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RESEARCH PAPER
Europrotomus (Mollusca: Caenogastropoda: Strombidae): a new
Middle Miocene European strombid genus (Revision
of Euprotomus Gill, 1870. Part 4)
Gijs C. Kronenberg
•
Mathias Harzhauser
Received: 15 March 2011 / Accepted: 12 September 2011
Springer-Verlag 2011
Abstract The medium-sized, strongly ornamented nom-
inal species Strombus schroeckingeri Ho
¨
rnes in Hoernes &
Auinger, 1884 is well known from Langhian deposits of the
Paratethys Sea. Its traditional generic affiliation in the
literature with Euprotomus Gill, 1870, implicated a
biogeographic relation of the Paratethys and the proto-
Mediterranean with the Neogene Indo-West-Pacific region.
This relation, however, is problematic because the Tethys
Seaway was already closed or a very shallow, strongly
evaporitic passage at that time. Despite its superficial
similarity with extant genera of the IWP region, the genus
is unrelated to all known strombid genera and represents a
new genus, which is introduced herein as Europrotomus
nov. gen. The genus comprises only one or maybe two
species, which occur in the European Middle Miocene
geological record. No ancestors are known so far. This
sudden occurrence is discussed as immigration from an
adjacent bioprovince rather than as a result of autochtho-
nous evolution. Hypothetically, such a biogeographic
source area might have been established along the coasts of
tropical West Africa—a scenario that is already docu-
mented for Pleistocene times.
Keywords Gastropoda Strombidae Euprotomus New
genus Miocene Biogeography Paratethys
Kurzfassung Die nominale Art Strombus schroeckingeri
Ho
¨
rnes in Hoernes & Auinger, 1884 ist ein stark orna-
mentierter Strombidae von mittlerer Gro
¨
ße, der aus dem
Langhium der Paratethys gut dokumentiert ist. Die tradi-
tionelle generische Zuordnung zu Euprotomus Gill, 1870
implizierte eine biogeographische Beziehung zwischen der
Paratethys und dem Proto-Mediterran sowie der neogenen
Indo-West-Pazifischen Region. Diese Verbindung ist je-
doch problematisch, da der Tethys-Seeweg zu dieser Zeit
bereits geschlossen war oder nur eine sehr seichte, stark
evaproritische Passage repra
¨
sentierte. Die Gattung zeigt
keine na
¨
here Verwandtschaft mit den bekannten Strom-
biden-Gattungen, trotz der oberfla
¨
chlichen A
¨
hnlichkeiten
mit einigen modernen Vertretern in der IWP-Region.
Daher wird hier fu
¨
r dieses Taxon Europrotomus nov. gen.
als neue Gattung eingefu
¨
hrt. Diese Gattung beinhaltet nur ein
oder zwei Arten, die ausschließlich aus dem europa
¨
ischen
Mittel-Mioza
¨
n bekannt sind. Bisher konnten keine Vor-
ga
¨
nger nachgewiesen werden, weshalb autochthone Evo-
lution als eher unwahrscheinlich betrachtet wird.
Stattdessen ko
¨
nnte dieses plo
¨
tzliche Auftreten durch
Immigration von einer benachbarten biogeographischen
Provinz erkla
¨
rt werden. Ein derartiges biogeograpohisches
Herkunftsgebiet ko
¨
nnte hypothetisch entlang der tropi-
schen Ku
¨
sten West-Afrikas etabliert gewesen sein—ein
Szenario, das auch aus dem Pleistoza
¨
n dokumentiert ist.
Schlu
¨
sselwo
¨
rter Gastropoda Strombidae Euprotomus
neue Gattung Mioza
¨
n Biogeographie Paratethys
Introduction
The wealth of taxa of the family Strombidae in the modern
Indo-West Pacific Region (IWP) has its roots in the
G. C. Kronenberg
NCBNaturalis, P.O. Box 9517, 2300 RA Leiden,
The Netherlands
e-mail: gijs.kronenberg@ncbnaturalis.nl
M. Harzhauser (&)
Geological-Paleontological Department, Natural History
Museum Vienna, Burgring 7, 1010 Vienna, Austria
e-mail: mathias.harzhauser@nhm-wien.ac.at
123
Pala
¨
ontol Z
DOI 10.1007/s12542-011-0121-1
Miocene. At that time a major radiation of genera and
species occurred there, which continued into Pliocene
times (Vredenburg 1925; Abbott 1960; Harzhauser 2007).
An important event in the evolutionary history of the
Tethyan lineages was the final closure of the Tethys Sea-
way around the Early/Middle Miocene boundary. At that
time the IWP lineages became cut off from the western
Tethyan area, which transformed into the proto-Mediter-
ranean Sea and the Paratethys (Harzhauser et al. 2002;
Harzhauser and Piller 2007). Several authors discuss a
short reconnection via Mesopotamia during the earliest
Middle Miocene (Ro
¨
gl 1998; Popov et al. 2004; Fig. 1).
Widespread Middle Miocene evaporites in the crucial area,
however, indicate a shallow and hypersaline seaway that
was probably not suitable for major faunal exchange.
Taxa with affinities with Indo-Pacific genera appearing
in the proto-Mediterranean during the phase of ceasing
connectivity are therefore highly interesting for discussions
on biogeographic and palaeogeographic reconnections
during the Middle Miocene (Grecchi 1978;Ro
¨
gl 1998).
One of these striking taxa is the nominal strombid species
Strombus schroeckingeri Ho
¨
rnes in Hoernes & Auinger,
1884, which is in some morphological characters remi-
niscent of representatives of the genus Euprotomus.
A Middle Miocene occurrence of this purely IWP genus in
the Mediterranean area would clearly imply a marine
passage between the Indo-Pacific and the Mediterranean
seas at that time. Moreover, it would even point to a
western origin of the genus, which is known so far in the
IWP only from Pliocene strata (Abbott 1960).
Thus, in the course of our research on the Paratethyan
Persististrombus lineages (for some preliminary notes, see
Harzhauser and Kronenberg 2008) and a continuation of
the revision of Euprotomus Gill, 1870 (Kronenberg 1998,
1999, 2002a, b), we examined a number of specimens of
the nominal taxon Strombus schroeckingeri Ho
¨
rnes in
Hoernes & Auinger, 1884.
Strombus schroeckingeri has usually been allocated to
Monodactylus Mo
¨
rch, 1852 (TS by SD (subsequent des-
ignation): Strombus adustus Chemnitz [not avail-
able] = Lambis aratrum Ro
¨
ding,
1798)byHo
¨
rnes in
Hoernes & Auinger (1884: 165) and Cossmann (1904: 8),
or Euprotomus Gill, 1870 (TS by monotypy Strombus
aurisdianae Linnaeus, 1758) by e.g.; Csepreghy-Meznerics
(1954: 398), Kojumdigieva and Strachimirov, (1960: 130),
Strausz (1966: 223) and Atanackovic (1985: 128). Bandel
(2007: 148) more or less explicitly advocated inclusion of
S. schroeckingeri in Lentigo Jousseaume, 1886 (TS by
monotypy Strombus lentiginosus Linnaeus, 1758). As
Strombus schroeckingeri lived about the same time as the
representatives of Persististrombus Kronenberg & Lee,
2007 [TS by OD (original designation] Strombus granul-
atus Swainson, 1822), in the southern basins of the Central
Paratethys one might also expect a close relationship
between S. schroeckingeri and one or more species allo-
cated to Persististrombus. Therefore, we will take the
opportunity to briefly discuss the taxonomic odyssey of the
taxa involved.
Analysis of the concepts of Monodactylus, Euprotomus
and Lentigo
Most authors allocated schroeckingeri Ho
¨
rnes in Hoernes
& Auinger, 1884,toMonodactylus Mo
¨
rch, 1852,orto
Euprotomus Gill, 1870. Based on shell characters, this
allocation appears to be far-fetched, yet in light of the
complex history of the genus-level taxon, this allocation is
not as strange as one might think.
