FASCICULI INSTITUTI GEOLOGICI HUNGARIAE
AD ILLUSTRANDAM NOTIONEM GEOLOGICAM ET PALAEONTOLOGICAM
GEOLOGICA
HUNGARICA
SERIES PALAEONTOLOGICA
FASCICULUS 56
Nonmarine mollusc fauna from the Lower and
Middle Miocene, Bakony Mts, W Hungary
by
József Kókay
BUDAPEST, 2006
© Copyright Geological Institute of Hungary (Magyar Állami Földtani Intézet), 2006
All rights reserved! Minden jog fenntartva!
Serial editor:
LÁSZLÓ KORDOS
Reviewer:
MARGIT BOHN-HAVAS
Translator:
PÁL MÜLLER
Linguistic reviewer:
PHILIP RAWLINSON
Technical editor:
OLGA PIROS
DEZSŐ SIMONYI
DTP:
ILDIKÓ TIEFENBACHER
Illustrator:
ANTAL PENTELÉNYI, ALTANCECEG VAD
Cover design:
DEZSŐ SIMONYI
The publication of this volume was sponsored by the grant No. T 14298 of the
National Research Found (OTKA) and the X. section of Earth Sciences of
the Hungarian Academy of Sciences
Published by the Geological Institute of Hungary — Kiadja a Magyar Állami Földtani Intézet
Responsible editor:
KÁROLY BREZSNYÁNSZKY
Director
HU ISSN 0374–1893
ISBN 963 671 254 9
Geologica Hungarica series Palaeontologica, fasciculus 56
3
CONTENTS
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Geological description of the studied area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lower Miocene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ottnangian . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Karpatian . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Badenian . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lower Badenian . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Middle Badenian . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Upper Badenian . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Sarmatian . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Evaluation of the studied faunas from the Bakony Mts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Palaeogeographic implications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Geological age and stratigraphy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Palaeoecological, salinity and facies questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Climatic condition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Systematic descriptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Classis GASTROPODA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia NERITIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Theodoxus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia CYCLOPHORIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Palaina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Cochlostoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia VIVIPARIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Viviparus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia VALVATIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Valvata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia POMATIASIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pomatias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia HYDROBIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Nematurella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Sellia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Microprososthenia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Martinietta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Bythinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pseudamnicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Amnicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Bithynia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Ferebithynia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Schuettemmericia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Stalioa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
7
9
9
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11
11
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17
21
21
22
23
25
27
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30
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4
JÓZSEF KÓKAY
Genus Staliopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia TRUNCATELLIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Sandbergerina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Staadtiellopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Emmericia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia ACICULIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Platyla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia MELANIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Brotia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Melanoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia MELANOPSIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Melanopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia ELLOBIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Carychium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Laemodonta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Melampus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pedipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Stolidoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Auriculastra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia LIMNAEIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Galba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Stagnicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Radix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Lymnaea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Aplexa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia PLANORBIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Anisus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Gyraulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Hippeutis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Segmentina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Planorbarius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia FERRISSIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Ferrissia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia ANCYLIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Ancylus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia ACROLOXIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pseudancylastrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia COCHLICOPIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Azeca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Cochlicopa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia VERTIGINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Negulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Acmopupa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Vertigo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia CHONGRINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Gastrocopta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia PUPILLIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pupilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Leiostyla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Argna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Enneopupa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Paracoryna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia VALLONIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Vallonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Acanthinula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Spermodea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Planogyra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
42
42
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46
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49
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50
50
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53
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58
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69
60
60
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61
61
61
61
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62
62
65
65
67
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69
70
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Nonmarine mollusc fauna from the Lower and Middle Miocene, Bakony Mts, W Hungary
Genus Strobilops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia ENIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Mastus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Ena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Napaeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia SUCCINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Succinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia ENDODONTIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Helicodiscus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Discus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia VITRINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Semilimax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia ZONITIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Vitrea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Nesovitrea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Aegopis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Retinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Aegopinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Oxychilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Daudebardia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Zonitoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Janulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia LIMACIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Limax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia MILACIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Milax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia ARIONIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Arion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia SUBULINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Fortuna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Opeas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia CLAUSILIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Triptychia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Serrulastra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Nordsieckia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Cochlodina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Canalicia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pseudidyla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Mactrogastra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Clausilia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia OLEACINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pseudoleacina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Palaeoglandina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Poiretia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia HELICIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Perforatella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Caracollina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Canariella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Galactochilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Tropidomphalus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Helicigona . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Klikia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Cepaea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Classis BIVALVIA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia DREISSENIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Dreissena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Congeria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
5
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75
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76
76
77
77
78
78
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79
79
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80
80
80
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81
82
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83
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86
86
86
87
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Familia MARGARITIFERIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Margaritifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia UNIONIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Unio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Familia SPHERIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Spherium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Pisidium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Appendix: Inventory of nonmarine Miocene molluscs from the Bakony Mts . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
95
95
95
95
97
97
97
101
107
191
Geologica Hungarica series Palaeontologica, fasciculus 56
7
INTRODUCTION
During many decades of fieldwork in the Bakony Mts, thousands of core drillings were studied, described and sampled. I also visited and surveyed the sequence of many outcrops and shafts and open air mines. As it is clear from the list
of my publications, my interest was mainly focused on Miocene stratigraphy and malacology. During this multiannual
work I encountered many outcrops with nonmarine (terrestrial and freshwater) molluscs. Such faunae are sporadic, similary to vertebrate ones, but they are of major significance for stratigraphic, evolutionary and palaeographical as well as
for climatological or palaeoenvironmental studies. So I descided to elaborate this nonmarine Miocene fauna, because
from the Carpathian Basin only a few sporadic findings of this sort had been described. From the studied area, earlier
only a few nonmarine fauna had been previously desribed (WENZ 1931; BODA 1959; KECSKEMÉTI-KÖRMENDY 1962;
KÓKAY 1966, 1967, 1992)
Determination of this enormous quantity of material required the consultation of a large number of publications.
Generally, one major site yielded about two or three dozens of terrestrial and/or freshwater taxa. In marine or brackish
fauna there frequently occur some sporadic specimens of nonmarine origin, washed in from the shoreline. Consequently,
in many publications, taxa having different origins may appear.
Names and the meaning of the individual stages and substages were used in accordance with the RCMNS decisions.
251 gastropod and 21 bivalve taxa have been determined. From these taxa the Ottnangian yielded 13, the Karpatian 4,
the Lower Miocene 59, the Lower Badenian 60, the Middle Badenian 19, the Upper Badenian 116, while the Sarmatian
141 taxa. Out of the total number of 272, 117 species were aquatic, while 155 terrestrial gastropods were determined. As
an example, it is can be stated that from the studied faunas 150 taxa are known from the North Alpine (brackish-)
Freshwater Molasse. This fact is crucial for a palaeogeographical evaluation of these regions.
Fortunately, the classic, although evidently outdated in some parts, work of WENZ (1923 to 1929) gives a catalogue
of all the known nonmarine Tertiary taxa.
It would be useless to give a quantitative analysis of the studied fauna, because most of the material originates from
drilling cores. A quantitative analyse of such analysis material is worthy only if:
— the core again is almost 100%;
— the diameter of the cores its identical;
— the fossil may be extracted entirely from the matrix;
— the whole material was used.
Evidently, the above cases are rare.
Geologica Hungarica series Palaeontologica, fasciculus 56
9
GEOLOGICAL DESCRIPTION
OF THE STUDIED AREA
On the enclosed map (Figure 1) the Miocene fossiliferous
localities of the Bakony Mts are depicted. The sites-yielding nonmarine fossils will be presented below in a chronological order,
and their stratigraphical setting and lithological characters will
be illustrated by sections of characteristic drillings.
The stratigraphic position of nonmarine layers were assessed
mainly according by their relation with nearby marine sediments.
In some instances the nonmarine deposits intercalate marine
ones, but quite often these represent the initial or closing stratum
of marine sequences. The dates given in this paper can be regarded as highly, due mainly to the extensive stratigraphic work carried out in the area.
Figure 1. Map of the localities
LOWER MIOCENE
Lower Miocene deposits were found in the forelands of the W Bakony Mts, from the
drilling Somlóvásárhely–1 (Svh–1), Ottnangian and Karpatian, but with no possibility of precise dating. The drilling, between the depths 144.4 and 273.5 m penetrated a heavily benthonic clay and silt series with pebbly and sandy intercalations (Figure 2).The clay often contains
biotite crystals.The series lays on the Oligocene Csatka Fm (BÁLDI 1986) and is capped by a
marine Lower Badenian sequence. At about at the middle part of clay series (between
188–193 m and 220–231 m) there are lacustrine to marsh deposits, yielding a rich nonmarine
mollusc fauna. The sequence is lithologically different from that of the Oligocene Csatka Fm.
Therefore SELMECZI (1989) separated it from the Csatka Fm under the name Somlóvásárhely
Fm. I am indebted to her for supplying me with the mollusc she collected from the mentioned
drilling. The mollusc-bearing layers contain calcareous algal oncoids characteristic for freshwater and brackish deposits, indicating a shallow, warm water environment.
The sequence yielded 59 mollusc species or subspecies, of which 56 are gastropods, 40
forms are terrestrial, and the rest are freshwater. 23 taxa of the assemblage were known from
Lower Miocene deposits. From the Lower Miocene of the drilling Somlóvásárhely–17 aquatic mollusc species were collected. Species of the genera Theodoxus, Brotia and Melanopsis
might also have survived in a low salinity, brackish environment, but the rest of the taxa point
unequivocally to a freshwater one. Consequently, we presume that the environment was not
at all under a marine influence.
The age of the mollusc-yielding deposit may be determined more precisely using data
about the tectogenetic processes of the mountain belt (KÓKAY 1996). Along some faults and
fault systems vertical movements took place, accompanied by intensive volcanism (KÓKAY
2002b). If there was a possibility, marine environments were established in depressions —
e.g. in the Várpalota Basin. In a depression in the W Bakony Mts. Fluviolacustrine sediments
accumulated with volcanosedimentary material. This latter has mostly been transformed to
Figure 2. Columnar section
of drilling Somlóvásárhely
Svh–1
1 — clay, 2 — clay marl, 3 —
gravel, 4 — sand, 5 — coaly
clay, 6 — bentonitic clay, 7 —
marine, 8 — snail
10
JÓZSEF KÓKAY
bentonite. This type of sequence was penetrated by the Somlóvásárhely drilling between the marine Lower Badenian and
the Oligocene Csatka Fm. This seems to point to a continuous sedimentation from the Ottnangian, on up to the Badenian
marine transgression. The sequence described herein includes the Ottnangian and Karpatian stages.
This dating is substantiated by the mollusc fauna as well. The most important mollusc taxa: Viviparus pachystoma
dissitus, Valvata homalogyra, Sellia miocaenica, Stalioa praeglobulina, Staliopsis puisseguri, Platyla callosa, P. subfusca, Brotia escheri, Gyraulus applanatus, Negulus suturalis suturalis, Acanthinula tuchoricensis, Strobilops fischeri,
Napaeus complanatus, Aegopis algiroides, Oxychilus denudatum var., Arion sp., Triptychia grandis, Canalicia cf. attracta, Pseudoleacina producta, Pseudoleacina confusa subsulcosa, Perforatella homalospira, Galactochilus mattiacum,
Tropidomphalus robustus, T. extinctus, Klikia osculum tenuis, Pisidium ex gr. casertanum.
Species of the genera Theodoxus, Brotia and Melanopsis might also have survived in a low salinity brackish environmental, but the rest of the taxa point unequivocally to a freshwater one. Consequetly, we presume that the environment
was not at all under a marine influence.
OTTNANGIAN
Ottnangian nonmarine fossils have only been found in the Várpalota Basin. The Ottnangian sea inundated the deep
parts of the collapsing basin (KÓKAY 1991, 1996). In these places the sedimentation began with terrestrial or freshwater,
sometimes with brackish deposits.
At the NW border of the Várpalota Basin, at Bántapuszta (Figure 3) the well drilling Öskü–2, between 19.5 m and
63.8 m penetrated sand and gravel layers with silty claymarl intercalations (Figure 4). In the lower part of this sequence
prints of the following nonmarine fossils were found: Pomatias bisulcatum, Amnicola? sp., Planorbarius cornu cornu.
Milax? sp., Pisidium sp. The first mentioned terrestrial form indicates a Miocene age.
The key section of the well V–133 was performated
in the centre of the basin (KÓKAY 1973). At the base of
the Ottnangian, between 208–226 m, a lignit-bearing,
silty claymarl of freshwater origin was found. This yielded fragments of Planorbarius and Characea oogonia.
The drilling Inota–138 penetrated a sandy and
pelitic sequence of freshwater (and, possibly partly of
brackish) origin between 240.5–263 m (Figure 5), with
Figure 3. Map of the Várpalota Basin
1 — Drilling, 2 — Border of the studied area
Figure 4. Columnar section of
drilling Várpalota Öskü–2
Figure 5. Columnar section of
drilling Várpalota I–138
1 — clay,2 — gravel, 3 — sand, 4
— snail
1 — clay marl, 2 — sand, 3 — silt, 4
— snail
Geological description of the studied area
11
the following: Theodoxus cyrtocelis, Staliopsis moravicus, Brotia escheri,
Acanthinula tuchoricensis, Aegopis algiroides, Triptychia vulgata, T. cf. grandis,
Perforatella cf. homalospira, Tropidomphalus cf. robustus, Klikia cf. devexa. The
first mentioned two and the last form prove a Lower Miocene age. The
Theodoxus and Staliopsis species are characteristic elements of the latest
Ottnangian regressive layers in the North Alpine Molasse Zone. At Várpalota
these species appear slightly earlier, but still within the Ottnangian four species
indicate a Lower Miocene age, none of them point to a younger one.
KARPATIAN
From the Karpatian, nonmarine molluscs are known from the western Bakony
Mts (drilling Nagygörbő–1 and wells at Herend, Figure 6). These were found in
terrestrial sediments, overlain by the transgressive Lower Badenian: Brotia
escheri and Unio sp.
At the basis of the marine sequence in the borehole Várpalota–219 (SE
Bakony Mts),a single specimen of Pomatias turonicum was found, which was
Figure 6. Map of the Herend Basin
Figure 7. Columnar section of drilling
Polgárdi Po–2
washed in from dryland. In the terrestrial sequence of the Karpatian from the
drilling Polgárdi Po–2 (Figure 7, KÓKAY 2002a) the following were found:
Planorbarius cornu solidus, Tropidomphalus extinctus and Cepaea bohemica.
1 — dacitic tuffite, bentonite, 2 — terrestrial,
3 — brachyhaline, 4 — lacustrine, 5 — snail
BADENIAN
Lower Badenian (Moravian, PAPP et al. 1978)
1. Western Bakony Mountains. The uppermost, benthonitic layers of the Karpatian variegated terrestrial clay and
gravel sequence (Somlóvásárhely Fm) were inundated from the west by the Early Badenian sea. The Lower Badenian
(Moravian, PAPP et al. 1978) sedimentation starts with lacustrine and paludal deposits, which seem to be slightly younger
at the western border of the Bakony Mts, than further to the East, in the centre of the mountains.
Drilling Nagygörb ő N g – 1. At the western border of the Bakony Mts, at the margin of the Little Hungarian Plain
(Kisalföld) a key drilling was deepened (Figure 8). Between 892–923 m there was a mainly pelitic and sandy freshwater
layer intercalated with brown coal-seams and coaly clays, just under a Lower Badenian silty marine sequence. The freshwater layer contained fossil rich beds, and the fossils were poorly or moderately well preserved. Bithynia subgracilis shells and
opercula were predominant; in the upper third of the sequence Melanopsis visianiana was frequent. I am indebted to Dr.
Margit Bohn-Havas, who was so kind to allow me to study her samples. 14 species were determined: Theodoxus sinjanus,
T. cyrtocelis, Bithynia subgracilis, B. dunkeri, Brotia escheri, Melanoides procurvicosta, Melanopsis visianiana,
Planorbarius cornu cornu, Unio larteti etc.
12
JÓZSEF KÓKAY
Figure 8. Columnar section of drilling
Nagygörbő Ng–1
Figure 9. Columnar section of drilling
Herend He–13
Figure 10. Columnar section of drilling
Várpalota Vk–384
1 — variegated clay, 2 — clay marl, 3 —
sand, 4 — marl, 5 — coaly clay, 6 — coal
seam, 7 — snail
1 — clay, 2 — variegated clay, 3 — clay
marl, 4 — coaly clay, 5 — coal seam, 6 —
dacitic tuffite, bentonite
1 — clay, 2 sand, 3 — sandstone, 4 — alginite,
5 — coal seam, 6 — snail
H e r e n d B a s i n . As it was demonstrated in the monograph about the region (KÓKAY 1966), the sea, situated west
of the mountains, entered with a fjord-like narrow basin into the centre of the mountains. In this narrow trough the basal
sediments were paludal to lacustrine with coal-seams. The total thickness of the Lower Badenian was 250 m, while the
coal-seams occurred in a 30 to 70 m thick basal layer. In the above mentioned monograph just a few terrestrial and freshwater taxa were mentioned, with a rich marine to brackish fauna. Here the coal-seam bearing sequence is demonstrated
by the drilling He–13 (Figure 9), but all available samples have been examined.
As I demonstrated (KÓKAY 1966), the marine sedimentation already prevailed is the western margin of the Bakony
Mts, coevally with a marsh environment in the Herend embayment. The temporal discrepancy between the Nagygörbő
and the Herend paludal-lacustrine sediment might not be significant, however.
The number of the known terrestrial and freshwater species is 28, of which 8 are terrestrial: Nematurella species predominate with Microprososthenia belchatowensis, Martinietta tumida, “Amnicola” pseudoglobulus, Brotia escheri,
Emmericia subpatula, Gyraulus trochiformis dealbatus, Planorbarius cornu mantelli, Theodoxus barakovici etc. There
is just one form common with the Nagygörbő fauna: Theodoxus cyrtocelis.
2. Eastern Bakony Mts:
Vá r p a l o t a B a s i n . Nonmarine forms were, washed into the marine beds of the “Szabó” sandpit in Várpalota.
This pit under conservation yielded more than 400 species, including 8 species of the family Ellobiidae (Pedipes,
Melampus, Stolidoma), which recently lived on dryland, near to tropical beaches. Brotia escheri and Theodoxus grateloupianus were also recorded. These usually live in fresh or highly diluted brackish waters.
Freshwater mollusc from the layers capping the marine Lower Badenian cycle. The best fauna was found in the NW
part of the Várpalota Basin, at the western margin of the Bántapuszta subbasin, in the drilling Vk–384. The uppermost
section of the marine Lower Badenian (at 69.3–70.4 m, Figure 10) contained freshwater molluscs mixed with marine and
Geological description of the studied area
13
brachyhaline ones. The freshwater forms were most probably carried in by a creek or small river, comprising altogether
10 species: Theodoxus grateloupianus, T. burdigalensis trifasciatus, Brotia escheri, Gyraulus amiculus, Ferrissia
deperdita, Margaritifera flabellata, Unio eseri, U. lorioli, Pisidium ex aff conventus, P. ex aff. nitidum, as well as coalfield plant debris.
Middle Badenian (Wielician, PAPP et al. 1978)
Middle Badenian (Wielician, PAPP et al. 1978) terrestrial deposits are known only in the Várpalota Basin (KÓKAY 1985,
1991): variegated clays and gravel, sporadically containing the remains of molluscs. I was found in the bentonitic clay under
the Upper Badenian coal-seam (drilling I–124 — Figure 11, I–129 — Figure 12, I–135, Cs–17 — Figure 13) terrestrial and
freshwater molluscs (mainly fragments), comprising altogether 19 species: Theodoxus cf. grateloupianus, Pomatias conica,
Platyla alta, P. cf. beatrcis, Brotia escheri, Carychium nouleti gibbum, Radix cf. dilatata, Anisus sp., Gyraulus amiculus,
Planorbarius sansaniensis, Enneopupa subcylindrella?, Vitrea procrystallina, Aegopis costatus, Oxychychylus? sp.,
Triptychia cf. suturalis, Clausilia ex aff. rolfbrandti, Palaeoglandina gracilis porrecta, Perforatella (M.) cf. punctigera,
Canariella bakonyensis, Klikia cf. coarctata, Margaritifera cf. flabellata, Pisidium ex aff. conventus.
Upper Badenian (Kosovian, PAPP et al. 1978)
Western Bakony region:
Ta p o l c a B a s i n . In the paper which dealt with the Badenian along Lake Balaton (KÓKAY 1986), the Tapolca Basin
Badenian was described in detail, based on drilling data. The well Hegymagos Hg–78/18 gave important data about the
nonmarine Upper Badenian mollusc fauna (Figure 14). The Late Badenian sedimentation ended in a freshwater environment, with fossil rich clays and claymarls between 154.6–158.8 m. Beneath it there were Upper, Middle and Lower
Badenian brackish and marine layers,
which were probably ranged in one
cycle. The regression during the Late
Badenian was followed by Sarmatian
restricted marine sediments.
In the rich fauna of the freshwater
part of the sequence Brotia escheri
turrita prevails. Numerous freshwater
and terrestrial forms (47 taxa) were
registered. In spite of the moderate
distance from the western Bakony
Mts, the number of common taxa is
low. For example Sandbergerina suc-
Figure 11. Columnar section of drilling
Várpalota I–124
Figure 12. Columnar section of drilling
Várpalota I–129
Figure 13. Columnar section of drilling
Várpalota Cs–17
1 — clay, 2 — silt, 3 — alginite, 4 — coal
seam, 5 — lime mud, 6 — marine, 7 — snail
1 — clay, 2 — variegated clay, 3 — clay
marl, 4 — gravel, 5 — sandstone, 6 — alginite, 7 — coaly clay, 8 — coal seam, 9 —
lime mud, 10 — dacitic tuffite, bentonite, 11
— terrestrial, 12 — freshwater, 13 — snail
1 — clay, 2 — variegated clay, 3 — clay marl,
4 — coaly clay, 5 — coal seam, 6 — lime mud,
7 — terrestrial, 8 — freshwater, 9 — mesohaline, 10 — marine, 11 — lacustrine, 12 — paludal, palustrine, 13 — snail
14
JÓZSEF KÓKAY
cineiformis is lacking from the Hegymagos fauna, which is coeval with the freshwater clay and which closes the cycle at Pusztamiske. Common taxa are present
among the terrestrial forms. Geological data also substain the assumption that
these two sedimentary basins were not directly connected.
In some levels the Hydrobia species occur within the sequence of freshwater
origin, pointing to some miohaline influence of short duration and just tolerable
for freshwater forms.
My sincere thanks are due to Dr. György Majoros for allowing me to study the
samples.
U p p e r B a d e n i a n l a g o o n . In the last four decades numerous wells were
drilled in the western Bakony Mts, especially reconnaissance-drillings for coal and
bauxite; some of these were “key wells”. Cores of the last mentioned types were
studied in many respects, yielding fundamental stratigraphical, faciological and
palaeogeographical information. The wells in the Pusztamiske area were the most
important from my point of view. These initiated the study of nonmarine Badenian
molluscs. On the other hand, they furnished key information for the understanding
of the Late Badenian geography. A study of wells around Pusztamiske (KÓKAY
1992) revealed that at the site of this village a NNW–SSE oriented 3 km wide
graben was formed syngenetically. This graben was inundated by the sea from
NNW, but the connections were restricted in a lagoonal environment and freshand lower brachyhaline water prevailed.
Towards the south (Nyirád) the carbonate content of the Upper Badenian strata increases; simultaneously, these indicate a decreasing salinity. The graben evidently had a branching towards the East, where, in the Herend Basin, there is a
benthonitic sequence essentially identical with that of Pusztamiske (KÓKAY 1966).
Thus, in the Late Badenian a branching lagoon system was formed in the Western
Bakony Mts; nowadays by now its deposits remain only as vestiges. The
Pusztamiske wells explored the middle part of the southern branch of the system.
— Pusztamiske region (Figure 15). A 15–25 m thick, well-rounded and sorted
gravel bed covers the marine Lower Badenian and it is probably of littoral origin.
It is most likely that this is the Middle Badenian, having been deposited between
Figure 14. Columnar section of drilling Lower and Upper Badenian layers. The Upper Badenian in this region is about 40
Tapolca Hg–78/18
m thick. It begins with a coal-seam, coally clay and a lacustrine to lagoonal clay1 — variegated clay, 2 — marl, 3 — coaly
containing
level. The sequence contains masses of Brotia and Theodoxus shells,
clay, 4 — lime mud, 5 — limestone, 6 —
freshwater, 7 — oligo-miohaline, 8 — mio- and in the coally clay there are concentric calcareous concretions, probably of
haline, 9 — pliohaline, 10 — brachyhaline, algal origin. Brotia escheri turrita and Theodoxus burdigalensis burdigalensis
11 — snail
dominate. The washed material included numerous small shells of freshwater,
oligo- and miohaline and terrestrial molluscs, partly belonging to unknown or rare
taxa: Nematurella schuetti, N.? aquensis, Pseudamnicola leognanensis, Stalioa globulina, Carychium nouleti gibbum,
Auriculastra badeniensis, Vertigo kochi, Canariella bakonyensis etc.
The coal-seam is capped by pelitic or sandy mollusc bearing beds of
lower brachyhaline origin and a littoral gravel with well-rounded pebbles
(KÓKAY 1992). The gravel is covered by a 3.5–6.6 m thick, Brotia-bearing
limestone. This yellowish-white, ocre-patched limestone is generally
porous, containing footprints, and cracks. Laminated beds are included. In
some levels abundant prints of Brotia escheri turrita, Gyraulus sp. and
Theodoxus burdigalensis burdigalensis were found. The limestone was best
developed in the well Pm–3 (Figure 16), between 18.1–24.7 m. In this interval various mollusc assemblages occurred, reflecting the changing salinity.
At the bottom there was a layered, rootprint-containing calcareous mud,
covered with a freshwater limestone bearing moulds of Bithynia glabra
(23.3–23.5 m). Between 21.4 and 21.5 m prints of Theodoxus pictus and
Hydrobia ventrosa are abundant, indicating a miohaline environment.
Between 20.0 and 20.3 m Brotia escheri turrita, Theodoxus burdigalensis
and Gyraulus sp. prints occur with terrestrial gastoropods: Galactochilus
sp., Poiretia taurinensis, Cepaea silvana. The limestone was of freshwater
Figure 15. Map of the Pusztamiske area
between 19.6 and 19.8 m, with abundant Lymnaeid taxa. At a depth of 19.3
Geological description of the studied area
15
Figure 16. Columnar section of drilling
Pusztamiske Pm–3
Figure 17. Columnar section of drilling
Pusztamiske Pm–2
Figure 18. Columnar section of drilling
Nyirád Nyirt–1
1 — clay, 2 — marl, 3 — gravel, 4 — lime mud,
5 — limestone, 6 — dacitic tuffite, bentonite, 7
— freshwater, 8 — oligohaline, 9 — miohaline,
10 — oligo-miohaline, 11 — mesohaline, 12 —
brachyhaline, 13 — snail
1 — clay, 2 — clay marl, 3 — gravel, 4 — sand,
5 — sandstone, 6 — coaly clay, 7 — coal seam, 8
— limestone, 9 — freshwater, 10 — oligohaline,
11 — brachyhaline, 12 — snail
1 — clay, 2 — clay marl, 3 — marl, 4 — silt,
5 — coaly clay, 6 — lime mud, 7 — limestone, 8 — dacitic tuffite, bentonite, 9 —
freshwater, 10 — oligo-miohaline, 11 —
oligohaline, 12 — miohaline, 13 — pliohaline, 14 — brachyhaline,
m an inch-thick layer of calcareous clay contains Pirenella and foraminifer remnants, indicating a lower brachyhaline
(14–16 promil salinity).
In the upper part of the limestone a 20–40 cm thick, strongly-weathered dacitic tuff layer was observed, covered by
a maximum 2.3 m thick (in the well Pm–2 — Figure 17) freshwater clay, claymarl and marl. The most characteristic mollusc was the abundant Sandbergerina succineiformis, as well as Brotia escheri turrita, Theodoxus burdigalensis,
Gyraulus microstatus and Bithynia glabra. The terrestrial gastropods are represented by the Carychium and Vertigo
species. The freshwater series passes into a brackish Sarmatian sequence (in the well Pm–2). At the beginning of the
Sarmatian sedimentation, Bithynia glabra still prevails, but upwards it disappears, while the typical Sarmatian taxa also
appear (see also the description of Bithynia glabra).
— Nyirád Basin. At the southern end of the Upper Badenian lagoon (south of the village of Nyirád), bauxite
prospecting wells and the Nyrt–1 reconnaissance drilling found a 24–29 m thick Upper Badenian sequence (Figure 18).
This consists of freshwater and oligo-miohaline lagoonal deposits. It contains more carbonate than the Pusztamiske
sequence, and its fauna points to a lower salinity. Associations indicating a lower brachyhaline environment are not present here. Hydrobia-, Stagnicola praebouilleti, Brotia-, Theodoxus-, Bithynia- and Sandbergerina-bearing associations
can all be recognized.
— Herend Basin. The Bánd bentonite seam is covered by a Brotia (“Melania”)-bearing clay sequence, which was
described in the monograph of the Herend Basin Miocene (KÓKAY 1966) as well as in the paper describing the Upper
Tortonian of the Bakony Mts (KÓKAY 1967). In the light of the data gathered since then from drillings and outcrops, it
16
JÓZSEF KÓKAY
Figure 19. Columnar section of drilling
Herend He–20
Figure 20. Columnar section of drilling
Várpalota I–86
1 — clay, 2 — gravelly clay, 3 — gravel, 4 —
limestone, 5 — dacitic tuffite, bentonite, 6
— freshwater, 7 — oligo-miohaline, 8 —
snail
1 — clay, 2 — variegated clay, 3 — gravel, 4
— sand, 5 — sandstone, 6 — silt, 7 — coaly
clay, 8 — dacitic tuffite, bentonite, 9 — terrestrial, 10 — freshwater, 11 — snail
is clear that it is closely related to the
coeval formations of the Pusztamiske and
Nyirád region in its fauna, geology and
sequence. Thus the Herend region may
be regarded as a part of the described
Late Badenian lagoon system (KÓKAY
1992).
Unfortunately, work in the open air
bentonite pit in Bánd was stopped much
earlier; in the collapsed pit collecting
work is no longer possible. A canalisation
trench was made at Herend not long ago,
parallel to the highway leading to Graz in
which the Upper Badenian “Brotia”bearing bentonitic sequence cropped out.
Using also the stored core samples of
another drilling (He–20 — Figure 19), a
good picture might be gained of the Late
Badenian mollusc fauna in this basin. The
approximately 30 m thick sequence is
essentially of freshwater origin, although
in its lower and middle parts there are
beds containing sporadically oligo- or
miohaline Hydrobioids. It contains terrestrial gastropods and shells of Brotia
escheri escheri and Brotia escheri turrita,
which were often encrusted with a concentric algal crust. Other, frequent species
are: Theodoxus grateloupianus, T. burdigalensis, Gyraulus microstatus as well as
the Nematurella species (including N.
aquensis), which is also frequent in the
Pusztamiske coal-seam. The Herend
Basin yielded 32 species, some of which
can only be determined at the generic
level.
Eastern Bakony Mts:
Vá r p a l o t a B a s i n , Upper Badenian lacustrine sediments in the SE Bakony Mts. The SE foreland of the Bakony
Mts tectonically collapsed at an earlier date (KÓKAY 1991, 1996). The basin, formed in this way, started as a marsh with
a coal formation; subsequently a lake occupied with a surface area of almost 100 km. It had no direct connection with
the sea, in contrast to the case of the lagoon in the western Bakony Mts. A subsurface exchange of water masses might
be presumed to a limited extent, as was suggested earlier (KÓKAY 1987, 1991; Gaudant personal communication).
According to J. Gaudant (pers. comm.), close relative of fishes found in the Upper Badenian alginites recently lived near
sea lakes and lagoons of the Indo-west Pacific realm.
A great amount of material was collected to help to determine the salinity conditions of the lake in the Várpalota
Basin. The lower some dm of the lacustrine sequence yielded abundant shells of three mollusc species: Congeria boeckhi, Theodoxus crenulatus varpalotensis and Ferebithynia vadaszi. These three forms seldom occur together; in a narrow
layer or in a small outcrop just one or two of them occur. The following three associations were observed at the beginning of the studies: Congeria-Theodoxus or Ferebithynia-Theodoxus or Congeria-Ferebithynia. The last-mentioned one
occurred only rarely, near shore limestone. These associations may reflect bottom conditions (Figures 20, 21). A more
detailed collection proved the existence of more types of associations. The coal-seam (generally huminitic clays or calcareous muds) yielded freshwater and terrestrial communities: Valvata moguntina, Pomatias conica, Bithynia glabra,
Brotia escheri, Radix dilatata, Anisus rousianus, Gyraulus amiculus, G. trochiformis kleini, Planorbarius cornu cf.
solidus, Aegopis costatus, Pisidium bellardii, Discus cf. pleuradrus, Triptychia sp., “Clausilia” sp., Palaeoglandina cf.
gracilis porrecta.
A max. 150 m thick sequence caps the coal-seam: claymarl, sand and limestone yielded the following species:
Theodoxus crenulatus varpalotensis, Ferebithynia vadaszi, Brotia escheri (and ssp. turrita ), Radix dilatata, Anisus rou-
Geological description of the studied area
sianus, Gyraulus verticillus, Ancylus wittmanni, A. moravicus, Congeria boeckhi
and Unio jaccardi.
Although the majority of these species tolerate a slightly increased salinity
(especially the Congeria species), the association as a whole must be regarded as
a freshwater one (see below).
The Várpalota Upper Badenian yielded altogether 34 non-marine mollusc
taxa. All forms of the coal-seam occurred in the western Bakony Mts and the
Tapolca Basin sequences as well, while from the fauna of the cover six taxa were
registered from these two areas.
M ó r g r a b e n , F e h é r v á r c s u r g ó . There is but one datum from the eastern margin of the Bakony Mts: the Mór graben. The key well Fehérvárcsurgó
Fcst–5 penetrated a thick terrestrial sequence between the Triassic basement and
the Pannonian (Upper Miocene). Based on its lithology and general geological
setting, the section between 153.8 and 179.9 m may be placed in the Upper
Badenian (Figure 22). A part of the sequence is lithologically similar to the
Várpalota alginitic layers. It is probable the Mór graben had an open connetion to
the Várpalota area. In the 168–169 m section the following molluscs were found:
Carychium nouleti gibbum, Anisus cf. rousianus, Gastrocopta nouletiana, Vitrea?
sp., Aegopinella? sp., Discus pleuradrus, Canariella bakonyensis, Klikia? sp. and
Cepaea sp. An alginite layer at 171.5–173.0 m yielded Radix dilatata and
Gyraulus cf. verticillus.
This small fauna is similar to other Upper Badenian associations in the
Bakony Mts. I am grateful to Dr. B. Bernhardt for kindly allowing me to study the
samples.
17
Figure 21. Columnar section of drilling
Várpalota V–318
1 — clay, 2 — coaly clay, 3 — coal seam, 4
— dacitic tuffite, bentonite, 5 — freshwater,
6 — snail
SARMATIAN
Sarmatian nonmarine fauna were found in both parts of the Bakony Mts, in
layers intercalated between marine ones, or as washed in specimens.
Western Bakony Mts:
Tapolca Basin. The drilling Hg–78/18 penetrated a huminitic claymarl at
152–152.5 m (Figure 14), yielding Bithynia glabra, Gyraulus trochiformis kleini,
Anisus sp., and Lymnaea ex gr. stagnalis. The fauna is sporadic and ill-preserved,
coming from a thin marsh deposit at the bottom of the Sarmatian marine sequence.
The environment at N y i r á d . In the hydrobioid clay and limestones from the
bauxite prospecting drillings the following taxa could be determined: Theodoxus
grateloupianus, Pomatias conica, Valvata sarmatica, Carychium suevicum,
Gyraulus nedici, Planorbarius cornu mantelli, Stagnicola praebouillet, S. palustris,
Galba dupuyana, Lymnaea turrita, Ferrissia deperdita, Gastrocopta acuminata,
Poiretia taurinensis etc., as well as carpus Celtis and Characea oogonia.
In pelitic and calcareous mud deposits of the 31.1–63.8 m section of the drilling
Nyirt–1 (Figure 18) the following nonmarine taxa were found (accompanied by
hydrobiids, except for the section between 63.6–63.8 m): Pomatias conica, Bithynia
glabra, Stagnicola armaniacensis, Radix socialis regularis, Lymnaea turrita,
Gyraulus nedici, G. trochiformis kleini, Segmentina larteteti, Semilimax intermedia
crassitesta, Poiretia taurinensis and Canariella bakonyensis etc.
P u s z t a m i s k e r e g i o n . The wells found the oldest Sarmatian marine layers with intercalated marsh deposits, containing terrestrial and freshwater molluscs. Generally these were mixed with Sarmatian marine forms and thus they
were probably washed-in. The following were found: Bithinya glabra, Carychium
suevicum, Galba dupuyana, Lymnaea turrita, Anisus hilgendorfi subcarinata,
Gyraulus pulici, G. trochiformis kleini, Segmentina larteti, Vertigo bakonyensis,
Strobilops subconoideus, Semilimax intermedia crassitesta and Canariella
bakonyensis etc.
Western Bakony Mts:
Vá r p a l o t a B a s i n . The Sarmatian is on average 150 m thick in this basin
(KÓKAY 1991). At the beginning, in the Kozardian stage, the tectonically subsiding
Figure 22. Columnar section of drilling
Fehérvárcsurgó Fcst–5
1 — clay, 2 — variegated clay, 3 — clay
marl, 4 — alginite, 5 — coaly clay, 6 — coal
seam, 7 — bentonitic clay, 8 — terrestrial, 9
— freshwater, 10 — snail
18
JÓZSEF KÓKAY
sedimentary basin was abruptly filled in
with fluvial-terrestrial deposits due to the
increased relief energy. Towards the south
this sequence was more and more frequently intercalated by Sarmatian brackish
sediments. The nearshore paludal and
lacustrine sediments contain great masses
of nonmarine molluscs and the conditions
of preservation were excellent here. On the
base of the Sarmatian, near to the Badenian boundary one well (drilling Inota–129
— Figure 12) yielded the following terrestrial molluscs: Cochlostoma septemspirale, Pomatias conica, Acanthinula trochulus, Canariella bakonyensis, Klikia sp.
From the deposits of the transgressing
older (Kozardian) Sarmatian sea, generally from pelitic sediments, 56 taxa could
be determined. The most important of
these are: Cochlostoma septemspirale,
Pomatias consobrinum, Bithynia glabra,
Carychium sandbergeri, Gyraulus trochiformis kleini, Planorbarius cornu cornu,
Negulus suturalis gracilis, Argna aperta,
A. oppoliensis, Paracryna pseudoennea,
Vitrea procrystallina steinheimensis,
Nesovitrea mendica, Daudebardia praecursor, Nordsieckia pontica, Canariella
bakonyensis, Dreissena dobrei, Pisidium
pseudospherium etc.
Nonmarine molluscs are most abunFigure 23. Columnar section of drilling
Figure 24. Columnar section of drilling
Várpalota Ö–11
Várpalota I–51
dant in the upper 10 to 30 m of the
1 — clay, 2 — variegated clay, 3 — clay
1 — clay, 2 — variegated clay, 3 — gravel, 4 Várpalota Sarmatian. These are generally
marl, 4 — marl, 5 — coaly clay, 6 — coal
— marl, 4 — gravel, 5 — terrestrial, 6 —
huminitic pelitic formations sporadically
seam, 7 — dacitic tuffite, bentonite, 8 — terfreshwater, 7 — paludal, 8 — snail
occurring
with calcareous mud levels
restrial, 9 — freshwater, 10 — paludal, 11 —
brachyhaline, 12 — snail
(Figures 23, 24). Even in these layers
there are brackish intercalations with rich
faunas (KÓKAY 1954). Some of the layers contains hydrobioids with nonmarine forms, indicating a mio- or oligohaline
salinity. The sediments contain various associations, wich may be described by their most frequent taxa, e.g: StagnicolaCepaea, Stagnicola-Planorbarius, Segmentina, Anisus, Bithynia, Valvata-Pisidium, Hydrobia-Cepaea, Klikia-Pisidium.
Some are dominated by terrestrial gastropods. From this upper Sarmatian sequence 124 non- marine forms were determined of which 75 are terrestrial gastropods. Some characteristic species include: Cochlostoma septemspirale, Valvata
moguntina, V. politioanei, Pseudamnicola steinheimensis, Bithynia glabra, Sandbergerina succineiformis var.,
Carychium pachychilus, C. suevicum, Stagnicola praebouilleti, S. armaniacensis glabraeformis, S. palustris, Radix
dilatata, R. hyaloleuca, Aplexa subhypnorum physaeformis, Anisus hilgendorfi subcarinata, Gyraulus trochiformis
denudatus, G. homalosomus, Hippeutis fasciatus, Segmentina larteti, Planorbarius saniensis, Pseudancylastrum decoratum, Vertigo callosa perarmata, V. angustior oecsensis, V. pusilla sarmatica, Gastrocopta acuminata larteti, G.
obstructa ferdinandi, Leiostystyla gottschicki, Argna suemeghyi, Vallonia lepida, Spermodea candida, Strobilops sandbergeri, S. pappi, Semilimax intermedia, Janulus supracostatus, Fortuna tertia, Triptychia leobersdorfensis sarmatica,
Nordsieckia pontica, Pseudidyla boettgeri, Palaeoglandina gracilis porrecta, Perforatella punctigera, Klikia kaeufeli, K.
goniostoma, Cepaea etelkae, Pisidium steinheimense, P. pseudospherium.
The uppermost, closing layer of the Várpalota Sarmatian is a 1–3 m thick Melanopsis impressa-bearing claymarl and
clay, rich in dacitic tuff and molluscs. Its fauna is transitional to an extent similar to the Pannonian s.str. one. The
Sarmatian forms are of a small size and contain many Rotalia beccarii forams. Its association is characteristic for the
level. In some cores it contained non-marine elements as well. These are of a Sarmatian type and are as follows: Valvata
moguntina, V. sarmatica, V. soceni wiesensis, Bythinella eugenii, Pseudamnicola tholosa, Platyla polita, Carychium
sandbergeri, C. pachychilus, Galba dupuyana, Stagnicola palustris, Radix dilatata, R. hyaloleuca, Lymnaea turrita,
Geological description of the studied area
19
Aplexa subhypnorum physaeformis, Anisus cf. rousianus, Gyraulus nedici, G. pulici, G. trochiformis kleini, Segmentina
larteti, Planorbarius cornu solidus, Pseudancylastrum decorata, Vertigo callosa, V. callosa diversidens, V. ovatula trolli, V. angustior oecsensis, Gastrocopta nouletiana, G. acuminata, G. suevica, G. infrapontica, Vallonia lepida, V. cf. subcyclophorella major, Strobilops costatus, S. sandbergeri, S. cf. tiarula, Discus pleuradrus, Nesovitrea mendica,
Zonitoides cfr. suevicus, Oxychylus procellarius, Janulus supracostatus, J.? sp., Limax crassus, Perforatella punctigera,
Canariella bakonyensis, Klikia cf. kaeufeli, K. giengensis, Cepaea etelkae, Pisidium bellardii, P. pseudosphaerium.
The Várpalota Sarmatian is unique with its rich nonmarine mollusc fauna of 134 taxa. Of these 85 are terrestrial, 45
are freshwater gastropods and 4 are bivalves. With respect to the molluscs, many Characea oogonia and Carpus remnants were collected. These will be studied by László Rákosi (Geological Institute of Hungary), and vertebrate and Celtis
fossils are to be, studied by László Kordos (Geological Institute of Hungary).
The nonmarine Sarmatian fauna of the western Bakony Mts is poorer than the Várpalota one, but they are close to
each other. Five terretrial forms were found in the western Bakony, and these were not present in Várpalota. Thus the
Bakony yielded altogether 141 nonmarine Sarmatian molluscs, of which 88 are terrestrial, 45 are freshwater gastropods,
while four are bivalves.
From Várpalota, BODA (1959) enumerated 20 nonmarine Sarmatian mollusc species. Howewer his findings are outdated, as is clear from the synonym lists.
Geologica Hungarica series Palaeontologica, fasciculus 56
21
EVALUATION OF THE STUDIED FAUNAS
FROM THE BAKONY
This chapter aims to summarise the results which are presented in detail in the descriptive part.
Altogether 272 taxa were recognised from the Miocene deposits of the Bakony Mts. In addition, several poorly preserved fragmentary forms were found: Azeca, Truncatellina, Clausilia, Succinea, Gastrocopta, Cepaea, Unio sp., and
scutum of three indeterminable slugs. Of these, 251 are gastropods, and 21 are bivalves. 96 snails are aquatic, and 153
are terrestrial. Some cores are extremely rich in fossils, and if these had been on the surface they would have yielded
incredibly rich associations.
From the Ottnangian of the Várpalota 13 forms were determined; from the Karpatian of the Bakony Mts 4; were distinguished and from the Lower Miocene of Somlóvásárhely 59 taxa were recognised. The Lower Badenian of the Bakony
Mts yielded 58 species, of which 36 originated from Herend, 18 from Várpalota and 15 from Nagygörbő. From the
Middle Badenian (Várpalota) 22 nonmarine mollusc forms were determined while from the Upper Badenian sediments
116 taxa were recognised. The figures for the following places are:
Pusztamiske 62, Nyirád 25, Herend 41, the Tapolca Basin 46, Várpalota 33, and Fehérvárcsurgó 5. The Bakony
Sarmatian yielded 142 nonmarine mollusc taxa: 4 from the Tapolca Basin, 32 from Nyirád, 26 from Pusztamiske, while
134 from Várpalota (see the faunalist).
PALAEOGEOGRAPHIC IMPLICATIONS
Out of the 272 taxa almost 55% (149) are known from the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) series or from the northern Alpine Foreland. There are forms characteristic for the older (Kirchberg) and for
the younger (Silvana) associations. Characteristic Kirchberg forms are: Theodoxus cyrtocelis, Nematurella scholli, N.
pappi, Amnicola pseudoglobulus, Bithynia dunkeri, B. glabra, Stagnicola armaniacensis, S. praebouilleti, Radix dilatata, Gyraulus trochiformis dealbatus, Ferrissia wittmanni, Margaritifera flabellata bavarica, Unio eseri.
Species known from the Silvana layers are the following: Pomatias consobrinum, P. conica, Bithynia glabra,
Nematurella flexilabris, N. schuetti, Sandbergerina succineiformis, Melanopsis cf. fritzei curta, Brotia escheri turrita,
Carychium nouleti gibbum, C. eumicron peneckei, Galba dupuyana, Stagnicola jaccardi, S. praebouilleti, Radix dilatata, R. socialis regularis, I. ymnaea turrita, Gyraulus trochiformis kleini, Segmentina larteti, Planorbarius cornu mantelli, Planorbarius cornu solidus, Ferrissia deperdita, Negulus suturalis, Strobilops uniplicata plana, Spermodea candida,
Helicodiscus subteres, Vitrea procrystallina, Nesovitrea subhammonis, Janulus moersingensis, Zonitoides suevicus,
Succinea minima, Opeas minutum, Triptychia suturalis, Tropidomphalus incrassatus, T. zelli, Palaeoglandina gracilis
porrecta, Klikia giengensis, K. coarctata, Cepaea silvana, Unio lorioli, U. hinschi, U. jaccardi.
The Steinheim am Aalbuch locality in the northern Alpine Foreland is well known for its rich freshwater and terrestrial mollusc fauna. According to recent investigations (GREGOR 1982, HEISSING 1989) it is coeval with the lowermost
Sarmatian of the Paratethys, and approximately coeval with about the Sarmatian – Upper Badenian boundary. Most of
the taxa enumerated below are known only from Steinheim and the Bakony localities: Pseudamnicola steinheimensis,
Amnicola pseudoglobulus, Stagnicola armaniacemsis palustriformis, S. armaniacensis glabraeformis, Radix socialis
praelongata, Lymnaea turrita, Aplexa subhypnorum physaeformis, Anisus hilgendorfi subcarinata, Gyraulus trochiformis kleini, G. trochiformis denudatus, Hippeutis fasciatus, Pseudancylastrum deperditolacustris, Cochlicopa subrimata loxostoma, C. subrimata procera, Vertigo callosa perarmata, Gastrocopta acuminata larteti, G. acuminata procera, Argna reperta, Strobilops subconoideus, Succinea minima, Vitrea procrystallina steinheimensis, Opeas minutum,
Klikia coarctata steinheimensis, Pisidium steinheimense.
