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Carnets de Voyages Paléontologiques
dans le Bassin Anglo-Parisien
Tome 3 (2017)
LRG-Éditions
CARNETS DE VOYAGES PALÉONTOLOGIQUES
DANS LE BASSIN ANGLO-PARISIEN
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Directeur de la publication : J. LE RENARD
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les conditions de soumission de leurs articles.
ISBN : 978-2-9538000-3-6
Dépôt légal : Octobre 2017 (Réédition de Mars 2021)
Copyright (including artwork) belongs to Jacques LE RENARD & Olivier GAIN, no part of this publication may be reproduced, stored in retrieval
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Jacques LE RENARD
78640 NEAUPHLE LE CHÂTEAU
lerenard@claranet.fr
Olivier GAIN
50130 CHERBOURG EN COTENTIN
lamarck@wanadoo.fr
APGC
Associat ion Paléontologiq ue et Géologique du Cote ntin
@: cossmannia@cossmann.org
Carnets de Voyages Paléontologiques dans le Bassin Anglo-Parisien
Tome 3 (2017)
Sommaire
_________
Steve TRACEY, Brian CRAIG, Laurent BELLIARD & Olivier GAIN : One, four or
forty species? - early Conidae (MOLLUSCA, GASTROPODA) that led to a
radiation and biodiversity peak in the late Lutetian Eocene of the Cotentin,
NW France. - -- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -- - - - p. 1-38
Laurent BELLIARD, Olivier GAIN & Jacques LE RENARD : Les genres Coluzea
et Fusinus (MOLLUSCA, GASTROPODA, NEOGASTROPODA) dans l’Éocène Moyen
du Cotentin (Manche, France). - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -- -- - - - - -p. 39-56
Olivier GAIN & Jacques LE RENARD : Occurrence du genre Coluzea
(MOLLUSCA, GASTROPODA, NEOGASTROPODA) dans l’Éocène Supérieur du
Cotentin (Manche, France). - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -p. 57-60
Olivier GAIN, Laurent BELLIARD & Jacques LE RENARD : Redéfinition
taxinomique de Fusus brasili Cossmann & Pissarro, 1905; “quand petit
‘Fusus’ deviendra grand”- - - - - - -- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - --- - - - - - - - - - - -p. 61-80
Malcolm Francis SYMONDS, Olivier GAIN & Laurent BELLIARD : New species
of Neritidae (MOLLUSCA, GASTROPODA, NERITIMORPHA) from the Middle
Eocene of the Cotentin (Manche, France). - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -p. 81-88
Olivier GAIN, Laurent BELLIARD & Jacques LE RENARD : La famille des
Cassidae (MOLLUSCA, GASTROPODA, CAENOGASTROPODA) dans l’Éocène
Moyen du Cotentin (Manche, France). - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - --- - - - - -- - p. 89-112
Laurent BELLIARD, Olivier GAIN & Jacques LE RENARD : Le genre Rimella
(MOLLUSCA, GASTROPODA, CAENOGASTROPODA) dans l’Éocène Moyen du
Cotentin (Manche, France). - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - -p. 113-123
Annexes - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - - - - - - - - - - - - - - - - - - - - p. 124-143
LRG-Éditions
@ le-renard.gain@lrg-editions.fr
ISBN: 978-2-9538000-3-6
Dépôt légal : Octobre 2017 (Réédition de Mars 2021)
Carnets de Voyages Paléontologiques dans le Bassin Anglo-Parisien Tome 3
Août 2017, pp. 1-38, figs. 1-132, pls. 1-9 —ISBN 978-2-9538000-3-6
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
One, four or forty species? - early Conidae (Mollusca, Gastropoda) that led
to a radiation and biodiversity peak in the late Lutetian Eocene of the
Cotentin, NW France.
Steve TRACEY(1), Brian CRAIG(2), Laurent BELLIARD(3) & Olivier GAIN(4)
1. Department of Earth Sciences, The Natural History Museum, London SW7 5BD, U.K. s.tracey@nhm.ac.uk
2. 25 Rodway Road, Bromley, BR1 3JJ, U.K. bcraig.amoria@googlemail.com
3. 94240 L’HAY-LES-ROSES, FRANCE. mololea@free.fr
4. 50130 CHERBOURG-OCTEVILLE, FRANCE. lamarck@wanadoo.fr
____________________
ABSTRACT - The earliest confirmed record of the family Conidae is the late Paleocene (Thanetian) Hemiconus
described herein as H. leroyi Tracey & Craig n. sp.. The genus Hemiconus appears to contain all described cone species
through the early to middle Ypresian. A second clade is first observed in the late Ypresian with the appearance of a
group of larger cones conveniently referred to Eoconus. Because of the extensive variation within Hemiconus, authors
have expressed different views on the number of recognisable species. The rapid radiation of Hemiconus observed in
the middle Lutetian of the Paris Basin, reached an acme in Normandy in the late Lutetian and this fauna is examined
from several sites in the Cotentin. The following new species are described by TRACEY & CRAIG: Hemiconus trisulcatus,
H. pissarroi, H. lateralis, H. constantinensis, and a new genus Papilliconus is proposed for P. papillatus n. sp. whose size
would suggest a relationship with Hemiconus but whose other characters do not agree. New species Papilliconus
radulfivillensis, Hemiconus auriculatus, and Eoconus veteratoris are described from Bartonian-Priabonian deposits at
Rauville-la-Place (Manche). A neotype is designated for Conus stromboides Lamarck, type species of Hemiconus, in
view of the confusion caused by the various interpretations of this species. The genera Hemiconus and Artemidiconus
are returned to the Conidae.
KEYWORDS : Mollusca, Gastropoda, Conoidea, Conidae, Hemiconus, Eoconus, Papilliconus, Artemidiconus, Cotentin,
Manche, France, Paleocene, Eocene, Lutetian, Bartonian-Priabonian.
TITRE - Une, quatre ou quarante espèces? – les Conidae (Mollusca, Gastropoda) précurseurs d'une ramification
ayant conduit à un pic de biodiversité dans l'Éocène Moyen terminal du Cotentin, au nord-ouest de la France.
RÉSUMÉ - Le premier représentant fossile confirmé de la famille des Conidae, H. leroyi Tracey & Craig n. sp.,
appartenant au genre Hemiconus, est signalé et décrit en provenance du Paléocène (Thanétien). Le genre Hemiconus
semble contenir toutes les espèces de cônes décrites dans l'Ypresien inférieure à moyen, mais un deuxième clade
émerge à la fin de l’Yprésien avec l'apparition d'un groupe de cônes plus grands, commodément désignés par
Eoconus. En raison de la grande variation au sein d’Hemiconus, les auteurs ont exprimé des points de vue différents
sur le nombre d'espèces reconnaissables. La rapide radiation des Hemiconus, observable d'abord dans le Lutétien
moyen du bassin de Paris, a atteint son apogée en Normandie à la fin du Lutétien. Cette faune provenant de différents
sites du Cotentin est ici étudiée. Les nouvelles espèces suivantes sont décrites par TRACEY & CRAIG: Hemiconus
trisulcatus, H. pissarroi, H. lateralis, H. constantinensis et un nouveau genre Papilliconus est proposé pour P. papillatus
n. sp., dont la taille suggère une parenté avec Hemiconus mais qui présente d’autres caractéristiques distinctives. Les
nouvelles espèces Papilliconus radulfivillensis, Hemiconus auriculatus et Eoconus veteratoris sont décrites dans les dépôts
tardifs de l’Éocène de Rauville-la-Place (Manche). Un néotype est désigné pour Conus stromboides Lamarck, espèce
type d’Hemiconus, compte tenu de la confusion engendrée par les différentes interprétations de cette espèce. Les
genres Hemiconus et Artemidiconus sont réintégrés dans les Conidae.
MOTS-CLÉS : Mollusca, Gastropoda, Conoidea, Conidae, Hemiconus, Eoconus, Papilliconus, Artemidiconus, Cotentin,
Manche, France, Paléocène, Éocène, Lutétien, Bartonien-Priabonien.
Type material figured is held in the Department of Earth Sciences at the Natural History Museum, London, England
(NHMUK).
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
INTRODUCTION
The earliest records of Conidae, currently considered
valid, emanate from France (see KOHN, 1990), in which
area this family apparently originated. KOHN (1990)
discussed and refuted most records of supposed cones
from Jurassic and Cretaceous deposits. He noted the
possible exception of Conus marticensis Matheron, 1843
from the late Cretaceous (Turonian) of Martigues,
France which has a spirally corded, flattened spire and
resembles some more modern cones. Further confir-
mation of this occurrence is desirable as the species
does not possess any of the characters that the fossil
record would suggest were plesiomorphic for the
Conidae. TUCKER & TENORIO (2009) referred C.
marticensis to their new genus Eoconus and cited
reports of the same species from the Oligocene of
Germany and Denmark without comment.
Text-fig. 1: “Conus” primitivus, Cretaceous (Albian), Madagascar,
dubiously assigned to Conidae (original figure after COLLIGNON,
1949).
Another dubious Cretaceous cone, Conus primitivus
Collignon, 1949 from the Albian of Madagascar (Text-
fig. 1), although based on a poorly preserved fossil,
lacking ornament and with sutures obscured by
matrix, was included in the extant genus Profundiconus
without explanation by TUCKER & TENORIO (2009). If
neogastropods were in fact present in the early
Cretaceous it is more likely that they were related to
the Buccinoidea, Cancellarioidea or Muricoidea
(TRACEY et al., 1993; TRACEY, 2010) rather than
Conoidea. In the absence of reliably confirmed fossil
records of true cones from the late Cretaceous or early
Paleocene, we follow KOHN (1990) in considering the
Conidae to have a Cenozoic origin, although we
demonstrate that the earliest record is the late
Thanetian species Hemiconus leroyi described herein
(see Appendix). This discovery, with an age of c. 58 Ma
(KING, 2016, Fig. 70), provides a new fossil calibration
date for generating chronograms from molecular
phylogenies. To date, molecular phylogenies of the
family Conidae have been calibrated using an earliest
fossil date of 55 Ma (Paleocene–Eocene boundary, now
56.0 Ma) from KOHN (1990) for the two early Eocene
(late Ypresian) species H. rouaulti (d’Archiac, 1850)
and H. concinnus (J. Sowerby, 1821), that he considered
to be the oldest well-authenticated cones. Subse-
quently, this date has been used, without revision, by
DUDA & KOHN (2005), CUNHA et al. (2005),
PUILLANDRE et al. (2015) and URIBE et al. (2017). The
single Paleocene species and two more known from
the middle Ypresian, Hemiconus bicoronatus
(Melleville, 1843) (Plate 1, Fig. 2) and Hemiconus fallax
Pacaud, 2016 share common characters in their spire
ornament and can be confidently referred to
Hemiconus. The late Ypresian saw H. rouaulti
(d’Archiac, 1850) (Plate 1, Fig. 4) and H. diastictus
(Cossmann, 1923) in the Aquitaine region and H.
concinnus (J. Sowerby, 1821) (Plate 1, Fig. 3) in the
London Basin. Two conid moulds recorded from the
Ypresian of India and referred with some doubt to
Neogene species by COSSMANN & PISSARRO (1909)
were too poorly preserved to be assignable to any
genus. Near the end of the Ypresian a second clade of
cones made its first appearance with
Cossmann, 1923 in the south of France. The clade was
characterised by larger shells, lacking a subsutural
row of beads but having sharp peripheral nodules on
the spire similar to those found in some modern Conus,
Conasprella and Profundiconus species. It continued in
the early Lutetian with Conus diadema Edwards, 1857
(non G.B. Sowerby, 1834) and C. cf. deperditus
Bruguière, 1792 in England and C. calvimontanus
Deshayes, 1865 in France. This clade expanded in the
middle Lutetian with a number of species worldwide,
including Conus diversiformis Deshayes, 1835 in France
(Text-fig 20) and C. sauridens Conrad, 1833 in
southeastern U.S.A. (Text fig. 2b), type species of
Eoconus (Tucker and Tenorio, 2009). Up until this
period only two distinct clades of cones, Hemiconus
and those we refer herein to Eoconus, can be
recognised. Two species in the middle Lutetian of
France and England, Conus parisiensis Deshayes, 1865
(Plate 1, Figs. 5a-b) and C. filifer Edwards, 1857 (Plate
1, Figs. 6a-b) both retained a multispiral protoconch
and developed a smooth, straight-sided body whorl,
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
and have been thought to represent Conilithes or other
genera (e.g. TRACEY & TODD, 1996). However, the
subsutural row of beads and the spire ornament of
tubercles crossed by spiral cords, best seen in imma-
ture shells, show that they should be referred to
Hemiconus. The English C. selseiensis Cossmann, 1896
(Plate 1, Fig. 8) has a pleated subsutural collar
recalling that of Hemiconus except that the pleats are a
result of regular, incised growth striae rather than
discrete rounded beads. With its low spire C. selseiensis
Cossmann, 1896 has a somewhat distant resemblance
to Hemiconus and so we refer it to this genus with a
query. Typical Hemiconus (e.g. H. stromboides
(Lamarck, 1802), Plate 1, Fig. 7) radiated in the middle
Lutetian of the Paris Basin and notably in the late
Lutetian of the Cotentin, and the resulting diversity is
discussed below.
PROBLEMS RELATING FOSSIL SHELL
MORPHOLOGIES TO MODERN GENERA
With over 800 living species considered valid the
family Conidae (Fleming, 1822), in the traditional
sense, is one of the most diverse molluscan families,
although in terms of shell morphology it is relatively
conservative. Cone shells, both fossil and Recent are
readily recognisable as such. Whereas once it was
common practice to refer all fossil cones to the genus
Conus (s. lat.), modern opinions have tended to divide
the living cones into various numbers of groups based
largely on shell morphology (da MOTTA, 1991),
radular characters (TUCKER & TENORIO, 2009) or
molecular phylogeny (PUILLANDRE et al., 2015), the
latter two methods being problematical for deter-
mining the taxonomy and nomenclature of Eocene
species. TUCKER & TENORIO (2009, 2013) proposed a
division of the Conidae into 115 genera spread
between 5 families. PUILLANDRE et al. (2015) modified
this arrangement to comprise a single family of living
cones with four genera within which a number of the
new genera of TUCKER & TENORIO (2009) were given
subgeneric status or found to be synonyms of other
genera. Two additional genera (Lilliconus and
Pseudolilliconus) were shown by URIBE et al. (2017) to
comprise deeply divergent clades in their molecular
phylogenetic analysis that also confirmed the
monophyly of the Conidae. One, perhaps unwelcome,
result of this is that most fossil species of uncertain
affinities can no longer be referred to Conus (s. lat.)
without further qualification, as this term now applies
only to species in the Large Major Clade (sensu DUDA
& KOHN, 2005) defined by Conus marmoreus Linnæus,
1758 and excludes the Small Major Clade comprising
Conasprella and its three sister genera, and the basal
living clade of Profundiconus (Uribe et al., 2017).
