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Abstract

In this paper we review the Neogastropoda (in part) of the Zanclean lower Pliocene assemblage of Le Pigeon Blanc, Loire-Atlantique department, France, which we consider the ‘type’ locality for Assemblage III of Van Dingenen et al. (2015). Twenty-six species are recorded, of which three are new: Euthria palumbina nov. sp., Bartschia (Agassitula) harasewychi nov. sp., Brocchinia pigeonblancensis nov. sp. Aplus aequicostatus (Bellardi, 1877) is considered a junior subjective synonym of Aplus scaber (Millet, 1865). Fusus (Aptyxis) rostratus ligerianus Peyrot, 1938 is considered a junior subjective synonym of Aptyxis omphale (Millet, 1864). The data presented here concurs with that discussed in previous parts of this monograph, suggesting 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.
Cainozoic Research, 17(1), pp. 23-61, June 2017 23
Introduction
In this paper we continue our studies on the Neogene
gastropod fossil assemblages of northwestern France
(see Ceulemans et al., 2014, 2016a, b; Van Dingenen et
al., 2014, 2015, 2016): Gastropods of the order Neogas-
tropoda (excluding Conoidea) 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). The Nassari-
idae of Assemblage III were revised in Van Dingenen et
al. (2015), although a summary and update are included
herein.
In his unpublished thesis, Brébion (1964) of the Centre
National de la Recherche Scientique, Paris recorded 20
species within the groups covered in this paper from Le
Pigeon Blanc and other Assemblage III localities, some
of which were described as new. However, as the thesis
was never published, the names do not comply with ar-
ticle 13 of the ICZN code (1999) and must be considered
nomina nuda.
The lower Pliocene gastropods of Le Pigeon Blanc (Loire-
Atlantique, Northwest France), 4*. Neogastropoda (in part)
Frank Van Dingenen1, Luc Ceulemans2 & Bernard M. Landau3, 4
1 Cambeenboslaan A 11, B-2960 Brecht, Belgium; email: fvd@telenet.be
2 Avenue Général Naessens de Loncin 1, B-1330 Rixensart, Belgium; email: luc.ceulem@skynet.be
3 Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, Netherlands; Instituto Dom Luiz da Universidade de
Lisboa, Campo Grande, 1749-016 Lisboa, Portugal; and International Health Centres, Av. Infante de Henrique 7,
Areias São João, P-8200 Albufeira, Portugal; email: bernielandau@sapo.pt
4 Corresponding author
Received 25 February 2017, revised version accepted 1 April 2017
In this paper we review the Neogastropoda (in part) of the Zanclean lower Pliocene assemblage of Le Pigeon Blanc, Loire-
Atlantique department, France, which we consider the ‘type’ locality for Assemblage III of Van Dingenen et al. (2015). Twenty-six
species are recorded, of which three are new: Euthria palumbina nov. sp., Bartschia (Agassitula) harasewychi nov. sp., Brocchinia
pigeonblancensis nov. sp. Aplus aequicostatus (Bellardi, 1877) is considered a junior subjective synonym of Aplus scaber (Millet,
1865). Fusus (Aptyxis) rostratus ligerianus Peyrot, 1938 is considered a junior subjective synonym of Aptyxis omphale (Millet,
1864). The data presented here concurs with that discussed in previous parts of this monograph, suggesting 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.
Key words: northwestern France, lower Pliocene, Neogastropoda, new taxa
Geological setting, Material and methods
(see Van Dingenen et al, 2015: 75-79, gures 1, 2).
Abbreviations:
MNHN.F Muséum national d’Histoire naturelle (collec-
tion 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).
Systematic palaeontology
Order Neogastropoda Wenz, 1938
Superfamily Buccinoidea Ranesque, 1815
Family Buccinidae Ranesque, 1815
Genus Aplus de Gregorio, 1885
Type species (by subsequent designation; Vokes, 1971)
Murex plicatus forma serzus de Gregorio, 1885, Neo-
gene, Italy.
* For nr 3 in this series see Vita Malacologica 15: 35-55.
24 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
1826 Anna Risso, p. 214. Type species (by monotypy):
Anna massena Risso, 1826, Pleistocene, France.
Anna massena Risso, 1826 is a nomen dubium
(see Brunetti & Della Bella, 2014).
1885 Aplus de Gregorio, p. 279.
NoteBrunetti & Della Bella (2014) argued that Anna
massena Risso, 1826, is a nomen dubium, possibly a
species of Raphitoma (family Raphitomidae), and so the
genus Anna could not be used. The next available ge-
neric taxon for this group of shells is Aplus de Gregorio,
1885. De Gregorio (1885) did not designate a type spe-
cies and included in his new genus Aplus, Murex plica-
tus ‘Brocchi’ (i.e., Murex plicatus Gmelin, 1791, sensu
Brocchi, 1814). Although Brunetti & Della Bella (2014)
designated Murex plicatus forma nilus de Gregorio,
1885 as the type species, the type species of Aplus had
already been designated by Vokes (1971, p. 83).
WoRMS (Bouchet, 2015a) synonymised Aplus with Pol-
lia (type species: Buccinum undosum Linnaeus, 1758).
Pollia undosa is an Indo-Pacic species with quite a dif-
ferent shell form and unlikely to be monophyletic with
the eastern Atlantic group here ascribed to Aplus. Simi-
larly, the tropical American Annaspecies also form a
distinct group, Ameranna Landau & Vermeij, 2012. Until
molecular data proves otherwise, we prefer to keep them
distinct.
Aplus scaber (Millet, 1865)
Plate 1, gs 1-3
1854 Fusus Scaber Millet, p. 162 (nomen nudum).
1865 Fusus scaber Millet, p. 591.
1873 Pollia aequicostata Bellardi, p. 182, pl. 12, g. 23.
1907 Fusus scaber Millet – Couffon, p. 185.
1964 Cantharus (Pollia) exsculpta Dujardin, 1837
Brébion (partim), p. 428 [non Aplus exsculptus
(Dujardin, 1837)].
1964 Cantharus (Pollia) aequicostata Bellardi, 1872
[sic] – Brébion, p. 429, pl. 10, gs 24, 25.
1981 Pollia aequicostata Bellardi, 1872 [sic] – Ferrero-
Mortara et al., p. 48, pl. 6, g. 8.
2014 Aplus aequicostatus (Bellardi, 1877) – Brunetti &
Della Bella, p. 18, gs 5A-F.
Material and dimensions Maximum height 14.6 mm.
NHMW 2015/0133/0369-71 (3: Pl. 1, gs 1-3 respective-
ly), NHMW 2015/0133/0372 (32), LC (40), FVD (35). Le
Pigeon Blanc, Le Landreau, Nantes area, Loire-Atlan-
tique department, NW France.
Discussion – Brunetti & Della Bella (2014) reviewed the
Italian Pliocene Aplus species. The size, shape, sculpture
and protoconch character of the Le Pigeon Blanc speci-
mens are most like that of Aplus aequicostatus (Bellardi,
1873). The protoconch of the French material consists of
about 2.5 convex whorls, with a medium-sized nucleus.
The protoconch shape is a little variable, inated and so-
mewhat uncoiled in some specimens. There is also con-
siderable variability in the shell prole seen in the series
illustrated here, with broader and more slender speci-
mens present. For comparison with congeners see Bru-
netti & Della Bella (2014).
Brébion (1964, p. 429) noted that Fusus scaber Mil-
let was a synonym of A. inaequicostatus. He probably
did not use Millet’s name, correctly considering Fusus
scaber Millet, 1854 to be a nomen nudum. However,
the name was later validated by the following descrip-
tion: ‘Fusus scaber, Millet. Coq. petite, un peu allongée,
ventrue inférieurement; composée de six à sept tours
de spire garnis de côtes étroites, saillantes qui se ren-
dent ainsi jusqu’à la suture; ces côtes sont couvertes
de stries serrées, rudes, si ce n’est la partie supérieure
de chaque tour qui n’en présente aucune. Ouverture
presque ovale, à canal très-court, et quelques légers
plis se font remarquer sur la partie intérieure du bord
droit, qui présente en dehors un fort bourrelet. Lon-
gueur: 10-11 millimètres; diamètre: 4-5 millimètres.
Sc, Th. (1865, p. 591)’. We cannot apply Article 23.9.1.2
(ICZN 1999) to consider Millet’s name a nomen obli-
tum as Couffon (1907, p. 185) used the name as a valid
taxon. Therefore, A. inaequicostatus (Bellardi, 1873) is
a junior subjective synonym of Aplus scaber (Millet,
1865).
Millet (1865, p. 591) recorded this species from Assem-
blage I localities of Sceaux d’Anjou and Thorigné, Bré-
bion (1964, p. 428) added St-Michel. Brébion also recor-
ded Cantharus (Pollia) exsculpta Dujardin, 1837 from Le
Pigeon Blanc. We have not found any specimens of Aplus
exsculptus at Le Pigeon Blanc, and the record probably
refers to A. inaequicostatus.
Distribution – Upper Miocene (Tortonian): Atlantic, NW
France (Millet, 1865; Brébion, 1964). Lower Pliocene:
Atlantic, NW France (Brébion, 1964). Upper Pliocene:
Italy (Brunetti & Della Bella, 2014).
Genus Euthria Gray, 1850
Type species (by subsequent designation; Petit, 2012) –
Murex corneus Linnaeus, 1758, present-day, Mediterra-
nean.
1850 Euthria Gray, p. 67
Note Hadorn & Fraussen (1999) synonymised Sipho-
nofusus Drivas & Jay 1990 with Euthria Gray, 1850.
Siphonofusus species are slender, with a longer sipho-
nal canal than usual for Euthria and live in deep water.
Moreover, they are from Japan and unlikely to be mono-
phyletic with Euthria, as envisaged here, which seems to
be a European group. We are unconvinced that any of the
Indo-Pacic species described (i.e. Fraussen & Hadorn,
2003) are monophyletic with the eastern Atlantic species.
We therefore provisionally exclude Siphonofusus from
the generic synonymy until we have more molecular data
on the group.
Cainozoic Research, 17(1), pp. 23-61, June 2017 25
Euthria palumbina nov. sp.
Plate 1, gs 4-6
1964 Buccinulum (Euthria) lecointrei Brébion, p. 420,
pl. 10, gs 15-16 (nomen nudum).
Type material Holotype NHMW 2015/0133/0363 (Pl.
1, g. 4), height 38.6 mm; paratype 1 NHMW 2015/
0133/0364 (Pl. 1, g. 5), height 25.8 mm; paratype 2
NHMW 2015/0133/0366, height 24.0 mm (Pl. 1, g. 6);
paratype 3 MNHN.F.A57934, height 24.2 mm; para-
type 4 MNHN.F.A57935, height 21.7 mm; paratype 5
MNHN.F.A57935, height 22.6 mm.
Other material Maximum height 39.2 mm. NHMW
2015/0133/0365 (25), LC (50+), FVD (50+).
Etymology From Latin palumbes, palumbis, noun,
wood-pigeon, ringdove; a reference to the type locality
of Le Pigeon Blanc (the white pigeon). Euthria gender
feminine.
Locus typicus Le Landreau, Le Pigeon Blanc, Loire-
Atlantique department, NW France.
Stratum typicum – Zanclean, lower Pliocene.
DiagnosisAn Euthria species of small size, relatively
slender fusiform in shape, with a protoconch consisting
of two whorls, teleoconch sculptured by 9-10 rounded
ribs that weaken on penultimate whorl and become obso-
lete, or almost so, on last whorl, very weak spiral sculp-
ture of alternate strength,an ovate aperture thickened by
labial varix, denticulate within,the parietal tooth strong-
est, and a moderately long siphonal canal, 19% of total
height.
DescriptionShell small to small for genus, solid, rela-
tively slender fusiform. Protoconch consisting of two
convex whorls, with medium-sized nucleus (dp = 1.0
mm; hp = 1.1 mm; dn = 270 µm). Protoconch boundary
sharp. Teleoconch of six convex whorls, with periphery
just below mid-whorl. Sculpture of 9-10 elevated, proso-
cline, rounded ribs, broadening towards abapical suture,
roughly equal in width to their interspaces, crossed
by very numerous ne cords of primary and second-
ary strength. Last whorl weakly inated, with narrow,
slightly concave subsutural ramp, shoulder high, round-
ed, whorl constricted at base, axial sculpture strongest
at shoulder, weakening to subobsolete over base, spiral
sculpture strengthening over base. Aperture ovate, anal
canal marked by adapical notch, siphonal canal of mod-
erate length, 19% of total height, open, narrow, slightly
posteriorly recurved. Outer lip thickened by labial varix,
sinuous in prole, edge bevelled, bearing a row of small
rounded denticles along entire lip height, placed at inner
edge of bevelled portion. Columella evenly excavated,
bearing row of irregular tubercles along entire length,
with stronger parietal tubercles adapically; tubercles sub-
obsolete mid-columella in some specimens. Columellar
callus thin, adherent, sharply delimited, poorly expand-
ed. Siphonal fasciole rounded, narrow.
DiscussionEuthria palumbina nov. sp. is fairly typical
for the genus as understood here (see note under generic
assignment above), although rather small shelled com-
pared to some of its congeners. The species is common in
the Le Pigeon Blanc assemblage and there is little vari-
ability. Euthria adunca (Bronn, 1831) from the Pliocene
Mediterranean is larger shelled, the shoulder is slightly
more angular, the axial ribs are wider, the spiral cords
stronger and the siphonal canal is longer. Euthria sub-
marginata (d’Orbigny, 1852) from the middle Miocene of
the Loire Basin is even smaller, squatter, thinner shelled,
with more numerous axial ribs (12) and stronger and less
numerous spiral cords, the primary cords forming elon-
gated tubercles where they cross the ribs. Euthria var-
icigera Peyrot, 1928 from the middle Miocene Aquitaine
Basin of France has less convex spire whorls, more nu-
merous axial ribs (15), stronger primary spiral sculpture
and a more elongate last whorl. Euthria pseudomargi-
nata Peyrot, 1928 from the lower Miocene Burdigalian
Aquitaine Basin is smaller shelled than E. palumbina and
like the previous species also has more numerous axial
ribs (15) and stronger spiral sculpture than E. palum-
bina. Euthria verrucifera Bellardi, 1873 from the upper
Mio cene of Italy is similar in shape to E. palumbina, but
differs in having broader axial ribs and stronger spiral
sculpture.
Brébion (1964, p. 421) recorded this species from the
middle Miocene Loire Basin (Mirebeau), Assemblage
I localities (Renauleau, Sceaux-d’Anjou, Thorigné, St-
Michel), Assemblage II (Apigné), Assemblage III (Le
Pigeon Blanc, Le Girondor, Palluau) and Assemblage IV
(Gourbesville).
Distribution Middle Miocene: Loire Basin (Brébion,
1964). Upper Miocene (Tortonian and Messinian): Atlan-
tic, NW France (Brébion, 1964). Lower Pliocene: Atlan-
tic, NW France (Brébion, 1964). Upper Pliocene-Pleis-
tocene: Atlantic, NW France (Brébion, 1964).