Klein (1753: 98) introduced the name Monodactylus as a
genus of his ‘‘classis’’ Alata and referred to illustrations in
Buonanni (1681), Petiver (1708 [not seen]), Lister (1685–
1692), and Rumphius (
1705) (Table 1). The figures in these
works represent specimens with one digit at the tip of the
outer wing, now considered to be in Euprotomus; Lobatus
Iredale, 1921 [TS by monotypy Strombus bituberculatus
Lamarck, 1822 (= Strombus raninus Gmelin, 1791)]; or
Tricornis Jousseaume, 1886 [TS by monotypy Strombus
tricornis Lamarck, 1816 (= Strombus tricornis Lightfoot,
Fig. 1 Occurrences of Europrotomus plotted on the palaeogeography
of the circum-proto-Mediterranean area during the early Middle
Miocene (after Harzhauser and Piller 2007). The Tethys Seaway
connecting the proto-Mediterranean Sea with the early Indo-Pacific
Ocean was either closed or extremely shallow and evaporitic (Ro
¨
gl
1998; Popov et al. 2004). Localities: 1 Weitendorf, Wetzelsdorf
(Austria), 2 Hrvac
´
ani (Bosnia and Herzegovina), 3 Sa
´
msonha
´
za,
Ma
´
rkha
´
za (Hungary), 4 Cos¸teiu de Sus, La
˘
pugiu de Sus, (Romania), 5
Tarnene, Pleven (Bulgaria), 6: Montjuic at Barcelona (Spain)
G. C. Kronenberg, M. Harzhauser
123
1786)] except for fig. 306 [mirror imaged] in Buonanni
(1681), which is definitely a strombid, but without a digit at
the tip of the wing. This specimen cannot be identified with
certainty, but the shape of the elongated anterior canal
suggests a specimen of Lobatus that has been tampered
with. As Klein (1753) is a pre-Linnaean work, his names
are not available anyway (International Commission on
Zoological Nomenclature (1999) Art. 3.1).
Mo
¨
rch (1852) was the first author to make the name
Monodactylus, attributed to Klein (1753), meet the require-
ments of the International Commission on Zoological
Nomenclature (1999). Unfortunately, the name was preoc-
cupied by de Lace
´
pe
`
de (1801) who had established it for a
perciform fish. In his list of species allocated to Monodacty-
lus,Mo
¨
rch (1852: 62) mentioned four species that are now
allocated to Euprotomus, but also the nominal taxa Strombus
lentiginosus Linnaeus, 1758 (the type species of Lentigo
Jousseaume, 1886) and S. fasciatus Born, 1778 (now allo-
cated to Conomurex Bayle in P. Fischer, 1884). Both these
species do not have a ‘‘monodactylus.’’ Adams and Adams
(1853) returned to Klein’s original concept, i.e., included all
‘‘monodactyl’’ species in Monodactylus, and excluded both S.
lentiginosus and S. fasciatus from that taxon. Although there
is an overlap with Euprotomus, it should be noted that the
concept of Monodactylus Mo
¨
rch (non Lace
´
pe
`
de) is not the
same as the concept of Euprotomus.
Tryon (1885) was probably one of the first authors who
used both Monodactylus and Euprotomus as valid genus
level taxa. Like Mo
¨
rch (1852) and Adams and Adams
(1853), Tryon was not aware that the name Monodactylus
was preoccupied as he selected S. pacificus Swainson, 1821
(= Lambis vomer Ro
¨
ding, 1798) as type species. Cossmann
(1904: 7) did not follow this designation, but selected yet
another species, viz. S. gallus Linnaeus, 1758, a species not
listed by Mo
¨
rch (1852), and therefore not eligible as type
species of Monodactylus. Cossmann (1904) further listed
two fossil species, viz. S. trigonus Grateloup and S. sch-
roeckingeri,inMonodactylus. For a summary of type
designations for Monodactylus, see Abbott (1960: 125).
Tryon (1885) designated Strombus laciniatus ‘‘Chem-
nitz’’ Dillwyn, 1817 (= Strombus sinuatus [Lightfoot],
1786) as type species of Euprotomus. This is an invalid
designation as Strombus aurisdianae was the only species
mentioned by Gill (1870). By his action, Tryon in fact
changed the whole concept of Euprotomus. Apart from the
type species, Tryon allocated five more species within
Euprotomus. Of these, two are now allocated to Lentigo;
the other four are now allocated to Sinustrombus Bandel,
2007, and Thersistrombus Bandel 2007. For a preliminary
discussion on these taxa, see Kronenberg (2009). None of
these species is currently allocated to Euprotomus.
In his influential work Abbott (1960) brought an end to the
confusion about Monodactylus and Euprotomus.Abbott
pointed out that Monodactylus
Mo
¨
rch was preoccupied and
restricted the concept of Euprotomus. Indeed, Euprotomus
sensu Abbott is a morphologically coherent group that is
probably monophyletic. This has been confirmed by molec-
ular data by Latiolais (2003, 2006). In the consensus tree as
presented by Latiolais et al. (2006), the nominal taxa Lambis
vomer (Ro
¨
ding, 1798), L. bulla (Ro
¨
ding, 1798)andStrombus
aurisdianae Linnaeus, 1758—all allocated to Euprotomus by
Abbott (1960)—plot out as a clade that is sister to the clade of
Panamic, western and eastern Atlantic strombids.
Apart from the retroactive allocation to Lentigo of its
possible synonym S. almerai (see Almera and Bofill y Poch
1885), only Bandel (2007) allocated this species to Lentigo
Jousseaume, 1886. Originally, this genus was introduced
with only one species allocated to it, viz. Strombus len-
tiginosus. Apart from the type species, Abbott (1960)
allocated four more recent species to Lentigo: the nominal
taxa L. pipus, S. fasciatus, S. latus and S. granulatus.
Moolenbeek and Dekker (1993) pointed out that, based on
shell morphology and characters of the radula, the position
of S. fasciatus in Lentigo is no longer tenable, and they
allocated this species to Conomurex. Subsequently, Kro-
nenberg and Lee (2007) allocated S. granulatus and S. latus
to their new genus Persististrombus, retaining only
S. lentiginosus and L. pipus in Lentigo. In their discussion,
Kronenberg and Lee (2007) implicitly redescribed Lentigo
on shell characters. They discriminated Lentigo from
Persististrombus by the two notches on the adapical part of
the outer lip, resulting in two lobes, of which the adaxial
one is attached to the spire; a more distinct posterior canal;
presence of a number of small triangular projections at the
abapical side of the outer lip on the flange between the
strombid notch and the anterior canal, and the columellar
callus that does not reach the base of the columella, yet
Table 1 References to figures and descriptions referred to by Klein
(1753) to his concept of Monodactylus
1. Lister 873/29 = Tricornis tricornis
2. Lister 871/25 = cf. Lobatus raninus
Petiver p. 189 no. 3
3. Lister 874/30 = Lobatus gallus
4. Buonanni 309; 310 = Lobatus gallus
5. a. Rumphius caption to pl. 37
Buonanni 301 = Euprotomus aurisdianae
Lister 872 = Euprotomus aratrum and E. bulla
b. Rumphius 111 # XI; text refers to E. aurisdianae and E.
bulla
c. Buonanni 302 = Euprotomus bulla
Figure in Klein (1753) pl. 6/106 = cf. Tricornis tricornis
d. Lister 871/26 = Euprotomus cf. aurisdianae
e. Buonanni 306 = ?? ,,Tricornis‘‘s.l.
The Petiver reference has not been checked
A new Middle Miocene European strombid genus
123
thickened at the abapical part but not forming a distinct
pad. This opinion was echoed by Landau and da Silva
(2010). For an overview of species’ allocations to the
genera mentioned above, see Table 2.
Abbreviations
GCKE Private collection Gijs C. Kronenberg,
Eindhoven, the Netherlands; to be deposited in
NCBNaturalis
NHMW Naturhistorisches Museum Wien, Austria
Systematic palaeontology
Family Strombidae Rafinesque, 1815
Genus Europrotomus Kronenberg & Harzhauser gen.
nov.
