A new Janulus species - Società Paleontologica Italiana

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A new Janulus species - Società Paleontologica Italiana
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Bollettino della Società Paleontologica Italiana, 50 (3), 2011, 165-173. Modena, xxx 2011
A new Janulus species (Gastropoda, Pulmonata, Gastrodontidae)
from the Zanclean (early Pliocene) of Tuscany (central Italy)
Giuseppe Manganelli, Ivan Martini & Andrea Benocci
G. Manganelli, Dipartimento di Scienze Ambientali, Università di Siena, Via Mattioli 4, I-53100 Siena; [email protected]
I. Martini, Dipartimento di Scienze della Terra, Università di Siena, Via Laterina 8, I-53100 Siena; [email protected]
A. Benocci, Dipartimento di Scienze Ambientali, Università di Siena, Via Mattioli 4, I-53100 Siena; [email protected]
KEY WORDS - Gastropods, Pliocene, Zanclean, Tuscany, Janulus spadinii n. sp.
ABSTRACT - Janulus spadinii n. sp. (Gastropoda, Pulmonata, Gastrodontidae) is described herein from the Zanclean (early Pliocene)
of Balze di Caspreno (Tuscany). It belongs to a Macaronesian genus occasionally reported from the Tertiary of Europe. The new species is
very similar to the Recent Madeiran Janulus stephanophorus by virtue of its shape, dorsal sculpture and radial rows of palatal teeth. It is also
apparently similar to some fossil European species, however, due to insufficient information regarding the latter, no well supported distinction
may be proposed at present. A brief survey of all the nominal taxa of the species-group assigned to Janulus is also provided.
RIASSUNTO - [Una nuova specie di Janulus (Gastropoda, Pulmonata, Gastrodontidae) dal Pliocene inferiore della Toscana (Italia centrale)]
- Janulus spadinii n. sp. è descritto sulla base di materiale raccolti nelle argille continentali del Pliocene inferiore delle Balze di Caspreno
(Toscana). La nuova specie appartiene ad un genere attualmente esclusivo della Macaronesia, ma conosciuto per il Terziario europeo. Janulus
spadinii n. sp. è molto simile a Janulus stephanophorus, una specie vivente a Madeira (le due specie condividono: forma, scultura dorsale
e file radiali di denti palatali) ma ricorda anche alcune entità descritte per il Terziario europeo. Una differenziazione adeguata rispetto a
questi taxa è al momento impossibile, mancando del tutto informazioni sulla struttura dei loro denti palatali. Conclude il lavoro l’elenco di
tutti i taxa nominali assegnati a Janulus, inclusivo di una breve discussione sul loro status.
INTRODUCTION
The genus Janulus Lowe, 1852 is regarded as a
European palaeoendemite of the Tertiary warm climatic
phases, associated with moist forest biotopes (Esu, 1999).
It survives in the Macaronesian refuges (Walden, 1983,
1984; Cameron et al., 2007) and includes only three Recent
species: J. bifrons (Lowe, 1831) and J. stephanophorus
(Deshayes, 1850) from Madeira and the probably recently
extinct J. pompilius (Shuttleworth, 1852) from La Palma,
Canary Islands (Riedel, 1980; Fontaine et al., 2007).
Another Madeiran species, J. calathoides (Lowe, 1863),
tentatively added by Riedel (1998), is actually a species
of Discus Fitzinger, 1833 (Bank et al., 2002). However,
there is no definitive evidence that the three species really
belong to the same genus, because only one of them (J.