Euthria turonensis Peyrot, 1938
Plate 1, g. 7
*1938 Euthria adunca var. turonensis Peyrot, p. 230, pl.
4, gs 11, 16, 19.
1952a Euthria adunca turonensis Peyrot, 1938 – Glibert,
p. 324, pl. 9, g. 4.
?1964 Buccinulum (Euthria) regulare Brébion, p. 423,
pl. 10, g. 19 (nomen nudum).
Material and dimensions Height 19.3 mm. NHMW
2015/0133/0368 (1: Pl. 1, g. 7). Le Pigeon Blanc, Le Lan-
dreau, Nantes area, Loire-Atlantique department, NW
France.
Revised descriptionShell small, solid, slender fusiform.
Protoconch not preserved. Teleoconch of ve weakly but
26 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
regulary convex whorls, shoulder and subsutural ramp
poorly delimited by weak angulation, with periphery
just below mid-whorl. Sculpture of elevated, orthocline,
rounded ribs, extending between sutures, broadening
slightly towards abapical suture, half the width of their
interspaces; 9 on rst whorl, 17 on last whorl, crossed
by numerous spiral cords of primary to tertiary strength.
Last whorl relatively elongate, regularly rounded, con-
stricted at base, axial sculpture strongest mid-whorl,
weakening over base. Aperture ovate, anal canal marked
by adapical groove, siphonal canal moderate length, open,
narrow, slightly posteriorly recurved. Outer lip thickened
by labial varix, edge bevelled, bearing a row of elongated
denticles along entire lip length, placed at inner edge
of bevelled portion, extending as lirae within aperture.
Columella evenly excavated, bearing two small abapi-
cal tubercles, one mid-aperture, one parietal. Columel-
lar callus adherent, sharply delimited, narrow. Siphonal
fasciole rounded, narrow.
Discussion Euthria turonensis Peyrot, 1938, which
seems to be excedingly rare at Le Pigeon Blanc, differs
from Euthria palumbina nov. sp. in being more slender-
shelled, the whorls are more weakly, but regularly con-
vex rather than subangular, it lacks a well-dened sub-
sutural ramp, the axial ribs are more numerous and the
spiral cords stronger and less numerous. Moreover, the
denticles placed within the outer lip are more numerous
and extend into the aperture as lirae rather than being
restricted to the inner edge of the bevelled outer lip.
We consider it conspecic with the specimens illustrated
by Glibert (1952a, pl. 9, g. 4) as Euthria adunca turonen-
sis Peyrot, 1938. The series illustrated by Glibert shows
some variability in the strength of the shoulder cord; his
g 4a is rather broader-shelled, with a stronger shoul-
der cord, like the specimen illustrated by Peyrot (1938,
pl. 4, gs 11, 16, 19), whereas Glibert’s g. 4c is more
elongated, with a weak shoulder, similar to the specimen
from Le Pigeon Blanc. As pointed out by Glibert (1952a,
p. 325) the north western French forms differ from the
Pliocene Mediterranean E. adunca (Bronn, 1831) in be-
ing much smaller sized and in having more numerous
axial ribs (15-17 vs. 10-11).
It also ts well with the description of a shell illustrated
by Brébion (1964) as Buccinulum (Euthria) regulare (no-
men nudum) from the older upper Miocene Assemblage
I locality of St-Michel: Long. 14mm, larg. 7mm. Forme
étroite à spire formée de 4 à 5 tours ornés d’une douzaine
de côtes sétendant sur toute leur hauteur et de 7 à 8 cor-
dons spiraux plus étroit égaux à leurs intervalles, plus
ns et serrées vers l’arrière des tours, l’ensemble étant
très régulier. Base entièrement couverte de cordons;
labre peu épaissi, muni d’une dizaine de denticules; bord
columellaire avec 2 denticules en avant et une faible dent
pariétale; canal court (1964, p. 423)’. Brébion drew at-
tention to the narrow shell shape, regular sculpture, with
the ribs extending between the sutures and the two tuber-
cles on the abapical portion of the columella. The only
differences between the Le Pigeon Blanc shell and that
from St-Michel is that the former has a greater number
of axial ribs on the last whorl (17, vs. 12) and more nu-
merous denticles within the outer lip (16 vs. 12). Brébion
(1964) wrote that this shell differed from E. turonensis
par la taille plus faible, la petitesse du canal, l’absence
de bourrelet’. However, as discussed above, these differ-
ences are not consistent.
Unfortunately, the single shell available from Le Pigeon
Blanc gives us no information on the intraspecic vari-
ability.
Distribution Middle Miocene (Langhian): Atlantic,
Loire Basin, France (Peyrot, 1938; Glibert, 1952a). ?Up-
per Miocene: Atlantic (Tortonian): north western France
(Brébion, 1964). Lower Pliocene: Atlantic, NW France
(this paper).
Family Colubrariidae Dall, 1904
Genus Bartschia Rehder, 1943
Type species (by original designation) Bartschia sig-
nicans Rehder, 1943, present-day, Florida, USA.
1943 Bartschia Rehder, p. 199.
Subgenus Agassitula Olsson & Bayer, 1972
Type species (by original designation) – Metula agassizi
Clench & Aguayo, 1941, present-day, Cuba.
1972 Agassitula Olsson & Bayer, p. 917.
Bartschia (Agassitula) harasewychi nov. sp.
Plate 1, g. 8; Plate 2, g. 11
1964 Buccinulum (Euthria) aenigmaticum Brébion, p.
425, pl. 10, g. 22 (nomen nudum).
Type material Holotype: NHMW 2015/0133/0367 (Pl.
1, g. 8), height 27.9 mm, width 10.3 mm; paratype 1
NHMW 2015/0133/0426 (Pl. 2, g.11), height 20.4 mm,
width 7.4 mm.
Other material – LC (3 fragments).
Etymology – Named after M. G. (Jerry) Harasewych, Re-
search Zoologist at the Smithsonian Institution, Washing-
ton, DC, USA, in recognition of his many contributions
to American malacology and particularly his work on the
family Colubrariidae. Bartschia gender feminine.
Locus typicus Le Landreau, Le Pigeon Blanc, Loire-
Atlantique department, NW France.
Stratum typicum – Zanclean, lower Pliocene.
Diagnosis – A Bartschia (Agassitula) species of medium
size, with a tall spire, covered by nely reticulated sculp-
ture, in which the axial component is slightly dominant,
Cainozoic Research, 17(1), pp. 23-61, June 2017 27
type species, Bartschia (A.) agassizi Clench & Aguayo,
1941 and Bartschia (A.) peartae Harasewych, 2014 from
the present-day Caribbean are both immediately sepa-
rated from the French species by their ner sculpture.
Bartschia (A.) guppyi Olsson & Bayer, 1972, also from
the present-day Caribbean and B. (A.) limonensis (Ols-
son, 1922) from the lower Pleistocene Moin Formation of
Costa Rica both have closer-set sculpture, with tubercles
developed at the sculptural intersections, giving the sur-
face a beaded rather than cancellated appearance.
Brébion (1964, p. 426) recorded this species from the
upper Miocene Tortonian Assemblage I localities (Tho-
rigné, Contigné). Unfortunately, we have been unable
to locate any of Brébion’s material reported as being
in ‘musée d’Angers’. In this paper we extend the strati-
graphic range for the species to the lower Pliocene Zan-
clean Assemblage III.
Distribution – Upper Miocene (Tortonian): Atlantic, NW
France (Brébion, 1964). Lower Pliocene: Atlantic, NW
France (Brébion, 1964).
Family Columbellidae Swainson, 1840
Genus Mitrella Risso, 1826
Type species (by subsequent designation; Cox, 1927)
M. aminea Risso, 1826 (= Murex scriptus Linnaeus,
1758), present-day, Mediterranean.
1826 Mitrella Risso, p. 247.
Note – The genus Mitrella Risso, 1826 as used here is in-
terpreted rather widely and is unlikely to be monophylet-
ic. We are unaware of any molecular studies on mitrellid
gastropods, and await these, hoping that they will shed
some light on relationships. Until then we do not think it
useful to give a generic synonymy.
Mitrella erythrostoma (Bellardi, 1848)
Plate 1, g. 9
*1848 Columbella erythrostoma Bellardi, Bonelli m.s.,
p. 9, pl. 1, gs 4-5.
1890 Columbella (Mitrella) erythrostoma Bon. – Sacco,
p. 40, pl. 2, g. 40.
1904 Mitrella erythrostoma (Bon.) var. compressula
Sacco, p. 93, pl. 19, gs 51-52.
1938 Columbella (Alia) ligeriana Peyrot, p. 204, pl. 4,
gs 45, 46.
1952a Columbella (Alia) erythrostoma Bonelli, 1825
Glibert, p. 320, pl. 8, g. 5.
1975 Mitrella erythrostoma var. compressula Sacco
Fekih, p. 126, pl. 37, g. 16.
1981 Columbella (Mitrella) erythrostoma Bellardi,
1848, Bonelli m.s. – Ferrero-Mortara et al., p. 183,
pl. 57, g. 12.
2011 Mitrella erythrostoma (Bellardi, 1848) – Landau
et al., p. 28, pl. 13, g. 19.
a moderately inated and broad last whorl, 56% of total
height, modestly thickened outer lip, nely denticulate
within and a short siphonal canal, 7% of total height.
DescriptionShell of medium size and thickness, fusi-
form, with elevated, conical spire. Protoconch not pre-
served. Teleoconch of 6.5 evenly convex whorls, lacking
shoulder, separated by shallow adpressed suture. Sharp
protoconch-teleoconch boundary preserved, marked by
the onset of ve spiral cords and narrow, close-set axial
ribs, 19 on rst whorl, crossed by narrow, close-set equal
spiral cords, increasing in number abapically, ten on pen-
ultimate whorl, forming dense, horizontally elongated
cancellated pattern, with axial sculpture predominant.
Single spiral thread intercalated in some interspaces on
penultimate whorl. A sudden change in sculpture on
second half of the penultimate whorl is probably due
to damage during life. Last whorl 56% of total height,
moderately inated (width/ height 0.66), unsculptured,
with irregular surface, crossed by coarse growth lines.
Aperture ovate, anal canal relatively shallow and broad,
siphonal canal short, open, dorsally reected. Outer lip
thickened by modest varix, with bevelled edge, bearing
a row of 11 short, subequal denticles along inner edge.
Siphonal canal of moderately short, 7% of total height,
open, weakly recurved. Columella smooth. Columellar
and parietal callus moderately thickened, adherent, poorly
delimited. Siphonal fasciole poorly developed, rounded.
Discussion Although many of the colubrariid genera/
subgenera related to Metula H. Adams & A. Adams, 1853
have now been synonymised with it (WoRMS; Bouchet,
2015b), we follow Harasewych (2014) in recognising
Bartschia Rehder, 1943 as a separate genus. Bartschia
species differ from Metula in having proportionally broad-
er shells, with more evenly rounded whorls, a shorter, wid-
er aperture with more pronounced denticles within the
outer lip and slightly coarser sculpture with spiral cords
predominant. The subgenus Agassitula Olsson & Bayer,
1972, differs from Bartschia in having more elongated
shells, in which the spire is half or more of the total shell
height and a longer siphonal canal (Harase wych, 2014,
p. 92). The genera/subgenera Agassitula and Bartschia
share a similar stratigraphical and geographical distribu-
tion and are not recognised by all (i.e. WoRMS; Bouchet,
2015c). The specimens from Le Pigeon Blanc t well
within the generic description of Bartschia (Agassitula),
except possibly in that the axial sculpture is slight domi-
nant. The holotype (Pl. 1, g. 8) from France seems to
have suffered some injury during life, possibly a crab
attack, from which it survived, but lost its sculpture on
whorls formed subsequent to the injury. The paratype (Pl.
2, g. 11) did not suffer injury and shows no change in
sculpture on the last whorl. However, it is not chosen as
the holotype as it is not fully grown and lacks the adult
apertural dentition.
Whilst the genus Metula is recognised in the European
Neogene, as far as we are aware, this is the rst record for
the subgenus Bartschia (Agassitula), or indeed Bartschia,
in Europe. Today the genus is tropical American. The
28 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
Material and dimensions Maximum height 22.7 mm.
NHMW 2015/0133/0373 (1: Pl. 1, g. 9), LC (5), FVD (2).
Le Pigeon Blanc, Le Landreau, Nantes area, Loire-Atlan-
tique department, NW France.
Discussion Mitrella erythrostoma (Bellardi, 1848) is
a relatively large-shelled mitrellid, characterised by its
weakly convex spire whorls, its moderately inated last
whorl, elongated aperture, with the outer lip pinched in-
wards just above mid-height, bearing teeth along its en-
tire inner edge, more strongly developed on the pinched
section, its poorly callused inner lip and short siphonal
canal. Some specimens have a shorter spire, which gave
rise to Sacco’s (1904) variety compressula. The speci-
men from Le Pigeon Blanc has a slightly taller spire than
usual for the species, but the last whorl characters are
identical. We would agree with Glibert (1952a) in syno-
nymising the French middle Miocene specimens from
the Loire Basin described by Peyrot (1938) as Columbella
(Alia) ligeriana.
The only species somewhat similar is the Mediterranean
Pliocene Mitrella turgidula (Brocchi, 1814), which also
has a pinched outer lip, but it is easily separated from
M. erythrostoma by its broader, regularly conical, more
pointed spire, shallower suture and broad last whorl,
which is more strongly constricted at the base.
Distribution Middle Miocene: Atlantic, Loire Basin,
France (Peyrot, 1938; Glibert, 1952a). Upper Miocene:
Proto-Mediterranean, Italy (Glibert, 1952a). Lower Plio-
cene: Atlantic, NW France (this paper); Guadalquivir Ba-
sin, Spain (Landau et al., 2011); central Mediterranean,
Tunisia (Fekih, 1975). Upper Pliocene: western Medi-
terranean, Estepona Basin, Spain (BL unpublished data);
central Mediterranean, Italy (Bellardi, 1848; Sacco, 1904).
Mitrella bruggeni van Aartsen, Menkhorst & Gitten-
berger, 1984
Plate 1, g. 10
1981 Pyrene broderipi (G.B. Sowerby I, 1844) – Sabelli
& Spada, unnumbered page 2, gs 7a-c. (non G.B.
Sowerby I, 1844).
1984 Mitrella maldonadoi Luque, p. 13 (nomen nu-
dum).
*1984 Mitrella bruggeni van Aartsen, Menkhorst & Git-
tenberger, p. 77, g. 176a.
1987 Mitrella maldonadoi Luque, 1984 Luque, p.
231, pl. 1, gs 3-5.
2003 Mitrella bruggeni van Aartsen, Menkhorst & Git-
tenberger Giannuzzi-Savelli et al., p. 254, gs
613-622.
Material and dimensions Maximum height 11.9 mm.
NHMW 2015/0133/0378 (1: Pl. 1, g. 10), LC (3). Le Pi-
geon Blanc, Le Landreau, Nantes area, Loire-Atlantique
department, NW France.
