GeolCarp_Vol66_No2_139

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, APRIL 2015, 66, 2, 139—156 doi: 10.1515/geoca-2015-0016

Introduction and taxonomic history

The  Turiec  Basin,  located  in  the  Western  Carpathians  in
northern  Slovakia  (Fig. 1),  harboured  a  closed,  long-lived
freshwater lake during the Late Miocene (Pipík et al. 2012).
Data quality about its fauna and flora is, however, quite var-
ied. The best known fossil group is the ostracods, documented
in  several  successive  papers  by  Pipík  (2002,  2004,  2005),
Pipík & Bodergat (2003a,b, 2004a,b, 2006, 2007, 2008), and
Pipík et al. (2012). The fauna revealed a high degree of ende-
micity, which makes the Lake Turiec a particularly interest-
ing  research  field  and  a  potential  equivalent  to  comparable
extant and fossil long-lived systems.

In contrast, the published data on the mollusc fauna of

Lake Turiec is mostly imprecise and contains many misiden-
tifications. The first work dates back to the 19

century and

only  mentions  the  bivalve  Congeria  triangularis  (Andrian
1865).  Based  on  determinations  by  Štúr  (1860)  –  Andrian
(1866)  gave  more  detailed  information  and  additionally  re-
corded  Planorbis  pseudoammonius,  Paludina  sadleri,  and
unidentifiable  remains  of  the  genera  Paludina,  Planorbis,
Melanopsis, and Valvata from the Bystrička locality (see also
Vetters  1910).  Half  a  century  later,  Vigh  (1915)  mentioned
viviparids, melanopsids, neritids, hydrobiids, planorbids, and
dreissenids  from  Martin  brickyard.  After  determinations  by
Wenz, who compared the fauna with the much older Moravian

Upper Miocene endemic lacustrine gastropod fauna of the

Turiec Basin: addressing taxonomic, paleobiogeographic

and stratigraphic issues

THOMAS A. NEUBAUER

, MATHIAS HARZHAUSER

and RADOVAN PIPÍK

Geological-Paleontological Department, Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria;

thomas.neubauer@nhm-wien.ac.at; mathias.harzhauser@nhm-wien.ac.at

Geological Institute, Slovak Academy of Sciences, Branch: Ďumbierska 1, 974 01 Banská Bystrica, Slovak Republic; pipik@savbb.sk

(Manuscript received September 11, 2014; accepted in revised form March 12, 2015)

Abstract: The present work displays the first detailed taxonomic study on the freshwater gastropod fauna of the Upper
Miocene Lake Turiec. Apart from several mentions of species and genus names in the literature, the mollusc fauna has
been poorly studied up to now. Some of the cited genera implied peculiar paleobiogeographic relationships, urging a
taxonomic investigation to either prove or revise such arising claims. Variable degrees of preservation, however, lim-
ited the possibility to identify all the fossils at species level. The fauna includes at least ten species, of which five turned
out to be new to science. Four of those were sufficiently well preserved to be described as new species, namely Viviparus
pipiki Neubauer & Harzhauser nov. sp., Melanopsis glaubrechti Neubauer & Harzhauser nov. sp., Tournouerina
turiecensis Neubauer & Harzhauser nov. sp., and Radix kovaci Neubauer & Harzhauser nov. sp. Additionally, the new
genus Popovicia Neubauer & Harzhauser nov. gen. is introduced for the primary homonym Metohia Popović, 1964 non
Absolon, 1927. Most importantly, this taxonomic study revises many of the names cited in the literature and proves
most of the alleged paleobiogeographic relationships wrong. The only biogeographic and stratigraphic surprise is the
record of Popovicia cf. compressa, a species described from lower Pliocene deposits of the Metohia Basin in Kosovo.
The majority of the fauna, however, has only been documented for the Turiec Basin, once more confirming the high
degree of its endemicity. The faunal relationships indicate a latest Middle to early Late Pannonian (Middle to Late
Tortonian) age, which is in agreement with available age models.

Key words: freshwater molluscs, taxonomy, endemicity, paleobiogeography, stratigraphy, Western Carpathians, Neogene.

“Oncophora  beds”  (today  known  as  the  Rzehakia  beds,
Čtyroký et al. 1973), Remeš (1923) recorded the species Vivi-
parus  oncophorae,  Melanoptychia  pseudoscalaria,  Theo-
doxis (Neritodonta) aff. crenulata, Hydrobia sp., Theodoxis
sp.,  Congeria  subclaviformis,  and  Oncophora  sp.  from  the
Hrby  locality  near  Bystrička.  Additionally  he  mentioned
Stalioia gracilis from the Dolina locality near Martin.

The first “real” taxonomic work with synonymy lists, short

descriptions  and  illustrations  was  performed  by    Andrusov
(1954).  He  clearly  discussed  previous  misidentifications  and
partly  synonymized  them  with  the  species  observed  by  him-
self, namely Succinea (Amphibina) primaeva, Viviparus neu-
mayri,  Melanoptychia  sp.,  Theodoxus  (Calvertia)  crenulatus
crenulatus,  and  Congeria  sp.  ex  gr.  C.  ornithopsis.  In  the
same  year  Pokorný  (1954)  reported  a  few  terrestrial  gastro-
pods and a sphaeriid bivalve with determinations on genus or
family level only. Further studies were carried out by A. On-
drejičková  and  M.  Rakús,  who  produced  manuscripts  that
were unfortunately never published (see Kováč et al. 2011).

Some genus names deriving from published as well as un-

published  data  were  mentioned  by  Kováč  et  al.  (2011)  and
Pipík et al. (2012), namely Viviparus, Theodoxus, Melanop-
sis, Hydrobia, Pyrgula,  Lymnaea, Ancylus, Gyraulus, Plan-
orbis,  Kosovia,  Congeria,  Unio,  and  Anodonta.  Especially
the record of the genus Kosovia was eye-catching, since it is
a known endemite for basins in Kosovo and southern Serbia.

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Krstić  et  al.  (2012)  determined  the  species  as  Kosovia  stro-
phostomopsis  Milošević,  1965,  which  was  originally  de-
scribed from the Late Pontian (Early Pliocene) of the Metohia
Basin.  Additionally  they  mentioned  Lithoglyphus  nanus
Roshka, 1973, a Maeotian species of the region SE of Odessa.

In summary, the few papers mentioning molluscs from the

Turiec Basin produce quite a motley collection of names de-
riving from various stratigraphic levels and geographical re-
gions.  On  the  one  hand,  many  of  these  names  might  raise
doubts  about  the  identifications,  given  the  deviating  paleo-
biogeographic distribution. On the other hand, if these mol-
luscs  were  correctly  determined,  these  findings  would
reshuffle  parts  of  the  Late  Miocene  paleobiogeographic
framework.  Consequently,  one  main  task  of  this  taxonomic
work  is  to  clarify  these  and  other  ambiguities  to  straighten
up the paleobiogeographic framework of the Late Miocene in
this region. The second point regards taxonomy itself – since

the fauna has never been studied in detail and rarely figured,
this paper presents an important contribution to our knowl-
edge of Miocene freshwater faunas and to our understanding
of endemic faunas in the Miocene of Europe.

Geological setting

The geological history of the Turiec Basin is well under-

stood,  supported  by  data  from  structural  geology,  sedimen-
tology,  paleoecology,  geochronology,  geophysics,  and
geomorphology  (Buday  1962;  Gašparík  et  al.  1974,  1995;
Konečný  et  al.  1983;  Hók  et  al.  1998;  Rakús  et  al.  2005;
Kováč et al. 2011; Pipík et al. 2012; Králiková et al. 2014).
The basin is a 40 km long and 10 km wide westward-dipping
halfgraben system with a sedimentary infill of up to 1200 m
thickness. It is enclosed by the Krivánska Malá Fatra Mts in
the  north,  the  Lúčanská  Malá  Fatra  Mts  in  the  west,  the
Ve ká Fatra Mts in the east, and the Kremnické vrchy Mts in
the  southeast.  Its  southwestern  part  is  delimited  from  the
Horná  Nitra  Basin  by  the  Žiar  Mts.  The  basement  consists
mainly  of  the  Mesozoic  complexes  of  the  Central  Western
Carpathian  paleo-Alpine  tectonic  units  and  in  its  northern
part  also  of  Paleogene  post-nappe  sedimentary  cover.  The
basin became restricted during the Middle Miocene as a re-
sult  of  rapid  uplift  of  the  surrounding  mountain  chains  and
volcanic activity.

Neogene sedimentation started during the late Middle

Miocene.  The  transtensional  to  extensional  tectonic  regime
during  that  time  resulted  in  the  subsidence  of  the  southern
part of the basin. The initial phase of sedimentation is char-
acterized by a volcano-sedimentary complex in the southern
region, lacustrine fine-grained sedimentation in the northern
region, and subsequent alluvial fan deposits during the Late
Badenian  to  Early  Pannonian  (Early  Serravallian  to  Early
Tortonian – Kováč et al. 2011). The main part of the Neo-
gene sediments is represented by the lacustrine Martin For-
mation,  reaching  from  the  late  Middle  Miocene  up  to  the
earliest  Pliocene  (Kováč  et  al.  2011).  Increased  subsidence
along  the  uplifted  mountains  led  to  accumulations  of  its
clayey infill. Most of the known freshwater biota, including
the herein presented molluscs, derives from this unit, reflect-
ing the deposits of an isolated freshwater lake. On the basis
of the rich ostracod assemblages Pipík et al. (2012) defined
six  paleoecological  communities,  allowing  the  ecological
and  bathymetrical  zonation  of  Lake  Turiec  during  the  Late
Miocene.  Rapid  uplift  of  the  surrounding  mountain  chains
during the latest Miocene and Early Pliocene resulted in the
deposition  of  alluvial  fans  along  the  western,  southwestern,
and  northwestern  basin  margins.  This  is  followed  by  Pleis-
tocene alluvial and river terrace deposits, which cover large
parts of the basin at present. For a detailed reconstruction of
the sedimentary history of the basin see Kováč et al. (2011).

Material and methods

The studied material is stored in the collection of the

Múzeum Andreja Kme a in Martin, Slovak Republic. It de-

Fig. 1. Geographical sketch of the Turiec Basin in northern Slovakia
with indication of sampled localities (modified after Kováč et al.
2011; Pipík et al. 2012). Grey areas display larger villages, filled
circles the localities mentioned in the text.

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rives from drill cores and several outcrops sampled by Miloš
Rakús  in  the  1950s  and  1990s  in  the  Turiec  Basin  (Fig. 1).
Consequently  any  quantitative  study  is  not  possible.  Some
specimens were cleaned from sediment particles with an ul-
trasonic  device.  The  SEM-photos  were  produced  with  a
JEOL JSM-6610LV at the Natural History Museum Vienna
using an output voltage of 15 kV.

Systematic paleontology

The systematical arrangement follows Bouchet & Rocroi

(2005), Jörger et al. (2010), Criscione & Ponder (2013), and the
FreshGEN database (Neubauer et al. 2014b).

Class: Gastropoda Cuvier, 1795

Subclass: Neritimorpha Golikov & Starobogatov, 1975

Order: Cycloneritimorpha Frýda, 1998

Superfamily: Neritoidea Rafinesque, 1815

Family: Neritidae Rafinesque, 1815

Subfamily: Neritininae Poey, 1852

Genus: Theodoxus Montfort, 1810

Type species: Theodoxus lutetianus Montfort, 1810

(currently considered as a synonym of Theodoxus fluviatilis
(Linnaeus, 1758)). Recent, Europe. Type by original desig-
nation (Welter-Schultes 2012).

Theodoxus cf. postcrenulatus Papp, 1953

(Fig. 2A—C)

1923 Theodoxis (Neritodonta) aff. crenulata Klein – Remeš, p. 113
?cf. 1953 Theodoxus (Theodoxus) postcrenulatus n. sp. Papp, p. 96,

pl. 2, figs. 1—3

1954 Theodoxus (Calvertia) crenulatus crenulatus (Klein) – An-

drusov, p. 257 [non Neritina crenulata Klein, 1853]

?cf. 1997 Theodoxus (Theodoxus) postcrenulatus Papp, 1953 – For-

dinál, p. 267, pl. 1, fig. 1

?cf. 2004 Theodoxus postcrenulatus Papp – Harzhauser & Tempfer, p. 60

Material: 10 specimens from Martin brickyard, coll. nos.

SNM 9/2011 (PZ-696), SNM 12/2011 (PZ-699a-d); 1 speci-
men from Trebostovo (Drill core GT-11), coll. no. SNM
171/2006 (PZ-573); height: 5.5 mm, width: 6.5 mm (largest
specimen).

Description: Shell small, sturdy, with ca. 2.5 whorls.

