A Remarkable Non-Marine Mollusc Fauna of Early Eocene Age from A

65: 31 – 76 2019

A remarkable non-marine mollusc fauna of Early Eocene age from a fissure infill in Karsdorf quarry (Sachsen-An- halt, Germany)1 Eine bemerkenswerte nichtmarine Molluskenfauna von früheozänem Alter aus einer Spaltenfüllung im Stein- bruch Karsdorf (Sachsen-Anhalt, Deutschland) 1

Dietrich Kadolsky

66 Heathhurst Road, Sanderstead, Surrey CR2 0BA, United Kingdom; [email protected]

Revision accepted February 11, 2020. Published online at www.senckenberg.de/geologica-saxonica on April 3, 2020.

Abstract Non-marine molluscs in excellent shell preservation are described and fgured from the infll of a large fssure exposed in the cement quarry Karsdorf (Germany, state Sachsen-Anhalt, ca. 49 km WSW of Leipzig) in Lower Muschelkalk (Triassic) limestones. The fauna consists of 36 mollusc species, of which 6 are freshwater inhabitants and the remaining 30 are terrestrial. Of the latter, 15 are only known from fragments insuffcient to recognize even the superfamily, but are reported and fgured nonetheless to indicate the degree of diversity of this fauna. 13 new species are named: Acroloxus aspis, Acroloxus korys, Carychium vagum, Carychium bachi, Turricarychium muelleri, Ovicarychium hennigeri, Ovicarychium ronlederi, Zuella venusta, Acanthinula (s.l.) karsdorfensis, Albinulopsis gibba, Afrodontops euro paea, Afrodontops comes, Palaeostoa costellata, and 5 new genera are proposed (Turricarychium (type species T. muelleri), Ovicarychium (type species O. ronlederi), Zuella (type species Z. venusta), Albinulopsis (type species A. gibba), Afrodontops (type species A. europaea)). Because of the remarkable diversity of the Carychiinae with 7 species the generic classifcation of the fossils in this subfamily is reviewed and in addition the genus Carychiopsina (type species Pupa schwageri Reuss 1868) is proposed. Of note is the frst record of the hitherto oldest known members of the Charopidae or Endodontidae, Afrodontops europaea and A. comes, which is also the frst record of this Gondwanan group in Europe. The family Palaeostoidae is suggested to be removed from Clausilioidea and assigned to Orthalicoidea. These assessments add to previously identifed biogeographical relationships between the Paleogene fauna of Europe with that of South America and the Caribbean. The fssure infll is interpreted as the deposit of a pond, which was not connected to the regional surface drainage system and may have formed in a depression caused by subrosion.When the fssure formed, these pond sediment slid into it.The land snails lived in a well vegetated and humid environment in a tropical to subtropical climate. Four species are conspecifc with, or closely related to species from the late Ypresian fauna of Grauves (France, Paris Basin): Carychium vagum is conspecifc, Carychium bachi is closely related to C. sp. aff. bachi, and Albinulopsis gibba is closely related to A. bonneti (Cossmann 1907); but Palaeostoa costellata is intermediate in characters between P. exarata (Michaud 1838) from Rilly-la-Montagne (late Thanetian) and P. cf. fontenayi (Sandberger 1871) from Grauves, indicating an age range for the mollusc fauna younger than the level of Rilly-la-Montagne (late Thanetian) and older than the level of Grauves, i.e. most likely in the range early to middle Ypresian (Early Eocene).

Kurzfassung Nichtmarine Mollusken in ausgezeichneter Erhaltung werden aus der Füllung einer großen Spalte in Kalksteinen des Unteren Muschel- kalkes (Trias), die im Zementsteinbruch Karsdorf (ca. 49 km WSW Leipzig, Sachsen-Anhalt, Deutschland) aufgeschlossen ist, beschrieben und abgebildet. Die Fauna besteht aus 36 Arten, von denen 6 Süßwasserbewohner und die anderen 30 Landschnecken sind. Von letzteren sind 15 nur durch Fragmente bekannt, die nicht einmal zur Identifzierung der Oberfamilie ausreichen. Sie sind hier dennoch documentiert, um die Diversität der Fauna zu illustrieren. 13 neue Arten werden benannt (Acroloxus aspis, Acroloxus korys, Carychium vagum, Cary

1 Contribution no. 14 by the author in the series “Non-marine and marginally marine mollusc faunas in the European Tertiary”. For no. 12 see Harzhauser et al. (2016), Paläontologische Zeitschrift, DOI 10.1007/sl2542-015-0277-1. For no. 13 see Harzhauser et al. (2016), Archiv für Molluskenkunde 145(1): 23 – 58.

ISBN 978-3-91000661-4 | ISSN 1617-8467 | DOI: 10.26049/GEOLSAX65-2019-04 31 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

chium bachi, Turricarychium muelleri, Ovicarychium hennigeri, Ovicarychium ronlederi, Zuella venusta, Acanthinula (s.l.) karsdorfensis, Albinulopsis gibba, Afrodontops europaea, Afrodontops comes, Palaeostoa costellata), und 5 neue Gattungen vorgeschlagen (Turricary chium (Typusart T. muelleri), Ovicarychium ( Typusart O. ronlederi), Zuella ( Typusart Z. venusta), Albinulopsis ( Typusart A. gibba), Afro dontops ( Typusart A. europaea)). Wegen der großen Diversität der Carychiinae mit 7 Arten werden die Gattungszuordnungen der fossilen Arten in dieser Subfamilie revidiert; zusätzlich wird die Gattung Carychiopsina (Typusart Pupa schwageri Reuss 1868) vorgeschlagen. Beachtenswert ist der erste Nachweis der bis jetzt ältesten bekannten Mitglieder der Charopidae oder Endodontidae, Afrodontops europaea und A. comes, die zugleich die ersten bekannt gewordenen Vertreter dieser in den Gondwana-Kontinenten beheimateten Gruppe in Europa sind. Es wird vorgeschlagen, die Palaeostoidae von den Clausiloidea zu den Orthalicoidea zu versetzen. Diese Bewertungen ergänzen die bereits bekannten biogeographischen Beziehungen zwischen der Paläogenen Fauna Europas mit der von Südamerika und der Karibik. Dier Spaltenfüllung wird als Teichsedimentiert interpretiert. Der Teich war nicht mit dem regionalen oberfächlichen Entwässerungssystem ver- bunden und könnte sich über einer Subrosionssenke gebildet haben. Als die Spalte sich bildete, rutschten die Teichsedimente in sie hinein. Die Landschnecken lebten in einem humiden Milieu mit reicher Vegetation in einem subtropischen bis tropischen Klima. Vier Arten sind entweder konspezifsch, oder nahe verwandt mit Arten der spätypresischen Fauna von Grauves (Pariser Becken, Frankreich): Carychium vagum ist konspezifsch, Carychium bachi ist nahe verwandt mit C. sp. aff. bachi von Grauves, Albinulopsis gibba ist nahe verwandt mit A. bonneti (Cossmann 1907). Palaeostoa costellata steht in ihren Merkmalen zwischen P. exarata (Michaud 1838) von Rilly-la-Montagne (spätes Thanetium) und P. cf. fontenayi (Sandberger 1871) von Grauves, was das Alter der Karsdorfer Molluskenfauna als jünger als das Niveau von Rilly-la-Montagne und älter als das von Grauves belegt, also höchstwahrscheinlich den Bereich frühes bis mittleres Ypresium (Früheozän)

Introduction Karsdorf quarry exposes a succession of basal Low- er Muschelkalk strata: Lower Wellenkalk and Zone of Oolith Beds (“Oolithbankzone”). Locally the overlying The fossil record in areas subjected to erosion is nec- Middle Wellenkalk and Terebratula Beds are also pre- essarily non-existent unless exceptional circumstances sent. The fssure infll also contains Muschelkalk mate- provide protection from erosion and destructive diagen- rial attributable to the Upper Muschelkalk (Trochiten- esis. One such circumstance is preservation in fssures kalk, Ceratites Beds). This proves that at the time of and other rock cavities in carbonate rocks. A large fssure karst fssure formation an almost complete Muschelkalk flled with Palaeogene sediments in the limestone quarry succession was present in Karsdorf, implying erosional of Karsdorf (Germany, Sachsen-Anhalt, ca. 51°16.5’ N, removal of some 130 m of carbonatic sediment since that 11°40.3’ E) had been investigated between 2005 and time (Henniger et al. 2011: Fig. 3). This implies a total 2012 by A. Müller and his team from Leipzig University. original depth of the karst fssure of some 200 m, assum- In this quarry limestone is extracted for cement produc- ing it extends down to the Röt Formation underlying the tion currently by the company Opterra GmbH, and earlier Muschelkalk. The quarry operations exposed only some by Lafargue Zement GmbH. The fssure is exposed at the 15 m of fssure infll below present-day land surface. A upper quarry face at the SE edge of the quarry; it runs stratigraphic differentiation of the karst fssure infll was NNW-ESE. The geological situation in which this fssure not possible, as the sediment did not show lithological was formed has been described and interpreted by Hen- differences. Likewise the fossil content appears to be niger et al. (2011) and is summarized here: biostratigraphically rather homogenous (Henniger et al. The karst fssure is located in limestones of the Low- 2011, and this study). Nonetheless it should be born in er Muschelkalk (Triassic: early part of Anisian). The mind that much material was recovered from scree gen- Muschelkalk is regionally present in the Querfurt Plateau erated by quarry operations. and continues southwards into the Naumburg Syncline, a Apart from Muschelkalk clasts and a small amount large low-relief depression (Fig. 1). The present-day area of quartz sand in all grain sizes, the sediment consists of the Querfurt Plateau was in pre-Lutetian times low rel- of dark bluish-grey to occasionally yellowish-green silts ative to the Geiseltal area. Salt movement in the underly- and clays with common plant debris. A diverse assem- ing Zechstein caused uplift of the Querfurt Plateau, and blage of seeds and spores had been recovered, as well as subsidence of the Geiseltal area (Fig. 1). The Geiseltal remains of molluscs, turtles, crocodiles, amphibians and Depression became a sedimentary basin for coal-bearing a single tooth of the rodent Paramyidae sp. The arago- sediments, which had been exploited in large open-cast nitic mollusc shells are well preserved. mines. This sedimentary succession has become famous Henniger et al. (2011) supposed that the Karsdorf karst for its Middle Eocene mammal faunas; other organisms fssure infll were of similar age as the Geiseltal Forma- including non-marine molluscs were also preserved due tion, i.e. Lutetian. They base this conclusion on the mol- to the infux of carbonatic groundwaters resulting from lusc fauna. The numerous plant remains indicate merely limestone dissolution in the Querfurt Plateau. an age range from the Eocene to the Miocene, with the The updoming of the Querfurt Plateau generated oldest age indicator being Tinospora wilkinsonii Chandler extensional stress which resulted in fracturing of the hitherto known from Early Eocene strata of England and Muschelkalk limestones. The fractures enabled meteor- France. ic waters to access the evaporites of the Röt Formation Henniger et al. (2011) fgured some of the molluscs, (Upper Buntsandstein, basal Anisian). Salt and gypsum mostly in open nomenclature. The entire mollusc fauna is dissolution in the Röt, and carbonate dissolution in the here described and documented, and re-analyzed for its Muschelkalk widened the fractures. palaeoenvironmental and biostratigraphical signifcance.

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Fig. 1. Geological sketch map and cross-section of the site region. Redrawn and modifed after Henniger et al. 2011 (Fig. 1a, 1b). Use of these fgures with permission of E.Schweizerbart’sche Verlagsbuchhandlung OHG. Abb. 1. Geologische Kartenskizze und Profl der Region mit der Fundstelle. Umgezeichnet und verändert nach Henniger et al. 2011 (Abb. 1a, 1b). Verwendung der Abbildungen mit Erlaubnis der E.Schweizerbart’schen Verlagsbuchhandlung OHG.

It should be noted that the material may be unique as the The descriptive shell terminology follows Cox (1960) chance of recovery of additional material from the fssure unless otherwise indicated. Measurements (in mm) are is uncertain and possibly low. height : width : (optional) apertural height, unless stated otherwise. The dimensions of defective shells are not restored to entire shells; such measurements are marked in Material and methods italics. The mollusc fauna is primarily grouped into aquatic and terrestrial molluscs. Within these ecological groups Sediment samples from the Karsdorf karst fssure had the suprageneric classifcations and their succession fol- been collected, washed, sieved and picked by A. Müller lows Bouchet & Rocroi (2010) for the bivalve, and Bou- and his team, consisting of M. Henninger, R. M. Leder chet et al. (2017) for the gastropods, unless otherwise and F. Bach, from Leipzig University. Drawings of speci- stated. Fragments of land snails which cannot even be mens were made by this author with a Wild M5 binocu- assigned to a superfamily are treated at the end of the lar microscope with drawing tube, and photographs with taxonomy chapter. a Canon EOS 500 camera. The material is stored in the To obtain numerical frequency data, the usually frag- Geological and Paleontological Collection of Leipzig mented specimens were counted by counting the same University (GPSL) with the inventory numbers TM 0188 shell part in each species indicative of an individual, to TM 0335), and a few specimens in the collection of typically the apex or the adult aperture, whichever gave the author (acronym DK). The taxonomic assessment larger numbers. While the shell preservation of the Kars- was supported by the study of reference material from dorf fauna is better than that of the studied materials from the following institutions: the Paris Basin, shells > 2 mm are always fragmented, possibly by sediment compaction as well as sample pro- MNHNP Muséum National d’Histoire Naturelle, Département Histoire de la Terre (Paris); cessing. The bulk of the shell infll is both clear sparite and dark silt matrix, but some gypsum crystallisation is SMF Naturmuseum und Forschungsinstitut Senckenberg (Frankfurt a.M.); unfortunately in evidence, where crystal growth has frac- MUWI Museum Wiesbaden (Wiesbaden); tured whorls and pushed part of the shell wall outwards. In the author’s initial notes, all species received an in- SMNS Staatliches Museum für Naturkunde Stuttgart (Stutt- gart); formal number prefxed with KA. In case a species iden- NHMUK Natural History Museum UK, Paleontology Depart- tifcation was not possible, these terms are here main- ment (London); tained. ZNSGH Zentralmagazin Naturwissenschaftlicher Sammlun- gen (Geiseltalsammlung), Martin-Luther Universität Halle-Wittenberg, Halle (Saale).

33 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

Taxonomic treatment (s.l.) aquensis Mathéron 1843 (: 215, pl. 36, fgs. 6 – 7; Lutetian: “environs d’Aix, au quartier de Montaiguet”), whose apical angle in fg. 6 is 55°, but in fg. 7 it is 45°. Aquatic mollusks This discrepancy indicates the diffculty of obtaining ac- curate measurements from published fgures showing insuffcient detail of the apical region of the shell. Fig.3 Class Gastropoda Cuvier 1795 shows Mathéron’s fg. 7 with the outline of the Karsdorf Subclass Heterobranchia Burmeister, 1837 specimen inset at the same scale (Fig. 3). The recent type species Galba truncatula lives in Infraclass Euthyneura Spengel, 1881 very shallow waters such as ditches, slow fowing brooks Cohort Tectipleura Schrödl et al. 2011 and the shallow zone of ponds. Subcohort Panpulmonata Jörger et al., 2010 Superorder Hygrophila Férussac, 1822 Physidae Fitzinger 1833 Lymnaeoidea Rafnesque 1815 Lymnaeidae Rafnesque 1815 Physidae sp. KA30 Lymnaeinae Rafnesque 1815 Pl. 1A, Fig. 7; Pl. 1B, Fig. 7

Material: Karsdorf quarry, fssure infll: GPSL TM 0265/1, GPSL TM 0337/1. 2 specimens from 2 out of 18 samples. Galba Schrank 1803 Dimensions: 1.30 : 0.96 (Pl. 1, Fig. 7) Type species (fxed in Opinion 1896 (ICZN 1998)): Buccinum trun catulum Müller 1774. Recent, Europe. Remarks: In western Europe the Physidae were much more diverse in the Palaeocene – Eocene period than in Galba sp. KA1 the Oligocene to present period. Unfortunately the dimen- Pl. 1A, Figs. 1 – 3; Pl. 1B, Figs. 1 – 2 sions and shape of the apical whorls of the named Paleo- v 2011 Galba cf. aquensis ssp. – Henniger et al.: 326, Pl. 3, Fig. 11 gene species is unsuffciently known. (Karsdorf quarry) [non Lymnaea aquensis Mathéron 1843].

Material: Karsdorf quarry, fssure infll: GPSL TM 0247/1, GPSL TM 0248/1, GPSL TM 0249/>368, GPSL TM 0250/1, GPSL Planorbidae Rafnesque 1815 TM 0251/5, GPSL TM 0252/1, GPSL TM 0253/19, GPSL TM 0254/377, GPSL TM 0255/16. In total ca. 790 specimens from 11 Planorbinae Rafnesque 1815 out of 18 samples. Pompholycodeini Lindholm 1927 Diagnosis: A species whose frst 4.4 whorls are convex and form a slender spire with an spire angle of 43°. Biomphalaria Preston 1910 Galba sp. KA1 resembles the recent Galba truncatula Type species: B. smithi Preston 1910 (original designation). East (O.F.Müller 1774) in the relatively small apical angle and Africa, recent. the strong convexity of the whorls. However, in Galba truncatula the whorls grow faster in height and width, Anatomical and molecular genetic data have shown (e.g. resulting in a slightly larger apical angle.An as yet un- DeJong et al. 2001, 2003; Morgan et al. 2002; Albrecht described Galba species from the late Rupelian Jacobs- et al. 2007) that the genus-group names Planorbina Hal- berg Formation (formerly „Süßwasserschichten“) of the deman 1843 (type species Planorbis olivaceus Wagner Mainz Basin differs by a much wider apical angle. & Spix in Spix & Wagner 1827 (subsequent designation by Dall 1905) from Brazil [= Planorbis glabratus Dimensions: 5.00 : 2.49 : 2.50; 4.4 whorls (Pl.1, Fig. 1), Say 1818]) and Australorbis Pilsbry 1934 (type species length of columella fragment: 6.5 (Pl.1 Fig. 2) Planorbis guadeloupensis G.B.Sowerby I 1822 (original designation) from Caribbean Islands [= P. glabratus Say Additional observations: The largest undeformed apex 1818]) are subjective synonyms of Biomphalaria. Con- fragment is fgured Pl. 1, Fig. 1, plus a columellar frag- sequently, the genus is today present in South America as ment showing the columellar torsion (Fig. 2). An identi- well as in Africa. The ICZN ruled in Opinion 735 (ICZN fcation on the species level is not attempted as the de- 1965) i.a., that the name Biomphalaria is to have priority tails of the early whorls of the numerous nominal species over Planorbina if considered congeneric. of Paleocene and Eocene age in Europe are not known. The genus Biomphalaria cannot consistently be iden- According to the original fgures, which, however, show tifed by the shell characters of the extant species. Wenz insuffcient detail, all appear to have consistently wider (1923) was the frst to place all Paleogene European large apical angles. This is defnitely the case for Lymnaea planorbids which did not closely resemble Planorbarius

34 GEOLOGICA SAXONICA — 65: 2019

in this genus, then named Planorbina. This classifcation Deshayes in 1863. Available for this study was Sandberg- is plausible, as many extant species, in particular B. gla er’s material (1870: pl. 9, fg. 12; 1872: 191) from Mont brata (Say 1818), differ from Planorbarius and resemble Bernon, preserved in the MUWI. many Paleogene European species by their whorls more slowly increasing in size, and a relatively lesser shell and aperture height than has Planorbarius. Shells attributed Acroloxidae Thiele 1931 to Biomphalaria have been reported from South America as old as Late Cretaceous (Salvador 2018) and in et al. Genus Acroloxus Beck 1838 North America at least as old as Paleocene (B. planocon vexa (Meek & Hayden 1861), fde Wenz 1923). Type species: Patella lacustris Linnaeus 1758 (subsequent designation by Herrmannsen 1846). Europe, recent.