TUCKER & TENORIO (2009) distinguished Hemiconus
from other Conidae principally by the lack of a
‘dentiform plait’ (i.e. top of the columella, discussed
below) and based largely on this character elevated it
to a separate family Hemiconidae. However,
Hemiconus appears to accord with other Conidae in
most respects and we have therefore adopted the more
pragmatic arrangement of PUILLANDRE et al. (2015) of
a single monophyletic family Conidae in which we
include Hemiconus. The problems of attempting to
establish direct relationships between Eocene and
Recent cones is exemplified by Conus edwardsi
Cossmann, 1889 from the English middle-late Lutetian
which has traditionally been referred to Conus
(Leptoconus) owing to a resemblance in shell-shape to
the living C. (L.) amadis Gmelin, 1791. In fact the shape
of C. edwardsi in mid-growth and adulthood is nearer
to those of the contemporaneous American Eoconus
sauridens (Conrad, 1833) and living Indian Conus
(Splinoconus) bayani Jousseaume, 1872. (Text-fig 2).
Text-fig. 2: Convergence of shell shape between genera.
a. Eoconus? edwardsi (Cossmann, 1889) middle to late Lutetian,
Bramshaw, Hampshire.
b. Eoconus sauridens (Conrad, 1833) Lutetian, Stone City Bluff,
Texas, U.S.A.
c. Conus (Splinoconus) bayani Jousseaume, 1872 living, India.
However, unlike these species the body whorl of
juvenile C. edwardsi is covered in punctate spiral
grooves, resembling adult Conasprella (Dalliconus)
species such as C. (D.) hopwoodi (Tomlin, 1937) and
particularly Conus (Turriconus) acutangulus Lamarck,
1810. There is also a close resemblance in the spire
ornaments of these three species, consisting of a sub-
sutural collar limited by a punctate groove and a row
of peripheral nodules delimited by another weaker
groove below (Text-fig. 3). On this basis C. edwardsi
might be considered as ancestral to both the Small and
Large Major Clades (sensu DUDA & KOHN, 2005) of
living Conidae (Conus and Conasprella) despite having
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
a very different overall profile, except that the orna-
ment of other Recent species of Conasprella (Dalliconus)
such as the West Atlantic C. (D.) sauros (Garcia, 2006)
and C. (D.) armiger (Crosse, 1858) (see Kaicher, 1976 as
Conus clarki) closely resemble the late Ypresian
Hemiconus rouaulti (d’Archiac, 1850) from southern
France (Plate 1, Figs. 4a, b) with a similar shell shape.
The interrelationship of all these species is therefore
problematic and C. edwardsi is perhaps best referred to
Eoconus with a query. We note also that the middle
Ypresian species Conus bicoronatus Melleville, 1843 has
been referred to Conasprella by PACAUD (2016),
although we do not recognise a great similarity to any
species of that genus, the characters of C. bicoronatus
being essentially those of Hemiconus in which we
retain it.
Text-fig. 3: Similar early ontogeny in different clades.
a. Eoconus? edwardsi (Cossmann, 1889) late Lutetian,
Bramshaw, Hampshire.
b. Conus (Turriconus) acutangularis Lamarck, 1810 living, Indo-
Pacific.
c. Conasprella hopwoodi (Tomlin, 1937) living, Indo-Pacific.
CHARACTERS USED TO ASSESS CONID SHELL
MORPHOLOGY IN FOSSILS
Protoconch : In the species reviewed two main
protoconch types have been observed, a) Multispiral :
up to 6 whorls, sinusigerous, conical with deep
sutures, embryonic whorl c. 150 µm. diameter; b)
Paucispiral: 1–2 whorls, growth lines orthocline,
tumid, embryonic whorl(s) 450 µm. to over 1 mm.
diameter. Intermediate forms are also found with
slightly larger embryonic shells and c. 2-4 larval
whorls, perhaps indicating shorter larval lives. Conid
protoconchs have been useful for distinguishing
fossils at the specific level, but they appear to have less
value above specific rank as a similar range is found in
each of the major clades of both fossil and living
Conidae. Furthermore, the range of larval shells of
Conidae is very similar to those of some other
conoidean families, e.g. Terebridae and Conorbidae
(TRACEY, in prep.).
Text-fig. 4: Shell shape terminology of early Conidae
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
DUDA & PALUMBI (1999) produced phylogenies
showing that transition from planktotrophic to leci-
thotrophic development in cones was a common
phenomenon and took place repeatedly in different
lineages. In all cases the transition involved loss of
planktotrophy while the regaining of a free-swimming
larva had apparently never been indicated. There
would therefore be no reason to create new conid
genera solely on the basis of the protoconch type.
Plate 9 highlights the range of protoconch sizes
observed in the genera covered here and the diverse
ornamentation of the spire whorls when seen in apical
view. Protoconch measurements given here follow the
methods outlined by LEAL (1991).
Adult shell-shape : Ignoring extreme forms, most
living cones have a conical body whorl with a spire
that is initially acute, enlarging rapidly to form an
obtusely angled or flattened top with varying degrees
of angularity on the shoulder. Distinguishing features
are often lost in many seemingly adult, shallow water
cones so that, if a number of Recent cones deprived of
their colour patterns were mixed together it is doubt-
ful whether they could confidently be sorted into
species-level, or even genus-level groups. The range of
shell shapes of living Conidae (see RÖCKEL et al. 1995)
are not particularly representative of Paleogene fossil
cones and Text-fig. 4 shows some of the commonest
profiles seen in the studied material.
Spire : The ontogeny of the spire whorls may be a
useful character for showing generic traits (although
see caveats above in relation to Eoconus? edwardsi). The
presence of peripheral nodules on the earliest teleo-
conch whorls is considered a synapomorphy of the
Conidae (TUCKER & TENORIO, 2009).
Shoulder nodules : The initial nodules typically
develop into relatively large, low and rounded tuber-
cles (Hemiconus) or smaller, more numerous nodules
limited by a sharp upper carina (Eoconus). Peripheral
nodules may persist to the last whorl or become fused
into an angular ridge at some stage of their develop-
ment, a character state commonly found in living
Conus, Conasprella and Profundiconus.
Subsutural collar : A row of beads immediately
below the suture, interspersed with growth striae but
not formed by them, is found in all early Hemiconus
and is here considered a plesiomorphic character of
the genus. These beads were obsolescent on many
Hemiconus by the late Lutetian and the Bartonian
although H. scabriculus (Solander, 1766) from the
Bartonian and H. granularis (Borson, 1820) from the
Miocene retained them. Without this key character,
referral of later species to Hemiconus is somewhat
subjective. A subsutural collar divided by regular
incised growth striae giving the appearance of elon-
gate beads as seen in the English middle Lutetian
Hemiconus? selseiensis (Cossmann, 1896) still occurs
rarely in living cones, e.g. Conus (Turriconus)
acutangulus and some Conasprella species. Another
Recent Indo-Pacific Conasprella, C. coromandelica
(Smith, 1894), displays true subsutural beading as in
Hemiconus. Whether or not this is the result of conver-
gence has yet to be determined.
Features of the inner lip : TUCKER & TENORIO
(2009) separated Eoconus and some other genera using
the presence or absence of a ‘dentiform plait’ on the
columella. This refers to the junction of the short,
thickly callused columella with the long, non-callused
paries (i.e. the previous whorl) which together
comprise the inner lip. We have examined many
Recent and fossil cones and find that the plait-like top
of the columella at this junction is more prominent in
broader, thick-shelled species (e. g. the living Conus
marmoreus) but less so in narrower ones with thinner
shells. The same variation occurs to a lesser extent in
Hemiconus. The ‘cicatrice pariétale’ used as a distingui-
shing character of Hemiconus by COSSMANN (1889,
1896) refers to a rounded groove located on the paries
at the top of and slightly inside the aperture and is
more apparent when the outer lip is broken away. The
groove extends for a short distance below an ena-
melled muscle scar that adjoins the interior suture and
may be shallow or deep, parallel with the suture and
then rising sharply towards the aperture. Examination
of a number of Recent and fossil species suggests that
it occurs in all Conidae, where its width and depth
varies between individuals, and it also appears in
typical Conorbidae (Conorbis). We therefore do not
consider either of these inner lip characters to have
any particular taxonomic significance within the
Conidae.
RANGE AND PALEOECOLOGY OF THE
HEMICONUS CLADE
It has generally been assumed that Hemiconus is now
extinct, but its widespread dispersal in the Neogene
would suggest that the genus might well have survi-
ved until today. Hemiconus granularis (Borson, 1820), a
species with granular ornament and similar shape to
H. scabriculus (Solander, 1766) is widely recorded
(under various varietal names and synonyms) from
the early to middle Miocene of Italy, Poland, Austria,
Czech Republic and France (SACCO, 1893; BALUK, 1997;
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
see synonymy in HARZHAUSER & LANDAU 2016). The
purported type locality of H. granularis in Valle
Andona, Italy (Pliocene) is thought to be an error, the
most likely age being late Miocene (Messinian). The
species was reassigned to the monotypic genus
Artemidiconus da MOTTA, 1991 (J.K. TUCKER in
HARZHAUSER & LANDAU, 2016, p. 39) which was
treated as a genus of Conorbidae by TUCKER &
TENORIO (2009). The somewhat contradictory charac-
ter assessments of Artemidiconus given in these two
publications need clarification here. The fact that both
Miocene H. granularis and living West Atlantic
Artemidiconus can show peripheral tubercles/nodules
on the earliest teleoconch whorls classifies both as
Conidae rather than Conorbidae. Furthermore, the
presence of a subsutural row of beads and lower row
of tubercles in H. granularis (see HARZHAUSER &
LANDAU, 2016, p. 39-40, fig. 3A) suggests that H.
granularis should be referred to Hemiconus, as defined
herein. If H. granularis is closely related to living
Artemidiconus as suggested by these authors, then
perhaps the Hemiconus clade lives on today. In view of
its granular body whorl ornament and 3-whorl
multispiral protoconch this species is more likely to be
derived from the lineage of H. scabriculus than from
any of the paucispiral Cotentin species described
below. In the Cotentin the genus displays a much
greater morphological variability than one would
infer from the diagnosis of its type species, and
exceeds the diversity seen in other conid clades at any
one time and place, except perhaps some living Indo-
Pacific faunas. Granular ornamentation of the last
whorl seems to be a character that shows interspecific
variation in a number of living species, although we
have considered it to be a useful feature for differen-
tiating some Hemiconus fossils. A number of living
species exhibit similar granular ornaments, including
some members of Conus and Conasprella, but conti-
nuing molecular studies would be required to inves-
tigate if this could be due to convergence. The facies
succession at the late Lutetian Cotentin localities
studied is of siliceous bioclastic limestone with
Campanile, fining upward to calcareous muds with
sea-grass vegetation (unit FHB-A of LE RENARD &
GAIN, 2012) in which Hemiconus occur commonly. The
overlying unit FHB-B comprises muddy sands with
Potamididae at Fresville and Hautteville or calcareous
sands with coral rubble in an adjoining basin at
Néhou. It is in these sheltered littoral habitats that the
late Lutetian radiation of Hemiconus occurred.
Here, for completion, we have also included three
related conid species encountered in the younger
(between Bartonian and late Priabonian) clayey silty
sands of Rauville-la-Place (‘Argiles à corbules’ of
VIEILLARD & DOLLFUS, 1875). The apparent reduction
of conid diversity at this time may have been a local
response to an apparently estuarine paleoenviron-
ment, but perhaps also reflects a general decline in
species numbers coterminous with global cooling after
the close of the late Lutetian.
Plate 1
Fig. 1 a-c : Hemiconus leroyi Tracey & Craig, n. sp. Holotype NHMUK PI TG 26839 (leg. S. TRACEY), Thanetian,
Cauroy-les-Hermonville (Marne) ;
Fig. 2 a-b : Hemiconus bicoronatus (Melleville, 1843) middle Ypresian, Saint-Gobain (Aisne) : a) MNHN
J.F.02897; b) (protoconch) NHMUK PI TG 26840 (leg. S. TRACEY) ;
Fig. 3 : Hemiconus concinnus (J. Sowerby, 1821), NHMUK PI TG 26833 (coll. WETHERELL), late Ypresian,
Highgate Archway, London ;
Fig. 4a-b : Hemiconus rouaulti (d’Archiac, 1850), late Ypresian, Tuilerie de Gan (Pyrénées-Atlantiques) : a)
NHMUK PI TG 26834 (leg. S. TRACEY), b) [protoconch] NHMUK PI TG 26841 (leg. S. TRACEY) ;
Fig. 5a-b : Hemiconus parisiensis (Deshayes, 1865): a) NHMUK PI TG 26842 (leg. S. TRACEY) middle Lutetian,
Cauvigny (Oise), b) NHMUK PI TG 26843 (leg. S. TRACEY) middle Lutetian, Fercourt (Oise) ;
Fig. 6a-b : Hemiconus filifer (Edwards, 1857) middle Lutetian, Bracklesham Bay (W. Sussex) : a) NHMUK PI
OR 71195(2) (coll. EDWARDS); b) (protoconch) ;
Fig. 7 : Hemiconus stromboides (Lamarck, 1802) NHMUK PI TG 26845 (leg. O. GAIN) late Lutetian, Néhou ;
Fig. 8 : Hemiconus? selseiensis (Cossmann, 1896) Lectotype NHMUK PI OR 71197(1) (coll. EDWARDS), middle
Lutetian, Bramshaw (Hants).
Scale: = 1cm. Figs: 1a-c, 2a, 3, 4a, 5a, 6a, 7, 8; ‡ = 1mm. Figs: 2b, 4b, 5b, 6b.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Plate 1
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Text-fig. 5: Schematic representation of relationships between ‘varieties’ of Hemiconus stromboides (Lamarck, 1802) proposed by PEZANT,
showing various combinations of characters involved (after PEZANT, 1910).
ONE, FOUR OR FORTY SPECIES ?