22
JÓZSEF KÓKAY
It is remarkable that in the Bakony Miocene species with southern (i.e. Croatian, Dalmatian, Bosnia-Herzegovinian)
Miocene (Ottnangian and Badenian) affinities the following occur: Theodoxus sinjanus, T. barakovici, Valvata homalogyra, Melanopsis visianiana, Radix hyaloleuca, Gyraulus nedici, G. pulici, Congeria boeckhi, C. hercegovinensis, C.
venusta, Pisiidium bellardii.
Affinities to southern France (Sansan and Dax) are also obviously present in the studied nonmarine fauna:
Theodoxus.burdigalensis, Nematurella aquensis, Pseudamnicola leognanensis, Schuettemmericia paulensis, Galba
dupuyana, Stagnicola armaniacensis, Anisus dupuyanus, A. omalus, A. rousianus, Gyraulus goussardianus, G. microstatus, G. callistus, Segmentina larteti, Planorbarius sansaniensis, Gastrocopta tapeina, Caracollina phacodes barreri,
Tropidomphalus extinctus, Unio larteti.
Taxa known from the Miocene of the Czech Republic include the following: Theodoxus cyrtocelis, Staliopsis moravicus, Staadtielliopsis rubeschi var., Gyraulus amiculus, Ancylus moravicus,Negulus suturalis suturalis, Acanthinula
tuchoricensis, Strobilops fischeri, Aegopis algiroides, Oxychilus denudatus var., Triptychia vulgata, Canalicia cf. attracta, Pseudoleacina producta, P. confusa subsulcosa, Tropidomphalus robustus, Klikia osculum tenuis, Klikia cf. devexa,
Cepaea bohemica.
Affinities to southern Poland are present with the following taxa (ŁOMNICKI 1886 and KADOLSKY-PIECHOCKI 2000.):
Microprososthenia belchatovensis, Martinietta tumida, Bithynia glabra subgracilis, Radix dilatata, Stagnicola armaniacensis, S. kreutzi, Lymnaea turrita, Planorbarius cornu mantellí, P. sansaniensis, Gyraulus trochiformis kleini,
Gastrocopta nouletiana, Strobilops sandbergeri, Caracollina phacodes barreri, Cepaea silvana.
Only a few taxa common with the Miocene of the Caucasus are present in the Bakony (STEKLOV 1966): Carychium
suevicum, Gastrocopta acuminata, G. nouletiana, G. nouletiana graciliden, Vertigo callosa, Strobilops costata, Opeas
minutum.
The nonmarine Miocene (mainly Pannonian = Upper Miocene) molluscs of the Carpathian and Vienna Basins coincide to the extent of about 25% with that of the Bakony Lower and Middle Miocene (BARTHA 1959; BODA 1959; GAÁL
1911;. LUEGER 1981; SCHLIKUM 1978, 1979). There are scarce data about the northern Italian nonmarine Lower or Middle
Miocene molluscs. SACCO (1886, 1897) mentions from the “Elveziano” the following species: Theodoxus grateloupianus
and Poiretia taurinensis.
The above mentioned areas are in fact faunal subprovinces, comparable to extant ones in Europe.To summarise, it can
be said that the Bakony Miocene nonmarine fauna is closest to that of the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”), given that they have the most species in common.
GEOLOGICAL AGE AND STRATIGRAPHY
The geological age of the localities is well established, based on the data presented above and in line with other references. Variations between coeval faunas do not point to age differences — rather, these reflect the complex systems of local
conditions, which are even more variable than in the case of the marine biota. There are unusual factors (e.g. the transport
of the shells into ecologically remote settings such as the transport of terrestrial gastropod shells into freshwater or marine
environments; or the washed-in fauna of a wet forest origin differs substantially from that of a dry grassland).
The abundance of a given species may not necessarily express the geological age. This parameter is not only controlled by what is optimal for the taxon living conditions, but also by the transportability of its shell. Small gastropod
shells with a dentate aperture (e.g. Vertigo, Gastrocopta, Carychium) retain the air and thus they may float much longer
than more simple shells, consequently, these may be carried much farther into the basin.
Using the references, 272 taxa were evaluated from the point of view of geological age that is, which forms are older
or younger than a given stage or substage.
— Taxa known from pre-Badenian formations: Theodoxus grateloupianus trigonulus, T. burdigalensis, T. burdigalensis trifasciatus, T. barakovici, T. cyrtocelis, T. sinjanus,Viviparus pachystoma dissitus, Valvata homalogyra, V.
moguntina, Pomatias bisulcatum,P. turonicum, Nematurella scholli, N. pappi, N. flexilabris, N. aquensis, Pseudamnicola
leognanensis, Schuettemmericia paulensis, Stalioa praeglobulina, Staliopsis moravicus, S. puisseguri, Staadtiellopsis
rubeschi, Ferebithynia vadaszi, Laemodonta bourgeoisi, Melampus turonensis, Pedipes myotis pisolina, Stolidoma
tournoueri, S. mayeri, Gyraulus amiculus, G. applanatus, G. trochiformis dealbatus, Ancylus moravicus, Negulus suturalis suturalis, Nesovitrea mendica, Acanrhinula tuchoricensis, Strobilops fischeri, Napaeus complanatus, Discus euglyphoides euglyphoides, Aegopis algiroides, Oxychilus denudatus, Triptychia vulgata, Canalicia cf. attracta,
Psedoleacina producta, P. confusa subsulcosa, Perforatella homalospira, Galactochilus mattiacum,Tropidomphalus
robustus,T. extinctus, Klikia osculum tenuis, K. cf. devexa var., Cepaea bohemica, Congeria boeckhi, C. hercegovinensis, C. venusta, Unio eseri.
— Taxa known from pre-Sarmatian formations: Theodoxus grateloupianus, T. burdigalensis, T. burdigalensis trifasciatus, T. barakovici, T. cyrtocelis, T. sinjanus, T. crenulatus varpalotensis, Palaina martensi, Pomatias bisulcatum,
Evaluation of the studied faunas from the Bakony
23
Valvata moguntina, Nematurella scholli, N. pappi, N. flexilabris, N. aquensis, Pseudamnicola leognanensis, Staliopsis
moravícus, Schuettemmericia paulensis, Amnicola punctum, Bithynia glabra, B. glabra subgracilis, B. dunkeri,
Ferebithynia vadaszi, Sandbergerina succineiformis var., S. hidasensis, Staadtielliopsis rubeschi, Platyla falkneri, Brotia
escheri, B. escheri turrita, Melanopsis visianiana, Laemodonta bourgeoisi, Melampus turonensis, Pedipes myotis pisolina, Stolidoma tournoueri, S. mayeri, S. dolioliforrnis, S. gracilis, Carychium nouleti, C. nouleti gibbum, C. eumicron, C.
eumicron peneckei, Galba dupuyana, Stagnicola jaccardi, S . praebouilleti, S. armaniacensis, S. kreutzi, Radix socialis
regularis, R. dilatata, R. hyaloleuca, Anisus dupuyanus, A. omalus, A. rousianus, Gyraulus goussardianus, G. amiculus,
G. applanatus, G. alienus, G. nedici, G. pulici, G. microstatus, G. callistus, G. trochiformis dealbatus, Segmentina
larteti, Planorbarius sansaniensis, Ferrissia wittmanni, Ancylus moravicus, Azeca cf peneckei. Negulus suturalis,
Vertigo ovatula miliiformis, Vertigo kochi var., Gastrocopta tapeina, Paracoryna pseudo ennea, Spermodea candida,
Planogyra nana, Strobilops uniplicata plana, S. subconoidea, S. costata bilamellata, S. sandbergeri, Succinea minima,
Helicodiscus subtetes, Semilimax intermedia crassitesta, Vitrea procrystallina, Nesovitre subhammonis, N. mendica,
Aegopis costatus, Daudebardia praecursor, Zo- nitoides suevicus, Janulus moersingensis, J. supracostatus, Opeas minutum, Triptychia vulgata, T. grandis, Serrulina ex gr. ptycholarynx, Pseudidyla boettgeri, Poiretia taurinensis,
Perforatella punctigera, Caracollina phacodes barreri, Tropidomphalus cf. incrassatus, T. zelli, Klikia giengensis, K. cf.
coarctata, Cepaea silvana, Congeria boeckhi, C. hercegovinensis, C. venusta, Margaritfera flabellata, Unio eseri, U.
lorioli, U. jaccardi, U. larteti, U. hinschi, Pisidium bellardii, P. annandalei.
— Taxa known from post-Badenian deposits only: Cochlostoma septemtrionale, Valvata sarmatíca, V. politioanei, V.
moesiensis, V. soceni wiesensis, Bythinella eugenii, Pseudamnicola tholosa, P. steinheimense, P. inflata, P. immutata,
Amnicola pseudoglobulus, Carychium sandbergeri, C. suevicum, C. pachychilus, Stagnicola palustris, S. armaniacensis
palustriformis, S. armaniacensis glabraeformis, Radix socialis praelongata, Aplexa subhypnorum physaeformis, Anisus
hilgendorfi subcarinata, Gyraulus trochiformis denudatus, G. verticillus, G. homalosomus, Hippeutis fasciatus,
Pseudancylastrum deperditolacustris, Cochlicopa subrimata loxostorna, C. subrimata procera, Vertigo callosa diversidens, V. callosa convergens, V. callosa cardiostma, V. callosa perarmata, V. protracta suevica, V. angustior ocsensis,
Gastrocopta acuminata larteti, G. acuminata procera, G. nouletiana gracilidens, G. suevica, Leiostyla gottschicki,
Argna suemeghyi, Strobilops tiarula, S. pappi, Mastus pupa sarmatica, Vitrea procrystallina steinheimensis, Zonitoides
schraieri, Fortuna tertia, Limax crassus, Milax lorentheyi, Triptychia cf. suturalis var., Nordsieckia pontica,
Mactrogastra cf. voesendorfensis, Galactochilus sarmaticum, Tropidomphalus zelli depressus, Helicigona wenzi, Klikia
kaeufeli, K. goniostoma, K. coarctata cf. steinheimensis, Cepaea etelkae, Dreissena dobrei, Pisidium steinheimense, P.
pseudospherium, P. ex aff. nitidum, P. ex aff. conventus.
— Taxa known from post-Sarmatian deposits: Cochlostoma septemspirale, Carychium pachychilus, Stagnicola
palustris, Zonitoides schraieri, Gyraulus verticillus, G. homalosomus, Vertigo angustior ocsensis, Argna suemeghyi,
Strobilops pappi, Fortuna tertia, Nordsieckia pontica, Mactrogastra cf. voesendorfensis, Tropidomphalus zelli depressus, Helicigona wenzi, Klikia kaeufeli, K. goniostoma, Dreissena dobrei, Pisidium pseudospherium, P. ex aff. nitidum, P.
ex aff. conventus.
— Still living, modern forms among the above mentioned post-Sarmatian taxa: Cochlostoma septemspirale,
Stagnicola palustris, and the four Pisidium species.
PALAEOECOLOGICAL, SALINITY AND FACIES QUESTIONS
The palaeoecological requirements of terrestrial gastropods may be estimated using the “closest living relative”
method, given that ecology is highly taxon dependent. The great majority of these gastropods live in or on forest litter,
on wet mossy rocks or under pebbles (e.g. Pomatias, Vertigo, Acanthinula). Some prefer wet meadows or grasslands
along shores or banks (Carychium). Others live on dry grasslands (Vallonia, Strobilops). Helicigona lives in dry woodlands, being a foliage eater. In riparian forests Triptychia is common, and there are some carnivorous taxa (Aegopis,
Poiretia, Palaeoglandina).
One of the most delicate questions is the estimation of the salinity. This may be connected not with just the aquatic
forms, but with species of the family Ellobiidae.These live close to sea shores, generally in tropical and subtropical areas,
except for two or three species. One species has been found in the Várpalota Ottnangian and 6 from the Lower Badenian
“Szabó” sandpit, evidently washed in from the shore. A new species was described from the Pusztamiske coal-seam bearing sequence. No data were found regarding how far from the seashore these taxa lived. Presumably this factor might
have been species-specific for these small gastropods, living on the litter of wet forests. From several tens of meters this
distance might attain a few kilometres, depending on the temporal marine inundations as well.
Assessing the salinity requrements of aquatic molluscs is even more difficult. The basic question is, what does the
term “freshwater mollusc” mean? Most molluscs of this type may tolerate a salinity of 3–4‰, or even a higher one.
However, this tolerance may be species specific within a given genus.
24
JÓZSEF KÓKAY
Forms living permanently in freshwater milieu should be regarded as freshwater taxa: Viviparus, Valvata,
Nematurella, Bithynella, Amnicola, Bithynia, Schuettemmericia, Stalioa, Stadtielliopsis, Sandbergerina, Emmericia,
Brotia, Melanoides, Galba, Stagnicola, Radix, Lymnaea, Aplexa, Anisus, Gyraulus, Hippeutis, Segmentina,
Planorbarius, Ferrissia, Ancylus, Pseudancylastrum, Margaritifera, Unio, Spherium, Pisidium, Dreissena.
Evidently, numerous species of the enumerated genera (and those of others, not occurring in the studied Miocene
fauna) may enter into waters of some salinity (the salinity is classified according to HILTERMANN 1966). Some of the most
striking examples are Bithynia tentaculata, and Radix ovata, which live even in pliohaline seawater. My observations
suggest that Bithynia glabra, which is close to B. tentaculata, was similarly tolerant towards salinity. Melaniidae (Brotia
and Melanoides) tolerate some ‰ of salinity, and so does Lymnaea stagnalis — up to 7‰. In the mio- to mezohaline
Pannonian Lake Viviparus, Valvata and Unio lived in masses. Congeria species, especially the small ones (C. basteroti,
C. amygdaloides, C. brairdii, C. sandbergeri, KÓKAY 1966), preferred mezo- to pliohaline environments, while the bigger ones lived rather in oligo- or miohaline waters, or in freshwater. Congeria species still live in freshwater. In the studied fauna the lymnaeid Galba dupuyana, which is a close relative of the extant G. truncatula, was genuine freshwater
form. This recent species lives in springs, small creeks and along banks of small streams. Bithynella species require similar conditions.
The salinity tolerance of the Theodoxus species varies significantly, depending on the particular species. For example Th. pictus never lived in freshwater, rather in mezo- and brachyhaline environments; this can be deduced from the
communities in which it is found. In contrast Th. crenulatus and its subspecies, preferred fresh- and oligohaline water.
Numerous modern Theodoxus species have been reported exclusively from freshwater. Stagnicola glabra, close to the
Miocene S. praebouilleti, is known only from freshwater. Some species of lymnaeids tolerate some permille of salinity.
Radix ovata is extreme in this respect, as it has been reported from water with a maximum — 14‰ of salinity (GLÖER et
al. 1980).
The planorbid Anisus planorbis tolerates slightly saline waters (SOÓS 1943). Planorbarius corneus is a close relative
of P. cornu and P. sansaniensis and is known exclusively from freshwater. Extant Gyraulus species are also regarded as
bound to freshwater. Accompanying species of fossil forms, however, seem to indicate that some Miocene Gyraulus
species tolerated oligo- or miohaline salinities. Segmentina is strictly a freshwater form. Similarly, the so-called capsnails (Ancylus, Ferrissia and Pseudancylastrum) strictly require freshwater. Amnicola pseudoglobulus and Emmericia
subpatula evidently remained freshwater forms, judging from their accompanying faunas and extant relatives. The
accompaniing species prove that Sandbergerina succineiformis lived exclusively in a freshwater environment. Several
Nematurella species appear in the “Brackwassermolasse” series, but they are still present on the North Alpine Freshwater
Molasse they are still present. Consequently these should be regarded as freshwater forms which tolerate oligo- to miohaline water.
The bivalve Margaritifera flabellata must be considered as fluvial — i.e. it is a freshwater species based on its extant
relatives. The European Unio species tolerate 3–5‰ salinity. The Pannonian taxa lived in a mio- to mezohaline environment, judged by their accompanying forms. Pisidium species are usually regarded as freshwater, but some forms tolerate 1–3‰ salinity (RICHNOVSZKY, PINTÉR 1979).
Regarding their salinity tolerance, Pseudamnicola species are the most doubtful. Possibly, some species (e.g. P. leognanensis) were able to get into freshwater, similarly to “Potamogyrus jenkinsi”, a species which immigrated from New
Zealand about a hundred years ago. This is a brackish water dwelling form, which tolerates a salinity up to 17‰, but is
regarded as a freshwater form by GLÖER et al. (1980). A similar case may also be supposed up to with respect to
Pseudamnicola leognanensis. However, it seems very probable that P. tholosa and P. immutata are freshwater forms,
capable of entering into a low salinity environment.
Evidently, we must expect that some fossils may have been washed in from environments different from that of the
sedimentation. If these sites differ significantly, the analysis does not pose difficulties. A typical case is when terrestrial snail shells are washed into marine or lacustrine deposits. Freshwater forms embedded into marine sediments may easily be identified. For example, the cycle closing layer from the drilling Várpalota Vk–384 well, in which a marine fauna
was mixed with freshwater elements, was most probably carried in by a river. The most problematical case might be if
freshwater forms were carried into an oligo- to miohaline depositional environment, making the assessment of the salinity almost impossible. A similarly difficult task is a reconstruction of an environment with quickly changing salinity and
slow deposition. In such cases a low salinity community might be mixed with a freshwater one. I regard the Hydrobia
species as tolerating, or even preferring an oligo-to miohaline environment. Considering all these difficulties, an attempt
will be made to reconstruct the ecological and salinity conditions of the above-described sites.
The base of the lower Badenian in the Nagygörbő Ng–1 locality should be regarded as of freshwater origin, although
some taxa might have tolerated a slight salinity. The presence of Characeae oogonia with opercula of Bithynia in some
pelitic intercalations further increases the probability of a freshwater environment. The Lower Badenian coal-seam bearing sequence of the Herend Basin contains coaly or lacustrine clay layers. The salinity of their depositional environment
may be analysed as follows: in the 184.0–188.5 m interval of the well He–13 there was a definite freshwater fauna:
Evaluation of the studied faunas from the Bakony
25
Bithynia glabra, Lymnaea sp., Segmentina larteti, Ferrissia deperdita, Discus? sp. Between 166.1–169.3 m: Brotia
escheri, Emmericia subpatula, Gyraulus sp., Planorbarius cornu mantelli, Vertigo sp. is similarly a freshwater association. A reddish alluvial mud from 215.9–219.8 m of the drilling He–20 also yielded also a freshwater fauna: Theodoxus
burdigalensis trifasciatus, Valvata moguntina, Amnicola pseudoglobulus, Brotia escheri, Emmericia subpatula,
Planorbarius cornu solidus, Characeae oogonia. These last mentioned remnants point to a freshwater milieu as well.
Another type of case was observed in the core 81.1–81.6 m of the well He–52, where a black coaly Brotia-bearing
clay yielded: Theodoxus barakovici, Valvata sp., Nematurella scholli, Pseudamnocola adiaphora parvula, Emmericia
subpatula, Brotia escheri, Ferrissia deperdita, Pisidium bellardii, P. steinheimense. This is a freshwater association,
although the species Theodoxus barakovici and Nematurella scholli were sometimes found with hydrobioids proving a
certain grade of salinity.
A clay from the 32.4–34.0 m interval of the drilling He–20 yielded Theodoxus grateloupianus, Nematurella pappi,
N. aquensis, Brotia escheri, Gyraulus microstatus, G. trochiformis kleini. The two Gyraulus and the Planorbarius point
to a freshwater milieu, while the rest of the forms could have tolerated a certain (oligohaline?) salinity. The entire association is regarded as a freshwater one.
The same drilling, from depth of 38.4–39.2 m, instead of Gyraulus and Planorbarius, contained Hydrobia ventrosa
and H. uiratamanensis shells, pointing to an oligohaline environment.
In the well Nyirt–1 at Nyirád a huminitic clay was found between 74.5 and 74.7 m, with Bithynia glabra, Brotia
escheri turrita, Galba dupuyana, Lymnaea sp., Gyraulus sp., Planorbarius cornu mantelli, and many Characeae oogonia. This is a typical freshwater community.
In the Tapolca Basin, the 157.0–157.7 m interval of the well drilling Hg–78/18 gave a rich mollusc fauna. Of the 20
aquatic forms 2 Hydrobia were identified, indicating a mio- mesohaline milieu, while the rest were freshwater forms; at
most some of them might have tolerated a slight salinity. According to Figure 4 of HILTERMANN (1966), this might indicate a salinity of 2–3‰.
The Várpalota hanging wall Upper Badenian alginite fauna has been mentioned above. In all probability this indicates a freshwater milieu. This is documented with the presence of Ancylus, Ferrissia, Gyraulus, Anisus, Planorbarius,
Unio and Pisidium taxa as well as with Potamogeton seeds, which are very frequent in some places.
CLIMATIC CONDITIONS
The general climatic conditions of the Early and Late Badenian have not been mentioned with the description of the
localities given that this topic is an extremely broad one (BÖHME 2003). It must be mentioned, however, that in the Lower
Badenian brackish and marine layers, the presence of a mangrove vegetation has been confirmed (NAGY, KÓKAY 1991).
This is further proof of a hot, tropical or subtropical climate, besides the coral reefs and other tropical and subtropical
faunal and floral elements. Coral reefs were present even in the Late Badenian sea.
In the studied fauna the following forms required warmth: Brotia, Melanoides, Laemodonta, Pedipes, Stolidoma,
Auriculastra, Opeas, and, probably, the Nematurella and Stalioa taxa as well — altogether 16 forms. Their disappearance at the Late Badenian/Sarmatian boundary might be best explained by a climatic deterioration. By this time an
important geohistorical event took place, namely the final closure of the connection between the Paratethys and the
Mediterranean (KÓKAY 1985). From this time on the Central Paratethys was only connected to the cooler and low salinity basins of the Eastern Paratethys. A cooling might mean that a former subtropical climate changed to a warm
Mediterranean type. The mentioned taxa could not survive this trauma and consequently they are missing from the
Sarmatian of the Western and Central Paratethys. The gastropod Brotia escheri s. l. is present in the Bulgarian Sarmatian
but this area was situated more to the south, thus warmer bays and lagoons might have existed there rather than in the
Pannonian area. The genus Brotia reappeared in the Pannonian area at the beginning of the Pannonian, together with subtropical vertebrates (e.g. Pongidae) and floral elements. This might have been due to the closure of connections with the
cooler Eastern Paratethys.
The disappearence of species of the nearshore family Ellobiidae cannot be explained by the decrease of the salinity
of the sea. Modern forms of this family live in mangroves and at the seaward sides of lagoons (the Auriculastra species,
described from Pusztamiske also lived along a lagoon.). Their vanishing may better be explained by the cooling of the
climate and by the disappearance of their main habitat, the mangrove.
Thus, at the Badenian/Sarmatian boundary there is a climatic break. Frequent and conspicious Badenian species such
as Brotia escheri are absent from the Sarmatian paludal, lacustrine or lagoonal deposits. At the same time the genus
Aplexa was present as it required being, less warmth.
Geologica Hungarica series Palaeontologica, fasciculus 56
27
SYSTEMATIC DESCRIPTIONS
Altogether 272 taxa were determined, including 251 gastropods and 21 bivalves. The figures for the taxa of the following periods are: Ottnangian 13, the Karpatian 4, the Lower Miocen 59, Lower Badenian yielded 58, the Middle
Badenian 22, the Upper Badenian 113 and the Sarmatian 142 taxa. 117 species and subspecies were aquatic, while the
number of terrestrial gastropods is 155.
The notions “palaeontological species” and “morphospecies” have to be emphasised as being in contrast to the biological species, although this problem is well known from textbooks. Not only transitional forms may be problematical,
but the palaeontological meaning of the term “population” inevitably differs from its use in biology. Shells found in one
single, even thin layer at a given locality might have belonged to animals living during a quite long interval (as the sedimentation might have been slow). In some cases a discrepancy of several thousand years might be assumed, enabling
significant changes in palaeoecological conditions. Consequently, the observed palaeontological “population” may lead
to erroneous deductions.
Making the case even worse, some new taxa were based only on fragmentary shells (e.g. the species of Melanoides)
instead of on the study of an “ideal” population.
Accordingly, taxa were regarded as different on the subspecies level if one or two important characters deviated and
on the species level if three or four traits were significantly different. If one or two characters diverged they were regarded as being of inferior importance and the form was regarded as a variety (“var.”).
For gastropods, the systematics of WENZ (1938–1944) was used, but in some cases the more up-to-date taxonomies
of LUEGER (1981), NORDSIECK (1981), PIECHOCKI, DYDUCH-F. (1993), BOETERS et al. (1989), KADOLSKY (1993) and others were taken into consideration. Only the more important synonyms were enumerated.
The bigger and more fragile fossils were cleaned by hand from the matrix, while the rest were washed. Moulds in
limestones and limy marls were cast with silicone.
New species, introduced in this work:
Ferebithynia nov. gen.
Conauricula nov. subgen.
Theodoxus grateloupianus trigonulus nov.ssp.
Viviparus pachystoma dissitus nov. ssp.
Sellia miocaenica nov. sp.
Stalioa praeglobulina nov. sp.
Stalioa globulina nov. sp.
Pseudamnicola (Staja) adiaphora parvula nov. ssp.
Staadtielliopsis bandensis nov. sp
Melanoides procurvicosta nov. sp..
Carychium (Carychiella) eumicron unidentata nov. sp.
Auriculastra (Conauricula ) badeniensis nov. sp.
Lymnaea stagnalis sarmatica nov. ssp.
Pseudancylastrum decorata nov. sp.
Vertigo pusilla sarmatica nov. ssp.
Vertigo bakonyensis nov. sp.
Argna (Aghardiella) sublamellata nov. sp
Enneopupa subcylindrella nov. sp.
28
JÓZSEF KÓKAY
Retinella applanata nov. sp.
Fortuna varpalotensis nov. sp.
Triptychia leobersdorfensis sarmatica nov. ssp,
Cochlodina (Miophaedusa) varpalotensis nov. sp.
Pseudidyla (Canaliciella) tricarinata nov. sp.
Canariella bakonyensis nov. sp.
Pisidium bakonyensis nov. sp
I would like to express my deepest gratitude to Dr. László Rákosi for permitting me to study palaeobotanical material (seeds), which was most useful in evaluating salinity conditions. My friend, Dr. Endre Krolopp helped me many times,
especially with suggestions regarding taxonomical questions and by pointing out significant and referent literature.
Classis: GASTROPODA
Familia: Neritidae
Genus: Theodoxus MONTFORT, 1810
Theodoxus grateloupianus (FÉRUSSAC), 1825
Plate I Figures 1–5
1874 Neritina Grateloupiana — SANDBERGER: p. 510, Pl. 25 Fig. 29
1896 Tripaloia? Grateloupiana — SACCO: parte XX: p. 52, Pl. 5 Fig. 61
1896 Tripaloia? Grateloupiana var. dertonensis — SACCO: parte XX: p. 53, Pl. 5 Fig. 62
1919 Neritina Grateloupiana — COSSMANN et PEYROT: Tome LXX: p. 54 Pl. 53 Figs 15–20
1929 Theodoxus (? Calvertia) grateloupianus — WENZ F. C.: p. 2951
1966 Neritina grateloupiana — STRAUSZ: p. 59, Fig. 35a, b, Pl. 49 Figs 21, 22
1967 Theodoxus grateloupianus dentatus — KÓKAY: p. 83, 89, Pl. 8 Figs 1, 2
Numerous specimens were collected from the uppermost Upper Badenian layers of the Hegymagos drilling as well
as from the Herend Upper Badenian. The whorls are well fitted, with a flat apical field. The columellar plate is strongly dentated, with furrows which often bifurcate. The shell is a unicoloured hazelnut brown or with triangular lightcoloured spots. Length: 6.2–9 mm, height 4.7–7 mm, H/L 76–82%.
Th. burdigalensis, which occurs abundantly at Pusztamiske, is easy to distinguish from this species (based on my
descriptions).
Specimens found in the Várpalota Sarmatian are bigger than the above-mentioned ones. The bentonite pit at Herend
exposed Brotia-bearing clays, from which spotted specimens originated large numbers (with small triangular spots and
with their blunt ends aperturally oriented). The upper Brotia-bearing limestone, found in the well Pusztamiske Pm–3,
was intercalated by an unconsolidated calcareous mud, yielding this species with Brotia escheri turrita, Hydrobia soceni,
Stagnicola praebouilleti, Lymnaea sp. and Gyraulus sp.
A specimen with a coloured pattern was found in the regressive section of the Várpalota Lower Badenian.
Earlier I distinguished a subspecies under the name “dentatus”, based on the dentated external aperture. However,
this view must be revised, as
— there are numerous transitional forms within a population (e.g. at Bánd, in the hanging wall of the bentonite),
— the literature also mentions dentated specimens without a subspecific distinction.
Therefore, I regard the subspecies Th. grateloupianus dentatus as synonymous with the nominal subspecies.
Theodoxus grateloupianus trigonulus nov. ssp.
Plate I Figures 6–7
Derivatio nominis: trigonulus (lat.) = trigonal
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 95
Locus typicus: Somlóvásárhely, drilling Svh–1, 188.2–189.0 m
Stratum typicum: Lower Miocene (Ottnangian to Karpatian) lime mud containing freshwater clay
Description: The height of the holotype is 5.8 mm and its width is 6.0 mm. It is a medium-sized shell, with a flat apical field. It widens strongly and consequently it is a triangular shape. On the whorl, on a brown base there are elongate
triangular light patches pointing backwards. The colour pattern and the densely grooved columellar plate are identical
with those on the nominal subspecies.
Systematic descriptions
29
Theodoxus burdigalensis burdigalensis (D’ORBIGNY), 1852
Plate I Figures 8–12
1883 Neritina burdigalensis — BENOIST: p. 381, Pl. 15 Figs 1, 2
1919 Neritina (Theodoxus) burdigalensis — COSSMANN et PEYROT: Tome LXX: p. 56, Pl. 8 Figs 25–28
This species is especially abundant in the Upper Badenian coaly clay intercalating with the seams in the drillings of
Pusztamiske as well as in the Nyirád drillings associated with Brotia escheri turrita. The species is also abundant in some
layers of the Upper Badenian bentonitic clay series of the Herend Basin, together with B. escheri turrita.
This is a relatively small form (length 7–8 mm, height 5.8–6.4 mm, H/L, 79–84%). The whorls are well closed, with
a sligthly convex apical field. The inner side of the external apertural margin is smooth and the columellar plate smooth,
but in some specimens the margin of the plate is slightly dentated. The specimens are dark brown and those originating
from coaly clays are blackish brown. After its accompanying forms it also tolerated freshwater environments.
The description of COSSMANN, PEYROT (1919) contradict their figures. On Figure 27 the columellar plate is slightly
but clearly dentated. Specimens in Figures 25–27 are unicoloured brown, instead of being patchy. In this way the
Hungarian specimens fully correspond to the figures of COSSMANN, PEYROT (1919); the latter originate from the
“Burdigalian–Helvetian” of the Aquitanian basin, roughly corresponding to our Karpatian and Badenian. The colouring
and the markings of the specimens from Herend are diverse. The most abundant ones are creamy white, the dark coloured
are more rare. Furthermore drop-shaped and helical markings also occur on them.
Its closest relative is Th. grateloupianus, but it is bigger, more elongate, and its columellar plate is strongly furrowed
with the furrows often being bifurcate.
Theodoxus burdigalensis trifasciatus (GRATELOUP), 1840
Plate I Figures 13, 14, 15
1840 Neritina fluviatilis — GRATELOUP: Atl. 1 Pl. 1 Figs 1, 2
1840 Neritina fluviatilis var. trifasciata — GRATELOUP: Atl. 1 Pl. 1 Fig. 3
The regressive closing layer of the Várpalota Lower Badenian (drilling VK–384) yielded one, slightly damaged specimen. It differs from the nominal subspecies in the following respects:
— it has more of a widening whorl, indicating fast growth; consequently, it is wider;
— it is a light ground colour and there is a zigzagging ornament arranged spirally.
Theodoxus barakovici (BRUSINA), 1902
Plate II Figures 1–8
1902 Neritodonta barakovici — BRUSINA: Pl. 14 Fig. 63–77
1929 Theodoxus (Calvertia) barakovici — WENZ F. C.: p. 2937
1966 Theodoxus crenulatus — KÓKAY: p. 32, Pl. 1 Figs 18–19
1979 Theodoxus (Neritaea) barakovici barakovici — JURIŠIĆ-POLŠAK: p. 17, 41, Pl. 1 Figs 13–16, 22
The species is frequent in the Herend Lower Badenian coal-seam bearing sequence and generally occurs in coaly
clays. The colour pattern the Hungarian specimens deviate from those of Brusina. This pattern is extremely variable in
Th. species, generally depending (quite consistently) on population (i.e. locality). Most of Brusina’s specimens from
Herzegovina are decorated longitudinally and bizonally, while most Herend specimens have a dark brown axial hachure
on a light brown ground. Most hachures are zigzagging with a smaller or bigger amplitude. About 5% of the specimens
have a clear bizonal pattern. Specimens found in layers of brackish origin (with Hydrobia) have a lighter colour.
Often the younger part of the spire slightly stands out and similar specimens occur in Herzegovina as well. The columellar plate is smooth in the Herend specimen, with the lower fourth of its rim being slightly turned outwards, sometimes with 2–4 small teeth.
The operculums are generally damaged and they seem to be slighly narrower than those figured on Brusina. In one
specimen the outstanding rib is preserved, its length is similar to that on Brusina’s figure. The diameter of the adult specimen is about 6–7 mm and it is similar to that of the Herzegovina specimens.
Being that most Theodoxus species are highly variable, it would seem unwise to base a new subspecies on this form.
The accompaniing fauna points to a freshwater or oligo- to miohaline milieu. Specimens originating from the last mentioned environment are generally bigger and the young part of their spire stands out more significantly than is the case
with the freshwater ones.
Brusina’s species is doubtlessly related to Th. pictus (Fér.), but it deviates due to its rounded outline, and by its
smooth or slightly dentated collumellar plate. Further, Th. pictus has a more complicated columellar edge, and a zonal
or scale-like colour pattern. On its operculum there are hornformed ridges. It did not live in freshwater.
30
JÓZSEF KÓKAY
So far it has not been reported from the Hungarian Miocene. Its palaeogeographical significance is, that it is known
from the Herzegovinan Karpatian and, possibly, the Lower Badenian.
Theodoxus cyrtocelis (KRAUSS), 1929
Plate II Figures 9–10
1929 Neritina cyrtocelis — KRAUSS: p. 145
1929 Theodoxus (Theodoxus) cyrtocelis — WENZ F. C.: p. 2991
1964 Theodoxus (Theodoxus) cyrtocelis — SCHLIKUM: p. 4, Pl. 1 Figs 3–6
1966 Theodoxus (Theodoxus) cyrtocelis — SCHLIKUM: p. 323, Pl. 12 Figs 1–5
1970d Theodoxus (Theodoxus) cyrtocelis — SCHLIKUM: p. 176, Pl. 3 Fig. 1
1989 Theodoxus (Theodoxus) cyrtocelis — REICHENBACHER: p. 154, Pl. 1 Figs 1–2
1997. Theodoxus cyrtocelis — PIECHOCKI: p. 105. Fig. 1
Specimens from the base of the Lower Badenian in the Nagygörbő–1 well belong here, as well as those from the
Lower Badenian coal-seam bearing sequence of Herend. On a brown ground there are irregularly placed spots and occasionally an elongated flow-pattern. On the upper part of the last whorl generally there is a slight rounded-angular part.
This is often absent from the Lower Miocene specimens. The columellar plate is arched, sometimes with 3–5 unconspicious folds. The accompanying fauna indicates an oligohaline or freshwater environments.
The species is well-known from the North Alpine “brackish” and “upper freshwater” deposits. It is also present in the
lowermost part of the Várpalota Ottnangian.
Theodoxus sinjanus (BRUSINA), 1884
Plate II Figure 11–13
1884 Neritodonta sinjana — BRUSINA: p. 82
1897 Neritodonta sinjana — BRUSINA: p. 28, Pl. 15 Figs 10–21
1929 Theodoxus (Calvertia) sinjanus — WENZ F. C.: p. 2980
1979 Theodoxus (Neritaea) sinjanus — JURIŠIĆ-POLŠAK: p. 19, Pl. 3 Figs 4–11
Some compressed specimens were found in the Lower Badenian base layers of the Nagygörbő–1 well. The edge of
the columellar plate is moderately dentated, but grooved and smooth ones are also present. The lower section of the columellar margin is slightly arched outwards. The colour pattern is variable, most frequently on a light- or coffee-brown
ground and there are axial and parallel dark-brown lines arranged densely or sparsely. Such specimens are similar to
BRUSINA’s (1897) Figure 14. There are specimens with a dense zigzag pattern similar to those on BRUSINA’s (1897) Figure
21. In several specimens it is visible that the originally axial lines turn to zigzag ones as growth advances; finally, the
pattern passes to a patchy one. A similar trend is visible on Brusina’s figures. Some specimens were unicoloured brown.
The height of the Nagygörbő specimens is generally 5–6 mm, but bigger ones (8–9 mm) were also found, similar in
size to Brusina’s ones.
Theodoxus sinjanus was originally described from the Middle Miocene freshwater deposits of Dalmatia.
Theodoxus crenulatus varpalotensis BARTHA, 1955
1955 Theodoxus (Calvertia) varpalotensis — BARTHA: p. 278
1956 Theodoxus crenulatus varpalotensis — BARTHA: p. 501, 502, Pl. 1 Figs 9–12
1967 Theodoxus crenulatus — KÓKAY: p. 77, Pl. 7 Figs 6–9
Bartha, describing the Várpalota Pannonian molluscs, erroneously included this new species into the Pannonian. He
studied specimens from a collection as is clear from his text. The material originated from the hangwall layers of the
Upper Badenian Várpalota coal-seam. In the dozen cm thick mollusc-bearing clay covering the seam, this species is often
present in masses, together with Congeria boeckhi and Ferebithynia vadaszi. It differs from the nominal subspecies in
having an outstanding apical (juvenile) part and in having, invariably, a marble-like zigzag colour pattern.
Familia: Cyclophoridae
Genus: Palaina SEMPER, 1865
Palaina martensi (ANDREAE), 1902
Plate II Figures 14, 15
1902 Adelopoma martensi — ANDREAE: p. 23, Fig. 11
1923 Palaina (Palaina) martensi — WENZ F. C.: p. 1764
1967 Palaina (Palaina) martensi — SCHÜTT: p. 200, Fig. 1
Systematic descriptions
31
Two damaged specimens were found in the lower part of the Várpalota Sarmatian. Their last one or two whorls are
missing. It is similar to the specimen — illustrated by SCHÜTT (1967) — from the Lower Sarmatian of the Vienna Basin.
ANDREAE (1902) described the species from Silesia (in Polish Opole; in German: Oppeln).
Genus: Cochlostoma JAN, 1830
Cochlostoma septemspirale (RAZUM.), 1789
Plate III Figures 2, 3
1897 Pomatias septemspiralis — WAGNER: p. 580, Pl. 2 Fig. 21
1983 Cochlostoma septemspirale — KERNEY et al.: p. 63, Pl. 1 Fig. 1
The upper part of the Lower Badenian coal-seam bearing series in Herend (Bánd–4 well) yielded one, almost entire
specimen. Unfortunately it is embedded in the matrix rather deeply and only its dorsal side is visible. Its height is 6 mm
and its width 3 mm. Together with the embrional one it is built of eight whorls and it has a dense axial rib system. The
ribs are sharp and the interspaces smooth.
Fragmentary specimens are frequent in the Várpalota Sarmatian with apex and whorl pieces being present.
Their form and ribs are identical with those of the Herend specimen. The aperture is encircled by a collar-like peristome, characteristic for the species, and its appearance in the calcareous areas of Southern Europe is recent. It is not mentioned in the literature as a fossil. The modern specimens are 7–8 mm high and 3.8 mm wide.
Familia: Viviparidae
Genus: Viviparus MONTFORT, 1810
Viviparus pachystoma dissitus nov. ssp.
Plate III Figures 4, 5
Derivatio nominis: dissitus (Lat.) = deviating
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 96
Locus typicus: drilling Somlóvásárhely Svh–1., 230,3–230,5 m
Stratum typicum: Lower Miocene (Ottnangian–Karpatian) freshwater clay, containing lime mud
Descriptio: Comparatively big shell. The holotype specimen is about 30 mm high and its width is about 25 mm. The
shell is damaged, but it is sufficiently complete to be able to state the differences between it and the nominal subspecies
V. pachystoma (Sandberger).
The shell consists of six step-like whorls. The upper part of the convex whorls are flat, especially in the juvenile specimens. On the whorls there are five strip-like spiral ribs of which the lowermost is the biggest. This is partly covered by
the subsequent whorls. Close to the aperture a sixth faint rib may be present over the lowermost one.
Diagnosis differentialis: The most important character differentiating this from the nominal subspecies is the presence
of the mentioned spiral ribs. The holotype specimen is bigger than Sandberger’s one, which is a maximum 24 mm high.
The new taxon is present throughout the Lower Miocene of Somlóvásárhely, generally as fragments. Its outline and
decoration is quite variable. In some specimens the juvenile whorls are almost angular. The top of the whorl is almost
horizontal and it joins the almost flat side at approximately a right angle. Some specimens have only four spiral keels, of
which the lowermost is the biggest. One specimen has much thicker ribs than the others.
FRANZ (1932: p. 42, Figure 30) found similar ribbed specimens in the lower part of the Lower Miocene (Aquitanian)
of the Mainz Basin. In his specimens the lowermost rib is the thickest, similar to that of the Hungarian ones.
Viviparus ex aff. sadleri (PARTSCH), 1856
Plate III Figure 1
1942 Viviparus sadleri — STRAUSZ: p. 38, 57, Pl. 1 Figs 1–9, 11–14, 20, 23–24
The lowermost part of the Nagygörbő Lower Badenian yielded one damaged specimen, without an apex and its posterior is visible. It seems to be close to the Upper Pannonian (Pontian) Viviparus sadleri and is widespread in the
Carpathian Basin. The shell of our specimen is smooth and it has not spiral cords.
Familia: Valvatidae
Genus: Valvata O. F. MÜLLER, 1774
Valvata moguntina (BOETTGER), 1884
Plate III Figures 6, 7
1884 Amnicola moguntina — BOETTGER: p. 276
1914a Pseudamnicola moguntina — FISCHER et WENZ: p. 126, Pl. 9 Fig. 45
32
JÓZSEF KÓKAY
1926 Amnicola (Amnicola) moguntina — WENZ F. C.: p. 2069
1932 Valvata moguntina — PEYROT: p. 11, Pl. 17 Figs 20–21
The species is present in the Várpalota Sarmatian, sometimes in masses. It conforms well with Boettger’s species, but
its growth-lines are weaker, and it is only visible under a significant magnification. The convexity and the number of
whorls, the depth of the sutures, and the shape of aperture coincide well. The axis of the aperture makes an angle of about
80° with the axis of the shell in both Boettger’s and the Hungarian specimens. The umbilical opening is also identical.
Average size: height 2.5 mm; width 2.2 mm. The width/height ratio varies between 80% and 100%. Specimens from the
Pusztamiske and Várpalota Upper Badenian are damaged and their status is doubtful.
WENZ (1926) mentions the species from the Lower Miocene of the Mainz Basin and France, while Boettger’s specimens are Middle Miocene.
Valvata sarmatica PAPP, 1954
Plate III. Figure 8
1954 Valvata (Turrivalvata) sarmatica — PAPP: p. 24, Pl. 4 Figs 3–4
1971 Valvata sarmatica — ŠVAGROVSKY: p. 228, Pl. 29 Figs 5–6
1974 Valvata sarmatica — PAPP: p. 333, Pl. 3 Figs 3–4
This small form was found in the Tapolca Basin and Pusztamiske Upper Badenian and in the Nyirád and Várpalota
Sarmatian.
PAPP (1954) described the species from the Sarmatian of the Vienna basin.
Valvata politioanei JEKELIUS, 1944
1944 Valvata politioanei — JEKELIUS: p. 55, Pl. 7 Figs 20–23
1959 Valvata politioanei — BODA: p. 633, Pl. 34 Fig. 4
Some samples from the Várpalota Sarmatian contain large numbers of this form. JEKELIUS (19944) described it from
the Banat (Eastern Rumania) Sarmatian.
Valvata moesiensis JEKELIUS, 1944
Plate III Figure 9
1944 Valvata moesiensis — JEKELIUS: p. 55, Pl. 7 Figs 11–14
1959 Valvata moesiensis — BODA: p. 632, 730, Pl. 34 Figs 1–3
The species, described from the Banat (Rumania) Sarmatian, was recorded by BODA (1959) from coeval formations
of Várpalota. It is similar to Valvata sarmatica PAPP, but it is bigger and its shell is thicker.
Valvata soceni wiesensis PAPP, 1954
Plate III Figure 10
1954 Valvata (Turrivalvata) soceni wiesensis — PAPP: p. 25, Pl. 3 Figs 23–24
1971 Valvata soceni wiesensis — ŠVAGROVSKY: p. 229, Pl. 29 Figs 7–16
1974 Valvata soceni wiesensis — PAPP: p. 333, Pl. 3 Figs 8–9
It is not rare in some samples from the upper part of the Várpalota Sarmatian. So far, it has been reported from the
Sarmatian of the Vienna Basin and of Slovakia.
Valvata homalogyra BRUSINA, 1874 var.
Plate III Figures 11, 12
1874. Valvata homalogyra — BRUSINA: p. 90
1897. Valvata hömalogyra — BRUSINA: p. 25. Pl. XIV. Figs 7–9
1928. Valvata (Valvata) homalogyra — WENZ F. C.: p. 2469
Thirtythree, mostly subjuvenile specimens, were collected from the lowest part of the Lower Miocene sequence of
the drilling Somlóvásárhely–1. The adult specimens have three whorls, which quickly increase in size with age. The shell
is quite flat, the sutures are deep and the umbilicus is deep and broad. The wall is thin, shiny and shows small growth
lines.
Brusina gives a 1.6 mm diameter and a 0.83 mm height for his specimen. The Somlóvásárhely ones are smaller: the
biggest ones are of 1.15 mm diameter and are 0.60 mm high; thus they are smaller than the other types. This may not be
Systematic descriptions
33
regarded as a specific character and thus the difference is designated with the word war “var.” Brusina described the
species he had from Sinj and Miočić and some other Dalmatian localities, which, according to JURIŠIĆ-POLŠAK (1979),
are „Helvetian” and Lower Badenian
Familia: Pomatiasidae
Genus: Pomatias STUDER, 1798
Pomatias bisulcatum (ZIETEN), 1846
Plate IV Figure 1
1845 Cyclostoma bisulcatum — THOMAE: p. 146, Pl. 4 Fig. 2
1847 Cyclostoma bisulcatum — KLEIN: p. 76, Pl. 1 Fig. 21
1923 Pomatias bisulcatum — WENZ F. C.: p. 1802
Some slightly compressed steinkerns originated from the Öskü–2 well, from the base of the Várpalota Ottnangian
sequence. These are not fully adult ones. They are adorned with spiral grooves and cords of differing size and this is characteristic for the species. THOMAE’s (1845) figure might represent even P. antiquum, (Brong.) while that mentioned by
LOCARD (1893) as “P. bisulcatum” probably belongs to P. consorbinum (Sandb.). It seems that these three forms are connected with transitions. P. bisulcatum is known from the Lower Miocene.
Pomatias turonicum WENZ, 1924
Plate IV Figure 2
1874. Cyclostomus turgidulus — SANDBERGER: p. 534. tab. XXVI. Fig. 23
1923. Pomatias turonicum — WENZ F. C: p. 1813
1923b. Pomatias turonicum — WENZ: (V.) p. 115
2002. Pomatias turonicum — BINDER: p. 163. Taf. 1. Figs 1–4; Taf. 10. Figs 2–4
The species has been found in the lower quarter of the marine Karpatian sequence of Várpalota (drilling V–219).
There are densely packed spiral ribs and acute growth lines forming a net. So far it has been found is the “Vindobonian”
(Manthelan) of Touraine, Central France, from the “Helvetian” of Württemberg (Germany), and from the Karpatium of
the Korneuburger Basin. A characteristic fragment was also detected in the Lower Miocene of the Somlóvásárhely–1
drilling.