Type species designated herein: Strombus (Monodacty-
lus) schroeckingeri Ho
¨
rnes in Hoernes & Auinger, 1884.
Diagnosis: Strombid genus encompassing shells of
medium size, with medium-sized spire, flaring outer lip
with sharp rim, short anterior canal, distinct row of
shoulder knobs on the last three whorls, becoming tri-
fid, and diverging cords on abapertural side of the last
whorl.
Description: See description of Europrotomus schroec-
kingeri below.
Table 2 Concepts of the genus level taxa Monodactylus, Euprotomus and Lentigo, by alphabetical arrangement of the species allocated to these
taxa by different authors
Genus Mo
¨
rch, 1852 Adams & Adams,
1853
Tryon, 1884 Cossmann, 1904 Abbott, 1960 Kronenberg
et al., 2010/
2011
Monodactylus
Adustus (= aratrum)
[Euprotomus]
aratrum (= vomer)
[Euprotomus]
aurisdianae
(= bulla)
[Euprotomus]
fasciatus
[Conomurex]
lentiginosus[Lentigo]
striatogranulatus
(= aurisdianae)
[Euprotomus]
Adustus(= aratrum)
[Euprotomus]
aratrum (= vomer)
[Euprotomus]
aurisdianae
[Euprotomus]
australis (= iredalei)
[Euprotomus]
costo-muricatus
(= raninus)
[Lobatus]
gallus [Lobatus]
guttatus (= bulla)
[Euprotomus]
peruvianus[Lobatus]
striatogranulatus
(= aurisdianae)
[Euprotomus]
tricornis [Tricornis]
Aurisdianae
(= aurisdianae ? bulla)
aurisdianae var.
melanostomus
(= aratrum)
bituberculatus
(= raninus)[Lobatus]
gallus[Lobatus]
pacificus(= vomer)
pacificus var. australis
(= iredalei)
peruvianus [Lobatus]
tricornis [Tricornis]
Gallus [Lobatus]
schroeckingeri
trigonus
[allocated
to Tricornis by
Abbott 1960]
Euprotomus
Laciniatus (= sinuatus)
[Sinustrombus]
latissimus [Sinustrombus]
lentiginosus [Lentigo]
papilio (= pipus)
[Lentigo]
ponderosus (= thersites)
[Thersistrombus]
taurus [Sinustrombus]
Laciniatus
(= sinuatus)
[Sinustrombus]
Aurisdianae
aratrum
aurisdianae
aurisdianae
bulla
vomer vomer
vomer hawaiensis
vomer iredalei
Aratrum
aurisdianae
aurora
bulla
chrysostomus
hawaiensis
iredalei
vomer
Lentigo Fasciatus
[Conomurex]
granulatus
[Persististrombus]
latus
[Persististrombus]
lentiginosus
pipus
Lentiginosus
pipus
Extinct taxa accompanied by a dagger (). Current species names between brackets (), current generic allocation between square brackets []. Type species,
when designated, underlined. See also text
G. C. Kronenberg, M. Harzhauser
123
Derivation of name: Compound noun of ‘‘Euro,’’ refer-
ring to the European distribution of this genus as known so
far, and ‘‘protomus’’ as allusion to the genus Euprotomus
Gill, 1870, to which the type species has been allocated.
Distribution: This fossil strombid genus is known so far
only from the European Middle Miocene and was restricted
to the Mediterranean Sea and the adjacent Paratethys Sea.
Remarks: Other species assigned to Europrotomus gen.
nov.: Strombus almerai Crosse, 1885; but see below.
Europrotomus schroeckingeri (Ho
¨
rnes in Hoernes &
Auinger, 1884) comb. nov.
Figs. 2a–f, 3a–g
*1884 Strombus (Monodactylus) schro
¨
ckingeri Ho
¨
rnes
in Hoernes & Auinger: 165, pl. 19, figs. 6–7
1904 Strombus (Monodactylus) schro
¨
ckingeri
Ho
¨
rnes.—Cossmann: 8
1954 Strombus (Euprotomus) schro
¨
ckingeri Ho
¨
rnes.—
Csepreghy-Meznerics: 398, pl. 2, figs. 1–2
1960 Strombus (Euprotomus) schroeckingeri
Ho
¨
rnes.—Kojumdgieva & Strachimirov: 130, pl.
35, figs. 5a–5b
1966 Strombus (Euprotomus) schro
¨
ckingeri Ho
¨
rnes.—
Strausz: 223, fig. 104
1985 Canarium (Euprotomus) schroeckingeri
(Hoernes).—Atanackovic: 128, Pl. 24,
figs. 10–11
1993 Strombus
(Strombus) coronatus Defrance.—
Nikolov (1993): 69, pl. 3, figs. 7–8
1993 Strombus (Strombus) nodosus subcancellata
(Grateloup).—Nikolov: 70, pl. 3, figs. 9–10
2003 Strombus (Euprotomus) schroeckingeri
(Ho
¨
rnes).—Harzhauser et al.: 333, fig. 8
2007 Strombus (Lentigo) schroeckingeri Ho
¨
rnes, 1880
[sic!].—Bandel: 148
2006 Strombus schro
¨
ckingeri [sic!].—Hiden (2006): 6
?1885 Strombus lentiginosus Linne
´
Var.—Almera &
Bofill y Poch: 32, pl. 1 figs 1, 2 [non Strombus
lentiginosus Linnaeus, 1758]
?1885 Strombus almerai Crosse: 242. nom nov. pro
Strombus lentiginosus Almera & Bofill y Poch,
non Linnaeus
?1886 Strombus almerae Crosse.—Almera & Bofill y
Poch: 402, pl. 10, figs. 1–2 [unjustified
emendation of S. almerai Crosse]
Lectotype: designated herein: NHMW 1867/XIX/72;
Fig. 2a; illustrated in Hoernes and Auinger (1884; pl.
19, fig. 7).
Locus typicus: Cos¸teiu de Sus, Romania; Transylvanian
Basin.
Stratum typicum: marly sandstone of the lower Badenian
(= lower Langhian).
Measurement: height: 43.4 mm.
Paralectotype 1: Cos¸teiu de Sus, Romania; Transylva-
nian Basin; height: 47.2 mm (Fig. 2b), NHMW
1855/XLIII/19b.
Paralectotype 2: Cos¸teiu de Sus, Romania; Transylva-
nian Basin; height: 53.8 mm (Fig. 2c), NHMW
1867/XIX/72; previously illustrated by Hoernes and
Auinger (1884; pl. 19, fig. 6).
Paralectotype 3:La
˘
pugiu de Sus, Romania; Transylva-
nian Basin; height: 52 mm (Fig. 3a); NHMW
1855/XLIII/19a
Paralectotype 4
:La
˘
pugiu de Sus, Romania; Transylva-
nian Basin; height: 52 mm (Fig. 3b); NHMW 1872/V/
25;
Additional specimens from the type locality Cos¸teiu de
Sus (leg. et don. Anton and Thomas Breitenberger):
NHMW 2007z0114/0005; Fig. 2d; height: 42 mm.
NHMW 2007z0114/0008; Fig. 2e; height: 52 mm.
NHMW 2007z0114/0007; Fig. 2f; height: 51.5 mm.
NHMW 2007z0114/0001; Fig. 3c; height: 23 mm
(juvenile).
NHMW 2007z0114/0002; Fig. 3d; height: 23 mm
(juvenile).
NHMW 2007z0114/0004; Fig. 3e; height: 30.5 mm
(dwarf specimen).
NHMW 2007z0114/0006; Fig. 3f; height: 41 mm.
NHMW 2007z0114/0009; Fig. 3g; height: 49.5 mm.
Additional specimens from La
˘
pugiu de Sus (leg. et don.
Anton and Thomas Breitenberger):
GCKE 6306.
