{"id":98,"date":"2021-05-21T19:58:26","date_gmt":"2021-05-21T19:58:26","guid":{"rendered":"https:\/\/hnords.de\/wordpress\/?page_id=98"},"modified":"2022-07-26T12:05:19","modified_gmt":"2022-07-26T10:05:19","slug":"fossil-clausiliidae","status":"publish","type":"page","link":"https:\/\/hnords.de\/wordpress\/fossil-clausiliidae\/","title":{"rendered":"Fossil Clausiliidae"},"content":{"rendered":"

Fossil Clausiliidae from Europe are known since the Upper Cretaceous and traced through the Tertiary and Quaternary.
\nIn 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.
\nRich representative faunas of fossil Clausiliidae allow conclusions to climate and vegetation; changes of clausiliid faunas are the result of climatic changes.
\nAs 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.
\nFossil 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.
\nNevertheless, there are several extinct clausiliid groups the relationships of which are unclear, especially if the material is scarce and essential characters cannot be examined.
\nIn the following two reports on my current work on fossil Clausiliidae are given:<\/p>\n

List of fossil Clausiliidae from western Palaearctic;<\/p>\n

Pliocene and Pleistocene Clausiliidae from central Europe north of the Alps.<\/p>\n

\"\"<\/a><\/p>\n

\"\"<\/a><\/p>\n

I. List of fossil Clausiliidae from western Palaearctic<\/b><\/p>\n

Hartmut Nordsieck (III.2021)<\/p>\n

See European Door Snails (Clausiliidae), I, chapter 10.<\/p>\n

II. Pliocene and Pleistocene Clausiliidae from central Europe north of the Alps<\/strong><\/p>\n

Hartmut Nordsieck (VII.2021, supplemented XI.2021)<\/p>\n

I. Faunas from Pliocene to early Middle Pleistocene<\/p>\n

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).
\nThe 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.
\nSpecies, which were not traced by myself, are listed in brackets.<\/p>\n

Pliocene:<\/p>\n

A, Neudegg: Serrulella austriaca<\/em>, Nordsieckia succedens<\/em>, Ruthenica filograna<\/em>, R<\/em>. cf. filograna<\/em>, Macrogastra densestriata<\/em>, Clausilia<\/em> stranzendorfensis<\/em>, C<\/em>. strauchiana geisserti<\/em>, Parafusulus neudeggensis<\/em>.<\/p>\n

A, Stranzendorf A: Clausilia stranzendorfensis<\/em>, C<\/em>. s<\/em>. geisserti<\/em>.<\/p>\n

A, Stranzendorf C: Serruluna pretiosa<\/em>, Serrulella<\/em>? sp. (apical fragments), Clausilia<\/em> stranzendorfensis<\/em>, C<\/em>. s<\/em>. geisserti<\/em>.<\/p>\n

Early Pleistocene:<\/p>\n

A, Stranzendorf D: Macrogastra sessenheimensis<\/em>? (apical fragments), Clausilia<\/em> dubia<\/em>, C<\/em>. pumila<\/em>.<\/p>\n

A, Stranzendorf F: Cochlodina laminata<\/em>, Ruthenica<\/em> filograna<\/em>, Macrogastra<\/em> plicatula<\/em>, M<\/em>. sessenheimensis<\/em>, C<\/em>. cruciata<\/em>, Clausilia<\/em> pumila<\/em>, C<\/em>. corynodes ornatula<\/em>.<\/p>\n

A, Stranzendorf G: C<\/em>. cruciata<\/em>, Clausilia<\/em> pumila<\/em>.<\/p>\n

A, Stranzendorf K: Macrogastra<\/em> densestriata<\/em>, M<\/em>. plicatula<\/em>, C<\/em>. cruciata<\/em>, Clausilia<\/em> pumila<\/em>.<\/p>\n

A, Stranzendorf K\/L: Macrogastra<\/em> ventricosa<\/em>? (apical fragments), M<\/em>. plicatula<\/em>, C<\/em>. cruciata<\/em>, Clausilia<\/em> pumila<\/em>, C<\/em>. c<\/em>. ornatula<\/em>.<\/p>\n

A, Stranzendorf L: Macrogastra<\/em> ventricosa<\/em>, Clausilia<\/em> cruciata<\/em>, C<\/em>. c<\/em>. ornatula<\/em><\/p>\n

D, Fischach: Macrogastra<\/em> ventricosa<\/em>, M<\/em>. sessenheimensis<\/em>, Clausilia<\/em> dubia<\/em>, C<\/em>. pumila<\/em>, C<\/em>. c<\/em>. ornatula<\/em>.<\/p>\n

D, Markt Buch 1-7: Macrogastra<\/em> ventricosa<\/em>, Clausilia<\/em> dubia<\/em>, C<\/em>. cruciata<\/em>, C<\/em>. pumila<\/em>, C<\/em>. c<\/em>. ornatula<\/em>, C<\/em>. dehmi<\/em>.<\/p>\n

D, Holzkirchen: Clausilia<\/em> pumila<\/em>, C<\/em>. rugosa<\/em> antiquitatis<\/em>, C<\/em>. c<\/em>. ornatula<\/em>.<\/p>\n

D, Dinkelscherben-Uhlenberg: Macrogastra<\/em> sessenheimensis<\/em>, Clausilia<\/em> dubia<\/em>, C<\/em>. cruciata<\/em>, C<\/em>. pumila<\/em>, C<\/em>. c<\/em>. ornatula<\/em>, C<\/em>. dehmi<\/em>.<\/p>\n

A, Krems-Schie\u00dfst\u00e4tte 11: Macrogastra<\/em> sessenheimensis<\/em>.<\/p>\n