Pomatias consorbinum (SANDBERGER), 1874
Plate IV Figure 3
1874 Cyclostoma consorbinum — SANDBERGER: p. 606, Pl. 29 Fig. 33a
1966 Pomatias consorbinus — KÓKAY: p. 33, Pl. 2 Fig. 4
1967 Pomatias consorbinum — SCHÜTT: p. 201
1976 Pomatias consorbinum — SCHLIKUM: p. 2, Pl. 1 Fig. 1
It has been recorded from the Herend Upper Badenian bentonitic layers. It is difficult to distinguish from Pomatias
bisulcatum because the spiral cords are very variable. The distinctive feature is that the cords are paired in P. bisulcatum.
Small specimens were found in the Várpalota Sarmatian as well. The species is present in the terrestrial Badenian and
Sarmatian as well as in the North Alpine “silvana”-bearing sequence.
Pomatias conica (KLEIN), 1853
Plate IV Figure 4
1853 Cyclostoma conicum — KLEIN: p. 217, Pl. 5 Fig. 14
1910 Cyclostoma (Tudora) conicum — ROLLIER: p. 133, Pl. 3
1981 Pomatias conica — LUEGER: p. 10, Pl. 1 Figs 11–12, Pl. 6 Fig. 3
An external mould was found in the Nyirád Upper Badenian freshwater limestone. The form is known in the North
Alpine coeval “silvana” layers and in the Vienna Basin Sarmatian and Pannonian. Fragmentary specimens were also
found in the Várpalota and in the W Bakony Middle Badenian and Sarmatian.
Familia: Hydrobiidae
During recent decades it was principally RADOMAN (1966, 1972, 1983) who documented (based on the anatomy of
extant hydrobiids) that shells may be morphologiaclly similar or almost identical remote taxonomic units. Evidently, the
34
JÓZSEF KÓKAY
present study might only be based on conchological features. In the near future, however, electronmicroscopic studies of
fine shell structure may help in clearing the taxonomic relationships of fossil hydrobiids.
The taxonomy of hydrobiids is often controversial and different authors, have different opinions. It is primarily
WENZ’s (1938–1944) system that is applied here, but there are more up-to-date works (e.g KABAT, HERSCHLER 1993,
SCHLIKUM 1960, 1961, 1965, 1968, 1970c and others). For instance the family Bulimidae and the genus Bulimus, based
on an ICZN opinion, is replaced by the names Bythiniidae and Bithynia, while the name Bulimus is used for a terrestrial snail belonging to the subclass Euthyneura. Schlikum uses the names Stalioa and Staliopsis; the last-mentioned one
is also used by ČTYROKY (1972). The subfamily position of Nematurella is dubious. KABAT, HERSCHLER (1993) use the
name Amnicola exclusively for the American species, while for similar European forms the name Pseudamnicola has
been proposed. There are, however, many European taxa, earlier ranged into Amnicola; these do not fit into the genus
Pseudamnicola. For these, I used the name Amnicola within quotation marks.
Subfamilia: H y d r o b i i n a e
Genus: Nematurella SANDBERGER, 1874
Nematurella flexilabris SANDBERGER, 1874
Plate IV Figure 5–7
1874 Nematurella flexilabris — SANDBERGER: p. 575, Pl. 20 Fig. 24
1893 Nematurella flexilabris — LOCARD: p. 202, Pl. 11 Fig. 4
1893 Nematurella Lugdunensis — LOCARD: p. 201, Pl. 11 Fig. 5
1926 Nematurella flexilabris — WENZ F. C.: p. 2007
1960 Nematurella flexilabris — SCHLICKUM: p. 203, Pl. 18 Fig. 1
The upper part of the Herend Lower Badenian coal-bearing sequence (an old opencast coal mine) yielded some well
preserved specimens. Characteristic for the species is the curved external labial swelling (“flexilabris”). In the studied
small palaeopopulation (Geological Institute of Hungary collection) a certain variability was observed in the
height/width ratio and in the relative height of the whorls. SANDBERGER (1874) described it from the “silvana” beds
(coeval with the Badenian).
Nematurella pappi SCHLICKUM, 1960
Plate IV Figure 8, 9
1960 Nematurella pappi — SCHLICKUM: p. 206, Pl. 18 Figs 4–5
1971 Nematurella pappi — SCHLICKUM: Pl. 3. Fig. 29
From the Brotia bearing cover of the upper Badenian (“Bánd”) bentonite of the Herend Basin well-preserved specimens were collected and these are identical with Schlickum’s species. It differs from N. flexilabris because although it
occurs in the same area (but in the coal-bearing level) it has:
— more convex whorls,
— deeper sutures,
— a smaller size,
— a right apertural edge.
Nematurella scholli SCHLICKUM, 1960 var.
Plate IV Figure 10–12
1960 Nematurella scholli — SCHLICKUM: p. 207, Pl. 18 Figs 6–7
1971 Nematurella scholli — SCHLICKUM: Pl. Fig. 4
It has been found frequently in the Lower Badenian coal-bearing level at Herend as well as at the eastern margin of
the basin: east of Márkó, from the well Máb–2, from the Lower Badenian thin coaly clay beds. The whorls are relatively flat and the shell is small. The specimens are at most 2.2 mm high, although SCHLIKUM (1960) mentions 2.9 mm high
specimens.
The types came from the “Viviparus” layers of the Bavarian Molasse, which corresponds roughly to the Ottnangian/
Karpatian boundary. It was found in freshwater and in low salinity layers.
Nematurella schuetti SCHLICKUM, 1960
Plate IV Figures 13–15, 16–17
1960 Nematurella schuetti — SCHLICKUM: p. 211, Pl. 19 Fig. 12
Systematic descriptions
35
One single specimen was found from the Upper Badenian part of the He–20 well in the Herend Basin, and from the
Pusztamiske Upper Badenian. This corresponds very well to Schlickum’s species, but is about half its size. To some
extent it is close to Nematurella aquensis as well. Schlickum recognised it from the Bavarian “Süssbrackwassermolasse”,
corresponding to the Upper Ottnangian and Karpatian.
Nematurella aquensis (DEGRANGE et TOUZIN), 1892 var.
Plate V Figures 1–11
1893 Stenotyra aquensis — DEGRANGE et TOUZIN: Tom VI: p. 183, Pl. 4 Fig. 7
1919 Bythinella aquensis — COSSMANN et PEYROT: p. 413, Pl. 16 Figs 37–38
1926 Stenotyra aquensis — WENZ F. C.: p. 2209
1960 Nematurella aquensis — SCHLICKUM: p. 206
Numerous specimens originate from the Upper Badenian coal-seam bearing sequence of Pusztamiske and from the
lower part of the Herend Upper Badenian (drilling He–20). The whorls are of low convexity and the suture is moderately deep. The apertural margin is thickened at the posterior of the aperture, giving a pillow-like appearance. This feature
is characteristic for the taxon. The degree of variation is high, especially regarding the form of the aperture and the proportions of the shell. In some specimens there is an umbilical slit. Some specimens coming from a depth of 53.6 m of
the well Pm–1 have a varix but transitions occur towards the smooth, typical form. The varices might be explained by
variations in the local ecological conditions. The Hungarian specimens are slightly smaller (2.0 mm high on average)
than those of COSSMANN, PEYROT (1919: 2.5 mm).
About two dozen shells were found in the Várpalota Sarmatian (drilling Inota–82). These are slightly smaller than
those from Pusztamiske. It occurred also in the Hidas (Mecsek Mts, SW Hungary) Middle Badenian coal-seam. It is
mentioned from the Aquitanian basin Lower Miocene.
Although SCHLICKUM (1960) placed it in the genus Nematurella but some of its characteristic features point rather to
the genera Stenothyra or Stenothyrella.
Genus: Sellia RAINCOURT, 1884
Sellia miocaenica nov. sp.
Plate VI Figures 1–10
Derivatio nominis: from the Miocene age
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 117
Locus typicus: Somlóvásárhely, drilling Svh–1, 230.3–230.5 m
Stratum typicum: Lower Miocene (Ottnangian to Karpatian) freshwater clay with lime mud
Descriptio: Height of the holotype 1.9 mm, width 1.3 mm. Height of the aperture 0.85 mm, width 0.8 mm. The small
shell has four and a half whorls, of which the first two and a half are convex. Further on, a slightly outstanding ridge does
appear on the upper third of the whorl. Beneath that the whorl becomes slightly convex and subparallel with the axis.
Over the ridge the upper field is straight or even slightly convex, inclining by 20 degrees towards the axis. This appearance give it a slight resemblance to a pagoda. The surface of the shell is covered with slight spiral lineations. At the base
there is a very narrow umbilical fissure. The aperture is oval and its upper margin is slightly pointed. The outer aperture
is sharp but does not thin outwards. It is quite outstanding at the mentioned ridge. The inner aperture is less sharp.
Variations within the population: in the Lower Miocene sequence of the drilling the species is not rare, especially in
the lower part of it. Altogether about 150 specimens were collected. In this set the following variability was observed:
The height/width ratio of the holotype is 1.46. There are wider and narrower specimens and generally their h/w ratio is
between 1.4 and 1.6. The ridge or keel on the whorls may be sharply margined in some specimens. The aperture may be
oval and elongate; in other cases it may be pointed at the upper margin, but it may even be round. Generally, the grade
of the slight longitudinal (spiral) lineation decreases remarkably going upwards in the sequence (i.e. with geological
time). In some specimens, originating from upper levels, the shell may be completely smooth.
Diagnosis differentialis:The, morphologically similar genus Pyrgula generally has no umbilicus and thus the identity with it may be ruled out. Neumayr suggested that the genus Pyrgula may have its origin in some Pannonian Hydrobia,
demonstrating this change with a broad set of transitional forms. The new form may be a late descendant of Sellia pulchra RAINCOURT, 1884 (see WENZ 1923–1930. p. 2040), known from the freshwater mid- and upper Eocene of France
and England. Raincourt has described the only known species of Sellia from the “Middle Sand” of the Paris Basin. Sellia
pulchra is quite similar to the Somlóvásárhely form. The most important differences are: the older species is bigger
(height 5.5 mm); the spiral ridge is similar in them, but the whorl beneath the ridge is not parallel to the axis in the Eocene
form. The shell of the Eocene form is subconical. The ridge runs along the middle of the whorl. Upward from it, the outline forms an angle of 45 degrees with the axis. The number of the whorls is 6.
36
JÓZSEF KÓKAY
Probably Hydrobia wenzi BOETTGER (WENZ 1923–1930. p. 1953), from the oldest Miocene of the Mainz Basin, may
belong to the same genus. It is slightly bigger than the Somlóvásárhely species. The whorls are decorated with lines and
in the middle there is a ridge. Beneath it, the lower part of the whorls is slightly conical and their upper part forms a 70degree angle with the axis. There is no transition to any of the Hydrobia species in the same association.
Genus: Microprososthenia KADOLSKY–PIECHOCKI
Microprososthenia belchatovensis KADOLSKYET–PIECHOCKI, 2000
Plate VII Figures 1–7
2000 Microprososthenia belchatovensis — KADOLSKY et PIECHOCKI: p. 219. Figs 1–4 Pl. 1. Figs 1–2
Many specimens of this small-sized snail were found in the freshwater Lower Badenian of Herend. Most of these are
damaged. Its shape is hydrobioid, number and it has four and a half to five whorls. Under a high magnification (SEM) a
granulation is visible on the embrional whorl. On some specimens there is a spiral groove. Umbilical fissures are hardly
visible or not present at all.The specimen from Poland is slightly bigger, its height is 1.5 to 2.2 mm, while the Herend
specimens are 1.5 to 1.75 mm high. The Polish locality (Belchatów) is of freshwater origin and its age corresponds to the
Karpatian and Lower Badenian.
Genus: Martinietta SCHICKUM, 1974
Martinietta tumida KADOLSKYET–PIECHOCKI, 2000
Plate VII Figures 8–11, Plate VIII Figure 1
2000 Martinietta tumida, KADOLSKY et PIECHOCKI, p. 221. Figs 6–12. Pl. 1. Fig. 3
Several specimens of this very small, hydrobioid shell were found in the Lower Badenian freshwater sequence of
Herend. It has four and a half slightly convex whorls. Under SEM it can be seen that the embrional whorl is granulate.
It is identical with the species found in Poland, in a freshwater series coeval with the Hungarian Karpatian and Lower
Badenian. The height of the Polish specimens is between 1.2–1.6 mm, while the ones form Herend are 1.3 to 1.5 mm
high.
Genus: Bythinella MOQUIN et TANDON, 1856
Bythinella eugenii JEKELIUS, 1944
Plate VIII Figure 2
1944 Bythinella eugenii — JEKELIUS: p. 62, Pl. 12 Figs 1–3
1959 Bythinella eugenii — BODA: p. 636, 733, Pl. 35 Fig. 1
The species was described from the Banatian (Rumania) Sarmatian. 15 specimens coming from the freshwater Upper
Badenian of the Hegymagos well are identical with the type.
The Hungarian shells are slightly smaller than Jekelius’ form. It is present in the Herend Lower Badenian coal-seam
bearing series. BODA (1959) recognised it in the Várpalota Sarmatian.
Genus: Pseudamnicola PAULUCCI, 1878
Pseudamnicola tholosa JEKELIUS, 1944
1944 Pseudamnicola tholosa — JEKELIUS: p. 64, Pl. 12 Figs 15–19
1959 Pseudamnicola tholosa — BODA: p. 636, 734, Pl. 35 Figs 2–3
1971 Pseudamnicola tholosa — ŠVAGROVSKY: p. 248, Pl. 36 Figs 1–6
The species, introduced by Jekelius from the Banat Sarmatian, was found in the Upper Badenian of Pusztamiske and
Várpalota as well as in the Várpalota Sarmatian.
The specimens — described as “Hydrobia (Pseudamnicola) friedbergi” (KÓKAY 1966) from the Lower Badenian limnobrackish layers at Herend — seem to be close to this species, but it is smaller and the whorls are less convex.
Systematic descriptions
37
Pseudamnicola steinheimensis (MILLER), 1900
Plate VIII Figures 3–7
1900 Bythinella steinheimensis — MILLER: p. 401, Pl. 7 Fig. 23
1920 Pseudamnicola pseudoglobulus forma steinheimensis — GOTTSCHICK: p. 187, Fig. 8c
1922 Amnicola pseudoglobulus steinheimensis — WENZ: p. 151, Fig. 84
1923 Pseudamnicola steinheimensis — WENZ F. C.: p. 2078
It is frequent in some layers of the upper part of the Várpalota Sarmatian. It has the follolowing dimensions: height,
1.2–1.9 mm; width, 0.9–1.2 mm. It is close to Jekelius’ form P. tholosa. This latter differs from Miller’s species in being
slightly bigger, having flatter whorls and, accordingly, the sutures are shallower, at least judging by JEKELIUS’ (1944) figures. The separation of the two forms seems to be dubious. P. steinheimensis has so far only been found at Steinheim.
Pseudamnicola leognanensis (COSSMANN et PEYROT), 1918
Plate VIII Figures 8, 9
1918 Pseudamnicola leognanensis — COSSMANN et PEYROT: p. 411, Pl. 16 Figs 55–57
This has been found at Pusztamiske in the coal-seam bearing coaly clay and is of high quantity. This Hydrobia-like
form is very close to that described from the Aquitanian basin. Its authors did not describe the variability of the species,
but their three figures reveal changing values of the height/width proportion, of the convexity of the whorls, and variations in the form of the aperture.This traits change in a similar way in the Pusztamiske specimens as well. The height is
between 2.5–3.0 mm and there are five whorls. Some lower and wider specimens are morphologically very close to the
species Pseudamnicola convexa convexa (SANDB.), which occurs in the Silvana layers.
Pseudamnicola inflata JEKELIUS, 1944
Plate VIII Figures 10, 11
1944 Pseudamnicola inflata — JEKELIUS: p. 64, Pl. 12 Figs 20–23
Two specimens were found in the Várpalota Sarmatian (M. III Shaft) and these are close to the ones of Jekelius’. The
shell from the Pusztamiske Upper Badenian coal-seam bearing series (well Pm–1) is also similar to the Rumanian taxon.
The observed deviations are:
— the Hungarian shell is stumpier and not so high;
— the whorls are rounded and angular instead of a blown-up convex type;
— the axial decoration is denser and finer.
It could belong to a new subspecies, although the variability of the form is unknown.
Pseudamnicola immutata (FRAUENFELD), 1856
1944 Pseudamnicola immutata — JEKELIUS: p. 62, Pl. 12 Figs 4–7
1966 Hydrobia immutata — KÓKAY: p. 34, Pl. 2 Fig. 10
1971 Pseudamnicola immutata — ŠVAGROVSKY: p. 250, Pl. 36 Figs 7–9
This form generally occours in limnobrackish facies, but probably tolerates the freshwater as well. It is frequent in
the Sarmatian, but also occurs in the Herend Lower Badenian.
Subgenus: S t a j a BRUSINA, 1897
Pseudamnicola (Staja) adiaphora parvula n. ssp.
Plate VIII Figure 12
Derivatio nominis: “parvula”, tiny
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 99
Locus typicus: Herend, well 52, 81.1–81.6 m depth
Stratum typicum: Lower Badenian coaly clay
Descriptio: Its appearence is “viviparoid” and the shell is moderately thick. There are four whorls and moderately
deep sutures. The shell surface is smooth, even shiny, with slight growth-lines. The height and width of the aperture are
identical, forming a rounded square (=„rotundo-quadrangularis” lat.). The vertical part of the backturning covers the narrow umbilical fissure. The upper section of the columellar margin thick and approximately horizontal. The sharp lower
edge of the aperture forms a blunt angle with the columellar one. The, similarly sharp, external margin forms a right
angle with the upper columellar one. The second specimen (paratype) lacks its apex — otherwise it is identical with the
38
JÓZSEF KÓKAY
holotype. Dimensions of the holotype: height 1.3 mm; width 0.9 mm; the width and height of the aperture are equally
0.6 mm.
Diagnosis differentialis: The species is close to Pseudamnicola (Staja) adiaphora BRUSINA (1897), described from
the Upper Pannonian of Slovenia. Brusina’s specimen is twice as big as the Hungarian one and its aperture is higher. The
external margin forms a sharp angle with the upper columellar one (BRUSINA: 20, Pl. 14 Figures 10–11).
Subfamilia: A m n i c o l i d a e ?
Genus: Amnicola? GOULD et HALDEMAN, 1841
“Amnicola” punctum (EICHWALD), 1853
1856 Paludina partschi — HÖRNES: p. 588, Pl. 47 Fig. 24
1923 Amnicola? (Amnicola) punctum — WENZ F. C.: p. 2079
1966 Hydrobia punctum — STRAUSZ: p. 65, Pl. 47 Fig. 9
Small specimens originate from the coal-seam bearing section of the well Pusztamiske Pm–3. Based on their character, they belong to the genus Amnicola rather than to Hydrobia. Amnicola might have lived in fresh- or oligohaline water.
It is known from Badenian and coeval formations.
“Amnicola” pseudoglobulus (D’ORBIGNY), 1853
Plate VIII Figure 13
1847 Paludina globulus — KLEIN: p. 88, Pl. 3 Fig. 13
1874 Gillia utriculosa — SANDBERGER: p. 635, Pl. 28 Fig. 1
1900 Gillia utriculosa var. elongata — MILLER: p. 401, Pl. 7 Fig. 22
1922 Amnicola pseudoglobulus pseudoglobulus — WENZ: p. 151, Fig. 83
1926 Amnicola (Amnicola) pseudoglobulus pseudoglobulus — WENZ F. C.: p. 2076
1964 Amnicola pseudoglobulus — SCHLICKUM: p. 8, Pl. 1 Figs 16–18
1998 Pseudamnicola pseudoglobulus — FINGER: p. 15. Taf. 5. Fig. D–E
Two specimens were found in the Herend Lower Badenian coalseam-bearing sequence (well He–20). One of them is
slightly damaged, the other one more so; both are close to the type specimens. The species has been found in the Lower
Miocene of the Somlóvásárhely drilling.
This form has also been found in the Bavarian “Upper Brackish” and “Freshwater Molasse” as well as from the
Steinheim layers.
Subfamilia: B i t h y n i i n a e
Genus: Bithynia LEACH, 1818
Bithynia glabra (ZIETEN), 1830
Plate VIII Figures 14–19, Plate IX Figures 1, 2–8
1847 Cyclostoma glabrum — KLEIN: p. 77, Pl. 1 Fig. 22
1874 Bithynia gracilis — SANDBERGER: p. 561, Pl. 28 Fig. 16
1928 Bulimus glabrus — WENZ F. C.: p. 2239
1967 Bithynia gracilis — KÓKAY: p. 85, Pl. 8 Fig. 8
1976 Bithynia (Bithynia) glabra — SCHLICKUM: p. 4, Pl. 1 Fig. 8
I mentioned this species earlier under the name “Bithynia gracilis”, which is a synonym of Bithynia glabra, according to SCHLICKUM (1976). In the Nyirád wells it is frequent in the Upper Badenian limestone (“Bithynia limestone”). In
the Pusztamiske drillings it occurs in the Upper Brotia bearing limestones and clays. The oldest Sarmatian sequence
transgrades gradually over these Badenian sequences. The lowermost (some decimetres thick) sequence contains this
species but it decreases upwards in number and dimensions. It is accompanied by the gradual appearance of Sarmatian
molluscs. The opercula are well preserved and they show up spiral (excentric) nuclei.
This species was found in the base layers of the Várpalota Upper Badenian coal measures and in the Sarmatian, generally as opercula. The species is frequent in the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) sequence.
Bithynia glabra subgracilis (ŁOMNICKI), 1886
Plate IX Figures 9–12
1886 Bithynia subgracilis — ŁOMNICKI: p. 72, Pl. 2 Fig. 26
1923 Bulimus glabrus subgracilis — WENZ F. C.: p. 2243
Systematic descriptions
39
A high number of fragile specimens were found in the well Nagygörbő Ng–1. Entire specimens are rare, but well preserved opercula are frequent. The specimens, especially the subjuvenile ones, are quite close to the nominal subspecies.
The whorls are similarly convex, and the sharp margin and the outline of the aperture are also similar, but their dimensions (5–7 mm) are superior to those of the Nagygörbő specimens. The grown-up specimens from Nagygörbő are superior to those of the nominal subspecies, and even to those from Galicia, being 8 to 10 mm high. The opercula from
Nagygörbő have lesser or bigger spiral (excentric) nuclei and their total size is greater than that of the nominal subspecies.
Some specimens do not show the spiral nuclei. The concentric growth-lines are generally quite fine, but may be strong
as well. The opercula are generally thin and sheet formed, but some are quite thick and hollowed. Łomnicki did not draw
or describe the opercula. Therefore, there is a certain degrea of doubt about the Nagygörbő specimens still exists. There
are some fragmentary shells which displaying an extremelyfine spiral groove system under a strong lens. These specimens may belong to a separate species or may represent an extreme variety.
The species occurs in the Galician “Tortonian”, i.e. in the Badenian.
Bithynia dunkeri (GUDE), 1913
Plate IX Figure 13
1851 Paludina ovata — DUNKER: p. 159, Pl. 21., Figs 10–11
1874 Bithynia ovata — SANDBERGER: p. 560, Pl. 28, Fig. 17
1893 Bithynia ovata — LOCARD: p. 200, Pl. 11, Fig. 13
1926 Bithynia dunkeri — WENZ F. C.: p. 2236
1966 Bithynia dunkeri — SCHLICKUM: p. 324, Pl. 12, Figs 19–21
1989 Bithynia dunkeri — REICHENBACHER: p. 151–152, Pl. 1, Figs 6–7
From the Nagygörbő well, two specimens of this form were detected characteristic for the North Alpine Freshwater
Molasse (“Süssbrackwassermolasse”) series. It was found in a 10 cm thick layer, accompanied by specimens of Bithynia
subgracilis. One of the specimens is excellently preserved. It may be differentiated from Bithynia subgracilis by its shallower sutures and by its less convex whorls. The quick growth is indicated by a shell dominated by the last whorl. The
total height is 9 mm while that of the last whorl is 7 mm. A big, 11 mm high specimen of Bithynia subgracilis, found
together with the mentioned form, has a similarly 7 mm high last whorl. No transitional forms were found between the
species.
In the literature Bithynia dunkeri is characterised as having a dense array of axial growthlines. According to DUNKER
(1851) these are rudimentary, while LOCARD (1893) describes them as occurring on the last whorls. On the Hungarian
specimen these are only visible under a lens on the dorsal side of the last whorl.
Ferebithynia nov. gen.
Derivatio nominis: Fere (lat.), almost (Bithynia)
Typus species: Bulimus vadaszi WENZ
Locus typicus: Geresd, Mecsek Mts
Stratum typicum: freshwater clay, Karpatian, Lower Miocene
Descriptio of the type specimen: oval, elongated shell, and a pointed apex. There are 5 ½ whorls, initially gradually
increasing in size, and then getting bigger. The last whorl makes about two-thirds of the total height. The whorls are moderately convex and they are delimited by a well-developed suture. The first (initial) whorl is smooth while the subsequent
ones are covered with a fine dense spiral hachure, visible under strong magnification. The last one and a half whorls are
almost smooth but the decoration is completely invisible even under a strong lense. The last two whorls bear slightly
curved growth-lines.
The aperture is egg-shaped, its upper edge is pointed and the apertural margins continuous. The pillar edge and the
umbilical fissure is widely covered by the aperture.The external part of the aperture is thin and folded outwards and it is,
ring-shaped in some specimens. Its upper part is curved upwards. The opercula are yellowish pale and horny in
appearence. They are of spiral (excentric) construction and the growth-lines are concentric near to the margin. These are
concave, and droplet shaped.
The dimensions of the described specimen are the following: height 9.5 mm, width 5.8 mm, height of the aperture,
5.3 mm, width of aperture — 4.0 mm.
Diagnosis differentialis: WENZ (1931) described the species “Bulimus vadaszi” from the Karpatian of the Mecsek
Mts, mentioning its presence in the Várpalota “Tortonian” (Upper Badenian) coal-seam cover beds. It was first given the
name “Bulimus”, but later a decision of the ICZN transferred this name to that of a terrestrial snail taxon, and renaming
it “Bithynia”. A closer study of the specimens of Wenz’s form revealed that it does not conform with the genus Bithynia
given that:
40
JÓZSEF KÓKAY
— there is a spiral lineation on the shell surface, especially on the upper whorls;
— the aperture margin is thick, especially at the newel (columella);
— the operculum has an excentrically placed nucleolusis which is definitely spiral in its structure (this is uncommon
in Bithynia species).
Thus, Wenz’s species may not be placed in the genus Bithynia.
The traits of the form ruled out placing it among any of the closely related genera:
The structure of the operculum is close to that of the genus Neumayria, but the characteristic small pitches — which
look like strikes from a hammer (“malleata”) are lacking from the shell surface, and the apertural margin also deviates.
The genus Gabbia also has a spiral operculum, but these species are low and small, with a smooth shell surface. The
Digoniostoma species are generally spirally lineated, but have umbilical fissures; furthermore, their opercula are concentric. The genus Parafossarulus similarly deviates from the proposed new genus. Hydrobioides forms have a smooth shell
surfaces, their apertures are thickened and ring-like, externally there is a varix, and they have concentric opercula (WENZ
1938, 1944).
The above-mentioned facts warrant the introduction of a new genus, but its name should reflect its close relationship
to Bithynia.
Ferebithynia vadaszi (WENZ), 1931
Plate X Figures 1–3, 4–7
1931 Bulimus vadaszi — WENZ: p. 118, Pl. 10 Fig. 2
1966 Bithynia vadaszi — STRAUSZ: p. 66, Pl. 47 Figs 17, 18
1967 Bulimus vadaszi — KÓKAY: p. 85, Pl. 9 Figs 1–7
1991 Bithynia vadaszi — KÓKAY: p. 52
As has been mentioned, in the description of the new genus, the species was described by WENZ (1931) from
Karpatian freshwater (or, possibly, of very low salinity) deposits of the Mecsek Mts, with a remark about its occurrence
in the “Tortonian” (=Badenian) coal-seam cover beds at Várpalota. On the juvenile whorls the spirally lineated surface,
as well as the thickened aperture, is more pronounced in the Mecsek specimens than in the Várpalota ones. The spiral
lineation is variably developed in the Várpalota specimens; on some shells, especially those coming from shoreline sandy
deposits, it is barely visible. Specimens form the interior of the basin generally have flatter whorls, while those found in
sandy shore deposits have more convex ones, even a zigzag; a spiral colour pattern is visible on some of the specimens
(KÓKAY 1987). The opercula usually show up the mentioned spiral growth-lines and the excentric nucleoli. In some cases
these features remain invisible. This explains why WENZ (1931) described the Mecsek opercula as having concentric
growth-lines. Opercula from Geresd (Mecsek Mts), preserved in the collection of the Hungarian Geological Institute,
also have spiral ornamentation. These are elongate, and droplet like.
Subfamilia: E m m e r i c i i n a e
Genus: Schuettemmericia SCHLICKUM, 1961
Schuettemmericia paulensis (DEGRANGÉ et TOUZIN), 1893 var.
Plate X Figures 8–10, Plate XI Figures 1–2
1893 Fossarulus paulensis — DEGRANGÉ et TOUZIN: p. 187, Pl. 187, Fig. 9
1919 Stalioa paulensis — COSSMANN et PEYROT: p. 418, Pl. 16, Figs 53, 54, 60–62
1926 Stalioa (Stalioa) paulensis — WENZ F. C.: p. 2188
1993 Schuettemmericia paulensis, KADOLSKY: p. 352. Figs 14–16
One well preserved adult and another subjuvenile specimen were found from the uppermost Upper Badenian deposits
of the well Pusztamiske Pm–1, as well as from the Lower Badenian of the drilling Herend 40. Taking into consideration
the variability of the species, the Hungarian form agrees well with the French one but is about half its size. Two well preserved specimens were found in the Lower Badenian coal-seam bearing series of the well Herend 20. One of these deviates in that it has a spiral lineation on its shell. This points to a similarity with the closely allied Stalioa lemani (Bast.)
which is, however, of greater height and has more whorls (COSSMANN et PEYROT 1919).
The species is recorded as being from the Langhian, and it corresponds to the Lower Badenian mentioned by
Cossmann and Peyrot. SCHLICKUM (1961: 62) further substantiated the possibility that the Lower Miocene S. subpyrenaica and S. paulensis may be the evolutionary precursors of the genus Emmericia. This appears (according to the mentioned authors) in the Pliocene and their opinion was based on the form of the shell and on the outward turned apertural margin. This apparently plausible proposition must be rejected, because:
Systematic descriptions
41
— the Lower Badenian of Herend yielded Emmericia subpatula (KÓKAY 1966);
— the species Emmericia pliocaenica Sacco (see BARTHA 1959) known from the Hungarian Pannonian, is in fact Late
Miocene instead of Pliocene, S. paulensis is of Langhian age and corresponds to the Early Badenian.
Thus, the genera Emmericia and Stalioa lived concurrently in the Badenian and they were even found together in
Herend. However,their close evolutionary relationship seems not to have been proven.
Genus: Stalioa BRUSINA, 1870
Stalioa globulina nov. sp.
Plate XI Figures 3–4, 5–11, Plate XII Figures 1–2
Derivatio nominis: globulina (lat.) = globule.
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 98
Locus typicus: drilling Pusztamiske Pm–3, 58.6–59.3 m
Stratum typicum: Upper Badenian coaly mollusc bearing clay
Dimensions: height of holotype, 1.7 mm, width, 1.5 mm
Descriptio: The small shell is quite thick-walled and looks as if it is built of small globuli. The spire is short, not steplike and it consists of 3½–4, „quickly growing” convex whorls separated by deep sutures. The shell is shiny and even
under a strong magnifying lens weak growth-lines are barely visible. The subsphaeric last whorl dominates the shell. At
the base there is a deep umbilicus of moderate size. The aperture is asymmetric, elliptic, and the external margin is more
curved than the inner one. The longitudinal axis of the aperture makes a 45° angle with the shell axis. The outer and
upper labium is turned outwards, the lower one less so, while the inner margin is straight and without a rim. On the external lip there is a slight elevation (“bourrelet”), which is visible only with tangential illumination. A narrow callus is present between the lower apertural margin and the left side of the umbilicus. Similarly, this is also exclusively visible with
appropriate illumination. The embrional whorl is strongly granulated.
Variatio populationis: almost 100 specimens were investigated. The above described species marks are not very variable. The spira height is 1.5–1.7 mm and its width 1.3–1.5 mm. The most variable feature is the form of the aperture: it
might vary from a crescent (in this case the columellar margin is strongly curved) to an oval outline. In some specimens
the upper ends of the external and internal margins meet at an angle. In other specimens the lower margin is bent outwards as well, but in some the upper margin resembles a “pouring lip”. A week peristomal elevation outward from the
external lip is present only in some adult specimens.
Diagnosis differentialis: The new species is the closest to the above-described S. paulensis (DEGRANGÉ et TOUZIN,
1893). The main differences are:
— The columellar lip is not outward turned and it does not cover the wide umbilicus.
— The last whorl is dominant over the shell and its subsphaerical form is conspicious.
— In S. paulensis the callus is lacking between the lower margin of the aperture and the left side of the umbilicus.
Specimens of the new taxon are of a bigger.
Stalioa praeglobulina nov. sp.
Plate XII Figures 3–4, 5–12, Plate XIII Figures 1–2
Derivatio nominis: praeglobulina (Lat.) = an ancestor of the species globulina
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 97
Locus typicus: Somlóvásárhely, Drilling Svh–1, 192.6 m
Stratum typicum: Lower Miocene (Ottnangian to Karpatian) freshwater clay
Descriptio: Size of the holotype: height 1.0 mm, width, 1.3 mm.
This is a small “valvatoid”shell, built of three and a half globules. The last whorl is significantly bigger than the previous one indicating to a fast growth. The shell is step-like, with convex whorls. The sutures are not very deep. On the
wall, especially on the adult part, there are growth lines. The embrional whorl, under high magnification (SEM), shows
granulae, or rather it is covered with small pits irregularly. The umbilicus is wide and deep. The columellar part of the
aperture is rounded while the rest is sharp and slightly bent outward, especially on the upper segment. Its outline is elliptical and on the top it is slightly pointed. From the lowest edge of the columellar aperture a narrow callus or fold is present, leading to the left side of the umbilicus.
Variatio populationis: Six adultive and several subjuvenile specimens are in the collection. From the adultive specimens the umbilicus is free of matrix in three. Apart from 1 to 2 mm deviations, the specimens are almost uniform in
shape.
Diagnosis differentialis: The new species is the closest to the above mentioned S. globulina form — in fact, it is the
42
JÓZSEF KÓKAY
evolutionary predecessor of it. The older taxon differs from the younger one in the following ways:
— it is smaller and wider: the height/width ratio is 1.133. This ratio for the holotype is 0.769. This is the main difference between the two species. The older Somlóvásárhely species is significantly wider, than the younger Pusztamiske one,
— the umbilicus is wider;
— the callus is more developed;
— the aperture is more round;
— on the outer margin of the aperture there is no pillow like protuberance.
Genus: Staliopsis RZEHAK, 1838
Staliopsis moravicus ŘZEHAK, 1893
Plate XIII Figure 3
1893 Staliopsis moravica — ŘZEHAK: p. 171, Pl. 2 Fig. 23
1972 Staliopsis (Staliopsis) moravicus — ČTYROKY: p. 67, Figs 5, 6, Pl. 2 Fig. 1, Pl. 3 Figs 1–3
A badly damaged specimen came from the lowest part of the Várpalota Ottnangian, from a freshwater coaly clay.
Hitherto it had been known only from the Western Paratethyan “Rzehakia”-bearing Upper Ottnangian.
The steplike whorls are entoured by spiral cords. {The identification was kindly checked by P. Čtyroky (Prague), who
is the best specialist of the Rzehakia beds.}
Staliopsis puisseguri (SCHLICKUM), 1964
Plate XIII Figures 4–6
1964 Euchilus dehmi, SCHLICKUM: p. 9. pars
1965 Euchilus (Staliopsis) puisseguri, SCHLICKUM: p. 103. Figs 2–3
The species is not rare in the Lower Miocene of the Somlóvásárhely drilling, but very few entire specimen have been
found. Its size is identical with Schlikum’s form. The thickening behind the external aperture is also consistent in them.
The Hungarian specimens deviate from Schlickum’s ones in having generally smooth shells; the rudimentary spiral lineation is only present on some specimens. Taking into consideration the fact that the members of the genus are highly
variable even within one and the same population, this deviation does not contradict the identity of the German and
Hungarian forms; Schlickum described his species from the Bavarian Upper Freshwater Molasse — that is from layers
more or less coeval with the Somlóvásárhely ones.
Familia: Truncatellidae
Genus: Sandbergerina KADOLSKY, 1993
Sandbergerina succineiformis (SANDBERGER), 1874 var.
Plate XIII Figures 7–12
1874 Euchilus succineiforme — SANDBERGER: p. 490, Pl. 25 Fig. 7
1921 Stalioa succineiformis — WENZ: p. 185, Pl. 27 Fig. 15
1926 Stalioa succineiformis — WENZ F. C.: p. 2177
1961 Nystia succineiformis — SCHLICKUM: p. 59, Pl. 4 Fig. 11
1970-c Nystia succineiformis — SCHLICKUM: p. 292, Fig. 9 (non 10!)
1993 Sandbergerina succineiformis — KADOLSKY: p. 384. Fig. 91.
The Upper Badenian pelitic layers of the Pusztamiske coal prospecting wells, as well as the “Nystia”-bearing limestone of the Nyirád wells (both layers of Late Badenian age) yielded specimens of this form in abundance. The
Pusztamiske specimens are excellently preserved.
It has a characteristic “hydrobioid” shape, with a slightly steplike whorl with varices, a thickened aperture, and
a peristome. The umbilicus is narrow, sometimes covered, and the shell surface is shiny with growth-lines, being
visible.
It has been described from the Mainz Basin “Landschneckenmergel” (“terrestrial snail bearing marl”), which is
coeval with the Hungarian Upper Badenian. Hitherto had not been reported from the Central Paratethys. The Hungarian
specimens are about half as big as those from the type locality. This may be explained by the fact that the accompanying
fauna suggests a slight salinity. This could be less favourable for the species than a freshwater environment. The abbreviation “var.” refers to its small size. The Upper Sarmatian beds of Várpalota also yielded some specimens. These are
slightly wider than the Badenian ones and have less varices. One subjuvenile specimen was collected from the Upper
Systematic descriptions
43
Badenian Herend Basin.
Sandbergerina hidasensis (KÓKAY), 1967
1967 Valvata hidasensis — KÓKAY: p. 84, 90, Pl. 8 Figs 3–6
It is non firmly accepted that the Hidas-type specimens (Mecsek Mts, Middle Badenian), as well as those from the
Nyirád Upper Badenian, belong to the genus Nystia. The thick margin of the aperture and the varices clearly point to this
genus and rule out the possibility of a Valvata.
S. succineiformis can be easily distinguished from N. hidasensis because the former is relatively higher it has a steplike spire and more convex whorls; consequently it has deeper sutures. Its stature is hydrobioid, and its aperture is more
marginated. The peristome better evolved and the varices are stronger than in S. succineiformis. The ratio of the height
of the last whorl and the total height is:
— 24–30% in S. succineiformis;
— 13–16% in S. hidasensis.
Specimens of S. succineiformis are bigger (height 2.5–3.0 mm) than those of N. hidasensis height 1.7–2.7 mm). This
species looks similar to the outline of the Viviparus species.
Genus: Stadtiellopsis SCHLICKUM, 1968
Staadtiellopsis rubeschi (REUSS), 1849 var.
Plate XIII Figures 13–14; Textfigure 1
1874 Euchilus? rubeschi — SANDBERGER: p. 423, Pl. 24 Fig. 31
1891 Euchilus rubeschi — KLIKA: p. 113, Fig. 112
1926 Stalioa rubeschi — WENZ F. C.: p. 2177
1961 Nystia rubeschi — SCHLICKUM: p. 61
1964 Nystia rubeschi — ČTYROKY et al.: p. 138, Figs 6–7
1968 Nystia (Stadtiellopsis) rubeschi — SCHLICKUM: p. 45, Pl. 5 Figs 10–11
1993 Stadtiellopsis rubeschi, KADOLSKY: p. 385. Fig. 92
This is a specimen from the Pusztamiske Upper Badenian, found in moulds from coeval
limestone in Nyirád. It is „hydrobioid” in stature, its whorls are convex, the base is perforated,
and there is a dense, barely outstanding axial ribbing. The latter does not represent growth- Textfigure 1. Staadtielloplines, although KLIKA (1891) denoted them this way. ČTYROKY et al (1964) produced a figure sis rubeschi (REUSS), 1849
var.
showing a smooth specimen. The aperture of this is damaged or it is not completely grown up;
however, remnants of the aperture are clearly bent outwards. The Hungarian specimens do not have a spiral lienage or any
other ornament, except for the last whorls on which some traces of ornamentation are visible. KLIKA’s (1891) figure shows
only axial elements. The Pusztamiske (from the well Pm–2) specimen is small being about half as big as the others.
Hitherto, the species was known only from the Lower Miocene coal-bearing formations in the Czech Republic.
Staadtiellopsis bandensis nov. sp.
Plate XIV Figures 1–2
Derivatio nominis: after Bánd, the type locality
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 100
Locus typicus: The Bánd bentonite mine, situated in the Herend Basin. The specimen was found in the matrix washed out of a
Brotia escheri shell, originating from the cover layers
Stratum typicum: Upper Badenian freshwater or oligohaline clay
Descriptio: The only, holotype specimen is 2.5 mm high, 1.2 mm wide, and its aperture is 1.0 mm high.
Small and with a hydrobiid-like stature it has five moderately convex whorls. The spire is slightly convex, with moderately deep sutural lines, and the apex is blunt. The shell is shiny, with fine growth-lines. On the penultimate whorl there
are two faint varix-like formations. At the base there is a narrow umbilical fissure. The aperture is crescent-formed, continuous, and its inner margin is gently arched (the outer one more so). The margins meet in an angle at the top. The columellar margin is turned out at the umbilical fissure. The rest of the margin is sharp, and below there is a very small
pouring lip. The external apertural margin bears a relatively wide, pillow-like swelling.
Under strong magnification a spiral furrow is visible on the centre of the last whorl, and a very faint second one is
situated on the dorsal side of the base.
Diagnosis differentialis: the new species is the closest to the species Nystia gracilis and N. lemani (SCHLIKUM 1961,
1970c). The apertural margin of the first-mentioned one is more bent outwards and its shell is smooth. The second one
has more convex whorls and more spiral furrows are visible on its surface.
44
JÓZSEF KÓKAY
Genus: Emmericia BRUSINA, 1897
Emmericia ? subpatula KÓKAY, 1966
1966 Emmericia subpatula — KÓKAY: p. 35, 107, Pl. 2 Fig. 13
This species, described from the Herend lower Badenian coal bearing sequence, retaines its original generic status
even after this revision. The revision of the Herend freshwater and terrestrial forms make it clear that this species is abundant in the coal-seam bearing layer, but due to its fragile shell generally only the thickened apertures were found. Study
of numerous fragments has revealed that the shell is often adorned with spiral lineae or grooves. The holotype is smooth,
but among the fragments there are many transitional forms. The spirally adorned specimens are close to the
Schuettemmericia subpyrenaicum (Noulet) form (SANDBERGER 1874: 513, Pl. 25 Figure 34; SCHLICKUM 1961: 62). The
Herend form differs from this because its outer apertural margin makes an angle with the columellar one — characteristically for E. species — while Noulet’s species has an elliptic ring-shaped aperture. One may suppose that the Herend
species is an extreme variety of Schuettemmericia subpyrenaicum, possibly marking the first evolutionary appearence of
the genus Emmericia. This hypothesis was made earlier by Cossmann et Peyrot as well as by Schlickum (see at the
description of Schuettemmericia paulensis).
The Herend form is also close to E. canaliculata (BRUSINA, 1874, 1897) described from the Dalmatian Miocene. The
Hungarian form is lower and wider, its whorls are more convex, its aperture is wider and vertically oriented, and furthermore it has no pouring lip-like edge on the lower part.The extant E. expansilabris Bourg. (RADOMAN, 1983), which lives
in Dalmatian streams, is also similar to the Herend form, but it is thinner and the axis of its droplet-shaped aperture is
vertical. The similarly modern species E. ventricosa Brusina (RADOMAN, 1983) is much wider and it has extremely convex whorls.
Familia: Aciculidae
Genus: Platyla MOUQUIN-TANDON, 1856
Platyla callosa (BOETTGER), 1870 var.
Plate XIV Figures 3–4
1870 Acicula callosa, BOETTGER: p. 284. Pl. 13. Fig. 1
1989 Platyla callosa, BOETERS et al.: p. 100. Fig. 92
Some specimens were found in the Lower Miocene of the Somlóvásárhely drilling. These are fragmentary: but the
last whorls have entire apertural margins. These are close to those of Boettger’s form, especially regarding their size.
Some specimens, however, are close to Platyla falkneri BOETERS et al. 1989, known from the Badenian of Opole (Oppeln
in Germ.), in Poland, as well as from the Upper Badenian of Herend.
On some specimens there is a rudiment of an earlike structure on the lower side of the columellar aperture. This
is a character of P. falkneri BOETERS et al. P. callosa has only been found in the Lower Miocene of Tuchorice, Czech
Republic.
Platyla alta (CLESSIN), 1911
Textfigure 2
1911 Acmea alta, CLESSIN: p. 107
1923 Acme (Acme) alta, WENZ F. C. p. 1855
1989 Platyla alta, BOETERS et al.: p. 94. Fig. 172
A specimen was collected from the Middle Badenian of Várpalota. The last two whorls were
preserved and these correspond well with those of Clessin’s species. The width of the shell is 1.0
mm, the height of the aperture is 0.75 mm, and its width is 0.55 mm. The margin of the aperture
is well-developed; from a lateral view it seems to be straight and at the base it is slightly widened.
The outline of the aperture is gently rounded-quadlangular.
The species is known from a formation of similar age at Undorf, Germany.
Platyla subfusca (FLACH), 1889
Plate XIV Figure 5
1889 Acme subfusca, FLACH: p. 71, 73. Pl. Fig. 1. (non vidi!)
1904 Acme callosiuscula, ANDREAE: p. 14. Fig. 13
1976 Acicula (Platyla) callosiuscula, SCHLICKUM: p. 3. Pl. 1. Fig. 4
1989 Platyla subfusca, BOETERS et al.: p. 169. Figs 170, 171
Textfigure 2. Platyla
alta (CLESSIN), 1911
Systematic descriptions
45
From the Lower Miocene of Somlóvásárhely two specimens of the last whorl were recognised and they have entire
apertures. Their size points to Flach’s species. Its range is from the Upper Oligocene to the Upper Badenian.
Platyla falkneri BOETERS, GITTENBERG et SUBAI, 1989
Plate XIV Figure 6
1989 Platyla falkneri, BOETERS et al.: p. 115. Figs 113, 116
The Upper Badenian of the well Herend-20 yielded a last whorl of this species. On the columellar part of the collar
encirling the aperture there is an ear-like wide expansion beneath the poorly developed umbilical callus. This is characteristic for the species. In the form P. polita this enlargement is smaller and the apertural margin together with the collar
is narrower. Therefore the Herend form can be identified as P. falkneri, which was described from the Tortonian of
Silesia. A small fragment was also found in the Lower Badenian of Herend.
Platyla polita (HARTMANN), 1840
Plate XIV Figure 7
1989 Platyla polita polita — BOETERS et al.: p. 145, Figs 145, 146, 151, 152, 17
Several, mostly fragmentary shells were collected from the Várpalota Sarmatian. The work referred to above gives a
complete synonymy. The species is stillpresent today from the Sarmatian.
Platyla cf. beatricis (GAÁL), 1911
Textfigure 3
1911 Acme beatricis, GAÁL: p. 43,50. Tab. 3. Fig. 1
1989 Platyla beatricis, BOETERS et al.: p. 215. Figs 214–215, 219
GAÁL (1911) described a species from the Sarmatian of Rákosd, Transsylvania. According to
Boeters et al., this form is taxonomically doubtful because the type has been lost. In collection three
incomplete specimens are preserved. Figures 214 and 215 of BOETERS et al. might correspond to
Gaál’s species, while Figure 213 shows most probably a specimen of P. polita.The Middle Badenian Textfigure 3. Platyla
cf. beatricis (GAÁL),
specimen from from Várpalota corresponds quite well to the figures of Boeters et al. The aperture 1911
and the external thickened part is relatively thin in both of the Hungarian and in the Rákosd specimen. The form and proportions of the aperture is also similar, but the Várpalota specimen is slightly smaller than
theTranssylvanian one: the widht of the whorl is 0.85 mm, the height of the aperture 0.70 mm, its widht is 0.55 mm. On
the figure of Gaál these values are even smaller.