PEZANT (1910), studying about 500 small cones from
la Ferme de Launay (Parnes, Oise) noticed that the
basic shell characters of Hemiconus appeared to recom-
bine in various ways which he construed as represen-
ting a single species with two tiers of varieties, mostly
connected by intermediate forms (Text-fig. 5). This was
a plausible conclusion to reach in view of the large
sample examined. Faced with the same problem in our
study of >1000 Hemiconus from Cotentin localities we
have found it useful to arrange the material into 4
loosely defined groups of species with common charac-
teristics, typified by the species H. stromboides
(Lamarck, 1802), H. tromelini (Vasseur, 1882), H.
peraratus Cossmann, 1897 and H. tremletti Le Renard,
1994. Specimens occur with apparently intermediate
characters that might question the validity of these
groups, but it seems as likely that the apparently
continuous variation represents mixing of chronolo-
gically distinct morphologies within the stratigraphi-
cally condensed sequences present. Further stratigra-
phically detailed collection from the exposed sections
might reveal a more accurate picture of both the taxo-
nomy and the evolution of these cones. For that reason
we maintain the above as informal groups, pending
further research. PUILLANDRE et al. (2011), in their
molecular study of cryptic species of Conasprella living
in the Indo-Pacific region, showed that species,
separated at genetic level within a complex, demons-
trated only minor differences in shell morphology and
colour. By analogy, the 17 fairly well defined species of
Hemiconus that we have identified in the Cotentin may
well prove to be a conservative estimate of the total
fauna in view of the morphological variability encoun-
tered. It is possible that more than twice as many
species may be present in the stratigraphical interval
investigated.
Plate 2
Figs. 9-13 : Hemiconus stromboides (Lamarck, 1802) : 9. Neotype NHMUK PI OR 31452(1) (coll. DESHAYES),
Calcaire Grossier, Grignon (Yvelines) ; 10a-b. NHMUK PI TG 26846 (leg. B. CRAIG) Fresville ; 11. NHMUK PI
TG 26847 (leg. S. TRACEY) Fresville ; 12. (protoconch) NHMUK PI TG 26848 (leg. B. CRAIG) Néhou ; 13.
(protoconch) NHMUK PI TG 26849 (leg. S. TRACEY) Ferme de l’Orme.
Figs. 14-18 : Hemiconus cryptoconoides Cossmann & Pissarro, 1901 : 14. NHMUK PI TG 26850 (leg. S. TRACEY)
Néhou; 15. NHMUK PI TG 26851 (leg. B. CRAIG) Néhou ; 16. NHMUK PI TG 26852 (leg. O. GAIN) Néhou ; 17a-
b. NHMUK PI TG 26853 (leg. O. GAIN) Néhou ; 18. (protoconch) NHMUK PI TG 26854 (leg. S. TRACEY) Néhou.
Figs. 19-22 : Hemiconus trisulcatus Tracey & Craig, n. sp : 19. Paratype NHMUK PI TG 26855 (leg. B. CRAIG)
Hautteville ; 20a-b. Paratype NHMUK PI TG 26856 (leg. S. TRACEY) Hautteville ; 21. Paratype NHMUK PI TG
26857 (leg. B. CRAIG) Néhou ; 22a-c. Holotype (22c. protoconch) NHMUK PI TG 26858 (leg. S. TRACEY)
Hautteville.
Scale: = 1cm. Figs: 9,10a-b, 11, 14, 15, 16, 17a-b, 19, 20a-b, 21, 22a-b; I = 500µm Figs: 12, 13, 18, 22c.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
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Plate 2
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
This level of diversity is lent credibility by studies of
conid populations living in coral reef habitats (e.g.
KOHN, 1983) and a recent personal observation of the
fauna of Tiwi Beach in Kenya by one of us (BC) who
found 27 species of Conus (sensu PUILLANDRE et al.,
2015) living in shallow water of the lower shore within
a 200 m2 area of back reef with patches of sea-grass.
This suggests that the biodiversity of some present-
day Indo-Pacific tropical reef environments bear com-
parison with those observed within the Cotentin area
studied (approx. 30 km.2). We were unable to obtain
enough specimens that showed significant colour
patterns under ultraviolet light to be of use in differen-
tiating species, although lines of small dots were
observed on some forms in the H. peraratus group. This
could be a useful field for future study.
COTENTIN LOCALITIES
(for details see DUGUÉ et al., 2012; LE RENARD & GAIN,
2012)
Middle Eocene (late Lutetian) Faluns de Hautteville-
Bocage :
Fresville : Ferme de Vauville, Fresville (Manche)
[49°25'51.67"N 1°21'55.87"W]
Gourbesville : Hameau Beauvais, N. of Gourbesville
(Manche) [49°25'47.04"N 1°24'35.77"W]
Hautteville-Bocage : Château de Hautteville-Bocage
et ferme de la Basse Cour, Hautteville-Bocage
(Manche) [49°25'2.24"N 1°28'42.76"W]
Néhou : Fosse Launay, Néhou (Manche)
[49°25'15.38"N 1°33'11.79"W]
Middle or late Eocene (Bartonian or Priabonian) ‘Argiles
à corbules’ (VIEILLARD & DOLLFUS, 1875) :
Rauville : Rauville-la-Place (Manche)
[49°23'38.86"N 1°29'36.88"W]
ABBREVIATIONS
NHM London/ NHMUK : Natural History Museum,
London, U.K.
MNHN : Muséum National d’Histoire Naturelle, Paris,
France.
COTENTIN SPECIES COVERED
Family : Conidae Fleming, 1822
Genus : Hemiconus Cossmann, 1889
a) Group of H. stromboides
Hemiconus stromboides (Lamarck, 1802)
Hemiconus cryptoconoides Cossmann & Pissarro, 1901
Hemiconus trisulcatus Tracey & Craig n. sp.
Hemiconus disjunctus (Deshayes, 1865)
Hemiconus granatinus (Deshayes, 1865)
Hemiconus constantinensis Tracey & Craig n. sp.
b) Group of H. tromelini
Hemiconus turbinopsis (Deshayes, 1865)
Hemiconus tromelini (Vasseur, 1882)
c) Group of H. peraratus
Hemiconus scabriculus (Solander, 1766)
Hemiconus peraratus Cossmann 1897
Hemiconus douvillei Cossmann & Pissarro, 1901
Hemiconus gouetensis Cossmann, 1897
Hemiconus pissarroi Tracey & Craig n. sp.
d) Group of H. tremletti
Hemiconus tremletti Le Renard, 1994
Hemiconus lateralis Tracey & Craig n. sp.
Hemiconus auriculatus Tracey & Craig n. sp.
Hemiconus angulifer Cossmann & Pissarro, 1901
Genus: Papilliconus Tracey & Craig n. gen.
Papilliconus papillatus Tracey & Craig n. sp.
Papilliconus radulfivillensis Tracey & Craig n. sp.
Genus: Eoconus Tucker & Tenorio, 2009
Eoconus derelictus (Deshayes, 1865)
Eoconus bareti (Vasseur, 1882)
Eoconus diversiformis (Deshayes, 1835)
Eoconus veteratoris Tracey & Craig n. sp.
Eoconus cf. sulciferus (Deshayes, 1835)
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
SYSTEMATIC DESCRIPTIONS
Hemiconus
Cossmann, 1889
Type species by OD: Conus stromboides Lamarck, 1802.
« Hemiconus nov. sect. Coquille stromboide,
sillonnée, à tours non étagés, noduleux, à spire aiguë,
terminée par un gros bouton dévié ; labre très arqué,
faiblement échancré en arrière. Type : C. stromboides,
Lamk. » (COSSMANN, 1889, p. 234) [Shell stromboid,
grooved, whorls not stepped, nodular, with acute
spire ending in a large deviated button; lip very
arched, feebly notched posteriorly. Type : C.
stromboides, Lamk.] »
Emended diagnosis : Shell small, usually below
25 mm. in height, biconical with relatively tall spire
occupying 30-40% of shell height; protoconch typically
paucispiral of c. 1.75 whorls or, less commonly,
multispiral of up to 5.5 whorls. Teleoconch with a
beaded cord immediately below the suture, randomly
intersected by incised growth striae, 2-5 fine cords on
the upper part of the whorl and rounded or triangular
nodules on lower half of early whorls that often extend
onto later whorls, overridden by 2-4 spiral cords
which may carry regularly spaced granules; a pair of
granular cords on the shoulder commonly persists on
to the last whorl; sinus below the suture forming a
shallow curve on the whorl top; lip gently curved, not
strongly projecting.
Discussion : A paucispiral protoconch has previ-
ously been considered a fundamental character of
Hemiconus, however we show here that an indispu-
table Hemiconus, H. scabriculus (Solander, 1766) from
the Bartonian of England has a long multispiral proto-
conch and the same is probably true of H. macrocentrus
from the Lutetian of the Paris Basin (unpublished
data) and several early Eocene (Ypresian) species. The
teleconch whorls of the single Paleocene Hemiconus, H.
leroyi Tracey & Craig, n. sp., and the Ypresian species
bicoronatus Melleville, fallax Pacaud, concinnus
Sowerby, diastictus Cossmann, and rouaulti d’Archiac,
all share a similar morphology, being ornamented
with a row of beads immediately below the suture, a
row of larger tubercles on the periphery (i.e. shoulder),
2-4 spiral cords overriding the tubercles, and 1-3
weaker cords on the upper part of the whorl. In view
of their similar apical ontogeny, it seems reasonable to
include all of the earliest cones in Hemiconus (Plate 1,
Figs. 1-7).
a) Group of H. stromboides.
Species with relatively large protoconchs and low
tubercles centrally arranged on spire whorls, crossed
by spiral cords.
Hemiconus stromboides
(Lamarck, 1802)
Text-fig. 6 ; Plate 1, Fig. 7 ; Plate 2, Figs. 9-13 ; Plate 9,
Fig. 114.
Conus (stromboïdes), LAMARCK – 1802 – p. 387, pl. 7, fig. 2.
Conus stromboides, DESHAYES – 1835 – p. 749, pl. 98, figs. 15, 16.
Conus stromboides, COSSMANN – 1889 – p. 234.
Hemiconus stromboides, COSSMANN – 1896 – p. 150.
Conus stromboides var, PEZANT – 1910 – pl. 13, fig. 6.
Hemiconus stromboides, COSSMANN & PISSARRO – 1913 – pl. 48, fig. 214 bis 1.
Hemiconus stromboides, TUCKER & TENORIO – 2009 – p. 157. Pl. 11, fig. 6.
Text-fig. 6: Hemiconus stromboides
a, b. original figure after LAMARCK (1802) pl. 7, figs. 2a, b.
c, d. after DESHAYES (1835) pl. 98, figs. 15, 16.
Holotype of Hemiconus stromboides (Lamarck,
1802) : Not in LAMARCK collection, Geneva (CLERC &
FAVRE, 1910-1918) or MNHN Paris (J.-M. PACAUD
pers. comm.) ; presumed lost
Neotype of Hemiconus stromboides (Lamarck,
1802) : (designated herein): NHMUK PI OR 31452(1)
(coll. DESHAYES) Grignon, Natural History Museum,
London.
Type stratum and locality of H. stromboides :
Middle Lutetian, Calcaire Grossier, Grignon
(Yvelines) France.
Distribution of H. stromboides : Middle Lutetian
to Bartonian, Paris Basin and northern France.
Grignon, Beyne, Parnes (DESHAYES, 1835) ; Fresville,
Acy, Le Fayel (COSSMANN, 1889).
Original description of H. stromboides : “Conus
(stromboïdes). Testa subfusiformis, infernè et versùs apicem
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
sulcato-punctata; spirâ acutâ, obsoletè nodosâ: anfractibus
obtusis. n. L. n. Grignon. Il est petit, strié, par-tout
transversalement, et ressemble à un strombe à demi-formé.
Mon cabinet.” (LAMARCK, 1802 : p. 387) [It is small,
spirally striate all over, and resembles a half grown
Strombus].
Additional material examined of H. stromboides
: 20 lots from Grignon ; Villiers-Saint-Fréderic ; La-
Ferme-de-l’Orme ; Fresville ; Hautteville-Bocage ;
Néhou in NHM London and private collections.
Emended diagnosis of H. stromboides : Fusiform,
weakly spirally striate Hemiconus ; height : width ratio
2.4-2.9:1 ; spire c. 40% of shell height, ornamented with
obscure low rounded central tubercles crossed by 3-4
spiral cords with 2-3 weaker spiral threads above ;
beaded subsutural cord weakly present throughout
growth ; body whorl covered in very close, fine spiral
threads and profile slightly concave basally ; columel-
la straight ; aperture widens towards the base;
protoconch large, diameter c. 760 µm., paucispiral, of
1.75 whorls. Neotype 17.0 x 5.7 mm. Cotentin
specimens reach 21.0 x 8.2 mm.
Discussion of H. stromboides : LAMARCK’s (1802)
ambiguous original figure (Text-fig. 6 a-b) and
description are insufficient to define this species and
subsequent identification has generally been based on
DESHAYES’s (1835) more accurately drawn version
(Text-fig. 6 c-d). COSSMANN & PISSARRO (1907) figured
as H. stromboides a shell that seems very similar to their
figure of H. disjunctus (Deshayes, 1865). Confusion has
since arisen from differing interpretations of H.
stromboides featured on internet websites. Figures of
shells missing part of the outer lip (e.g. TUCKER &
TENORIO, 2009) appear more narrowly spindle-shaped
than when complete. For these reasons and in accor-
dance with ICZN (1999) Article 75.3, we consider it
advisable to designate a neotype. Specimens
corresponding to DESHAYES’s concept of H. stromboides
occur uncommonly in the Cotentin (Plate 2, Figs. 10-
12).
Hemiconus cryptoconoides
Cossmann & Pissarro, 1901
Text-fig. 7 ; Plate 2, Figs. 14-18 ; Plate 9, Fig. 118.
Hemiconus cryptoconoides, COSSMANN & PISSARRO – 1901 – p. 67, pl. 7, figs.
19, 20.
Hemiconus cryptoconoides, PEZANT – 1910 – p. 188.
Hemiconus cryptoconoides, LE RENARD & PACAUD – 1995 – p. 122.
Holotype of Hemiconus cryptoconoides
Cossmann & Pissarro, 1901 : Coll. de l’École des Mines
(COSSMANN & PISSARRO, 1901).
Type stratum and locality of H. cryptoconoides :
Late Lutetian, Faluns de Hautteville-Bocage, unit
FHB-A, Hautteville-Bocage (Manche).
Distribution of H. cryptoconoides :Hautteville-
Bocage, Fresville (COSSMANN & PISSARRO, 1901) ; Les
Vignettes [Hérouval] (PEZANT, 1909).