A, Krems-Schie\u00dfst\u00e4tte 9: Serruluna aptyx<\/em>, Serrulella<\/em>? sp. (apical fragment), Ruthenica<\/em> filograna<\/em>, (Macrogastra<\/em> densestriata<\/em>), (M<\/em>. sessenheimensis<\/em>), Clausilia<\/em> dubia<\/em>, C<\/em>. pumila<\/em>, C<\/em>. c<\/em>. ornatula<\/em>.<\/p>\n

PL, Kielniki 3A: Serruluna<\/em> biptyx<\/em>, Cochlodina<\/em> cerata<\/em>, C<\/em>. orthostoma<\/em>, Ruthenica<\/em> filograna<\/em>, Macrogastra<\/em> borealis<\/em>, M<\/em>. plicatula<\/em>, Clausilia<\/em> dubia<\/em>, C<\/em>. pumila<\/em>.<\/p>\n

A, Radlbrunn: Clausilia<\/em> dubia<\/em>, C<\/em>. r<\/em>. antiquitatis<\/em>.<\/p>\n

A, Deutsch-Altenburg 30A: Macrogastra<\/em> ventricosa<\/em>, M<\/em>. plicatula<\/em>, Clausilia<\/em> dubia<\/em>, C<\/em>. cruciata<\/em>, C<\/em>. pumila<\/em>, C<\/em>. c<\/em>. ornatula<\/em>.<\/p>\n

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

Middle Pleistocene:<\/p>\n

D, Wei\u00dfenburg 7: Cochlodina<\/em> laminata<\/em>, Macrogastra<\/em> ventricosa<\/em>, M<\/em>. densestriata<\/em>, M<\/em>. plicatula<\/em>, Clausilia<\/em> dubia<\/em>, C<\/em>. cruciata<\/em>, C<\/em>. pumila<\/em>, Fusulus<\/em> interruptus<\/em>.<\/p>\n

PL, Kozi Grzbiet: Cochlodina<\/em> laminata<\/em>, C<\/em>. cerata<\/em>, C<\/em>. orthostoma<\/em>, Ruthenica<\/em> filograna<\/em>, Macrogastra<\/em> densestriata<\/em>, M<\/em>. tumida<\/em>, M<\/em>. plicatula<\/em>, C<\/em>. cruciata<\/em>, Clausilia<\/em> pumila<\/em>, Strigillaria<\/em> cana<\/em>.<\/p>\n

A, Hundsheim: Cochlodina<\/em> laminata<\/em>, Ruthenica<\/em> filograna<\/em>, Macrogastra<\/em> ventricosa<\/em>, M<\/em>. tumida<\/em>, M<\/em>. plicatula<\/em>, Clausilia<\/em> dubia<\/em>, C<\/em>. pumila<\/em>, C<\/em>. c<\/em>. ornatula<\/em>.<\/p>\n

D, Wiesbaden-Mosbach III: Ruthenica<\/em> filograna<\/em>, Macrogastra<\/em> ventricosa<\/em>, M<\/em>. plicatula<\/em>, Clausilia<\/em> dubia<\/em>, C<\/em>. cruciata<\/em>, C<\/em>. pumila<\/em>, C<\/em>. r<\/em>. antiquitatis<\/em>, C<\/em>. c<\/em>. ornatula<\/em>.<\/p>\n

F, Achenheim: Clausilia<\/em> dubia<\/em>, C<\/em>. pumila<\/em>, C<\/em>. r<\/em>. antiquitatis<\/em>, C<\/em>. c<\/em>. ornatula<\/em>.<\/p>\n

D, Wiesbaden-Mosbach IV: Clausilia<\/em> pumila<\/em>, C<\/em>. r<\/em>. parvula<\/em>, C<\/em> c<\/em>. ornatula<\/em>.<\/p>\n

Notes:<\/p>\n

Neudegg (Frank & Rabeder 1996, D\u00f6ppes & Rabeder 1997: 106-110):
\nLike the fauna of micromammals (D\u00f6ppes & Rabeder: 108)\u00a0 the clausiliid fauna is clearly of Pliocene age (Nordsieck 2000: 2, foot note 2; 2007: 138-139), because all species except of Macrogastra<\/em> densestriata<\/em> are extinct. An age higher than 2.6 Ma is also proved by the occurrence of a Triptychia<\/em> species (T<\/em>. neudeggensis<\/em> Schnabel).
\nThe species given by Frank (in D\u00f6ppes & Rabeder: 106) as Macrogastra<\/em> sp. is Parafusulus neudeggensis<\/em> (Nordsieck 2007: 173); the species determined as Serrulina<\/em> serrulata<\/em> is Serrulella austriaca<\/em> (Nordsieck: 160). As concerns Ruthenica<\/em> species, a complete specimen of R<\/em>. filograna<\/em> and an apertural fragment somewhat differing from that species (R<\/em>. cf. filograna<\/em>) are present.<\/p>\n

Stranzendorf (D\u00f6ppes & Rabeder 1997: 130-139):
\nAs 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 D\u00f6ppes & Rabeder: 132, 136) concerning the continued occurrence of these Pliocene Clausilia<\/em> species (C.<\/em> strauchiana<\/em>, C<\/em>. stranzendorfensis<\/em>) in Early Pleistocene layers of Stranzendorf, which later on (2006: 366) was repeated, is based on confusion with similar Clausilia<\/em> species (C<\/em>. pumila<\/em>, C<\/em>. cruciata<\/em>).
\nAs 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<\/em> species in Sd layers by Frank (: 132) may refer to M<\/em>. sessenheimensis<\/em> and (or) M<\/em>. plicatula<\/em>. In contrast to her information (: 132) I could not find M<\/em>. badia <\/em>and M. tumida<\/em> in any Sd fauna, M<\/em>. densestriata<\/em> only as one fragment in Sd K. My determination of Clausilia<\/em> r<\/em>. antiquitatis<\/em>? from Sd F (Nordsieck 2007: 139) was wrong, caused by confusion with C<\/em>. cruciata<\/em>.<\/p>\n

