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Cainozoic Research, 17(2), pp. 75-166, December 2017 75
The upper Miocene gastropods of northwestern France, 1.
Patellogastropoda and Vetigastropoda
Bernard M. Landau1, 4, Frank Van Dingenen2 & Luc Ceulemans3
1 Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The 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-8200-261 Albufeira, Portugal; email: bernardmlandau@gmail.com
2 Cambeenboslaan A 11, B-2960 Brecht, Belgium; email: fvd@telenet.be
3 Avenue Général Naessens de Loncin 1, B-1330 Rixensart, Belgium; email: luc.ceulem@skynet.be
4 Corresponding author.
Received 7 July 2017, revised version accepted 9 September 2017
Seventy-six species are recorded of which 26 are new to science: Diodora sancticlementensis nov. sp., Sinezona geigeri nov.
sp., Anatoma redoniana nov. sp., Clanculus (Clanculus) brebioni nov. sp., Clanculus (Clanculopsis) sancticlementensis nov. sp.,
Clanculus (Clanculopsis) umbilicovadus nov. sp., Jujubinus coronatus nov. sp., Jujubinus redoniensis nov. sp., Jujubinus sceauxensis
nov. sp., Gibbula brebioni nov. sp., Gibbula clanculiforma nov. sp., Gibbula conicomagus nov. sp., Gibbula marianae nov. sp.,
Phorcus gallicophorcus nov. sp., Calliostoma gibbuliforme nov. sp., Calliostoma lamellatum nov. sp., Calliostoma michaeli nov. sp.,
Calliostoma microgemmatum nov. sp., Calliostoma quaggaoides nov. sp., Calliostoma presselierense nov. sp., Calliostoma spinosum
nov. sp., Calliostoma verrucosum nov. sp., Thysanodonta chauvereauensis nov. sp., Pareuchelus dautzenbergi nov. sp., Dikoleps
insulsa nov. sp., Lodderena redferni nov. sp., Parviturbo rubioi nov. sp., Pseudorbis beugnonensis nov. sp., Skenea minuticostata nov.
sp. and Skenea wareni nov. sp.
We highlight the rst description of a member of the subfamily Thysanodontinae in the Europe faunas. Six are homonyms and receive
replacement names: Trochus heliciformis Millet, 1865 is a junior homonym of T. heliciformis von Zieten, 1832, and is renamed
Calliostoma biangulatum nov. nom. Trochus millegranus Millet, 1864, is a is a junior homonym of T. millegranus Philippi, 1836, and
is renamed Callisotoma milletigranum nov. nom. Trochus torulosus Millet, 1865 is a junior homonym of T. torulosus Philippi, 1845,
and is renamed Calliostoma miotorulosum nov. nom. Trochus tumidus Millet, 1865 is a junior homonym of T. tumidus Montagu, 1803,
and is renamed Calliostoma miotumidum nov. nom. Trochus echinatus Millet, 1865 is placed in the genus Calliostoma, rendering C.
echinatum Dall, 1881 a secondary homonym, which is renamed C. caribbechinatum nom. nov.
The following are new subjective synonyms: Emarginula dujardini Dollfus & Dautzenberg, 1886 is considered a junior subjective
synonym of Emarginula imbricata Millet, 1865. Gibbula (Colliculus) sosensis Cossmann & Peyrot, 1917 is considered a junior
subjective synonym of Trochus insignis Millet, 1854, Monodonta (Oxystele) amedei turoniensis Glibert, 1949 is considered a junior
subjective synonym of Pitonellus trochiformis Millet, 1865.
This is the most speciose European Neogene Vetigastropod assemblage known, with 31 different genera/subgenera represented. The
fauna is highly endemic, with 63% of the species known only from the northwestern French Assemblage I localities. The generic
composition is similar to that seen at these latitudes today; a few genera are extinct, such as Lucapinella, Paroxystele and Pareuchelus.
Few thermophilic elements are present. The composition of the assemblage represents a hard-bottomed littoral fauna.
Key words: northwestern France, upper Miocene, Gastropoda, new taxa
Introduction
The rich fossil gastropod assemblages of northwestern
France have been the object of study by our group for sev-
eral years. To date only the lower Pliocene Assemblage
III fauna (of Van Dingenen et al., 2015) has been studied
in part, and a systematic account of their content given by
Van Dingenen et al. (2015; Nassariidae), Van Dingenen
et al. (2016; Caenogastropoda), Ceulemans et al. (2016a;
Patellogastropoda and Vetigastropoda), Ceulemans et al.
(2016b; Muricidae) and Van Dingenen et al. (2017; Neo-
gastropoda, in part). Our aim is to mirror these papers
with revisions of the equivalent taxonomic groups in
the upper Miocene Tortonian Assemblage I deposits. In
this paper we cover the same groups as those covered by
Ceulemans et al. (2016a), the Patellogastropoda and Veti-
gastropoda. In order to avoid repetition of generic and
species synonymies and discussions, we frequently refer
back to that paper, which should be used in conjunction.
It is with some trepidation that we commence the sys-
tematic taxonomic revision of the gastropods in what
was previous known as the ‘Redonien inférieur’ (Bré-
76 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
bion, 1964) or ‘Redonien chaud’ (Lauriat-Rage, 1981), as
this is the fourth attempt to do so (Landau et al., 2016).
Millet (1854, 1865) erected many new species described
from material from northwestern France. He prepared
four lithographed plates meant to accompany his ‘Pa-
léontologie de Maine-et-Loire’ (Millet, 1854). However,
these plates were never published, as he decided to pre-
pare plates with photographed specimens. Millet’s plates
were found by chance by Couffon (1909) and have subse-
quently been lost. Couffon (1915) was the next to attempt
the task. He wrote ‘Nous avons d’ailleurs entrepris une
Iconographie du Miocène supérieur, en Anjou, qui ne
comprendra moins de 40 planches, in 4°; les maquettes
des première 25 planches sont déjà prêtes pour la pho-
totypie et les événements actuels ont seuls empêché leur
apparition’. These ‘current events’ most likely refer to
World War I, and this work was never completed. The
third effort was that of Brébion (1964), who prepared
his thesis on the ‘Redonian’ gastropods, describing and
erecting many new species. Until now, this was the only
work illustrating the fauna and has become the ‘bible’ of
the ‘Redonian’. Unfortunately, Brébion never published
his thesis and therefore all the taxa erected are nomina
nuda. This is, therefore, the fourth attempt to produce an
illustrated systematic account of the gastropods.
Geological setting
The localities studied herein of La Presselière (Sceaux-
d’Anjou), Le Grand Chauvereau (Saint-Clément-de-la-
Place), Renauleau and Beugnon (Doué-en-Anjou) (Text-
g. 1) were included in the ‘Redonien’ of Brébion (1964)
and are localities in the ‘Redonien chaud’ of Lauriat-Rage
(1981) in the Maine-et-Loire department. The numerous
problems surrounding the term ‘Redonien’ were high-
lighted by Van Dingenen et al. (2015), who concluded
that it was best to restrict the term to the ‘Redonien Stra-
totypique’ of Néraudeau et al. (2003). Van Dingenen et
al. (2015) recognised four major fossil assemblages in the
post mid-Miocene of northwestern France, characterised
by distinct gastropod faunas and designated them as As-
semblages I – IV. Van Dingenen et al. (2015) correlated
the stratigraphic sequences these asemblages occurred in
with the upper Miocene to lower Pleistocene formations
of Cornwall and East Anglia in England, and Belgium
(2015, p. 78, g. 2). The localities studied here are the As-
semblage I localities of Van Dingenen et al. (2015).
Le Grand Chauvereau (Saint-Clément-de-la-Place,
Maine-et-Loire)
A deposit occurring along a 3 km wide irregular area,
composed of ‘Redonian’ sands, marls and clays, rang-
ing from La Poissardière in the NW to La Bellangerie
in the SE. Most of it overlies the ‘Orthogneiss de Saint-
Clément-de-la-Place’, a deposit outcropping along a 10
km long area oriented WNW-ESE. The most southern
part overlies the ‘Granite de Bécon et de Saint-Lam-
bert-la-Potherie’, outcropping along a 9 km long and 2
km area E-W oriented. This granite intrudes the ortho-
gneissic basement of Saint-Clément-de-la-Place. Multi-
ple NNE-SSW oriented microgranite veins, up to 10 m
in thickness, cut through both these geological units. At
Le Grand Chauvereau the ‘Chauvereau-Naizance’ vein,
in the form of solid rock and individual boulders, is ex-
posed in a limited area on the shore of the pond, indi-
rectly dening the pond contours. We can assume that
this microgranite vein was part of the local Tortonian pal-
aeogeography (Cavet et al., 1976; Janjou et al., 1998; and
second author’s personal observations).
In this ‘Redonian’ deposit, several wide fossiliferous lens-
es are present, of which Le Grand Chauvereau is the best
known, consisting of ne sands and clayey sands with
small to medium-sized pebbles. The fossiliferous layer
occurs 3-4 m below the surface (depending on the loca-
tion), approximately 1 m thick. Within this fossiliferous
lense a 20-30 cm whitish homogeneous highly fossilifer-
ous sandy layer occurs. Mollusc fossils are accompanied
by enormous numbers of bryozoans, scattered corals and
sh teeth. The molluscs at St-Clément-de-la-Place are the
most diverse and best preserved. We therefore consider
this the ‘type’ locality for Assemblage I. The gastropod
fossils are almost all small in size, with very few shells
attaining more than 20 mm in height. In contrast, the bi-
valve fossils show a much greater size range, with some
relatively large specimens found with articulated valves,
suggesting a low degree of post-mortem transport. Of
note at St-Clément-de-la-Place is the assemblage of Poly-
placophora (chitons), which is probably the most abun-
dant and diverse in the European Neogene and will be
described in a separate paper (Dell’Angelo et al. in prep.).
La Presselière (Sceaux-d’Anjou, Maine-et-Loire)
The famous 100 m wide fossiliferous outcrop is sur-
rounded by colluvions and alluvial deposits, in the NW
by Plio-Quaternary silt and residual (or reworked ?)
sandy gravel formations (Brossé et al., 1989).
The deposit consists of ne sands with small to medium-
sized pebbles. The fossiliferous layer occurs 1-1.5 m be-
low the surface, at some places showing a layer of clay.
The section continues down for 1.5 m containing fossils
which are frequently agglutinated to irregular sandy con-
cretions, increasing in size towards the base. The assem-
blage is similar to that of St-Clément-de-la-Place, with
small gastropod shells strongly predominant, but the fos-
sils are somewhat worn, suggesting some degree of post-
mortem transport.
Renauleau (Doué-en-Anjou, Maine-et-Loire)
The limited exposure at Renauleau and Beugnon (900 x
500 m) is located north of the village Brigné-sur-Layon.
The site of Renauleau is located 60 m NE of the ‘Fontaine
de Renauleau’. At this locality the Tortonian overlies the
middle Cenomanian, represented by black clays. A pos-
Cainozoic Research, 17(2), pp. 75-166, December 2017 77
sible source for the older Cretaceous reworked fossil bra-
chiopods, oysters and rare shark teeth found mixed wih
the Miocene fossils is the upper Cenomanian ‘Marnes à
Ostracées‘, which occurs 250 m NE of the site (Blaise et
al., 1986).
The Tortonian deposits consist of ne sands and clayey
sands, with numerous small to medium-sized pebbles
and clay nodules. The fossiliferous layer occurs 6.5 m
below the surface, approximately 3 m thick and covered
by a 30-40 cm sandstone layer, gently dipping towards
the south. In the upper part of this lense, white-bluish
pockets with higher shell concentrations occur. At this
deposit more fragments of larger shells such as muricids
and coniids may be found, but invariably fragmented and
showing signs of transport. In contrast large articulated
bivalves occur in living position between 1.5 and 2 m
below the sandstone. As with Sceaux-d’Anjou, the shells
are usually somewhat worn.
Beugnon (Doué-en-Anjou, Maine-et-Loire)
The southern section of the Beugnon-Renauleau depos-
it overlies the ‘Complexe de Saint-Georges-sur-Loire’
(schist-sandstone and volcanic Upper Ordovician to
Lower Devonian).
Named after the ‘Fontaine du Beugnon’ located 250 m
SSW of the outcrop, the abandoned Beugnon quarry ex-
poses at least 4 m of brownish consolidated calcareous
beds, alternating with ne white sandy layers (Blaise et
al., 1986).
The gastropod assemblage in these sandy beds is similar
to that of Renauleau, although some species are found in
only one or other of the localities.
Rectications/clarications regarding the Beugnon-
Renauleau area
1. Brébion used in his unpublished thesis the misspelled
locality name ‘Reneauleau’ instead of Renauleau. The
spelling was probably adopted following Millet (1854),
who also used the spelling ‘Reneauleau’. This spelling
error has long been overlooked, although it might have
been intentional. Consulting historical maps we have
found that the map of Cassini, the rst general map of the
French territory, published from 1756 to 1815, uses the
spelling ‘Reneauleau’. The ‘Carte de l’état major’, pro-
duced between 1817 and 1866 on the orders of Napoleon
I, is the rst using the modern spelling Renauleau. All
subsequent maps and administrative entries to date retain
this later spelling.
2. The rst author has found during his work in the
Naturalis collection, labels that indicate the location
‘Brigneau’. Fortunately GPS coordinates were known.
After these were plotted, it turned out that they corre-
spond exactly to the site of Beugnon, named after the
nearest landmark, the ‘Fontaine du Beugnon’. The most
plausible answer is that researchers who were unaware
Figure 1. Geographic location of the localities sampled. 1. La Presselière, Sceaux-d’Anjou; 2. Le Grand Chauvereau, St-Clément-
de-la-Place; 3. Renauleau; 4. Beugnon, Maine-et-Loire, France.
78 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of nor thwestern France, 1. Patellogastropoda and Vetigastropoda
of the fontaine, named the quarry after the nearest land-
mark known to them: Brigneau, a small suburb north of
the village of Brigné-sur-Layon. To avoid confusion, we
continue to use Beugnon.
3. The former commune of Brigné-sur-Layon was merged
this year into the newly formed commune Doué-en-An-
jou. In order to follow the current municipal administra-
tion we will from now on only refer to Doué-en-Anjou.
Other Assemblage I localities are:
• Thorigné-d’Anjou (Maine-et-Loire),
• Contigné (Maine-et-Loire),
• Les Pierres Blanches and St. Anne (Chalonnes-sur-
Loire, Maine-et-Loire),
• Saint-Michel-et-Chanveaux (Maine-et-Loire),
• Beaulieu (Mayenne) (Brébion, 1964, p. 15).
Material and methods
Large collections of northwestern French material are
present in the Naturalis Biodiverity Center in Leiden
(The Netherlands), the accumulation of several important
collections collected by A.W. Janssen (1987) and kindly
donated by W. Groeneveld (2004), C. Deerenberg (2010),
F.A.D. van Nieulande (2012), H.P.J. Keukelaar (2015) and
P. Hessel (2015). These were consulted and the material
catalogued herein. We note that the locality given in the
RGM collections of Brigneau or Brigné are more cor-
rectly here called Beugnon. These historical collections
were augmented by new collections accumulated by the
authors.
It is intended that this series of monographs be used
in conjunction with those published by the authors in
Cainozoic Research (i.e. Ceulemans et al., 2016a, b; Van
Ding enen et al., 2015, 2016, 2017) describing the strati-
graphically younger Assemblage III fauna of northwest-
ern France. Where the generic synonymy and species
chresonymy were covered in the Assemblage III series,
they are not repeated here. In the chresonymy we have in-
cluded the works of Millet (1854, 1864, 1865), but in most
cases have excluded the records given by Couffon (i.e.
1909, 1915), as these are species lists without descrip-
tions or illustrations and are not possible to verify. In the
Assemblage III series we used Brébion’s specimens as
holotypes wherever possible. This has not been followed
in this Assemblage I series for practical reasons. Almost
all the species are small to minute. In order to describe
them adequately we have chosen the best specimen as
holotype, often selected from numerous specimens and
almost invariably better preserved than the specimens
gured by Brébion (1964) in his unpublished thesis, in
which the specimens were photographed, but no light or
scanning microscopy was performed.
Type material was deposited in the Muséum national
d’Histoire naturelle, Paris (collection de Paléontologie),
the Naturhistorisches Museum Wien, Vienna, Aus-
tria and the Naturalis Biodiversity Center, Leiden, The
Netherlands. Further material is present in the personal
collections of Luc Ceulemans, Rixensart (Belgium) and
Frank Van Dingenen, Brecht (Belgium).
Abbreviations:
FVD Frank Van Dingenen private collection (Brecht,
Belgium).
LC Luc Ceulemans private collection (Rixensart,
Belgium).
MHNN.P Muséum d’Histoire naturelle de Nantes, Pal-
aeontology collection (Nantes, France).
MNHN.F Muséum national d’Histoire naturelle, collec-
tion de Paléontologie (Paris, France).
NHMW Naturhistorisches Museum Wien collection
(Vienna, Austria).
RGM Naturalis Biodiversity Center, collection Cain-
ozoic Mollusca (Leiden, The Netherlands).
Systematic palaeontology
Subclass Patellogastropoda
Superfamily Patelloidea Ranesque, 1815
Family Patellidae Ranesque, 1815
Genus Patella Linnaeus, 1758
1758 Patella Linnaeus, p. 780. Type species (by sub-
sequent designation, Fleming, 1818): Patella vul-
gata Linnaeus, 1758, present-day, Europe.
For generic synonymy see Ceulemans et al. (2016a, p. 53).
Note – Based on molecular data, Koufopanou et al. (1999)
showed that the typical European Patella species formed
a monophyletic clade, within which there was little mo-
lecular divergence. Moreover, their analysis conrmed
that Ansates pellucidum (Linnaeus, 1758) belonged in the
clade Patella s.s., as suggested by Ridgway et al. (1998).
Therefore Ansates G.B. Sowerby II, 1839 and Patellastra
Monterosato, 1884a, b, used in Landau et al. (2003), are
synonymised with Patella Linnaeus, 1758.
Patella protea Doderlein, 1862
Plate 1, gs 1-3.
1854 Patella Alternata Millet, p. 166 (nomen nudum
and junior homonym of P. alternata Say, 1826).
*1862 Patella protea Doderlein, p. 98.
1865 Patella alternata Millet, p. 599 (junior homonym
of P. alternata Say, 1826).
1896b Patella protea Dod. – Sacco, p. 23, pl. 2, gs 87-
88.
1964 Patella protea Doderlein, 1862 – Brébion, p. 66,
pl. 1, gs 13, 14.
Material and dimensions – Maximum diameter 12.6
Cainozoic Research, 17(2), pp. 75-166, December 2017 79
mm; height 3.1 mm. St-Clément-de-la-Place: NHMW
2016/0103/0001-0003 (3), NHMW 2016/0103/0004 (50+),
RGM.1309474 (50+), RGM.1309475 (6), RGM.1309493
(50+), LC (50+), FVD (50+). Sceaux-d’Anjou: NHMW
2016/0103/0645 (2), RGM.1348018 (1).
Discussion – Brébion (1964, pl. 1, gs 13, 14) illustrated
a small patellid from Assemblage I localities (Sceaux-
d’Anjou, St-Michel, Contigné) as P. protea Doderlein,
1862, which was originally described from the upper
Miocene Tortonian of Italy and said to be common at
Montegibbio. Numerous specimens at hand from the As-
semblage I locality of St-Clément-de-la-Place resemble
tiny specimens of P. caerulea, but differ in having an an-
terior primary rib placed at six o’clock, when the dorsum
is viewed on its antero-posterior axis, posterior border
placed at the top, whereas in P. caerulea the anterior ribs
are placed at ve o’clock and seven o’clock. They are also
considerably smaller: according to Brébion 10-15 mm di-
ameter vs. 20-50 mm diameter for present-day P. caeru-
la. The specimens at hand from St-Clément-de-la-Place
are smaller, similar in size to specimens of P. protea from
the type locality of Montegibbio (3-6.5 mm; Sacco, 1986,
p. 23), with one exceptionally large specimen attaining
12.6 mm in maximum diameter.
Patella protea in the NW French Neogene is found in
several Assemblage I localities. Millet (1854, p. 166;
1865, p. 599) recorded it from Sceaux-d’Anjou, Brébion
(1964, p. 67) added St-Michel and Contigné, to which we
add St-Clément-de-la-Place, the only locality in which it
is abundant.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet; 1854, 1865; Brébion, 1964); Proto-Medi-
terranean, Italy (Doderlein, 1862; Sacco, 1896b).
Superfamily Lottioidea Gray, 1840
Family Lottiidae Gray, 1840
Genus Tectura Gray, 1847
1847b Tectura Gray, p. 158. Type species (by original des-
ignation): Patella parva da Costa, 1778 [= Tectura
virginea (Müller, 1776)], present-day, British Isles.
Tectura virginea (Müller, 1776)
Plate 2, gs 1-2.
*1776 Patella virginea Müller, p. 43.
1778 Patella parva da Costa, p. 7, pl. 8, g. 11.
1835 Patella pulchella Forbes, p. 591, g. 61.
1844 Lottia unicolor Forbes, p. 135.
1848 Tectura virginea var. conica Jeffreys – Wood, p.
161, pl. 18, g. 6c.
1865 Tectura virginea var. conica Jeffreys, p. 249.
1865 Tectura virginea var. lactea Jeffreys, p. 249.
1896b Tectura virginea (Müll.) – Sacco, p. 19, pl. 2, gs
46-49.
1896b Tectura virginea var. parvoligustica Sacco, p. 20,
pl. 2, gs 50-52.
1925 Acmaea virginea (Müller) – Harmer, p. 875, pl.
65, g. 31.
1925 Acmaea virginea var. conica S.V. Wood [sic] –
Harmer, p. 876, pl. 65, g. 32.
1949 Patelloidea (Tectura) cf. virginea Müller, 1776 –
Glibert, p. 31, pl. 1, g. 14.
1964 Acmaea (Tectura) cf. virginea Müller, 1776 – Bré-
bion, p. 69.
2003 Acmaea (Tectura) virginea (O.F. Müller, 1776) –
Landau et al., p. 8, pl. 5, g. 3 (cum syn.).
2004 Tectura virginea (Müller O.F., 1776) – Chirli, p.
20, pl. 7, gs 4-11.
Plate 1. Patella protea Doderlein, 1862; 1. NHMW 2016/0103/0001, diameter 8.5 mm, height 4.1 mm. 2. NHMW 2016/0103/0002,
diameter 7.0 mm, height 2.8 mm. 3. NHMW 2016/0103/0003, diameter 5.6 mm, height 1.9 mm. Le Grand Chauvereau, St-
Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
80 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Material and dimensions – Maximum diameter 7.6
mm; height 3.8 mm. St-Clément-de-la-Place: NHMW
2016/0103/0005-0006 (2), NHMW 2016/0103/0007
(50+), RGM.1309477 (10), RGM.1309494 (50+),
RGM.1309565 (15), RGM.1348012 (50+), LC (50+), FVD
(50+). Sceaux-d’Anjou: NHMW 2016/0103/0009 (10),
RGM.1309476 (10), RGM.1309502 (25), RGM.1309521
(7), RGM.1347948 (14), RGM.1348022 (11), LC (12),
FVD (3). Renauleau: NHMW 2016/0103/0008 (16), LC
(8), FVD (5). Beugnon: RGM.1309566 (1), LC (1).
Discussion – In the French Assemblage I localities
Tectura virginea (Müller, 1776) is highly variable in
height, with almost at (Pl. 2, g 1) and high arched
(Pl. 2, g. 2) specimens found together at St-Clément-
de-la-Place, where this species is extremely abundant.
Many shells retain their original colour pattern, simi-
lar to that seen in present-day specimens (see Fretter &
Graham, 1976, p. 20, g. 15), although their maximum
diameter does not reach that recorded from the extant
populations (10 mm; Fretter & Graham, 1976; Poppe &
Goto, 1991). The smaller dimensions of the Assemblage
I specimens compared to conspecic populations found
in other strata is a recurrent theme we will encounter in
this series.
Distribution – Middle Miocene: Atlantic, Loire Basin,
France (Glibert, 1949). Upper Miocene: Atlantic (Torton-
ian), NW France (Brébion, 1964). Lower Pliocene: At-
lantic, Coralline Crag, England (Harmer, 1925); cen-
tral Mediterranean, Italy (Chirli, 2004), Tunisia (Fekih,
1975). Upper Pliocene: Atlantic, Red Crag, England (S.V.
Wood, 1848; Harmer, 1925), Mondego Basin, Portugal
(Silva, 2001); western Mediterranean, Estepona Basin, S.
Spain (Landau et al., 2003); central Mediterranean, Italy
(Sacco, 1896b). Pleistocene: Atlantic, England (Harmer,
1925); western Mediterranean, Balearic Islands (Cuerda
Barceló, 1987); central Mediterranean, Italy (Cerulli-
Irelli, 1916; Ruggieri & Greco, 1965; Taviani et al., 1998).
Present-day: Atlantic, northern Scandinavia to Cape
Verde Islands, 0-100m depth attached to hard substrates
(Poppe & Goto, 1991), Mediterranean (Giannuzzi-Savelli
et al., 1994).
Subclass Vetigastropoda
Superfamily Fissurelloidea Fleming, 1822
Family Fissurellidae Fleming, 1822
Subfamily Emarginulinae Children, 1834
Tribe Diodorini Odhner, 1932
Genus Diodora Gray, 1821
1821 Diodora Gray, p. 233. Type species (by monotypy):
Patella apertura Montagu, 1803, present-day, Brit-
ish Isles.
For generic synonymy see Ceulemans et al. (2016a, p. 54).
Diodora graeca (Linnaeus, 1758)
Plate 3, gs 1-4
*1758 Patella graeca Linnaeus, p. 784.
1854 Fissurella Exorata Millet, p. 166 (nomen nudum).
1854 Fissurella Labiatoides Millet, p. 166 (nomen nu-
dum).
1864 Fissurella labiatoides Millet, p. 680, footnote 2.
1865 Fissurella exorata Millet, p. 598.
1964 Diodora italica Defrance, 1820 – Brébion, p. 62
[non D. italica (Defrance, 1820)].
1964 Diodora apertura Montagu, 1803 – Brébion, p.
63, pl. 1, g. 2.
2016a Diodora graeca (Linné, 1758) – Ceulemans et al.,
p. 54, pl. 1, gs 4, 5 (cum syn.).
Material and dimensions – Maximum diameter 28.5
mm; height 9.0 mm. St-Clément-de-la-Place: NHMW
2016/0103/0077-0078 (2), NHMW 2016/0103/0079 (5),
RGM.1309481 (50+ juveniles), RGM.1309482 (12 sub-
adult and juveniles), RGM.1309487 (5 adult and juveniles),
RGM.1309492 (2), RGM.1309495 (8 subadult and juve-
niles), RGM.1309780 (7), RGM.1309781 (50+ subadult
and juveniles), LC (50+), FVD (50+). Sceaux-d’Anjou:
NHMW 2016/0103/0080 (9), RGM.1309478 (1 large adult),
RGM.1309479 (20), RGM.1309480 (4), RGM.1309483
(50+ subadult and juveniles), RGM.1309484 (10 sub-
adult and juveniles), RGM.1309485 (38), RGM.1309488
(1), RGM.1309491 (12), RGM.1309501 (35 subadult and
Plate 2. Tectura virginea (Müller, 1776); 1. NHMW 2016/0103/0005, diameter 7.5 mm, height 2.8 mm. 2. NHMW 2016/0103/0006,
diameter 7.0 mm, height 2.8 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper
Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 81
juveniles), LC (50+), FVD (50+). Renauleau: NHMW
2016/0103/1438 (11), LC (50+), FVD (40). Beugnon:
RGM.1309486 (25 subadult and juveniles), RGM.1309489
(7 subadult and juveniles), RGM.1309490 (22), RGM.
1309495 (11), RGM.1309496 (5), LC (5), FVD (2).
Discussion – In the Assemblage I localities we encoun-
ter the same problem with Diodora graeca (Linnaeus,
1758) outlined by Ceulemans et al. (2016a, p. 55). Some
specimens have more regular sculpture (Pl. 3, g. 4) and
had been described in the literature as Diodora aper-
tura (Montagu, 1803), although intermediate specimens
are also present (Pl. 3, g. 3). We therefore continue to
consider them as extreme forms of a single species. For
discussion see Landau et al. (2003) and Ceulemans et al.
(2016a, p. 55).
Brébion (1964, p. 65) reported this species (as D. aper-
tura) in localities representing Assemblages II (Apigné,
Le Temple du Cerisier, St-Jacques, Carcé), III (Le Pi-
geon Blanc, Le Girondor, La Gauvinière, Palluau, La
Dixmérie) and IV (St-Jean-la-Poterie, Gourbesville).
This is probably also the species recorded, but not illus-
trated, by Brébion (1964, p. 62) as D. italica (Defrance,
1820). Whilst the shell shape and relative smoothness of
the sculpture are reminiscent of this species, the close-up
of the sculpture (Pl. 3, g. 4c) shows it to be that of D.
graeca: strong primary cords with one medium-strength
secondary cord intercalated and one tertiary cord in each
of the interspaces, as opposed to the alternating primary
and secondary cords seen in D. italica. We therefore add
the Assemblage I localities given by Millet (1854, 1864,
1965) and Brébion (1964) for this species (Renauleau,
Sceaux-d’Anjou, Thorigné, St-Michel, St-Clément-de-la-
Place).
Distribution – Middle Miocene: Paratethys, Badenian,
Poland (Bałuk, 1975); Hungary (Strausz, 1966). Upper
Miocene: Atlantic (Tortonian and Messinian), NW France
(Millet, 1854, 1864; Brébion, 1964); Tethys, Tortonian:
Po valley, Italy (Sacco, 1896b). Lower Pliocene: Atlan-
tic, NW France (Brébion, 1964; Ceulemans et al., 2016a);
Coralline Crag, England (Harmer, 1923), Luchtbal For-
mation, Belgium (Marquet & Landau, 2006); central
Mediterranean, Italy (Chirli, 2004). Upper Pliocene: Red
Crag, England (Harmer, 1923); western Mediterranean,
Estepona, S. Spain (Landau et al., 2003); France (Chirli &
Richard, 2008); central Mediterranean, Italy (Malatesta,
1974; Caprotti, 1976; Cavallo & Repetto, 1992). Pliocene
(unspecied): central Mediterranean, Italy (Brambilla &
Lualdi, 1988); Atlantic Morocco (Lecointre, 1952). Up-
per Pliocene-lower Pleistocene: Atlantic, NW France
(Brébion, 1964); central Mediterranean, Italy (Cerulli-
Irelli, 1916; Malatesta, 1960). Pleistocene (unspecied):
western Mediterranean, Balearic Islands, (Cuerda Bar-
Plate 3. Diodora graeca (Linnaeus, 1758); 1. NHMW 2016/0103/0077, diameter 23.4 mm, height 9.0 mm; 2. NHMW 2016/0103/0078,
diameter 24.1 mm, height 8.5 mm; 3. NHMW 2016/0103/0081, diameter 23.2 mm, height 8.0 mm; 4. NHMW 2016/0103/0081,
diameter 26.1 mm, height 7.1 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, up-
per Miocene.
82 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
celó, 1987); Atlantic, Morocco (Lecointre, 1952). Upper
Pleistocene: England, Ireland (Harmer, 1923), The Neth-
erlands (Wesselingh & Pouwer, 2011). Present-day: At-
lantic British Isles to Canaries, Mediterranean and Black
Sea, rocky shores under stones and rocks, especially
where a little silt occurs, living near sponges, on which it
feeds (Poppe & Goto, 1991).
Diodora multida (Deshayes, 1830)
Plate 4, g. 1
*1830 Fissurella multida Deshayes, p. 136.
1830 Fissurella mitis Deshayes, p. 136.
1938 Fissurella (Lucapina) mitis Deshayes – Peyrot, p.
15, pl. 1, gs 10-12.
1849 Diodora multida Deshayes, 1831 [sic] – Glibert,
p. 28, pl. 1, g. 12.
1964 Diodora multida var. mitis Deshayes, 1832 [sic]
– Brébion, p. 65.
Material and dimensions – Maximum diameter 17.9 mm;
height 8.9 mm. Sceaux-d’Anjou: NHMW 2016/0103/0228
(1), NHMW 2016/0103/0229 (3), LC (8), FVD (8).
also have a more convex anterior slope than the multida
form, also seen in Glibert’s gure (1949, pl. 1, g. 12e).
Diodora multida differs from D. graeca (Linnaeus,
1758) with which it co-occurs in being smaller shelled,
in having a relatively more elevated dorsum and in hav-
ing ner sculpture. Diodora sancticlementensis nov. sp.,
found so far only at St-Clément-de-la-Place where we
have not found D. multida, has a maximimum size only
one-third that of D. multida, is more depressed, and has
smooth axial ribs.
Brébion (1964, p. 66) only recorded this species from the
upper Pliocene-Pleistocene Assemblage IV locality of
Gourbesville. This is the rst record in the Tortonian As-
semblage I of northwestern France.
Distribution – Middle Miocene: Atlantic, Loire Basin,
France (Peyrot, 1938; Glibert, 1949). Upper Miocene: At-
lantic (Tortonian), NW France (this paper). Upper Pliocene-
Pleistocene: Atlantic, NW France (Brébion, 1964).
Diodora sancticlementensis nov. sp.
Plate 5, gs 1-4
Type material – Holotype MNHN.F.A57687, diameter
5.2 mm; height 2.0 mm; paratype 1 MNHN.F.A57688,
diameter 6.9 mm; height 2.4 mm; paratype 2 NHMW
2016/0103/0083, diameter 5.8 mm; height 1.9 mm; para-
type 3 NHMW 2016/0103/0084, diameter 6.4 mm; height
2.0 mm; paratype 4 RGM.1309498, height 1.9 mm, diam-
eter 5.9 mm; paratype 5 RGM.1309499, height 1.6 mm,
diameter 5.4 mm.
Other material – Maximum diameter 6.7 mm. St-
Clément-de-la-Place: NHMW 2016/0103/0085 (50+),
RGM.1309497 (50+), RGM.1309500 (25), RGM.1309523
(50+), LC (50+), FVD (50+).
Etymology – Named after the type locality of St-Clé-
ment-de-la-Place. Diodora gender femimine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Diodora species of small size, low domed,
sculptured by about 70 narrow, ne, smooth axial ribs of
irregular strength, spiral sculpture inconspicuous.
Description – Shell small, patelliform, oval-elongated,
relatively low, with apical aperture placed just posterior
to mid length. Apical aperture ovate, thickened by callus
on inner surface. Both shorter posterior and longer ante-
rior slopes almost straight in prole. Sculpture of about
70 narrow axial ribs of irregular strength; in places of
alternate strength, which are not interrupted by growth
lines, resulting in almost smooth axial ribs.
Discussion – Diodora sancticlementensis nov. sp. is a
Plate 4. Diodora multida (Deshayes, 1830); 1. NHMW 2016/
0103/0228, diameter 17.9 mm, height 8.9 mm. La Presse-
lière, Sceaux-d’Anjou, Maine-et-Loire, NW France, Torton-
ian, upper Miocene.
Discussion – Both Fissurella multida Deshayes, 1830
and F. mitis Deshayes, 1830 were described from middle
or upper Miocene assemblages of northwestern France.
We have scant material attributed to this species, so we
accept Glibert’s (1949, p. 28) position in considering them
extreme forms of a single species, the mitis form having
ner sculpture in which the cords are all of roughly equal
strength. A similar variability in sculpture occurs in
many congeners (see D. graeca above). Having said that,
material at hand from Sceaux-d’Anjou is all of the mitis
variety. Apart from the ner sculpture, our specimens
Cainozoic Research, 17(2), pp. 75-166, December 2017 83
distinctive species, characterised by its small size for the
genus and sculpture of numerous, ne, smooth axial ribs.
In some specimens the ribs are of alternate strength at the
anterior and posterior margins and subequal on the lat-
eral slopes (Pl. 5, g. 3), but this feature is variable with
some specimens having every 5th or 6th cord stronger (Pl.
5, g. 2), or all cords subequal (Pl. 5, g. 4). In most Eu-
ropean Diodora species growth lines interrupt the axial
ribs forming a cancellate or squamate surface sculpture.
However, this is not the case in Diodora sancticlementen-
sis, which has smooth ribs. We can nd no other European
fossil or present-day species with this type of sculpture.
In this series covering the Assemblage I localities of
northwestern France we are struck by the number of gen-
era or species that have ‘dwarf’ forms in these deposits.
Diodora sancticlementensis is a good example of this. At
rst it would seem that these shells represent juveniles.
However, juvenile Diodora shells have an apical ‘beak’
on the dorsum at the anterior edge of the apical aperture,
which disappears with maturity. Moreover, D. sancticle-
mentensis is represented at St-Clément-de-la-Place by
hundreds of individuals, none of which have a greater
maximum diameter than 6.7 mm. This species is found
only at the type locality.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Tribe Emarginulini Children, 1834
Genus Emarginula Lamarck, 1801
1801 Emarginula Lamarck, p. 69. Type species (by
monotypy): Emarginula conica Lamarck, 1801
[= Emarginula ssura (Linnaeus, 1758)], present-
day, Europe.
For generic synonymy see Ceulemans et al. (2016a, p. 55).
Emarginula adriatica Costa, 1830
Plate 6, gs 1-2
*1830 Emarginula adriatica, Costa, p. 11.
1854 Emarginula Proclinata Millet, p. 166 (nomen nu-
dum).
1865 Emarginula proclinata Millet, p. 599.
1938 Emarginula elongatoides Peyrot, p. 18, pl. 1, gs
15, 16.
1949 Emarginula clathrataeformis Eichwald, 1830 –
Glibert, p. 18, pl. 1, g. 6 (?non Eichwald, 1830).
1964 Emarginula clathrataeformis Eichwald, 1830 –
Brébion, p. 55 (?non Eichwald, 1830).
2003 Emarginula adriatica O.G. Costa, 1829 [sic] –
Landau et al., p. 19, pl. 3, gs 1-3 (cum syn.).
Material and dimensions – Maximum diameter 7.4
mm; height 3.4 mm. St-Clément-de-la-Place: NHMW
2016/0103/0095-0096 (2), NHMW 2016/0103/0097 (3),
RGM.1309545 (2), RGM.1309612 (7), LC (6), FVD
(7). Sceaux-d’Anjou: NHMW 2016/0103/0098 (18),
RGM.1309607 (2), RGM.1309614 (5), RGM.1348015 (8),
LC (2), FVD (15).
Discussion – Emarginula adriatica Costa, 1830 is vari-
able in shape, with the apex more or less recurved and
the axial sculpture stronger or equal in strength to the
concentric ribs. Constant features are the oval-elongated
shape and the papillae formed at the intersections of the
sculpture. The nomenclature of this group is confused in
the literature, making fossil records unreliable. Glibert
(1949) illustrated shells we consider to be conspecic
under the name E. clathrataeformis Eichwald, 1830. We
are unsure if the Paratethyan shell is indeed conspecic,
but the middle Miocene Loire Basin shells illustrated by
Glibert are the same species as those gured here (Pl.