Text-fig. 7: Hemiconus cryptoconoides, original figure after COSSMANN
& PISSARRO (1901) pl. 7, figs. 19, 20. Hautteville.
Material examined of H. cryptoconoides : 25 ex.
from Cotentin sites in NHM London and in private
collections.
Diagnosis of H. cryptoconoides : Narrowly
fusiform, weakly spirally striate Hemiconus ; Height :
width ratio 3.0-3.4:1 ; spire almost 40% of shell height,
without tubercles but with slightly swollen spire
whorls, crossed by 3-4 spiral cords with 2-3 weaker
raised spiral threads on the ramp ; beaded subsutural
cord obscure or absent on later whorls ; body whorl
straight-sided and covered in somewhat spaced, fine
spiral threads ; columella straight; protoconch large,
diameter c. 700 µm., paucispiral, of 1.5 whorls with a
very shallow suture. Cotentin specimens reach 18.2 x
5.7 mm.
Discussion of H. cryptoconoides : PEZANT (1910)
considered that the variation shown by H.
cryptoconoides from Les Vignettes in the Paris Basin
was continuous with that of H. stromboides (Lamarck,
1802), and that the two were conspecific, not even
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
worthy of varietal distinction. Here we conserve
Cossmann’s name for this distinctively narrow
Hemiconus, its relationship with H. stromboides needing
further research.
Hemiconus trisulcatus
Tracey & Craig, n. sp
Plate 2, Figs. 19-22 ; Plate 9, Fig. 117.
Holotype of Hemiconus trisulcatus n. sp. :
NHMUK PI TG 26858 (leg. S. TRACEY), Hautteville-
Bocage
Type stratum and locality of H. trisulcatus n. sp.
: Late Lutetian, Faluns de Hautteville-Bocage, unit
FHB-A, Hautteville-Bocage (Manche).
Paratypes of H. trisulcatus n. sp.: NHMUK PI TG
26856 (leg. S. TRACEY), Hautteville-Bocage ; NHMUK
PI TG 26855 (leg. B. CRAIG), Hautteville-Bocage ;
NHMUK PI TG 26857 (leg. B. CRAIG), Néhou.
Distribution of H. trisulcatus n. sp. : Only known
from Hautteville-Bocage and Néhou (Manche).
Etymology of H. trisulcatus n. sp. : Latin adjective
referring to the three characteristic subsutural
grooves.
Diagnosis of H. trisulcatus n. sp. : Narrowly
fusiform Hemiconus with tall spire; whorls weakly
angular to flat sided with 3 punctate grooves below
the suture and fine spiral threads covering the
remainder of the shell.
Description of H. trisulcatus n. sp. : Small
narrowly turriform-fusiform shell ; spire 37-46% of
shell height ; Height : width ratio 3 - 3.5:1 ; protoconch
large, diameter c. 730 µm., paucispiral of 1.75 whorls ;
Early whorls flat sided with flush sutures, typically
ornamented with 7-8 low, rib-like nodules per whorl
which may be obscure or even absent ; later whorls
tending to develop a blunt angularity in mid-whorl
that becomes a rounded shoulder on the last whorl ;
subsutural collar beaded, often only weakly, and
underscored by 3 regularly spaced spiral grooves
which are made punctate by the raised growth lines
within them ; body whorl straight-sided and covered
in numerous close, fine spiral threads becoming more
prominent towards the base, crossed by almost
vertical growth striae ; sinus wide and very shallow,
outer lip gently curved ; aperture very narrow with
parallel sides ; columella straight, callus hidden from
view. Most angular form 20.5 x 6.9 mm., narrowest
14.6 x 4.1 mm.
Discussion of H. trisulcatus n. sp. : The more
acute, flat-sided early whorls, triple grooved
ornament, very close, fine spiral striae and tendency to
develop an angular whorl profile in some individuals
separates this from other slender Hemiconus such as H.
cryptoconoides Cossmann & Pissarro, 1901.
Hemiconus disjunctus
(Deshayes, 1865)
Text-fig. 8 ; Plate 3, Figs. 23-29 ; Plate 9, Fig. 119.
Conus disjunctus, DESHAYES – 1865 – p. 419, pI. 100, figs. 17-19.
Hemiconus disjunctus, COSSMANN & PISSARRO – 1901 – p. 63, pl. 7, fig. 15.
Type stratum and locality of Hemiconus
disjunctus (Deshayes, 1865) : Calcaire Grossier,
Grignon, Ully-Saint-George, Damery, Cumières.
Material examined of H. disjunctus : 15 lots from
Paris Basin and Cotentin localities in NHM London
and private collections.
Diagnosis of H. disjunctus : Small, narrowly
turriform Hemiconus with prominent, rounded tuber-
cles in mid-whorl on the spire, crossed by 3-5 spiral
cords, the uppermost tending to give the tubercles
sharp upper edges ; protoconch large, diameter c. 680
µm., paucispiral of 1.5-1.75 whorls, domed with
almost flush sutures ; last whorl straight to gently
concave in outline ; ornament of close spiral threads,
often obsolescent. Spire up to 43% of shell height. 16.7
x 5.9 mm.
Discussion of H. disjunctus : Like H. stromboides
(Lamarck, 1802) this species has central rounded
tubercles on the whorls but these persist further down
the spire and are sharper and more prominent in H.
disjunctus. It also resembles H. granatinus from which
it is differentiated by the narrower profile, stronger
tubercles and the spiral threads not being granulated.
The specimen of Hemiconus disjunctus (Deshayes, 1865)
whose colour pattern under UV light was illustrated
by CAZE et al. (2012, p. 46, pl. 20, fig F) appears to be
an example of H. parisiensis (Deshayes, 1865).
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Text-fig. 8: Hemiconus disjunctus, original figure after DESHAYES
(1865) pl. 100, figs. 17, 18, 19.
Hemiconus granatinus
(Deshayes, 1865)
Text-fig. 9 ; Plate 3, Fig. 30.
Conus granatinus, DESHAYES – 1865 – p. 419, pl. 100, figs. 22-23.
Hemiconus granatinus, COSSMANN & PISSARRO – 1901 – p. 64 [in part], pl. 7, fig.
13.
Hemiconus dumasi, COSSMANN & PISSARRO – 1901 – p. 64, pl. 7, fig. 18.
Type stratum and locality of Hemiconus
granatinus (Deshayes, 1865) : Calcaire Grossier
supérieur ; Sables moyens : Chambors, Caumont.
Distribution of H. granatinus : Western Paris
Basin : Trye-Château, Chambors, Caumont ; Cotentin
: Fresville, Néhou.
Material examined of H. granatinus : 5 lots from
Chambors and Fresville in NHM London and private
collections.
Diagnosis of H. granatinus : Small biconical
Hemiconus developing rounded tubercles on the
whorls, often becoming elongate on the last whorl and
crossed by 4 or more spiral cords ; protoconch large,
paucispiral of 1.75 whorls with a very shallow suture;
ramp with 3-4 spiral cords ; last whorl straight-sided
with rounded shoulder ; ornament of more than c. 25
granulated spiral threads.
Discussion of H. granatinus : Uncommon in the
Cotentin. COSSMANN & PISSARRO’s (1901, pl. 7) figure
of H. dumasi seems to show another example of H.
granatinus while one of the shells figured (fig. 14) as H.
granatinus (Deshayes, 1865) has the flat-sided whorls
that characterise H. peraratus Cossmann, 1897 and H.
douvillei.
Remark 1 Constantinus : Gallo--Romanic name of the region, derived from
that of the city of Coutances (Latin Constantia) linked to Gaius Flavius Valerius
Constantius Caesar, Roman emperor from 293 to 306 AD.
Text-fig. 9: Hemiconus granatinus
a, b. original figure of Conus granatinus after DESHAYES (1865) pl.
100, figs 22, 23.
c. NHMUK PI TG 26837(1) (coll. WRIGLEY) Chambors (Oise),
France
Hemiconus constantinensis
Tracey & Craig, n. sp.
Plate 3, Figs. 31-36 ; Plate 9, Fig. 116.
Holotype of Hemiconus constantinensis n. sp. :
NHMUK PI TG 26869 (leg. B. CRAIG), Natural History
Museum, London.
Type stratum and locality of H. constantinensis
n. sp. : Late Lutetian, Faluns de Hautteville-Bocage,
upper part of unit FHB-A, Fresville (Manche).
Paratypes of Hemiconus constantinensis n. sp. :
NHMUK PI G 15511(1-2) (coll. PISSARRO) Fresville;
NHMUK PI TG 26867, 26868 (leg. B. CRAIG) Fresville;
NHMUK PI TG 26870, 26871, 26872 (leg. S. TRACEY)
Néhou.
Distribution of H. constantinensis n. sp. :
Fresville and Néhou (Manche).
Etymology of H. constantinensis n. sp. : Taken
from the Latin name for the Cotentin region. [Remark 1].
Additional material examined of H.
constantinensis n. sp. : 5 examples in private
collections.
Diagnosis of H. constantinensis n. sp. : Small
Hemiconus with a large protoconch, stepped spire,
smooth upper whorls, bicarinate nodular periphery
and spaced, granulated spiral cords.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Plate 3
Figs. 23 – 29 : Hemiconus disjunctus (Deshayes, 1865) : 23. NHMUK PI TG 26859 (leg. B. CRAIG) Néhou ; 24.
NHMUK PI TG 26860 (leg. B. CRAIG) Néhou ; 25. NHMUK PI TG 26861 (leg. L. BELLIARD) Néhou ; 26. NHMUK
PI TG 26862 (leg. S. TRACEY) Néhou ; 27. [spire] NHMUK PI TG 26863 (leg. S. TRACEY) Néhou ; 28. [protoconch]
NHMUK PI TG 26864 (leg. S. TRACEY) Néhou ; 29. (protoconch) NHMUK PI TG 26865 (leg. S. TRACEY) Villiers-
St-Frédéric (Yvelines).
Fig. 30 : Hemiconus granatinus (Deshayes, 1865), NHMUK PI TG 26866 (leg. B. CRAIG) Fresville.
Figs. 31 – 36 : Hemiconus constantinensis Tracey & Craig, n. sp. : 31a-b. Paratype NHMUK PI TG 26867 (leg.
B. CRAIG) Fresville ; 32. Paratype NHMUK PI TG 26868 (leg. B. CRAIG) Fresville ; 33. Holotype NHMUK PI TG
26869 (leg. B. CRAIG) Fresville ; 34. Paratype NHMUK PI TG 26870 (leg. S. TRACEY) Néhou ; 35. Paratype
(protoconch) NHMUK PI TG 26871 (leg. S. TRACEY) Néhou ; 36. Paratype (protoconch) NHMUK PI TG 26872
(leg. S. TRACEY) Néhou.
Scale: = 1cm. Figs: 23, 24, 25, 26, 27, 30, 31a-b, 32, 33, 34; I= 500µm. Figs: 28, 29, 35, 36.
Plate 3
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Description of H. constantinensis n. sp. : Small
biconical shell, spire 34% of shell height; protoconch
large, c. 725 µm., paucispiral of 1.75 whorls ; whorl
profile concave above, smooth with faint growth striae
and a beaded subsutural collar ; rounded tubercles
below crossed and truncated by two prominent spiral
cords; tubercles persist to the last whorl where they
form a nodular biangular shoulder. Last whorl slightly
sinuous in profile, typically ornamented with 11
spaced, granulated spiral threads c. 9 plain spirals on
the base. Sinus wide and shallow, aperture narrow
with parallel sides ; inner lip slightly concave centra-
lly, outer lip gently curved. Up to 15.4 x 7.1 mm.
Discussion of H. constantinensis n. sp. : This is
broader than H. disjunctus (Deshayes, 1865) which also
lacks the granulated ornament. Differs from H.
granatinus (Deshayes, 1865) in the stepped profile, the
smooth and concave upper part of the whorls, the
biangular peripheral cords and by typically having
fewer granulated spirals on the last whorl ; Néhou
examples may have up to 30 spirals below the
biangular shoulder due to the development of
intermediary threads (Plate 3, fig. 34).
b) Group of H. tromelini
Species with broad, relatively smooth whorls and
peripheral areas, with obscure nodules, variably
developed.
Hemiconus turbinopsis
(Deshayes, 1865)
Text-fig. 10 ; Plate 4 Figs. 37-40 ; Plate 9 Fig. 113
Conus turbinopsis, DESHAYES – 1865 – p. 425, pl. 100, figs. 10-11.
Hemiconus turbinopsis, COSSMANN – 1896 – p. 152.
Hemiconus tromelini, COSSMANN & PISSARRO – 1901 – p. 65, pl. 7, fig. 21 (non
Vasseur, 1882).
Conus noduloso-turbinopsis mi-parti, PEZANT – 1910 – pl. 13, fig. 5.
Type stratum and locality of Hemiconus
turbinopsis (Deshayes, 1865) : Middle Calcaire
Grossier, Grignon, Chaussy, St-Félix, Hérouval.
Distribution of H. turbinopsis : Paris Basin and
Cotentin (Néhou, Gourbesville).
Material examined of H. turbinopsis : NHMUK
PI TG 26837 Mouchy (coll. DESHAYES) including 4
juvenile shells of dubious identity ; NHMUK PI TG
26873 Gourbesville (leg O. GAIN) ; Néhou (3 ex.) ; 6
additional shells from Néhou in private collections.
Remark 2 This is the largest species of Hemiconus from the Cotentin and the
first record of this species from the area.
Text-fig. 10: Hemiconus turbinopsis
a, b. original figure of Conus turbinopsis after DESHAYES (1865) pl.
100, figs 10, 11.
c. NHMUK PI TG 26837 (1)(coll. DESHAYES), Mouchy-le-Châtel
(Oise), France. (Scale bar 10 mm).
Diagnosis of H. turbinopsis : Relatively large,
ovately biconical Hemiconus without tubercles on the
adult whorls which are, however, slightly convex in
the lower part; subsutural cord divided into numerous
somewhat elongate beads ; spire 33% of shell height,
protoconch large, up to diameter 1 mm, paucispiral of
1.75 whorls with a very shallow suture ; spire profile
slightly convex (cyrtoconoid), shoulder obtusely
rounded, last whorl straight to mildly convex in adults
; ornament of spiral threads, usually weak or effaced,
but more marked in smaller shells. Columella more or
less straight above, concave below. Up to 25 x 11 mm
(DESHAYES, 1865); Cotentin specimens reach 25.2 x 11.2
mm. [Remark 2].