Dinkelscherben-Uhlenberg and other sites of the Upper Swabian Deckschotter (Dehm 1979, Nordsieck 1990: 153, R\u00e4hle 1995):
\nWithin 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 (R\u00e4hle: 114). The species formerly treated as subspecies of Macrogastra<\/em> plicatula<\/em> (R\u00e4hle: 111) is M<\/em>. sessenheimensis<\/em>, 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 R\u00e4hle (: 111) Clausilia<\/em> r<\/em>. antiquitatis<\/em> was found by me only in the material from Holzkirchen.<\/p>\n

Krems-Schie\u00dfst\u00e4tte (D\u00f6ppes & Rabeder 1997: 28-34):
\nFrom the numerous layers of Krems only small samples from Kr 7, 7\/2, 9 and 11 were available (the material examined by Lo\u017eek 1976, 1978 could not be revised).
\nFrom Kr 9 Lo\u017eek (1978: 30) mentioned an \u201cIphigena<\/em> sp. (kleine Form)\u201d, which is probably Macrogastra sessenheimensis<\/em>. From Kr 12 he gave the same species (\u201ckleine Iphigena<\/em>\u201d) and M<\/em>. densestriata<\/em>.
\nThe record of Macrogastra<\/em> plicatula<\/em> in Kr 9 by Frank (in D\u00f6ppes & Rabeder: 30) could also refer to M<\/em>. sessenheimensis<\/em>. The other Macrogastra<\/em> species given by her for that layer (determined as M<\/em>. lineolata<\/em> and cf. latestriata<\/em>) might be M<\/em>. densestriata<\/em>.
\nThe species mentioned by Lo\u017eek (: 28-29) as Serrulina<\/em> aff. serrulata<\/em> from Kr 5 (and Kr 6, with question mark) belongs with high probability to the genus Serrulella<\/em>. 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.<\/p>\n

Kielniki (Stworzewicz 1981):
\nIn 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).<\/p>\n

Radlbrunn (D\u00f6ppes & Rabeder 1997: 114-116):
\nAs to the age of the clausiliid fauna see Nordsieck 1990 (: 153). This fauna is remarkably poor; it includes only Clausilia<\/em> dubia<\/em> and a great amount of C<\/em>. rugosa antiquitatis<\/em>.<\/p>\n

Deutsch-Altenburg (D\u00f6ppes & Rabeder 1997: 238-270):
\nThe 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<\/em> species of these layers given by Frank (in D\u00f6ppes & Rabeder: 243, 262, 265) as M<\/em>. densestriata<\/em> and M<\/em>. badia<\/em> are probably M<\/em>. plicatula<\/em>; only in DA 4B one fragment of a new subspecies of M<\/em>. densestriata<\/em> could be found. The Baleini species given by her (: 243, Strigillaria cana<\/em>, Laciniaria plicata<\/em>, Alinda biplicata<\/em>) could not be traced in the extensive material available for me; only the presence of S<\/em>. cana<\/em> is probable.<\/p>\n

Middle Pleistocene faunas:
\nThe early Middle Pleistocene faunas, which were examined more thoroughly, Wei\u00dfenburg 7 (Dehm 1971), Kozi Grzbiet (Stworzewicz 1981), Hundsheim (Frank in D\u00f6ppes & Rabeder 1997: 271) and Wiesbaden-Mosbach III and IV (Gruner & Gruner 2009, 2011), were taken for comparison with the older faunas.
\nAs concerns the age of Wei\u00dfenburg 7, the information \u201eAltpleistoz\u00e4n\u201c 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).
\nThe clausiliids from Wei\u00dfenburg 7 have already been listed by Dehm (: 81-82). The material, which was determined by him as Clausilia<\/em> cruciata<\/em> n. subsp.?, contained two species, C<\/em>. cruciata<\/em> and C<\/em>. pumila<\/em> (see part III). C<\/em>. r<\/em>. parvula<\/em> does not occur in Wei\u00dfenburg 7 (Dehm w. c.).
\nThe determination of the clausiliids from Kozi Grzbiet by Stworzewicz has already been revised (Nordsieck 2007: 140).
\nMacrogastra<\/em> cf. densestriata<\/em> given by Frank (: 271) for Hundsheim may be M<\/em>. plicatula<\/em>. Clausilia<\/em> schlickumi<\/em> described from Hundsheim is a form of C<\/em>. corynodes<\/em> ornatula<\/em> (Nordsieck 2007:174).
\nFrom 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<\/em> bidentata<\/em> is C<\/em>. r<\/em>. antiquitatis<\/em>.<\/p>\n

II. Records of Pliocene and Pleistocene Clausiliidae from central Europe<\/p>\n

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).
\nThe FADs of the species concerned are given as follows:<\/p>\n

Serrulina serrulata<\/em>: Records in layers of central Europe probably based on confusion with Serrulella<\/em> species (see part I).<\/p>\n

Cochlodina laminata<\/em>: 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).<\/p>\n

C<\/em>. orthostoma<\/em>: Kielniki 3A (Early Pleistocene). The record as cf. in the older layer Sd C is doubtful (Frank in D\u00f6ppes & Rabeder 1997: 132).<\/p>\n

