Fossil Clausiliidae

Fossil Clausiliidae

Fossil Clausiliidae from Europe are known since the Upper Cretaceous and traced through the Tertiary and Quaternary.
In contrast to several other fossil stylommatophoran groups, relationships of fossil Clausiliidae to other ones and to extant groups can be recognized with high probability, because they are highly diverse and rich in characters, especially of their closing apparatus.
Rich representative faunas of fossil Clausiliidae allow conclusions to climate and vegetation; changes of clausiliid faunas are the result of climatic changes.
As a rule, because of their high evolutionary rate, fossil Clausiliidae have a small stratigraphic range. Therefore, in comparison with other stylommatophoran groups, they have a high biostratigraphic value.
Fossil Clausiliidae give keys for the reconstruction of the evolution of the family, because relationships to extant species can be recognized (see 2.) and appearance dates (first and last ones) can be stated.
Nevertheless, there are several extinct clausiliid groups the relationships of which are unclear, especially if the material is scarce and essential characters cannot be examined.
In the following two reports on my current work on fossil Clausiliidae are given:

List of fossil Clausiliidae from western Palaearctic;

Pliocene and Pleistocene Clausiliidae from central Europe north of the Alps (with discussion of their first appearance dates).

List of fossil Clausiliidae from western Palaearctic

Hartmut Nordsieck (III.2021)

A revised genus list of fossil Clausiliidae from western Palaearctic (except Anatolia) with stratigraphic ranges is given as follows:

Phaedusinae A. J. Wagner 1922

Former Serrulininae

Serrulellini H. Nordsieck 2007
Serrulella H. Nordsieck (in Zilch) 1978 (Burdigalian – Early Pleistocene)
Serrulastra H. Nordsieck 1981 (Aquitanian – Zanclean)
(S. (Serrulastra), S. (Serruplica) H. Nordsieck 1981, S. (Serrustigma) H. Nordsieck 1981)

Nordsieckiini H. Nordsieck 2007
Nordsieckia Truc 1972 (Tortonian – Piacenzian)
Note: Nordsieckia has a much plesiomorphic shell with columellar lamellae in the upper whorls (Nordsieck 2015: fig. 8).

New tribe
Serruluna H. Nordsieck 1981 (Piacenzian – Early Pleistocene)
Note: According to the completed diagnosis (Nordsieck 2013) Serruluna is not related to Nordsieckia (contrary to Nordsieck 2007).

Serrulinini Ehrmann 1927
Pravispira Lindholm 1924 (Early Pleistocene)

Ciscaucasian former Serrulininae (see Steklov 1966)

Uncertain tribe
Euxinophaedusa Likharev 1962 (Serravallian – Tortonian)

Serrulinini Ehrmann 1927
Serrulina Mousson 1873 (Tortonian)
Pravispira Lindholm 1924 (Tortonian)
(P. (Pravispira), Pravispira sensu Likharev 1962 = P. n. subgen.)
Pontophaedusa sensu Likharev 1962 = n. gen. (Tortonian)
Notes: Two species described by Likharev (: 303-306) are different from the genera or subgenera, respectively, with which he has affiliated them.
Serrulina (Pravispiranazranica looks like a Pravispira species, but differs from the other species of the genus by the decollate shell and the presence of only two palatal plicae; its clausilium plate is without inner fold. For it a new subgenus is erected.
Pontophaedusa praefuniculum differs much from Pfuniculum by a higher superior and less protruding inferior lamella, the presence of three palatal plicae (lower one misinterpreted as lunella by Likharev) and the gutter-like clausilium plate. Because in the Caucasian region the plicae type of lunellar is only found in Serrulinini species, the new genus is temptatively classified with that tribe.

Laminiferinae Wenz 1923

Laminiferini
Laminifera O. Boettger 1863 (Thanetian – Burdigalian)
(L. (Palaeophaedusa) Wenz 1920, L. (Laminifera))
Omanifera H. Nordsieck 2015 (Priabonian)
Baboria Cossmann 1898 (Burdigalian)
Laminiplica H. Nordsieck 1978 (Zanclean – Piacenzian)

Oospiroidini H. Nordsieck 2007
Oospiroides Wenz 1920 (Thanetian – Ypresian)

Polloneriini H. Nordsieck 2007
Polloneria Sacco 1886 (Piacenzian – Early Pleistocene)

Eualopiinae H. Nordsieck 1978

Rillyini H. Nordsieck 1985
Proalbinaria O. Boettger (in Oppenheim) 1895 (Maastrichtian)
Rillyopsis H. Nordsieck 1985 (Thanetian)
Rillya Munier-Chalmas (in P. Fischer) 1883 (Thanetian – Ypresian?)
Pararillya H. Nordsieck 2002 (Thanetian)
Rillyella H. Nordsieck 2002 (Thanetian)
Rillya sensu auct. incertae sedis (Thanetian – Lutetian)
Neniopsis Wenz 1920 (Thanetian – early Rupelian)
Rillyarex H. Nordsieck 1985 (Priabonian)
Omanyllia H. Nordsieck 2015 (Priabonian)
Neniopsilla H. Nordsieck 2020 (Priabonian)

