximum height 31.3 mm.
NHMW 2015/0133/0328 (1, Pl. 3 Fig. 7), NHMW 2015/0133/
0329 (26); LC (50+). Le Pigeon Blanc, Le Landreau, Nantes
area, Loire-Atlantique department, NW France.
Distribution. — Lower-Middle Miocene: Middle Miocene:
northeastern Atlantic (Langhian): Loire Basin, France
(Glibert, 1952); Paratethys (Langhian-Serravallian): Austria
(Hörnes, 1853; Hoernes & Auinger, 1885; Schultz, 1998),
Hungary (Strausz, 1966), Poland (Bałuk, 1995), Romania
(Csepreghy-Meznerics, 1956); Proto-Mediterranean Sea
(Serravallian): Karaman Basin, Turkey (Landau et al., 2013).
Upper Miocene: Atlantic (Tortonian and Messinian): Atlantic,
NW France (Brébion, 1964). Lower Pliocene: Atlantic, NW
France (Brébion, 1964); western Mediter ranean, France (Goret
et al., 2013); western Mediterranean, Estepona Basin, Spain
(Landau et al., 2007); central Mediterranean, Italy (Chirli,
2000), Tunisia (Fekih, 1975). Lower-upper Pliocene: central

VITA MALACOLOGICA 15: 49
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
includes the Atlantic from the Bay of Biscay to NW Africa
and the Mediterranean as well as the Canaries and Azores.
Oliverio & Gofas (2006) questioned whether all the specimens
illustrated by Bouchet & Warén (1985) as H. basileus were
indeed conspecic. Comparing the NW French shells to H.
basileus they have a slightly lower spire and a rounded shoul-
der, placed lower than in typical H. basileus, as described by
Oliverio & Gofas (2006).
Finding this H. basileus-like form in the Pliocene of NW
France makes us also wonder if these are not all forms of a sin-
gle species or if this is an ancestral form to the two extant spe-
cies. We have insufcient fossil material to be certain of the
specic identity of these shells, however, they clearly belong
to this group of coralliophilid gastropods.
Bouchet & Warén (1985) suggested that H. squamosus
evolved from the Mediterranean Pliocene species H. bractea-
tus (Brocchi, 1814), but this species differs more strongly from
the Le Pigeon Blanc species in being larger, with a far taller,
more scalate spire, in having a sharp shoulder and in having
the primary spiral cords narrower and more elevated.
Brébion (1964: 395) recorded this species only from the
Assemblage III locality of Le Pigeon Blanc.
DISCUSSION
In this paper we record 14 muricid species from the
Assemblage III locality of Le Pigeon Blanc, of which two are
described as new Hexaplex (Trunculariopsis) ledoni spec. nov.
and Ocinebrina lauriatrageae spec. nov., and two are left in
open nomenclature. Eleven muricid genera are represented.
The name Murex lineatus Millet, 1865, a junior homonym of
Murex lineatus Gmelin, 1791, is replaced by Favartia milleti
nom. nov.
In the lower Pliocene four species: Hexaplex (Trun-
culariopsis) ledoni spec. nov., Favartia milleti nom. nov.,
Ocinebrina lauriatrageae spec. nov., Hadriania brevituba
(Millet, 1854) are found only in the north western French
assemblages, although some of them extend their range into
the upper Miocene north western French Assemblage I local-
ities. This will be discussed more detailed in a subsequent
paper. The connections with the Mediterranean Pliocene
assemblages, which share 6 (43%) species with Le Pigeon
Blanc are much stronger than with the more northern English
eastern assemblages, which share 2 (14%) species and only 1
(7%) also occurs in the Pliocene North Sea Basin assemblages.
As in other parts in this series (Ceulemans et al., 2016; Van
Dingenen et al., 2015), at generic level, the assemblage shows
a more thermophilic character compared to that found in the
geographical region today. Hexaplex (Trunculariopsis) has a
distribution today in the Mediterranean and adjacent Atlantic,
including the southern coast of Portugal; Dermomurex,
Muricopsis, Hadriania and Typhinellus have a Mediterranean
distribution, but are found in the Atlantic only south of the
European coast (Houart, 2001). Jaton today is a West African
genus. Similarly, coralliophilid species are not found today
the Chattian upper Oligocene to Burdigalian lower Miocene
C. burdigalensis and the lower Pliocene shells gured herein,
and the poor state of preservation, we hesitate to synonymise
the two. Coralliophila panormitana (Monterosato, 1869) from
the present-day Mediterranean and Atlantic Iberian coast also
has similar ne spiral sculpture, but the specimens seen differ
in having a more dilated aperture and the umbilicus is nar-
rower than in the specimens from Le Pigeon Blanc.
Hirtomurex Coen, 1922
Hirtomurex Coen, 1922: 69. Type species (by original desig-
nation): Fusus lamellosus Philippi, 1836, present-day,
Mediterranean.
Hirtomurex aff. squamosus
(Bivona Ant. in Bivona And., 1838)
Plate 3 Fig. 9
aff. Fusus squamosus Bivona Ant. in Bivona And., 1838: 320,
g. 22.
aff. Coralliophila nov. sp. Brébion, 1964: 394, pl. 9, g. 19.
aff. Coralliophila squamosa (Bivona, 1838) – Bouchet &
Warén, 1985: 153, gs 357-361 (with synonymy).
aff. Coralliophila basileus (Dautzenberg & H. Fischer, 1896)
– Bouchet & Warén, 1985: 154, gs 362-366.
Material and dimensions. — Maximum height 14.0 mm.
NHMW 2015/0133/0330 (1, Pl. 3 Fig. 9); LC (4). Le Pigeon
Blanc, Le Landreau, Nantes area, Loire-Atlantique depart-
ment, NW France.
Distribution. — Lower Pliocene: Atlantic, NW France
(Brébion, 1964).