C<\/em>. cerata<\/em>: Kielniki 3A (Early Pleistocene).<\/p>\n

Ruthenica filograna<\/em>: Neudegg (Piacenzian).<\/p>\n

Macrogastra ventricosa<\/em>: Sd K\/L (Early Pleistocene)?, Sd L.<\/p>\n

M<\/em>. tumida<\/em>: Kozi Grzbiet, Hundsheim (Middle Pleistocene). The record as cf. in Sd F is doubtful (Frank in D\u00f6ppes & Rabeder: 132). Record as cf. in Kr 8\/2 (Lo\u017eek 1976) not checked.<\/p>\n

M<\/em>. densestriata<\/em>: Frechen-Fortuna (Piacenzian).<\/p>\n

M<\/em>. badia<\/em>: No fossil record. The record in Deutsch-Altenburg = DA 4B is probably wrong (see part I).<\/p>\n

M<\/em>. borealis<\/em>: Kielniki 3A (Early Pleistocene). The record as M<\/em>. cf. latestriaia<\/em> in Krems = Kr 9 (Frank in D\u00f6ppes & Rabeder: 30) is probably wrong (see part I).<\/p>\n

M<\/em>. attenuata<\/em>: (Holocene). The record as M<\/em>. lineolata<\/em> in Kr 9 (Frank in D\u00f6ppes & Rabeder: 30) is probably wrong (see part I).<\/p>\n

M<\/em>. sessenheimensis:<\/em> Sessenheim (Piacenzian). For the last appearance date = LAD see part III.<\/p>\n

M<\/em>. plicatula<\/em>: Sd F (Early Pleistocene).<\/p>\n

Clausilia baudoni<\/em>: Moncucco Torinese (Messinian). Record in Amb\u00e9rieu-le-Bugey (Tortonian) not checked.<\/p>\n

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

C<\/em>. strauchiana<\/em>: Eichkogel (Tortonian). Record in Doln\u00e9 Ore\u0161any (Tortonian) not checked. For the LAD see part I. 2.<\/p>\n

C<\/em>. pumila<\/em>: Sd D (Early Pleistocene).<\/p>\n

C<\/em>. stranzendorfensis<\/em>: Neudegg, Sd A (Piacenzian). For the LAD see part I. 2.<\/p>\n

C<\/em>. cruciata<\/em>: Sd F (Early Pleistocene).<\/p>\n

C<\/em>. dubia<\/em>: Sd D (Early Pleistocene).<\/p>\n

C<\/em>. rugosa antiquitatis<\/em>: 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<\/em>. r<\/em>. parvula<\/em> see part III.<\/p>\n

C<\/em>. corynodes ornatula<\/em>: 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<\/em>. corynodes <\/em>subspecies.<\/p>\n

Fusulus interruptus<\/em>: Wei\u00dfenburg 7 (Middle Pleistocene).<\/p>\n

Strigillaria cana<\/em>: DA 4 (Early Pleistocene) (see part I).<\/p>\n

Laciniaria plicata<\/em>: Gombasek (Middle Pleistocene). The record in DA 4B is probably wrong (see part I).<\/p>\n

Alinda biplicata<\/em>: (Late Pleistocene). The record in DA 4B is probably wrong (see part I).<\/p>\n

III. Pleistocene (and related Pliocene) taxa<\/p>\n

Extant species means subspecies of the respective species occurring in the region of the locality.
\nThe given values of rib density (R1<\/sub>) are means of samples.<\/p>\n

Cochlodina<\/em> (C<\/em>.) laminata<\/em> (Montagu 1803)<\/p>\n

C<\/em>. laminata<\/em>\u00a0from Early Pleistocene (Sd F, Kr 7\/2, DA 4B) like that from Middle Pleistocene (Wei\u00dfenburg 7, Hundsheim) and extant species; lowermost palatal plica distinct. = C<\/em>. l<\/em>. laminata<\/em>.<\/p>\n

Ruthenica<\/em> filograna<\/em> (Rossm\u00e4ssler 1836)<\/p>\n

R<\/em>. filograna<\/em> 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<\/em>. filograna<\/em>.<\/p>\n

Macrogastra<\/em> (M<\/em>.) ventricosa<\/em> (Draparnaud 1801)<\/p>\n

M. ventricosa <\/em>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<\/em>. v<\/em>. cf. major<\/em> Rossm\u00e4ssler.
\nM<\/em>. ventricosa<\/em> from Middle Pleistocene (3 localities) like extant species; in Wiesbaden-Mosbach lower fold of inferior lamella weakened. = M<\/em>. v<\/em>. ventricosa<\/em>.<\/p>\n

M<\/em>. (n. subgen.) borealis<\/em> (O. Boettger 1878)<\/p>\n

M<\/em>. borealis<\/em> from Kielniki like extant species, with relatively dense ribbing (R1 <\/sub>6.3). = M<\/em>. b.<\/em> bielzi<\/em> H. Nordsieck.<\/p>\n

M<\/em>. (n. subgen.) densestriata<\/em> (Rossm\u00e4ssler 1836)<\/p>\n

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

M<\/em>. (Pyrostoma<\/em>) sessenheimensis<\/em> (H. Nordsieck 1974)<\/p>\n

Smaller than M<\/em>. plicatula<\/em>; 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.
\nForms from Sessenheim, Pliocene (type form), and Sd F more densely ribbed than \u00a0form from Deckschotter localities (Fischach, Uhlenberg).
\nExtinct at the end of\u00a0 early Early Pleistocene (latest occurrence in Krems 6, Lo\u017eek 1978: 29, if \u201cIphigena<\/em> sp.\u201d = M<\/em>. sessenheimensis<\/em>).<\/p>\n