Disjunctariini H. Nordsieck 2014
Note: Unlike former Serrulininae, Disjunctariini have an adnate peristome like the Eualopiinae. Therefore and because of their age (Middle Eocene) they are classified with that subfamily. Recent findings of well-preserved shells in the Roncà layers (Zamberlan w. c.) confirm this classification, because one of the species contained (Dsatyrus Oppenheim) is very similar to Eualopia.
Disjunctaria O. Boettger 1877 (Bartonian)

Eualopiini
Eualopia O. Boettger 1877 (Aquitanian)

New tribe
Monoptychia H. Nordsieck 1972 (Piacenzian)
(M. (Monoptychia), M. (Protoptychia) H. Nordsieck 2007)
Note: The differences in shell characters (Nordsieck 2007: 173) and the large stratigraphical gap to other Eualopiinae speak for a tribe separation of Monoptychia.

Constrictinae H. Nordsieck 1981
Constricta O. Boettger 1877 (Chattian – Serravallian)
Regiclausilia H. Nordsieck 1981 (Serravallian)

Alopiinae A. J. Wagner 1913

Alopiini
Protriloba H. Nordsieck 2014 (Serravallian – Tortonian)
Note: The shell characters of Protriloba would also fit the tribe Macedonicini.

Cochlodinini Lindholm 1925 (1923)
Cochlodina A. Férussac 1821(Burdigalian – recent)
(C. (Miophaedusa) H. Nordsieck 1972, C. (Procochlodina) H. Nordsieck 1969, C. (Cochlodina), C. (Paracochlodina) H. Nordsieck 1969)

Clausiliinae

Mentissoideini
Mentissoidea sensu Likharev 1962 (Tortonian)
Elia H. Adams & A. Adams 1855 (Pliocene)
(E. (Caucasica) O. Boettger 1877)

Emarginariini H. Nordsieck 2007
Emarginaria O. Boettger 1877 (Burdigalian – Zanclean)

Clausiliini
Truciella H. Nordsieck 1978 (Messinian – Zanclean)
Ruthenica Lindholm 1924 (Piacenzian – recent)
Canalicia O. Boettger 1863 (Rupelian?, Chattian – Burdigalian)
Pseudidyla O. Boettger 1877 (Burdigalian – Serravallian)
(P. (Pseudidyla), P. (Canaliciella) H. Nordsieck 1981)
Trolliella H. Nordsieck 1981 (Serravallian)
Macrogastra Hartmann 1841 (Serravallian – recent)
(M. (Macrogastra), M. (Macrogastrella) n. subgen., M. (Pyrostoma) Vest 1867)
Clausilia Draparnaud 1805 (Serravallian – recent)
(C. (Hollabrunnella) H. Nordsieck 2007, C. (Clausilia), C. (Lombardiella) H. Nordsieck 2013, C. (Neostyriaca) A. J. Wagner 1920)
Parafusulus H. Nordsieck 2007 (Piacenzian)
Pliodiptychia H. Nordsieck 1978 (Piacenzian)
Fusulus Fitzinger 1833 (Piacenzian – recent)
(F. (Pliofusulus) H. Nordsieck 2013, F. (Fusulus))

Baleini A. J. Wagner 1913
Quadriplicata O. Boettger 1878 (Tortonian – Pliocene)
Laeviphaedusa sensu Likharev & Steklov 1965 = n. gen. (Tortonian)
Note: Laeviphaedusa miocaenica Likharev & Steklov 1965 belongs certainly not to Laeviphaedusa, but to a genus of its own. Its smaller and ribbed shell is not semiapostrophic like in Lhyrcanica. The lunellar is much different: lower palatal plica with posterior part, anterior part well-developed, anterior upper palatal plica and palatal callus present. It does not resemble to any group of former Serrulininae. Because of its prominent anterior lower palatal plica, keel furrow and palatal callus it is tentatively classified with the Baleini.
Laciniaria Hartmann 1842 (Middle Pleistocene – recent)
Alinda H. Adams & A. Adams 1855 (Late Pleistocene – recent)
Strigillaria Vest 1867 (Early Pleistocene – recent)
(S. (Strigillaria))

References

Likharev, I. M. (1962): Klauziliidy (Clausiliidae). – In: Fauna SSSR, Molljuski, 3 (4): 317 pp. Moskva & Leningrad.

Nordsieck, H. (2007): Worldwide Door Snails (Clausiliidae), recent and fossil. – 214 pp., 20 pls. Hackenheim (ConchBooks).

Nordsieck, H. (2013): Clausiliidae (Gastropoda, Stylommatophora) from the Villafranchian site Monte Serampoli (Lower Valdarno, Tuscany, Italy). – Archiv für Molluskenkunde, 142 (2): 237-244.