Description: Medium-sized shells of eight teleoconch
whorls. Protoconch strongly abraded in all available
specimens and seems to consist of about 2.5 rather high,
moderately convex whorls. Up to three varix-like swellings
appear on earliest teleoconch whorls aside from numerous
oblique axial ribs, soon passing into pointed knobs or
nodes. Spire whorls bear a prominent keel, situated vari-
ably in the area from the middle of the whorl to the anterior
suture. Concave sutural ramp bearing distinct growth lines
that are crossed by stronger spiral threads. Spiral threads
accompanied by 8–10 blunt spiral ridges on the body
whorl, which bears irregular nodes of variable strength;
most prominent and pointed ones appear along the shoulder
of the body whorl.
A new Middle Miocene European strombid genus
123
G. C. Kronenberg, M. Harzhauser
123
Outer lip expanding, attaches up to the 4th and 5th spire
whorl; thickened posterior to the strombid notch but thin
anterior to it. In the anterior 2/3 of the outer lip a swelling
with strong lirae is developed about 5–7 mm deep in the
aperture. Three (Hoernes and Auinger 1884: pl. 19 fig. 7a,
b here refigured Fig. 2a) to seven (Hoernes and Auinger
1884: pl. 19 fig. 6a, b, here refigured Fig. 2c) broad, low
axial lobes occur especially in its posterior part, but also
extending to the lateral part as well, bordered by a narrow,
moderately deep canal where the wing attaches to the spire.
These lobes are a continuation of well-developed spiral
cords on the dorsal side of the shell. The two to three most
adapical of these cords develop only at the point where the
outer lip starts to expand, but the others, when present, are
a continuation of strong, knob-bearing, spiral cords already
present on the dorsal side of the shell. Columellar callus
extending on the base without covering it completely.
Further lirae occur in the very anterior part and the pos-
terior termination of the columella.
Comparison of shell characters: the shells show a
broad, glossy columellar callus partially covering the base.
In this feature it is quite close to the modern representatives
of Euprotomus. However, Europrotomus is readily distin-
guished from Euprotomus by the difference in the much
shorter and the less bent anterior canal, and the absence of
the finger-like digit on the outer lip. The sculpture of the
outer lip has some counterpart in the shells of the modern
Euprotomus vomer-hawaiensis-iredalei complex. Even the
lirated axial ridge in the inner side of the outer lip is,
although much weaker, still presented in the modern spe-
cies group. Another similarity is the narrow posterior canal,
the presence of varices on the early teleoconch whorls. The
paucispiral protoconch of the extant Euprotomus iredalei
(Abbott 1960) resembles that of Europrotomus schroec-
kingeri in its rather low, bulbous shape (about 2–2.5
whorls).
As far as Lentigo is concerned, the general appearance
of Europrotomus is quite similar, and this is also true for
the dorsal sculpture. In Lentigo there are however only two
lobes on the posterior part of the outer lip, contrary to the
three to seven in Europrotomus Moreover, in Europroto-
mus these lobes are clearly a continuation of the well-
developed spiral cords, which is not the case in Lentigo.
Also, the triangular projections on the strombid notch and
flange between the strombid notch and anterior canal
present in Lentigo are missing in Europrotomus.
In Persististrombus Kronenberg & Lee, 2007, the edge
of the outer lip has no lobes at all, and the outer lip is
attached to the shell much more abapically.
The presence of the lobes on the rim of the outer lip is
slightly reminiscent of the lobes as seen in Sinustrombus
sinuatus ([Lightfoot], 1786), but in the latter species the
lobes are much better developed, the outer lip is attached
much closer to the apex, and the flange between the
strombid notch and the anterior canal. Also, in S. sinuatus
the rim of the outer lip is bent sharply towards the colu-
mella when reaching adulthood, a character not observed in
Europrotomus.
The continuation of the spiral cords into the lobes at the
rim of the outer lip is somewhat reminiscent of the way the
digits are formed in Lambis and Harpago.InLambis and
Harpago, however, these digits are much longer and are
formed by distinct lobes at the mantle edge, where the
lateral sides of the mantle edge are bent to reach other, thus
forming a hollow cone that is subsequently filled with shell
material. The place where the rims of the lobes touched
each other remains visible as a very narrow furrow.
Moreover, the anterior canal in Lambis is very much
elongated.
Based on shell morphology, we conclude that S. sch-
roeckingeri cannot be allocated to Persististrombus, nor
can it be allocated to either Euprotomus or Lentigo. The
previous taxonomic confusion is largely based on the
strongly changing concepts of Monodactylus Mo
¨
rch (non
Lace
´
pe
`
de) and Euprotomus. As no other genus level taxon
within the family Strombidae appears to fit to accommo-
date this species, Europrotomus is introduced as a new
genus.
Remarks: A poorly preserved strombid from the Langh-
ian and lower Serravallian of Spain, erroneously identified as
Strombus lentiginosus (Almera & Bofill y Poch, 1885), was
subsequently renamed by Crosse (1885)asS. almerai and
accepted by Almera and Bofill y Poch (1886)asS. almerae.
The change in spelling of the specific epithet is however an
unjustified emendation of S. almerai (International Com-
mission on Zoological Nomenclature (1999) Art. 33.2.3).
This species closely resembles S. schroeckingeri in size and
ornamentation. Unfortunately, the sculpture of the aperture
is poorly described, except for some comments on granula-
tions of the columella. Additionally, its wing is described as
expanding and undulated in its posterior part, and a faint
posterior canal is also mentioned by Almera and Bofill y
Poch (1885, 1886). Differences, however, are the stout spire
and the shoulder of the last whorl, which seems to be closer to
the suture than in Europrotomus gen. nov. schroeckingeri.
Hence, this specimen might be conspecific with the Para-
tethyan Europrotomus. schroeckingeri, but a clear decision
depends on the availability of more material.
Fig. 2 Europrotomus schroeckingeri (Ho
¨
rnes in Hoernes & Auinger,
1884) comb. nov. from Cos¸teiu de Sus in Romania (Langhian).
a Lectotype, NHMW 1867/XIX/72, b Paralectotype 1, NHMW
1855/XLIII/19b, c Paralectotype 2 NHMW 1867/XIX/72, d NHMW
2007z0114/0005, e NHMW 2007z0114/0008, f NHMW 2007z0114/
0007
b
A new Middle Miocene European strombid genus
123
G. C. Kronenberg, M. Harzhauser
123
Palaeoecology: All specimens are found in marly silty
sandstone in association with a diverse subtropical mollusc
fauna pointing to shallow marine soft bottom environments
with normal salinity. It is unknown from the Middle
Miocene reefs and coral carpets of the Paratethys (own
observation). It is only very rarely found associated with
other Stromboidea except for Tibia dentata (Grateloup
1827).
Distribution: Weitendorf, Wetzelsdorf (Styrian Basin,
Austria); Sa
´
msonha
´
za, Ma
´
rkha
´
za (Cserha
´
t Mountains,
Hungary); Cos¸teiu de Sus, La
˘
pugiu de Sus (Transylvanian
Basin, Romania); Hrvac
´
ani (southern Pannonian Basin,
Bosnia and Herzegovina), Tarnene and Pleven (Carpathian
Basin, Bulgaria) (see Harzhauser et al. 2003 for map).
Discussion
A guest from the east or a home-made western Tethyan
element?
Europrotomus schroeckingeri displays a remarkable dis-
tribution. From literature references, it is seemingly con-
fined to the Central Paratethys, which is possibly rather a
matter of a lack of investigations in adjacent regions.