Apex  very  low,  in  some  specimens  fully  immersed.  Apical
region slightly flattened to convex; grades over a highly con-
vex shoulder into the steep, straight base. Aperture inclined
with about 60°; perfectly semicircular in shape. Callus mod-
erately  thickened,  with  weakly  granulated  surface;  forms  a
shallow  ramp  towards  a  sharp,  densely  but  weakly  serrated
edge,  which  lies  below  the  plane  formed  by  the  aperture.
This  inclination  produces  weak  incisions,  where  the  callus
touches  the  apertural  margin  (adapically  as  well  as  abapi-
cally).  In  a  single  specimen  colouring  is  preserved  as  dark,
broad, widely-spaced zigzag lines on white ground.

Remarks: It is extremely difficult to distinguish the many

Theodoxus species-group taxa that have been introduced for
the Miocene of the Pannonian region (e.g. Handmann 1887;

Brusina 1892, 1902; Jekelius  1944;  Papp  1953).  Since many
of  these  were  established  based  on  different  colour  patterns
only, which has been shown to be not taxonomically relevant
(see e.g. Welter-Schultes 2012), many may actually represent
junior  synonyms  of  others.  The  present  species  fits  morpho-
logically  (as  well  as  concerning  the  colouring)  well  with  the
specimens described and illustrated by Papp (1953) as Theo-
doxus postcrenulatus. It corresponds in the presence of small
teeth and a weakly granulated callus. The main difference is
the  distinctly  larger  size  of  the  Austrian  specimens  (ca.
10 mm  –  Papp  1953).  Therefore,  the  identification  remains
somewhat uncertain. It could be that the few available speci-
mens are not fully grown individuals.

The present species differs from the similar T. intracarpati-

cus Jekelius, 1944, which has a more elongated shell, a more
thickened callus and less distinct teeth (if at all), and T. soceni
Jekelius, 1944, which has a higher apex and thus appears
bulkier (see also Harzhauser et al. 2002).

Andrusov (1954) mixed up the species with Theodoxus

crenulatus (Klein, 1853), a wide-spread species throughout the
Middle Miocene of Central Europe (Wenz 1929; Schlickum
1976; Bartha 1979; Binder 2003; Harzhauser et al. 2012),
which differs in its slightly more elevated spire.

Distribution: Up to now only recorded from Late Mio-

cene localities of the Vienna Basin (Götzendorf, Austria,
Pannonian F, Papp 1953; Pezinok, Slovakia, Pannonian E,
Fordinál 1997).

Subclass: Caenogastropoda Cox, 1960

Superfamily: Viviparoidea Gray, 1847

Family: Viviparidae Gray, 1847

Subfamily: Viviparinae Gray, 1847

Genus: Viviparus Montfort, 1810

Type species: Viviparus fluviorum Montfort, 1810 (cur-

rently considered as a synonym of Viviparus viviparus (Lin-
naeus, 1758)). Recent, Northern Eurasia, Europe, Anatolia
and Northern America. Type by original designation (Welter-
Schultes 2012).

Viviparus pipiki Neubauer & Harzhauser nov. sp.

(Fig. 2D—J)

1866 Paludina Sadleri – Andrian, p. 196 [non Vivipara sadleri Neu-

mayr, 1869]

1923 Viviparus oncophorae Rzehak – Remeš, p. 113 [non Vivipara on-

cophorae Rzehak, 1893]

1954 Viviparus neumayeri [sic] (Brusina) – Andrusov, p. 257, pl. 16,

figs. 12—13 [non Vivipara neumayri Brusina, 1874]

Material: More than 100 specimens from Martin brick-

yard,  coll.  nos.  SNM  1/2011  (PZ-688a—o),  SNM  2/2011
(PZ-689a—ch),  SNM  3/2011  (PZ-690a,b),  SNM  4/2011
(PZ-691a—k),  SNM  5/2011  (PZ-692a—c),  SNM  11/2011
(PZ-698), SNM 16/2011 (PZ-703a—e).

Holotype: SNM 11/2011 (PZ-698); height: 28.8 mm,

width: 22.3 mm (Figs. 2D—E).

Paratype: SNM 11/2011 (PZ-698); height: 32.5 mm,

width: 23.7 mm (Figs. 2F—G).

Stratum typicum: Martin Formation, Upper Miocene.

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Type locality: Brickyard near the town of Martin, Turiec

Basin, Slovak Republic; 49°04’ N, 18°53’ E.

Name: In honour of Radovan Pipík (Slovak Academy of

Sciences, Banská Bystrica), who greatly contributed to our
knowledge of Lake Turiec.

Diagnosis: Broad, conical viviparid with up to 6 whorls;

spire angle distinctly decreases during ontogeny, while
whorl convexity increases towards highly convex whorls in
later ontogeny.

Description: Protoconch depressed, almost planispirally

coiled, covered by weak axial wrinkles; termination indiscern-
ible,  whorl  number  unknown.  Both  degree  of  convexity  and
spire  angle  change  distinctly  during  ontogeny:  early  whorls
broadly  conical  (spire  angle  of  ca.  120°),  weakly  convex,
tightly  coiled,  with  widely  ovoid  aperture  (Fig. 2J).  Adult

Fig. 2.  A  –  Theodoxus  cf.  postcrenulatus  Papp,  1953  (SNM 12/2011,  PZ-699);  B  –  T.  cf.  postcrenulatus  (SNM 9/2011,  PZ-696);
C  –  T.  cf. postcrenulatus (SNM 12/2011, PZ-699); D—E – Viviparus pipiki nov. sp., holotype (SNM 11/2011, PZ-698); F—G –  V. pipiki,
paratype (SNM 11/2011, PZ-698); H – V. pipiki, protoconch (SNM2/2011, PZ-689); I – V. pipiki, protoconch (SNM 2/2011,  PZ-689);
J – V. pipiki, juvenile specimen (SNM 3/2011, PZ-690). All specimens from Martin brickyard. Scale bars correspond to 100 µm (H—I),
1 mm (A—C, J), and 10 mm (D—G).

shells conical to weakly ovoid (ca. 50—60°) with strongly con-
vex, entirely non-stepped whorls. Height-width ratio of adult
shells slightly variable: shells usually rather broad, occasion-
ally  more  slender  specimens  occur  (Fig. 2F—G).  Shell  com-
prises  up  to  6  whorls.  Last  whorl  attains  70—75 %  of  total
height.  Aperture  broadly  drop-shaped,  with  small  posterior
notch.  Distinct  sigmoidal  growth  lines  (adapically  opistho-
cyrt, abapically prosocyrt) cover shell.

Remarks: Owing to its thick shell this species is among

the most common and best preserved ones in the Turiec Ba-
sin. It looks very similar to Viviparus neumayri Brusina,
1874 from the Pliocene “Paludina beds” of Croatia, match-
ing in degree of convexity and shell shape (see also Neumayr
& Paul 1875). However, this species is distinctly smaller and
whorl height increases more slowly than in the present spe-

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cies, resulting in a relatively shorter spire. Another compara-
ble species is Viviparus loczyi Halaváts, 1903, from the Late
Pannonian of the Balaton region, which has a similar shape,
but it has less convex whorls and tends to be more slender.

Andrian (1866) confused the present species with Vivi-

parus sadleri Neumayr, 1869, which differs in its distinctly
stepped spire. Remeš (1923) in turn determined it as V. onco-
phorae  Rzehak,  1893,  which  was  described  from  the  Early
Miocene of Moravia and likewise has a stepped spire. Wenz
(1928)  erroneously  introduced  the  replacement  name  Vivi-
parus oncophoriorum for Rzehak’s taxon to separate it from
Vivipara ( = Viviparus) oncophora Brusina, 1874. This error
is  probably  based  on  a  reading  error  by  Wenz,  who  cited
Rzehak’s species also as “Vivipara oncophora” and thus re-
ferred to it as a primary homonym. This nomenclatural act,
however, is invalid. The Code clearly states that a one-letter
difference between species-group names combined with the
same generic name is sufficient to prevent homonymy (ICZN
1999,  Art. 57.6).  The  exceptions  to  this  rule  mentioned  in
Article 58 do not apply here. Hence, the name Vivipara onco-
phorae  was  validly  introduced  by  Rzehak  (1893).  Viviparus
oncophoriorum Wenz, 1928 is an objective synonym of Vivi-
parus oncophorae Rzehak, 1893.

Distribution: Endemic to the Turiec Basin.

Genus: Popovicia Neubauer & Harzhauser nov. gen.

1964 Metohia nov. gen. – Popović, p. 47
1968 Metohia nov. gen. – Popović, p. 206

Type species: Metohia levantica Popović, 1964.

Pliocene (Early Dacian), Kosovo.

Other included species: Kosovia compressa Pavlović,

1931 (Late Pontian), Metohia turriculoidea Popović, 1964
(Late Dacian).

Name: In honour of Radmila Popović (formerly in the In-

stitute of Geological and Geophysical Research, Belgrade),
who intensively studied the Viviparidae of the Metohia Basin.

Diagnosis: Shell planispiral to trochiform, sinistrally

coiled, attaining up to 4 whorls. Apex shallow to deeply im-
mersed. Umbilicus immersed in planispiral shells, but raised
and flattened in trochiform shells, producing pseudo-dextral
appearance. Whorls regularly convex to angulated, but never
resulting in offset keel; ramp above shoulder straight, whorl
portion below convex. Sutures deep. Sculpture may be present
as weak to distinct spiral keels.

Remarks: Several nomenclatorial issues concerning the

alleged viviparid genera Metohia and Kosovia have strained
the literature so far. First of all, Metohia Popović, 1964 is a
primary homonym of Metohia Absolon, 1927 (Ostracoda),
which is why we introduce Popovicia as a replacement name
here. The second genus, Kosovia, with which the species dis-
cussed below was originally affiliated, is not available from
the original publication (Pavlović 1931), since no type spe-
cies was fixed (ICZN 1999, Art. 13.3). The first to indicate a
type species, namely Kosovia ornata Pavlović, 1931, was
Atanacković (1959). He clearly referred to Pavlović’s work
and description, which is sufficient as indication for the ge-
nus (Art. 13.1.2) and makes him the author of Kosovia. Zilch

(1959—1960), who claimed to fix the type species as well,
was published later (17 July, 1959).

The subfamily Kosoviinae as found in the literature was

never  described  and  is  therefore  unavailable.  Atanacković
(1959) first uses the name (as “Kosovinae”, but see Art. 29 of
the Code), but did not describe it or added a bibliographic ref-
erence as indication. The only other mention in the literature is
by Milošević (1978), who did not describe it either. Since we
do not aim at a systematic revision, a solution to the system-
atic  placements  of  Kosovia  and  Popovicia  is  still  pending.
Following  Atanacković  (1959),  we  place  it  preliminarily  in
the Viviparidae.

Distribution: Only known from the Metohia Basin, Ko-

sovo, and the Turiec Basin.

Popovicia cf. compressa (Pavlović, 1931)

(Fig. 5B)

cf. 1931 Kosovia compressa nov. spec. Pavlović, p. 21, pl. 11, figs. 11—13
cf. 1964 Metohia compressa (Pavl.) – Popović, p. 50, 53, pl. 3, fig. 1,

pl. 4, fig. 1

cf. 1965 Kosovia strophostomopsis strophostomopsis nov. spec. Milo-

šević, p. 116, text-fig. 3, pl. 1, fig. 1

cf. 1965 Kosovia strophostomopsis compressiforma nov. s. spec. Milo-

šević, p. 120, pl. 1, fig. 2

cf. 1965 Kosovia compressa Pavl. – Milošević, pl. 1, fig. 4
cf. 1967 Kosovia strophostomopsis V. Miloš. – Milošević, text-fig. 4a
cf. 1967 Kosovia s. compressiforma V. Miloš. – Milošević, text-fig. 4b
cf. 1968 Metohia compressa (Pavl.) – Popović, p. 208, 216, pl. 3,

fig. 1, pl. 4, fig. 1

cf. 1970 Kosovia strophostomopsis V. Milošević – Milošević, text-fig. 2B
2011 Kosovia – Kováč et al., p. 371
2012 Kosovia stromphostomopsis [sic] – Krstić et al., p. 44

Material: 1 specimen from Trebostovo (Drill core GT-11),

coll. no. SNM 161/2006 (PZ-563); height: 4.5 mm, diame-
ter: 11 mm (Fig. 5B).

Description: Small sinistral, planorbiform shell com-

posed  of  3.25  whorls.  Protoconch  unknown.  Early  whorls
apically flattened, forming a weakly convex plane; traces of
weak striation present. Last whorl overgrows previous ones,
forms a distinct crest at its uppermost part, which separates
an inner concave portion and an outer convex portion. Inner
concave portion shows weak striation. In profile a weak an-
gulation  occurs  slightly  below  whorl  midline.  In  latest  on-
togeny  the  apertural  margin  starts  to  expand  funnel-like:
while the peristome becomes strongly inflated, the actual ap-
ertural opening inside retains its original width.