Nordsieck, H. (2015): Fossil Clausilioidea in space and time, with special emphasis on Cretaceous and pre-Oligocene Cenozoic Clausiliidae (Gastropoda: Stylommatophora). – Archiv für Molluskenkunde, 144 (1): 83-97.

Steklov, A. A. (1966): Nazemnye molljuski neogena predkavkasja i ih stratigrafičeskoe značenje (Terrestrial Neogene mollusks of Ciscaucasia and their stratigraphic importance). – Transactions of the Academy of Sciences of the USSR, Geological Institute, 163: 262 pp., 14 pls. Moskva.

 

Pliocene and Pleistocene Clausiliidae from central Europe north of the Alps

Hartmut Nordsieck (VII.2021, supplemented XI.2021)

I. Faunas from Pliocene to early Middle Pleistocene

The main Pliocene and Pleistocene clausiliid faunas from central Europe north of the Alps, which have been studied more thoroughly, are listed in the following. They are ordered chronologically; for more detailed information on their age see Nordsieck 2007 (: 139-140) and the literature mentioned there (: 142-143).
The formerly studied Pliocene faunas from western and central Europe (F, Celleneuve; D, Frechen-Fortuna; F, Sessenheim; F, Cessey-sur-Tille) are treated in former papers (see References); they are not considered in this article.
Species, which were not traced by myself, are listed in brackets.

Pliocene:

A, Neudegg: Serrulella austriaca, Nordsieckia succedens, Ruthenica filograna, R. cf. filograna, Macrogastra densestriata, Clausilia stranzendorfensis, C. strauchiana geisserti, Parafusulus neudeggensis.

A, Stranzendorf A: Clausilia stranzendorfensis, C. s. geisserti.

A, Stranzendorf C: Serruluna pretiosa, Serrulella? sp. (apical fragments), Clausilia stranzendorfensis, C. s. geisserti.

Early Pleistocene:

A, Stranzendorf D: Macrogastra sessenheimensis? (apical fragments), Clausilia dubia, C. pumila.

A, Stranzendorf F: Cochlodina laminata, Ruthenica filograna, Macrogastra plicatula, M. sessenheimensis, C. cruciata, Clausilia pumila, C. corynodes ornatula.

A, Stranzendorf G: C. cruciata, Clausilia pumila.

A, Stranzendorf K: Macrogastra densestriata, M. plicatula, C. cruciata, Clausilia pumila.

A, Stranzendorf K/L: Macrogastra ventricosa? (apical fragments), M. plicatula, C. cruciata, Clausilia pumila, C. c. ornatula.

A, Stranzendorf L: Macrogastra ventricosa, Clausilia cruciata, C. c. ornatula

D, Fischach: Macrogastra ventricosa, M. sessenheimensis, Clausilia dubia, C. pumila, C. c. ornatula.

D, Markt Buch 1-7: Macrogastra ventricosa, Clausilia dubia, C. cruciata, C. pumila, C. c. ornatula, C. dehmi.

D, Holzkirchen: Clausilia pumila, C. rugosa antiquitatis, C. c. ornatula.

D, Dinkelscherben-Uhlenberg: Macrogastra sessenheimensis, Clausilia dubia, C. cruciata, C. pumila, C. c. ornatula, C. dehmi.

A, Krems-Schießstätte 11: Macrogastra sessenheimensis.

A, Krems-Schießstätte 9: Serruluna aptyx, Serrulella? sp. (apical fragment), Ruthenica filograna, (Macrogastra densestriata), (M. sessenheimensis), Clausilia dubia, C. pumila, C. c. ornatula.

PL, Kielniki 3A: Serruluna biptyx, Cochlodina cerata, C. orthostoma, Ruthenica filograna, Macrogastra borealis, M. plicatula, Clausilia dubia, C. pumila.

A, Radlbrunn: Clausilia dubia, C. r. antiquitatis.

A, Deutsch-Altenburg 30A: Macrogastra ventricosa, M. plicatula, Clausilia dubia, C. cruciata, C. pumila, C. c. ornatula.

A, Deutsch-Altenburg 4B: Serrulella ultima, S.? sp. (apical fragment), Cochlodina laminata, Macrogastra ventricosa, M. densestriata, M. plicatula, Clausilia dubia, C. cruciata, C. pumila, C. c. ornatula, (Strigillaria cana).

Middle Pleistocene:

D, Weißenburg 7: Cochlodina laminata, Macrogastra ventricosa, M. densestriata, M. plicatula, Clausilia dubia, C. cruciata, C. pumila, Fusulus interruptus.

PL, Kozi Grzbiet: Cochlodina laminata, C. cerata, C. orthostoma, Ruthenica filograna, Macrogastra densestriata, M. tumida, M. plicatula, C. cruciata, Clausilia pumila, Strigillaria cana.

A, Hundsheim: Cochlodina laminata, Ruthenica filograna, Macrogastra ventricosa, M. tumida, M. plicatula, Clausilia dubia, C. pumila, C. c. ornatula.