Remarks: The European Tertiary freshwater limpets Biomphalaria sp. KA2 group as two independently evolved clades: Acroloxi- Pl. 1A, Figs. 4 – 6; Pl. 1B, Figs. 4 – 6 dae with the genus Acroloxus and Planorbidae: Ancyli- v 2011 Australorbis cf. pseudammonius – Henniger et al.: 326, nae with the genera Ancylus Müller 1773 and Ferrissia Pl. 3, Fig.7 [tantum, non Pl. 3, F 14 (= Afrodontops europaea Walker 1903. They are easily distinguishable by the po- n. sp.); non Helicites pseudoammonius Schlotheim 1820]. sition and orientation of their apices, which are pointing backwards. In Acroloxus the apex is positioned and is Material: Karsdorf quarry, fssure infll: GPSL TM 0256/1, GPSL pointing to the left, and in the Ancylinae is positioned TM 0257/1, GPSL TM 0258/1, GPSL TM 0259/>11, GPSL TM 0260/1, GPSL TM 0261/1, GPSL TM 0262/2, GPSL TM 0263/52, centrally or slightly to the right. The distinction beween GPSL TM 0264/2. In total ca. 70 specimens from 12 out of 18 sam- the two taxa was, at least on the genus or subgenus level, ples, common in only 1 sample. Usually fragmented, only one in- already made by most 19th century authors, but genus- tact juvenile specimen and several protoconchs. level nomenclature was confused by the long-held assumption that the type species of Ancylus was Patella Dimensions: 0.81 : 1.35 : 0.80 (Pl. 1, Fig. 4) lacustris Linnaeus. 0.57 : 0.49 : 0.45 (Pl. 1, Fig. 5) The distinction between the named Paleocene and width 0.68 (Pl. 1, Fig. 6) Eocene Acroloxus species is currently tentative, as very few specimens were available for study. Therefore, in- Observations: The fragments of mature shells indicate traspecifc variability and potential growth allometry that the species is a large planorbid of Planorbarius or could not be assessed. Nonetheless two new species are Biomphalaria habitus. The only intact specimen is ju- proposed below, with reasons for this procedure given in venile, having 2.4 whorls at a breadth of 1.35 mm and the Remarks. a height/width ratio of 0.60. Due to gentle deformation the last half whorl is laterally compressed, resulting in the narrow appearance of the aperture. The protoconch Acroloxus aspis n.sp. is shaped like a miniature Physa with fattened sides. It Pl. 2A, Figs. 1 – 2; Pl. 2B, Figs. 1 – 2 is much smaller than the similarly shaped protoconch of v 2011 Ancylus cf. dutemplei – Henniger et al.: 326, Pl. 3, Fig. 9 – Oligocene Planorbarius sp., see Kadolsky 2015a (: 56, 10 [non Ancylus dutemplei Deshayes 1863] fgs. 5 – 6) with rounded whorls. The size of this specimen differs markedly from that of Holotype: GPS Leipzig TM 0266 (Pl. 2, Fig. 1); length 1.66 mm; Biomphalaria pseudoammonia (Schlotheim 1820), which width 1.22 mm; height 0.42 mm; Fig. 1. at 2.4 whorls is ca. 1.7 mm wide. This was established Paratypes: GPSL TM 0267/1, GPSL TM 0268/14DK/1. In total 16 in several specimens with preserved shells from the type specimens from 5 out of 18 samples. locality Bouxwiller (SMF 273139, coll. Schlickum). This juvenile from Karsdorf is more similar to B. sub Locus typicus and stratum typicum: Germany, Sach ovata (Deshayes 1825), but B. subovata has a larger nu- sen-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle cleus (0.23 mm compared to 0.17 mm in the Karsdorf (Saale), ca. 51°16′26″ N, 11°41′09″ E. Fissure infll: cal specimen). The size and growth rates of the early wells careous and carbonaceous silts and clays mixed with are very similar, but the last preserved whorl in the Kars- Triassic limestone debris; age: Early Eocene. dorf species increases more in width. Due to slight shell deformations, the juvenile stage of the Karsdorf speci- Derivatio nominis: Ασπίς (Greek): a shield. This word men and the observed differences the Karsdorf species is a noun in apposition and hence immutable. cannot be positively identifed with B. subovata. The type locality of B. subovata is Mont-Bernon Diagnosis: A species of Acroloxus characterized by the near Epernay, being the sole original locality. Deshayes absence of radial striae, an ovoid outline (ratio length : (1825: 741) gave only Epernay; his later (1863: 741) cita- width 1.36), a rounded and not protruding apex situat- tion of “Mont-Bernon près Epernay” is a precision of the ed only slightly to the left-hand side of the shell, which same locality. Wenz (1923: 1517 – 1518) cited incorrectly is only moderately elevated (ratio height:length 0.25 – Jonchery as type locality, which was frst mentioned by 0.27).

35 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

Dimensions: 0.43 : 1.65 (Pl. 2, Fig. 1, holotype) Acroloxus korys n. sp. 0.52 : 2.11 (Pl. 2, Fig. 2) Pl. 2A, Figs. 3 – 4; Pl. 2B, Figs. 3 – 4 Additional observations: The shells are usually pre- Holotype: GPS Leipzig TM 0269, ex sample KD2382. Pl. 2, Fig. 3. served as fragments, as they are rather fragile. The great- Paratypes: GPSL TM 0270/1, GPSL TM 0271/19DK/1. In total 22 est length of a damaged shell was measured as 2.61 mm, specimens from 1 out of 18 samples. without missing parts restored. Locus typicus and stratum typicum: Germany, Sachsen- Remarks: The named Paleocene and Eocene Acroloxus Anhalt, Karsdorf cement quarry ca. 33 km SW Halle species can be grouped according to their traits as fol- (Saale), ca. 51°16′26″ N, 11°41′09″ E. Infll of a large fs- lows: sure (Henniger et al. 2011): calcareous and carbonaceous silts and clays mixed with Triassic limestone debris; age: Shell outline ovoid (ratio length : width = 1.29 – 1.40), Early Eocene. radial striae present: A. lemoinei (Cossmann 1889), A. be rellensis (Laubrière & Carez 1881), A. matheroni (Boissy Derivatio nominis: Kóruς (Greek): a helmet. The word 1848). is a noun in apposition and hence immutable. Shell outline ovoid (ratio l:w = 1.36 – 1.43), radial striae absent: A. aspis n.sp., A. korys n.sp. Diagnosis: A species of Acroloxus with a relatively high (ratio height : length 0.35 – 0.36) and elongate-ovoid shell Shell outline elongate-ovoid (ratio l : w = 1.50 – 1.54), (ratio length : width approximately 1.43 – 1.57), a round- radial striae present: A. arenarius (Cossmann 1889), A. ed and strongly eccemtric but not overhanging apex, and dutemplei (Deshayes 1863), Acroloxus sp. Krumbiegel lack of radial sculpture. 1962. A. dutemplei (Deshayes 1863) from the Calcaire grossier Dimensions: 0.70 : 1.90 (Pl. 2, Fig. 3, holotype) of Boursault of Lutetian age differs from A. aspis n.sp. by 0.52 : 1.47 (Pl. 2, Fig. 4) its more elongate shell (ratio length : width 1.52 according to the original fgure), well developed radial striae Remarks (see also under A. aspis): A. korys n.sp. differs and a protruding and overhanging apex. in two uncorrelated characters from the synpatric A. as pis: the strongly elevated shell and the much more excen- Acroloxus sp. from the Geiseltal area (Lutetian, lig- tric apex. Further, the shell is slightly more elongate and nite pit Cecilie), which has also been identifed with the long sides run subparallel to each other, resulting in a A. dutemplei by Wenz (1932) and Krumbiegel (1962), more oblong outline. The ratio shell height : shell length differs from A. aspis n.sp. by its elongate shape (ratio is higher than in any named Eocene Acroloxus species. length : width = 1.54), but is similar by the not protrud- Intermediates with the much fatter A. aspis n.sp. have ing and not overhanging apex. Its shell shape is similar to not beem observed. that of A. dutemplei. As radial striae were mentioned by Wenz (1932), but are not recognizable on Krumbiegel’s photographs of two specimens, it may be assumed that they are very subtly developed. Acroloxus arenarius Bivalvia Linnaeus 1758 (Cossmann 1889) from Chenay (Thanétien, Dép. Marne) has a less elongate shape (ratio l : w = 1.50), but differs Subclass Autobranchia Grobben 1894 by the strongly protruding and overhanging apex and the Superorder Heteroconchia Gray 1854 presence of radial sculpture over most of the shell. Thus, A. aspis n. sp., A. korys n. sp. and A. sp. from Clade Heterodonta Neumayr 1884 Geiseltal differ from the remaining known Paleocene Order Venerida Gray 1854 and Eocene species by their rounded and not overhanging apex and their weak or absent radial sculpture. These Sphaerioidea Deshayes 1855 (1820) traits are not due to them being immature: radial sculp- Sphaeriidae Deshayes 1855 (1820) ture which, when present, is present from a very early Sphaeriinae Deshayes 1855 (1820) stage of shell growth onwards. The type specimen of Ac roloxus berellensis (Laubrière & Carez 1881), which is only a little larger than the Karsdorf material (length 2.0 Sphaeriidae sp. indet. KA15 mm according to Laubrière & Carez 1881), has strong radial striae beginning close to the apex, which is slightly Pl. 2A, Figs. 5 – 8; Pl. 2B, Figs. 5 – 8 overhanging. Material: Karsdorf quarry, fssure infll: GPSL TM 0272/1, GPSL TM 0273/1, GPSL TM 0274/1, GPSL TM 0275/1, GPSL TM 0276/1, GPSL TM 0277/1. Fragments counted in total as 6 specimens from 5 out of 18 samples.

36 GEOLOGICA SAXONICA — 65: 2019

The outer surface has widely spaced growth riblets and Megalomastomatidae Blanford 1864 fner growth striae. Of the hinge of the right valve the ?Megalomastomatinae Blanford 1864 posterior laterals P1 and P3 and the larger part of the ligament area are preserved. The laterals are broadly tri- angular in profl, of similar size, parallel to each other Megalomastomatinae? sp. KA14 and reach their highest points simultaneously. The outer lateral carries several transverse wrinkles. The ligament Pl. 2A, Fig. 9; Pl. 2B, Fig. 9 area is elongate. Of the hinge of the left valve the poste- Material: Karsdorf quarry, fssure infll: GPSL TM 0278/1. 1 speci men from 1 out of 18 samples. rior part is partly preseved, showing part of the ligament area and the lateral P2. A large “pit” below P2 is formed Dimensions: Aperture height: 3.88 (Pl. 2, Fig. 9). The by the hinge plate, possibly im response to the large P3 aperture is almost perfectly circular and only slightly at- of the right valve. tached to the preceding whorl. A high terminal varix is offset from the peristome which is expanded and slightly Dimensions: 3.84 : 4.52 (Pl. 2, Fig. 5) refected. The shell has a narrow umbilicus. The orna- greatest dimension: 1.62 (Pl. 2, Fig. 6) mentation consists of growth striae only; surface corro- greatest dimension: 2.77 (Pl. 2, Fig. 7) sion may have oblitterated fner details. greatest dimension: 2.42 (Pl. 2, Fig. 8) The circular aperture is characteristic for the Megalo- In the genus Sphaerium Scopoli 1777 the laterals don’t mastomatinae, but no comparable species is known in the have visible wrinkles, and in the right valve the dorsal Tertiary of Europe. The recent species live on Caribbean posterior lateral (P3) is markedly smaller than the ventral Islands and in Central America. The Chattian Ventriculus lateral (P1). In the left valve a posterior “pit” is not nor- pupa (M. Braun 1838) (Family Pupinidae) has also a cir- mally developed, as the hinge plate is rather narrow. A cular aperture, which is larger (4.4 mm) and has a very very small “pit” was observed in one Pisidium amnicum weakly developed terminal varix close to the peristome (Müller 1774) population, and larger “pits” in Polyme which is not expanded. soda subarata (Schlotheim 1820), family Cyrenidae.

Cochlostomatinae Kobelt 1902 Terrestrial gastropods

Cochlostoma Jan 1830 Class Gastropoda Cuvier 1795 Type species: Cyclostoma maculatum Draparnaud 1805 (SD Wenz 1923). Recent, France. Caenogastropoda Cox 1960 Grade Architaenioglossa Haller 1892 Cochlostoma sp. KA16 Cyclophoroidea Gray 1847 Pl. 3A, Figs. 1 – 2; Pl. 3B, Figs. 1 – 2

Cyclophoroidea sp. 31 Material: Karsdorf quarry, fssure infll: GPSL TM 0279/1, GPSL TM 0280/1, GPSL TM 0281/>3, GPSL TM 0282/6. In total 12 Pl. 3A, Fig. 3 specimens (apical fragments) from 4 out of 18 samples.

Material: 1 protoconch fragment, sample KD 803; lost ex GPSL. The protoconch comprises 1.2 whorls and is 0.72 mm wide; the nucleus is 0.20 mm wide. Ornamentation is not Greatest dimension: 1.09 (Pl. 3, Fig. 3). perceptible. The early teleoconch whorls (1.4 preserved) have widely spaced, regular collabral ribbing. The relatively strong spiral striae exclude other land snail groups except the Pomatiidae, which are, however, not Dimensions: 1.48 : 1.42 (Pl. 3, Fig. 1) known from the Early Eocene in Europe. width: 0.90 (Pl. 3, Fig. 2) The apex is much broader than in the Rupelian species Cochlostoma sp. Ha 7 (Kadolsky 2015), while the protoconch of the Chattian C. labellum (Thomä 1845) has very similar dimensions.

37 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

Subclass Heterobranchia Burmeister, 1837 We restrict Carychiopsis to fusiform shells with strong Infraclass Euthyneura Spengel, 1881 ribbing and a parietal fold. Other than the type species, only C. michaudi (Boissy 1848) is included, albeit ten- Cohort Tectipleura Schrödl et al. 2011 tatively. Subcohort Panpulmonata Jörger et al., 2010 Clade Eupulmonata J. Morton 1955 Carychiopsina n.gen. Order Ellobiida Van Mol 1967 Type species: Pupa schwageri Reuss 1868 (Tuchořice, Early Mio cene). Ellobioidea L.Pfeiffer 1854 (1822) Shell ventricose, ovate-conical, with ribs and fne spiral Ellobiidae L.Pfeiffer 1854 (1822) sculpture; 2 palatal folds, 1 parietal fold. Also included Carychiinae Jeffreys 1830 are C. costulata (Sandberger 1862) (syn. Carychium eurabdium Bourguignat 1864, Carychiopsis schwageri hochheimensis Wenz 1923), Carychiopsina prisyazhnyu In the family Ellobiidae a conspicuous lamella sitting at ki (Stworzewicz 1999) and possibly C.? surai (Stworze- the junction of columellar and parietal margin is in the wicz 1999) which lacks the parietal knob. This taxon literature (e.g. Zilch 1959 (: 64), Strauch 1977, Stworze- comprises Late Oligocene and Early Miocene species wicz 1999) often identifed as parietal. However, this fold habitually attributed to Carychiopsis. In addition to their is attached to the columella, as many published fgures, shell characters they are are separated from the Eocene as well as our Figs. (Pl. 3 Figs. 5 and 11, Pl. 4 Figs. 2, 3, Carychiopsis species by a time span of up to 18 ma 5, 9, and 11) clearly demonstrate, and is therefore here (43 – 24 ma. BP, Bartonian 1 to early Chattian). termed supracolumellar, even though in apertural view its position often seems to be shifted towards the parietal wall. In describing the apertural dentition raised struc- Turricarychium n.gen. tures extending far into the shell’s interior are termed la- Type and only known species: Turricarychium muelleri n.sp.from mellae, while short protuberances are named tubercles or Karsdorf. knobs if they are roundish in outline, and folds when they are elongate. Multiple palatal lamellae are lirae. Diagnosis: Shell turriform, spira initially conical to In Europe a remarkable hiatus exists in the temporal slightly coeloconoid, last 1½ whorls not increasing in distribution of diversity and character range between the width resulting in an overall fusiform shape of the spire; Paleocene to Middle Eocene species (mainly known from surface with fne riblets; aperture with 2 palatal folds, a the Paris Basin and now also from Karsdorf) and the mid- columellar and a strong suparacolumellar lamella; the Oligocene to recent species. Eocene species are much latter is internally extended and undulating inside the last more diverse in their habitus: they include much larger whorl; no parietal fold. and more solid shells with stronger dentition and often more lamellae and/or folds than the post-Eocene lineages, Ovicarychium n.gen. as well as a greater range of shell shapes. Of the Eocene species only few can be attributed to the genus Carychi Type species: Ovicarychium ronlederi n.sp. um (see C. vagum and C. bachi below). The other Eocene Shell ovate-elongate to ovate-conical, with weak ribs or lineages have become extinct before the beginning of the Oligocene. The Oligocene is depleted in carychiids: Car just growth lines. Apertural folds variable: 0 – 1 parietal, ychium moenanum Wenz 1917 from the latest Rupelian 1 – 3 palatal and 0 – 1 basal folds. Columellar and supra- to earliest Chattian; Carychiopsina costulata (Sandberger columellar lamellae always present. 1862) from the latest Chattian and Carychiella eumicra Included: O. bigeminatum (Deshayes 1863), O. car (Bourguignat 1857) from the latest Chattian or Early Mi- bonarium (Wenz 1932), O. hennigeri n.sp., O. interfer ocene (Aquitanian), all from the Mainz Basin. Carychium ens (Deshayes 1863), O. polysarcum (Cossmann 1914), extends to the present, while Carychiella and Carychi O. quadridens (Andreae 1884), O. remiense (Boissy 1848), opsina didn’t survive the Early Miocene. O. ronlederi n.sp., O. sparnacense (Deshayes 1863). The Eocene species have customarily been grouped in the genus Carychiopsis Sandberger 1871. As this as- Remarks: This group contains species with rather vary- semblage is morphologically rather heterogeneous, a ing apertural dentitions. These appear to be of limited more differentiated generic classifcation of the Carychii- diagnostic value for supraspecifc classifcations, as pala- dae is here proposed. This classifcation is outlined in the following diagnoses and key. 1 A single species is reported from Bartonian strata: Carychium nincki Cossmann in Cossmann & Pissarro 1913 from Marines Carychiopsis Sandberger 1871 in the Paris Basin. Its barrel-shaped shell and the presence of an Type species (by monotypy; see Sandberger 1871: 155): Pupa dhor- angular lamella indicate that it may be a Pupilloidean, possibly ni Deshayes 1863. a gastrocoptid.