Discussion A few specimens from Le Pigeon Blanc
are ascribed to the present-day species Mitrella bruggeni
van Aartsen, Menkhorst & Gittenberger, 1984. It is one
of the few shells from Le Pigeon Blanc with colour pat-
tern preserved, showing spiral rows of spots. This type
of pattern is present in several present-day European
Mitrella species, of which M. bruggeni is one. Mitrella
gervillii (Payraudeau, 1826) is another, but differs in be-
ing more slender, with slightly more convex spire whorls
and usually with more numerous teeth within the outer
lip. Mitrella broderipi (G.B. Sowerby II, 1844) is another
western Mediterranean species with spotted colour pat-
tern, but differs in being smaller, squatter, with slightly
more convex whorls and a less constricted base. Lastly,
M. svelta Kobelt, 1889 (= Columbella lanceolata Locard,
1886; non G.B. Sowerby I, 1832a) is the largest and most
slender of the group, with a taller, more straight-sided
spire than any of the preceding species. As far as we are
aware, this is the rst fossil record for the species. A more
northern distribution during the lower Pliocene than
found today is consistent with many other thermophilic
elements in the Assemblage III fauna.
Distribution – Lower Pliocene: Atlantic, NW France (this
paper). Present-day, western Mediterranean, Canaries,
Madeira (van Aartsen, Menkhorst & Gittenberger, 1984).
Mitrella vialensis (Sacco, 1890)
Plate 2, gs 1, 2
*1890 Columbella (Clinurella) vialensis Sacco, p. 46, pl.
2, g. 53.
1964 Mitrella (Atilia) inedita Bellardi, 1890 Brébion
(partim: Pigeon Blanc records), p. 127, pl. 9, g.
28 (only) [non Mitrella inedita (Sacco, 1890)].
1981 Columbella (Clinurella) vialensis Sacco in Bel-
lardi, 1890 – Ferrero-Mortara et al., p. 183, pl. 57,
g. 12.
2002 Mitrella vialensis (Sacco in Bellardi, 1890)
Chirli, p. 13, pl. 7, gs 3-8.
2010 Mitrella minima (Sacco, 1890) – Sosso & Dell’-
Angelo, p. 59 unnumbered g. top left (non Sacco,
1890).
Material and dimensions Maximum height 11.8 mm.
NHMW 2015/0133/0374 (1: Pl. 2, g. 1), NHMW 2015/
0133/0375 (1: 9.0 mm), NHMW 2015/0133/0376 (Pl. 2, g.
1); NHMW 2015/0133/0377 (50+), LC (50), FVD (50+).
Le Pigeon Blanc, Le Landreau, Nantes area, Loire-Atlan-
tique department, NW France.
Discussion Mitrella vialensis (Sacco, 1890) is charac-
terised by its tall, slender, non-scalate, coeloconoid spire,
weakly rounded spire whorls, relatively low last whorl,
which is strongly constricted at the base and the presence
of denticles within the outer lip. No protoconch descrip-
tion of Italian specimens is available, but the specimens
from Le Pigeon Blanc have a tall multispiral protoconch
of about 3-3.5 smooth convex whorls (Pl. 2, g. 2). Ital-
ian specimens at hand (NHMW coll.) have a multispiral
Cainozoic Research, 17(1), pp. 23-61, June 2017 29
protoconch similar to that of the French specimens. The
only possible difference between the two populations is
that in the Italian specimens the spiral cords on the base
are slightly stronger and cover more of the lower half of
the last whorl than they do in the shells from Le Pigeon
Blanc. However, we consider this difference of minor sig-
nicance.
Mitrella vialensis belongs to a group of closely simi-
lar species including Mitrella borsoni (Bellardi, 1848),
which is widespread in the middle and upper Miocene
Atlantic and Mediterranean Sea (Landau et al., 2013).
This species differs from M. vialensis in having a more
scalate spire and a less constricted base. Mitrella mini-
ma (Sacco, 1890), also from the Pliocene of Italy, is very
similar to M. vialensis but smaller, more constricted at
the base and has a smooth inner lip without denticles.
The strength of the denticles is rather variable in M. via-
lensis and M. minima may be a subadult specimen as
the syntype illustrated by Ferrero Mortara et al., (1981,
pl. 57, g. 10) lacks the thickened labial varix typical of
the genus. Either way, the specimen illustrated by Sosso
& Dell’Angelo (2010, p. 59) as M. minima has strong la-
bial dentition and should be ascribed to ++M. vialensis.
Mitrella scalaris (Sacco, 1890) from the middle Mio-
cene of Italy, is of similar size and shape, but differs in
having an outer lip that is more ared and somewhat
alate adapically.
Brébion (1964, p. 401) recorded, but did not gure, M.
borsoni from numerous Assemblage I, III and IV locali-
ties, but not Le Pigeon Blanc. Until we review the As-
semblage I localities we cannot be certain to which of
this closely similar group of species the reference refers
to. He did gure one shell from Le Pigeon Blanc under
the name Mitrella (Atilia) inedita Bellardi, 1890 (1964,
pl. 9, g. 28), which we have included in the synonymy
of M. vialensis. Mitrella inedita (Sacco, 1890) from the
upper Miocene of Italy is also similar, but differs in hav-
ing a regularly conical rather than coeloconoid spire and
the base is less constricted resulting in a broader siphonal
canal, which is also slightly shorter than in M. vialensis
and less posteriorly reected.
Distribution – Lower Pliocene: Atlantic, NW France (this
paper); central Mediterranean, Italy (Sacco, 1890; Chirli,
2002). Upper Pliocene: central Mediterranean, Italy (Sos-
so & Dell’Angelo, 2010).
Genus Anachis H. Adams & A. Adams, 1853
Type species (by subsequent designation, Tate, 1868)
Columbella scalarina G.B. Sowerby I, 1832, present-day,
Panamic Pacic.
1853 Anachis H. Adams & A. Adams, p. 184.
Note – This European Neogene group of tall-spired, slen-
der columbellids with cancellate sculpture have been
placed by all authors in the genus Anachis H. Adams &
A. Adams, 1853. The Panamic Pacic type species, Co-
lumbella scalarina G.B. Sowerby I, 1832 is quite different
with predominantly axial sculpture and a solid, inated
last whorl that is large in relation to the spire. We doubt
that the two groups are monophyletic. Unfortunately the
European group has no modern representatives, so their
relationship cannot be tested using molecular phyloge-
netics. The shell shape of this group is far closer to the
tropical American Atlantic genus Suturoglypta Radwin,
1968, but this also has strongly predominant axial sculp-
ture. We hesitate to erect a genus for this group as some
European Neogene species such as Anachis hoernesi
(Mayer, 1869) and A. haueri (Hoernes & Auinger, 1880)
seem to have intermediate features.
Anachis milleti nov. sp.
Text-g. 1/1-4, Plate 2, g. 3
1964 Anachis fannyae (Dollfus mss.) – Brébion, p. 410,
pl. 10, gs 5, 6 (nomen nudum).
Type material Holotype: MNHN.F.A57685 (Text-g.
1/1), height 6.8 mm, width 2.6 mm; paratype 1 NHMW
2016/0103/0063 (Text-g. 1/2), height 5.8 mm, width 2.6
mm; paratype 2 MNHN.F.A57686, height 6.8 mm, width
2.7 mm; paratype 3 NHMW 2016/0103/0064, height 7.2
mm, width 2.9 mm; St-Clément-de-la-Place. Paratype 4
MNHN.F.A57656 (pl. 10, g. 5 of Brébion, 1964; herein
Text g. 1/3), height 5.6 mm, Sceaux-d’Anjou (Maine-et-
Loire, France).
Other material St-Clément-de-la-Place: maximum height
7.2 mm. NHMW 2016/0103/0065 (40), LC (50+), FVD
(50+). Sceaux d’Anjou: maximum height 7.0 mm. NHMW
2016/0103/0066 (50+), LC (50+), FVD (50+). Le Pigeon
Blanc: maximum height 11.3 mm. NHMW 2015/0133/0379
(1: Pl. 2, g. 3), NHMW 2015/0133/0380 (5). Palluau
(Vendée, France): MNHN.F.A57657 (pl. 10, g. 6 Brébion,
1964; herein Text-g. 1/4), MNHN.F.A57658 (4)
Etymology Named after Pierre-Amié Millet de la Tur-
taudière, (1783-1873), Secrétaire Général de la Société
d’Agriculture d’ Angers, in recognition of his pioneering
work on the palaeontology of Maine-et-Loire. Anachis
gender feminine.
Locus typicus Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – An Anachis species of small size, with a slen-
der fusiform shell shape, a tall spire and weakly inated
last whorl, bearing moderately dense and ne reticulated
sculpture with axial and spiral elements of roughly equal
in strength, forming small tubercles at the intersections,
and a tall multispiral protoconch.
Description Shell small, slender fusiform. Protoconch
multispiral, two smooth convex whorls preserved. Teleo-
30 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
conch of 5.5 straight-sided to weakly convex whorls,
with periphery at or just above abapical suture. Suture
shallow. Sculpture on rst teleoconch whorl of two nar-
row, raised spiral cords placed adjacent to sutures. On
second whorl a third cord develops below adapical cord,
with a further cord added below adapical cord on subse-
quent whorls. Axial sculpture of prosocline ribs, 15-16
on pen ultimate whorl, equal in strength to spiral cords,
forming moderately dense reticulated surface sculpture.
Small tubercles develop at sculptural intersections. Last
Text-gure 1. Assemblage I Anachis species: 1-4. Anachis milleti nov. sp.; 1: holotype, MNHN.F.A57685, height 6.8 mm; 2:
paratype 1, NHMW 2016/0103/0063, height 5.8 mm; 3: paratype 4, MNHN.F.A57656, height 5.6 mm (photo Jocelyn Fal-
connet; MNHN); 4: MNHN.F.A57657 (photo Jocelyn Falconnet; MNHN); 5-6. Anachis hordacea (Millet, 1865). 5: NHMW
2016/0103/0072, height 6.9 mm; 6: NHMW 2016/0103/0073, height 6.0 mm; 7-8. Anachis collyrata (Millet, 1865); 7. NHMW
2016/0103/0067, height 9.0 mm; 8. NHMW 2016/0103/0068, height 11.7 mm.
Figures 1-2, 5-8 from Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France; Figure 3 from Sceaux-
d’Anjou, Maine-et-Loire, France; Figure 4 from Palluau, Vendée, France. All Tortonian, upper Miocene.
Cainozoic Research, 17(1), pp. 23-61, June 2017 31
Family Fasciolariidae Gray, 1853
Subfamily Fusininae Wrigley, 1927
Genus Aptyxis Troschel, 1868
Type species (by monotypy) Murex syracusanus Lin-
naeus, 1758, present-day, Mediterranean.
1868 Aptyxis Troschel, p. 61, 64.
1882 Aptysis Bucquoy et al., p. 16, 35. Incorrect subse-
quent spelling.
Aptyxis omphale (Millet, 1864)
Plate 2, gs 4, 5
1854 Fusus Rostratus Sismonda [sic] – Millet, p. 162
(non Fusus rostratus Olivi, 1792; non Solander,
1766 = Fusinus sanctaluciae (von Salis-Marsch-
lins, 1793).
1854 Fusus Omphale Millet, p. 162 (nomen nudum).
1854 Fusus Vicinus Millet, p. 162 (nomen nudum).
1854 Fusus Ventricosus Millet, p. 162 (nomen nudum).
*1864 Fusus omphale Millet, p. 674.
1865 Fusus vicinus Millet, p. 590.
1865 Fusus ventricosus Millet, p. 590 (non Lesson,
1842; Menke, 1843 and others).
1938 Fusus (Aptixis [sic]) rostratus var. ligeriana Pey-
rot, p. 233, pl. 4, gs 13, 14.
1938 Fusus (Aptixis [sic]) rostratus var. simplicior Pey-
rot, p. 234.
1938 Fusus (Aptixis [sic]) turonensis Peyrot, p. 235, pl.
4, gs 28, 34 (not g. 16 as stated in text; lapsus).
1952a Fusus (Aptyxis) rostratus ligerianus Peyrot, 1938
– Glibert, p. 349, pl. 11, g. 3.
1964 Fusinus (Aptyxis) ligerianus Peyrot, 1938 – Bré-
bion, p. 478.
Material and dimensions Maximum height 23.1 mm.
NHMW 2015/0133/0381 (1: Pl. 2, g. 4), NHMW 2015/
0133/0382 (Pl. 2, g. 5), NHMW 2015/0133/0383 (12), LC
(20), FVD (15). Le Pigeon Blanc, Le Landreau, Nantes
area, Loire-Atlantique department, NW France.
Discussion The enormous shell variability found in
this species was discussed at length by Glibert (1952a),
who synonymised the various forms described by Peyrot
(1938). The specimens from Le Pigeon Blanc are small
compared to some of those found in the middle Miocene
Loire Basin, similar to the smaller specimens illustrated
by Glibert (1952a, pl. 11, gs 3c, 3e). Like in the older
Miocene Loire Basin population, the shell width and
the strength of the shoulder cord, making the whorls
rounded or slightly angular, is variable. The protoconch
is paucispiral, composed of about 1.5 whorls, with a large
bulbous nucleus (Pl. 2, g. 5). However, Millet (1854)
listed several of the forms described by Peyrot (1938)
under the names Fusus rostratus Sismonda, F. omphale
Millet, F. vicinus Millet and F. ventricosus Millet. Mil-
let’s (1854) epithets are all nomina nuda (Landau et al.,
2016b), but F. omphale was validated in 1864 by the fol-
whorl weakly inated, not shouldered, with periphery
just below mid-whorl, roundly constricted at base, bear-
ing ten spiral cords and 18-19 ribs that weaken over base.
Aperture small, ovate, outer lip strongly thickened by
labial varix bearing 4-5 denticles within, adapical two
denticles more strongly developed. Siphonal canal open,
of moderate length, narrow, posteriorly recurved. Colu-
mella straight, bearing four stout folds. Columellar cal-
lus thickened, narrow, sharply delimited, parietal portion
poorly developed. Siphonal fasciole attened, not delim-
ited from base, bearing spiral cords.
Discussion – Although Anachis milleti nov. sp. occurs in
the lower Pliocene Assemblage III locality of Le Pigeon
Blanc, it is far more abundant in the upper Miocene As-
semblage I deposits. We therefore take the opportunity
to describe this species, but nominate St-Clément-de-la-
Place for the type stratum and locality.
This group of Anachis species with tall spires and reticu-
lated sculpture are well represented in the French Atlan-
tic Miocene. Anachis degrangei Dollfus in Degrange-
Touzin, 1894 from the middle Miocene Aquitaine and
Loire basins of France is the most similar to A. milleti in
its tall slender shape and relatively weakly inated last
whorl, but this species has fewer ribs and cords and the
two subsutural cords are more strongly developed, form-
ing a subsutural collar, resulting in a somewhat scalate
appearance to the spire. This subsutural collar is even
more strongly developed in A. clathrata (Dujardin, 1837),
which is lower spired than the preceding species and has
predominantly axial sculpture. This group is well repre-
sented in the NW French Assemblage I localities, where
two further species occur: A. collyrata (Millet, 1865)
(Text-g. 1/7-8) and A. hordacea (Millet, 1865) (Text-g.