Within the Paratethys it is restricted to the southern basins
(Harzhauser et al. 2003). The rare shell is found in
Romania, Bulgaria, Hungary, Bosnia and Herzegovina and
in the Styrian Basin in Austria. It is unknown from the
North Alpine Foreland Basin, the Vienna Basin and the
more northern Carpathian Foredeep. During the early
Middle Miocene, the southern Paratethys basins, e.g., the
Styrian and Pannonian basins, had a direct connection to
the proto-Mediterranean Sea, while the northern basins,
e.g., the Vienna and Carpathian basins, were fed by a more
eastern connection. Both water masses were probably
separated by emerged areas that now form the Danube
Basin. This assumption is supported by geochemical data
on stable isotope composition of mollusc shell aragonite,
which clearly indicates strongly deviating water chemistry
between these two branches of the Paratethys Sea (Latal
et al. 2005). Therefore, the strombid was hindered on its
way to the north. In respect to the generally low endemicity
of the southern Paratethyan mollusc faunas at that time
(Harzhauser and Piller 2007), it may be expected that
Europrotomus schroeckingeri was also distributed in the
proto-Mediterranean area. Its absence from the fossil
record may be explained by the fact that coeval Langhian
deposits in the Mediterranean area are rare and mainly
represented by deep water deposits (e.g., Mourik et al.
2011). None of the rare Langhian shallow water faunas
from the Monte dei Cappuccini in the Turin Mountains
(Sacco 1893) and from the Mut Basin in Turkey (Mandic
et al. 2004) yielded Europrotomus schroeckingeri.
Only the occurrence of Europrotomus almerai (Crosse
1885) in the Middle Miocene of Spain suggests a wider
distribution of the genus. Nevertheless, its sudden appear-
ance and the lack of obvious ancestors might point to
immigration from a nearby bioprovince.
Immigration from western Africa would be a realistic
scenario. This hypothesis, however, is a hard test as there are
no fossil-bearing Lower and Middle Miocene deposits
known along the West African coast. The Paleocene mol-
luscan fauna from Nigeria has been discussed by Newton
(1905, 1922), Eames (1957) and Adegoke (1977), and from
the Gold Coast (= Ghana) by Cox (1952), but although there
are stromboideans discussed (Rimellidae. Rostellariidae and
Seraphsidae) in those papers, no strombids are mentioned. It
is quite unlikely that Europrotomus schroeckingeri
arose
from one of these stromboidean families. Therefore, no
potential West African ancestor can be shown.
Immigration from the East is unlikely as there is also no
evidence of E. schroeckingeri in the contemporaneous but
always lowly diversified faunas of the Eastern Paratethys
(Ilyina 1993). Hence, it would be tempting to explain this
erratic appearance by immigration from either the Ameri-
cas in the west or the proto-Indo-West-Pacific Region in
the east. An arrival of Europrotomus from the western
biogeographic units of the Central Americas, however, is
extremely unlikely in respect to the fossil record (e.g., Jung
and Heitz 2001). A connection of the Mediterranean or the
Paratethys with the early Indo-Pacific existed during the
Early Miocene (Ro
¨
gl 1998; Harzhauser et al. 2007).
Hypothetically, this marine connection would have allowed
the immigration of Europrotomus. Nonetheless, it has to be
kept in mind that all Oligocene-Miocene mollusc faunas of
Arabia, Eastern Africa, Pakistan, India, Java and Borneo
lack any clear evidence for a Europrotomus-Euprotomus-
related strombid (see Vredenburg 1925; Beets 1941;
Abbott 1960; Hoek Ostende et al. 2002; Harzhauser 2007,
2009; Harzhauser et al. 2009). Thus, there is little reason to
hypothesize a westward immigration into the Mediterra-
nean area. Moreover, during the Middle Miocene when
Europrotomus appears, the eastern route via the Tethys
Seaway was already closed or hard to cross (Ro
¨
gl 1998,
Harzhauser et al. 2007).
Fig. 3 a–b Europrotomus schroeckingeri (Ho
¨
rnes in Hoernes &
Auinger, 1884) comb. nov. from La
˘
pugiu de Sus in Romania
(Langhian). a Paralectotype 3, NHMW 1855/XLIII/19a, b Paralecto-
type 4, NHMW 1872/V/25, c–g Europrotomus schroeckingeri
(Ho
¨
rnes in Hoernes & Auinger, 1884) comb. nov. from Cos¸teiu de
Sus in Romania. c Juvenile specimen NHMW 2007z0114/0001,
d juvenile specimen, NHMW 2007z0114/0002, e dwarf specimen
NHMW 2007z0114/0004, f strongly sculptured specimen, NHMW
2007z0114/0006, g specimen with reduced sculpture, NHMW
2007z0114/0009
b
A new Middle Miocene European strombid genus
123
All the occurrences are concentrated in the southern
basins of the Paratethys Sea, which was a northern gulf of
the proto-Mediterranean Sea (Ro
¨
gl 1998; Popov et al.
2004; Harzhauser and Piller 2007). Soon after, the genus
vanishes completely from the Paratethys Sea and is
unknown from deposits younger than 14–13 Ma. This
range coincides conspicuously with the mid-Miocene Cli-
matic Optimum (Zachos et al. 2001), while its sudden
decline coincides with the Mid-Miocene climate transition
(Shevenell et al. 2004). This climate-related stratigraphic
and biogeographic pattern was documented for a wide
range of Paratethyan taxa of gastropods and foraminifers
(Harzhauser and Piller 2007) as well as bivalves (Harzha-
user et al. 2003).
Hypothetical relations and roots
Although Europrotomus can easily be distinguished from
other genus level taxa, morphological elements of the last
whorl suggest some affinities with the genera to which it is
compared above, e.g., Euprotomus, Lentigo, Persisti-
strombus, Sinustrombus, Harpago and Lambis. In his thesis,
Latiolais (2003: fig. 1) illustrates a maximum likelihood
tree constructed from 325 bp of nuclear histone H3 for
numerous species of Strombidae. Unfortunately, in the tree
constructed from 640 bp of mitochondrial COI (Latiolais
2003: fig. 2), some of the species are not represented,
so these species are also missing in the consensus tree
(Latiolais 2003: fig. 3; Latiolais et al. 2006). Based upon the
nuclear histone H3 results, Europrotomus gen. nov. may be
derived from a Canarium-like ancestor and at the root of the
clade that leads to all recent American and West African
genera (Lobatus; Persististrombus; Strombus) and also the
Indo-Pacific genera Conomurex, Euprotomus, Gibberulus,
Harpago, Lambis, Lentigo, Sinustrombus, Thersistrombus
and Tricornis (names following Kronenberg et al. 2010/
2011). This may be an indication of a Miocene ‘‘explosion’’
of genus level taxa (Williams and Duda 2008).
The proposed relation with a Canarium
-like ancestor,
however, is highly speculative, as Europrotomus appears
abruptly in the fossil record and is restricted to a rather
narrow time span during the Middle Miocene. Only few
strombid genera are present in the Early Miocene of the
Western Tethys. Of these, only Persististrombus is recor-
ded from numerous localities, documenting a continuous
occurrence of the lineage from the Oligocene up to the
Pliocene (Lozouet and Maestrati 1986; Harzhauser and
Kronenberg 2008). A close relation of Persististrombus
with Europrotomus, as discussed above, is very unlikely.
Similarly, a phylogenetic relation with Dilatilabrum
Cossmann, 1904, which occurs as a Palaeogene relic in
the Lower Miocene of the Aquitaine Basin in France,
can be ruled out. Representatives of Dilatilabrum are
characterized by extremely solid shells with wing-like
outer lips with straight edges. Finally, a third—still
unnamed—strombid genus is known from the Lower
Miocene of the Aquitaine Basin in France and the Turin
Mountains in Italy. It is represented only by the species
‘‘Strombus’’ mitroparvus Sacco, 1893. This small elongate
strombid with strong nodes and a narrow wing might be the
earliest species of a lineage that is represented in the
Miocene to Pleistocene of the IWP by ‘‘Strombus’’ micklei
Ladd, 1972, and ‘‘Strombus’’ blanci Tro
¨
ndle
´
& Salvat,
2010 (Ladd 1972; Tro
¨
ndle
´
and Salvat 2010). ‘‘Strombus’’
mitroparvus, with its very narrow aperture and an insig-
nificant wing that does not reach the spire, differs funda-
mentally from Europrotomus and is a very unlikely
ancestor candidate.