Familia: Melaniidae
Genus: Brotia H. ADAMS, 1866
Brotia escheri turrita (KLEIN), 1847
Plate XIV Figures 8–11 (nominal sp.), Figures 12–13, 14 (subsp.)
1847 Melania turrita — KLEIN: p. 81, Pl. 2 Fig. 2
1852 Melania turrita — KLEIN: p. 159. Tab. III. Fig. 10
1874 Melania escheri var. rotundata — SANDBERGER: p. 572, Pl. 20 Fig. 19
1893 Melania escheri var. rotundata — LOCARD: p. 168, Pl. 9 Fig. 11
1929 Brotia escheri turrita — WENZ F. C.: p. 2599
1966 Brotia escheri — KÓKAY: p. 40, Pl. 3 Figs 20–21 (!)
1976 Brotia (Tinnyea) escheri — SCHLICKUM: p. 4, Pl. 1 Fig. 9
1992 Brotia escheri turrita — KÓKAY: p. 177, Pl. II. Figs 1–2
The form is a subspecies of Brotia escheri escheri and it occurs from the Oligocene to the Upper Badenian in freshwater and oligohaline formations. It differs from the nominal subspecies in that it has simple, straight ribs instead of spiny
or knobby ones. These are denser in B. e. turrita. On the last whorl the axial ribs may even disappear. The nominal form
also occurs in the upper Part of the Várpalota Upper Badenian as well as from the Ottnangian of Várpalota, the Lower
Miocene of Somlóvásárhely and from the Lower Badenian of Várpalota and Herend. Transitional forms toward B. e.
aquitanica (NOULET 1846) and B. e. laurae (MATHERON 1842) were also found.
The form occurs in appropriate facies of the Lower and Middle Miocene of Central and Western Europe. My observations show that it does not reach the Sarmatian in the Central Paratethys, probably due to climatic changes.
46
JÓZSEF KÓKAY
Genus: Melanoides ELIVIER, 1804
Melanoides procurvirostra n. sp.
Plate XV Figures 1, 2–4
Derivatio nominis: precursor of the species curvirostra
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 101
Locus typicus: Well Nagygörbő Ng–1, 901 m
Stratum typicum: Base of Lower Badenian freshwater series; fine sandy claymarl
Descriptio: dimensions of the holotype — height: 31 mm (originally it might have been about 39 mm); width: 12 mm.
It is a medium-sized acicular shell, and it is damaged, with the apex and the external and lower margin of the aperture missing. Thus the holotype consists of six moderately convex whorls. The missing apical part might similarly have
consisted of 6 additional whorls. There are spiral and axial decorations on the surface. An important character is that the
axial ribs are curved backwards; this is more accentuated on the juvenile part. The specimen could not be extracted completely from the matrix, due to its fragility. Consequently, the number of these axial ribs may only be estimated as 15 per
whorl. On the ribs there are 4 (on the lower part 5) knots, the size of which increases upwards. These are arranged along
spiral lines and have blunt ends. The biggest knot is the highest, immediately under the suture line. On the last whorl the
knots are weaker, appearing as slightly undulating spiral cords. On the base there are 8 such cords. The columellar side
of the aperture is thicke than the other parts.
There are apical fragments from specimens other than the holotype damaged and in this way the holotype may be
reconstructed. On these specimens the curves and knots on the ribs are more conspicuous. The uppermost 3 or 4 whorls
are smooth and slightly convex. (Remark: on species of the related genus Brotia the axial ribs appear after the first,
smooth 3 or 5 whorls; these are straight and smooth through some subsequent whorls.)
Variatio populationis: after the study of numerous damaged specimens it may be stated that the species has a wide
range of variety, similar to other species of this genus. The curved axial ribs are typical for all varieties. However there
is a great diversity in the number of these ribs. In some specimens it may exceed 20, but these ribs are much less developed than those of the holotype. The number of spiral cords may be 8 or 10. On the lower half of the whorls these are
continuous, while on the upper part there are small flat knots on the intercrossings with ribs. A specimen was found on
which 20 axial ribs crossed 8 spiral cords, and at the crossings there are knots on the whole surface.
Diagnosis differentialis: the new form is the closest to the south European Upper Miocene to Lower Pleistocene
Melanoides curvirostra (DESH.) (ESU et GIROTTI 1974), but it differs in the following mays:
1. it is twice as big;
2. its whorls, even the juvenile ones, are flatter;
3. after the smooth juvenile whorls the curved and knotted ribs and cords appear suddenly, while in M. curvirostra
there is a transition: the first two or three spiral cords (and, eventually the straight axial lineae) appear on the convex
whorls. It must be mentioned that the difference between Melanoides curvirostra and the modern M. tuberculata
(Müller) seems to be over emphasised, especially if the populations of many localities are taken into consideration.
Hungarian Middle Pleistocene specimens of M. tuberculata from Tata may be as big as those of M. procurvirostra and
they demonstrate a very large number of variations. The initial whorls of M. tuberculata are similar to those of M. curvirostra. Both species require warmth. The genus Melanoides prefers the area from the warm Mediterranean to tropical
freshwater, or even slightly saline environments.
The earlier literature does not mention the genus Melanoides from the Middle Miocene. Hitherto it was mentioned
from the Palaeogene and Upper Miocene or younger layers.
Familia: Melanopsidae
Genus: Melanopsis FÉRUSSAC, 1807
Melanopsis visianiana BRUSINA, 1874
Plate XV Figures 5–7
1874 Melanopsis Visianiana, BRUSINA: p. 37. Pl. I Fig. 7–8
1897 Melanopsis Visianiana, BRUSINA: p. 11. Pl. Fig. 5
1929 Melanopsis visianiana visianiana, WENZ F. C.: p. 2851
This species is quite frequent in the Lower Badenian freshwater deposits of the Nagygörbő drilling. Generally it is
not well-preserved. It resembles M. kleini KURR (1856) of the North Alpine Molasse deposits, but it is much bigger
(height 15 to 21 mm) than the Hungarian ones. These latter are 11 to 13 mm high and 5 to 5.5 mm wide. The last whorl
is 7 to 8 mm high and 5 to 5.5 mm wide; its last whorl is 7 to 8 mm high and the total number of whorls is 7 or 8.
Systematic descriptions
47
The Nagygörbő specimens are very close to, but slightly (by some tenth of mm) larger than those described by
Brusina from coeval sediments of Miocic, Dalmatia. On some of the Hungarian specimens there are some uneven structures on the surface of the shell (ridges or depressions); consequently, the sutures may be undulating.
Melanopsis cf. fritzei curta LOCARD, 1893
Plate XV Figure 8
1893. Melanopsis callosa var. curta, LOCARD: p. 181. Pl. IX Fig. 20.
In the upper part of the Lower Miocene of the Somlóvásárhely drilling some fragments and an entire subjuvenile
entire specimen of a Melanopsis form were collected. The last-mentioned specimen is much wider and thicker than
Locard’s species. The latter was collected from the ”Helvetian” of the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) in Switzerland. Wenz included this form in the species M. kleini KURR (1856). However, I cannot agree
with him, given that the size and proportions are quite different in the mentioned two taxa. The Lower Miocene name
“callosa” should be regarded as a junior synonim of “fritzei”.
Familia: Ellobiidae
Genus: Carychium O. F. MÜLLER, 1774
Out of the genera of this family, this is the only one which is not restricted to the shorelines of warm seas. Numerous
specimens were found in the washed up residues of pelitic sediments. In some samples it was extremely abundant. Its
small size promoted the washing of its shells from the shore into the areas of sedimentation. Two specimens were found
in the Lower Badenian of the Herend Basin while the rest are from the Upper Badenian and Sarmatian deposits. Strauch
(1977) based a specific system on the columellar wrinkles. However, this line of thought seems to have been carried to
excess. The columellar plates or wrinkles are variable features which reflect populations (in the palaeontological sense);
they do not appear to reflect subspecific differences. There are many other important traits as well.
Carychium nouleti nouleti BOURGUIGNAT, 1857
Plate XV Figure 9
1874 Carychium nouleti — SANDBERGER: p. 543, Pl. 28 Fig. 29
1881 Carychium nouleti — BOURGUIGNAT: p. 101, Pl. 6 Figs 176–179
1977 Carychium (Sarafia) nouleti — STRAUCH: p. 162, Pl. 15 Figs 24–27, Pl. 18 Fig. 61, Pl. 20 Fig. 83
2004 Carychium nouleti — BINDER: p. 194. Pl. 3. Fig. 2
One, excellently preserved shell was found in the Lower Badenian Coal measure of the Herend well. It has been found in
the Central and Western European Miocene — i.e. in the Lower Miocene (STYRIA, BINDER 2004) and in the Middle Miocene.
Carychium nouleti gibbum SANDBERGER, 1874
Plate XV Figures 10, 11
1891 Carychium nouleti var. gibbum — FLACH: p. 57, Pl. 3 Fig. 6
1923 Carychium nouleti gibbum — WENZ F. C.: p. 1197
1972 Carychium nouleti gibbum — GALL: p. 14
It is frequent in the Upper Badenian deposits, especially at Pusztamiske and the Tapolca Basin. It differs from the
nominal form in having a thicker, more marginated aperture and a more or less visible axial lineation.
It occurs in the Bavarian “silvana” beds at Undorf, which are coeval with the Hungarian localities.
Carychium sandbergeri HANDMANN, 1887
Plate XV Figures 12, 13
1907 Carychium Sandbergeri — TROLL: p. 67, Pl. 2 Figs 6, 7
1923 Carychium sandbergeri — WENZ F. C.: p. 1199
1977 Carychium (Sarafia) sandbergeri — STRAUCH: p. 167, Pl. 16 Figs 36, 38
The Upper Badenian of the Tapolca and the Pusztamiske Basins as well as the Várpalota Sarmatian yielded specimens of this species. It differs from the other C. species by its higher, more cylindrical form and straighter whorls. The
literature mentions that it is from the Central European “Tortonian” (=Badenian) and Lower Pannonian.
48
JÓZSEF KÓKAY
Carychium suevicum BOETTGER, 1877
Plate XV Figure 14, Plate XVI Figure 1
1977 Carychium suevicum — STRAUCH: p. 163, Fig. 4, Pl. 13 Fig. 8, Pl. 18 Fig. 59, Pl. 20 Fig. 89
A part of the Sarmatian specimens from Nyirád and Várpalota belongs here, based on the columellar wrinkles. C.
nouleti, occurring in layers older than Sarmatian, is close to this species. The type locality is Steinheim.
Subgenus: S a r a p h i a
Carychium (Saraphia) pachychilus SANDBERGER, 1875
Plate XVI Figures 2–4
1959 Carychium minimum — BODA: p. 637, 735, Pl. 35 Figs 13, 14
1977 Carychium (Sarafia) pachychilus — STRAUCH: p. 164, Pl. 15 Figs 31–35, Pl. 18 Figs 60, 62, 63, Pl. 20 Figs 87, 88
1981 Carychium (Sarafia) pachychilus — LUEGER: p. 14, Fig. 1, Pl. 1 Figs 5–10
It is frequent in the Várpalota Sarmatian. About one third of the specimens show a parietal dental plate, albeit zo varying degrees. This feature is more developed in the geologically younger C. berthae HALAVÁTS 1903; this was fused by
Lueger with Sandberger’s form,based on the columellar plate.The species is widespread in the Sarmatian, Pannonian and
Pontian of the Pannonian Basin.
Subgenus: C a r y c h i e l l a
Carychium (Carychiella) eumicron eumicron BOURGUIGNAT, 1857
Plate XVI Figure 5
1970 Carychium nanum var. major — BOETTGER: p. 297, Pl. 13 Fig. 8
1977 Carychium (Carychiella) eumicron — STRAUCH: p. 159, Pl. 14 Figs 13–15, Pl. 17 Fig. 53, Pl. 19 Fig. 76
This form has been found in the Lower Miocene series of the Somlóvásárhely drilling as well as is the the lower part
of the Sarmatian of Várpalota. It is also known from the Silesian Opole (Poland, German name: Oppeln).
Carychium (Carychiella) eumicron peneckei FLACH, 1891
Plate XVI Figure 6
1891 Carychium minutissimum Al. Br. mut. Peneckei — FLACH: p. 57, Pl. 3 Fig. 7
1923 Carychium eumicron peneckei — WENZ F. C.: p. 1190
1972 Carychium eumicron peneckei — GALL: p. 14
The form was described from the Bavarian Upper Freshwater Molasse (Middle Miocene). One specimen was found
in the Lower Badenian coal-seam bearing series of Herend. The whorls of this small type (1.3 mm high and 0.6 mm wide)
are very convex. Itdoes not have a lower columellar tooth. The right outer apertural margin is slightly damaged. The specimen corresponds well to the figure of FLACH (1891).
Carychium (Carychiella) eumicron unidentata nov. ssp.
Plate XVI Figures 7, 8
Derivatio nominis: unidentata (Lat.) = with one tooth
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 102
Locus typicus: drilling Somlóvásárhely Svh–1. 192.0–192.2 m
Stratum typicum: Lower Miocene (Ottnangian to Karpatian) freshwater clay with lime mud.
Descriptio: The small-sized (height 1.45 mm, width 0.7 mm), cylindrical shell has four and a half whorls and is terraced. The shell is smooth with slight growth lines present only on the last whorl.The aperture is semi-angular. Its columellar and palatal sides are straight, slightly bent outwards, and marginated. There is one small pointed columellar tooth
situated quite deeply. On the palatal side there is no tooth or ridge.
Diagnosis differentialis: I collected nine specimens from the cores. The individuals are essentially uniform in shape.
The smallest adultive specimen is 1.2 mm high, and 0.55 mm wide. In one specimen it was possible to study the spindle
plate, because the last whorl was broken off. This part hardly widens at all even less developed than in C. eumicron
(BOURG. 1857). This is the species that stands closest to the Hungarian one. This latter has, however, a more developed
columellar tooth, and there is a tooth or protuberance on the inner side of the external apertural margin.
Systematic descriptions
49
Genus: Laemodonta PHILIPPI, 1846
Subgenus: P r o p l e c o t r e m a
Laemodonta (Proplecotrema) bourgeoisi TOURNOUER, 1878 var.
Plate XVI Figure 9
1872 Plecotrema Bourgeoisi — TOURNOUER: p. 104, Pl. 4 Fig. 6
1916 Plecotrema (Plecotremopsis) Bourgeoisi — MORGAN: p. 34, Figs 39–42
1923 Laemodonta (Plecotremopsis) bourgeoisi — WENZ F. C.: p. 1162
1952 Laemodonta (Plecotremopsis) bourgeoisi — GLIBERT: p. 400, Pl. 15 Fig. 14
Two specimens originated from the Várpalota marine Ottnangian. This terrestrial snail lived in the nearshore zone
(probaly in the Mangrove zone). It indicates a warm climate. In contrast to earlier descriptions, the spiral elements are
dominant on the shell; consequently, the knots on the ribs are not conspicious. According to MORGAN (1916), the ornamentation shows a range of variety. It is known from the Western European Lower and Middle Miocene.
Genus: Melampus MONTFORT, 1810
Melampus turonense striata KÖRMENDY, 1962
1962 Melampus (Melampus) turonense striata KECSKEMÉTI-KÖRMENDY: p. 95, 98, Pl. 11 Figs 10, 11
Found in the Lower Badenian of the “Szabó” sandpit, Várpalota.
Genus: Pedipes BRUGUIÉRE, 1792
Subgenus: N e a l e x i a WENZ, 1920
Pedipes (Nealexia) myotis pisolina DESHAYES, 1831
1966 Pedipes (Nealexia) myotis pisolina — STRAUSZ: p. 484, Pl. 46 Figs 38–42
Found in the Lower Badenian of the “Szabó” sandpit, Várpalota.
Genus: Stolidoma DESHAYES, 1863
Stolidoma tournoueri RAINCOURT, 1876
1962 Stolidoma tournoueri — KECSKEMÉTI-KÖRMENDY: p. 94, Pl. 11 Fig. 7
Found in the Lower Badenian of the “Szabó” sandpit, Várpalota.
Stolidoma mayeri TOURNOUER, 1870
1962 Stolidoma mayeri — KECSKEMÉTI-KÖRMENDY: p. 94, Pl. 11 Fig. 8
Found in the Lower Badenian of the “Szabó” sandpit, Várpalota.
Stolidoma dolioliformis KÖRMENDY, 1962
1962 Stolidoma dolioliformis — KECSKEMÉTI-KÖRMENDY: p. 94, 98, Pl. 11 Fig. 14
Found in the Lower Badenian of the “Szabó” sandpit, Várpalota.
Stolidoma gracilis KÖRMENDY, 1962
1962 Stolidoma gracilis — KECSKEMÉTI-KÖRMENDY: p. 94, 99, Pl. 11 Fig. 15
Numerous specimens were found in the Lower Badenian of the “Szabó” sandpit, Várpalota.
Genus: Auriculastra MARTENS, 1880
C o n a u r i c u l a nov. subgenus
Derivatio nominis: Conical: auricula (lat.)
Typus species: C. badeniensis nov. sp.
Stratum typicum: Upper Badenian coal bearing clay
Descriptio: dimensions of the holotype: height — 2.6 mm; width — 1.7 mm. The height of the aperure is 1.6 mm
while its is width 0.6 mm. It is a small shell if compared to other genera of the family and the shell itself is thin. Its stature
and spire look rather like a wide Conus species. Five stepwise superponed whorls constitute the shell.
50
JÓZSEF KÓKAY
The aperturte is elongate and its upper rim joins the whorl at an angle. The external margin is simple, thin, internally smooth and it is without teeth. On the internal part of the lower apertural margin there is a columellar wrinkle (plate)
while above it there is a parietal plate. There is no internal cover on the inner side of the aperture (which is a feature of
the nominal subgenus Auriculastra). The surface is decorated by spirally arranged, extremely small pits which are only
visible under powerful magnification. Within the familiy Ellobiidae, only two genera — Melampus and Ellobium —
have been reported with similar decoration.
Auriculastra (Conauricula) badeniensis n. sp.
Plate XVI Figures 10–11, 12, 13
Derivatio nominis: from the Badenian stage
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 103
Locus typicus: Pusztamiske, well Pmt–3, 52.0–52.3 m
Stratum typicum: Upper Badenian coal bearing clay
Descriptio: see the description of the subgenus
Variatio populationis: Eight specimens were found in the Pusztamiske sample. The holotype is not the bigges: there
is one damaged specimen which is 3 mm high. From the well Pm–3 (52.7–53.1 m) two specimens were collected. One
is compressed and it is 3.3 mm high; another, entire one, is much smaller.
The spiral punctuation is not equally developed on a the specimens. In one specimen a poorly developed plate was
found above the parietal dental plate.
The genus is known from the European Miocene. Right up until recent times it lived along seashores in the Indo-west
Pacific realm. The Pusztamiske specimens probably lived along a low salinity lagoon, under a subtropical, humid climate.
Familia: Lymnaeidae
Genus: G a l b a SCHRANK, 1803
Galba dupuyana (NOULET), 1854
Plate XVI Figures 14–16
1874 Limnaeus Laullardianus — SANDBERGER: p. 543, Pl. 28 Fig. 27
1881 Limnaea dupuyana — BOURGUIGNAT: p. 118, Pl. 6 Figs 202–204
1881 Limnaea eumicron — BOURGUIGNAT: p. 119, Pl. 6 Figs 205–207
1916 Limnaea Dupuyi — DOLLFUS: p. 370, TextFig. 6
1923 Radix (Radix) dupuyana — WENZ F. C.: p. 1242
This is a small 3–5 mm high snail of lymnaeid type. It is frequent in the western Bakony Upper Badenian freshwater and low salinity deposits and occurs in the Várpalota Badenian as well. The Hungarian specimens look quite similar
to SANDBERGER’s (1874) and Dollfus’ figures. It occurs in the Middle- and West European Middle Miocene.
Genus: Stagnicola LEACH, 1830
Stagnicola jaccardi (MAILLARD), 1892
1892 Limnaea jaccardi — MAILLARD: p. 99, Pl. 6 Figs 24, 25
1923 Galba (Galba) jaccardi — WENZ F. C.: p. 1370
A specimen was collected from the Hegymagos well; specimen this is narrow and elongate. Its surface is covered
with faint spiral ridges. With the growth-lines and wrinkles these form a slightly net-like pattern. It has been reported
from formations comparable in age to the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) in Switzerland.
Stagnicola armaniacensis (NOULET), 1857
Plate XVII Figures 1
1874 Limneus armaniacensis — SANDBERGER: p. 581, Pl. 28 Fig. 25
1881 Limnea armaniacensis — BOURGUIGNAT: p. 116, Pl. 6 Fig. 195
1892 Limnea armaniacensis — MAILLARD: p. 115, Pl. 7 Fig. 20
1976 Stagnicola (Stagnicola) armaniacensis — SCHLICKUM: p. 5, Pl. 1 Fig. 12
Systematic descriptions
51
From the Hegymagos well one specimen and one mould were collected. SCHLICKUM’s (1976) specimen differs from
these and from other, earlier collected ones in being wider stouter. It occurs in the “silvana” beds.One specimen, found
in the Várpalota Sarmatian, is slightly bigger.
Stagnicola armaniacensis palustriformis (GOTTSCHICK), 1911
Plate XVII Figure 2–3
1911 Limnaea (Limnophysa) palustriformis — GOTTSCHICK: p. 512, Pl. 7 Figs 7–9
1920 Limnaea (Limnophysa) armaniacensis palustriformis — GOTTSCHICK: p. 113, Pl. 2 Fig. 8c
1923 Galba (Galba) armaniacensis palustriformis — WENZ F. C.: p. 1352
One, subjuvenile specimen was found in the uppermost, freshwater layers of the Upper Badenian in the Pusztamiske
Pm–3 well. Its shell may be smooth but it may also be adorned with a spiral net-like rib pattern.
It differs from the modern Stagnicola palustris (Plate XVII Figures 3, 5.) in that it is smaller in size; however, a 21
mm high mould was found in the Pm–3 well and one also in the Várpalota Sarmatian which were of the same size.
Stagnicola armaniacensis glabraeformis (GOTTSCHICK et WENZ), 1911
1911 Limnaea (Limnophysa) glabraeformis — GOTTSCHICK et WENZ: p. 513, Pl. 7 Figs 10–11
1920 Limnaea (Limnophysa) armaniacensis glabraeformis — GOTTSCHICK: p. 113, Pl. 1 Figs 8a, 8b
1923 Galba (Galba) armaniacensis glabraeformis — WENZ F. C.: p. 1352
Elongate and narrow specimens from the Várpalota Sarmatian correspond well to coeval ones from Steinheim.
Stagnicola praeboulleti SCHLICKUM, 1970
Plate XVII Figures 4–8
1967 Limnaea (Stagnicola) cfr. syrtica — KÓKAY: p. 86, Pl. 8 Fig. 7
1970a Stagnicola (Stagnicola) praeboulleti — SCHLICKUM: p. 92, Figs 12–17
1970b Stagnicola (Stagnicola) praeboulleti SCHLICKUM: p. 179, Pl. 3 Figs 8–9
1976 Stagnicola (Stagnicola) praeboulleti SCHLICKUM: p. 5, Pl. 1 Fig. 13
It was found in large numbers in the cores of the Nyirád wells, generally as moulds in freshwater limestones. For my
1967 publication I could not determine the specimens, nor could I describe them as a new taxon. Slightly later SCHLICKUM
(1970a) described a new species, based on well-preserved specimens from the Bavarian “Upper Helvetian”, “Upper
Brackish Molasse” lacustrine layers. Schlickum’s form agrees well with the Hungarian one with respect to its size and
proportions, but the author does not mention, the following:
— the shell varices and strong wrinkles are often visible as a result of the recurrent thickening of the aperture during growth (although these features are visible on his figures);
— (caused by the same trend) the further growth of the shell is frequently “gonoid”rather than “circoid” — i.e. there
is a slight angle on it;
— the thickenings of the aperture are visible on the inner side as well being present on the internal moulds as prints
of varices.
The Nyirád and Várpalota Sarmatian Stagnicola specimens may also be placed into S. praeboulleti.
Stagnicola kreutzii (ŁOMNICKI), 1886
Plate XVII Figure 9
1886 Limnaea (Limnophysa) KREUTZII — ŁOMNICKI: p. 81
1886 Limnaea (Limnophysa) KREUTZI var. subflavida — ŁOMNICKI: p. 81, Pl 2 Fig. 40
1886 Limnaea (Limnophysa) KREUTZI var. subfusca — ŁOMNICKI: p. 81, Pl 2 Fig. 41
One small, well-preserved specimen from the Várpalota Sarmatian agrees well with ŁOMNICKI’s (1886) figures It may
be regarded as a transition between the two subspecies of the mentioned author. It is know from the Middle Miocene
(“Tortonian”) of Southern Poland (Galicia).
Stagnicola palustris (O. F. MÜLLER), 1774
Plate XVII Figure 10
1923 Galba (Galba) palustris palustris — WENZ F. C.: p. 1391
1943 Stagnicola palustris — SOÓS: p. 83, Pl. 4 Fig. 15
1979 Stagnicola palustris — RICHNOWSZKY et PINTÉR: p. 75, Fig. 26b
52
JÓZSEF KÓKAY
In the Brotia-bearing limestone of the Pusztamiske Pm–3 well a mould was found, from which silicone casts were
made. Its size: height — 21 mm; width, 9 mm, height of aperture —10.5 mm; width of aperture — 5.8 mm.
On the surface of the last whorl a coarse net-like ornament with spiral elements is visible and this is characteristic for
the species. The species is occurs whith a large number of variations especially with respect to the proportions of the
shell. In the literature it is subdivided into numerous forms. The narrow form of the Hungarian specimens is similar to
the forms S. turricula (HELD.) or clessiniana (HAZ.). It may have existed in oligohaline environments, but it is more likely that it occurred in shallow, marsh like ones. The Pusztamiske specimen was found in a limestone, containing abundant evidence of rootprints.
The Várpalota Sarmatian yielded a narrow specimen with a preserved shell. Its height is 24 mm; width — 9 mm;
height of aperture — 10 mm; width of aperture 4.3 mm. The shell surface is similar to that from Pusztamiske and it
resembles the mentioned two forms.
Hitherto, the first known appearance of the species was from theUpper Pontian (Upper Miocene) of Hungary
(BARTHA 1959).
Genus: Radix MONTFORT, 1810
Radix socialis regularis (MAILLARD), 1892
Plate XVII Figures 12–13
1892 Limnaea dilatata var. regularis — MAILLARD: p. 112, Pl. 7 Figs 12–13
1923 Radix (Radix) socialis regularis — WENZ F. C.: p. 1289
In its regular outline and size some specimens from Várpalota and Hegymagos wells agree well with those described
from the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) of Switzerland. It is also known from the
Sarmatian of Várpalota.
Radix socialis praelongata (GOTTSCHICK et WENZ), 1916
Plate XVII Figure 11
1916 Limnaea (Radix) ilatata var. praelongata — GOTTSCHICK et WENZ: p. 99, Pl.5. Figs 15–17
1923 Radix (Radix) socialis praelongata — WENZ F. C.: p. 1289
This is a relatively thinner, but higher form, found in some wells at Nyirád. It has been described from the coeval “silvana” beds.
Radix dilatata (NOULET), 1854
Plate XVII Figure 14
1881 Lymnaea dilatata — BOURGUIGNAT: p. 112, Pl. 6 Fig. 193
1881 Lymnaea sphaerogyra — BOURGUIGNAT: p. 113, Pl. 6 Fig. 194
1874 Limnaeus dilatatus — SANDBERGER: p. 523, Pl. 28 Fig. 24
1892 Limnaea pachygaster — MAILLARD: p. 109, Pl. 7 Figs 6, 7
1892 Limnaea dilatata — MAILLARD: p. 110, Pl. 7 Figs 8, 11
This is a rather big, thin shelled form with strong growth-lines and wrinkles. It is found in cores from the SW part of
the Várpalota coal field, from the base of the Upper Badenian coal-seam from the Upper Badenian of Nyirád, and from
the Várpalota Sarmatian.
It is known from the lower part of the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) in Swiss and
from France (Sansan). Both of these are coeval with the Karpatian and the Badenian.
Radix hyaloleuca (BRUSINA), 1902
Plate XVII Figure 15
1902. Lymnaea hyaloleuca — BRUSINA: Pl. 1. Figs 36–39
1907 Limnaea hyaloleuca — BRUSINA: p. 211
1923 Radix (Radix) hyaloleuca — WENZ F. C.: p. 1250
Small specimens were found in the Sarmatian of Várpalota. The species has also been found in the Middle Miocene
deposits of Herzegovina.
Systematic descriptions
53
Genus: Lymnaea LAMARCK, 1799
Lymnaea turrita (KLEIN), 1855
Plate XVII Figure 16
1853 Limnaeus turritus — KLEIN: p. 220, Pl. 5 Fig. 17
1874 Limnaeus turritus — SANDBERGER: p. 581, Pl. 28 Fig. 26
1881 Limnaea turrita — BOURGUIGNAT: p. 117, Pl. 6 Figs 200, 201
1923 Lymnaea turrita — WENZ F. C.: p. 1227
1976 Lymnaea turrita — SCHLICKUM: p. 5, Pl. 1 Fig. 15
KLEIN’s (1853) figure shows a small and narrow specimen. SCHLICKUM’s (1976) specimen is bigger and thicker and
is the one which deviates most from the type. The Tapolca specimens also demonstrate the best agreement with Klein’s
type with respect to size. It is also found in the Sarmatian of Várpalota.
It is known from the Middle Miocene of Central and Western Europe.
Lymnaea stagnalis sarmatica nov. sp.
Plate XVII Figure 17
Derivatio nominis: alluding to the Sarmatian stage
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 104
Locus typicus: Várpalota coal basin, Shaft M. III, 133.5 m
Stratum typicum: Sarmatian freshwater clay
Descriptio: it is a tower-shaped shell with thin a wall, concave sides, and 9 moderately convex whorls. The last one
appears to increase in suddenly size forming three-quarters of the total height. The shell surface is covered with fine, parallel growth-lines resulting in a silky shine. On the lower whorls a faint spiral lineation is visible. The aperture resembles a reversed human ear, bearing a parietal plate on its upper part.
Out of the great number of fragmentary specimens only one was complete and this was chosen as the holotype. Its
dimensions: height 32 mm; width — 14.5 mm; height of last whorl — 24 mm; height of aperture — 17 mm.
Diagnosis differentialis: this form does not differ much from the still living and common Lymnaea stagnalis but there
are the following variations:
— more whorls — i.e. 7 or 8;
— big, grown-up specimens are bigger, with their height being as much as 40 mm;
— the ornament of the new subspecies is characteristic. The spiral ornament is present on the upper whorls in the
modern (living) species while the ornament is on the lower whorls in the case of the Hungarian specimens.
Some fragments were found in the Upper Badenian of the Pusztamiske and Tapolca Basins.
Genus: Aplexa FLEMMING, 1820
Aplexa subhypnorum physaeformis GOTTSCHICK, 1920
1920 Aplexa subhypnorum physaeformis — GOTTSCHICK: (part III) p. 116, Pl. 1 Figs 9b, 9b1
1923 Aplexa subhypnorum physaeformis — WENZ F. C.: p. 1719
1959 Aplexa subhypnorum — BODA: p. 638, 736, Pl. 37 Figs 5, 6
Several fragments were found in the Sarmatian of Várpalota. It differs from the nominal subspecies due to its flatter
and wider stature. Gottschick described it from Steinheim, where it occurs together with the nominal subspecies. These
are more or less coeval with the Hungarian occurrences.
Familia: Planorbidae
Genus: Anisus STUDER, 1820
Anisus dupuyanus (NOULET), 1854
Plate XVII Figure 18
1874 Planorbis Dupuyanus — SANDBERGER: p. 542, Pl. 28 Fig. 22
1881 Planorbis dupuyanus — BOURGUIGNAT: p. 131, Pl. 7 Figs 225, 228
2000 Anisus (Anisus) dupuyianus — FISCHER: p. 136, Fig. 6
54
JÓZSEF KÓKAY
Tis species is found in the uppermost part of the Upper Badenian of the Pusztamiske P–2 well, and in the upper part
of the Várpalota Sarmatian. Most of the specimens are damaged. Their size, the clearly visible growth-lines and the density of the whorls agrees well with Noulet’s species.
WENZ (1923: p. 1532) included strongly deviating forms into Noulet’s species. However, we cannot agree with him.
The differences are also clearly visible on the figures of BOURGUIGNAT (1881).
This species is known from the Langhian of Sansan in France.
Anisus dupuyanus omalus (BOURGUIGNAT), 1881
Plate XVII Figures 19–21
1881 Planorbis omalus — BOURGUIGNAT: p. 134, Pl. 7 Figs 237–240
2000 Anisus (Disculifer) omalus — GINSBURG: p. 137, Fig. 7
This form originates from the Upper Badenian freshwater clays in the Hegymagos well and from the Saramatian of
Várpalota. WENZ (1923: p. 1532) clusters into one species (“Paraspira” dupuyana) several forms from the group
described by BOURGUIGNAT (1881). There are, however, important variations between these forms and thus it is safe to
distinguish them under the subspecific name “omalus”.
There is a characteristic blunt ridge along the whorls which becomes acute close to the aperture. The aperture is
placed asymmetrically and is bent downwards toward the base. Our specimen is 3 mm in diameter; it is therefore much
smaller than the type (9 mm) but a closely related form (“P.” gyreligmus BOURGUIGNAT) has a diameter of only 5 mm.
Anisus rousianus (NOULET), 1854
Plate XVIII Figures 1–4
1881 Planorbis rousianus (BOURGUIGNAT): 132, Pl. 7 Figs 225–232
One well-preserved and one damaged specimen were found in the Hegymagos well. The five whorls (circular in
cross-section) are placed assymetrically. It is 5 mm in diameter, similar to Bourguignat’s type; the latter originated
from deposits at Sansan in France and it is coeval with the Badenian. A complete subjuvenile specimen was collected from the uppermost Upper Badenian of the Pusztamiske Pm–1 well. Two damaged specimens, which are 6.5 mm
in diameter, came from the Pm–2 well, from the same level. In the cover of the Várpalota Upper Badenian coal-seam
there is an “Anisus”-bearing thin layer with ill-preserved specimens of this form. It occurs in the Sarmatian of
Várpalota as well.
Anisus hilgendorfi subcarinata (GOTTSCHICK), 1920
Plate XVIII Figures 5, 6
1911 Planorbis (Tropidiscus? Gyrorbis?) Hilgendorfi var. — GOTTSCHICK: p. 514, Fig. 3b, Pl. 7 Fig. 12
1920 Gyrorbis hilgendorfi subcarinatus — GOTTSCHICK: p. 163
1923 Paraspira (Paraspira) hilgendorfi subcarinata — WENZ F. C.: p. 1534
On this species there is an acute rim or crest, separating the lower and upper part of the shell. It is the form which is
closest to the extant species A. vortex (LINNAEUS) (RICHNOWSZKY et PINTÉR 1979). The fossil form is less flat its whorls
are more convex and the rim is higher and wider.
It occurs in the Upper Badenian and Sarmatian of the western Bakony Mts, as well as in the Várpalota Sarmatian and
in some places it is abundant. It is also present in the Steinheim beds.
Genus: Gyraulus CHARPENTIER, 1837
Gyraulus alienus (ROLLE), 1862
Plate XVIII Figures 7–10
1862 Planorbis alienus — ROLLE: p. 210, Pl. 1 Figs 6, 7
1923 Gyraulus (Gyraulus) alienus — WENZ F. C.: p. 1542
This is present at the base of the Sarmatian in the Pusztamiske Pm–3 well and there are numerous specimens; they
are highly involute. Relatively thick (high) whorls and an almost circular, blunt subtriangular and elongate aperture are
characteristic for the species.
ROLLE (1862) reported this specimen from the Middle Badenian of Hidas (Mecsek Mts).
Systematic descriptions
55
Gyraulus pulici (BRUSINA), 1897
Plate XVIII Figures 11–16
1897 Planorbis pulici — BRUSINA: p. 6, Pl. 2 Figs 8–10
1902 Planorbis pulici — BRUSINA: Pl. 3 Figs 22–24
1923 Gyraulus (Gyraulus) pulici — WENZ F. C.: p. 1571
1959 Gyraulus pavlovici — BODA: p. 736, Pl. 35 Figs
Thisis frequent in the Várpalota Sarmatian. It is close to G. trochiformis kleini (GOTT. et WENZ). The aperture of this
last mentioned form is, however, asymmetric and is slightly downturned G pulici. also has a blunt keel which is close to
the midline, but points slightly downwards from it. It is possible that there are transitions between these forms. Brusina
described the form from the Middle Miocene of Herzegovina.
Gyraulus nedici (BRUSINA), 1902
Plate XIX Figure 1–3
1902 Planorbis nedici — BRUSINA: Pl. 3 Figs 25–27
1923 Gyraulus (Gyraulus) multicingulatus — WENZ F. C.: p. 1364
1959 Gyraulus solenoides — BODA: p. 737, Pl. 35 Fig. 10
This is quite common in the Sarmatian of Várpalota. This species deviates from G. microstatus (BOURG. 1881) (which
occurs in the Western Bakony Lower Sarmatian) in having a circular or almost circular aperture. The latter is slightly
downturned, while that of G. microstatus is rather upturned.
It seems that there are intermediate (symmetric) forms as well, as on the figure of BODA (1959). G. nedici is slightly
smaller than G. microstatus. It occurs in the Nyirád Sarmatian as well.
Brusina described the form the from pre-Pannonian strata of Herzegovina.
Gyraulus microstatus (BOURGUIGNAT), 1881
Plate XIX Figure 4–5
1881 Planorbis microstatus — BOURGUIGNAT: p. 141, Pl. 8 Figs 268–272
This species is frequent in the Upper Badenian of the Pusztamiske and Hegymagos wells and in some layers of
Herend; furthermore, some subjuvenile specimens were found in the Lower Miocene of the Somlóvásárhely drilling. The
whorls, which are circular in cross-section encircled by the aperture asymmetrically and in a more flat manner on the
base side. The shape of the aperture varies considerably, even within one population. The form of the aperture shows
transitional shapes towards that of G. trochiformis kleini GOTTSCHICK et WENZ. This type is 2 mm in diameter while the
Hungarian specimens are slightly bigger being between 2–3 mm.
WENZ (1923: p. 1555) regards it as a subspecies of G. goussardianus, which seems not to be the case, as G. microstatus is:
— smaller,
— more involute,
— less flattened,
— its aperture is wider and less elongate.
The species was recognised in the Sansan of Southern France and it occurred in freshwater strata coeval with those
of the Hungarian localities.
Gyraulus amiculus (ŘZEHAK), 1893
Plate XIX Figure 6–7
1893 Planorbis amiculus — ŘZEHAK: p. 175, Pl. 2 Fig. 9
In the regressive Lower Badenian and Middle Badenian series of Várpalota some specimens were found which seem
to be indentical with Rzehak’s species.
The whorls are rather involute, fast growth is indicated and they are inflated and convex. The aperture is almost circular and it embraces the previous whorl slightly asymmetrically; it is gently turned towards the base. It is small being
1.5 mm in diameter, and 0.8 mm high. The proportions are identical with those of the type.
WENZ (1923: p. 1593) regarded it as a synonym of Gyraulus trochiformis dealbatus, although their separation seems
to be justified.
The type was found in Lower Miocene beds in their south of the Czech Republic.
56
JÓZSEF KÓKAY
Gyraulus callistus (BOURGUIGNAT), 1881
Plate XIX Figures 8–11
1881 Planorbis callistus — BOURGUIGNAT: p. 145, Pl. 8 Figs 258–262
The Pusztamiske and Hegymagos specimens may be identified with those from Sansan, France. The elliptic aperture
embraces symmetrically the previous convex whorl. On some specimens faint radial bands are visible. The biggest specimen is 1.6 mm in diameter, while Bourguignat mentions a dimension of 1.5 mm, thus the correspondence is good.
WENZ F. C. (1923: p. 1626) identifies the species with the G. lenapalus (BOURGUIGNAT) form, but this does not seem
to be reasonable as this latter is much more flat and its aperture is asymmetric.
Gyraulus verticillus (BRUSINA), 1902 var.
Plate XIX Figure 12
1902 Planorbis verticillus — BRUSINA: p. 8, Pl. 4 Figs 36–39
1902 Planorbis verticillus — LŐRENTHEY: p. 187, Pl. 13, Figs 12, 14
1923 Gyraulus (Gyraulus) verticillus — WENZ F. C.: p. 1624
This form originates from the Lower Badenian freshwatwer base beds of the Nagygörbő drilling. A complete, but
slightly subjuvenile specimen and a fragment of a last whorl of an adult were collected, as well as some compressed
examples. It is very small in size: the subjuvenile one is 1.5 mm in diameter, the adult 2–3 mm. It is a stout, planorboid
shell. In the median plane and on both sides there are 3 keels. There are 3 whorls, on the adult 3½–4. The keels are slightly blunt. The umbilicus is gently deeper at the base side. The aperture is triangular and downturned slightly to the base.
It is visible on some bigger specimens and, — especially on the figures of LŐRENTHEY (1902) — on the adult whorl there
are two furrow-like depressions behind the keels.
Lőrenthey’s specimens are flatter than Brusina’s and those from Nagygörbő. The Nagygörbő shells are more symmetric (the aperture is not downturned), similar to those of LŐRENTHEY (var.”ptychodes”, 1902 Figure 13). This last-mentioned is more flat than the Nagygörbő ones. Based on the last whorl of the mentioned adult it seems that during growth
the shell becomes more and more flat and the keels blunter.
Based on figures and descriptions of BRUSINA (1892, 1902) and LŐRENTHEY (1902) the species demonstrates a number of variations. Thus it seems reasonable to identify the Nagygörbő form with Brusina’s one specimen.
Hitherto this form was known only from the Lower Pannonian of Croatia and the Budapest area (Tinnye). However,
from the hanging wall of the Várpalota Upper Badenian coal-seam a well preserved specimen was collected. In the latter the central keel is missing which is a consequence of it being of the adult age (diameter — 3.0 mm; width — 2.0
mm). The variability of the species accounts for such differences. The only specimen from the Várpalota Sarmatian is
relatively flat and it is similar to Lőrenthey’s figured specimen.
Gyraulus geminus (BRUSINA), 1897
1869 Planorbis applanatus, NEUMAYR: p. 368. Pl. XII Fig. 22
1874 Planorbis applanatus, BRUSINA: p. 101
1897 Planorbis geminus, BRUSINA: p. 5. Pl. II. Figs 11–13
1923 Giraulus geminus, WENZ F. C.: p. 1554
In the Pusztamiske Pm–2 drilling two specimens were found in the base of the Upper Badenian. One of these is excellently preserved. It has four flat whorls and, medially, a rounded keel. Its aperture is elongate, oval and it has well-developed grooving lines. It is known from Dalmatia, from layers coeval with the Karpatian and Badenian. Its occurrence in
the Pannonian or Pontian is questionable.
Gyraulus applanatus (THOMAE), 1845
Plate XIX Figure 13–14
1874 Planorbis (Gyraulus) declivis, SANDBERGER: p. 370. Pl. 25. Fig. 9
1891 Planorbis declivis, KLIKA: p. 107. Fig. 104
1893 Segmentina declivis, LOCARD: p. 136. Pl. 8. Fig. 3
1921 Gyraulus applanatus, WENZ: p. 176. Pl. 25. Fig. 15–16
1923 Gyraulus (Gyraulus) trochiformis applanatus, WENZ F. C.: p. 1579
1997 Gyraulus applanatus — PIECHOCKI: p. 111. Fig. 8
Some small speciemns were found in the lowest part of the Lower Miocene of the Somlóvásárhely drilling. The shells
are flat, especially at their lower part. In the lower third of the shell there is a more or less rounded keel. The aperture is
elongate. The specimens must be regarded as subjuvenile ones; the biggest one is less then 2 mm in diameter and has
Systematic descriptions
57
three whorls. The above-quoted papers mention a diameter of 4–6 mm and 5–6 whorls. It is well known from the Lower
Miocene of the North Alpine foreland and the Czech Republic.
Gyraulus trochiformis dealbatus (BRAUN), 1851
Plate XIX Figure 15
1874 Planorbis (Gyraulus) dealbatus — SANDBERGER: p. 492, Pl. 25 Fig. 10
1893 Planorbis dealbatus — LOCARD: p. 154, Pl. 9 Fig. 2
1923 Gyraulus (Gyraulus) trochiformis dealbatus — WENZ F. C.: p. 1591
1972 Gyraulus (Gyraulus) trochiformis dealbatus — ČTYROKY: p. 79 Pl. 5 Figs 2–9
1997 Gyraulus dealbatus — PIECHOCKI: p. 113. Fig. 9
This form is known from the Lower and Middle Miocene of Central and western Europe; it occurs in the Herend
Lower Badenian coal-seam series, as well as in the closing member of the Várpalota Lower Badenian.
Gyraulus trochiformis kleini GOTTSCHICK et WENZ, 1921
Plate XIX Figure 16–17
1874 Planorbis (Gyraulus) laevis — SANDBERGER: p. 578, Pl. 28 Fig. 21
1916 Gyraulus multiformis Kleini — GOTTSCHICK et WENZ: p. 101, Fig. 3
1921 Gyraulus kleini — WENZ: p. 189, Pl. 27 Figs 18, 19
1922 Gyraulus trochiformis kleini — WENZ: p. 142, Figs 2–4
1923 Gyraulus (Gyraulus) trochiformis kleini — WENZ F. C.: p. 1595
1997 Gyraulus kleini — PIECHOCKI: p. 113. Fig. 10.
1998 Gyraulus kleini — FINGER: p. 9. Taf. 2, 3, 4
This form has been collected from the Upper Badenian of Herend, Pusztamiske, Várpalota and the Tapolca Basin. Its characters are: a slightly involute shell, convex whorls and a slightly downturned aperture. The grade of this downturning and the
form of the external apertural margin are variable. It seems that there are transitional forms toward G. applanatus (THOMAE),
G. dealbatus (BRAUN) and G. microstatus (BOURG.). The taxon was found in the Várpalota and W Bakony Sarmatian.
It occurs in the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) series and in the base layer of Steinheim.
Gyraulus trochiformis denudatus (HILGENDORF), 1866
1885 Planorbis denudatus — QUENSTEDT: p. 24, Pl. 49 Fig. 11
1900 Gyraulus (Gyraulus) Zieteni var. denudatus — MILLER: p. 404, Pl. 7 Fig. 30
1922 Gyraulus trochiformis denudatus — WENZ: p. 149, Fig. 69
1923 Gyraulus Gyraulus trochiformis denudatus — WENZ F. C.: p. 1619
1959 Valvata simplex — BODA: p. 632, 729, Pl. 23 Fig. 21
Hitherto the species was only known from Steinheim, but a damaged specimen was collected from the Sarmatian of
Várpalota.
Gyraulus goussardianus (NOULET), 1854
Plate XIX Figures 18–20
1881 Planorbis goussardianus — BOURGUIGNAT: p. 138, Pl. 8 Figs 254–257
partim! 1923 Gyraulus (Gyraulus) goussardianus — WENZ F. C.: p. 1555
2000 Gyraulus (Gyraulus) goussardianus — FISCHER: p. 138. Figs 9a–c
Specimens from the Upper Badenian of Pusztamiske wells may be identified with this form. The shell is comparatively flat and it is built of convex, slightly involute whorls. The aperture is moderately elongate; it embraces the previous whorl almost symmetrically, but generally it is slightly downturned toward the base. In this way the basal side is
deeper and concave.
It occurs in France, in coeval layers of Sansan.
Gyraulus homalosomus (BRUSINA), 1902
Plate XX Figure 1
1902 Planorbis homalosomus — BRUSINA: Pl. 5 Figs 34–36
1921 Gyraulus rhytidophorus homalosomus — WENZ: p. 76
1923 Gyraulus (Gyraulus) homalosomus homalosomus — WENZ F. C.: p. 1556
Both the lower and upper sides are flat. On the upper side the last whorl bears curved growth ribs. One specimen was
found in the Várpalota Sarmatian (i.e. well Inota–135). It is known from the Leobersdorf (Austria) and the Zagreb Sarmatian.