1/5-6), both of which differ from A. milleti in having less
dense reticulated sculpture, with fewer ribs and cords on
the spire whorls. Like in A. milleti, in A. collyrata fur-
ther cords develop below the adapical cord on the spire
whorls, whereas in A. hordacea the abapical cord devel-
ops rst, followed by the middle cord and adapical cord
last. These species will be discussed further in the rel-
evant paper.
Anachis milleti is far less common in the Le Pigeon Blanc
assemblage than it is in the Assemblage I localities. How-
ever, the few specimens available are all about half as
large again as the largest from Assemblage I. The small
size of the Assemblage I species is a theme we will visit
repeatedly during this series of papers.
Brébion (1964, p. 411) recorded A. milleti (as A. fannyae)
from the middle Miocene Loire Basin (Manthelan), As-
semblage I localities (Sceaux-d’Anjou, St-Clément-de-la-
Place, Beaulieu), Assemblage II (Apigné, Le Temple du
Cerisier), Assemblage III (Le Pigeon Blanc, Palluau, La
Gauvinière).
Distribution Middle Miocene: Loire Basin (Brébion,
1964). Upper Miocene: Atlantic (Tortonian), NW France
(Brébion, 1964). Lower Pliocene: Atlantic, NW France
(Brébion, 1964).
32 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
partment, NW France.
Discussion This remarkable fusinid cannot be con-
fused with any other shell. It is the largest shell found at
Le Pigeon Blanc, characterised by sharply carinate later
whorls and axial sculpture restricted to the early teleo-
conch whorls. Whilst juvenile specimens are not uncom-
mon, adult shells are extremely difcult to nd. For full
discussion see Ceulemans et al. (2014).
Distribution Lower Pliocene: Atlantic, NW France
(Cossmann, 1901; Brébion, 1964; Ceulemans et al., 2014).
Subfamily Peristerniinae Tryon, 1880
Genus Polygona Schumacher, 1817
Type species (by monotypy) – Polygona fusiformis Schu-
macher, 1817, present-day, Caribbean.
1817 Polygona Schumacher, p. 241.
Polygona bellardii (Michelotti, 1847)
Plate 2, g. 7
*1847 Turbinella Bellardii Michelotti, p. 264, pl. 8, g. 2.
1884 Latirus Bellardii (Michtti.) – Bellardi, p. 24, pl. 1,
g. 19.
1935 Lathyrus (Ascolathyrus) Bellardii (Michtti) – Mon-
tanaro, p. 66, pl. 5, gs 7, 8.
1964 Latirus bellardii Michelotti, 1847 – Brébion, 473,
pl. 12, gs 3, 4.
Material and dimensions Maximum height 29.6 mm.
NHMW 2015/0133/0384 (1: Pl. 2, g. 7), NHMW 2015/
0133/0385 (30), LC (50+), FVD (50+). Le Pigeon Blanc,
Le Landreau, Nantes area, Loire-Atlantique department,
NW France.
Discussion Following Vermeij & Snyder’s (2006) re-
cent revision of the Latirus-group, we place this spe-
cies in the genus Polygona Schumacher, 1817. It ts well
within the Polygona angulata group of Vermeij & Sny-
der (2006) characterised by shells with a stepped spire, a
distinct shoulder angulation, nodose axial ribs, an adapi-
cal sinus present on the outer lip, the outer lip convex in
the central portion and lirae within the outer lip usually
beaded, but smooth in some small species. Polygona bel-
lardii (Michelotti, 1847) is one of the smaller species in-
cluded in this genus, and very similar in size, shape and
sculpture to some of the extant western Atlantic species
such as P. lactea (Matthews-Cascon, Matthews & Ro-
cha, 1991). The outer lip appears smooth in the Le Pigeon
Blanc specimen illustrated, but it is weakly lirate within,
the lirae stopping a short distance before the lip margin.
We would agree with Brébion that we cannot separate
the shells from Le Pigeon Blanc from that illustrated by
Bellardi (1884, pl. 1, g. 19) as Latirus bellardii (Micht-
ti.). There is some intraspecic variation in the French
lowing description: ‘Coq. allongée, fusiforme, composée
de sept à huit tours de spire, le dernier terminé en une
longue queue; tous garnis de côtes saillantes, couvertes,
comme tout le reste de la coquille, de stries élevées, al-
ternant en grosseur. Cette coquille atteint 6 centimètres
et demi de longeur, et 2 centimètres et plus de diamètre
(1864, p. 678)’, which applies to this species. It would have
been benecial to retain Peyrot’s (1938) name, which has
been widely used in the literature, but we cannot satisfy
the requirements of Article 23.9.1.2 (ICZN 1999) to con-
sider Millet’s name a nomen oblitum. Therefore Fusus
(Aptyxis) rostratus ligerianus Peyrot, 1938 must be con-
sidered a junior subjective synonym of Fusus omphale
Millet, 1864.
Brébion (1964, p. 479) recorded this species from Assem-
blage I localities (Renauleau, Sceaux d’Anjou, Tho rigné,
St-Clément-de-la-Place, Contigné, St-Michel, Cha lonnes,
Beaulieu), Assemblage II (Apigné, Moulin de Carcé), As-
semblage III (Le Pigeon Blanc, La Dixmérie), and As-
semblage IV (Gourbesville).
DistributionMiddle Miocene: Atlantic (Langhian) Loire
Basin, France (Glibert, 1952a). Upper Miocene: Atlantic
(Tor tonian and Messinian): north western France (Brébi-
on, 1964). Lower Pliocene: Atlantic, NW France (Brébion,
1964).
Genus Carinofusus Ceulemans, Landau & Van Ding-
enen, 2014
Type species (by monotypy) – Clavella neogenica Coss-
mann, 1901, Pliocene, north western France.
2014 Carinofusus Ceulemans, Landau & Van Dingenen,
p. 25.
Carinofusus neogenicus (Cossmann, 1901)
Plate 2, g. 6
*1901 Clavella neogenica Cossmann, p. 21.
2014 Carinofusus neogenicus (Cossmann, 1901) – Ceu-
lemans et al., p. 26, pl. 1 gs 1-4, pl. 2, gs 1-4
(cum syn).
Material and dimensions NHMW 2014/0288/0001
(Pl. 2, g. 6), height 87.2 mm, width 31.2 mm; NHMW
2014/0288/0002, height 101.5 mm, width 32.2 mm;
NHMW 2014/0288/0003, height 54.9 mm, width 27.0
mm; NHMW 2014/0288/0004 incomplete juvenile,
height 24.4 mm; NHMW 2014/0288/0005, outer lip and
spire fragment of large adult, height: 72.5 mm (frag-
ment; reconstructed height at least 110 mm); NHMW
2014/0288/0006 (9 juveniles + adult fragments); FVD (2
incomplete adults + 63 subadult/juvenile); LC (45 sub-
adult/juvenile); MNHN.F.A51235 (Brébion 1964, pl.
12, g. 6) and MNHN.F.A51236 (Brébion 1964, pl. 12,
g. 7), specimens from MNHN’s collection. Le Pigeon
Blanc, Le Landreau, Nantes area, Loire-Atlantique de-
Cainozoic Research, 17(1), pp. 23-61, June 2017 33
shells; in some specimens the axial ribs broaden and
are more widely spaced on the last whorl, becoming
subobsolete on the last half whorl. The protoconch is
paucispiral consisting of 1.5 smooth whorls with a bul-
bous nucleus (Pl. 2, g. 7c). Polygona spinifera (Bel-
lardi, 1884), also from the upper Miocene Tortonian of
Italy is closely similar to P. bellardii, but has a slightly
wider shell and develops small spines on the tubercles
at the whorl periphery. This might just be a variety of
P. bellardii, but we provisionally keep them distinct as
we have not seen any spiny specimens in the Le Pigeon
Blanc assemblage.
Brébion (1964, p. 474) recorded this species from Assem-
blage III (Le Pigeon Blanc, La Dixmérie, Palluau), and
Assemblage IV (Gourbesville).
Distribution – Upper Miocene: Proto-Mediterranean (Tor -
tonian): central Mediterranean, Italy (Michelotti, 1847;
Bel lardi, 1884; Montanaro, 1935). Lower Pliocene: Atlan-
tic, NW France (Brébion, 1964). Upper Pliocene-Pleis-
tocene: NW France (Brébion, 1964).
Family Nassariidae Iredale 1916 (1835)
Subfamily Nassariidae Iredale 1916 (1835)
Genus Tritia Risso, 1826
Type species (by subsequent designation, Gray, 1847)
Buccinum reticulatum Linnaeus, 1758, present-day, Eu-
rope.
1799 Nassa Lamarck, 1799. Type species (by monotypy):
Buccinum mutabile Linnaeus, 1758, present-day,
Mediterranean. Junior homonym of Nassa Röding,
1798.
1826 Tritia Risso, p. 172.
1826 Cyclope Risso, p. 169. Type species (by mono-
typy): Cyclope neritoidea Risso, 1826 (substitute
name for Buccinum neriteum Linnaeus, 1758),
present-day, Mediterranean.
1840 Cyclonassa Swainson, p. 300. Type species (by
monotypy): Buccinum neriteum Linnaeus, 1758,
present-day, Mediterranean. Junior objective syn-
onym of Cyclope, with the same type species.
1847 Hinia Leach in Gray, p. 269. Type species (by sub-
sequent designation, Cossmann, 1901): Buccinum
reticulatum Linnaeus, 1758, present-day, Mediter-
ranean.
1853 Telasco H. Adams & A. Adams, p. 119. Type
species (by subsequent designation: Bucquoy et
al., 1882): Buccinum variabile Philippi, 1836,
present-day, Mediterranean.
1853 Uzita H. Adams & A. Adams, p. 120. Type species
(by subsequent designation, Cossmann, 1901):
Buccinum miga Bruguière, 1789, present-day,
West Africa.
1853 Neritula H. Adams & A. Adams, p. 122. Type spe-
cies (by subsequent designation, Cernohorsky,
1984): Buccinum neriteum Linnaeus, 1758,
present-day, Mediterranean.
1853 Amycla H. Adams & A. Adams, p. 186. Type spe-
cies (by subsequent designation: Bucquoy et al.,
1882): Buccinum corniculum Olivi, 1792, present-
day, Mediterranean.
1912 Gussonea Monterosato, p. 295. Type species (by
original designation): Buccinum tinei Maravigna,
1840, present-day, Mediterranean.
1918 Amyclina Iredale, p. 28, 31. Type species (by orig-
inal designation): Buccinum corniculum Olivi,
1792, present-day, Mediterranean. Unnecessary
substitute name for Amycla H. Adams & A. Ad-
ams, 1853.
1920 Hannonia Pallary, p. 36. Type species (by mono-
typy): Nassa tingitana Pallary, 1901, present-day,
Mediterranean.
1929 Naytiopsis Thiele, p. 324. Type species (by mono-
typy): Buccinum granum Lamarck, 1822, present-
day, Mediterranean.
1929 Proneritula Thiele, p. 324. Type species (by mono-
typy): Cyclope westerlundi Brusina, 1900 [= Tritia
neritea (Linnaeus, 1758)], present-day, Mediterra-
nean.
NoteWe draw attention to an important paper by Galin-
do et al. (2016) on nassariid phylogeny, which showed
that shell characters fail to accurately dene supraspe-
cic taxa within Nassariinae. They suggested that all
the European and West African species should be placed
under Tritia Risso, 1826 (type species Buccinum reticu-
latum Linnaeus, 1758, present-day, Europe), although
surprisingly some southern Australian and New Zealand
species also t within the Tritia molecular clade. They
also recognised the genus Naytia H. & A. Adams in West
Africa (type species Strombus glabratus G.B. Sowerby
II, 1842, present-day, West Africa). The western Atlantic
and eastern Pacic species should be placed in Phron-
tis H. & A. Adams, 1853 (type species Buccinum tiarula
Kiener, 1841, present-day, Pacic coast of America). The
genus Nassarius Duméril, 1805 (type species Buccinum
arcularia Linnaeus, 1758, present-day, Indo-Pacic) was
considered strictly Indo-Pacic, together with the genus
Reticunassa Iredale, 1936 (type species, Nassa paupera
Gould, 1850, present-day, Indo-Pacic). Therefore, all
the species described and discussed by Van Dingenen
et al. (2015) under the genus Nassarius should be reas-
signed to Tritia.
Van Dingenen et al. (2015) described and recorded 11
nassariid species from Le Pigeon Blanc:
Tritia brebioni (Van Dingenen, Ceulemans, Landau &
Silva, 2015)
Tritia crebresulcata (Bellardi, 1882)
Tritia landreauensis (Van Dingenen, Ceulemans, Lan-
dau & Silva, 2015)
Tritia merlei (Van Dingenen, Ceulemans, Landau & Sil-
va, 2015)
Tritia spectabilis vandewouweri (Glibert, 1959)
Tritia pacaudi (Van Dingenen, Ceulemans, Landau &
Silva, 2015)
Tritia palumbis (Van Dingenen, Ceulemans, Landau &
34 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
north-east Spain (Martinell, 1982; Gili, 1991), southern
France (Martinell & Domènech, 1986; Gili, 1991); cen-
tral Mediterranean, Italy (Bellardi, 1882; Chirli, 1988,
2000), Tunisia (Fekih, 1975). Upper Pliocene: western
Mediterranean, Estepona Basin, Spain (Landau et al.,
2009), Alpes Maritimes (Chirli & Richard, 2008); central
Mediterranean, Italy (Bellardi, 1882; Malatesta, 1974;
Cavallo & Repetto, 1992). Pliocene indet.: central Medi-
terranean, Italy (Sacco, 1904).
Tritia nitida (Jeffreys, 1867)
Plate 2, g. 9
*1867 Nassa nitida Jeffreys, p. 349.
2009 Nassarius nitidus (Jeffreys, 1867) – Landau et al.,
p. 31, pl. 6, gs 3-5 (cum syn.).
Material and dimensions Height 10.2 mm. NHMW
2015/0133/0411 (1: Pl. 2, g. 9). Le Pigeon Blanc, Le Lan-
dreau, Nantes area, Loire-Atlantique department, NW
France.