M<\/em>. (P.<\/em>) plicatula<\/em> (Draparnaud 1801)<\/p>\n

M<\/em>. plicatula<\/em> from Sd F coarsely ribbed (only one incomplete fragment available).
\nM<\/em>. plicatula<\/em> from later Early Pleistocene (from Sd K to DA 4B, 6 localities) smaller than that from Wei\u00dfenburg 7 and extant species; more densely ribbed than most extant subspecies (R1 <\/sub>7.5-9.0); with two folds of inferior lamella; in part anterior lower palatal plica present. = M<\/em>. plicatula<\/em> cf. aprutica<\/em> H. Nordsieck.
\nM. plicatula<\/em> from Middle Pleistocene (3 localities) like extant species (R1<\/sub> 5.5-6.5). = M<\/em>. p<\/em>. plicatula<\/em>.<\/p>\n

Clausilia<\/em> (C<\/em>.) dubia<\/em> Draparnaud 1805<\/p>\n

C<\/em>. dubia<\/em> from Sd D ventricose, with short apical part; inferior lamella like C<\/em>. d<\/em>. vindobonensis <\/em>A. Schmidt. = C<\/em>. d<\/em>. cf. vindobonensis<\/em>.
\nC. dubia<\/em> from Sd I, L\/M, Kr (2 localities) and DA (4 localities) inferior lamella and clausilium plate, if present, like C<\/em>. d<\/em>. vindobonensis<\/em>. = C.<\/em> d<\/em>. vindobonensis<\/em>.
\nC<\/em>. dubia<\/em> 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<\/em>. d<\/em>. vindobonensis<\/em> and C<\/em>. d<\/em>. dubia<\/em>.
\nC<\/em>. dubia<\/em> from Middle Pleistocene of Germany (2 localities) inferior lamella and clausilium plate like C<\/em>. d<\/em>. dubia<\/em>. = C<\/em>. d<\/em>. dubia<\/em>.
\nC<\/em>. dubia<\/em> from Hundsheim = C<\/em>. d<\/em>. vindobonensis<\/em>.
\nAll Pleistocene forms are normally ribbed (R1<\/sub> between 7.5 and 10).<\/p>\n

\u00a0<\/em>C<\/em>. (C<\/em>.) stranzendorfensis<\/em> H. Nordsieck 1990<\/p>\n

Assumed to be closely related to C<\/em>. cruciata<\/em>. Differs from that species mainly by the development of the inferior lamella (upper fold in part reduced).
\nForms from 3 Pliocene localities (Neudegg, Sd A, Sd C) somewhat different (Nordsieck: 162-163), if subspecies?
\nOn average larger than C<\/em>. strauchiana<\/em> geisserti<\/em>; 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<\/em>. s<\/em>. geisserti<\/em> (R1<\/sub> 8.0-8.2).<\/p>\n

C<\/em>. (C<\/em>.) cruciata<\/em> (Studer 1820)<\/p>\n

C<\/em>. cruciata<\/em> from Early Pleistocene (from Sd F to DA 4B, 8 localities) relatively densely ribbed (R1 <\/sub>6.9-7.9); outer corner of clausilium plate upbent. = C<\/em>. cruciata<\/em> cf. bonellii<\/em> (form from Abruzzo).
\nC<\/em>. cruciata<\/em> from Wei\u00dfenburg 7 like those from Early Pleistocene; = C<\/em>. c<\/em>. cf. bonellii<\/em>. Compared with C<\/em>. pumila<\/em> from the same site smaller, more densely ribbed (R1 <\/sub>6.9, pumila<\/em> 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.
\nC<\/em>. cruciata<\/em> from Wiesbaden-Mosbach like extant species; outer corner of clausilium plate right-angled, less upbent. = C<\/em>. c<\/em>. cruciata<\/em>.
\nRemark: C<\/em>. cruciata<\/em> and C<\/em>. pumila<\/em> 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<\/em>. cruciata<\/em> more s-like ascending, with more distinct folds in front; subcolumellar lamella more bent below. Because C<\/em>. cruciata<\/em> and C<\/em>. pumila<\/em> from the Middle Pleistocene deposit Wei\u00dfenburg 7 are clearly separable, it is assumed that also in Early Pleistocene deposits both species are present.<\/p>\n

\u00a0<\/em>C<\/em>. (C<\/em>.) strauchiana<\/em> geisserti<\/em> H. Nordsieck 1976<\/p>\n

Assumed to be closely related to C<\/em>. pumila<\/em>.
\nSmaller than C<\/em>. pumila<\/em>; more densely ribbed (R1<\/sub> 8.0-8.7); subcolumellar lamella strongly bent; anterior lower palatal plica and palatal hump strongly developed; outer corner of clausilium plate upbent, pointed.
\nSpecimens from 5 Pliocene localities not much different.<\/p>\n

C<\/em>. (C<\/em>.) pumila<\/em> C. Pfeiffer 1828<\/p>\n

C<\/em>. pumila<\/em> from Early Pleistocene (from Sd D to DA 4B, 11 localities) on average smaller, more densely ribbed than Middle Pleistocene and extant species (R1 <\/sub>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<\/em>. p<\/em>. cf. pumila<\/em>.
\nC<\/em>. pumila<\/em> from Middle Pleistocene (8 localities) with superior lamella continuous with spiral lamella, more coarsely ribbed (R1 <\/sub>5.8-6.8):form from Wei\u00dfenburg 7 like extant species, outer corner of clausilium plate upbent;
\nform from Wiesbaden-Mosbach and other western localities (5 localities) corresponding, outer corner of clausilium plate less upbent;
\nboth = C<\/em>. p<\/em>. pumila<\/em>.
\nC<\/em>. p<\/em>. sejuncta<\/em> M\u00f6rch 1864 could not be traced in the Middle Pleistocene deposits.<\/p>\n