D, Wiesbaden-Mosbach III: Ruthenica filograna, Macrogastra ventricosa, M. plicatula, Clausilia dubia, C. cruciata, C. pumila, C. r. antiquitatis, C. c. ornatula.

F, Achenheim: Clausilia dubia, C. pumila, C. r. antiquitatis, C. c. ornatula.

D, Wiesbaden-Mosbach IV: Clausilia pumila, C. r. parvula, C c. ornatula.

Notes:

Neudegg (Frank & Rabeder 1996, Döppes & Rabeder 1997: 106-110):
Like the fauna of micromammals (Döppes & Rabeder: 108)  the clausiliid fauna is clearly of Pliocene age (Nordsieck 2000: 2, foot note 2; 2007: 138-139), because all species except of Macrogastra densestriata are extinct. An age higher than 2.6 Ma is also proved by the occurrence of a Triptychia species (T. neudeggensis Schnabel).
The species given by Frank (in Döppes & Rabeder: 106) as Macrogastra sp. is Parafusulus neudeggensis (Nordsieck 2007: 173); the species determined as Serrulina serrulata is Serrulella austriaca (Nordsieck: 160). As concerns Ruthenica species, a complete specimen of R. filograna and an apertural fragment somewhat differing from that species (R. cf. filograna) are present.

Stranzendorf (Döppes & Rabeder 1997: 130-139):
As to the clausiliids of the Pliocene layers of Stranzendorf (Sd A, Sd C) see Nordsieck (1990: 162-163; 2007: 138-139). The hypothesis of Frank (in Döppes & Rabeder: 132, 136) concerning the continued occurrence of these Pliocene Clausilia species (C. strauchiana, C. stranzendorfensis) in Early Pleistocene layers of Stranzendorf, which later on (2006: 366) was repeated, is based on confusion with similar Clausilia species (C. pumila, C. cruciata).
As to the clausiliids of the Early Pleistocene layers (Sd D-L) only a part of the material was available for me. The determination of certain Macrogastra species in Sd layers by Frank (: 132) may refer to M. sessenheimensis and (or) M. plicatula. In contrast to her information (: 132) I could not find M. badia and M. tumida in any Sd fauna, M. densestriata only as one fragment in Sd K. My determination of Clausilia r. antiquitatis? from Sd F (Nordsieck 2007: 139) was wrong, caused by confusion with C. cruciata.

Dinkelscherben-Uhlenberg and other sites of the Upper Swabian Deckschotter (Dehm 1979, Nordsieck 1990: 153, Rähle 1995):
Within the examined clausiliid faunas, of which the four richest ones are listed here, nearly the same species.have been found (Nordsieck: 153, foot note 22); therefore their age may not be much different (Rähle: 114). The species formerly treated as subspecies of Macrogastra plicatula (Rähle: 111) is M. sessenheimensis, which is known from Pliocene to lower Early Pleistocene. Its occurrence is a proof that the layers of the Deckschotter cannot be younger than lower Early Pleistocene. In contrast to the information of Rähle (: 111) Clausilia r. antiquitatis was found by me only in the material from Holzkirchen.

Krems-Schießstätte (Döppes & Rabeder 1997: 28-34):
From the numerous layers of Krems only small samples from Kr 7, 7/2, 9 and 11 were available (the material examined by Ložek 1976, 1978 could not be revised).
From Kr 9 Ložek (1978: 30) mentioned an “Iphigena sp. (kleine Form)”, which is probably Macrogastra sessenheimensis. From Kr 12 he gave the same species (“kleine Iphigena”) and M. densestriata.
The record of Macrogastra plicatula in Kr 9 by Frank (in Döppes & Rabeder: 30) could also refer to M. sessenheimensis. The other Macrogastra species given by her for that layer (determined as M. lineolata and cf. latestriata) might be M. densestriata.
The species mentioned by Ložek (: 28-29) as Serrulina aff. serrulata from Kr 5 (and Kr 6, with question mark) belongs with high probability to the genus Serrulella. These layers, the age of which is questionable because of lack of micromammals (Rabeder 1981: 334-336), can therefore be dated as Early Pleistocene (see Nordsieck 2007: 141); thus, they may not be much younger than Kr 7-12.

Kielniki (Stworzewicz 1981):
In Nordsieck 1990 (: 154, fig. 6) and 2000 (: 3, foot note 4) the age of the clausiliid fauna of Kielniki 3A was given as too high; according to Nadachowski (1990: 217-218) it is Early Pleistocene, biostratigraphically lower Biharian, Mokra phase. The determination of the clausiliids from Kielniki by Stworzewicz has already been revised (Nordsieck 2000: 3, foot note 4).

Radlbrunn (Döppes & Rabeder 1997: 114-116):
As to the age of the clausiliid fauna see Nordsieck 1990 (: 153). This fauna is remarkably poor; it includes only Clausilia dubia and a great amount of C. rugosa antiquitatis.