Discussion – Rolán & Luque (1994) showed that T. re-
ticulata (Linnaeus, 1758) and T. nitida (Jeffreys, 1867)
were separable on shell, soft parts and egg capsule mor-
phology. The shell of T. nitida differs in having a some-
what scalate spire and fewer axial ribs (11-19 in T. nitida
vs. 16-23 in T. reticulata). The mid-apertural denticles
are often more strongly developed in T. reticulata, and T.
nitida has a protoconch with less than two whorls, T. re-
ticulata more than two. The single shell from Le Pigeon
Blanc is squat with only nine strongly elevated axial ribs,
but probably belongs within the range of variability of
T. nitida. Unfortunately, the protoconch is not preserved
in the French fossil shell. Tritia antiqua (Bellardi, 1882)
also belongs within the T. reticulata species complex. It is
closely similar to T. nitida (Jeffreys, 1867), but differs in
having a squatter shell than T. nitida, and having strong
tubercles on the columella, which extend along the whole
length of the inner lip. Tubercles on the columella are
sometimes more numerous, but not as strongly developed
in either T. nitida or T. reticulata. The shell from Le Pi-
geon Blanc has two small abapical columellar tubercles
and a slightly stronger parietal tubercle, all far weaker
than those seen in T. antiqua.
Distribution Lower Pliocene: western Mediterranean,
north-eastern Spain (Martinell, 1982; Gili, 1991); central
Mediterranean, Italy (Bellardi, 1882; Chirli, 2000), Tuni-
sia (Fekih, 1975). Upper Pliocene: Atlantic, NW France
(this paper), Mondego Basin, central west Portugal (Sil-
va, 2001); western Mediterranean, Estepona, southern
Spain (Landau et al., 2009), Alpes Maritimes (Chirli &
Richard, 2008); central Mediterranean, Italy (Bellardi,
1882; Ruggieri et al., 1959). Pleistocene: Atlantic, United
Kingdom (Harmer, 1916); central Mediterranean, Italy
(Cerulli-Irelli, 1911). Recent: Atlantic, Mediterranean
(Rolán & Luque, 1994).
Silva, 2015)
Tritia columbina (Van Dingenen, Ceulemans, Landau &
Silva, 2015)
Tritia turpis (Van Dingenen, Ceulemans, Landau & Sil-
va, 2015)
Tritia turonensis (Deshayes, 1844)
Tritia sp. A.
To this list we add two further nassariid species:
Tritia gibbosula pliopergibbosa (Sacco, 1904)
Plate 2, g. 8
*1904 Nassa (Arcularia) gibbosula var. pliopergibbosa
Sacco, p. 63, pl. 15, gs 34-35.
non 1964 Arcularia gibbosula Linné, 1766 [sic] – Brébion,
p. 434, pl. 10, gs 32, 33 [non Nassarius gibbosu-
lus (Linnaeus, 1758) = Tritia gendryi (Van Ding-
enen, Ceulemans, Landau & Silva, 2015)].
2009 Nassarius gibbosulus pliopergibbosus (Sacco,
1904) – Landau et al., p. 60, pl. 12, gs 11-12, pl.
22, g. 2 (cum syn.).
Material and dimensions Height 12.4 mm. NHMW
2015/0133/0410 (1: Pl. 2, g. 8). Le Pigeon Blanc, Le Lan-
dreau, Nantes area, Loire-Atlantique department, NW
France.
Discussion Tritia gibbosula pliopergibbosa (Sacco,
1904) differs from the present-day Mediterranean and
adjacent Atlantic Tritia gibbosula (Linnaeus, 1758) in the
proportions of their shells. The fossil shells are less tall
than the present-day shells, but of similar diameter. This
gives the shell a squatter, more quadrate appearance, es-
pecially the last whorl. In the fossil shells the mean ratio
of the diameter against height of the shell is 75%, whereas
in the Recent shells it is only 69%. For further discussion
see Landau et al. (2009).
This species is represented in the Le Pigeon Blanc as-
semblage by a single, well preserved shell. It is typical
in size and shape for the species. Brébion’s (1964, p. 434)
records for Arcularia gibbosula in the Assemblage IV
localities of north western France represent Tritia gen-
dryi (Van Dingenen, Ceulemans, Landau & Silva, 2015).
This species is much larger than T. gibbosula or T. g. plio-
pergibbosa, it has subobsolete axial folds on the dorsum,
but no distinct dorsal gibbosity, the venter is rounded
and not depressed or dimpled as in T. gibbosula or T. g.
pliopergibbosa, and the parietal tooth is more strongly
developed, below which the columella is more deeply ex-
cavated in the mid-portion.
Distribution – Middle Miocene: Atlantic, Aquitaine Ba-
sin, France (Peyrot, 1925). Late Miocene: Atlantic, Ca-
cela, southern Portugal (NHMW coll.); Proto-Mediter-
ranean, Italy (Bellardi, 1882; Montanaro, 1939; Venzo
& Pelosio, 1963). Lower Pliocene: Atlantic, NW France
(this paper), Guadalquivir Basin, Spain (González-Del-
gado, 1989; Landau et al., 2011); western Mediterranean,
Cainozoic Research, 17(1), pp. 23-61, June 2017 35
data and is mainly limited to a repeat of Bellardi’s Latin
descriptions accompanied by a new description in Italian.
The mitriid material from the lower Pliocene Assemblage
III locality of Le Pigeon Blanc is usually incomplete and
invariably has the apex abraded and the protoconch miss-
ing. We recognise three species in the assemblage. They
all have spiral sculpture on early teleo conch whorls, they
all have almost at-sided to weakly concave spire whorls
and they all lack spiral sculpture over the base. We have
compared them to similar described forms, but note that
the identications must be considered provisional until
the family is reviewed. It is also possible that further spe-
cies lurk amongst the broken mitriid material at hand.
Mitra sp. 1
Plate 2, g. 10
Material and dimensions Maximum height 27.2 mm.
NHMW 2015/0133/0395 (1: Pl. 2, g. 10); NHMW
2015/0133/0421 (3), LC (3), FVD (1). Le Pigeon Blanc,
Le Landreau, Nantes area, Loire-Atlantique department,
NW France.
Description Shell small for genus, slender fusiform.
Apex abraded in all specimens. Teleoconch of six almost
at-sided relatively tall whorls separated by supercial,
markedly oblique linear suture. Early teleoconch whorls
bearing 6-7 narrow attened cords separated by punctate
grooves; cords weaken abapically and disappear on third
whorl. Last whorl slender, 60% of total height, hardly
constricted at base, devoid of spiral sculpture over base.
Aperture elongate, narrow, 40% of total height, outer lip
simple, siphonal canal moderate length, weakly recurved.
Columella with four oblique folds, weakening abapically.
Columellar callus sharply delimited, poorly expanded,
forming narrow callus rim, thickened abapically, form-
ing medial border of narrow, shallow umbilical slit.
Discussion Mitra sp. 1 has a slender fusiform shell
shape, with almost at-sided spire whorls, separated by
a shallow oblique suture. It is closely similar to Mitra
infundibulum Bellardi, 1887, described from the up-
per Miocene of Italy. These shell features led Bellardi
(1887a, p. 12) to erect a 6ª Seriefor this species alone.
Despite the similarity in shell shape, M. infundibulum
was described as having ve columellar folds (Bellardi,
1887a, p. 12), whereas all the French fossil specimens at
hand have four.
Distribution – Lower Pliocene: Atlantic, NW France (this
paper).
Mitra gravis Bellardi, 1887
Plate 3, gs 1-3
*1887a Mitra gravis Bellardi, p. 11, pl. 1, g. 6.
1964 Mitra gravis Bellardi, 1887 – Brébion, p. 501, pl.
12, gs 31, 32.
Superfamily Muricoidea Ranesque, 1815
Family Mitridae Swainson, 1829
Subfamily Mitrinae Swainson, 1829
Genus Mitra Lamarck, 1798
Type species (by tautonymy) Voluta mitra Linnaeus,
1758, present-day, Indo-Pacic.
1798 Mitra Lamarck, p. 369.
1798 Mitra Röding, p. 135. Type species (by subsequent
designation, Winckworth, 1945): Voluta episco-
palis Linnaeus, 1758, present-day, Indo-Pacic.
Homonym and junior objective synonym of Mitra
Lamarck, 1798.
1815 Mitraria Ranesque, p. 145. Invalid: established
as a substitute name for Mitra Lamarck, 1798
without diagnosis or included species.
1853 Chrysame H. Adams & A. Adams, p. 171. Type
species (by subsequent designation, Cox, 1927):
Mitra coronata Lamarck, 1811, present-day, West
Pacic.
1900 Fuscomitra Pallary, p. 263. Type species (by sub-
sequent designation, Cox, 1936): Mitra fusca Pal-
lary, 1900 (= M. cornea Lamarck, 1811), present-
day, Mediterranean.
1917 Episcomitra Monterosato, p. 26. Type species (by
monotypy): Mitra zonata Marryat, 1818, present-
day, Mediterranean.
Note Landau et al. (2011, 2013) argued that the the
European Neogene Mitraspecies of the M. fusiformis
group and present-day Mediterranean species should
be placed in separate genus [Episcomitra Monterosato,
1917; 2011: Fuscomitra Pallary, 1900; 2013]. Apart from
the unbarbed outer lip, the European species tend to have
spiral sculpture restricted to the base on the last whorl,
whereas many of the Pacic Mitra species have a totally
spirally striate teleoconch.
The molecular phylogeny presented by Fedosov et al.
(2015) included one Mediterranean species Mitra cor-
nicula (Linnaeus, 1758) and found it was nestled amongst
the core Mitridae (2015, p. 347, g. 3). Therefore the shell
features highlighted by Landau et al. (2011, 2013), seem
not to be of generic value. Similarly, Landau et al. (2016a)
separated the American tropical eastern Pacic spe-
cies and placed them in the genus Atrimitra Dall, 1918.
Atri mitra is also synonymised with Mitra by WoRMS
(Rosenberg, 2015). In view of the above, this might well
be true, however, no eastern Pacic species were included
in the molecular phylogeny of Fedosov and until proven
we exclude Atrimitra from the synonymy of Mitra.
The European fossil Mitra species are in need of review.
There has been no critical review of the Miocene-Pliocene
Mediterranean species following the standard works on
the family by Bellardi (1887a, b, 1888). Cernohorsky
(1976) suggested that many of Bellardi’s species were
synonyms, but neither performed a critical review, nor
formally synonymised Bellardi’s taxa. Chirli (2002) g-
ured numerous Italian Pliocene species attributing them
to Bellardi’s species, but his text adds little comparative
36 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in par t)
abrasion, separated by punctate grooves; cords weaken
abapically and disappear on third whorl. Last whorl
moderately inated, 64% of total height, weakly con-
stricted at base, devoid of spiral sculpture over base.
Aperture elongate, narrow, 43% of total height, outer
lip simple, siphonal canal moderate length, weakly re-
curved and twisted. Columella with ve oblique folds,
weakening abapically. Columellar callus sharply de-
limited, poorly expanded, forming narrow callus rim,
thickened abapically, forming medial border of narrow,
shallow umbilical slit.
Discussion This species seems to belong to Bellardi’s
(1887a, p. 14) ‘8ª Serie’, characterised by their regularly
fusiform shape, the presence of cords on the early whorl
and absence of them over the base in most species, ve
columellar folds and the siphonal canal slightly twisted
to the left. Bellardi used Mitra turricula De Cristofori &
Jan as reference species for the group, which differs from
Mitra atava Bellardi, 1887 in having teleoconch whorls
strongly shouldered just below the suture. Some of the
other species included in this group are closely similar
to M. atava and, like the rest of the group, need to be
revised. This is the largest of the Le Pigeon Blanc mitrids
and also bears some resemblance to Mitra fusiformis
(Brocchi, 1814), which has 5-6 columellar folds, but the
latter differs in having shouldered whorls and the early
spire whorls lack spiral sculpture.
Distribution – Lower Pliocene: Atlantic, NW France (this
paper).
Family Cystiscidae Stimpson, 1865
Subfamily Persiculinae G. A. Coovert & H. K. Coovert,
1995
Genus Gibberula Swainson, 1840
Type species (by monotypy) – G. zonata Swainson, 1840
(= Volvaria oryza Lamarck, 1822), present-day, eastern
Atlantic.
1840 Gibberula Swainson, p. 323.
1875 Granula Jousseaume, p. 167. Type species (by
subsequent designation; Coan, 1965): Marginel-
la bensoni Reeve, 1865, present-day, South Af-
rica.
1951 Kogomea Habe in Kuroda, p. 103. Type species
(by original designation): Erato novemprovincia-
lis Yokoyama, 1928, Pleistocene, Japan.
1957 Epiginella Laseron, p. 279. Type species (by origi-
nal designation): E. ablita Laseron, 1957, present-
day, Australia.
1957 Phyloginella Laseron, p. 280. Type species (by
original designation): P. compressa Laseron, 1957,
present-day, Coral Sea.
1962 Diluculum Barnard, p. 14. Type species (by origi-
nal designation): D. inopinatum Barnard, 1962,
present-day, South Africa.
Material and dimensions Maximum height 28.4
mm. NHMW 2015/0133/0417 (1: Pl. 3, g. 1); NHMW
2015/0133/0418 (1: Pl. 3, g. 2); NHMW 2015/0133/0419
(1: Pl. 3, g. 3); NHMW 2015/0133/0420 (15), LC (50+),
FVD (18). Le Pigeon Blanc, Le Landreau, Nantes area,
Loire-Atlantique department, NW France.
Description Shell small to medium-sized for genus,
fusiform with tall conical spire. Apex abraded in all
specimens. Teleoconch of 6-7 weakly convex whorls
separated by impressed linear suture. Early teleoconch
whorls bearing seven narrow attened cords separated by
punctate grooves; cords weaken abapically and disappear
on third whorl. Last whorl weakly inated, 60% of total
height, hardly constricted at base, devoid of spiral sculp-
ture over base. Aperture elongate, narrow, 38% of total
height, outer lip simple, siphonal canal short. Columella
with four oblique folds, weakening abapically. Columel-
lar callus sharply delimited, poorly expanded, forming
narrow callus rim, thickened abapically closing umbili-
cus completely in most specimens.
Discussion We would agree with Brébion (1964, p.
501) in considering this species closely similar to Mitra
gravis Bellardi, 1887 described from the upper Miocene
of Italy. These mitrids form part of Bellardi’s (1887a, p.
14) ‘5ª Serie’, characterised by their relatively long spire
in relation to the last whorl, giving them a somewhat tur-
riculate rather than fusiform contour, cords on the early
whorl and absence of them over the base, four columellar
folds and a short siphonal canal. Of the species included
in this group by Bellardi, the Le Pigeon Blanc shells are
most like M. gravis, and although we have not seen this
species and it was not re-illustrated by Ferrero Mortara
et al. (1981) nor Davoli (2000), we provisionally consider
them conspecic.
Distribution Upper Miocene: Proto-Mediterranean
(Tortonian), Italy (Bellardi, 1887a). Lower Pliocene: At-
lantic, NW France (Brébion, 1964).
Mitra cf. atava Bellardi, 1887
Plate 3, gs 4-5
cf. *1887a Mitra atava Bellardi, p. 15, pl. 1, g. 12.
cf. 1981 Mitra atava Bellardi, 1887 Ferrero Mortara et
al., p. 147, pl. 41, g. 3.
Material and dimensions Maximum height 51.4
mm. NHMW 2015/0133/0393 (1: Pl 3, g. 4), NHMW
2015/0133/0394 (7), LC (9), FVD (3). Le Pigeon Blanc,
Le Landreau, Nantes area, Loire-Atlantique department,
NW France.