bifrons) is known anatomically. Janulus has a holopod
foot and distal genitalia with a penial dart-sac and an
accessory duct connecting penis and free oviduct (Pilsbry,
1947). Holopod snails with apertural palatal teeth, a
penial dart-sac and an accessory duct connecting penial
complex and female genitalia (duct of bursa copulatrix or
free oviduct) are assigned to the gastrodontids, a mainly
Nearctic family including a few other genera besides
Janulus [Gastrodonta Albers, 1850, Nastia Riedel, 1989,
Poecilozonites Boettger, 1884, Pilsbryna Baker, 1929,
Striatura Morse, 1864 and Zonitoides Lehmann, 1862
(Riedel, 1998; Slapcinsky & Cole, 2004)]. Although
the accessory duct joining penial complex and female
genitalia has been regarded as a unique autapomorphy
supporting the monophyly of the gastrodontids (Hausdorf,
1998), a robust phylogeny of these land snails is still
ISSN 0375-7633
lacking. Moreover, this duct may not be homologous: it is
absent in Pilsbryna and connects penial dart-sac and duct
of bursa copulatrix in Gastrodonta, penis and free oviduct
in Janulus, and penis and duct of bursa copulatrix in all the
others (however, in Zonitoides, a second accessory duct
joins duct of bursa copulatrix and free oviduct according
to Pilsbry, 1946; Riedel, 1980; Gittenberger et al., 1984).
Consequently the monophyly of the family, as currently
conceived, is not certain, like the number of genera/
subgenera it includes (see different accounts by Hausdorf,
1998; Riedel, 1998; Schileyko, 2003; Slapcinsky & Cole,
2004).
Recent Janulus species have a discoidal to trochiform
shell, angled or shouldered at periphery, with small
umbilicus, narrow ribbing on dorsal surface of teleoconch
whorls and peristome simple and not reflexed. One
species (J. bifrons) is larger, with less tightly coiled and
less narrowly ribbed whorls and peristome internally
thickened (it strongly recalls the shells of hygromiids
such as Trochulus Chemnitz, 1786, Monachoides Gude
& Woodward, 1921, etc.). Finally, at least one species (J.
stephanophorus) has radial rows of palatal teeth at 120
degrees to each other, each row consisting of three-four
teeth: one adaxial, very small and knob-like, one medial,
short and pleat-like and one abaxial, larger and inverted
comma-like or sometimes V-like, split into a longer outer
branch and a shorter inner branch (Pl. 1, fig. 9).
Although the fossil record is much poorer than it
was formerly thought (see for example Wenz, 1923),
sure reports spanning from the Oligocene (e.g. Janulus
schottleri Wenz, 1922) to the end of the Pliocene and
ecology of Recent species (Seddon, 2008) are in line with
doi:10.4435/BSPI.2011.15
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Bollettino della Società Paleontologica Italiana, 50 (3), 2011
Fig. 1 - Location map of the study area. a) Structural setting of southern Tuscany. b) Geological sketch of the Montaperti-Pianella area. Numbers
(1-6) indicate sites sampled during the 1990s. Most of these outcrops are now covered by recent alluvial deposits, vegetation or dumped rubble.
the palaeoecological meaning attributed to the genus in
the literature.
In this paper we describe a new Janulus species from
the Zanclean of Balze di Caspreno (Tuscany), already
reported as Janulus suttonensis (Wood, 1872) by De
Stefani (1880). We also shortly survey all the nominal
taxa of the species-group assigned to this genus, many of
which may belong to other groups of land snails.
geological setting
The Siena Basin is a post-collisional basin of the inner
Northern Apennines, a collisional belt formed during
the Cenozoic in response to the interaction between the
Adria and the Corso-Sardinian microplates (Carmignani
et al., 2001). It corresponds to approximately the middle
part of an elongated depression (Fig. 1a) extending for
about 200 km in a NNW-SSE direction. The tectonic
setting responsible for the origin and evolution of this
basin is still debated, and has been explained in different
ways: according to many authors (Costantini et al., 1982;
Martini & Sagri, 1993; Pascucci et al., 1999; Carmignani
et al., 2001; Brogi, 2011) its origin is connected with an
extensional regime acting until the late Miocene, while
other researchers postulate a dominantly compressional
tectonic setting (Boccaletti & Sani, 1998; Finetti et al.,
2001; Bonini & Sani, 2002).
Regardless of its structural origin, the Siena Basin
was interested by a continental and marine deposition
during the Neogene, and to limited and discontinuous
sedimentation of alluvial deposits (Aldinucci et al., 2007)
and travertine (Brogi et al., 2010) during the Quaternary.
