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Cainozoic Research, 19(2), pp. 117-120, December 2019 117
Introduction
Even in relatively well-studied fossil faunas, unexpected
discoveries continue to be made. A case in point involves
three species from the Lutetian and the Priabonian (Mid-
dle and Late Eocene) of France. In our examination of
specimens from the Paris Basin, together with a reap-
praisal of relevant literature, we uncovered a new clade of
gastropods with a previously unrecognized labral tooth.
Here we describe this group as the new genus Denticulo-
fusus and compare its characteristics to those of related
fossil and living genera.
Material Examined
MNHN.F = Muséum national d’Histoire naturelle, Col-
lection de Paléontologie (Paris, France)
Parnes, Carrière aux Hommes (Oise, France) MNHN. F.
B64775 (Sarazin coll.): 3 sp.
Parnes, Grande Cronière (Oise, France) MNHN.F.A71168
(Pacaud coll.): 5 sp.
Chaussy, Les Garennes (Val d’Oise, France) MNHN. F.
A71167 (Schtrock coll.): 5 sp.
Cahaignes, Requiécourt (Eure, France) MNHN.F.A71166
(Braillon coll.): 1 sp.
Cahaignes, Requiécourt (Eure, France) MNHN.F.B64776
(Lhomme coll.): 2 sp.
Fontenay-en-Vexin, Bois du But (Eure, France) MNHN -
.F. A71165 (Faullummel coll.): 5 sp.
Fontenay-en-Vexin, Bois du But (Eure, France) MNHN. F.
A71162 (Pacaud coll.): 4 sp.
Fontenay-en-Vexin, Bois du But (Eure, France) MNHN. F.
A71163 (Pons coll.): 16 sp.
Villiers-Saint-Frédéric, Butte Saint-Léonard (Yvelines,
France) MNHN.F.A71164 (Pons coll.): 3 sp.
Systematic Palaeontology
Superfamily Turbinelloidea Swainson, 1835
Family Columbariidae Tomlin, 1928
Genus Denticulofusus, new genus
Type species – Fusus gothicus Deshayes, 1834, Lutetian,
Middle Eocene, Paris Basin (Figure 1)
Etymology – Combination of Latin denticula (small tooth)
and fusus (spindle).
Diagnosis – Shell narrowly fusiform, with relatively low
spire, and a long, very narrowly open siphonal canal;
protoconch multispiral, high conical, consisting of 4 con-
vex whorls; rst three whorls smooth and fourth whorl
with strong, widely arched folds parallel to protoconch/
teleoconch transition; sutures channeled; spiral sculpture
consisting of cords and a peripheral shoulder angulation;
A new Eocene genus of Columbariidae (Gastropoda:
Turbinelloidea) with a labral tooth
Geerat J. Vermeij1,3 and Jean-Michel Pacaud2
1 University of California, Davis; Department of Earth and Planetary Science, One Shields Avenue, Davis, CA, 95616,
USA; e-mail: gjvermeij@ucdavis.edu
2 CR2P Centre de Recherche en Paléontologie – Paris; Muséum national d’Histoire naturelle, Sorbonne Université,
CNRS; 8 rue Buon, CP 38, 75005 Paris (France); e-mail: pacaud@mnhn.fr
3 corresponding author
Received: 8 May 2019, revised version accepted 27 July 2019
A previously unrecognized new genus, Denticulofusus (type species: Fusus gothicus Deshayes, 1834, Lutetian of the Paris Basin,
France) is proposed for Middle Eocene columbariine turbinellid gastropods with a labral tooth situated at the outer lip at the end
of a spiral groove. Additional species, both from the Lutetian and Priabonian of the Cotentin Basin of Normandy, France, are
respectively D. monicae (Belliard, Gain & Le Renard, 2017) and D. cavelieri (Gain & Le Renard, 2017). The presence of the labral
tooth distinguishes Denticulofusus from the Eocene to Recent genus Coluzea Finlay in Allan, 1926, and the Eocene genus Falsifusus
Grabau, 1904.