58
JÓZSEF KÓKAY
Genus: Hippeutis CHARPENTIER, 1837
Hippeutis fasciata GOTTSCHICK, 1920
Plate XX Figure 2–4
1920 Hippeutis fasciata — GOTTSCHICK: p. 171, Pl. 7 Fig. 10
1923 Hippeutis (Hippeutis) fasciata — WENZ F. C.: p. 1637
1998 Hippeutis fasciata — FINGER: p. 15. Taf. 6. Figs G–I
Some specimens of this extraordinary form originated from the upper part of the Sarmatian of Várpalota. It conforms
well with the type from Steinheim. In the Hungarian specimens the keel is slightly better developed and the first two or
three whorls are smooth.
Genus: Segmentina FLEMING, 1817
Segmentina larteti (NOULET), 1854
Plate XX Figure 5
1874 Planorbis (Segmentina) Larteti — SANDBERGER: p. 542, 579, Pl. 28 Fig. 23
1881 Segmentina Larteti — BOURGUIGNAT: p. 123, Pl. 8 Figs 281–284
1893 Segmentina Larteti — LOCARD: p. 133, Pl. 8 Fig. 1
1923 Segmentina larteti larteti — WENZ F. C.: p. 1663
1976 Segmentina larteti larteti — SCHLICKUM: p. 6 Pl. 1 Fig. 18
1998 Segmentina larteti — FINGER: p. 15. Taf. 7. Figs A–C
Numerous damaged, broken specimens originate from the Lower Badenian of Herend and from the Upper Badenian
of Pusztamiske, Nyirád and Fehérvárcsurgó. In some layers of the Sarmatian Várpalota it is frequent. It is also known
from Sansan of Southern France and from the North Alpine “silvana” beds.
Genus: Planorbarius FRORIEP, 1806
Planorbarius cornu cornu (BRONGNIART), 1810
Plate XX Figure 6
1874 Planorbis cornu — SANDBERGER: p. 374, 370, Pl. 18 Fig. 12, Pl. 20 Fig. 26
1874 Planorbis cornu var. subteres — SANDBERGER: p. 452, Pl. 21 Fig. 5
1893 Planorbis cornu — LOCARD: p. 141, Pl. 8 Fig. 8
1910 Planorbis (Coretus?) cornu — ROLLIER: p. 130, Pl. 2 Figs 17–24
1921 Planorbis cornu — WENZ: p. 128, 176, Pl. 25 Figs 13–14
This snail existed over a long range of time: from the Middle Eocene to the Late Miocene (“Pannonian” s. str). It was
detected in the Lower Badenian of Nagygörbő, in the Upper Badenian of Hegymagos, Pusztamiske and Várpalota. WENZ
(1923: p. 1426) regards several subspecies mentioned the literature as synonymous, because transitional forms exist. This
is just one example of the everlasting problem of morphospecies.
Planorbarius cornu solidus (THOMAE), 1845
1845 Planorbis corniculum — THOMAE: p. 154, Pl. 4 Fig. 7
1862 Planorbis incrassatus — RAMBUR: vol. 10 p. 177, vol. 8 Pl. 8 Figs 3–4
1863 Planorbis solidus — SANDBERGER: p. 71, Pl. 7 Fig. 8
1874 Planorbis cornu var. solidus — SANDBERGER: p. 524, Pl. 26 Fig. 16
1881 Planorbis solidus — BOURGUIGNAT: p. 127, Pl. 7 Figs 211–213
1893 Planorbis solidus — LOCARD: p. 139, Pl. 8 Figs 6, 7
This form deviates from the nominal subspecies in being less flattened, the aperture is less marginated, and a longitudinal lineation is well-developed. It differs from the subspecies P. cornu mantelli (DUNKER) in that its whorls follow
each other symmetrically, and it is similar to those of the nominal form. It is rather involute.
It occurred in the Lower Badenian coal-seam bearing series of the well Herend H–20, in an alluvial mud-like layer.
It is known from the Middle Miocene freshwater formations of Central and Western Europe. WENZ (1923) did not differentiate it from the nominal subspecies.
Some fragments from the base of the Várpalota coal-seam probably belong here. The margins on the aperture are
clearly visible on these.
Systematic descriptions
59
Planorbarius cornu mantelli (DUNKER), 1848
Plate XX Figure 7
1847 Planorbis pseudammonius — KLEIN: p. 77, Pl. 1 Fig. 23
1853 Planorbis platystomai — KLEIN: p. 219, Pl. 5 Fig. 16
1874 Planorbis Mantelli — SANDBERGER: p. 577, Pl. 28 Fig. 18
1893 Planorbis Mantelli — LOCARD: p. 142, Pl. 8 Figs 9–11
1910 Planorbis (Coretus) solidus — ROLLIER: p. 133, Pl. 3 Figs 6–8, 11–13
1921 Planorbis cornu mantelli — WENZ: p. 185, 189, Pl. 27 Figs 20–21
1932 Planorbis (Coretus) cornu var. mantelli — PEYROT: p. 255, Pl. 15 Figs 23–25
1954 Planorbis cornu mantelli — CSEPREGHY-MEZNERICS: p. 60, Pl. 8 Fig. 10
1976 Planorbarius cornu mantelli — SCHLICKUM: p. 7, Pl. 1 Fig. 19
Dunker’s subspecies deviates from the nominal form in having an asymmetric last whorl that is slightly downturned,
a spiral lineation that is frequent on the surface, and the umbilicus is deeper.
An adult specimen was found in the Hegymagos well and it is also present in the freshwater Badenian at Herend and
Nyirád. Numerous specimens were collected from the Sarmatian of Tapolca, Pusztamiske and Várpalota.
It is also known from the Middle Miocene of Central and Western Europe.
Planorbarius sansaniensis (NOULET), 1854
Plate XX Figures 8–13
1881 Planorbis sansaniensis — BOURGUIGNAT: p. 128, Pl. 7 Figs 216–218
1881 Planorbis telaeus — BOURGUIGNAT: p. 129, Pl. 7 Figs 214–215
1881 Planorbis anabaeus — BOURGUIGNAT: p. 129, Pl. 7
1886 Planorbis (Hemisoma) sansaniensis — ŁOMNICKI: p. 83, Pl. 2 Fig. 48
1916 Planorbis sansaniensis — DOLLFUS: p. 372, Pl. 6 Figs 19–22
1923 Planorbarius sansaniensis — WENZ F. C.: p. 1476
1932 Planorbis (Coretus) sansaniensis — PEYROT: p. 256, Pl. 15 Figs 18, 26–28
2000 Planorbarius sansaniensis — FISCHER: p. 139. Figs 5a–c
A relatively thick, strongly involute form, reflecting a fast pace of growth. The shell surface is sometimes decorated
with a spiral lineation forming a reticular pattern with the growthlines. The spiral lienation may be undulate. Its shell is
not as high as that of Planorbarius thiollieri (MICH.). Its aperture is slightly marginated; the umbilicus is quite deep due
to the thick whorls.
From the uppermost Badenian of the Hegymagos drilling (Tapolca Basin) and the Pusztamiske (Pm–2) well damaged
adults and entire subjuvenile specimens were collected. The best one came from the Várpalota Sarmatian.
It has also been found in France (Sansan), in the “Tortonian” of Southern Poland (“Galicia”) and there is some evidence that it may be present in the North Alpine Molasse Zone (Lower and Middle Miocene).
Familia: Ferrissiidae
Genus: Ferrissia WALKER, 1903
Ferrissia wittmanni (SCHLIKUM), 1964
Plate XX Figure 14
1964 Ancylus wittmanni — SCHLICKUM: p. 15, Pl. 2 Figs 36–38
1966 Ancylus wittmanni — SCHLICKUM: p. 326, Pl. 13 Fig. 29
1970d Ancylus wittmanni — SCHLICKUM: p. 180, Pl. 3 Figs 11–13
1989 Ferrissia wittmanni — REICHENBACHER: p. 144, Pl. 1 Fig. 8
One sizeable specimen was found in the mollusc-bearing hangwall of the Várpalota Upper Badenian coal-seam. It
has the following dimensions: length — 10.5 mm; width — 7.0 mm; height — 2.5 mm. It is slightly more flattened (l/h=
4.2) than Schlikum’s specimens (l/h= 3.8-3.9), but this deviance is within the range of variability of the species. Its outline is oval and its posterior margin is blunter than the anterior one. Its apex is in the midline, slightly behind the centre
and it is gently bent rightwards. The fine radial ribs increase in strength from the apex toward the anterior margin. The
specimens from the Nagygörbő Lower Badenian are smoother. A less flattened form occurs in the Middle Badenian of
Hidas (Mecsek Mts).
It deviates from F. deperdita (DESM.) because of its wider oval outline and its bigger size. It occurs in the North Alpine
Freshwater Molasse (“Süssbrackwassermolasse”) which is coeval with the Ottnangian–Badenian Paratethyan stages.
60
JÓZSEF KÓKAY
Ferrissia deperdita (DESMAREST), 1814
Plate XX Figure 15, Plate XXI Figure 1
1877 Ancylus Senckenbergianus — BOETTGER: p. 200, Pl. 29 Fig. 7
1921 Ancylus deperditus senckenbergianus — WENZ: p. 189, Pl. 27 Fig. 17
1923 Pseudancylus senckenbergianus — WENZ F. C.: p. 1696
1959 Acroloxus deperditus — BODA: p. 639, 737, Pl. 37 Fig. 8
1975 Ferrissia senckenbergiana — WAUTIER: p. 425, Pl. 42 Figs 1–9
1976 Ferrissia deperdita — SCHLICKUM: p. 7, Pl. 1 Fig. 20
Numerous specimens were collected from the Lower Miocene of Somlóvásárhely, and the Badenian and Sarmatian
of the Bakony Mts. In some layers it is frequent. Its outline is elongate oval. Its apex is situated slightly towards the right
side, and around it there is a minute radial lineation. It is widespread in the Miocene (approximately the equivalent of
the Karpatian–Sarmatian) of Central and Western Europe.
Familia: Ancylidae
Genus: Ancylus O. F. MÜLLER, 1774
Ancylus moravicus ŘZEHAK, 1893
Plate XXI Figures 2–3, 4
1972 Ancylus moravicus — ČTYROKY: p. 83, Figs 14–16, Pl. 6 Figs 4–10
There is a slightly fragmented specimen from the Upper Badenian hangwall of the Várpalota coal-seam which best
agrees with Rzehak’s species. It is a relatively wide shell (length — 5.9 mm; width — 4.3 mm; w/l — 73%). Its outline
approaches a slightly rounded square and its apex is gently placed towards the rightward. Čtyroky’s figures and descriptions reveal that its outline length/width ratio (61–83%) and height/width ratio (30–45%) are quite variable. The lastmentioned ratio is 37% in the Hungarian specimen. The specimens from the Chech Republic (Moravia) are characterised
by strong, concentric (sometimes step-like) growth-lines. The Várpalota specimen is smoother but similarly smooth
examples occur in Řzehak’s material as well. Traces of radial ribs are visible only under a strong lens, at the right side
of the apex. This species (forma obtusus ŘZEHAK 1893) has been found in the Lower Miocene of the Somlóvásárhely
drilling.
Earlier, it was known only from the Rzehakia-beds of Ottnangian age.
Familia: Acroloxidae
Genus: Pseudancylastrum LINDHOLM, 1909
Pseudancylastrum deperditolacustris (GOTTSCHICK), 1911
Plate XXI Figure 5
1911 Ancylus (Acroloxus) deperdito-lacustris — GOTTSCHICK: p. 521, Pl. 7 Fig. 14
1923 Ancylus deperditolacustris — WENZ F. C.: p. 1686
This form has been identified as being separate from the extant Acroloxus lacustris (LINNAEUS). However, this does
not seem to be well-justified. The fossil form is slightly lower, more elongate and its leftward-turned apex is more acute.
The specimen collected from the Lower Badenian coal-seam bering series is fragmented, and its margin is damaged.
Around its apex, under a strong magnification, a radial rib system is visible and this is a feature of the genus. The shell
is glass-like and translucent.The form has been identified in the Steinheim beds.
Pseudancylastrum decorata n. sp.
Plate XXI Figures 6, 7, 8
Derivatio nominis: decoratum: decorated (Latin)
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 105
Locus typicus: Inota, well Inota–82, 122.2–123.8 m
Stratum typicum: Sarmatian freshwater clay
Descriptio: The dimensions of the holotype are the following: length — 2.7 mm; width — 1.9 mm; height — 0.7 mm.
Systematic descriptions
61
The shell is oval and if widens gently towards the anterior. The apex is close to the posterior margin and it is slightly towards the turned left. Its point is slightly depressed. The shell is thin, fragile, with a clearly visible radial ribbing.
Its concentric ribs are better described as growth-lines. The species is frequent, but entire specimens are exceptional and
even these are not completely developed. The adult specimens may be 4–5 mm long.
Diagnosis differentialis: the closest relative is Pseudancylastrum deperditolacustris (GOTTSCHICK). It differs due to
its elongate form, its less developed radial ribbing, and its glass-like, translucent shell. The species generally occurs within freshwater faunas.
Subjuvenile specimens of this species have been recognised in the Lower Miocene of the Somlóvásárhely drilling.
Familia: Cochlicopidae
Genus: Azeca LEACH, 1831
Azeca cf. peneckei ANDREAE, 1908
Plate XXI Figure 9
1891 Azeca boettgeri — PENECKE: p. 364, Pl. 21 Fig. 8
1902 Azeca Peneckei — ANDREAE: p. 15
1923 Azeca (Azeca) peneckei — WENZ F. C.: p. 1095
In the Upper Badenian of Herend a last whorl was found, with a slightly damaged aperture. The form of the aperture
is an elongate triangle, with dental plates in the pharynx. The upper part of the whorl is axially wrinkled.
The species is known also from the “Tortonian” (= Badenian?) beds of Styria and of Silesia, S Poland (Opole).
Genus: Cochlicopa RISSO, 1826
Cochlostoma subrimata loxostoma (KLEIN), 1853
Plate XXI Figure 10
1853 Achatina loxostoma — KLEIN: p. 214, Pl. 5 Fig. 12
1916 Cochlicopa subrimata var. loxostoma — GOTTSCHICK et WENZ: p. 71, Pl. 1 Figs 13, 14
1923 Cochlicopa subrimata loxostoma — WENZ F. C.: p. 1107
1981 Cochlicopa subrimata loxostoma — LUEGER: p. 16, Pl. 1 Fig. 4
From the upper part of the Várpalota Sarmatian a last whorl was found. The species is known from coeval beds in
Steinheim as well as from the Lower Pannonian of the Vienna Basin.
Cochlostoma subrimata cf. procera GOTTSCHICK, 1920
Plate XXI Figure 11
1920 Cochlicopa subrimata var. Procera — GOTTSCHICK: p. 63
1923 Cochlicopa subrimata procera — WENZ F. C.: p. 1108
Two last whorls were found in the Upper Badenian (well He–20). The apertural margin of these whorls is damaged.
The height of the last whorl indicates a total height of about 2.2 mm and so this is a small form. According to WENZ
(1923) the subspecies of Cochlicopa subrimata occur in the Alpine–Carpathian Miocene. Among these, C. s. procera is
the smallest, having the most convex whorls. In the latter respect the two fragments agree with GOTTSCHICKS’s (1920)
description. Unfortunately, no figure has been published showing this subspecies. It also in the Steinheim a. A.
Familia: Vertinigidae
Genus: Negulus BOETTGER, 1889
Negulus suturalis suturalis (SANDBERGER), 1858
Plate XXI Figures 12, 13
1863 Pupa (Pupilla) suturalis — SANDBERGER: p. 54. Pl. 6. Fig. 1
1914a Negulus suturalis — FISCHER et WENZ: p. 92. Pl. 5. Fig. 13
1923 Negulus suturalis — WENZ F. C.: p. 1024
62
JÓZSEF KÓKAY
Several specimens have been recognised in the Lower Miocene of the Somlóvásárhely drilling. Its stratigraphic range
is from the Upper Oligocene to layers coeval with the Badenian in the Northern Alpine foreland.
Negulus suturalis gracilis GOTTSCHICK et WENZ, 1919
Plate XXI Figure 14
1919 Negulus suturalis gracilis — GOTTSCHICK et WENZ: p. 9. Pl. 1. Figs 11–13
1923 Negulus suturalis gracilis — WENZ F. C. p. 1027
1981 Negulus suturalis gracilis — LUEGER: p. 18. Pl. Fig. 2
1996 Negulus gracilis — FORDINAL: p. 6. Pl. 1. Figs 3–4
From the nominal subspecies this form deviates because of its smaller and narrower shell. It has been found in the
Upper Badenian of the Hegymagos drilling as well as in the Sarmatian and Upper Badenian of Várpalota. It is also reported from Sarmatian and Pannonian deposits.
Genus: Acmopupa BOETTGER, l889
Acmopupa? nov. sp.
Plate XXI Figure 15
Two small shells were found in the Lower Miocene of the Somlóvásárhely drilling.The better-preserved one is 2.3
mm high and about 1.3 mm wide. Its last whorl is 1.5 mm high. The shell is conical with four whorls. The apex is obtuse.
The whorls are slightly convex and the sutures are moderately deep. The base is not perforated, and instead there is a
small depression. The shell is completely smooth and shiny, similar to white china. The external margin of the aperture
is missing and the columellar margin and the left lower margin are slightly bent outwards, indicating an adult age. The
beginning of the basal margin is thin suggesting a sharp external margin; unfortunately the latter is missing from our
specimens. The specimens may not be taxonomically well-placed. They are perhaps the closest to the genus Acmopupa
(BOETTGER 1889: p. 271, FISCHER, WENZ 1914a: p. 93, WENZ, ZILCH 1959: p. 148) having an obtuse apex, an aperture
without teeth, and a low number of convex whorls. On the other hand, this form differs significantly from the nominal
form because there is no axial lineation and the base is not perforated. It is possible that it represents a new subgenus.
Genus: Vertigo O. F. MÜLLER, 1774
Subgenus: Ve r t i g o s . str.
Vertigo kochi BOETTGER, 1889 var.
Plate XXII Figures 1–4
1889 Vertigo (Alaea) kochi — BOETTGER: p. 304, Pl.? Fig. 9
1902 Vertigo kochi — ANDREAE: p. 19 (No. 18)
1914a Vertigo (Vertigo) kochi — FISCHER et WENZ: p. 101, Pl. 6 Fig. 26
1921 Vertigo (Vertigo) kochi — WENZ: p. 147, Pl. 18 Figs 36, 37
1923 Vertigo (Vertigo) kochi — WENZ F. C.: p. 995
Numerous extremely well-preserved specimens have been recovered from the Upper Badenian of the Pusztamiske
well. One specimen was found in the coal-seam series of the well Pm–4, two in the well Pm–1 and another one in the
drilling Pm–2, in the uppermost Badenian. Apart from the Hungarian specimens, the form has up to now only been found
in the Upper Oligocene of the Mainz Basin and from the “Tortonian” of Opole (Silesia, Poland). For this reason a detailed
comparison with Hungarian speciemns will be given below.
The Mainz specimens have 8 teeth:
— 3 parietal teeth: the median is long, the lateral pair smaller, and of the latter infraparietal one is the smallest;
— two columellar teeth of which the lower one is the smaller;
— 3 parietal teeth: the uppermost is the smallest; the second one strong and high; the third one is in a deep setting
— it is very strong, wide and flattened.
In the specimen from the well Pm–4 the three parietal teeth are similar to those described above. The lower columellar tooth is smaller than the upper. A tiny basal teeth is in excess if compared to the Mainz specimens. The three palatal
teeth are similar to those of Boettger’s species. Specimens from the well have a similar dentation but the tiny parietal
tooth is missing, in contrast to of the Mainz, Pm–2 and Pm–4 ones. The small basal tooth is also in excess. Thus the
Hungarian specimens possess 8 or 9 teeth.
The apertures of the Pusztamiske specimens are a rounded triangular shape. The specimen from the well Pm–1 has
a narrow and sharp apertural margin around its aperture due to its completely adult state. Deviations in the dentation
from the specimens of Boettger are not significant, given that similar variations occur within one and the same popula-
Systematic descriptions
63
tion. One trait is different in the Pusztamiske form: behind the aperture there is a swelling. Before this, one or two short
furrows are visible, running parallel to the suture. This feature is not mentioned by Boettger. This seems not to warrant
a specific difference and therefore the deviation is regarded as a variation (“var.”). The Hungarian specimens are slightly bigger than the German and Polish ones: height — 1.6 mm; width — 1.0 mm.
Vertigo protracta suevica GOTTSCHICK et WENZ, 1919
Plate XXII Figures 5–7
1919 Vertigo (Alaea) protracta suevica — GOTTSCHICK et WENZ: p. 21, Pl. 1 Figs 40, 41
1923 Vertigo (Vertigo) protracta suevica — WENZ F. C.: p. 1001
1981 Vertigo (Vertigo) protracta suevica — LUEGER: p. 21, Pl. 2 Figs 14, 15
This species was found in the Upper Badenian of Pusztamiske and the Tapolca Basin, as well as in the Sarmatian of
Várpalota. The only deviation from the type is the presence of an extra small infraparietal tooth.
The closely Vertigo callosa (REUSS) is generally bigger, relatively wider and more inflated, and its dentation is stronger.
This form is known from Steinheim and from the Pannonian and Pontian of the Vienna Basin.
Vertigo callosa (REUSS), 1860
Plate XXII Figures 8–10
1860 Pupa (Vertigo) callosa — REUSS: p. 72, Pl. 2 Figs 6, 7
1874 Pupa (Vertigo) callosa — SANDBERGER: p. 400, Pl. 34 Fig. 10
1919 Vertigo (Alaea) callosa — GOTTSCHICK et WENZ: p. 13, Pl. 1 Figs 26–28
1921 Vertigo (Vertigo) callosa — WENZ: p. 983
1981 Vertigo (Vertigo) callosa — LUEGER: p. 20, Pl. 2 Figs 3–5
This species illustrates a number of variations, ranging from the Upper Oligocene to the Upper Miocene of Central
and Western Europe. Some authors, especially Lueger and Wenz, clustered many of its forms as synonyms.
The form occurred in the Upper Badenian of Hegymagos and in the Sarmatian of Várpalota. The species has been
recognised in the Lower Miocene of the Somlóvásárhely drilling.
Vertigo callosa diversidens (SANDBERGER) 1874
Plate XXIII Figures 2–3, 9
1874 Pupa (Vertigo) diversidens — SANDBERGER p. 549. Pl. XXIX. Fig. 23
1919 Vertigo (Alea) callosa diversidens — GOTTSCHICK et WENZ 15. Pl. 1. Fig. 31
1923 Vertigo (Vertigo) callosa diversidens — WENZ F. C. 988
1998 Vertigo callosa diversidens — FINGER Pl. 9. Fig. E
The Upper Badenian of the well Pm–1 and the Sarmatian of Várpalota yielded examples of the subspecies V. callosa
diversidens.
Vertigo callosa convergens (BOETTGER), 1889
Plate XXIII Figure 4
1889 Vertigo (Alaea) callosa mut. convergens — BOETTGER: p. 295, 297, 311. Pl.7. Fig. 4
1923 Vertigo (Vertigo) callosa convergens — WENZ F. C.: p. 987. in synonymy
Individual specimens from Hegymagos and Pusztamiske (Pm–4), respectively, of the Upper Badenian series seem to
belong to the subspecies V. callosa convergens.
Vertigo callosa cardiostoma (SANDBERGER), 1874
Plate XXIII Figure 5
1874 Pupa cardiostoma — SANDBERGER: p. 600
1891 Pupa (Alaea) cardiostoma — FLACH: p. 56. Pl. 3. Figs 5
1919 Vertigo (Alea) callosa cardiostoma — GOTTSCHICK et WENZ: p. 14. Pl. 1. Fig. 30
In the Upper Badenian of the well Pm–1 the form V. callosa cardiostoma was also found.
Vertigo callosa perarmata GOTTSCHICK et WENZ, 1919
Plate XXIII Figures 1, 6–8
1919 Vertigo (Alaea) callosa perarmata — GOTTSCHICK et WENZ: p. 15, Pl. 1 Figs 34, 35
1923 Vertigo (Vertigo) callosa perarmata — WENZ F. C.: p. 993
1998 Vertigo callosa perarmata — FINGER: Pl. 9. Fig. F
64
JÓZSEF KÓKAY
The Várpalota Sarmatian yielded this form. It has more teeth than the nominal form, having altogether nine or ten. It
is also smaller, and is known from the Steinheim beds.
Vertigo ovatula trolli WENZ, 1914
Plate XXIV Figures 1–2
1914a Vertigo trolli — FISCHER et WENZ: p. 102, Pl. 7 Fig. 27
1923 Vertigo (Vertigo) ovatula trolli — WENZ F. C.: p. 1000
1981 Vertigo (Vertigo) ovatula trolli — LUEGER: p. 21, Pl. 2 Figs 6, 7
Specimens from the Upper Badenian of the Hegymagos well and in the Sarmatian of Várpalota have slightly deeper
situated columellar teeth than the typical form. The Várpalota ones have a deep-situated basal dental slat. In the
Hegymagos specimens these are parallel.
This form is known from the Central European Middle and Upper Miocene. The locality outside and closest to
Hungary is that of the Pannonian of Leobersdorf (LEUGER 1981).
Vertigo ovatula miliiformis BOETTGER, 1884
Plate XXIV Figure 3
1884 Vertigo (Alata) ovatula var. miliiformis — BOETTGER: p. 270, Pl. 4 Fig. 9
1889 Vertigo (Alata) ovatula mut. miliiformis — BOETTGER: p. 302
1923 Vertigo (Vertigo) ovatula miliiformis — WENZ F. C.: p. 999
In the Upper Badenian of the Hegymagos this species, can easily be identified, although an axial lineation on the shell
surface is barely visible. This feature is just vaguely depicted on BOETTGER’s (1884) figure.
This form has also been found in the “Landschneckenmergel” (= terrestrial snail-bearing marl) near Frankfurt a. M.,
and this is coeval with the Badenian.
Vertigo pusilla sarmatica nov. sp.
Plate XXIV Figure 6
Derivatio nominis: Sarmatian = sarmatica (Lat.)
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 106
Locus typicus: Várpalota, Shaft M. III, 112 m depth
Stratum typicum: upper part of the Sarmatian, freshwater clay
Descriptio: Dimensions of the holotype: height — 2.0 mm; width — 1.2 mm; width/height ratio: 0.6.
The shell is sinistrally coiled, it has five whorls, and is barrel-shaped. The whorls are gently convex, and the growthlines are barely visible. The aperture is heart-shaped. The external apertural margin, is medially, strongly infolded. In the
aperture there are two parietal, two columellar and two palatal teeth; furthermore there is a tiny extra tooth at the columellar angle of the parietal part. The teeth are quite wide, strong, and well developed.
Descriptio differentialis: the new form is the closest to the modern V. pusilla MÜLLER. The differences indicate a separation on the subspecies level:
— V. pusilla is narrower, its has an average height of 2.0 mm, and its width 1.0–1.1 mm.
— The teeth are narrower, and filigree. The dentation shows a number of variations within and between populations.
Its sixth teeth are usually visible, but often there is a basal dental plate as well. The small infraparietal teeth, clearly visible on the new subspecies, is lacking from the nominal form.
— The aperture of V. pusilla is horseshoe-shaped. At the medium of the palatal side there is no depression; however, this is well-developed in the Miocene subspecies.
Another, damaged specimen is similar to the holotype.
V. pusilla moedlingensis WENZ et EDLAUER (1942: p. 89, Pl. 4 Figure 9) has been described from the Pontian of the
Vienna Basin. It is smaller than the Várpalota or the modern form (height — 1.6 mm; width — 1.0 mm) and its dentation is different (it has a basal tooth).
Vertigo bakonyensis nov. sp.
Plate XXIV Figure 4–5; Textfigure 4
Derivatio nominis: it originates from the Bakony Mts
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 107
Locus typicus: Pusztamiske, well Pm–3, 14.7–14.8 m
Stratum typicum: Sarmatian, brackish coaly clay
Systematic descriptions
65
Descriptio: Dimensions of the holotype: height — 1.5 mm; width — 0.8 mm.
It is a cylindrical shell, gently convex with five moderately convex whorls, and it has growthlines. The aperture is slightly thickened, its margin is turned outwards, and it forms a peristom.
There is no occiputal swelling on the last whorl. There is a depression along the middle of the
whorl, paralled with the suture, starting at the aperture. It is almost as deep as the suture and its
length is equal to one quarter of the whorl. The outline of the aperture is obliquely oval, and there
is a slight embayement at the external margin. There is a thin anguloparietal dental plate in the
aperture, which may be followed up to the parietal margin, which extends deeply inward. A less
developed parallel dental plate does not reach the parietal margin, but extends deeply into the Textfigure 4. Vertigo
pharynx. There is a sharp parietal plate inside, corresponding to the depression on the last whorl. bakonyensis nov. sp.
This does not reach the aperture. On the columellar side, deep inside, there is a tooth like tubercle, turned inwards.
Diagnosis differentialis: The new species is the closest to the extant V. alpestris ADLER 1838, but is slightly smaller.
Its whorls are more flat, the suture less deep, and the dentation is quite different. The outline of the aperture is similar.
Only one specimen of this new form has so far bean found.
Subgenus: Ve r t i l l a
Vertigo (Vertilla) angustior ocsensis (HALAVÁTS), 1903
1923 Vertigo (Vertilla) angustior öcsensis — WENZ F. C.: p. 1007
1942 Vertigo (Vertilla) angustior öcsensis — WENZ et EDLAUER: p. 90, Pl. 4 Fig. 10
1959 Vertigo (Vertilla) angustior öcsensis — BARTHA: p. 79, Pl. 15 Figs 9, 10
1959 Vertigo (Vertilla) angustior — BODA: p. 640, 738, Pl. 36 Fig. 1
1981 Vertigo (Vertilla) angustior öcsensis — LUEGER: p. 22, Pl. 2 Fig. 8
1974 Vertigo (Vertilla) angustior öcsensis — PAPP: p. 383, Pl. 17 Fig. 6
Numerous specimens were found in the upper part of the Várpalota Sarmatian. It occurs in the Sarmatian, Pannonian and
Pontian of the Vienna Basin, as well as in the Transdanubian Pontian. BODA (1959) detected it in the Várpalota Sarmatian.
Familia: Chondrinidae
Genus: Gastrocopta WOLLASTON, 1878
Subgenus: A l b i n u l a
Gastrocopta (Albinula) acuminata acuminata (KLEIN), 1846
Plate XXIV Figure 7
1923 Gastrocopta (Albinula) acuminata acuminata — WENZ F. C.: p. 916
1974 Gastrocopta (Albinula) acuminata acuminata — PAPP: p. 384, Pl. 17 Fig. 11
1981 Gastrocopta (Albinula) acuminata acuminata — LUEGER: p. 23, Pl. 2 Fig. 10
This form is known from the Central European Middle and Upper Miocene. It is a newly-discovered in the type from
Hungarian Badenian from the Hegymagos well in the Tapolca Basin.
Gastrocopta (Albinula) acuminata larteti DUPUY, 1850
1850 Pupa larteti — DUPUY: p. 307, Pl. 15 Fig. 5
1919 Leucochila acuminata larteti — GOTTSCHICK et WENZ: p. 11
1923 Gastrocopta acuminata larteti — WENZ F. C.: p. 919
1959 Gastrocopta acuminata larteti — BARTHA: p. 79, Pl. 15 Fig. 1
1959 Gastrocopta acuminata larteti — BODA: p. 639, 738, Pl. 35 Figs 15, 16
1981 Gastrocopta (Albinula) acuminata larteti — LUEGER: p. 24, Pl. 2 Fig. 11
BODA (1959) found this form in the Várpalota Sarmatian. It is wider than the previous species and has a barrel-like
shape. It is widespread in the Middle and Upper Miocene of Central and Western Europe, and it also occurs in the Pontian
of Várpalota.
Gastrocopta (Albinula) acuminata procera (GOTTSCHICK et WENZ), 1916
Plate XXIV Figure 8
1919 Leucochila acuminata procera — GOTTSCHICK et WENZ: p. 11, Pl. 1 Figs 18, 19
66
JÓZSEF KÓKAY
It has an elongate cylindrical, smooth form, with a horseshoe formed aperturte. One fragment — a last whorl — from
the Várpalota Sarmatian, agrees well with the Steinheim type specimens. It has been found in the Sarmatian of Várpalota.
Subgenus: S i n a l b i n u l a
Gastrocopta (Sinalbinula) nouletiana nouletiana (DUPUY), 1850
Plate XXIV Figures 9, 10, b, Plate XXV Figure 1
1923 Gastrocopta (Sinalbinula) nouletiana nouletiana — WENZ F. C.: p. 930
1974 Gastrocopta (Sinalbinula) nouletiana nouletiana — PAPP: p. 385, Pl. 17 Fig. 8
1981 Gastrocopta (Sinalbinula) nouletiana nouletiana — LUEGER: p. 25, Pl. 2 Figs 16–19, 22
The species is not rare in the Central European Middle and Upper Miocene. No Badenian occurrence has been found
so far. It has been found in the Upper Badenian of the Pusztamiske, Hegymagos and Fehérvárcsurgó wells, as well as
from the Sarmatian of Várpalota.
The species described by DUPUY (1850: p. 309, Pl. 15 Figure 6) deviates from our specimens and from those described
in the above references. The main difference is in the angulo-parietal main dental plate, which reaches higher onto the
parietal plate in Dupuy’s specimen and is less confluent with the external upper apertural margin.
Gastrocopta (Sinalbinula) nouletiana gracilidens (SANDBERGER), 1874
Plate XXV Figures 3, 4
1884 Leucochilus Nouletianum var. gracilidens — BOETTGER: p. 272. T. IV. Figs 12–13
1885 Pupa gracilidens — CLESSIN: p. 84, Pl. 7 Fig. 11
1923 Gastrocopta (Sinalbinula) nouletiana gracilidens — WENZ F. C.: p. 934
1979 Gastrocopta nouletiana gracilidens — KROLOPP: p. 291, Pl. 4 Fig. 1
The main deviation from the nominal subspecies is that its main parietal dental plate is narrower and slimmer, and
its outline is more cylindrical. It has been found in the Hegymagos drillings.
It is present in the Central European Miocene in levels coeval with the Badenian, and also in the Sarmatian of
Várpalota.
Gastrocopta (Sinalbinula) nouletiana tapeina (BOURGUIGNAT), 1881
Plate XXV Figure 2
1881 Vertigo tapeina — BOURGUIGNAT: p. 80, Pl. 4. Figs 110–113
The species has been described the from Sansan. It has been found in the Sarmatian of Várpalota (drilling I–118) as
well. WENZ (1923) includes seven different taxa of Bourguignat into Gastrocopta nouletiana. Although Bourguignat’s
figures are somewhat sketchy, it seems that Wenz’s procedure is not well founded. The specimen is very similar to the
type. Both have well developed anguloparietal lamelliform teeth. On the Sansan specimen there is a well visible axial rib
system, while the shell of Várpalota one is smooth.
Bourguignat’s specimens came from layers coeval with the Badenian.
Gastrocopta (Sinalbinula) suevica (BOETTGER), 1875
1900 Pupa (Leucochilus) suevica — MILLER: p. 398, Pl. 7 Fig. 16
1919 Leucochilus suevica — GOTTSCHICK et WENZ: p. 13, Pl. 1 Figs 24, 25
1923 Gastrocopta (Sinalbinula) suevica — WENZ F. C.: p. 937
1967 Gastrocopta (Sinalbinula) suevica — SCHÜTT: p. 208, Fig. 13
1974 Gastrocopta (Sinalbinula) suevica — PAPP: p. 385, Pl. 17 Fig. 9
1979a Gastrocopta (Sinalbinula) suevica — SCHLIKUM: p. 408, Pl. 23 Fig. 5
Some last whorls were found in the Upper Badenian of the Puszatmiske. The angular teeth are quite jagged and have
an inward bent branch. It is smaller than G. nouletiana, its aperture is more circular, and the teeth are smaller.
It is known from the Middle and Upper Miocene of the Paratethys.
Gastrocopta (Sinalbinula) ferdinandi (ANDREAE), 1902
Plate XXV Figure 5
1902 Leucochilus ferdinandi — ANDREAE: p. 18, Fig. 9
1923 Gastrocopta (Sinalbinulina) obstructa ferdinandi — WENZ F. C.: p. 929
1981 Gastrocopta (Sinalbinula) obstructa ferdinandi — LUEGER: p. 26, Pl. 2 Fig. 13
2004 Gastrocopta (Sinalbinula) ferdinandi — HARZHAUSER et BINDER: p. 127, pl. 8. Figs 12–14
Systematic descriptions
67
Two, excellently preserved specimens and some last whorls were found in the upper part of the Várpalota Sarmatian.
It has an elongate cylindrical form with moderately convex whorls. Its aperture is horseshoe shaped. The deth agree with
the figures, in the literature. The masin parietal dental plate is well-developed or gently jagged.
So far it has been found in the layers of Opole (S Poland, Silesia), which are coeval with the Upper Badenian, and
from the Pontian of the Vienna Basin.
Gastrocopta (?Sinalbulina) infrapontica WENZ, 1927
1927 Gastrocopta (Sinalbulina) fissidens infrapontica — WENZ: p. 47. Pl. 2. Fig. 8
1967 Gastrocopta (Sinalbulina) ferdinandi — SCHÜTT: p. 207. Fig. 11
1979a Gastrocopta (Sinalbulina) fissidens infrapontica — SCHLICKUM: Pl. 23. Fig. 6
1981 Gastrocopta (? Sinalbulina) fissidens infrapontica — LUEGER: p. 29. Pl. 2. Figs 20, 21
1996 Gastrocopta (? Sinalbulina) infrapontica — FORDINAL: p. 7. Pl. 1. Fig. 7
One specimen was found in the uppermost part of the Várpalota Sarmatian. It has also been found in the
Badenian, Sarmatian and Pannonian–Pontian layers. Wenz described the species from the Pannonian sediments of
Leobersdorf.
Familia: Pupillidae
Genus: Pupilla LEACH in TURTON, 1828
Pupilla iratiana suevica GOTTSCHIK et WENZ, 1919
Plate XXV Figure 7
1919 Pupilla iratiana suevica — GOTTSCHIK et WENZ: p. 5. Pl. 1. Figs 4–5
1923 Pupilla (Primipupilla) iratiana suevica — WENZ F. C.: p. 960
About a dozen damaged specimens were found in the Sarmatian of Várpalota (well Ősi–69). It is certain that it
belongs to the genus Pupilla. In most cases the aperture missing. The most complete specimen has at least six whorls;
these are only slightly convex and the sutures are shallow. The columellar axis of the shell is narrow and tube-like. Its
external appearance relates it to this genus. There are diagonal growth-lines on the surface. The most entire specimen is
2.3 mm high and 1.5 mm wide. The shell wall is apparently thick, but this is due to a subsequent, diagenetic infilling,
which is clear in one specimen in which the apical part is filled with calcite. Where the shell is broken off of this cement,
its thinness is obvious.
It is the closest to BOETTGER’s species (1889) which was found in “Landschneckenmergel” near to Frankfurt a. Main.
Genus: Leiostyla LOWE, 1852
Leiostyla gottschicki (WENZ), 1922
Plate XXV Figure 6
1922 Lauria (Leiostyla) gottschicki — WENZ: p. 107, Fig. 1
1923 Lauria (Leiostyla) gottschicki — WENZ F. C.: p. 1036
1967 Leiostyla (Leiostyla) gottschicki — SCHÜTT: p. 207, Fig. 10
One entire and several damaged specimens were collected from the Sarmatian of Várpalota (drilling Inota–135). All
have apertures which are filled in with a matrix, but the main teeth and dental plates are visible and agree well with
SCHÜTT’s (1967) description and figure.The Pannonian species L. austriaca WENZ 1921 has a more pronounced axial ribbing, more convex whorls and less deep sutures. The columellar teeth of these two species are very different.
The species has been reported from Steinheim and from the Sarmatian of Hollabrunn, Austria.
Genus: Argna COSSMANN, 1889
Argna oppoliensis (ANDREAE), 1902
Plate XXV Figure 8, Plate XLI Figure 5
1902 Coryna oppoliensis — ANDREAE: p. 16, Fig. 8a
1923 Agardhia oppoliensis — WENZ F. C.: p. 1038
1976 Argna oppoliensis — SCHLICKUM: p. 10, Pl. 2 Fig. 28
68
JÓZSEF KÓKAY
Fragmentary specimens were found in the Sarmatian of Várpalota (Shaft M. III, drillings Berhida Bh–3, Ősi–63 and
Cs–13). Its smooth shell surface is a typical characteristic. In some specimens there are minute growth wrinkles on the
upper part of the last whorl. In the aperture there is a sizeable parietal dental plate and a columellar plate. Deep in the
pharynx there are two or three palatal dental ridges. In the Lower and Upper Badenian of Herend some fragments of apertures were found.
It is known from Opole (S Poland) from layers coeval with the Badenian, as well as from the “silvana” beds of the
North Alpine Freshwater Molasse (“Süssbrackwassermolasse”).
Argna sumeghyi (BARTHA), 1956
Plate XXV Figure 9
1956 Agardia sümeghyi — BARTHA: p. 519, Pl. 4 Figs 3, 4, 7, 8
1959 Agardia oppoliensis turrita — BARTHA: p. 81, Pl. 15 Fig. 23
1959 Agardia sümeghyi — BARTHA: p. 81, Pl. 15 Fig. 17
1978 Argna oppoliensis — SCHLICKUM: p. 252, Pl. 19 Fig. 10
1981 Argna suemeghyi — LUEGER: p. 32, Pl. 19 Figs 9–11
Several last whorls were found in the upper part of the Várpalota Sarmatian and these show up the characteristic features. Besides the main parietal and columellar teeth there are two or three palatal wrinkles (blunt dental ridges) deep in
the aperture, as was also pointed out by LUEGER (1981). At the base there is a fissure. The shell is covered with faint and
irregular growth ribs. This ribbing is more regular and stronger in some specimens (new subspecies?). It has been found
in the Pontian of the Bakony Mts and the Vienna Basin. It is possible that a last whorl from the Upper Badenian of Herend
also belongs here.
Subgenus: A g a r d h i e l l a
Argna (Agardhiella) reperta (SANDBERGER), 1895
Plate XXV Figure 10
1921 Aghardia aperta — GOTTSCHICK et WENZ: p. 21, Fig. 1
1923 Aghardia reperta — WENZ F. C.: p. 1039
From the deeper part of the Sarmatian of Várpalota (well Ősi–69) one, almost entire specimen was collected. Another
relatively wider specimen was found from the Sarmatian of Várpalota (drilling Csór–13.). The parietal plate and the columellar apertural margin are slightly damaged. The external apertural margin is partly obscured by the matrix and the
aperture is evidently toothless. The umbilical fissure is extremely narrow such as to be almost nonexistent. According to
the original figure of GOTTSCHICK and WENZ (1921), the shell is built of five slightly convex whorls. The shell surface is
covered with a diagonal, sharp, dense and fine system of ribs. The dimensions and proportions also correspond to those
of the type form (height — 2.1 mm; width — 1.2 mm). It has also been found in the beds at Steinheim.
Argna (Agardhiella) sublamellata nov. sp.
Plate XXV Figures 11–12
Derivatio nominis: alluding to its similarity to A. lamellata
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 108
Locus typicus: Várpalota Basin, well Inota–135, 116–124.6 m
Stratum typicum: Sarmatian, freshwater clay
Descriptio: Tower like shell, thin-walled, with a cylindrical lateral outline and low whorls. The whorls are moderately convex, and the sutures gently deepened. The number of whorls and the shape of the juvenile examples are unknown.
Due to the lack of an entire specimen, one with the two last whorls was designated as the holotype. The shell surface is
covered with sharp, elevated axial and parallel ribs. The interribs are twice or three times wider than the ribs. On the base
there is a narrow umbilical fissure.
The aperture is elongate heart-shaped, with a slightly outward-bent margin. In the aperture there are four tooth-like
elevations. The uppermost of these is a parietal one on the middle of the frontal plate. On the margin of the external apertural margin there is a strong tooth, or marginal swelling. Externally, this part is marked with a small pit or embayment.
There are two weak columellar teeth on the left side of the straight apertural margin, which can be followed inside. These
are the endings of two spiral columellar plates. Pharyngal or palatal plates do not exist within the apertural margin. This
is a subgeneric feature.
Dimensions of the holotype: inferred height — 3–4 mm; width — 0.9 mm.
Systematic descriptions
69
Diagnosis differentialis: The new species is closest to the extant A lamellata (CLESSIN) (KERNEY et al. 1983: p. 124).
In this form the upper columellar tubercle is wider while the lower one is close to the base. Its umbilicus is closed. The
ribs on the surface are similar, but arranged obliquely rather than axially. It lives in wet environment, under boulders and
in crevices.
Genus: Enneopupa BOETTGER, 1889
Enneopupa subcylindrella nov. sp.
Plate XXV Figure 13
Derivatio nominis: alluding to its similarity to E. cylindrella
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 109
Locus typicus: Pusztamiske, well Pm–1, 52.7–52.8 m
Stratum typicum: Upper Badenian, coal-seam bearing beds
Descriptio: A last whorl, the aperture and a part of the whorl is missing. In spite of these facts the species may be
described using fragments from the Herend Basin. Its main characteristics are the following:
— it has a relatively flat whorl and upwards it narrows conically,
— the shell surface is regularly ribbed axially and the ribs may be followed to the basal pit,
— on the base there is a narrow, tube-like pit with a wider rim,
— externally, on the lower part of the whorl, close to the suture, there is a gentle depression or furrow,
— in the aperture there are two columellar dental plates (the upper one is longer and reaches forward to a greather
extent),
— behind it (deeper) there is one subcolumellar and one supercolumellar tooth,
— on the broken surface the cross-section of three palatal teeth or a wrinkle is visible,
— the parietal part of the aperture is missing,
— the whorl is 1.6 mm in diameter.
Descriptio supplementi: Similar fragments were found in the Herend coal-seam sequence (well Bánd–2). On these
the aperture, corresponding to that of other species of the genus, is visible. Supplementary to those characteristics visible on the holotype are the following:
— the pear-shaped aperture is marginated. There is a tooth on the middle of the columellar margin. In the inner, upper
corner of the aperture there is a marginal parietal tooth. On the external margin there is a palatal protuberance,
— behind the external margin of the aperture there are five elongate palatal teeth, respectively.
The dental wrinkles are parallel to each other. The uppermost one is elongate, less developed and it is just below the
suture. The fourth is also elongate and should be regarded as a dental plate. Behind these there are three hooked teeth,
standing across (the holotype specimen probably breaks along these):
— on a columellar fragment the teeth and plates correspond well to those visible in the holotype from Pusztamiske,
— the axial ribs are weaker on the Herend fragments than on the holotype.
Descriptio differentialis: the described species is very characteristic for this rare genus. It is the closest to the typespecies of this form, — that is, the Lower Miocene (from Hocheim) E. cylindrella (SANDBERGER). (WENZ 1923: p. 1012,
WENZ, ZILCH 1959: p. 172, Figure 592). Sandberger’s form has two columellar, two parietal and one angular teeth; inside
there are 3–4 palatal wrinkles or dental plates.
Genus: ?Paracoryna FLACH, 1890
Paracoryna pseudoennea (FLACH), 1890
Plate XXV Figure 14
1891 Pupa (Coryna) pseudoennea — FLACH: p. 51, Pl. 3 Fig. 3
1894 Ennea pseudoennea — CLESSIN: p. 26, Pl. 1 Fig. 5
1923 Agardhia pseudoennea — WENZ F. C.: p. 1039
Several last whorls were found in the Sarmatian of Várpalota. It has just one parietal tooth, but in some specimens
even this is lacking. Deep in the pharynx there are two palatal dental wrinkles whose direction indicates a backward tendency. The shell is realtively slim, with an umbilical fissure and smooth surface. Its generic position has not been determined for certain, (WENZ, ZILCH 1959, p. 172) but it certainly cannot be ranged into the subgenus Agardhiella, as this
does not possess palatal pharyngal plates.
It is known from the upper part of the Lower Miocene in the Czech Republic as well as from the “Undorf” coal-seam
sequence in the northern foreland of the Alps, which is about coeval with the Upper Badenian.