Discussion. — Hirtomurex squamosus (Bivona Ant. in
Bivona And., 1838) is characterised by its strongly squamous
spiral cords of alternate strength and well-developed shoul-
der. Having said this, the shell can be highly variable, which
has led to the large number of synonyms listed by Bouchet &
Warén (1985: 153). Our shells from the lower Pliocene of Le
Pigeon Blanc are small compared to the specimens illustrated
by Bouchet & Warén (1985: gs 357-361) and Giannuzzi-
Savelli et al. (2003: gs 237-243) and the spire is less sca-
late. They are also closely similar to the shells illustrated by
Bouchet & Warén (1985: gs 362-366) as Hirtomurex basileus
(Dautzenberg & Fischer, 1896), which was said to differ from
H. squamosus in being smaller-shelled and in having a broader
siphonal canal, which is less deviated from the vertical axis.
Comparing our shells with specimens illustrated by Oliverio
& Gofas (2006: g. 7), H. basileus tends to have a less rounded
shoulder than the French Pliocene shells, which is placed
higher. Bouchet & Warén (1985) questioned if these species
were indeed distinct, but pointed out that they had different
distributions; H. basileus only occurs off the Canaries and
Azores, whereas H. squamosus has a wider distribution that

VITA MALACOLOGICA 15: 50
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
REFERENCES
ADAMS, H. & ADAMS, A., 1853-58. The genera of Recent
Mollusca; arranged according to their organization. 2 vols. [Vol.
1: 1-256, pls. 1-32 (1853); 257-484; Vol. 2: 1-92, pls. 33-72 (1854);
93-284, pls. 73-96 (1855); 285-412, pls. 97-112 (1856); 413-540,
pls. 113-128 (1857); 541-660, pls. 129-138 (1858).], John van
Voorst, London.
ANDREOLI, G. & MARSIGLI, S., (1991) 1992. Contributo alla
conosenza della malacofauna pliocenica modenese (Introduzione
e parte 1 – famiglia Muricidae). ― Natura Modenese 1: 3-20.
BAŁUK, W., 1995. Middle Miocene (Badenian) gastropods from
Korytnica, Poland, 2. ― Acta Geologica Polonica 45: 1-255.
BARCO, A., CLAREMONT, M., REID, D.G., HOUART, R.,
BOUCHET, P., WILLIAMS, S.T., CRUAUD, C., COULOUX, A.
& OLIVERIO, M., 2010. A molecular phylogenetic framework
for the Muricidae, a diverse family of carnivorous gastropods. —
Molecular Phylogenetics and Evolution 56: 1025-1039.
BELLARDI, L., 1873. I molluschi dei terreni terziarii del Piemonte
e della Liguria, 1. Cephalopoda, Pteropoda, Heteropoda, Gas-
north of the Bay of Biscay. This would suggest lower Pliocene
SSTs for the region of north western France at least as warm as
those found today off southern Portugal.
ACKNOWLEDGEMENTS
We would li ke to th a nk Ca rlos Marque s da Si l v a of the Universit y
of Lisbon, Portugal, for his advice and help with graphics.
Thanks also to the referees: Geerat J. Vermeij, Department of
Geology, University of California at Davis, USA and Roland
Houart, Institut Royal des Sciences naturelles de Belgique,
Brussels, Belgium for their helpful comments. Thanks also to
Jean-Michel Pacaud (MNHN) for his help on the Brébion’s col-
lection and for advises. The photos of the MNHN materiel were
taken by Peter Massicard (e-recolnat project MNHN). Special
thanks should be given to the families Provost (Le Pigeon
Blanc, Le Landreau, France) and Madeleineau (L’Errière, Le
Landreau, France) for allowing us to excavate on their proper-
ties, without them this publication would not have been possible.
Fig. 3. Geography, stratigraphy and distribution of species found in the Pliocene of Le Pigeon Blanc. For geographic distribution 1 = North Sea
Basin, 2 = Atlantic coasts British Isles, 3 = NW France, 4 = Mediterranean. For stratigraphic distribution black signied Atlantic distribution
(A), grey Mediterranean distribution (M).

VITA MALACOLOGICA 15: 51
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
ordre de matières. Histoire naturelle des vers, 1-2. 1: xviii + 1-344
(1789); 2: 345-757 (1792), Paris (Panckoucke).
BUCQUOY, F. & DAUTZENBERG, P.G., 1882. Les mollusques
marins du Rousillon, 1. Gastropodes, avec atlas de 66 planches
photographées d’apres nature. 1-84, Paris (J.B. Baillière & Daut-
zenberg).
CAPROTTI, E., 1976. Malacofauna dello stratotipo piacenziano
(Pliocene de Castell’Arquato). — Conchiglie 12: 1-56.
CAVALLO, O. & REPETTO, G., 1992. Conch iglie fossili del Roero.
Atlante iconograco. — Associazione Naturalistica Piemontese
Memorie (Associazione Amici del Museo ‘Federico Eusebio’) 2:
1-251.
CERULLI-IRELLI, S., 1911. Fauna malacologica mariana, 5.
Cancellariidae, Marginellidae, Mitridae, Fusidae, Chryso-
domidae, Buccinidae, Nassidae, Columbellidae, Muricidae,
Trito nidae, Cassididae, Cypraeidae, Chenopodidae. — Paleonto-
graphia Italica 17: 280 -325.
CEULEMANS, L., LANDAU, B.M. & VAN DINGENEN, F., 2014.
Carinofusus gen. nov. from the Mio-Pliocene transition of west-
ern France. — Vita Malacologica 12: 23-30.
CEULEMANS, L., VAN DINGENEN, F. & LANDAU, B.M.,
2016. The lower Pliocene gastropods of Le Pigeon Blanc
(Loire Atlantique, Northwest France). Patellogastropoda and
Vetigastropoda. — Cainozoic Research 16(1): 51-100.