38 GEOLOGICA SAXONICA — 65: 2019

tal, parietal and basal folds occur in varying numbers and Early Miocene strata of the Mainz Basin) has a shell sim- combinations. The character in common is the relatively ilar to that of Carychium, but is very small and with re- broad habitus of the shell, unlike Carychium s.str., Tur duced lamellae and tubercles: palatal tubercle vestigial or ricarychium, Carychiopsis and Carychiopsina. The spe- absent, supracolumellar lamella internally not expanded cies O. bigeminatum, O. carbonarium and O. ronlederi and not undulating. This appears to be a sideline of Cary have a terminal varix. chium with reduced shell size and apertural armatures. Its extent is uncertain, as reductions in size and internal folds could have occurred repeatedly. Zuella n.gen. The type species of the nominal genus Saraphia Ris- Type and sole species: Zuella venusta n.sp. from Karsdorf. so 1826 (S. uniplicata Risso 1826, designated by Gray 1847: 176) is not an ellobiid (shell smooth, shiny; 6 Derivatio nominis: Zuella is the diminutive form of Zua whorls; one palatal fold; height 3 mm; in water holes); Turton 1831, a synonym of Cochlicopa Férussac 1821, Strauch (1977) and subsequent authors had overlooked alluding to the similar habitus and much smaller size; this type designation and and based their assignment of gender feminine. Saraphia as a subgenus of Carychium on Pilsbry’s designation of Saraphia tridentata Risso 1826 (1948: 1051) Diagnosis: A small shell of elongate pupoid shape (re- as type species which is an invalid designation because it sembling Cochlicopa), with a smooth and glossy surface, postdates that of Gray (1847). a slightly expanded peristome and a rounded supracolu- The troglobiont genera Zospeum Bourguignat 1856 mellar lamella. and Koreozospeum Jochum & Prozorova in Jochum 2015 are not considered further here. They are only Remarks: This taxon is tentatively placed in Carychiinae et al. known in the recent fauna. Their shells differ by being on account of the presence of a supracolumellar lamella. broader and more conical. Shell size and shape are consistent with the range known from carychiids, but Zuella is dissimilar to all other known carychiids with respect to the glossy surface and Key to the carychiine genera the absence of both the columellar lamella and any palatal fold. No other superfamilies of pulmonate land snails 1. Shell with only a supracolumellar lamella, glossy .. seem to be appropriate for this taxon. For example, Coch- ...... Zuella licopidae do have a similar shell shape with a glossy sur- – Shell with a supracolumellar and a columellar lamel- face, but are much larger and lack a supracolumellar fold. la ...... 2 2. Shell around 1 mm high, no or vestigial palatal knob, Carychium O. F. Müller 1773 supracolumellar lamella internally not extended and Type species: Carychium minimum O. F. Müller 1774. Recent, Eu not undulating ...... Carychium (Carychiella) rope. – Shell with at least one well developed palatal knob, larger than 1 mm ...... 3 Shell ovate-elongate, with riblets or growth lines. The 3. Shell relatively broad, ovate-elongate to ovate-coni- supracolumellar lamella and a palatal fold are always de- cal ...... 4 veloped; the columellar lamella may be present or absent. – Shell more slender, ± elongate ...... 5 Species attributable to Carychium are known from the 4. Shell ovate-elongate to ovate-conical, with weak ribs Early Eocene to the present. We place two Eocene spe- or just growth lines. 0 – 1 parietal, 1 – 3 palatal and cies from the Paris Basin and Karsdorf, C. bachi n.sp. and 0 – 1 basal knobs, some species with terminal varix C. vagum n.sp. in Carychium; C. hypermeces Cossmann (Eocene) ...... Ovicarychium 1889 and C. cylindroides Staadt in Cossmann & Pissarro – Shell ovate-conical, costulate, with 1 parietal and 1913 probably belong here, too, as do forms identifed by 1 – 2 palatal knobs (Late Oligocene and Early Mio- Krumbiegel (1962) as Carychium minimum [non Müller cene) ...... Carychiopsina 1774] and C. cf. nouleti [non Bourguignat 1857] from the 5. Shell moderately elongate, with one palatal knob Lutetian Geiseltal Formation. In the Oligocene only C. only ...... Carychium moenanum Wenz 1917 from the Late Rupelian Sulzheim – Shell fusiform or turriform ...... 6 Formation of the Mainz Basin is known. The frequency 6. Shell fusiform, strongly ribbed, with 1 parietal and and diversity of Carychium in European land snail assem- one palatal knob ...... Carychiopsis blages increased since Early Miocene times. – Shell turriform, weakly ribbed, with 2 palatal knobs A subgenus Tainocarychium Schütt 1988 was pro- only ...... Turricarychium posed for Carychium euboicum Schütt 1988 from Late Miocene strata of Euboea (= Evvia, Greece), because its shell has an elongate palatal fold in the frst half of the Key to the species from Karsdorf last whorl. Carychiella Strauch 1977 (type species: Carychium 1. Shell with terminal varix, collabral ribbing reduced, eumicrum Bourguignat 1857 from latest Oligocene or surface glossy; 6 folds or lamellae: 2 palatales, 1 pari-

39 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

etalis, 1 basalis, 1 supracolumellaris, 1 columellaris: from Karsdorf. Many specimens from Grauves agree al- ...... Ovicarychium ronlederi n.sp. most completely with the holotype from Karsdorf (Pl. 3A – Shell without terminal varix, 1 – 5 lamellae or folds .... 2 Figs. 5, 7 and 8). 2. Shell ovate, glossy, only a supracolumellar lamella ...... Zuella venusta n.gen. n.sp. Dimensions: 1.43 : 0.54 : 0.53 (Pl. 3, Fig. 4, holotype) – Shell shape variable, at least with supracolumellar la- 1.21 : 0.51 : 0.51 (Pl. 3, Fig. 5; mella and one palatal fold, not glossy ...... 3 from Grauves) 3. Only supracolumellar lamella and one palatal fold 1.32 : 0.51 : 0.42 (Pl. 3, Fig. 7; visible in aperture ...... 4 from Grauves) – 4 – 5 lamellae or folds visible in aperture ...... 5 1.40 : 0.55 : 0.48 (Pl. 3, Fig. 8; 4. Shell cylindrical ...... Carychium bachi n.sp. from Grauves) – Shell elongate, rounded conical...... 1.28 : 0.50 : 0.43 (from Grauves) ...... Carychium vagum n.sp. 5. Shell turriform-conical, last quarter whorl trumpet- Remarks: 1. The lot from Grauves (MNHNP, originally like expanded, with columellar and supracolumellar inventarized as J11799; from the Cossmann collection) lamella, and 2 palatal folds...... was identifed by Cossmann as Carychium hypermeces ...... Turricarychium muelleri n.sp. Cossmann 1889. This lot of 38 specimens contains 2 – Shell elongate-ovoid, broad, last quarter whorl hardly non-carychiine shells, 22 Carychium sp. aff. bachi n.sp. expanded, with strong columellar and supracolumel- and 14 C. vagum n.sp. None of them agrees with the lar lamella ...... 6 original fgure (see pl. 3A Fig. 6). See further comments 6. Shell 1.44 mm high with 4.75 whorls; aperture with 3 under C. bachi n.sp. palatal and 1 basal tubercle...... Ovicarychium hennigeri n.sp. – Shell (estimated) 1.0 mm high with 4.5 whorls (in- Carychium bachi n.sp. complete preservation) ..... Ovicarychium? sp. KA34 Pl. 3A, Figs. 10 – 11; Pl. 3B, Figs. 10 – 11

v 2011 Carychiopsis sp. 2 – Henniger et al.: 326, Fig. 4.5 – 4.6 Carychium vagum n.sp. [non Pl. 3, Fig. 13 = Turricarychium muelleri n.sp. (KA6)]. Pl. 3A Figs. 4, 5, 7, 8; Pl. 3B, Figs. 4, 5, 7, 8 Holotype (Pl. 3, Fig. 10): GPS Leipzig TM 0287. 1.54 : 0.49 : 0.48 mm, 5.2 whorls. Holotype: GPSL TM 0283; Pl. 3 Fig. 4. Paratypes: GPSL TM 0288/1, GPSL TM 0289/22, GPSL TM Paratypes from the locus typicus: GPSL TM 0284/2, GPSL TM 0290/1, GPSL TM 0291/17, GPSL TM 0292/14, GPSL TM 0293/3, 0285/1, GPSL TM 0286/1 (4 specimens). Total from Karsdorf is 5 DK/2. specimens from 4 out of 18 samples. In total 60 specimens from 9 out of 18 samples. Locus typicus and stratum typicum: Germany, Sach sen-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle Locus typicus: Germany, Sachsen-Anhalt, Karsdorf ce- (Saale), ca. 51°16′26″ N, 11°41′09″ E. Fissure infll: cal- ment quarry ca. 33 km SW Halle (Saale), ca. 51°16′26″ N, careous and carbonaceous silts and clays mixed with Tri- 11°41′09″ E. Fissure infll: calcareous and carbonaceous assic limestone debris; age: Early Eocene. silts and clays mixed with Triassic limestone debris; age: Early Eocene. Other paratypes: Grauves (France, Département Marne), late Ypresian: 14 specimens; MNHNP F.J17530 – F.J17535 Derivatio nominis: In honour of Frank Bach, Kustos and F.J17546 – F.J17547, coll. Cossmann. 3 specimens (curator) at the Institute for Geophysics and Geology at from this lot are fgured here (Pl. 3 Figs. 5, 7, 8). Leipzig University.

Derivatio nominis: Lat. vagus, -a, -um: roaming; refer- Diagnosis: A relatively small Carychium species with ring to the occurrences in the Paris Basin and in Sachsen- a slender cylindrical shell of 5.2 whorls; surface glossy, Anhalt. with growth lines only; in the aperture only the palatal tubercle and supracolumellar lamella are visible. Diagnosis: A relatively small Carychium species with a slender roundedly conical shell; surface glossy, with growth Dimensions: 1.54 : 0.49 : 0.48 (Pl. 3, Fig. 10, holotype) lines only; palatal tubercle and supracolumellar lamella 1.34 : 0.43 (Pl. 3, Fig. 11) well developed, the latter inside simple (i.e.not extended Dimensions of Carychium sp. aff. bachi from Grauves: nor undulating); columellar lamella not recognizable. 1.36 : 0.49 : 0.42 (Pl. 3, Fig. 9).

Additional observations: The sample from Grauves con- Relationships: The closest known relative is a species tains specimens which are broader than those from Kars- from Grauves (Département Marne, France; late Ypre- dorf. This may well be an effect of the small sample size sian), here fgured Pl. 3, Fig. 9. Its whorls are in their size

40 GEOLOGICA SAXONICA — 65: 2019

and shape completely congruent with the outline of the Diagnosis: as for the genus. whorls of C. bachi (see overlay, Fig. 9, abapertural view). A constant difference is the number of whorls: the spe- Dimensions: 1.89 : 0.80 : 0.67; 5.2 whorls (Pl. 4, F. 1, cies from Grauves terminates growth about half a whorl holotype) earlier than does C. bachi. The species from Grauves 0.93 : 0.52 (Pl. 4, F. 2) is therefore referred to as Carychium sp. aff. bachi. 22 1.75 : 0.69 (Pl. 4, F. 3). specimens of this species were encountered in a lot from the Cossmann collection (MNHNP, inventory numbers Additional observations: The abapical palatal fold is F.J11799/21 and F.J17529/1), mixed with 14 Carychium smaller than the adapical one and lacks a corresponding vagum (see above) and 2 non-carychiine shells. Coss- depression on the shell outside. The supracolumellar la- mann had this lot identifed as Carychium hypermeces mella points towards the abapical palatal fold. – Whorls Cossmann 1889, but the identifcation is highly uncer- 2 ½ – 4 (count beginning with the protoconch) increase tain: Cossmann’s fgure (1889: pl.12, fg. 33), which rep- much faster in width then the preceding ones, resultung resents a shell from early Ypresian strata of Mont Bernon, in a very weakly concave (coeloconoid) spira outline. shows a regularly conical shell shape which is not nor- This effect may have become somewhat exaggerated in mally encountered in carychiines. Therefore the drawing some specimens by the crystallization of gypsum, push- appears to be faulty, all the more because another im- ing shell walls outwards, but is also present in unaffected probable feature is the suture which runs at a right angle specimens. – The last ¼ whorl increases again in width. – to the shell axis. C. hypermeces was described from 4 Complete resorption of internal shell walls and the colu- specimens in the collection de Laubrière whose wherea- mella in the adapical part of the spire was observed in bouts are unknown. Cossmann & Pissarro (1913: pl.57, apical fragments. fg. 255-5) copied the original fgure, here reproduced pl. 3A, Fig. 6. Additional material from the type local- Relationships: A similar or related species has not been ity possibly referable to C. hypermeces is not known; a identifed. single specimen from Mont Bernon and also identifed by Cossmann as C. hypermeces is very different from the original fgure and from C. sp. aff. bachi as well as from Ovicarychium hennigeri n. sp. C. vagum. In addition to the incorrect original fgure, the Pl. 4A, Figs. 4 – 5; Pl. 4B, Figs. 4 – 5 size of C. hypermeces is uncertain: Cossmann 1889 gives as height 1.25 mm, but the scale given by Cossmann & v 2011 Carychiopsis cf. carbonaria- Henniger et al.: 326, Pl.3 Pissarro 1913: pl.57, fg. 255-5 implies a height of 1.86 Fig.12; Fig. 4.1-4.2 [non Wenz 1932]. mm. C. hypermeces has to be considered unidentifable until the species is redescribed from its syntypes or at Holotype: GPS Leipzig TM 0305; 2.03 : 0.99 : 0.93 mm, 5.5 least from topotypical material. whorls; Pl. 4 Fig. 4. Paratypes: GPSL TM 0306/1, GPSL TM 0307/18, GPSL TM 0308/24, GPSL TM 0309/2, GPSL TM 0310/3, GPSL TM 0311/1, GPSL TM 0312/2, GPSL TM 0313/16, DK/2. Turricarychium muelleri n. sp. In total 70 specimens from 10 out of 18 samples. Pl. 4A, Figs. 1 – 3; Pl. 4B, Figs. 1 – 3 Locus typicus and stratum typicum: Germany, Sachs- v 2011 Carychiopsis sp. 2 – Henniger et al.: Pl. 3, Fig.13 [non Figs. en-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle 4.5 – 4.6 (= Carychium bachi)]. (Saale), ca. 51°16′26″ N, 11°41′09″ E. Fissure infll: cal- v 2011 Carychiopsis sp. 1 – Henniger : 326, Fig. 4.3 – 4.4. et al. careous and carbonaceous silts and clays mixed with Tri- assic limestone debris; age: Early Eocene. Holotype: GPSL TM 0294; Pl. 4 Fig. 1. Paratypes: GPSL TM 0294/1, GPSL TM 0295/1, GPSL TM 0296/1, Derivatio nominis: Dedicated to Dr. Matthias Henniger, GPSL TM 0297/167, GPSL TM 0298/72, GPSL TM 0299/146, one of the researchers of the Karsdorf site. GPSL TM 0300/5, GPSL TM 0301/7, GPSL TM 0302/5, GPSL TM 0303/53, GPSL TM 0304/23, DK/2. Diagnosis: An Ovicarychium species with a broad, ovate- In total 483 specimens from 14 out of 18 samples. oblong shell; surface with riblets and fner spiral striation; aperture with two palatal and one parietal tubercle; Locus typicus and stratum typicum: Germany, Sachs- both palatal tubercles correspond to furrows on the out- en-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle side of the shell, which extend for ¼ whorl (unlike the (Saale), ca. 51°16′26″ N, 11°41′09″ E. Fissure infll: cal- tubercles themselves); supracolumellar lamella points to careous and carbonaceous silts and clays mixed with Tri- the interval between the palatal folds. The outer apertural assic limestone debris; age: Early Eocene. margin is strongly thickened and expanded, but a terminal varix is not developed. Derivatio nominis: dedicated to Prof. Dr. Arnold Müller (Leipzig), who researched the Karsdorf site and trusted Dimensions: 2.03 : 0.99 : 0.93 (Pl. 4 F. 3, holotype) the molluscs to the author for study. apertural height 0.85 (Pl. 4 F. 4)