70
JÓZSEF KÓKAY
Paracoryna praeambula (FLACH), 1891
Textfigure 5
1891 Coryna praeambula, FLACH: p. 2. Pl. III Fig. 2
1894 Coryna praeambula, CLESSIN: p. 26. I Fig. 4
1923 Aghardia praeambula, WENZ F. C.: p. 1038
The last whorls of two specimens, with entire apertural margins, were found in the
Sarmatian of Várpalota. It has an acute supracolumellar tooth and an also acute parietal one,
as well as two palatal plates; these characteristic for the species.
The discoverer of the species found the types in the Miocene at the North Alpine foreland (Undorf), coeval with the Badenian.
Textfigure 5. Paracoryna
praeambula (FLACH), 1891
Familia: Valloniidae
Genus: Vallonia RISSO, 1826
Vallonia lepida (REUSS), 1852
1959 Vallonia subpulchella — BODA: p. 640, 739, Pl. 37 Fig. 1
1996 Vallonia lepida — GERBER: p. 88, Figs 3d, 29a–29g, 30a–30d, 31a–31f, 32a–32c, 33
According to GERBER (1996) the species of Boda from the Sarmatian of Várpalota belongs to V. lepida, and Gerber
also gives a synonym list of this species. It is known from the Oligocene to the end of the Miocene.
Genus: Acanthinula BECK, 1847
Acanthinula tuchoricensis (KLIKA), 1891
Plate XXV Figure 15
1891 Helix (Acanthinula) tuchoricensis — KLIKA: p. 42. Fig. 35
1923 Acanthinula tuchoricensis — WENZ F. C.: p. 978.
From the lower part of the freshwater Lower Miocene of the Somlóvásárhely drilling a part of the last whorl of this
form was found with the apertural margin, as well as a highly damaged specimen.The whorls are quite flat, the shell moderately wide and there is a thread-like axial ridge system. All this is characteristic for the genus and, indead, for Klika’s
species. It is characteristic for the Lower Miocene of the northern part of the Czech Republic. The species was also
reported from Opole in Silesia (Oppeln in German) by ANDRAE (1902) from layers coeval with the Lower Badenian.
Acanthinula trochulus (SANDBERGER), 1874
Plate XXVI Figure 1
1874 Pupa (Modicella) trochulus — SANDBERGER: p. 601, Pl. 29 Fig. 25
1923 Acanthinula trochulus — WENZ F. C.: p. 977
1976 Acanthinula trochulus — SCHLICKUM: p. 11, Pl. 2 Fig. 30
1981 Acanthinula trochulus — LUEGER: p. 34, Pl. 3 Fig. 14
This species has been recognised in the Lower Miocene of the Somlóvásárhely drilling as well as from the Upper
Badenian coal-seam in the Pusztamiske Pm–4 well and from the Várpalota Sarmatian. It has quite a deep suture and
strongly convex whorls with a blunt midline ridge; furthermore a tube-like hollowed base is also characteristic. The fine
axial ribs are just about visible on the last whorl. As with SCHLICKUM’s (1976) figure the shell surface is essentially
smooth. The ribs are better developed on the younger Várpalota specimens.
Schlickum described it from the “silvana” beds, while Lueger mentioned it from Pannonian and Pontian beds.
Genus: Spermodea WESTERLUND, 1902
Spermodea candida FALKNER, 1974 var.
Plate XXVI Figure 2
1974 Spermodea candida — FALKNER: p. 231, Pl. 10 Figs 1, 2, Pl. 11 Figs 5, 6
Two complete and two damaged specimens were found in the Várpalota Sarmatian (well Inota–135). These are smallsized, (width — 1.4–1.45 mm; height — 1.3–1.35 mm). The form agrees well with Falkner’s form. The Hungarian spec-
Systematic descriptions
71
imens deviate only from the type form in being relatively lower — shorter in height and their aperture is more rounded
at its lower part. The size and density of ribs are similar. The initial whorls have a granulated surface.
Falkner found the species in the Undorf open air coal mine near Regensburg, Germany. These layers are approximately coeval with the Upper Badenian.
Genus: Planogyra MORSE, 1864
Planogyra nana (A. BRAUN), 1843
Plate XXVI Figures 3–4
1874 Planogyra nana — SANDBERGER: p. 374 Pl. 22 Fig. 14
1891 Helix (Acanthinula) nana — KLIKA: p. 41 Fig. 34
1923 Acanthinula nana — WENZ F. C.: p. 972
1974 Planogyra nana — FALKNER: p. 238 Pl. 10 Fig. 4, Pl. 11 Figs 9–10
One well-preserved specimen has been recognised in the Lower Miocene of the Somlóvásárhely drilling. It has been
reported from the northern Alpine Foreland from Upper Oligocene layers to levels coeval with the Upper Badenian, as
well as from the Lower Miocene of the Czech Republic. In the Sarmatian of Várpalota a damaged specimen has been
found which has an umbilicus slightly narrower than those of the older ones.
Genus: Strobilops Pilsbry, 1893
Strobilops fischeri WENZ, 1915
Plate XXVI Figures 5–6
1891 Strobilus diptyx — KLIKA: p. 34 Fig. 26
1915 Strobilops (Strobilops) fischeri — WENZ: p. 78 Fig. 6, Pl. IV Fig. 5
1923 Strobilops (Strobilops) fischeri — WENZ F. C. p. 1047.
One complete specimen has been recognised in the Lower Miocene of the Somlóvásárhely drilling. It corresponds
well with the species of Wenz. It is known from the Lower Miocene of the northern Czech Republic.
Strobilops uniplicata plana (CLESSIN), 1885
Plate XXVI Figures 7–8
1885 Strobilus planus — CLESSIN: p. 80, Pl. 7 Fig. 8
1915 Strobilops (Strobilops) uniplicata depressa — WENZ: p. 77, Pl. 4 Fig. 10
1923 Strobilops uniplicata plana — WENZ F. C.: p. 1059
1976 Strobilops uniplicata plana — SCHLICKUM: p. 11, Pl. 2 Fig. 31
This form has a relatively low, steplike spire. The last whorl is widest at its upper quarter, and thus its cross-section
is trapezoidal. On the smooth base there is a moderate-sized umbilicus. On the upper part of the whorls there is a faint
ribbing. It was found in the Hegymagos drilling. It is known from the “silvana” beds.
Strobilops subconoidea (JOOSS), 1912
Plate XXVI Figures 9–10
1912 Strobilops subconoidea — JOOSS: p. 34, Pl. 2 Fig. 4
1915 Strobilops (Strobilops) subconoidea — WENZ: p. 81, Pl. 4 Fig. 3
1923 Strobilops (Strobilops) subconoidea — WENZ F. C.: p. 1055
Some specimens were found in the Sarmatian of Pusztamiske and Várpalota and one fragment of a last whorl from
the Lower Badenian coal-seam of Herend (well Bánd–2). These are characterised by:
— a smooth base with growth-lines,
— the side is similarly smooth, while on the upper part of the whorls there are radial ribs,
— where the ribs appear, there is a faint angle, resulting in a blunt keel on the last whorl,
— it has a moderately developed umbilicus with a narrow pit.
The literature does not mention the basal plates. On the Hungarian specimens there are three dental plates inside on
the bottom of the base; the outer plate is narrow and moderately high and it is the longest. The median plate is half as
long but it is the highest and strongest. The inner one is the shortest, lowest and weakest and is placed almost as a columellar plate.
72
JÓZSEF KÓKAY
On the Pusztamiske specimens the radial ribs are poorly developed but are densely packed. It is also found in the
Steinheim beds.
Strobilops costata (CLESSIN), 1885
Plate XXVI Figures 11–13
1885 Strobilus costata — CLESSIN: p. 79, Pl. 7 Fig. 10
1902 Strobilus costatus — ANDREAE: p. 11, 26, Fig. 6
1915 Strobilops (Strobilops) costata — WENZ: p. 79, Fig. 7, Pl. 9 Figs 15, 16
1923 Strobilops (Strobilops) costata — WENZ F. C.: p. 1041
1967 Strobilops (Strobilops) costata — SCHÜTT: p. 213, Fig. 15
This form occurs in the closing bed of the Upper Badenian freshwater sequence in the Pusztamiske Pm–2 well and in
the Sarmatian of Várpalota. It is moderately high, pagoda-shaped and the whorls follow each other in a step-like fashion.
There are two outstanding parietal teeth and the inner one is the smallest. The basis is perforated by a moderate sized hole.
There are faint growth-lines on it and these become slightly stronger towards the aperture. There is a well developed radial ribbing on the upper half of the whorls. The closest related form is Strobilops subconoidea, which deviates in having:
— a funnel-shaped umbilical hole,
— the shell is lower and the spire is less steplike,
— the base is slightly more convex and smoother,
— the radial ribs on the whorls are fainter and denser,
— on the last whorl the blunt keel is more marked.
This form occurs in the “Tortonian” of the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) and in the
Sarmatian of the Vienna Basin.
Strobilops costata bilamellata (CLESSIN), 1885
Plate XXVI Figures 14–15
1885 Strobilus bilamellatus — CLESSIN: p. 79, Pl. 7 Fig. 9
1923 Strobilops (Strobilops) costatus — WENZ F. C.: p. 1041 (partim!)
WENZ, in his monograph (1915) about the genus Strobilops, fused the species S. bilamellata with S. costata; however, there are significant differences between them, as it is clear from Clessin’s descriptions and figures. From the
Sarmatian of Várpalota an entire specimen was found and this agrees well with S. bilamellata. On the last whorl of S.
costata there is a more or less developed keel, the sutures are shallower and the umbilicus is narrow. S. bilamellata, on
the other hand, has convex whorls, and this particularly true of the last one. Consequently, the basis is more convex and
the aperture more circular. The sutures are deeper and the umbilicus is wider. Thus I cannot agree with the fusion made
by WENZ (1923); a subspecific separation, at least, is warranted.
CLESSIN described the species from Undorf, from strata which are more or less coeval with the Upper Badenian.
Strobilops sandbergeri (ŁOMNICKI), 1886
Plate XXVI Figures 16–18
1886 Strobilops Sandbergeri — ŁOMNICKI: p. 58, Pl. 1 Fig. 3
1923 Strobilops (Strobilops) sandbergeri — WENZ F. C.: p. 1054
This species has been described from strata in Southern Poland (“Galicia”), which are coeval with the Badenian. Its
width is slightly superior to its height. There is a moderately developed ribbing on the whorls and the base is smooth. It
has a very narrow umbilicus, which is not covered by the apertural margin. This latter is curved and semicircular. The
external parietal tooth is well-developed, while the inner one is faint.
Strobilops pappi SCHLICKUM, 1970
Plate XXVII Figures 1–3
1970a Strobilops (Strobilops) pappi — SCHLICKUM: p. 84, Figs 2, 3
1981 Strobilops (Strobilops) pappi — LUEGER: p. 36, Figs 3b, 3c, Pl. 4 Fig. 1
1985 Strobilops (Strobilops) pappi — LUEGER: p. 351, Pl. 45 Figs 4–6
2004 Strobilops (Strobilops) pappi — HARZHAUSER et BINDER: p. 124 Pl. 7 Figs 4–6
It is not rare in the Sarmatian of Várpalota, although most of the specimens are broken. There are well-developed ribs
on the whorls. On the last one there is a blunt keel. The base is almost smooth, but towards the aperture wrinkles appear.
The umbilicus is narrow. It is known from the Pannonian of the Vienna Basin.
Systematic descriptions
73
Strobilops tiarula (SANDBERGER), 1886
Plate XXVII Figures 4–5
1915 Strobilops (Strobilops) tiarula — WENZ: p. 81, Pl. 4 Fig. 13
1959 Strobilops tiarula — BODA: p. 640, 739, Pl. 36 Fig. 4
1981 Strobilops (Strobilops) tiarula — LUEGER: p. 35, Pl. 4 Figs 2, 3
1985 Strobilops (Strobilops) tiarula — LUEGER: p. 350, Pl. 45 Figs 1–3
The species was found in the Upper Badenian of Hegymagos, in the Sarmatian of Várpalota and — albeit not with
certainty — in the Pusztamiske coal-bearing series. It is a relatively high, pagoda-shaped shell without an umbilicus as
the navel is generally covered by the apertural margin. Well-developed growth-wrinkles and ribs are present even on the
base. There are two parietal teeth and the external one is the stronger. The apertural margin is moderately thickened. In
some specimens the growth-lines are less developed, and the base may almost be smooth. This points to the possibility
that the ancestor of this species might be S. elasmodonta (REUSS), 1860.
Strobilops sp.
Plate XXVII Figures 6–7
Some Strobilops shells from the Sarmatian of Várpalota (Shaft M. III, 112 m) could not be identified with any species
in the literature. This form is small-sized (max 1.5 mm in diameter), which may indicate specimens that it is not completely grown up. The last quarter of the whorl is missing and the summit is slightly compressed.
— Both the upper and lower sides of the shell are smooth and on the upper side faint growth-lines are visible.
— The last whorl is downturned forming a rounded keel at the upper quarter of its height.
— The umbilicus is extremely narrow.
— With respect to the parietal plates, the outer one is more developed.
The form closest to this is S. uniplicata plana (JOOSS, 1985). The Hungarian specimen deviates from this because
Jooss’ form has radial ribs on the surface of the whorls and its umbilicus is also bigger.
Familia: Enidae
Genus: Mastus BECK, 1837
Mastus pupa sarmatica PAPP, 1974
Plate XXVII Figure 8
1974 Mastus pupa sarmatica — PAPP: p. 386, Figs 66/1, 66/2
One specimen originates from the Sarmatian of Várpalota. The first four (apical) whorls have been preserved while
the fifth is an external mould. Its form and size identifies it with the form described from the Sarmatian of the Vienna
Basin.
Genus: Ena TURTON, 1831
Ena sp.
Plate XXVII Figure 9
1981 Ena sp., LUEGER: p. 37 Pl. 4 Fig. 8
A half — probably the last — of this form whorl has been recognised in the Lower Miocene of the Somlóvásárhely
drilling and it corresponds very well to Lueger’s specimen. The whorl is bent downwards at a sharp angle and the growth
lines are clearly visible. There is no possibility for a more precise determination. Lueger reports it from Pontian sediments. It seems to be the closest to the extant species Ena montana (DRAP.) (KERNEY et al. 1983)
Genus: Napaeus ALBERS, 1850
Napaeus complanatus (REUSS), 1852
Plate XXVII Figures 10, 11, 12
1852 Bulimus complanatus — REUSS: p. 11, 12, 29. Pl. III Fig. 4
1874 Bulimus (Petraeus) complanatus — SANDBERGER: p. 433 Pl. 24 Fig. 11
1891 Bulimus (Medea?) complanatus — KLIKA: p. 69 Fig. 64
1891 Bulimus (Petraeus) turgidulus — KLIKA: p. 69 Fig. 65
1923 Ena (Napaeus) complanatus — WENZ F. C.: p. 1074
74
JÓZSEF KÓKAY
Three broken specimens of this species have been recognised in the Lower Miocene of the Somlóvásárhely
drilling. In spite of their state these can be easily identified. The juvenile whorls are well-preserved and the apical
angle is identical in the specimens. The external margin of the aperture of the adult whorls are bent outward in a similar way to the figures of Reuss’ species. On the top of the last whorl there is a narrow depression just under the suture.
The growth lines are strongest below the suture. The species is characteristic for the Lower Miocene of the north
Czech Republic.
Familia: Succineidae
Genus: Succinea DRAPARNAUD, 1801
Succinea minima KLEIN, 1853
Plate XXVII Figure 13
1874 Succinea minima — SANDBERGER: p. 601, Pl. 29 Fig. 26
1892 Succinea minima — MAILLARD: p. 90, Pl. 6 Fig. 14
1923 Succinea (Amphibina) minima — WENZ F. C.: p. 893
1976 Succinea (Hydrotropa?) minima — SCHLICKUM: p. 11, Pl. 2 Fig. 34
This species has been recognised from the Sarmatian of Várpalota as well as in the Lower Miocene of the
Somlóvásárhely drilling. It agrees best with Maillard’s figure, but it is wider and has more convex whorls. More elongate specimens were also found and these are similar to that of Sandberger. The Lower Badenian coal-measure of Herend
also yielded a fragment. It has also been found in the North Alpine “silvana” beds and in the Steinheim beds.
Familia: Endodontidae
Genus: Helicodiscus MORSE, 1864
Subgenus: H e b e t o d i s c u s H. B. BAKER, 1929
Helicodiscus (Hebetodiscus) subteres (CLESSIN), 1885
Plate XXVII Figures 14–16
1885 Patula subteres — CLESSIN: p. 76, Pl. 7 Fig. 6
1923 Pyramidula subteres — WENZ F. C.: p. 1062
1976 Helicodiscus (Hebetodiscus) subteres — SCHLICKUM: p. 11, Pl. 2 Fig. 35
1979b Helicodiscus (Hebetodiscus) subteres — SCHLICKUM: p. 68, Fig. 2
A subjuvenile and an adult specimen were found in the Hegymagos well, agreeing with the type form. It is known
from the “silvana” beds of the Bavarian Molasse and is coeval with the Hungarian occurrence.
Genus: Discus FITZINGER, 1833
Discus euglyphoides (SANDBERGER), 1874
Plate XXVIII Figures 1–2
1874 Patula (Charopa) euglyphoides — SANDBERGER: p. 583 Pl. 31 Fig. 1
1892 Patula euglyphoides — MAILLARD: p. 16 Pl. I Fig. 19
1911 Patula (Charopa) euglyphoides — GAÁL: p. 58 Pl. III Fig. 6
1923 Goniodiscus (Goniodiscus) euglyphoides euglyphoides — WENZ F. C.: p. 329
1976 Discus (Discus) euglyphoides — SCHLICKUM: p. 12 Pl. 2 Fig. 36
An uncommonly sizable specimen, about 7.5 mm in diameter, has been found in the Lower Miocene of the
Somlóvásárhely drilling. Its conical spire is broken and compressed, but otherwise the specimen is quite well-preserved.
On the upper margin of the whorls there is a rounded keel. On the last whorl 80 well-developed transversal ridges can be
seen. These ridges are more and more densely packed toward the umbilicus. The umbilicus is not especially wide.
Specimens of D. pleuradrus in the same assemblage are about half as big as this one and their whorls are convex. The
number of their transversal ridges is about 70 and the umbilicus is wider. The range of D. euglyphoides is from the
Ottnangian to the Sarmatian in the North Alpine Molasse region and in the “silvana” beds.
Systematic descriptions
75
Discus pleuradrus (BOURGUIGNAT), 1881
Plate XXVIII Figures 3–4
1881 Helix pleuradra — BOURGUIGNAT: p. 53, Pl. 3 Figs 67–72
1881 Helix dasypleura — BOURGUIGNAT: p. 55, Pl. 3 Figs 73–77
1892 Patula (Janulus) supracostata — MAILLARD: p. 17, Pl. 1 Fig. 20
1918 Piramidula (Gonyodiscus) silvana — JOOSS: p. 290
1923 Gonyodiscus (Gonyodiscus) pleuradra pleuradra — WENZ F. C.: p. 341
1959 Goniodiscus costatus — BODA: p. 642, 740, Pl. 36 Figs 5, 6
1976 Discus (Discus) pleuradrus — SCHLICKUM: p. 12, Pl. 2 Fig. 37
1981 Discus (Discus) pleuradrus — LUEGER: p. 40, Pl. 4 Figs 6, 7
This species has been recognised in the Lower Miocene of the Somlóvásárhely drilling as well as from the Upper
Badenian of the Herend and Tapolca Basins, from the base of the Várpalota coal-seam, and from the Upper Badenian of
the Fehérvárcsurgó well. It is most frequent in the Sarmatian of Várpalota.
The lower and upper sides of the initial whorls are smooth and the transversal ribbing appears only later, being
expressed on the base. The ribs generally weaken abruptly towards the base, and thin around the umbilicus they become
slightly stronger again.
This form is widespread in the Middle and Upper Miocene of Central and Western Europe.
Familia: Vitrinidae
Genus: Semilimax AGASSIZ, 1845
Semilimax intermedia (REUSS), 1852
Plate XXVIII Figure 5
1852 Vitrina intermedia — REUSS: p. 18, Pl. 1 Fig. 4
1891 Vitrina intermedia — KLIKA: p. 23, Fig. 15/1
1923 Vitrina intermedia intermedia — WENZ F. C.: p. 216
1981 Semilimax intermedius — LUEGER: p. 41, Pl. 5 Figs 1–3
One specimen was found in the Upper Badenian of the Herend Basin and also in upper part of the Várpalota
Sarmatian. The shell surface is smooth and there are no major growth-lines visible on it, in contrast to REUSS’ (1852) figure. Lueger’s figure also depicts a smooth specimen. It is known from almost the entire area covered by the MiddleEuropean Miocene.
Semilimax intermedia crassitesta (ANDREAE), 1902
Plate XXVIII Figures 6–7
1891 Vitrina intermedia — KLIKA: Pl. 24 Figs 5/2a, 15/2b
1902 Semilimax intermedia var. crassitesta — ANDREAE: No 18: p. 26
1923 Semilimax intermedia crassitesta — WENZ F. C.: p. 217
At the base of the Pusztamiske and Nyirád Sarmatian one entire and three damaged specimens were collected. This
form deviates from the nominal type due to its higher shell and thicker wall. It is known from the Lower Miocene of the
Czech Republic and from the nonmarine beds at Opole (Southern Poland).
Familia: Zonitidae
Genus: Vitrea FITZINGER, 1836
Vitrea procrystallina (ANDREAE), 1902
Plate XXVIII Figures 8–9
1902 Hyalina (Vitrea) procrystallina — ANDREAE: No. 18: p. 10, Fig. 4
1923 Vitrea procrystallina procrystallina — WENZ F. C.: p. 293
1976 Vitrea procrystallina — SCHLICKUM: p. 12, Pl. 2 Fig. 38
1981 Vitrea procrystallina procrystallina — LUEGER: p. 43
Two juvenile and one subjuvenile specimens were found in the Upper Badenian of the Tapolca Basin and one juvenile specimen in fhe Middle Badenian of the Várpalota Basin. The upper part of the last whorl is angulate. In cross-section is trapezoidal.
76
JÓZSEF KÓKAY
The form is known from the North Alpine Molasse Zone, from the “silvana” beds, and from Opole (S Poland). These
layers are coeval with the Badenian.
Vitrea procrystallina steinheimensis GOTTSCHICK, 1920
Plate XXVIII Figures 10–12
1920 Vitrea procrystallina steinheimensis — GOTTSCHICK: p. 37
1923 Vitrea procrystallina steinheimensis — WENZ F. C.: p. 294
1981 Vitrea (Vitrea) procrystallina steinheimensis — LUEGER: p. 42, Pl. 3 Figs 4, 7
Specimens from the upper part of the Várpalota Sarmatian agree well with this form. It is bigger than the nominal
subspecies. It has been found in the Steinheim beds. Recognised in the Pannonian and Pontian of the Vienna Basin.
Genus: Nesovitrea C. M. COOKE, 1921
Subgenus: Pe r p o l i t a BAKER, 1928
Nesovitrea (Perpolita) boettgeriana (CLESSIN), 1885
Plate XXVIII Figures 13–14, 15
1885 Hyalina Boettgeri — CLESSIN: p. 75, Pl. VII Fig. 7
1923 Zonitoides (Zonitoides) boettgerianus — WENZ F. C.: p. 296
1975 Perpolita boettgeriana — SCHLICKUM: p. 40, Pl. 3 Fig. 3
1999 Nesovitrea boettgeriana — KÓKAY: Fig. 40
Several, mostly subjuvenile examples of this species were found in the Lower Miocene of the Somlóvásárhely drilling.
Their shells are slightly smoother than the shell of Clessin’s species. It has been found in layers coeval with the Upper
Badenian in the Northern Alpine Foreland, at Undorf. In the Upper Badenian of Mátraszőlős one specimen was found.
Nesovitrea (Perpolita) mendica SLAVIK, 1869
Plate XXIX Figures 1–5
1869 Helix (Hyalina) mendica — SLAVIK: p. 262, Pl. 4 Figs 7, 8
1891 Hyalina mendica — KLIKA: p. 31, Fig. 23
1902 Hyalina mendica — ANDREAE: p. 8
1923 Zonitoides (Hyalina) mendicus — WENZ F. C.: p. 296
Some damaged specimens were collected from the upper part of the Várpalota Sarmatian. KLIKA (1891) gives 3.5 mm
as a maximum diameter. The biggest, damaged Hungarian specimen is 2.3 mm in diameter. Otherwise the agreement is
good. Damaged specimens from the Herend Upper Badenian are also similar to these.
Nesovitrea (Perpolita) subhammonis GOTTSCHICK, 1928
Plate XXIX Figures 6–9
1928 Zonitoides subhammonis — GOTTSCHICK: p. 146, Pl. 2 Fig. 6
1975 Perpolita subhammonis — SCHLICKUM et STRAUCH: p. 40, Pl. 3 Fig. 4
Several damaged juvenile specimens were collected from the Hegymagos and Pusztamiske wells. The species has
been described from sediments comparable to the sequence of the mentioned drillings, in the Northern Alps.
Genus: Aegopis FITZINGER, 1833
Aegopis algiroides (REUSS), 1852
Plate XXIX Figure 10
1852 Helix algiroides — REUSS: p. 11, 19 Pl. I Fig. 5
1852 Helix Haidingeri — REUSS: p. 11, 20 Pl. I Fig. 6
1852 Helix semiplana — REUSS: p. 20 Pl. I Fig. 7–8
1874 Archeozonites Haidingeri — SANDBERGER: p. 404, 443 Pl. 24 Fig. 26
1874 Archeozonites semiplanus — SANDBERGER: p. 442 Pl. 24 Fig. 25
1891 Archeozonites haidingeri — KLIKA: p. 25 Fig. 17
1891 Archeozonites haidingeri var. reussi — KLIKA: p. 27 Fig. 18
1892 Archeozonites subverticillus — MAILLARD: p. 11 Pl. I Fig. 15
1923 Aegopis algiroides — WENZ F. C.: p. 249
A small fragment of a whorl has been found in the Lower Miocene of the Somlóvásárhely drilling. On its upper side
there are well-developed growth lines. The lower part of the whorl is covered by faint growth lines only. Its lower mar-
Systematic descriptions
77
gin forms a sharp angle with the next whorl. This characterises the juvenile specimens which were described by Reuss
as “semiplanus”. Juvenile specimens of A. costatus (SANDB.) are similar to A. algiroides, but the ribs are rounded.
Its known range extends from the Upper Oligocene to the Burdigalian (North Czech Republik)
Aegopis costatus (SANDBERGER), 1874
Plate XXIX Figures 11, 12
1874 Archeozonites costatus — SANDBERGER: p. 604
1892 Archeozonites subcostatus — MAILLARD: p. 13, Pl. 1 Fig. 16
1916 Zonites (Aegopis) costatus — GOTTSCHICK et WENZ: p. 21, Pl. 1 Fig. 1
1923 Zonites (Aegopis) costatus — WENZ F. C.: p. 254
1976 Archeozonites costatus — SCHLICKUM: p. 18, Pl. 5 Fig. 66
1981 Archeozonites costatus — LUEGER: p. 44
SANDBERGER (1874) described the species A. costatus — mentioning A. subcostatus as well — from the North Alpine
Molasse Zone. The first-mentioned form does not have a keel, and is adorned with flat ribs and wrinkles, especially on
the base. The second one is close to the first, having an extremely faint spiral keel and dense subtle riblets. BAUMBERGER
(1927) regarded the second one as a subspecies of the first, while WENZ (1923) fused them as synonyms, under the name
A. costatus. GOTTSCHICK and WENZ (1916) suggested that A. subcostatus specimens are adults of the other species. A half
whorl was found in the upper part of the Upper Badenian of Pusztamiske (well Pm–2). This has a deep and well-developed umbilicus, while in the well Pmt–3 an apical fragment was found. The wrinkles are poorly developed, as is visible
on the figure of SCHLICKUM (1976). Some fragments were recorded from the Hegymagos well, from the Lower Badenian
of Herend (drilling Bánd–2) and from the Várpalota Sarmatian.
SCHLICKUM (1976) mentions the form as being present in the “silvana” beds.
Genus: Retinella FISCHER, 1877
Retinella applanata nov. sp.
Plate XXIX Figure 13–15
Derivatio nominis: applanata (Lat.) = flattened
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 110
Locus typicus: Várpalota Basin, drilling I–135, 111.6–124.6 m
Stratum typicum: Sarmatian freshwater clay
Descriptio: the flattened, planispiral shell is built of four whorls. The sutures are slightly deepened and the whorls
are moderately convex. The last whorl encircles the whole base and the umbilicus is medium-sized. On the shell (under
a higher magnification) a densely-packed, regularly placed radial rib system is visible. The side of the whorl is smooth.
On the base there are slight growth-lines and hardly visible spiral lineae. The aperture is elongate and oval. Its diameter
is 2.5 mm and its height 0.8 mm.
Diagnosis differentialis: gives its characteristics, the form may only be placed in the genus Retinella. It differs from
other members of the genus due to its small size. The question emerges: is the specimen an adult? Even if it is not, the
planispiral whorls characterise the form accurately. In this respect, it stands quite close to the small-sized subgenus
“Gyralina”, with its type species Retinella (Gyralina) circumlineata (KÜSTER). The Hungarian form differs from this,
being even more flattened and having fainter spiral decoration. It is possible to range it in a new subgenus. However, giving that only one specimen is known, it would be risky to propose one.
Genus: Aegopinella LINDHOLM, 1926
Aegopinella denudata (REUSS), 1852 var.
Plate XXX Figures 1–2
1852 Helix denudata — REUSS: p. 11, 21 Pl. I Fig. 9
1874 Hyalina denudata — SANDBERGER: p. 441 Pl. 24 Fig. 23
1891 Hyalina denudata — KLIKA: p. 28 Fig. 19
1914 Hyalina (Hyalina) denudata — FISCHER et WENZ: p. 45 Pl. II Fig. 3
1923 Oxychilus (? Oxychilus) denudatum denudatum — WENZ F. C.: p. 29
2004 Aegopinella denudata — BINDER: p. 197 Pl. 3 Fig. 4
A quite well-preserved specimen has been recognised in the Lower Miocene of the Somlóvásárhely drilling. Its flat,
plate-like shell, with a diameter of 13 mm, is built of four and a half whorls. The shell is covered with small growth folds,
visible even with bare eyes. At the base the shell is incomplete, having a deep and moderately narrow umbilicus. It dif-
78
JÓZSEF KÓKAY
fers from Reuss’ species in having one less whorl and by its slightly narrower umbilicus. According to Klika, the species
is 12 to 14 mm wide. If the Hungarian specimen was even wider, it could even have five whorls. This might be regarded
as an unimportant deviation from Reuss’species. Therefore the difference is indicated with the word “var.”
The species is a characteristic member of the Lower Miocene of the North Alpine Foreland and the Czech Republic.
BINDER (2004) mentioned its presence the Lower Miocene of Austria (Styria).
Genus: Oxychilus FRITZINGER, 1833
Oxychilus procellarius (JOOSS), 1918
1923 Oxychilus (Oxychilus) procellarius — WENZ F. C.: p. 279
1959 Oxychilus procellarius — BARTHA: Pl. 17 Figs 8–10
1981 Oxychilus (Oxychilus) procellarius — LUEGER: p. 46, Pl. 6 Fig. 2
This compressed specimen has been recognised in the Lower Miocene of the Somlóvásárhely drilling as well as from
the Sarmatian of Várpalota (Shaft S. III and drilling Inota–51). It seems that sometimes there are less whorls on these
than on Jooss’ species. The shell surface is shiny, with faint growth-lines.
It is present in the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) from the “silvana” beds up to the
end of the Miocene, as well as from the Pannonian and Pontian of the Vienna Basin and Transdanubia (W Hungary).
Genus: Daudebardia HARTMANN, 1821
Daudebardia praecursor ANDREAE, 1902
Plate XXX Figures 3, 4
1902 Daudebardia praecursor — ANDREAE: p. 2, Fig. 1
1923 Daudebardia (Daudebardia) praecursor — WENZ F. C.: p. 307
This form has been found in the lower and in the upper part of the Sarmatian of Várpalota. One of specimens is almost
entire. ANDREAE (1902) mentioned its occurence Opole (S Poland, Silesia) from layers coeval with the Badenian.
Genus: Zonitoides LEHMANN, 1862
Zonitoides schraieri SCHLICKUM, 1978 var.
Plate XXX Figures 5, 6
1978 Zonitoides (Zonitoides) schraieri — SCHLICKUM: p. 255, Pl. 19 Fig. 15
1981 Zonitoides schraieri — LUEGER: p. 47, Pl. 5 Fig. 6
One specimen was collected from the Várpalota Sarmatian (Shaft M. III). It deviates from the species in having a
lower summit and the spiral keel is blunter. It is known from the Pontian of the Balaton region and Vienna Basin.
Zonitoides suevicus (JOOSS), 1918 var.
Plate XXX Figures 7–8, 9–11
1918 Polita suevica — JOOSS: p. 2 89
1923 Zonitoides (Zonitoides) suevicus — WENZ F. C.: p. 289
1976 Zonitoides (Zonitoides) suevicus — SCHLICKUM: p. 18, Pl. 5 Fig. 67
Two specimens were collected from the Upper Badenian of Herend. These deviate from the species in the following:
— they are smaller (2.1 mm in diameter, while for the type it is 5.2 mm);
— on the figure of Schlikum the heigtht represents 57% of the width. In the Herend specimens this ratio is 62%,
while in the Várpalota ones it is 60%. This variation may be within the range of the species, or it may be explained by
the fact that it represents a subjuvenile state. A deviation, not, exceeding a subspecific one, is thus stated. It is known
from the “silvana” beds.
Genus: Janulus LOWE, 1852
Janulus supracostatus (SANDBERGER), 1874
Plate XXX Figure 12
1923 Janulus supracostatus — WENZ F. C.: p. 304
1976 Janulus supracostatus — SCHLICKUM: p. 13, Pl. 3 Fig. 43
Systematic descriptions
79
Several fragmentary specimens were collected from the Sarmatian of Várpalota. The last whorl is missing from all
these specimens. The first whorls are smooth, the ribs appear gradually, and disappear at the base. The biggest, incomplete specimen is 6.5 mm in diameter with seven whorls. It has been found in the “silvana” beds.
Janulus moersingensis JOOSS, 1918
1918 Janulus mörsingensis — JOOSS: p. 289
1923 Janulus moersingensis — WENZ F. C.: p. 303
1976 Janulus moersingensis — SCHLICKUM: p. 14, Pl. 3 Fig. 44
A specimen was collected from the base of the Lower Badenian in the Nagygörbő well. The last whorl is incomplete.
Its base is smooth and hollowed. The initial whorls are smooth but later on the transversal ribs are faint and dense (the
Hungarian specimen ends here). Finally they become stronger and farther apart from each other. From the Upper
Badenian of the Herend Basin and the Hegymagos well several fragmentary specimens were collected.
Its occurence has also been reported from the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) “silvana” beds.
Janulus? sp.
Plate XXX Figures 13–15
Several gastropod fragments were found in the Sarmatian of Várpalota and these are characterisd by:
— well-pronounced, curved ribs on the upper part of the whorls which disappear towards the base;
— the diameter of the shell may be estimated as 8–10 mm;
— a wide umbilicus;
— on the inner side of the whorls there are sporadically elongate teeth.
Consequently, the specimens most probably belong to the genus Janulus, taking into consideration the description of
WENZ, ZILCH (1959: p. 262). It is bigger than Janulus supracostatus and its radial ribs are more pronounced.
Familia: Limacidae
Genus: Limax LINNAEUS, 1758
Limax crassus CLESSIN, 1894
1911 Limax crassa — GAÁL: p. 62, Pl. 3 Fig. 16
1959 Limax crassus — BODA: p. 741, Pl. 36 Fig. 8
The shield (scutum) from the Upper Badenian of Pusztamiske and from the Sarmatian of Várpalota are resemble a
rounded brick-shaped form, with a notch on the upper margin. The concentric growth-lines are situated around the apex,
which is situated at the right upper angle. The lines follow this brick-like outline. The lower side is concave.
It is present in the Sarmatian of Rákosd, Transylvania (Rumania), and in coeval beds of the North Alpine Freshwater
Molasse (“Süssbrackwassermolasse”).
Limax sp.
Several scutum have been recognised in the Lower Miocene of the Somlóvásárhely drilling and these are typical for
the genus. It has no stratigraphic significance.
Familia: Milacidae
Genus: Milax GRAY, 1855
Milax lorentheyi (GAÁL), 1910
1911 Milax Lőrentheyi — GAÁL: p. 61, Pl. 3 Fig. 18
1923 Milax lorentheyi — WENZ F. C.: p. 315
1959 Milax lorentheyi — BODA: p. 642, 741, Pl. 36 Figs 9, 10
This shield (scutum) occurs in the Sarmatian of Várpalota. It is known exclusively from Sarmatian deposits.
80
JÓZSEF KÓKAY
Familia: Arionidae
Genus: Arion FÉRUSSAC, 1819
Arion sp.
With some reserve, I range here some scutum of slugs which are somewhat smaller and more oval than the previously mentioned ones. They are thin with a watch-glass shape on one side, and slightly convex on the other. These were
found in the Lower Miocene of the Somlóvásárhely drilling. The genus has no stratigraphic significance.
Familia: Subulinidae
Genus: Opeas ALBERS, 1850
Opeas minutum (KLEIN), 1853
Plate XXX Figure 16
1853 Bulumus minutus — KLEIN: p. 212, Pl. 5 Fig. 9
1874 Subulina minuta — SANDBERGER: p. 596, Pl. 29 Fig. 16
1886 Subulina minuta — ŁOMNICKI: p. 66, Pl. 1 Fig. 16
1891 Stenogyra (Opeas) minuta var. reunensis — PENECKE: p. 365, Pl. 21 Fig. 9
1892 Stenogyra (Subulina) minuta — MAILLARD: p. 87, Pl. 6 Fig. 11
1925 Opeas minutum — WENZ F. C.: p. 872
1966 Micromelania? sp. — KÓKAY: p. 35, Pl. 2 Fig. 14
1976 Opeas minutum — SCHLICKUM: p. 14, Pl. 3 Fig. 46
One well-preserved shell was collected from the Upper Badenian of Herend and it is 5.3 mm high. It is moderately
convex with relatively densely-packed whorls, (5.5 in total), and the apex is blunt. There are densely-packed, faint
growth-wrinkles on the surface. The aperture has a curved, sharp external margin, which forms a point with the columellar side. This last mentioned side has a rolled shape and behind it there is a narrow pit on the base. All this facts point to
the genus Opeas.
It is also known from the younger “silvana” beds (about coeval with the Badenian). Extant species prefer warm climates.
Genus: Fortuna SCHLICKUM et STRAUCH, 1979
Fortuna tertia SCHLICKUM et STRAUCH, 1979
Plate XXX Figures 17, 18, 19; Textfigure 6a
1972 Fortuna sp.— SCLICKUM et STRAUCH: p. 73 Abb. 5
1979 Fortuna tertia — SCHLICKUM et STRAUCH: p. 53, Pl. 5 Fig. 70
This is a small terrestrial gastropod. Numerous specimens were found in the upper part of the Sarmatian at Várpalota.
It agrees well with the named species. It is also present in the Pontian of Burgenland (Austria). The genus occurs in the
Upper Miocene and Pliocene.
Fortuna varpalotensis nov. sp.
Plate XXX Figure 20; Textfigure 6b
Derivatio nominis: from the type locality Várpalota
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 111
Locus typicus: A Várpalota Basin, drilling I–135, 111.6–124.6 m
Stratum typicum: Sarmatian freshwater clay
Descriptio: this is a conical shell which widening strongly with four whorls. The
apex is rounded. The whorls are moderately convex and the suture is also not very
deep.The last whorl dominates the shell, being as high as three-quarters of the total
height, pointing to a fast pace of growth. Its size is: height — 3.3 mm; width — 1.7
mm The height of the last whorl 2.4 mm and the height of the aperture 1.8 mm. The
conic angle is 44 degrees. The aperture is elongate and is pointed at the top and bottom ends. The external aperture is sharp and on the columellar side it is twisted (as
in other species of the genus). At the columellar margin there is a slight umbilical
fissure. The shell is smooth and under a microscope some slight growthlines are visible. The base is convex.
Textfigure 6. a) Fortuna tertia
SCHLICKUM et STRAUCH, 1979; b)
Fortuna varpalotensis nov. sp.
Systematic descriptions
81
Diagnosis differentialis: one complete specimen and two apical parts have been found. In the Lower Miocene of the
Somlóvásárhely drilling an entire juvenile specimen was collected: it had two and a half whorls and an aperture similar
to the described form. The closest known species is Fortuna tertia, which occurs in the Sarmatian of Várpalota . However,
it is easy to distinguish them from one another. F. tertia is more slender and its conic angle is only 33 degrees . The last
whorl is much lower, forming just 60 percent of the total height. Its aperture is also not as high.
The base and the last whorl form an obtuse angle. Other species of the genus are even more unlike the new taxon.
Familia: Clausiliidae
Genus: Triptychia SANDBERGER, 1874
Triptychia vulgata (REUSS), 1852
Plate XXXI Figure 1
1852 Clausilia vulgata — REUSS: p. 34, Pl. 4 Fig. 1
1860 Clausilia vulgata — REUSS: p. 74, Pl. 2 Fig. 1
1891 Triptychia (Plioptychia) vulgata — KLIKA: 76, Fig. 73
A fragmentary specimen with four whorls was collected from the freshwater beds at the base of the Ottnangian in
Várpalota. The density of the ribs and the low whorls agree well with the type from the Czech Republic Lower
Miocene.
Triptychia grandis (KLEIN), 1847
Plate XXXI Figures 2, 3, 4
1847 Clausilia grandis — KLEIN: p. 73 Pl. I Fig. l6
1867 Clausilia grandis — QENSTEDT: p. 485, Pl. XLI. Fig. 38
1877 Clausilia (Triptychia) grandis — BOETTGER: p. 20, Pl. I Figs 11–14
1885 Clausilia grandis — QENSTEDT: p. 622, Pl. 48 Fig. 70
1892 Clausilia (Triptychia) grandis — MAILLARD: p. 83, Pl. VI Figs 5–7
1923 Triptychia (Triptychia) grandis — WENZ F. C.: p. 808
1976 Triptychia (Triptychia) grandis — SCHLICKUM: p. 14, Pl. 3 Fig. 47
Several fragments and apical parts were found in the Lower Miocene of the Somlóvásárhely drilling, including the
lower half of a compressed specimen with its aperture. The side of the whorls is straight. There is an array of dense,
minute and sharp axial ribs which are quite low. The ribs are triangular in cross-section and generally quite uneven. In
some cases they are branching. In adult specimens the ribs are slightly opystocyrtal. The interrib spaces are as wide as
the ribs. On the adult whorls there are narrow fissures running parallel with the sutures. Their number may be as high
as 8 or 10. The total height can be up to 30–32 mm and the, width about 9 mm. The last whorl is 11 mm high. Boettger
mentions a total height of 30 to 35 mm, but other papers mention taller specimens as well. Its stratigraphic range
extends from the Upper Brackish Molasse (Maillard) to the end of the Badenian or — albeit with some doubt — to the
Sarmatian.
Triptychia cf. suturalis (SANDBERGER), 1874
Plate XXXI Figures 5–6
1847 Clausilia antiqua — KLEIN: p. 74 Pl. 1 Fig. 17
1874 Clausilia (Triptychia) suturalis — SANDBERGER: p. 652 Pl. 28 Fig. 11
1877 Clausilia (Triptychia) suturalis — BOETTGER: p. 20
1885 Clausilia antiqua — QENSTEDT: p. 622 Pl. 48 Fig. 69
1892 Clausilia (Triptychia) suturalis — MAILLARD: p. 85 Pl. 6 Fig. 8
1923 Triptychia suturalis — WENZ F. C.: p. 819
In the lowest Sarmatian of Pusztamiske an almost entire and a broken last whorl of this form were recognised, while
some fragments were found in the Sarmatian of Nyirád. The last whorl is about 13 mm high. This value, if extrapolated, indicates a shell with a height of about 32 mm. A dense array of (but less dense than in T. grandis) sharp and steep
sided axial ribs can be seen on the surface. The ribs are undulating and their upper part (below the suture) slightly
prosocyrtal. The width of the ribs is about the same as that of the interrib space. There are one or two narrow furrows
on the upper part of the last whorl. Based on these traits, the specimens most probably belong to T. suturalis.The latter
has also been recorded from sediments in the North Alpine Foreland, which are coeval with the Upper Badenian and
the Sarmatian
82
JÓZSEF KÓKAY
Triptychia leobersdorfensis sarmatica nov. ssp.
Plate XXXI Figures 7–8, 9; Plate XLI Figures 6, 7
1959 Triptychia cfr. suturalis, BODA: p. 740 Pl. 36 Figs 2–3
Derivatio nominis: sarmatica (Lat.) = sarmatian
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 112
Locus typicus: Várpalota, drilling Csór–17, 128 m
Stratum typicum: Upper Sarmatian freshwater clay
Descriptio: this is a slender, towered shell. The apical part is missing but the last five whorls have been preserved. It
is slightly compressed but otherwise well preserved. The whorls are slightly convex. Their height is less than their width.
On the surface there is a dense array of tiny parallel and axial ribs.On the last whorls these ribs are slightly prosocyrtous.
On the upper part of the whorls there are four to six, hardly visible, spiral furrows forming a reticulum with the axial ribs.
The aperture is pyriform, elongate. At the base there is a faint rib parallel with the margin. On the columellar side there
is a well developed upper plate which does not reach the apertural margin, similar to the spiral and subcolumellar plates.
By extrapolating the proportions, the total height of the shell can be reconstructed as about 42 mm, its width is 10
mm and the height of the last whorl is 16 mm.
Diagnosis differentialis: the closest known relative might be the Pannonian T. leobersfensis (TROLL) (LUEGER l981)
recorded from the Vienna Basin and western Hungary. The new taxon is different in that it is slightly bigger (the
„Troll”species may be 35 to 40 mm high, according to Lueger) as well as by the spiral groove system on the whorls. The
upper plate is bigger and the tiny ribs are axially arranged in the new form. The apex is similar in the two species. In the
„Troll’’ form the axial ribs are denser (10 to 15 ribs on 2 mm) but less dense (6 to 7) in the new subspecies (I am indebted to Dr. Th. Schnabel for the kind informations).
Genus: Serrulastra NORDSIECK, 1981
Serrulastra ex gr. ptycholarynx (BOETTGER), 1877
Textfigure 7
1877 Clausilia (Serrulina) ptycholarynx — BOETTGER: p. 74, Pl. 2 Fig. 27
1923 Serrulina ptycholarynx ptycholarynx — WENZ F. C.: p. 780
2002 Serrulastra ptycholarinx — BINDER: p. 171, Taf. 3 Fig. 6; Taf. 9 Figs 3–4
Fragments of this form were found in the upper part of the Pusztamiske Upper
Badenian (well Pm–2) and from the Sarmatian of Nyirád. Their size, the density of the
narrow ribs and, especially, the dentation of the apertural margin indicate the vicinity of
this species. It has been described from the Vienna Basin Sarmatian, and from the
Karpatian of the Korneuburger Basin.
Textfigure 7. Serrulastra ex gr.
ptycholarynx (BOETTGER), 1877
Genus: Nordsieckia TRUĆ, 1972
Nordsieckia pontica LUEGER, 1978
Plate XXXI Figures 10, 11
1981 Nordsieckia fischeri pontica — LUEGER: p. 92, Pl. 7 Figs 7–12
1981 Nordsieckia pontica — NORDSIECK: p. 81, Pl. 9 Figs 32, 33
1996 Nordsieckia pontica — FORDINÁL: p. 10, Pl. 2 Fig. 9
Numerous specimens of this small species were found in the Sarmatian of Várpalota. Unfortunately, there is no entire
example but the older figures also show damaged specimens. It has a smooth, bobbin-shaped shell. The last whorl is
slightly coiled out. On the last quarter of the last whorl there are axial ribs which grow stronger towards the aperture. The
aperture is simple pear shaped from outside, slightly swelling out towards the columellar side.It has a well developed sinule and an upper plate.
It has also been found in the Pannonian and Pontian of the Vienna Basin and southern Slovakia.
Genus: Cochlodina FÉRUSSAC, 1821
Subgenus: M i o p h a e d u s a
Cochlodina (Miophaedusa) varpalotensis nov. sp.