CHENU, J.C., 1859. Manuel de conchyliologie et de paleontologie
conchyliologique. 1. vii + 508 pp., Paris (Masson).
CHIRLI, C., 2000. Malacofauna Pliocenica Toscana, 2 Super-
famiglia Muricoidea. 142 pp., Firenze (C. Chirli).
CLENCH, W.J. & PÉREZ FARFANTE , I. 1945. The genus Murex in
the Western Atlantic. — Johnsonia 1(17): 1-58, pl. 1-29.
COEN, G., 1922. Del genere Pseudomurex (Monterosato, 1782). —
Atti della Società italiana di scienze naturali 61: 68-71.
COSSMANN, M., 1903. Essais de paléoconchologie comparée 5.
215 pp., 9 pls., Paris (Cossmann).
COSSMANN, M., 1909. Rectication de nomenclature. — Revue
Critique de Paléozoologie et Paléophytologie 13: 67-68.
COSSMANN, M., 1921. Rectications de nomenclature. — Revue
Critique de Paléozoologie et Paléophytologie 25(2): 79-80.
COSSMANN, M. & PEYROT, A., 1909-1935 (after 1924 continued
by A. Peyrot). Conchologie néogénique de l’Aquitaine. — Actes
de la Société Linnéenne de Bordeaux 63: 73-293 (1909); 64: 235-
400 (1910), 401-445 (1911); 65: 51-98 (1911). 99-333 (1912); 66:
121-232 (1912), 233-324 (1913); 68: 5-210, 361-435 (1914); 69:
157-365 (1917); 70: 5-180 (1918), 181-491 (1919) 73: 5-321 (1922);
74: 257-342 (1923); 75: 71-318 (1924); 77: 51-256 (1925); 78: 199-
256 (1926); 79: 5-263 (1928); 82: 73-126 (1931); 83: 5-116 (1931);
84: 5-288 (1933); 85: 5-71 (1933); 86: 257-353 (1935). Also pub-
lished as a 6 volume book with different pagination as Édition
in-8°, Extrait des Actes de la Société Linnéenne de Bordeaux
(‘Ouvrages couronnés par l’Académie des Sciences, Arts et
Belles-Lettres de Bordeaux’) 1: 1-220 (1909); 221-428 (1911);
429-718 (1912); 2: 1-204 (1913); 205-496 (1914); 3: 1-384 (1917);
385-695 (1919); 4: 1-322 (1922); 323-610 (1924); 5: 1-206 (1927);
207-465 (1928); 6: 1-294 (1931); 295-541 (1932).
COUFFON, O., 1907. Le Miocène en Anjou. — Bulletin de la
Société d’Études Scientiques d’Angers N.S. 36 (1906): 157-196.
tero poda (Muricidae e Tritonidae). — Memorie della Reale Acc-
ademia delle Scienze di Torino (2)27 (1873): 33-294 (reprint 264
pp.) (June 10, 1873).
BENOIST, E.A., 1880. Etude sur les espèces de la sous-famille des
Muricinae obser vées dans le Miocène du Sud-Ouest de la France.
— Actes de la Société Linnéenne de Bordeaux 34: 145-173.
BEV ILAQUA, A. 1928. Studi sulla fauna fossile marina plioce-
nica e quaternaria dell’Isola di Rodi (Egeo). — Atti della Società
Italian a di Scienze Natu rale e del Museo Civico di Stor ia Naturale
di Milano 67: 150-178 .
BIVONA-BERNARDI, A., 1832. Caratteri di un nuovo genere di
conchiglie fossili, estratti dalle Collettanee di Storia naturale,
opera manoscritta del barone Antonino Bivona Bernardi. —
Effemeridi Scientiche e Letterarie per la Sicilia 1: 55-62, pl. 1;
2: 3-8, pl. 2.
BIVONA-BERNARDI, A., 1838. Generi e specie di molluschi
descritti dal Barone Antonio Bivona e Bernardi. Lavori postumi
pubblicati dal glio Andrea dottore in medicina con note ed aggi-
unte. — Giornale di Scienze Lettere e Arti per la Sicilia 61: 211-
227 [stated date march 1838]; 63: 319-324 [stated date september
1838].
BLAINVILLE, H.M.D. de, 1828-1830. Malacozoaires ou animaux
mollusques. [in] Faune Française. 320 p., 48 pl. [livr. 18 (1828):
1-80; livr. 2 (1829): 81-176; livr. 23 (1829): 177-240; livr. 28
(1830): 241-320], Paris (Levrault).
BOETTGER, O., 1902. Zur Kenntnis der Fauna der mittelmiocä-
nen Schichten von Kostej im Krassó-Szörényer Komitat.Mit
einem Situationsplan der Fundpunkte, 2. — Verhandlungen und
Mitteilungen des Siebenbürgischen Vereins für Naturwissen-
schaften zu Hermannstadt 51 (1901): 1-200.
BOETTGER, O., 1906-1907. Zur Kenntnis der Fauna der mit-
telmiocänen Schichten von Kostej im Krassó-Szörényer
Komitat. Gasteropoden und Anneliden, 3. — Verhandlungen
und Mitteilungen des Siebenbürgischen Vereins für Natur-
wissenschaften zu Hermannstadt 54/55: i-viii, 1-99 (1906); 101-
244 (1907).
BORSON, S. 1820-1825. Saggio di orittograa piemontese. —
Memorie della Reale Academia di Scienze di Torino 25: 180-299
(1820); 26: 297-364 (1821); 29:, 251-318 (1825).
BOUCHET, P., HOUART, R. & GOFAS, S., 2015. Hadriania
craticulata Bucquoy, Dautzenberg & Dollfus, 1882. [In]
MolluscaBase 2015. Accessed through: World Register of Marine
Species at http://www.marinespecies.org/aphia.php?p=taxdetails
&id=387245 on 2016-04-13.