Remarks: The specimen from Turiec is highly reminiscent

of  Popovicia  compressa  (Pavlović,  1931),  which  was  de-
scribed from the Late Pontian (earliest Pliocene) of the Meto-
hia Basin in Kosovo. It matches perfectly concerning general
shape and apertural inflation. The main difference is the pres-
ence  of  seven  distinct  spiral  keels  in  the  Kosovan  species,
which could not be observed to such an extent on the Turiec
specimen;  this,  however,  could  be  a  result  of  the  moderate
preservation.  Moreover,  like  the  mode  of  apertural  inflation
this  feature  seems  to  be  quite  variable  (Popović  1964).  The
Turiec  specimen  is  also  generally  smaller  and  probably  re-
flects  a  subadult  specimen  (3.25  whorls  at  11 mm  diameter
versus 4 whorls at ca. 15 mm for typical P. compressa).

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The taxonomic history of the genus Metohia Popović,

1964 ( = Popovicia) and the included species is very compli-
cated  and  needs  some  discussion  here  in  order  to  avoid  fu-
ture  confusion.  Popović  (1964)  described  the  new  genus
Metohia to separate planorbiform and low trochiform shells
among  the  Kosoviinae  from  the  high  trochiform,  sinistral
Kosovia species (see also Pavlović 1931; Atanacković 1959).
Additionally, she described the two new species M. levantica
and M. turriculoidea. One year later Milošević (1965) intro-
duced  the  new  species  Kosovia  strophostomopsis  and  two
new  subspecies,  K.  strophostomopsis  compressiforma  and
K.  strophostomopsis  disjuncta,  all  of  which  are  strikingly
similar to the two species of Popović. Despite citing the paper
of  Popović  in  the  reference  section,  Milošević  did  not  dis-
cuss it at all. Only two years later Milošević (1967) referred
to  the  topic  again,  when  discussing  the  systematic  position
of  Kosovia  compressa.  Again  he  did  not  comment  on  the
morphological  similarities  between  Popović’s  and  his  spe-
cies. To complete the confusion Popović (1968) re-published

exactly the same paper as in 1964 in a different journal. Un-
fortunately, all the names were again marked as new, mis-
leadingly pretending first descriptions.

Subsequently, Popović (1969) synonymized K. strophosto-

mopsis with Metohia levantica and regarded both K. strophos-
tomopsis  compressiforma  and  K.  strophostomopsis  disjuncta
as intermediate stages between Metohia levantica and M. tur-
riculoidea.  This  taxonomic  act  was  rejected  by  Milošević
(1970)  for  two  reasons.  First,  he  differentiated  between  his
“pseudo-sinistral”  Kosovia  forms  and  the  “dextral”  Metohia
species.  Given  the  full  correspondence  of  morphologies  this
differentiation  is  untenable.  Second,  Milošević  (1970)  stated
stratigraphical  discrepancies:  while  K.  strophostomopsis  and
its  two  subspecies  are  described  from  Late  Pontian  (earliest
Pliocene)  strata,  M.  levantica  and  M.  turriculoidea  were  re-
corded from Dacian deposits (middle Early Pliocene). He did
not agree with the idea of Popović (1969) that all these species
may  actually  derive  from  Dacian  layers.  More  precisely,  he
stated  that  his  species  come  from  sediments  above  the  main

Fig. 3. A, D – Melanopsis glaubrechti nov. sp., paratype (SNM 10/2011, PZ-697), specimen digitally recombined from two separate im-
ages; B – M. glaubrechti, juvenile shell (SNM 10/2011, PZ-697); C — M. glaubrechti, holotype (SNM 16/2011, PZ-703g); E – Tournouerina
turiecensis nov. sp., paratype 1 (SNM 10/2011, PZ-697); F – T. turiecensis, holotype (SNM 9/2011, PZ-696); G – T. turiecensis,
paratype 2 (SNM 10/2011, PZ-697); H – T. turiecensis, paratype 3 (SNM 10/2011, PZ-697); I – T. turiecensis, protoconch (SNM 10/2011,
PZ-697). All specimens from Martin brickyard. Scale bars correspond to 100 µm (I), 1 mm (D—H), and 5 mm (A—C).

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coal seam. However, the latest stratigraphical investigation by
Elezaj et al. (2010) indeed confirmed the Dacian age of the de-
posits containing the Metohia ( = Popovicia) species, with P.
levantica marking Lower Dacian deposits and P. turriculoidea
occurring in Upper Dacian layers.

Given the different extent of Pontian, Dacian, and Roma-

nian  deposits  in  the  Metohia  Basin  it  is  most  likely  that
Milošević misjudged the age of the deposits. The type locali-
ties  of  Popović’s  Dacian  species  are  located  in  the  south
(Orahovac)  and  southwest  (Trakanić  brook  at  Çabrat  hill
near  Gjakovë)  of  the  Metohia  Basin,  where  Pontian  sedi-
ments  are  very  restricted  (S)  to  absent  (SW)  (Atanacković
1990). In contrast, the type locality of Milošević’s species is
right in the centre of the basin (Deiq), where Pontian to Ro-
manian  deposits  have  been  documented  (Popović  1970;
Milošević 1983; Atanacković 1990). Geographically, all the
type localities involved lie within a 30-km radius.

In summary, we fully agree with Popović (1969) concern-

ing  the  stratigraphical  attribution  to  the  Dacian.  However,
we rather tend to synonymize K. strophostomopsis stropho-
stomopsis  and  K.  strophostomopsis  compressiforma  with
Popovicia  compressa  and  K.  strophostomopsis  disjuncta
with  Popovicia  levantica.  Since  the  evolutionary  change
throughout the Late Pontian and Dacian seems continuous, it
is anyway difficult to draw a line between taxonomic entities
(compare, e.g. Neubauer et al. 2013a,b). More detailed work
on type material would be necessary to clarify this issue.

Distribution: Up to now only recorded for Upper Pon-

tian deposits of the Metohia Basin (Pavlović 1931, 1932;
Popović 1964) and the Turiec Basin.

Superfamily: Cerithioidea Fleming, 1822

Family: Melanopsidae H. Adams & A. Adams, 1854

Subfamily: Melanopsinae H. Adams & A. Adams, 1854

Genus: Melanopsis Férussac, 1807

Type species: Melania costata Olivier, 1804. Recent,

Europe. Subsequent designation by Gray (1847, p. 153).

Melanopsis glaubrechti Neubauer & Harzhauser nov. sp.

(Fig. 3A—D)

1923 Melanoptychia pseudoscalaria Sdbg. – Remeš, p. 113 [non

Melanoptychia pseudoscalaria Sandberger, 1886]

1954 Melanoptychia sp. – Andrusov, p. 257, pl. 16, figs. 14—17

Material: More than 30 specimens and fragments from

Martin brickyard, coll. nos. SNM 5/2011 (PZ-692), SNM
9/2011 (PZ-696), SNM 10/2011 (PZ-697), SNM 13/2011
(PZ-700), SNM 16/2011 (PZ-703a, f—g, ch).

Holotype: SNM 16/2011 (PZ-703g); height: 12.5 mm,

width: 5 mm (Fig. 3C).

Paratype: SNM 10/2011 (PZ-697), juvenile shell; height:

8.8 mm, width: 4.6 mm (Fig. 3A,D).

Stratum typicum: Martin Formation, Upper Miocene.
Type locality: Brickyard near the town of Martin, Turiec

Basin, Slovak Republic; 49°04’ N, 18°53’ E.

Name: In honour of Matthias Glaubrecht (Museum für

Naturkunde Berlin), an expert for the living species of Mela-
nopsidae.

Diagnosis: Small ovoid melanopsid with distinct ribs,

which descend adapically over a small knob down to the su-
ture.  The  ribs  and  the  relief  of  the  ribs  of  the  penultimate
whorl,  which  are  clearly  visible  in  early  ontogeny,  are  not
regularly  aligned,  producing  irregular  sculpture  pattern  be-
low the sutures.

Description: Slender, drop-like shell, attaining its maxi-

mum  diameter  on  the  penultimate  whorl  and  maintains  this
on the last whorl, which attains almost 50 % of total height.
Protoconch  poorly  preserved  but  small  and  smooth.  Includ-
ing the protoconch, the shell comprises up to 9 whorls. The
typical  features  are  the  strong  axial  ribs,  which  start  faintly
on the 3

to 4

whorl. They form a strong crest of uniform

elevation in the lower (abapical) three quarters of the whorls.
Above, the ribs terminate in a small knob; in the uppermost
quarter,  the  ribs  rapidly  slope  down  from  the  knob  towards
suture. In early ontogeny this produces a thin, weakly sculp-
tured  band  in  the  uppermost  part  of  the  whorl.  This  weak
sculpture is not produced by down-sloping ribs, but by ribs
of  the  penultimate  whorl,  whose  abapical  portions  can  be
traced beyond a thin shell layer. Since the number of ribs is
not equal on each whorl, ribs and relief of ribs of penultimate
whorl are not regularly aligned (Fig. 3D). In such cases, where
“pseudo-ribs” run between two ribs, they may reach down to
the  level  of  knobs  but  never  beyond.  Aperture  ovoid,  some-
times with columellar fold; no thickened callus present.

Remarks: This species is reminiscent of specimens from

the Pannonian F of Götzendorf in the Vienna Basin illustrated
by Papp (1953, pl. 12, figs. 16—17), determined as Melanopsis
bouei sturii Fuchs, 1873. A very similar specimen as shown
by  Papp  was  documented  by  Fordinál  (1993,  pl. 12,  fig. 4)
from the Pannonian of the Bratislava area. Both correspond
to M. glaubrechti concerning the general outline and the ar-
rangement of the ribs with terminating adapical knobs. How-
ever,  direct  comparison  with  material  from  Götzendorf
shows subtle but consistent differences: M. sturii has a highly
regular  morphology,  with  even  suture,  smooth  subsutural
band,  and  often  intentions  of  a  second  row  of  weak  knobs
near the whorl base. In contrast, M. glaubrechti has a fairly
irregular  suture  line,  the  typical  irregular  sculpture  pattern
below  the  sutures,  and  just  one  row  of  knobs.  The  overall
similarity still hints at a close phylogenetic relationship be-
tween the two species.

The present species also differs from “M. boettgeri Hala-

váts, 1903”, which has a conical shell and distinctly stronger
knobs forming a spruce-like outline. This species name is a pri-
mary homonym of M. boettgeri Klika, 1891 and was replaced
by Cossmann (1909) with the new name M. balatonensis.
Both Wenz (1929) and Papp (1953), who obviously overlooked
this, synonymized this species with M. sturii. The similar spe-
cies M. pseudaustriaca Sauerzopf, 1952 differs in its distinctly
conical outline and the entirely smooth subsutural band.

Remeš (1923) refers to this species as Melanoptychia pseu-

doscalaria, which was described from the Lower Miocene
“Oncophora beds” (Rzehakia beds) of Moravia. This species
can be distinguished based on its much higher last whorl and
the considerably thickened callus. Moreover, the ribs termi-
nate directly at the upper suture (Rzehak 1893, pl. 2, fig. 7).

Distribution: Endemic to the Turiec Basin.

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Order: Littorinimorpha Golikov & Starobogatov, 1975

Superfamily: Truncatelloidea Gray, 1840

Family: Hydrobiidae Stimpson, 1865

Subfamily: Hydrobiinae Stimpson, 1865

Genus: Tournouerina Schlickum, 1971

Type species: Nematurella lugdunensis Tournouër, 1879.

Pliocene, France. Type by original designation.

Tournouerina turiecensis Neubauer & Harzhauser nov. sp.

(Fig. 3E—I)

?2012 Lithoglyphus nannus [sic] – Krstić et al., p. 44 [non Lithoglyphus

nanus Roshka, 1973]

Material: 35 specimens from Martin brickyard, coll. nos.

SNM 9/2011 (PZ-696), SNM 10/2011 (PZ-697), SNM 13/2011
(PZ-700).

Holotype: SNM 9/2011 (PZ-696); height: 4.0 mm,

width: 2.2 mm (Fig. 3F).

Paratype 1: SNM 10/2011 (PZ-697); height: 4.8 mm,

width: 3.0 mm (Fig. 3E).

Paratype 2: SNM 10/2011 (PZ-697); height: 4.5 mm,

width: 2.1 mm (Fig. 3G).

Paratype 3: SNM 10/2011 (PZ-697); height: 4.2 mm,

width: 2.1 mm (Fig. 3H).