Its Neogene basin-fill history dates back to the latest
G. Manganelli et alii - A new species of Janulus from Zanclean of Tuscany
Miocene (Messinian) with the deposition of a thick
fluvio-lacustrine succession that unconformably overlies
pre-Neogene bedrock. Most of these Miocene deposits
are buried beneath Pliocene sediments and crop out in
limited western marginal areas of the basin (Lazzarotto
& Sandrelli, 1977; Bossio et al., 2000). Marine conditions
in the Siena Basin began at the base of the Zanclean
(Costantini et al., 1982; Bossio et al., 1993), with the
deposition of a thick sedimentary succession, ranging
from nearshore (sands and conglomerates) to offshore
(mud) environments (Costantini et al., 1982; Manganelli
et al., 2010; Martini et al., 2011). Marine sedimentation
persisted until the late Piacenzian, when uplifting of
southern Tuscany caused emergence of the Siena Basin
(Costantini et al., 1982; Bossio et al., 1993). Local
episodes of continental sedimentation have been reported
within the marine deposits in the southwestern sector of
the basin (Bossio et al., 1992, 1993).
Janulus material was collected along the Arbia
River (Montaperti-Pianella area, northern sector of
the Siena Basin), where continental deposits crop out
discontinuously in small scattered outcrops (Fig. 1b),
most of which are now covered by recent alluvial deposits,
vegetation or rubble from quarries. The best exposures
of continental deposits crop out in natural cliffs on the
left bank of the Arbia river (the “Balze di Caspreno”,
fossiliferous sites 5 and 6 in Fig. 1b). According to
Martini et al. (2011) these sediments are indicative of a
generic fluvial setting, with dominantly clayey floodplain
silts (locally containing peat horizons) and fine sands,
with subordinate lens-shaped bodies of fluvial gravelly
sand. The age of these deposits has long been debated:
De Stefani (1880), who first described this fossiliferous
site, attributed these deposits to the lower part of the
Pliocene, on the basis of lithological and palaeontological
similarities with analogous sediments exposed close to
Siena, where they are interbedded with certainly Pliocene
marine strata (De Stefani, 1876). De Stefani (1880), and
later De Castro & Pilotti (1933), also described fossil
remains of brackish gastropods in these muddy sediments,
implying connection with a nearby marine area. Since
marine conditions in the Siena Basin started in the earliest
Pliocene, De Stefani (1880) and De Castro & Pilotti (1933)
took this fact as an indirect evidence of the Pliocene age
of these deposits. The deposits of the Balze di Caspreno
area were later attributed to a generic Neogene (Signorini,
1966, 1967) and recently assigned to the Messinian
(Lazzarotto et al., in press), due to strong lithological
similarities with the “Argille del Casino” Fm. cropping out
in the nearby Casino Basin (Lazzarotto & Sandrelli, 1977).
Partially in contrast with this attribution, Manganelli et al.
(2007) reported some continental molluscs (Strobilops cf.
romani Wenz, 1915 and Eostrobilops aloisii Manganelli,
Delle Cave & Giusti, 1989), which in Europe have been
recorded only from the Zanclean.
Finally, Martini et al. (2011) reported the discovery
of cobbles showing evidence of bio-erosion (e.g.
borings) in gravelly fluvial facies, indicating provenance
from recycled marine nearshore gravels. According to
the geological history of the Siena Basin (Costantini
et al., 1982; Lazzarotto et al., in press) these cobbles
could only be derived from unknown Pliocene deposits
or at least from the Serravallian Ponsano Sandstone
167
Fm. exposed in the northern Casino Basin (Bossio et
al., 1998). However, micropalaeontological analysis
carried out in fine floodplain sediments (Bambini, pers.
comm.) shows a lack of recycled Serravallian calcareous
nannofossils, while bedrock-derived nannofossils are
common. These findings suggest that the deposits date
back to the Pliocene. According to Martini et al. (2011),
integrated palaeontological and stratigraphical analysis
of the continental and marine succession exposed in the
surrounding area also enables assignment of these deposits
to the MPl3 Zone of planktonic foraminifera zonation
(zonal scheme of Foresi et al., 2001).