Deutsch-Altenburg (Döppes & Rabeder 1997: 238-270):
The examined clausiliid faunas of the Deutsch-Altenburg layers of Early Pleistocene age (DA 30A, 2C1, 37, 35, 4B), of which only two are listed here, include nearly the same species. The Macrogastra species of these layers given by Frank (in Döppes & Rabeder: 243, 262, 265) as M. densestriata and M. badia are probably M. plicatula; only in DA 4B one fragment of a new subspecies of M. densestriata could be found. The Baleini species given by her (: 243, Strigillaria cana, Laciniaria plicata, Alinda biplicata) could not be traced in the extensive material available for me; only the presence of S. cana is probable.

Middle Pleistocene faunas:
The early Middle Pleistocene faunas, which were examined more thoroughly, Weißenburg 7 (Dehm 1971), Kozi Grzbiet (Stworzewicz 1981), Hundsheim (Frank in Döppes & Rabeder 1997: 271) and Wiesbaden-Mosbach III and IV (Gruner & Gruner 2009, 2011), were taken for comparison with the older faunas.
As concerns the age of Weißenburg 7, the information „Altpleistozän“ given by Koenigswald (1971) has according to the micromammals listed by him (: 122) to be corrected to Middle Pleistocene, biostratigraphically upper Biharian, Templomhegy phase (Nordsieck 2007: 140).
The clausiliids from Weißenburg 7 have already been listed by Dehm (: 81-82). The material, which was determined by him as Clausilia cruciata n. subsp.?, contained two species, C. cruciata and C. pumila (see part III). C. r. parvula does not occur in Weißenburg 7 (Dehm w. c.).
The determination of the clausiliids from Kozi Grzbiet by Stworzewicz has already been revised (Nordsieck 2007: 140).
Macrogastra cf. densestriata given by Frank (: 271) for Hundsheim may be M. plicatula. Clausilia schlickumi described from Hundsheim is a form of C. corynodes ornatula (Nordsieck 2007:174).
From Wiesbaden-Mosbach (Mosbach layers) not only the material of Gruner, but also that from older collections (SMF) was available. The species determined by former authors as Clausilia bidentata is C. r. antiquitatis.

II. Records of Pliocene and Pleistocene Clausiliidae from central Europe

Because the records of Clausiliidae, especially their first appearance dates = FADs, are important for the calibration of DNA trees, in order to get chronograms, they should be reliable. This is not the case for some records in Harzhauser & Neubauer (2021, supplementary table 1), which will be used for that purpose. In this table the age of all records from Stranzendorf = Sd is given as Piacenzian, though only Sd A – Sd C have Pliocene age (see part I).
The FADs of the species concerned are given as follows:

Serrulina serrulata: Records in layers of central Europe probably based on confusion with Serrulella species (see part I).

Cochlodina laminata: Sessenheim (Piacenzian)?, Sd F (Early Pleistocene). The record in the Sessenheim layer is uncertain, because only one fragment was found, no one in the material got later on (see Nordsieck 1976).

C. orthostoma: Kielniki 3A (Early Pleistocene). The record as cf. in the older layer Sd C is doubtful (Frank in Döppes & Rabeder 1997: 132).

C. cerata: Kielniki 3A (Early Pleistocene).

Ruthenica filograna: Neudegg (Piacenzian).

Macrogastra ventricosa: Sd K/L (Early Pleistocene)?, Sd L.

M. tumida: Kozi Grzbiet, Hundsheim (Middle Pleistocene). The record as cf. in Sd F is doubtful (Frank in Döppes & Rabeder: 132). Record as cf. in Kr 8/2 (Ložek 1976) not checked.

M. densestriata: Frechen-Fortuna (Piacenzian).

M. badia: No fossil record. The record in Deutsch-Altenburg = DA 4B is probably wrong (see part I).

M. borealis: Kielniki 3A (Early Pleistocene). The record as M. cf. latestriaia in Krems = Kr 9 (Frank in Döppes & Rabeder: 30) is probably wrong (see part I).

M. attenuata: (Holocene). The record as M. lineolata in Kr 9 (Frank in Döppes & Rabeder: 30) is probably wrong (see part I).

M. sessenheimensis: Sessenheim (Piacenzian). For the last appearance date = LAD see part III.

M. plicatula: Sd F (Early Pleistocene).

Clausilia baudoni: Moncucco Torinese (Messinian). Record in Ambérieu-le-Bugey (Tortonian) not checked.

C. bidentata crenulata: Monte Serampoli (Early Pleistocene). FAD of C. b. bidentata in central Europe in Middle Pleistocene (see Nordsieck 1990). The record in Sessenheim is caused by admixture of an extant fragment (see Nordsieck 1976).

C. strauchiana: Eichkogel (Tortonian). Record in Dolné Orešany (Tortonian) not checked. For the LAD see part I. 2.

C. pumila: Sd D (Early Pleistocene).

C. stranzendorfensis: Neudegg, Sd A (Piacenzian). For the LAD see part I. 2.

C. cruciata: Sd F (Early Pleistocene).

C. dubia: Sd D (Early Pleistocene).

C. rugosa antiquitatis: Tegelen C5 (Early Pleistocene) (see Nordsieck 1990). The record in Sd F is based on a wrong determination (see part I). For the LAD and the relation to C. r. parvula see part III.