KEY WORDS: Eocene, Columbariidae, Denticulofusus, new genus, labral tooth
118 Vermeij & Pacaud - A new Eocene genus of Columbariid ae with a labral tooth
Figure 1. A: Adult specimen : Lutetian of Par nes, Grande-Cronière (Oise), MNH N.F.A71161 (leg. Pacaud); B: Juvenile specimen,
protoconch: Lutetian of Fontenay-en-Vexin, Bois du But (Eure), MNHN.F.A71180 (leg. Pacaud); C: Juvenile specimen : Lutetian
of Fontenay-en-Vexin, Bois du But (Eure), MNHN.F.A71180 (leg. Pacaud). Photographs Gaëlle Doitteau (e-recolnat Project,
MNH N)
axial sculpture consisting of about twelve angular ribs;
aperture small; outer lip with broad adapical sinus, its
edge abapically with three small labral teeth, the largest
of which is situated between the lowest two cords; in-
ner side of outer lip smooth; inner lip adherent; adapical
ridge on inner lip absent.
Included species – Fusus gothicus Deshayes, 1834; Colu-
zea cavelieri Gain & Le Renard, 2017; Coluzea monicae
Belliard, Gain & Le Renard, 2017.
Cainozoic Research, 19(2), pp. 117-120, December 2019 119
Remarks – While examining several French and English
Eocene species that have often been considered to belong
to the fasciolariid subfamily Fusininae (e.g. Wrigley,
1927), we discovered a previously overlooked feature on
the shell of a species originally named Fusus gothicus
by Deshayes (1834; 1835). This feature, a blunt, small
labral tooth located at the end of a groove between the
lowest two cords on the last whorl, is not an artifact of
preservation, for it can also be discerned at the same po-
sition at earlier apertural positions. Inspection of illustra-
tions further revealed that a similar tooth is present at the
same position and at earlier growth stages in two recently
described species from the Cotentin Basin of Norman-
dy, France, originally named Coluzea cavelieri and C.
monicae. The tooth was not mentioned in the description
of any of these three species.
We assign Denticulofusus to the Columbariidae largely
because of the presence of a peripheral keel on adult
whorls. The Eocene fusinine fasciolariid genus Eofusus
Vermeij & Snyder, 2018 resembles Denticulofusus but it
has an enlarged peripheral cord rather than a keel, and
the outer lip tends to be somewhat convex rather than
straight as in Denticulofusus and other columbariids. No
fossil or living member of the Fusininae has a labral tooth
(Vermeij & Snyder 2018).
All three taxa for which we establish the new genus Den-
ticulofusus were assigned previously to Coluzea Finlay
in Allan, 1926 (Pacaud & Le Renard, 1995; Belliard et
al., 2017; Gain & Le Renard, 2017). Other Eocene Eu-
ropean species that have been assigned to Coluzea lack
the labral tooth, as in C. dissimilis (Deshayes,1834), C.
erecta (von Koenen, 1889), C. londini (Wrigley, 1927),
C. multilirata (von Koenen, 1889), C. regnorum (Wrig-
ley, 1927), C. unicarinata (Deshayes, 1834) and C. ytenae
(Wrigley, 1927). The ontogeny of the primary cords in
these species without a labral tooth diers from that in
Denticulofusus and from that in living species of Coluzea
as well as from that in the type of Coluzea, the Middle
Miocene C. dentata (Hutton, 1877) from New Zealand
(see Beu & Maxwell, 1990, for a redescription). In the
living fauna, species assigned to Coluzea are distributed
in deep waters in New Zealand, tropical Australia and
southeastern Africa (Darragh, 1969; 1987; 1997; Har-
asewych, 2011). In New Zealand the genus extends back
to the Early Eocene (Beu & Maxwell, 1990). The name
Coluzea should perhaps be restricted to the Late Eocene
and younger species of Coluzea in New Zealand. These
younger species, unlike the earlier Eocene taxa in New
Zealand and living species in Africa and Australia, have
nodes instead of open spines on the peripheral keel (see
also Harasewych, 2019). A new generic name would then
be needed for the open-spined species previously attrib-
uted to Coluzea. The only European species in that group
is Fusus serratus Deshayes, 1825, from the Lutetian of
the Paris Basin.