Hemiconus tromelini
(Vasseur, 1882)
Text-fig. 11 ; Plate 4 Figs. 41-47 ; Plate 9 Fig. 115.
Conus tromelini, VASSEUR – 1881 – p. 245 (nomen nudum).
Conus tromelini, VASSEUR – 1882 – pl. 2, fig. 31.
Hemiconus tromelini, COSSMANN – 1897 – p. 206, pl. 4, figs. 24, 25, 30.
Hemiconus tromelini, COSSMANN & PISSARRO – 1901 – p. 65, pl. 7, fig. 21.
Hemiconus lennieri, COSSMANN & PISSARRO – 1901 – p. 66, pl. 7, fig. 16-17.
Hemiconus tromelini, COSSMANN & VASSEUR – 1917 – pl. 2, fig. 31
Syntype of Hemiconus tromelini (Vasseur, 1882) :
MNHN.F.J03982.
Type stratum and locality of H. tromelini : Late
Lutetian or Bartonian, Le Bois-Gouët (Loire-
Atlantique).
Distribution of H. tromelini : Loire-Atlantique
and Cotentin.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Material examined of H. tromelini : NHMUK PI
G 15494(1) (coll. PISSARRO) Fresville; >30 ex. in NHM
London and private collections.
Diagnosis of H. tromelini : Small biconical
Hemiconus; spire 35-40% of shell height; subsutural
cord prominent with beads on earliest whorls, later
obscure or fused ; upper part of early whorls
comprising a narrow, shallow groove crossed by close,
regular growth striae followed by 2-3 spiral cords
which become obsolete on later whorls ; lower part of
spire whorls convex, often developing low rounded
tubercles which may persist to the shoulder ; shoulder
somewhat rounded on last whorl, peripheral area
smooth, rather polished and close spiral threads on the
base; slightly concave in outline ; protoconch large,
paucispiral, diameter c. 780 µm., of 1.75 whorls with a
very shallow suture; large examples reach 13.3 x 6 mm.
Discussion of H. tromelini : The original figures
of H. tromelini (Text-fig.11) from the Loire-Inférieure
all show a flat-sided or slightly concave spire and a
subangular shoulder. We have been unable to separate
these from Cotentin specimens referred to H. lennieri
Cossmann & Pissarro, 1901, or indeed from juvenile
shells in the NHM London recorded from Mouchy by
DESHAYES and referred to H. turbinopsis. COSSMANN
compared both H. lennieri Cossmann & Pissarro, 1901
and H tromelini (Vasseur, 1882) to H. defrancei
(Deshayes, 1865) of the Paris Basin.
Text-fig. 11: Hemiconus tromelini (Vasseur, 1882)
a. copy of original figure, after COSSMANN & VASSEUR (1917)
pl. 2, fig. 31 Le Bois-Gouët.
b. abnormal specimen, after COSSMANN & PISSARRO (1901) pl.
7, fig. 21. Fresville.
c, d. H. lennieri original figure after COSSMANN & PISSARRO (1901) pl.
7, figs. 16, 17. Hautteville.
c) Group of H. peraratus
Species with small protoconchs and spirally corded
whorls, the lowest cord being produced into flanges,
appearing as a double cord on the shoulder of the last
whorl.
Hemiconus scabriculus
(Solander, 1766)
Plate 4, Figs. 48-53.
Conus scabriculus, SOLANDER – 1766 – p. 15, pl. 1, fig. 21.
Conus lineatus, SOLANDER – 1766 – p. 15, pl. 1, fig. 22.
Conus corculum, SOWERBY (J. de C.) – 1841 – p. 27, pl. 623, figs. 8-9.
Conus scabriculus, EDWARDS – 1857 – p. 198, pl. 24, fig. 9.
Conus lineatus, EDWARDS – 1857 – p. 199, pl. 24, fig. 10.
Type stratum and locality of Hemiconus
scabriculus (Solander, 1766) : Bartonian, “…cliffs by
the sea coast, between Christ Church and Lymington,
but more especially about the cliffs by the village of
Hordwell…” (SOLANDER, 1766), i.e. including Barton,
Hampshire.
Distribution of H. scabriculus : (H. scabriculus s.
s.) Barton Clay and Becton Sand formations of Barton,
England ; (H. cf. scabriculus) Lutetian of Grignon, Paris
Basin (DESHAYES), Fresville, Cotentin, and ?
Bracklesham, England ; Bartonian of Monneville
(DESHAYES), Berville and Chavançon (D’ORBIGNY).
Material examined of H. scabriculus : Numerous
examples in NHM London and private collections
from Barton (Hants) U.K.
Diagnosis of H. scabriculus : Small biconical
Hemiconus, spire 30-35% of total shell height; body
whorl rather convex above, briefly concave at base.
Typical ornament of widely spaced, fine, sharp spiral
cords on the whorls, raised into regular, narrow,
lobate tubercles; two such cords appear on the spire
whorls, the lower resting in the suture or concealed by
it, and these become the bicarinate shoulder; last
whorl with c.6 cords below the shoulder and c.6
smoother cords on base, developing finer interme-
diary spiral threads between them with growth. Spiral
sculpture variable, extreme forms having numerous
closer, finer spiral cords throughout, sometimes
devoid of tubercles or granules, although this orna-
ment may occur at any time during growth; proto-
conch multispiral, maximum diameter c. 670 µm., of
5.5 convex whorls with deep sutures; initial embryonic
whorl c. 150 µm. diameter with a granular surface that
continues with the addition of a few oblique, then
linear, spiral microstriae, fading and becoming
smooth by the 4th protoconch whorl; larval lip with a
shallow sinus and a strongly projecting lower lobe.
Discussion of H. scabriculus : EDWARDS (1857)
followed SOLANDER (1766) in retaining both H.
scabriculus and H. lineatus as valid species, although he
noted their overlapping descriptions and recorded
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
multispiral protoconchs of up to 3 whorls in both. It is
probable that EDWARDS did not see a perfectly preser-
ved example with its 5.5 whorls, such as that shown
for the first time here (Plate 4, Fig. 50). Collections of
this species from Barton display a variable range of
ornaments including intermediate forms and those in
which the juvenile shell with lineatus-style ornament
develops the spaced, dentiform spirals of the typical
form on later whorls (Plate 4, Fig. 49). H. corculum
(Sowerby), although often recorded in Paris Basin
faunas (COSSMANN & PISSARRO, 1913; LE RENARD &
PACAUD, 1995), was correctly synonymised with H.
lineatus [= H. scabriculus] by EDWARDS (1857). The
holotype of H. corculum (Plate 4, Fig. 53) has a similar
matrix and preservation to shells from Barton, and its
locality data may be erroneous. Specimens from
Grignon, France with particularly spaced granular
spiral ornament have been referred to H. scabriculus in
the literature although the protoconch characters of
these examples have not yet been recorded. Here we
provisionally refer shells from Fresville with a few
more spiral cords than the typical form (Plate 4, Figs.
54-55) to H. scabriculus, awaiting protoconch details
from further material.
Hemiconus peraratus
Cossmann, 1897
Text-fig. 12 ; Plate 5, Figs. 56, 57, 59 – 67 ; Plate 9 Fig.
125.
Conus granatinus, VASSEUR – 1882 – pl. 2, fig. 29-30 (non Deshayes, 1865).
Hemiconus peraratus, COSSMANN – 1897 – p. 206 [70], pl. 4, figs. 8, 22 & 23.
Hemiconus peraratus, COSSMANN & VASSEUR – 1917 – pl. 2, figs. 29-30.
Holotype of Hemiconus peraratus Cossmann,
1897 : MNHN.F. J04871 (coll. COSSMANN).
Type stratum and locality of H. peraratus : Late
Lutetian, Coislin (Loire-Atlantique).
Distribution of H. peraratus : Loire-Atlantique
and Cotentin.
Additional material examined of H. peraratus :
NHMUK: PI G 15492 (coll. Pissarro) 5 ex ; PI G 58414
(coll. de GERVILLE, leg. SOWERBY) 1 ex ; >500 additional
specimens from Cotentin localities in private
collections.
Diagnosis of H. peraratus : Small biconical
Hemiconus with flat-sided whorls, spire 35% of total
height; below the beaded subsutural cord are up to 4
spiral cords increasing in strength downwards, the
lowest being the most prominent and produced into
14 or so narrow, dentiform nodules per whorl, more
distinct on the early whorls; underlying suture more
or less canaliculate; another similar cord emerges from
the suture on the last whorl to form a relatively
inconspicuous bicarinate shoulder; protoconch small,
diameter c. 500 µm., paucispiral of 1.75 whorls with an
impressed suture; last whorl straight-sided,
ornamented by numerous low, close, rounded spiral
cords, wider than their interspaces.
Plate 4
Figs. 37 – 40. Hemiconus turbinopsis (Deshayes, 1865) : 37. NHMUK PI TG 26873 (leg. O. GAIN) Gourbesville ;
38. NHMUK PI TG 26874 (leg. B. CRAIG) Néhou ; 39. NHMUK PI TG 26875 (leg. O. GAIN) Néhou ; 40a-b. NHMUK
PI TG 26876 (leg. B. CRAIG) Néhou.
Figs. 41 – 47. Hemiconus tromelini (Vasseur, 1882) : 41. H. lennieri Cossmann & Pissarro, 1901 NHMUK PI G
15494(1) (coll. PISSARRO) Fresville ; 42. NHMUK PI TG 26877 (leg. S. TRACEY) Néhou ; 43. NHMUK PI TG 26878
(leg. S. TRACEY) Hautteville ; 44. (protoconch) NHMUK PI TG 26879 (leg. B. CRAIG) Néhou ; 45. NHMUK PI TG
26880 (leg. S. TRACEY) Néhou ; 46. NHMUK PI TG 26881 (leg. S. TRACEY) Néhou ; 47. NHMUK PI TG 26882 (leg.
S. TRACEY) Hautteville.
Figs. 48 – 53. Hemiconus scabriculus (Solander, 1766) : 48a-b. Typical form, NHMUK PI TG 26883 (leg. S. TRACEY)
Barton, Bed H ; 49. Specimen showing transition from lineatus to scabriculus ornament, NHMUK PI TG 26884
(leg. S. TRACEY) Barton Bed H ; 50. (protoconch) Barton Bed H ; 51. ‘var.’ lineatus Solander, 1766)NHMUK PI TG
26886 (leg. S. TRACEY) Barton bed F ? ; 52. Conus lineatus Solander, 1766 NHMUK PI OR 71204(1) (coll. EDWARDS)
Barton ; 53. Conus corculum J. de C. Sowerby, 1841, Holotype NHMUK PI OR 71205 (coll. EDWARDS)
‘Bracklesham’.
Figs. 54 – 55. Hemiconus cf. scabriculus (Solander, 1766) : 54a-b. NHMUK PI TG 26887 (leg. B. CRAIG) Fresville ;
55. NHMUK PI TG 26835 (coll. PISSARRO, as H. douvillei) Fresville.
Scale: . = 1cm. Figs: 37, 38, 39, 40a-b; .. = 1cm. Figs: 41, 42, 43, 45, 46, 47; … = 1cm. Figs: 48a-b, 49, 51, 52, 53,
54a-b, 55; I = 500µm. Figs: 44, 50.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Plate 4
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Discussion of H. peraratus : This is the most
abundant Hemiconus in the Cotentin and variable in
height to width ratio. Typically the shoulder is some-
what angular but examples with a swollen periphery
occur (Plate 5, Fig. 65). In a narrower morph from
Néhou, the dentiform nodules are more pronounced,
the upper part of the whorls concave and initially
smooth and the spiral threads more distanced and
sometimes granulated (Plate 5, Figs. 66, 67). This
might well represent a different taxon except that it
seems to be linked to typical H. peraratus Cossmann,
1897 by a series of intermediate forms. Broader shells
with more or less granulated cords on the spire could
also be considered to intergrade with H. douvillei
Cossmann & Pissarro, 1901 (Plate 5, Figs. 63-64).
Text-fig. 12: Hemiconus peraratus Cossmann, 1897, original figure
after Cossmann (1897) pl. 4 figs. 22-23. Coislin (Loire-Atlantique)
Hemiconus douvillei
Cossmann & Pissarro, 1901
Text-fig. 13 ; Plate 5, Figs. 68–72.
Hemiconus douvillei, COSSMANN & PISSARRO – 1901 – p. 65, pl. 7, figs. 11-12.
Hemiconus granatinus, COSSMANN & PISSARRO – 1901 – p. 64 [in part], pl. 7, figs.
14 (non Deshayes).
Type stratum and locality of Hemiconus douvillei
Cossmann & Pissarro, 1901 : Late Lutetian,
Hautteville.
Distribution of H. douvillei : Hautteville-Bocage,
Fresville (COSSMANN & PISSARRO).
Material examined of H. douvillei : NHMUK PI G
15559 (coll. PISSARRO) Fresville; > 20 ex. from Cotentin
localities in NHM London and in private collections.
Diagnosis of H. douvillei : Small, broadly
biconical, thick-shelled Hemiconus with flat-sided
whorls, ornamented with close, regular, rounded
granules along the spiral striae on the last whorl below
a strong double cord around the shoulder, and 2-4
granular cords on the upper part of the whorls.
Discussion of H. douvillei : COSSMANN &
PISSARRO (1901, pl. 7) figure two Hautteville-Bocage
specimens of which their fig. 11 in particular closely
matches the present diagnosis. PISSARRO’s material at
NHM London consists of a similar specimen (Plate 5,
Fig. 68) and another more elongate shell with deeper
sutures and spaced spiral cords which we have
referred to H. cf. scabriculus (Plate 4, Fig. 55).
Text-fig. 13: Hemiconus douvillei original figure after Cossmann &
Pissarro (1901) pl. 7, figs 11, 12. Hautteville
Hemiconus gouetensis
Cossmann, 1897
Text-fig. 14 ; Plate 5, Figs. 58, 73–75 ; Plate 9, Fig. 122.
Hemiconus peraratus var. gouetensis, COSSMANN – 1897 – p. 207 [71], pl. 4, figs.
28 & 29.
Holotype of Hemiconus gouetensis Cossmann,
1897 : MNHN.F. J04871.
Type stratum and locality of H. gouetensis : Late
Eocene, Bartonian (Auversian), Le Bois-Gouët.
Distribution of H. gouetensis : Loire-Atlantique
and Néhou, Cotentin.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Material examined of H. gouetensis : NHMUK PI
TG 26903, 26904 (leg. O. GAIN) Néhou; PI TG 26905
(leg. B. CRAIG) Néhou; and c. 20 ex. in private
collections from Néhou.