C. corynodes ornatula: Sd F (Early Pleistocene). The record in Sessenheim is caused by admixture of an extant fragment (see Nordsieck 1976). The record in Unterparschenbrunn (Pliocene) is wrong. No fossil record of extant C. corynodes subspecies.

Fusulus interruptus: Weißenburg 7 (Middle Pleistocene).

Strigillaria cana: DA 4 (Early Pleistocene) (see part I).

Laciniaria plicata: Gombasek (Middle Pleistocene). The record in DA 4B is probably wrong (see part I).

Alinda biplicata: (Late Pleistocene). The record in DA 4B is probably wrong (see part I).

III. Pleistocene (and related Pliocene) taxa

Extant species means subspecies of the respective species occurring in the region of the locality.
The given values of rib density (R1) are means of samples.

Cochlodina (C.) laminata (Montagu 1803)

C. laminata from Early Pleistocene (Sd F, Kr 7/2, DA 4B) like that from Middle Pleistocene (Weißenburg 7, Hundsheim) and extant species; lowermost palatal plica distinct. = C. l. laminata.

Ruthenica filograna (Rossmässler 1836)

R. filograna from Early Pleistocene (Sd F, Kr 9) similar to those from Middle Pleistocene (Wiesbaden-Mosbach, Hundsheim) and extant species. Pleistocene specimens relatively small, exhibiting a distinct palatal callus (less in Wiesbaden-Mosbach). = R. filograna.

Macrogastra (M.) ventricosa (Draparnaud 1801)

M. ventricosa from Early Pleistocene (Sd L, DA (4 localities)) on average somewhat smaller than those from Middle Pleistocene and extant species; both folds of inferior lamella about equally strong; in part with rudiment of anterior lower palatal plica on palatal callus. = M. v. cf. major Rossmässler.
M. ventricosa from Middle Pleistocene (3 localities) like extant species; in Wiesbaden-Mosbach lower fold of inferior lamella weakened. = M. v. ventricosa.

M. (n. subgen.) borealis (O. Boettger 1878)

M. borealis from Kielniki like extant species, with relatively dense ribbing (R1 6.3). = M. b. bielzi H. Nordsieck.

M. (n. subgen.) densestriata (Rossmässler 1836)

Only few records of M. densestriata from Early Pleistocene (Sd K, Kr 12, 9?).
M. densestriata n. subsp. from DA 4B.
M. densestriata from Middle Pleistocene (5 localities) with rib density like M. d. densestriata (R1 7.4-8.2), except samples from Kozi Grzbiet with coarser ribbing like M. d. gredleri H. Nordsieck (R1 6.7).

M. (Pyrostoma) sessenheimensis (H. Nordsieck 1974)

Smaller than M. plicatula; superior lamella smoothly continuous with spiral lamella; lower interlamellar fold shifted inwards; only one fold of inferior lamella, if present; subcolumellar lamella receding; anterior lower palatal plica present, high in position, not elongated inwards.
Forms from Sessenheim, Pliocene (type form), and Sd F more densely ribbed than  form from Deckschotter localities (Fischach, Uhlenberg).
Extinct at the end of  early Early Pleistocene (latest occurrence in Krems 6, Ložek 1978: 29, if “Iphigena sp.” = M. sessenheimensis).

M. (P.) plicatula (Draparnaud 1801)

M. plicatula from Sd F coarsely ribbed (only one incomplete fragment available).
M. plicatula from later Early Pleistocene (from Sd K to DA 4B, 6 localities) smaller than that from Weißenburg 7 and extant species; more densely ribbed than most extant subspecies (R1 7.5-9.0); with two folds of inferior lamella; in part anterior lower palatal plica present. = M. plicatula cf. aprutica H. Nordsieck.
M. plicatula from Middle Pleistocene (3 localities) like extant species (R1 5.5-6.5). = M. p. plicatula.

Clausilia (C.) dubia Draparnaud 1805

C. dubia from Sd D ventricose, with short apical part; inferior lamella like C. d. vindobonensis A. Schmidt. = C. d. cf. vindobonensis.
C. dubia from Sd I, L/M, Kr (2 localities) and DA (4 localities) inferior lamella and clausilium plate, if present, like C. d. vindobonensis. = C. d. vindobonensis.
C. dubia from Kielniki, Kr 7/2 and Radlbrunn corresponding, but outer corner of clausilium plate right-angled, less or not bent up. = forms intermediate between C. d. vindobonensis and C. d. dubia.
C. dubia from Middle Pleistocene of Germany (2 localities) inferior lamella and clausilium plate like C. d. dubia. = C. d. dubia.
C. dubia from Hundsheim = C. d. vindobonensis.
All Pleistocene forms are normally ribbed (R1 between 7.5 and 10).