SYSTEMATIC PALAEONTOLOGY
Janulus spadinii n. sp.
(Pl. 1, figs 2-7)
Diagnosis - A species of gastrodontid Janulus with
shell very similar to that of Recent Madeiran Janulus
stephanophorus by virtue of its shape, dorsal sculpture and
radial rows of palatal teeth (three internal radial rows of
teeth at 120 degrees to each other; each row consisting of
three teeth, the more external of which larger and inverted
comma-like), but differing by virtue of its lesser size and
the more depressed shell.
Description - Shell (Pl. 1, figs 2-7) dextral, very small
in size, lenticular in shape, with about six, very narrow
and tightly coiled whorls, separated by deep sutures; last
whorl angled at periphery; umbilicus small, narrow and
cylindrical; aperture semilunar, with light parietal callus
and peristome not thickened or reflexed; protoconch
sculpture unknown; teleoconch with many strong, regular,
sligthly prosocline ribs, stopping just below periphery
(in one specimen: 54 ribs in first teleoconch whorl, 61 in
second, 63 in third, Pl. 1, fig. 2); internally, three radial
rows of palatal teeth at 120 degrees to each other; each
row consisting of three teeth: one adaxial, very small
and knob-like, one medial, small and knob-like and one
abaxial, larger and inverted comma-like.
Dimensions - The largest specimen, an incomplete
shell, is 4.2 mm wide. The holotype, a juvenile shell, is
1.4 mm high and 2.8 mm wide.
Type locality and horizon - Balze di Caspreno, along
Arbia River (Montaperti-Pianella area, east of Siena),
lacustrine grey clays of Zanclean age (MP13 Zone of
planktonic foraminifera zonation of Foresi et al., 2001).
For collecting sites, see Fig. 1.
Type material - Holotype (Pl. 1, fig. 2), a juvenile
shell from site 4 (Manganelli collection, Dipartimento
di Scienze Ambientali, University of Siena, no. 39588);
about 60 paratypes consisting of 110 shell fragments
(corresponding to at least 55 specimens) from Site
2 (Manganelli collection, Dipartimento di Scienze
Ambientali, University of Siena, no. 39587); 3 shells from
Site 4 (Manganelli collection, Dipartimento di Scienze
Ambientali, University of Siena, no. 39589); 7 shell
fragments (corresponding to at least 5 specimens) from
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Bollettino della Società Paleontologica Italiana, 50 (3), 2011
Site 6 (Manganelli collection, Dipartimento di Scienze
Ambientali, University of Siena, no. 39590).
Derivation of name - The new species is named
after our dear friend Valeriano Spadini from Lucignano
(Arezzo), passionate naturalist and scholar of fossil corals
and molluscs.
Remarks - Shell features of the species from the
Zanclean of Balze di Caspreno are sufficient to establish
that it really belongs to Janulus: it is very similar to the
Recent J. stephanophorus (the type species of the genus)
with which it shares shape, dorsal sculpture and radial
rows of palatal teeth.
In Fossilium Catalogus, Wenz (1923, pp. 300-306)
listed 12 Janulus species, one of which - Janulus striatus
(Eichwald, 1830) - only tentatively assigned to this genus.
Subsequently, Schlickum (1979), Schlickum & Geissert
(1980), Yü & Zhang (1982) and Harzhauser & Binder
(2004) described four other putative Janulus species
(see Appendix 1). Unfortunately information about
most of them, especially on the presence and structure
of the internal radial rows of palatal teeth, is scarce or
absent and this prevents sure generic assignment. Only
the late Miocene J. gottschicki (Jooss, 1912) and J.
schottleri Wenz, 1922 are known to have palatal teeth but
information on the former is inadequate (see Jooss, 1912).
On the contrary the pattern of palatal teeth of J. schottleri
is consistent with that of the Recent J. stephanophorus and
consequently its inclusion in Janulus is well supported.