6, gs 1-2). Glibert described the characteristic tubercles
formed at the sculptural intersections in his material:
Plate 5. Diodora sancticlementensis nov. sp.; 1. Holotype MNHN.F.A57687, diameter 5.2 mm, height 2.0 mm; 2. Paratype 1
MNHN.F.A57688, diameter 6.9 mm, height 2.4 mm; 3. Paratype 2 NHMW 2016/0103/0083, diameter 5.8 mm, height 1.9
mm; 4. Paratype 3 NHMW 2016/0103/0084, diameter 6.4 mm, height 2.0 mm. Le Grand Chauvereau, St-Clément-de-la-Place,
Maine-et-Loire, NW France, Tortonian, upper Miocene.
84 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
‘Les côtes principals sont également ornées d’une série
de tubercules pointus raprochés qui leur donnent une
apparence dentelée (1949, p. 19)’. We are unsure if the
middle Miocene Paratethyan shell described by Eichwald
(1830) is conspecic. If it were, E. adriatica might be-
come a junior synonym of E. clathrataeformis, as Costa’s
work was not published before November 1830 and not
1829 as stated on the title page (Fasulo, 2013). The date
of the front page of Eichwald’s work is given as January
10th 1830, which means that the ofcial version of the
book was read by the censor in 1829. In the preface Eich-
wald dates his letter 23rd March 1830 and in the epilogue
he refers to a paper published in May 1830, which he had
just seen during the nal phase of the book’s preparation.
Therefore a date before May/June 1830 is not possible,
but we cannot get closer to a publication date. There is
no further information on the library stamp of the copy
at the NHMW (M. Harzhauser, personal communication
June 2016).
Emarginula octaviana Coen, 1939, also found in the As-
semblage I deposits, has a similarly elongated depressed
shell with a strongly recurved apex and a reticulate sculp-
ture, but in that species the lamellae on the selenizone,
which are relatively coarser, are raised above the level of
the reticulate sculpture.
Millet (1854) recorded this species from Assemblage I
localities of Sceaux-d’Anjou and Thorigné, to which Bré-
bion (1964, p. 55) added St-Clément-de-la-Place and Les
Pierres Blanches.
Distribution – Middle Miocene: Atlantic, Loire Basin
(Peyrot, 1938; Glibert, 1949). Upper Miocene: Atlantic
(Tortonian), NW France (Millet, 1854; Brébion 1964).
Upper Pliocene: western Mediterranean, Estepona Basin,
S. Spain (Landau et al., 2003). Pliocene (indeterminate):
central Mediterranean, Italy (Piani, 1984). Lower Pleis-
tocene: central Mediterranean, Italy (Cerulli-Irelli, 1916;
Borghi & Vecchi, 1998). Present-day: Atlantic coasts of
Europe north to France, Morocco and Mediterranean, cir-
calittoral, on corals from 20-400 m depth (Piani, 1984).
Emarginula imbricata Millet, 1865
Plate 7, gs 1-2
1854 Emarginula Imbricata Millet, p. 166 (nomen nu-
dum).
*1865 Emarginula imbricata Millet, p. 599.
1886 Emarginula Dujardini Dollfus & Dautzenberg, p.
142.
1964 Emarginula dujardini Dollfus & Dautzenberg –
Brébion, p. 60, pl. 1, gs 10, 11.
2016a Emarginula dujardini Dollfus & Dautzenberg,
1886 – Ceulemans et al., p. 56, pl. 2, g. 1 (cum
syn.).
Material and dimensions – Maximum diameter 20.2
mm, height 9.1 mm. St-Clément-de-la-Place: NHMW
2016/0103/0094 (1), RGM.1309503 (1), LC (3), FVD (8).
Sceaux-d’Anjou: NHMW 2016/0103/1426 (1), LC (1),
FVD (1). Renauleau: NHMW 2016/0103/1604 (1 frag-
ment), LC (5), FVD (3). Beugnon: RGM.1309504 (1 frag-
ment), RGM.1309505 (1 fragment).
Discussion – This is the largest emarginulid in the upper
Miocene-Pleistocene northwestern French assemblages.
When we recorded the species in the Assemblage III local-
ity of Le Pigeon Blanc (Ceulemans et al., 2016a), we failed
to spot that Emarginula dujardini Dollfus & Dautzen berg,
1886 is a subjective junior synonym of E. imbricata Mil-
let, 1865. The description given by Millet (1865, p 599)
that validated the earlier nomen nudum (Millet, 1854, p.
166) undoubtedly refers to this species ‘Cette espèce, la
plus grande de Maine-et-Loire, se distingue encore de
celles-ci, en ce qu’elle présente à sa base plutôt un ovale
qu’une ellipse, que son sommet, qui est à peine courbé,
est placé aux deux tiers de la longueur totale de la co-
quille, que la fente marginale n’est pas suivie d’un canal
et que toutes les stries ou très-petites côtes rayonnantes
qui la couvrent sont imbriquées. Longueur: 13 milli-
mètres; diamètre: 10 millimètres; hauteur: 8 millimètres.
Sc., Th., Ren. (1865, p 599)’ This species has rarely been
Plate 6. Emarginula adriatica Costa, 1830; 1. NHMW 2016/0103/0095, diameter 7.4 mm, height 3.2 mm; 2. NHMW 2016/0103/0096,
diameter 7.4 mm, height 3.4 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper
Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 85
recorded in the literature and we cannot therefore satisfy
the requirements of Article 23.9.1.2 (ICZN 1999a) and
consider Millet’s name a nomen oblitum.
Emarginula imbricata is widespread, but always uncom-
mon. Millet (1854, 1865) recorded it from the Assemblage
I localities of Sceaux-d’Anjou, Thorigné and Renauleau,
to which Brébion (1964, p. 61) added St-Clément-de-la-
Place and Chalonnes, Assemblage II (Apigné) and As-
semblage III (Le Girondor, La Gauvinière); to which
Ceulemans et al. (2016a) added Le Pigeon Blanc. For fur-
ther discussion see Ceulemans et al. (2016a, p. 56).
Distribution – Middle Miocene: Atlantic, Loire Basin,
France (Dollfus & Dautzenberg, 1886; Peyrot, 1938;
Glibert, 1949). Upper Miocene: Atlantic (Tortonian and
Messinian), NW France (Brébion, 1964). Lower Pliocene:
Atlantic, NW France (Brébion, 1964; Ceulemans et al.
2016a).
Emarginula octaviana Coen, 1839
Plate 8, gs 1-2
1830 Emarginula elongata, Costa, p. 10 (non Defrance,
1819; non G.B. Sowerby I, 1823; non Gray, 1825).
*1839 Emarginula octaviana Coen, p. 71.
1854 Emarginula Ornata Millet, p. 166 (nomen nu-
dum).
1865 Emarginula ornata Millet, p. 599.
1949 Emarginula elongata Da Costa – Glibert, p. 20, pl.
1, g. 9.
2016a Emarginula octaviana Coen, 1839 – Ceulemans et
al., p. 57, pl. 2, g. 3 (cum syn.).
Material and dimensions – Maximum diameter 12.8
mm; height 5.4 mm. St-Clément-de-la-Place: NHMW
2016/0103/0086-0087 (2), NHMW 2016/0103/0088 (29),
RGM.1309605 (5), RGM.1309609 (20), RGM.1309613
(6), LC (11), FVD (10). Sceaux-d’Anjou: NHMW 2016/
0103/0089 (30), RGM.1309604 (2), RGM.1309606 (20),
RGM.1309608 (40), RGM.1309610 (9), RGM.1309615
(13), RGM.1309623 (6), RGM.1348016 (11), LC (18), FVD
(50+). Renauleau: NHMW 2016/0103/1549 (7), LC (34),
FVD (7). Beugnon: LC (3).
Discussion – The salient characters of this species were
described by Glibert (1949, p. 20; as E. elongata): the
low, elongated shell shape, coarse reticulate sculpture
and most importantly the lamellae on the selenizone,
Plate 7. Emarginula imbricata Millet, 1865; 1. RGM.1309503, diameter 17.9 mm, height 7.0 mm. Le Grand Chauvereau, St-Clément-
de-la-Place. 2. NHMW 2016/0103/1426, diameter 17.2 mm, height 8.0 mm. La Presselière, Sceaux-d’Anjou, Maine-et-Loire, NW
France, Tortonian, upper Miocene.
Plate 8. Emarginula octaviana Coen, 1839; 1. NHMW 2016/0103/0086, diameter 8.2 mm, height 3.1 mm; 2. NHMW 2016/0103/0087,
diameter 9.3 mm, height 4.9 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper
Miocene.
86 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
which are relatively coarse and raised above the level of
the reticulate sculpture. The shell height is variable in the
specimens from St-Clément-de-la-Place (Pl. 8, gs 1b,
2b). For further discussion see Ceulemans et al. (2016a,
p. 58).
Millet (1854, 1865) recorded this species from Renauleau,
Sceaux-d’Anjou, Thorigné and St-Clément-de-la-Place,
to which Brébion (1964, p. 59; as E. elongata) added
Chalonnes and a single specimen from Assemblage IV
(Gourbesville). The maximum size of the specimens
from Sceaux-d’Anjou is considerably greater than that of
the specimens from St-Clément-de-la-Place (maximum
diameter 12.7 mm vs. 9.3 mm). Ceulemans et al. (2016a,
p. 57) added the Assemblage III locality of Le Pigeon
Blanc.
Distribution – Upper Miocene: Atlantic (Tortonian),
NW France (Millet, 1854, 1865; Brébion, 1964). Lower
Pliocene: Atlantic, NW France (Brébion, 1964; Ceule-
mans et al. 2016a). Upper Pliocene-lower Pleistocene:
NW France (Brébion, 1964). Present-day: Atlantic Ca-
naries, Portugal, Morocco and Mediterranean (Poppe &
Goto, 1992).
Emarginula rosea Bell, 1824
Plate 9, gs 1-2
*1824 Emarginula rosea Bell, p. 52, pl. 4, gs 1, 2.
1854 Emarginula Rostrata Millet, p. 166 (nomen nu-
dum).
1865 Emarginula rostrata Millet, p. 599.
1923 Emarginula rosea T. Bell – Harmer, p. 779, pl. 62,
g. 10.
1949 Emarginula conica Lamarck, 1801 – Glibert, p.
17, pl. 1, g. 3 [non Lamarck, 1801 = E. ssura
(Linnaeus, 1758)].
1954 Emarginula conica Lamarck, 1801 – van Regteren
Altena, p. 57, pl. 1, g. 3 [non Lamarck, 1801 = E.
ssura (Linnaeus, 1758)].
1964 Emarginula reticulata Sowerby, 1813 – Brébion
(partim, Assemblage I specimens only), p. 55, pl.
1, g. 5 (non Sowerby, 1813 = E. ssura (Linnae-
us, 1758)].
1964 Emarginula reticulata var. rosea Bell, 1824 – Bré-
bion, p. 57, pl. 1, g. 6.
2003 Emarginula rosea Bell, 1824 – Landau et al., p.
21, pl. 3, g. 7 (cum syn.).
2011 Emarginula rosea Bell, 1824 – Wesselingh & Pou-
wer, p. 136, gs 21, 22.
Material and dimensions – Maximum diameter 7.5
mm; height 7.0 mm. St-Clément-de-la-Place: NHMW
2016/0103/0091-0092 (2), NHMW 2016/0103/0092 (50+),
RGM.1309554 (1), RGM.1309611 (50+), RGM.1309620
(50+), LC (50+), FVD (50+). Sceaux-d’Anjou: NHMW
2016/0103/0093 (50+), RGM.1309557 (6), RGM.1309616
(9), RGM.1309619 (17), RGM.1309621 (1), RGM.1309622
(25), RGM.1348017 (21), LC (20), FVD (9). Renauleau:
LC (4). Beugnon: LC (2).
Discussion – This group of emarginulids from the Mio-
cene of northwestern France is problematic. Glibert
(1949) described and illustrated specimens from the mid-
dle Miocene Loire Basin of France as Emarginula re-
ticulata J. Sowerby, 1813 (= E. ssura Linnaeus, 1758).
Brébion (1964) recorded E. reticulata and E. rosea Bell,
1824 from the ‘Redonian’ assemblages, and considered
rosea a variety of E. reticulata. Ceulemans et al. (2016a)
recognised E. ssura in the lower Pliocene Assemblage
III deposit of Le Pigeon Blanc. Emarginula rosea is dis-
tinguished by its very elevated shell, usually smaller in
size than E. ssura, coarse sculpture and the posterior
position of the apex, which is slightly more recurved.
The problem lies in the identication of the Assemblage
I specimens. Some of the larger specimens are taller than
they are broad, and are typical for E. rosea, whilst many
of the smaller ones are lower and at rst glance similar to
E. ssura. However, examination of the numerous speci-
mens at hand from St-Clément-de-la-Place and Sceaux-
d’Anjou shows that the apex is always more posteriorly
placed than it is in E. ssura. This is also true of the shell
illustrated by Brébion (1964, pl. 1, g. 5) from Beauleau.
Moreover, the size range of the Assemblage I shells is
similar to that of E. rosea, E. ssura being twice as large
or more. We therefore consider all the Assemblage I shells
Plate 9. Emarginula rosea Bell, 1824; 1. NHMW 2016/0103/0090, diameter 5.2 mm, height 5.4 mm; 2. NHMW 2016/0103/0091,
diameter 5.4 mm, height 4.5 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper
Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 87
seen E. rosea and therefore, E. rostrata Millet, 1865 is a
junior subjective synonym of E. rosea rather than E. re-
ticulata, as suggested by Brébion (1964, p. 56). It seems
that the shell increases in height with growth, the tall-
est being the most gerontic (Pl. 9, g. 1). The specimens
from the middle Miocene Loire Basin described by Glib-
ert (1949) as E. reticulata (= E. ssura) have a more cen-
trally placed apex and do indeed seem to be that species.
It seems, therefore, that both species were present in the
middle-upper Miocene of northwestern France, and may
have exploited different habitats as we have not found the
two species together, although Fretter & Graham (1976,
p. 10) suggested the two species shared a similar habitat
today.
Emarginula rosea is the commonest emarginulid in
the Assemblage I localities and, as discussed above, we
consider the Assemblage I references to E. reticulata of
Brébion (1964, p. 57) to be E. rosea (Renauleau, Sceaux-
d’Anjou, Thorigné, St-Michel, St-Clément-de-la-Place,
Les Pierres Blanches, Beaulieu). The Assemblage II
records need to be veried. Brébion (1964, p. 58) record-
ed E. rosea from the Assemblage III localities of Le Gi-
rondor, La Gauvinière and Palluau, although Ceulemans
et al. (2016a) did not nd it at Le Pigeon Blanc.
Distribution – Upper Miocene: Atlantic (Tortonian),
NW France (Millet, 1854, 1865; Brébion, 1964). Lower
Pliocene: North Sea Basin, Coralline Crag, England
(Harmer, 1923), Kattendijk Formation, Belgium (Mar-
quet, 1998); Atlantic, NW France (Brébion, 1964). Upper
Pliocene: North Sea Basin, Red Crag, England (Harmer,
1923); western Mediterranean, Estepona Basin, S. Spain
(Landau et al., 2003), North Sea Basin (Marquet, 1995);
central Mediterranean, Italy (Piani, 1984). Pliocene (in-
determinate): North Sea Basin, The Netherlands (van
Regteren Altena, 1954; Wesselingh & Pouwer, 2011).
Lower Pleistocene: central Mediterranean, Italy (Cer-
ulli-Irelli, 1916). Upper Pleistocene: North Sea Basin,
Netherlands (Wesselingh & Pouwer, 2011). Present-day:
Atlantic coasts of Europe north to the British Isles and
Mediterranean, 0-90 m (Piani, 1984).
Tribe Fissurellideini Pilsbry, 1890
Genus Lucapinella Pilsbry, 1890
Type species (by subsequent designation (Pilsbry, 1890) –
Clypidella callomarginata Dall, 1871, present-day, Cali-
fornia.
Lucapinella clypeata (Grateloup, 1828)
Plate 10, gs 1-2
1828a Fissurella depressa Grateloup, p. 79, no. 9 [non
F. depressa Lamarck, 1822 (= F. crassa Lamarck,
1822)].
*1828a Fissurella clypeata Grateloup, p. 79, no. 10.
2013 Lucapinella clypeata (Grateloup, 1828) – Landau
et al., p. 23, p. 1, g. 1 (cum syn.).
Material and dimensions – Maximum diameter 4.6 mm.
St-Clément-de-la-Place: NHMW 2016/0103/1626-1627
(2). Sceaux-d’Anjou: RGM.1348014 (1). Renauleau:
NHMW 2016/0103/1628 (5)
Discussion – Cossmann & Peyrot (1917) considered Fis-
surella depressa Grateloup, 1828 (non Lamarck, 1822,
renamed F. aquensis by d’Orbigny, 1852) described from
the early Oligocene Rupelian of Gaas, France, and speci-
mens from the early Miocene Aquitanian-Burdigalian
of the Aquitaine Basin, France, described as Fissurella
clypeata Grateloup,1828 to represent a single highly vari-
able species. The specimens from the Assemblage I de-
posits are very small, but otherwise t within the wide
range of variability.
Plate 10. Lucapinella clypeata (Grateloup, 1828); 1. NHMW
2016/0103/1626, height 2.0 mm, length 4.6 mm; 2. NHMW
2016/0103/1627, height 1.9 mm, length 4.1 mm. Le Grand
Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW
France, Tortonian, upper Miocene.
Distribution – Lower Oligocene: Atlantic (Rupelian):
Aquitaine Basin, France (Grateloup, 1828). Lower Mi-
ocene: Atlantic (Aquitanian and Burdigalian): Aquitaine
Basin, France (Cossmann & Peyrot, 1917; Lozouet et al.,
2001a); Proto-Mediterranean Sea (Burdigalian): Colli
Torinesi, Italy (Sacco, 1896b). Middle Miocene: north-
eastern Atlantic (Langhian): Aquitaine Basin, France
(Cossmann & Peyrot, 1917), Loire Basin, France (Glib-
ert, 1949); Paratethys (Langhian-Serravallian): Vienna
Basin, Austria (Hörnes, 1856), Poland (Bałuk, 1975),
Hungary (Csepreghy-Meznerics, 1954; Strausz, 1954,
1966), Bulgaria (Kojumdgieva & Strachimirov, 1960),
Romania (Popa & Ianoliu, 2000), Bosnia (Atanacković,
1985); Proto-Mediterranean Sea (Serravallian): Karaman
Basin, Turkey (Landau et al., 2013). Upper Miocene: At-
lantic (Messinian): NW France (this paper); Proto-Medi-
terranean Sea (Tortonian): Po valley, Italy (Sacco, 1896b).
Superfamily Haliotoidea Ranesque, 1815
Family Haliotidae Ranesque, 1815
Genus Haliotis Linnaeus, 1758
88 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Type species (by subsequent designation, de Montfort,
1810) – Haliotis asinina Linnaeus, 1758, present-day,
Indo-Pacic.
1758 Haliotis Linnaeus, p. 779.
1854 Teinotis H. Adams & A. Adams, p. 442. Type spe-
cies (by subsequent designation, Fischer, 1885):
Haliotis asinina Linnaeus, 1758, present-day,
Indo-Pacic.
1856 Schismotis Gray, p. 147. Type species (by
monotypy): Schismotis excisa Gray, 1856, present-
day, Australia.
1885 Tinotis Fischer, p. 845. Incorrect subsequent spell-
ing of Teinotis.
1908 Teinotus Schepman, 1908. Incorrect subsequent
spelling of Teinotis.
1929 Sanhaliotis Iredale, p. 270. Type species (by origi-
nal designation): Haliotis varia Linnaeus, 1758,
present-day, Australia.
1933 Exohaliotis Cotton & Godfrey, p. 16. Type spe-
cies (by original designation): Haliotis cyclobates
Péron, 1816, present-day, South Australia.
1943 Ovinotis Cotton, p. 175, 179. Type species (by
original designation): Haliotis ovina Gmelin,
1791, present-day, Indo-Pacic.
1964 Eurotis Habe & Kosuge, p. 6. Type species (by
original designation): Haliotis tuberculatus Lin-
naeus, 1758, present-day, Europe.
1964 Usahaliotis Habe & Kosuge, p. 8. Type species
(by original designation): Haliotis cracherodii
Leach, 1814, present-day, north eastern Pacic.
Haliotis cf. tuberculata coccinea Reeve, 1846
Plate 11, g. 1
cf. *1846 Haliotis coccinea Reeve, 1846, p. 55.
cf. 1994 Haliotis coccinea Reeve, 1846 – Giannuzzi-
Savelli et al., p. 54, g. 104.
cf. 2000 Haliotis tuberculata coccinea Reeve, 1846 – Gei-
ger & Poppe, p. 90, pl. 7.
cf. 2002 Haliotis tuberculata coccinea Reeve, 1846 – Ávi-
la et al., p. 348, g. 5.
cf. 2012 Haliotis tuberculata coccinea Reeve, 1846 – Gei-
ger & Owen, p. 135, pl. 69, gs 1-14.
Material and dimensions – Maximum diameter 16.8
mm (fragment). St-Clément-de-la-Place: NHMW 2016/
0103/0147 (1), NHMW 2016/0103/0148 (26 fragments),
RGM.1309555 (2 fragments), RGM.1309564 (3 frag-
ments), LC (12 juveniles), FVD (1 fragment). Renauleau:
NHMW 2016/0103/1439 (1 larger fragment), LC (1 juve-
nile).
Discussion – The genus is represented in the St-Clément-
de-la-Place assemblage by a number of small specimens
that we consider to be juvenile, as none of the tremata
(respiratory openings) are open. They are sculptured by
seven wavy narrow axial ribs between the suture and
the tremata, narrower than their interspaces. The shoul-
der rib is wider, splitting and surrounding the tremata.
Three further ribs are placed between the shoulder and
the sharp, elevated peripheral keel. The base bears eight
regular ribs.
Similar axial sculpture composed of wavy ribs is present
in juvenile shells of the present-day Haliotis tuberculata
coccinea Reeve, 1846, but with slightly fewer cords (3-5
vs. 7). We therefore hesitate in considering the shells from
St-Clément-de-la-Place conspecic with the present-day
H. t. coccinea, but the two are nevertheless very similar.
Haliotis coccinea is now considered a subspecies of H.
tuberculata Linnaeus, 1758 (Geiger, 1998, p. 97; Geiger
& Poppe, p. 90; Geiger & Owen, 2012, p. 135). Today
this species has a more southern distribution, found in the
Azores and Canary Islands. We have Pleistocene fossil
specimens at hand from Santa Maria Island in the Azores
(NHMW coll.; Ávila et al., 2002).
Distribution – Upper Miocene (Tortonian): Atlantic, NW
France (this paper).
Superfamily Scissurelloidea Gray, 1847
Note – For this section we have followed the terminology
and descriptive terms adopted by Geiger (2012). We are
greatly endebted to that author for his advice and guid-
ance with this superfamily.
Family Scissurellidae Gray, 1847
Subfamily Scissurellinae Gray, 1847
Genus Scissurella d’Orbigny, 1824
Type species (by subsequent designation, Gray, 1847b)
– Scissurella laevigata d’Orbigny, 1824 (= S. costata
d’Orbigny, 1824), present-day, Mediterranean.
1824 Scissurella d’Orbigny, p. 341, 343.
1856 Schismope Jeffreys, p. 321. Type species (by
monotypy): Scissurella striatula Philippi, 1844,
Plate 11. Haliotis cf. tuberculata coccinea Reeve, 1846; 1.
NHMW 2016/0103/0147, diameter 3.0 mm x 2.1 mm. Le
Grand Chauvereau, St-Clément-de-la-Place, Maine-et-
Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 89
present-day, Italy.
1861 Woodwardia Crosse & Fischer, p. 160. Type spe-
cies (by monotypy): Scissurella elegans d’Orbigny,
1824, Pliocene, Italy.
1998 Maxwellella Bandel, p. 19. Type species (by origi-
nal designation): Scissurella annulata Ravn, 1933,
Paleocene, Denmark.
1998 Reussella Bandel, p. 44. Type species (by original
designation): Scissurella depressa Reuss, 1860,
Miocene, Bohemia. Junior homonym of Reussella
Galloway, 1933 [Foraminifera].
1998 Praescissurella Lozouet, p. 66. Type species (by
or ig inal designation): Scissurella depontaillieri Coss-
mann, 1879, Oligocene, France.
Scissurella transylvanica Reuss, 1860
Plate 12, g. 1
*1860 Scissurella transylvanica Reuss, p. 266, pl. 7, g.
6.
2012 Scissurella transylvanica Reuss, 1860 – Geiger, p.
363, gs 259-263 (cum syn.).
2013 Scissurella transylvanica Reuss, 1860 – Landau et
al., p. 23, pl. 54, g. 2 (cum syn.).
Material and dimensions – Maximum diameter 1.4 mm.
St-Clément-de-la-Place: NHMW 2016/0103/1429 (3),
RGM.1309568 (30). Sceaux-d’Anjou: RGM.1309575 (1),
RGM.1309589 (1). Renauleau: NHMW 2016/0103/1429
(1), NHMW 2016/0103/1440 (10). Beugnon: RGM.
1309552 (2).
Discussion – Scissurella transylvanica Reuss, 1860 is
characterised by its strongly axially sculptured proto-
conch (Pl. 12, g 1d) and strong, close-set axial ribs on
the teleoconch. Little can be added to the excellent revi-
sion on the group by Geiger (2012), except to say that this
occurrence in the Assemblage I deposits of NW France is
the stratigraphically youngest for the species.
Distribution – Upper Oligocene: Aquitaine Basin, France
(Bandel, 1998). Lower Miocene: northeastern Atlantic
(Aquitanian): Aquitaine Basin, France (Benoist, 1875; Lo-
zouet et al., 2001). Middle Miocene: northeastern Atlantic
(Langhian): Loire Basin, France (de Morgan, 1915); Para-
tethys (Langhian-Serravallian): Hungary (Kecskemétiné
Körmendy, 1962; Kókay, 1966; Strausz, 1966), Poland
(Friedberg, 1938; Bałuk, 1975), Romania (Reuss, 1860);
Proto-Mediterranean Sea (Serravallian): Karaman Basin,
Turkey (Landau et al., 2013). Upper Mio cene (Tortonian):
Atlantic, NW France (this paper).
Genus Sinezona Finlay, 1926
Type species (by original designation) – Schismope brevis
Hedley, 1904, present-day, New Zealand.
1926 Sinezona Finlay, p. 341, 346.
1998 Daizona Bandel, p. 57. Type species (by original
designation): Sinezona doliolum Herbert, 1986,
present-day, South Africa.
1998 Ariella Bandel, p. 63. Type species (by original
designation): Ariella haliotimorpha Bandel, 1998,
Oligocene, France.
Sinezona geigeri nov. sp.
Plate 13, g. 1
Type material – Holotype MNHN.F.A57395, diameter 0.9
mm; height 0.75 mm; paratype 1 NHMW 2016/0103/0010;
paratype 2 NHMW 2016/0103/0011; paratype 3 NHMW
2016/0103/0011; paratype 4 RGM.1309567.
Other material – St-Clément-de-la-Place: NHMW 2016/
0103/1394 (1).
Etymology – Named after Daniel L. Geiger, curator at the
Santa Barbara Museum of Natural History, California,
USA in recognition of his enormous contribution to the
the knowledge of the superfamily Scissurelloidea. Sine-
zona gender feminine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Sinezona species of small size, with no axial
sculpture on shoulder of teleoconch II, base with seven
raised ribs that end abruptly at edge of base and deep um-
bilicus bordered by raised funiculus.
Plate 12. Scissurella transylvanica Reuss, 1860; 1. NHMW 2016/0103/1429, diameter 1.4 mm; 1d, detail of protoconch (SEM im-
age). Renauleau, Maine-et-Loire, NW France, Tortonian, upper Miocene.
90 Landau, Van Dingenen & Ceulemans. The upper Miocene gast ropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Description – Shell minute, depressed trochiform. Pro-
toconch of 1.25 whorls, bearing strong axial sculpture on
the outer half of the whorl, not connecting to embryonic
cap. Teleoconch I of 0.65 whorl, bearing six strong, wi-
dely spaced axial ribs, no spiral in position of selenizone.
Teleoconch II of 0.7 whorl, suture little impressed, spi-
ral sculpture above selenizone starting with three cords,
increasing abapically to eleven at outer lip; cords of ir-
regular strength and position. Last whorl below seleni-
zone straight-sided to weakly concave, bearing irregular
ne cords. Base convex, with seven sharp, elevated ribs,
which start abruptly at the edge of base. Selenizone just
below apex of shell, slit closed to form triangular oblong
foramen, keels moderately elevated. Umbilicus open,
deep, bordered by elevated funiculus.
Discussion – Sinezona geigeri nov. sp. is separated from
most of its European congeners by the absence of axial
sculpture on the shoulder. There are relatively few eastern
Atlantic congeners with which it can be compared. The
Mediterranean and West African Pliocene to present-day
Sinezona cingulata (O.G. Costa, 1861) has far denser axial
sculpture on the protoconch and axial sculpture that ex-
tends between the sutures on teleoconch II. Sinezona semi-
costata Burnay & Rolán, 1990 from the Canary and Cape
Verde Islands has a strongly depressed shell and strong
axial sculpture. Sinezona terquemi (Deshayes, 1865) from
the lower Miocene Aquitaine Basin of France has dense
axial sculpture, weak spiral sculpture restricted to the ad-
umbilical quarter of the base, and a broad umbilicus that
is not delimited by a funiculus.
Distribution – Upper Miocene (Tortonian): Atlantic, NW
France (this paper).
Family Anatomidae McLean, 1989
Genus Anatoma Woodward, 1859
Type species (by monotypy) – Scissurella crispata Flem-
ing, 1828, present-day, British Isles.
1859 Anatoma Woodward, p. 204.
1877 Schizotrochus Monterosato, p. 416. Type species
(by monotypy): Scissurella crispata Fleming,
1828, present-day, British Isles. Objective syno-
nym.
1998 Hainella Bandel, p. 36. Type species (by origi-
nal designation): Scissurella euglypta Pelseneer,
1903, present-day, Antarctic.
1998 Thieleella Bandel, p. 35. Type species (by original
designation): Scissurella amoena Thiele, 1912,
present-day, Antarctic.
1998 Pagodella Bandel, p. 2 (nomen nudum).
Anatoma redoniana nov. sp.
Plate 14, g. 1
Type material – Holotype NHMW 2016/0103/1470, di-
ameter 1.4 mm, height 0.9 mm, St-Clément-de-la-Place.
Paratype 1 NHMW 2016/0103/1624, diameter 1.2 mm,
height 0.7 mm, Renauleau.
Other material – Renauleau: NHMW 2016/0103/1625 (2
fragments).
Etymology – Named after the ‘Redonian’ stage, the name
used until recently for these NW French post-middle
Mio cene assemblages. Anatoma gender feminine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Anatoma species of small size, low-turreted
to at dorsum, with protoconch bearing occulent sculp-
ture, teleoconch with narrow sutsel, slightly narrower
than selenizone, sculptured by ne, close-set sinuous
axial ribs on both dorsum and venter, no spiral sculpture,
keeled base.
Description – Shell small, somewhat turreted to at dor-
sum. Protoconch of 1.75 whorls, sculpture occulent, no
varix. Teleoconch I of 0.5 whorl bearing close-set, sinu-
ous axials. Teleoconch II of one whorl, suture moderately
impressed, sutsel (portion of the base between the suture
and the lower keel of the selenizone above it; Geiger,
2012) slightly narrower than selenizone. Shoulder bear-
Plate 13. Sinezona geigeri nov. sp.; 1. Holotype MNHN.F.A57395, diameter 0.9 mm, height 0.75 mm (SEM image). Le Grand Chau-
vereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 91
ing numerous close-set, sinuous axial ribs. Spiral sculp-
ture absent. Base not constricted below selenizone, mod-
erately strongly keeled mid-base, with similar pattern
of sinuous axials. Umbilicus open, deep, continuously
sloping from base; funiculus curved away from apertural
margin. Selenizone at periphery; lunules distinct; keels
strong, elevated, external surface with many strong axial
lines; slit open. Aperture subquadrate, somewhat ared,
particularly towards umbilicus, where apertural margin
fuses with funiculus.
Discussion – Despite being represented by scant mate-
rial, Anatoma redoniana nov. sp. is so unlike its Europe-
an congeners that it merits description. The most salient
characters of this new species are the dense sinuous axial
sculpture covering both the dorsum and the venter and the
keeled base. We note that the spire in all three specimens
from Renauleau is completely at and not slightly elevat-
ed as seen in the holotype (Pl. 14, g. 1c), but the rest of
the characters and sculpture are like that of the holotype
from St-Clément-de-la-Place. Anatoma umbilicata (Jef-
freys, 1883), with which it is most similar in the European
faunas differs in having far less dense axial sculpture and
in not having a keeled base. The shells identied as A.
umbilicata by Landau et al. (2003) from the lower upper
Pliocene of Estepona, southern Spain are not this spe-
cies, but Anatoma eximia (Seguenza, 1880), which has a
more turreted shell shape, much wider sutsel at any given
whorl count and the selenizone is placed higher on the
whorl (Geiger, 2012, p. 1123). Anatoma crispata (Flem-
ing, 1828) has a more globose shell with ne axial and
spiral sculpture and a wider sutsel. The most similar spe-
cies in shape and sculpture is A. aupouria (Powell, 1937)
from present-day New Zealand, but this species differs
in having a few spiral cords at the shoulder and within
the umbilicus.
Distribution – Upper Miocene (Tortonian): Atlantic, NW
France (this paper).
Superfamily Trochoidea Ranesque, 1815
Family Trochidae Ranesque, 1815
Subfamily Trochinae Ranesque, 1815
Tribe Trochini Ranesque, 1815
Genus Clanculus de Montfort, 1810
1810 Clanculus de Montfort, p. 191. Type species (by
original designation): Trochus pharaonius Lin-
naeus, 1758, present-day, Red Sea.
For generic synonymy see Ceulemans et al. (2016a).
Note – In this section we follow the terminology of Her-
bert (1993). The works of Chirli (2004) and Spadini (2006)
showed the Mediterranean Pliocene Clanculus fauna was
far more diverse than previously thought. Until now only
one species, Clanculus (Clanculopsis) baccatus (Defrance,
1824) was known from the middle and upper Miocene of
northwestern France. In this work we show this also to be
an underestimate of Clanculus diversity at the time.
Clanculus (s.s.) brebioni nov. sp.
Plate 15, g. 1; Plate 16, gs 1-2
Type material – Holotype NHMW 2016/0103/0246,
height 11.9 mm, width 12.3 mm; paratype 1 NHMW
2016/0103/0246, height 10.4 mm, width 11.1 mm; para-
type 2 MNHN.F.A66713, height 10.5 mm, width 11.4 mm
(incomplete); paratype 3 MNHN.F.A66714, height 9.4
mm, width 9.5 mm (incomplete).
Other material – St-Clément-de-la-Place: LC (1), FVD
(1).
Etymology – Named after Philippe Brébion of the
Muséum National d’Histoire Naturelle, Paris, in recogni-
tion of his work on the French Redonian assemblages.
Clanculus gender masculine.
Plate 14. Anatoma redoniana nov. sp.; 1. Holotype NHMW 2016/0103/1470, diameter 1.4 mm; height 0.9 mm; 1d, detail of proto-
conch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
92 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Clanculus (s.s) species of medium size, with
moderately elevated conical spire, bearing reticulated
sculpture on rst two teleoconch whorls, axial sculpture
that weakens abapically, so that sculpture on last two
whorls of low beaded cords, four primary cords on last
whorl with single smooth or beaded secondary cord in
each interspace, surface covered in extremely ne, close-
set axial growth lines and spiral threads forming tiny
rows of dots in interspaces, and strong apertural denti-
tion.
Description – Shell medium size for genus, solid, tro-
chiform, with a moderately elevated conical spire. Pro-
toconch not preserved. Teleoconch of ve whorls, with
periphery at abapical suture. Suture impressed. First
teleoconch whorl with two low spiral cords. On second
whorl a third cord develops below adapical suture. Nar-
row, close-set, prosocline ribs, visible in interspaces be-
tween cords, forming reticulated sculpture with small,
low, rounded tubercles at intersections. Abapically axial
sculpture weakens and disappears; spiral cords remain
low, bearing small beads. Last whorl bearing four rows
of primary beads, with a single secondary cord in inter-
spaces, developed to a varying degree; in holotype only
secondary cord between 3rd and 4th primaries beaded, in
paratype all secondary cords beaded. Surface covered
in extremely ne and close-set prosocline growth lines.
Fine spiral threads in interspaces between primary and
secondary spiral sculpture interrupted by growth lines
forming tiny spiral rows of dots. Base weakly convex,
bearing 6-7 beaded cords, equal in strength and spacing.
Umbilicus deep, surrounded by inductural callus, peri-
umbilical edge smooth bearing stout, irregular umbilical
dentition. Mature apertural dentition moderately strongly
developed; anal tooth stout, six labial ridges; upper colu-
mellar tooth stout; bid lower columellar tooth. Columel-
la between upper and lower teeth smooth. Parietal callus
Plate 15. 1. Clanculus (s.s.) brebioni nov. sp., holotype NHMW 2016/0103/0246, height 11.9 mm, width 12.3 mm. Le Grand Chau-
vereau, St-Clément-de-la-Place. 2. Clanculus (Clanculopsis) baccatus (Defrance, 1824), NHMW 2016/0103/0239, height 7.5
mm, width 7.9 mm. Renauleau. 3. Clanculus (Clanculopsis) sancticlementensis nov. sp., holot ype MNHN.F.A57700, height
8.9 mm, width 9.0 mm. Le Grand Chauvereau, St-Clément-de-la-Place. 4. Clanculus (Clanculopsis) umbilicovadus nov. sp.,
holotype NHMW 2016/0103/0248, height 4.1 mm, width 5.1 mm. Le Grand Chauvereau, St-Clément-de-la-Place. All: Maine-
et-Loire, NW France, Tortonian, upper Miocene.
Plate 16. Clanculus (s.s.) brebioni nov. sp.; 1. Paratype 1 NHMW 2016/0103/0246, height 10.4 mm, width 11.1 mm; 2. Holotype
NHMW 2016/0103/0246, height 11.9 mm, width 12.3 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW
France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 93
moderately developed, bearing 3-4 parietal ridges. Col-
our pattern of interrupted vertical ammules, dark dots
on many beads.
Discussion – Clanculus (s.s.) brebioni nov. sp. is the
largest shelled and most solid Clanculus species in the
Assemblage I fauna. We have attributed it to Clanculus
(s.s.), as the holotype has a bid lower columellar tooth.
We note that the paratype appears to have a single lower
columellar tooth, but this may be artefact, as the labial
ridges on the abapical half of the inside of the aperture
are abraded, the abraded area extending to the abapi-
cal portion of the lower columellar tooth. Although C.
(s.s.) brebioni has the same number of primary cords as
Clanculus (Clanculopsis) sancticlementensis nov. sp.,
in the latter the reticulated early sculpture persists fur-
ther along the teleoconch, the suture is canaliculated,
the cords are raised and the beading is more prominent.