Description Shell medium-sized for genus, broad
fusiform with conical spire. Apex abraded in all speci-
mens. Teleoconch of 6-7 convex whorls separated by
impressed linear suture. Early teleoconch whorls bear-
ing narrow attened cords, difcult to quantify due to
Cainozoic Research, 17(1), pp. 23-61, June 2017 37
Gibberula miliaria (Linnaeus, 1758)
Plate 3, g. 6
*1758 Voluta miliaria Linnaeus, p. 730.
1952a Persicula (Gibberula) miliaria Linné, 1766
Glibert, p. 369, pl. 12, g. 8.
1964 Gibberula miliaria Linné, 1766 – Brébion, p. 533.
1987 Gibberula miliaria (Linné, 1758) – Gofas, p. 125,
gs 14-16, pl. 1, g. d-d’.
2003 Gibberula miliaria (Linné, 1758) Giannuzzi-
Savelli et al., p. 276, gs 699-704.
Material and dimensions Maximum height 7.5 mm.
NHMW 2015/0133/0396 (1: Pl. 3, g. 6), NHMW 2015/
0133/0397 (8), LC (12), FVD (5). Le Pigeon Blanc, Le
Landreau, Nantes area, Loire-Atlantique department,
NW France.
Discussion Two Gibberula species occur in the Le Pi-
geon Blanc assemblage, the larger of the two is ascribed
to Gibberula miliaria (Linnaeus, 1758). The spire is low
in most specimens, the outer lip denticulate in all, and
the columella bears 5-6 folds, weakening adapically. It is
similar to the Pliocene Mediterranean G. proxima Lan-
dau, La Perna & Silva, 2006 from the Estepona Basin,
Spain, but the latter is broader-shelled, with the shoulder
placed higher, G. miliaria is more slender and regularly
cylindrical in shape, and the labial varix is thicker, espe-
cially abapically in G. proxima.
Gibberula miliaria was also recorded by Glibert (1952a)
from the middle Miocene Loire Basin and by Brébion
(1964) in the Assemblage I, II and IV deposits in north-
western France (Brébion, 1964). These records will be
discussed in a subsequent paper. Interestingly, the pres-
ence of this species in the Pliocene Mediterranean and
Iberian coast has not been conrmed in the recent lit-
erature (Chirli, 2002; Landau et al., 2006b; Silva et al.,
2011). We have not included older references, which need
to be conrmed.
Silva et al. (2011) discussed the biogeographical im-
plications of the European shallow marine marginel-
liform gastropods and considered them a relatively
thermophilic group. They demonstrated the increase in
generic and specic diversity with a decrease in latitude
(southwards), and the southwards shift in diversity since
Pliocene time (2011, tables 1, 2, gure 7). The Le Pigeon
Blanc marginellid assemblage suggests that Gibberula is
the most tolerant to cooler waters, as it is the only genus
present in the lower Pliocene of north western France and
we are not aware of any Pliocene records of marginel-
lids further north. It is also interesting that the two spe-
cies found here; G. miliaria and G. philippii (Montero-
sato, 1878) are now extant in the Mediterranean, whereas
the Pliocene cohort of species (Chirli, 2002; Landau et
al., 2006b; Silva et al., 2011) are extinct. Landau et al.
(2006b) and Silva et al. (2011) traced the diverse origins
of the Pliocene Mediterranean marginellid assemblage
and recognised a strong West African inuence to the
fauna. It is possible that the cooling of SST (sea surface
temperture) since the Pliocene resulted in the extinction
of the more thermophilic West African inuenced cohort
of species, which were then replaced by more tempera-
ture tolerant northern forms.
Brébion (1964, p. 503) recorded this species from Assem-
blage I localities (Renauleau, Thorigné, Sceaux d’Anjou,
St-Clément-de-la-Place, St-Michel, Beaulieu), Assem-
blage II (Apigné), Assemblage III (Le Pigeon Blanc, Le
Girondor), and Assemblage IV (Gourbesville). In the
distribution we have only included the Assemblage III
records. The others will be discussed in subsequent pa-
pers.
Distribution Lower Pliocene: Atlantic, NW France
(Brébion, 1964). Present-day: Mediterranean and adja-
cent Atlantic (Gofas, 1987).
Gibberula philippii (Monterosato, 1878)
Plate 3, g. 7
*1878 Marginella philippi [sic] Monterosato, p. 109.
(nom. nov. pro Marginella minuta Philippi, 1844
non Pfeiffer, 1840 = Gibberula pfeifferi Faber,
2004; non Marginella minima Gray, 1826).
1964 Gibberula philippi [sic] Monterosato, 1878 – Bré-
bion, p. 534, pl. 13, gs 15-17.
1987 Gibberula philippii (Monterosato, 1878) – Gofas,
p. 129, gs 18-20, pl. 2, gs b-e.
2003 Gibberula philippii (Monterosato, 1878) Gian-
nuzzi-Savelli et al., p. 276, gs 713-714.
Material and dimensions Maximum height 3.5 mm.
NHMW 2015/0133/0398 (1: Pl. 3, g. 7), NHMW 2015/
0133/0399 (20), LC (50+), FVD (50+). Le Pigeon Blanc,
Le Landreau, Nantes area, Loire-Atlantique department,
NW France.
Discussion The smaller of the two Gibberula spe-
cies found at Le Pigeon Blanc we attribute to Gibberula
philippii (Monterosato, 1878). The French fossil speci-
mens agree with the description and gures given by Go-
fas (1987) for the species, except that the denticulation at
the inner edge of the outer lip is variable: well developed
in some specimens, absent in others. Interestingly, the
presence of this species in the Pliocene Mediterranean
and Iberian coast has not been conrmed in the recent
literature (Chirli, 2002; Landau et al., 2006b; Silva et
al., 2011), for further discussion see under G. miliaria
(above). We have not included older references, which
need to be conrmed.
Brébion (1964, p. 503) recorded this species from the
middle Miocene Loire Basin (La Beurelière), Assem-
blage I localities (Thorigné, St-Clément-de-la-Place, St-
Michel), Assemblage II (Apigné), Assemblage III (Le Pi-
geon Blanc, Palluau), and Assemblage IV (Gourbesville).
In the distribution we have only included the Assemblage
III records. The others will be discussed in subsequent
papers.
Distribution Lower Pliocene: Atlantic, NW France
38 Van Dingenen, Ceulemans & Landau. The lower Pliocene gast ropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
inant medium-sized gastropod found in Le Pigeon Blanc,
with hundreds of specimens present.
The presence of Amalda in vast numbers in the Le Pigeon
Blanc assemblage is interesting. Landau & Silva (2006)
pointed out that today Amalda is a temperate genus and
that it occurs in tropical and subtropical zones only in
places where there is upwelling of cooler, nutrient-rich
waters. They suggested that in the fossil assemblages, the
presence of large numbers of Amalda, in assemblages of
tropical or subtropical character, was an indicator of up-
welling. Thus, the great number of Amalda found in the
Le Pigeon Blanc assemblage, that is subtropical in char-
acter, suggests upwelling in the area. This hypothesis is
further supported by the presence of Patella pellucida
Linnaeus, 1758 (Ceulemans et al., 2016, p. 53). Patella
pellucida is a north-western Atlantic shallow water graz-
ing gastropod typically living and feeding on laminar-
ian algae (Silva et al., 2006). Today the species is usually
considered to be primarily a northern cooler water spe-
cies, inhabiting the Atlantic coasts of Europe, but rarely
found off the Atlantic NW African coast and western-
most Mediterranean. At higher sea surface temperatures
laminarian algae become physiologically stressed and are
found only where seasonal upwelling of cooler nutrient-
rich waters occurs. Such oceanographic conditions ex-
ist today, e.g., off the western coast of Iberia. Therefore,
the presence of P. pellucida in Le Pigeon Blanc suggests
laminarian algae surviving in subtropical waters, and
hence the presence of upwelling.
Brébion (1964, p. 483) recorded A. glandiformis morpho-
type elongata from Assemblage I localities (Thorigné, St-
Michel), Assemblage II (Apigné, Lillion), and Assemblage
III (Corcoué-sur-Logne, Les Cléons, Le Pigeon Blanc, Le
Girondor, La Gauvinière, La Dixmérie, Palluau).
Distribution Middle Miocene: Atlantic (Langhian),
Loire Basin, France (Glibert, 1952a); Aquitanian Basin,
France (Glibert, 1960); Paratethys, Hungary (Strausz,
1966), Bulgaria (Kojumdgieva & Strachimirov, 1960);
Proto-Mediterranean: Karaman Basin, Turkey (Landau
et al., 2013). Upper Miocene: Atlantic, Cacela, Portugal
(Pereira da Costa, 1867), NW France (Glibert, 1960; Bré-
bion, 1964); Proto-Mediterranean: Italy (Sacco, 1904; Da-
voli, 1989). Lower Pliocene: Atlantic, Guadalquivir Basin
(González-Delgado, 1992; Landau et al., 2011). Upper
Pliocene: Atlantic, Mondego Basin, Portugal (Cox, 1941;
Zbyszewski, 1959; Brébion, 1971; Silva, 2001, 2002), Bou
Regreg Basin, NW Morocco (Glibert, 1960); western
Mediterranean: Estepona Basin (Landau & Silva, 2006).
Subfamily Scaphellinae Gray, 1857
Genus Euroscaphella Van Dingenen, Ceulemans & Lan-
dau, 2014
Type species (by original designation) – Voluta lamberti
J. Sowerby, 1816, North Sea Basin, Pliocene, England.
2014 Euroscaphella Van Dingenen, Ceulemans, & Lan-
dau, p. 104.
(Brébion, 1964). Present-day: Mediterranean and adja-
cent Atlantic (Gofas, 1987).
Superfamily Olivoidea Latreille, 1825
Family Olividae Latreille, 1825
Genus Amalda H. Adams & A. Adams, 1853
Type species (by subsequent designation, Cossmann,
1899) Ancillaria tankervilii Swainson, 1825, present-
day, Caribbean.
1853 Amalda H. Adams & A. Adams, p. 148.
Amalda glandiformis (Lamarck, 1810) morphotype
elongata
Plate 3, g. 8
*1810 Ancillaria glandiformis Lamarck, p. 305.
1964 Ancilla (Baryspira) glandiformis Lamarck, 1810
Brébion, p. 482.
2006 Amalda glandiformis (Lamarck, 1810) morpho-
type elongata Landau & Silva, p. 6, pl. 1, gs
1-8 (cum syn.).
2013 Amalda glandiformis (Lamarck, 1810) – Landau
et al., p. 222, pl. 32, gs 9, 10, pl. 68, g. 4, pl. 80,
g. 6.
Material and dimensions Maximum height 51.2 mm.
NHMW 2015/0133/0386 (1: Pl. 3, g. 8), NHMW 2015/
0133/0387 (12), LC ( 25), FVD (50+). Le Pigeon Blanc,
Le Landreau, Nantes area, Loire-Atlantique department,
NW France.
Discussion This species was discussed in detail by
Landau & Silva (2006) and Landau et al. (2013). The
elongata morphotype of Amalda glandiformis (Lamarck,
1810) is characterised by its tall narrow spire and thin
callus. Specimens of the form elongata are very similar
to Amalda obsoleta (Brocchi, 1814), generally fusiform
in shape, with a tall narrow spire and thin spire and pari-
etal callusses, but are separated based on the width of the
ancillid band, which is much wider in A. obsoleta than in
A. glandiformis forma elongata. As discussed by Landau
& Silva (2006), all the Pliocene Iberian Atlantic reports
for the genus Amalda correspond to A. glandiformis mor-
photype elongata, although not always identied as such.
Cox (1941) erected the name Ancilla marinhensis for the
Portuguese Pliocene form, without justifying the sepa-
ration, apart from saying they were different from the
French and Italian Miocene specimens. Brébion (1971,
p. 130) synonymised the Portuguese shells with Amal-
da glandiformis. Zbyszewski (1959), on the other hand
considered them A. obsoleta. All these shells fall within
the range of A. glandiformis form elongata as dened by
Glibert (1952a). The same is true in the Le Pigeon Blanc
assemblage. Amalda glandiformis forma elongata is
the only form of the species found in the north-western
French Assemblage III localities. Moreover, it is the dom-
Cainozoic Research, 17(1), pp. 23-61, June 2017 39
Bivetia Jousseaume, 1887b, p. 193, non 1887a, p.
163.
1949 Bivetiella Marks, p. 456. Type species (by original
designation): Cancellaria similis G. B. Sowerby
I, 1833, present-day, north western Africa. Junior
objective synonym and junior homonym of Bive-
tiella Wenz, 1943.
Bivetiella cancellata (Linné, 1767)
Plate 4, g. 1
*1767 Voluta cancellata Linné, p. 1191.
2006a Cancellaria (Bivetiella) cancellata (Linnaeus, 1767)
– Landau et al., p. 63, pl. 1, gs 1, 2 (cum syn.).
2007 Bivetiella cancellata (Linnaeus, 1767) – Verheck-
en, p. 295, gs 11, 13-18, 21A, 22, 23A, 24A, 25.
2010 Cancellaria cancellata (Linnaeus, 1767) Sosso
& Dell’Angelo, p. 43, unnumbered g. p. 59 3rd
row middle.
2011 Bivetiella cancellata (Linnaeus, 1767) Brunetti
et al., p. 88, gs 1C-G, 2A-C.
Material and dimensions Maximum height 10.5 mm
(incomplete juvenile). LC (2 juveniles, most complete; Pl.
4, g. 1). Le Pigeon Blanc, Le Landreau, Nantes area,
Loire-Atlantique department, NW France.
Discussion This species was fully discussed by Land-
au et al. (2006a) and Brunetti et al. (2011). It seems to
be very uncommon in the Le Pigeon Blanc assemblage,
where it is represented by a few juvenile specimens. Bive-
tiella cancellata (Linné, 1767) today is found along the
coast of West Africa and into the Mediterranean, but not
the southern Atlantic Iberian coast. This record in the
lower Pliocene of north western France again demon-
strates a range contraction for the species due to cooling
since Pliocene time.
Distribution Lower Pliocene: Atlantic, NW France
(this paper), Guadalquivir Basin, Spain (González-Del-
gado, 1992; Landau et al., 2011); Morocco (Lecointre,
1952; González-Delgado et al., 1999); western Mediter-
ranean, northern Spain (Almera & Boll, 1884; Mar-
tinell, 1979); Roussillon, France (Fontannes, 1882);
central Mediterranean, Italy (Chirli, 2002; Brunetti et
al., 2011); Tunisia (Fekih, 1975). Upper Pliocene: Atlan-
tic, Mondego Basin, Portugal (Zbyszewski, 1959; Silva,
2001), Estepona Basin, Spain (Vera-Peláez et al., 1995;
Landau et al., 2006a); central Mediterranean, Italy
(Sacco, 1894; Glibert, 1960; Caretto, 1963; Malatesta,
1974; Pavia, 1975; Caprotti, 1976; Chirli, 1988; Cavallo
& Repetto, 1992; Sosso & Dell’Angelo, 2010; Brunetti
et al., 2011). Pleistocene: western Mediterranean, Mo-
rocco (Lecointre, 1952), Balearic Islands (Cuerda Bar-
celó, 1987); central Mediterranean, Italy (CerulliIrelli,
1911; Glibert, 1960; Malatesta, 1960). Present-day:
West Africa, Morocco-Angola, Canary, São Tomé and
Principe islands, western Mediterranean, Alboran Sea
(Verhecken, 2007).