BOUCHET, P. & WARÉN, A., 1985. Revision of the Northeast
Atlantic bathyal and abyssal Neogastropoda excluding Turridae
(Mollusca, Gastropoda). — Bollettino Malacologico Suppl. 1:
121-296.
BRÉBION, P., 1964. Les gastéropodes du Redonien et leur signi-
cation, 1-2. — Thèse de doctorat ès-Sciences. Paris (Faculté des
Sciences de l’Université de Paris: 775 pp., 15 pls. (27 June 1964,
unpublished).
BROCCHI, G., 1814. Conchiologia fossile subapennina, con oss-
ervazioni geologiche sugli Apennini e sul suolo adiacente, 1-2.
1-240 (1); 241-712 (2), 16 pls., Milano (Stamperia Reale).
BRUGUIÈRE, J.G., 1789-1792. Encyclopédie méthodique ou par

VITA MALACOLOGICA 15: 52
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
naturae, secundum classes, ordines, genera, species, cum char-
acteribus, disserentis, synonymis, locis etc. Editio decima tertia,
aucta, reformata, cura J.F. Gmelin, 1(6). Vermes testacea. 3021-
4120, Lipsiae (G.E. Beer).
GORET, B., LEDON, D. & PONS, J., 2013. Les Muricidae
(Gastropoda, Muricoidea) du Pliocène Inférieur de Catalogne
(France, Espagne). — Palaeontos 23: 1-52.
GORET, B. & PONS, J., 2013. Les Muricidae (Gastropoda,
Muricoidea) du Miocène de Montpeyroux (Languedoc, France).
— Palaeontos 23: 53-70.
GRANT, U.S., IV & GALE, H.R., 1931. Catalogue of the marine
Pliocene and Pleistocene Mollusca of California and adjacent
regions. — San Diego Society of Natural History Memoirs 1:
1-1036, pls. 1-32.
GRATELOUP, J.P.S. de, 1828-35. Tableau des coquilles fossiles
qu’on rencontre dans les terrains calcaire tertiaires (faluns)
des environs de Dax, dans le Département des Landes, 1-12.
— Bulletins d’Histoire Naturelle de la Société Linnéenne de
Bordeaux 2(9): 72-109 (1828a) (1); 2(10): 123-158 (1828b) (2);
2(10): 192-204 (1828c) (3). — Actes de la Société Linnéenne de
Bordeaux 5(27): 192-204 (1832a) (4); 5(29): 263-282 (1832b) (5);
5(30): 314-344 (1832c) (6); 6(32): 31-48 (1833a) (7); 6(33): 90-100
(1833b) (8); 6(34): 159-164 (1833c) (9); 6(35): 188-212 (1834a)
(10); 6(37): 270-320 (1834b) (11); 7(39): 101-114 (1835) (12).
GRATELOUP, J.P.S. de, 1845-1847. Conchyliologie fossile
des terrains tertiaires du Bassin de l’Adour (environs de
Dax), 1. Univalves. Atlas. Bordeaux (Th. Lafargue): All plates
published 1845, except plates 2, 4, 11 (1847). Note: For dates of
the plates we follow Lesport, J.F., Cluzaud, A. & Verhecken,
A. 2012. Les publications du Docteur Jean-Pierre Sylvestre de
Grateloup sur les mollusques fossiles du Bassin d’Aquitaine (S.-
O. France): dates de par utions et commentaires. — Bulletin de la
Societé Linéenne de Bordeaux (n.s.) 40: 417-485.
GRAY, J.E., 1847. A list of the genera of Recent Mollusca, their
synonyma and types. — Proceedings of the Zoological Society
of London (1847): 129-219 [November].
HARMER, F.W., 1914-1925. The Pliocene Mollusca of Great
Britain, being supplementary to S.V. Wood’s monograph of the
Crag Mollusca, 1. — Monographs of the Palaeontographical
Society 1(1): 1-200 (1914); 1(2): 201-302 (1915), 1(3): 303-461
(1918), 1(4): 463-483 (1919), 2(1): 485-652 (1920), 2(2): 653-704
(1921), 2(3): 705-856 (1923), 2(4): 857-900 (1925).
HÖRNES, M., 1851-1870. Die fossilen Mollusken des Tertiär-
Beckens von Wien. — Abhandlu ngen der Kaiserlich-Königlichen
Geologischen Reichsanstalt 3-4: 1-42, pl. 1-5 (1851), 43-208, pl.
6-20 (1852), 209-296, pl. 21-32 (1853), 297-382, pl. 33-40 (1854),
383-460, pl. 41-45 (1855), 461-736, pl. 46-52 (1856) (3); 1-479, pls
1-85 (1870) (4).
HOERNES, R. & AUINGER, M., 1879-91. Die Gasteropoden
der Meeres-Ablagerungen der ersten und zweiten Miocänen
Mediterran-Stufe in der Österreichisch-Ungarischen Monar -
chie. — Abhandlungen der Kaiserlich-Königlichen Geo logischen
Reichsanstalt 12: 1-382, 50 pls. Published in parts: 1-52, pls 1-6
(1879); 53-112, pls 7-12 (1880); 113-152, pls 13-16 (1882); 153-192,
pls 17-22 (1884); 193-232, pls 23-28 (1885); 233-282, pls 29-36
(1890); 283-330, pls 37-42 (1891); 331-382, pls 43-50 (1891).
COUFFON, O., 1908. Le Miocène en Anjou. (Supplément). —
Bulletin de la Société d’Études Scientiques d’Angers N.S. 37
(1907): 50 -58.