Conclusions
The Neogene strombid fauna of the proto-Mediterranean
and Paratethys seas was dominated by representatives of
the Persististrombus lineage. Apart from Dilatilabrum and
‘‘ Strombus’’ mitropravus from the Early Miocene, the only
other genus present was Europrotomus known solely from
the Middle Miocene. It remained rather rare and did not
radiate as it is known so far only from a single species,
although a second one, viz. Strombus almerai, might have
existed in the western Mediterranean.
This taxon has no obvious roots in the fossil record and
is unrelated to the known Early Miocene strombids of the
entire Tethys Realm. Moreover, it has no direct relation
with the modern genera to which it has been assigned in the
literature so far. The lack of potential ancestor species in
the Lower Miocene deposits of Eurasia might indicate that
Europrotomus schroeckingeri is an immigrant, which
expanded its tropical distribution area toward the north
during the warm Langhian. As the connection into the early
Indo-Pacific was already ceasing during the Middle Mio-
cene, this passage is an unlikely immigration route. Hence,
an arrival from the west would be more realistic. Especially
the tropical coasts of West Africa might have acted as
source area. This ad-hoc hypothesis, however, is hard to
test because of the lack of Miocene faunas in West Africa.
Some support for this hypothesis is a Pleistocene pendent
scenario within the
Persististrombus lineage. With the
Pliocene cooling, this thermophilic strombid retreated from
the Mediterranean Sea and survived as Persististrombus
latus (Gmelin 1791) along the coasts of West Africa. From
there, it re-entered the Mediterranean during the Marine
Isotope Stages 7 and 5 (De Torres et al. 2009) and probably
also during MIS 3 (Ro
¨
gl et al. 1997; Zazo et al. 1984).
The case of Europrotomus, with its superficial similar-
ities with extant IWP-genera, clearly documents the
G. C. Kronenberg, M. Harzhauser
123
negative impact of outdated genus-concepts on biogeog-
raphy and palaeogeography conclusions.
Acknowledgments We would like to thank Anton and Thomas
Breitenberger, Bad Vo
¨
slau and Pottenstein, Austria, for donating
material to both NHMW and Gijs C. Kronenberg; this latter specimen
will be donated to NCBNaturalis at some point in the future. Dr.
Bernard Landau, Albufeira, Portugal, and an anonymous reviewer
carefully read the manuscript and provided valuable suggestions for
improvement.
References
Abbott, R.T. 1960. The genus Strombus in the Indo-Pacific. Indo-
Pacific Mollusca 1(2): 33–146.
Adams, H., and A. Adams. 1853–1858. The Genera of Recent
Mollusca; arranged according to their organization. London:
John van Voorst [for details see Trew, 1992].
Adegoke, O.S. 1977. Stratigraphy and Paleontology of the Ewekoro
formation (Paleocene) of southwestern Nigeria. Bulletins of
American Paleontology 71(295): 1–379.
Almera, J., and A. Bofill y Poch. 1885. Moluscos fo
´
siles de los
terrenos terciarios superiores de Catalun
˜
a. Memorias de la Real
Academia de Ciencias de Barcelona 2: 25–45, pls 1–2.
Almera, J., and A. Bofill y Poch. 1886. Moluscos fo
´
siles de los
terrenos terciarios superiores de Catalun
˜
a. Familia Estro
´
mbidos
d’Orbigny. Boletino de Comision de la Mapa geolo
´
gico del
Espana 13: 394–437.
Atanackovic, M.A. 1985. Mekusci Morskog Miocena Bosne. Geo-
logija Bosne I Hercegovine, Fosilna Fauna I Flora 1: 1–308.
Bandel, K. 2007. About the larval shell of some Stromboidea,
connected to a review of the classification and phylogeny of the
Strombimorpha (Caenogastropoda). Pala
¨
ontologie, Stratigra-
phie, Fazies (15), Freiberger Forschungshefte C 524: 97–206.
Beets, C. 1941. Eine Jungmioza
¨
ne Mollusken-Fauna von der
Halbinsel Mangkalihat, Ost-Borneo (nebst Bemerkungen u
¨
ber
andere Faunen von Ost-Borneo; die Leitfossilien-Frage). Ver-
handelingen van het Geologisch-Mijnbouwkundig Genootschap
voor Nederland en Kolonien, Geologische Serie 13: 1–282.
Born, I. 1778. Index rerum naturalium Musei Cæsarei Vindobonensis.
Pars I.ma. Testacea. Verzeichniß der natu
¨
rlichen Seltenheiten
des k. k. Naturalien Cabinets zu Wien. Erster Theil. Schalthiere.
[1–40], 1–458, [1–82]. Vindobonae: Krauss.
Buonanni, F. 1681. Ricreatione dell’Occhio e della Mente, nell’os-
seruation’ delle chiocciole, propost a’ curiosi delle opere della
Natura, dal (…). Con quattrocentro e cinquanta figure di
testacei diuersi, sopra cui si speigano molti curiosi problemi.
[14], 1–384, [15], 1–20, 1–96, 319 figs, Roma.
Cossmann, M. 1904. Essais de Pale
´
oconchologie Comparee
´
. Livrai-
son 6: 1–151. Paris, France.
Cox, L.R. 1952. Cretaceous and Eocene fossils from the Gold Coast.
Gold Coast Geological Survey bulletin 17: 1–68.
Crosse, H. 1885. Mollusca fossilia stratum tertiariorum supernorum
Cataluniae descripta a doctore D. Jacobo Almera et D. Arthuro
Bofill et Poch.—Proemium. Strombidae: Strombus, Rostellaria
(1). Journal de Conchyliologie 3e se
´
r 25: 242–243.
Csepreghy-Meznerics, I. 1954. Helvetische und tortonische Fauna aus
dem o
¨
stlichen Cserha
´
tgebirge. Annales de l’Institut Ge
´
ologique
de Hongrie 41: 1–185.
de Grateloup, J.P.S. 1827. Description de plusieurs espe
`
ces de
coquilles fossiles des environs de Dax (Landes). Bulletin
d’Histoire Naturelle de la Socie
´
te
´
Linne
´
enne de Bordeaux
2(7): 3–26.
de Lace
´
pe
`
de, B.G.E. 1801. Histoire naturelle des poissons. 3: i–lxvi,
1–558, pls. 1–34. Plassan: Paris.
De Torres, T., J.E. Ortiz, I. Arribas, A. Delgado, R. Julia
`
, and J.A.
Martı
´
n-Rubı
´
. 2009. Geochemistry of Persististrombus latus
Gmelin from the Pleistocene Iberian Mediterranean realm.
Lethaia 43: 149–163.
Dillwyn, L.W. 1817. A descriptive catalogue of recent shells
arranged according to the Linnaean method with particular
attention to the synonymy. Vol. 2, 581–1092 ? [29] index.
Cornhill, London: J. & A. Arch.
Eames, F.E. 1957. Eocene Mollusca from Nigeria, a revision. Bulletin
of the British Museum (Natural History) Geology 3(2): 25–70.
Fischer, P. 1884. Manuel de conchyliologie et de pale
´
ontologie
conchyliologique ou Histoire naturelle des mollusques vivants et
fossils 7: 609–688.
Gill, T. 1870. On the Pterocerae of Lamarck, and their mutual
relations. American Journal of Conchology 5: 120–139.
Gmelin, J.F. 1791. Caroli a Linne
´
Systema naturae per regna tria
naturae. Editio decima tertia. v. 1; Vermes: 3021–3910.
Grecchi, G. 1978. Problems connected with the recorded occurrence
of some Mollusks of Indo-Pacific affinity in the Pliocene of the
Mediterranean area. Rivista Italiana di Paleontologia e Stratig-
rafia 84: 797–812.
Harzhauser, M. 2007. Oligocene and Aquitanian Gastropod Faunas
from the Sultanate of Oman and their biogeographic implications
for the early western Indo-Pacific. Palaeontographica 280:
75–121.