C<\/em>. (C<\/em>.) rugosa<\/em> antiquitatis<\/em> H. Nordsieck 1990<\/p>\n

Examined from deposits of Early Pleistocene (Deckschotter, Radlbrunn, DA, 4 localities) and Middle Pleistocene (only western localities, 6).
\nIn comparison with other subspecies of C<\/em>. rugosa<\/em> densely ribbed (type form R1<\/sub> 10.5-11.7); with both folds of inferior lamella or only the upper one; outer corner of clausilium plate \u00b1 upbent and pointed.
\nType form from Radlbrunn with more weakened folds of inferior lamella.
\nSamples from late early Middle Pleistocene (3 localities) more densely ribbed (R1<\/sub> >13), tending to C<\/em>. r<\/em>. parvula<\/em>. Sample from Wiesbaden (Mosbach IV) transitional to C<\/em>. r<\/em>. parvula<\/em> (R1<\/sub> 18.0; clausilium plate partly like in r<\/em>. parvula<\/em>).<\/p>\n

C<\/em>. (C<\/em>.) rugosa<\/em> parvula<\/em> (A. F\u00e9russac 1807)<\/p>\n

Replacing C<\/em>. r<\/em>. antiquitatis<\/em> in late early Middle Pleistocene (Nordsieck 1990: 156, foot-note 23)..<\/p>\n

C<\/em>. (Neostyriaca<\/em>) corynodes<\/em> ornatula<\/em> Andreae 1884<\/p>\n

C<\/em>. schlickumi<\/em> and C<\/em>. c<\/em>. austroloessica<\/em> (both Klemm 1969) are forms of C. c<\/em>. ornatula<\/em> (Nordsieck 2007: 174). Frank (2006: 376-378) used both names for C.<\/em> c<\/em>. ornatula<\/em> from Austria, the name austroloessica<\/em> (contrary to Krolopp 1994: 8) also for C. corynodes<\/em> from Late Pleistocene loesses.
\nC<\/em>. c<\/em>. ornatula<\/em> 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 <\/sub>6.0-8.6).
\nForm from Deckschotter (Uhlenberg) larger, more coarsely ribbed (R1<\/sub> 5.2).
\nIn late early Middle Pleistocene (3 localities) more densely ribbed (R1<\/sub> 9.3-9.5).<\/p>\n

C<\/em>. (N.<\/em>) dehmi<\/em> (H. Nordsieck 2007)<\/p>\n

Deckschotter (2 localities).
\nSmaller than C<\/em>. c<\/em>. ornatula<\/em> of Deckschotter localities, more densely ribbed (R1<\/sub> 9.0); subcolumellar lamella receding (ending near to lunellar).<\/p>\n

Fusulus<\/em> interruptus<\/em> (C. Pfeiffer 1828)<\/p>\n

F<\/em>. interruptus<\/em> from Middle Pleistocene (3 localities) like extant species.<\/p>\n

Strigillaria <\/em>cana<\/em> (Held 1836)<\/p>\n

Probably present in DA 4 deposits.<\/p>\n

IV. Faunal and species changes<\/p>\n

Faunal changes:<\/p>\n

In the Pliocene layers only extinct species (except of Macrogastra<\/em> densestriata<\/em>) are present. The most radical faunal change is that at the Plio-\/Pleistocene boundary (~ 2.6 Ma ago).
\nIn the early Early Pleistocene layers nearly only extant \u00a0species have been found. Exceptions are Macrogastra<\/em> sessenheimensis<\/em> (recorded from Pliocene to early Early Pleistocene, latest known occurrence Kr 6) and Clausilia<\/em> dehmi<\/em> (only from early Early Pleistocene, Deckschotter sites).
\nIn older layers of Early Pleistocene only species of Cochlodina<\/em> and Clausiliini occur. The forms of these species are in part evaluated as subspecies of the extant species.
\nIn contrast to the older layers, in younger layers of Early Pleistocene also species of \u201eSerrulininae\u201c (Serrulella<\/em>, Serruluna<\/em>) have been traced (from Kr 9 to DA 4B). If this is confirmed, \u201cSerrulininae\u201d 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).
\nThe forms of\u00a0 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<\/em>).
\nSince Middle Pleistocene the forms of most species cannot be distinguished from the extant subspecies of the same region. Exceptions are Clausilia<\/em> rugosa<\/em> antiquitatis<\/em> and C<\/em>. \u00a0corynodes<\/em> ornatula<\/em>.
\nC<\/em>. rugosa<\/em> antiquitatis<\/em> became extinct or was replaced by C<\/em>. r<\/em>. parvula<\/em> before late Middle Pleistocene.
\nC<\/em>. corynodes<\/em> ornatula<\/em> has also been found in late Middle Pleistocene and Late Pleistocene layers, at least in those from Lower Austria (Frank in D\u00f6ppes & Rabeder 1997, as Neostyriaca corynodes austroloessica<\/em>) and S. W. Germany (Upper Rhine valley; own investigations). The hypothesis that C<\/em>. c<\/em>. ornatula<\/em> has been replaced in late Middle Pleistocene by C<\/em>. corynodes<\/em> with smooth shells like the extant subspecies (Nordsieck 1990: 156, foot-note 24) has thus not been confirmed.<\/p>\n