Stratum typicum: Martin Formation, Upper Miocene.
Type locality: Brickyard near the town of Martin, Turiec

Basin, Slovak Republic; 49°04’ N, 18°53’ E.

Name: After the Turiec Basin.
Diagnosis: Small, ovoid hydrobiid with up to 6 convex

whorls, separated by moderately deep sutures, a large last
whorl, and an inclined, broadly drop-shaped aperture with a
thickened posterior tip.

Description: Shell small, composed of 6 whorls. Shell

shape is variable, reaching from rather broadly ovoid to slen-
der,  almost  conical.  Protoconch  low  trochiform,  consisting
of exactly 1 highly convex whorl with a diameter of 300 µm;
termination  indicated  by  indistinct  growth  rim;  surface
throughout  covered  with  small,  densely  arranged,  weak  but
consistent wrinkles, which partly merge into irregular, indis-
tinct axial riblets. Teleoconch whorls distinctly and regularly
convex, separated by moderately deep sutures. Last whorl at-
tains  ca.  70 %  of  total  height.  Aperture  broadly,  regularly
drop-shaped, weakly inclined (20—30° to axis), with uninter-
rupted,  slightly  reflected  peristome.  Inner  lip  weakly  pro-
truding  below  posterior  tip.  Aperture  growing  increasingly
in  anterior  direction  in  latest  stage,  producing  a  thickened
posterior tip, with former shell layers distinctly visible. Um-
bilicus narrow; in slender specimens almost fully covered by
aperture. Faint, weakly prosocyrt growth lines cover shell.

Remarks: After detailed literature review and study of

fossil material, we are not aware of a single hydrobiid spe-
cies in the Mio-Pliocene of Europe that might be conspecific
with the present one. The characteristic ovoid aperture with
the thickened posterior tip and the narrow umbilicus argues
for an affiliation with the genus Tournouerina (e.g. Schlickum
1971, 1978; Schlickum & Puisségur 1977, 1978; Meijer

1989). Even the curvature of the aperture in side-view
matches perfectly with the illustrations of Schlickum (1978).
Especially the type species T. lugdunensis (Tournouër, 1879)
from the Pliocene of Miribel (Ain, France) is very similar in
terms of outline shape, but it has less convex whorls and a
less bulbous last whorl.

Some species of the genus Prososthenia Neumayr, 1869

also exhibit certain similarities, like P. sepulcralis Partsch
in Cžjžek, 1848 from Lake Pannon. This species differs in
its  clearly  less  convex  whorls,  the  less  inclined  aperture,
and the weak angle at the base, producing a conical outline
(Papp  1953;  Harzhauser  &  Binder  2004).  The  Pannonian
Hydrobia  testulata  Papp,  1953,  which  was  synonymized
with Hydrobia pseudocornea Brusina, 1902 by Harzhauser
&  Binder  (2004),  has  similar  shape  and  size,  but  clearly
differs in the aperture, which lacks the thickened posterior
tip, and the more convex whorls. Another very similar Pan-
nonian  species  is  Caspia  boeckhi  (Lörenthey,  1902)  from
Tinnye,  Hungary,  which  can  be  distinguished  by  the  more
conical  outline,  formed  by  a  distinct  angle  on  the  last
whorl, and the whorl convexity, which lies below the median.
The  quite  variable  Middle  Miocene  Prososthenia  neutra
Brusina, 1897 from Miočić, Croatia, has a less bulbous last
whorl  and  a  stronger  inclined  aperture  resulting  in  an  al-
most covered umbilicus. The likewise variable Dacian spe-
cies  Hydrobia  arminiensis  Jekelius,  1932  from  the  Bra ov
Basin  has  a  wider  aperture  that  almost  or  fully  covers  the
umbilicus, in most cases a bigger last whorl, and more con-
vex whorls.

Most likely the identification of Krstić et al. (2012) as

Lithoglyphus nanus Roshka, 1973 [misspelt as “nannus”] ac-
tually  reflects  the  present  species.  Lithoglyphus  nanus  is
a  Late  Miocene  taxon  from  the  Eastern  Paratethys  and  has
a  much  bulkier  shell  with  bigger  last  whorl  and  strongly
inclined aperture. Whether or not the records of Hydrobia sp.
from  Hrby  and  “Stalioia  [sic]  gracilis”  from  Dolina  by
Remeš  (1923)  correspond  to  the  present  species  could  not
be  verified.  The  material  from  Dolina  available  to  us  com-
prises  lithified  freshwater  limestones  with  unidentifiable
fragments  of  shelly  impressions  and  molds.  As  correctly
pointed  out  by  Andrusov  (1954),  the  name  Stalioa  gracilis
has to be replaced with Stalioa rzehaki. Wenz (1925) consid-
ered  Staliopsis  gracilis  Rzehak,  1893  and  Euchilus  gracile
Sandberger,  1875  congeneric,  making  the  former  a  second-
ary  homonym  of  the  latter.  For  Staliopsis  gracilis  he  intro-
duced  the  replacement  name  Stalioa  rzehaki  Wenz,  1925.
Currently  it  is  again  affiliated  with  Staliopsis  (Čtyroký
1972).

Distribution: Endemic to the Turiec Basin.

Clade: Panpulmonata Jörger et al., 2010

Order: Hygrophila Férussac, 1822

Superfamily: Lymnaeoidea Rafinesque, 1815

Family: Lymnaeidae Rafinesque, 1815

Subfamily: Lymnaeinae Rafinesque, 1815

Genus: Radix Montfort, 1810

Type species: Helix Auricularia Linnaeus, 1758. Recent,

Europe. Type by original designation (Welter-Schultes 2012).

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Fig. 4. Radix kovaci nov. sp. A – Holotype (SNM 114/2006, PZ-517), Mošovce (GT-14); B – Juvenile specimen (SNM 148/2006, PZ-550),
Horná Štubňa (GHŠ-1); C – Paratype 1 (SNM 155/2006, PZ-557), Horná Štubňa (GHŠ-1); D – Paratype 3 (SNM 121/2006, PZ-524a),
Mošovce (GT-14); E – Paratype 2, (SNM 153/2006, PZ-555), Horná Štubňa (GHŠ-1); F — Sediment infills of apertures of two specimens
(SNM 118/2006, PZ-521), Mošovce (GT-14). Scale bars correspond to 10 mm.

Radix kovaci Neubauer & Harzhauser nov. sp.

(Fig. 4A—F)

Material: Several tens of specimens in different ontoge-

netic  stages,  from  following  localities:  Mošovce  (Drill  core
GT-14), coll. nos. SNM 114/2006 (PZ-517), SNM 117-119/
2006  (PZ-520-522),  SNM 121/2006  (PZ-524b),  SNM 120/
2006  (PZ-523); Horná Štubňa (Drill core GHŠ-1), coll. nos.
SNM 144-145/2006

(PZ-546-547),

SNM 146-154/2006

(PZ-548-561), SNM 136-143/2006 (PZ-538-545); Koš any
nad Turcom (Drill core GT-13), coll. nos. SNM 107-113/2006
(PZ-510-516).

Holotype: SNM 114/2006 (PZ-517); height: 14 mm, dia-

meter: 16 mm (Fig. 4A).

Paratype 1: SNM 155/2006 (PZ-557), from drill core

GHŠ-1, Horná Štubňa; height: 10.8 mm, diameter: 8.5 mm
(Fig. 4C).

Paratype 2: SNM 153/2006 (PZ-555), from drill core

GHŠ-1, Horná Štubňa; height: 11.4 mm, diameter: 11.2 mm
(Fig. 4E).

Paratype 3: SNM 121/2006 (PZ-524a), from type locality;

sediment infill of aperture; height: 15.5 mm, diameter:
16.6 mm (Fig. 4D).

Stratum typicum: Martin Formation, Upper Miocene

(drilling depth 229.5—229.6 m).

Type locality: Mošovce (Drill core GT-14), Turiec Basin,

Slovak Republic; 48°54’ N, 18°53’ E.

Name: In honour of Michal Kováč (Univerzita Komen-

ského, Bratislava), who intensively worked on the sedimen-
tary evolution and stratigraphy of the Pannonian Basin
System and adjacent areas.

Diagnosis: Moderately sized Radix with rapidly expand-

ing whorls towards an extremely large aperture, finally over-
growing the spire.

Description: Most of the material does not allow a proper

description. The few complete or nearly so specimens show
a  shell  with  ca.  4  whorls,  a  very  small  and  short  spire,  and
an extremely rapidly expanding last whorl. The direction of
expansion  forms  an  angle  of  usually  around  55°  to  the  col-
umella.  Aperture  prominent,  elliptical  to  almost  round,  ter-
minates  in  sharp  peristome.  Where  it  touches  base  of
penultimate whorl it forms small notch. Degree and direction
of apertural expansion as well as apertural shape highly vari-
able.  Apparently  expansion  continues  throughout  ontogeny,
so that aperture even overgrows entire spire in latest ontogeny;
in  abapertural  view  this  produces  bulky,  cap-like  appear-
ance. Some specimens show elongate shape due to stronger
lateral expansion of the aperture.

Remarks: After careful review of the rich literature we

are still not aware of any lymnaeid species of the Mio-

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Pliocene of Europe that might be conspecific with this shell.
The uniquely formed aperture distinguishes it from all other
known  species.  Most  of  the  available  specimens  are  pre-
served  as  mere  impressions  or  sediment  fillings  of  the  last
aperture, which form cap-like molds. The alleged records of
Ancylus  mentioned  in  the  literature  probably  refer  to  these
fillings as indicated by the museum labels. The final growth
stage with the inflated aperture is also reminiscent of species
of the genus Hiscerus Gorjanović-Kramberger, 1923, a lym-
naeid  with  cap-shaped,  oval  shells  (see  Gorjanović-Kram-
berger  1923;  Moos  1944).  These,  however,  are  much  more
bulbous and usually have a wrinkled surface.

The only rather similar taxon is Radix kobelti (Brusina,

1884) from the Pannonian of the Zagreb area, which also ex-
hibits a very wide aperture, but differs in the lower spire and
the almost horizontally oriented and stronger expressed ex-
pansion. Moreover, shells of R. kobelti are distinctly larger
(see also Brusina 1897; Gorjanović-Kramberger 1923).

Distribution: Endemic to the Turiec Basin.

Radix sp.
(Fig. 5A)

Material: 1 specimen from Martin (Drill core ZGT-3),

coll. no. SNM 183/2006 (P-520a); height: 27 mm, width
15.5 mm.

Remarks: A second Radix species is documented by a

single, moderately preserved specimen. It can be easily dis-
tinguished from the co-occurring R. kovaci by its huge but

slender shell. A determination at species level is not possible
as the aperture is covered with sediment.

Superfamily: Planorboidea Rafinesque, 1815

Family: Planorbidae Rafinesque, 1815

Genus: Planorbarius Duméril, 1806

Type species: Helix cornea Linnaeus, 1758. Recent,

Europe. Subsequent monotypy by Froriep (1806, p. 165).

Planorbarius nov. sp.

(Fig. 5C)

?1866 Planorbis pseudoammonius – Andrian, p. 196 [non Helicites

pseudoammonius Schlotheim, 1820]

Material: Several fragments and impressions from the fol-

lowing  localities:  Horná  Štubňa  (Drill  core  GHŠ-1),  coll.  no.
SNM  160/2006  (PZ-562a,b);  Martin  (Drill  core  ZGT-3),  coll.
no.  SNM  190/2006  (PZ-579);  Martin  brickyard  (compressed),
coll. no. SNM 13/2011 (PZ-700); Turčiansky Peter (Drill core
GT-6, 37.2—37.3 m), coll. no. SNM 180/2006 (PZ-517); height:
6.5 mm, diameter: 18 mm (most complete specimen, Fig. 5C).

Description: Most complete specimen attains 4.5 whorls

at a maximum diameter of 18 mm. Whorls appear regularly
convex and increase regularly in diameter. Up to penultimate
whorl apical region flat, not depressed; only last whorl dis-
tinctly elevated. Umbilical region unknown. Growth lines
but no horizontal striation visible.

Fig. 6. Bivalves. A—C – Mytilopsis? sp. (SNM 9/2011, PZ-696), juvenile specimen, Martin brickyard; D – Pisidium sp. (SNM 13/2011,
PZ-700b), Martin brickyard; E – Unio cf. atavus Hörnes, 1865 (SNM 14/2011, PZ-701), Martin brickyard; F – U. cf. atavus (SNM 7/2011,
PZ-694c), Martin brickyard; G – U. cf. atavus (SNM 7/2011, PZ-694a), Martin brickyard; H – Mytilopsis? sp. (SNM 166/2006, PZ-568),
Martin brickyard; I – Lymnocardiinae gen. et sp. indet. 1 (SNM 380/1968, G-129), Martin-Záturčie (BJ-2); J – Lymnocardiinae gen. et
sp. indet. 1 (SNM 380/1968, G-129), Martin-Záturčie (BJ-2); K – Lymnocardiinae gen. et sp. indet. 2 (SNM 380/1968, G-129), Martin-
Záturčie (BJ-2). Scale bars correspond to 200 µm (C), 1 mm (A—B, D), 5 mm (I—K), and 10 mm (E—H).