 C. (C.) stranzendorfensis H. Nordsieck 1990

Assumed to be closely related to C. cruciata. Differs from that species mainly by the development of the inferior lamella (upper fold in part reduced).
Forms from 3 Pliocene localities (Neudegg, Sd A, Sd C) somewhat different (Nordsieck: 162-163), if subspecies?
On average larger than C. strauchiana geisserti; inferior lamella more strongly s-like ascending, in part upper fold of inferior lamella reduced; subcolumellar lamella more strongly bent below; outer corner of clausilium plate upbent, but differently distinct. Ribbing like C. s. geisserti (R1 8.0-8.2).

C. (C.) cruciata (Studer 1820)

C. cruciata from Early Pleistocene (from Sd F to DA 4B, 8 localities) relatively densely ribbed (R1 6.9-7.9); outer corner of clausilium plate upbent. = C. cruciata cf. bonellii (form from Abruzzo).
C. cruciata from Weißenburg 7 like those from Early Pleistocene; = C. c. cf. bonellii. Compared with C. pumila from the same site smaller, more densely ribbed (R1 6.9, pumila 5.8); inferior lamella more s-like ascending, folds in part more distinct; subcolumellar lamella more bent below; outer corner of clausilium plate more often pointed.
C. cruciata from Wiesbaden-Mosbach like extant species; outer corner of clausilium plate right-angled, less upbent. = C. c. cruciata.
Remark: C. cruciata and C. pumila from Early Pleistocene deposits can hardly be distinguished, because they agree in size, sculpture, upper lamellae and plicae. The only differences are those of the lower lamellae: inferior lamella in C. cruciata more s-like ascending, with more distinct folds in front; subcolumellar lamella more bent below. Because C. cruciata and C. pumila from the Middle Pleistocene deposit Weißenburg 7 are clearly separable, it is assumed that also in Early Pleistocene deposits both species are present.

 C. (C.) strauchiana geisserti H. Nordsieck 1976

Assumed to be closely related to C. pumila.
Smaller than C. pumila; more densely ribbed (R1 8.0-8.7); subcolumellar lamella strongly bent; anterior lower palatal plica and palatal hump strongly developed; outer corner of clausilium plate upbent, pointed.
Specimens from 5 Pliocene localities not much different.

C. (C.) pumila C. Pfeiffer 1828

C. pumila from Early Pleistocene (from Sd D to DA 4B, 11 localities) on average smaller, more densely ribbed than Middle Pleistocene and extant species (R1 6.3-8.5); superior lamella continuous with spiral lamella; outer corner of clausilium plate upbent (Sd, 3 localities) or not or less upbent (Uhlenberg, DA, 2 localities). = C. p. cf. pumila.
C. pumila from Middle Pleistocene (8 localities) with superior lamella continuous with spiral lamella, more coarsely ribbed (R1 5.8-6.8):form from Weißenburg 7 like extant species, outer corner of clausilium plate upbent;
form from Wiesbaden-Mosbach and other western localities (5 localities) corresponding, outer corner of clausilium plate less upbent;
both = C. p. pumila.
C. p. sejuncta Mörch 1864 could not be traced in the Middle Pleistocene deposits.

C. (C.) rugosa antiquitatis H. Nordsieck 1990

Examined from deposits of Early Pleistocene (Deckschotter, Radlbrunn, DA, 4 localities) and Middle Pleistocene (only western localities, 6).
In comparison with other subspecies of C. rugosa densely ribbed (type form R1 10.5-11.7); with both folds of inferior lamella or only the upper one; outer corner of clausilium plate ± upbent and pointed.
Type form from Radlbrunn with more weakened folds of inferior lamella.
Samples from late early Middle Pleistocene (3 localities) more densely ribbed (R1 >13), tending to C. r. parvula. Sample from Wiesbaden (Mosbach IV) transitional to C. r. parvula (R1 18.0; clausilium plate partly like in r. parvula).

C. (C.) rugosa parvula (A. Férussac 1807)

Replacing C. r. antiquitatis in late early Middle Pleistocene (Nordsieck 1990: 156, foot-note 23)..

C. (Neostyriaca) corynodes ornatula Andreae 1884

C. schlickumi and C. c. austroloessica (both Klemm 1969) are forms of C. c. ornatula (Nordsieck 2007: 174). Frank (2006: 376-378) used both names for C. c. ornatula from Austria, the name austroloessica (contrary to Krolopp 1994: 8) also for C. corynodes from Late Pleistocene loesses.
C. c. ornatula from Early Pleistocene (from Sd F to DA 4B, 10 localities, except 2 localities from Deckschotter) and Middle Pleistocene (among others from Hundsheim and Wiesbaden-Mosbach, 6 localities) more or less densely ribbed (R1 6.0-8.6).
Form from Deckschotter (Uhlenberg) larger, more coarsely ribbed (R1 5.2).
In late early Middle Pleistocene (3 localities) more densely ribbed (R1 9.3-9.5).