Some other species may belong to Janulus on the basis
of their shell shape and dorsal sculpture, but we lack
information about radial rows of palatal teeth. They are:
J. angustiumbilicatus (Sacco, 1886, sensu Ferrero Mortara
et al., 1994), J. austriacus Harzhauser & Binder, 2004, J.
olisipponensis (Roman, 1907) and J. suttonensis (Wood,
1872). On the contrary nothing sure can be said about all
the other species, most of which may belong to completely
different genera. For example, according to Lueger (1981),
J. joossi Schlickum, 1970 is a species of Discus (Discidae),
namely Discus pleuradras (Bourguignat, 1881).
Among fossil species which belong or may belong
to this genus, Caspreno Janulus matches the alleged
syntype of Patula angustiumbilicata illustrated by Ferrero
Mortara et al. (1984: Pl. 54, figs 1a-b), Janulus austriacus
Harzhauser & Binder, 2004 (Pl. 7, figs 12-15) from the late
Miocene of Lower Austria, Helix suttonensis Wood, 1872
(Pl. 1, figs 2a-c) from the Pliocene of England and Patula
olisipponensis Roman, 1907 (fig. 8) from the Neogene
of Portugal. Unfortunately, at the present state of the art,
no well supported distinction can be proposed until more
detailed descriptions of these taxa become available. As
far as we know, it differs from Janulus suttonensis, of
which we examined a detailed drawing of the holotype by
Gordon Riley, kindly made available by Richard Preece,
and from Janulus austriacus by virtue of its lenticular
shape (trochiform shape in Janulus suttonensis; discoidal
shape in Janulus austriacus). It is very similar to the shells
of “Janulus” angustiumbilicatus published by Ferrero
Mortara et al. (1984), but they are different from Sacco’s
description and figures of this taxon (see Appendix).
Finally it differs from Janulus olisipponensis, on the basis
of Roman’s (1907) figure only, by virtue of its lenticular
shape and slightly prosocline ribs (discoidal shape and
markedly prosocline ribs in Janulus olisipponensis).
AcknowledgEments
Helen Ampt revised the English. Mikaela Bernardoni
and Florence Conti (Library of the Faculty of Sciences of the
University of Siena, Italy) helped with bibliographic research.
Elena Gavetti and Daniele Ormezzano (Museo Regionale di
Scienze Naturali, Turin, Italy) supported with information on L.
Bellardi and F. Sacco’s type-material, Richard Preece (University
EXPLANATION OF PLATE 1
Fig. 1
-
Janulus stephanophorus (Deshayes, 1850). SEM apical view of a shell from Ilha da Madeira: Ribeira da Porto Novo, A.H.
Walden leg. 11.11.1972 (Göteborgs Naturhistoriska Museum 72-13.386); scale bar: 2 mm.
Figs 2-7 - Janulus spadinii n. sp. from Balze di Caspreno.
2 - Holotype (a juvenile shell) from Site 4 (Manganelli collection, Dipartimento di Scienze Ambientali, University of Siena, no.
39588); SEM apical (a) and apertural (b) views; scale bar: 2 mm.
3 - A juvenile shell from Site 4 (Manganelli collection, Dipartimento di Scienze Ambientali, University of Siena, no. 39589); SEM
apical view; scale bar: 2 mm.
4-7-Shell fragments from Site 2 (Manganelli collection, Dipartimento di Scienze Ambientali, University of Siena, no. 39590); SEM
views of palatum illustrating the internal radial rows of palatal teeth; scale bar: 2 mm.
Fig. 8
-
Janulus bifrons (Lowe, 1831): apical (a), apertural (b) and umbilical (c) views of a shell from Ilha da Madeira: Garajau, A.H.
Walden leg. 2.4.1983 (Göteborgs Naturhistoriska Museum 83-16.180); scale bar: 4 mm.
Fig. 9
-
Janulus stephanophorus (Deshayes, 1850): apical (a), apertural (b) and umbilical (c) views of a shell from Ilha da Madeira:
Ribeira da Porto Novo, A.H. Walden leg. 11.11.1972 (Göteborgs Naturhistoriska Museum 72-13.386); scale bar: 4 mm.