Fig. 5. A – Radix sp. (SNM 183/2006, PZ-520), Martin (ZGT-3); B – Popovicia cf. compressa (Pavlović, 1931) (SNM 161/2006, PZ-563),
Trebostovo (GT-11); C – Planorbarius nov. sp. (SNM 160/2006, PZ-562a), Horná Štubňa (GHŠ-1). Scale bars correspond to 10 mm.

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Distribution: So far only known from the Turiec Basin.

Discussion

Faunal composition

The freshwater gastropod fauna consists of at least 10 spe-

cies (Table 1). In many cases, the rather poor preservation of
the  fossils  prevents  precise  determination.  Therefore,  some
taxa could not be identified at the species level. Many fossils
from  drill  cores  are  strongly  compressed,  deformed  or  pre-
served as shelly impressions only. The best preserved speci-
mens  derive  from  the  Martin  brickyard,  including  most  of
the thick-shelled Viviparus and the small  Theodoxus, Mela-
nopsis,  and  Tournouerina.  Most  of  the  species  that  have
been determined at the species level are unknown from other
European  freshwater  systems.  This  again  confirms  the  high
degree of endemicity in Lake Turiec, as was already shown
for the ostracod fauna (Pipík et al. 2012).

The composition above genus level is quite diverse with

7  families  present  (Viviparidae,  Neritidae,  Melanopsidae,
Bithyniidae,  Hydrobiidae,  Lymnaeidae,  and  Planorbidae).
This rather high number, however, must not be overestimated,
since this diversity is never found in a single locality but is
the sum of all species recorded from Upper Miocene depos-
its  of  the  entire  lake.  The  varied  composition  is  certainly
reflected in the paleoecological and/or stratigraphical differ-
ences  throughout  the  basin.  The  ecological  model  estab-
lished  by  Pipík  et  al.  (2012)  based  on  the  ostracod  fauna
indicates  the  presence  of  quite  diverse  habitats  in  Lake  Tu-
riec. Most of the molluscs derive from marsh/wetland habi-
tats  or  littoral  to  sublittoral  areas  in  the  north  and  are
preserved  as  accumulations  in  sublittoral  sediments  (e.g.  in
Martin brickyard). More precise data, however, is not avail-
able, excluding reconstruction of the original habitats of the
single species.

In addition to the freshwater gastropods, several fragments

of terrestrial gastropods were found in the studied material, in-
cluding a few opercula of Pomatias and remnants of helicids.
None of these, however, allowed proper identification, in most
cases not even at the family level. Bivalves are present as well
and display a great systematic diversity similar to that of the
gastropods. They are represented by at least one slender, elon-
gate, triangular species of Mytilopsis? sp. (Dreissenidae),
two single valves of Pisidium sp. (Sphaeriidae), several im-
prints of two species of Lymnocardiinae, and several shells
and fragments of Unio cf. atavus Hörnes, 1865 (Unionidae)
(Fig. 6). The overall rather poor preservation of the material
makes more precise identifications difficult.

Stratigraphy

The earliest studies correlated the deposits with the Pan-

nonian  (“Congerienschichten”)  based  on  elements  of  the
mollusc  fauna  Štúr  1860;  Andrian  1866;  Vetters  1910;
Fig. 7). Vigh (1915) in turn suggested a “Pliocene to Levan-
tinian”  age.  Remeš  (1923),  in  contrast,  indicated  an  Early
Miocene age, based on the findings of the genus Oncophora
( = Rzehakia) which is restricted to the late Ottnangian (mid-
dle Burdigalian) (e.g. Čtyroký 1972). This implied a tempo-
ral  shift  of  more  than  7 Ma  and  naturally  led  to  continuing
discussions  among  later  authors  (Andrusov  1938;  Čechovič
1948,  1954).  Since  not  a  single  follow-up  study  could  con-
firm the record of Rzehakia, it was and is considered a misi-
dentification. Based on his determinations, Andrusov (1954)
supported  the  former  opinion  and  suggested  an  Early  Pan-
nonian age. More precise age constraints are provided from
K-Ar radiometric dating by Konečný et al. (1983). They give
an age of ca. 12.4—10.7 Ma (Late Sarmatian/Early Pannonian
or Late Serravallian/Early Tortonian) for the rhyolitic forma-
tion  of  the  Central  Slovakia  Volcanic  Field  underlying  the
Martin Formation, bounding the beginning of the Martin For-
mation and the main evolutionary episode of the endemic fauna

Species Martin

brickyard

Horná

Štubňa

(GHŠ-1)

Turčiansky

Peter

(GT-6)

Trebostovo

(GT-11)

Mošovce

(GT-14)

Košťany

nad

Turcom
(GT-13)

Martin

(ZGT-3)

Martin-

Záturčie

(BJ-2)

Theodoxus cf. postcrenulatus Papp, 1953

Viviparus pipiki Neubauer & Harzhauser nov. sp.

Popovicia cf. compressa (Pavlović, 1931)

Melanopsis glaubrechti Neubauer & Harzhauser
nov. sp.

Tournouerina turiecensis Neubauer & Harzhauser
nov. sp.

Bithynia sp.

Radix kovaci Neubauer & Harzhauser nov. sp.

x x

Radix sp.

Gyraulus sp.

Planorbarius nov. sp.

Lymnocardiinae gen. et. sp. indet. 1

Lymnocardiinae gen et. sp. indet. 2

Mytilopsis? sp.

x x

Pisidium sp.

Unio cf. atavus Hörnes, 1865

Table 1: Mollusc species present in the Turiec Basin. Counts are avoided here as the specimens derive from several non-quantitative col-
lections.

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of Lake Turiec. An age of 6.0 Ma (Late Pannonian, earli-
est Pontian or Late Messinian) is indicated by Konečný
et  al.  (1983)  and  Kováč  et  al.  (2011)  for  the  Abramová
Member, which is correlated to the top of the formation.
A  similar  age,  documenting  the  upper  boundary  of  the
formation  and  an  approximate  end  of  Lake  Turiec,  de-
rives from radiometric data from zircon and apatite fission
track thermochronology of the rhyolite tuffs and breccias
from  the  south  margin  of  the  basin  (Králiková  et  al.
2014). Detailed lithostratigraphic divisions were provided
by Rakús et al. (2005) and Kováč et al. (2011).

The given age constraints are in good agreement with

the  majority  of  the  fauna  of  the  Martin  Formation,  in-
cluding  the  species  Theodoxus  cf.  postrenulatus,  Vivi-
parus  pipiki  nov.  sp.,  Melanopsis  glaubrechti  nov.  sp.,
Tournouerina turiecensis nov. sp., Planorbarius nov. sp.,
Mytilopsis?  sp.,  and  Unio  cf.  atavus.  In  neighbouring

Fig. 7. Compiled stratigraphic chart based on the Geological Time Scale 2012 (Gradstein et al. 2012) with correlation of regional units and
biozones. Regional correlations follow Sacchi & Horváth (2002), Krstić (2003), Popov et al. (2006), Harzhauser & Mandic (2008), Krijgsman
et al. (2010), Vasiliev et al. (2011), Magyar & Geary (2012), Andreescu et al. (2013), Stoica et al. (2013), and ter Borgh et al. (2013). The
boundaries of the intervals used by Wenz (1923—1930) are adapted to those of the current stages. Ranges of former and present stratigraphic
classifications of the Martin Formation are given to the right.

Fig. 8.  Map  of  sediment  distributions  of  Late  Miocene  lakes
referred to in the discussion. Lake Pannon: extent for the early
late  Pannonian  (ca.  8 Ma),  after  Magyar  et  al.  (1999).  Lake
Metohia:  extent  for  the  Pontian,  after  Atanacković  (1990),
Elezaj et al. (2010), and Neubauer et al. (2015). Lake Turiec:
after Pipík et al. (2012). Note that the lake outlines do not nec-
essarily correspond to the basin boundaries.

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Lake Pannon Theodoxus postcrenulatus is recorded for Pan-
nonian biozones E—F, corresponding to the upper Lymnocar-
dium  conjungens  Zone  to  Mytilopsis  neumayri/zahalkai
Zone  or  to  the  Middle  Pannonian  (Papp  1953;  Fordinál
1997;  for  a  biostratigraphic  correlation  chart  see  Magyar  et
al. 1999 and Harzhauser et al. 2004). Likewise, the first vivi-
parids  occur  in  Lake  Pannon  during  zone  F  (e.g.  Strausz
1942; Papp 1953). Melanopsis sturii, probably a close rela-
tive or even ancestor of the here described M. glaubrechti, is
documented for the Pannonian F as well (Papp 1953). Unio
atavus is an Early-Middle Pannonian species (Harzhauser et
al.  2007).  This  pattern  suggests  a  similar  temporal  range  of
the deposits from Martin brickyard. The genus Tournouerina
is unknown so far from Miocene localities but is widespread
during  the  Pliocene  (Schlickum  1978).  In  contrast,  Popovi-
cia  cf.  compressa  rather  indicates  a  younger  age.  This  spe-
cies  was  described  from  Upper  Pontian  deposits  of  the
Metohia Basin (Pavlović 1931; Popović 1969), which can be
roughly correlated with the latest Pannonian, and so earliest
Pliocene (Elezaj et al. 2010). Since the fauna of the Metohia
Basin  shows  a  highly  endemic  development  (e.g.  Pavlović
1931,  1932;  Milošević  1962;  Popović  1964,  1970;  Atanac-
ković & Stevanović 1990), a reliable biostratigraphical corre-
lation  is  not  feasible.  Absolute  datings  are  also  not  available
for the Metohia Basin, lowering the biostratigraphic value of
its fauna. At present, this discrepancy remains unsolved.

Based on these considerations, the mollusc-bearing layers

of the Martin Formation can be roughly correlated with the
latest Middle or early Late Pannonian, corresponding to the
Middle and Late Tortonian of the International Timescale.

Paleobiogeography

One of the most important results of this taxonomic work is

the revision of species and genus names erroneously stated for
the  Turiec  Basin.  Names  like  Ancylus,  Hydrobia,  Kosovia,
Melanoptychia,  Lithoglyphus,  Lymnaea,  and  Pyrgula  greatly
biased the picture of factual biodiversity, paleobiogeography,
and  paleoecology.  Most  of  the  available  determinations  and
names  in  the  literature  represent  misidentifications,  partly
even on the genus level. “Hydrobia” and “Pyrgula” as well as
“Lithoglyphus nannus” mentioned by Krstić et al. (2012) most
likely reflect the here described Tournouerina turiecensis nov.
sp.;  “Lymnaea”  corresponds  to  one  of  the  described  Radix
species; “Ancylus” turned out to be sedimentary infills of the
inflated apertures of Radix kovaci nov. sp.; “Planorbis pseudo-
ammonius”  most  probably  reflects  Planorbarius  sp.;  the  al-
leged occurrence of the Early Pannonian endemic Papyrotheca
was based on a misidentified fragment of a dreissenid bivalve.

Certainly, the most interesting fact is the record of Popovi-

cia  cf.  compressa.  This  species  belongs  to  an  evolutionary
lineage seemingly endemic to the Metohia Basin in Kosovo
(Fig. 8). Probably originating from the Middle-Late Miocene
Kosovia  lineage,  the  planorbiform  P.  compressa  evolved
during the Late Pontian (earliest Pliocene), followed by the
planorbiform to low trochiform P. levantica in the Early Da-
cian,  and  terminating  with  the  high  trochiform  P.  turricu-
loidea in the Late Dacian (middle Early Pliocene) (Popović
1969; Elezaj et al. 2010). It remains unclear how a member

of this lineage could enter the Turiec Basin. Direct dispersal
is  impossible,  since  neither  the  Lake  Turiec  nor  the  Lake
Metohia had any hydrological connections to the interjacent
Lake  Pannon,  where  this  genus  is  absent.  One  possible  ex-
planation is transport via birds, which seems to account for
several  present  and  past  distribution  patterns  of  freshwater
gastropods  (Wesselingh  et  al.  1999;  Figuerola  &  Green
2002; van Leeuwen et al. 2012).