Diagnosis of H. gouetensis :Small, narrowly
fusiform Hemiconus differing from H. peraratus not
only in relative width but also in the lack of nodular
development, the 2 prominent spiral cords on the
ramp and the deeply canaliculate sutures; subsutural
beads often obscure or absent; protoconch small,
diameter c. 485 µm., of 1.75 whorls; last whorl
ornament as in H. peraratus Cossmann, 1897. Size 12.5
x 5.2 mm.
Text-fig.14: Hemiconus gouetensis Cossmann, 1897, original figure of
H. peraratus var. gouetensis after Cossmann (1897) pl. 4. figs. 28, 29.
Coislin (Loire-Atlantique).
Discussion of H. gouetensis : COSSMANN regarded
this as a variety of H. peraratus Cossmann, 1897 and it
is possible that the two forms constitute a continuous
series. We have not been able to compare topotypical
examples from the Loire Valley and so we prefer to
treat COSSMANN’s gouetensis as a full species.
Hemiconus pissarroi
Tracey & Craig n. sp
Text-fig 15 ; Plate 5, Figs. 76 – 79 ; Plate 9 Fig. 126.
Conus (Conospira) lebruni, COSSMANN & PISSARRO – 1901 – p. 68, pl. 7, fig.
24 (non Deshayes, 1865).
Holotype of Hemiconus pissarroi n. sp. : NHMUK
PI TG 26906 (leg S. TRACEY), Natural History Museum,
London.
Type stratum and locality of H. pissarroi n. sp. :
Late Lutetian, Faluns de Hautteville-Bocage, unité 1
(DUGUÉ et al., 2012), Fresville (Manche), France.
Paratype of Hemiconus pissarroi n. sp. : PI G
15493(1) (coll. PISSARRO) Fresville; NHMUK PI TG
26907, 26908, (leg. B. CRAIG) Néhou.
Distribution of H. pissarroi n. sp. : Fresville,
Néhou
Etymology of H. pissarroi n. sp. : Named for
Georges Manzana PISSARRO, co-author with Maurice
COSSMANN on several classical palaeontological
works. Some of his Cenozoic Mollusca collection was
purchased by the Natural History Museum, London
in April 1904.
Diagnosis of H. pissarroi n. sp. : Small Hemiconus
with relatively long, rather smooth body whorl,
granulated cords on the spire and a single row of
sharp tubercles above the suture, emerging as a
double row on the last whorl.
Description of H. pissarroi n. sp. : Shell small,
biconical, spire 30% of shell height ; specimens exami-
ned eroded but protoconch small, paucispiral of less
than 2 whorls ; spire profile slightly concave, sutures
somewhat canaliculate, spire whorls flat-sided,
ornamented with a prominent beaded subsutural cord
followed by 3 finely beaded cords on the ramp and a
row of pointed tubercles resting on the lower suture
until the last whorl where a lower row of tubercles
emerges to form a narrow but pronounced double row
around the periphery ; body whorl straight-sided with
numerous close, faint spiral striae, coarser and stron-
ger on the base; aperture parallel-sided, paries with a
shallow depression at the top and a marked discon-
tinuity where it meets the short columella basally; anal
sinus shallow, lip gently curved. Up to 12.7 x 5.3 mm.
Discussion of Hemiconus pissarroi nov. sp. :
Despite a certain resemblance to some members of the
H. peraratus complex, this species is always easily
separated by eye, which is perhaps why COSSMANN &
PISSARRO (1901) considered it to belong to a separate
genus, Conus (Conospira). However, it is surprising
that they confused this species with Conus lebruni,
described and figured by DESHAYES (1865) and
(correctly) by themselves (COSSMANN & PISSARRO,
1913), which is a squat species with prominent square
tubercles, and is unlikely to be a Hemiconus. H. pissarroi
n. sp. most resembles H. peraratus Cossmann, 1897
from which it is distinguished by its prominent,
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
bisected tubercles on the shoulder, its more elongate
body whorl and finer, weaker and more numerous
spiral threads.
Text-fig. 15: Hemiconus pissarroi Tracey & Craig n. sp., as Conilithes
lebruni after COSSMANN & PISSARRO (1901) pl. 7, fig. 24, Fresville.
d) Group of H. tremletti
Species with prominent rounded nodules just above
the suture; spiral cords subordinate or absent until the
base.
Hemiconus tremletti
Le Renard, 1994
Text-fig. 16 ; Plate 6, Figs. 80-84 ; Plate 9 Fig. 123
Conus nodulosus, DESHAYES – 1865 – p. 416, pI. 100, figs. 24-26. (non G.B.
Sowerby, 1864).
Conus lineatus var. nodulosa, COSSMANN – 1889 – p.234. (non G.B. Sowerby,
1864).
Hemiconus tremletti, LE RENARD – 1994 – p. 38.
Hemiconus lineatus tremletti, LE RENARD & PACAUD – 1995 – p. 122.
Type stratum and locality of Hemiconus tremletti
Le Renard, 1994 : Calcaire Grossier, Grignon, Mouy.
Distribution of H. tremletti : Middle to late
Lutetian, Paris Basin (uncommon) and Cotentin
(common at Néhou, less so at Fresville and
Hautteville-Bocage).
Material examined of H. tremletti : >50 ex. from
the Cotentin in NHM London and private collections.
Diagnosis of H. tremletti : Small, biconical
Hemiconus, spire c. 37% of shell height; protoconch
Plate 5
Figs. 56, 57, 59-67. Hemiconus peraratus Cossmann, 1897 : 56. (protoconch) NHMUK PI TG 26888 (leg. B.
CRAIG) Néhou ; 57. (protoconch) NHMUK PI TG 26889 (leg. S. TRACEY) Néhou ; 59. (protoconch) NHMUK PI
TG26896 (leg. S. TRACEY) Fresville ; 60. NHMUK PI G 15492(1) (coll. PISSARRO) Fresville ; 61. NHMUK PI TG
26892 (leg. B. CRAIG) Néhou ; 62. NHMUK PI TG 26893(leg. B. CRAIG) Néhou ; 63. NHMUK PI TG 26894 (leg.
O. GAIN) Néhou ; 64. NHMUK PI TG 26895 (leg. O. GAIN) Néhou ; 65. NHMUK PI TG 26896 (leg. S. TRACEY)
Fresville; 66. NHMUK PI TG 26897 (leg. O. GAIN) Néhou ; 67. NHMUK PI TG 26898 (leg. B. CRAIG) Néhou.
Figs. 68-72. Hemiconus douvillei Cossmann & Pissarro, 1901 : 68. NHMUK PI G 15559 (coll. PISSARRO)
Fresville ; 69. NHMUK PI TG 26899 (leg. S. TRACEY) Néhou ; 70. NHMUK PI TG 26900 (leg. S. TRACEY) Néhou
; 71. NHMUK PI TG 26901 (leg. S. TRACEY) Néhou ; 72. NHMUK PI TG 26902 (leg. B. CRAIG) Fresville.
Figs. 58, 73-75. Hemiconus gouetensis Cossmann, 1897 : 58. (protoconch) NHMUK PI TG26890 (leg. B. CRAIG)
Néhou ; 73. NHMUK PI TG 26903 (leg. O. GAIN) Néhou ; 74. NHMUK PI TG 26904 (leg. O. GAIN) Néhou ; 75.
NHMUK PI TG 26905 (leg. B. CRAIG) Néhou.
Figs. 76-79. Hemiconus pissarroi Tracey & Craig, n. sp. 76. Holotype NHMUK PI TG 26906 (leg. S. TRACEY)
Fresville ; 77. Paratype NHMUK PI G 15493(1) (coll. PISSARRO as Conus lebruni) Fresville ; 78. Paratype
NHMUK PI TG 26907 (leg. B. CRAIG) Néhou ; 79. Paratype NHMUK PI TG 26908 (leg. B. CRAIG) Néhou
.
Scale: = 1cm. Figs: 60-79; I = 500µm. Figs: 56-59.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Plate 5
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
small, diameter c. 510 µm., paucispiral, of 1.75 whorls
with distinct sutures; teleoconch more or less smooth
with 10-12 prominent tubercles per whorl, central on
the first whorl but low on the remaining spire whorls,
typically resting on and partly obscured by the follo-
wing whorls; subsutural cord rather irregularly and
obscurely beaded; 2-4 spiral threads on the ramp; last
whorl straight-sided to slightly sinuous with close fine
cords becoming stronger basally. Size 11 x 5 mm.
(DESHAYES, 1865). Cotentin examples reach 11.4 x 5.2
mm.
Discussion of H. tremletti : Shell more distinctly
biconical and nodules smaller than the sharp tubercles
of H. disjunctus (Deshayes, 1865), from which it also
differs by its smaller protoconch.
Text-fig. 16: Hemiconus tremletti Le Renard, 1994, as Conus
nodulosus after DESHAYES (1865) pl. 100, figs. 24, 25, 26.
Hemiconus lateralis
Tracey & Craig, n. sp.
Plate 6, Figs. 85-86 ; Plate 9 Fig. 120.
Holotype of Hemiconus lateralis n. sp. :
NHMUK PI TG 26914 (leg. S. TRACEY). Natural
History Museum London).
Type stratum and locality of H. lateralis n. sp. :
Late Lutetian, Faluns de Hautteville-Bocage, upper
part of unit FHB-A, Fresville (Manche).
Paratypes of Hemiconus lateralis n. sp. : NHMUK
PI G 15573(1) (coll. PISSARRO);
Distribution of H. lateralis n. sp. : Only known
with certainty from the type locality.
Etymology of H. lateralis n. sp. : Latin adjective
lateralis, spreading sideways, i.e. broad.
Additional material examined of H. lateralis n.
sp. : 5 ex. in private collections.
Diagnosis of H. lateralis n. sp. : Small, broadly
biconical Hemiconus, with a smooth profile and
angular shoulder. Lacking subsutural beads.
Description of H. lateralis n. sp. : Shell small,
spire 32% of shell height ; protoconch large, diameter
645 µm., paucispiral of 1.75 whorls ; Spire whorls flat-
sided, slightly extraconic at first, with a narrow row of
nodules imbricated over the lower suture, and a more
or less smooth subsutural collar ; upper part of whorl
initially almost smooth, developing c. 5 weak, spiral
threads on later whorls ; peripheral nodules divided
by a central sulcus on the last whorl, where they dete-
riorate to form a weakly nodular shoulder overridden
by a double spiral cord. Last whorl covered in close,
rounded spiral cords becoming more prominent
towards the base. Sinus wide and shallow, aperture
narrow with parallel sides ; columella straight, outer
lip gently curved. Up to 10.8 x 5.4 mm.
Plate 6.
Figs. 80 – 84. Hemiconus tremletti Le Renard, 1994 : 80. NHMUK PI TG 26909 (leg. B. CRAIG) Néhou ; 81.
NHMUK PI TG 26910 (leg. B. CRAIG) Néhou ; 82. NHMUK PI TG 26911 (leg. B. CRAIG) Néhou ; 83. NHMUK
PI TG 26912 (leg. B. CRAIG) Néhou ; 84. (protoconch) NHMUK PI TG 26913 (leg. S. TRACEY) Néhou.
Figs. 85 – 86. Hemiconus lateralis n. sp., 85a-d. Holotype (85c protoconch, 85d apical) NHMUK PI TG 26914
(leg. S. TRACEY) Fresville ; 86. Paratype NHMUK PI G 15573(1) (coll. PISSARRO as Conus parisiensis) Fresville.
Figs. 87 – 88. Hemiconus angulifer Cossmann & Pissarro, 1901, 87. NHMUK PI TG 26915 (leg. B. CRAIG) Néhou
; 88a-b. (88b protoconch) NHMUK PI TG 26916 (leg. B. CRAIG) Néhou.
Figs. 89 – 90. Hemiconus auriculatus n. sp., 89a-c. Holotype (89b protoconch, 89c apical) NHMUK PI TG
26917 (leg. S. TRACEY) Rauville-la-Place ; 90a-b. Paratype (90b protoconch) NHMUK PI TG 26918 (leg. S.
TRACEY) Rauville-la-Place.
Scale: = 1cm. figs: 80, 81, 82, 83, 85a-b, 86, 87, 88a, 89a, 90a; = 2mm. figs: 85d, 89c; I = 500µm. figs: 84,
85c, 88b, 89b, 90b.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Discussion of H. lateralis n. sp.: The broad
biconical aspect of the shell, flattened whorl profile
and subdued ornament are quite different from most
Hemiconus and recall the shapes of some living
Conidae. However, the double perphiperal cord and
vestigial subsutural collar link it to Hemiconus. This
species most closely resembles H. tremletti Le Renard,
1994 but differs in having no beads on the subsutural
cord, a larger protoconch and a narrower row of nodu-
les placed lower on the spire whorls, becoming a
narrow double cord crossing the nodules on the last
whorl. It differs from H. auriculatus n. sp. by its smooth
profile, lower and less prominent nodules and greater
breadth.
Hemiconus auriculatus
Tracey & Craig, n. sp
.
Plate 6 Figs. 89-90 ; Plate 9 Fig. 121.
Holotype of Hemiconus auriculatus nov. sp. :
NHMUK PI TG 26917 (leg. S. TRACEY), Natural
History Museum, London.
Type stratum and locality of H. auriculatus n. sp.
: Late Eocene, ‘Argiles à corbules’ (VIEILLARD &
DOLLFUS, 1875), Rauville-la-Place (Manche).
Paratype of Hemiconus auriculatus n. sp. :
NHMUK PI TG 26918 (leg. S. TRACEY).
Distribution of H. auriculatus n. sp. : Recorded
only from the type locality and horizon.
Etymology of H. auriculatus n. sp. : Latin adj.
auriculatus, having ear-like protuberances, referring to
the appearance of the tubercles in apical view.
Additional material examined of H. auriculatus
n. sp. : 7 fragmentary examples in private collections.
Diagnosis of H. auriculatus n. sp. : Small
biconical Hemiconus, lacking subsutural beads, with
three spiral cords on the periphery of the last whorl,
the uppermost strongest and developing flat-topped,
semicircular tubercles.
Description of H. auriculatus n. sp. : Shell very
small, protoconch large, diameter c. 640 µm.,
paucispiral of a little less than 2 whorls, larval lip
slightly curved ; spire whorls stepped, subsutural
collar absent, upper part of whorls smooth or with 1-3
weak spiral threads; lower half of whorls dominated
by a prominent row of narrow tubercles, appearing as
semicircular lobes in apical view (Plate 6, Fig. 89c)
joined by two subordinate lower nodular cords on the
shoulder of the last whorl, which is ornamented with
numerous close weak spiral cords, increasing in
strength basally. Sinus wide and shallow, somewhat
inequilateral; aperture narrow with parallel sides;
columella straight, outer lip gently curved. Holotype
9.3 x 3.6 mm., largest paratype 7.5 x 3.2 mm.