Plate XXXI Figures 12–13, 14–19
Derivatio nominis: alluding to Várpalota, the type locality
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 113
Systematic descriptions
83
Locus typicus: well Inota–135, 111.6–124.6 m
Stratum typicum: freshwater clays with brackish intercalations of the upper part of the Sarmatian
Descriptio: the dimensions of holotype are: it has a last whorl with a height of 4 mm; the height of the aperture is
2.8 mm; the width of the whorl is 2 mm.
The last whorl has strong growth-lines. At its end there are strong wrinkles and ribs. Before the aperture there is a
pillow-like swelling and a poorly-developed occiput. The aperture is pear-shaped and its rim is bent outwards. The upper
plate is well-developed. The lower one is also quite strong, but does not reach the margin; instead bifurcates and its lower
branch reaches the margin. Below it there is a subcolumellar plate which also reaches the margin. Dorsally there is the
beginning of the principal wrinkle which does not run out to the palatal margin.
Below this there is a lunelle situated transversally, making an angle of 20–30° with the principal wrinkle. There is no
pharyngal wrinkle or interlamelle. The end of the twisted clausilium plate forms a bay between the two curved peaks, so
it is kidney-shaped.
Diagnosis differentialis: seven last whorls were found within a palaeopopulation. The apertural margins are generally damaged but the columellar margins are entire. They agree well with the holotype. In some specimens a second, poorly-developed bifurcating branch of the lower plate may also reach the columellar plate. In some specimens there is a
pharyngal wrinkle between the principal one and the suture, close to the palatal margin.
The new species is closest to C. oppolensis (NORDSIECK 1981), but deviates because a pharyngal cover and the clausilium is cut out deeper in a symmetric way. The deep embayment of the clausilium is characteristic for the subfamily
Alopiinae and, especially, for the genus Cochlodina.
Genus: Canalicia BOETTGER, 1877
Canalicia cf. attracta (BOETTGER), 1870
Textfigure 8
1870 Clausilia attracta — BOETTGER: p. 294 Pl. XIII Fig. 6
1877 Clausilia (Canalicia) attracta — BOETTGER: p. 83
1891 Clausilia (Canalicia) attracta — KLIKA: p. 84 Fig. 81
1923 Canalicia attracta — WENZ F. C.: p. 784
1981 Canalicia attracta — NORDSIECK: p. 107
From the Lower Miocene of the Somlóvásárhely drilling an damaged last whorl
Textfigure 8. Canalicia cf. attracta
and some fragments have been found. The surface is covered with a dense array of
(BOETTGER), 1870
axial ribs which are most developed, dichotomic and bifurcating on the last whorl. At
the bottom, behind the columellar part of the aperture, there is a well-developed ridge. Behind it there is a depression.
The columellar apertural margin forms an angle with the upper margin. A bifurcating branch of the basal plate proceeds
onto the inner (right) apertural margin. The narrow and curved rhomboidical clausilium plate is visible at the upper part
of the whorl, which is characteristic for the species (KLIKA 1891, Figure 81e). Its end is bent upwards. Unfortunately, the
external and lower apertural margin is missing and a great part of the aperture is filled with matrix. All this seems to
point to Boettger’s species, which is also present in the coeval formations of the Czech Republic.
Genus: Pseudidyla BOETTGER, 1877
Subgenus: C a n a l i c i e l l a NORDSIECK, 1981
Pseudidyla (Canaliciella) boettgeri NORDSIECK, 1981
Plate XXXI Figure 20, Textfigure 9
1902 Clausilia (Canalicia) n. sp. — ANDREAE: No 18: p. 20
1981 Pseudidyla (Canaliciella) boettgeri — NORDSIECK: p. 73, Pl. 8 Figs 13, 14
One single last whorl has been found in the upper part of the Várpalota
Sarmatian (well Csór–17, 124.1–127.6 m). Its aperture is partly filled with
matrix. However, it is visible that its shell is covered with sharp ribs, similar to
that on Nordsieck’s figure. The sinule is also similarly narrow, the lower plate
reaches the columellar margin, a backward turning gutter is on the base and the
palatal side is rim-like and bent outwards. It has also been reported from Opole
(Oppeln in German), Poland, from freshwater marls coeval with the Upper
Badenian stage.
Textfigure 9. Pseudidyla (Canaliciella) boettgeri NORDSIECK, 1981
84
JÓZSEF KÓKAY
Genus: Pseudidylas str.?
Pseudidyla tricarinata nov. sp.
Plate XXXII Figures 1–2, 3, Textfigure 10
Derivatio nominis: having three rims = tricarinata (Lat.)
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary,
Budapest), Inv. No MÁFI, Gy. 114
Locus typicus: Várpalota Basin, well Csór–8, 128.8–133 m
Stratum typicum: Sarmatian freshwater clay
Descriptio: In this of the bobbin-shaped shell the two last whorls have been preserved. These are gently convex with sutures that are slightly depressed. The surface
is covered with sharp axial ribs, which are not quite evenly developed. Some are
Textfigure 10. Pseudidyla tricarinata
weaker, curved, fading out or branching. Their number on the penultimate whorl is nov. sp.
35. The interrib depressions are about three times wider than the ribs. The aperture is
slightly damaged at its lower margin. Its outline is pearshaped and the columellar side is bent outward and is curved. The
parietal side is depressed below the sinule, which is normally developed. The upper plate is perpendicular to the margin.
Close to it, a slight interlamella reaches the margin. Below this the strong lower plate may be followed deep into the pharynx. After a bifurcation it reaches the margin. Immediately below the lower plate there is a furrow in the columellar margin. The rim is prolonged towards the basally-situated canal deep into the pharynx. The canal forms a ridge or keel outside of the shell and crosses the ribs. Parallel to the canal an internal ridge is placed; externally this appears as a depression. Parallel to this the lower palatal side of the aperture is curved. This curve may be followed towards the pharynx.
Above it, in upper palatal situation, there is a round pharyngal tubercle, corresponding to an outside pit. Above this pit
the palatal side of the sinule is swelling out forming the third ridge. The name “tricarinata” alludes to these three ridges.
The total height of the holotype may be estimated as 10–12 mm.
Diagnosis differentialis: The new species, which has three ridge on the neck, deviates from other species of the genus
Pseudidyla. NORDSIECK (1981) introduced a new subgenus, Canaliciella, but the Hungarian form does not fit into this. It
could represent a new subgenus. The closest form is P. polyptyx (BOETTGER 1877) and its subspecies P. p. supragonyptyx
SCHÜTT (NORDSIECK 1981). However, with regard to the upper palatal swelling and the threefold ridge, there is considerable deviation from the new species.
Genus: Mactrogastra HARTMANN
Subgenus: M a c t r o g a s t r a
Mactrogastra (Mactrogastra) cf. voesendorfensis (PAPP et THENIUS), 1953
Plate XXXII Figure 4–5; Textfigure 11
1953 Pseudidyla vösendorfensis — PAPP et THENIUS: p. 22, Pl. 4 Fig. 8
1981 Mactrogastra (Mactrogastra) voesendorfensis — NORDSIECK: p. 80, Pl. 9 Figs 26–28
This form from the Sarmatian of Várpalota (well Bh–3, 269.1–270.2 m) has a long
neck, well-developed growth-ribs and a pear-shaped aperture. This aperture is filled
with matrix, and consequently the identity can only be based upon the external characters, the marginal teeth and plates.
It is also present in the older Pannonian and Sarmatian of the Vienna Basin.
Textfigure 11. Mactrogastra
(Mactrogastra) cf. voesendorfensis (PAPP et THENIUS), 1953
Genus: Clausilia DRAPARNAUD
Clausilia ex aff. rolfbrandti (SCHLICKUM), 1969
Textfigure 12
1969 Canalicia rolfbrandti — SCHLICKUM: p. 198 Fig. 6
1972 Clausilia rolfbrandti — NORDSIECK: p. 176 Figs 8–9; Pl. 10 Figs 16–18,
1975 Clausilia rolfbrandti — SCHLICKUM: p. 64, Tab. 6 Fig. 45
1981 Clausilia rolfbrandti — NORDSIECK: p. 100, 109
Some fragments were found in the Middle Badenian of Várpalota. These belong to
a small species which has relatively thick upper and lower columellar apertural margins
that are sharply bent. It is similar to Schlickum’s species. The lower plate and interlamella are poorly developed. The species is known from the Pliocene of Germany and
France.
Textfigure 12. Clausilia ex aff.
rolfbrandti (SCHLICKUM), 1969
85
Systematic descriptions
Familia: Oleacinidae
Genus: Pseudoleacina WENZ, 1914
Subgenus: P s e u d o l e a c i n a WENZ, 1914
Pseudoleacina (Pseudoleacina) producta (REUSS), 1852 var.
Plate XXXII Figures 6, 7
1852 Achatina producta — REUSS: p. 11, 32 Pl. III Fig. 12
1874 Oleacina producta — SANDBERGER: p. 444 Pl. 24 Fig. 29
1891 Oleacina producta — KLIKA: p. 23 Fig. 14
1923 Poiretia (Pseudoleacina) producta producta — WENZ F. C.: p. 861
1929 Pseudoleacina producta — PFEFFER: p. 213
With represent to this form, fragments of the last whorl were found in the lacustrine Lower Miocene of the
Somlóvásárhely drilling, as well as an apex. On the apical fragment it is clearly visible that below the suture a well-developed fine axial rib system appears, but this fades out abruptly downwards. This is also clearly visible on Reuss’ original
figure. Some fragments of the columellar margin were also detected. The last whorl is higher in the Hungarian specimens than on Reuss’ figures. Several columellar apertural margin fragments have been
found. The last whorl is taller than that of the figured specimens. Taking into consideration this fact and the proportions of the adult shell, the Hungarian specimens could be
bigger than those from the Czech Republic. Nevertheless, there is insufficient to evidence describe a new taxon. The form is also known from the Lower Miocene in the
Czech Republic.
Pseudoleacina (Pseudoleacina) rakosdensis (GAÁL)
Plate XXXII Figures 12, 13, Textfigure 13
1911 Oleacina rákosdensis — GAÁL: p. 52 Pl. III Fig. 2
1923 Poiretia (Pseudoleacina) rákosdensis — WENZ F. C.: p. 863
A juvenile specimen and a fragment of an adult one have been found in the
Sarmatian of Várpalota. Characteristically, below the suture line the growth-lines are
better developed and more dense. GAÁL (1911) described the species from the Sarmatian
of Rákosd, Transylvania.
Textfigure 13. Pseudoleacina
(Pseudoleacina)
rakosdensis
(GAÁL)
Subgenus: Paraglandina PFEFFER, 1929
Pseudoleacina (Paraglandina) confusa subsulcosa (THOMAE), 1845
1845 Achatina subsulcosa — THOMAE: p. 152, Pl. III Fig. 12
1863 Glandina subsulcosa — SANDBERGER: p. 46, Pl. V Fig. 3
1874 Oleacina subsulcosa — SANDBERGER: p. 410
1923 Poiretia (Pseudoleacina) subsulcosa — WENZ F. C.: p. 865
1959 Pseudoleacina (Paraglandina) subsulcosa — WENZ et ZILCH: p. 458, Fig. 1629
This species has been found in the Lower Miocene of the Somlóvásárhely drilling. The most complete specimen consists of three and a half whorls,which are moderately convex.The protoconch is smooth and the next whorls are axially
ribbed. The ribs are densely-placed, convex, and rather regular. In some cases they bifurcate or dichotomate. On fragments of the last whorl it can be stated that the size of the ribs decreases towards the base. On the second whorl the ribs
are scarcer but better developed. Below the suture there is a slight furrow. The apical angle is low and consequently the
apex is not convex or dome-shaped. The species P. confusa PFEFFER (1929) has a better developed wider but sparser ridge
system. The apex of P. producta (REUSS) is more convex and its shell is thinner. Its axial ridges are finer and denser. The
apex of the species Palaeoglandina gracilis porrecta (GOBANZ, 1854) is convex, wide dome-like and bigger. P. subsulcosa has been found is the North Alpine foreland from the Upper Oligocene to the Lower Miocene.
Genus: Palaeoglandina WENZ, 1914
Palaeoglandina gracilis porrecta (GOBANZ), 1854
Plate XXXII Figures 8, 9
1854 Achatina porrecta — GOBANZ: p. 196, Pl. 3 Fig. 5
1874 Glandina inflata var. porrecta — SANDBERGER: p. 605, Pl. 29 Fig. 32
86
JÓZSEF KÓKAY
1892 Glandina inflata var. porrecta — MAILLARD: p. 5, Pl. 1 Fig. 4
1900 Glandina porrecta — MILLER: p. 401, Pl. 7 Fig. 25
1923 Poiretia (Palaeoglandina) gracilis porrecta — WENZ F. C.: p. 846
1976 Palaeoglandina gracilis porrecta — SCHLICKUM: p. 19, Pl. 5 Fig. 69
Fragments and initial whorls from the Sarmatian of Várpalota may be identified with this species. There are fine axial
ribs and wrinkles on the convex shell surface. Spiral elements are absent, as is clear from MILLER’s (1900) figure.
It has been reported from the “silvana” and Steinheim beds of the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”).
Genus: Poiretia FISCHER, 1883
Poiretia taurinensis (SACCO), 1886
Plate XXXII Figure 10
1886 Glandina taurinensis — SACCO: p. 459, Pl. 2 Fig. 3
1897 Glandina taurinensis — SACCO: XXII: p. 56 Pl. 4 Figs 83, 84
1923 Poiretia (?Poiretia) taurinensis — WENZ F. C.: p. 854
1959 Oleacina sp. — BODA: p. 641, 740, Pl. 36 Fig. 7
2002 Palaeoglandina taurinensis — BINDER: p. 172 Taf. 2 Figs 4–6; Taf. 3 Fig. 3
This species has been found in the Brotia-bearing limestone of the Pusztamiske Pm–3 well. There are two fragmentary moulds. The surface is covered with faint axial growth-lines, and with deeper furrows on the last whorl; this us in
accord with the generic traits. The proportions of the last whorl point to Sacco’s species as described from the
“Elveziano”. Some fragments were collected from the closing beds of the Upper Badenian of the well Pm–1 and from
the coal-seam of the well Pm–4. The “Oleacina” form, found in the Várpalota Sarmatian and described by Boda on the
evidence of an entire specimen, agrees well with this species.
It has also been found in the Karpatien of the Korneuburger Basin.
Poiretia ex aff. taurinensis (SACCO), 1886
Plate XXXII Figure 11
A large-sized, moderately well-preserved specimen was found in the Sarmatian of Várpalota. However, a new species
cannot be identified given the available evidence. Its initial whorls are missing and its aperture is obscured by the matrix.
It seems to be close to Sacco’s species, but it is bigger (its height about 37 mm) and the last whorl is relatively low (70%
of the total height). It has a well-pronounced axial rib system.There are faint growth-lines between the ribs.
Familia: Helicidae
Genus: Perforatella SCHLÜTER, 1838
Subgenus: M o n a c h o i d e s GUDE et WOODWARD, 1921
Perforatella (Monachoides) homalospira (REUSS), 1860
Plate XXXIII Figure 1
1860 Helix (Fruticicola) homalospira — REUSS: p. 65, Pl. I Fig. 3
1874 Helix homalospira — SANDBERGER: p. 429, Pl. 24 Fig. 6
1891 Helix homalospira — KLIKA: p. 53, Fig. 48
1923 Monacha (Monacha) homalospira — WENZ F. C.: p. 413
The species is not rare in the Lower Miocene of the Somlóvásárhely drilling. Many fragments and compressed or
porous fragile specimens have been found. The most complete specimen has 5 slightly convex whorls. The apex is moderately high. The base side is quite convex. The umblicus is narrow. The aperture has a margin that is bent outwards.
There are growth folds, especially on the first three or four whorls. There are small protuberances arranged along oblique
lines that are visible under high magnification. The apical part is delimited toward the base by a blunt ridge. This ridge
is not so pronounced in the Hungarian specimens comparated to the Czech ones. However, this small difference does not
warrant the introduction of a new taxon. As has been indicated above, the species is also known from the Czech Republic,
from layers coeval with the Hungarian ones.
Systematic descriptions
87
Perforatella (Monachoides) punctigera (THOMAE), 1845
Plate XXXIII Figure 2
1874 Helix (Fruticicola) punctigera — SANDBERGER: p. 499, Pl. 25 Fig. 19
1959 Monacha punctigera — BODA: p. 642, 741, Pl. 36 Fig. 11
This species occurs frequently in the Sarmatian of Várpalota, but usually only small fragments are encountered. It is
also present in the Lower and Upper Badenian of Herend. It has been reported from the Lower and Middle Miocene of
Central Europe.
Genus: Caracollina BECK, 1837
Caracollina phacodes barreri (BOURGUIGNAT), 1881
Plate XXXIII Figure 3
1881 Helix Barreri — BOURGUIGNAT: p. 58, Pl. 3 Figs 57–61
1886 Helix (Gonostoma) tenuispirata — ŁOMNICKI: p. 60, Pl. 1 Fig. 5
1923 Caracollina phacodes barreri — WENZ F. C.: p. 465
1976 Caracollina phacodes barreri — SCHLIKUM: p. 16, Pl. 4 Fig. 55
Several damaged specimens originate from the upper part of the Várpalota Sarmatian.
These are covered with strong growth-lines rather than the ribs that are present on Schlikum’s specimens.. It is also
evident in Central and Western European formations, coeval with the Badenian.
Genus: Canariella HESSI, 1918
Canariella bakonyensis nov. sp.
Plate XXXIII Figures 4–5, 6
Derivatio nominis: the type locality belongs to the Bakony Mts
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 115
Locus typicus: Várpalota Basin, well Csór–17, 124.1–127 m
Stratum typicum: Sarmatian limy clay
Descriptio: The dimensions of the holotype: diameter — about 15 mm; height — about 5 mm.
It is a flat shell with six whorls. The whorls are barely convex and the sutures are extremely shallow. The last whorl
encircles the previous ones. The upper half of the whorls is covered with a dense array of radial, gently-curved ribs.
The ribs make an angle of 70–80° with the suture-line. The outer margin of the whorls is slightly keeled and this keel
is separated from the rest by a small furrow. There are some faint cords parallel to the margin, the number of which
decreases as they progress outwards. At the crossings of the cords and ribs there are small knobs. Except for the protoconch, the juvenile whorls are covered with spiral cords on their total width. On the last whorls there are only one to
three cords, parallel to the marginal keel. On the adult whorls the curved ribs vary in size and some are even branching. On the base there is a funnel-shaped umbilicus, covered with wrinkles. The base is smooth and at most some
growth-lines are visible.
Diagnosis differentialis: based on its features, the new species belongs to the genus Canariella (WENZ 1959: p. 695)
and it is characterised by spiral elements. WENZ (1924) ranged his new form — C. disciformis — here, although the spiral elements are missing from it; it is also much taller. The Hungarian specimens are quite close to MAILLARD’s (1892)
Helix nummulina (regarded by Wenz as a synonym) especially with regard to its flatness and the curved ribs. The characteristic keel, however, is lacking from the Swiss form.
Fragments and damaged specimens were found at Pusztamiske, Nyirád, Tapolca Basin and Fehérvárcsurgó (all Upper
Badenian). Some fragments were also collected from the Pusztamiske and Nyirád Sarmatian and from the Várpalota
Middle Badenian (drilling I–124.). Its stratigraphic range extends from the Middle Badenian to the Sarmatian.
Canariella nov. sp.
Plate XXXIII Figures 7–8
Fragments showing characters of this genus have been found in the Lower Miocene of the Somlóvásárhely drilling.
The apical part of the shell is strongly tuberculate. The flat tubercles are radially arranged. Tiny papillae are also present, being arranged along curves. Externally, their size increases slightly. This trait points to the species Canariella disciformis WENZ (1924). However, there is a significant difference: the external margin of the whorls consists of a thick
88
JÓZSEF KÓKAY
and slightly upwardly bent shelf. The upper side of the whorls is flat instead of being convex. In the literature no similar form could be found. The fragments are insufficiently preserved for the introduction of a new taxon. Wenz’ species
is known from the Silvana beds (SCHLICKUM 1976).
Genus: Galactochilus SANDBERGER, 1874
Galactochilus mattiacum (STEININGER), 1835
Plate XXXIII Figures 9, 10, 11
1845 Helix Mattiaca — THOMAE: p. 129, Pl. II Fig. 2
1863 Helix (Xerophila) mattiaca — SANDBERGER: p. 23, Pl. III Fig. 2
1874 Helix (Galactochilus) mattiacum — SANDBERGER: p. 498, Pl. 25 Fig. 17
1921 Galactochilus inflexum mattiacum — WENZ: p. 175, Pl. 25 Figs 4–5
1923 Galactochilus inflexum mattiacum — WENZ F. C.: p. 493
1929 Galactochilus mattiacum — PFEFFER: p. 71
Fragents of this sizable species have been found in the Lower Miocene of the Somlóvásárhely drilling. Its open
umbilicus is covered by a callus of the inner lip. This is completely consistent with the figures and descriptions of the
above-quoted papers. There are rudimentary granulated ridges on the first one and a half whorls. These give place on the
subsequent whorls to unevenly thick growth ridges. These do not completely encircle the wall of the whorls. Some thicker ones are double, especially near the base. Between the ridges there are small papillae of uneven size. The apertural
margin is wide and thick. Given these characters the form may be regarded as identical with the named species which
has also been reported from the Lower Miocene (Aquitanian) of the Northern Alpine foreland.
Galactochilus silesiacus (ANDREAE), 1902
1902 Galactochilus silesiacus — ANDREAE: p. 2, 4, Fig. 3–4
1911 Galactochilus silesiacum — GAÁL: p. 53, 82 Fig. 16
1974 Galactochilus silesiacus — PAPP: p. 388 TextFig. 67, Pl. 18 Fig. 1
1981 Galactochilus silesiacus — LUEGER: p. 58
From the drilling at Hegymagos, (Tapolca Basin) a significantly damaged specimen was found and, by preclusion, it
may best be identified with the named form. From the time span in question only silesiacus and G. sarmaticus GAÁL are
known. The smaller size and the more slender stature points unequivocally to Andreae’s species. Most probably, this
species was the ancestor of G. sarmaticum and G. leobersdorfensis TROLL, 1907.
Galactochilus sarmaticum GAÁL, 1910
Plate XXXIV Figures 1–2
1911 Galactochilus sarmaticum — GAÁL: p. 60, Pl. 1 Figs 1–5, Pl. 85 Fig. 17
1974 Galactochilus sarmaticum — PAPP: p. 387 Fig. 67
Almost entire, large specimens were collected from the Sarmatian of Várpalota. The umbilicus is covered with a thick
and big callus, which indicates this species.
Galactochilus leobersdorfensis (TROLL), 1907
Plate XXXIV Figures 3–4
1907 Helix Leobersdorfensis — TROLL: p. 74, Pl. 2 Fig. 10
1923 Galactochilus leobersdorfensis — WENZ F. C.: p. 494
1957 Galactochilus leobersdorfensis — PAPP: p. 87
1981 Galactochilus leobersdorfensis — LUEGER: p. 57, Pl. 13 Fig. 5
1985 Galactochilus leobersdorfensis — LUEGER: p. 357, Pl. 47 Figs 1–3
One specimen of this taxon has been identified from the Sarmatian of Várpalota. It has also been reported from the
Pannonian s. str. layers of the Vienna Basin. It is slightly compressed and the aperture is missing, but it may be identified with Troll’s taxon. The form of the spire and the smooth upper whorls point to this species. Other Galactochilus
species have a fine granulated and hatched ridge system on the upper whorls. The shell is relatively high and slightly
ovoid. The umbilicus is deep and moderately wide. The taxa described so far have an incompletely covered umbilicus.
On the shell there are well-developed growthlines. The species has also been reported from the Pannonian and Pontian
sediments. In Várpalota, it is associated with G. sarmaticum GAÁL.
Systematic descriptions
89
Genus: Tropidomphalus PILSBRY, 1895
Subgenus: P s e u d o c h l o r i t i s BOETTGER, 1908
Tropidomphalus (Pseudochloritis) robustus (REUSS), 1852
Plate XXXIV Figures 5–6, Textfigure 14
1852 Helix robustus — REUSS: p. 11, 25, Pl. II Fig. 7
1852 Helix trichophora — REUSS: p. 11, 25, Pl. II Fig. 8
1874 Helix (Hemicycla) robusta — SANDBERGER: p. 431, Pl. 25 Fig. 10
1891 Helix (Chloritis) robusta — KLIKA: p. 60 Fig. 56
1923 Tropidomphalus (Pseudochloritis) robustus — WENZ F. C.: p. 517
Some shell fragments of this form were found in the Lower Miocene of the
Somlóvásárhely drilling. From the thicker shell fragments, by the character of the
cylindrical umbilicus and the protoconch, it may be stated that it belongs to the
Textfigure 14. Tropidomphalus (Pseudochloritis) robustus (REUSS), 1852
named genus and respectively the subgenus. The shell fragments are covered with
well-developed, partly dichotomising growth folds. On these folds there are flat
protuberances. These are generally round but some are oval or angulate. These elevations are generally arranged along
the growth-lines with shorter or longer intermittent spaces. This type of shell surface characterises the species T. robustus (which has been described by Klika in details). The shell fragments can be identified with this species. One entire,
but compressed specimen has also been found. The form has also been reported from the Lower Miocene of the Czech
Republic.
Tropidomphalus (Pseudochloritis) extinctus (RAMBUR), 1862
Plate XXXIV Figure 7
1862 Helix extincta — RAMBUR: p. 172, Pl. VII Figs 5–6
1874 Helix (Campylaea) extincta — SANDBERGER: p. 531, Pl. 26 Fig. 20
1892 Helix (Campylaea) extincta — MAILLARD: p. 25, Pl. II Fig. 10
1911 Helix (Campylaea) extincta — COLLOT: p. 98, Pl. XII Fig. 25
1923 Tropidomphalus (Pseudochloritis) extinctus — WENZ F. C.: p. 508
Some damaged specimens and fragments have been found in the Lower Miocene of the Polgárdi and Somlóvásárhely
drillings. A half whorl from the Somlóvásárhely drilling agrees well with that on the figure of Collot in that the upper
side of the whorl is flattened and thus it is asymmetric. On the shell surface there are small protuberances arranged along
oblique lines. On the whorls there are growth folds and short cutting-like depressions. At the base there is a well-developed umbilicus. It is known from middle France and Swiss, from layers coeval with the Somlóvásárhely specimens.
Tropidomphalus (Pseudochloritis) gigas PFEFFER, 1929
Plate XXXIV Figures 8–9
1929 Tropidomphalus (Pseudochloritis) gigas — PFEFFER: p. 226
1951 Tropidomphalus (Pseudochloritis) gigas — PAPP: p. 63, 64
1967 Tropidomphalus (Pseudochloritis) gigas — SCHÜTT: p. 219, Fig. 23
1974 Tropidomphalus (Pseudochloritis) gigas — PAPP: p. 389, Pl. 18 Fig. 2
1981 Tropidomphalus (Pseudochloritis) gigas — LUEGER: p. 58, Pl. 12 Fig. 4, Pl. 13 Fig. 4, Pl. 16 Fig. 5
Some fragments and one big specimen can be identified with this species, collected from the Bánd bentonite pit in
the Herend Basin. Although the last half whorl is missing, it agrees well with the named species. Originally, it might have
been 35 mm in diameter. The closest form is T. exstinctus (RAMBUR), which is unlike T. gigas in the following respects:
— the shell is lower and slightly steplike,
— the sutures are deeper,
— the umbilicus is wider,
— the growth-lines and wrinkles are more developed,
— it is somewhat smaller with a maximum height of 30 mm.
The identical features are:
— a depressed spire which is moderately convex at its top,
— the wall is relatively thick.
T. gigas is possibly a descendant of T. exstinctus. The main difference between them is that T. gigas has about one
and a half more whorls.
90
JÓZSEF KÓKAY
The generic features may be easily recognised under a strong lens:
— there are small wart-like knobs on most whorls (except the last one) and especially on the initial ones,
— on the last whorl there is a very faint spiral groove system. These grooves are not continuous: they are regularly
interrupted by growth-lines and wrinkles.
The form is reported from the Upper Badenian, Sarmatian and Pannonian (s.str.) of Austria. Fragments from the
Pusztamiske Pm–1 well also belong here, as well as specimens from the Sarmatian of Várpalota.
Tropidomphalus (Pseudochloritis) zelli (KURR), 1856
1874 Helix (Campylaea) Zellii — SANDBERGER: p. 591, Pl. 29 Fig. 9
1892 Helix (Campylaea) Zelli — MAILLARD: p. 28, Pl. 2 Fig. 12
1910 Helix (Campylaea) Zelli — ROLLIER: p. 133, Pl. 3 Fig. 10
1923 Tropidomphalus (Pseudochloritis) zelli — WENZ F. C.: p. 519
1976 Tropidomphalus (Pseudochloritis) zelli — SCHLICKUM: p. 16
1981 Tropidomphalus (Pseudochloritis) zelli — LUEGER: p. 60
Some fragments of this form were found in the Lower Badenian coal-bearing sequence of the Herend Basin (well
Bánd–2). The apical part, with its initial whorls, is very characteristic. Lueger published a photo of the initial whorls of
the species T. zelli depressus (1981: Pl. 16 Figure 4). This is very similar to the Herend specimen. The species deviates
by its more convex and out-standing whorls, in agreement with Lueger’s description. The initial whorls are covered by a
rugged surface and the knoblets seem to be arranged radially. There is a well expressed growth-line system on the surface. It has also been found in the “silvana” beds of the North Alpine Molasse Zone.
Tropidomphalus (Pseudochloritis) zelli depressus WENZ, 1927
Plate XXXIV Figures 10–11
1927 Tropidomphalus (Pseudochloritis) zelli depressus — WENZ: p. 45, Pl. 2 Fig. 1
1981 Tropidomphalus (Pseudochloritis) zelli depressus — LUEGER: p. 59, Pl. 11 Fig. 1, Pl. 12 Fig. 5, Pl. 16 Fig. 4
A compressed specimen was found in the Sarmatian at Várpalota (drilling Csór–17). The best preserved part is its
apical zone. It has also been found in the Vienna Basin and Transdanubian Pannonian and Pontian.
Tropidomphalus (Pseudochloritis) cf. incrassatus (KLEIN), 1853
Plate XXXIV Figures 12, 13, 14
1853 Helix incrassata — KLEIN: p. 208, Plate 5 Fig. 6
1874 Helix (Campylaea) inflexa — SANDBERGER: p. 589, Pl. 29 Fig. 8
1923 Tropidomphalus (Pseudochloritis) incrassatus incrassatus — WENZ F. C.: p. 510
1929 Pseudochloritis incrassatus — PFEFFER: p. 225
1976 Tropidomphalus (Pseudochloritis) incrassatus incrassatus — SCHLIKUM: p. 16, Pl. Fig. 56
Characteristic fragments were collected from wells Pm–2 and Pm–4. Thiny round setiferous pits are only visible
under a strong lens. These are arranged along diagonal and parallel rows. Given the aboves a specific determination can
be made, as:
— in T. zellii (Kurr) the pits are bigger, they are oval, and they are wider spaced,
— T. zellii is widespread in the North Alpine Molasse Zone, from beds coeval with the Hungarian ones.
Genus: Helicigona RISSO, 1826
Helicigona wenzi SOÓS, 1934
Plate XXXV Figure 1
1934 Helicigona wenzi — SOÓS: p. 210, Fig. 12
1955 Helicigona wentzi — BARTHA: p. 311, Pl. 2 Figs 7, 14, 15
1959 Helicigona wentzi — BARTHA: Pl. 17 Figs 6, 7
1981 Helicigona wenzi — LUEGER: p. 66, Pl. 8 Figs 9, 10, Pl. 16 Fig. 8
An apical part has been collected from the coal-bearing sequence in the Pusztamiske Pm–3 well. The flat (but not
compressed by overburden) shell and the surface (including the initial whorls) are covered with small knobs; these facts
indicate the species and they are in agreement with Lueger’s figure. Radial growth-wrinkles are fainter in our specimen.
The whorls are moderately convex. The closest form, H. atava WENZ 1927 has a less depressed apical part. Similar ini-
Systematic descriptions
91
tial whorls originate from other Pusztamiske, Herend and Nyirád wells (from the Upper Badenian). Compressed, damaged specimens were also collected from the Sarmatian of Várpalota.
This form is also present from the Karpatian to the Pontian (Upper Pannonian s.l.).
Genus: Klikia PILSBRY, 1895
Subgenus: K l i k i a
Klikia (Klikia) osculum tenuis (KLIKA), 1891
Plate XXXV Figures 2–3, 4–7
1891 Helix osculum var. tenuis — KLIKA: p. 49 Fig. 43
1911 Klikia osculum THOM. var. tenuis — WENZ: p. 84
1923 Klikia (Klikia) osculum var. tenuis — WENZ F. C.: p. 552
Several specimens have been found in the Lower Miocene of the Somlóvásárhely drilling. These seem to be transitional between the Lower Miocene K. osculum (THOMAE) and the Middle Miocene K. giengensis (KLEIN) taxa. No entire
specimen has been found but it is clear that the surface is finely granulate in the elongated forms. The granulae are most
conspicuous on the first three or four whorls, but here they are more spars than on the rest of the shell.The whorls are
flatter in the nominal subspecies but the form of the umbilicus and the aperture is essentially the same as that of the
new form. Around the umbilical part of the aperture there is a callus. Our specimens are 8.5 to 10.5 mm wide and thus
they are smaller than the nominal form but greater than K. giengensis (KLEIN). The species K. osculum villosella
(THOMAE, 1854) has a wider umbilicus and the shell is more flattened and its decoration is finer. Given these details
the Hungarian form agrees well with Klika’s species, which has been reported from the Lower Miocene of the Czech
Republic.
Klikia (Klikia) ex gr. osculum (THOMAE), 1845
Plate XXXV Figure 8
1845 Helix osculum — THOMAE: p. 137, Pl. III Fig. 4
1863 Helix (Fruticicola) osculum — SANDBERGER: p. 19, Pl. III Fig. 13
1874 Helix (Gonostoma) osculum — SANDBERGER: p. 377, 427, 455, Pl. 22 Fig. 18
1891 Helix osculum — KLIKA: p. 47 Fig. 41
1891 Helix osculum var. ornata — KLIKA: p. 49 Fig. 49
1911 Gonostoma (Klikia) osculum — WENZ: p. 79, 97, Pl. IV Figs 1–5
1923 Klikia (Klikia) osculum osculum — WENZ F. C.: p. 547
1929 Klikia osculum — PFEFFER: p. 86
Two fragments of this form were found in the Lower Miocene of the Somlóvásárhely drilling. These are parts of specimens that are 12 to 13 mm in size. The umbilicus is moderately wide while the apertural margin is moderately thickened and bent. The most characteristic deviation from Klikia’s form is that the decoration is not reticulate and it is not
rice-grain (i.e. elongated) in its form. Small, round protuberances (papillae) are arranged along oblique lines sparsely but
regularly. Pfeffer mentions such a feature on the shell. This form can in all likelihood be regarded as a member of the
nominal subspecies or one of the other subspecies. K. osculum is has also been found in the Upper Oligocene to the
Lower Miocene in the Czech Republic.
Klikia (Klikia) giengensis (KLEIN), 1847
Plate XXXV Figure 9
1847 Helix Giengensis — KLEIN: p. 69, Pl. 1 Fig. 9
1874 Helix (Gonostoma) osculum var. giengensis — SANDBERGER: p. 585, Pl. 29 Fig. 4
1892 Helix (Gonostoma) osculum var. giengensis — MAILLARD: p. 68, Pl. 5 Fig. 12
1910 Helix (Gonostoma) Giengensis — ROLLIER: p. 133, Pl. 3 Figs 48–51
1911 Klikia giengensis — WENZ: p. 85, Pl. 4 Figs 6–10
1923 Klikia giengensis — WENZ F. C.: p. 539
1959 Helicigona aff. leptoloma apicalis — BODA: p. 643, 742, Pl. 37 Fig. 2
1976 Klikia (Klikia) giengensis — SCHLICKUM: p. 16, Pl. 4 Fig. 58
Its an entirely preserved specimen of this form was collected from the Sarmatian of Várpalota (Shaft M. III). Its
dimensions (width — 9 mm; height — 5.3 mm), proportions, the aperture and the moderate-sized umbilicus all point to
this species. A few broken specimens were found in the Upper Badenian coal sequence of Pusztamiske (Pm–1). These
pieces enable a determination to be made (i.e. backward-bent aperture, open umbilicus, size). Around the umbilicus and
behind the apertural margin the shell surface shows the characters of this species.
It is also known from the North Alpine and Western European “Tortonian” (coeval with the Badenian).
92
JÓZSEF KÓKAY
Klikia (Klikia) kaeufeli WENZ, 1927
Plate XXXV Figure 10
1927 Klikia (Klikia) kaeufeli — WENZ: p. 45, Pl. 2 Fig. 5
1981 Klikia (Klikia) kaeufeli — LUEGER: p. 67, Pl. 10 Fig. 2
This form is widespread in the Várpalota Sarmatian, and entire specimens have been found (in the shaft M. III). It is
generally bigger and lower than Klikia giengensis. It has also been found in the Pannonian of the Vienna Basin.
Subgenus: A p u l a BOETTGER, 1909
Klikia (Apula) goniostoma (SANDBERGER), 1874
Plate XXXV Figure 17
1874 Helix (Fruticicola) goniostoma — SANDBERGER: p. 702, Pl. 32 Fig. 12
1934 Helicigona (Kosicia) Pelissae — SOÓS: p. 199, Fig. 8
1934 Helicigona (Campylaea) Gaali — SOÓS: p. 200, Fig. 10
1959 Helicigona pontica — BARTHA: Pl. 17 Figs 1, 4, 5
1981 Klikia (Apula) goniostoma — LUEGER: p. 68, Pl. 10 Fig. 3
2004 Klikia (Apula) goniostoma — HARZHAUSER et BINDER: p. 134, pl. 10 Figs. 6–9
One specimen of this form has been recovered from the upper part of the Várpalota Sarmatian. This is an internal
mould, and it is almost completely decorticated. A notable characteristic is its big size (19 mm in diameter) and the narrow umbilicus which is greatly covered by the outwardly bent aperture. The spire is realtively low (8–9 mm). It is close
to the form described by Soós as Helicigona pelissae. Based on this, it may be identified with Sandberger’s species. So
far, it has been reported only from Pannonian and Pontian layers.
Klikia (Apula) cf. devexa (REUSS) var., 1860
Plate XXXV Figures 11–15
1860 Helix devexa — REUSS: p. 65, Pl. I Fig. 4
1874 Helix (Fruticicola) devexa — SANDBERGER: p. 429, Pl. 24 Fig. 5
1891 Helix devexa — KLIKA: p. 50, 54 Fig. 45
1911 Klikia devexa — WENZ: p. 94, Pl. IV Figs 26–28
1923 Klikia (Apula) devexa — WENZ F. C.: p. 537
1929 Klikia (Apula) devexa — PFEFFER: p. 87
An almost entire but compressed specimen has been found in the lower part of the Lower Miocene of the
Somlóvásárhely drilling. This may be identical with the mentioned form. It is a moderately flat form. On the last whorl
there is a (hardly) visible blunt keel. Growth-lines and radial folds are well-developed on the basal side, near the umbilicus. There is a fine granulation on the shell surface, especially near the apex and close to the umbilicus. The aperture is
bent externally but its umbilical part is missing. The diameter of this specimen is 11.5 mm, but another specimen is 13
mm. On two, compressed, specimens it is clear that before the damaged apertural margin there is a slight embayment,
characteristic for the species. SANDBERGER (1874) mentions a bigger specimen with a more open umbilicus. KLIKA (1891)
also refers to the same specimen. However, they did not give any figures. It is possible that our specimens belong to the
same taxon. The quite wide umbilicus is a character of BABOR’s (1898) subspecies prominens too. The diameter of the
Babor’s subspecies is only 10 mm. The subspecies of the form, described by BABOR (1898) has been reported from the
Lower Miocene of the Czech Republic.
Klikia (Apula) cf. coarctata (KLEIN), 1853
1853 Helix coarctata — KLEIN: p. 206, Pl 5 Fig. 3
1874 Helix (Monacha) coarctata — SANDBERGER: p. 586, Pl. 29 Fig. 5
1892 Helix (Monacha) coarctata — MAILLARD: p. 71, Pl. 5 Fig. 14
1923 Klikia (Apula) coarctata — WENZ F. C.: p. 534
1976 Klikia (Apula) coarctata — SCHLICKUM: p. 17, Pl. 4 Fig. 60
Some fragments of this form were collected from the Lower Badenian of the Herend Basin (drilling Bánd–4). Under
a strong lens a very small, densely-packed network of protuberances is visible. This given the size of the fragments and
the depressed umbilicus, indicates the named species. The Upper Badenian of Pusztamiske (drilling Pm–3) also yielded
similar fragments.
This form has also been reported from the “silvana” beds of the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) and from the Vienna Basin Sarmatian.
Systematic descriptions
Klikia (Apula) cf. coarctata steinheimensis JOOSS, 1918
Plate XXXV Figure 16
1918 Klikia coarctata var. steinheimensis — JOOSS: p. 294
1923 Klikia (Apula) coarctata steinheimensis — WENZ F. C.: p. 537
1927 Klikia (Apula) coarctata steinheimensis — WENZ: p. 46, Pl. 2 Fig. 4
1981 Klikia (Apula) coarctata steinheimensis — LUEGER: p. 69, Pl. 10 Fig. 4
A damaged internal mould was found in the Várpalota Sarmatian (drillings Küngös–2, Csór–8). These
are 14 mm in diameter. The print of the apertural margin points to the subgenus Apula. It is characterised
by a narrow umbilicus a depressed umbilical area and a flat shell. These features strongly point to this
species. Another, compressed specimen is 12 mm in diameter.
The form has been recorded from Steinheim, and also from the Pannonian (s.str.) of the Vienna Basin.
Genus: Cepaea HELD, 1857
Cepaea bohemica (BOETTGER), 1870
Plate XXXVI Figure 1
1852 Helix rostrata — REUSS: p. 27 Pl. II Fig. 9
1870 Helix (Macularia) bohemica — BOETTGER: p. 290 Pl. XIII Fig. 4
1874 Helix (Coryda) bohemica — SANDBERGER: p. 432 Pl. 24 Fig. 8
1891 Helix (Coryda) bohemica — KLIKA: p. 57 Fig. 54
1923 Cepaea bohemica — WENZ F. C.: p. 609
A slightly compressed specimen has been found in the Lower Miocene of the Somlóvásárhely drilling.
It is closely related to C. silvana, but deviates by its smaller size (diameter — 18 mm, while C. silvana is
20 to 25 mm wide). The apertural margin of the Hungarian specimen is bent outwards smoothly, while in
C. silvana this turn is generally abrupt and the margin is thicker. A part of the growth-lines show a forklike branching.
From the Polgárdi drilling (KÓKAY 2001.) an internal mould was collected with some shell remnants.
This is 16 mm in diameter. On the second and third whorl there are fine arched growth lines. The external
aperture is bent outwards at a right angle, similar to that on Boettger’s Figure 4. This form is frequent in
the Lower Miocene of the Czech Republic. It has been also found in the Middle Badenian of Sámsonháza
(two specimens).
Cepaea silvana (KLEIN), 1847
Plate XXXVI Figure 2–3
1847 Helix sylvestrina var. — KLEIN: p. 67, Pl. 1 Fig. 4
1851 Helix silvestrina — DUNKER: p. 161, Pl. 21 Figs 3–5
1853 Helix silvana — KLEIN: p. 205, Pl. 5 Fig. 2
1874 Helix (Macularia) sylvana — SANDBERGER: p. 592, Pl. 29 Fig. 13
1910 Helix (Macularia) silvana — ROLLIER: p. 134, Pl. 3 Figs 64–118
1923 Cepaea silvana silvana — WENZ F. C.: p. 667
1976 Cepaea silvana silvana — SCHLICKUM: p. 17, Pl. 4 Figs 62, 63
1989 Cepaea silvana silvana — REICHENBACHER: p. 165, Pl. 2 Figs 17–19
One compressed specimen was collected from the Pusztamiske Upper Badenian (drilling Pm–3). Its
main features are in a good state — i.e. the strongly marginated aperture, the lip and the umbilicus. The
shell is generally adorned with coloured bands, but unicoloured specimens are also frequent (such as the
Pusztamiske one). A complete external mould was collected from a Brotia-bearing limestone of the well
Pm–3.
The form has also been reported from the “sylvana” beds of the North Alpine Freshwater Molasse
(“Süssbrackwassermolasse”, and this is the original its name). Its range extends from the Karpatian to the
Sarmatian, in Paratethyan terms. So far it has not been found in the Karpatian and Vienna Basins.
Cepaea etelkae (HALAVÁTS), 1923
1959 Cepaea silvestrina etelkae — BODA: p. 643, 742, Pl. 37 Fig. 4
1959 Cepaea sylvestrina etelkae — BARTHA: p. 82, Pl. 16 Figs 3–4
1959 Cepaea etelkae — BARTHA: Pl. 16 Figs 2, 5
1981 Cepaea (Cepaea) etelkae — LUEGER: p. 72, 742, Pl. 13 Figs 1, 2, Pl. 14 Figs 1–7
93
94
JÓZSEF KÓKAY
This form is frequent in the Sarmatian of Várpalota, but entire specimens are rare. Boda reported it from the
Sarmatian of this and other regions. It is widespread in the Pannonian and Pontian of the Carpathian and Vienna
Basins.
Classis: BIVALVIA
Familia: Dreissenidae
Genus: Dreissena VAN BENEDEN, 1835
Dreissena dobrei (BRUSINA), 1902
Plate XXXVI Figure 4
1902 Dreissensia dobrei — BRUSINA: Pl. 21 Figs 58–63
1959 Dreissena dobrei — BARTHA: p. 84, Pl. 8 Figs 1, 2
One single specimen from the middle part of the Várpalota Sarmatian (drilling Berhida–3) agrees well with Brusina’s
species. Its ventral side is slightly steeper, but the variability of the species covers this feature. It is reported as being 9–12
mm long. The Várpalota specimen is 9 mm. It has also been found in the Pontian around lake Balaton. The genus might
have lived in fresh- or slightly brackish waters.
Genus: Congeria PARTSCH, 1835
Congeria boeckhi WENZ, 1931
Plate XXXVI Figure 5
1931 Congeria böckhi — WENZ: p. 117, Pl. 10 Fig. 1
1967 Congeria böckhi — KÓKAY: p. 77
1978 Congeria boeckhi — KOCHANSKY-DEVIDE et ŠLISKOVIĆ: p. 47, Pl. 5 Figs 1–14
It occurs in masses in the immediate hanging wall of the Várpalota Upper Badenian coal-seam as well as in the
“Congeria-limestone”; the latter is a nearshore heteropic facies of alginite. There are concentric or zigzag brown patterns
on many specimens. In Transdanubia (W Hungary) it has been reported from the Ottnangian to the Upper Badenian.
WENZ (1931) described it from the Karpatian (“Helvetian”) of the Mecsek Mts and also mentioned its occurrence in the
Várpalota “Tortonian” (= Badenian). Kochansky-Devide and Šlisković report its presence in the Ottnangian and
Karpatian of Croatia and Northern Bosnia.
Congeria venusta KOCHANSKY-DEVIDÉ et ŠLISKOVIĆ, 1978 var.
Plate XXXVI Figure 6
1978 Congeria venusta KOCHANSKY-DEVIDE et ŠLISKOVIĆ: p. 69, 96, Pl. 13 Figs 20–23, Pl. 14 Figs 24–40, Fig. 3
A small-sized, double-shelled, opened Congeria was found in the marly hanging wall of the Várpalota coal-seam.
With some hesitation, I identified this with Kochansky-Devide’s form. The ventral side of the type is curved outward,
while in the Hungarian specimen it is straight. Other features — i.e. size, position of the beak, the gently curved crest
agree well with those of the type series, although most of them have a sharper crest. The agreement is best with figures
30, 31, 35, and 36 in KOCHANSKY-DEVIDE et ŠLISKOVIĆ (1978), Pl. 13.
Congeria brardii BRONGNIART — reported from the Lower Badenian of Herend (KÓKAY 1966: p. 73, Pl. 13 Figs 5, 6)
— is bigger than the above-described Várpalota form and its crest is blunter and the ventral side is steeper.