Especially in this light it is notable that apart from the

somewhat  doubtful  record  of  Theodoxus  postcrenulatus
none  of  the  species  was  found  in  the  nearby,  coeval  Lake
Pannon. In contrast to the situation for Lake Turiec, the fauna
of  Lake  Pannon,  whose  shore  line  was  some  160 km  to  the
southeast  during  the  Late  Pannonian  (Magyar  et  al.  1999;
Fig. 8), is particularly well known. Presently a statement on
such discrepancies and paleobiogeographical relationships in
general would be premature, since our knowledge about the
Turiec mollusc fauna is still incomplete. The collection and
evaluation  of  more  and  better  preserved  fossils  would  cer-
tainly  allow  a  more  detailed  analysis  of  the  paleobiogeo-
graphical  picture  as  well  as  a  reconstruction  of  the  lake’s
paleoecological settings. Nevertheless, the present work con-
tributes to the still fragmentary puzzle of the actual biodiver-
sity of the Late Miocene Lake Turiec – and once more the
high degree of its endemicity has been proven.

Acknowledgments: Our thanks go to the following persons
for  help:  Andrej  Bendík  (Múzeum  Andreja  Kme a,  Martin)
for providing access to the here studied material stored in the
paleontological  collection  of  the  museum  in  Martin;  Anita
Eschner (NHM Vienna) for access to the malacological collec-
tion  of  the  NHM  Vienna;  Alice  Schumacher  (NHM  Vienna)
for  making  the  majority  of  the  macro-photographs;  Oleg
Mandic (NHM Vienna) for help with translation of the Ser-
bian literature; Andreas Kroh (NHM Vienna) for discussion
of  nomenclatural  issues;  Sonja  Herzog-Gutsch  and  Andrea
Kourgli (NHM Vienna) for help with literature search. Finally,
we are grateful for the inputs by two anonymous reviewers,
who improved an earlier draft of the manuscript. This work
contributes to the FreshGEN Project financed by the Austrian
Science  Fund  (FWF  Project P25365-B25:  “Freshwater  sys-
tems  in  the  Neogene  and  Quaternary  of  Europe:  Gastropod
biodiversity, provinciality, and faunal gradients”).

References

Adams H. & Adams A. 1853—1858: The genera of Recent Mollusca

arranged according to their organizations. Van Voorst, London,
2 vol. of text (661 pp.), 1 vol. of plates.

Andreescu I., Codrea V., Lubenescu V., Munteanu T., Petculescu

A., tiucă E. & Terzea E. 2013: New developments in the Up-
per Pliocene-Pleistocene stratigraphic units of the Dacian Ba-
sin (Eastern Paratethys), Romania. Quatern. Int. 284, 15—29.

Andrian F.v. 1865: Die Zusammensetzung des Thuroczer Tertiär-

beckens. Verh. k.k. Geol. R.-A. 1865, 2, 91—92.

Andrian F.v. 1866: Bericht über die im Sommer 1864 ausgeführten

Detail-Aufnahmen des Thuroczer und der angrenzenden Theile
des Trentschiner Comitates. Jb. k.k. Geol. R.-A. 16, 2, 182—200.

Andrusov D. 1938: Karpaten-Miozän und Wiener Becken. Petro-

leum 34, 27, 1—9.

153

ENDEMIC UPPER MIOCENE LACUSTRINE GASTROPODS OF THE TURIEC BASIN

GEOL

EOL

EOLOGICA CARPA

OGICA CARPA

OGICA CARPATHICA

THICA

THICA, 2015, 66, 2, 139—156

Andrusov D. 1954: On the age of the Turiec depression infill and the

Pliocene evolution of Central Slovakia. Geol. Sbor. Slov. Akad.
Vied 5, 1—4, 255—269 (in Slovak).

Atanacković M. 1959: Pliocène du Bassin de Kosovo (Serbie

méridionale). Geol. Glasnik 3, 257—377 (in Serbian).

Atanacković M. 1990: Kosovo-Metohia Becken (Süd-Serbien) und

sein limnisches Pont. In: Stevanović P., Nevesskaya L.A., Mari-
nescu F., Sokač A. & Jámbor Á. (Eds.): Chronostratigraphie und
Neostratotypen. Neogen der Westlichen (“Zentrale”) Paratethys,
Bd. VIII, Pl1. Pontien. Verlag der Jugoslawischen Akademie der
Wissenschaften und Künste und der Serbischen Akademie der
Wissenschaften und Künste, Zagreb, Beograd, 1—952.

Atanacković M. & Stevanović P. 1990: Pontische limnische Mol-

lusken aus Kosovo und Metohia Becken (Südserbien). In: Ste-
vanović  P.,  Nevesskaya  L.A.,  Marinescu  F.,  Sokač  A.  &
Jámbor  Á.  (Eds.):  Chronostratigraphie  und  Neostratotypen.
Neogen der Westlichen (“Zentrale”) Paratethys, Bd. VIII, Pl1.
Pontien.  Verlag  der  Jugoslawischen  Akademie  der  Wissen-
schaften und Künste und der Serbischen Akademie der Wissen-
schaften und Künste, Zagreb, Beograd, 1—952.

Bartha F. 1979: Upper Pannonian mollusk fauna from the south-west-

ern Balaton area. Földt. Közl. 109, 1—12 (in Hungarian).

Binder H. 2003: Terrestrial and freshwater Gastropods of the Kar-

patian (Lower Miocene). In: Brzobohatý R., Cicha I., Kováč
M. & Rögl F. (Eds.): The Karpatian – a Lower Miocene Stage
of the Central Paratethys. Masaryk University, Brno, 213—215.

Bouchet P. & Rocroi J.-P. (Eds.) 2005: Classification and nomen-

clator of Gastropod Families. Malacologia 47, 1—2, 1—397.

Brongniart A. 1810: Sur des Terrains qui paroissent avoir été formés

sous l’eau douce. Ann. Mus. Nat. Hist. Natur. 15, 357—405.

Brusina S. 1874: Fossile Binnen-Mollusken aus Dalmatien, Kroatien

und Slavonien nebst einem Anhange. Actienbuchdruckerei,
Agram, 1—138.

Brusina S. 1884: Die Fauna der Congerienschichten von Agram in

Kroatien. Beitr. Palaeont. Geol. Österr.-Ung. Orients 3, 125—187.

Brusina S. 1892: Fauna fossile terziaria di Markuševec in Croazia.

Con un elenco delle Dreissensidae della Dalmazia, Croazia e
Slavonia. Glasn. Hrv. Naravosl. Društva 7, 113—210.

Brusina S. 1897: Material for Neogene malacological faunas of

Dalmatia, Croatia and Slavonia with some species from Bosnia
and Herzegovina and Serbia. Djela Jugosl. Akad. Znanosti
Umjetnosti 18, 1—43 (in Croatian).

Brusina S. 1902: Iconographia Molluscorum Fossilium in tellure ter-

tiaria Hungariae, Croatiae, Slavoniae, Dalmatiae, Bosniae,
Herzegovinae, Serbiae and Bulgariae inventorum. Officina Soc.
Typographicae, Agram, 30 plates.

Buday T. 1962: Neogene of the Turiec Basin. Sborn. Ústř. Úst.

Geol., Svazek XXVII – Geológia, Vyd. ČAV, Praha, 475—502
(in Czech).

Cossmann M. 1909: Essais de Paléoconchologie Comparée. Huitieme

Livraison, Chez l’auteur, Paris, 1—248.

Cox L.R. 1960: Thoughts on the classification of the Gastropoda.

Proc. Malacol. Soc. London 33, 239—261.

Criscione F. & Ponder W.F. 2013: A phylogenetic analysis of ris-

sooidean and cingulopsoidean families (Gastropoda: Caeno-
gastropoda). Mol. Phylogenet. Evol. 66, 1075—1082.

Cuvier G. 1795: Second Mémoire sur l’organisation et les rapports

des animaux à sang blanc, dans lequel on traite de la structure
des Mollusques et de leur division en ordre, lu à la société
d’Histoire Naturelle de Paris, le 11 prairial an troisième. Mag.
Encycl. 2, 433—449.

Cžjžek J. 1848—1849: Erläuterungen zur Geognostischen Karte

der Umgebungen von Wien. Wilhelm Braumüller, Wien,
xxx + 104 + 75.

Čechovič V. 1948: A finding of the Oncophora Beds in the Pannon-

ian Basin. Práce Štát. Geol. Úst. 17, 73—86 (in Slovak).

Čechovič V. 1954: Conditions of formation and stratigraphic position

of the Oncophora Beds. Geol. Sbor. Slov. Akad. Vied 5, 1—4,
269—311 (in Slovak).

Čtyroký P. 1972: Die Molluskenfauna der Rzehakia – (Oncophora)

– Schichten Mährens. Ann. Naturhist. Mus. Wien 76, 41—141.

Čtyroký P., Seneš J., Strauch F., Papp A., Kantorová V., Ondrejíč-

ková A., Vass D. & Bohn-Havas M. 1973: Die Entwicklung
der Rzehakia (Oncophora) Formation – M

2c—d

– in der zen-

tralen Paratethys. In: Papp A., Rögl F. & Seneš J. (Eds.): Chro-
nostratigraphie und Neostratotypen. Miozän der zentralen
Paratethys, Bd. III, M2. Ottnangien. Vydavatel’stvo Slovenskej
akadémie vied, Bratislava, 89—113.

Duméril A.M.C. 1806: Zoologie analytique, ou méthode naturelle

de classification des animaux, rendue plus facile à l’aide de
tableaux synoptiques. Allais, Paris, 1—344.

Dunker W. 1848: Über die in der Molasse bei Günzberg unfern Ulm

vorkommenden Conchylien und Pflanzenreste. Palaeontogr. 1,
155—168.

Elezaj Z., Tmava A., Pashko P. & Vaso P. 2010: The Neogene of

the Dukagjini Basin. Muz. Olt. Craiova. Olt. Stud. comun.

tiin . Nat. 26, 1, 291—295.

Figuerola J. & Green A.J. 2002: Dispersal of aquatic organisms by

waterbirds: a review of past research and priorities for future
studies. Freshwater Biol. 47, 483—494.

Fleming J. 1822: The philosophy of zoology, a general view of the

structure, functions and classification of animals. 2. Constable
& Co., Edinburgh, 1—618.

Fontannes F. 1876: Le vallon de la Fuly et les sables à Buccins des

environs d’Heyrieu (Études stratigraphiques et paléontologiques
pour servir a l’histoire de la période tertiaire dans le bassin du
Rhône. I.). Ann. Soc. Agricult. Hist. Nat. Arts Util. Lyon., Quatr.
Sér. 8, 13—72.

Fordinál K. 1993: Genus Melanopsis (Gastropoda) in Upper Mio-

cene sediments in the Bratislava area. Západ. Karpaty Sér.
Paleont. 17, 57—67.

Fordinál K. 1997: Molluscs (gastropoda, bivalvia) from the Pan-

nonian deposits of the western part of the Danube Basin (Pezi-
nok-clay pit). Slovak. Geol. Mag. 3, 4, 263—283.

Froriep L.F. 1806: C. Dumeril’s Analytische Zoologie. Aus dem

Französischen mit Zusätzen. Landes-Industrie-Comptoir, Wei-
mar, vi + 1—344.

Frýda J. 1998: Higher classification of the Paleozoic gastropods in-

ferred from their early shell ontogeny. In: Bieler R. &
Mikkelsen P.M. (Eds.): 13th International Malacological Con-
gress, Abstracts, Washington D.C., 1—108.

Fuchs T. 1873: Beiträge zur Kenntniss fossiler Binnenfaunen. VI.

Neue Conchylienarten aus den Congerien-Schichten und aus
Ablagerungen der sarmatischen Stufe. Jb. k.k. Geol. R.-A. 23,
19—26.

Férussac A.E.J.P.J.F.d’Audebard de 1821—1822: Tableaux systéma-

tiques des animaux mollusques classés en familles naturelles,
dans lesquels on a établi la concordance de tous les systèmes;
suivis d’un prodrome général pour tous les mollusques ter-
restres ou fluviatiles, vivants ou fossiles. Bertrand Sowerby,
Paris, Londres, 1—110.

Férussac J.B.L. d’Audebard de & Férussac A.E.J.P.J.F. d’Audebard

de 1807: Essai d’une méthode conchyliologique appliquée aux
mollusques fluviatiles et terrestres d’après la considération de
l’animal et de son test. Nouvelle édition augmentée d’une syn-
onymie des espèces les plus remarquables, d’une table de con-
cordance systématique de celles qui ont été décrites par
Géoffroy, Poiret et Draparnaud, avec Müller et Linné, et ter-
minée par un catalogue d’espèces observées en divers lieux de
la France. Delance, Paris, xvi + 1—142.