Harzhauser, M. 2009. Aquitanian gastropods of coastal Tanzania and
their biogeographic implications for the early western Indo-
Pacific. Palaeontographica 289: 123–156.
Harzhauser, M., and G.C. Kronenberg. 2008. A note on Strombus
coronatus Defrance, 1827 and Strombus coronatus Ro
¨
ding, 1798
(Mollusca: Gastropoda). The Veliger 50(2): 120–128.
Harzhauser, M., and W.E. Piller. 2007. Benchmark data of a changing
sea—Palaeogeography, Palaeobiogeography and events in the
Central Paratethys during the Miocene. Palaeogeography,
Palaeoclimatology, Palaeoecology 253: 8–31.
Harzhauser, M., W.E. Piller, and F.F. Steininger. 2002. Circum-
Mediterranean Oligo-Miocene biogeographic evolution—the
gastropods’ point of view. Palaeogeography, Palaeoclimatolo-
gy, Palaeoecology
183: 103–133.
Harzhauser, M., O. Mandic, and M. Zuschin. 2003. Changes in
Paratethyan marine molluscs at the Early/Middle Miocene
transition—diversity, paleogeography and paleoclimate. Acta
Geologica Polonica 53: 323–339.
Harzhauser, M., A. Kroh, O. Mandic, W.E. Piller, U.B. Go
¨
hlich,
M. Reuter, and B. Berning. 2007. Biogeographic responses to
geodynamics: A key study all around the Oligo-Miocene
Tethyan Seaway. Journal of Comparative Zoology 246:
241–256.
Harzhauser, M., M. Reuter, W.E. Piller, B. Berning, A. Kroh, and
O. Mandic. 2009. Oligocene and Early Miocene gastropods from
Kutch (NW-India) document an early biogeographic switch from
Western Tethys to Indo-Pacific. Pala
¨
ontologische Zeitschrift 83:
333–372.
Hiden, H. 2006. Der Vulkan von Weitendorf. Hengist 2006(2): 4–9.
Hoek Ostende, L.W. van den, J. Leloux, F.P. Wesselingh, and C.F.
Winkler Prins. 2002. Cenozoic Molluscan types from Java
(Indonesia) in the Martin Collection (Division of Cenozoic
Mollusca), National Museum of Natural History, Leiden. NNM
Technical Bulletin 5: 1–130.
Hoernes, R., and M. Auinger. 1884. Die Gastropoden der Meeresab-
lagerungen der ersten und zweiten Mioza
¨
nen Mediterranstufe in
der o
¨
sterreichischen-ungarischen Monarchie. Abhandlungen der
k.k. geologischen Reichsanstalt 12: 153–192.
A new Middle Miocene European strombid genus
123
Ilyina, L.B. 1993. Handbook for identification of marine Middle
Miocene gastropods of Southwestern Eurasia [in Russian]. Trudy
Paleontologic
ˇ
eskogo Instituta 255: 1–149.
International Commission on Zoological Nomenclature. 1999. Inter-
national code of zoological nomenclature. Ed. 4, i-xxix ?
1–306. London, UK.
Iredale, T. 1921. Molluscan nomenclatural problems and solutions.
Proceedings of the Malacological Society of London 14:
198–208.
Jousseaume, F. 1886. Coquilles marines des cotes d’Abyssinie et de
Zanzibar, recueilis par M. Raffray en 1873 et en 1874. Le
Naturaliste Se
´
rie 1(3): 220–222.
Jung, P., and A. Heitz. 2001. The subgenus Lentigo (Gastropoda:
Strombidae) in tropical America, fossil and living. The Veliger
44: 20–53.
Klein, J.T. 1753. Tentamen methodi ostracologicae: sive Dispositio
naturalis cochlidum et concharum in suas classes, genera et
species, iconibus singulorum generum aeri incisis illustrata;
accedit lucubratiuncula de formatione, cremento et coloribus
tum commentariolum in locum Plinii Hist. nat. libri. IX. cap.
XXXIII. de concharum differntiis. Denique sciagraphia methodi
ad genus serpentium ordinate digerendum. [8], 1–177, [27], pls.
1–102, [8], 1–44, 1–16, [1] Lugduni Batavorum.
Kojumdigieva, E., and B. Strachimirov. 1960. Tortonien. Le
´
s fossiles
de Bulgarie. Academie des Sciences de Bulgarie 7: 1–320. (in
Bulgarian).
Kronenberg, G.C. 1998. Revision of Euprotomus Gill, 1870. 1. The
systematic position of Strombus listeri Gray, 1852 (Gastropoda
Prosobranchia: Strombidae). Vita Marina 45(3–4): 1–6.
Kronenberg, G.C. 1999. Revision of Euprotomus Gill, 1870. 2. On the
identity of Strombus hirasei Kuroda, 1942 (Gastropoda Proso-
branchia: Strombidae). The Festivus 31(6): 63–67.
Kronenberg, G.C. 2002a. Addendum to a revision of Euprotomus Gill,
1870. 2a. On the identity of Strombus hirasei Kuroda, 1942
(Gastropoda Prosobranchia: Strombidae). The Festivus 34(9): 110.
Kronenberg, G.C. 2002b. Revision of Euprotomus Gill, 1870. 3.
Description of Euprotomus aurora spec. nov. from the Indian
Ocean (Gastropoda, Strombidae). Vita Malacologica 1: 55–60.
Kronenberg, G.C. 2009. A note on Bandel’s 2007 review of the
classification and phylogeny of the Strombidae (Caenogastro-
poda). Basteria 73(1–3): 65–67.
Kronenberg, G.C., U. Wieneke, and H. Stoutjesdijk. 2010/2011.
Superfamily Stromboidea. World Register of Marine Species.
http://www.marinespecies.org/aphia.php?p=taxdetails&id=23123
[Submitted 2010, partially published 2010/2011, not complete yet].
Last accessed: 4 Aug 2011.
Kronenberg, G.C., and H.G. Lee. 2007. Genera of American strombid
gastropods (Gastropoda: Strombidae) and remarks on their
phylogeny. The Veliger 49(4): 256–264.
Ladd, H.S. 1972. Cenozoic fossil mollusks from western Pacific
islands; Gastropods (Turritellidae through Strombidae). Geolog-
ical Survey Professional Paper 532: 1–79.
Lamarck, J.B.P.A. de Monet de, 1816. Tableau encyclope
´
dique et
me
´
thodique des trois re
`
gnes de la nature… Vingttroisie
`
me
partie. Mollusques et polypes divers. Vol. 1: Liste des objets
repre
´
sente
´
s dans les planches de cette livraison, 16 pp, pls. 1–95;
Vol. 2: pls. 96–314; Vol. 3: pls. 391–488 (incl. 431bis and
431bis*), Paris, Mme Veuve Agasse.
Lamarck, J.B.P.A. de Monet de, 1822. Histoire Naturelle des
Animaux sans Verte
`
bres, pre
´
sentant des characte
`
res ge
´
ne
´
raux
et particuliers de ces animaux, leur distribution, leurs classes,
leurs familles, leurs genres, et la citation des pricipales espe
`
ces
qui s’y rapportent, pre
´
ce
´
de
´
e 7: 1–711, Paris.
Landau, B.M., and C.M. da Silva. 2010. Early Pliocene gastropods of
Cubagua, Venezuela: taxonomy, palaeobiogeography and ecos-
tratigraphy. Palaeontos 19: 1–221.
Latal, C., W.E. Piller, and M. Harzhauser. 2005. Shifts in oxygen and
carbon isotope signals in marine molluscs from the Central
Paratethys (Europe) around the Lower/Middle Miocene transi-
tion. Palaeogeography, Palaeoclimatology, Palaeoecology 231:
347–360.
Latiolais, J.M. 2003. The phylogenetic underpinnings for spatial
patterns ofmorphological disparity: analyses using strombid
gastropods. Unpublished thesis. i–iv, 1–34, Louisiana State
University.
Latiolais, J.M., M.S. Taylor, K. Roy, and M.E. Hellberg. 2006. A
molecular phylogenetic analysis of strombid gastropod morpho-
logical diversity. Molecular Phylogenetics and Evolution 41:
436–444.