There are also separated by the subgeneric character of
the lower columellar tooth; bid in Clanculus (s.s.), sim-
ple in C. (Clanculopsis). They also seem to have a differ-
ent colour pattern, with dark dots on many of the beads
in C. (s.s.) brebioni.
This new species is more closely similar to C. (s.s.) bon-
ttoi Chirli, 2004 from the Italian Pliocene, but that spe-
cies is smaller shelled (maximum height 7.7. mm, width
7.4 mm; Spadini, 2006), and the spiral sculpture forms
far more prominent beads; the beads in C. (s.s.) brebioni
are rather subdued for the genus and the adapically bead-
ed cord placed close below the suture is more strongly
leading to a canaliculated suture, as in C. (C.) sancticle-
mentensis, but not in C. (s.s.) brebioni. The Pliocene to
present day European C. (s.s.) corallinus (Gmelin, 1791)
differs in having more numerous beaded cords. The nu-
merous present-day Eastern Atlantic species revised by
Rubio & Rolán (2002) all differ in having more numer-
ous beaded cords of roughly equal strength on the last
whorl, rather than four primary cords above the basal
cord, with secondaries intercalated, as seen in C. (s.s.)
brebioni.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Subgenus Clanculopsis Monterosato, 1880
1880 Clanculopsis Monterosato, p. 222. Type species
(by subsequent designation, Sacco, 1896a, p. 21):
Trochus cruciatus Linnaeus, 1758, present-day,
Mediterranean.
Clanculus (Clanculopsis) baccatus (Defrance, 1824)
Plate 15, g. 2; Plate 17, gs 1-2
*1824 Monodonta baccata Defrance, 1824, p. 475.
1854 Monodonta Baccata Defr. – Millet, p. 157.
1964 Clanculus baccatus (Defrance, 1824) – Brébion,
p. 122.
2016a Clanculus (Clanculopsis) baccatus (Defrance, 1824)
– Ceulemans et al., p. 58, pl. 2, g. 4 (cum syn.).
Material and dimensions – Maximum height 11.0 mm,
width 10.9 mm. St-Clément-de-la-Place: RGM.1309556
(17), LC (5), FVD (5). Sceaux-d’Anjou: NHMW 2016/
0103/0242 (25), RGM.1309543 (10 +15 juveniles), RGM.
1309544 (50+), RGM.1309550 (15), RGM.1309573 (22),
RGM.1309580 (23 adults and subadults), LC (50+),
FVD (50+). Renauleau: NHMW 2016/0103/0239-
0240 (2), NHMW 2016/0103/0241 (27), LC (50+), FVD
(50+). Beug non: RGM.1309541 (2), RGM.1309542 (8
+ 10 juveniles), RGM.1309563 (17), RGM.1309594 (5),
RGM.1309743 (4), FVD (1).
Discussion – Clanculus (Clanculopsis) baccatus (De-
france, 1824) differs from the Pliocene to present-day C.
(s.s.) corallinus (Gmelin, 1791) by the presence of a single
lower columellar tooth; in C. (s.s.) corallinus the lower
columellar tooth is bid. The specimens of C. (C.) bac-
catus from Assemblage I have a second columellar tooth
below the strong single tooth, but it is far less strongly
developed than in C. corallinus. Clanculus (C.) baccatus
has six subequal primary beaded cords on the last whorl
above the base, plus a well-developed secondary cord be-
tween the adapical cord and the second cord, which is
almost equal in strength to the primary cords.
The specimens of C. (C.) baccatus from the Assemblage
I localities are small compared to those of the middle
Miocene Loire Basin of France (maximum height 10.9;
Plate 17. Clanculus (Clanculopsis) baccatus (Defrance, 1824); 1. NHMW 2016/0103/0239, height 7.5 mm, width 7.9 mm; 2. NHMW
2016/0103/0240, height 7.7 mm, width 7.8 mm. Renauleau, Maine-et-Loire, France, Tortonian, upper Miocene.
94 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Glibert, 1949, p. 67) and slightly higher-spired than those
from the lower Pliocene Assemblage III locality of Le
Pigeon Blanc, which are wider and more depressed (Ceu-
lemans et al., 2016a, pl. 2, g. 4). Clanculus (C.) baccatus
has six subequal primary beaded cords on the last whorl
above the base, plus a well-developed secondary cord be-
tween the adapical cord and the second cord.
Millet (1854) recorded this species from Assemblage
I localities of Sceaux-d’Anjou, Thorigné, Renauleau,
Saint-Michel and St-Clément-de-la-Place. Brébion (1964)
recorded this species from numerous localities, repre-
senting Assemblage I, III and IV. However, this is the
only Clanculus species he recorded from the French ‘Re-
donian’, which is shown here to be incorrect. Therefore,
his records must be veried.
For further discussion see Ceulemans et al. (2016a, p. 58).
Distribution – Middle Miocene: Atlantic (Langhian),
Loire Basin, France (Glibert, 1949). Upper Miocene: At-
lantic (Tortonian), NW France (Millet, 1854; Brébion,
1964). Lower Pliocene: Atlantic, NW France (Brébion,
1964; Ceulemans et al., 2016a). Upper Pliocene-lower
Pleistocene: Atlantic, NW France (Brébion, 1964).
Clanculus (Clanculopsis) sancticlementensis nov. sp.
Plate 15, g. 3; Plate 18, gs 1-4
Type material – Holotype MNHN.F.A57700, height 8.9
mm, width 9.0 mm; paratype 1 MNHN.F.A57700, height
8.5 mm, width 8.5 mm; paratype 2 MNHN.F.A57701,
height 9.6 mm, width 9.2 mm; paratype 3 NHMW
2016/0103/0244, height 8.1 mm, width 8.2 mm; paratype
4 RGM.1309546, height 8.7, width, 8.4 mm; paratype 5
RGM.1309547, height 86.5, width, 7.1 mm¸ paratype 6
RGM.1309548, height 6.6, width, 7.0 mm.
Other material – Maximum height 9.5 mm, width 9.3
mm. St-Clément-de-la-Place: NHMW 2016/0103/0245
(47), RGM.1309549 (25 juveniles), LC (50+), FVD (50+).
Etymology – Named after the type locality of St-Clé-
ment-de-la-Place. Clanculus gender masculine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Clanculus (Clanculopsis) species of medium
size, with moderatey elevated conical spire, bearing four
primary spiral cords on last whorl, crossed by narrow,
close-set axial ribs, visible in interspaces, small tubercles
developed at intersections and moderately weak aper-
tural dentition.
Description – Shell medium size for subgenus, solid, tro-
chiform, with moderately elevated conical spire. Proto-
conch consists of 1.5 smooth, convex whorls. Teleoconch
of four whorls, with periphery at abapical suture. Suture
impressed, canaliculate. First teleoconch whorl with two
elevated spiral cords. On second whorl a third cord de-
velops below adapical suture, gaining in strength. Third
whorl with three subequal spiral cords, fourth cord devel-
ops above abapical suture. About 28 narrow prosocline
ribs, elevated in interspaces between cords, forming small
Plate 18. Clanculus (Clanculopsis) sancticlementensis nov. sp.; 1. Holotype MNHN.F.A57700, height 8.9 mm, width 9.0 mm; 2.
Paratype 1 MNHN.F.A57700, height 8.5 mm, width 8.5 mm; 3. Paratype 2 MNHN.F.A57701, height 9.6 mm, width 9.2 mm;
4. Paratype 3 NHMW 2016/0103/0244, height 8.1 mm, width 8.2 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-
et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 95
rounded tubercles at intersections. Last whorl convex,
bearing four primary cords, with secondary cords de-
veloped in some of the interspaces, in some specimens.
Base weakly convex, bearing six smooth to beaded
cords, equal in strength and spacing. Umbilicus deep,
surrounded by inductural callus, periumbilical edge
smooth in most specimens; bearing weak umbilical den-
tition in some specimens. Mature apertural dentition
moderately weakly developed; anal tooth only slightly
stronger than labial ridges; upper columellar tooth
small, simple; single lower columellar tooth, with one
or two small tubercles below. Columella between upper
and lower teeth smooth. Parietal callus moderately de-
veloped, smooth or bearing 1-2 parietal ridges. Colour
pattern of interrupted vertical ammules is preserved in
most specimens.
Discussion – This new species is unequivocably placed
in the subgenus Clanculopsis Monterosato, 1880, char-
acterised by its single lower columellar tooth. It is with
surprise that we note that the vast majority of Clanculus
(Clanculopsis) specimens from St-Clément-de-la-Place
are not conspecic with those found in Renauleau and
Sceaux-d’Anjou, but belong to a distinct species; Clancu-
lus (Clanculopsis) sancticlementensis nov. sp. It differs
from Clanculus (Clanculopsis) baccatus (Defrance,
1824) in having only four primary spiral cords on the last
teleoconch whorl, whereas C. (C.) baccatus (Defrance,
1824) has six. Moreover, the mature apertural dentition,
even in the shells in which it is most strongly developed,
is always weaker than in C. (C.) baccatus. The sculpture
in C. (C.) sancticlementensis is similar to that of C. (s.s.)
bonttoi Chirli, 2004 from the Italian Pliocene, but that
species has a bid lower columellar tooth and therefore
belongs within Clanculus (s.s.). The numerous present-
day Eastern Atlantic species revised by Rubio & Rolán
(2002) all differ in having more numerous beaded cords
of roughly equal strength on the last whorl, rather than
four strong primary cords above the basal cord seen in C.
(C.) sancticlementensis.
We have so far only found C. (C.) sancticlementensis at
St-Clément-de-la-Place.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Clanculus (Clanculopsis) umbilicovadus nov. sp.
Plate 15, g. 4; Plate 19, g. 1
Type material – Holotype NHMW 2016/0103/0248, height
4.1 mm, width 5.1 mm.
Other material – Known only from holotype.
Etymology – Compound name from Latin ‘umbilicus’,
meaning navel and ‘vadus, -a, -um’, adjective meaning
shallow, describing the shallow umbilicus for the genus.
Clanculus gender masculine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Clanculus (Clanculopsis) species of small
size, with depressed trochiform shell shape, sculpture
composed of poorly developed narrow spiral cords and
crowded, strongly prosocline axial growth lines, shallow,
smooth-edged umbilicus, and weak apertural dentition.
Description – Shell small for subgenus, of medium thick-
ness, depressed trochiform, with low spire. Protoconch
consists of 1.5 smooth, convex whorls. Teleoconch of
three depressed whorls, with periphery at abapical suture.
Suture impressed, linear. First teleoconch whorl bearing
two low, narrow spiral cords. Abapically spiral sculpture
weakens further, four subobsolete cords on penultimate
whorl. Axial sculpture of very ne, close-set, strongly
prosocline growth lines covers entire shell. Last whorl
strongly convex, bearing four narrow, poorly developed
primary spiral cords, with even weaker secondary and
tertiary cords developed in interspaces. Base weakly
convex, bearing 11 smooth cords of irregular strength
and position, with secondaries developed in some inter-
spaces. Umbilicus shallow, surrounded by inductural
callus, periumbilical edge smooth, devoid of umbilical
dentition. Apertural dentition (?mature) weakly devel-
oped; anal tooth equal in strength to labial ridges; upper
columellar tooth small, simple; single lower columellar
tooth. Columella between upper and lower teeth smooth.
Parietal callus thickened, smooth.
Discussion – Although represented by a single incom-
Plate 19. Clanculus (Clanculopsis) umbilicovadus nov. sp.; 1. Holotype NHMW 2016/0103/0248, height 4.1 mm, width 5.1 mm. Le
Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
96 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
plete specimen, Clanculus (Clanculopsis) umbilicovadus
nov. sp. is unlike any of its European Eastern Atlantic or
Mediterranean fossil to present-day congeners in having
a shallow umbilicus. It is also the most weakly sculptured
of its congeners. The Pliocene to present-day C. (C.) jus-
sieui (Payraudeau, 1826) is most similar in having weak
sculpture and weak apertural dentition, but differs in
having a deep umbilicus.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Subfamily Cantharidinae Gray, 1857
Genus Jujubinus Monterosato, 1884
Type species (by subsequent designation, Crosse, 1885,
p. 140) – Trochus matonii Payraudeau, 1826 [= Jujubi-
nus exasperatus (Pennant, 1777), present-day, Mediter-
ranean.
1884a Jujubinus Monterosato, p. 109
(For generic synonymy see Ceulemans et al., 2016a, p.
59).
Jujubinus coronatus nov. sp.
Plate 20, gs 1-3
Type material – Holotype MNHN.F.A57695, height 4.3
mm, width 3.9 mm; paratype 1 NHMW 2016/0103/0189,
height 4.8 mm, width 4.3 mm; paratype 2 NHMW
2016/0103/0190, height 4.6 mm, width 3.9 mm; para type
3 NHMW 2016/0103/0191, height 4.7 mm, width 3.9 mm;
paratype 4 NHMW 2016/0103/0192 (juvenile).
Other material – St-Clément-de-la-Place: NHMW
2016/0103/0193 (21), LC (17), FVD (15).
Etymology – Latin ‘coronatus’, meaning crowned, refer-
ring to the prominently beaded adapical cord. Jujubinus
gender masculine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis –Jujubinus species of small size, with cyrto-
conoid spire, sculptured by spiral cords, adapical cord
coarsely beaded giving whorls coronate appearance, and
peripheral cord on last whorl stronger, every fourth bead
further strengthened to give periphery crenulated ap-
pearance, intermediate cords narrow, non-beaded or 1-2
cords adjacent to sutures nely beaded.
Description – Shell small, of medium thickness, conical,
with moderately high cyrtoconoid spire, attened base.
Protoconch paucispiral, composed of 1.3 smooth whorls
(dp = 320 µm, dn = 135 µm). Teleoconch consisting of
about ve straight-sided to weakly concave whorls, with
periphery at abapical suture. Suture linear, impressed.
Sculpture on spire whorls of one coarsely beaded broader
adapical cord placed immediately below suture, giving
whorls coronate appearance. Below lie narrow subequal
Plate 20. Jujubinus coronatus nov. sp.; 1. Holotype MNHN.F.A57695, height 4.3 mm, width 3.9 mm; 2. Paratype 4 NHMW
2016/0103/0192, detail of protoconch (SEM image); 3. Paratype 1 NHMW 2016/0103/0189, height 4.8 mm, width 4.3 mm. Le
Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 97
cords separated by narrow grooves, six on penultimate
whorl; cords non-beaded or 1-2 cords closest to sutures
nely-beaded. Last whorl with coronate subsutural cord,
6-8 narrow cords below, the abapical two cords strength-
ened to form peripheral band delimiting base and become
beaded, every fourth bead further strengthened to give
periphery crenulated appearance. Base imperforate, at-
tened, bearing about 14 concentric cords interrupted by
prosocline axial growth lines. Aperture tangential; peri-
stome discontinuous, outer lip not thickened, angled at
periphery, smooth within. Columella straight, bearing
a weak denticle mid-columella. Colour pattern is pre-
served, consisting of whitish blotches placed below the
coronate adapical cord and mid-whorl (Pl. 20, g. 3), in
some specimens coalescent to form almost continuous
white band (Pl. 20, g. 1) and white blotches at periphery
of last whorl coinciding with crenulations.
Discussion – This species is placed in the genus Jujubi-
nus Monterosato, 1884 based on its smooth paucispiral
protoconch (Pl. 20, g. 2) and the presence of a small
columellar tooth. Jujubinus coronatus nov. sp. is sepa-
rated from all of its European Miocene to present-day
congeners by the character of its coronate spire whorls.
A swollen or beaded abapical cord is not uncommonly
developed in members of this genus [i.e. the present day
J. montagui (W. Wood, 1828); J. exasperatus (Pennant,
1777); J. striatus (Linnaeus, 1758)], but we can nd no
other congener with a beaded adapical cord. The most
similar species is Jujubinus redoniensis nov. sp., from the
same assemblage, which also has a beaded adapical cord;
for comparison see under that species.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Jujubinus aff. exasperatus (Pennant, 1777)
Plate 21, gs 1-4
aff.*1777 Trochus exasperatus Pennant, p. 126.
aff. 1923 Trochus (Calliostoma) exasperatus (Pennant) –
Harmer, p. 723, pl. 58, g. 18.
1964 Calliostoma cf. exasperatum Pennant, 1777 – Bré-
bion, p. 77.
aff. 2003 Jujubinus exasperatus (Pennant, 1777) – Landau
et al., p. 42, pl. 9, g. 6.
aff. 2004 Jujubinus exasperatus (Pennant, 1777) – Chirli, p.
79, pl. 32, gs 10-12, pl. 33, gs 1-3 (?non g 4).
Material and dimensions – St-Clément-de-la-Place:
Maximum height 6.3 mm, width 4.8 mm. NHMW
2016/0103/0234-0237 (4), NHMW 2016/0103/0238 (25),
LC (20), FVD (17). Sceaux-d’Anjou: RGM.1309732 (2).
Discussion – We echo Brébion’s (1964, p. 77) problem
in the identication and placement of this small spe-
cies. Whilst Brébion is correct in saying that the aper-
tural characters are not those of Jujubinus Monterosato,
1884 – i.e., that they lack the columellar fold or swelling
– some of our specimens do have a weak swelling mid-
columella, or just below (Pl. 21, g. 1a). We note that the
other small Jujubinus species from the Assemblage I
localities also have the columellar swelling poorly deve-
loped; i.e., J. coronatus nov. sp. and J. redoniensis nov.
sp. More importantly, the paucispiral protoconch (Pl. 21,
g. 4) lacks the honeycomb surface sculpture that is so
characteristic to the genus Calliostoma Swainson, 1840.
Therefore placement in Jujubinus is more appropriate.
Jujubinus aff. exasperatus is smaller shelled than the
Pliocene to present day J. exasperatus (maximum height
6.3 mm vs. 9-15 mm for present-day specimens; Gian-
nuzzi-Savelli et al., 2004). Both the adapical and abapi-
Plate 21. Jujubinus aff. exasperatus (Pennant, 1777); 1. NHMW 2016/0103/0234, height 6.0 mm; 2. NHMW 2016/0103/0235, height
4.9 mm; 3. NHMW 2016/0103/0236, height 5.3 mm; 4. NHMW 2016/0103/0237, detail of protoconch (SEM image). Le Grand
Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
98 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
cal cords are beaded, the abapical more strongly so, in
some specimens leading to a crenulated periphery to the
last whorl. The cords between these are smooth to nely
beaded. This type of sculpture could t within the enor-
mous sculptural variability seen in the present-day speci-
mens (Giannuzzi-Savelli et al., 2004, gs 289-302). Some
colour pattern is preserved in the fossil material, with
white blotches present at the periphery, but this is not an
uncommon colour pattern in Jujubinus species and not
helpful as a specic character. Indeed the intraspecic
colour pattern in J. exasperatus itself is highly variable
(Giannuzzi-Savelli et al., 2004, gs 289-302).
However, we doubt they are conspecic. The Assemblage
I specimens are smaller, a feature common to many As-
semblage I taxa and we hesitate to use that on its own
to differentiate species, but they also have the columel-
lar swelling far less well developed and the crenulation
of the periphery of the last whorl is a feature we have
not seen in J. exasperatus. Nevertheless, we refrain from
erecting a new taxon based on this material.
Brébion (1964, p. 78) recorded this species from Assem-
blage I localities of Renauleau and Sceaux-d’Anjou, to
which we add St-Clément-de-la-Place.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Brébion, 1964).
Jujubinus proximus (Millet, 1865)
Plate 22, gs 1-3
1854 Trochus Proximus Millet, p. 157 (nomen nudum).
*1865 Trochus proximus Millet, p. 582.
1964 Calliostoma proximum Millet 1854 [sic] – Bré-
bion, p. 93, pl. 2, g. 15.
Type material – Syntypes: Thorigné, Sceaux-d’Anjou and
Renauleau, Musée d’Angers, France (de Brébion, 1964,
p. 93).
Material and dimensions – Maximum height 21.1 mm;
width 15.1 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0149 (1), NHMW 2016/0103/0151 (40), RGM.1309674
(3), RGM.1309760 (6), LC (50+), FVD (50+). Sceaux-
d’Anjou: NHMW 2016/0103/0153 (50+), RGM.1309650
(18), RGM.1309695 (9), RGM.1309714 (13), RGM.1309725
(26), RGM.1309774 (27), RGM.1309779 (20), LC (50+),
FVD (50+). Renauleau: NHMW 2016/0103/0154-0155
(2), NHMW 2016/0103/0152 (50+), LC (50+), FVD (50+).
Beugnon: RGM.1309676 (5), LC (4), FVD (1).
Original description – ‘Trochus proximus, Millet. Coq.
en cône assez allongé, composée de 8-10 tours de spire
aplatis, chacun d’eux recouvert par quatre lets légère-
ment perlés, dont celui de la base et celui du sommet sont
plus gros et séparés par la suture. Dessous à peine con-
cave, couvert de stries non perlées. Hauteur: 15-16 mil-
limètres; diamètre à la base: 12 millimètres. Th., Sc, Ren.
(Millet, 1865, p. 582)’
Revised description – Shell of medium size and thick-
ness, with high regularly conical spire. Protoconch pau-
cispiral, surface smooth. Teleoconch consisting of about
seven weakly convex to straight-sided whorls, with pe-
Plate 22. Jujubinus proximus (Millet, 1865); 1. NHMW 2016/0103/0154, height 10.0 mm, diameter 6.5 mm; 2. NHMW 2016/0103/0155,
height 10.1 mm, width 6.1 mm. Le Renauleau. 3. NHMW 2016/0103/0149, height 10.1 mm, width 7.3 mm. Le Grand Chauvereau,
St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 99
riphery just above abapical suture. Suture linear, super-
cial, Sculpture of ve beaded cords of subequal strength,
with a secondary beaded cord between cords 1 and 2 on
the last whorl in some specimens. Last whorl roundly
angled at periphery. Base not sharply delimited, bearing
about 12 similarly beaded, occasionally smooth cords.
Aperture tangential; peristome discontinuous, outer lip
not thickened, rounded at periphery, bearing an internal
ridge bordering the broad, shallow, rounded siphonal ca-
nal. Columella sloping abaxially, bearing a weak colu-
mellar tooth.
Discussion – As Jujubinus proximus (Millet, 1865) is one
of the most abundant Jujubinus species in the Assem-
blage I fauna and the original description is incorrect in
certain details, we offer a revised description. The most
important error in Millet’s text is in describing four cords
per whorl, whereas, as commented by Brébion in his re-
vised description (1964, p. 93), it has ve. Both Millet and
Brébion described the basal cords as smooth, but in some
secimens they are weakly beaded (Pl. 22, g. 3c).
This species could be confused with Calliostoma deshay-
esi (Mayer, 1862), described from the older middle Mio-
cene deposits of the Loire basin. They share the same tall
conical shell shape (Pl. 22, gs 1, 2), although the apical
angle can vary slightly and seems slightly wider on aver-
age than the Loire Basin shell, with some considerably
broader shells also present (Pl. 22, g. 3). However, the
Assemblage I species differs in having ve beaded spiral
cords as opposed to the four described by Glibert (1949, p.
42) for the Loire Basin specimens, the base has more nu-
merous concentric cords and the columella slopes abaxi-
ally, whereas in C. deshayesi it is almost vertical. Most
importantly, the Assemblage I species does not belong
within the genus Calliostoma, as it has a smooth and not
honeycomb-sculptured protoconch and the apertural char-
acters; columellar tooth and internal ridge bordering the
siphonal canal, are characters we associate with the genus
Jujubinus. We have not seen specimens of C. deshayesi
from the Loire Basin and Glibert gives no description of
the protoconch, but based on the shell illustrated, place-
ment in the genus Calliostoma is probably correct.
Jujubinus proximus is abundant and the largest Jujubinus
species in the Assemblage I deposits. It is particularly
common at Renauleau, where it reaches a greater maxi-
mum size than at the other localities. There is some con-
siderable degree of intraspecic variability in slender-
ness and basal angulation. The presence of ve primary
cords per whorl is consistent, although an occasional spi-
ral thread is intercalated in the interspaces; these second-
ary cords gaining in strength in gerontic specimens and
almost equal to the primary cords. In the very occasional
specimen the spiral cords are weakly beaded or almost
smooth.
Millet (1854, 1865) and Brébion (1964, p. 31) recorded
this species from the Assemblage I localities of Thorigné,
Sceaux-d’Anjou and Thorigné.
Distribution – Upper Miocene: Atlantic, Tortonian,
Messinian, NW France (this paper).
Jujubinus cf. proximus (Millet, 1865)
Plate 23, g. 1
cf.*1865 Trochus proximus Millet, p. 582.
Matrial and dimensions – NHMW 2016/0103/0194, height
14.6 mm, width 9.5 mm, Le Grand Chauvereau, St-Clé-
ment-de-la-Place, Maine-et-Loire, NW France.
Plate 23. Jujubinus cf. proximus (Millet, 1865); 1. NHMW
2016/0103/0194, height 14.6 mm, width 9.5 mm; Le Grand
Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW
France, Tortonian, upper Miocene.
Description – Shell of medium size and thickness, with
high conical spire. Protoconch paucispiral, surface
abraded. Teleoconch consisting of about eight weakly
convex whorls, with periphery just above abapical su-
ture. Suture linear, supercial, Sculpture of nely beaded
cords of irregular strength, with an occasional secondary
beaded cord intercalated, 6-7 primary cords on penulti-
mate whorl. Last whorl slightly concave adapically, swol-
len and rounded at periphery. Base not sharply delimited,
bearing similar beaded sculpture. Aperture tangential;
peristome discontinuous, outer lip not thickened, round-
ed at periphery, bearing an internal ridge bordering the
broad, shallow, rounded siphonal canal. Columella deep-
ly excavated in mid-portion, bearing single, prominent,
basal columellar tooth.
Discussion –The shape is closely similar to that of the
largest specimens of Jujubinus proximus (Millet, 1865),
except that the last whorl is somewhat swollen abapically,
but this might be due to abnormal repair from crab attack
(Pl. 23, g. 1b). The rst two whorls have ve primary spi-
ral cords, like J. proximus, but a secondary cord appears
on the third whorl that gains in strength to become equal
to the primary cords and further cords appear abapically.
On later whorls the cords are subsequently narrower and
more crowded than in J. proximus. The columellar tooth
is strongly developed and the siphonal canal prominently
ridged within the aperture, both more strongly developed
than usual for J. proximus. We have insufcient material
100 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
to conclude whether this is an abnormal specimen of J.
proximus or a distinct species.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Jujubinus redoniensis nov. sp.
Plate 24, gs 1-2
1964 Calliostoma couffoni Brébion, p. 90, pl. 2, g. 11
(nomen nudum).
Type material – Holotype MNHN.F.A57696, height 4.8 mm,
width 3.6 mm; paratype 1 MNHN.F.A57697, height 4.8 mm,
width 3.6 mm; paratype 2 NHMW 2016/ 0103/ 0205, height
5.0 mm, width 3.8 mm; paratype 3 NHMW 2016/ 0103/ 0206,
height 4.8 mm, width 3.4 mm; paratype 4 NHMW 2016/
0103/0207 (juvenile); paratype 5 RGM. 1309667, height 5.6
mm, width 4.4 mm; paratype 6 RGM. 1309668, height 6.5
mm, width 4.3 mm; paratype 7 NHMW 2016/ 0103/ 0254,
height 7.8 mm, width 6.2 mm; paratype 8 NHMW 2016/
0103/0255, height 7.0 mm, width 5.7 mm.
Other material – Maximum height 6.1 mm, width 4.5
mm. Sceaux-d’Anjou: NHMW 2016/0103/0208 (35), RGM.
1309669 (6); LC (20), FVD (26).
Etymology – Named after the ‘Redonian’ stage, the name
used until recently for these NW French post-middle Mi-
ocene assemblages. Jujubinus gender masculine.
Locus typicus – La Presselière, Sceaux-d’Anjou, Maine-
et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Jujubinus species of small size, with a regu-
larly conical shell shape, nely reticulated sculpture on
the rst two teleoconch whorls, later whorls bearing a
single beaded adapical cord, smooth cords below, abapi-
cal cord more strongly developed, a attened base bear-
ing eight irregular cords, and a very weak columellar
swelling.
Description – Shell small, of medium thickness, with
tall regularly conical spire. Protoconch paucispiral, com-
posed of 1.5 smooth whorls. Teleoconch consisting of
about ve straight-sided whorls, with periphery at abapi-
cal suture. Suture linear, impressed. Sculpture on rst
teleoconch whorls consists of two spiral cords, three on
second whorl, crossed by narrow prosocline axial ribs,
forming reticulated surface sculpture with small tuber-
cles developed at intersections. On third whorl axial
ribs disappear, sculpture consisting of a beaded adapi-
cal cord, beads close-set and of medium strength, cords
below smooth, rounded, three on third whorl, four on
penultimate, the abapical cord more strongly developed.
Last whorl with beaded adapical cord, three cords below
to roundly angular periphery, formed by two stronger
cords. Base attened, imperforate, bearing eight cords
of irregular strength, tending to be subobsolete mid-base
and stronger towards centre. Aperture tangential; peri-
stome discontinuous, outer lip not thickened, angled at
periphery, smooth within. Columella straight, bearing
very weak thickening mid-columella.
Discussion – Jujubinus redoniensis nov. sp. belongs to
a group of small Jujubinus species present in the As-
semblage I deposits with weak columellar dentition for
the genus. This species is characterised by its crowded
narrow cords, beaded on early whorls, smooth, except
for the adapical cord, on later whorls, and its swollen
abapical cord. The regularly conical shell shape is con-
stant, although the apical angle is slightly broader in
some specimens and the usually acute angle at the base
is slightly more rounded in some specimens. It is most
similar to J. coronatus nov. sp. (see above) in having a
beaded adapical cord, but differs in 1) having a regularly
conical instead of cyrtoconoid spire, 2) having nely re-
ticulated sculpture on the rst two teleoconch whorls, 3)
although both have a beaded adapical cord, the beads in
J. coronatus are stronger and more widely spaced, 4) the
spiral cords below are less numerous, broader, stronger
and rounded in J. redoniensis and 5) in having fewer ba-
sal cords.
Jujubinus aff. exasperatus (Pennant, 1777) differs in hav-
ing the ad- and abapical cords more strongly beaded, which
perisists onto the late adult whorls and having fewer
Plate 24. Jujubinus redoniensis nov. sp.; 1. Holotype MNHN.F.A57696, height 4.8 mm, width 3.6 mm; 2. Paratype 4 NHMW
2016/0103/0207, detail of protoconch (SEM image). La Presselière, Sceaux-d’Anjou, Maine-et-Loire, NW France, Tortonian,
upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 101
cords that are again wider than their interspaces. Juju-
binus striatus (Linnaeus, 1758) has fewer cords, again
wider than their interspaces. Jujubinus sceauxensis nov.
sp. differs in having nely beaded sculpture.
Brébion (1964, p. 90) recorded this species from several
Assemblage I localities (Sceaux-d’Anjou, Renauleau, St-
Clément-de-la-Place).
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Brébion, 1964).
Jujubinus sceauxensis nov. sp.
Plate 25, gs 1-3
1964 Jujubinus (Strigosella) andegavensis Brébion, p.
121, pl. 3, g. 13 (nomen nudum).
Type material – Holotype NHMW 2016/0103/0676, height
4.0 mm, width 4.0 mm; paratype 1 NHMW 2016/ 0103/0197,
height 7.7 mm, width 6.1 mm; paratype 2 NHMW 2016/
0103/0669, height 6.5 mm, width 6.5 mm; paratype 3
NHMW 2016/0103/0671, height 5.1 mm, width 5.3 mm;
paratype 4 NHMW 2016/0103/0200, height 6.7 mm, width
6.3 mm; paratype 5 RGM.1309653, height 7.4 mm, width
6.0 mm; paratype 6 RGM.1309654, height 7.6 mm, width
6.4 mm; paratype 7 RGM.1309720, height 7.3 mm, width
6.3 mm; paratype 8 MNHN.F.A57937, height 6.7 mm,
width 5.9 mm; paratype 9 MNHN.F.A57938, height 4.9
mm, width 4.8 mm.
Other material – St-Clément-de-la-Place: LC (2), FVD
(5). Sceaux-d’Anjou: NHMW 2016/0103/0204 (9), NHMW
2016/0103/0687 (6), RGM.1309655 (1), RGM. 1309687 (4),
RGM.1309713 (2), RGM.1309716 (1), RGM. 1309721 (6),
RGM.1309729 (4), RGM.1309769 (9), RGM.1348009 (2),
FVD (6).
Etymology – Named after the type locality of Sceaux-
d’Anjou. Jujubinus gender masculine.
Locus typicus – La Presselière, Sceaux-d’Anjou, Maine-
et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Jujubinus species of small size, with pointed
apex, low cyrtoconoid spire, sculptured by nine beaded
spiral cords of equal strength, sharply angled last whorl,
attened base bearing 14 cords and narrow, shallow um-
bilicus.
Description – Shell small, solid, with pointed apex and
moderately low cyrtoconoid spire. Protoconch paucispi-
ral; surface abraded, but seemingly smooth. Teleoconch
consisting of ve straight-sided whorls, with periphery at
abapical suture. Suture linear, supercial. Sculpture con-
sists of about nine, narrow, close-set, beaded spiral cords
of equal strength. Last whorl sharply angled at base. Base
attened, bearing about 14 concentric cords interrupted
by prosocline axial growth lines, with a shallow, narrow,
Plate 25. Jujubinus sceauxensis nov. sp.; 1. Holotype NHMW 2016/0103/0676, height 4.0 mm, width 4.0 mm; 2. Paratype 1
NHMW 2016/0103/0197, height 7.7 mm, width 6.1 mm; 3. Paratype 2 NHMW 2016/0103/0669, height 6.5 mm, width 6.5 mm;
4. Parat ype 3 NHMW 2016/0103/0671, height 5.1 mm, width 5.3 mm. La Presselière, Sceaux-d’Anjou, Maine-et-Loire, NW
France, Tortonian, upper Miocene.
102 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwester n France, 1. Patellogastropoda and Vetigastropoda
smooth umbilicus. Aperture strongly tangential; peris-
tome discontinuous, outer lip not thickened, angled at
periphery, smooth within. Columella oblique, bearing a
broad thickening below mid-columella.
Discussion – Jujubinus sceauxensis nov. sp. is a very pe-
culiar little species. The nely beaded sculpture makes
it supercially similar to some of the nely beaded cal-
liostomiids found in the Assemblage I deposits of NW
France, like Calliostoma gratiosum (Millet, 1865) and
C. microgemmatum nov. sp., but the apertural characters
show it to be a canthariid rather than a calliostomiid. The
protoconch is abraded in all specimens, but seems to be
smooth, devoid of the typical honeycomb pattern seen in
Calliostoma species. There is no European fossil or ex-
tant species with which this new species can be usefully
compared.
Brébion (1964, p. 122) recorded this species from the As-
semblage I localities of Sceaux-d’Anjou and Renauleau.
Despite intensive collecting at Renauleau, we have only
found it at St-Clément-de-la-Place and Sceaux-d’Anjou.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Jujubinus striatus (Linnaeus, 1758)
Plate 26, gs 1-2
*1758 Trochus striatus Linnaeus, p. 759.
2016a Jujubinus striatus (Linnaeus, 1758) – Ceulemans
et al., p. 62, pl. 4, g. 5 (cum syn.).
Material and dimensions – St-Clément-de-la-Place:
Maximum height 6.1 mm, width 5.1 mm. NHMW
2016/0103/0127-0129 (3), NHMW 2016/0103/0130 (17),
LC (15), FVD (15).
Discussion – Like the specimens from the Assemblage III
locality of Le Pigeon Blanc, the specimens of Jujubinus
striatus (Linnaeus, 1758) from St-Clément-de-la-Place
are small compared to those found in European seas to-
day (7-13 mm in height; Giannuzzi-Savelli et al., 1994,
gs 303-314). Colour pattern is preserved in some speci-
mens consisting of darker dots at the suture and vertical
ammules, also seen in some present day specimens (i.e.
Giannuzzi-Savelli et al., 1994, g. 311). The whorl coiling
is somewhat disjunct in some specimens as seen in the
series illustrated (Pl. 26, g. 1). As far as we are aware,
this is the stratigraphically oldest record for the species.
For further discussion see Ceulemans et al. (2016a, p. 6).
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper). Lower Pliocene: Atlantic, NW France
(Ceulemans et al., 2016a), Guadalquivir Basin, S. Spain
(Landau et al., 2011); western Mediterranean, NE Spain
(Martinell, 1978); central Mediterranean, Italy (Chirli,
2004). Upper Pliocene: Atlantic, Red Crag, England
(Harmer, 1923); western Mediterranean, Estepona, S.
Spain (Landau et al., 2003); central Mediterranean, Italy
(Spadini, 1986; Cavallo & Repetto, 1992). Pleistocene: At-
lantic, Britain (Harmer, 1923); The Netherlands (Pou wer
& Wesselingh, 2012); western Mediterranean, Balearic
Islands, (Cuerda Barceló, 1987); central Mediterranean,
Italy (Malatesta, 1960; Taviani et al., 1998). Present-day:
Atlantic, Isle of Man to Canaries, Madeira, Azores and
Mediterranean, from the extreme low tide to 200m deep
on seaweeds and small stones (Poppe & Goto, 1991).
Genus Colliculus Monterosato, 1888
1888 Colliculus Monterosato, p. 171. Type species (sub-
sequent designation, Bucquoy, Dautzenberg &
Doll fus, 1898, p. 773): Trochus adansonii Payrau-
deau, 1826, present-day, Mediterranean.
For generic synonymy see Ceulemans et al. (2016a).
Note – We follow Lozouet et al. (2001, p. 18) in giving
full genus rank to Colliculus Monterosato, 1888. For fur-
ther discussion see Ceulemans et al. (2016a).
Colliculus biangulatus (Eichwald, 1830)
Plate 27, g. 1
*1830 Trochus biangulus Eichwald, p. 221.
Plate 26. Jujubinus striatus (Linnaeus, 1758); 1. NHMW 2016/0103/0127, height 5.5 mm, width 4.3 mm; 2. NHMW 2016/0103/0128,
height 5.5 mm, width 3.5 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper
Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 103
1856 Trochus biangulus Eichw. – Hörnes, p. 460, pl. 45,
g. 15.
non 1917 Gibbula (Colliculus) biangulata (Eichwald) – Coss-
mann & Peyrot, p. 124, pl. 4, gs 20-22 [=Collicu-
lus aquitanicus (Cossmann & Peyrot, 1917)].
1928 Gibbula biangulata Eichw. – Friedberg, p. 486, pl.
30, g. 20.
1949 Gibbula biangulata Eichwald, 1830 – Glibert, p.
58, pl. 3, g. 8.
1964 Gibbula (Colliculus) biangulata Eichwald, 1830 –
Brébion, p. 105.
1966 Gibbula biangulata Eichwald, 1830 – Strausz
(?partim), p. 35, pl. 52, gs 21-23 (?pl. 53, gs
1-3).
1975 Gibbula (Gibbula) cf. varia (Linnaeus) – Bałuk, p.
35, pl. 3, g. 1.
1975 Gibbula (Gibbula) cf. varia (Linnaeus, 1766 [sic])
– Bałuk, p. 35, pl. 3, g. 1.