41 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

Additional observations: The abapical palatal furrow Diagnosis: A species of Ovicarychium of ovate-conical creates the impression of a crest encircling the umbilical shape, a glossy surface with growth lines, an aperture area. A narrow umbilicus is probably present, but par- with terminal varix, a well developed parietal fold, three tially covered by the expanded columellar margin. The palatal knobs of which the middle one is strong and the original shape of the entire shell is a little more oblong two others very weak, the abapical one being pointed to than in the holotype, because of a tendency of the shells by the supracolumellar lamella, and a basal knob. Exter- to have the apical part pushed inwards, as it is broken off nal depression corresponding to palatal fold absent. the penultimate whorl because of its diminished mechan- ical strength due to the resorption of the the columella Additional observations: The terminal varix is strong- and the inner walls of the apical whorls. est developed in its abapical part. The peristome is almost straight and only modestly expanded. The last whorl does Remarks: O. carbonarium (Wenz 1932) is of similar size fare only very little before the aperture. The supracolu- and shape (see Pl. 4, Fig. 7), but differs by the duplicated mellar lamella is strongly developed (Pl. 4, Fig. 9). peristome, the lack of an external infrapalatal furrow, the middle palatal tubercle being narrower, the infrapalatal Dimensions. 1.44 : 0.76 : 0.76 (Pl. 4, Fig. 8, holotype) knob being very small and in a more abapical position, 1.20 : 0.65 (Pl. 4 Fig. 9) and the columellar lamella terminating further inside than Remarks: O. bigeminatum is the only other known spe- in O. hennigeri. cies with a terminal varix. It differs from O. ronlederi O. bigeminatum is smaller, more slender and lacks by lacking a basal fold and having two palatal folds of the external furrow corresponding to two palatal knobs, similar strength. but has a well-developed terminal varix. O. michelini is usually larger, more conical-elongate, lacks the parietal and abapical palatal knob, has a single Ovicarychium? sp. KA34 weak external depression and lacks a terminal varix. Pl. 4A, Fig. 6; Pl. 4B, Fig. 6 O. polysarcum is very broad, with a conical spire, a single palatal fold almost opposite the supracolumellar Material: Karsdorf quarry, fssure infll: GPSL TM 0320/1, GPSL lamella corresponding to a weak external depression and TM 0321/3, GPSL TM 0322/4. In total 8 specimens from 2 out of no terminal varix. 18 samples. O. sparnacense is smaller, more slender, without parietal and lower palatal fold and terminal varix. Dimensions: 0.90 : 0.55 : 0.45 (Pl. 4 Fig. 6) O. quadridens is smaller (Andreae 1884, internal mould: 1.6 : 0.9 mm), and has a relatively strong parietal fold. Remarks: Shell conical-ovoid, with well developed su- O. ronlederi is smaller, has a weak infrapalatal fold, a pracolumellar and columellar lamellae. As the adult ap- basal fold and no palatal external depression, as its place ertural margin was not preserved, its characters are un- is taken by a terminal varix. known, which precludes proposal of a new named species and even renders the genus attribution provisional. The shell is similar to that of O. ronlederi, but much smaller: Ovicarychium ronlederi n. sp. the fgured shell is 0.90 mm high at 4.25 whorls; compare O. ronlederi Fig. 9: height 1.20 mm at 3.75 whorls. The Pl. 4A, Figs. 8 – 9, Pl. 4B, Figs. 8 – 9 overlay of the contours of the holotype of O. ronlederi Holotype: GPS Leipzig TM 0314; Pl. 4 Fig. 8. (Pl. 4A Fig. 6) shows that the early teleoconch whorls of O. ronlederi are less wide, but the last two whorls of Paratypes: GPSL TM 0315/1, GPSL TM 0316/6, GPSL TM 0317/2, GPSL TM 0318/1, GPSL TM 0319/3, DK/1. In total 15 specimens O. ronlederi exceed in width those of O. sp. KA34. from 6 out of 18 samples.

Locus typicus and stratum typicum: Germany, Sach Zuella venusta n.sp. sen-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle Pl. 4A, Figs. 10 – 11; Pl. 4B, Figs. 10 – 11 (Saale), ca. 51°16′26″ N, 11°41′09″ E. Fissure infll: calcareous and carbonaceous silts and clays mixed with Tri- Holotype: GPS Leipzig TM 0323 (Pl. 4 Fig. 10), 1.36 : 0.64 : 0.54 assic limestone debris; age: Early Eocene. mm, 4.5 whorls. Paratypes: GPSL TM 0324/1, GPSL TM 0325/3, GPSL TM Derivatio nominis: Dedicated to Dr. Ronny M. Leder, 0326/2, GPSL TM 0327/4, GPSL TM 0328/2, DK/1. one of the researchers of the Karsdorf site. 2 In total 14 specimens from 5 out of 18 samples.

Locus typicus and stratum typicum: Germany, Sachs- en-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle 2 The name Carychium lederi Boettger 1880 (: 383) exists al ready, dedicated to the Austrian explorer Hans Leder (1843 – (Saale), ca. 51°16′26″ N, 11°41′09″ E. Fissure infll: cal- 1921) (https://www.biographien.ac.at/oebl/oebl_L/Leder_ careous and carbonaceous silts and clays mixed with Tri- Hans_1843_1921.xml). assic limestone debris; age: Early Eocene.

42 GEOLOGICA SAXONICA — 65: 2019

Derivatio nominis: Lat. venustus, -a, -um: lovely, beauti- Remarks: A large angular fold welded to a large palatal ful; referring to the evenly proportioned and well round- fold is also present in Albinula, but here this composite ed outline of the glossy whorls. fold (“anguloparietal fold”, Manganelli & Giusti 2000) occupies at most half the apertural height, whereas in Al Diagnosis: as for the genus, of which it is currently the binulopsis it takes 2/3 of the apertural height. Albinula sole species. species are also more elongate, develop about one more whorl and their shell height is about double that of Al Description: Shell very small, elongate-pupoid, not um- binulopsis. The dimensions of the whorls are from the bilicate; surface without recognizable growth lines, glossy. protoconch onwards larger than in Albinulopsis. The col- 4.5 whorls in total, protoconch/ teleoconch boundary not umellar fold is in Albinula less prominent and often lo- recognizable. Whorls convex, sutures moderately deep, cated more inside. The palatal folds differ only in detail. a narrow and weak subsutural cord is present.Last half The shell characters, and in particular the develop- whorl pulled inward, just before the peristome expand- ment of the angular fold, are well within the range of ing slightly, not descending or ascending on the preced- characters present in Gastrocoptidae. ing whorl. Aperture pear-shaped, peristome prosocline- Gastrocoptid species were hitherto known since the straight, sharp, but with a rounded transition into the in- Priabonian (Cox 1925b). Of these, “Albinula” kennardi terior, and a very weak and narrow lip slightly set back Cox 1925, is much smaller than younger Albinula spe- from the apertural margin. A rounded supracolumellar cies, viz. 1.9: 1.3 mm. In this respect it may be transition- lamella is present. al to Albinulopsis gibba, but it is still more elongate, and the angular lamella is fairly small. The other Priabonian Dimensions. 1.36 : 0.64 : 0.54 (Pl. 4 Fig. 10, holotype) gastrocoptid species, Gastrocopta (Sinalbinula) matura 1.19 : 0.56 (Pl. 4 Fig. 11). Cox 1925, agrees in its shape, size and dentition with the range present in later Gastrocopta species. “Carychium” nincki Cossmann in Cossmann & Pissarro 1913 from Bartonian strata of Marines in the Paris Basin may be Order Stylommatophora Schmidt 1855 another gastrocoptid species. Suborder Helicina Schileyko 1979 Infraorder Pupilloidei Schileyko 1979 (Orthu Albinulopsis gibba n.sp. rethra Pilsbry 1900) Pl. 5A, Figs. 1 – 3; Pl. 5B, Figs. 1 – 3

Pupilloidea Turton 1831 v 2011 Vertiginidae? gen.indet., sp.1 –Henniger et al.: 326, Fig.4.7. Gastrocoptidae Pilsbry 1918 Holotype: GPS Leipzig TM 0330, here Pl. 5, Fig. 1, 1.47 : 1.15 : 0.67 mm; 4.1 whorls. Paratypes: GPSL TM 0331/1, GPSL TM 0332/1, GPSL TM 0333/4, Albinulopsis n.gen. GPSL TM 0334/1, GPSL TM 0335/1, DK/1. Type species: Albinulopsis gibba n.sp. In total 9 specimens from 6 out of 18 samples.

Derivatio nominis: Alluding to the similarity with Albi Locus typicus and stratum typicum: Germany, Sachs- nula; gender: feminine. en-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle (Saale), ca. 51°16′26″ N, 11°41′09″ E. Fissure infll: cal- Diagnosis: Shell very small, conical-pupoid, with the careous and carbonaceous silts and clays mixed with Tri- last whorl slightly ascending on the preceding one, creat- assic limestone debris; age: Early Eocene. ing a relatively large umbilicus; terminal varix present; peristome expanded; aperture with a very large angular Derivatio nominis: Lat. gibba, a hump, alluding to the fold attached to an even larger parietal fold, up to four stout habitus of the shell. The name gibba is a noun and palatal folds, one basal fold and one columellar lamella. therefore invariable.

Other included species: Pupa bonneti Cossmann 1907 Diagnosis: A. gibba is slightly larger than A. bonneti, has (Pl. 5, Figs. 4 – 5) 3. a more faring aperture and a much stronger developed

3 Cossmann (1907: 283, pl. 10, fg. 272-3) described “Pupa Bonneti ” from a single specimen in coll. Bonnet, whose current location is unknown. As its sediment- flled aperture prevented recognition of the apertural barriers, Cossmann was unable to interpret its relationships. A new fgure of the specimen by Cossmann & Pissarro (1913: pl. 59, fg. 272-3) added no knowledge about the species, so that Wenz (1923: 1063) could do nothing but classify the species as “Pupillidae incertae sedis”. Two specimens from the type locality Grauves exist in coll. Cossmann (MNHNP J17537 – J17538; here Pl. 5, Figs. 2 – 3) which have the apertural dentition partially exposed, showing their close relationship with A. gibba n.sp. from Karsdorf. Cossmann must have acquired these after 1913.

43 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany) terminal varix. Of the four palatal folds the abapical two Valloniidae Morse 1864 are fused; basal fold large. Acanthinulinae Steenberg 1917 Description: Shell conical-pupoid, with moderately con vex whorls. Nucleus ca. 0.16 mm wide. Boundary to the teleoconch not recognizable. Teleoconch surface with Acanthinula Beck 1847 fne riblets. The last whorl grows in width much less than Type species (designated by v.Martens 1860): Helix aculeata Mül- the preceding ones, and ascends gently on the preced- ler 1774. Recent, Europe. ing whorl. The last 1/8 whorl expands again, and the apertural outer margin is faring. A broad terminal varix is strongest developed abapically, where it is separated Acanthinula (s.lat.) karsdorfensis n.sp. from the outer apertural margin by an indistinct furrow. Pl. 4A, Fig. 12; Pl. 4B, Fig. 12 In the aperture, the angular fold reaches the peristome and is welded to the parietal fold which is located more Holotype and sole specimen: GPSL TM 0329. 1 com- inside. This anguloparietal fold complex takes about 2/3 plete shell, strongly deformed: apical part pushed into of the apertural height. Four small palatal folds are pre- last whorl and tilted. 1.52 : 1.33 mm (height not restored), sent, of which the two abapical ones are fused with each aperture height 0.64 mm; ca. 3 whorls. other and correspond to a short elongate depression on the outside. The second palatal fold (counted from near- Locality: Germany, Sachsen-Anhalt, Karsdorf cement est to the apex) is the largest one. The basal fold is set quarry ca. 33 km SW Halle (Saale), ca. 51°16′26″ N, back from the apertural margin and is fairly large. The 11°41′09″ E. Fissure infll: calcareous and carbonaceous columellar fold is large and slightly undulating, which silts and clays mixed with Triassic limestone debris; age: results in it pointing downward in apertural view. The Early Eocene. palatal and columellar folds sit on a lip formed at a short distance from the peristome. A large umbilical depres- Diagnosis: A species of Acanthinula s.lat. smaller than sion exist, but due to incomplete preservation it cannot be all other known Palaeogene species, ovate- conical, with ascertained whether this is a pseudumbilicus (i.e. caused an open umbilicus and a refected peristome. only by the confguration of the last whorl) or not. Description: Overall shell shape was probably ovate- Dimensions: 1.44 : 1.13 : 0.67 (Pl. 5, Fig. 1, holotype). conical, with a rounded apex. Protoconch relatively large, Dimensions of A. bonneti: granulate, at least 1.25 whorls, on last 0.2 whorls also 1.38 : 1.09 : 0.75 (Pl. 5, Fig. 2) subtle collabral striae. Teleoconch with distant collabral 1.40 : 1.06 : 0.62 (Pl. 5, Fig. 3) thread-like ribs, many weaker ones in the interstices and even weaker spiral striae. Peristome refected. Umbilicus Remarks: The shell is slightly deformed so that it ap- narrow, open. pears to be broader in apertural view than it would in its intact state. In lateral view the shell is accordingly nar- Dimensions: 1.55 : 1.32 : 0.64 (Pl. 4, Fig. 12, holotype). rower. As this deformation is slight, it does not affect the differentiation from A. bonneti. Remarks: Species KA 23 (Pl. 9, Figs. 8 – 9) has similar The two specimens of A. bonneti from Grauves dif- sculpture but without spiral striae, the fragments are larg- fer by their slightly lesser size and narrower terminal er and the peristome is less expanded but may have an in- varix. Amongst themselves they show some variability: ternal weak lip and a weak and broad terminal varix. Spe- In specimen MNHNP J17537 (Pl. 5 Fig. 4) the last 1/8 cies KA13 (Pl. 9, Fig. 1) likewise has a similar sculpture whorl is more strongly ascending on the preceding whorl, including spiral striae, but is larger, and the peristome is and the peristome is stronger faring and the terminal var- much more expanded and sports an internal lip. ix is closer to the peristome than in specimen MNHNP Species attributable to Acanthinula s.lat. are present J17538 (Pl. 5 Fig. 5), which is also more slender. Due to throughout the Paleocene and Eocene of France only: sediment infll in the aperture the palatal folds are only Acanthinula (s.lat.) archiaci (Boissy 1848), A. armoricen partially visible and the basal fold not at all. Nonetheless sis (Cossmann 1902), A. cenchridium (Cossmann 1902), the two fused palatal folds of A. gibba seem to be absent A. dumasi (Boissy 1848), A. geslini (Boissy 1848), A. stueri in A. bonneti, and the corresponding external depression (Cossmann 1892) (with synonym bouryi Cossmann 1889, is absent. non Morgan 1885). All of them are signifcantly larger than A. karsdorfensis, the smallest (A. dumasi (Boissy 1848) already being 2.5 mm high and 2 mm wide (Boissy 1848, Deshayes 1863). The refected peristome of A. kars dorfensis indicates a mature specimen despite its very small size.

44 GEOLOGICA SAXONICA — 65: 2019

Infraorder Clausilioidei tending ¼ whorl into the interior, and up to four palatal folds. The palatal folds do not reach the peristome, and Superfamily Clausilioidea Gray 1855 are elongate, with their long axis at right angles to the peristome.

Clausilioidea sp. KA26 Remarks: The families Charopidae and Endodontidae Pl. 5A, Fig. 6; Pl. 5B, Fig. 6 cannot consistently be distinguished by shell characters. Attribution of several subfamilies to Charopidae or Material: Karsdorf quarry, fssure infll: 1 apex fragment out of 18 Endodontidae differs signifcantly between Schileyko samples. GPSL TM 0188. (2001) and Bouchet et al. (2017); furthermore, the latter authors highlight unresolved issues in phylogeny and Description: Preserved are 2.2 sinistral whorls, of which classifcation. The genus Afrodonta is placed in Endo- 1.7 constitute the protoconch. Nucleus 0.36 mm wide, dontidae: Endodontinae by Schileyko (2001: 902), but in protoconch diameter 1.21 mm. All whorls are moderate- Charopidae by Herbert & Kilburn (2004: 246). ly convex; the suture is distinct and simple. Teleoconch The habitus of Afrodontops, in particular the de- with collabral riblets which fade away on the umbilical pressed shape with a wide umbilicus and four subequal side. Shell probably non-umbilicate. Columella twisted, and almost equidistant palatal folds is strikingly similar to without lamellae. that of many charopid and endodontid genera described mainly from Pacifc Islands, but also from eastern Indo- Dimensions: 1.79 : 1.25 (Pl. 5, Fig. 6). nesia, New Zealand, Australia, subsaharan Africa, South Atlantic Islands (see synopsis by Schileyko 7, 2001) Remarks: The shell habitus could be that of a Triptychia and fossil from South America (Salvador et al. 2018). (family Filholiidae) or a clausiliid (family Clausiliidae, However, all of them are larger, with breadths ranging both in Clausilioidea). Triptychia species are known from 1.0 to 10 mm, and lack a large parietal lamella or since the Lutetian (Schnabel 2006a). Unfortunately lamellae. The recent genera have typically several pari- characters of the earliest whorls which could possibly etal folds or lamellae of smaller dimensions; if only one serve to distinguish between Clausiliidae and Filholiidae parietal fold is developed, it is usually in the context of an are very insuffciently known. In the Oligocene species overall reduction of the dentition, and such fold is much T. kadolskyi Schnabel 2007, at a shell width of 1.24 mm smaller than that of Afrodontops europaea. Moreover, a hollow columella with two lamellae is visible. The ab- numerous genera without apertural dentition exist. sence of columellar lamellae in sp. KA26 argues against A single genus, Afrodonta Melvill & Ponsonby 1908 an attribution of this fragment to the Filholiidae. How- is recognized in Africa (South Africa, Mozambique, ever, a shell fragment with a hook-shaped palatal fold Kenya). Its type species, A. bilamellaris Melvill & Pon- (sp. KA 35, see below; Pl. 9 Fig. 10) could represent a sonby 1908, and another species, A. connollyi Solem Triptychia species, which would be an older occurrence 1970, resemble Afrodontops by their relatively strong than hitherto known. parietal lamella, a well developped columellar tubercle and a surface ornamentation of growth lines only. They differ however by their larger shell size (average breadth of A. bilamellaris 1.54 – 1.65 mm, and of A. connollyi Superfamily unassigned to an infraorder 1.48 – 1.55 mm). The lectotype of Afrodonta bilamellaris is here illustrated (Pl. 7, Fig. 1) at the same scale as the Punctoidea Morse 1864 Afrodontops specimens (Pl. 6, Figs. 7 – 11). Other spe- Charopidae Hutton 1884 or Endodontidae cies attributed to Afrodonta exhibit a larger range of shell Pilsbry 1895 shapes and dentitions, e.g. two parietal folds or none, and 0 – 7 palatal folds (Burnup 1912, Solem 1970, Herbert & Kilburn 2004). Solem (1970) supposed the genus in its current scope to be polyphyletic. It is tempting to postu- Afrodontops n.gen. late a relationship between the Recent A.-bilamellaris- Type species: Afrodontops europaea n.sp. connollyi-group and Afrodontops on the basis of the strong parietal lamella, but in view of the huge spatial Additional included species: Afrodontops comes n.sp. and temporal distance between them this at present no more than a hypothesis. Derivatio nominis: Similar to Afrodonta; gender femi- The present-day distribution of the Charopidae + En- nine. dodontidae is exclusively Gondwanan, with “excursions” into the Pacifc islands resulting in spectacular radiations. Diagnosis: Shell very small (width 0.84 mm), with a The presence of the family in South America is proven very low spire and wide umbilicus; whorls well rounded, by fossils back to the Barremian/Aptian (see checklist by ornamentation consisting of growth lines; aperture with a Salvador et al. 2018). The discovery of Afrodontops in simple peristome; one or two large parietal lamellae ex- the Eocene of Europe is a frst for the group. Possibly the