Vermeij & Snyder (2018) propose that other European
species of “Coluzea” without a labral tooth (other than
C. serrata) bear a very close resemblance to the Eocene
American genus Falsifusus Grabau, 1904 (type spe-
cies: Fusus ottonis Aldrich, 1897). The only dierence
between Falsifusus and the European taxa is that the
protoconch of the former is multispiral, whereas that
of the European taxa is paucispiral. Given the frequent
transitions from multispiral to paucispiral protoconchs
in many gastropod lineages, we consider the European
species without a labral tooth to represent derived condi-
tions within Falsifusus, a genus that, as Vermeij & Sny-
der (2018) indicate, belongs to the Columbariidae and not
to the Fusininae.
The only other Palaeogene European genus that appears
to belong to the Columbariidae is Clavogyra Leroy, 2018
(type species: C. ledeei Leroy, 2018) from the Cuisian
(Late Ypresian, Early Eocene) of the Paris Basin. Its shell
is exceptionally large (up to 120 mm), more than twice
the length of species of Denticulofusus; and its spiral
sculpture features two keels. There is no mention of a
labral tooth in this genus.
Within Columbariidae, a labral tooth also occurs in the
genus Columbarium von Martens, 1881. In that genus,
the tooth is situated at the end of a basal cord or keel
(Darragh, 1969; Vermeij, 2001) rather than between
cords as in Denticulofusus. Although the earliest spe-
cies of Columbarium, C. antecedens Pacaud, 2015, is
known from the Danian (Early Palaeocene) of Belgium
and France, our examination of specimens at the Insti-
tut Royal des Sciences Naturelles de Belgique in Brus-
sels and at the Muséum national d’Histoire naturelle in
Paris failed to reveal a labral tooth, calling into question
the generic placement of this species. Leroy (2018) has
pointed out similarities between this species and Palaeo-
gene members of Coluzea. The early fossil record of Co-
lumbarium is otherwise conned to Australasia, where it
is known from the Palaeocene onward (Dar ragh, 1969;
Harasewych, 2011). The early fossil record of Columbar-
ium is otherwise conned to Australia and New Zealand,
where it is known from the Palaeocene onward (Darragh,
1969; 1987; 1991). Fusus rugatus Aldrich, 1886 from the
Late Palaeocene Gregg’s Landing Formation of Alabama,
attributed by Darragh (1969) to Columbarium, was said
by Darragh to be unlike any early Australian species, and
instead is more likely a member of the Palaeocene to Re-
cent New World genus Fulgurofusus Grabau, 1904 (see
also Grabau, 1904). In the living fauna, Columbarium
occurs in Australasia, the South China Sea, Japan, and
southern Africa (Darragh, 1969; Harasewych, 2011). Co-
lumbarium therefore appears always to have been a genus
in the western Pacic and Indian Oceans. Its distribution
does not overlap with that of Denticulofusus, and we sur-
mise that the labral tooth evolved independently in Den-
ticulofusus and Columbarium.
Acknowledgements
We thank Jonathan A. Todd (Natural History Museum),
who showed the senior author specimens of the type
species of our new genus; Gaëlle Doitteau (MNHN)
and Lilian Cazes (MNH N/CNRS) for their assistance
in photographic work, and Alyssa Henry for her techni-
cal assistance. This work beneted from a grant from
120 Vermeij & Pacaud - A new Eocene genus of Columbar iidae w ith a labral tooth
the French state managed by the Agence nationale de la
Recherche via the program “Investissements d’avenir”
(ANR-11-INBS-0004-RECOLNAT).
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