Gašparík J., Brestenská E., Forgáč J., Franko O., Hajošová M.,

Hanáček J., Marková M., Matkulčík E., Planderová E. & Sitár

154

NEUBAUER, HARZHAUSER and PIPÍK

GEOL

EOL

EOLOGICA CARPA

OGICA CARPA

OGICA CARPATHICA

THICA

THICA, 2015, 66, 2, 139—156

V. 1974: Structural drill GHŠ-1 (Horná Štubňa). Reg. Geol.
Západ. Karpát 3, 1—97 (in Slovak).

Gašparík J., Halouzka R., Miko O., Gorek J., Rakús M., Bujnovský

A., Lexa J., Panáček A., Samuel O., Gašparíková V., Planderová
E., Snopková P., Fendek M., Hanáček J., Modlitba I., Klukanová
A., Žáková E., Horniš J. & Ondrejičková A. 1995: Explanations
to the geological map of the Turiec Depression 1 : 50 000. Štátny
Geologický Ústav Dionýza Štúra, Bratislava, (in Slovak).

Golikov A.N. & Starobogatov Y.I. 1975: Systematics of proso-

branch gastropods. Malacologia 15, 1, 185—232.

Gorjanovic-Kramberger C. 1923: Die Valenciennesiden und einige

anderen Limnaeiden der pontischen Stufe des unteren Pliocaens
in ihrer stratigraphischen und genetischen Bedeutung. Glasn.
Hrv. Prirodosl. Društva 35, 87—114.

Gradstein F.M., Ogg J.G., Schmitz M.D. & Ogg G.M. (Eds.) 2012:

The Geologic Time Scale 2012. 2 vol. Elsevier, Oxford, 1—1176.

Gray J.E. 1840: Shells of molluscous animals. In: Synopsis of the

contents of the British Museum. G. Woodfall, London, 105—152.

Gray J.E. 1847: A list of the genera of recent Mollusca, their syn-

onyma and types. Proc. Zool. Soc. London 15, 129—242.

Halaváts G.v. 1903: Die Fauna der pontischen Schichten in der

Umgebung des Balatonsees. Res. Wissenschaft. Erforsch. Bala-
tonsees 1, 1, 1—80.

Handmann R. 1887: Die fossile Conchylienfauna von Leobersdorf

im Tertiärbecken von Wien. Aschendorff, Münster, 1—47.

Harzhauser M. & Binder H. 2004: Synopsis of the Late Miocene

mollusc fauna of the classical sections Richardhof and Eichko-
gel in the Vienna Basin (Austria, Pannonian, MN9—MN11).
Arch. Molluskenkd. 133, 1—2, 1—57.

Harzhauser M. & Mandic O. 2008: Neogene lake systems of Cen-

tral and South-Eastern Europe: Faunal diversity, gradients and
interrelations. Palaeogeogr. Palaeoclimatol. Palaeoecol. 260,
417—434.

Harzhauser M. & Tempfer P.M. 2004: Late Pannonian Wetland

Ecology of the Vienna Basin based on Molluscs and Lower
Vertebrate Assemblages (Late Miocene, MN 9, Austria). Cour.
Forsch-Inst. Senckenberg 246, 55—68.

Harzhauser M., Kowalke T. & Mandic O. 2002: Late Miocene (Pan-

nonian) Gastropods of Lake Pannon with Special Emphasis on
Early Ontogenetic Development. Ann. Naturhist. Mus. Wien,
Ser. A 103, 75—141.

Harzhauser M., Daxner-Höck G. & Piller W.E. 2004: An integrated

stratigraphy of the Pannonian (Late Miocene) in the Vienna
Basin. Austrian J. Earth Sci. 95/96, 6—19.

Harzhauser M., Latal C. & Piller W.E. 2007: The stable isotope archive

of Lake Pannon as a mirror of Late Miocene climate change.
Palaeogeogr. Palaeoclimatol. Palaeoecol. 249, 335—350.

Harzhauser M., Neubauer T.A., Gross M. & Binder H. 2014a: The

early Middle Miocene mollusc fauna of Lake Rein (Eastern
Alps, Austria). Palaeontogr. Abt. A 302, 1—6, 1—71.

Harzhauser M., Neubauer T.A., Georgopoulou E. & Harl J. 2014b:

The Early Miocene (Burdigalian) mollusc fauna of the North
Bohemian Lake (Most Basin). Bull. Geosci. 89, 4, 819—908.

Harzhauser M., Neubauer T.A., Mandic O., Zuschin M. & Ćorić S.

2012: A Middle Miocene endemic freshwater mollusc assem-
blage from an intramontane Alpine lake (Aflenz Basin, Eastern
Alps, Austria). Palaeontol. Z. 86, 1, 23—41.

Hók J., Kováč M., Rakús M., Kováč P., Nagy A., Kováčová-Slam-

ková M., Sitár V. & Šujan M. 1998: Geologic and tectonic evo-
lution of the Turiec depression in the Neogene. Slov. Geol.
Mag. 4, 165—176.

Hörnes M. 1851—1856: Die fossilen Mollusken des Tertiär-Beckens

von Wien. I. Univalven. Abh. Geol. R.-A. 3, 185—296 (1853),
385—460 (1855), 461—736 (1856).

ICZN 1999: International Code of Zoological Nomenclature. Inter-

national Trust for Zoological Nomenclature, London, 1—306.

Jekelius E. 1932: Fauna Neogenă a României. Die Molluskenfauna

der dazischen Stufe des Beckens von Bra ov. Mem. Inst. Geol.
Rom. 2, 1—118.

Jekelius E. 1944: Sarmat und Pont von Soceni (Banat). Mem. Inst.

Geol. Rom. 5, 1—167.

Jörger K.M., Stöger I., Kano Y., Fukuda H., Knebelsberger T. &

Schrödl M. 2010: On the origin of Acochlidia and other enig-
matic euthyneuran gastropods, with implications for the sys-
tematics of Heterobranchia. BMC Evol. Biol. 10, 323.

Klein A. 1853: Conchylien der Süsswasserkalkformation Württem-

bergs. Jahresh. Ver. vaterl. Naturkd. Württemb. 9, 203—223.

Klika G. 1891: Die Tertiären Land- und Süsswasser-Conchylien des

nordwestlichen Böhmen. Arch. Naturwiss. Landesdurchf. Böh-
men 7, 1—122.

Konečný V., Lexa J. & Planderová E. 1983: Stratigraphy of the

Central Slovakia Volcanic Field. [Stratigrafické členenie neo-
vulkanitov stredného Slovenska.] Západ. Karpaty, Geol. 9,
1—203 (in Slovak with English abstract).

Kováč M., Hók J., Minár J., Vojtko R., Bielik M., Pipík R., Rakús

M., Krá  J., Šujan M. & Králiková S. 2011: Neogene and Qua-
ternary  development  of  the  Turiec  Basin  and  landscape  in  its
catchment: a tentative mass balance model. Geol.  Carpathica
62, 4, 361—379.

Králiková S., Vojtko R., Andriessen P., Kováč M., Fügenschuh B.,

Hók  J.  &  Minár  J.  2014:  Late  Cretaceous—Cenozoic  thermal
evolution  of  the  northern  part  of  the  Central  Western  Car-
pathians  (Slovakia):  revealed  by  zircon  and  apatite  fission
track thermochronology. Tectonophysics 615—616, 142—153.

Krijgsman W., Stoica M., Vasiliev I. & Popov V.V. 2010: Rise and

fall of the Paratethys Sea during the Messinian Salinity Crisis.
Earth Planet. Sci. Lett. 290, 183—191.

Krstić N. 2003: Paludinian beds of Voivodina. In: Papaianopol I.,

Marinescu F., Krstić N. & Macale R. (Eds.): Chronostratigra-
phie und Neostratotypen. Neogen der Zentrale Paratethys, Bd.
X, Pl2. Romanien. Editura Academiei Române, Bucure ti,
1—527.

Krstić N., Savić L. & Jovanović G. 2012: The Neogene lakes of the

Balkan land. Ann. Géol. Pénins. Balk. 73, 37—60.

Linnaeus C. 1758: Systema naturae per regna tria naturae, secun-

dum classes, ordines, genera, species, cum characteribus, dif-
ferentiis, synonymis locis. Tomus 1. Laurentius Salvius,
Holmiae, iii + 1—824.

Lörenthey E. 1902: Die Pannonische Fauna von Budapest. Palaeon-

togr. 48, 137—256.

Magyar I. & Geary D.H. 2012: Biostratigraphy in a Late Neogene

Caspian-Type Lacustrine Basin: Lake Pannon, Hungary. In:
Baganz O.W., Bartov Y., Bohacs K. & Nummedal D. (Eds.):
Lacustrine sandstone reservoirs and hydrocarbon systems.
AAPG Memoir 95, 255—264.

Magyar I., Geary D.H. & Müller P. 1999: Paleogeographic evolu-

tion of the Late Miocene Lake Pannon in Central Europe.
Palaeogeogr. Palaeoclimatol. Palaeoecol. 147, 151—167.

Meijer T. 1989: Notes on Quaternary freshwater Mollusca of the

Netherlands, with descriptions of some new species. Meded.
Werkgr. Tert. Kwart. Geol. 26, 4, 145—181.

Michaud G. 1855: Descriptions de coquilles fossiles découvertes

dans les environs de Hauterive (Drôme). Ann. Soc. Linn. Lyon,
N.S. 2, 33—64.

Milošević V. 1962: Systematic overview of the type specimens from

the paleontological collection of the Natural History Museum
in Belgrade. Bull. Mus. Hist. Nat. Belgrade, Sér. A 16—17, 3—44
(in Serbian).

Milošević V. 1965: Kosovia strophostomopsis nov. spec. from the

freshwater Neogene of the Metohija depression. Bull. Mus.
Hist. Nat. Belgrade, Sér. A 19—20, 113—127 (in Serbian).

Milošević V.M. 1967: On the systematic position of the gastropod

155

ENDEMIC UPPER MIOCENE LACUSTRINE GASTROPODS OF THE TURIEC BASIN

GEOL

EOL

EOLOGICA CARPA

OGICA CARPA

OGICA CARPATHICA

THICA

THICA, 2015, 66, 2, 139—156

species Kosovia compressa P. Pavlović. Bull. Mus. Hist. Nat.
Belgrade, Sér. A 22, 3—15 (in Serbian).

Milošević V.M. 1970: Paleontological and stratigraphical relations

of the gastropod species ex gr. Kosovia strophostomopsis V.
Milošević and Metohia levantica (R. Popović) from lacustrine
Neogene deposits of the Metohija depression. Bull. Mus. Hist.
Nat. Belgrade, Sér. A 25, 71—81 (in Serbian).

Milošević V.M. 1978: Kosovia (Kosovia) striata n. sp. from Lower

Pliocene deposits of the Metohija Depression (Kosovo com-
plex). Bull. Mus. Hist. Nat. Belgrade, Sér. A 33, 175—182 (in
Serbian).

Milošević V.M. 1983: Mollusk freshwater Neogene fauna from Me-

tohija and a genus Theodoxus. Bull. Mus. Hist. Nat. Belgrade,
Sér. A 38, 135—150 (in Serbian).

Montfort P.D.d. 1810: Conchyliologie systématique et classifica-

tion méthodique de coquilles; offrant leurs figures, leur ar-
rangement générique, leurs descriptions caractéristiques, leurs
noms; ainsi que leur synonymie en plusieurs langues. Ouvrage
destiné à faciliter l’étude des coquilles, ainsi que leur disposi-
tion dans les cabinets d’histoire naturelle. Coquilles univalves,
non cloisonnées. 2. Schoell, Paris, 1—676.

Moos A. 1944: Neue Funde von Lymnaeiden, insbesondere von Va-

lenciennesiiden im Pannon Kroatiens. Vjestn. Hrv. Državn.
Geol. Zav. Hrv. državn. Geol. Muz. 2, 3, 341—390.

Neubauer T.A., Harzhauser M. & Kroh A. 2013a: Phenotypic evo-

lution in a fossil gastropod species lineage: evidence for adap-
tive radiation? Palaeogeogr. Palaeoclimatol. Palaeoecol. 370,
117—126.

Neubauer T.A., Harzhauser M. & Mandic O. 2013b: Phenotypic

evolution in a venerid bivalve species lineage from the late
Middle Miocene Central Paratethys Sea: a multi-approach
morphometric analysis. Biol. J. Linnean Soc. 110, 2, 320—334.

Neubauer T.A., Harzhauser M., Kroh A., Georgopoulou E. & Man-

dic O. 2014a: Replacement names and nomenclatural com-
ments for problematic species-group names in Europe’s
Neogene freshwater Gastropoda. Part 2. ZooKeys 429, 13—46.

Neubauer T.A., Harzhauser M., Kroh A., Georgopoulou E. & Man-

dic O. 2014b: The FreshGEN Database: Freshwater Gastro-
pods of the European Neogene.