C. (N.) dehmi (H. Nordsieck 2007)

Deckschotter (2 localities).
Smaller than C. c. ornatula of Deckschotter localities, more densely ribbed (R1 9.0); subcolumellar lamella receding (ending near to lunellar).

Fusulus interruptus (C. Pfeiffer 1828)

F. interruptus from Middle Pleistocene (3 localities) like extant species.

Strigillaria cana (Held 1836)

Probably present in DA 4 deposits.

III. Faunal and species changes

Faunal changes:

In the Pliocene layers only extinct species (except of Macrogastra densestriata) are present. The most radical faunal change is that at the Plio-/Pleistocene boundary (~ 2.6 Ma ago).
In the early Early Pleistocene layers nearly only extant  species have been found. Exceptions are Macrogastra sessenheimensis (recorded from Pliocene to early Early Pleistocene, latest known occurrence Kr 6) and Clausilia dehmi (only from early Early Pleistocene, Deckschotter sites).
In older layers of Early Pleistocene only species of Cochlodina and Clausiliini occur. The forms of these species are in part evaluated as subspecies of the extant species.
In contrast to the older layers, in younger layers of Early Pleistocene also species of „Serrulininae“ (Serrulella, Serruluna) have been traced (from Kr 9 to DA 4B). If this is confirmed, “Serrulininae” disappeared from central Europe at the beginning of Pleistocene (~ 2.6 Ma ago) and reappeared in the Early Pleistocene (about 1.5 Ma ago), to disappear again in the later Early Pleistocene (last known occurrence in DA 4B, about 1.2 Ma ago).
The forms of  the clausiliid species of late Early Pleistocene are in part evaluated as subspecies of the extant species. In the faunas of the eastern localities species with Carpathian range appeared, among others also the first species of Baleini in DA sites (Strigillaria cana).
Since Middle Pleistocene the forms of most species cannot be distinguished from the extant subspecies of the same region. Exceptions are Clausilia rugosa antiquitatis and C.  corynodes ornatula.
C. rugosa antiquitatis became extinct or was replaced by C. r. parvula before late Middle Pleistocene.
C. corynodes ornatula has also been found in late Middle Pleistocene and Late Pleistocene layers, at least in those from Lower Austria (Frank in Döppes & Rabeder 1997, as Neostyriaca corynodes austroloessica) and S. W. Germany (Upper Rhine valley; own investigations). The hypothesis that C. c. ornatula has been replaced in late Middle Pleistocene by C. corynodes with smooth shells like the extant subspecies (Nordsieck 1990: 156, foot-note 24) has thus not been confirmed.

Chronosubspecies:

Fossil subspecies preceding the extant ones of the same region (= chronosubspecies), which were unknown until now, have been found within the following species:
Macrogastra ventricosa, M. plicatula, Clausilia dubia, C. pumila, C. cruciata.
C. rugosa antiquitatis and C. corynodes ornatula are chronosubspecies within the species concerned which were already known.

Plesiomorphic characters within species:

Characters, which in extant subspecies have been regarded as plesiomorphic (Macrogastra, Nordsieck 2006, Clausilia, Nordsieck 1990, 2002), have been traced in the following chronosubspecies:
Macrogastra ventricosa cf. major (in part anterior lower palatal plica present), M. plicatula cf. aprutica (anterior lower palatal plica present), Clausilia cruciata cf. bonellii and C.  rugosa antiquitatis (clausilium plate with upbent outer corner).
C. dubia vindobonensis and C. p. cf. pumila from Early Pleistocene have the plesiomorphic characters folds of inferior lamella distinct, clausilium plate with upbent outer corner.
This is regarded as evidence of a correct character analysis in the cited papers.
C. corynodes ornatula has the plesiomorphic character shell ribbed.
The chronosubspecies of several Macrogastra and Clausilia species with plesiomorphic characters are similar to extant ones, which have the same characters and occur in S., E. or S. E. Europe, as is shown by the following list:
M. ventricosa: v. major Rossmässler (S. E. Alps to Crna Gora and Serbia);
M. plicatula: p. aprutica H. Nordsieck (S. Italy), also similar to p. licana (A. J. Wagner 1912) (S. E. Alps to Croatia);
C. dubia: d. vindobonensis Schmidt 1856 (E. and S. E. Europe);
C. cruciata: c. bonellii (S. Italy, especially to undescribed form from Abruzzo);
C. pumila: p. pumila (E. and S. E. Europe).
C. rugosa: antiquitatis H. Nordsieck 1990 similar to C. r. pinii Westerlund 1878 (central Italy);
The chronosubspecies (except C. r. antiquitatis) are classified here with the similar extant subspecies. It is clear that this classification is unsatisfactory, because with regard to their high age they were certainly genetically different. Therefore, the qualifying abbreviation cf. is added. The shell similarities, on which the classification is based, do not allow an alternative, especially no description as new subspecies, because these could only be distinguished from the similar extant subspecies by their age.

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