45 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

very small size of most species made passive dispersal Afrodontops comes n.sp. easier, but simultaneously made them prone to be over- Pl. 5A, Fig. 11; Pl. 5B, Fig. 11 looked by researchers. Holotype and sole specimen: GPS Leipzig TM 0199; Pl. 5 Fig. 11. Afrodontops europaea n.sp. Pl. 5A, Figs. 7 – 10; Pl. 5B, Figs. 7 – 10 Locus typicus and stratum typicum: Germany, Sachs- en-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle v 2011 Australorbis cf. pseudammonius – Henniger et al.: 326, (Saale), ca. 51°16’26″ N, 11°41’09″ E. Fissure infll: cal- Pl. 3, Fig. 14 [tantum, non Pl. 3, F 7 (= Biomphalaria sp. KA2); careous and carbonaceous silts and clays mixed with Tri- non Schlotheim 1820] assic limestone debris; age: Early Eocene. Holotype: GPS Leipzig TM 0189; Pl. 5, Fig. 8. Derivatio nominis: Lat. comes, a companion (of A. euro Paratypes: GPSL TM 0190/1, GPSL TM 0191/1, GPSL TM 0192/1, GPSL TM 0193/4, GPSL TM 0194/2, GPSL TM 0195/1, GPSL TM paea); the epithet is a noun in apposition and hence im- 0196/1, GPSL TM 0197/4, GPSL TM 0198/1, DK/1. mutable. In total 18 specimens from 8 out of 18 samples. Diagnosis: Afrodontops comes is an Afrodontops species Locus typicus and stratum typicum: Germany, Sachs- with two parietal lamellae, an adapical palatal knob and en-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle a columellar knob. The shell shape and shell size are like (Saale), ca. 51°16’26″ N, 11°41’09″ E. Fissure infll: cal- those of A. europaea. careous and carbonaceous silts and clays mixed with Tri- assic limestone debris; age: Early Eocene. Dimensions: 0.43 : 0.83 : 0.39; 2.2 whorls (Pl. 5, Fig. 11). Diagnosis: Afrodontops europaea is an Afrodontops species with one strong parietal lamella, three palatal knobs Remarks: The differences between A. europaea and and a columellar knob. A. comes are comparable to the differences between modern Afrodonta species (Herbert & Kilburn 2004) Description: Shell very depressed, spire very little el- which are likewise of similar size and habitus and have evated above the last whorl; 2.3 whorls, well rounded; 0 – 2 parietal lamellae, 1 columellar knob or fold and 0 – 8 umbilicus wide; nucleus of protoconch convex, dome- palatal folds. shaped, boundary to teleoconch indistinct; teleoconch whorls convex, suture deepened; ornamentation consists of growth lines of variable strength, slightly prosocline- Infraorder Orthalicoidei Hausdorf & Bouchet concave. Last whorl not descending towards the aperture. 2017 Aperture nearly circular, at the parietal wall cut out in a very shallow arc. Peristome simple; set back from the per- Orthalicoidea v. Martens 1860 istome in the interior are three palatal knobs, a columellar Palaeostoidae Nordsieck 1986 knob and one parietal lamella; the palatal and columellar knobs are all of similar shape: elongate with their long axes perpendicular to the peristome. The parietal lamella The species of the genus Palaeostoa have early on been is large and extends ca. 1/4 whorl into the shell’s interior, placed in the South American modern genus Megaspira where it develops a furrow on its crest. Lea in Jay 1836 ((Boissy 1848, Deshayes 1863, Sand- berger 1870 – 71, Gardner 1885, Newton 1891, New- Dimensions: 0.43 : 0.83 : 0.39 (Pl. 5, Fig. 8, holotype) ton & Harris 1894) or as a separate genus Palaeostoa 0.41 : 0.69 (Pl. 5, Fig. 7) within the family Megaspiridae (Pilsbry 1904, Wenz 0.44 : 0.83 : 0.40 (Pl. 5, Fig. 9) 1923, Zilch 1959). The type species of Megaspira is M. 0.44 : 0.84 : 0.39 (Pl. 5, Fig. 10). ruschenbergiana Lea 1836, which is a junior subjective synonym of Pupa elatior Spix in Spix & Wagner 1827 Remarks: As the shells are lightly corroded, any or- from Brazil; its current classifcation is in superfamily namentation fner than the growth lines could possibly Orthalicoidea, family Megaspiridae (Simone 2006; Bou- have been eroded. The palatal and columellar knobs are chet et al. 2017). Nordsieck (1986) proposed to classify set back from the peristome and are therefore easily ob- Palaeostoa and possibly the Turonian Ptychicula Tausch scured by rock matrix (Pl. 5, Fig. 10). 1886 in a separate family Palaeostoidae, differentiated from Megaspiridae by the presence of three subequal columellar lamellae (in Megaspira the two abapical ones are weak and lie in the adapical half of the columella), the presence of numerous palatal lirae (none in Meg aspira), the absorption of the early internal lamellae and

46 GEOLOGICA SAXONICA — 65: 2019

lirae, and biogeography. Nordsieck placed the Palaeos- careous and carbonaceous silts and clays mixed with Tri- toidae in Clausilioidea (1986: 98, 109, 112; 2015), pre- assic limestone debris; age: Early Eocene. sumably because of some similarities with the Filholii- dae: the turreted shell, and the presence of columellar Diagnosis: A Palaeostoa species in which the frst 1.4 and parietal lamellae, and (in Triptychia) some palatal whorls of the teleoconch bear fne collabral riblets be- plicae. Thus both families have in common the putative coming stronger on the following whorls; they are con- plesiomorphous condition of the Stylommatophora as tinuing to the peripheral angularity of the whorls and are deduced by Nordsieck (1986: Fig. 1), which renders the fner and more numerous than in P. exarata and P. elon interpretation of relationships ambiguous. The internal gata, and of equal strength across their length, unlike P. lamellae system of the Clausiliidae is more derived and fontenayi from Grauves and Bouxwiller. bears in detail little resemblance to that of the Filholiidae and Palaeostoidae. Schnabel (2006) described and sum- Description: Protoconch 1.1 whorls, convex, glossy, with marized the differences between Filholiidae and Clausi- very fne spirals and growth lines. The latter are strong- liidae and doubted whether a superfamily Clausilioidea est near the adapical suture. Nucleus ca. 290 µm broad, which included Filholiidae and Palaeostoidae were truly whole protoconch ca. 1.1 mm broad. Apex dome-shaped. monophyletic. I propose to place at least the Palaeos- Flanks of protoconch whorls somewhat fattened. toidae in Orthalicoidea, within which the closest rela- Teleoconch whorls moderately convex, with stepped tionship would be with Megaspiridae. The differences suture; frst 1.4 whorls with indistinct collabral ribbing, between Palaeostoa and Megaspira, as emphasized by strongest adapically, resulting in a crenellated suture. All Nordsieck, are to be expected due to separate evolution following whorls with strong and regular collabral rib- of Megaspira on a different continent since the middle bing, ribs narrower than their interstices; a subsutural fur- Cretaceous. The biogeographical counterargument does row is present. The ribs terminate at an obtuse peripheral not hold, as faunal exchanges between South America angle. Umbilical side with subtle growth lines and some and Europe + North Africa are well established, see spiral furrows. Umbilicus narrow but open. Aperture not chapter “Discussion and Conclusions” below. preserved. Fragments show three strong columellar folds and a much weaker one in supracolumellar position, Palaeostoa Andreae 1884. Type species: Pupa fontenayi one parietal lamella and 8 – 9 palatal lirae of subequal Sandberger 1871 (subsequent designation by Cossmann strength. These lirae are already developed on the frst 1905: 59). Lutetian. Bouxwiller, Alsace. This designation teleoconch whorl. On larger whorl fragments three mi- has precedence over the designation of Clausilia crenata nor lirae being intercalated between the larger ones have Sandberger 1871 by Wenz (1923: 737). Andreae (1884) been observed. synonymized the two taxa and gave fontenayi precedence over crenata; Wenz’s (1923: 737) later reversal of prec- Dimensions: edence (crenata over fontenayi) was thus invalid. P. costellata Synonym: Eomegaspira Pilsbry 1904. Type species 4.58 : 2.12; 5.7 whorls (Pl. 6, Fig. 2; holotype) by original designation: “Megaspira exarata (Michaud) largest dimension 1.29 (Pl. 6, Fig. 3) Deshayes”, here corrected to Pyramidella exarata 7.29 : 3.38 (Pl. 7, Fig. 1) Michaud 1838. Rilly-la-Montagne, Département Marne, width 3.83 (Pl. 7, Fig. 2) late Thanetian. 3.58 : 2.32 (Pl. 7, Fig. 3) 1.90 : 1.80 (Pl. 7, Fig. 4) 1.22 : 1.62 (Pl. 7, Fig. 5) Palaeostoa costellata n.sp. width 1.16 (Pl. 7, Fig. 6) Pl. 6, Figs. 2 – 3; Pl. 7, Figs. 1 – 6; Pl. 8A, Fig. 3; P. elongata Pl. 8B, Fig. 3 32.36 : 8.31 : 8.26 (Pl. 8, Fig. 1; apical part only) v 2011 Vertiginidae? gen.indet., sp.2 – Henniger et al.: 326, Fig. 4.8 P. exarata Holotype: GPSL TM 0200, here Pl.6 Fig. 2 and Pl. 8 Fig. 3. 3.25 : 2.15 (Pl. 8, Fig. 2) Paratypes: GPSL TM 0201/1, GPSL TM 0202/1, GPSL TM 0203/1, P. cf. fontenayi GPSL TM 0204/1, GPSL TM 0205/1, GPSL TM 0206/1, GPSL TM height 4.50 (Pl. 8, Fig. 4) 0207/1, GPSL TM 0208/2, GPSL TM 0209/1, GPSL TM 0210/2, GPSL TM 0211/1, GPSL TM 0212/1, GPSL TM 0213/3, GPSL TM P. fontenayi 0214/1, TM 0234/1, DK/1. width 3.25 (Pl. 8, Fig. 5, holotype of Fragments counted in total as 20 specimens from 12 out of 18 sam- Clausilia crenata Sand- ples. berger) Locus typicus and stratum typicum: Germany, Sachs- Remarks: P. costellata n.sp. can be described as a ‘miss- en-Anhalt, Karsdorf cement quarry ca. 33 km SW Halle ing link’ in a lineage leading from the Thanetian P. elon (Saale), ca. 51°16’26″ N, 11°41’09″ E. Fissure infll: cal- gata (Melleville 1843) (Pl. 8, Fig. 1) to the Lutetian

47 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

P. fontenayi (for the age and stratigraphic occurrences Helicoid land snail sp. KA12 of Thanetian and Ypresian species see Textfg. 2). In the Pl. 8A, Figs. 7 – 9; Pl. 8B, Figs. 7 – 9 older species P. elongata (from Châlons-sur Vesle) and P. exarata (Michaud 1838) (from Rilly-la-Montagne, Material: Karsdorf quarry, fssure infll: GPSL TM 0216/1, GPSL late Thanetian) (Pl. 8 Fig. 2) strong collabral ribbing ex- TM 0217/1, GPSL TM 0218/1. In total 3 specimens from 3 out of ists from the beginning of the teleoconch onwards; in the 18 samples. younger P. cf. fontenayi (Grauves, late Ypresian) (Pl. 8, Fig. 4) and P. fontenayi (Bouxwiller, Lutetian) (Pl. 8, Shell probably with a low spire and rounded, convex Fig. 5) collabral ribbing is greatly reduced: ribs are con- whorls. Umbilicus open, wide. Ornamentation is fne spicuous only in the subsutural zone, giving a crenellated riblets on apical and umbilical sides, no spirals. The appearance to the suture. whorls of this shell are larger than in helicoid species In P. costellata the collabral ribs are very weak on the KA19 and 25. frst 1.4 teleoconch whorls, and on the following whorls less strong than in P. elongata and P. exarata but more Dimensions: Width 2.41; 2.6 whorls (Pl. 8 Fig. 7) numerous. Width 0.79 (Pl. 8 Fig. 8) Width 1.04 (Pl. 8 Fig. 9).

Indeterminable land snails Land snail sp. KA19 Pl. 8A, Fig. 10; Pl. 8B, Fig. 10

A. Fragments with protoconch Material: GPSL TM 0219 (Pl. 8, Fig. 10: 1 apical fragment), TM 0220 (1 fragment).

Helicoid land snail sp. KA3 The frst 1.6. whorls are preserved. Their shape is similar Pl. 8A, Fig. 6; Pl. 8B, Fig. 3 to that of sp. KA25, but the whorls are larger and are descending, indicating an elevated spire. They are well rounded. The umbilicus is open but narrow. This proto- v 2011 Archaegopis sp. – Henniger et al.: 326, Pl. 3, Fig. 8 conch is similar to that of Cochlostoma sp. KA16, but it Material: GPSL TM 0215: 1 apical fragment from Karsdorf ce- is smaller. ment quarry. Dimensions: 0.55 : 0.70 : 0.31 (Pl. 8 Fig. 10). Description: Protoconch 0.6 whorls without recognizable ornamentation, 0.85 mm wide, nucleus 0.27 mm wide. Teleoconch : only the frst 1.6 whorls are preserved, Helicoid land snail sp. KA25 having a diameter of 2.69 mm. Shell shape helicoid, Pl. 8A, Fig. 11; Pl. 8B, Fig. 11 spira moderately elevated, apex rounded. Whorls with peripheral angle. Teleoconch ornamentation consists Material: GPSL TM 0221/1, GPSL TM 0222/1, GPSL TM 0241/1. of collabral riblets which tend to be regular but change In total 3 specimens from 3 out of 18 samples. strength and density at growth interruptions. Corrosion of the apical side may have oblitterated microsculpture, The frst 1.4 whorls are preserved. Their shape and size but traces of fne spirals can still be seen on the last ¼ are similar to those of Afrodontops europaea, but the um- whorl. Umbilical side not preserved: a small umbilicus is bilicus is narrower and the aperture lacks internal barri- present; ornamentation consists of growth lines and spi- ers. rals of similar strength. Dimensions: 0.42 : 0.68 (Pl. 8 Fig. 11). Remarks: The incomplete preservation, coupled with the lack of diagnostic shell characters (at least in the preserved part of the shell) precludes assignment of the Elongate sinistral land snail sp. KA29 specis to a particular genus or even family. Externally Pl. 8A, Figs. 12 – 13; Pl. 8B, Figs. 12 – 13 similar species are signifcantly larger; their diameter at 2.2 whorls is: Material: GPSL TM 0223/1, GPSL TM 0224/1, GPSL TM 0225/1. Phacozonites luna (Michaud 1837) (Merfy, Thanetian): In total 3 specimens from 2 out of 18 samples. 3.3 mm The shells are sinistral. The apex is rounded. The nucleus Phacozonites? ceciliae (Wenz 1932) (Geiseltal, Lute- is 0.13 mm wide. Preserved is a total of 3.75 whorls; the tian): 7.3 mm protoconch/ teleoconch boundary is not recognizable. Archaegopis discus (Thomä 1845) (Hochheim, Chat- Whorl outline is convex, becoming fatter with age, and tian): 5.5 mm. are stepped near the suture. Teleoconch sculpture con-

48 GEOLOGICA SAXONICA — 65: 2019

sists of broad fattened growth striae with narrower inter- Land snail species KA 38 stices. No umbilicus. Columella twisted. Pl. 9, Fig. 2

Dimensions: 1.09 : 0.51 (Pl. 8, Fig. 12) Material: GPSL TM 0230: 1 fragment. 0.96 : 0.43 (Pl. 8, Fig. 13) The fragment shows distinct ribs on the outside, which The sinistral coiling and the twisted columella combined are mostly regularly spaced and narrower than their in- with the absence of an umbilicus are characters shared terstices. The peristome is strongly expanded and bears with some species of Scalaxis Pilsbry 1909, which to an internal lip which is separated from the sharp apertural date are only known from the Thanetian limestone of margin by 2 furrows. Rilly-la-Montagne. Due to incomplete preservation we refrain from attribution to this or any other genus. Land snail sp. KA 22 Elongate sinistral land snail sp. KA37 Pl. 9, Fig. 7

Pl. 8A, Fig. 14; Pl. 8B, Fig. 14 Material: GPSL TM 0238 (Pl. 9, Fig. 7): 1 apertural fragment.

Material: GPSL TM 0226: 1 apical fragment from one sample out The shell has a narrow umbilicus and widely spaced col- of 18. labral ribs without spiral striae on the umbilical side. The columellar margin is straight, expanded and refected. It The shell is sinistral and broader than that of sp. KA29. does not bear denticles (the ‘knobs’ on the specimen are 2.75 whorls are preserved. The nucleus is 0.09 mm wide. matrix rock). Sculpture is not recognizable, nor is the shape of the columella. The umbilicus is open. Greatest dimension: 1.52 (Pl. 9, Fig. 2)

Dimensions: 0.79 : 0.51 (Pl. 8 Fig. 14). Species KA13 shows some similarity, but is much larger and has spiral striae. This species may be related to sp. KA29; the open umbilicus is compatible with Cylindrellina Munier-Chalmas 1884, but sinistral species are hitherto not known in this Land snail sp. KA23 genus. Also in this species the preservation is too in- Pl. 9, Figs. 8 – 9 complete for a defnitive genus attribution. Cylindrellina species are known from Danian strata of Belgium and Material: GPSL TM 0239/1, GPSL TM 0240/1, GPSL TM 0242/1, Thanetian to Lutetian strata of France. GPSL TM 0243/1. Fragments counted in total as 4 specimens from 3 out of 18 samples.