Lightfoot, J. 1786. A catalogue of the Portland Museum, lately the
property of the Duchess Dowager of Portland deceased Which
wil… i–viii, 1–194, London.
Linnaeus, C. 1758. Systema naturae, per Regna tria Naturae
secundum Classes, Ordines, Genera, Species, cum characteri-
bus, differentiis, synonymis, locis. Ed. 10 1: 1–823, Holmiae.
Lister, M. 1685–1692. Historiae sive synopsis methodicae Conchy-
liorum quorum omnium picturae. Issued in 4 books, total 8 p.
and 494 leaves of plates. London.
Lozouet, P., and P. Maestrati. 1986. Le Strombus granulatus
Swainson, 1822 une relique Mesogeenne. Xenophora, Bulletin
de l’Association Francaise de Conchyliologie 31: 11–15.
Mandic, O., M. Harzhauser, J. Schlaf, W.E. Piller, F. Schuster,
U. Wielandt-Schuster, J.H. Nebelsick, A. Kroh, F. Ro
¨
gl, and
P. Bassant. 2004. Palaeoenvironmental Reconstruction of an
Epicontinental Flooding - Burdigalian (Early Miocene) of the
Mut Basin (Southern Turkey). Courier Forschungsinstitut Senc-
kenberg 248: 57–92.
Moolenbeek, R.G., and H. Dekker. 1993. On the identity of Strombus
decorus and Strombus persicus, with the description of Strombus
decorus masirensis n. ssp. and a note on Strombus fasciatus. Vita
Marina 42(1): 3–10.
Mo
¨
rch, O.A.L. 1852. Catalogus Conchyliorum quae reliquit D.
Alphonso d’Aguirra & Gadea Comes de Yoldi, Regis danae
cubiculariorum princeps, ordinis dannebrogici in prima classe
& ordinis Caroli Tertii Eques (…) fasciculus primus. Cephalo-
phora (…). 1-172, Hafniae, Ludovicus Klein.
Mourik, A.A., H.A. Abels, F.J. Hilgen, A. Di Stefano, and W.J.
Zachariasse. 2011. Improved astronomical age constraints for the
middle Miocene climate transition based on high-resolution
stable isotope records from the central Mediterranean Maltese
Islands. Paleoceanography 26: PA1210. doi:10.1029/2010
PA001981.
Newton, R.B. 1905. Eocene Shells from Nigeria. Annals and
Magazine of Natural History. Serie 7(15): 83–91.
Newton, R.B. 1922. Eocene Mollusca from Nigeria. Geological
Survey of Nigeria Bulletin 3: 1–114.
Nikolov, P.I. 1993. Some molluscs from the Badenian (middle
Miocene) west of Pleven (Central northern Bulgaria). I. Gastrop-
oda: orders Archaeogastropoda and Mesogastropoda. Geologica
Balcanica 23(6): 61–72.
Popov, S.V., F. Ro
¨
gl, A.Y. Rozanov, F.F. Steininger, I.G. Shcherba,
and M. Kova
´
c
ˇ
. 2004. Lithological-Paleogeographic maps of
Paratethys. 10 Maps Late Eocene to Pliocene. Courier Fors-
chungsinstitut Senckenberg 250: 1–46.
Rafinesque, C.S. 1815. Analyse de la nature ou tableau d’universet
des corps organise
´
s. [portrait] [title page] [dedication] [1]
5–224, incl. additions and corrections. Jean Barravecchia,
Palermo.
Ro
¨
ding, P.F. 1798. Museum Boltenianum sive catalogus cimeliorum e
tribus regnis naturae que olim collegerat Joa. Fried Bolten,
M.D.p.d. per XL annos proto physicus Hamburgensis. Pars
Secunda continens Conchylia sive Testacea univalvia, bivalvia
G. C. Kronenberg, M. Harzhauser
123
& multivalvia. I-VIII, 1–199, Hamburg, Johan. Christi. Trappii.
[reprint American Malacological Union, 1986].
Ro
¨
gl, F. 1998. Palaeogeographic Considerations for Mediterranean
and Paratethys Seaways (Oligocene to Miocene). Annalen des
Naturhistorischen Museums in Wien 99A: 279–310.
Ro
¨
gl, F., W. Antl-Weiser, F. Brandsta
¨
tter, M.D. Dermitzakis, W.
Papesch, W.E. Piller, O. Schultz, N.K. Symeonidis, M.V. Triant
Aphyllou, and V. Tsapralis. 1997. Late Pleistocene marine
cycles in Corfu. Annales Ge
´
ologiques des Pays Helle
´
niques 37:
663–767.
Rumphius, G.E. 1705. D’Amboinsche rariteirkamer, Behelzende eene
Beschryvinge van allerhande zoo weeke als harde schaaldieren,
te weeten Krabben, Kreeften, en diergelyke Zeedieren, alsmede
allerhande Hoorntjes en Schulpen, die men in d’Amboinsche Zee
vindt: Daar beneven zimmige Mineraalen, Gesteenten, en
soorten van Aarde, die in d’Amboinsche, en zommige omlegg-
ende Eilanden. Verdeelt in drie boeken, En met nodige
Printverbeeldingen, alle naar het leven getekent, voorzien.
Beschreven door (…). i-xxviii, 1–340 [43], pls 1–60,
Amsterdam.
Sacco, F. 1893. I Molluschi dei terreni terziarii del Piemonte e della
Liguria. Parte XIV. (Strombidae, Terebellidae, Chenopidae ed
Halidae). Memoire della Reale Accademia delle Scienze di
Torino 14: 1–38.
Shevenell, A.E., J.P. Kennett, and D.W. Lea. 2004. Middle Miocene
Southern Ocean cooling and Antarctic cryosphere expansion.
Science 305: 1766–1770.
Strausz, L. 1966. Die Mioza
¨
n-Mediterranen Gastropoden Ungarns,
1–692. Budapest: Akade
´
miai Kiado
´
.
Swainson, W. 1821. Exotic Conchology; or figures and descriptions
of rare, beautiful, or undescribed shells, drawn on stone from the
most selected specimens; the descriptions systematically
arranged on the principles of mm. Cuvier and De Lamarck,
with references to the Linnaean calassification (…). Vol. 1
[12] ? 16 pls. London.
Swainson, W. 1822. Appendix. Description of several new shells, and
remarks on others, contained in the collection of the late Mrs.
Bligh. [in] A catalogue of the rare and valuable shells, which
formed the celebrated collection of the late Mrs. Bligh 20 pp.
London.
Trew, A. 1992. Henry and Arthur Adams’s new molluscan names,
1–63. Wales, UK: National Museum of Cardiff.
Tro
¨
ndle
´
, J., and B. Salvat. 2010. La thanatoce
´
nose du lagon de l’atoll
Niau (Polyne
´
sie franc¸aise) avec la description d’une nouvelle
espe
`
ce de Strombus (Mollusca, Gastropoda, Strombidae). Zoo-
systema 32: 613–623.
Tryon, G.W. 1885. Family Strombidae. Manual of Conchology 2(7):
99–152, pls 1–12.
Vredenburg, E. 1925. Description of Mollusca from the Post—Eocene
Tertiary formations of North - Western India. 1. Memoirs of the
Geological Survey of India 50: 1–350.
Williams, S.T., and T.F. Duda. 2008. Did tectonic activity stimulate
speciation in the Oligo-Miocene Indo-West Pacific? Evolution
62: 1618–1634.
Zachos, J., M. Pagani, L. Sloan, E. Thomas, and K. Billups. 2001.
Trends, rhythms, and aberrations in global climate 65 ma to
present. Science 292: 686–693.
Zazo, C., J.L. Goy, and E. Aguierre. 1984. Did Strombus survive the
last interglacial in the western Mediterranean Sea? Mediterra-
nea, seria Geologica 3: 131–137.
A new Middle Miocene European strombid genus
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