2006 Gibbula (Gibbula) varia (Linnaeus, 1766 [sic]) –
Bałuk, p. 35, pl. 3, g. 1 [non Gibbula varia (Lin-
naeus, 1758)].
Material and dimensions – Maximum height 8.4 mm,
width 7.8 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0036 (1), NHMW 2016/0103/0037 (50+), RGM.
1309524 (50+), RGM.1309532 (30), RGM.1309591 (1),
LC (50+), FVD (50+). Sceaux-d’Anjou: NHMW 2016/
0103/0038 (8), RGM.1309509 (5), LC (2), FVD (13). Re-
nauleau: NHMW 2016/0103/0117 (50+), LC (50+), FVD
(50+). Beugnon: RGM.1309533 (1), RGM.1309742 (20),
FVD (1).
Discussion – Colliculus biangulatus (Eichwald, 1830),
which is widespread in the middle and upper Miocene
NW French assemblages, is most like the older Atlan-
tic lower Miocene C. aquitanicus (Cossmann & Peyrot,
1917) from the Aquitaine Basin of France, but lower-
spired and more strongly biangular than the Aquitanian
species, with narrower and more numerous spiral cords
between the angulations.
Brébion (1964, p. 106) recorded this species from numer-
ous Assemblage I deposits (Sceaux-d’Anjou, St-Michel,
Les Pierres Blanches, Thorigné, St-Clément-de-la-Place).
Distribution – Middle Miocene: Atlantic, Loire Basin,
France (Glibert, 1949); Paratethys, Austria (Hörnes,
1856), Hungary (Strausz, 1966), Poland (Friedberg, 1928;
Bałuk, 1975, 2006). Upper Miocene: Atlantic (Tortoni-
an), NW France (Brébion, 1964).
Colliculus insignis (Millet, 1854)
Plate 28, g. 1
*1854 Trochus Insignis Millet, p. 156.
1865 Trochus insignis Millet – Millet, p. 582.
1917 Gibbula (Colliculus) sosensis Cossmann & Pey-
rot, p. 127, pl. 4, gs 33-36.
1964 Gibbula (Colliculus) sosensis Cossmann & Pey-
rot, 1916 – Brébion, p. 106, pl. 2, gs 34, 35.
Material and dimensions – Maximum height 7.3 mm,
width 8.7 mm. St-Clément-de-la-Place: NHMW 2016/
Plate 27. Colliculus biangulatus (Eichwald, 1830); 1. NHMW 2016/0103/0037, height 5.7 mm, width 5.5 mm. Le Grand Chauvereau,
St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Plate 28. Colliculus insignis (Millet, 1854); 1. NHMW 2016/0103/0039, height 5.5 mm, width 5.9 mm. Le Grand Chauvereau, St-
Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
104 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigast ropoda
0103/0039 (1), NHMW 2016/0103/0040 (50+), RGM.
1309600 (4), RGM.1309618 (1), LC (50+), FVD (50+).
Sceaux-d’Anjou: NHMW 2016/0103/0041 (50+), RGM.
1309510 (37), RGM.1309511 (50+), RGM.1309516 (39),
RGM.1309518 (50+), RGM.1309726 (5), RGM.1347852
(45), RGM.1347916 (12), RGM.1347996 (47), LC (50+),
FVD (50+). Renauleau: NHMW 2016/0103/0116 (50+),
LC (50+), FVD (50+). Beugnon: NHMW 2016/0103/0042
(3), RGM.1309527 (1), RGM.1309537 (34), RGM.1309738
(13), RGM.1309539 (2), RGM.1309629 (6).
Discussion – This species is easily recognised by its
sculpture composed of strongly developed shoulder and
peribasal cords, which make the whorl prole sharply bi-
angular, with two weaker cords running between these
major cords. The axial sculpture of close-set lamellar
growth lines, which cross the ribs elevated, give the
surface a somewhat scabrous appearance. However, the
name Gibbula (Colliculus) sosensis Cossmann & Peyrot,
1917 must give way to the older name Trochus insignis
Millet, 1854. The original description highlights the
strong spiral sculpture: ‘Insignis, Millet. – Sceaux, Tho-
rigné. – Cette espèce a quelques rapports avec le genre
Dauphinula [incorrect spelling of Delphinula Gray,
1847], par sa bouche, dont les bords sont comme réunis,
à raison de la décurrence des bords columellaires (1854,
p. 156).’ This description was later enlarged: ‘Trochus
insig nis, Millet. Coq. petite, courte, composée de 5 tours
de spire, chacun d’eux creusé en gouttière spirale est en-
touré de quatre liserets arrondis, dont deux plus minces,
placés au centre. Dessous bombé, couvert de nes stries.
Ombilic lisse, assez profond. Hauteur et diamètre, 6 à 7
millimètres. Sceaux, Thorigné, Reneauleau (Millet, 1865,
p. 582)’. The description leaves little doubt that it refers to
this species. We cannot satisfy the requirements of Arti-
cle 23.9.1.2 (ICZN 1999a) to consider Millet’s name a no-
men oblitum. Therefore Colliculus insignis (Millet, 1854)
must take priority over Colliculus sosensis (Cossmann &
Peyrot, 1917).
The most similar species is Colliculus neraudeaui Ceu-
lemans, Van Dingenen & Landau, 2016, from the lower
Pliocene Assemblage III locality of Le Pigeon Blanc,
Loire-Atlantique, NW France, but C. insignis differs in
being smaller-shelled, in having two cords between the
stronger shoulder and basal cords instead of one in C.
neraudeaui, in having more numerous but ner cords on
the base and a narrower, shallower umbilicus. The lamel-
lar axial growth lines seen in C. insignis seem to be ab-
sent in C. neraudeaui, although the surface of the latter,
which is known from a single complete specimen plus
several fragments, is somewhat abraded.
Millet (1854, 1865) recorded this species from the As-
semblage I localities of Sceaux-d’Anjou, Renauleau and
Thorigné, to which Brébion (1964, p. 107) added St-Clé-
ment-de-la-Place.
Distribution – Middle Miocene: Atlantic, Aquitaine Ba-
sin (Cossmann & Peyrot, 1916). Upper Miocene: Atlan-
tic (Tortonian), NW France (Millet, 1854, 1865; Brébion,
1964).
Genus Gibbula Risso, 1826
1826 Gibbula Risso, p. 134. Type species (by subse-
quent designation, Herrmannsen, 1847, p. 473):
Trochus magus Linnaeus, 1758, present-day, Med-
iterranean.
Gibbula brebioni nov. sp.
Plate 29, gs 1-2
Type material – Holotype MNHN.F.A57689, height 8.7
mm, width 7.5 mm; paratype 1 NHMW 2016/0103/0108,
height 9.7 mm, width 8.0 mm; paratype 2 NHMW
2016/0103/0109, height 8.2 mm, width 8.0 mm; paratype
3 NHMW 2016/0103/0110, height 9.5 mm, width 8.1 mm;
paratype 4 RGM.1309753, height 7.4 mm, width 6.4 mm;
paratype 5 RGM.1309772, height 7.4 mm, width 6.7 mm.
Other material – St-Clément-de-la-Place: NHMW
2016/ 0103/ 0111 (10 + 50 juveniles), RGM.1309754 (10),
RGM. 1309773 (9), LC (11), FVD (14). Sceaux-d’Anjou:
NHMW 2016/ 0103/0216 (8), RGM.1309657 (4), FVD (2).
Re nau leau: NHMW 2016/0103/1431 (1), LC (1).
Etymology – Named after Philippe Brébion of the
Muséum National d’Histoire Naturelle, Paris, in recogni-
tion of his work on the French Redonian assemblages.
Gibbula gender feminine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Gibbula species of small size, with regularly
conical spire, shallow suture, sculpture of ne, close-set
spiral cords, much narrower than their interpaces, axial
sculpture restricted to prosocline growth lines, and weak
columellar fold.
Description – Shell small, trochiform, with regularly
conical spire. Protoconch paucispiral, composed of 1.5
smooth whorls. Teleoconch of 5.5 whorls, prole of early
whorls convex, weaker so abapically, periphery at abapi-
cal suture. Suture weakly impressed, linear. Sculpture
of ne, close-set spiral cords, about one quarter width
of their interspaces, irregularly interrupted by strongly
prosocline growth lines. Last whorl angled at base. Base
not sharply delimited, bearing narrow, shallow umbili-
cus and about 14 concentric cords. Aperture subtrigonal,
outer lip simple, bevelled edge, roundly angled at periph-
ery. Peristome incomplete. Columella bearing weak fold
placed just below mid-height. Parietal callus poorly de-
veloped, restricted to a thin callus wash. Colour pattern
of vertical ammules is preserved.
Discussion – At rst glance Gibbula brebioni nov. sp.
can be confused with Gibbula striatellata (Millet, 1865);
both are about the same size and both have a regularly
Cainozoic Research, 17(2), pp. 75-166, December 2017 105
conical shell shape, however, G. brebioni has ner spiral
sculpture and lacks the swollen suprasutural cord present
in G. striatellata, moreover, the suture is deeper in G.
striatellata. The character of the base is similar in both
species; a shallow, narrow umbilicus and dense spiral
cords.
In the present-day faunas the Mediterranean G. spratti
(Philippi, 1844) also has dense spiral sculpture and a nar-
row, shallow umbilicus, but differs in having a less regu-
larly conical prole, the whorls are more convex, espe-
cially the last whorl, which is regularly rounded and not
angled at the base. Gibbula adriatica (Philippi, 1844) has
a similar shell prole, but coarser spiral sculpture. In the
fossil assemblages none of the Gibbula species gured
by Cossmann & Peyrot (1917) from the lower and middle
Miocene Aquitaine Basin of France have such ne spiral
sculpture.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Gibbula clanculiforma nov. sp.
Plate 30, g. 1
Type material – Holotype NHMW 2016/0103/0249,
height 5.5 mm, width 5.8 mm.
Other material – Known only from holotype.
Etymology – Named reecting the supercial resem-
blance to shells of the genus Clanculus. Gibbula gender
feminine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Gibbula species of small size, with low coni-
cal spire, sculpture composed of numerous beaded spiral
cords on both dorsum and base, adapical cord slightly
more strongly developed, last whorl with rounded periph-
ery in which base is not strongly delimited from dorsum,
narrow, but deep umbilicus, and single weak columellar
tubercle.
Description – Shell small, solid, trochiform, with low
conical spire. Protoconch not preserved. Teleoconch of
four weakly convex whorls, with periphery at abapi-
cal suture. Suture impressed, shallowly canaliculate.
Sculpture on rst teleoconch whorl of three cords, with
a fourth appearing on second half of rst whorl below
adapical suture, crossed by close-set, prosocline axial
ribs forming ne reticulated pattern, with small tuber-
cles developed at intersections. Abapically, axial sculp-
ture weakens, visible only in interspaces between cords,
cords increase in number, irregular in strength and be-
come strongly beaded, the adapical cord more strongly
developed. Last whorl depressed, slightly concave be-
low suture, strongly rounded at periphery, bearing eight
beaded cords above periphery. Base poorly delimited,
Plate 29. Gibbula brebioni nov. sp.; 1. Holotype MNHN.F.A57689, height 8.7 mm, width 7.5 mm; 2. Paratype 1 NHMW
2016/0103/0108, height 9.7 mm, width 8.0 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France,
Tortonian, upper Miocene.
106 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
bearing 13 beaded cords, narrowly but deeply imbilicate.
Peristome incomplete. Outer lip simple, slightly concave
adapically, rounded and somewhat ared abapically,
smooth within. Columella excavated, bearing weak tu-
bercle placed just below mid-height. Columella callus
erect, forming medial border of umbilicus. Parietal cal-
lus poorly developed.
Discussion – Although represented by a single specimen,
Gibbula clanculiforma nov. sp. is so unlike any of its
congeners that is merits description. At rst glance the
sculpture is similar to that of shells of the genus Clancu-
lus de Montfort, 1810, especially the strongly beaded spe-
cies like Clanculus (Clanculopsis) baccatus (Defrance,
1824) (see above), but Clanculus species have a very
characteristic columellar and umbilical structure (see Pl.
15, gs 1-4), absent in G. clanculiforma. Based on shell
shape and sculpture, placement in the Calliostomatidae
Thiele, 1924 could also be a possibility, but these species
have a smooth columella. The small very solid nature
of the shell and the small but well developed columel-
lar tubercle lead us to include this small shell within the
Cantharidinae Gray, 1857, genus Gibbula Risso, 1826.
There are no Neo gene to present-day eastern Atlantic or
Mediterranean species with which this species can be
compared. Some of the Gibbula species from the Plioce-
ne North Sea Basin such as G. octosulcata (Nyst, 1835)
and G. beetsi van Regteren Altena, 1954 are similar in
shape and have beaded spiral cords, but are clearly not
conspecic.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Gibbula conicomagus nov. sp.
Plate 31, gs 1-2
Type material – Holotype NHMW 2016/0103/1654,
height 14.1 mm, width 11.7 mm; paratype 1 NHMW
2016/0103/1655, height 14.8 mm, width 11.4 mm (incom-
plete).
Other material – Maximum height 13.7 mm, width 13.3
mm (incomplete). Renauleau: FVD (2).
Etymology – Name reecting the sculpture, similar to
that of the G. magus species group and the conical shell
shape. Gibbula gender feminine.
Locus typicus – Renauleau, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Gibbula species of medium size, with regu-
larly conical shell shape, rugose sculpture of low axial
folds crossed by six spiral cords, abapical cord reinforced
forming periphery, ne scabrous axial sculpture in inter-
spaces between cords, last whorl angled at peribasal
cord, base attened bearing about six cords and narrow
umbilicus.
Description – Shell medium-sized, solid, trochiform,
with regularly conical spire. Protoconch not preserved.
Teleoconch of 5.5 almost at-sided whorls, with periph-
ery at abapical suture. Suture deeply impressed, linear.
Sculpture of weakly developed, low, opisthocline axial
rugae and six spiral cords; abapical cord strengthened
forming periphery. Numerous ne axial lamellae visible
in interspaces between cords, so that sculpture grossly
rugose, nely scabrous. Last whorl angled at base. Base
attened, sharply delimited by rounded peribasal cord,
bearing narrow, shallow umbilicus and about six concen-
tric cords. Aperture subtrigonal, outer lip simple, bevel-
led edge, angled at periphery. Peristome incomplete. Col-
umella bearing weak fold placed just below mid-height.
Parietal callus not developed.
Discussion – The sculpture of Gibbula conicomagus
nov. sp. composed of opisthocline axial rugae, crossed
by spiral cords with ne axial lamellae present in the
interspaces between the cords suggests that it belongs to
the G. magus species group, although it differs from G.
magus (Linnaeus, 1758), G. sagus (Defrance, 1828) and
G. fanulum (Gmelin, 1791) in having a regularly conical
spire with almost at sided whorls. Moreover, unlike its
congeners mentioned above, the axial rugae are hardly
raised. In these characters it is similar to the species
from the middle Miocene Karaman Basin of Turkey re-
corded as G. cf. sagus by Landau et al. (2013), which also
has a regular conical spire and weak axial rugae, but the
Plate 30. Gibbula clanculiforma nov. sp.; 1. Holotype NHMW 2016/0103/0249, height 5.5 mm, width 5.8 mm. Le Grand Chau-
vereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 107
Turkish species has a broader apical angle and ner and
more irregular spiral sculpture.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Gibbula fanulum (Gmelin, 1791)
Plate 32, g. 1
*1791 Trochus fanulum Gmelin, p. 3573.
1964 Gibbula (Forskalena) fanulum Gmelin in Linné,
1790 [sic] – Brébion, p. 113, pl. 3, g. 7.
2003 Gibbula (Forskalena) fanulum (Gmelin, 1791) –
Landau et al., p. 49, pl. 11, g. 1 (cum syn.).
2004 Gibbula (Forskalena) fanulum (Gmelin in L.,
1790) [sic] – Chirli, p. 70, pl. 28, gs 11, 12, pl.
29, gs 1-5.
Material and dimensions – Maximum height 10.6 mm,
width 11.2 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0099 (1), NHMW 2016/0103/0100 (50+), RGM.
1309506 (50+), RGM.1309507 (30), RGM.1309522 (50+
juveniles), RGM.1309559 (2), LC (50+), FVD (50+).
Sceaux-d’Anjou: NHMW 2016/0103/0103 (13), LC (24),
FVD (19). Renauleau: NHMW 2016/0103/1414 (32), LC
(22), FVD (25).
Discussion – Gibbula fanulum (Gmelin, 1791) is a very
characteristic species, with its strongly nodular sculpture
at the shoulder and deep groove between the shoulder and
strong, rounded basal cord. This characteristic sculpture
makes it difcult to confuse G. fanulum with any of its
congeners. Pracchia (1998) considered the presence or
absence of the strong axial nodules on the periphery in
Italian Pliocene populations of G. fanulum an ecophe-
notypic character, the nodular form inhabiting rocky
environment and the smoother form living on sandy sub-
strate, without intermediate forms. The specimens from
the Assemblage I locality of St-Clément-de-la Place are
Plate 31. Gibbula conicomagus nov. sp.; 1. Holotype NHMW 2016/0103/1654, height 14.1 mm, width 11.7 mm; 2. Paratype 1
NHMW 2016/0103/1655, height 14.8 mm, width 11.4 mm. Renauleau, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Plate 32. Gibbula fanulum (Gmelin, 1791); 1. NHMW 2016/0103/0099, height 9.4 mm, width 9.5 mm. Le Grand Chauvereau, St-
Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
108 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
strongly nodular, suggesting a rocky environment near-
by. As repeatedly encountered in this monograph, the As-
semblage I specimens are considerably smaller than the
Mediterranean Pliocene and present-day shells, which at-
tain up to 21 and 19 mm in height respectively (Landau et
al., 2003; Poppe & Goto, 1991).
Brébion (1964, p. 113) recorded this species from only
Assemblage I localities (Sceaux-d’Anjou, Renauleau,
Thorigné, St-Clément-de-la-Place, St-Michel). These are
the stratigraphically oldest records for the species.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Brébion, 1964). Lower Pliocene: central Mediter-
ranean, Italy (Chirli, 2004), Tunisia (Fekih, 1975). Up-
per Pliocene: western Mediterranean, Estepona Basin,
Spain (Landau et al., 2003); central Mediterranean, Italy
(Sacco, 1896a; Malatesta, 1974; Spadini, 1986; Pracchia,
1988). Pleistocene: western Mediterranean, Balearic Is-
lands, (Cuerda Barceló, 1987); central Mediterranean,
Italy (Cerulli-Irelli, 1916; Malatesta, 1960; Brambilla et
al., 1988; Dell’Angelo & Forli, 1995). Present-day: south-
ern Portugal and Mediterranean, shallow waters, where it
prefers Posidonia elds (Poppe & Goto, 1991).
Gibbula marianae nov. sp.
Plate 33, g. 1
Type material – Holotype NHMW 2016/0103/0677, height
6.1 mm, width 6.4 mm; paratype 1 NHMW 2016/ 0103/ 0677,
height 6.0 mm, width 5.7 mm; paratype 2 MNHN. F.
A57941, height 6.2 mm, width 5.8 mm, Sceaux-d’Anjou.
Paratype 3 RGM.1309601, height 4.2 mm, width 4.5 mm,
St. Clément-de-la-Place.
Other material – Sceaux-d’Anjou: RGM.1348010 (4),
FVD (2).
Etymology – Named after Mariana Mihai, lawyer and
wife of the second author, in recognition of her support
and patience. Gibbula gender feminine.
Locus typicus – La Presselière, Sceaux-d’Anjou, Maine-
et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Gibbula species of small size, with low coni-
cal shell shape, strong spiral sculpture composed of el-
evated cords, strongly beaded by close-set axial lamellae;
on last whorl two stronger cords above periphery, with
two weaker cords between, periphery marked by thick-
ened peribasal cord, attened imperforate base bearing
six cords, and thickened columella bearing poorly delim-
ited tooth.
Description – Shell small, low trochiform. Protoconch
paucispiral, surface smooth. Teleoconch of four de-
pressed whorls separated by shallow suture. Sculpture on
rst teleoconch whorl composed of three narrow, elevat-
ed cords. Abapically, ad- and abapical cords strengthen;
abapical cord strongly dominant. On penultimate whorl
fourth cord appears above suture. Close-set, strongly
prosocline axial lamellae cross cords forming beads,
most strongly developed on ad- and abapical cords. Last
whorl acutely angled at base, bearing strongly beaded
cords 1 and 3, cord 2 weaker, with a further weak cord
intercalated between cords 1 and 2. Peribasal cord de-
limiting base strongly developed, weakly beaded. Base
attened, bearing six narrow equal cords; umbilicus ab-
sent. Aperture subquadrate. Outer lip with bevelled edge,
angled at periphery. Columella thickened, sloping abaxi-
ally, bearing poorly-delimited tooth abapically. Parietal
callus not developed.
Discussion – Gibbula marianae nov sp. is easily sepa-
rated from its congeners by its strongly beaded spiral
sculpture. This sculpture is reminiscent of the Collicu-
lus species discussed previously [i.e. C. insignis (Millet,
1854)], but the shell lacks the deep umbilicus that we as-
sociate with that genus. We cannot nd any European
fossil or extant species with which to usefully compare
this species, which has so far only been found at Sceaux-
d’Anjou and St. Clément-de-la-Place.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Plate 33. Gibbula marianae nov sp.; 1. Holotype NHMW 2016/0103/0677, height 6.1 mm, width 6.4 mm. La Presselière, Sceaux-
d’Anjou, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 109
Gibbula provosti Ceulemans, Van Dingenen & Landau,
2016
Plate 34, gs 1-2
1964 Gibbula (Colliculus) varia Linné, 1766 [sic] var.
termieri nov. sp. Dollfus mss., (emend.) – Bré-
bion, p. 109, pl. 3, g. 3 (nomen nudum).
*2016a Gibbula provosti Ceulemans, Van Dingenen &
Landau, p. 62, pl. 4, g. 6; pl. 5, g. 1.
Material and dimensions – Maximum height 7.5 mm,
width 8.6 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0112-0113 (2), NHMW 2016/0103/0114 (9 + 30 juve-
niles), RGM.1309756 (45 juveniles), RGM.1309771 (50+
juveniles), LC (3), FVD (20). Sceaux-d’Anjou: NHMW
2016/0103/0218 (21), RGM.1309691 (2), RGM.1309703
(1), RGM.1309731 (2), RGM.1309762 (13), RGM.1309765
(10), RGM.1347917 (1), LC (10), FVD (12). Renauleau:
LC (1).
Discussion – The species described and illustrated by
Brébion (1964, p. 109, pl. 3, g. 3) as Gibbula (Colliculus)
varia var. termieri Dollfus mss. (emend) (nomen nudum)
was formally described as Gibbula provosti Ceulemans,
Van Dingenen & Landau, 2016, based on material from
the lower Pliocene Assemblage III locality of Le Pigeon
Blanc. However, the species is commoner and better pre-
served at the upper Miocene Assemblage I locality of St-
Clément-de-la-Place. Gibbula provosti is characterised
by its strongly depressed spire, its surface covered by
ne, narrow spiral cords, its strongly oblique columella
bearing a relatively welldeveloped tooth, its base bear-
ing prominent cords and growth lines, and its narrow
smooth umbilicus. The plentiful material at St-Clément-
de-la-Place allows us to make several further observa-
tions not included in the original description. There is an
important change in shape with ontogeny. Juvenile shells
are even more strongly depressed and lack the concave
adapical half to the last whorl seen in adult specimens
(Pl. 34, g. 2). This can be seen just developing at the
aperture in the juvenile shell illustrated (Pl. 34, g. 1).
Secondly, a colour pattern is preserved, especially in the
juvenile shells consisting of spiral rows of dots or circles
similar to that seen in several present-day European Gib-
bula species [i.e. G. umbilicaris (Linnaeus, 1758)]. This
colour pattern was not commented on in the original de-
scription, but can be seen in paratype 3 from Le Pigeon
Blanc (NHMW 2015/0133/0068; Ceulemans et al., 2016a,
pl. 5, g. 1).
Gibbula provosti is similar to G. varia (Linnaeus, 1758)
from the Pliocene to present-day Mediterranean, but
differs in 1) being smaller in size, 2) having a more de-
pressed shell, especially the last whorl, 3) having nely
beaded spiral cords, 4) having more prominent sculpture
on the base, 5) having a more oblique columella, 6) hav-
ing a more strongly developed columellar tooth, and 7)
having a narrower umbilicus.
A similar species was gured by Brébion (1964, pl. 3, gs
1, 2) under the name G. varia var. monodontoides (Mil-
let, 1854) (nomen nudum; made available by Millet, 1865)
from localities in Assemblages I and II. This species has
a less depressed shell than Gibbula provosti and a bian-
gular whorl prole, the last whorl angled at the shoulder
and base. It is similar to G. varia, but was said to differ
in the complete absence, or almost so, of the umbilicus,
which is covered by a columellar callus. Unfortunately,
no basal view was offered by Brébion and we have found
no further specimens of this species.
Plate 34. Gibbula provosti Ceulemans, Van Dingenen & Landau, 2016; 1. NHMW 2016/0103/0112, height 4.6 mm, width 6.7 mm;
2. NHMW 2016/0103/0113, height 6.7 mm, width 8.2 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW
France, Tortonian, upper Miocene.
110 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Brébion (1964, p. 110) recorded G. provosti from the As-
semblage III localities of Le Pigeon Blanc and Le Giron-
dor, to which we add the Assemblage I locality of St-Clé-
ment-de-la-Place.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Brébion, 1964). Lower Pliocene: Atlantic, NW
France (Brébion, 1964; Ceulemans et al., 2016a).
Gibbula sagus (Defrance, 1828)
Plate 35, gs 1-3
*1828 Trochus sagus Defrance, p. 478.
1854 Trochus Sagus Millet, p. 156.
1903 Gibbula saga Defrance – Dollfus et al., p. 7, pl.
32, gs 3, 4.
1917 Gibbula pseudomagus d’Orbigny – Cossmann &
Peyrot, p. 110, pl. 3, gs 63-65.
1938 Gibbula saga Defrance – Peyrot, p. 26, pl. 1, gs
22-23.
1949 Gibbula sagus Defrance, 1828 – Glibert, p. 50, pl.
3, g. 4.
1964 Gibbula sagus Defrance, 1826 [sic] – Brébion, p.
102.
Material and dimensions – Maximum height 12.0 mm,
width 12.1 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0102 (1), NHMW 2016/0103/1653 (1), NHMW
2016/0103/0100 (6), RGM.1309517 (31 juveniles), LC
(50+), FVD (50+). Sceaux-d’Anjou: NHMW 2016/
0103/0104 (8), RGM.1309512 (50+ juveniles), RGM.
1309513 (13 juveniles), RGM.1309514 (8 juveniles),
RGM.1309519 (3), RGM.1309520 (5), LC (50+), FVD
(50+). Renauleau: NHMW 2016/0103/1652 (1), NHMW
2016/0103/0115 (50+), LC (40), FVD (23). Beugnon:
NHMW 2016/0103/0667 (4), RGM.1309508 (50+),
RGM.1309538 (5), RGM.1309741 (50+, mainly juveniles),
LC (6).
Discussion – Gibbula sagus (Defrance, 1828) from the
northwestern French Atlantic middle Miocene is simi-
lar to G. magus (Linnaeus, 1758), but differs in having
a less depressed shell, with a less angular last whorl pe-
riphery and a less depressed base. The basal cords are
more strongly developed and elevated in comparison to
G. magus, in which the basal sculpture is reduced to low
concentric ridges. These differences are evident when
middle Miocene populations from the Loire Basin are
compared with present day G. magus. The upper Mio-
cene specimen at hand from the Asemblage I localities
Plate 35. Gibbula sagus (Defrance, 1828); 1. NHMW 2016/0103/0101, height 10.7 mm, width 10.2 mm. Le Grand Chauvereau, St-
Clément-de-la-Place; 2. NHMW 2016/0103/1652, height 9.9 mm, width 8.9 mm. Renauleau. 3. NHMW 2016/0103/1653, height
9.9 mm, width 11.0 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 111
are here assigned to G. sagus; although smaller than
specimens from the Loire Basin, the spire is relatively
tall, the base not strongly depressed and the basal cords
are strongly developed, as typical for G. sagus (Pl. 35,
gs 1-2). Occasional specimens are more depressed with
a more strongly developed peripheral cord (Pl. 35, g. 3),
as seen in G. magus. However, we consider these to be G.
sagus, as the axial folds on the last whorl are developed
until the peripheral cord, whereas in G. magus the folds
tend to be restricted to the shoulder, and the basal sculp-
ture is strong. Nevertheless, the two species are undoubt-
edly closely similar and may represent an evolutionary
series.
This species was recorded from the lower and middle
Miocene Aquitaine Basin of France as G. pseudomagus
(d’Orbigny, 1852), which Cossmann & Peyrot (1917, p.
110) said differed from G. magus in being higher spired,
the last whorl less angular at the periphery and the um-
bilicus narrower, less clearly delimited than in the extant
species. These are the same differences as those between
G. sagus and G. magus and we agree with Glibert (1949,
p. 51) in considering G. pseudomagus (d’Orbigny, 1852)
and junior subjective synonym of G. sagus.
Brébion (1964, p. 103) recorded this species from Assem-
blage I localities (Sceaux-d’Anjou, Renauleau, Thorigné,
St-Clément-de-la-Place, Beaulieu). He also noted that
Dollfus (1905) had listed it from the Assemblage IV lo-
cality of Gourbesville, however, we provisionally exclude
this record from the distribution until it is veried.
Distribution – Lower Miocene: Atlantic, Aquitaine Ba-
sin (Cossmann & Peyrot, 1917). Middle Miocene: Atlan-
tic, Aquitaine Basin (Cossmann & Peyrot, 1917), Loire
Basin, France (Peyrot, 1938; Glibert, 1949). Upper Mi-
ocene: Atlantic (Tortonian), NW France (Millet, 1854;
Brébion, 1964), Cacela Basin, Portugal (Dollfus et al.,
1903).
Gibbula striatellata (Millet, 1865)
Plate 36, gs 1-2
1854 Trochus Striatellatus Millet, p. 156 (nomen nu-
dum).
*1865 Trochus striatellatus Millet, p. 582.
1964 Gibbula striatellata Millet, 1854 [sic] – Brébion,
p. 104, pl. 2, gs 32, 33.
Type material – Syntypes: Sceaux-d’Anjou, Musée
d’Angers, France ( de Brébion, 1964, p. 104).
Material and dimensions – Maximum height 12.5 mm,
width 11.0 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0105 (1), NHMW 2016/0103/0106 (29 + 20 juveni-
les), RGM.1309597 (8), RGM.1309752 (13), RGM. 1309755
(23), LC (50+), FVD (50+). Sceaux-d’Anjou: NHMW
2016/0103/0107 (22), RGM.1309632 (4), RGM.1309656 (4),
RGM.1309727 (3), RGM.1309763 (5), RGM.1309766 (7),
LC (8), FVD (10). Renauleau: NHMW 2016/0103/1501
(1), NHMW 2016/0103/1432 (4), LC (8), FVD (7).
Discussion – The original description reects well the
character of this species: ‘Trochus striatellatus, Millet.
Coq. en cône court, composée de 5 tours de spire légère-
ment striés, séparés par une suture légèrement renée en
cordon net sur ses bords. Dessous ombiliqué et couvert
de nes stries. Ouverture angulaire, garnie d’un léger
bourrelet intérieur. Hauteur et diamètre: 7 millimètres.
Plate 36. Gibbula striatellata (Millet, 1865); 1. NHMW 2016/0103/0105, height 10.8 mm, width 9.9 mm. Le Grand Chauvereau,
St-Clément-de-la-Place. 2. NHMW 2016/0103/1501, height 11.2 mm, width 13.2 mm. Renauleau, Maine-et-Loire, NW France,
Tortonian, upper Miocene.
112 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of nor thwestern France, 1. Patellogastropoda and Vetigastropoda
Sceaux (Millet, 1865, p. 582)’. The height of the shell
is somewhat variable, as seen in the series illustrated,
which reects the two extremes (Pl. 36, gs 1-2). On
the spire whorls lie about six attened, subequal cords
above the broader, elevated suprasutural cord, which is
also covered with secondary cords. The suture is deeply
impressed. The suprasutural cord continues along the
periphery of the last whorl, delimiting the base. The
basal angulation is variable, roundly angled in some
specimens (Pl. 36, g. 1), sharply in others (Pl. 36, g.
2). The base bears a narrow, shallow umbilicus and is
covered in irregular cords. The columella bears a weak
fold just below mid-columella. This species is most like
the Pliocene to present-day eastern Atlantic and Medi-
terranean G. cineraria (Linnaeus, 1758), but that species
differs in not having a swollen suprasutural cord and in
having a deeper umbilical perforation.
The spiral sculpture is not unlike that of G. provosti Ceu-
lemans, Van Dingenen & Landau, 2016 from the lower
Pliocene Assemblage III locality of Le Pigeon Blanc, but
that species is more depressed and lacks the swollen su-
prasutural cord.
Millet (1854, 1865) recorded this species from the As-
semblage I localities of Sceaux-d’Anjou and Thorigné, to
which Brébion (1964, p. 105) added Renauleau and St-
Michel and we add St-Clément-de-la-Place. We note that
the specimens from Renauleau are somewhat atter with
the peripheral cord more strongly swollen than seen in
the St-Clément-de-la-Place specimens, however, we con-
sider them conspecic.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Genus Paroxystele Schultz, 1969
Type species (by original designation) – Trochus patulus
Brocchi, 1814, Neogene, Italy.
1969 Paroxystele Schultz, p 217.
Paroxystele trochiformis (Millet, 1865)
Plate 37, g. 1
1854 Pitonellus trochiformis Millet, p. 157 (nomen nu-
dum).
*1865 Pitonellus trochiformis Millet, p. 584.
1949 Monodonta (Oxystele) amedei turoniensis Glibert,
p. 63, pl. 3, g. 10.
2016a Paroxystele turoniensis (Glibert, 1949) – Ceule-
mans et al., p. 66, pl. 6, g. 2.
Material and dimensions – Maximum height 11.1 mm,
width 17.6 mm. Renauleau: NHMW 2016/0103/1425
(1), LC (4), FVD (3). Beugnon: RGM.1309515 (7 juve-
niles), RGM.1309525 (1), RGM.1309526 (1 juvenile),
RGM.1309540 (2 juveniles).
Discussion – The specimens from Renauleau are typical
for Paroxystele turoniensis (Glibert, 1949), which is char-
acterised by its relatively low-spire, subcarinate base, and
having the whorls ornamented by six to eight spiral cords,
without secondary sculpture, subsutural folds are absent,
and the umbilical callus is broad and closely adherent,
lling the umbilicus. Ceulemans et al. (2016a) recorded
this species from the younger lower Pliocene Assemblage
III locality of Le Pigeon Blanc and noted that this was the
stratigraphically youngest record for the species and for
the typical ‘Miocene’ Paroxystele species-group, which
differ from those in the Pliocene in having a completely
closed umbilicus and only few Miocene species have
subsutural folds, which are usually present in the Plio-
Pleistocene Paroxystele patulum (Brocchi, 1814).
However, in Ceulemans et al. (2016a) we failed to no-
tice that Glibert’s name is a junior synonym of Pitonellus
trochiformis Millet, 1865. Millet’s original description
‘Pitonellus trochiformis, Millet. Coq. comme discoîdale
ou en cône racourci, composée de 6-7 tours de spire ar-
rondis en dessus, bien séparés les uns des autres, garnis
chacun de 6 lets arrondis, dont le plus gros, au sommet,
touche la suture. Le dessous, légèrement concave et lé-
gèrement strié, porte un ombilic complètement recouvert
par une callosité lisse, qui ne dépasse pas la hauteur de
ses bords. Diamètre: 28 millimètres; longueur: 20-22
millimètres (1865, p. 584)’ leaves little doubt that he was
referring to this species. Although Glibert’s name is bet-
ter known, we cannot satisfy the requirements of Article
23.9.1.2 (ICZN 1999a) and therefore Paroxystele trochi-
formis (Millet, 1865) must take priority.
Plate 37. Paroxystele trochiformis (Millet, 1865); 1. NHMW 2016/0103/1425, height 11.1 mm, width 17.6 mm. Renauleau, Maine-et-
Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 113
The stratigraphically older Aquitanian and Burdigalian
Atlantic species P. burdigalensis (Cossmann & Peyrot,
1917) differs in being lower spired and having secondary
spiral sculpture between the primary spiral cords. Par-
oxystele orientale (Cossmann & Peyrot, 1917) from the
middle Miocene eastern Mediterranean and Paratethys
differs in having more numerous spiral cords, the base
is more rounded and the umbilical callus is narrower, not
completely lling the umbilicus.
Millet (1865) recorded this species from the Assemblage
I localities of Renauleau, Doué, Sceaux-d’Anjou, Tho-
rigné, to which Brébion (1964, p. 117) added St-Michel,
Les Pierres Blanches and Beaulieu.
Distribution – Middle Miocene: Atlantic, Loire Basin,
France (Glibert, 1949). Upper Miocene: Atlantic (Torton-
ian), NW France (this paper). Lower Pliocene: Atlantic,
NW France (Ceulemans et al., 2016a).
Genus Phorcus Risso, 1826
Type species (by subsequent designation, Bucquoy et al.,
1885) – Phorcus margaritaceus Risso, 1826, present-day,
France (Mediterranean).
1826 Phorcus Risso, p. 133.
1847 Osilinus Philippi, p. 20. Type species (by subse-
quent designation, Opinion 1930, 1999b): Trochus
turbinatus Born, 1778, present-day, Mediterrane-
an.
1852 Neptheusa Leach in Gray, p. 146, 174. Type spe-
cies (by mo notypy): Trochus crassus Pulteney,
1799, present- day, British Isles.
1852 Trochocochlea Mörch, p. 142. Type species (by
subsequent designation, Bucquoy et al., 1885):
Trochus turbinatus Born, 1778, present-day, Med-
iterranean.
1884b Caragolus Monterosato, p. 43. Type species (by
subsequent designation, Crosse, 1885): Trochus
turbinatus Born, 1778, present-day, Mediterra-
nean.
Phorcus gallicophorcus nov. sp.
Plate 38, gs 1-2
Type material – Holotype MNHN.F.A57690, height
5.5 mm, width 5.3 mm; paratype 1 MNHN.F.A57691,
height 5.1 mm, width 4.0 mm; paratype 2 NHMW
2016/0103/0119, height 4.7 mm, width 4.1 mm; paratype 3
NHMW 2016/0103/0120, height 4.9 mm, width 4.2 mm;
paratype 4 RGM.1309747, height 3.2 mm, width 3.2 mm;
paratype 5 RGM.1309748, height 3.7 mm, width 3.1 mm.
Other material – St-Clément-de-la-Place: Maximum
height 5.1 mm, width 4.6 mm. NHMW 2016/0103/0121
(34), LC (20), FVD (25). Sceaux-d’Anjou: NHMW
2016/0103/0217 (8).
Etymology – Compound name reecting the Latin name
for the Province of Gaul, ‘Gallia’, which now includes
France, and the genus Phorcus. Phorcus gender mascu-
line.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Phorcus species of small size, with solid
shell, relatively tall spire, convex spire whorls, sculptured
Plate 38. Phorcus gallicophorcus nov. sp.; 1. Holotype MNHN.F.A57690, height 5.5 mm, width 5.3 mm; 2. Paratype 1
MNHN.F.A57691, height 5.1 mm, width 4.0 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France,
Tortonian, upper Miocene.