Chronosubspecies:<\/p>\n

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

Plesiomorphic characters within species:<\/p>\n

Characters, which in extant subspecies have been regarded as plesiomorphic (Macrogastra<\/em>, Nordsieck 2006, Clausilia<\/em>, Nordsieck 1990, 2002), have been traced in the following chronosubspecies:
\nMacrogastra<\/em> ventricosa<\/em> cf. major<\/em> (in part anterior lower palatal plica present), M<\/em>. plicatula<\/em> cf. aprutica<\/em> (anterior lower palatal plica present), Clausilia<\/em> cruciata<\/em> cf. bonellii<\/em> and C<\/em>. \u00a0rugosa<\/em> antiquitatis <\/em>(clausilium plate with upbent outer corner).
\nC<\/em>. dubia<\/em> vindobonensis<\/em> and C<\/em>. p<\/em>. cf. pumila<\/em> from Early Pleistocene have the plesiomorphic characters folds of inferior lamella distinct, clausilium plate with upbent outer corner.
\nThis is regarded as evidence of a correct character analysis in the cited papers.
\nC<\/em>. corynodes ornatula<\/em> has the plesiomorphic character shell ribbed.
\nThe chronosubspecies of several Macrogastra<\/em> and Clausilia<\/em> 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:
\nM<\/em>. ventricosa<\/em>: v<\/em>. major<\/em> Rossm\u00e4ssler (S. E. Alps to Crna Gora and Serbia);
\nM<\/em>. plicatula<\/em>: p<\/em>. aprutica<\/em> H. Nordsieck (S. Italy), also similar to p<\/em>. licana<\/em> (A. J. Wagner 1912) (S. E. Alps to Croatia);
\nC<\/em>. dubia<\/em>: d<\/em>. vindobonensis<\/em> Schmidt 1856 (E. and S. E. Europe);
\nC<\/em>. cruciata<\/em>: c<\/em>. bonellii<\/em> (S. Italy, especially to undescribed form from Abruzzo);
\nC<\/em>. pumila<\/em>: p<\/em>. pumila<\/em> (E. and S. E. Europe).
\nC<\/em>. rugosa<\/em>: antiquitatis<\/em> H. Nordsieck 1990 similar to C<\/em>. r<\/em>. pinii<\/em> Westerlund 1878 (central Italy);
\nThe chronosubspecies (except C<\/em>. r<\/em>. antiquitatis<\/em>) 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.<\/p>\n

References<\/strong><\/p>\n

Dehm, R. (1971): Eine altpleistoz\u00e4ne Spaltenf\u00fcllung von Wei\u00dfenburg in Bayern und ihre Molluskenfauna. \u2013 Mitteilungen der Bayerischen Staatssammlung f\u00fcr Pal\u00e4ontologie und historische Geologie, 11<\/strong>: 77-85, pl. 8.<\/p>\n

Dehm, R. (1979): Artenliste der altpleistoz\u00e4nen Molluskenfauna vom Uhlenberg bei Dinkelscherben. \u2013 Geologica Bavarica, 80<\/strong>: 123-125.<\/p>\n

D\u00f6ppes, D. & Rabeder, G. (eds.) (1997): Plioz\u00e4ne und pleistoz\u00e4ne Faunen \u00d6sterreichs. Ein Katalog der wichtigsten Fossilfundstellen und ihrer Faunen. \u2013\u00a0 Mitteilungen der Kommission f\u00fcr Quart\u00e4rforschung der \u00f6sterreichischen Akademie der Wissenschaften, 10<\/strong>: 411 pp.<\/p>\n

Frank, C. (2006): Plio-pleistoz\u00e4ne und holoz\u00e4ne Mollusken \u00d6sterreichs. \u2013\u00a0 Mitteilungen der Pr\u00e4historischen Kommission, 62<\/strong>, part 1: I-XV, 1-395, part 2: I-XV, 396-860, 62 pls.<\/p>\n

Frank, C. & Rabeder, G. (1996): Kleins\u00e4uger und Landschnecken aus dem Mittel-Plioz\u00e4n von Neudegg (Nieder\u00f6sterreich). \u2013 Beitr\u00e4ge zur Pal\u00e4ontologie, 21<\/strong>: 41-49.<\/p>\n

Gruner, M. & Gruner, H. (2009): Neue Untersuchungen zur mittelpleistoz\u00e4nen Gastropoden-Fauna der Mosbach-Sande von Wiesbaden-Am\u00f6neburg. \u2013\u00a0 Mainzer Naturwissenschaftliches Archiv, 47<\/strong>: 347-373.<\/p>\n

Gruner, M. & Gruner, H. (2011): Nachweis einer Glazialfauna in den Mainablagerungen des mittleren Mittelpleistoz\u00e4n (Mosbach IV\/T2) im Dyckerhoff-Steinbruch Wiesbaden-Am\u00f6neburg. \u2013\u00a0 Mainzer Naturwissenschaftliches Archiv, 48<\/strong>: 129-141, 1 pl.<\/p>\n

Harzhauser, M. & Neubauer, T. A. (2021): A review of the land snail faunas of the European Cenozoic \u2013 composition, diversity and turnovers. \u2013 Earth Science Review, 217<\/strong>: 31 pp. https:\/\/doi.org\/10.1016\/j.earscirev.2021.103610.<\/p>\n

Klemm, W. (1969): Das Subgenus Neostyriaca<\/em> A. J. Wagner 1920, besonders der Rassenkreis Clausilia<\/em> (Neostyriaca<\/em>) corynodes<\/em> Held 1836. \u2013 Archiv f\u00fcr Molluskenkunde, 99<\/strong> (5\/6): 285-311.<\/p>\n