Euroscaphella namnetensis Van Dingenen, Ceule-
mans, & Landau, 2014
Plate 3, g. 9
1908 Voluta Lamberti Sow. – Dumas, p.121.
1964 Scaphella lamberti (Sowerby, 1816) – Brébion, p.
517, pl. 13, g. 2 [non Euroscaphella lamberti (J.
Sowerby, 1816)].
1989 Scaphella lamberti (Sowerby, 1816) Lauriat-
Rage et al., p. 131, pl. 8, g. 19 [non Euroscaphel-
la lamberti (J. de C. Sowerby, 1816)].
*2014 Euroscaphella namnetensis Van Dingenen, Ceule-
mans, & Landau, p. 104, pl. 1, gs 1-10.
Material and dimensions Holotype NHMW 2014/
0287/0001 (Pl. 3, g. 9), length 82.3 mm, width 30.2
mm, protoconch diameter 5.2 mm; paratype 1 NHMW
2014/0287/0002, length 96.6 mm, width 31.3 mm (in-
complete: outer lip damaged); paratype 2 NHMW 2014/
0287/0003, length 84.3 mm, width 27.4 mm (incomplete:
outer lip damaged); paratype 3 NHMW 2014/0287/0004,
length 63.3 mm, width 23.6 mm (subadult: outer lip dam-
aged); paratype 4 NHMW 2014/0287/0005, length 46.9
mm, width 17.4 mm (juvenile), NHMW 2014/0287/0006
(11 juveniles); FVD (4 + 50+ subadult and juveniles); LC
(3 + 45 subadult and juveniles), Le Pigeon Blanc, Le Lan-
dreau, Nantes area, Loire-Atlantique department, NW
France.
Discussion Euroscaphella namnetensis Van Dingenen,
Ceulemans, & Landau, 2014 is characterised by its small
size for the genus, its solid, fusiform shell, strongly at-
tened protoconch, short spire, with whorls rapidly in-
creasing in height, in having a slender, weakly shouldered
last whorl, an outer lip with a well-developed palatal cal-
lus, and a short siphonal canal. It is most similar to the
slightly older E. miocenica Fischer & Tournouër, 1879
from the middle Miocene Langhian of the Loire Basin,
which also has a relatively solid shell and a well-devel-
oped palatal callus. It differs, however, in being smaller
in size, in having a more slender last whorl, which is less
shouldered than in E. miocenica and in having the base of
the last whorl hardly constricted, and therefore the sipho-
nal fasciole is not well-developed, as it is in E. miocenica.
For further discussion see Van Dingenen et al. (2014).
Distribution Lower Pliocene: Atlantic, NW France
(Van Dingenen et al., 2014).
Superfamily Cancellarioidea Forbes & Hanley, 1851
Family Cancellariidae Forbes & Hanley, 1851
Subfamily Cancellariinae Forbes & Hanley, 1851
Genus Bivetiella Wenz, 1943
Type species (by original designation of Bivetia Jous-
seaume, 1887b) Cancellaria similis G. B. Sowerby I,
1833, present-day, north western Africa.
1943 Bivetiella Wenz, p. 1356. Replacement name for
40 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
Genus Brocchinia Jousseaume, 1887
Type species (by monotypy) – Voluta mitraeformis Broc-
chi, 1814, Pliocene, Italy.
1887b Brocchinia Jousseaume, p. 221.
Brocchinia pigeonblancensis nov. sp.
Plate 4, gs 2-4
1964 Narona (Brocchinia) mitraeformis Brocchi, 1814
– Brébion (partim), p. 531, pl. 13, gs 12, 13 (non
Voluta mitraeformis Brocchi, 1814, non Lamarck,
1811 = Brocchinia tauroparva Sacco, 1894) (As-
semblage III records only).
Type material Holotype MNHN.F.A57681, height
12.0 mm (specimen illustrated by Brébion, 1964, pl.
13, gs 12, 13; herein Pl. 3, g. 2); paratype 1 NHMW
2015/0133/0388 (Pl. 3, g. 3), height 11.1 mm; paratype
2 NHMW 2015/0133/0414, height 12.2 mm; paratype
3 NHMW 2015/0133/0415, height 10.5 mm; paratype
4 NHMW 2015/0133/0416, height 10.4 mm; paratype 5
NHMW 2015/0133/0389 (juvenile; Pl. 3, g. 4).
Other material Maximum height 12.3 mm. NHMW
2015/0133/0390 (50+); LC (50+), FVD (50+). Le Pigeon
Blanc, Le Landreau, Nantes area, Loire-Atlantique de-
partment, NW France.
Etymology Named after the type locality of Le Pigeon
Blanc. Brocchinia gender feminine.
Locus typicus Le Landreau, Le Pigeon Blanc, Loire-
Atlantique department, NW France.
Stratum typicum – Zanclean, lower Pliocene.
Diagnosis A Brocchinia species of medium size for
genus, with a paucispiral protoconch composed of 1.5-
1.75 whorls, a tall, slender spire, sculpture consisting of
eight broad rounded axial ribs, somewhat nodular mid-
whorl, weakening abapically, obsolete, or almost so, on
last whorl, weak spiral sculpture, and a thin, somewhat
ared outer lip, lirate within.
Description – Shell of medium size for genus, of medium
thickness, fusiform, elongated, with an elevated coni-
cal spire. Protoconch of 1.5-1.75 relatively tall, smooth
whorls, with an impressed suture, nucleus medium-sized.
Junction with teleoconch sharply delimited by proso-
cline scar. Teleoconch of four shouldered whorls, with
steep, concave subsutural ramp, separated by weakly
impressed linear suture. Spiral sculpture weak, begins
at transition zone, consisting of ve narrow spiral cords,
narrower than their interspaces, Abapically number of
cords increases to 9-10 on penultimate whorl. Axial
sculpture starts on rst teleoconch whorl, consisting
of eight broad, prosocline, rounded ribs, obsolete on
subsutural ramp, nodular at shoulder, weakening again
towards abapical suture. Last whorl about 60% total
height. Sculpture on last whorl variable, in most speci-
mens axial ribs strong mid-whorl, in some ribs weaken
on last whorl, subobsolete towards outer lip. Aperture
ovate, about 40% of total height. Outer lip not thickened,
regularly convex, somewhat ared, lirate within. Sipho-
nal canal open, short, wide. Columella bearing two ob-
lique folds of equal strength. Parietal callus thin, sharply
delimited, closely adherent and hardly expanded. Um-
bilicus closed.
Discussion The taxonomy of the European Neogene
Brocchinia species becomes ever more complex. Malat-
esta (1974) noted Voluta mitraeformis Brocchi, 1814 to
be a junior primary homonym of Voluta mitraeformis
Lamarck, 1811 and suggested Cancellaria pusilla H. Ad-
ams, 1869 as a replacement name, although he considered
the Recent and Neogene populations to be separable and
stated that the name Cancellaria cerithiopsis Almera &
Boll, 1887 [sic] was available for the fossil specimens.
Petit (1986b) pointed out that Cancellaria pusilla H.
Adams, 1869 was not available, being a junior primary
homonym of Cancellaria pusilla G.B. Sowerby I, 1832
and that C. cerithiopsis dates from 1898, not 1887. Petit
(1986a) suggested Brocchinia parvula parvula (Beyrich,
1856) for the populations from the Miocene North Sea
Basin and Paratethys and Brocchinia parvula tauroparva
Sacco, 1894 for the Italian Miocene to Pliocene shells.
Davoli (1982, 1995) argued that Brocchi’s name was
available as Brocchi pointed out that his species belonged
to Lamarck’s genus Cancellaria and should be retained
in order to guarantee a stable nomenclature. On p. 645
Brocchi clearly proposed the species Voluta mitrae-
formis’, although in the text under the description he does
say ‘Questa Voluta ha la forma di una Mitra di Lamarck,
ma appartiene non per tanto al genere Cancellaria di
questo naturalista’. Under the ICZN rules this is of no
signicance. Its original introduction as Voluta mitrae-
formis makes it a junior primary homonym and thus una-
vailable. Landau et al. (2006a) pointed out that the genus
was represented in the European Neogene by a group of
related, but distinct species. They highlighted the impor-
tance of the protoconch morphology and showed that the
north western French upper Miocene-Pleistocene forms
were not conspecic with those found in the Mediterra-
nean. Brunetti et al. (2011) reviewed the Italian Pliocene
species. These authors rejected Petit’s (1986a) conclusion
that Brocchinia parvula tauroparva was typical of the
Pliocene Italian forms, as Sacco (1894) based this sub-
species on middle Miocene specimens, which Brunetti et
al. (2011, p. 120) said were different, without clearly stat-
ing what these differences were. These authors argued
that the rst available name for the Pliocene form was
B. mitraeformis depressiplicata Sacco, 1894. However,
they recognised three species in the Italian Pliocene:
B. depressiplicata, B. subanodosa Sacco, 1894 and B.
crassinodosa Sacco, 1894. Apart from teleoconch sculp-
tural details separating these three species, they differ in
the number of protoconch whorls, having 2.0: 2.75: 2.5
Cainozoic Research, 17(1), pp. 23-61, June 2017 41
Mediterranean: B. depressiplicata, B. subanodosa Sac-
co, 1894 and B. crassinodosa Sacco, 1894, one species
occurs in lower Pliocene of north western France: B.
pigeonblancensis and a further, probably undescribed,
species in the Pliocene North Sea Basin of England. One
species occurs in the upper Pliocene-Pleistocene of north
western France: B. rissoiaeformis. Today the genus has
a more restricted distribution in the Atlantic, found only
south of central Portugal.
Brébion (1964, p. 532) recorded Narona (Brocchinia) mi-
traeformis from numerous assemblages in north western
France, but in view of the discussion above we consid-
er only the Assemblage III (Le Pigeon Blanc, Palluau)
records as B. pigeonblancensis.
Distribution – Lower Pliocene: Atlantic, NW France (this
paper).
Genus Trigonostoma Blainville, 1827
Type species (by monotypy) – Delphinula trigonostoma
Lamarck, 1822 [= Trigonostoma scalare (Gmelin, 1791)],
present-day, Indo-Pacic.
1827 Trigonostoma Blainville, p. 652.
Trigonostoma umbilicare (Brocchi, 1814)
Plate 4, g. 5
*1814 Voluta umbilicaris Brocchi, p. 312, pl. 3, gs 10-
11.
non 1874 Cancellaria umbilicaris ? Brocchi – Wood, p. 182,
addendum plate, g. 10. (non Voluta umbilicaris
Brocchi) (= Trigonostoma bellardii De Stefani &
Pantanell, 1879).
non 1878 Cancellaria umbilicaris Brocch. – Nyst, pl. 1, g.
5, pl. 28, g. 8. (non Voluta umbilicaris Brocchi)
(= Trigonostoma bellardii De Stefani & Pantanell,
1879).
non 1882 Cancellaria umbilicaris Brocchi – Nyst, p. 8. (non
Voluta umbilicaris Brocchi) (= Trigonostoma bel-
lardii De Stefani & Pantanell, 1879).
non 1916 Trigonostoma umbilicare (Brocchi) Harmer,
p. 400, pl. 40, gs 3, 4. (non Voluta umbilicaris
Brocchi) (= Trigonostoma bellardii De Stefani &
Pantanell, 1879).
2006a Trigonostoma (Trigonostoma) umbilicare (Broc-
chi, 1814) – Landau et al., p. 68, pl. 3, g. 1 (cum
syn.).
2009 Trigonostoma umbilicare (Brocchi, 1814) Bru-
netti et al., p. 112, gs 16A-D.
2011 Trigonostoma umbilicare (Brocchi, 1814) – Land-
au et al., p. 30, pl. 15, g. 12.
Material and dimensions Height 34.3 mm. NHMW
2015/0133/0413 (1 incomplete; Pl. 4, g. 5), LC (1 in-
complete, 5 fragments). Le Pigeon Blanc, Le Landreau,
Nantes area, Loire-Atlantique department, NW France.
whorls respectively.
The revision by Brunetti et al. (2011) conrms the state-
ment made by Landau et al. (2006a) that the north west-
ern French Brocchinia species were not conspecic with
those found in the Italian Pliocene. When this statement
was made, the junior author (BL) was comparing mate-
rial from the Assemblage I locality of Sceaux d’Anjou.
This is the material described by Millet (1854, p. 160)
as Cancellaria auriculoides. However, on examination of
the plentiful material at hand from Assemblage I locali-
ties of Sceaux d’Anjou, Renauleau and St-Clément-de-
la-Place (NHMW coll.), the Le Pigeon Blanc specimens
differ in being slightly larger, but thinner shelled, with a
higher spire and in having the axial ribs much broader
and somewhat nodular mid-whorl. In most specimens
of Brocchinia pigeonblancensis nov. sp. the axial ribs
continue relatively strongly at least onto the rst half of
the last whorl, whereas in B. auriculoides the axial ribs
weaken much earlier on the second or third teleoconch
whorl. The spiral sculpture is weak in both species, but
even more so in B. pigeonblancensis. We note that there
might be a second Brocchinia species in the Assemblage
I material, which will be explored in a separate paper.
Cossmann (1899, p. 20) described B. rissoiaeformis from
the Upper Pliocene-Pleistocene Assemblage IV locality
of Gourbesville. The holotype is poorly preserved, but
the specimen illustrated by Brébion (1964, p. 532, pl. 13,
g. 14) from the type locality shows a rather small (height
7 mm), squat species, with a protoconch composed of two
whorls, about 12 spiral cords and no axial sculpture.
Harmer (1918, p. 396, pl. 39, gs 40, 41) described and
illustrated specimens of Brocchinia from the Coralline
and Red Crags of eastern England. Unfortunately he did
not describe the protoconch type. However, two speci-
mens at hand from the lower Pliocene Coralline Crag of
Gedgrave (NHMW coll.) have a paucispiral protoconch
similar to the shells from Le Pigeon Blanc, but the Eng-
lish specimens are larger and more solid than the French
shells, the spiral sculpture is coarser, the axial sculpture
is weaker so that the last whorl is devoid of ribs, and the
aperture is more dilated, with stronger lirations within
the outer lip. We provisionally consider them distinct.
In the Italian Pliocene B. pigeonblancensis is most like B.
depressiplicata, but differs in having a paucispiral proto-
conch of only 1.5-1.75 whorls instead of two and in hav-
ing fewer axial ribs (8 vs. 10-11), which are broader and
more prominent mid-whorl, giving the whorls a shoul-
dered appearance. Brocchinia crassinodosa Sacco, 1894,
apart from having a multispiral protoconch composed of
2.5 whorls, has even fewer (7 vs. 8), more elevated ribs
than B. pigeonblancensis, which extend between the su-
tures, as this species does not have a distinct subsutural
ramp.