COUFFON, O., 1915. Contribution à l’étude des faluns de l’Anjou,
4. Miocène supérieur, gisement de Saint-Michel-et-Chanveaux.
— Bulletin de la Société d’Études Scientiques d’Angers N.S.
44 (1914): 31-56.
CRISTOFORI, J. de & JAN, G., 1832. Catalogus in quatuor sec-
tiones divisus rerum naturalium in museo exstantium Josephi de
Christofori et Georgio Jan Plurinum Acad. Scient. et Societ. Nat.
Cur. Sodalium ecc. Sectio II. Pars I. Conchylia fossilia ex for-
matione telluris tertiaria in collectione nostra exstantia. 16 pp.,
Parma (Carmignani).
CROSSE, H. 1869. Diagnoses molluscorum novor um. — Journal de
Conchyliologie 17: 408-410.
CSEPREGHY-MEZNERICS, I., 1956. Die Molluskenfauna von Szob
und Let kés. — Magyar Állami Földtani I ntézet Évkönvye (Annales
de l’Institut de Géologie Publique de Hungarie) 45: 361-477.
CU ER DA BARCELÓ, J. 1987. Molluscos marinos y salobres del
Pleistoceno Balear. Palma de Mallorca (Publ. Caja de Baleares
‘Sa Nostra’): 421 pp.
DOLLFUS, G., 1905. Faune Malacologique du Miocène Supérieur
de Gourbesville (Manche) – Étage Rédonien. — L’Association
Française pour l’Avancement des Sciences. Congrès de
Cherbourg 1905: 358-371.
DOLLFUS, G., 1906. Faune malacologique du miocène supérieur de
Beaulieu (Mayenne) – Étage Rédonien. — L’Association Française
pour l’Avancement des Sciences. Congrès de Lyon 1906: 304-315.
DAUTZENBERG, P. & FISCHER, H., 1896. Dragages effectués pa r
l’Hirondelle et par la Princesse Alice 1888-1895. 1. Mollusques
Gastropodes. — Mémoires de la Société Zoologique de France
9: 395-498, pl. 15-22.
DUJARDIN, F., 1837. Mémoire sur les couches du sol en Touraine
et description des coquilles de la craie et des faluns. — Mémoire
de la Société Géologique de la France 2: 211- 311.
FEKIH, M., 1975. Paleoecologie du Pliocène marin au nord de la
Tunisie. — Annales des Mines et de la Géologie 27: 1-195.
FRIEDBERG, W., 1911-28. Mięczaki mioceńskie ziem Polskich
(Mollusca Miocaenica Poloniae), 1. Ślimaki i łódkonogi, 1.
Gastropoda et Scaphopoda. 631 pp. (issued in parts: 1, 1-112, pls
1-5 (1911); 2, 113-240, pls 6-14 (1912); 3, 241-360, pls 15-20 (1914);
4, 361-440, pls 21-26 (1923); 5, 441-631, pls 27-38 (1928), Lwow
(Muzeum Imienia Dzieduszyckich). Reprinted 1951-55 with
slightly different title and pagination, Warszawa (Wydawnictwa
Geologicz ne).
GERTMAN, R.L., 1969. Cenozoic Typhinae (Mollusca:
Gastropoda) of the Western Atlantic region. — Tulane Studies in
Geology and Paleontology 7: 143-191.
GIANNUZZI-SAVELLI, R., PUSATERI, F., PALMERI, A. &
EBREO, C., 2003. Atlante delle conchiglie marine del Medi-
terraneo, 4(1) (Neogastropoda: Muricoidea). 297 pp., Roma
(Edizione de ‘La Conchiglia’).
GLIBERT, M., 1952. Gastropodes du Miocène moyen du Bassin
de la Loire, 2. — Memoires de l’Institut Royal des Sciences
Naturelles de Belgique 2 (46): 241-450.
GMELIN, J.F., 1791. Caroli a Linnei systema natura per regna tria

VITA MALACOLOGICA 15: 53
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
LAMARCK, J.B.P.A. de M., 1822. Histoire naturelle des animaux
sans vertèbres, présentant des caractères généraux et partic-
uliers de ces animaux, leur distribution, leurs classes, leurs
familles, leurs genres, et la citation des principales espèces
qui s’y rapportent, précédée dúne introduction offrant la déter-
mination des caractères essentiels de l’animal, sa distinction
du végétal et des autres corps naturels; enn, l’exposition des
principes fondamentaux de la zoologie, 7. 711 pp., Paris (de
Lamarck).
LANDAU, B.M., HARZHAUSER, M., İSLAMOĞLU, Y. &
SILVA, C.M. da, 2013. Systematics and palaeobiogeography of
the gastropods of the middle Miocene (Serravallian) Karaman
Basin, Turkey. — Cainozoic Research 11-13: 3-584.
LANDAU, B.M., HOUART, R. & SILVA, C.M. da, 2007. The early
Pliocene Gastropoda (Mollusca) of Estepona, southern Spain, 7.
Muricidae. — Palaeontos 11: 1-87.
LINNAEUS, C., 1758. Systema naturae per regna tria naturae,
secundum classes, ordines, genera, species, cum characteri-
bus, differentiis, synonymis, locis, 1. Editio decima, reformata.
824 pp., Holmiae (Laurentii Salvii) [facsimile reprint, British
Museum (Natural History), 1956].
LOZOUET, P., LESPORT, J.F., & RENARD, P., 2001. Révision des
Gastropoda (Mollusca) du stratotype de l’Aquitanien (Miocène
inf.): site de Saucats ‘Lariey’, Gironde, France. — Cossmanniana
(hors série) 3: 189 pp.
LOZOUET, P. & RENARD, P., 1998. Les Coralliophilidae,
Gastropoda de l’Oligocène et du Miocène inférieur d’Aqui-
taine (Sud-Ouest de la France): systématique et coraux hôtes. —
Geobios 31(2): 171-184.