Koenigswald, W. von (1971): Die altpleistoz\u00e4ne Wirbeltierfaunula aus der Spaltenf\u00fcllung Wei\u00dfenburg 7 (Bayern). \u2013\u00a0 Mitteilungen der Bayerischen Staatssammlung f\u00fcr Pal\u00e4ontologie und historische Geologie, 11<\/strong>: 117-122.<\/p>\n

Krolopp, E. (1994): A Neostyriaca<\/em> g\u00e9nusz a magyarorsz\u00e1gi pleisztoc\u00e9n k\u00e9pz\u00f6dm\u00e9nyekben. \u2013 Malakol\u00f3giai t\u00e1j\u00e9koztat\u00f3 (M\u00e1tra M\u00fazeum), 13<\/strong>: 5-8.<\/p>\n

Lozek, V. (1976): Krems-Schie\u00dfst\u00e4tte. Malakologie. \u2013\u00a0 Mitteilungen der Kommission f\u00fcr Quart\u00e4rforschung der \u00f6sterreichischen Akademie der Wissenschaften, 1<\/strong>: 84-87.<\/p>\n

Lozek, V. (1978): Krems-Schie\u00dfst\u00e4tte. Malakologie. \u2013\u00a0 Mitteilungen der Kommission f\u00fcr Quart\u00e4rforschung der \u00f6sterreichischen Akademie der Wissenschaften, Erg\u00e4nzung zu Band 1<\/strong>: 27-31.<\/p>\n

Nadachowski, A. (1990): Lower Pleistocene rodents of Poland: faunal succession and biostratigraphy. \u2013\u00a0 Quart\u00e4rpal\u00e4ontologie, 8<\/strong>: 215-223.<\/p>\n

Nordsieck, H. (1972): Fossile Clausilien, I. Clausilien aus dem Plioz\u00e4n W-Europas. \u2013 Archiv f\u00fcr Molluskenkunde, 102<\/strong> (4\/6): 165-188, pls. 9-10, 10a.<\/p>\n

Nordsieck, H. (1974): Fossile Clausilien, II. Clausilien aus dem O-Plioz\u00e4n des Elsa\u00df. \u2013 Archiv f\u00fcr Molluskenkunde, 104<\/strong> (1\/3): 29-39, pl. 1.<\/p>\n

Nordsieck, H. (1976): Fossile Clausilien, III. Clausilien aus dem O-Plioz\u00e4n des Elsa\u00df, II (mit Bemerkungen zur systematischen Stellung von Triptychia<\/em>). \u2013 Archiv f\u00fcr Molluskenkunde, 107<\/strong> (1\/3): 73-82, pls. 10, 10a.<\/p>\n

Nordsieck, H. (1981): Fossile Clausilien, V. Neue Taxa neogener europ\u00e4ischer Clausilien, II. \u2013 Archiv f\u00fcr Molluskenkunde, 111<\/strong> (1\/3): 63-95, pls. 7-9.<\/p>\n

Nordsieck, H. (1990): Revision der Gattung Clausilia<\/em> Draparnaud , besonders der Arten in SW-Europa (Das Clausilia<\/em> rugosa<\/em>-Problem) (Gastropoda: Stylommatophora: Clausiliidae). \u2013 Archiv f\u00fcr Molluskenkunde, 119<\/strong> (4\/6): 133-179, 3 pls.<\/p>\n

Nordsieck, H. (2000): Annotated check-list of the fossil (pre-Pleistocene) Clausiliidae (Gastropoda: Stylommatophora) from central and western Europe. \u2013 Mitteilungen der deutschen malakozoologischen Gesellschaft, 65<\/strong>: 1-15.<\/p>\n

Nordsieck, H. (2002): The subspecies classification of Clausilia dubia <\/em>Draparnaud (Gastropoda: Stylommatophora: Clausiliidae), a critical revision. – Mitteilungen der deutschen malakozoologischen Gesellschaft, 68<\/strong>: 37-44.<\/p>\n

Nordsieck, H. (2006): Systematics of the genera Macrogastra<\/em> Hartmann 1841 and Julica<\/em> Nordsieck 1963, with the description of new taxa (Gastropoda: Stylommatophora: Clausiliidae). – Archiv f\u00fcr Molluskenkunde, 135<\/strong> (1): 49-71, 2 pls.<\/p>\n

Nordsieck, H.\u00a0 (2007): Worldwide Door Snails (Clausiliidae), recent and fossil. \u2013 214 pp., 20 pls. Hackenheim (ConchBooks).<\/p>\n

Rabeder, G. (1981): Die Arvicoliden (Rodentia, Mammalia) aus dem Plioz\u00e4n und dem \u00e4lteren Pleistoz\u00e4n von Nieder\u00f6sterreich. \u2013 Beitr\u00e4ge zur Pal\u00e4ontologie von \u00d6sterreich, 8<\/strong>: 1-373.<\/p>\n

R\u00e4hle, W. (1995): Altpleistoz\u00e4ne Molluskenfaunen aus den Zusamplattenschottern und ihrer Flu\u00dfmergeldecke vom Uhlenberg und Lauterbrunn (Iller-Lech-Platte, Bayerisch Schwaben). \u2013 Geologica Bavarica, 99<\/strong>: 103-117.<\/p>\n

Stworzewicz, E. (1981): Early Pleistocene land snails from Kielniki and Kozi Grzbiet (Poland). \u2013 Folia Quaternaria, 54<\/strong>: 43-77, 10 pls.<\/p>\n

 <\/p>\n

 <\/p>\n

 <\/p>\n","protected":false},"excerpt":{"rendered":"

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…<\/p>\n

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