The whorls are well rounded. The ornamentation consists B. Fragments of teleoconch with apertural of widely spaced raised collabral riblets, between which parts numerous growth lines of variable strength are developed. All collabral structures are slightly sinuous. The peristome is weakly expanded and bears a weak internal lip. Elongate landsnail sp. KA13 Greatest dimension of peristome 1.22 (Pl. 9, Fig. 8) Pl. 9, Fig. 1 Greatest dimension of peristome 1.00 (Pl. 9, Fig. 9) Material: GPSL TM 0227/1, GPSL TM 0228/1, GPSL TM 0229/1. In total 3 specimens from 3 out of 18 samples.

The aperture is oval, with a strongly expanded and re- C. Parts of teleoconch only fected peristome and a lip which is somewhat offset from the peristome. The shell has a narrow umbilicus. The last whorl is well rounded. The ornamentation con- Helicoid land snail sp. KA 20 sists of distant raised collabral ribs, between which spiral Pl. 9, Figs. 3 – 4 striation is well developed on both the adapical and the umbilical sides. This form may be a member of either the Material: GPSL TM 0231/1, GPSL TM 0232/1, GPSL TM 0233/1, Cyclophoroidea or Pupilloidea. GPSL TM 0235/1. Fragments counted in total as 4 specimens from 3 out of 18 samples. Dimensions: Apertural height 1.28 (Pl. 9, Fig. 1). All fragments show regular strong collabral ribbing. On the adapical fragments the ribs are regularly curved. The

49 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

umbilical fragment indicates a wide open umbilicus into Boettger 1877 (Schnabel 2006a: fg. 3 and 2006b: pl. 10 which the ribbing continues. Inside the umbilicus the ribs fgs. 114, 119), but these are of Miocene and Pliocene are crossed by spiral lirae which weaken gradually to- age. To date Triptychia is not known before the Lutetian. wards the outer rim of the umbilicus. This fragment belongs potentially to Clausilioidea sp. KA26. Due to lack of defnitive evidence it is preliminar- Greatest dimension (diagonally) 1.87 (Pl. 9, Fig.3) ily listed separately. Greatest dimension (along margin) 1.20 (Pl. 9, Fig. 4)

This type of ornamentation is reminiscent of the genus Land snail sp. KA36 Discus, but in Discus the collabral ribs weaken in the um- Pl. 9, Fig. 12 bilical side, and the umbilicus lacks spiral ornamentation. Material: GPSL TM 0246: 1 Fragment. This fragment of part of a middle whorl is well rounded, Elongate land snail sp. KA 21 with stronger curvature abapically. The sculpture con- Pl. 9, Figs. 5 – 6 sists of fne, fairly regular growth striae; further, the entire surface is covered by a very fne granulation. Material: GPSL TM 0236/1, GPSL TM 0237/1. In total 2 fragments counted as 1 specimen from 1 out of 18 samples. Greatest dimension (width) 2.64 (Pl. 9 Fig. 12).

Ornamentation of broad, fattened rounded ribs, wider than their interstices. No spirals. A subsutural furrow seems to exists, marked by irregularities in the ribs near Discussion and conclusions the adapical suture. On the inside along the suture there is a remnant of shell of the preceding whorl; its angle with the shell wall indicates an elongate shell. Faunal associations and paleoecology

Greatest dimension 2.23 (Pl. 9, Fig. 5) As all samples were taken from scree, their spatial relationships are unknown. Therefore the author attempted Elongate land snail sp. KA24 to deduce from the composition of individual samples Pl. 9, Fig. 11 differences which could be attributed to paleocological and/or stratigraphical causes. This was only possible in Material: GPSL TM 0245/1, GPSL TM 0336/1. In total 2 speci- a limited way because of the variances in the number pf mens from 2 out of 18 samples. recovered individuals per sample, which in turn resulted in apparent differences in species diversity per sample The shape is reminiscent of Oleacinidae species, but the (Fig. 2). The conspicuous correlation for terrestrial mol- remnant is insuffcent for a frm family identifcation. luscs suggests that samples with low numbers of spec- The ornamentation consists of gently sinuous collabral cimens would most likely had yielded larger numbers of ribs with narrower interstices into which growth lines can species if a larger rock sample had been taken and picked. be intercalated. The peristome is largely chipped off, but It follows that, the lower the recovery of specimens from an elongate palatal tubercle is still recognizable. There is a sample is, the less signifcant is the absence of a spe- no subsutural furrow. cies, i.e. the ‘absent’ criterion can only in a limited way be utilized in interpretations. Greatest dimension 2.91 (Pl. 9, Fig. 11). The possibility of grouping the samples into different associations (whether due to paleoecological infuences or signifcant age differences) was therefore approached Land snail sp. KA35 qualitatively (Table 1). Pl. 9, Fig. 10 Amongst the aquatic molluscs, the two most frequent species are Galba sp. KA1 and Biomphalaria sp. KA2. Material: GPSL TM 0244: 1 Fragment. They occur in 9 out of 18 samples together, whereby the two frequency maxima of Galba sp. KA1 are also the The fragment shows dichotomising collabral ribs, whose frequency maxima of Biomphalaria sp. KA2. The re- shape is somewhat similar to that of sp. KA21. The inside maining species co-occur in low numbers with these two, bears a remarkable long hook-shaped fold. with the exception of Acroloxus aspis, which in Sample no. 1 is the only aquatic species, and Acroloxus korys, Greatest dimension 1.20 (Pl. 9, Fig. 10). which occurs in only one sample (KD2382) where it is frequent.It follows that a single aquatic mollusc associa- Palatal folds of variable, and sometimes similar shape oc- tion can be recognized with confdence, characterized by cur in species of the subgenus Triptychia (Plioptychia) Galba sp. KA1 and Biomphalaria sp. KA2. Their habitat

50 GEOLOGICA SAXONICA — 65: 2019

Fig. 2. Species recovery per sample vs. sample size. Abb. 2. Artenzahl per Probe in Abhängigkeit von der Probengröße. was the photic zone of a lake or pond, where they lived and high energy aqueous transport mechanisms. The only on aquatic plants and on soft substrates up to the shore- coarse component is Muschelkalk debris, which is de- line. The more irregularly distributed Acroloxus species rived from the walls of the karst fssure. Remarkable is indicate probably a particular sub-environment. Modern the absence of recognizable sediment layering and the Acroloxus species live on reed, i.e. a frm substrate. By relative uniformity of the mollusc fauna as recovered analogy, A. korys and A. aspis likewise may have lived from various samples of sediment. The original depth on frm substrates such as stones, dead shells, branches of the karst fssure at the sampling level may have been fallen off from trees and larger aquatic plants, but not on about 130 m (Henniger et al. 2011). For an interpretation the lake substrate. of the depositional environment the following facts need The terrestrial mollusc associations are also fairly also be considered: homogenous. Out of the 18 samples, 4 are either com- The aquatic molluscs are, with the exception of the pletely or almost completely devoid (few fragments at- Sphaeriidae, pulmonates, i.e. air-breathing snails which tributed to undetermined land snails in samples 5 and 6) lived preferentially in shallow waters up to the water sur- of land snails. In the remaining 14 samples, Turricary face, and are often found on aquatic plants. This makes it chium muelleri occurs in all 14, Palaeostoa costellata unlikely that they lived at the bottom of a ca. 130 m deep in 12, Ovicarychium hennigeri in 10, Carychium bachi fssure, which would have been dark and hence devoid in 9, and Afrodontops europaea in 8 samples. All land of living plants. Large bottom-dwelling molluscs such snail species co-occur with Turricarychium muelleri as Viviparidae and Unionoidea are absent. Remarkably, and Palaeostoa costellata, and almost all of them with fshes are also absent, suggesting the pond was not con- Ovicarychium hennigeri and Afrodontops europaea, too. nected to the regional drainage system. It follows that only one terrestrial gastropod association None of the aquatic molluscs is an indicator of in- is recognizable, which is characterized by the 5 above- creased salinity. named species. Land snails: It is improbable that all or most would The diversity of the land snail fauna points to a well have lived on the steep walls of the karst fssure, because vegetated environment which would require a suffcient- of the likely darkness which prevented the growth of ly humid climate. This is consistent with the presence of a plants which are food base for many species, and because rich carpofora. Some land snails belong to groups known of the improbability that all or most of the species were today only from tropical to subtropical climates: Megalo- rock dwellers. More likely they were swept into a pond mastomatinae, Endodontidae/Charopidae, Orthalicoidea. from the adjacent land surface. The good preservation The species of the genus Carychium, which is the only and absence of obvious traces of weathering and wear by one still existing today occur world-wide and are ground- transport indicates very short-distance transports. dwellers in a variety of more or less moist habitats. The The good preservation of aragonitic shells requires remaining land snails are inconclusive due to lack of the waters to be alkaline, which, considering the pres- close living relatives. The fora also indicates a tropical ence of limestone formations around the karst fssure is to subtropical climate (Henniger et al. 2011). not surprising. It also must have been poorly oxygenated to preserve the numerous land plant remains. We propose therefore the hypothesis that the sedi- Depositional environment ment infll originated in a pond at the land surface, which may have formed in a depression above the incipient fs- The rather fne-grained, poorly sorted and not oxidized sure. This pond was not connected to the regional drain- nature of the sediment matrix excludes both wind-blown age system; the aquatic molluscs in it were only those

51 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany) 1 1 2 1 1 1 1 1 3 2 1 1 8 5 2 1 1 16 22 70 12 Sum 1 4 1 4 1 3 1 3 1 1 3 15 KD 2379 1 1 3 1 2 1 6 6 1 18 1 1 9 1 5 5 2 14 1 1 1 6 1 22 52 2 71 16 3 61 KD 2382

KD 807 1 1 + 1 + + 12 KD 806

1 1 1 1 1 3 1 20

1 1 4 1 2 16 5 7 5 53 23 483 KD 805

1 1 1 4 1 5 1 19 379 16 789 KD 2328 2 1 1 2 1 1

2 3 3 1

1? KD 2365

1 2 1 1 1

26 2 KD 803

1 1 1 2 1 17

+ 2 Karsdorf samples 2 1 1

2 2 1 1 1 + + 1 1 1 6 2

3 43 3 74 146

+ ++ 2

1 1 1

3 4 3 6 1 2 3 13 18 114

1 6 Composite sample

+ +

1 + ++ 10 + + + 3 ++ ++++ 358

1 1

1 2 3

2 1 2 1 3 2 2 7

6 2 1 2 3 4 5 6 7 KD2 KD6a KD15

n.sp. n.sp. n.sp. n.sp. n.sp. n.sp. sp. KA2 (s.lat.) karsdorfensis n.sp. Land snail sp. KA22 Land snail sp. KA23 Elongate land snail sp. KA24 Land snail sp. KA35 Land snail sp. KA36 spp. frgm. Pulmonata div. Elongate land snail sp. KA21 Helicoid land snail sp. KA20 Land snail sp. KA38 Elongate land snail sp. KA37 Elongate land snail sp. KA13 Helicoid land snail sp. KA12 Land snail sp. KA19 Helicoid land snail sp. KA25 Elongate land snail sp. KA29 Helicoid land snail sp. KA3 Afrodontops comes n.sp. Afrodontops Palaeostoa costellata Clausiliidae sp. KA26 n.sp. europaea Afrodontops Albinulopsis gibba Zuella venusta Acanthinula Ovicarychium ? sp. KA34 Acroloxus korys n.sp. Acroloxus Sphaeriidae sp. KA15 Acroloxus aspis n.sp. Acroloxus Ovicarychium ronlederi Turricarychium muelleri Turricarychium Ovicarychium hennigeri Carychium bachi n.sp. Biomphalaria Carychium vagum n.sp. Physidae sp. KA30 Cochlostoma sp. KA16 Megalomastomatinae? sp. KA14 Galba sp. KA1 Cyclophoroidea sp. KA34 Taxon

Distribution of mollusc species in samples. Estimated frequencies for numerical evaluations: + = 1. ++ 3. ++++ 30. Estimated frequencies species in samples. Distribution of mollusc Bewertungen: + = 1. ++ 3. ++++ 30. Probe. Geschätzte Häufgkeiten für numerische Molluskenvorkommen per

Terrestrial 1. 1. water Fresh

group Ecological Tabelle Table

52 GEOLOGICA SAXONICA — 65: 2019

Fig. 3. Stratigraphic position of the Karsdorf fauna in relation to the Paris Basin succession, to the European Land Mammal Ages (ELMA) and mammal zones, to the nannoplankton zonation, the chronostratigraphy and the absolute ages. First six columns adapted from Ogg & Gradstein (2017), the seventh after Aubry et al. (2005), BiochroM’97 (1997), Duprat (1997), Hooker (1996), King (2016), Legendre & Lévêque (1997), Mégnien et al. (1980), and Schmidt-Kittler (1987). Arrow in right-hand column indicates the possible age range of the Karsdorf mollusc fauna. Abb. 3. Stratigraphische Position der Karsdorfer Fauna in Bezug auf die Abfolge des Pariser Beckens, die Europäischen Landsäuger- alter (ELMA) und Säugerzonen, die Nannoplanktonzonierung, Chronostratigraphie und absolute Alter. Die ersten 6 Spalten nach Ogg & Gradstein (2017), die 7. nach Aubry et al. (2005), BiochroM’97 (1997), Duprat (1997), Hooker (1996), King (2016), Legendre & Lévêque (1997), Mégnien et al. (1980), und Schmidt-Kittler (1987). Der Pfeil in der rechten Spalte gibt den möglichen Altersbereich der Karsdorfer Molluskenfauna an. species most likely being accidentally picked up and P. elongata and P. exarata, and the late Ypresian P. cf. transported by waterfowl. When the karst fssure opened, fontenayi from Grauves. P. exarata has also been record- the pond sediments in which also the terrestrial fossils ed from the early Ypresian Epernay Formation of Mont had accumulated, slid into it. Bernon (Wenz 1923: 740), but in the absence of voucher material and the possibility of confusion with the similar P. costellata or P. exarata this record is shown as Pal Biostratigraphy aeostoa sp. in Fig. 2 and omitted from further considera- tions. Four species of land snails are conspecifc with, or are While the frst three species suggest a close stratigra closely related to species known from the Paris Basin; phic relationship with the fauna from Grauves, Palae see Fig. 3 for the age and stratigraphic origin of the men- ostoa costellata is closer to the Thanetian P. exarata tioned species: and P. elongata, suggesting an age of the Karsdorf fs- Carychium vagum occurs also in Grauves (Marnes à sure fauna between the levels of Rilly-la-Montagne and Unios and Térédines, late Ypresian, mammal zone MP10). Grauves. The stratigraphic assessment is possibly biased Carychium bachi is closely related to Carychium sp. towards the level of Grauves because of much richer ma- aff. bachi from Grauves. terials were available from Grauves than from the other Albinulopsis gibba n.sp. is closely related to Albinu important land snail bearing locality Mont Bernon near lopsis bonneti (Cossmann 1907) from Grauves. Epernay. Thus, an age as old as the level of Mont Ber- Palaeostoa costellata n.sp. The shell characters of non can currently not be excluded, although it cannot be this species are intermediate between the Thanetian proven either.

53 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

Table 2. Comparison of Geiseltal (Lutetian) mollusc species with putative “equivalent” species from Karsdorf (Ypresian). The term “equivalent species” is here used to comprise identical as well as closely related species. The Geiseltal species are not revised, and their nomenclature is that of Krumbiegel (1962). The earlier assumption (Henniger et al. 2011) of close faunal similarities between Karsdorf and Geiseltal is not confrmed in this study. Tabelle 2. Vergleich von Mollusken des Geiseltales (Lutetium) mit vermuteten „äquivalenten“ Arten con Karsdorf (Ypresium). Der Aus- druck „äquivalente Arten“ schließt hier identische und nahe verwandte Arten ein. Die Arten des Geiseltals sind nicht revidiert; Krumbiegels Nomenklatur (1962) wird benutzt. Die bisherige Annahme (Henniger et al. (2011)) enger faunistischer Beziehungen zwischen Karsdorf und Geiseltal wird in dieser Studie nicht bestätigt.