MARQUET, R., 1997. Pliocene gastropod faunas from Kallo (Oost-
Vlaanderen, Belgium), 3-4. Caenogastropoda: Aporrhaidae to
Muricidae (3); Buccinidae to Helicidae (4). — Contributions to
Tertiary and Quaternary Geology 34: 69-149.
MA LAT ESTA, A. 1960. Malac ofauna pleistoce nica di Gra mmichele
(Sicilia). — Memorie per Ser vire alla Car ta Geologica d’Italia 12:
1-196.
MALATESTA, A., 1974. Malacofauna pliocenica Umbra. —
Memorie per Servire alla Carta Geologica d’Italia 13: 1-498.
MAYER, K., 1869. Description de coquilles fossiles des terrains
tertiaires supérieurs (suite). — Journal de Conchyliologie (3)9:
82-88, 282-287.
MERLE, D., 1999. La radiation des Muricidae (Gastropoda:
Neogastropoda) au Paléogène: approche phylogénétique et évo-
lutive. Paris. Unpublished thesis, Muséum national d’Histoire
naturelle : i-vi, 499 pp.
MERLE, D., 2001. The spiral cords and the internal denticles of
the outer lip in the Muricidae: terminology and methodological
comments. — Novapex 2: 69-71.
MERLE, D., 2005. The spiral cords of the Muricidae (Mollusca:
Gastropoda): importance of ontogenetic and topological corre-
spondences for delineating structural homologies. — Lethaia 38:
367-379.
MERLE, D., GARRIGUES, B. & POINTIER, J.P., 2011. Fossil
and Recent Muricidae of the World. Part Muricinae. 648 pp.,
Hackenheim (ConchBooks).
MICHELOTTI, G., 1841. Monograa del genere Murex ossia
HOUART, R., 2001. A review of the Recent Mediterranean and
northeastern Atlantic species of Muricidae. 227 pp., Rome
(Evolver srl.).
HOUART, R. 2015. Description of a new species
of Typhinellus (Gastropoda: Muricidae: Typhinae) from the
Western Atlantic. — Zootaxa 4007(3): 427-432.
HOUART, R., GORI, S. & RYALL, P., 2011. New record of
Typhinellus labiatus (Cristofori & Jan, 1832) (Gastropoda: Muri-
cidae) from São Tomé and Principe and discussion about its clas-
sication and geographical distribution. — Novapex 12: 91-9 7.
ICZN [International Commission on Zoological Nomen clature],
1957. Opinion 479. Validation under the plenary powers of spe-
cic names for nine species of the Class Gastropoda occurring in
the New Zealand area as published by Martyn (T.) in 1784 in the
work entitled T he Universal Conchologist (Opinion supplemen-
tary to Opinion 456). — Opinions and Declarations Rendered by
the International Commission on Zoological Nomenclature 16:
365 - 416.
ICZN [International Commission on Zoological Nomen clature],
1970. Opinion 911. Six misidentied type-species in the super-
family Muricacea (Gastropoda). — Bulletin of Zoological
Nomenclatu re 27(1): 20-26.
ICZN [International Commission on Zoological Nomen clature],
1999. Opinion 1623. Risomurex Olsson & McGinty, 1958
(Mollusca, Gastropoda): Ricinula deformis Reeve, 1846desig-
nated as t he type species. — Bulleti n of Zoological Nomenclature
48 (1): 6 6 - 67.
ICZN [International Commission on Zoological Nomen clature],
1999. International Code of Zoological Nomen clature. Fourth
Edition: xxix + 306 pp., London (International Trust for
Zoological Nomenclature).
IREDALE, T., 1915. A commentary on Suter’s Manual of the New
Zealand Mollusca. — Transactions and Proceedings of the New
Zealand Institute 47: 417-497.
JOUSSEAUME, F.P., 1880. Division méthodique de la famille des
purpuridés. — Le Naturaliste 42: 335-336.
JUNG, P., 1969. Miocene and Pliocene mollusks from Trinidad. —
Bulletins of American Paleontology 55(247): 29 3-657.
KIENER, L.C., 1835-1837. Spécies général et iconographie des
coquilles vivantes. Vol. 8. Famille des Pur purifères. Pre mière
partie. Genres Cassidaire, (Cassidaria), Lamarck, pp. 1-10, pl.
1-2 [all (1835)]; Casque (Cassis), Bruguière, pp. 1-40, pl. 1-16 [all
(1835)]; Tonne (Dolium), Lamarck, pp. 1-16, pl. 1-5 [all (1835)];
Harpe (Harpa), Lamarck, pp. 1-12, pl. 1-6 [all (1835)]; Pourpre
(Purpura), Lamarck, pp. 1-151, pl. 1-46 [pp. 1-32 (1835), 33-80
(1836), 81-151 (1837); pl. 1-21 (1835), 22-39 (1836), 40-46 (1837)],
Paris, Rousseau & J.B. Baillière.
KOENEN, A. von, 1867. Das marine Mittel-Oligocän Nord-
Deutschlands und seine Mollusken-Fauna. — Palaeontographica
16(2): 53-127.
KÓKAY, J., 1966. Geologische und paläontologische Untersuchung
des Braunkohlengebietes von Herend-Márkó (Bakony-Gebirge,
Ungarn). — Geologic a Hungar ica, Series Palaeontologica 36: 1-147.
LAMARCK, J.B.P.A. de M. 1803. Suite des mémoires sur les fossils
des environs de Paris. — Annales du Muséum National d’His-
toire Naturelle de Paris 2: 217-227.

VITA MALACOLOGICA 15: 54
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
including the latest discoveries. 4 pp. + unpaginated captions to
61 pls., London (William Miller).