Fig. 10
-
Janulus pompilius (Shuttleworth, 1852): apical (a), apertural (b) and umbilical (c) views of a shell from La Palma, Canary
Islands, B.F. Blauner leg. 1851 (syntype, Naturhistorisches Museum Bern 18777). Photo by Eike Neubert (© Centre Suisse
de Cartographie de la Faune, reproduced by permission); scale bar: 2 mm.
G. Manganelli et alii - A new species of Janulus from Zanclean of Tuscany
169
Pl.
1
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Bollettino della Società Paleontologica Italiana, 50 (3), 2011
Museum of Zoology, Cambrdrige, UK) with a drawing of the
holotype of Janulus suttonensis, Henrik Walden and Ted von
Proschwitz (Göteborgs Naturhistoriska Museum, Göteborg,
Sweden) loaned material, Eike Neubert (Naturhistorisches
Museum der Burgergemeinde Bern, Switzerland) provided photos
of a syntype of Janulus pompilius, publication of which was
approved by GBIF Switzerland (GBIF Switzerland, Centre Suisse
de Cartographie de la Faune, Neuchâtel).
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Editor Alessandro Ceregato
Appendix - putative fossil Janulus species
Patula (Janulus) angustiumbilicata Sacco, 1886: 190-1, Pl. 1, figs
17a-b.
Type locality: “Alluvioni plioceniche […] di Fossano” (Sacco,
1886: 169).
Status: uncertain. According to Sacco’s description and figures,
the species has a punctiform umbilicus, but Ferrero Mortara et
al. (1984: Pl. 54, figs 1a-c) figured a putative syntype with a
small umbilicus. The material reported by Ferrero Mortara et al.
(1984) consists of about ten specimens, all consistent with the one
illustrated. However, there is no evidence that they constitute the
actual type material because the only label accompanying them dates
back to the 1970s (Daniele Ormezzano, pers. comm., September
2010). Although no information is available on radial rows of palatal
teeth, the specimen illustrated by Ferrero Mortara et al. (1984) is
similar to the Recent Janulus stephanophorus (Deshayes, 1850)
and the Caspreno species.
Janulus austriacus Harzhauser & Binder, 2004: 23, Pl. 7, figs 12-15.
Type locality: “Richardhof (sample RH A/2) close to Mödling
in Lower Austria (Vienna Basin); marls deposited in a Pannonian
(late Miocene) swampy lake adjacent to Lake Pannon; mammal
zone MN9.”
Status: probably a Janulus species: although no information
is available on radial rows of palatal teeth, it is very similar to the
Recent Janulus stephanophorus (Deshayes, 1850).
Janulus decorosus Yü & Zhang, 1982: 64-5, Pl. 7, figs 4-6.
Type locality: Baoyue formation, Sanshui Basin, Guangdong;
Middle Eocene.
Status: uncertain. Based on very poorly preserved specimens.
Patula densestriata Klika, 1891: 40, figs 32a-c.
Type locality: “bei Wärzen” (Bohemia).
Status: uncertain. Probably a Janulus species: although no
information is available on radial rows of palatal teeth, it is very
similar to Janulus schottleri Wenz, 1922.
Helix disparilis Deshayes, 1863: 818, Pl. 52, figs 12-15.
Type locality: “Côte Saint-Martin, près d’Etampes … Calcaire
de Beauce.”
Status: uncertain. Probably not a Janulus, but a discid species.
Janulus germainae Schlickum & Geissert, 1980: 237-8, Pl. 14,
fig. 31.
T ype locality : “Oberes Pliozän, Triptychia-Mergel …
Kiesgrube Mary Kocher, Sessenheim, Unterelsass.”
Status: uncertain. Probably not a Janulus, but a discid species.
Niederhöfer & Falkner (2004) stated that the type material could
include more than one species.
Patula gottschicki Jooss, 1912: 32-3, Pl. 2, fig. 2.
Type locality: “Pharion’s Sandgrube in Steinheim, Pupaschicht.”
Status: uncertain. Jooss’s description and figure are consistent
with a Janulus species (especially due to palatal teeth), but reexamination is needed.