Geiseltal Karsdorf Krumbiegel 1962 Henniger et al. 2011 this study Biomphalaria sp. KA2 Australorbis pseudoammonius pseudoammonius Australorbis cf. pseudo-ammonius Afrodontops europaea n.sp. Galba (Galba) aquensis michelini Galba cf. aquensis Galba sp. KA1 Ancylus dutemplei Ancylus cf. dutemplei Acroloxus aspis n.sp. Carychiopsis carbonaria Carychiopsis cf. carbonaria Ovicarychium hennigeri n.sp. Carychiopsis sp. 1 Turricarychium muelleri n.sp. Carychium bachi n.sp. Carychiopsis sp. 2 Turricarychium muelleri n.sp. Vertiginidae? gen.indet. sp. 1 Albinulopsis gibba n.sp. Vertiginidae? gen.indet. sp. 2 Palaeostoa costellata n.sp. Archaegopis (Phacozonites) ceciliae Archaegopis sp. Helicoid land snail sp. KA 3

Henniger et al. (2011) assumed a Middle Eocene While some differences between these assemblage (Lutetian) age of the Karsdorf fossil lagerstätte. This is may be due to ecological conditions, the degree of dif- mainly based on mollusc species supposedly in common ferences between the faunal assemblages of the Geiseltal with or closely related to those of the Lutetian Geiseltal Formation and of Karsdorf is consistent with a sizeable Formation. Table 2 summarizes the species reported by biostratigraphical difference as discussed above. Consist- Krumbiegel (1962) from the Geiseltal Formation which ent with a younger age of the Geiseltal fauna is the fact, were supposed to be related to species from Karsdorf. that most of the genera listed as present in Geiseltal but The nomenclature of the Geiseltal molluscs is unrevised not Karsdorf continue in Europe into the present or at from Krumbiegel (1962), and is for the Karsdorf mol- least the Neogene: Gyraulus, Acicula, Strobilops, Val luscs that of Henniger et al. (2011) and of this study. lonia, Palaeoglandina, giving a much younger aspect to Without a re-examination of the Geiseltal mollusc the Geiseltal fauna than to the Karsdorf fauna. fauna, its exact relationship to the Karsdorf fauna cannot be reliably assessed. The Lutetian age of the Geiseltal Formation is reliably based on mammal biostratigraphy Timing of the structural development and palynostratigraphy. Likewise the early to middle Ypresian age range of the Karsdorf mollusc fauna is well The new dating of the fssure infll facilitates some preci- established by the molluscs Carychium vagum, C. bachi, sions of the tectonic history. While the fssure was generated Albinulopsis gibba and Palaeostoa costellata, of which by the updoming of the Querfurt Plateau, the pond which no “equivalents” are known from Geiseltal. Also sig- supplied the infll sediments needed a high groundwater nifcant is that the Palaeostoa species from Karsdorf is level, i.e.it was formed in a relatively low-lying area, which closer to P. exarata than to P. cf. fontenayi from Grauves would have been the case before updoming. The Kars- and the Lutetian P. fontenayi s.str . dorf mollusc fauna is older than the fauna from Grauves, Species which may be identical with or closely relat- i.e older than mammal zone MP10. Sedimentation in the ed to Geiseltal species cannot reliably be identifed until Geiseltal Basin began in the (late?) Grauvian (late Ypre- the Geiseltal fauna is restudied. To date this author has sian, mammal zone MP10) and continued up to mammal only examined Ovicarychium carbonarium (Wenz 1932) zone MP14 (Haubold & Hellmund 1997), suggesting the and two smaller carychiine species which are all distinct salt withdrawal from the Geiseltal area and updoming of from the Karsdorf carychiines. the Querfurt Plateau began in the late Ypresian, leading to Signifcant differences exist also in the overall composi- the opening of the Karsdorf fssure causing the destruction tion of the faunal assemblages: In Karsdorf the following of the pond and slide of its sediments into this fssure. genera or (sub)families are present, which are absent in the Geiseltal Formation: Sphaeriidae, Megaloma sto ma tinae, Cochlostoma, Turricarychium, Zuella, Albinulopsis, Afro Palaeobiogeography dontops, Palaeostoa. Conversely, the genera Gyraulus, Ac icula, Strobilops, Vallonia, Cylindrovertilla?, Phacozonites, On a regional scale, the faunal affnities with the Paris Palaeoglandina, Canalicia? are present in the Geiseltal Basin are consistent with the presence of a continuous Formation, but lack in Karsdorf (generic assignments of land mass between Karsdorf and the Paris Basin (e.g., Geiseltal taxa quoted from Krumbiegel 1962, 1963). Ziegler 1990, Gibbard & Lewin 2016).

54 GEOLOGICA SAXONICA — 65: 2019

On a global scale, the biogeographical characteristics ly Barremian (e.g. Vidal et al. 2003, Assine et al. 2008), of the families are as follows: Amongst the freshwater but a land connection persisted in the Equatorial Atlantic molluscs the Lymnaeidae, Planorbidae, Physidae and Shear Zone until the Aptian (e.g. Sapin et al. 2016). Sphaeriidae occur worldwide and are present throughout the Cenozoic in Europe and elsewhere. The Acroloxidae are only palearctic and also present in Europe throughout Acknowledgments the Cenozoic.

The biogeographic affnities of the landsnails can be I wish to express my sincere gratitude to the members of the Insti- characterized as follows: Megalomastomatinae are di- tut für Geophysik und Geologie of Leipzig University, who carried out the sampling, washing and most of the picking of the fssure verse in the Paleogene of Europe, but are now living in infll: Prof. Arnold Müller, Dr. Ronny M. Leder and Dr. Matthias the Caribbean. Henniger; for granting access to reference material I thank Jon Ab- Cochlostomatinae are a palearctic group through to the lett (NHMUK-Malacology), Dr. Ronald Janssen (SMF), Prof. Mi- chael Rasser (SMNS), Dr. Fritz Geller-Grimm (MUWI), Dr. Oliver present. Wings (ZNSGH) and Jean-Michel Pacaud (MNHN). I am particu- Carychiinae: the Paleogene diversity in Europe points to larly grateful to Prof. A. Müller for encouraging this study, placing a palearctic origin. the material at my disposal and help with Textfgure 1. 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55 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

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(2015): Nichtmarine und perihaline Molluskenfaunen 12(11): 3041 – 3056. im Tertiär Europas, 11. Eine Molluskenfauna aus der Unteren Deshayes, G. P. (1824-1837): Description des coquilles fossiles Bunten Molasse des Alpenvorlandes (Bayern, Oligozän): Die des environs de Paris., 2 [Gastéropodes]: 1824: 1 – 178; 1833: Fundstelle Habach 5. – Geologica Saxonica, 61(1): 53 – 72. 179 – 306; 1834: 307 – 434; 1835: 435 – 562; 1836: 563 – 690; King, C. (edited posthumously by Gale, A. S.& Barry, T. L.) (2016): 1837a: 691 – 814; 1837b: Atlas: 51 pp., 106 pl. Paris (Deshayes; A revised correlation of Tertiary rocks in the British Isles and Béchet Jeune; Baudouin; Treuttel & Wurtz). adjacent areas of NW Europe. Geological Society Special Re- Deshayes, G. P. (1861 – 1863): Description des animaux sans vertè port, 27: iii+719 pp. bres découverts dans le bassin de Paris, 2 (Livr. 21, 22: 1 – 120, Krumbiegel, G. 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(1885): On the land Mollusca of the Eocenes [sic]. – chroM ’97. Mémoires et Travaux de l’Iinstitut de Montpellier. The Geological Magazine, N.S., (3) 2 (VI): 241 – 251, pl. 6. École Pratique des Hautes Études [also given as “Mém. Trav. Gibbard, P. L. & Lewin, J. (2016): Filling the North Sea Basin: Ce- E.P.H.E., Inst. Montpellier”], 21: 461 – 473. Montpellier. nozoic sediment sources and river styles. – Geologica Belgica, Manganelli, G. & Giusti, F. (2000): The gastrocoptids of the Fos- 19(3 – 4): 201 – 217. sil Forest of Dunarobba (Central Italy). With a preliminary re- Gray, J. E. (1847): A list of the genera of recent mollusca, their vision of the European Neogene nominal species of Albinula synonyma and their types. – Proceedings of the Zoological So- and Vertiginopsis (Gastropoda Pulmonata: Gastrocoptidae). – ciety of London, 15 (179): 129 – 219. Bollettino della Società Paleontologica Italiana, 39(1): 55 – 82. Hammouda, S. A., Kadolsky, D., Adaci, M., Mebrouk, F., Bensalah, Mathéron, P. (1842 – 1843): Catalogue méthodique et descriptif M., Mahboubi, M. & Tabuce, R. (2017): Taxonomic review of des corps organisés fossiles du Département des Bouches-du- the “Bulimes”, terrestrial gastropods from the continental Eo- Rhône et lieux circonvoisins; précédé d’un mémoire sur les ter- cene of the Hamada de Méridja (northwestern Sahara, Alge- rains supérieurs au Grès Bigarré du S.E.de la France [Preprint ria) (Mollusca: Stylommatophora: Strophocheilidae?), with a from:] Répertoire des Travaux de la Société de Statistique de

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Marseille, 6: Livr. 1: 1 – 95, pls. 1 – 13 (1842); Livr. 2: 96 – 269, Schileyko, A. A. (2001): Treatise on recent terrestrial pulmonate pls. 14 – 41 (1843, post May). Marseille. molluscs: Ruthenica, Supplement, 2(7): 881 – 1034. Moscow. Mégnien, F., Barta, L., Laurain, M., Leplat, J., Pomerol, B. & Wyns, Schmidt-Kittler, N. E. (1987a): International symposium on mam- R. (1980): Paléocène. – In: Mégnien, C.& Mégnien, F. (co- malian biostratigraphy and paleoecology of the European ord.): Synthèse géologique du bassin de Paris. 1. Stratigraphie Paleogene. – Münchner Geowissenschaftliche Abhandlungen, et paléogéographie. Mémoire du BRGM, 101: 337 – 350. Series A, 10: 1 – 311. Menning, M. & Hendrich, A. (eds.; with contributions by authors’ Schnabel, T. (2006a): Die känozoischen Filholiidae Wenz 1923. collective) (2016): Stratigraphische Tabelle von Deutschland Teil 2: Die pliozänen Triptychien, nebst Bemerkungen zum 2016: Table 100 × 141 cm. Potsdam (Deutsche Stratigraphische Typusmaterial und zur systematischen Stellung der Filholiidae Kommission). (Gastropoda: Pulmonata: Clausilioidea). – Archiv für Mollus Morgan, J. A. T., DeJong, R. J., Jung, Y., Khallaayoune, K., Kock, kenkunde, 135(1): 23 – 47. S., Mkoji, G. M. & Loker, E. S. (2002): A phylogeny of planor- Schnabel, T. (2006b): Die känozoischen Filholiidae Wenz 1923. bid snails, with implications for the evolution of Schistosoma Teil 3: Die miozänen Vertreter der Gattung Triptychia (Gas- parasites. – Molecular Phylogenetics and Evolution, 25(3): tropoda, Pulmonata, Clausilioidea). – Archiv für Mollusken- 477 – 488. kunde, 135 (2): 133 – 203. Newton, R. B. (1891): Systematic list of the Frederick E. Edwards Schnabel, T. (2007): Die känozoischen Filholiidae Wenz 1923. 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C. (1981): The occurrence of Proserpina in the British Ziegler, P. A. (1990): Geological Atlas of Western and Central Eu- Tertiary, with the description of a new species. – Archiv für rope 1990. Second and completely revised edition. – Interna- Molluskenkunde, 111(1/3): 49 – 54. tional Lithosphere Program Publication, 148: 239 pp., 56 encls. Risso, A. (1826): Histoire naturelle des principales productions de (Shell Internationale Petroleum Maatschappij BV) [ISBN 90- l’Europe méridionale et particulièrement de celles des environs 6644-125-9]. de Nice et des Alpes Maritimes, 4: 438 pp., 12 pls. Paris (Lev- Zilch, A. (1959 – 1960): Gastropoda. Euthyneura. – In: Schinde rault). wolf (Ed.): Handbuch der Paläozoologie, 6(2): 1: 1 – 200 Salvador, R. B., Cabrera, F., Martinez, S., Miquel, S. E., Simone, L. (17.6.1959); 2: 201 – 400 (25.11.1959); 3: 401 – 600 (30.3. R. L. & Cunha, C. M. (2018): Annotated catalogue of the fos- 1960); 4: 601 – 835, I – XII (15.8.1960). 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57 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

Plate legends The majority of fgured specimens is illustrated as both drawings and photos. Plates showing the same specimens are given the same plate number, with the suffx A denoting plates with drawings, and the suffx B denoting plates with photos. The origin of a specimen is the Early Eocene fssure infll in Karsdorf quarry, unless otherwise stated.

Tafelerklärungen Die Mehrzahl der abgebildeten Exemplare ist sowohl als Zeichnung als auch als Foto dargestellt. Tafeln, die die gleichen Exemplare zeigen, haben die gleiche Nummer, wobei ein angehängtes A Tafeln mit Zeichnungen, und ein angehängtes B Tafeln mit Fotos bezeichnet. Der Fundort der Exemplare ist die früheozäne Spaltenfüllung im Steinbruch Karsdorf, wenn nicht anders angegeben..

Plates 1A (drawings) and 1B (photos). Fig. 1: Galba sp. KA1. Composite sample. GPSL TM 0247. Figured by Henniger et al. (2011) (: Pl. 3, Fig. 11, as Galba cf. aquensis ssp.) Fig. 2 (Pl. 1B only): Galba sp. KA1. Composite sample. GPSL TM 0248. Fig. 3 (Pl. 1A only): Copy of original fgure of Lymnaea aquensis Mathéron (1843: 215, Pl. 36, Fig. 7) from „quartiers de Montaiguet“ (France, Département Bouches-du-Rhône; Lutetian) with the outline of the specimen of Galba sp. KA1 illustrated in Fig. 1 inset at the same scale. Height : width of Mathéron’s fgure: 40 : 15 mm. Fig. 4: Biomphalaria sp. KA2. Composite sample. GPSL TM 0256. Fig. 5: Biomphalaria sp. KA2. Sample KD 806. GPSL TM 0257. Fig. 6: Biomphalaria sp. KA2. Sample KD 806. GPSL TM 0258. Fig. 7: Physidae sp. KA30. Sample KD 2328. GPSL TM 0265.

Tafeln 1A (Zeichnungen) und 1B (Fotos). Fig. 1: Galba sp. KA1. Sammelprobe. GPSL TM 0247. Abgebildet von Henniger et al. (2011: Taf. 3 Fig. 11, als Galba cf. aquensis ssp.) Fig. 2 (nur Taf. 1B): Galba sp. KA1. Sammelprobe. GPSL TM 0248. Fig. 3 (nur Taf. 1A): Kopie der Originalfgur von Lymnaea aquensis Mathéron (1843: Taf. 36, Fig. 7) von „quartiers de Montaiguet“ (Frankreich, Département Bouches-du-Rhône; Lutetium) mit dem Umriss des Exemplars von Galba sp. KA1, hier abgebildet als Fig. 1, einkopiert im gleichen Maßstab. Höhe : Breite von Mathérons Abbildung: 40 : 15 mm. Fig. 4: Biomphalaria sp. KA2. Sammelprobe. GPSL TM 0256. Fig. 5: Biomphalaria sp. KA2. Probe KD 806. GPSL TM 0257. Fig. 6: Biomphalaria sp. KA2. Probe KD 806. GPSL TM 0258. Fig. 7: Physidae sp. KA30. Probe KD 2328. GPSL TM 0265.

Plates 2A (drawings) and 2B (photos). Fig. 1: Acroloxus aspis n. sp. Holotype. Composite sample. Ypresian. GPSL TM 0266. Fig. 2: Acroloxus aspis n. sp. Paratype. Sample 2382. Ypresian. GPSL TM 0267. Fig. 3: Acroloxus korys n. sp. Holotype. Sample KD 2382. GPSL TM 0269. Fig. 4: Acroloxus korys n. sp. Paratype. Sample KD 2382. GPSL TM 0270. Fig. 5: Sphaeriidae sp. KA15. Sample KD 2328. GPSL TM 0272. Fig. 6: Sphaeriidae sp. KA15. Sample KD 2382. GPSL TM 0273. Fig. 7: Sphaeriidae sp. KA15. Composite sample. GPSL TM 0274. Fig. 8: Sphaeriidae sp. KA15. Sample KD 2382. GPSL TM 0275. Fig. 9: Megalomastomatinae? sp. KA14. Composite sample. GPSL TM 0278.

Tafeln 2A (Zeichnungen) und 2B (Fotos). Fig. 1: Acroloxus aspis n. sp. Holotypus. Sammelprobe. Ypresium. GPSL TM 0266. Fig. 2: Acroloxus aspis n. sp. Paratypus. Probe 2382. Ypresium. GPSL TM 0267. Fig. 3: Acroloxus korys n. sp. Holotypus. Probe KD 2382. GPSL TM 0269. Fig. 4: Acroloxus korys n. sp. Paratypus. Probe KD 2382. GPSL TM 0270. Fig. 5: Sphaeriidae sp. KA15. Probe KD 2328. GPSL TM 0272. Fig. 6: Sphaeriidae sp. KA15. Probe KD 2382. GPSL TM 0273. Fig. 7: Sphaeriidae sp. KA15. Sammelprobe. GPSL TM 0274. Fig. 8: Sphaeriidae sp. KA15. Probe KD 2382. GPSL TM 0275. Fig. 9: Megalomastomatinae? sp. KA14. Sammelprobe. GPSL TM 0278.

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Plates 3A (drawings) and 3B (photos). Fig. 1: Cochlostoma? sp. KA16. Sample KD 803. GPSL TM 0279. Fig. 2: Cochlostoma? sp. KA16. Sample KD 803. GPSL TM 0280. Fig. 3: Cyclophoroidea sp. KA31. Sample KD 803. Specimen lost before photos were made; ex GPSL. Fig. 4: Carychium vagum n.sp. Holotype. Composite sample. GPSL TM 0283. Fig. 5: Carychium vagum n.sp. Paratype. Grauves (France, Departement Marne). Late Ypresian. Cossmann collection, MNHN F.J17530. Fig. 6 (Plate 3A only): Carychium hypermeces Cossmann 1889. Copy of Cossmann & Pissarro 1913: pl. 57, fg. 255-5 which is a copy of the original fgure by Cossmann (1889: pl. 11 fg. 33). Scale refers to the height measurement given originally by Cossmann (1889): 1.25 mm. The scale given by Cossmann & Pissarro (1913) implies a height of 1.83 mm, however. Fig. 7: Carychium vagum n.sp. Paratype. Grauves (France, Departement Marne). Late Ypresian. Cossmann collection, MNHN F.J17534. Fig. 8: Carychium vagum n.sp. Paratype. Grauves (France, Departement Marne). Late Ypresian. Cossmann collection, MNHN F.J17531. On Plate 3A the outline of the holotype of Carychium vagum (Fig. 4) is superposed in blue. Fig. 9: Carychium sp. aff. bachi n.sp. Grauves (France, Departement Marne). Late Ypresian. Cossmann collection, MNHN F.J17529. On Plate 3A the outline of the holotype of Carychium bachi is superposed in blue Fig. 10: Carychium bachi n.sp. Holotype. Composite sample. GPSL TM 0287. Fig. 11: Carychium bachi n.sp. Paratype. Composite sample. GPSL TM 0288.