PEYROT, A., 1938. Les mollusques testacés univalves des depots
Helvétiens du Bassin Ligérien. Catalogue critique, descriptive et
illustré. — Actes de la Société Linnéenne de Bordeaux 89: 5-361.
PHILIPPI, R.A., 1836. Enumeratio molluscorum siciliae cum
viventium tum in tellure tertiaria fossilium quae in itinere suo
observavit auctor. xiv + 268 pp., Berolini.
PUSCH, G.G., 1836-1837. Polens Paläontologie oder Abbildung und
Beschreibung der vorzüglichsten und der noch unbeschriebenen
Petrefakten aus den Gebirgsformationen in Polen, Volhynien
und den Karpathen nebst einigen allgemeinen Beiträgen zur
Petrefaktenkunde und einem Versuch zur Vervollständigung der
Geschichte des europäischen Auer-Ochsen. 1-80, pls 1-10 (1836);
i-xiii, 81-218, pls 11-16 (1837), Stuttgart (E. Schweizerbart’s
Verlagshandlung).
RADWIN, G.E. & D’ATTILIO, A., 1971. Muricacean supraspecic
taxonomy based on the shell a nd the radula. — The E cho 4 : 55-67.
RADWI N, G.E. & D’ATTI LIO, A., 1975. A catalogue of Mur icacean
generic taxa. — Transactions of the San Diego Society of Natural
History 17: 279-292.
RAFINESQUE, C.S., 1815. Analyse de la nature ou tableau de
l’univers et des corps organisés. 223 pp., Palerme.
REEVE, L. 1846. Monograph of the genus Ricinula. Conchologia
Iconica 3 pls. 1-6 and unpaginated text. L. Reeve & Co., London.
ROBINSON, D.G., 1991. The systematics and paleoecology of the
prosobranch Gastropods of the Pleistocene Moín Formation of
Costa Rica. 748 pp., 28 pls., Unpublished PhD thesis, Dept. of
Geology, Tulane University.
RÖDING, P.F., 1798. Museum Boltenianum, sive catalogus cime-
liorum e tribus regnis naturae quae olim collgera Joa. Fried.
Bolten, M.D.p.d. per XL annos proto physicus Hamburgensis, 2.
Conchylia sive Testacea univalvia, bivalvia et multivalvia. 199
pp., Hamburgi (Johan. Christi. Trappii).
ROUAULT, A., 1850. Description des fossiles du terrain éocène
des environs de Pau. — Mémoires de la Société Géologique de
France série 2, 3(2): 457-502.
RUGGIERI, G., BRUNO, F. & CURTI, G., 1959. La malacofauna
pliocenica di Altavilla (Palermo), 1. — Atti dell’Academia di
Scienze Lettere e Arti di Palermo 18: 1-97.
RUGGIERI, G. & GRECO, A. 1965. Studi geologici e paleontolo-
gici su Capo Milazzo con particolare riguardo al Milazziano. —
Geologica Romana 4: 41-88.
SACCO, F., 1904. I Molluschi dei terreni terziari del Piemonte
e della Liguria, 30. Aggiunte e correzioni (con 1400 gure).
Considerazioni generali. Indice generale dell’opera. 203 + xxxvi
pp., 31 pls., Torino (C. Clausen).
SCACCHI, A., 1836. Catalogus conchyliorum regni neapolitani
quae usque reperit. 18 pp., Neapoli (Typis Filiatre-Sebetii).
SCHULTZ, O., 1998. Tertiärfossilien Österreichs, Wirbellose,
niedere Wirbeltiere und marine Säugetiere; schöne, interessante,
häuge und wichtige Makrofossilien aus den Beständen des
Naturhistorischen Museums Wien und Privatsammlungen; eine
Bilddokumentation. 159 pp., Wien (Golschneck-Verlag).
SOWERBY, G.B. II, 1834-41. Murex. [In] The conchological illus-
trations, or coloured illustrations of all the hitherto ungured
enumerazione delle principali specie dei terreni sopracretacei
dell’Italia. Vicenza (Tipograa Tremeschin): 28 pp.
MILLET [DE LA TURTAUDIÈRE], P.A., 1854. Paléontologie de
Maine-et-Loire. 144 pp., Angers (Cosnier et Lachèse).
MILLET DE LA TURTAUDIÈRE, P.A., 1865. Indicateur du
Maine-et-Loire ou indications par commune de ce que chacune
d’elles renferme. Tome 2. 616 pp., Angers (Cosnier et Lachèse).
MON TAGU, G. 1803. Testacea Britannica, or natural history of
British shells, marine, land and the fresh-water, including the
most minute: systematically arranged and embellished with g-
ures. London (Romsey): xxxvii + 606 pp.
MONTANARO, E., 1935. Studi monograci sulla malacolo-
gia Miocenica Modenese. Parte I. I molluschi Tortoniani di
Montegibbio. — Paleontographia Italica 35 (n.s. V): 1-84.
MONTEROSATO, T.A. di, 1869. Testacei nuovi dei mari di
Sicilia. 18 pp + 1 pl., Palermo.
MONTEROSATO, T.A. di, 1884. Nomenclatura generica e specica
di alcune conchiglie mediterranee. 152 pp., Paler mo, Virzi.
MONTEROSATO, T.A. di, 1890. Conchiglie della profondità del
mare di Palermo. — Naturalista Siciliano 9(6): 140-151; 9(7):
157-166; 9(8): 181-191.
MONTEROSATO, T.A. di, 1917. Molluschi viventi e quaternari
raccolti lu ngo le coste della Tripolitania dall’Ing. Camillo Crema.
— Bollettino della Società Zoologica Italiana (3)4: 1-28.
OLIVERIO, M. & GOFAS, S., 2006. Coralliophiline diversity at
mid-Atlantic seamounts (Neogastropoda, Muricidae, Corallio-
philinae). — Bulletin of Marine Science 79(1): 205-230.