114 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
by rounded cords, beaded in adapical portion, imperfo-
rate base covered by rounded cords, and subobsolete
columellar thickening.
Description – Shell small, solid, trochiform. Protoconch
paucispiral, composed of 1.5 smooth whorls. Teleoconch
of ve relatively tall convex whorls, periphery at abapical
suture. Suture impressed, linear. Sculpture of close-set
spiral cords, slightly wider than their interspaces, beaded
by strongly prosocline growth lines to a variable degree
in adapical portion, especially last whorl. Last whorl in-
ated, evenly rounded to roundly angled at base. Base
not sharply delimited, imperforate, bearing about eight
rounded concentric cords. Aperture subtrigonal, outer lip
simple, bevelled edge, rounded and slightly ared abap-
ically. Peristome incomplete. Columella sloping strongly
abaxially, bearing subobsolete thickening just below
mid-height. Columella callus narrow, sharply delimited;
parietal callus poorly developed.
Discussion – Although much smaller shelled than its
present-day European congeners, the very thick shell,
rounded whorls and apertural characters, with a strongly
thickened outer lip, shallowly abaxially sloping columel-
la with a subobsolete thickening lead us to place this spe-
cies in the genus Phorcus Risso, 1826 rather than Gib-
bula. These shells are not juveniles, as the outer lip is
strongly thickened and numerous specimens available to
us are all similar.
It resembles a minute P. turbinatus (Born, 1778), but is
slightly taller spired and has narrower spiral cords.
Phorcus gallicophorcus nov. sp. is supercially similar
to a group of Pliocene North Sea Basin Gibbula species,
characterised by G. obconica (S.V. Wood, 1848), which
also has a small, solid shell (Marquet, 1998, g. 20), but
that species has narrower spiral cords and ner and more
conspicuous beading and more numerous cords on the
base. Gibbula beetsi van Regteren Altena, 1954 from
the Pliocene of Belgium and The Netherlands differs
similarly in having ner and stronger beading, but also
differs in having a more depressed, broader shell, with
a canaliculated suture and a small umbilicus (Pouwer
& Wesselingh, 2012, g 12), almost completely closed
in some specimens (Marquet, 1998, g. 18b). Gibbula
nehalenniae van Regteren Altena, 1954, also from the
Pliocene of Belgium and The Netherlands, was consid-
ered a forma or subspecies of G. obconica by Marquet
(1995, 1998, respectively), but we agree with Pouwer &
Wesselingh (2012, p. 156) in considering that species
closer to G. beetsi. Most importantly, all these North Sea
Basin Gibbula species differ in having an almost verti-
cal columella, whereas in Phorcus is it strongly abaxially
sloping.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Superfamily Turbinoidea Ranesque, 1815
Family Turbinidae Ranesque, 1815
Genus Bolma Risso, 1826
Type species (by monotypy) – Turbo rugosus Linné,
1767, present-day, Mediterranean.
1767 Bolma Risso, 1826, p. 117.
Note – We have not repeated here the list of generic syno-
nyms given by Beu & Ponder (1979), as we do not believe
the genus, as envisaged by these authors, to be mono-
phyletic. This is supported by the molecular phylogenetic
studies presented by Williams & Ozawa (2006). Landau
et al. (2003) argued that Ormastralium Sacco, 1896 spe-
cies should be considered a separate subgenus, elevated
to genus by Landau et al. (2013).
Bolma granosa (Borson, 1821)
Plate 39, gs 1-2
*1821 Trochus granosus Borson, p. 67, pl. 2, g. 6.
1827 Turbo calcar Defrance, p. 520 (non Linnaeus,
1758).
1854 Turbo Calcar Defr. – Millet, p. 158.
Plate 39. Bolma granosa (Borson, 1821); 1. NHMW 2016/0103/0230, height 11.5 mm, width 12.6 mm; 2. NHMW 2016/0103/0231,
height 11.3 mm, width 11.9 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper
Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 115
1938 Bolma miocalcar Peyrot, p. 36, pl. 1, gs 29-30.
1964 Astraea (Bolma) granosa Borson, 1821 – Brébion,
p. 135.
2003 Bolma (Bolma) granosa (Borson, 1821) – Landau
et al., p. 30, pl. 7, g. 2 (cum syn.).
Material and dimensions – Maximum height about 33.0
mm (incomplete). St-Clément-de-la-Place: NHMW
2016/0103/0230-0231 (2), NHMW 2016/0103/0232
(4 + 50 juveniles), RGM.1309595 (20 adults and sub-
adults), RGM.1348013 (9), LC (20), FVD (27). Sceaux-
d’Anjou: NHMW 2016/0103/0233 (12 + 15 juveniles),
RGM.1309576 (12 subadult + juveniles and oper-
cula), RGM.1309578 (50+ subadults and juveniles),
RGM.1309577 (50+ adults and subadults), RGM.1309579
(5 juveniles), RGM.1309587 (10+ subadults and ju-
veniles), LC (50+), FVD (50+). Renauleau: NHMW
2016/0103/1433 (4 incomplete), LC (40), FVD (23).
Discussion – Bolma granosa (Borson, 1821) is character-
ised by its nely beaded spiral sculpture on the subsutural
ramp and three strong cords mid-whorl on the last whorl,
the adapical cord forming short open spines. Bolma gra-
nosa is a small shelled species for the genus, and most of
the specimens from the Assemblage I localities are even
smaller than those from the middle Miocene Loire Basin
of France reported by Glibert (1949, p. 76) (maximum
most specimens Assemblage I height 11.5 mm vs. Loire
Basin 18.0 mm). The exception is one fragmentary speci-
men from Renauleau that is considerably larger than the
rest and suggests a maximum height of about 33 mm.
Landau et al. (2003) reported the last occurrence for the
species from the lower upper Pliocene of the Estepona
Basin, southern Spain, although these Pliocene speci-
mens are not identical to those from Assemblage I; they
are larger shelled (maximum 23.9 mm) and have a few
weak beaded spiral cords on the base as opposed to close-
set wavy axial ribs in the Assemblage I specimens. How-
ever, one of Glibert’s (1949, pl. 4, g. 7b) specimens from
the Loire Basin also has beaded cords on the base.
Brébion (1964, p. 136) recorded B. granosa widely from
the Assemblage I localities (Sceaux-d’Anjou, Thorigné,
St-Clément-de-la-Place, Les Pierres Blanches, Beaulieu,
Chalonnes, Contigné, St-Jacques).
Distribution – Middle Miocene: Atlantic, Loire Basin
(Peyrot, 1938; Glibert, 1949); Proto-Mediterranean, Italy
(Sacco, 1896a; Pavia, 1976). Upper Miocene: Atlantic
(Tortonian), NW France (Millet, 1854, 1865; Brébion,
1964); Proto-Mediterranean, Italy (Sacco, 1896a). Up-
per Pliocene: western Mediterranean, Estepona Basin, S.
Spain (Landau et al., 2003).
Bolma cf. meynardi (Michelotti, 1847)
Plate 40, gs 1-3
cf. *1847 Turbo Meynardi Michelotti, p. 177, pl. 7, g. 4.
1854 Turbo Trochleatus Millet, p. 158 (non zu Münster,
1841).
cf. 2013 Bolma meynardi (Michelotti, 1847) – Landau et
al., p. 30, text-g. 15/1, pl. 1, gs 9-12 (cum syn.).
1964 Astraea (Bolma) trochleata Millet, 1854 – Brébi-
on (partim), p. 131, pl. 3, gs 18, 19 (Assemblage
1 records only).
Material and dimensions – Maximum height 29.7 mm
(incomplete), width 37.6 mm. St-Clément-de-la-Place:
RGM.1309690 (1). Sceaux-d’Anjou: FVD (9). Re-
nauleau: NHMW 2016/0103/1434-1436 (3), NHMW
2016/0103/1437 (36 adult fragments and juveniles), LC
(50+), FVD (50+).
Plate 40. Bolma cf. meynardi (Michelotti, 1847); 1. NHMW 2016/0103/1434, height 29.7 mm, width 37.6 mm; 2. NHMW
2016/0103/1436, height 14.3 mm, width 20.3 mm; 3. NHMW 2016/0103/1439, height 31.8 mm. Renauleau, Maine-et-Loire, NW
France, Tortonian, upper Miocene.
116 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Discussion – The specimens illustrated here belong to
the Bolma rugosa species-group. Landau et al. (2013,
p. 30) considered the Miocene populations of ‘Bolma
rugosa’to represent Bolma meynardi (Michelotti, 1847)
and restricted B. rugosa (Linné, 1767) to the Pliocene to
present-day populations. Bolma meynardi differs from B.
rugosa in having less stepped early teleoconch whorls re-
sulting in a lower spire, in having less nodular rugae, the
rugae extending to the sutures, and in B. rugosa the basal
callus is less extensive than in B. meynardi, but more ex-
cavated by a central depression, which ends as a double
denticle at the columellar abapical extremity, not present
in B. meynardi. Furthermore, the suture on the second
half of the last whorl in B. meynardi, when viewed aper-
turally, is placed between the base and mid-whorl height,
whereas in B. rugosa the suture is at or below the base.
The Pliocene and Recent forms, although somewhat dif-
ferent, cannot be separated consistently (see Landau et
al., 2013).
This species group is represented in the Assemblage I de-
posits by adults from Renauleau and one specimen from
St-Clément-de-la-Place, where all specimens found are
either incomplete or worn, suggesting transport prior
to deposition. Juveniles are found at other sites, such as
Sceaux-d’Anjou. Nevertheless, this upper Miocene Re-
nauleau population is interesting, as the shells show in-
termediate features between the two species compared
above. Like the typical Miocene B. meynardi, the shell is
low-spired (Pl. 40, gs 1a, 2a), the rugae are not nodular
and extend to the suture (Pl. 40, gs 1b, 2b), and the basal
callus does not end as a double denticle at the columel-
lar abapical extremity (Pl. 40, gs 1a, 2a, 3). Conversely,
with B. rugosa these Renauleau specimens share a rela-
tively narrow basal callus, which is strongly excavated
centrally in fully grown specimens (Pl. 40, gs 1c) and
in adult specimens the suture on the second half of the
last whorl is placed at the base (Pl. 40, gs 1a, 3). One
further difference is the strikingly bicarinate periphery,
the whorl prole concave between the two (Pl. 40, g.
1a), which persists onto the last whorl in adult specimens
(Pl. 40, g. 2a). Although both B. meynardi and B. rugosa
can have the periphery marked by stronger cords, they
are neither as strongly developed, nor is the whorl prole
between the cords as concave.
It is not possible to attribute the Renauleau specimens
to B. meynardi or B. rugosa. Millet (1854) erected the
name Turbo trochleatus for juvenile shells from Sceaux-
d’Anjou, but this is a junior homonym of T. trochleatus
zu Münster, 1841. The specimen illustrated by Brébion
(1964, pl. 3, gs 18-19) show a shell almost identical to
the juvenile illustrated here (Pl. 40, g. 2). Based on the
material at hand, we consider it undesirable to erect a
new species/subspecies. We prefer to leave the Renauleau
population in open nomenclature as B. cf. meynardi as
they are Miocene and the shell shape and sculpture are
closer to the Miocene form.
Landau et al. (2013, p. 31, text-g. 15/1, 32) illustrated a
large specimen from the upper Miocene Tortonian At-
lantic of Cacela (southern Portugal) that was typical for
B. meynardi and attributed all upper Miocene records
to this species. These intermediate forms from the more
northern Tortonian of northwestern France suggest the
modern species may have arisen at these more northern
latitudes.
Brébion (1964, p. 45) recorded this species [as A. (B.)
trochleata)] from Assemblage I localities of Sceaux-
d’Anjou, Thorigné, Les Pierres Blanches and Beaulieu.
Here we add St-Clément-e-la-Place and Renauleau. We
have not included his Assemblage III records which were
shown by Ceulemans et al. (2016a, p. 66) to be B. rugosa.
Brébion (1964, p. 134) recorded and illustrated specimens
ascribed to A. (B.) italica (Sacco, 1896) from Assemblag-
es I and II. Astraea (B.) italica was described as a sub-
species of Trochus muricatus Dujardin, 1837, which is a
junior synonym of Turbo baccatus Defrance, 1827 (Gli-
bert, 1949, p. 73) from the middle Miocene Loire Basin
of France. The specimens illustrated by Brébion lack the
tubercles characteristic of the species and we therefore
doubt this record. We have not seen any northwestern
French upper Miocene material we would ascribe to B.
baccata. It is more likely that the specimens illustrated
by Brébion are unusual juvenile specimens of Bolma cf.
meynardi, which is highly variable.
Distribution –Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Family Calliostomatidae Thiele, 1924
Subfamily Calliostomatinae Thiele, 1924
Genus Calliostoma Swainson, 1840
Subgenus Calliostoma s. str.
Type species (by subsequent designation, Herrmannsen,
1846) – Trochus conulus Linnaeus, 1758, present-day,
Europe.
1840 Calliostoma Swainson, p. 218.
For generic synonymy see Ceulemans et al. (2016a, p.
68).
Note – The abundance and diversity of calliostomids in
the upper Miocene Assemblage I localities of northwest-
ern France is one of the most striking aspects of this fau-
na. The species are almost all small shelled for the genus,
but outstandingly variable in shape, from depressed len-
tiform to tall conical with a coeloconoid spire, the whorls
regularly convex to strongly angular or biangular and the
surface almost unsculptured to strongly sculptured and
spiney. There is a far greater variation in shape than in
any other European Neogene fossil assemblage, and the
species shapes are certainly more varied than the more
or less regularly conical shell shape with spiral or beaded
sculpture living in European waters today. Indeed, the
only character all these forms share is the honeycomb
microsculpture on the protoconch, typical for the genus
Calliostoma (Hickman & McLean, 1990, p. 109). This
strikingly diverse shell form seen in the radiation of Cal-
liostoma species from Assemblage I suggests that gener-
Cainozoic Research, 17(2), pp. 75-166, December 2017 117
ic/subgeneric characters in calliostomids, based on shell
form, may not be an indication of monophyletic groups
and we therefore refrain from using them. We are also not
aware of any molecular studies supporting the division of
European calliostomids into genera/subgenera.
Although the genus was relatively well represented in the
lower Pliocene northwestern French Assemblage III lo-
calities discussed by Ceulemans et al. (2016a), these spe-
cies are similar in shape and size to Pliocene to present-
day congeners from Europe and share little in common
with the earlier Assemblage I calliostomids. Therefore,
the discussion and comparison following most of the new
species is relatively short due to the absence of similar
species with which to compare.
Calliostoma alternatum (Millet, 1865)
Plate 41, gs 1-2
1854 Trochus Alternatus Millet, p. 157 (nomen nudum).
*1865 Trochus alternatus Millet, p. 583.
1964 Calliostoma alternatum Millet, 1854 [sic] – Bré-
bion, p. 78, pl. 1, gs 22-24.
Type material – Syntypes: Sceaux-d’Anjou, lost (de
Brébion, 1964, p. 78).
Material and dimensions – Maximum height 6.4 mm;
width 4.8 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0058 (1), NHMW 2016/0103/0059 (1), NHMW
2016/0103/0060 (50+), LC (50+), FVD (50+). Sceaux-
d’Anjou: NHMW 2016/0103/0062 (48), RGM.1309652
(1), RGM.1309682 (20), RGM.1309694 (4), RGM.1347998
(12), LC (50+), FVD (50+). Renauleau: NHMW 2016/
0103/0061 (21), LC (40), FVD (23).
Discussion – Calliostoma alternatum (Millet, 1865) is
characterised by its small size, relatively tall narrow con-
ical spire and its sculpture composed of beaded cords,
three subequal cords below suture, the 4th cord stronger,
with some of the beads strengthened to form tubercles
followed by a 5th cord of equal strength to the three sub-
sutural cords. The base is imperforate and bears nine
subequal cords, slightly wider than their interspaces.
Axial growth lines are most strongly developed on the
base, weakly beading the basal cords in some speci-
mens. Although the most striking feature of this species
is the tuberculate 4th cord, its development is quite vari-
able, with some specimens hardly developing tubercles.
Millet (1854, 1865) recorded the species only from the
Assemblage I locality of Sceaux-d’Anjou, to which Bré-
bion (1964, p. 79) added St-Clément-de-la-Place and Les
Pierres Blanches. Renauleau is added herein.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Calliostoma baccatum (Millet, 1865)
Plate 42, gs 1-2
1854 Trochus Baccatus Millet, p. 157 (nomen nudum).
*1865 Trochus baccatus Millet, p. 582.
2016a Calliostoma baccatum (Millet, 1865) – Ceule-
mans et al., p. 70, pl. 9, gs 1, 2 (cum syn.).
Material and dimensions – Maximum height 15.0 mm;
width 12.8 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0156-0157 (2), NHMW 2016/0103/0159 (20), RGM.
1309761 (5), LC (10), FVD (27). Sceaux-d’Anjou: 2016/
0103/0158 (17), RGM.1309649 (18), RGM.1309700 (4),
RGM.1309775 (12), LC (15), FVD (17).
Original description – ‘Trochus baccatus, Millet. Coq.
en cône assez allongé, composée, de 8-9 tours de spire,
quelquefois un peu bombés et couverts de 5 rangs de
perles nes. Hauteur: 11-12 millimètres; diamètre à la
base: 8 millimètres. Elle présente une variété plus allon-
gée, dont les tours de spire sont bombés. Th., Sc, Ren.
(Millet, 1865, p. 582)’.
Revised description – Shell of medium size and thick-
ness; spire moderately elevated, conical. Protoconch
paucispiral, covered in honeycomb surface sculpture.
Teleoconch consisting of about 7-8 straight-sided whorls,
with periphery just above abapical suture. Suture linear,
supercial. Sculpture of ve beaded cords of irregular
strength, the two abapical cords become fused on later
Plate 41. Calliostoma alternatum (Millet, 1865); 1. NHMW 2016/0103/0058, height 6.4 mm, width 4.8 mm; 2. NHMW 2016/0103/0058,
detail of protoconch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, up-
per Miocene.
118 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
whorls, forming a raised bid peripheral band. Second-
ary cords intercalated in some to all interspaces on last
whorl. Last whorl roundly angled at periphery. Base not
sharply delimited, bearing about 8 narrow beaded cords.
Aperture tangential; peristome discontinuous, outer lip
not thickened, rounded at periphery. Columella sloping
weakly abaxially, bearing a weak columellar thickening
abapically.
The two specimens gured illustrate extremes in devel-
opment of the spiral sculpture, which can be nely, but
distinctly beaded to smooth, or almost so. Most speci-
mens show a beaded or sub-obsoletely beaded sculpture
intermediate between the two gured.
Discussion – Ceulemans et al. (2016a, p. 70) pointed out
that the established name for this species, Callio stoma
tauromiliare Sacco, 1896 was a junior subjective syno-
nym of Calliostoma baccatum (Millet, 1865). The As-
semblage III material at hand to Ceulemans et al. did
not have the protoconch preserved, however, the speci-
men from St-Clément-de-la-Place gured here (NHMW
2016/0103/0156; Pl. 42, g. 1) has a perfectly preserved
protoconch showing a paucispiral nucleus covered in a
honeycomb pattern, conrming its placement in the ge-
nus Calliostoma.
Brébion (1964, p. 75) recorded this species from locali-
ties belonging to Assemblage I-III and commented on
the great variability seen. In his description he noted that
the base could be convex or attened and the basal cords
beaded or not. Ceulemans et al. (2016a) considered the
Le Pigeon Blanc population not to be so variable. This
is not true of the Assemblage I populations which vary
greatly in apical angle, width and attening of the base,
just as described by Glibert (1949) and Brébion (1964).
Calliostoma baccatum is the only Calliostoma species
found both in Assemblage I and Assemblage III deposits.
Calliostoma baccatum (Millet, 1865) is supercially sim-
ilar to Jujubinus proximus (Millet, 1865). Both species
are of similar size and have ve rows of beaded primary
cords. In perfectly preserved specimens the protoconch
separates the two, as Jujubinus species have smooth pro-
toconchs. However, in most specimens the protoconch
surface sculpture is abraded. The teleoconch shell of
C. baccatum differs from J. proximus in usually hav-
ing a wider apical angle, the shell is less solid, especially
the outer lip, which is thinner in C. baccatum. On later
whorls the abapical two cords fuse and form a periph-
eral raised band in C. baccatum, which is not the case
in J. proximus, and there is more secondary sculpture,
which starts earlier. In J. proximus there is at most 1-2
spiral threads between the uppermost cords on the last
whorl. Finally, C. baccatum lacks the interal ridge within
the aperture bordering the siphonal canal and although
it does have a slight columella swelling abapically, it is
weaker than in J. proximus. For further discussion see
Ceulemans et al. (2016a, p. 70).
Distribution – Lower Miocene: Burdigalian, Italy (Sacco,
1896a); Paratethys, Austria (Harzhauser et al., 2015).
Middle Miocene: Loire Basin, France (Glibert, 1949). Up-
per Miocene: Atlantic, Tortonian, Messinian, NW France
(Millet, 1854, 1865; Brébion, 1964). Lower Pliocene:
Atlantic, NW France (Brébion, 1964; Ceulemans et al.,
2016a).
Calliostoma biangulatum nov. nom.
Plate 43, gs 1-2
1854 Trochus Heliciformis Millet, p. 156 (nomen nu-
dum).
1865 Trochus heliciformis Millet, p. 582 (non von Zie-
ten, 1832).
1964 Jujubinus (Strigosella) heliciformis Millet, 1854
[sic] – Brébion, p. 120, pl. 3, g. 13 (non T. helici-
formis von Zieten, 1832).
Type material – Syntypes: Thorigné, Sceaux-d’Anjou and
Renauleau, Musée d’Angers, France (de Brébion, 1964,
p. 120).
Material and dimensions – Maximum height 11.3 mm,
Plate 42. Calliostoma baccatum (Millet, 1865); 1. NHMW 2016/0103/0156, height 12.0 mm, width 9.0 mm; 2. NHMW 2016/0103/0157,
height 14.6 mm, width 11.8 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper
Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 119
width 9.2 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0131-0134 (4), NHMW 2016/0103/0135 (4), RGM.
1309598 (3), LC (1), FVD (5). Sceaux-d’Anjou: NHMW
2016/0103/0183 (18), RGM.1309596 (5), RGM.1309603
(14), RGM.1309701 (1), RGM.1309734 (1), RGM.1347918
(1), RGM.1347999 (7), LC (5), FVD (8).
Etymology – Name reecting the biangular shape of the
last whorl. Calliostoma gender neuter.
Locus typicus – Given as Sceaux-d’Anjou, Renauleau
and St-Clément-de-la-Place.
Stratum typicum – Tortonian, upper Miocene.
Original description – ‘Trochus heliciformis, Millet. Coq.
conique, composée de 7-8 tours de spire disposés en vis
et couverts de rangs de perles qui se touchent récipro-
quement. Dessous légèrement concave, marqué d’une
fente ombilicale et revêtu de rangs de perles alternant de
grosseur. Longueur: 13 millimètres; diamètre à la base,
10 millimètres. Sc, Th., Ren. (Millet, 1865, p. 582)’.
Revised description – Shell small, trochiform. Proto-
conch paucispiral, of 1.2 whorls bearing honeycomb sur-
face sculpture, the holes in the sculpture relatively large
(dp = 435 µm, dn = 170 µm). Junction with teleoconch
sharply delimited by elevated scar. Teleoconch of four
whorls. Spire whorls with at subsutural ramp, sharply
angled at shoulder, straight-sided to abapical suture.
Sculpture of ne spiral cords; two on subsutural ramp,
paired, closer-set cords at shoulder and above suture,
one further cord between cord-pairs at shoulder and su-
ture. Crowded prosocline lamellae cross cords, forming
short pointed tubercles at intersections, giving sculpture
spiney appearance. Last whorl biangular at shoulder
and base, prole slightly concave between angulations,
sculpture of two cords on subsutural platform, paired
cords forming shoulder and peribasal carina; two further
cords between carinae. Base sharply angled, strongly
depressed, bearing eight primary concentric cords, with
secondary threads in some interspaces. Umbilicus shal-
low, narrow. Aperture ovate, outer lip simple, not ared,
columella smooth. Columellar callus hardly developed,
parietal callus thin.
Discussion – With this species we offer a revised de-
scription to supplement the short original one given by
Millet (1865, p. 582). Unfortunately, Trochus heliciformis
Millet, 1865 is a junior homonym of T. heliciformis von
Zieten, 1832 and therefore invalid. We erect the replace-
ment name Calliostoma biangulatum nov. nom.
This species is very characteristic with its highly sculp-
tured surface and biangular last whorl. There are no Eu-
ropean fossil or present-day congeners with which it can
be usefully compared. Brébion (1964) placed it in the ge-
nus/subgenus Jujubinus (Strigosella), but the honeycomb
patterned protoconch shows it to be a calliostomiid (Pl.
43, g. 2).
Millet recorded this species from the Assemblage I local-
ities of Sceaux-d’Anjou, Renauleau, Thorigné, to which
Brébion (1964, p. 121) added St-Clément-de-la-Place. We
note that Brébion did not rend the species at Renauleau
and despite intensive collecting neither did we.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma contractum (Millet, 1865)
Plate 44, gs 1-7
1854 Trochus Depressus Millet, p. 156 (non Gmelin,
1791).
1854 Trochus Contractus Millet, p. 157 (nomen nudum).
1865 Trochus depressus Millet, p. 582 (non Gmelin,
1791).
*1865 Trochus contractus Millet, p. 583.
Plate 43. Calliostoma biangulatum nov. nom.; 1. NHMW 2016/0103/0131, height 5.8 mm, width 5.5 mm; 2. NHMW 2016/0103/0134,
detail of protoconch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, up-
per Miocene.
120 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
1964 Calliostoma contractum Millet, 1854 [sic] – Bré-
bion, p. 97, pl. 2, g. 22.
1964 Gibbula (Steromphala) depressa Millet, 1854
[sic] – Brébion, p. 111, pl. 3, g. 6 (non Trochus
depressus Gmelin, 1791).
Type material – Syntypes: Thorigné and Sceaux-d’Anjou,
lost (de Brébion, 1964, p. 98).
Material and dimensions – Maximum height 15.2 mm;
width 14.6 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0167-0169 (3), NHMW 2016/0103/0256-0259 (4),
NHMW 2016/0103/0170 (50+), LC (1), FVD (22). Sceaux-
d’Anjou: NHMW 2016/0103/0171 (45), RGM.1309630
(1), RGM.1309665 (3), RGM.1309678 (7), RGM.1309689
(10), RGM.1309692 (2), RGM.1309704 (6), RGM.1309730
(7), RGM.1309768 (10), RGM.1348000 (19), LC (10),
FVD (27). Renauleau: NHMW 2016/0103/0172 (12), LC
(5), FVD (3).
Discussion – Millet (1865) validated his earlier (1854)
name Trochus contractus with the following description:
Plate 44. Calliostoma contractum (Millet, 1865); 1. NHMW 2016/0103/0167, height 5.0 mm, width 5.6 mm; 2. NHMW 2016/0103/0168,
height 7.4 mm, width 8.8 mm; 3. NHMW 2016/0103/0169, detail of protoconch (SEM image); 4. NHMW 2016/0103/0256, height
8.8 mm, width 8.5 mm; 5. NHMW 2016/0103/0257, height 8.1 mm, width 8.2 mm; 6. NHMW 2016/0103/0258, height 6.7 mm,
width 7.9 mm; 7. NHMW 2016/0103/0259, height 6.2 mm, width 7.7 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-
et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 121
‘Trochus contractus. Millet. Coq. en cône racourci, aiguë
au sommet, composée de 6 tours de spire disjoints par
une suture assez profonde, couronnant un rang de petites
perles. Chaque tour en outre présente 5 petits cordonnets.
Dessous légèrement concave, couvert de stries nes. Hau-
teur et diamètre: 8 à 9 millimètres. Sc, Th. (1865, p. 583)’.
Although this is a fair description of the predominant
form, this is probably the most variable of the Assemblage
I calliostomiids. The shell can be strongly depressed (Pl.
45, gs 6-7) to regularly conical elevated (Pl. 45, gs 4-5),
although as described by Millet, in all forms the apex is
pointed. Only the most adap ical cord is strongly beaded,
with the second cord weakly beaded in most specimens
(Pl. 45, g. 1), although more abapical cords are beaded
in some specimens (Pl. 45, gs 2, 6) and occasionally all
cords are beaded (Pl. 45, gs 5, 7). The basal angulation
can be roundly (Pl. 45, gs 1, 4) to acutely angled (Pl. 45,
gs 2, 6, 7). The basal sculpture of close-set, non-beaded
concentric cords is well developed in most specimens, al-
though the thickness of the cords is variable, and in some
specimens the cords are obsolete mid-base (Pl. 45, g. 7).
Millet erected another taxon Trochus depressus Millet,
1854 (nomen nudum) described later in Millet (1865) (jun-
ior homonym of Trochus depressus Gmelin, 1791) as: ‘Tro-
chus depressus, Millet. Coq. orbiculaire, légèrement co-
nique, composée de 5 tours de spire, le dernier plus grand
que tous les autres ensemble, caréné inférieurement, por-
tant 6 stries, dont la plus élevée, plus grosse que les autres,
est couverte de petites perles. La partie inférieure munie
d’un trou ombilical, est couverte de très-petites stries.
Hauteur: 6 à 7 millimètres; diamètre: 8-9 millimètres.
Sceaux. Rare (1865, p. 582). The main difference between
Millet’s two species is that T. contractus has an imperfo-
rate base and T. depressus has a small umbilicus. In our
opinion T. depressus applies to the extremely attened
form of this species, but numerous intermediates occur,
with a small deep umbilicus (Pl. 45, gs 1, 2, 6), shallow
(Pl. 45, gs 7) to imperforate (Pl. 45, g. 4).
There is little with which to compare this species. Bré-
bion (1964, p. 98) compared it to C. conuloides (Lamarck,
1822), which is now considered a junior subjective syn-
onym of C. zizyphinum (Linnaeus, 1758), but this is a
much larger species, with a less depressed shell and a
more regularly conical spire, and the adapical cords are
not usually beaded, and if so less strongly.
Millet (1854, 1865) recorded his two species T. depres-
sus and T. contractus, herein synonymised from the As-
semblage I localities of Sceaux-d’Anjou and Thorigné, to
which Brébion added Renauleau, St-Clément-de-la-Place
and Les Pierres Blanches.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Calliostoma gibbuliforme nov. sp.
Plate 45, gs 1-3
1964 Calliostoma multistriatum var. andegavensis Bré-
bion, p. 79, pl. 1, g. 25 (nomen nudum).
Type material – Holotype MNHN.F.A57692, height
3.5 mm, width 4.4 mm; paratype 1 MNHN.F.A57693,
height 3.7 mm, width 5.0 mm; paratype 2 NHMW
2016/0103/0123, height 4.4 mm, width 5.1 mm; paratype
3 NHMW 2016/0103/0124, height 4.0 mm, width 5.1
mm; paratype 4 NHMW 2016/0103/0125, height 2.3 mm,
width 3.2 mm; paratype 5 RGM.1309749, height 3.1 mm,
width 3.6 mm; paratype 6 RGM.1309750, height 3.2 mm,
width 3.6 mm.
Other material – Maximum height 3.2 mm, width 4.3
mm. St-Clément-de-la-Place: NHMW 2016/0103/0126
(28), RGM.1309751 (12), RGM.1309757 (6), LC (10),
FVD (14). Sceaux-d’Anjou: NHMW 2016/0103/0219 (14),
RGM.1348001 (3), LC (5), FVD (8). Renauleau: LC (1).
Etymology – Name reecting the shell shape resembling
a Gibbula species. Calliostoma gender neuter.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Calliostoma species of small size, low trochi-
form shell shape, with spiral sculpture of which only rst
and sometimes second cords are beaded, spiral sculpture
weakening abapically, strongly convex last whorl, round-
ed or slightly angular at peripheral cord, attened base
with weak concentric sculpture restricted to periumbili-
cal area, and deep, narrow umbilicus.
Description – Shell small, depressed, low trochiform.
Protoconch paucispiral, of 1.2 whorls bearing honey-
comb surface sculpture with small pits (dp = 435 µm, dn
= 215 µm). Junction with teleoconch sharply delimited
by abrupt end to protoconch sculpture. Teleoconch of 3.5
low convex whorls, with periphery at abapical suture,
separated by impressed suture. Sculpture on rst tele-
oconch whorl of three rounded spiral cords, increasing in
number to four on second teleoconch whorl, ve on pe-
nultimate whorl. Cords strongly developed and rounded
on some specimens, attened in other, subobsolete on the
last teleoconch whorl in most specimens. The adapical
cord is conspicuously beaded by prosocline axial growth
lines; in some specimens the second cord is also weakly
beaded, abapical cords smooth, one or two most abapi-
cal cords more strongly developed. Last whorl depressed,
rounded or slightly angular at peripheral cord, base at-
tened, smooth, except for 3-4 weak cords adjacent to,
and strengthening towards umbilicus. Umbilicus nar-
row, deep. Aperture ovate, outer lip simple, rounded to
somewhat angled at periphery, slightly ared abapically.
Columella short, smooth. Columellar callus thickened,
sharply delimited. Colour pattern is preserved in some
specimens consisiting of tiny white dots over the cords
(Pl. 46, g. 3).
Discussion – This new species, together with Calliosto-
ma quaggaoides nov. sp. (see below), form a small group
122 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
of Calliostoma Swainson, 1840 species that at rst sight
would be placed in the Cantharidinae Gray, 1857, genus
Gibbula Risso, 1826, but they lack a columellar fold and,
more importantly, they have the honeycomb protoconch
microsculpture so typical of the genus Calliostoma.
Calliostoma gibbuliforme nov. sp. is variable in shape;
the last whorl periphery is rounded in some specimens, a
bit angular in others. The number of spiral cords is con-
stant, but their strength is strongly variable. One, or less
frequently the two most adapical cords, are beaded and
one, or less frequently two of the most abapical cords, are
more strongly developed forming one or two peripheral
carinae. The character of the base and aperture are rela-
tively constant.
The only species with which C. gibbuliforme could be
confused is C. quaggaoides, which differs in having an
even more depressed shell, with a single subsutural cord,
which is not beaded and having colour pattern preserved
in almost all specimens consisting of axial stripes rather
than spots as seen in C. gibbuliformis.
Brébion (1964, p. 79) described this species based on two
specimens from Renauleau as var. andegavensis of Cal-
liostoma multistriatum (Wood in Kendall & Bell, 1886)
from the Gelasian Pleistocene of St Erth, England, how-
ever, that species is even more depressed with a lower
spire and weaker spiral sculpture.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma echinatum (Millet, 1865)
Plate 46, g.1
1854 Trochus Echinatus Millet, p. 157 (nomen nudum).
*1865 Trochus echinatus Millet, p. 583.
1911 Eucyclus echinatus Millet – Cossmann, p. 328.
1964 Eucyclus (?) echinatus Millet, 1854 [sic] – Bré-
bion, p. 70, pl. 1, g. 17.
Type material – Syntypes: Sceaux-d’Anjou and Thorigné,
lost (de Brébion, 1964, p. 71).
Material and dimensions – Maximum height 7.4 mm;
width 6.2 mm. Sceaux-d’Anjou: NHMW 2016/0103/0144
(1), NHMW 2016/0103/0145 (9), RGM.1309590 (2), RGM.
1348019 (2 fragments), LC (2), FVD (3). Renauleau:
NHMW 2016/0103/0146 (1), LC (2).
Discussion – Millet (1865) validated his earlier (1854)
nomen nudum with this description: ‘Trochus echinatus,
Millet. Coq. petite, conique, aiguë , composée de 7 tours
de spire couverts d’expansions lamellaires, placées par
rangées et disposes en tubes plus ou moins ouverts en
dessous et tronqués au sommet. Ces expansions ou tubes
qui s’emboîtent les uns dans les autres en se relevant au
sommet, présentent un rang plus saillant qui occupe le
centre de chaque tour. Hauteur: 7 millimètres; diamètre:
5-6 millimètres. Sc., Ren. Très-remarquable et très rare
espèce (1865, p. 583).’ This description highlights the
Plate 45. Calliostoma gibbuliforme nov. sp.; 1. Holotype MNHN.F.A57692, height 3.5 mm, width 4.4 mm; 2. Paratype 4 NHMW
2016/0103/0125, detail of protoconch (SEM image); 3. Paratype 2 NHMW 2016/0103/0123, height 4.4 mm, width 5.1 mm. Le
Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 123
most outstanding character of this species, which is the
long open lamellar spines that develop at the intersec-
tions where the axial lamellae cross the primary cords.
Cossmann (1918, p 328) placed this shell in the genus
Eucyclus Eudes-Deslongchamps, 1860, member of the
Eucyclidae Koken, 1896. However, in our opinion the re-
semblance to shells of this genus is supercial. Although
not gured as all of the specimens have an abraded proto-
conch, remnants of the typical calliostomiid honeycomb
pattern are preserved in some shells. The only species
with a similar shell is Calliostoma lamellatum nov. sp.
(see below), which has the same tall conical trochiform
shell shape and type of sculpture, but differs in having
more numerous spiral cords and axial lamellae that are
scabrous at the sculptural intersections, but not spinous as
they are in C. echinatum. Although, as discussed above,
the protoconch surface is abraded in the specimens of
C. echinatum at hand, it is well-preserved in one of our
specimens of C. lamellatum and shows the calliostomi-
id honeycomb surface pattern (Pl. 48, g. 2). Therefore
we place both of these species in the genus Calliostoma
Swainson, 1840.
Both Millet (1854, 1865) and Brébion (1964, p. 71) re-
corded this species from Assemblage I localities of
Sceaux-d’Anjou and Renauleau.
We note that the placement of Millet’s species in the ge-
nus Calliostoma renders the present-day Caribbean C.
echinatum Dall, 1881 a secondary homonym. We there-
fore propose the name C. caribbechinatum nom. nov. for
Dall’s species.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Calliostoma gratiosum (Millet, 1865)
Pl. 47, gs 1-4
1854 Trochus Gratiosus Millet, p. 156 (nomen nudum).
*1865 Trochus gratiosus Millet, p. 582.
1964 Calliostoma (Ampullotrochus) laureatus Mayer,
1874 – Brébion, p. 98, pl. 2, gs 23-26 (non May-
er, 1874).
Material and dimensions – Maximum height 23.9 mm,
width 21.4 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0161 (1), NHMW 2016/0103/0150 (1), NHMW 2016/
0103/0162 (16), RGM.1309672 (7), RGM.1309707 (4), LC
(3), FVD (19). Sceaux-d’Anjou: NHMW 2016/0103/0163
(1), NHMW 2016/0103/0164 (50+), RGM.1309651 (9),
RGM.1309670 (6), RGM.1309696 (7), RGM.1309706 (2),
RGM.1309718 (19), RGM.1309777 (8), LC (10), FVD (15).
Renauleau: NHMW 2016/0103/174 (15), LC (10), FVD
(8).