Discussion of H. auriculatus n. sp. : This differs
from H. tremletti Le Renard, 1994 in having flatter
nodules, 3 peripheral cords and in lacking subsutural
beads. Conus plicatilis Von Koenen, 1890 from the early
Oligocene of Lattorf appears to be closely related but
differs in having a more angular sinus, emphasised by
strong, regular, raised growth striae covering the
upper part of the whorls, and only a single row of
more rounded shoulder nodules.
Hemiconus angulifer
Cossmann & Pissarro, 1901
Text-fig. 17 ; Plate 6, Figs. 87-88 ; Plate 9, Fig. 124.
Hemiconus angulifer, COSSMANN & PISSARRO – 1901 – p. 63, pl.7, fig. 8.
Type stratum and locality of Hemiconus angulifer
Cossmann & Pissarro, 1901 : Fresville.
Distribution of H. angulifer : Fresville and Néhou,
Cotentin.
Material examined of H. angulifer : NHM UK PI
TG 26915, 26916 (leg. B. CRAIG).
Text-fig. 17: Hemiconus angulifer, original figure after COSSMANN &
PISSARRO (1901) pl. 7, fig. 8. Fresville
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Diagnosis of H. angulifer: Moderately small,
turriform Hemiconus, spire 41% of shell height, with
small nodules on the first two whorls becoming a
central angular carina on lower whorls ; 2-3 strong
cords on initial whorls increasing in number on lower
spire whorls ; protoconch small, diameter c. 500 µm.,
paucispiral of 1.75 whorls ; sutures deep with a
strongly beaded subsutural collar. Up to 11.9 x 4.5
mm.
Discussion of H. angulifer : The small protoconch
and prominent sharp angle of the later whorls
differentiate this species from most other nodular
species of Hemiconus. The strong cords on the apical
whorls are not present on H. tremletti (Le Renard, 1994)
which also has prominent nodules rather than the
angulated whorls found on H. angulifer. This is a
poorly known species and further material is needed
to assess its variability.
Papilliconus
Tracey & Craig, n. gen.
Type species P. papillatus Tracey & Craig, n. sp..
Diagnosis : Small, up to 10 mm. in height, broadly
ovately biconical, height : width ratio 2.0-2.4:1, smooth
to faintly spirally striate shells, with some degree of
angulation on the whorl periphery ; sinus relatively
deep ; protoconch small, somewhat cylindrical compa-
red to the rounded shell profile, with a dispropor-
tionally large initial whorl ; colour pattern of c. 14
narrow, dark spiral bands.
Discussion : The type species of Papilliconus
appears to have few characters (apart from small size)
in common with any other contemporaneous genus,
and so a new name at generic rank is indicated.
Papilliconus differs from Hemiconus by its ovate shape,
mostly smooth surface, papillate protoconch and
deeper sinus, and in lacking a beaded collar and
peripheral nodules.
Papilliconus papillatus
Tracey & Craig, n. sp.
Plate 7 Figs. 91-95 ; Plate 9, Fig. 128.
Holotype of Papilliconus papillatus n. sp. :
NHMUK PI TG 26920 (leg. B. CRAIG), Natural History
Museum, London.
Type stratum and locality of P. papillatus n. sp. :
Late Lutetian, Faluns de Hautteville-Bocage, unit
FHB-A, Néhou (Manche).
Paratypes of Papilliconus papillatus n. sp. :
NHMUK PI TG 26919 (leg S. TRACEY) ; NHMUK PI TG
26921 26922, 26923 (leg. B. CRAIG).
Distribution of P. papillatus n. sp. : Known only
from the type locality.
Etymology of P. papillatus n. sp. : Latin adjective
papillatus, nipple-like, referring to the protoconch.
Additional material examined of P. papillatus n.
sp. : 18 examples in private collections.
Diagnosis of P. papillatus n. sp. : Very small,
ovately biconical, more or less smooth shell, with a
slight rounded angulation at the periphery ; spiral
colour bands moderately close and irregularly
spotted.
Description of P. papillatus n. sp. : Shell very
small, spire 38% of shell height ; protoconch small,
papillate, diameter 510 µm., paucispiral of 2 whorls ;
whorl profile straight or slightly concave above; first
teleoconch whorl with small obscure nodules which
fuse to form a bluntly subangular swelling over-
hanging the suture on later whorls, becoming obsolete
by the last whorl ; spire whorls more or less smooth
although some specimens show 2-3 weak spiral
grooves on the upper part ; last whorl with regular,
faint spiral threads which are seen to correspond to a
residual colour pattern, occasionally preserved, of c.
14 narrow brown bands on the body whorl bearing
rather irregular circular spots like a pearl necklace, the
band above the periphery being somewhat wider than
the others ; vestigial nodules on the spire whorls also
show dark coloration. Body whorl sinuous in outline ;
sinus rather deep, slightly inequilateral ; outer lip
broadly rounded, aperture lenticular ; columella
slightly concave. Holotype 8.5 x 3.5 mm., paratypes up
to 9.1 x 3.8 mm.
Discussion of P. papillatus n. sp. : The
protoconch of P. papillatus shows some resemblance to
that of H. lineatus sensu Cossmann & Pissarro (1913,
figs. 214 bis-3, non Solander, 1766) although shells of
the latter have the usual subsutural beads and spirally
corded ornament that characterise Hemiconus.
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Papilliconus radulfivillensis
Tracey & Craig, n. sp.
Plate 7 Figs. 96-99 ; Plate 9, Fig. 127.
Holotype of Papilliconus radulfivillensis n. sp. :
NHMUK PI TG 26924 (leg. S. TRACEY). Natural
History Museum London.
Type stratum and locality of P. radulfivillensis n.
sp. : Late Eocene, ‘Argiles à corbules’ (VIEILLARD &
DOLLFUS, 1875), Rauville-la-Place (Manche).
Paratypes of Papilliconus radulfivillensis n. sp. :
NHMUK PI TG 26925, 26926, 26927, 26928 (leg. S.
TRACEY).
Distribution of P. radulfivillensis n. sp. : Known
only from the type locality and horizon.
Etymology of P. radulfivillensis n. sp. : After the
village of Rauville-la-Place (Manche) where the
species was excavated. [Remark 3].
Additional material examined of P.
radulfivillensis n. sp. : Several examples in private
collections.
Diagnosis of P. radulfivillensis n. sp. : Very small
ovate, more or less smooth shell, with a single or
double keel at the periphery ; spiral colour bands
somewhat spaced.
Description of P. radulfivillensis n. sp. : Small
ovate-biconical shell, spire 35% of shell height ; proto-
conch small, papillate, diameter 620 µm., paucispiral
of c. 1.75 whorls ; whorl profile concave above with a
more or less prominent blunt keel below, becoming a
double cord of variable strength on the periphery of
the last whorl. Body whorl sinuous to slightly convex.
Residual colour pattern of a subsutural dark band and
2 peripheral bands ; Sinus moderately deep, some-
what inequilateral ; outer lip gently rounded, aperture
broad, lenticular ; columella straight. Large examples
reach 10.0 x 4.8 mm.
Discussion of P. radulfivillensis n. sp. : This
differs from P. papillatus n. sp. by the spiral carination
and the shallower sinus and less convex lip. We also
figure a specimen (Plate 7 fig. 100) from the same
locality which lacks a keel or cord and shows only a
blunt angulation of the periphery, differing from P.
papillatus n. sp. in its broader, somewhat biconical
shape. This may represent a variation of P.
radulfivillensis n. sp., to which we refer it with some
doubt until a larger range can be examined.
Plate 7
Figs. 91 – 95. Papilliconus papillatus Tracey & Craig, n. gen., n. sp. : 91a-b. Paratype (91b–protoconch) NHMUK PI
TG 26919 (leg. S. TRACEY) Néhou; 92a-c. Holotype (92b with residual colour pattern enhanced) NHMUK PI TG 26920
(leg. B. CRAIG) Néhou ; 93. Paratype NHMUK PI TG 26921 (leg. B. CRAIG) Néhou ; 94. Paratype NHMUK PI TG
26922 (leg. B. CRAIG) Néhou ; 95. Paratype (apical) NHMUK PI TG 26923 (leg. B. CRAIG) Néhou.
Figs. 96 – 99. Papilliconus radulfivillensis Tracey & Craig, n. sp. : 96a-c.
Holotype (96b protoconch. 96c apical)
NHMUK PI TG 26924 (leg. S. TRACEY) Rauville-la-Place ; 97. Paratype NHMUK PI TG 26925 (leg. S. TRACEY)
Rauville-la-Place ; 98. (with residual colour pattern enhanced) Paratype NHMUK PI TG 26926 (leg. S. TRACEY)
Rauville-la-Place ; 99. Paratype NHMUK PI TG 26927 (leg. S. TRACEY) Rauville-la-Place.
Fig. 100. Papilliconus cf. radulfivillensis n.
sp., variant without peripheral cord, NHMUK PI TG 26928 (leg. S.
TRACEY) Rauville-la-Place.
Scale: = 1cm. Figs: 91a, 92a-b, 93, 94, 96a, 97, 98, 99, 100; ‡= 1mm. Figs: 95, 96c; I = 500µm. figs:
91b, 92c,
96b.
Remark 3 RAUVILLE: formerly known under the names Rodulfi villa at
about the year 1000, Radulfivilla in 1042 and Radulphivilla in 1332. The
place-name derives from a germanic anthroponym such as Radulf or
Radulfus and from old French ville in its original sense of « rural domain »,
from the Latin villa. (Source WIKIPEDIA).
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Plate 7
Eoconus
Tucker & Tenorio, 2009
Type species: Conus sauridens Conrad, 1833.
Diagnosis : « Shell is conical to elongated conical
in shape ; nodules may persist or become obsolete ; a
dentiform plait is well developed...; whorl tops have
cords; the anal notch is deep; the protoconch is
multispiral. » (TUCKER & TENORIO, 2009, p.142).
Remarks : To this diagnosis may be added: shell
relatively large, height up to 65 mm. ; spire initially
extraconic, becoming turreted, low or flat ; nodules
generally small and sharp ; body whorl more or less
straight sided with well-marked carinate or subca-
rinate shoulder and spiral cords mainly confined to
the base. The great similarity between the North
American E. sauridens (Conrad, 1833) and the contem-
poraneous E. diversiformis (Deshayes, 1835) at all
stages of growth, makes this the most appropriate
genus for both species, as noted by TUCKER & TENORIO
(2009). No single character of Eoconus is sufficient to
assign species to, or exclude them from any extant
groups of cones. The dentiform plait is a non-
diagnostic character, as is the multispiral protoconch
(see discussion above). The early teleoconch whorls of
E. diversiformis (Deshayes, 1835), E. derelictus
(Deshayes, 1835) and also E. deperditus (Bruguière,
1792) from the Paris Basin are ornamented with a
central row of nodules, spiral cords and raised growth
striae. In E. bareti (Vasseur, 1882) and E. veteratoris n.
sp. however, the nodules are weakly developed at first
and soon become fused into a blunt carina, while
spiral cords are only developed at a later stage. This
may be related to the diminishing protoconch size of
the latter two species. Since the genus Conus is no
longer appropriate for Eocene cones we use Eoconus as
a genus of convenience whose limits are yet to be
determined.
Eoconus derelictus
(Deshayes, 1865)
Text-fig 18 ; Plate 8, Figs. 101-103 ; Plate 9, Fig. 129.
Conus derelictus, DESHAYES – 1865 – p. 422, pl. 100, figs. 1, 2.
Conus (Leptoconus) derelictus, COSSMANN & PISSARRO – 1913 – pl. 48, fig. 214-
9.
Conus (Leptoconus) derelictus, LE RENARD & PACAUD –1995 – p. 63, pl.7, fig. 8.
Conus derelictus, LE RENARD & GAIN – 2012 – p. 53, fig. 48e.
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Text-fig. 18: Eoconus derelictus (Deshayes, 1865), original figure of
Conus derelictus after Deshayes (1865) pl. 100, figs. 1, 2.
Type stratum and locality of Eoconus derelictus
(Deshayes, 1865) : Calcaire Grossier: Chaussy,
Grignon.
Distribution of E. derelictus : Middle Lutetian,
central Paris Basin; late Lutetian, Néhou, Hautteville-
Bocage.
Material examined of E. derelictus : 3 lots from
Hautteville-Bocage and Néhou in NHM London; 35
additional specimens in private collections.
Diagnosis of E. derelictus : Moderately large,
turreted cone with a regularly uncoiling spire, early
whorls with c. 3 distinct cords that persist to adul-
thood and rather sharp tubercles on the shoulder ;
protoconch of 2.5-3 whorls ; body whorl typically with
close, thick, rounded spiral cords, developed to a
greater or lesser extent. Protoconch large, multispiral,
diameter c. 860 µm., of 2.5-3 whorls with a shallow
suture ; larval shell smooth but ending with several
fine orthocline growth striae. Large shells reach 52 x
32 mm (Deshayes, 1865). [Remark 4].
Eoconus bareti
(Vasseur, 1882)
Text-fig 19 ; Plate 8, Figs. 104-106 ; 132, Plate 9, Fig.
132.
Conus bareti, VASSEUR – 1881 – p. 245 (nomen nudum).
Conus bareti, VASSEUR – 1882 – pl.2, figs. 57, 59-62.
Conus bareti, COSSMANN – 1897 – p. 208, pl. 4, figs. 20-21.
Conus bareti, COSSMANN & PISSARRO – 1901 – p. 68, pl. 7, fig. 28.
Conus bareti, COSSMANN & VASSEUR – 1917 – pl. 2, figs. 57, 59-62.
Remark 4E. derelictus (Deshayes, 1865) is more common in the Cotentin
than in the Paris Basin. Shells from Néhou tend to have reduced spiral
ornament on the last whorl.
Syntypes of Eoconus bareti (Vasseur, 1882) :
MNHN.F.J03959, J03960.
Type stratum and locality of E. bareti : Late
Eocene, Bartonian (Auversian), Le Bois-Gouët.
Distribution of E. bareti : Loire-Inférieure and
Cotentin, Le Bois-Gouët, Coislin, Arthon, Fresville,
Hautteville-Bocage.