OLIVER IO, M. & MARIOTT INI, P., 2001. A Mole cular Framework
for the Phylogeny of Coralliophila and Related Muricoids. —
Journal of Molluscan Studies 67: 215-224.
OR BIGN Y, A. d’ 1852. Prodrome de paléontologie stratigraphique
universelle des animaux mollusques et rayonnés, faisant suite au
cours élémentaire de paléontologie et de géologie stratigraphique,
3. Paris (Victor Masson): 1-196, index 1-189.
PALLA, P., 1967. Gasteropodi pliocenici della Bassa Val d’Elsa
(Toscana Occidentale). — Rivista Italiana di Paleontologia e
Stratigraa 73: 931-1020.
PAYRAUDEAU, B.C., 1826. Catalogue descriptif et méthodique
des Annelides et des Mollusques de l’île de Corse. 218 pp. + 8
pl., Paris.
PEREIRA DA COSTA, F.A., 1866-1867. Molluscos fosseis.
Gasteropodes dos depositos terciarios de Portugal. — Memória
Commissão Geologica de Portugal 4(1): 1-116 (1866); (2): 117-252
(1867).
PER RILLIAT, C.M. 1972. Monograa de los moluscos del Mioceno
Medio de Santa Rosa, Veracruz, Mexico. Parte I. (Gasterópodos:
Fissurellidae a Olividae). — Paleontologia Mexicana 32: 1-130.
PERRY, G., 1810-1811. Arcana; or the museum of natural history:
containing the most recent discovered objects. Embellished with
colored plates, and corresponding descriptions; with extracts
related to animals, and remarks of celebrated travellers; combin-
ing a general survey of nature. Pls. 1-48 (= 1810), pls. 49-84 (=
1811), 348 unnumbered text pages. London.
PERRY, G., 1811. Conchology, or the natural history of shells: con-
taining a new arrangement of the genera and species, illustrated
by colored engravings executed from the natural specimens, and

VITA MALACOLOGICA 15: 55
Ceulemans et al. – Lower Pliocene gastropods of Le Pigeon Blanc
— Tulane Studies in Geology and Paleontology 29: 69-118.
VOKES, E.H., 1964. Supraspecic groups in the subfamilies
Muricinae and Tritonaliinae (Gastropoda: Muricidae). —
Malacologia 2: 1-41.
VOKES, E.H., 1972. Pterynopsis, new genus of Trophoninae
(Gastropoda). — Bulletin de l’Institut Royal des Sciences
Naturelles de Belgique. Sciences de la Terre 48(9): 1-7, 2 pls.
VOKES, E.H., 1989. Neogene paleontology in the northern
Dominican Republic, 8. The family Muricidae (Mollusca:
Gastropoda). — Bulletins of American Paleontology 97(332):
5-94.
WIEGMANN, A.F.A. & RUTHE, J.F., 1832. Handbuch der Zoolo-
gie. 621 pp., Lüderitz, Berlin.
WINCKWORTH, M.A., 1945. The types of the Boltenian genera.
— Proceedings of the Malacological Society of London 26: 136-
148.
WOOD, S.V., 1872-1874. Supplement to the monograph of the Crag
Mollusca, with descriptions of shells from the upper Tertiaries
of the east of England, 3. Univalves and bivalves, with an intro-
ductory outline of the geology of the same district, and map.
— Monographs of the Paleontographical Society: i-xxxi + 1-99
(1872), 100-231 (1874).
WOOL ACOTT, L. 1957. Notes on Austral ian Shells. — Procee dings
of the Royal Zoological Society of New South Wales (1955-1956):
112-117
ZILCH, A ., 1934. Zur Fauna des Mit tel-Miocäns von Kost ej (Banat).
Typus-Bestimmung und Tafeln zu O. Boettger’s Bearbeitungen.
— Senkenbergiana 16(4-6): 193-302.
Recent shells. pls 58-67 (1834), pls 187-199 and catalogue, pp. 1-9
(1841), G.B. Sowerby, London.
STRAUSZ, L., 1966. Die Miozän-Mediterranen Gastropoden
Ungarns. 692 pp., Budapest (Akadémiai Kiadó).
SWAINSON, W., 1820-1833. Zoological illustrations or original
gures and descriptions of new, rare or interesting animals. Ser.
1: pls. 1-8 (1820); pls. 19-83 (1821), pls. 84-134 (1822), pls. 135-
182 (1823); ser. 2: pls. 1-30 (1829), pls. 31-45 (1830), pls. 46-85
(1831), pls. 86-96 (1832), pls. 97-136 (1833), London (Baldwin &
Cradock).
TOURNOUËR, R., 1875. Etude sur quelques espèces de Murex
fossiles du falun de Pont-Levoy en Touraine. — Journal de
Conchyliologie 23: 144 -167.
VAN DINGENEN, F., CEULEMANS, L. & LANDAU, B.M., 2014.
Euroscaphella nov. gen. (Gastropoda: Volutidae) in the Neogene
of Europe, with the description of a new species: Euroscaphella
namnetensis nov. sp. from the Mio-Pliocene transition of north-
western France. — Cainozoic Research 14: 101-111.
VAN DINGENEN, F., CEULEMANS, L., LANDAU, B.M. &
SILVA, C.M. da, 2015. The family Nassariidae (Gastropoda:
Buccinoidea) from the late Neogene of northwestern France. —
Cainozoic Research 15: 75-122.
VENZO, S. & PELOSIO, G. 1963. La malacofauna Tortoniana del
Colle di Vigoleno (Preappenino Piacentino). — Palaeontographia
Italica 58: 43-213.
VERMEIJ, G.J. & VOKES, E.H., 1997. Cenozoic Muricidae of the
western Atlantic region, 12. The subfamily Ocenebrinae (in part).
Accepted: 10 June 2016