In the extant faunas the eastern Atlantic Brocchinia
clenchi Petit, 1986, found off Portugal, Selvagem Grande,
Canary Islands, Azores, and Western Sahara (Verhecken,
2007), also has a paucispiral protoconch of 1.5 whorls,
but differs in having fewer, wider spaced spiral cords and
weaker axial ribs.
In summary, in the Pliocene three species occur in the
42 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogast ropoda (in part)
Discussion – This species is represented by two large in-
complete specimens from Le Pigeon Blanc. Only part of
the spire and the outer lip are preserved, but they are typi-
cal for this rather large Trigonostoma species, which until
now was known only from the Pliocene Mediterranean
and adjacent Atlantic. This is the most northern Pliocene
record for the species. The North Sea Basin records of T.
umbilicare (Wood, 1874; Nyst, 1878, 1882; Harmer, 1916)
represent T. bellardii De Stefani & Pantanell, 1879.
Distribution – Lower Pliocene: Atlantic, NW France (this
paper), Guadalquivir Basin, Spain (Landau et al., 2011);
central Mediterranean, Italy (Sacco, 1894; Chirli, 2002;
Brunetti et al., 2009). Upper Pliocene: western Mediter-
ranean, Estepona Basin, Spain (Vera-Peláez et al., 1995;
Landau et al., 2006a); central Mediterranean, Italy (Sac-
co, 1894; Glibert, 1960; Cavallo & Repetto, 1992).
Trigonostoma sp.
Plate 4, g. 6
Material and dimensions Height 16.2 mm, width 10.5
mm. NHMW 2015/0133/0412 (1; Pl. 4, g. 6). Le Pigeon
Blanc, Le Landreau, Nantes area, Loire-Atlantique de-
partment, NW France.
Description Shell small to medium sized, solid, fusi-
form with scalate spire. Protoconch dome-shaped, con-
sists of three convex whorls. Teleoconch of four barrel-
shaped whorls, with moderately broad, attened, concave
sutural gutter. Suture supercial, undulating. Axial
sculpture consisting of eight relatively broad, elevated,
rounded, prosocline ribs, forming broad nodes at shoul-
der. Ribs cross sutural gutter, narrowing towards adapi-
cal suture, dividing gutter into large pits. Spiral sculpture
overrides axial ribs, consists of six narrow primary cords
below shoulder, with secondary and tertiary threads in-
tercalated in interspaces on later whorls. Spiral sculpture
most evident where it crosses ribs, subobsolete between
ribs on later whorls. Last whorl moderately inated,
with deeply concave sutural gutter, convex below shoul-
der, with ten primary spiral cords, basal cord delimiting
deep, moderately wide umbilicus, smooth within, except
inconspicuous growth lines. Aperture subtrigonal, ap-
proximately 40% of the total height, Outer lip thickened
by labial varix, lirate within, with small siphonal spout
abapically. Columella shallowly excavated, bearing three
oblique columellar folds, weakening abapically. Colu-
mellar callus thickened, erect, forming medial border of
umbilicus, parietal callus sharply delimited, thickened,
weakly expanded.
Discussion One well preserved specimen of a Trigo-
nostoma Blainville, 1827 species may represent an un-
described species. It is most similar to Pliocene Atlantic
and Mediterranean and T. bellardii De Stefani & Panta-
nell, 1879, but most specimens of this species have more
numerous axial ribs, which are narrower, less elevated
and spinous at the shoulder, the spiral sculpture, although
also weak, is stronger than in the shell from Le Pigeon
Blanc and does not weaken in the interspaces between
the ribs as it does in the French shell, and the umbilicus
is wider. Trigonostoma parvotriangula Sacco, 1894 from
the Pliocene Mediterranean has similar axial sculpture
composed of broad elevated ribs that form tubercles at the
shoulder, but this species has a broader shell shape and a
much wider umbilicus, which bears sculpture within. We
await further material to better characterise this species.
Distribution – Lower Pliocene: Atlantic, NW France (this
paper).
Subfamily Admetinae Troschel, 1865
Genus Pseudobabylonella Brunetti, Della Bella, Forli &
Vecchi, 2009
Type species (by original desiganation) Cancellaria
minima Reeve, 1856, present-day, West Africa.
2009 Pseudobabylonella Brunetti, Della Bella, Forli &
Vecchi, p. 65.
Pseudobabylonella fusiformis (Cantraine, 1835)
Plate 4, g. 7
*1835 Cancellaria fusiformis Cantraine, p. 391.
2006a Babylonella fusiformis (Cantraine, 1835) Lan-
dau et al., p. 86, pl. 9, gs 1-4 (cum syn.).
2008 Babylonella costillifera [sic] (Sowerby, 1818)
Chirli & Richard, p. 53, pl. 10, g. 7.
2009 Pseudobabylonella fusiformis (Cantraine, 1835) –
Brunetti et al., p. 66, gs 7B-L, 8A-D.
2009 Pseudobabylonella subangulosa (Wood, 1848)
Brunetti et al., p. 69, gs 9A-F, 12B.
2009 Pseudobabylonella aplicata Brunetti et al., p. 70,
gs 10C-H, 11A-E, 12A.
2010 Pseudobabylonella aplicata Brunetti, Della Bella,
Forli & Vecchi, 2009 Sosso & Dell’Angelo, p.
43, p. 60 unnumbered g. top row right.
2010 Pseudobabylonella fusiformis (Cantraine, 1835)
– Sosso & Dell’Angelo, p. 43, p. 60 unnumbered
g. middle row left.
Material and dimensions Height 8.7 mm. NHMW
2015/0133/0391 (1; Pl. 4, g. 7), LC (4). Le Pigeon Blanc,
Le Landreau, Nantes area, Loire-Atlantique department,
NW France.
Discussion – Landau et al. (2006a) and Verhecken (2007)
highlighted how uncomfortably Cancellaria fusiformis
Cantraine, 1835 and Cancellaria minima Reeve, 1856 t-
ted within the cancellariid generic groups recognised at
the time. Brunetti et al. (2009) erected a new supraspe-
cic taxon for these species: Pseudobabylonella, a solu-
tion with which we agree. The most outstanding charac-
ter of the genus is the ne cancellate sculpture covering
the protoconch.
Cainozoic Research, 17(1), pp. 23-61, June 2017 43
In the same paper these authors revised the representa-
tives of their new genus in the Plio/Pleistocene of Italy
and recognised three species: P. fusiformis, P. subangu-
losa (Wood, 1848) and P. aplicata Brunetti, Della Bella,
Forli & Vecchi, 2009 differing in shell prole and sculp-
tural detail. The variability of these species is tabulated
in their paper (Brunetti et al., 2009, p. 78).
In our opinion the authors are over-splitting. The reasons
for this conclusion are: 1) All three have exactly the same
protoconch. 2) All three have the same stratigraphic dis-
tribution (Brunetti et al., 2009, p. 79, tab. 3). 3). All three
species are widely distributed geographically, based on
the synonymy given by the authors (i.e. all three species
occur in the North Sea Basin and Mediterranean (the
record of B. fusiformis subangulosa (Wood) given by
Marquet (1998) is included in the synonymy of both P.
fusiformis and P. subangulata with a question mark). 4)
In their description, P. aplicata has ve cords on the sub-
sutural ramp and six cords from the shoulder to the abapi-
cal suture (2009, p. 73). The authors do not distinguish
primary from secondary cords. The development of sec-
ondary spiral sculpture in many cancellariids is variable,
and it is clear that their gs 11A-E have four primary
cords below the shoulder (including shoulder cord), with
a secondary thread developed in some of the interspaces
(compare their gs 11A, B vs. C, D). This is even more
clearly demonstrated by the two specimens illustrated
by Landau et al. (2006a, pl. 9, gs 2, 3) from the upper
Pliocene Estepona Basin of Spain, which where both syn-
onymised with P. aplicata by Brunetti et al. (2009). 5)
The present-day West African representative of the genus
B. minima (Reeve, 1856) differs in having a paucispiral
protoconch. The interspecic variability was illustrated
by Verhecken (2007, p. 293, gs 9A-F). The variabil-
ity in shell prole and sculpture mirrors that seen in its
Neogene ancestor P. fusiformis. Brunetti et al. (2009, p.
66) countered this argument by saying that these gures
probably represented various different species.
If we consider the shell characters of the specimen from
Le Pigeon Blanc, the elongated, slender shell prole is
that of P. aplicata, however, the whorls are regularly
rounded, as the shoulder cord is not outstanding. On the
penultimate whorl there are four primary cords from the
shoulder, with a single secondary thread intercalated
in the interspaces. This would t within the diagnostic
criteria for P. aplicata. However, there are 16 axial ribs,
which is considerably more than the 10-12 recognised by
Brunetti et al. (2009, p. 78) for P. aplicata. The proto-
conch is identical to that of the rest of the group (Pl. 4,
g. 7c).
At present we do not nd it useful to erect yet another spe-
cic taxon within Pseudobabylonella for the Le Pigeon
Blanc specimens. We prefer to consider P. fusiformis a
stratigraphically and geologically widespread species
characterised by its multispiral protoconch covered in
cancellate microsculpture and teleoconch with cancellate
sculpture composed of a variable number of axial and
spiral elements. This is probably the predecessor of the
present-day B. minima, which has a similarly sculptured
paucispiral protoconch and variable teleoconch sculp-
ture. If future genetic testing of B. minima should show it
to represent a species complex rather than a single taxon,
the above discussion could be reviewed.
Distribution Middle Miocene: North Sea Basin, Ger-
many (Kautsky, 1925; Glibert, 1960; Anderson, 1964;
Wien rich, 2001), Denmark (Ravn, 1907; Sorgenfrei, 1958;
Rasmussen, 1956, 1968), Netherlands (Glibert, 1960; Nord-
sieck, 1972; Janssen, 1984a & b), Belgium (Glibert, 1952b);
Atlantic, Aquitaine Basin, France (Cossmann, 1899; Pey-
rot, 1928). Upper Miocene: Proto-Mediterranean, Italy
(Sacco, 1894; Davoli, 1982, 1995). Lower Pliocene: At-
lantic: NW France (this paper); North Sea basin, Eng-
land (Wood, 1848; Harmer, 1916), Belgium (Marquet,
1997, 1998); central Mediterranean, Italy (Sacco, 1894;
Pavia, 1975; Chirli, 2002; Brunetti et al., 2009). Upper
Pliocene: western Mediterranean, Estepona Basin, Spain
(Landau et al., 2006a), Alpes Maritimes (Chirli & Rich-
ard, 2008); central Mediterranean, Italy (Sacco, 1894;
Cavallo & Repetto, 1992; Brunetti et al., 2009; Sosso &
Dell’Angelo, 2010).
Pseudobabylonella? sp.
Plate 4, g. 8
1964 Narona (Sveltia) elongata Brébion, p. 528, p. 13,
g. 8 (nomen nudum).
Material and dimensions Height 11.3 mm. NHMW
2015/0133/0392 (1: Pl. 4, g. 8). Le Pigeon Blanc, Le Lan-
dreau, Nantes area, Loire-Atlantique department, NW
France.
Description Shell small, moderately fragile, slender
fusiform. Protoconch and early teleoconch whorls miss-
ing. Five relatively tall, regularly convex teleoconch
whorls preserved. Sculpture of narrow orthocline axial
ribs, 15 on penultimate whorl, crossed by seven narrow
spiral cords, slightly narrower and more crowded on sub-
sutural ramp, shoulder cord slightly stronger, subequal
below shoulder. Last whorl elongate with 18 axial ribs,
secondary spiral threads intercalated between primaries
mid-whorl, base weakly constricted. Aperture ovate, outer
lip damaged, smooth within. Columella weakly excavated,
bearing two narrow oblique folds. Columella callus hardly
developed, poorly delimited; parietal callus not developed.
Discussion – This is undoubtedly the same species de-
scribed by Brébion (1964, p. 528, p. 13, g. 8) as Narona
(Sveltia) elongata Brébion (nomen nudum). However, we
are uncertain of the generic placement. Species included
in the genus Sveltia Jousseaume, 1887 are larger, more
solid, and have a better developed inner lip callus. The
slender, rather delicate shell, and weak inner lip callus are
reminiscent of Pseudobabylonella Brunetti, Della Bella,
Forli & Vecchi, 2009, but we would need to see a proto-
conch for the characteristic reticulated microsculpture to
be certain. We await more material to better characterise
this species.
44 Van Dingenen, Ceulemans & Landau. The lower Pliocene gastropods of Le Pigeon Blanc, 4. Neogastropoda (in part)
Brébion (1964, p. 528) recorded it from Assemblage III
(Le Pigeon Blanc) only.
Discussion
In this paper we record 26 neogastropod species, of
which four are left in open nomenclature, representing
17 genera. Three species are described as new: Euthria
palumbina nov. sp., Bartschia (Agassitula) harasewychi
nov. sp. and Brocchinia pigeonblancensis nov. sp. Just
under half of the species (46%) occur exclusively in
north western French Assemblage I-III deposits and are
therefore restricted stratigraphically and geographically.
Stratigraphically (see Text-g. 2), eight (30%) of the spe-
cies found at Le Pigeon Blanc are found in the middle
Miocene Langhian of the Loire Basin (see Glibert, 1949).
Six (23%) are also present in the Assemblage I (sensu Van
Dingenen et al., 2015) of north western France. Fourteen
(53%) are also recorded in the Miocene to present-day
Mediterranean and only one species is shared with the
North Sea Basin and one with the Atlantic coasts of the
British Isles.
As with previous parts of this series (Ceulemans et al.,
2016; Van Dingenen et al., 2016), an important number
of genera such as Aplus, Euthria, Aptyxis, Bivetiella and
Trigonostoma are still found in European waters, but with
a more southern distribution, again suggesting that aver-
age Sea Surface Temperatures were higher than they are
at these latitudes today, possibly similar to those found
off the southern Portuguese coasts at present time.
Acknowledgements
We would like to thank Jean-Michel Pacaud of the
MNHN (Paris) for making Brébion’s collection available
to us, also to Jocelyn Falconnet (MNHN, Paris) for tak-
ing photographs of the Brébion material in Paris. To M.G.
(Jerry) Harasewych, Research Zoologist at the Smithso-
nian Institution, Washington, DC, USA for some taxo-
nomic advice. Carlos Marques da Silva of the University
of Lisbon, Portugal, for his advice and help with graphics.
Thanks also to the referees: Ingemann Schnetler (Stevn-
strup, Denmark) and Robert Marquet (Antwerp, Belgium)
for their helpful comments. Special thanks should be
given to the families Provost (Le Pigeon Blanc, Le Lan-
dreau, France) and Madeleineau (L’Errière, Le Landreau,
France) for allowing us to excavate on their properties,
without them this publication would not have been pos-
sible.
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