Discussion – Millet (1865) validated his earlier (1854)
name Trochus gratiosus with the following description:
‘Trochus gratiosus. Millet. Coq. en cône très-large à sa
base, compose de 7-8 tours de spire aplatis, couverts de
nes stries perlées, séparées les unes des autres par un
let saillant placé au-dessus de la suture: let également
perlé, et portant en outre un certain nombre de perles
plus grosses, éloignées les unes des autres d’environ 3
millimètres. Hauteur: 14-15 millimètres; diamètre à la
base: 13-14 millimètres. Th., Sc, Ren. (1865, p. 582)’
This description characterises the shell relatively well;
we would add that both the adapical and abapical cords
bordering the suture are more strongly developed, ac-
centuating the suture and making it deeply impressed
and that the ne beaded cords are of alternate strength
on both the dorsum and the base. As noted by Millet, on
the abapical cord every 3 mm or so three beads strength-
en, the central one most, giving the periphery a slightly
crenulated appearance. The periphery of the last whorl
is delimited by two strengthened cords of roughly equal
weight. The protoconch is paucispiral and typical for cal-
liostomids, covered in a honeycomb pattern (Pl. 47, g.
3). One specimen from Renauleau (Pl. 47, g. 4) is twice
the size of specimens from other Assemblage I localities
sampled, has ner sculpture and the peripheral cords on
Plate 46. Calliostoma echinatum (Millet, 1865); 1. NHMW 2016/0103/0144, height 5.8 mm, width 4.4 mm. La Presselière, Sceaux-
d’Anjou, Maine-et-Loire, NW France, Tortonian, upper Miocene.
124 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
the last whorl are weaker. However, we consider it to t
within the intraspecic variability for Calliostoma gra-
tiosum (Millet, 1865).
Brébion (1964, p. 98) considered Millet’s species conspe-
cic with Trochus laureatus Mayer, 1874 described from
the upper Pliocene of Italy, giving priority to the junior
name. We have not seen Mayer’s type, but from the de-
scription and his own discussion he likens it to C. granu-
latum (von Born, 1778), saying it differs in being smaller,
more pointed, with ner spiral sculpture and a crenulated
base. The description and comparison also place it close
to two other Mediterranean Pliocene species illustrated
by Landau et al. (2003) belonging to the same species
group: C. opisthetonus (Fontannes, 1880) and C. scuti-
formis (Sacco, 1896). All these species differ from C.
gratiosum in being larger shelled, in having more coelo-
conoid spires and therefore a more pointed apex, coarser,
less strongly beaded spiral sculpture and a more attened
base. As mentioned above, we have not seen Mayer’s
species, but the shell illustrated does not seem conspe-
cic with C. gratiosum. The shells illustrated by Glibert
(1952, pl. 1, g. 4) as C. laureatum from the North Sea
Basin Belgian Miocene differ from C. gratiosum in the
same characters discussed above. The shell illustrated by
Janssen (1984, pl. 44, g. 3) as C. laureatum from the
Dutch Miocene differs slightly from other illustrations
of that species in having a rounded periphery on the last
whorl not delimited by strengthened basal cords.
Millet (1854, 1865) recorded this speces from the As-
semblage I localities of Thorigné, Sceaux-d’Anjou and
Renauleau, to which Brébion (1964, p. 99) added St-Clé-
ment-de-la-Place and Contigné.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Calliostoma lamellatum nov. sp.
Plaate 48, gs 1-2
Type material – Holotype NHMW 2016/0103/0139,
height 4.5 mm, width 3.4 mm; paratype 1 NHMW
2016/0103/0140, height 4.2 mm, width 3.3 mm; paratype
2 NHMW 2016/0103/0141, height 4.4 mm, width 2.5 mm
(juvenile); paratype 3 NHMW 2016/0103/0142, height
3.4 mm, width 2.9 mm (juvenile); St-Clément-de-la-
Place. Paratype 4 RGM.1309625, height 5.1 mm, width
3.4 mm; paratype 5 RGM.1309626, height 4.0 mm, width
2.9 mm; Sceaux-d’Anjou.
Other material – Maximum height 6.9 mm, width 4.5
mm. Sceaux-d’Anjou: NHMW 2016/0103/0143 (9),
RGM.1309627 (1 juvenile), RGM.1348002 (2). Re-
nauleau: NHMW 2016/ 0103/0160 (5).
Plate 47. Calliostoma gratiosum (Millet, 1865); 1. NHMW 2016/0103/0161, height 14.9 mm, width 12.9 mm. Le Grand Chauvereau,
St-Clément-de-la-Place; 2. NHMW 2016/0103/0163, height 12.6 mm, width 11.6 mm. La Presselière, Sceaux-d’Anjou; 3. NH-
MW 2016/0103/0150, detail of protoconch (SEM image); 4. NHMW 2016/0103/1497, height 21.5 mm, width 20.1 mm. Renauleau,
Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 125
Etymology – Name reecting the characteristic raised
axial lamellae seen in this species. Calliostoma gender
neuter.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Calliostoma species of small size, with
tall turbiniform shell shape, spiral sculpture of narrow
cords with stronger peripheral and peribasal cords on last
whorl, entire surface covered in prosocline axial lamel-
lae, giving surface roughly scabrous appearance, and
attened, imperforate base.
Description – Shell small, tall trochiform, with coni-
cal spire. Protoconch paucispiral, of 1.3 whorls, bearing
honeycomb surface sculpture. Junction with teleoconch
sharply delimited by protoconch lip. Teleoconch of four
whorls. Suture linear, supercial. First teleoconch whorl
bearing two elevated spiral cords of roughly equal strength
and raised, prosocline axial lamellae forming an evenly
reticulate sculpture. On second whorl, a third weaker cord
develops below adapical suture; abapical cord placed just
above abapical suture strengthens forming periphery. On
third whorl a fourth weaker cord develops again just be-
low adapical suture. Last whorl with two narrow cords
above stronger peripheral cord, below which lies an even
stronger peribasal cord, delimiting base. Base attened,
imperforate, bearing eight narrow spiral cords, strength-
ening slightly towards centre. Entire basal surface cov-
ered with raised axial lamellae, which cross spiral sculp-
ture, giving surface a roughly squamous appearance.
Aperture rounded, outer lip simple, rounded at periphery,
columella smooth. Columellar callus hardly developed,
parietal callus not developed.
Discussion – As mentioned above, Calliostoma lamella-
tum nov. sp. is similar to Calliostoma echinatum (Millet,
1865), but differs in having more numerous spiral cords
above the peripheral cord, in having the prominent pe-
ripheral and basal cords less strongly developed than in
C. echinatum, and in having more numerous axial lamel-
lae which are scabrous at the sculptural intersections, but
do not form spines.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma michaeli nov. sp.
Plate 49, gs 1-2
Type material – Holotype NHMW 2016/0103/1498, height
15.0 mm, width 12.5 mm; paratype 1 NHMW 2016/
0103/1499, height 15.6 mm, width 12.4 mm; paratype 2
MNHN.F.A57939, height 14.7 mm, width 12.3 mm.
Other material – Maximum height 16.0 mm, width 13.3
mm. NHMW 2016/0103/1500 (4), LC (5), FVD (6).
Etymology – Named after Michael Van Dingenen, son of
the second author. Calliostoma gender neuter.
Plate 48. Calliostoma lamellatum nov. sp.; 1. Holotype NHMW 2016/0103/0139, height 4.5 mm, width 3.4 mm. 2. Paratype 3
NHMW 2016/0103/0142, detail of protoconch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire,
NW France, Tortonian, upper Miocene.
126 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Locus typicus – Renauleau, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Calliostoma species, of medium-size, with
relatively tall spire, whorls becoming progressively more
convex, rounded last whorl, sculpture four beaded pri-
mary cords with secondary beaded cord in interspaces on
last whorl, and rounded imperforate base bearing broad
concentric cords.
Description – Shell of medium size and thickness, tro-
chiform, with relatively tall, somewhat scalate spire. Pro-
toconch paucispiral, of 1.3 whorls; surface abraded in all
specimens. Teleoconch of ve whorls, with periphery at
suture. Suture linear, supercial. First half teleoconch
whorl bearing two spiral cords and prosocline axial ribs,
slightly weaker than cords, forming reticulate sculpture.
Second half of rst whorl, a third cord develops below
adapical suture, rapidly gaining in strength, becoming
predominant; axial ribs disappear, cords become beaded.
Second whorl, fourth beaded cord appears above suture.
Third whorl bearing four beaded cords; stronger subsu-
tural cord and three weaker subequal cords below. Pe-
nultimate and last whorls becoming progressively more
convex, with secondary beaded cords intercalated in the
interspaces. Last whorl convex, bearing beaded cords of
alternate strength, rounded at periphery. Peribasal cord
bid, base rounded, imperforate, bearing 6-7 prominent
irregular, subobsoletely beaded cords. Aperture sub-
quadrate, outer lip simple, rounded at periphery, colu-
mella smooth. Columellar callus thickened and everted,
parietal callus thin.
Discussion – Calliostoma michaeli nov. sp. is one of the
larger Calliostoma species in the Assemblage I fauna. Its
sculpture of four primary beaded cords on the later whorls
is somewhat similar to that of C. spinosum nov. sp., but
the secondary cords are far more developed; in C. spino-
sum secondary sculpture is absent or restricted to a single
thread between cords 1 and 2 on the last whorl. Moreo-
ver, C. michaeli is larger shelled, the later whorls are more
convex and the basal cords are not as clearly beaded as
they are in C. spinosum. Calliostoma baccatum (Millet,
1865), also from the Assemblage I fauna, has a more reg-
ularly and broadly conical spire, less convex later spire
whorls, and an angled base with ner basal sculpture.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma microgemmatum nov. sp.
Plate 50, g. 1
Type material – Holotype NHMW 2016/0103/0136,
height 5.0 mm, width 4.4 mm; paratype 1 NHMW
2016/0103/0137, height 4.1 mm, width 4.3 mm ( juvenile);
paratype 2 NHMW 2016/0103/0138, height 3.1 mm, width
3.4 mm (juvenile); St-Clément-de-la-Place. Paratype 3
RGM.1309680, height 3.8 mm, width 3.9 mm; paratype
4 RGM.1309733, height 4.9 mm, width 5.4 mm; Sceaux-
d’Anjou. Paratype 5 MNHN.F.A57942, height 4.7 mm,
width 4.1 mm; paratype 6 MNHN.F.A57943, height 4.4
mm, width 4.1 mm; Renauleau.
Other material – Maximum height 6.5 mm, width 5.3
mm St-Clément-de-la-Place: LC (1). Sceaux-d’Anjou:
RGM.1348003 (6), LC (1). Renauleau: NHMW 2016/
0103/0198 (18), LC (5), FVD (7).
Etymology – Compound name from Latin ‘micro-’ prex
meaning tiny, and ‘gemmatum’, adjective meaning be-
jewelled, describing the sculpture of nely beaded cords.
Calliostoma gender neuter.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Plate 49. Calliostoma michaeli nov. sp.; 1. Holotype NHMW 2016/0103/1498, height 15.0 mm, width 12.5 mm; 2. Paratype 1
NHMW 2016/0103/1499, height 15.6 mm, width 12.4 mm. Renauleau, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 127
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Calliostoma species of small size, with con-
cave whorl prole from second teleoconch whorl, sculp-
tured by close-set, nely beaded spiral cords, with prom-
inent suprasutural cord delimiting base on last whorl,
and attened, imperforate base bearing similarly nely
beaded cords.
Description – Shell small, trochiform. Protoconch pau-
cispiral, of 1.3 whorls bearing honeycomb surface sculp-
ture. Junction with teleoconch sharply delimited by end
of protoconch sculpture. Teleoconch of four whorls. Su-
ture linear, impressed. First teleoconch whorl convex,
bearing eight narrow spiral cords, close-set in adapical
half of whorl, strengthening slightly and wider spaced
towards lower suture. From second whorl, prole con-
cave. Spiral cords increase in number, ten on penultimate
and last whorl, irregular in strength. A suprasutural cord
becomes swollen, rounded and elevated, forming periph-
ery. Axial growth lines increase in strength from second
whorl, nely beading spiral sculpture. Last whorl regu-
larly concave to prominent peribasal cord. Base imperfo-
rate, strongly depressed, bearing about 13 nely beaded
cords, strengthening and wider-set towards centre. Aper-
ture subquadrate, outer lip simple, angled at periphery,
columella smooth. Columellar callus hardly developed,
parietal callus reduced to thin callus wash.
Discussion – Calliostoma microgemmatum nov. sp. is
uncommon in the Assemblage I localities. It is a char-
acteristic species with its concave teleoconch whorls
and nely beaded spiral sculpture. The protoconch was
too worn to image in all specimens, but remnants of
the typical calliostomiid honeycomb surface sculpture
are present. It is difcult to nd similar congeners with
which to compare this species; small specimens of Cal-
liostoma gratiosum (Millet, 1865), also from the Assem-
blage I fauna are similar in having very nely gemmate
sculpture, but the whorls prole is less concave, and a
spiral on either side of the suture becomes reinforced,
whereas in C. microgemmatum only the suprasutural
cord becomes swollen.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma milletigranum nov. nom.
Plate 51, gs 1-3
1854 Trochus Millegranus Millet, p. 157 (nomen nudum).
1865 Trochus millegranus Millet, p. 584 (non Philippi,
1836).
1964 Calliostoma milleti Brébion, p. 91, pl. 2, gs 12-
13. (nom. nov. pro T. millegranus Millet, 1865,
non Philippi, 1836) (nomen nudum).
Type material – Syntypes: Sceaux-d’Anjou, lost (de
Brébion, 1964, p. 91).
Material and dimensions – Maximum height 13.7
mm; width 13.1 mm. Sceaux-d’Anjou: NHMW 2016/
0103/1491 (1), NHMW 2016/0103/0201-202 (2), NHMW
2016/0103/0203 (2), RGM.1309664 (2), RGM.1309684
(2), RGM.1309770 (2), LC (1). Renauleau: NHMW
2016/0103/1483 (2 adult + 6 juveniles), LC (2), FVD (1).
Etymology – Compound name reecting Brébion’s (1964)
intention of naming the species after Millet and Millet’s
original epiphet describing the beaded sculpture. Callio-
stoma gender neuter.
Locus typicus – Sceaux-d’Anjou, Maine-et-Loire, NW
France.
Stratum typicum – Tortonian, upper Miocene.
Original description – ‘Trochus millegranus, Millet.
Coq. en cône court, composée de 5 à 6 tours de spire;
les premiers saillants, arrondis, les derniers, sur un plan
oblique, présentent une surface plate; et tous sont cou-
verts de petits grains plus ou moins arrondis, qu’on ne
peut voir qu’au foyer d’une forte loupe. Le dessous légè-
rement bombé est couvert de stries nes, peu apparentes.
Hauteur: 9-10 millimètres; diamètre: 8 millimètres. Sc.
Très-rare. (Millet, 1865, p. 584)’.
Plate 50. Calliostoma microgemmatum nov. sp.; 1. Holot ype NHMW 2016/0103/0136, height 5.0 mm, width 4.4 mm. Le Grand
Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
128 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Discussion – Millet’s (1865, p. 583) description faithful-
ly describes this species and validates his earlier nomen
nudum (Millet, 1854, p. 157). Unfortunately it is a junior
homonym of T. millegranus Philippi, 1836, which is it-
self a junior subjective synonym of the Pliocene present-
day Mediterranean Clelandella miliaris (Brocchi, 1814).
This is probably the most nely sculptured of the As-
semblage I calliostomids, with the greatest number of
spiral cords, about 14 on the penultimate whorl, which
are very nely beaded by the prosocline growth lines.
The height of the spire and the basal angulation are quite
variable, as seen in the series illustrated (Pl. 51, gs 1-3).
The base is poorly delimited and similarly sculptured by
about 18 cords.
Millet (1854, 1865) recorded the species from the As-
semblage I locality of Sceaux-d’Anjou, to which Brébion
(1964, p. 92) added Renauleau.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Calliostoma miotorulosum nov. nom.
Plate 52, gs 1-3
1865 Trochus torulosus Millet, p. 583 (non Philippi,
1845).
1964 Calliostoma torulosum Mayer, 1874 – Brébion, p.
92, pl. 2, g. 14. (non T. torulosus Philippi, 1845).
Type material – Syntypes: Sceaux-d’Anjou, Musée d’An-
gers, France ( de Brébion, 1964, p. 93).
Material and dimensions – Maximum height 9.8 mm;
width 7.1 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0175 (1), NHMW 2016/0103/0180 (1), NHMW 2016/
0103/0176 (15), RGM.1309745 (2), LC (1), FVD (16).
Sceaux-d’Anjou: NHMW 2016/0103/0177 (1), NHMW
2016/0103/0178 (15), RGM.1309631 (7), RGM.1309663
(4), RGM.1309681 (11), RGM.1309697 (1), RGM.1309712
(4), RGM.1309717 (1), RGM.1309722 (3), RGM.1309728
(4), RGM.1309778 (1), RGM.1348004 (3), LC (5), FVD
(8). Renauleau: NHMW 2016/0103/0179 (11), LC (5),
FVD (10). Beugnon: RGM.1309677 (1).
Etymology – Honouring Millet’s original trivial name
‘torulosus’, adding the prex ‘mio-’ reecting the Mio-
cene period in which it lived. Calliostoma gender neu-
ter.
Locus typicus – Sceaux-d’Anjou, Maine-et-Loire, NW
France.
Stratum typicum – Tortonian, upper Miocene.
Original description – ‘Trochus torulosus, Millet. Coq.
conique, aiguë, composée de 7-8 tours de spire cou-
verts de stries nes, coupées ou ciselées obliquement
par des stries plus nes encore. Chaque tour, en outre,
est bordé inférieurement par un fort bourrelet arrondi,
ciselé verticalement, et sur lequel on remarque çà et là
quelques renements en forme de perle. Dessous an-
guleux, couvert de stries simples. Hauteur: 10-11 mil-
limètres; diamètre: 9-10 millimètres. Sceaux. (Millet,
1865, p. 583)’.
Discussion – Millet’s (1865, p. 583) description faith-
fully describes this species. Unfortunately it is a junior
homonym of T. torulosus Philippi, 1845, which is itself
a junior subjective synonym of the present-day south
Plate 51. Calliostoma milletigranum nov. nom.; 1. NHMW 2016/0103/1491, height 10.8 mm, width 8.4 mm; 2. NHMW 2016/0103/0201,
height 8.1 mm, width 7.8 mm; 3. NHMW 2016/0103/0202, height 7.7 mm, width 7.4 mm. La Presselière, Sceaux-d’Anjou, Maine-
et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 129
eastern Pacic Tegula quadricostata (W. Wood, 1828).
This small, but strongly sculptured calliostomiid, with its
coeloconoid spire and strongly developed crenulated su-
prasutural cord, which on the last whorl forms the acutely
angular periphery to the base is difcult to confuse with
any of it congeners. The protoconch is typically callios-
tomiid with a honeycomb sculpture composed of rather
narrow netting and wide holes in between (dp = 465 µm,
dn= 215 µm).
Millet (1865, p. 583) recorded the species only from
Sceaux-d’Anjou, but it is widely distributed in the As-
semblage I localities (St-Clément-de-la-Place, Chalon-
nes, Renauleau; Brébion, 1964, p. 93).
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1865; Brébion, 1964).
Calliostoma miotumidum nov. nom.
Plate 53, g. 1
1854 Trochus Tumidus Millet, p. 157 (nomen nudum).
1865 Trochus tumidus Millet, p. 583 (non T. tumidus
Montagu, 1803).
1964 Calliostoma tumidum Millet, 1854 [sic] – Brébi-
on, p. 96, pl. 2, gs 19-20 (?21).
Type material – Syntypes: Thorigné and Sceaux-d’Anjou,
Musée d’Angers, France ( de Brébion, 1964, p. 97).
Material and dimensions – Maximum height 14.5 mm,
width 14.2 mm. Sceaux-d’Anjou: NHMW 2016/ 0103/1504
(1), NHMW 2016/0103/1505 (2), LC (2), FVD (1).
Etymology – Honouring Millet’s original trivial name
‘tumidus’, adding the prex ‘mio-’ reecting the Miocene
period in which it lived. Calliostoma gender neuter.
Locus typicus – Sceaux-d’Anjou, Maine-et-Loire, NW
France.
Stratum typicum – Tortonian, upper Miocene.
Original description – ‘Trochus tumidus, Millet. Coq. en
cône raccourci, aiguë au sommet, composée de 7 tours
de spire arrondis et comme gonés, surtout les derniers
tours; tous sont couverts de stries nes et rapprochées
entre elles et présentent à la base un petit let que dé-
tache la suture. Dessous bombé, lisse ou marqué seule-
ment de quelques stries légères vers remplacement de
l’ombilic. Hauteur et diamètre: 14 millimètres. Sc, Th.’
(Millet, 1865, p. 583).
Revised description – Shell of medium size and thick-
ness, low trochiform, with wide apical angle. Protoconch
not preserved. Teleoconch of up to seven whorls; apex
pointed, early whorls straight sided, last two whorls in-
ated. Suture supercial, linear. First two teleoconch
whorls sculptured by ve narrow cords, adapical cord
Plate 52. Calliostoma miotorulosum nov. nom.; 1. NHMW 2016/0103/0175, height 6.2 mm, width 5.1 mm; 2. NHMW 2016/0103/0180
detail of protoconch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place. 3. NHMW 2016/0103/0177, height 8.1 mm,
width 6.1 mm, La Presselière, Sceaux-d’Anjou, Maine-et-Loire, NW France, Tortonian, upper Miocene.
130 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
becoming beaded on second whorl. Third whorl ad- and
abapical cords strengthen, beaded, with ve ner cords
in between. Fourth whorl adapical cord becomes bid,
six weaker nely beaded cords between ad- and abapical
cords. Last whorl rounded at periphery, with secondary
cords intercalated in all interspaces between primaries.
Base not sharply delimited, imperforate, bearing numer-
ous subequal smooth cords. Aperture rounded, outer lip
simple, rounded at periphery. Columella callus strongly
thickened and everted.
Discussion – Millet name Trochus tumidus is unfortu-
nately a junior homonym of T. tumidus Montagu, 1803
[= Gibbula tumida (Montagu, 1803)], we propose the re-
placement name Calliostoma miotumidum nov. nom.
We have offered a revised description of this species, as
Millet’s (1865, p. 583) text does not describe the spiral
sculpture adequately. The supercial suture is bounded
and accentuated above by a single beaded abapical cord
of the preceding whorl and below by a bid beaded adap-
ical cord of the subsequent whorl. It is this character most
importantly that separates it from Calliostoma milleti-
granum nov. nom., which is also sculptured by numerous
nely beaded cords, but the ones closest to the suture are
not strengthened. The base is also nely beaded in C. mil-
letigranum, with less numerous cords than seen in this
species, in which the crowded basal cords are smooth.
As commented by Brébion (1964), this species is similar
to one illustrated by Harmer (1923, p. 714, pl. 58, gs 1-3)
as Trochus (Calliostoma) bullatus (Philippi, 1844). We
have not seen this species, but it seems to be larger and
have fewer and slightly stronger beaded cords and in the
Sicilian specimen illustrated by Philippi (1844, pl. 18, g.
8) the cords bordering the suture are not strengthened.
Millet (1854, 1865) recorded this species from the As-
semblage I localities of Sceaux-d’Anjou and Thorigné,
to which Brébion (1964, p. 97) added Renauleau and St-
Clément-de-la-Place.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1865; Brébion, 1964).
Calliostoma planospirum (Millet, 1865)
Plate 54, g. 1
1854 Trochus Planospirus Millet, p. 156 (nomen nudum).
*1865 Trochus planospirus Millet, p. 582.
1964 Calliostoma planospira Millet, 1854 [sic] – Bré-
bion, p. 80, pl. 1, gs 26-28.
Type material – Syntypes: St-Clément-de-la-Place, Tho-
rigné and Sceaux-d’Anjou, Musée d’Angers, France ( de
Brébion, 1964, p. 81).
Material and dimensions – Maximum height 26.2 mm;
Plate 53. Calliostoma miotumidum nov. nom.; 1. NHMW 2016/0103/1504, height 14.5 mm, width 14.2 mm. La Presselière, Sceaux-
d’Anjou, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Plate 54. Calliostoma planospirum (Millet, 1865); 1. NHMW 2016/0103/1495, height 26.2 mm; width 23.3 mm. La Presselière,
Sceaux-d’Anjou, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 131
width 23.3 mm (incomplete). Sceaux-d’Anjou: NHMW
2016/0103/1495-1496 (2), RGM.1309666 (7 fragments),
LC (3), FVD (1).
Original description – ‘Trochus planospirus, Millet. Coq.
en cône régulier, aigu, de moyenne taille; composée de
11-12 tours de spire aplatis, et couverte de stries ou petits
lets circulaires. Hauteur: 30 millimètres; diamètre: 24
millimètres. Saint-Clément, Sc, Th. (Millet, 1865, p. 582)’.
Discussion – Calliostoma planospirum (Millet, 1865) is
the largest Calliostoma species in the Assemblage I fauna.
There is little to add to Millet’s description, except to say
that the whorls are narrowly swollen abapically, slightly
overhanging the suture. This species is closely similar
to the Pliocene to present-day Calliostoma zizyphinum
(Linnaeus, 1758), but differs in having a taller spire and
a more depressed base. The sculpture in C. zizyphinum
is somewhat variable, but most specimens have broader
cords than C. planospirum and a thickened abapical cord
is usually present (see Giannuzzi-Savelli et al., 1994, gs
158-168). In C. planospirum the cords are very ne and
equal in width over the entire whorl.
Brébion (1964, p. 81) described a similar form, based on
a couple of shells from Contigné, as var. saccoi (nomen
nudum), differing from the type in having coarser beaded
cords, in one specimen the cords of alternating strength.
We have not come across this form, but doubt they are
conspecic with C. planospirum.
Millet (1854, 1865) and Brébion (1964, p. 81) recorded
this species from the Assemblage I localities of St-Clé-
ment-de-la-Place, Sceaux-d’Anjou and Thorigné.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Calliostoma quaggaoides nov. sp.
Plate 55, gs 1-2
Type material – Holotype MNHN.F.A57682, height 3.3
mm, width 4.8 mm; paratype 1 MNHN.F.A57683, height
2.3 mm, width 4.0 mm; paratype 2 NHMW 2016/0103/
0048, height 2.6 mm, width 5.3 mm; paratype 3 NHMW
2016/0103/0049, height 2.6 mm, width 4.5 mm; paratype
4 NHMW 2016/0103/0122, height 2.4 mm, width 4.2
mm; St-Clément-de-la-Place. Paratype 5 RGM.1309702,
height 5.5 mm, width 6.7 mm; Sceaux-d’Anjou.
Other material – Maximum height 4.6 mm, width 5.6
mm. St-Clément-de-la-Place: NHMW 2016/0103/0050
(49). Sceaux-d’Anjou: NHMW 2016/0103/0051 (10),
RGM. 1309633 (4), RGM.1309683 (3), RGM.1309710 (2),
RGM. 1309723 (1), RGM.1348020 (2), LC (2). Renauleau:
NHMW 2016/0103/ 0118 (2), LC (1).
Etymology – Named after the extinct quagga zebra, refer-
ring to the zebra-like stripes. Calliostoma gender neuter.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Calliostoma species of small size, lenti-
form shell shape, with smooth whorls, except for single
suprasutural cord, angular periphery, narrow umbili-
cus bordered by cord and colour pattern of white radial
stripes.
Description – Shell small, depressed, lentiform. Proto-
conch paucispiral, of 1.2 whorls bearing honeycomb sur-
face sculpture (dp = 360 µm, dn = 160 µm). Junction with
teleoconch sharply delimited by abrupt end to protoconch
sculpture. Teleoconch of three low convex whorls, with
periphery at abapical suture, separated by impressed su-
ture. Sculpture restricted to a single suprasutural cord.
Last whorl depressed, angular at peripheral cord, base
attened, smooth, except for 3-4 weak cords adjacent to,
and strengthening towards periumbilical cord. Umbilicus
narrow, deep, smooth within. Aperture ovate, outer lip
simple, angled at periphery. Columella weakly thickened,
Plate 55. Calliostoma quaggaoides nov. sp.; 1. Holotype MNHN.F.A57682, height 3.3 mm, width 4.8 mm; 2. Paratype 4 NHMW
2016/0103/0122, detail of protoconch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France,
Tortonian, upper Miocene.
132 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
sloping abaxially. Colour pattern preserved in almost all
specimens of ne, radial white stripes.
Discussion – At rst sight this small lentiform species
resembles a species of Gibbula, however, the paucispiral
protoconch covered by honeycomb sculpture places it in
Calliostoma. Calliostoma gibbuliforme nov. sp. also has
a gibbuliform rather than trochiform shell shape, but is
less depressed, has weak spiral cords, absent in C. quag-
gaoides, and colour pattern of small white spots at the
periphery rather than axial stripes. We cannot nd any
other eastern Atlantic fossil or present-day faunas even
remotely similar to this shell to compare.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma pagodulum (Millet, 1865)
Plate 56, gs 1-4
1854 Trochus Pagodulus Millet, p. 157 (nomen nudum).
*1865 Trochus pagodulus Millet, p. 583.
1964 Calliostoma (Eucasta) pagodulum Millet, 1854
[sic] – Brébion, p. 101, pl. 2, gs 29, 30.
Type material – Holotype: Thorigné or Sceaux-d’Anjou,
Musée d’Angers, France ( de Brébion, 1964, p. 102).
Material and dimensions – Maximum height 13.5 mm,
width 10.0 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0043 (1), NHMW 2016/0103/0045 (1), NHMW
2016/ 0103/0044 (8), RGM.1309599 (2 fragments), LC (8),
FVD (5). Sceaux-d’Anjou: NHMW 2016/0103/0046 (19),
RGM. 1309536 (10), RGM.1309593 (1), RGM.1309671
(8), RGM.1347854 (5), RGM.1348005 (3), LC (8), FVD
(12). Renauleau: NHMW 2016/0103/1502-1503 (2),
NHMW 2016/0103/0047 (3), LC (15), FVD (9). Beugnon:
RGM.1309534 (11), RGM.1309535 (4), RGM.1309739 (2),
LC (2).
Discussion – Calliostoma pagodulum (Millet, 1865) is
probably the most striking calliostomiid found in As-
semblage I, characterised by having two strongly raised,
weakly undulating cords on each whorl, with the inter-
spaces deeply recessed. The cords are crossed by axial
lamellae that interrupt the cords giving them a beaded/sq-
uamous appearance. Further weaker cords are present on
the shoulder. The base bears a strong cord a short distance
from the crenulated peribasal cord, with about eight fur-
ther weaker beaded cords medially. The sculpture of Cal-
liostoma pagodulum (Millet, 1865) is unlike that of any
eastern Atlantic fossil or present-day Calliostoma species.
Millet (1854, 1865) described this species from the As-
semblage I localities of Sceaux-d’Anjou and Thorigné,
to which Brébion (1964, p. 102) added St-Clément-de-la-
Place.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Plate 56. Calliostoma pagodulum (Millet, 1865); 1. NHMW 2016/0103/1502, height 11.3 mm, width 8.9 mm; 2. NHMW
2016/0103/1503, height 13.5 mm, width 10.0 mm; Renauleau; 3. NHMW 2016/0103/0043 (subadult), height 7.7 mm, width 6.3
mm; 4. NHMW 2016/0103/0045, detail of protoconch (SEM image), Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-
Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 133
Calliostoma presselierense nov. sp.
Plate 57, gs 1-2
Type material – Holotype NHMW 2016/0103/0668,
height 7.1 mm, width 3.9 mm; paratype 1 NHMW
2016/0103/1461, height 3.3 mm (juvenile); paratype 2
MNHN.F.A57940, height 4.7 mm, width 2.5 mm.
Other material – LC (1).
Etymology – Named after the farm in which the locality
is situated, La Presselière, Sceaux-d’Anjou. Calliostoma
gender neuter.
Locus typicus – La Presselière, Sceaux-d’Anjou, Maine-
et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Calliostoma species of small size, with tall
slender trochiform shell shape, spiral sculpture of two
elevated narrow cords, with third weaker cord below
suture, and about 25 prosocline axial lamellae forming
pointed tubercles at intersections, attened imperforate
base delimited by smooth peribasal cord, bearing irregu-
lar cords without axial sculpture.
Description – Shell small, tall slender trochiform, with
conical spire. Protoconch paucispiral, composed of 1.3
whorls, bearing honeycomb pattern. Teleoconch of six
whorls. Suture linear, impressed. First teleoconch whorl
bearing two narrow elevated spiral cords, abapical one
slightly stronger forming periphery. Cords crossed by
about 25 strongly prosocline axial lamellae forming
pointed tubercles at intersections. On third whorl, third
weaker cord develops below adapical suture. Last whorl
with weak subsutural cord, two narrow subequal spinous
cords mid-whorl, strong smooth peribasal cord delimiting
base. Base attened, imperforate, bearing ve narrow spi-
ral cords of irregular strength. Aperture rounded, outer lip
simple, rounded at periphery, columella smooth. Columel-
lar callus narrow, thickened, parietal callus not developed.
Discussion – Although represented by a single adult
specimen and two juveniles with the protoconch pre-
served, this species is so distinctive it warrants descrip-
tion. Calliostoma presselierense nov. sp., C. lamellatum
nov. sp. and C. echinatum (Millet, 1865) form a species
group with similar sculpture composed of narrow elevat-
ed cords and axial lamellae forming spines or pointed
tubercles at the intersections. Calliostoma presselierense
is the tallest and most slender of the group, the spines
developed at the intersections are pointed tubercles rather
than long open spines as in C. echinatum or short spines
as in C. lamellatum. Moreover, the base in C. presselie-
rense is devoid of axial sculpture, whereas there is some
axial sculpture reticulating the basal cords in the other
two species. Calliostoma presselierense has so far only
been found at Sceaux-d’Anjou.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma spinosum nov. sp.
Plate 58, gs 1-6
1964 Calliostoma deshayesi Mayer, 1862 – Brébion, p.
76 (non Mayer, 1862).
Type material – Holotype MNHN.F.A57698, height
5.4 mm, width 4.5 mm; paratype 1 MNHN.F.A57699,
height 5.4 mm, width 4.3 mm; paratype 2 NHMW 2016/
0103/0209, height 6.0 mm, width 5.1 mm; paratype 3
NHMW 2016/0103/0210, height 6.8 mm, width 5.7 mm;
paratype 4 NHMW 2016/0103/0211, height 6.3 mm, width
4.5 mm; paratype 5 NHMW 2016/0103/0214, height 5.8
mm, width 5.4 mm; paratype 6 NHMW 2016/0103/0212
(juvenile); paratype 7 RGM.1309602, height 3.7 mm,
width 3.2 mm; St-Clément-de-la-Place. Paratype 8 RGM.
1309660, height 7.2 mm, width 5.8 mm; paratype 9 RGM.
1309661, height 6.9 mm, width 6.2 mm; Sceaux-d’Anjou.
Plate 57. Calliostoma presselierense nov. sp.; 1. Holot ype NHMW 2016/0103/0668, height 7.1 mm, width 3.9 mm; 2. Paratype
1 NHMW 2016/0103/1461, detail of protoconch (SEM image). La Presselière, Sceaux-d’Anjou, Maine-et-Loire, NW France,
Tortonian, upper Miocene.
134 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogast ropoda and Vetigastropoda
Other material – Maximum height 5.8 mm, width 4.7
mm. St-Clément-de-la-Place: NHMW 2016/0103/0213
(50+), RGM.1309560 (1 juvenile), RGM.1309673 (14),
RGM.1309746 (17), RGM.1309759 (18), LC (4), FVD (12).
Sceaux-d’Anjou: NHMW 2016/0103/0220 (50+), RGM.
1309662 (18), RGM.1309686 (17), RGM.1309693 (25), RGM.
1309708 (1), RGM.1309711 (41), RGM.1309764 (11), RGM.
1309776 (26), RGM.1348006 (41), LC (20), FVD (18).
Renauleau: NHMW 2016/0103/0215 (50+), LC (50), FVD
(34). Beug non: RGM.1309675 (20), RGM.1309744 (1).
Etymology – Latin ‘spinosus, -a, -um’, adjective, mean-
ing spiny; reecting the sharp tubercles developed at the
sculptural intersections. Calliostoma gender neuter.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Calliostoma species of small size, with de-
pressed, slightly scalate spire, sculptured by four primary
spiral cords bearing sharp tubercles, angular depressed
base bearing eight cords, and narrow umbilicus.
Description – Shell small, trochiform, with low, some-
Plate 58. Calliostoma spinosum nov. sp.; 1. Holotype MNHN.F.A57698, height 5.4 mm, width 4.5 mm; 2. Paratype 6 NHMW
2016/0103/0212, detail of protoconch (SEM image); 3. Paratype 2 NHMW 2016/0103/0209, height 6.0 mm, width 5.1 mm; 4.
Paratype 3 NHMW 2016/0103/0210, height 6.8 mm, width 5.7 mm; 5. Paratype 4 NHMW 2016/0103/0211, height 6.3 mm,
width 4.5 mm; 6. Paratype 5 NHMW 2016/0103/0214, height 5.8 mm, width 5.4 mm. Le Grand Chauvereau, St-Clément-de-la-
Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 135
what scalate spire. Protoconch paucispiral, of 1.1 whorls
bearing honeycomb surface sculpture (dp = 390 µm, dp =
160 µm). Junction with teleoconch sharply delimited by
end of protoconch sculpture. Teleoconch of four whorls.
Suture linear, impressed. First teleoconch whorl bearing
two elevated spiral cords of roughly equal strength and
about 18 axial ribs, slightly weaker than cords, forming re-
ticulate sculpture. On second whorl, a third cord develops
above the abapical suture; on third whorl a fourth again
just above abapical suture. On penultimate whorl adapical
cord placed just below suture, second cord forms shoulder,
third cord placed midway between shoulder and lower su-
ture, fourth cord placed just above suture. Axial sculpture
weakens abapically, but sharp pointed tubercles developed
at intersections, most strongly developed on adapical cord.
Last whorl bearing four primary spinous cords, second-
ary threads developed in interspaces between cords 1 and
2, and 2 and 3. Base sharply angled, attened, delimited
by bid, non-spinous peribasal cord, bearing about eight
irregular elevated cords beaded mid-base by prosocline
growth line, narrowly umbilicate. Aperture rounded, outer
lip simple, rounded at periphery, columella smooth. Colu-
mellar callus sharply delimited, parietal callus thin.
Discussion – As can be seen from the series illustrated,
we have interpreted this species as being highly variable.
The most distinctive character common to all these forms
is the presence of four primary spiral cords on the dor-
sum of the penultimate and last whorls. Having said this,
almost all the other characters are variable. The descrip-
tion offered above is based on what we consider the ‘typi-
cal’ form. Rather than include every variable factor in
the description we will describe below some of the most
important varieties.
Paratype 2 (Pl. 58, g. 3) has the primary spiral cords
strongly elevated, with denser, but less pointed tubercles,
on the last whorl the secondary cord between primaries 1
and 2 is almost as strong as the primaries, the base is less
attened and the cords on the base are strongly raised and
strongly beaded towards the umbilicus.