Material examined of E. bareti : 9 lots from
Fresville and Hautteville-Bocage in NHM London and
private collections.
Text-fig. 19: Eoconus bareti (Vasseur, 1882), as Conus bareti after
COSSMANN (1897) pl. 4, figs. 20, 21. Le Bois-Gouët.
Diagnosis of E. bareti : Moderately large, turreted
cone with a regularly uncoiling spire ; earliest whorls
smooth with sinuous growth lines and weakly deve-
loped nodules on the shoulder which are usually
fused to form a thick, rounded central keel ; body
whorl with fine spaced spiral cords and c. 3 finer
threads in the interspaces, shoulder ramp developing
3 or more spiral cords on later whorls ; protoconch
small, multispiral, diameter c. 560 µm., of c. 2 whorls
with a deep suture and a shallow sinus in the larval
lip. Up to 34 x 17 mm. (Cossmann, 1807); Cotentin
specimens up to 48 x 23 mm. [Remark 5]
Eoconus diversiformis
(Deshayes, 1835)
Text-fig 20 ; Plate 8, Figs. 110-111 ; Plate 9, Fig. 130.
Conus diversiformis, DESHAYES – 1835 – p. 747, pl. 98, fig. 9.
Conus diversiformis, COSSMANN – 1889 – p. 236.
Conus (Lithoconus) diversiformis, COSSMANN & PISSARRO – 1901 – p. 68, pl. 7,
figs. 25, 26.
Eoconus diversiformis, CAZE & al. – 2012 – p. 46, pl.20, fig 3.
Remark 5The small protoconch and the fused ridge on the spire whorls
readily distinguish juvenile specimens of E. bareti from E. derelictus.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Type locality of Eoconus diversiformis
(Deshayes, 1835) : Chaumont, les Groux, Brasles,
Parnes, Mouchy, Saint-Félix, Chaussy, Grignon,
Gomerfontaine, Montmirel, Auvers, Valmondois,
Mary, Jaignes, Betz, Acy, Caumont; La Palarea;
Bracklesham (England).
Distribution of E. diversiformis : Middle to late
Lutetian, central and western Paris Basin and the
Cotentin.
Text-fig. 20: Eoconus diversiformis (Deshayes, 1835)
a. original figure of Conus diversiformis after DESHAYES
(1835) pl. 98, fig. 11.
b. NHMUK PI TG 26836 (coll. WRIGLEY), Chambors (Oise),
France.
Material examined of E. diversiformis : NHMUK
PI TG 26836 Chambors (coll. WRIGLEY); NHMUK PI G
15495(1) Fresville (coll. PISSARRO).
Diagnosis of E. diversiformis : Moderately large,
smooth cone with a coeloconoid, rapidly uncoiling
spire becoming regularly turreted or flattened when
fully grown, rather sharp tubercles on the shoulder of
early whorls and a multispiral protoconch of 2.25
whorls.
Discussion of E. diversiformis : This has a great
resemblance to E. sauridens (Conrad), type species of
Eoconus, differing principally in having a shorter
protoconch. SOWERBY’s (1841) record of Conus
diversiformis from Bracklesham was later referred to
his new species Conus diadema by Edwards (1857).
CAZE et al. (2012, pl. 20, fig. 3) revealed the UV colour
pattern of E. diversiformis and remarked on its
similarity to that of E. deperditus (Bruguière, 1792).
Eoconus veteratoris
Tracey & Craig, n. sp.
Plate 8, Figs. 107-109 ; Plate 9, Fig. 131.
Holotype of Eoconus veteratoris n. sp. : NHM PI
TG 26936, (leg. S. TRACEY) Natural History Museum,
London.
Type stratum and locality of E. veteratoris n. sp.
: Bartonian-Priabonian, ‘Argiles à corbules’ (Vieillard
& Dollfus, 1875), Rauville-la-Place (Manche)
Paratypes of Eoconus veteratoris n. sp. : NHM PI
TG 26935,26937 (leg. B. CRAIG), Natural History
Museum, London.
Distribution of E. veteratoris n. sp. : Recorded
only from the type locality and horizon.
Etymology of E. veteratoris n. sp.: Latin adjective,
veteratoris, of the wise old fox. Named in honour of
Jacques LE RENARD, distinguished palaeontologist and
pioneer of research on the type horizon.
Additional material examined of E. veteratoris n.
sp. : 10, mostly fragmentary examples in private
collections.
Diagnosis of E. veteratoris n. sp. : Moderately
small, rather smooth conical shell with sharp shoulder
with small rounded nodules, a low, somewhat
coeloconoid, more or less nodular spire and a
paucispiral protoconch.
Description of E. veteratoris n. sp. : Protoconch
diameter c. 730 µm., paucispiral, of 2 smooth, convex
whorls with a sinuous lip ; initial whorl globular,
diameter 300 µm.. Spire profile somewhat coelo-
conoid, flattening and becoming more regularly turre-
ted later ; spire whorls ornamented above with raised
growth lines tracing a deep semicircular sinus and 4-5
subordinate fine spiral cords ; lower part of whorls
with c. 12 short, rounded tubercles resting on the
lower suture, which may become fused with growth
to form an angular shoulder. Last whorl slightly
sinuous in profile, smooth but with fine spiral cords
towards the base. Aperture parallel sided,
columella/paries junction shallowly indented Outer
lip slightly flared, gently curved. Holotype 30.5 x 13
mm.
Discussion of E. veteratoris n. sp. : Similar in
general appearance to E. diversiformis (Deshayes, 1835)
but distinguished from that and other known Eoconus
and Conilithes species by the shorter protoconch, and
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
from Conilithes by the spiral ornament on the whorl
tops.
Eoconus cf. sulciferus
(Deshayes, 1835)
Text-fig. 21; Plate 8, Figs. 112 a, b.
cf. Conus sulciferus, DESHAYES – 1835 – p. 748.
cf. Conus sulcifer, DESHAYES – 1835 – pl. 98, fig. 3-4.
cf. Conus sulciferus, DESHAYES – 1865 – p. 417.
Type stratum and locality of Eoconus cf.
sulciferus (Deshayes, 1835) : Late Eocene, Sables
moyens, Monneville, Ver, le Guépelle.
Distribution of E. cf. sulciferus : Late Lutetian in
the Cotentin and Bartonian of the northwestern Paris
Basin.
Material examined of E. cf. sulciferus :NHMUK
PI TG 26938 (leg. L. BELLIARD) Néhou.
Diagnosis of E. cf. sulciferus : Rather small, thick-
shelled Conus with low spire, whorls with close,
rounded tubercles persisting to the shoulder of the last
whorl, which is ornamented with numerous strong,
close, raised spiral threads. [Remark 6].
Text-fig. 21: Eoconus sulciferus (Deshayes, 1835), original figure of
Conus sulcifer after DESHAYES (1835) pl. 98, figs. 3, 4.
Plate 8
Figs. 101 - 103. Eoconus derelictus (Deshayes, 1865) : 101a-b. NHMUK PI TG 26929 (leg. S. TRACEY) Hautteville
; 102. NHMUK PI TG 26930 (leg. L. BELLIARD) Néhou ; 103. NHMUK PI TG 26931 (leg. S. TRACEY) Néhou ;
Figs. 104 -106. Eoconus bareti (Vasseur, 1882) : 104a-b. NHMUK PI TG26932 (leg. S. TRACEY) Hautteville ; 105.
NHMUK PI TG 26933 (leg. B. CRAIG) Hautteville ; 106. NHMUK PI TG 26934 (coll. D. CURRY ; leg. S. TRACEY)
Hautteville;
Figs. 107 - 109. Eoconus veteratoris Tracey & Craig, n. sp. : 107. Paratype NHMUK PI TG 26935 (leg. B. CRAIG)
Rauville-la-Place ; 108. Holotype NHMUK PI TG 26936 (leg. S. TRACEY) Rauville-la-Place ; 109a-b. Paratype
(109a protoconch, 109b apical) NHMUK PI TG 26937 (leg. B. CRAIG) Rauville-la-Place ;
Figs. 110 – 111. Eoconus diversiformis (Deshayes, 1835) : 110. NHMUK PI G 15495(1) (coll. PISSARRO) Fresville
; 111. NHMUK PI TG 26836 (coll. WRIGLEY) Chambors ;
Figs. 112a-b. Eoconus cf. sulciferus (Deshayes, 1835) NHMUK PI TG 26938 (leg. L. BELLIARD) Néhou.
Scale: = 1cm. Figs: 101a-b, 104a-b, 107, 108, 112; = 5mm. Figs: 102, 105, 110; ‡= 1mm. Fig: 109b;
I = 500µm. Figs: 103, 106, 109a, 111
Remark 6 A similar shell was recorded from Bois-Gouët as “Hemiconus sp.
voisin de Conus sulciferus Desh.” by COSSMANN & VASSEUR (1917, pl. 2, fig.
56.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Plate 8
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
CONCLUSIONS
The earliest confirmed Conidae appear to be from the
Paleocene of France with a possible ancestor in the
Cretaceous. The earliest conid species has all the
characters of Hemiconus. By the late Ypresian two
clades are clearly distinguished, typified by Eoconus
and Hemiconus. A study of the abundant conid faunas
in late Lutetian strata of the Cotentin has shown that
the few characters of shell morphology possessed by
cones can appear in multiple combinations, confusing
differences between species. Whereas the species of
Eoconus are rather easily separated, the
interrelationships of species of Hemiconus present
many possible interpretations. It seems that a division
into four informal groups (which may increase with
further research) is useful for classifying the Lutetian
Hemiconus of this area. However, the precise
relationship of Paleogene to Recent cones is not
conclusively supported by similarities in their few
sculptural characters, and a concentrated study of
morphology of shells from the Neogene would
perhaps solve some of these problems.
ACKNOWLEDGEMENTS
We would like to thank Gérard BARBE, Jean-Pierre
BOUILLON, Maurice GUILLIOU, Allan LAWSON, Gilles
LELONG, William POCOCK, Christian ROMANEK and
Malcolm SYMONDS for assistance and the donation of
specimens; our gratitude also to Alex BALL, Tomasz
GORAL and Bruno CASANOVA (NHM London) for help
with SEM imaging and photography, Diana CLEMENTS
(NHM London) for curation and reviewing the
manuscript, John K. TUCKER (Illinois Natural History
Survey), Christopher GARVIE (University of Texas,
Austin) and Jacques LE RENARD for useful discussion;
our particular thanks to Jon TODD (NHM London)
whose critical review considerably improved the
paper.
Plate 9
Apical views showing relative protoconch size
Fig. 113. Hemiconus turbinopsis, Néhou ; Fig. 123. H. tremletti, Néhou ;
Fig. 114. H. stromboides, La Ferme de l’Orme ; Fig. 124. H. angulifer, Néhou ;
Fig. 115. H. tromelini, Néhou ; Fig. 125. H. peraratus, Fresville ;
Fig. 116. H. constantinensis, Néhou ; Fig. 126. H. pissarroi, Néhou ;
Fig. 117. H. trisulcatus, Hautteville ; Fig. 127. Papilliconus radulfivillensis, Rauville ;
Fig. 118. H. cryptoconoides, Néhou ; Fig. 128. P. papillatus, Néhou ;
Fig. 119. H. disjunctus, Néhou ; Fig. 129. Eoconus derelictus, Néhou ;
Fig. 120. H. lateralis; Fresville ; Fig. 130. E. diversiformis, Chambors ;
Fig. 121. H. auriculatus, Rauville ; Fig. 131. E. veteratoris, Rauville ;
Fig. 122. H. gouetensis, Néhou ; Fig. 132. E. bareti, Hautteville.
Scale: I = 500µm.
One, four or forty species ? – early Conidae that led to a radiation and biodiversity peak in the late
Lutetian Eocene of the Cotentin, NW France.
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
Plate 9
S.TRACEY, B. CRAIG, L. BELLIARD & O .GAIN
Carnets de Voyages Paléontologiques dans le bassin Anglo-Parisien. Tome 3. Octobre 2017
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APPENDIX
Hemiconus leroyi
Tracey & Craig, n. sp.
Plate 1, Fig. 1a, b, c.
Hemiconus bicoronatus (Mellv.), COSSMANN – 1913 – p. 191 (non Melleville,
1843).
Hemiconus bicoronatus (Melleville), TRACEY et al.,- 1993 - p. 152, fig. 8.5 (non
Melleville, 1843).
Hemiconus sp., LEROY et al.– 2014 – pp. 25, 108, pl. 28, figs. 15, 16.
Holotype of Hemiconus leroyi n. sp. : NHMUK PI
TG 26839 (coll. C. ROMANEK, leg. S. TRACEY).
Type stratum and locality of H. leroyi n. sp. :
Paleocene (mid Thanetian), Châlons-sur-Vesle
Formation, niveau 4 of LEROY et al. (2014), Cauroy-lès-
Hermonville (Marne) France.
Distribution of H. leroyi n. sp. : Known only from
the type locality and from Jonchery-sur-Vesle
(COSSMANN, 1913) where a similar horizon was formerly
exposed.
Etymology of H. leroyi n. sp.: Named in honour of
Arnaud LEROY, discoverer of the type locality and the
first to illustrate an example of this species.
Additional material examined of H. leroyi n. sp. : 3
ex. in private collections.
Description of H. leroyi n. sp. : Moderately small,
narrowly biconical shell, spire c. 32% of shell height ;
whorl profile slightly convex with 8-9 large, low
rounded tubercles crossed by 2 spiral cords and a promi-
nent and strongly beaded subsutural collar with 3-4
finer spiral threads below it. Tubercles persist to the last
whorl where they form a broad, rounded, nodular
shoulder. Last whorl covered in close, fine spiral
threads. Sinus wide and shallow, aperture narrow with
parallel sides ; paries long and straight, columella short.
Holotype 14.1 x 6.1 mm. The larger example figured by
LEROY et al. (2014) is c. 2.25 x 10.0 mm.
Discussion of H. leroyi n. sp. : Although COSSMANN
first recorded what was probably this species in 1913,
that record has not been substantiated and has been
overlooked since. Subsequently the earliest conids have
long been considered to date from the early Eocene
(middle Ypresian) (KOHN, 1990).
Although the whereabouts of the specimen recorded by
COSSMANN is unknown, it is assumed, in view of the
rarity of cones in the Paleocene, that this was in fact
another example of H. leroyi n. sp.. Recently, LEROY et al.
(2014) figured this species and observed that it reached
about twice the size of the Ypresian H. bicoronatus
(Melleville, 1843) but they did not formally describe it as
new. H. leroyi n. sp. differs from H. bicoronatus<