Helix gyrorbis Klein, 1846: 72, Pl. 1, figs 14a-c.
Type locality: “… bei Oepfingen in der Nähe von Ehingen...”
Status: uncertain. Probably a Discus species. Sandberger (1872:
Pl. 21, figs 7a-7b) depicted a completely different species.
Janulus joossi Schlickum, 1979: 410 (depicted by Schlickum, 1978:
Pl. 19, fig. 17).
Type locality: “Oberpannon, Süsswassermargel … Öcs,
Komitat Veszprém, Ungarn.”
Status: uncertain. According to Lueger (1981) a junior synonym
of Discus pleuradras (Bourguignat, 1881).
Janulus mörsingensis Jooss, 1918: 289-90.
T ype locality : “… unteren Obermiocän – den SylvanaSchichten – von Mörsingen.”
Status: uncertain. Probably not a Janulus, but a punctid or a
discid species. According to Lueger (1981), the specimen assigned
to this species by Schlickum (1978: Pl. 19, fig. 16) belongs to Discus
pleuradras (Bourguignat, 1881).
Patula (Janulus) olisipponensis Roman, 1907: 82-3, text-fig. 8.
Type locality: “Casaes de Valle d’Obidos près Rio Maior. –
Etage Pontique.”
Status: probably a Janulus species: although no information
is available on radial rows of palatal teeth, it is very similar to the
Recent Janulus stephanophorus (Deshayes, 1850).
Helix rysa Wood, 1848: 4, Pl. 1, figs 1a-c.
Type locality: “Red Crag, Walton Naze.”
Status: uncertain. First assigned to Janulus by Sandberger
(1875), it recalls J. bifrons from which it differs by virtue of a
wider umbilicus, more reflexed peristome and blunt peripheral keel
(absent in J. bifrons).
Janulus schottleri Wenz, 1922: 48, text-figs 1a-d.
T ype locality : “Häufig in den Süsswasserschichten am
Streitberg bei Treis a. d. Lumda und bei Ilschhausen.”
Status: very similar to Janulus stephanophorus (Deshayes,
1850). The species has radial row of palatal teeth at 90 degrees to
each other; each row consisting of three teeth: adaxial and medial
teeth in the form of a short fold; abaxial tooth larger and V-like.
Apparently very similar to Patula densestriata Klika, 1891.
Helix striata Eichwald, 1830: 215, 300 (depicted by Eichwald,
1852: Pl. 11, figs 9a-d).
T ype locality : “près de Holowczynce, aux environs de
Mendzibosh, dans un calcaire tertiaire d’eau douce” (Eichwald,
1853).
Note: junior primary homonym of Helix striata Müller, 1774,
now Helicopsis striata (Pulmonata, Hygromiidae).
Status: uncertain. Eichwald’s (1852-53) description and figures
may be consistent with a Janulus species, but re-examination is
needed.
G. Manganelli et alii - A new species of Janulus from Zanclean of Tuscany
Patula (Janulus) supracostata Sandberger, 1872: Pl. 29, figs 2a-2c
(described by Sandberger, 1875: 584).
Type locality: “Vermes bei Delsberg (Ct. Bern), Birk bei
Mörsingen (v. Zell) Georgsgemünd und Hasenmühle bei Eichstädt
(Gümbel) im Kalke mit Helix sylvana, Undorf bei Regensburg in
gleichaltem Braunkohlen-Thone (Gümbel), an beiden Orten selten”
(Sandberger, 1875).
Status: uncertain. Sandberger’s description and figure may be
consistent with a Janulus species, but re-examination is needed.
According to Moser et al. (2009) the specimen figured by Sandberger
173
(1872), representing the holotype, cannot be identified. However
they thought that it may belong to the “Discus-convergent Janulus
germainae group”.
Helix suttonensis Wood, 1872: 2, Pl. 1, figs 2a-c.
Type locality: “Coralline Crag, Sutton.”
Status: probably a Janulus species: although no information
is available on radial rows of palatal teeth, it is very similar to the
Recent Janulus stephanophorus (Deshayes, 1850).