Tafeln 3A (Zeichnungen) und 3B (Fotos). Fig. 1: Cochlostoma? sp. KA16. Probe KD 803. GPSL TM 0279. Fig. 2: Cochlostoma? sp. KA16. Probe KD 803. GPSL TM 0280. Fig. 3: Cyclophoroidea sp. KA31. Probe KD 803. Exemplar verloren, bevor Fotos gemacht werden konnten; ex GPSL. Fig. 4: Carychium vagum n.sp. Holotypus. Sammelprobe. GPSL TM 0283. Fig. 5: Carychium vagum n.sp. Paratypus. Grauves (France, Departement Marne). Spätes Ypresium. Sammlung Cossmann, MNHN F.J17530. Fig. 6 (nur Tafel 3A): Carychium hypermeces Cossmann 1889. Kopie von Cossmann & Pissarro 1913: Taf. 57, Fig. 255-5, die eine Kopie der Originalabbildung Cossmanns ist (1889: Taf. 11 Fig. 33). Der Maßstab basiert auf der Höhenangabe Cossmanns (1889): 1.25 mm. Der von Cossmann & Pissarro (1913) gegebene Maßstab impliziert jedoch eine Höhe von 1.83 mm. Fig. 7: Carychium vagum n.sp. Paratypus. Grauves (France, Departement Marne). Spätes Ypresium. Sammlung Cossmann, MNHN F.J17534. Fig. 8: Carychium vagum n.sp. Paratypus. Grauves (France, Departement Marne). Spätes Ypresium. Sammlung Cossmann, MNHN F.J17531. In Tafel 3A der Umriss des Holotyps von Carychium vagum (Fig. 4) ist in Blau überlegt. Fig. 9: Carychium sp. aff. bachi n.sp. Grauves (France, Departement Marne). Spätes Ypresium. Sammlung Cossmann, MNHN F.J17529. Auf Tafel 3A ist der Umriss des Holotyps von Carychium bachi (Fig. 10) in Blau überlegt. Fig. 10: Carychium bachi n.sp. Holotypus. Sammelprobe. GPSL TM 0287. Fig. 11: Carychium bachi n.sp. Paratypus. Sammelprobe. GPSL TM 0288.

Plates 4A (drawings) and 4B (photos). Fig. 1: Turricarychium muelleri n.sp. Holotype. Composite sample. GPSL TM 0294. Fig. 2: Turricarychium muelleri n.sp. Paratype. Composite sample. GPSL TM 0295. Fig. 3: Turricarychium muelleri n.sp. Paratype. Composite sample. GPSL TM 0296. Fig. 4: Ovicarychium hennigeri n.sp. Holotype. Composite sample. GPSL TM 0305. Fig. 5: Ovicarychium hennigeri n.sp. Paratype. Sample KD 805. GPSL TM 0306. Fig. 6: Ovicarychium sp. KA34. Contours of the holotype of O. ronlederi n.sp. overlain. Sample KD 2382. GPSL TM 0320. Fig. 7: Ovicarychium carbonarium (Wenz 1932). Lectotype. Sachsen-Anhalt, Geiseltal area, lignite open cast mine Cecilie, Trichter CÏ I (lower level). Lutetian. Geiseltal-Museum Halle/S. G1926/C1 typ. Fig. 8: Ovicarychium ronlederi n.sp. Holotype. Composite sample. GPSL TM 0314. Fig. 9: Ovicarychium ronlederi n.sp. Paratype. Composite sample. GPSL TM 0315. Fig. 10: Zuella venusta n.sp. Holotype. Composite sample. GPSL TM 0323. Fig. 11: Zuella venusta n.sp. Paratype. Composite sample. GPSL TM 0324. Fig. 12: Acanthinula (s.l.) karsdorfensis n.sp. Holotype. Sample KD 2382. GPSL TM 0329.

Tafeln 4A (Zeichnungen) und 4B (Fotos). Fig. 1: Turricarychium muelleri n.sp. Holotypus. Sammelprobe. GPSL TM 0294. Fig. 2: Turricarychium muelleri n.sp. Paratypus. Sammelprobe. GPSL TM 0295. Fig. 3: Turricarychium muelleri n.sp. Paratypus. Sammelprobe. GPSL TM 0296. Fig. 4: Ovicarychium hennigeri n.sp. Holotypus. Sammelprobe. GPSL TM 0305. Fig. 5: Ovicarychium hennigeri n.sp. Paratypus. Probe KD 805. GPSL TM 0306. Fig. 6: Ovicarychium sp. KA34. Umriss des Holotypus von O. ronlederi n.sp. einkopiert. Probe KD 2382. GPSL TM 0320. Fig. 7: Ovicarychium carbonarium (Wenz 1932). Lectotypus. Sachsen-Anhalt, Geiseltalgebiet, Braunkohlentagebau Cecilie, Trichter CÏ I (unteres Niveau). Lutetium. Geiseltal-Museum Halle/S. G1926/C1 typ. Fig. 8: Ovicarychium ronlederi n.sp. Holotypus. Sammelprobe. GPSL TM 0314. Fig. 9: Ovicarychium ronlederi n.sp. Paratypus. Sammelprobe. GPSL TM 0315. Fig. 10: Zuella venusta n.sp. Holotypus. Sammelprobe. GPSL TM 0323. Fig. 11: Zuella venusta n.sp. Paratypus. Sammelprobe. GPSL TM 0324. Fig. 12: Acanthinula (s.l.) karsdorfensis n.sp. Holotypus. Probe KD 2382. GPSL TM 0329.

59 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

Plates 5A (drawings) and 5B (photos). Fig. 1: Albinulopsis gibba n.sp. Holotype. Composite sample. GPSL TM 0330. Figured by Henniger et al. (2011: Fig. 4.7a – 7d, as Ver- tiginidae? gen. indet., sp. 1) Fig. 2: Albinulopsis gibba n.sp. Paratype. Sample KD 2365. GPSL TM 0331. Fig. 3: Albinulopsis gibba n.sp. Paratype. Sample KD 2382. GPSL TM 0332. Fig. 4: Albinulopsis bonneti (Cossmann 1907). Grauves (Département Marne). Late Ypresian. Cossmann collection, MNHN J 17537. Late Ypresian. Fig. 5 ( Pl. 5B only): Albinulopsis bonneti (Cossmann 1907). Grauves (Département Marne). Late Ypresian. Cossmann collection, MNHN J17538. Late Ypresian. Fig. 6: Clausiliidae sp. KA26. Sample KD 803. GPSL TM 0186. Fig. 7: Afrodontops europaea n.sp. Paratype. Composite sample. GPSL TM 0190. Fig. 8: Afrodontops europaea n.sp. Holotype. Sample KD 2382. GPSL TM 0189. Fig. 9: Afrodontops europaea n.sp. Paratype. Composite sample. GPSL TM 0191. Figured by Henniger et al. (2011:326, pl. 3, fg. 14, as juvenile Australorbis aff. pseudoammonius). Fig. 10: Afrodontops europaea n.sp. Paratype. Composite sample. GPSL TM 0192. Fig. 11: Afrodontops comes n.sp. Holotype. Sample KD 2379. GPSL TM 0199.

Tafeln 5A (Zeichnungen) und 5B (Fotos). Fig. 1: Albinulopsis gibba n.sp. Holotypus. Sammelprobe. GPSL TM 0330. Abgebildet bei Henniger et al. (2011: Fig. 4.7a – 7d, als Ver- tiginidae? gen. indet., sp. 1) Fig. 2: Albinulopsis gibba n.sp. Paratypus. Probe KD 2365. GPSL TM 0331. Fig. 3: Albinulopsis gibba n.sp. Paratypus. Probe KD 2382. GPSL TM 0332. Fig. 4: Albinulopsis bonneti (Cossmann 1907). Grauves (Département Marne). Spätes Ypresium. Sammlung Cossmann, MNHN J 17537. Fig. 5 ( nur Taf. 5B): Albinulopsis bonneti (Cossmann 1907). Grauves (Département Marne). Spätes Ypresium. Sammlung Cossmann, MNHN J17538. Fig. 6: Clausiliidae sp. KA26. Probe KD 803. GPSL TM 0186. Fig. 7: Afrodontops europaea n.sp. Paratypus. Sammelprobe. GPSL TM 0190. Fig. 8: Afrodontops europaea n.sp. Holotypus. Probe KD 2382. GPSL TM 0189. Fig. 9: Afrodontops europaea n.sp. Paratypus. Sammelprobe. GPSL TM 0191. Figured by Henniger et al. (2011: Taf. 3, Fig. 14, als juve- niler Australorbis aff. pseudoammonius). Fig. 10: Afrodontops europaea n.sp. Paratypus. Sammelprobe. GPSL TM 0192. Fig. 11: Afrodontops comes n.sp. Holotypus. Probe KD 2379. GPSL TM 0199.

Plate 6 (photos). Fig. 1: Afrodonta bilamellaris (Melvill & Ponsonby 1908). Lectotype. Natal: Edendale Falls. Recent. NHMUK 1908.12.14.54. Fig. 2: Palaeostoa costellata n.sp. Holotype. Composite sample. GPSL TM 0200. Figured by Henniger et al. 2011 (: 326, Fig. 4.8a – 8d, as Vertiginidae?, gen. indet., sp. 2). Fig. 3: Palaeostoa costellata n.sp. Paratype. Composite sample. GPSL TM 0201. This fragment is fgured to show that the palatal lirae were already developed in the earliest teleoconch whorls.

Tafel 6 (Fotos). Fig. 1: Afrodonta bilamellaris (Melvill & Ponsonby 1908). Lectotypus. Natal: Edendale Falls. Rezent. NHMUK 1908.12.14.54. Fig. 2: Palaeostoa costellata n.sp. Holotypus. Sammelprobe. GPSL TM 0200. Abgebildet von Henniger et al. (2011: Fig. 4.8a – 8d, als Vertiginidae?, gen. indet., sp. 2). Fig. 3: Palaeostoa costellata n.sp. Paratypus. Sammelprobe. GPSL TM 0201. Dieses Fragment ist abgebildet, um zu zeigen, dass die pa- latalen Lirae bereits in den frühesten Teleoconchumgängen vorhanden waren.

Plate 7 (photos). Fig. 1: Palaeostoa costellata n.sp. Paratype. Sample KD 803. GPSL TM 0202. Fig. 2: Palaeostoa costellata n.sp. Paratype. Composite sample. GPSL TM 0203. Fig. 3: Palaeostoa costellata n.sp. Paratype. Sample KD 803. GPSL TM 0204. Fig. 4: Palaeostoa costellata n.sp. Paratype. Sample KD 803. GPSL TM 0205. Fig. 5: Palaeostoa costellata n.sp. Paratype. Sample KD 803. GPSL TM 0206. Fig. 6: Palaeostoa costellata n.sp. Paratype. Sample KD 807. GPSL TM 0207.

Tafel 7 (Fotos). Fig. 1: Palaeostoa costellata n.sp. Paratypus. Probe KD 803. GPSL TM 0202. Fig. 2: Palaeostoa costellata n.sp. Paratypus. Sammelprobe. GPSL TM 0203. Fig. 3: Palaeostoa costellata n.sp. Paratypus. Probe KD 803. GPSL TM 0204. Fig. 4: Palaeostoa costellata n.sp. Paratypus. Probe KD 803. GPSL TM 0205. Fig. 5: Palaeostoa costellata n.sp. Paratypus. Probe KD 803. GPSL TM 0206. Fig. 6: Palaeostoa costellata n.sp. Paratypus. Probe KD 807. GPSL TM 0207.

60 GEOLOGICA SAXONICA — 65: 2019

Plate 8A (drawings) and 8B (photos). Fig. 1: Palaeostoa elongata (Melleville 1843). Département Marne: Châlons-sur-Vesle. Late Thanetian. Cossmann collection, MNHN F.J03383. Apical part of specimen fgured by Cossmann & Pissarro 1913 (: Pl. 60, Fig. 275-1, as Palaeostoa exarata). Fig. 2: Palaeostoa exarata (Michaud 1838). Département Marne, Rilly-la-Montagne. Late Thanetian. Jooss collection, SMNS 66.878. Fig. 3: Palaeostoa costellata n.sp. Holotype. Composite sample. GPSL TM 0200. See also Pl. 6, Fig. 2. Fig. 4: Palaeostoa cf. fontenayi (Sandberger 1871). Département Marne, Grauves. Late Ypresian. Cossmann collection, MNHN F.J17540. Fig. 5: Palaeostoa fontenayi (Sandberger 1871). Holotype of Clausilia crenata Sandberger 1871. Alsace: Bouxwiller. Bouxwiller Forma- tion, Lutetian. MUWI. Fig. 6: Helicoid land snail sp. KA3. Composite sample. GPSL TM 0215. Figured by Henniger et al. (2011: Pl. 3, Fig. 8, as Archaegopis sp.). Fig. 7: Helicoid land snail sp. KA12. Composite sample. GPSL TM 0216. Fig. 8: Helicoid land snail sp. KA12. Composite sample. GPSL TM 0217. Fig. 9: Helicoid land snail sp. KA12. Sample KD 803. GPSL TM 0218. Fig. 10: Land snail sp. KA19. Composite sample. GPSL TM 01219. Fig. 11: Helicoid land snail sp. KA25. Composite sample. GPSL TM 0221. Fig. 12: Elongate sinistral land snail sp. KA29. Sample KD 2365. GPSL TM 0223. Fig. 13: Elongate sinistral land snail sp. KA29. Sample KD 2379. GPSL TM 0224. Fig. 14: Elongate sinistral land snail sp. KA37. Sample KD 2382. GPSL TM 0226.

Tafel 8A (Zeichnungen) und 8B (Fotos). Fig. 1: Palaeostoa elongata (Melleville 1843). Département Marne: Châlons-sur-Vesle. Spätes Thanetian. Sammlung Cossmann, MNHN F.J03383. Apex des von Cossmann & Pissarro (1913: Taf. 60, Fig. 275-1) als Palaeostoa exarata abgebildeten Exemplars. Fig. 2: Palaeostoa exarata (Michaud 1838). Département Marne, Rilly-la-Montagne. Spätes Thanetian. Jooss collection, SMNS 66.878. Fig. 3: Palaeostoa costellata n.sp. Holotypus. Sammelprobe. GPSL TM 0200. Siehe auch Taf. 6, Fig. 2. Fig. 4: Palaeostoa cf. fontenayi (Sandberger 1871). Département Marne, Grauves. Spätes Ypresium. Sammlung Cossmann, MNHN F.J17540. Fig. 5: Palaeostoa fontenayi (Sandberger 1871). Holotypus von Clausilia crenata Sandberger 1871. Elsass: Bouxwiller. Bouxwiller-For- mation, Lutetium. MUWI. Fig. 6: Helicoide Landschnecke sp. KA3. Sammelprobe. GPSL TM 0215. Figured by Henniger et al. (2011: Taf. 3, Fig. 8, als Archaego- pis sp.). Fig. 7: Helicoide Landschnecke sp. KA12. Sammelprobe. GPSL TM 0216. Fig. 8: Helicoide Landschnecke sp. KA12. Sammelprobe. GPSL TM 0217. Fig. 9: Helicoide Landschnecke sp. KA12. Probe KD 803. GPSL TM 0218. Fig. 10: Landschnecke sp. KA19. Sammelprobe. GPSL TM 01219. Fig. 11: Helicoide Landschnecke sp. KA25. Sammelprobe. GPSL TM 0221. Fig. 12: Elongate linksgewundene Landschnecke sp. KA29. Probe KD 2365. GPSL TM 0223. Fig. 13: Elongate linksgewundene Landschnecke sp. KA29. Probe KD 2379. GPSL TM 0224. Fig. 14: Elongate linksgewundene Landschnecke sp. KA37. Probe KD 2382. GPSL TM 0226.

Plate 9 (photos). Fig. 1: Elongate land snail sp. KA13. Composite sample. GPSL TM 0227. Fig. 2: Land snail sp. KA38. GPSL TM 0230. Fig. 3: Helicoid land snail sp. KA20. Composite sample. GPSL TM 0231. Fig. 4: Land snail sp. KA20. Composite sample. GPSL TM 0232. Fig. 5: Elongate land snail sp. KA21. Composite sample. GPSL TM 0236. Fig. 6: Elongate land snail sp. KA21. Composite sample. GPSL TM 0237. Fig. 7: Land snail sp. KA22. Composite sample. GPSL TM 0238. Fig. 8: Land snail sp. KA23. Composite sample. GPSL TM 0239. Fig. 9: Land snail sp. KA23. Composite sample. GPSL TM 0240. Fig. 10: Land snail sp. KA35. Sample KD 2365. GPSL TM 0244. Fig. 11: Elongate land snail sp. KA24. Composite sample. GPSL TM 0245. Fig. 12: Land snail sp. KA36. Sample KD 805. GPSL TM 0246.

Tafel 9 (Fotos). Fig. 1: Elongate Landschnecke sp. KA13. Sammelprobe. GPSL TM 0227. Fig. 2: Landschnecke sp. KA38. GPSL TM 0230. Fig. 3: Helicoide Landschnecke sp. KA20. Sammelprobe. GPSL TM 0231. Fig. 4: Landschnecke sp. KA20. Sammelprobe. GPSL TM 0232. Fig. 5: Elongate Landschnecke sp. KA21. Sammelprobe. GPSL TM 0236. Fig. 6: Elongate Landschnecke sp. KA21. Sammelprobe. GPSL TM 0237. Fig. 7: Landschnecke sp. KA22. Sammelprobe. GPSL TM 0238. Fig. 8: Landschnecke sp. KA23. Sammelprobe. GPSL TM 0239. Fig. 9: Landschnecke sp. KA23. Sammelprobe. GPSL TM 0240. Fig. 10: Landschnecke sp. KA35. Probe KD 2365. GPSL TM 0244. Fig. 11: Elongate Landschnecke sp. KA24. Sammelprobe. GPSL TM 0245. Fig. 12: Landschnecke sp. KA36. Probe KD 805. GPSL TM 0246.

61 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

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Plate 1A (drawings) / Tafel 1A (Zeichnungen).

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Plate 1B (photos) / Tafel 1B (Fotos).

63 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

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RV groove between ligament area P2 and margin of hinge plate 1 mm

8 P2

to umbo LV Ligament area ligament area

ligament area P2 P3 to umbo P1

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Plate 2A (drawings) / Tafel 2A (Zeichnungen).

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1 mm

6 7

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Plate 2B (photos) / Tafel 2B (Fotos).

65 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

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Plate 3A (drawings) / Tafel 3A (Zeichnungen).

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Plate 3B (photos) / Tafel 3B (Fotos).

67 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

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Plate 4A (drawings) / Tafel 4A (Zeichnungen).

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Plate 4B (photos) / Tafel 4B (Fotos).

69 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

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Columellaris Basalis

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Plate 5A (drawings) / Tafel 5A (Zeichnungen).

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4 3 2 0.5 mm

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Plate 5B (photos) / Tafel 5B (Fotos).

71 D. Kadolsky: A remarkable non-marine mollusc fauna of Early Eocene age from a fssure infll in Karsdorf quarry (Sachsen-Anhalt, Germany)

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Plate 6 (photos) / Tafel 6 (Fotos).

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Plate 7 (photos) / Tafel 7 (Fotos).

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Plate 8A (drawings) / Tafel 8A (Zeichnungen).

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Plate 8B (photos) / Tafel 8B (Fotos).

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Plate 9 (photos) / Tafel 9 (Fotos).