Paratype 3 (Pl. 58, g. 4) has a more regular conical spire,
no secondary spiral sculpture is developed and the base is
strongly attened, the basal cords are less strongly devel-
oped, but all of them are beaded.
Paratype 4 (Pl. 58, g. 5) has a more slender regular coni-
cal spire. On the last whorl the tubercles on the whorls are
closer-set than usual and only ve cords cover the base.
Paratype 5 (Pl. 58, g. 6) is thickened and gerontic, the
tubercles are close-set and lamellar and on the last whorl
the secondary cord between primaries 1 and 2 is almost
equal in strength to the primaries.
As well as these extreme morphs described above, in the
occasional shell the basal cords are reduced to ne threads,
and in some the tubercles seem less sharp, although they
may be abraded. Between these morphs and the typical
form lie numerous intermediate specimens. Despite this
wide range of variability, Calliostoma spinosum nov. sp.
is a distinctive species that can only be compared to C.
deshayesi (Mayer, 1862) from the middle Miocene Loire
Basin of France. They both have whorls bearing four pri-
mary spiral cords, but C. deshayesi is far higher spired,
the beads on the cords are not spinous and the base bears
fewer cords that are more nely or more weakly beaded.
Brébion (1964, p. 76) recorded this species from Assem-
blage I localities of Renauleau and St-Michel, to which
we add St-Clément-de-la-Place.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma umbellum (Millet, 1865)
Plate 59, g. 1
1854 Trochus Umbella Millet, p. 157 (nomen nudum).
*1865 Trochus Umbella Millet, p. 583.
1964 Calliostoma umbella Millet, 1854 [sic] – Brébion,
p. 95, pl. 2, gs 17, 18.
Plate 59. Calliostoma umbellum (Millet, 1865); 1. NHMW 2016/0103/0165, height 10.0 mm, width 11.0 mm. Le Grand Chauvereau,
St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
136 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwester n France, 1. Patellogastropoda and Vetigastropoda
Type material – Syntypes: Thorigné, Sceaux-d’Anjou and
Renauleau, Musée d’Angers, France (de Brébion, 1964,
p. 96).
Material and dimensions – Maximum height 10.5 mm,
width 12.2 mm (incomplete). St-Clément-de-la-Place:
NHMW 2016/0103/0165-0166 (2), LC (1). Sceaux-d’An-
jou: NHMW 2016/0103/0173 (4), RGM.1309588 (4), RGM.
1309659 (1), RGM.1309685 (3), RGM.1309705 (1), RGM.
1309737 (1), LC (4). Renauleau: NHMW 2016/ 0103/1484
(11), LC (8), FVD (6).
Discussion – Millet (1865) validated his earlier (1854)
name Trochus umbella with the following description:
‘Trochus umbella, Millet. Coq. assez grande, en cône
court, composée de 6-7 tours de spire portant des stries
à peine marquées, mais présentant a la base de chaque
tour un let qui accompagne la suture et formant carène
sur le dernier tour, ce dernier tour semble pouvoir s’en-
gaîner avec celui qui le touche et présente, d’ailleurs,
selon la variété, une surface plane ou légèrement creu-
sée vers le centre. Dessous bombé, lisse, étant marqué
seulement de quelques stries courtes vers le bord collu-
mellaire. Hauteur et diamètre: 15 millimètres. Th., Sc.,
Ren. (1865, p. 583)’ There is little to add to this descrip-
tion except to draw attention to the change in spire pro-
le with ontogeny from coeloconoid to cyrtoconoid. It
shows certain resemblance to the Pliocene to present-
day European C. zizyphinum (Linnaeus, 1758), but this
species is larger, with a taller regularly conical spire
and stronger spiral sculpture. It is also somewhat simi-
lar to C. planospirum (Millet, 1865), but this species is
higher spired with a at base and has a different shell
prole.
Brébion (1964, p. 96) recorded this species from Assem-
blage I localities (Sceaux d’Anjou, Thorigné and Renau-
leau), to which we add St-Clément-de-la-Place.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (Millet, 1854, 1865; Brébion, 1964).
Calliostoma verrucosum nov. sp.
Plate 60, gs 1-3
Type material – Holotype MNHN.F.A57694, height 4.4
mm, width 4.4 mm; paratype 1 NHMW 2016/0103/0184,
height 6.2 mm, width 5.3 mm; paratype 2 NHMW
2016/0103/0185, height 5.4 mm, width 5.0 mm; paratype
3 NHMW 2016/0103/0186, height 5.1 mm, width 4.8 mm;
paratype 4 NHMW 2016/0103/0187, height 3.4 mm, width
2.9 mm (incomplete juvenile); St-Clément-de-la-Place.
Paratype 5 RGM.1309719, height 5.9 mm, width 5.7 mm;
paratype 6 RGM.1309724, height 5.8 mm, width 5.4 mm;
Sceaux-d’Anjou.
Other material – St-Clément-de-la-Place: NHMW 2016/
0103/0188 (3), LC (1), FVD (2). Sceaux-d’Anjou: RGM.
1309699 (1), LC (1).
Etymology – Latin ‘verrucosus, -a, -um’ adjective, mean-
ing warty, rough, reecting the character of the sculpture.
Calliostoma gender neuter.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Plate 60. Calliostoma verrucosum nov. sp.; 1. Holotype MNHN.F.A57694, height 4.4 mm, width 4.4 mm; 2. Paratype 4 NHMW
2016/0103/0187, detail of protoconch (SEM image); 3. Paratype 1 NHMW 2016/0103/0184, height 6.2 mm, width 5.3 mm. Le
Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 137
Diagnosis – Calliostoma species of small size, with low
turbiniform shell shape, whorls strongly shouldered, con-
cave in prole, with coarsely beaded shoulder cord, de-
limiting narrow subsutural platform and peribasal cord,
nely beaded cords on rest of dorsum, and angular im-
perforate base covered in beaded cords.
Description – Shell small, depressed trochiform, with
scalate spire. Protoconch paucispiral, of 1.3 whorls bear-
ing honeycomb surface sculpture (dp = 560 µm, dn =
255 µm). Junction with teleoconch sharply delimited by
end of protoconch sculpture. Teleoconch of four whorls.
Suture linear, supercial. First teleoconch whorl bearing
ve spiral cords of roughly equal strength and prosocline
close-set axial lamellae. On second half of rst whorl
adapical cord strengthens to form shoulder. Abapically,
whorls shouldered, prole above and below shoulder
concave, with shallow, narrow subsutural ramp bearing
four nely beaded narrow cords, shoulder angled, abap-
ical cord strengthens, three minor nely beaded cords
between coarsely beaded shoulder and abapical cords.
Close set axial lamellae raised between cords on shoul-
der. Last whorl with shoulder placed high, peribasal cord
bid, strongly beaded, delimiting base. Base depressed,
imperforate, bearing eight beaded spiral cords. Aperture
ovate, outer lip simple, roundly angled at periphery, colu-
mella smooth. Columellar callus hardly developed, pari-
etal callus not developed.
Discussion – Calliostoma verrucosum nov. sp. is similar
in size and in having a bicarinate last whorl to C. bian-
gulatum nov. nom. (see above), but differs in having a
more depressed spire, the sculpture is strongly tubercu-
late, whereas in C. biangulatum the sculpture is lamel-
late, the shoulder and peribasal cords in C. verrucosum
are broader and more elevated and the basal cords are
thicker. Finally, the holes in the honeycomb protoconch
surface sculpture are bigger in C. biangulatum.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Calliostoma vibrayanum (Dollfus & Dautzenberg, 1886)
Plate 61, gs 1-2
*1886 Trochus (Ziziphinus) Vibrayanum Dollfus & Daut-
zenberg, p. 141.
1949 Calliostoma vibrayanum (Tournouër mss.) Doll-
fus & Dautzenberg, 1886 – Glibert, p. 32, pl. 2,
g. 4.
Material and dimensions – Maximum height 5.2 mm, St-
Clément-de-la-Place: LC (5). Sceaux-d’Anjou: NHMW
2016/0103/0673-0674 (2), NHMW 2016/0103/0675 (5),
RGM.1309658 (2), RGM.1309715 (6), RGM.1309767 (6),
RGM.1348011 (1), LC (2), FVD (2).
Discussion – Calliostoma vibrayanum (Dollfus & Dautz-
enberg, 1886) is characterised by its broad conical spire
with straight-sided to slightly coeloconoid prole com-
posed of at-sided whorls. The early teleoconch whorls
bear weakly beaded cords. Abapically only the suprasu-
tural cord strengthens; the cords above weaken, or in
most specimens disappear completely. The base is imper-
forate, attened, sharply delimited by a strong, rounded
peribasal cord, and almost smooth, except for a few weak
cords placed peripherally and medially. The aperture is
ovate and the columella oblique and relatively strongly
thickened abapically.
Other European fossil calliostomiids with a coeloconoid
spire prole and beaded cords that weaken or become ob-
solete on the later teleoconch whorls are C. audebardi (de
Basterot, 1825) from the lower Miocene Aquitaine Basin of
Plate 61. Calliostoma vibrayanum (Dollfus & Dautzenberg, 1886); 1. NHMW 2016/0103/0673, height 5.1 mm, width 4.4 mm; 2.
NHMW 2016/0103/0674, height 5.1 mm, width 4.6 mm, La Presselière, Sceaux-d’Anjou, Maine-et-Loire, NW France, Tortonian,
upper Miocene.
138 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
France, which differs in having more concave whorls, the
cords weaken but persist subobsoletely beaded on the last
whorl, and the cords both above and below the suture are
more strongly developed, C. benoisti Cossmann & Peyrot,
1917 from the middle Miocene Aquitaine Basin which has
a more pointed spire and a crenulated peribasal cord, and
C. xavieri (Dollfus, Cotter & Gomes, 1903) from the upper
Miocene Cacela Basin of southern Portugal, which differs
in having the spiral sculpture persisting stronger on the
last whorl and a more prominent peribasal cord. All three
have stronger basal sculpture than C. vibrayanum.
Distribution – Middle Miocene: Atlantic, Loire Basin,
France (Glibert, 1949). Upper Miocene: Atlantic (Torton-
ian), NW France (this paper).
Calliostoma cf. zizyphinum (Linnaeus, 1758)
Plate 62, gs 1-2
cf. *1758 Trochus zizyphinus Linnaeus, p. 759.
cf. 2014 Calliostoma zizyphinum (Linné, 1758) – Pouwer
& Wesselingh, p. 160, gs 31, 32.
cf. 2016a Calliostoma zizyphinum (Linnaeus, 1758) – Ceu-
le mans et al., p. 71, pl. 9, gs 3, 4.
Material and dimensions – Maximum height 18.7 mm,
width 15.5 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0180-0181 (5), NHMW 2016/0103/0182 (1), LC (1),
FVD (3). Sceaux-d’Anjou: RGM.1309698 (4). Renau-
leau: NHMW 2016/0103/1506 (4).
Discussion – The shells illustrated here as Calliostoma
cf. zizyphinum (Linnaeus, 1758) from St-Clément-de-la-
Place are relatively large and solid shelled compared to
other calliostomiids in the Assemblage I deposits. They
are very close in shape to the more strongly sculptured
forms of Calliostoma zizyphinum (Linnaeus, 1758), es-
pecially specimen NHMW 2016/0103/0181 (Pl. 62, g.
2), which also shows remnants of colour pattern on the
peripheral cord similar to that illustrated by Pouwer &
Wesselingh (2014, g. 32), which also has strong spiral
sculpture. However, we are unsure if the specimens from
St-Clément-de-la-Place are conspecic with the Pliocene
to present-day C. zizyphinum, as the apical angle is nar-
rower (especially specimen NHMW 2016/0103/0180, Pl.
62, g. 1). Moreover, there seem to be fewer cords on
the base. Ceulemans et al. (2016a) recorded the smoother
form of C. zizyphinum from the lower Pliocene Assem-
blage III. Calliostoma planospirum (Millet, 1865), also
from the Assemblage I fauna, is larger shelled, with ner
spiral sculpture that does not form a thickened abap-
ical cord and the base is atter, again with weaker spi-
ral sculpture. The Pliocene to present-day C. conulus
(Linnaeus, 1758) is also similar, but has beaded early
teleoconch whorls. Calliostoma lauguieri (Payraudeau,
1826) belongs to the same group of calliostomiid with
regular conical spires. It has a narrower apical angle than
C. zizyphinum and C. conulus, similar to the shells from
St-Clément-de-la-Place, but differ in also having beaded
early teleoconch whorls, although these disappear earlier
than they do in C. conulus.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Subfamily Thysanodontinae Marshall, 1988
Genus Thysanodonta Marshall, 1988
Type species (by original designation) – Thysanodonta
auklandiana Marshall, 1988, present-day, New Zealand.
1988 Thysanodonta Marshall, p. 217.
Thysanodonta chauvereauensis nov. sp.
Plate 63, gs 1-5
Type material – Holotype NHMW 2016/0103/0250,
Plate 62. Calliostoma cf. zizyphinum (Linnaeus, 1758); 1. NHMW 2016/0103/0180, height 18.7 mm, width 15.5 mm; 2. NHMW
2016/0103/0181, height 15.8 mm, width 13.0 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France,
Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 139
height 4.6 mm, width 3.7 mm; paratype 1 NHMW
2016/0103/0251, height 3.6 mm, width 3.2 mm; paratype
2 NHMW 2016/0103/0252, height 3.4 mm, width 2.8
mm; St-Clément-de-la-Place. Paratype 3 RGM.1309709,
height 3.1 mm, width 2.6 mm; Sceaux-d’Anjou.
Other material – St-Clément-de-la-Place: NHMW
2016/0103/0253 (7), LC (1), FVD (3). Sceaux-d’Anjou:
NHMW 2016/0103/1492-1493 (2), RGM.1309592 (1),
RGM. 1348007 (2).
Etymology – Named after the farm on which the locality
is situated, Le Grand Chauvereau. Thysanodonta gender
feminine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Thysanodonta species of small size, with tall
conical spire, in which rst teleoconch whorl bears axial
ribs, and single spinous shoulder cord that rapidly weak-
ens abapically, spiral sculpture weakens abapically, last
whorl with sharply beaded cords; 1-3 weaker adapically
and two more prominent abapically, and imperforate,
weakly sculptured, attened base.
Description – Shell small, of medium thickness, with
moderately high conical spire, attened base. Protoconch
paucispiral, composed of 1.5 whorls bearing honeycomb
sculpture. Teleoconch consisting of four whorls. Suture
linear, impressed. First teleoconch whorl sharply an-
gular, bearing weak prosocline axial ribs, single spiral
cord at shoulder forming spinous tubercles at intersec-
tions. Whorl prole straight to weakly concave above and
below shoulder. On second teleoconch whorl a weaker
beaded cord appears just below and just above sutures.
Abapically, shoulder cord rapidly weakens, two abapical
cords strengthen and further beaded cords appear above
in some specimens. Last whorl straight-sided, sharply an-
gled at base, bearing two strong abapical cords, the lower
Plate 63. Thysanodonta chauvereauensis nov. sp.; 1. Holotype NHMW 2016/0103/0250, height 4.6 mm, width 3.7 mm, 1c, detail
of protoconch (SEM image); 2. Paratype 1 NHMW 2016/0103/0251, height 3.6 mm, width 3.2 mm; 3. Paratype 2 NHMW
2016/0103/0252, height 3.4 mm, width 2.8 mm. Le Grand Chauvereau, St-Clément-de-la-Place. 4. NHMW 2016/0103/1492,
height 4.3 mm, width 3.5 mm; 5. NHMW 2016/0103/1493, height 4.1 mm, width 3.3 mm. La Presselière, Sceaux-d’Anjou,
Maine-et-Loire, NW France, Tortonian, upper Miocene.
140 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwester n France, 1. Patellogastropoda and Vetigastropoda
bordering the periphery and 1-3 weaker cords above. Base
attened, imperforate, bearing up to seven ne non-bead-
ed cords of irregular strength, obsolete mid-base in some
specimens. Aperture tangential; peristome discontinu-
ous, outer lip not thickened, angled at periphery, smooth
within. Columella straight, weakly thickened abapically.
Discussion – Marshall (1988) described a small group of
calliostomids that differed mainly in their extremely dis-
tinctive jaw and radular characters; the slender teeth far
longer than they are broad, indeed, the most slender of any
known gastropod. The subfamily Thysanodontinae was
erected for this group. Marshall (1988) suggested that they
were suctorially feeding, possibly on cnidarians. Marshall
(1988) also highlighted shell characters peculiar to the
subfamily; although they have a paucispiral protoconch
covered by honey-comb microsculpture like other callio-
stomids, the rst 1.5-2.5 teleoconch whorls are unicarinate
and the primary spiral cords all appear almost immedi-
ately at similar size. This is not seen in Calliostomatinae.
Thysanodontinae were originally described from South
Africa, Australia, New Zealand and New Caledonia, and
Marshall commented on their vicariant distribution.
The only two subsequent records including Thysanodont-
inae were from the same area (Marshall, 1995; Vilvens &
Maestrati, 2006).
This is the rst report of the subfamily in the European
faunas. It is possible that further species occur, but re-
main unrecognised or misidentied as Calliostoma spe-
cies. Three genera were described within the Thysano-
dontinae, of which the species from the upper Miocene
of France is best included in the genus Thysanodonta
Marshall, 1988. The spiral sculpture on the last whorl is a
little different from that of the antipodean Thysanodonta
species, which have three alternating primary and three
secondary cords intercalated, but we are reluctant to erect
a new genus for this species.
Thysanodonta chauvereauensis nov. sp. is a curious lit-
tle species. It is the smallest shelled calliostomid in the
Assemblage I deposits. A small patch of the honeycomb
surface sculpture is preserved just above the suture (Pl.
63, g. 1c). The most notable character in this species is
the drastic change in sculpture between the rst teleo-
conch whorl, which is unicarinate, and the later whorls,
which are straight-sided and predominated by two
strong abapical cords. The sculptural transition on the
second teleoconch whorl is rapid. Sculpture on the last
teleoconch whorl is rather variable; 1-3 beaded cords lie
above the upper of the two stronger abapical cords that
can be almost as strong as the abapical cords to rather
weak. Equally, the basal sculpture is variable, composed
of raised narrow cords, which can be subobsolete or
missing mid-base. Thysanodonta chauvereauensis dif-
fers from all its antipodean congeners (i.e. T. aucklandica
Marshall, 1988, T. wairua Marshall, 1988, T. boucheti
Marshall, 1988, T. eucosmia Marshall, 1995, T. festiva
Marshall, 1995, T. cassis Vilvens & Maestrati, 2006, T.
diadema Vilvens & Maestrati, 2006, T. pileum Vilvens &
Maestrati, 2006) in being smaller shelled and in having
the two abapical cords more strongly developed.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Family Liotiidae Gray, 1850
Genus Pareuchelus Boettger, 1907
Type species (by original designation) – Euchelus excel-
lens Boettger, 1907, Miocene, Romania.
1907 Pareuchelus Boettger, p. 186.
Pareuchelus lecointreae (Dollfus & Dautzenberg, 1899)
Plate 64, gs 1-3
1854 Delphinula Radiata Millet, p. 157 (nomen nu-
dum).
1865 Delphinula radiata Millet, p. 584 (non Kiener,
1838).
*1899 Turbo lecointreae Dollfus & Dautzenberg, p. 218,
pl. 9, gs 5, 6 (nom. nov. pro. Delphinula radiata
Millet 1865; non Kiener, 1838).
1938 Leptothyra? Lecointrae [sic] – Peyrot, p. 37.
1949 Turbo lecointreae Dollfus & Dautzenberg, 1899 –
Glibert, p. 77.
1964 Pareuchelus lecointreae Dollfus et Dautzenberg,
1899 – Brébion, p. 129, pl. 3, g. 17.
Material and dimensions – Maximum height 9.5 mm,
width 8.5 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0032 (1), 2016/0103/0033 (5), LC (3). Sceaux-d’An-
jou: NHMW 2016/0103/0035 (4), RGM.1309528 (3);
RGM.1309584 (1); RGM.1309617 (1), RGM.1347851 (3),
LC (6), FVD (6). Renauleau: NHMW 2016/0103/1423-
1424 (2), NHMW 2016/0103/0034 (14), LC (50), FVD
(15). Beugnon: RGM.1309529 (1); RGM.1309530 (1);
RGM.1309531 (2); RGM.1309740 (2), LC (1).
Discussion – Pareuchelus lecointreae (Dollfus & Dautz-
enberg, 1899) is similar to the older P. fossariopsis Coss-
mann & Peyrot, 1917, but this species from the Atlantic
lower Miocene Aquitanian of France has fewer spiral
cords (6 vs. 7), which are stronger. In the middle Mio-
cene Paratethys P. excellens Boettger, 1906 has only four
spiral cords that are even stronger, P. heres Boettger,
1906 seems to have the same number of cords as P. fos-
sariopsis. The holotype is incomplete (Zilch, 1934, pl. 4,
g. 55) and difcult to separate from the French lower
Miocene species, however, it differs from P. lecointreae
in the same way as P. fossariopsis; fewer and stronger
spiral cords.
In P. lecointreae the primary axials are often interrupt-
ed by the spirals and are not continuous axially (see Pl.
64, g. 1a). One interesting sculptural feature, which is
probably a generic character, is that there are anything
between two and six secondary axials between the pri-
mary axials that are only developed over the cords and
are short, not extending into the interspaces. The number
of these secondary axial riblets increases abapically.
Cainozoic Research, 17(2), pp. 75-166, December 2017 141
Millet (1854) recorded this species only from the Assem-
blage I locality of Sceaux-d’Anjou, but it is widespread
in the deposits; Brébion (1964, p. 130) added Thorigné,
St-Clément-de-la-Place and Beaulieu, to which we add
Renauleau.
Trochus sublimbatus d’Orbigny, 1852 from the Mediter-
ranean Pliocene has similar sculpture, but the similarity
is supercial. It belongs in the genus Danilia Brusina,
1865 (placed in Seguenzoidea: Chilodontidae). Danilia
species are solid-shelled, Pareuchelus Boettger, 1907
have thin shells. The aperture in Danilia bears a labial
varix, which is denticulate within, whereas the outer lip
in Pareuchelus is thin, lacking a labial varix, and smooth
within, so that specimens with a complete aperture are
scarce.
Distribution – Middle Miocene: Atlantic (Langhian)
Loire Basin, France (Peyrot, 1938). Upper Miocene: At-
lantic, Tortonian, NW France (Millet, 1854, 1865; Bré-
bion, 1964).
Pareuchelus dautzenbergi nov. sp.
Plate 65, g. 1
Type material – Holotype RGM.1309551, height 5.6
mm, width 5.1 mm; paratype 1 NHMW 2016/0103/1592,
height 6.0 mm, width 5.8 mm; paratype 2 NHMW
2016/0103/1593, height 4.6 mm, width 4.5 mm; paratype
3 NHMW 2016/0103/1594, height 3.5 mm, width 3.6 mm.
Other material – Maximum height 4.6 mm (incom-
plete), diameter 6.4 mm. St- Clément-de-la-Place:
NHMW 2016/0103/1595 (3 juveniles), LC (1), FVD (2).
Sceaux-d’Anjou: RGM.1309782 (1); RGM.1309783 (1),
RGM.1347997 (1), LC (1), FVD (2).
Etymology – Named after Philippe Dautzenberg (1849-
1935), Belgian malacologist, in recognition of his enor-
mous contribution to the eld. Pareuchelus gender mas-
culine.
Locus typicus – La Presselière, Sceaux-d’Anjou, Maine-
et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Pareuchelus species of small size, with a rel-
atively tall turibiform shell, highly sculptured teleoconch
whorls bearing narrow elevated cords of primary and
secondary strength and crowded axial lamellae, scabrous
of nely beading axials; spirals bearing twice as many
axials as interspaces between spirals, convex last whorl
bearing 8-10 strong cords.
Description – Shell small, relatively elevated turbiniform.
Protoconch paucispiral, composed of about 1.5 whorls,
with large nucleus. Teleoconch of four convex whorls
with periphery above abapical suture. Suture deeply im-
pressed, narrowly canaliculated. First teleoconch whorl
with three elevated spiral cords. On second whorl fourth
cord develops on subsutural platform. On penultimate
whorl further weaker cords above and below suture and
single secondary cord between 1st and 2n d primary cords.
Last whorl inated, convex bearing 8-10 strongly devel-
oped, narrow, elevated cords plus secondary cords in
some interspaces. The most adapical one or two primary
spirals turned upwards, lower three turned downwards.
Plate 64. Pareuchelus lecointreae (Dollfus & Dautzenberg, 1899); 1. NHMW 2016/0103/1423, height 7.2 mm, width 6.6 mm; 2.
NHMW 2016/0103/1424, height 7.2 mm, width 6.5 mm. Renauleau; 3. NHMW 2016/0103/0032, height 9.5 mm, width 8.5 mm.
Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
142 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Axial sculpture of continuous, crowded lamellae, scab-
rous over, or nely beading spiral cords. In interspaces
between ribs, cords bear a further spiral that disappears
in the interspaces between cords, so that cords bear twice
as many ribs as interspaces. Base not delimited, bearing
narrow umbilical chink. Aperture ovate, outer lip round-
ed, edge crenulated by spiral sculpture, smooth within.
Peristome complete. Columellar callus thickened abap-
ically bearing small ridge bordering siphonal canal. Pari-
etal callus thin, erect over umbilical chink.
Discussion – This species seems to have been overlooked
and lumped together with Pareuchelus lecointreae (Doll-
fus & Dautzenberg, 1899) by all workers, including our-
selves until late in the preparation of this monograph. It
differs from P. lecointreae in being smaller with a more
elevated spire, in having more numerous primary spiral
cords with secondaries developed in some of the inter-
spaces; P. lecointreae does not develop secondary spi-
rals. The difference in number of spirals is not just due
to a greater developement of secondaries, but is evident
from the rst teleoconch whorl, which only has two spi-
rals in P. lecointreae, three in Pareuchelus dautzenbergi
nov. sp. The two species also differ in their axial sculp-
ture, which consists of far more crowded, but less promi-
nent and less interrupted axials in P. dautzenbergi. This
is particularly evident on the subsutural platform. Fur-
thermore, the secondary axial sculpture (see above under
P. lecointreae) in P. dautzenbergi differs in only having
one secondary axial between the spirals that is longer and
extends a short distance into the interspaces. Pareuche-
lus fossariopsis Cossmann & Peyrot, 1917 from the
Atlantic lower Miocene Aquitanian of France has even
fewer spiral cords (6 vs. 8-10). Pareuchelus dautzenbergi
is found together with P. lecointreae at Sceaux-d’Anjou
and St-Clément-de-la-Place, although less frequent. At
Renauleau, where the genus is most abundant, we have
only found P. lecointreae.
Distribution – Upper Miocene: Atlantic, Tortonian, NW
France (this paper).
Family Skeneidae Clark, 1851
Genus Dikoleps Høisaeter, 1968
Type species (by original designation) – Margarita pu-
silla Jeffreys, 1847 [= Dikoleps nitens (Philippi, 1844)],
present-day, British Isles.
1968 Dikoleps Høisaeter, p. 47.
Dikoleps cutleriana (Clark, 1849)
Plate 66, g. 1
1849 Skenea Cutleriana Clark, p. 424.
2016a Dikoleps cutleriana (Clark, 1849) – Ceulemans et
al., p. 73, pl. 10, g. 1 (cum syn.).
Material and dimensions – Maximum diameter 1.3
mm, height 1.1 mm. St-Clément-de-la-Place: NHMW
2016/0103/0020 (1), NHMW 2016/0103/0021 (50+), RGM.
1309558 (11), RGM.1309645 (10), RGM.1347961 (2), LC
(5). Sceaux-d’Anjou: RGM.1309628 (10), LC (2). Re-
nauleau: NHMW 2016/0103/1400 (29), LC (5).
Discussion – We see no signicant difference between
the extant specimens and those from St-Clément-de-
la-Place. This is the stratigraphically oldest record for
Plate 65. Pareuchelus dautzenbergi nov. sp.; 1. Holotype RGM.1309551, height 5.6 mm, width 5.1 mm; La Presselière, Sceaux-
d’Anjou, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 143
Dikoleps cutleriana (Clark, 1849). The genus is known
in the Atlantic of France since the upper Oligocene;
Dikoleps sublaevis Lozouet, 1999 differs in having a
smooth surface, without any spiral sculpture and D.
aldradensis Lozouet, 1999 from the lower Miocene has
spiral sculpture restricted to the base.
For further discussion see Ceulemans et al. (2016a, p. 73).
Distribution – Upper Miocene: Atlantic, Tortonian, NW
France (Ceulemans et al., 2016a). Lower Pliocene: NW
France (Ceulemans et al., 2016a); Luchtbal Sand, Bel-
gium (Marquet & Landau, 2006). Present-day: Atlantic,
SW England (Fretter & Graham, 1977) southwards into
Mediterranean, Corsica (van Aartsen et al., 1984).
Dikoleps insulsa nov. sp.
Plate 67, g. 1
Type material – Holotype 2016/0103/1590, height 460 µm,
diameter 960 µm; paratype 1 NHMW 2016/0103/1591,
height 510 µm, diameter 1.0 mm.
Other material – Known from type series only.
Etymology – Latin ‘insulsus, -a, -um’, adjective meaning
boring, referring to the lack of shell characters. Dikoleps
gender femimine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Dikoleps of small size, with smooth pau-
cispiral protoconch, teleoconch of 1.5 whorls, devoid of
sculpture, except for occasional growth lines accentuated
towards aperture, convex, non-delimited base, broad,
smooth, deep umbilicus, columellar and parietal calluses
only slightly thickened.
Description – Shell minute, planorbid. Protoconch of one
smooth whorl, with large nucleus, (dp = 180 µm, dn = 120
µm). Junction with teleoconch marked by scar. Teleoconch
of 1.5 depressed, strongly convex whorls, smooth, except
for occasional growth lines becoming accentuated towards
aperture. Suture linear, impressed. Last whorl depressed,
strongly convex. Base not sharply delimited, convex, bear-
ing wide, smooth, deep umbilicus. Aperture ovate, proso-
cline in prole, outer lip simple, Peristome complete, with
only weakly tickened columellar and parietal callus.
Discussion – Although we are fairly certain this is a ske-
neiid, we struggle to assign it to a genus. We have placed
it provisionally in Dikoleps Høisaeter, 1968, which have
depressed to strongly depressed shells and can lack spiral
sculpture on the dorsum. However, its European congeners
reviewed by Rubio et al. (2004) all have umbilical ribs,
although in D. nitens (Philippi, 1844) they can be much re-
duced (see Rubio et al., 2004, p. 118, gs 3, 4). Dikoleps in-
sulsa nov. sp. further differs from all of its European con-
geners in having an even more depressed shell. Cirsonella
Angas, 1877 species are also smooth, but less depressed
and have an umbilical callus pad. Moel leriopsis Bush, 1897
species also have similarly planorbid shells, but the proto-
conch has ne spiral sculpture and the umbilicus is keeled.
Two relatively deep-water European species described by
Rubio & Rolán (2013) under Trenchia Knudsen, 1964; T.
biangulata and T. anselmoi differ in having an umbilical
keel. We note that although originally described in Skene-
idae, it is now transferred to Seguenzioidea [unassigned]
(Bouchet et al., 2013; WoRMS). Skenea bogii Chirli, 2004
from the Italian lower Pliocene also lacks spiral sculpture,
but is less depressed, with more numerous whorls separat-
Plate 66. Dikoleps cutleriana (Clark, 1849); 1. NHMW 2016/0103/0020, diameter 1.3 mm, height 1.1 mm; 1d, detail of protoconch
(SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Plate 67. Dikoleps insulsa nov. sp.; 1. Holotype NHMW
2016/0103/1590, height 460 µm, diameter 960 µm (SEM
image). Le Grand Chauvereau, St-Clément-de-la-Place,
Maine-et-Loire, NW France, Tortonian, upper Miocene.
144 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
ed by a deeper suture, and more accentuated axial growth
lines. Skenea pelagia Nofroni & Valenti, 1987 from the
present-day Mediterranean is similar in shape, especially
the character of the base and umbilicus, but differs in hav-
ing spiral sculpture on the teleoconch.
Distribution – Upper Miocene: Tortonian, NW France
(this paper).
Genus Leucorhynchia Crosse, 1867
Type species (by monotypy) – Leucorhynchia caledonica
Crosse, 1867, present-day, New Caledonia.
1867 Leucorhynchia Crosse, p. 319.
Leucorhynchia rotellaeformis (Grateloup, 1832)
Pl. 68, g. 1
*1832a Delphinula rotellaeformis Grateloup, p. 203.
1845 Delphinula rotellaeformis Grat. – Grateloup, pl. 12.
1854 Delphinula Plicatella Millet, p. 156.
1856 Delphinula rotellaeformis Grat. – Hörnes, p. 473,
pl. 46, g. 6.
1865 Delphinula plicatella Millet – Millet, p. 584.
1917 Leucorhynchia rotellaeformis (Grateloup) – Coss-
mann & Peyrot, p. 205, pl. 6, gs 42-44.
non 1954 Leucorhynchia rotellaeformis Grat. – Strausz, p.
8, pl. 9, g. 161 (= Leucorhynchia zboroviensis
Friedberg, 1928).
non 1966 Leucorhynchia rotellaeformis Grateloup (1832)
1840 – Strausz, p. 47, pl. 50, gs 10-15 (= Leuco-
rhynchia zboroviensis Friedberg, 1928).
1964 Leucorhynchia rotellaeformis Grateloup, 1832 –
Brébion, p. 125, pl. 3, g. 16.
1975 Leucorhynchia rotellaeformis (Grateloup, 1840
[sic]) – Bałuk, p. 52, pl. 5, g. 9.
Material and dimensions – Maximum diameter 3.3
mm, height 1.9 mm. St-Clément-de-la-Place: NHMW
2016/0103/0027 (1), 2016/0103/0028 (26), RGM. 1309641
(2), LC (10). Sceaux-d’Anjou: NHMW 2016/0103/0030
(22), RGM.1309679 (20), RGM.1309758 (9), RGM.1347959
(2), RGM.1348008 (2), LC (15). Renauleau: NHMW
2016/0103/0029 (50+), LC (50+), FVD (50+). Beugnon:
NHMW 2016/0103/0031 (8).
Discussion – This is undoubtedly the same species de-
scribed by Millet as Delphinula plicatella: ‘Plicatella,
Millet. – Sceaux. – Reneauleau. – Cette espèce a quelque
analogie avec les Delphinula callifera, Desh., et D. ca-
nalifera, Lamk.; mais les petits plis rayonnants qui par-
tent de l’ombilic dans notre espèce, sont sufsants pour
la faire distinguer (Millet, 1854, p. 156)’. The specimen
gured (Pl. 68, g. 1) illustrated the spiral sculpture that
was not commented on by Cossmann & Peyrot (1917).
Leucorhynchia miorotelloides Sacco, 1896 was described
from the lower Miocene Proto-Mediterranean of Italy. It is
difcult to compare with the small gures given by Sacco
(1896a, pl. 4, g. 67) and the syntype illustrated by Fer-
rero Mortara et al. (1894, pl. 50, g 7), which is poorly
preserved. Sacco’s description (1896a, p. 53) does not help
either, as he admits this may just be a form of L. rotellae-
formis. We refrain from synonymising the two, as the colu-
mellar callus seems to be even more strongly developed
in the Italian shell, but more material would be required
to be certain. Leucorhynchia zboroviensis Friedberg, 1928
from the middle Miocene Paratethys of Poland, is quite
different, with an angular periphery and a wider umbilicus
bounded by a raised periumbilical cord. Leucorhynchia iri-
color Boettger, 1907 described from the middle Miocene
of Kostej, is probably a synonym of L. rotellaeformis as
suggested by Bałuk (1975, p. 53), but we have not seen
specimens from the type locality. Today the genus in the
Atlantic has a more southern distribution with three spe-
cies described from West Africa (Rolán & Rubio, 2012).
Millet (1854, 1865) recorded this species from the As-
semblage I localities of Sceaux-d’Anjou and Renauleau,
to which Brébion (1964, p. 126) added Beaulieu and we
add St-Clément-de-la-Place.
Plate 68. Leucorhynchia rotellaeformis (Grateloup, 1832); 1. NHMW 2016/0103/0027, diameter 3.3 mm, height 1.9 mm; 1d, detail
of protoconch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper
Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 145
Distribution – Lower Miocene: Atlantic (Burdigalian),
Aquitaine Basin, France (Cossmann & Peyrot, 1917).
Middle Miocene: Atlantic, Aquitaine Basin, France
(Cossmann & Peyrot, 1917); Paratethys, Austria (Hörnes,
1856), Poland (Bałuk, 1975). Upper Miocene: Atlantic
(Tortonian), NW France (Millet, 1854; Brébion, 1964).
Genus Lodderena Iredale, 1924
Type species (by original designation) – Liotia minima
Tennison-Woods, 1878, present day, South Australia.
1924 Lodderena Iredale, p. 182, 233.
1958 Cyclostremiscus (Pachystremiscus) Olsson &
McGinty, p. 52. Type species (by original desig-
nation): Pachystremiscus pulchellus Olsson &
McGinty, 1958, present-day, Caribbean.
Lodderena redferni nov. sp.
Plate 69, gs 1-2
Type material – Holotype NHMW 2016/0103/1462, di-
ameter 1.2 mm, height 645 µm; paratype 1 NHMW
2016/0103/1463, diameter 1.3 mm, height 625 µm, St-Clé-
ment-de-la-Place. Paratype 2 NHMW 2016/0103/1611,
diameter 1.3 mm, height 650 µm, Renauleau.
Other material – Known only from type series.
Etymology – Named after Colin Redfern of Boca Raton,
Florida (USA) in recognition of his work on the genus.
Lodderena gender feminine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Lodderena species of minute size, with quad-
ricarinate last whorl, carinae raised and wavey, lacking
mid-basal cord, nely rugose surface, rugae strengthen-
ing towards aperture, and moderately wide, deep umbili-
cus delimited by further wavey cord.
Description – Shell minute, almost planispiral. Proto-
conch paucispiral, composed of just over one whorl, with
deformed nucleus (dp = 170 µm, dn = 65 µm). Teleoconch
of 1.5 quadricarinate whorls, separated by deeply canal-
iculated suture. Carinae formed by raised wavey cords;
adapical carina placed mid-dorsum, abapical carina de-
limiting base, with two further slightly closer-spaced
carinae placed at periphery either side of mid-line, fth
weaker carina delimiting moderately wide, deep um-
bilicus. Mid-basal cord absent; spiral sculpture absent.
Whorl prole between carinae concave. Surface nely
rugose; rugae coarser on base adjacent to aperture. Aper-
ture round, peristome complete, outer lip simple.