Accessed at http://www.marinespecies.org/freshgen on 2014-12-16
Neubauer T.A., Harzhauser M., Kroh A., Georgopoulou E. & Man-

dic O. 2015: A gastropod-based biogeographic scheme for the
European Neogene freshwater systems. Earth Sci. Rev. 143,
98—116.

Neumayr M. 1869: Beiträge zur Kenntniss fossiler Binnenfaunen.

II. Jb. k.k. Geol. R.-A. 19, 355—382.

Neumayr M. & Paul C.M. 1875: Die Congerien- und Paludien-

schichten Slavoniens und deren Faunen. Abh. k.k. Geol. R.-A.
7, 3, 1—106.

Olivier G.A. 1804: Voyage dans l’Empire Othoman, l’Égypte et la

Perse, fait par ordre du gouvernement, pendant les six
premières années de la République. Tome second. Agasse, Paris,
1—466.

Papp A. 1953: Die Molluskenfauna des Pannon des Wiener Beckens.

Mitt. Geol. Gesell. Wien 44, 85—222.

Pavlović P.S. 1931: On the fossil mollusk fauna from the Skopje

Basin. Glasn. Skopsk. Naučn. Društva, Odeljenje Prir. Nauka
9, 3, 1—28 (in Serbian).

Pavlović P.S. 1932: New data on the fossil faunas from the Kosovo

and Metohija depressions. Bull. Serv. Géol. Roy. Yougosl. 1, 2,
231—253 (in Serbian).

Pipík R. 2002: Ostracode fauna of the Blážovce and Martin forma-

tions (Turiec basin). Miner. Slovaca 34, 105—112.

Pipík R. 2004: Freshwater ostracods (Ostracoda) and Upper Mio-

cene paleobiotopes of the northern part of the Turiec Depres-
sion (Slovakia). Miner. Slovaca 36, 87—100 (in Slovak).

Pipík R. 2005: Upper Miocene freshwater ostracods (Ostracoda) of

the littoral deposits of the Turiec Formation. Miner. Slovaca 37,
27—40 (in Slovak).

Pipík R. & Bodergat A.-M. 2003a: Ostracodes du Miocène

supérieur du Bassin de Turiec (Slovaquie): familles Darwinul-
idae et Ilyocyprididae. Rev. Esp. Micropal. 35, 345—355.

Pipík R. & Bodergat A.-M. 2003b: Upper Miocene Ostracods of the

Turiec Basin (Slovakia) – sub-family Cyclocypridinae. Ann.
Limnol. – Int. J. Lim. 39, 347—361.

Pipík R. & Bodergat A.-M. 2004a: Euxinocythere (Ostracoda,

Cytheridae, Leptocytherinae) du Miocène supérieur du Bassin
de Turiec (Slovaquie): taxonomie et paléoécologie. Rev. Mi-
cropaleont. 47, 36—52.

Pipík R. & Bodergat A.-M. 2004b: Cyprididae (Ostracoda) du Mio-

cène supérieur du Bassin de Turiec (Slovaquie): Taxonomie et
Paléoécologie. Rev. Micropaleont. 47, 225—242.

Pipík R. & Bodergat A.-M. 2006: Groupe de Candona clivosa, nou-

veau groupe de Candoninae (Crustacea, Ostracoda) et sa diver-
sification dans le Bassin de Turiec (Slovaquie) au Miocène
supérieur. Geobios 39, 3, 394—414.

Pipík R. & Bodergat A.-M. 2007: Candoninae trapézoidales (Crus-

tacea, Ostracoda) du Bassin de Turiec (Slovaquie) du Miocène
supérieur – systématique, écologie et évolution. Geobios 40,
5, 645—676.

Pipík R. & Bodergat A.-M. 2008: Nouvelle faune d’ostracodes

d’eau douce du Miocène supérieur de la Paratéthys Centrale et
description d’espèces appartenant aux genres Pseudocandona,
Fabaeformiscandona et Candonopsis. Palaeontogr. Abt. A
286, 89—121.

Pipík R., Bodergat A.-M., Briot D., Kováč M., Krá J. & Zielinski

G. 2012: Physical and biological properties of the late Mio-
cene, long-lived Turiec Basin, Western Carpathians (Slovakia)
and its paleobiotopes. J. Paleolimnol. 47, 233—249.

Poey F. 1852: Introduction to Cyclostomates with general descrip-

tion of gastropods and special emphasis to terrestrial opercu-
lated gastropods. Mem. Hist. Nat. Cuba 1, 8, 77—96 (in Spanish).

Pokorný V. 1954: Paleontological investigation of the Neogene de-

posits in the vicinity of Sučany—K ačany. Věst. Ústř. Úst. Geol.
29, 81—84 (in Czech).

Popov S.V., Shcherba I.G., Ilyina L.B., Nevesskaya L.A., Para-

monova N.P., Khondkarian S.O. & Magyar I. 2006: Late Mio-
cene to Pliocene palaeogeography of the Paratethys and its
relation to the Mediterranean. Palaeogeogr. Palaeoclimatol.
Palaeoecol. 238, 91—106.

Popović R. 1964: New genus Metohia from the Levantinian of the

Metohija Basin. Nafta 15, 3, 45—53 (in Serbian).

Popović R. 1968: Metohia n. gen. from the Levantinian of the Me-

tohija Basin. Vesn. Zav. Geol. Geofiz. Istr. N.R. Srbije, Ser. A
26, 203—218 (in Serbian).

Popović R. 1969: On a presence of Dacian equivalents in the Meto-

hija Basin. Vesn. Zav. Geol. Geofiz. Istr. N.R. Srbije, Ser. A 27,
105—120 (in Serbian).

Popović R. 1970: Nouveaux représentants du genre de gastéropodes

Viviparus des sédiments levantins du bassin de Metohja. Bull.
Sci., Sect. A 15, 9—10, 317—319.

Rafinesque C.S. 1815: Analyse de la nature ou tableau de l’univers

et des corps organisés. Privately published by author, Palermo,
1—223.

Rakús M., Hók J., Krá J. & Kotulová J. 2005: The Turčianska kot-

lina Depression an example of the Strike-Slipe Basin. Geolines
19, 97—98.

Remeš M. 1923: Oncophora beds near Turčiansky Sv. Martin in Slo-

vakia. Čas. Mineral. Geol. 1, 113 (in Czech).

Roshka V.K. 1973: Maeotian mollusks from the north-western part

of the Black Sea. Shtiintsa, Kishinev, 1—284 (in Russian).

Rzehak A. 1893: Die Fauna der Oncophora Schichten Mährens.

156

NEUBAUER, HARZHAUSER and PIPÍK

GEOL

EOL

EOLOGICA CARPA

OGICA CARPA

OGICA CARPATHICA

THICA

THICA, 2015, 66, 2, 139—156

Verh. Naturforsch. Ver. Brünn 31, 142—192.

Sacchi M. & Horváth F. 2002: Towards a new time scale for the Up-

per Miocene continental series of the Pannonian basin (Central
Paratethys). EGU Stephan Mueller Spec. Publ. Ser. 3, 79—94.

Sandberger C.L.F. 1870—1875: Die Land- und Süßwasser-Con-

chylien der Vorwelt. C. W. Kreidel, Wiesbaden, livr. 1—3: 1—96
(1870); livr. 4—5: 97—160 (1871); livr. 6—8: 161—256 (1872);
livr. 9—10: 257—352 (1873); livr. 11: 353—1000 (1875).

Sandberger C.L.F. 1886: Bemerkungen über einige Binnencon-

chylien des Wiener Beckens. Verh. k.k. Geol. R.-A. 1886, 5,
118—119.

Sauerzopf F. 1952: Beitrag zur Entwicklungsgeschichte des südbur-

genländischen Pannons. Burgenl. Heimatbl. 14, 1, 1—16.

Schlickum W.R. 1971: Zur Systematik fossiler Hyrobiiden. 1. Arch.

Molluskenkd. 101, 159—167.

Schlickum W.R. 1976: Die in der pleistozänen Gemeindekiesgrube

von Zwiefaltendorf a. d. Donau abgelagerte Molluskenfauna
der Silvanaschichten. Arch. Molluskenkd. 107, 1—3, 1—31.

Schlickum W.R. 1978: Die Gattung Tournouerina Schlickum. Arch.

Molluskenkd. 108, 237—243.

Schlickum W.R. & Puisségur J.-J. 1977: Die Molluskenfauna des

Altpleistozäns von St. Bernard (Département Côte-d’Or).
Arch. Molluskenkd. 107, 4, 6, 273—283.

Schlickum W.R. & Puisségur J.-J. 1978: Die Molluskenfauna der

Schichten mit Viviparus burgundinus und Pyrgula nodotiana
von Montagny-les-Beaune (Dép. Côte-d’Or). Arch. Molluskenkd.
109, 1, 3, 1—26.

Schlickum W.R. & Strauch F. 1979: Die Land- und Süßwassermol-

lusken der pliozänen Deckschichten der rheinischen Braunkohle.
Abh. Senckenberg. Naturforsch. Gessell. 536, 1—144.

Stimpson W. 1865: Researches upon the Hydrobiinae and allied

forms: chiefly made from materials in the Museum of the
Smithsonian Institution. Smithson. Misc. Collect. 7, 1—59.

Stoica M., Lazăr I., Krijgsman W., Vasiliev I., Jipa D. & Floroiu A.

2013: Paleoenvironmental evolution of the East Carpathian
foredeep during the late Miocene-early Pliocene (Dacian Ba-
sin; Romania). Glob. Planet. Change 103, 135—148.

Strausz L. 1942: Viviparen aus dem Pannon Mittel-Transdanubiens.

M. Kir. Földt. Int. Évk. 36, 1, 1—63.

Štúr D. 1860: Bericht über die geologische Uebersichts-Aufnahme

des Wassergebietes der Waag und Neutra. Jb. k.k. Geol. R.-A.
11, 17—151.

ter Borgh M., Vasiliev I., Stoica M., Knežević S., Matenco L., Krijg-

sman W., Rundić L. & Cloetingh S. 2013: The isolation of the
Pannonian basin (Central Paratethys): New constraints from
magnetostratigraphy and biostratigraphy. Glob. Planet. Change
103, 99—118.

Tournouër R. 1879: Conchyliorum fluviatilium fossilium, quae in

stratis tertiariis superioribus Rumaniae Dr. Gregorio Stefa-
nesco collegit, novae species. J. Conchyl. 27, 261—264.

van Leeuwen C.H.A., van der Velde G., van Lith B. & Klaassen M.

2012: Experimental quantification of long distance dispersal
potential of aquatic snails in the gut of migratory birds. PLoS
ONE 7, 3, e32292.

Vasiliev I., Iosifidi A.G., Khramov A.N., Krijgsman W., Kuiper K.,

Langereis C.G., Popov V.V., Stoica M., Tomsha V.A. & Yudin
S.V. 2011: Magnetostratigraphy and radio-isotope dating of
upper Miocene-lower Pliocene sedimentary successions of the
Black Sea Basin (Taman Peninsula, Russia). Palaeogeogr.
Palaeoclimatol. Palaeoecol. 310, 163—175.

Vetters H. 1910: Beiträge zur Geologie des Zjargebirges und

des angrenzenden Teiles der Mala Magura in Oberungarn.
Denkschr. Kaiserl. Akad. Wiss. Math-Naturw. Kl. 85, 1—60.

Vigh G. 1915: Geological observations between the border-moun-

tains of Nitra, Turiec, and Trenčín counties. M. Áll. Földt. Int.
Évi Jel. 1914, 65—96 (in Hungarian), 71—106 (in German).

Welter-Schultes F.W. 2012: European non-marine molluscs, a

guide for species identification. Planet Poster Editions, Göt-
tingen, 1—679.

Wenz W. 1923—1930: Fossilium Catalogus I: Animalia. Gastropoda

extramarina tertiaria. W. Junk, Berlin, I: 1—352 (1923), II:
353—736 (1923), III: 737—1068 (1923), IV: 1069—1420 (1923),
V: 1421—1734 (1923), VI: 1735—1862 (1923), VII: 1863—2230
(1926), VIII: 2231—2502 (1928), IX: 2503—2886 (1929), X:
2887—3014 (1929), XI: 3015—3387 (1930).

Wenz W. 1928: Zur Systematik tertiärer Land- und Süßwassergas-

tropoden. VIII. Senckenbergiana 10, 220.

Wesselingh F.P., Cadée G.C. & Renema W. 1999: Flying high: on

the airborne dispersal of aquatic organisms as illustrated by the
distribution histories of the gastropod genera Tryonia and Pla-
norbarius. Geol. En Mijnb. 78, 165—174.

Zilch A. 1959—1960: Gastropoda. Teil 2: Euthyneura. In: Schinde-

wolf O.H. (Ed.): Handbuch der Paläozoologie. Band 6. Verlag
Gebrüder Borntraeger, Berlin, xii + 1—834.