Discussion – Lodderena species have shells character-
ised by being planispiral or almost planispiral, spirally
and/or axially striated, often with one or more prominent
peripheral keels, an almost round aperture and a pau-
cispiral protoconch with a deformed nucleus (Moolen-
beek, 1996). Warén (1992, p. 155) placed the present-day
Plate 69. Lodderena redferni nov. sp.; 1. Holotype NHMW 2016/0103/1462, diameter 1.2 mm, height 645 µm; 1d, detail of proto-
conch (SEM image); 2. Paratype 1 NHMW 2016/0103/1463, diameter 1.3 mm, height 625 µm (SEM image). Le Grand Chau-
vereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
146 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwester n France, 1. Patellogastropoda and Vetigastropoda
Mediterranean Cyclostrema catenoides in the genus Lod-
derena, but it was transferred to Skenea Fleming, 1825 by
WoRMS (Gofas, 2015). The new species from France is
undoubtedly a Lodderena species and the rst conrmed
fossil record for the genus in Europe. It is characterised
by its quadricarinate last whorl, the carinae formed by
wavey lines. Lodderena redferni nov. sp. is uncommon
and so far only found at St-Clément-de-la-Place. We note
that the outer lip is described at simple, non-thickened.
This is unusual for the genus and might indicate that the
two specimens at hand are not fully mature.
The French Miocene species is most similar to some of
the present-day Caribbean species, such as L. ornata (Ols-
son & McGinty, 1958), also recorded in the Cape Verde
Archipelago by Rolán (2005), which also has wavey cari-
nae, but it differs in having two further cords, one placed
between the apex and the mid-dorsal cord and a second
placed mid-base, and the surface is covered by ne spi-
ral cords. Another present-day Caribbean species; Lod-
derena pulchella (Olsson & McGinty, 1958), also has a
quadricarinate periphery to the last whorl, plus a raised
central basal cord, but the cords are linear and not wavey,
the axial sculpture is much stronger than in L. redferni and
the outer lip is thickened by varix. Lodderena janetmayae
Rubio, Rolán & Redfern, 1998 from the Bahamas is simi-
lar in number of carinae, which are also wavey, but differs
in having the abapical cord placed mid-base and having
axial ribs between the two peripheral cords. Lodderena
bunnelli Redfern & Rolán, 2005 from the Bahamas is
quite different with a honeycomb pattern at the periphery.
Distribution – Upper Miocene: Tortonian, NW France
(this paper).
Genus Parviturbo Pilsbry & McGinty, 1945
Type species (by original designation) – Parviturbo reh-
deri Pilsbry & McGinty, 1945, present-day, Florida, USA.
Parviturbo rubioi nov. sp.
Plate 70, gs 1-2
Type material – Holotype NHMW 2016/0103/0026, di-
ameter 940 µm, height 840 µm; paratype 1 NHMW
2016/0103/1469, diameter 1.1 mm, height 980 µm; para-
type 2 RGM.1309642, diameter 930 µm, height 830 µm.
Other material – Maximum diameter 1.1 mm, height 980
µm. St-Clément-de-la-Place: NHMW 2016/0103/0712
(1).
Etymology – Named after Federico Rubio of Valencia,
Spain in recognition of his excellent review of the genus.
Parviturbo gender masculine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis –Parviturbo species of small size, with rela-
tively elevated shell, moderately dense regularly reticu-
lated surface pattern consisting of three spiral cords on
rst whorl, ve on last whorl and 18 axial ribs on rst
whorl, 21 on last, in which axial and spiral elements are
of equal strength, and strong periumbilical cord.
Plate 70. Parviturbo rubioi nov. sp.; 1. Holotype NHMW 2016/0103/0026, diameter 940 µm, height 840 µm (SEM image); 2. Para-
type 1 NHMW 2016/0103/1469, diameter 1.1 mm, height 980 µm (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place,
Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 147
Description – Shell minute, turbiniform. Protoconch
of just over one smooth whorl, with large nucleus (dp =
250 µm, dn = 95 µm). Junction with teleoconch sharp,
marked by beginning of spiral sculpture. Teleoconch of
1.8 convex whorls, separated by impressed suture. Three
spiral cords on spire whorl, ve on last whorl, about one-
quarter width of their interspaces. Axial sculpture of 18
prosocline ribs on rst whorl, 21 on last whorl, equal in
width to spiral sculpture, forming evenly reticulated sur-
face pattern. Interspaces slightly concave, bearing irreg-
ular micropustules. Last whorl convex, bearing ve sub-
equal spiral cords, plus stronger peribasal and strongest
periumbilical cord. Base deeply umbilicate, bearing con-
spicuous growth lines within. Aperture rounded, outer
lip simple, columella thickened, slightly reected at base.
Discussion – Parviturbo rubioi nov. sp., with its reticu-
lated surface sculpture in which neither the spiral nor ax-
ial elements are strongly overwhelming, is typical of the
Eastern Atlantic species group as illustrated by Rubio et
al. (2015, p. 171). It is most similar to the extant P. rolani
Engl, 2001 from the Canary Islands, but this species has
only two spiral cords on the spire whorl as opposed to
three in the French shell and the axial ribs are thinner and
more numerous than in P. rubioi. Other European species
with regularly reticulated surface sculpture are P. elegan-
tulus (Philippi, 1844) from the Plio-Pleistocene to present
day Mediterranean, which has ner reticulated sculpture
with four spiral cords on the spire whorl; P. alboranensis
Peñas & Rolán in Peñas et al. 2006, which has the same
number of cords on the spire whorl, but the cords mid-
whorl on the last whorl are zig-zag, leading to the inter-
spaces being hexagonal honeycomb shaped; P. ergasticus
Rubio, Rolán & Gofas in Rubio et al., 2015 from the Bay
of Biscay, which has much ner reticulated sculpture with
more numerous ribs and cords. Several present-day Euro-
pean species such as P. fenestratus (Chaster, 1896) and P.
azoricus Rubio, Rolán & Segers in Rubio et al., 2015 are
separated by their predominantly spiral sculpture.
Distribution – Upper Miocene (Tortonian): Atlantic, NW
France (this paper).
Genus Pseudorbis Monterosato, 1884
Type species (by monotypy) – Fossarus granulum Bru-
gnone, 1873, present day, Mediterranean.
1884 Pseudorbis Monterosato, p. 109.
Pseudorbis beugnonensis nov. sp.
Plate 71, g. 1
Type material – Holotype MNHN.F.A57703, height 720
µm, width 770 µm; paratype 1 NHMW 2016/0103/0273,
height 710 µm, width 750 µm.
Other material – Beugnon: NHMW 2016/0103/0274 (20).
Etymology – Named after the type locality of Beugnon.
Pseudorbis gender masculine.
Locus typicus – Beugnon, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis –Pseudorbis species of small size, with regu-
larly globose shape, solid shelled, bearing sculpture of
seven broad, strap-like cords on last whorl, and circular
non-dilated aperture bordered by thick outer lip.
Description –Shell minute, solid, globose turbiniform.
Protoconch paucispiral, of 1.5 smooth whorls, with a
large rounded nucleus (dp = 260 µm, dp = 85 µm, dp1
= 170 µm). Junction with teleoconch not sharply delim-
ited. Teleoconch of 1.2 whorl separated by broadly canal-
iculated suture. Sculpture of broad, attened, strap-like
cords of equal strength, 2-3 times the width of their inter-
spaces, three on rst half teleoconch whorl. Last whorl
globose, regularly convex, bearing seven spiral cords.
Base continuous with dorsum, imperforate. Aperture cir-
cular, peristome complete, narrow. Outer lip thick, regu-
larly rounded, not dilated, anal and siphonal canals not
developed. Columella thickened, narrow, smooth.
Plate 71. Pseudorbis beugnonensis nov. sp.; 1. Holotype MNHN.F.A57703, height 720 µm, width 770 µm; 1d, detail of protoconch
(SEM image). Beugnon, Maine-et-Loire, NW France, Tortonian, upper Miocene.
148 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Discussion – Pseudorbis beugnonensis nov. sp. is easily
separated from Pseudorbis granulum (Brugnone, 1873),
with which it co-occurs, by its sculpture of broad, strap-
like cords as opposed to the rather sharp cords seen in
P. granulum. It is in much more similar to the strati-
graphically slightly older middle Miocene P. falunica
Lozouet, 1999 from the Loire Basin of France, which is
also sculptured by broad cords, but in that species they
are more numerous; ve on the spire whorl, and 14 on the
last whorl, including the basal cords. In P. falunica the
apertural portion on the last whorl is dilated, whereas in
P. beugnonensis it is not. Pseudorbis carinifera Lozouet
1999 from the Atlantic upper Oligocene of France differs
from P. beugnonensis in the same characters as P. granu-
lum (see above). Pseudorbis jameonsis Rubio & Babío,
1990 from the Canary Islands differs in having eight pri-
mary spiral cords mid-whorl, which are much narrower
than in P. beugnonensis.
Distribution – Upper Miocene: Atlantic (Tortonian), NW
France (this paper).
Pseudorbis granulum (Brugnone, 1873)
Plate 72, g. 1
*1873 Fossarus granulum Brugnone, p. 13.
1990 Pseudorbis granulum (Brugnone, 1873) – Rubio
& Babío, p. 204, gs 1:1-5.
1992 Pseudorbis granulum (Brugnone, 1873) – Warén,
p. 160, g. 40E.
1994 Pseudorbis granulum (Brugnone, 1873) – Gian-
nuzzi-Savelli et al., gs 365, 366.
2015 Pseudorbis granulum (Brugnone, 1873) – Rubio et
al., p. 244, gs 45, 46A-G, 47A-E, 48A-H, 49A-F.
Material and dimensions – Maximum diameter 1.4
mm, height 1.3 mm. St- Clément-de-la-Place: NHMW
2016/0103/0017 (1), NHMW 2016/0103/0018 (50+),
RGM. 1309634 (50+), LC (8). Sceaux-d’Anjou: RGM.
1309586 (2), RGM.1347974 (1). Renauleau: NHMW
2016/0103/1316 (15), LC (2). Beugnon: NHMW
2016/0103/0019 (14), LC (1).
Discussion – Pseudorbis granulum (Brugnone, 1873) is
a very characteristic species, with its prominent spiral
cords forming elevated carinae, six on the last whorl, with
a further three on the base. The protoconch is paucispiral,
consisting of just 1.25 whorls, with a large nucleus. The
crest seen on the last quarter protoconch whorl (Pl. 72, g.
1d) is also seen in the present-day specimen illustrated by
Rubio & Babío (1990, p. 206, g. 1:5). The only small dif-
ference between the French fossil shell and that illustrated
by Rubio & Babío (1990) is that it lacks the very ne axial
folds seen adjacent to the crest. The authors describe these
as ribs, although in our opinion they are not true ribs. We
consider this difference insignicant. Pseudorbis jameon-
sis Rubio & Babío, 1990 from the Canary Islands differs
in having eight primary spiral cords mid-whorl.
Lozouet (1999, p. 12) described P. carinifera from the
Atlantic upper Oligocene of France. This species is al-
most identical to P. granulum, with which it was not
compared in the original description. Rubio et al. (2015)
stated that ‘the number of spiral cords of P. carinifera
is intermediate between P. granulum and P. jameoensis
(2015, p. 250). This is not correct. The number of cords
is the same: six primaries on the last whorl, three weaker
ones on the base. Judging from the single shell illustrat-
ed from Saint-Paul-lès-Dax, the cords on the Oligocene
shell are lower and the coiling seems regular, whereas in
P. granulum the spire is slightly tilted in respect to the
last whorl, so that the suture is more oblique. The cords in
this species group seem to increase in strength over time.
Lozouet (1999) noted that in the Atlantic southwestern
France lower Oligocene specimen from Gaas were weak-
er sculptured than the upper Oligocene shells and that the
lower Miocene ones were similar in sculpture to the up-
per Oligocene ones; these are all considered P. carinifera
(Lozouet, 1999). We note that by the upper Miocene Tor-
tonian the sculpture is the same as that seen in the extant
Pseudorbis granulum. As far as we are aware, this is the
rst fossil record for the species.
Distribution – Upper Miocene (Tortonian): Atlantic, NW
France (this paper). Present-day: Atlantic, Canary Islands
(Rubio & Babío, 1990) to Senegal (Rubio et al, 2015),
Mediterranean (Warén, 1992).
Plate 72: Pseudorbis granulum (Brugnone, 1873); 1. NHMW 2016/0103/0017, diameter 1.4 mm, height 1.3 mm; 1d, detail of proto-
conch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 149
Genus Skenea Fleming, 1825
Type species (by subsequent designation, Gray, 1847) –
Helix serpuloides Montagu, 1808, present-day, British
Isles.
1825 Skenea Fleming, p. 246.
1827 Delphinoidea Brown, p. 32. Type species (by sub-
sequent designation, Gray, 1847b): Helix unispi-
ralis Montagu, 1803, present-day, British Isles.
Skenea dautzenbergi (Glibert, 1949)
Plate 73, g. 1
1949 Circulopsis dautzenbergi Glibert, p. 71, pl. 5, g. 1.
Material and dimensions – Maximum diameter 1.6 mm,
height 1.1 mm. St-Clément-de-la-Place: NHMW 2016/
0103/0022 (1), 2016/0103/0023 (50+), RGM.1309553 (15),
RGM.1309561 (6), RGM.1309624 (50+), LC (10). Sceaux-
d’Anjou: RGM.1347975 (2), Renauleau: NHMW 2016/
0103/0024 (42), LC (10). Beugnon: NHMW 2016/ 0103/
0025 (16).
Revised description – Shell small, planorbid. Protoconch
of one smooth whorl, with large nucleus (dp= 190 µm,
dn = 110 µm). Junction with teleoconch marked by ne
growth lines and appearance of single ne cord just
above suture. Teleoconch of 2.2 depressed, strongly con-
vex whorls, separated by impressed suture. On second
half of rst teleoconch whorl further spiral cords appear
on abapical half of whorl; adapical half remains smooth,
except for ne spiral line at shoulder. Last whorl de-
pressed, strongly convex, bearing ne spiral sculpture
below shoulder and on base. Base bearing wide umbili-
cus bordered by peri-umbilical cord, three strong cords
run within umbilicus. Aperture ovate, prosocline in pro-
le, outer lip simple. Peristome complete, attened.
Discussion – This species is supercially similar to the
lower Pliocene to present-day European Moelleriopsis
messanensis (Seguenza, 1876), which occurs in the lower
Pliocene Assemblage III localities of NW France. Indeed,
in Ceulemans et al. (2016a, p. 74) the Tortonian Assem-
blage I localities of northwestern France were included
in the distribution of that species. However, close inspec-
tion of the specimens under SEM imaging shows them
not to be a Moelleriopsis species, but a Skenea species.
The protoconch has no spiral sculpture and the peristome
is attened. In our opinion this is the same species as
described by Glibert (1949) under the name Circulopsis
dautzenbergi. Although the original description does not
mention any spiral sculpture on the teleoconch, this is
only visible under SEM. We therefore consider it useful
to include a revised description.
Skenea dautzenbergi (Glibert, 1949) is characterised by
its ne spiral sculpture on the abapical half of the last 1.5
teleoconch whorls and the few, but strong, cords running
in the umbilicus. There is a suggestion of some micro-
pustules on the second half protoconch whorl adjacent to
the suture, but it is too abraded to be certain. The French
fossil species differs from the type species S. serpuloides
(Montagu, 1808), from the present-day eastern Atlantic
and Mediterranean S. pelagia Nofroni & Valenti, 1987
and S. giemellorum Romani, Bogi & Bartolini, 2015 in
that these three species have stronger spiral sculpture,
which covers the entire teleoconch surface and the um-
bilical cords are not strongly developed. We note that
whilst the generic description of Skenea given by Warén
(1992, p. 155) describes the protoconch as perfectly
smooth, in S. giemellorum it is sculptured by a couple
of spiral threads. Skenea dautzenbergi is most similar to
the French Atlantic upper Oligocene S. indubitabilis Lo-
zouet, 1999, which also has stronger periumbilical keels
than the present-day European species, but in that species
the teleoconch is smooth, devoid of spiral sculpture and
four cords run within the umbilicus as opposed to three
(periumbilical keel not included) in S. dautzenbergi. Ske-
nea multicostellata Lozouet, 1999, also from the French
Atlantic upper Oligocene, like the present-day species,
has spiral sculpture covering the entire whorl surface.
Distribution – Middle Miocene: Atlantic, Loire Basin,
France (Glibert, 1949). Upper Miocene: Tortonian, NW
France (this paper).
Plate 73. Skenea dautzenbergi (Glibert, 1949); 1. NHMW 2016/0103/0022, diameter 1.6 mm, height 1.1 mm; 1d, detail of protoconch
(SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
150 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwester n France, 1. Patellogastropoda and Vetigastropoda
Skenea minuticostata nov. sp.
Plate 74, g. 1
Type material – Holotype NHMW 2016/0103/0056, di-
ameter 850µm; paratype 1 NHMW 2016/0103/0057,
diameter 750µm; paratype 2 NHMW 2016/0103/0711,
diameter 900µm; paratype 3 RGM.1309646; paratype 4
RGM.1309647.
Other material – St-Clément-de-la-Place: NHMW 2016/
0103/1395 (1), RGM.1309648 (1 fragment).
Etymology – Latin compound name from ‘minutus, -a,
-um’, adjective, meaning small or tiny and ‘costatus, -a,
-um’, adjective, meaning ribbed; reecting the sculpture
of very ne, close-set ribs. Skenea gender feminine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Skenea species of small size, with paucispi-
ral protoconch bearing single cord mid-whorl, teleoconch
last whorl angular at shoulder, periphery and base, an-
gulations marked by broad cord interrupted by irregular,
close-set axial ripple-like ribs that cover entire whorl and
continue into deep umbilicus.
Description – Shell small, planorbid. Protoconch of one
whorl, with large nucleus, bearing a single spiral cord mid-
whorl (dp= 210 µm, dn = 120 µm). Junction with teleo-
conch marked by beginning of axial sculpture. Teleo conch
of two depressed whorls, separated by deeply impressed
suture. Sculpture on rst whorl of two poorly delimited
spiral cords placed below suture and at shoulder. Irreg-
ular ripple-like axial ribs radiate from cords, but leav-
ing mid-portion of whorl smooth. Abapically, axial ribs
strengthen, broaden and become elevated, axial ribs coa-
lesce mid-whorl, so that on second teleoconch whorl axial
ribs extend across dorsum, elevated, roughly equal in
width to their interspaces. Last whorl angled at shoulder,
periphery and base, which are delimited by broad cords,
interrupted by the irregular, elevated axial sculpture that
extends onto base and into wide umbilicus. Aperture
ovate, prosocline in prole, outer lip simple. Peristome
complete.
Discussion – It is difcult to nd any species that can be
usefully compared in the European fossil and present-
day faunas. The axial sculpture of ripple-like axial ribs is
unique for the genus. It is most similar to some Lodderena
Iredale, 1924 species such as the present-day species L.
ornata (Olsson & McGinty, 1958), L. omanensis Moolen-
beek, 1996 and L. tanae Moolenbeek, 1996 from Oman
and L. formosa Powell, 1930 and L. nana Powell, 1930
from New Zealand have predominantly spiral sculpture,
whereas in Skenea minuticostata nov. sp. the sculpture
is predominantly axial. Lodderena pachy nepion (Pilsbry
& Olsson, 1945) from the Pacic coast of Colombia does
have axial sculpture, but this is much less dense than in
S. minuticostata. The most similar species we could nd
is L. pulchella Olsson & McGinty, 1958 from the present
day Caribbean, which has very similar sculpture on the
dorsum, but the axial ribs fade over the base and there is
an extra inner cord on the base, absent in L.minuticostata.
An undescribed extant Lodderena species from Mauri-
tius illustrated on the website of the National Museum of
Wales (http://www.museumwales.ac.uk/curatorial/bio-
syb/mollusca/research/interpretation/accessed 17.052016)
has very similar sculpture to the French Miocene species.
Distribution – Upper Miocene: Tortonian, NW France
(this paper).
Plate 74. Skenea minuticostata nov. sp.; 1. Holotype NHMW 2016/0103/0056, diameter 850µm; 1d, detail of protoconch (SEM im-
age). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 151
Skenea wareni nov. sp.
Plate 75, g. 1
Type material – Holotype MNHN.F.A57684, diameter 1.4
mm; height 950 µm; paratype 1 NHMW 2016/0103/0052;
paratype 2 NHMW 2016/0103/0053; paratype 3 NHMW
2016/0103/0054; paratype 4 RGM.1309636; paratype 4
RGM.1309637; paratype 5 RGM.1309638.
Other material – St-Clément-de-la-Place: NHMW 2016/
0103/0055 (11), RGM.1309639 (18 fragments), LC (3). Re-
nauleau: NHMW 2016/0103/1399 (7), LC (2).
Etymology – Named after Anders Warén of the Zoolo-
giska Institutionen, Götenborg (Sweden), in recognition
to his enormous contributions to malacology, and espe-
cially skeneimorph gastropods. Skenea gender femimine.
Locus typicus – Le Grand Chauvereau, St-Clément-de-
la-Place, Maine-et-Loire, NW France.
Stratum typicum – Tortonian, upper Miocene.
Diagnosis – Skenea species of small size, with paucispi-
ral protoconch bearing spiral threads, teleoconch bear-
ing strong spiral cords crossed by lamellar growth lines,
giving surface beaded or scabrous appearance, sculpture
continuing onto base, broad umbilicus.
Description – Shell small, planorbid. Protoconch of one
whorl, with large nucleus, bearing irregular spiral threads
(dp = 240 µm, dn = 140 µm). Junction with teleoconch
marked by the beginning of three strong spiral cords
placed adjacent to each suture and at shoulder, crossed by
close-set lamellar growth lines, which cut cords, giving
them nely beaded or scabrous appearance. Teleoconch
of 2.6 depressed, strongly convex whorls, separated by
deeply impressed suture. After 0.75 teleoconch whorls
secondary cords appear in interspaces, rapidly gaining in
strength, with further tertiary cords developing in inter-
spaces on last half whorl. Entire teleoconch surface cov-
ered in raised lamellar growth lines, giving interspaces
between beaded cords scabrous appearance. Last whorl
depressed, strongly convex, bearing about 12 beaded
spiral cords of subequal strength; alternate strength
mid-whorl. Base bearing wide umbilicus, sculptured by
prominent beaded cords of alternating strength. Aperture
ovate, prosocline in prole, outer lip simple, Peristome
complete.
Discussion – We had initially placed Skenea minuticos-
tata nov. sp. and Skenea wareni nov. sp. in the genus Lod-
derena Iredale, 1924. Warén (1992, p. 155) considered the
genus Lodderena to included skeneimorph gastropods
with spirally striated shell with three or more scaley per-
iumbilical ribs [cords]. The present-day Mediterranean
Cyclostrema catenoides Monterosato, 1877, which was
placed in the genus Lodderena by Warén, is most like S.
wareni but differs in having only the periumbilical cords
strongly developed, the rest of the teleoconch surface is
covered with ne, close-set spiral cords. Warén (1992)
noted that the differences between Lodderena and Ske-
nea species were small and that they might turn out to
be congeneric. Indeed, L. catenoides was placed in the
genus Skenea Fleming, 1825 by WoRMS (Gofas, 2015).
We therefore follow WoRMS in placing these species
in Skenea and use Lodderena in a more restricted sense
following Rubio et al. (1998). Skenea wareni is similar
to the type species of Skenea, S. serpuloides (Montagu,
1808), but the latter does not have stronger periumbilical
cords and the axial sculpture is not lamellar.
Distribution – Upper Miocene: Tortonian, NW France
(this paper).
Genus Skeneoides Warén, 1992
Type species (by original designation) – Delphinula ex-
ilissima Philippi, 1844, present-day, Italy.
1992 Skeneoides Warén, p. 156.
Plate 75. Skenea wareni nov. sp.; 1. Holotype MNHN.F.A57684, diameter 1.4 mm, height 950 µm; 1d, detail of protoconch (SEM
image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
152 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
Skeneoides crassistriata Lozouet, 1999
Plate 76, g. 1
1999 Skeneoides crassistriata Lozouet, p. 12, pl. 7, gs
1-4.
Material and dimensions – Maximum diameter 1.9
mm, height 1.1 mm. St-Clément-de-la-Place: NHMW
2016/0103/0013 (1), NHMW 2016/0103/0014 (10),
RGM.1309640 (5), LC (2). Renauleau: NHMW 2016/
0103/0015 (4). Beugnon: NHMW 2016/0103/0016 (14).
Discussion – Skeneoides crassistriata Lozouet, 1999 is a
striking species, characterised by the extremely strongly
developed and elevated axial ribs. It is most similar to
the present-day Mediterranean S. jeffreysii (Monterosato,
1872), but this species has weaker primary axial ribs, sec-
ondary ribs in the interspaces and about ve spiral cords
mid-whorl. Skeneoides exilissima (Philippi, 1844), also
extant in the Mediterranean, again has weaker, more nu-
merous primary ribs and stronger spiral cords delimiting
the shoulder and base. Skeneoides tenuistriata Lozouet,
1999 from the Oligocene of France has ne reticulated
surface sculpture. This species was described based on
material from the middle Miocene of Ferrière-Larçon,
Indre-et-Loire, France.
We note that ICZN (1999a) Art. 30.1.4.4 rules that ge-
nus names ending –oides are masculine, and accordingly
that the name should be crassistriatus. But there is an
exception; that when treated as feminine in the original
description it should remain so. Warén (1992, p. 156)
nominated Delphinula exilissima Philippe, 1844 as type
species for Skeneoides, and used the combination Skene-
oides exilissima, meaning that he considered Skeneoides
to be feminine. Therefore, Lozouet (1992) was quite cor-
rect in keeping the trivial name for his new species femi-
nine.
Distribution – Middle Miocene: Atlantic (Langhian)
Loire Basin (Lozouet, 1999). Upper Miocene (Tortoni-
an): Atlantic, NW France (this paper).
Family Phasianellidae Swainson, 1840
Subfamily Tricoliinae Woodring, 1928
Genus Tricolia Risso, 1826
Type species (by subsequent designation, Gray, 1847b)
– Turbo pullus Linnaeus, 1758, present-day, Mediterra-
nean.
1826 Tricolia Risso, p. 122.
1847a Thicolia Leach in Gray, p. 271. Incorrect subse-
quent spelling.
1847b Thicolea Gray, p. 144. Incorrect subsequent spell-
ing.
1847b Tricolea Gray, p. 144. Incorrect subsequent spell-
ing.
1852 Eudora Leach in Gray, p. 147, 199. Type species
(by monotypy): Eudora varians Leach in Gray,
1852, present-day, Europe. Junior homonym of
Eudora Péron & Lesueur, 1810 [Cnidaria] and
several others.
1853 Chromotis H. & A. Adams, p. 19. Type species (by
monotypy): Phasianella neritina Dunker, 1846,
present-day, South Africa.
1884a Tricoliella Monterosato, p. 110. Type species (by
subsequent designation, Pilsbry, 1888): Turbo
pullus Linnaeus, 1758, present-day, Mediterra-
nean. Junior objective synonym of Tricolia.
Plate 76. Skeneoides crassistriata Lozouet, 1999; 1. NHMW 2016/0103/0013, diameter 1.8 mm, height 1.1 mm; 1c, detail of proto-
conch (SEM image). Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France, Tortonian, upper Miocene.
Cainozoic Research, 17(2), pp. 75-166, December 2017 153
1891 Steganomphalus Harris & Burrows, p. 112. Type
species (by typication of replaced name): Eudo-
ra varians Leach in Gray, 1852, present-day, Eu-
rope. Nom. nov. pro Eudora Leach in Gray, 1852,
non Péron & Lesueur, 1810 [Cnidaria].
1920 Epheriella Pallary, p. 48. Type species (by original
designation): Epheriella algoidea Pallary, 1920,
present-day, Morocco.
1973 Eutricolia Nordsieck, p. 10. Type species (by orig-
inal designation): Turbo speciosus Megerle von
Mühlfeld, 1824, present-day, Mediterranean.
Tricolia pullus s.l. (Linnaeus, 1758)
Plate 77, gs 1-6
*1758 Turbo pullus Linnaeus, p. 1233.
1854 Phasianella Turbinoides Lamk. – Millet, p. 158
(non Lamarck, 1804).
1925 Phasianella pulla var. pulchella Récluz – Harmer,
p. 870, pl. 65, g. 26.
1964 Tricolia pullus Linné, 1766 [sic] – Brébion, p.
136, pl. 3, gs 22-23.
2003 Tricolia pullus pullus (Linnaeus, 1758) – Landau
et al., p. 34, pl. 8, gs 1, 2 (cum syn.).
2004 Tricolia pullus (Linné, 1758) – Chirli, p. 42, pl.
13, gs 10-13, pl. 14, gs 1-5.
2010 Tricolia pullus (Linnaeus, 1758) – Sosso & Dell’
Angelo, p. 19, p. 30 unnumbered g. bottom right.
Material and dimensions – Maximum height 10.3
mm, width 5.7 mm. St-Clément-de-la-Place: NHMW
2016/0103/0518-0524 (7), NHMW 2016/0103/0525 (50+),
RGM.1309569 (3), RGM.1309570 (50+), RGM.1309574
(30 juveniles), RGM.1347889 (50+), LC (50), FVD (50+).
Sceaux-d’Anjou: RGM.1309581 (13), RGM.1309583 (1
adult + 3 juveniles), RGM.1347853 (50+), RGM.1347956
(26), RGM.1348021 (21), LC (10). Renauleau: NHMW
2016/0103/1482 (10), LC (10), FVD (3). Beugnon:
RGM.1309582 (41 adults and subadults).
Discussion – In the present-day faunas there are two sub-
species living off the shores of continental Europe. Trico-
lia pullus pullus (Linnaeus, 1958) found in the Mediter-
ranean, up to the Straits of Gibraltar and T. pullus picta
(Da Costa, 1778), which is predominantly an Atlantic
subspecies, ranging from the British Isles to the north,
southwards to Morocco. In the western Mediterranean
transitional morphologies can be found (Gofas, 1982).
Usually the Atlantic subspecies is thinner shelled, with
the upper part of the whorls slightly more attened, with
a zigzag colour pattern predominating, especially on the
base of the last whorl.
The Assemblage I specimens from St-Clément-de-la-
Place illustrated here (Pl. 77, gs 1-6) cover the entire
range of shell shape and pattern illustrated by Gofas
(1982, gs 1-13) for Mediterranean and Atlantic forms,
with globose and more slender forms, striped and spot-
ted forms, and we would hesitate to say if they are of
the pullus or picta morphotype. It is possible that in the
upper Miocene this distinction was not present and that
the ancestral form exhibited both morphotypes. Despite
several attempts at SEM imaging to illustrate the proto-
conch, none of the ve specimens examined had a clearly
delimited protoconch/teleoconch boundary. We therefore
provisionally consider the Assemblage I population to be
T. pullus sensu lato. We note that specimens from Re-
nauleau are much larger, up to 10 mm in height.
The middle Miocene Loire Basin shell illustrated by
Glibert (1949, p. 78, pl. 4, g. 16) as Tricolia millepunc-
Plate 77. Tricolia pullus (Linnaeus, 1758); 1. NHMW 2016/0103/0518, height 4.2 mm; 2. NHMW 2016/0103/0519, height 5.3 mm;
3. NHMW 2016/0103/0520, height 4.5 mm; 4. NHMW 2016/0103/0521, height 4.3 mm; 5. NHMW 2016/0103/0522, height 4.4
mm; 6. NHMW 2016/0103/0523, height 4.3 mm. Le Grand Chauvereau, St-Clément-de-la-Place, Maine-et-Loire, NW France,
Tortonian, upper Miocene.
154 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwestern France, 1. Patellogastropoda and Vetigastropoda
tata (Benoist, 1873) is umbilicate and therefore probably
a distinct species. The Loire Basin shell identied by
the same author as T. eichwaldi (Hörnes, 1855) is also
probably correct, and differs from T. pullus in having a
relatively expanded last whorl, low spire and a concave
subsutural collar (see also Landau et al., 2013, p. 34).
The Pliocene to present-day Mediterranean T. tenuis
(Michaud, 1829) differs from T. pullus in having a higher
spire, atter whorls and a predominance of dotted pat-
tern, with the ammules restricted to area immediately
below the suture.
Brébion (1964, p. 137) recorded this species from As-
semblage I (Renauleau, Thorigné, Sceaux-d’Anjou, St-
Clément-de-la-Place, Beaulieu).
Distribution – Upper Miocene (Tortonian): Atlantic, NW
France (Brébion, 1964). Lower Pliocene: central Mediter-
ranean, Tunisia (Fekih, 1975), Italy (Pavia, 1975; Chir-
li, 2004; Sosso & Dell’Angelo, 2010). Upper Pliocene:
western Mediterranean, Estepona, Spain (Landau et al.,
2003); central Mediterranean, Italy (Sacco, 1896a). Low-
er Pleistocene: Atlantic, St Erth, England (Harmer, 1925).
Pleistocene: western Mediterranean, Balearic Islands,
(Cuerda Barceló, 1987); central Mediterranean, Italy
(Cerulli-Irelli, 1916; Taviani et al., 1998). Present-day:
western coasts of British Isles, south into the Mediterra-
nean, intertidal to 35m deep in the north, in the Mediter-
ranean often found in Posidonia elds in shallow water
(Poppe & Goto, 1991).
Remarks
In this work on the Patellogastropoda and Vetigastropoda
of the upper Miocene Tortonian Assemblage I localities
(sensu Van Dingenen et al., 2015) 76 species are record-
ed, of which 26 are new to science, six are homonyms and
receive replacement names and ve are left in open no-
menclature. This is an important increase in the diversity
compared to the 44 species recorded by Brébion (1964).
We were able to identify all the vetigastropods recorded
by Brébion except for four:
Calliostoma cf. conulum [sic] Linnaeus, 1758 (1964, p.
82, pl. 2, g. 1). Recorded at Renauleau (1) and Sceaux-
d’Anjou (2), the specimen illustrated is poorly preserved
and only half the height of present-day specimens. This
record is doubtful and requires conrmation.
Calliostoma hybridum Millet, 1854 (1964, p. 89, pl. 2, g.
10). This is a nomen nudum, later formally described by
Millet (1865, p. 583). Unfortunately this is a junior homo-
nym of Trochus hybridus Linnaeus, 1758. Recorded from
Renauleau (2) and Sceaux-d’Anjou (2), we were unable to
nd any specimens that we could ascribe to this species.
The specimen illustrated by Brébion might be a form of
Calliostoma gratiosum (Millet, 1865).
Calliostoma couffoni Brébion (1964, p. 90, pl. 2, g. 11)
nomen nudum. Despite being recorded from numerous
Assemblage I deposits by Brébion, we have not found any
specimens that t this species.
Gibbula (Colliculus) varia var. monodontoides Millet,
1854 (1964, p. 108, pl. 3, gs 1-2). This is a nomen nudum,
later formally described and validated by Millet (1865,
p. 582). This species seems to be abundant at St-Michel,
which was not sampled and Apigné, which is a Messinian
Assemblage II locality. Hopefully we will come across
this species later in the series.
We highlight the rst description of a member of the sub-
family Thysanodontinae in the European faunas and the
rst fossil record, with the description of Thysanodonta
chauvereauensis nov. sp. Marshall (1988) described the
subfamily as having a vicariant distribution in South Af-
rica, Australia, New Zealand and New Caledonia. Sub-
sequent reports and additions of new species have not
increased this geographic distribution (Marshall, 1995;
Vilvens & Maestrati, 2006). The description of the sub-
family in Europe suggests a wider distribution, and it is
quite possible further members remain unrecognised or
described as Calliostoma species.
A full synthesis of the Assemblage I fauna will be given
at the end of the series, but already some preliminary
observations are worthy of discussion. The Assemblage
I vetigastropod fauna is remarkable in several respects.
As far as we are aware, it is the most speciose European
Neogene gastropod assemblage known, with 76 species
representing 33 different genera/subgenera. In contrast
with this specic diversity, almost no species attains a
height greater than a couple of centimetres. Even gen-
era usually represented by medium sized shells such as
Diodora, Gibbula and Calliostoma are represented in
Assemblage I by small species or dwarf forms of spe-
cies found in other assemblages. The composition of the
fauna suggests shallow water environment, littoral zone
and the presence of genera such as Patella, Tectura,
Diodora, Gibbula, Jujubinus, and Calliostoma amongst
others, and the abundance of specimens in these genera,
suggest a hard bottom environment, which are usually
under-represented in the fossil record.
The fauna is also highly endemic, with 48 of the 76 (63%)
species known only from the northwestern French As-
semblage I localities (Fig. 2). Of these 75 species only 9
(12%) survived into the lower Pliocene Assemblage III
fauna of northwestern France. Thirty-eight (51%) also oc-
cur in the earlier middle Miocene Langhian assemblage
of the Loire Basin, but relationships with other European
basins are weak; six (8%) also occur in the Miocene-
present-day North Sea Basin, 14 (19%) in the Mediterra-
nean and one (1.3%) in the Pleistocene of St. Erth (Corn-
wall, England).
Amongst the vetigastropods, the generic composition
is similar to that seen today; a few genera are extinct,
such as Lucapinella, Paroxystele and Pareuchelus. There
are a few elements suggesting thermophilia; the genus
Leucorhynchia occurs today in a more southern distribu-
tion, found off the coast of West Africa, and the species
Haliotis tuberculata coccinea Reeve, 1846 and Gibbula
fanulum (Gmelin, 1791).
Further remarks will be made as the series develops.
Cainozoic Research, 17(2), pp. 75-166, December 2017 155
Figure 2. Geography, stratigraphy and distribution of species found in the upper Miocene Tortonian Assemblage I localities of
northwestern France. For geographic distribution 1 = North Sea Basin, 2 = Atlantic coasts British Isles, 3 = NW France, 4 =
Mediterranean. For stratigraphic distribution black signies Atlantic distribution (A), grey Mediterranean distribution (M).
(Continued next pages).
156 Landau, Van Dingenen & Ceulemans. The upper Miocene gastropods of northwester n France, 1. Patellogastropoda and Vetigastropoda
Cainozoic Research, 17(2), pp. 75-166, December 2017 157
Acknowledgements
We would like to thank Philippe Bouchet, Jean-Michel
Pacaud and Pierre Lozouet of the Muséum national
d’Histoire naturelle (Paris) for their helpful suggestions,
cooperation and allowing us access to the Brébion’s col-
lection in Paris. Daniel Geiger, Curator of Malacology,
Santa Barbara Museum of Natural History, California,
for his advice on the Haliotidae and Scissurellidae. An-
ders Warén, Department of Invertebrate Zoology, Swed-
ish Museum of Natural History, Stockholm, Sweden,
for his advice on the skeneimorph gastropods. Carlos
Marques da Silva of the University of Lisbon, Portugal,
for his advice and help with graphics. Ronald Pouwer,
fossil mollusc curator at the Naturalis Biodiveristy Cent-
er in Leiden (The Netherlands), for making the RGM
collections available to us and helping with some of the
photography. Renate Helwerda, also from Naturalis, for
help with photography and taxonomic suggestions. To
our reviewers Renate Helwerda (see above), Alan Beu,
Paleontology Department, GNS Science, Lower Hutt,
New Zealand, and Bruce Marshall, National Museum of
New Zealand, Wellington, New Zealand for their sugges-
tions and advice, to Mr. Hammoneau, Mr. Justeau, fam-
ily Lherbette for allowing us to collect on their property,
and last but certainly not least a special thanks to Arie
W. Janssen, also from Naturalis, for taking the time to
pre-review this work and make many useful suggestions.
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