It has also been reported from the Ottnangian and Karpatian freshwater deposits of Croatia and Bosnia-Herzegovina
Congeria hercegovinensis KOCHANSKY-DEVIDÉ et ŠLISKOVIĆ, 1978
Plate XXXVI Figure 7
1978 Congeria hercegovinensis — KOCHANSKY-DEVIDE et ŠLISKOVIĆ: p. 38, 87, Pl. 2 Figs 1, 18, Fig. 4
A small-sized, Mytilus-shaped Congera-species was found in the hanging wall of the Upper Badenian coal-seam of
Várpalota. It has a sharp crest which tends towards the ventral side. The ventral side is steep while the dorsal side is semicircular. These features agree with the mentioned species, but its authors mention specimens in which the dorsal upper
and the lower margin meet at an angle. Our single specimen agrees with Kochansky-Devide et Šlisković Pl. 2 Figure 1.
It has also been found is deposits in Bosnia-Herzegovina and these are coeval with the Ottnangian, Karpatian and
Badenian.
Systematic descriptions
95
Familia: Margaritiferidae HAAS, 1940
Genus: Margaritifera SCHUMACHER, 1816
Margaritifera flabellata (GOLDFUSS), 1837
Plate XXXVI Figures 8–10
1863 Unio flabellatus — GOLDFUSS: p. 173, Pl. 132 Fig. 4
1863 Unio costatus — GOLDFUSS: p. 173, Pl. 132 Fig. 5
1851 Margaritana wetzleri — DUNKER: p. 162, Pl. 21 Figs 25, 26
1938 Margaritifera flabellata — MODELL: p. 148
1964 Margaritifera flabellata — SCHLICKUM: p. 22, Pl. 3 Fig. 48
Fragmentary specimens of this form were collected from the closing beds of the Várpalota Lower Badenian. The main
features of these specimens are:
— an elongate form, length about 50 mm, height 23 mm,
— the beak is shifted forwards, it is flat and not prominrnt,
— the shell is thick and the lower margin is straight,
— on the beak region there are undulate zigzag wrinkles, while on the areal field it is covered with oblique ridges
running to the upper posterior margin,
— the cardinal teeth are relatively well-developed.
Ecologically it is not easy to interpret the presence of this form given that the extant European Margaritifera margaritifera (LINNAEUS) lives exclusively in clear, oxygen rich, cool freshwater, depleted of calcium-hydrocarbonate (i.e.
low hardness). These conditions clearly were not fulfilled in the Miocene paludal environment, but the big size of the
bivalve indicates conditions close to optimal ones.
The shell is characteristically kidney-shaped — i.e. the lower margin is embayed — but this is not universal. The figures of Goldfuss, Dunker and Schlikum show straight margins. Schlickhum’s figure shows a subjuvenile specimen, similar to the Várpalota ones. The species lived in a fluvial environment and the shells were washed into a marine environment, together with those of other freshwater molluscs.
Some fragments have been in the Upper Badenian of Herend found. A right valve, preserved around the beak, is covered with undulate wrinkles and it has a strong cardinal tooth.
The species is widespread in the Lower and Middle Miocene of the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”).
Margaritifera flabellata bavarica MODELL, 1938
Plate XXXVII Figure 1
1874 Unio flabellatus — SANDBERGER: p. 568 Pl. 30 Fig. 1
1893 Unio flabellatus — LOCARD: p. 237 Pl. 11 Fig. 16
1938 Margaritifera flabellata bavarica — MODELL: p. 148
1940 Margaritifera fabellata bavarica — MODELL: p. 92, 94
1957 Margaritifera flabellata bavarica — MODELL: p. 96 Fig. 1
Modell proposed a new subspecies for this big form (length — 93 mm; width — 49 mm). A similarly big specimen
was collected from the Lower Badenian coal bearing series of Herend. It is accompanied by freshwater molluscs: Brotia
escheri (frequent), Theodoxus barakovici (frequent), Nematurella scholli (frequent), Microprososthenia belchatovensis
(not rare), Valvata moguntina, Gyraulus trochiformis dealbatus, Ferrissia deperdita, Pisidium bellardii.
Modell stated that the range of the taxon is from the Lower Badenian to the Sarmatian.
Familia: Unionidae FLEMING, 1828
Genus: Unio PHILIPSSON, 1788
Unio eseri KRAUSS, 1852
Plate XXXVII Figure 3
1852 Unio Eseri — KRAUSS: p. 153, Pl. 3 Fig. 6
1874 Unio eseri — SANDBERGER: p. 556, Pl. 30 Fig. 3
1964 Unio eseri — SCHLICKUM: p. 22, Pl. 3 Figs 49, 50
The closing beds of the Várpalota Lower Badenian cycle yielded a right valve of a unionid the posterior part of which
is missing. In spite of the incomplete outline it agrees well with the mentioned species, taking into consideration the high
96
JÓZSEF KÓKAY
degree of variability of the unionids, especially with regard to dentation. The cardinal tooth is well-developed and it narrows downwards; there is a notch between it and the inner margin for the posterior cardinal tooth of the left valve.
The anterior muscle attachment is deep and strong, and it is separated into anterior and posterior parts by a small wall,
in agreement with Krauss’ figure.
The species has hitherto been reported only from the “Helvetian” and “Sarmatian” of the North Alpine Freshwater
Molasse (“Süssbrackwassermolasse”). One fragment from the base of the Lower Badenian in the Nagygörbő well may
also belong here.
Unio jaccardi LOCARD, 1892
Plate XXXVII Figure 2
1893 Unio Jaccardi — LOCARD: p. 241, Pl. 11 Fig. 19, Pl. 12 Fig. 11
1938 Unio jaccardi — MODELL: p. 150
1967 Crassunio (Crassus) jaccardi — MODELL: p. 862, Pl. 14 Figs 14, 15
Poorly preserved specimens from the upper part of the Várpalota Upper Badenian pelitic sequence agree well with
Locard’s species with respect to outline and size.
It has also been found in the coeval formations of the Swiss Molasse („Oeningien”) MODELL (1967) found it at Hessen
(„Torton”), Germany.
Unio larteti NOULET, 1881
Plate XXXVII Figures 4–5
1881 Unio Larteti — BOURGUIGNAT: p. 154, Pl. 8 Fig. 302
1943 Elliptio larteti — MODELL: p. 110, Pl. 1 Figs 1, 2
Two fragmentary left valves originate from the Lower Badenian of Nagygörbő. Their outline, proportions as well as
the concentric growth-lines (ribs) point to Noulet’s species. The latter was found in coeval freshwater deposits of Sansan,
southern France. Unfortunately, due to the poor state of preservation it is not possible to recognize the fine-wrinkled
ornament on the beak area. This is, however, present in several Unio species. In one of the valves there is a dental lathe,
parallel to the posterior margin.
The Hungarian specimens are smaller than the French ones, but the proportions are similar. According to Bourguignat
the length is 41 mm and the height is 19 mm. In the Nagygörbő specimen the dimensions are: 29 mm, and 14.5 mm,
respectively. The beak is placed similarly. The Hungarian specimens seem to be close to the Lower Miocene Unio favrei
of Swiss as well (LOCARD 1893: p. 246, Pl. 12 Figure 14), but it is much taller (length, 28, height, 18 mm). Based on upto date systematic research (HAAS 1969) it appears that it does not belong to the genus Elliptio.
Unio hinschi MODELL, 1967
Plate XXXVIII Figure 1
1967 Unio hinschi — MODELL: p. 864 Tab. 14. Figs 20–22
One specimen of this form was collected from the uppermost layers of the Upper Badenian of the Várpalota Basin,
and it can be identified with Modell‘ s species. It is close to the form on Figure 22. It also has an elongated triangular
outline with a forward-pushed obtuse umbo. Its posterior margin is less pointed and it is similar to Figure 21 of Modell.
On the surface the are strong ribs. Its lenght/height ratio is 1.82 on Figure 22, while the ratio of the Várpalota specimen
is 1.88. According to Modell the species has a variable outline.
Modell described the taxon from the “ Sarmatian Unio layers “ at the river Rhine (Pfungstadt, Germany).
Unio lorioli LOCARD, 1892
Plate XXXVIII Figures 2–3
1892 Unio Lorioli — LOCARD: p. 243, Pl. 11 Fig. 18
A damaged left valve and a beak of a right valve were found in the closing beds of the Lower Badenian of Várpalota.
Its outline is close to that of Locard’s species, although the posterior and lower margin meet in a more angular way. The
lower margin is curved. Its beak is placed in a more posterior position than that of Locard’s species. The beak region is covered with undulate zigzag wrinkles, similar to that of the Alpine form. Locard did not mention the teeth. The cardinal tooth
of the left valve is similar to that of the living European U. crassus RETZIUS, 1783. The anterior cardinal tooth is a flat plate
which barely stands out from the margin. The posterior plate is triangular and protrudes outwards significantly. There is a
narrow notch between the two cardinal teeth which provides a place for the cardinal tooth of the right valve. The mentioned
Systematic descriptions
97
insignificant deviations are within the range of most unionids and, therefore the specimen can be placed alongside Locard’s
species. The length/width ratio is 66% in the Helvetian form, while it is 63% in the Hungarian one.
It has also been found in the North Alpine Freshwater Molasse (“Süssbrackwassermolasse”) in Swiss, which is similar in age to the Hungarian locality.
Unio cf. blumrichi MODELL, 1940
1940 Unio blumrichi — MODELL: p. 91, Pl. 4. Figs 3–5
1976 Unio cf. blumrichi — SCHLICKUM: p. 17, Pl. 5 Fig. 64
Some umbo fragments were found in the Lower Miocene of the Somlóvásárhely drilling. These may be best ranged
into Modell’s species. On the surface there are jagged and platy ribs which are identical with those on Schlickum’s figure. The species reported from freshwater sediments of the North Alpine foreland, coeval with the Badenian.
Familia: Spheriidae
Genus: Spherium SCOPOLI, 1777
Spherium sp.
Many fragments of this thin-shelled species have been recognised in the Lower Miocene of the Somlóvásárhely
drilling. An entire shell could have been up to 6 mm or 8 mm long. There is no way to determine this form specifically.
Genus: Pisidium PFEIFFER, 1828
Pisidium bellardii BRUSINA, 1897
Plate XXXVIII Figures 4–5
1897 Pisidium bellardii — BRUSINA: p. 36, Pl 21 Figs 43–45
1955 Pisidium bellardii — BARTHA: p. 312, Pl. 2 Figs 6, 8
This form has been found in the lower parts of the Lower and Upper Badenian of Herend, and from the uppermost,
freshening bed of the Várpalota Lower Badenian (V–297 well); it has also been reported from from the Sarmatian of the
same area (Shaft M. III). The specimens are well-preserved and agree well with Brusina’s examples from the Miocene
of Dalmatia. Bartha reported the form from the “Upper Pannonian” (=Pontian) of Várpalota.
Pisidium pseudosphaerium SCHLESCH, 1947 var.
Plate XXXVIII Figures 6–8, Plate XXXIX Figures 1–4, 6
1979 Pisidium pseudosphaerium — RICHNOVSZKY et PINTÉR: p. 174, Fig. 73
1980 Pisidium pseudosphaerium — GLÖER et al.: p. 60 (with figure)
1993 Pisidium pseudosphaerium — PIECHOCKI et DYDUCH-FALNIOWSKA: p. 152, Figs 81, 82e
This form demonstrates a high variability and is frequent in the Sarmatian of Várpalota. The description of
RICHNOVSZKY, PINTÉR (1979) fits well to these specimens, but its variability is much higher with respect its outline, thickness and the development of its dentation. It is quite flat and the beak protrudes only slightly but these features are variable, too. The shell surface is smooth, generally bearing one to four growth-lines. The smooth shell is the feature which
deviates most from the extant species. So far it has not been found in a fossil state. It lives exclusively in freshwater of
considerable hardness.
Pisidium steinheimense GOTTSCHICK, 1920
Plate XXXIX Figure 5
1920 Pisidium steinheimense — GOTTSCHICK: Pl. 2 Fig. 14
1921 Pisidium steinheimense — GOTTSCHICK: p. 177
This form was found in the Upper Badenian of the drilling Nyrt–1 in the Lower and Upper Badenian of Herend as
well as in the Sarmatian of Várpalota (Shaft M. III). These examples agree well with the named species with respect to
the oval outline, the dense and faint growth-lines, the position of the beak and the shape. The main feature unifying these
forms is the embayment of the margin of the valves along the lateral teeth.
It has also been found is coeval freshwater deposits at Steinheim, Germany.
98
JÓZSEF KÓKAY
Pisidium annandalei (PRASHAD), 1925
Plate XXXIX Figure 7; Textfigure 15
1997 Pisidium annandalei — PIECHOCKI: p. 114 Fig. 13
One, partly damaged, specimen was found in the Upper Badenian beds of Herend.
It is characteristically tall and triangle-shaped with an evolved dental margin. This
species is very widespread in the Miocene of Europe, in Turkey, and especially in the
Middle Miocene of Poland.
Textfigure 15. Pisidium annandalei (PRASHAD), 1925
Pisidium bakonyensis nov. sp.
Plate XL Figures 1–2, 3–4
Derivatio nominis: the type locality belongs to the Bakony Mts
Holotypus: deposited in the Geological Museum of Hungary (Geological Institute of Hungary, Budapest), Inv. No MÁFI, Gy. 116
Locus typicus: Pusztamiske, well Pm–1, 52 m
Stratum typicum: Upper Badenian coaly clay
Descriptio: in the holotype the right valve is entire and it is relatively thick-shelled. Its beak is not pointed but reaches over the upper margin; the beak is well-developed and shifted posteriorly. The shell is covered with a dense concentric ribbing. The dental margin is well-developed and arched. The lateral teeth and dental pits are clearly visible; the cardinal (C3) tooth is weak, and gently curved. The dimensions of the right valve are the following: length — 1.6 mm; height
— 1.4 mm.
The lower margin of the left valve (belonging to another specimen) is missing. Its length is 1.7 mm. The anterior lateral tooth (p2) and the C2 cardinal tooth are strong.
Diagnosis differentialis: it is the closest to the extant P. tenuilineatum STELFOX (RICHNOVSZKY, PINTÉR 1979, GLÖER
et al. 1980) especially with respect to its outline and the position of the beak. The new form deviates in its dental apparatus. Below the C3 cardinal tooth of the right valve there is a deep dental pit. The posterior lateral tooth p1 is weaker
but longer than in P. tenuilineatum. The pits between the lateral teeth are deeper. On the left valve the cardinal teeth are
situated beneath the anterior end of the beak instead of being centrally placed. On the left valve the anterior lateral tooth
is more developed than in the extant form. The C2 cardinal tooth is longer. The beak is slightly bigger and more outstanding.
Pisidium ex gr. conventus CLESSIN, 1877
Plate XL Figures 5–8
1980 Pisidium conventus — GLÖER et al.: p. 66 (with figure)
1993 Pisidium conventus — PIECHOCKI et DYDUCH-FALNIOWSKA: Figs 102, 82c
This form is small sized (length max. 2.5 mm), thin-shelled and it has a variable outline (from rhomboidal to elliptical). Pisidium shells were encountered in the closing bed of the Várpalota Lower Badenian and in the terrestrial Middle
Badenian, showing traces of a regression. One subjuvenile valve was found in the Upper Badenian of Herend. These
shells were accompanied by other molluscs, evidently washed in fluvially. A large number of the specimens are juvenile
or subjuvenile. The shell surface is slightly and irregularly lineated, generally showing growth-lines and sometimes concentric lamellae. The beak region is characteristically flat, the dental margin is weak. The cardinal teeth are poorly-developed lathes. It is frequent that the C4 tooth of the left valve is barely visible.
It may belong to a new species.
Pisidium ex gr. nitidum JEYNS, 1832
Plate XL Figures 9–10
1943 Pisidium (Eupisidium) nitidum — SOÓS: p. 442, Pl. 30 Fig. 17
1979 Pisidium nitidum — RICHNOVSZKY et PINTÉR: p. 171, Fig. 71g
1980 Pisidium nitidum — GLÖER et al.: p. 62 (with figure)
The form has been found in the Lower Badenian of Várpalota. Its characteristic features are: more convex valves, a
more rhomboidal outline, a slightly thicker shell a more developed dental margin, and the lateral teeth are particularly
stronger than in Jeyn’s form.
It seems to be close to Pisidium nitidum, which demonstrates considerable variations with respect to several described
varieties.
Systematic descriptions
99
Pisidium ex gr. casertanum (POLI), 1791
Plate XL Figure 11, Plate XLI Figures 1–4
1979 Pisidium casertanum — RICHNOVSZKY et PINTÉR: p. 177 Fig. 75
1980 Pisidium casertanum — GLÖER et al.: p. 65
1993 Pisidium casertanum — PIECHOCKI et DYDUCH-FALNIOWSKA: p. 165 Fig. 91
Several fragments have been found in the lower part of the Lower Miocene of the Somlóvásárhely drilling. These vary
from subjuvenile to adult specimens. They are 2 mm to 4 mm long. Their outline deviates from those on Poli’s figure in
having a more angular posterior part of the shell. The character of the hinge and the teeth is very close to those on Poli’s
species. Their stage of development strongly depends on the age of the individual. This is especially true for the cardinal teeth and even more for the C4, which may be strongly outstanding. Because of the bad state of the specimens it would
be risky to base a new species on them.
Geologica Hungarica series Palaeontologica, fasciculus 56
101
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Plates
PLATES
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108
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Plate I
1.
2.
3–4.
5.
6–7.
8–9.
10.
11.
12.
13–14.
15.
Theodoxus grateloupianus (FÉR.) — Várpalota, Lower Badenian. 2.5×, p. 28.
Theodoxus grateloupianus (FÉR.) — Herend (Bánd), Upper Badenian. 6×, p. 28.
Theodoxus grateloupianus (FÉR.) — Herend (Bánd), Upper Badenian. 6×, p. 28.
Theodoxus grateloupianus (FÉR.) — Várpalota, Sarmatian. 7×, p. 28.
Theodoxus grateloupianus trigonulus nov. ssp. holotypus — MÁFI, Gy. 95., Somlóvásárhely, Lower Miocene. 7×, p. 28.
Theodoxus burdigalensis burdigalensis (D’ORB.) — Pusztamiske, Upper Badenian. 6×, p. 29.
Theodoxus burdigalensis burdigalensis (D’ORB.) — Pusztamiske, Upper Badenian. 7×, p. 29.
Theodoxus burdigalensis burdigalensis (D’ORB.) — Pusztamiske, Upper Badenian. 6×, p. 29.
Theodoxus burdigalensis burdigalensis (D’ORB.) — Herend (Bánd), Upper Badenian. 10×, p. 29.
Theodoxus burdigalensis trifasciatus (D’ORB.) — Herend, Lower Badenian. 7×, p. 29.
Theodoxus burdigalensis trifasciatus (D’ORB.) — Várpalota, Lower Badenian. 10×, p. 29.
Plates
109
110
JÓZSEF KÓKAY
Plate II
1–2.
3–6.
7–8.
9–10.
11–13.
14–15.
Theodoxus barakovici (BRUS.) — Herend, Lower Badenian. 6×, p. 29.
Theodoxus barakovici (BRUS.) — Herend, Lower Badenian. 7×, p. 29.
Theodoxus barakovici (BRUS.) opercula — Herend, Lower Badenian. 12×, p. 29.
Theodoxus cyrtocelis (KRAUSS) — Várpalota, Ottnangian. 5×, p. 30.
Theodoxus sinjanus (BRUS.) — Nagygörbő, Lower Badenian. 40×, p. 30.
Palaina martensi (ANDREAE) — Várpalota, Sarmatian. 40×, p. 30.
Plates
111
112
JÓZSEF KÓKAY
Plate III
1. Viviparus ex aff. sadleri (PARTSCH) — Nagygörbő, Lower Badenian. 4×, p. 31.
2. Cochlostoma septemspirale (RAZUM.) — Várpalota, Sarmatian. 7×, p. 31.
3. Cochlostoma septemspirale (RAZUM.) — Várpalota, Sarmatian. 14×, p. 31.
4. Viviparus pachystoma dissitus nov. ssp. holotype — MÁFI, Gy. 96., Somlóvásárhely, Lower Miocene. 1.6×, p. 31.
5. Viviparus pachystoma dissitus nov. ssp. — Somlóvásárhely, Lower Miocene. 7×, p. 31.
6. Valvata moguntina (BOETTGER) — Várpalota, Sarmatian. 20×, p. 31.
7. Valvata moguntina (BOETTGER) — Várpalota, Sarmatian. 18×, p. 31.
8. Valvata sarmatica PAPP — Várpalota, Sarmatian. 23×, p. 32.
9. Valvata moesiensis JEK. — Várpalota, Sarmatian. 23×, p. 32.
10. Valvata soceni wiesensis PAPP — Várpalota, Sarmatian. 23×, p. 32.
11–12. Valvata homalogyra BRUS. var. — Somlóvásárhely, Lower Miocene. 40×, p. 32.
Plates
113
114
JÓZSEF KÓKAY
Plate IV
1.
2.
3.
4.
5–7.
8.
9.
10–12.
13–15.
16– 17.
Pomatias bisulcatum (ZIETEN) — Várpalota, Ottnangian. 4×, p. 33.
Pomatias turonicum (WENZ) — Várpalota, Karpatian. 2.5×, p. 33.
Pomatias consobrinum (SANDB.) — Herend, Upper Badenian. 2×, p. 33.
Pomatias conica (KLEIN), silicon mould — Nyirád, Upper Badenian. 3×, p. 33.
Nematurella flexilabris SANDB. — Herend, Lower Badenian. 20×, p. 34.
Nematurella pappi SCHLICK. — Herend, Upper Badenian. 22×, p. 34.
Nematurella pappi SCHLICK. — Herend, Upper Badenian. 20×, p. 34.
Nematurella scholli SCHLICK. — Herend, Lower Badenian. 17×, p. 34.
Nematurella schuetti SCHLICK. — Pusztamiske, Upper Badenian. 23×, p. 34.
Nematurella cf. schuetti SCHLICK. — Pusztamiske, Upper Badenian. 28×, p. 34.
Plates
115
116
JÓZSEF KÓKAY
Plate V
1.
2–5.
6.
7–8.
9.
10.
11.
Nematurella ? aquensis (DEGR. et TOUZIN) — Pusztamiske, Upper Badenian. 25×, p. 35.
Nematurella aquensis (DEGR. et TOUZIN) — Pusztamiske, Upper Badenian. 30×, p. 35.
Nematurella aquensis (DEGR. et TOUZIN) — Herend, Upper Badenian. 30×, p. 35.
Nematurella aquensis (DEGR. et TOUZIN) — Pusztamiske, Upper Badenian. 30×, p. 35.
Nematurella aquensis (DEGR. et TOUZIN) — Várpalota, Sarmatian. 30×, p. 35.
Nematurella aquensis (DEGR. et TOUZIN) — Várpalota, Sarmatian. 33×, p. 35.
Nematurella aquensis (DEGR. et TOUZIN) — Pusztamiske, Upper Badenian. 600×, p. 35.
Plates
117
118
JÓZSEF KÓKAY
Plate VI
1.
2–4.
5–7.
8–10.
Sellia miocaenica nov. sp. holotypus — Somlóvásárhely, Lower Miocene. 26×, p. 35.
Sellia miocaenica nov. sp. — Somlóvásárhely, Lower Miocene. 25×, p. 35.
Sellia miocaenica nov. sp. — Somlóvásárhely, Lower Miocene. 26×, p. 35.
Sellia miocaenica nov. sp. — Somlóvásárhely, Lower Miocene. 30×, p. 35.
Plates
119
120
JÓZSEF KÓKAY
Plate VII
1–4.
5.
6.
7.
8–11.
Microprososthenia belchatovensis KAD. et PIECH. — Herend, Lower Badenian. 30×, p. 36.
Microprososthenia belchatovensis KAD. et PIECH. — Herend, Lower Badenian. 60×, p. 36.
Microprososthenia belchatovensis KAD. et PIECH. — Herend, Lower Badenian. 180×, p. 36.
Microprososthenia belchatovensis KAD. et PIECH. — Herend, Lower Badenian. 650×, p. 36.
Martinietta tumida KAD. et PIECH. — Herend, Lower Badenian. 33×, p. 36.
Plates
121
122
JÓZSEF KÓKAY
Plate VIII
1.
2.
3–7.
8.
9.
10.
11.
12.
13.
14–16.
17.
18–19.
Martinietta tumida KAD. et PIECH. — Herend, Lower Badenian. 650×, p. 36.
Bythinella eugenii JEK. — Tapolca, Upper Badenian. 33×, p. 36.
Pseudamnicola steinheimensis (MILLER) — Várpalota, Sarmatian. 25×, p. 37.
Pseudamnicola leognanensis COSSM. et PEYR. — Pusztamiske, Upper Badenian. 21×, p. 37.
Pseudamnicola leognanensis COSSM. et PEYR. — Pusztamiske, Upper Badenian. 17×, p. 37.
Pseudamnicola inflata (JEK.) var. — Pusztamiske, Upper Badenian. 24×, p. 37.
Pseudamnicola inflata (JEK.) var. — Várpalota, Sarmatian. 20×, p. 37.
Pseudamnicola (Staja) adiaphora parvula nov. ssp. holotypus — MÁFI, Gy. 99., Herend, Lower Badenian. 30×, p. 37.
“Amnicola” pseudoglobulus (D’ORB.) — Herend, Lower Badenian. 9×, p. 38.
Bithynia glabra (ZIETEN) — Pusztamiske, Sarmatian. 8×, p. 38.
Bithynia glabra (ZIETEN) — Pusztamiske, Sarmatian. 6.4×, p. 38.
Bithynia glabra (ZIETEN), silicon mould — Nyirád, Upper Badenian. 5×, p. 38.
Plates
123
124
JÓZSEF KÓKAY
Plate IX
1.
2–5.
6.
7.
8.
9.
10.
11.
12.
13.
Bithynia glabra (ZIETEN) — Nyirád, Upper Badenian. 8×, p. 38.
Bithynia glabra (ZIETEN) opercula — Nyirád, Upper Badenian. 18×, p. 38.
Bithynia glabra (ZIETEN) opercula — Nyirád, Upper Badenian. 15×, p. 38.
Bithynia glabra (ZIETEN) opercula — Nyirád, Upper Badenian. 18×, p. 38.
Bithynia glabra (ZIETEN) opercula — Nyirád, Upper Badenian. 23×, p. 38.
Bithynia glabra subgracilis (LOMN.) — Nagygörbő, Lower Badenian. 7×, p. 38.
Bithynia glabra subgracilis (LOMN.) — Nagygörbő, Lower Badenian. 4.5×, p. 38.
Bithynia glabra subgracilis (LOMN.) — Nagygörbő, Lower Badenian. 5.2×, p. 38.
Bithynia glabra subgracilis (LOMN.) opercula — Nagygörbő, Lower Badenian. 14×, p. 38.
Bithynia dunkeri GUDE — Nagygörbő, Lower Badenian. 5×, p. 39.
Plates
125
126
JÓZSEF KÓKAY
Plate X
1. Ferebithynia vadaszi (WENZ) — Várpalota, Upper Badenian. 6.5×, p. 40.
2. Ferebithynia vadaszi (WENZ) — Várpalota, Upper Badenian. 5.2×, p. 40.
3. Ferebithynia vadaszi (WENZ) — Várpalota, Upper Badenian. 5.4×, p. 40.
4. Ferebithynia vadaszi (WENZ) — Várpalota, Upper Badenian. 5.5×, p. 40.
5. Ferebithynia vadaszi (WENZ) — Várpalota, Upper Badenian. 5.8×, p. 40.
6. Ferebithynia vadaszi (WENZ) — Várpalota, Upper Badenian. 12×, p. 40.
7. Ferebithynia vadaszi (WENZ) opercula — Várpalota, Upper Badenian. 20×, p. 40.
8. Schuettemmericia paulensis (DEGR. et TOUZIN) — Herend, Lower Badenian. 24×, p. 40.
9–10.Schuettemmericia paulensis (DEGR. et TOUZIN) — Várpalota, Sarmatian. 23×, p. 40.
Plates
127
128
JÓZSEF KÓKAY
Plate XI
1–2.
3–4.
5–6.
7.
8.
9.
10–11.
Schuettemmericia paulensis (DEGR. et TOUZIN) — Várpalota, Sarmatian. 21×, p. 40.
Stalioa globulina nov. sp. holotypus — MÁFI, Gy. 98., Pusztamiske, Upper Badenian. 28×, p. 41.
Stalioa globulina nov. sp. — Pusztamiske, Upper Badenian. 30×, p. 41.
Stalioa globulina nov. sp. — Pusztamiske, Upper Badenian. 37×, p. 41.
Stalioa globulina nov. sp. — Pusztamiske, Upper Badenian. 31×, p. 41.
Stalioa globulina nov. sp. — Pusztamiske, Upper Badenian. 30×, p. 41.
Stalioa globulina nov. sp. — Pusztamiske, Upper Badenian. 24×, p. 41.
Plates
129
130
JÓZSEF KÓKAY
Plate XII
1.
2.
3.
4.
5–6.
7.
8.
9.
10.
11–12.
Stalioa globulina nov. sp. — Pusztamiske, Upper Badenian. 10.3×, p. 41.
Stalioa globulina nov. sp. apex — Pusztamiske, Upper Badenian. 650×, p. 41.
Stalioa praeglobulina nov. sp. holotypus — MÁFI, Gy. 97., Somlóvásárhely, Lower Miocene. 40×, p. 41.
Stalioa praeglobulina nov. sp. holotypus — MÁFI, Gy. 97., Somlóvásárhely, Lower Miocene. 44×, p. 41.
Stalioa praeglobulina nov. sp. — Somlóvásárhely, Lower Miocene. 40×, p. 41.
Stalioa praeglobulina nov. sp. — Somlóvásárhely, Lower Miocene. 30×, p. 41.
Stalioa praeglobulina nov. sp. — Somlóvásárhely, Lower Miocene. 37×, p. 41.
Stalioa praeglobulina nov. sp. — Somlóvásárhely, Lower Miocene. 33×, p. 41.
Stalioa praeglobulina nov. sp. — Somlóvásárhely, Lower Miocene. 37×, p. 41.
Stalioa praeglobulina nov. sp. — Somlóvásárhely, Lower Miocene. 42×, p. 41.
Plates
131
132
JÓZSEF KÓKAY
Plate XIII
1.
2.
3.
4–5.
6.
7–8.
9–10.
11–12.
13–14.
Stalioa praeglobulina nov. sp. apex — Somlóvásárhely, Lower Miocene. 200×, p. 41.
Stalioa praeglobulina nov. sp. apex — Somlóvásárhely, Lower Miocene. 600×, p. 41.
Staliopsis moravicus RZEHAK — Várpalota, Ottnangian. 13×, p. 42.
Staliopsis puisseguri (SCHLICK.) — Somlóvásárhely, Lower Miocene. 15×, p. 42.
Staliopsis puisseguri (SCHLICK.) — Somlóvásárhely, Lower Miocene. 13×, p. 42.
Sandbergerina succineiformis (SANDBERGER) var. — Pusztamiske, Upper Badenian. 19×, p. 42.
Sandbergerina succineiformis (SANDBERGER) var. — Várpalota, Sarmatian. 18×, p. 42.
Sandbergerina succineiformis (SANDBERGERINA) var. — Pusztamiske, Upper Badenian. 19×, p. 42.
Staadtiellopsis rubeschi (REUSS) var. — Pusztamiske, Upper Badenian. 18×, p. 43.
Plates
133
134
JÓZSEF KÓKAY
Plate XIV
1–2.
3–4.
5.
6.
7.
8–9.
10–11.
12–13.
14.
Staadtiellopsis bandensis nov. sp. holotypus — MÁFI, Gy. 100., Herend (Bánd), Upper Badenian. 19×, p. 43.
Platyla callosa (BOETTGER) — Somlóvásárhely, Lower Miocene. 19×, p. 44.
Platyla subfusca (FLACH) — Somlóvásárhely, Lower Miocene. 27×, p. 44.
Platyla falkneri B.-G.-S. — Herend, Upper Badenian. 19×, p. 45.
Platyla polita (HARTMAN) — Várpalota, Sarmatian. 19×, p. 45.
Brotia escheri turrita (BRONGT.) — Várpalota, Lower Badenian. 2×, p. 45.
Brotia escheri turrita (BRONGT.) — Herend, Lower Badenian. 1.5×, p. 45.
Brotia escheri turrita (KLEIN) — Pusztamiske, Upper Badenian. 2.3×, p. 45.
Brotia escheri turrita (KLEIN) — Tapolca, Upper Badenian. 0.7×, p. 45.
Plates
135
136
JÓZSEF KÓKAY
Plate XV
1.
2.
3.
4.
5–7.
8.
9.
10.
11.
12.
13.
14.
Melanoides procurvicosta nov. sp. holotypus — MÁFI, Gy. 101., Nagygörbő, Lower Badenian. 1.7×, p. 46.
Melanoides procurvicosta nov. sp. — Nagygörbő, Lower Badenian. 2.4×, p. 46.
Melanoides procurvicosta nov. sp. — Nagygörbő, Lower Badenian. 6×, p. 46.
Melanoides procurvicosta nov. sp. — Nagygörbő, Lower Badenian. 2×, p. 46.
Melanopsis visianiana BRUSINA — Nagygörbő, Lower Badenian. 4.2×, p. 46.
Melanopsis cf. fritzei curta LOCARD — Nagygörbő, Lower Badenian. 15×, p. 47.
Carychium nouleti nouleti BOURG. — Herend, Lower Badenian. 36×, p. 47.
Carychium nouleti gibbum SANDB. — Pusztamiske, Upper Badenian. 30×, p. 47.
Carychium nouleti gibbum SANDB. — Pusztamiske, Upper Badenian. 32×, p. 47.
Carychium sandbergeri HANDM. — Várpalota, Sarmatian. 28×, p. 47.
Carychium sandbergeri HANDM. — Várpalota, Sarmatian. 25×, p. 47.
Carychium suevicum BOETTGER — Nyirád, Upper Badenian. 28×, p. 48.
Plates
137
138
JÓZSEF KÓKAY
Plate XVI
1.
2–4.
5.
6.
7.
8.
9.
10–11.
12.
13.
14–16.
Carychium suevicum BOETTGER — Nyirád, Upper Badenian. 30×, p. 48.
Carychium pachychilus SANDB. — Várpalota, Sarmatian. 24×, p. 48.
Carychium eumicron eumicron BOURG. — Várpalota, Sarmatian. 40×, p. 48.
Carychium eumicron peneckei FLACH — Herend, Lower Badenian. 40×, p. 48.
Carychium eumicron unidentata nov. sp. holotypus — MÁFI, Gy. 102., Somlóvásárhely, Lower Miocene. 34×, p. 48.
Carychium eumicron unidentata nov. sp. — Somlóvásárhely, Lower Miocene. 40×, p. 48.
Laemodonta bourgeoisi TOURN. var. — Várpalota, Ottnangian. 10×, p. 49.
Auriculastra (Conauricula) badeniensis nov. sp. holotypus — MÁFI, Gy. 103., Pusztamiske, Upper Badenian. 18×, p. 50.
Auriculastra (Conauricula) badeniensis nov. sp. — Pusztamiske, Upper Badenian. 22×, p. 50.
Auriculastra (Conauricula) badeniensis nov. sp. — Pusztamiske, Upper Badenian. 18×, p. 50.
Galba dupuyana (NOULET) — Tapolca, Upper Badenian. 10×, p. 50.
Plates
139
140
JÓZSEF KÓKAY
Plate XVII
1.
2.
3.
4–6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19–21.
Stagnicola armaniacensis (NOULET) — Tapolca, Upper Badenian. 3.5×, p. 50.
Stagnicola armaniacensis palustriformis (GOTT.), silicon mould — Pusztamiske, Upper Badenian. 0.8×, p. 51.
Stagnicola armaniacensis palustriformis (GOTT.), silicon mould — Pusztamiske, Upper Badenian. 3.5×, p. 51.
Stagnicola praebouilleti SCHLICK., silicon mould — Nyirád, Upper Badenian. 4×, p. 51.
Stagnicola praebouilleti SCHLICK., silicon mould — Nyirád, Upper Badenian. 3×, p. 51.
Stagnicola praebouilleti SCHLICK., silicon mould — Nyirád, Upper Badenian. 2×, p. 51.
Stagnicola kreutzi (LOMN.) — Várpalota, Sarmatian. 5×, p. 51.
Stagnicola palustris (O. F. MÜLLER), silicon mould — Pusztamiske, Upper Badenian. 4.2×, p. 51.
Radix socialis praelongata (GOTT. et WENZ), silicon mould — Nyirád, Upper Badenian. 1.8×, p. 52.
Radix socialis regularis (MAILLARD), silicon mould — Nyirád, Upper Badenian. 4.5×, p. 52.
Radix socialis regularis (MAILLARD), silicon mould — Nyirád, Upper Badenian. 2.2×, p. 52.
Radix dilatata (NOULET) — Várpalota, Sarmatian. 1.4×, p. 52.
Radix hyaloleuca (BRUSINA) — Várpalota, Sarmatian. 8×, p. 52.
Lymnaea turrita (KLEIN) — Tapolca, Upper Badenian. 8×, p. 53.
Lymnaea stagnalis sarmatica nov. ssp. holotypus — MÁFI, Gy. 104., Várpalota, Sarmatian. 1.3×, p. 53.
Anisus dupuyanus (NOULET) — Pusztamiske, Upper Badenian. 11×, p. 53.
Anisus dupuyanus omalus (BOURG.) — Tapolca, Upper Badenian. 12×, p. 54.
Plates
141
142
JÓZSEF KÓKAY
Plate XVIII
1–2.
3–4.
5.
6.
7–10.
11–13.
14–16.
Anisus rousianus (NOULET) — Várpalota, Sarmatian. 11×, p. 54.
Anisus rousianus (NOULET) — Pusztamiske, Upper Badenian. 17×, p. 54.
Anisus hilgendorfi subcarinata (GOTT.) — Várpalota, Sarmatian. 4×, p. 54.
Anisus hilgendorfi subcarinata (GOTT.) — Várpalota, Sarmatian. 6×, p. 54.
Gyraulus alienus (ROLLE) — Pusztamiske, Upper Badenian. 24×, p. 54.
Gyraulus pulici (BRUSINA) — Várpalota, Sarmatian. 16×, p. 55.
Gyraulus pulici (BRUSINA) — Várpalota, Sarmatian. 16×, p. 55.
Plates
143
144
JÓZSEF KÓKAY
Plate XIX
1–3.
4.
5.
6–7.
8–10.
11.
12.
13.
14.
15.
16–17.
18.
19–20.
Gyraulus nedici (BRUSINA) — Nyirád, Sarmatian. 20×, p. 55.
Gyraulus microstatus (BOURG.) — Pusztamiske, Upper Badenian. 18×, p. 55.
Gyraulus microstatus (BOURG.) — Pusztamiske, Upper Badenian. 22×, p. 55.
Gyraulus amiculus (RZEHAK) — Várpalota, Lower Badenian. 27×, p. 55.
Gyraulus callistus (BOURG.) — Pusztamiske, Upper Badenian. 20×, p. 56.
Gyraulus callistus (BOURG.) — Pusztamiske, Upper Badenian. 22×, p. 56.
Gyraulus verticillus (BRUS.) var. — Nagygörbő, Lower Badenian. 28×, p. 56.
Gyraulus applanatus (THOMAE) — Somlóvásárhely, Lower Miocene. 18×, p. 56.
Gyraulus applanatus (THOMAE) — Somlóvásárhely, Lower Miocene. 23×, p. 56.
Gyraulus trochiformis dealbatus (BRAUN) — Herend, Lower Badenian. 18×, p. 57.
Gyraulus trochiformis kleini GOTT. et WENZ — Várpalota, Sarmatian. 17×, p. 57.
Gyraulus goussardianus (NOULET) — Pusztamiske, Upper Badenian. 13×, p. 57.
Gyraulus goussardianus (NOULET) — Pusztamiske, Upper Badenian. 20×, p. 57.
Plates
145
146
JÓZSEF KÓKAY
Plate XX
1.
2.
3–4.
5.
6.
7.
8–9.
10–12.
13.
14.
15.
Gyraulus homalosomus (BRUSINA) — Várpalota, Sarmatian. 16×, p. 57.
Hippeutis fasciata GOTTSCHICK — Várpalota, Sarmatian. 23×, p. 58.
Hippeutis fasciata GOTTSCHICK — Várpalota, Sarmatian. 20×, p. 58.
Segmentina larteti (NOULET) — Várpalota, Sarmatian. 10×, p. 58.
Planorbarius cornu cornu (BRONGT.) — Nagygörbő, Lower Badenian. 3.8×, p. 58.
Planorbarius cornu mantelli (DUNKER) — Várpalota, Sarmatian. 1.5×, p. 59.
Planorbarius sansaniensis (NOULET) — Tapolca, Upper Badenian. 3×, p. 59.
Planorbarius sansaniensis (NOULET) — Várpalota, Sarmatian. 2×, p. 59.
Planorbarius sansaniensis (NOULET) — Nyirád, Upper Badenian. 3.5×, p. 59.
Ferrissia wittmanni SCHLICK. — Várpalota, Upper Badenian. 4.6×, p. 59.
Ferrissia deperdita (DESM.) — Várpalota, Upper Badenian. 7.7×, p. 60.
Plates
147
148
JÓZSEF KÓKAY
Plate XXI
1.
2.
3.
4.
5.
6.
7–8.
9.
10.
11.
12.
13.
14.
15.
Ferrissia deperdita (DESM.) — Somlóvásárhely, Lower Miocene. 13×, p. 60.
Ancylus moravicus ŘZEHAK (forma „obtusa”) — Somlóvásárhely, Lower Miocene. 18×, p. 60.
Ancylus moravicus ŘZEHAK (forma „obtusa”) — Somlóvásárhely, Lower Miocene. 23×, p. 60.
Ancylus moravicus ŘZEHAK — Várpalota, Upper Badenian. 6.5×, p. 60.
Pseudancylastrum deperditolacustris (GOT.) — Herend, Lower Badenian. 25×, p. 60.
Pseudancylastrum decorata nov. sp. holotypus — MÁFI, Gy. 105., Várpalota, Sarmatian. 16×, p. 60.
Pseudancylastrum decorata nov. sp. — Várpalota, Sarmatian. 12×, p. 60.
Azeca cf. peneckei ANDREAE — Herend, Upper Badenian. 25×, p. 61.
Cochlostoma subrimata loxostoma (KLEIN) — Várpalota, Sarmatian. 12×, p. 61.
Cochlostoma subrimata cf. procera GOTT. — Herend, Upper Badenian. 28×, p. 61.
Negulus suturalis suturalis (SANDB.) — Somlóvásárhely, Lower Miocene. 23×, p. 61.
Negulus suturalis suturalis (SANDB.) — Várpalota, Sarmatian. 26×, p. 61.
Negulus suturalis gracilis GOTT. et WENZ — Várpalota, Sarmatian. 28×, p. 62.
Acmopupa ? nov. sp. — Somlóvásárhely, Lower Miocene. 24×, p. 62.
Plates
149
150
JÓZSEF KÓKAY
Plate XXII
1.
2.
3.
4.
5.
6–7.
8.
9.
10.
Vertigo kochi BOETTGER var. — Pusztamiske, Upper Badenian. 31×, p. 62.
Vertigo kochi BOETTGER var. — Pusztamiske, Upper Badenian. 27×, p. 62.
Vertigo kochi BOETTGER var. — Pusztamiske, Upper Badenian. 60×, p. 62.
Vertigo kochi BOETTGER var. — Pusztamiske, Upper Badenian. 29×, p. 62.
Vertigo protracta suevica GOTT. et WENZ — Tapolca, Upper Badenian. 31×, p. 63.
Vertigo protracta suevica GOTT. et WENZ — Várpalota, Sarmatian. 31×, p. 63.
Vertigo callosa (REUSS) — Somlóvásárhely, Lower Miocene. 26×, p. 63.
Vertigo callosa (REUSS) — Várpalota, Sarmatian. 30×, p. 63.
Vertigo callosa (REUSS) — Várpalota, Sarmatian. 25×, p. 63.
Plates
151
152
JÓZSEF KÓKAY
Plate XXIII
1.
2–3.
4.
5.
6–7.
8.
9.
Vertigo callosa perarmata GOTT. et WENZ — Várpalota, Sarmatian. 35×, p. 63.
Vertigo callosa diversidens (SANDB.) — Várpalota, Sarmatian. 40×, p. 63.
Vertigo callosa convergens BOETTGER — Tapolca, Upper Badenian. 37×, p. 63.
Vertigo callosa cardiostoma (SANDB.) — Pusztamiske, Upper Badenian. 35×, p. 63.
Vertigo callosa perarmata GOTT. et WENZ — Várpalota, Sarmatian. 41×, p. 63.
Vertigo callosa perarmata GOTT. et WENZ — Várpalota, Sarmatian. 36×, p. 63.
Vertigo callosa diversidens GOTT. et WENZ — Pusztamiske, Upper Badenian. 33×, p. 63.
Plates
153
154
JÓZSEF KÓKAY
Plate XXIV
1.
2.
3.
4–5.
6.
7.
8.
9.
10.
Vertigo ovatula trolli WENZ — Tapolca, Upper Badenian. 27×, p. 64.
Vertigo ovatula trolli WENZ — Várpalota, Sarmatian. 24×, p. 64.
Vertigo ovatula miliiformis BOETTGER — Tapolca, Upper Badenian. 26×, p. 64.
Vertigo bakonyensis nov. sp. holotypus — MÁFI, Gy. 107., Pusztamiske, Sarmatian. 33×, p. 64.
Vertigo pusilla sarmatica nov. ssp. holotypus — MÁFI, Gy. 106., Várpalota, Sarmatian. 25×, p. 64.
Gastrocopta acuminata acuminata (KLEIN) — Tapolca, Upper Badenian. 25×, p. 65.
Gastrocopta acuminata procera (GOTT. et WENZ) — Várpalota, Sarmatian. 31×, p. 65.
Gastrocopta nouletiana nouletiana (DUPUY) — Várpalota, Sarmatian. 24×, p. 66.
Gastrocopta nouletiana nouletiana (DUPUY) — Várpalota, Sarmatian. 27×, p. 66.
Plates
155
156
JÓZSEF KÓKAY
Plate XXV
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11–12.
13.
14.
15.
Gastrocopta nouletiana nouletiana (DUPUY) — Várpalota, Sarmatian. 24×, p. 66.
Gastrocopta nouletiana tapeina (BOURG.) — Várpalota, Sarmatian. 26×, p. 66.
Gastrocopta nouletiana gracilidens (SANDB.) — Pusztamiske, Upper Badenian. 40×, p. 66.
Gastrocopta nouletiana gracilidens (SANDB.) — Várpalota, Sarmatian. 23×, p. 66.
Gastrocopta ferdinandi (ANDREAE) — Várpalota, Sarmatian. 18×, p. 66.
Leiostyla cf. gottschicki (WENZ) juv. — Várpalota, Sarmatian. 20×, p. 67.
Pupilla iratiana suevica GOTT. — Várpalota, Sarmatian. 20×, p. 67.
Argna oppoliensis (ANDREAE) — Várpalota, Sarmatian. 20×, p. 67.
Argna suemeghyi (BARTHA) — Várpalota, Sarmatian. 20×, p. 68.
Argna reperta (SANDB.) — Várpalota, Sarmatian. 20×, p. 68.
Argna sublamellata nov. sp. holotypus — MÁFI, Gy. 108., Várpalota, Sarmatian. 34×, p. 68.
Enneopupa subcylindrella nov. sp. holotypus — MÁFI, Gy. 109., Pusztamiske, Upper Badenian. 19×, p. 69.
Paracoryna pseudoennea (FLACH) — Várpalota, Sarmatian. 23×, p. 69.
Acanthinula tuchoricensis (KLIKA) — Somlóvásárhely, Lower Miocene. 40×, p. 70.
Plates
157
158
JÓZSEF KÓKAY
Plate XXVI
1.
2.
3–4.
5–6.
7–8.
9–10.
11–13.
14–15.
16–18.
Acanthinula trochulus (SANDB.) — Várpalota, Sarmatian. 13×, p. 70.
Spermodea candida FALKNER — Várpalota, Sarmatian. 25×, p. 70.
Planogyra nana BRAUN — Somlóvásárhely, Lower Miocene. 30×, p. 71.
Strobilops fischeri WENZ — Somlóvásárhely, Lower Miocene. 18×, p. 71.
Strobilops uniplicata plana (CLESSIN) — Tapolca, Upper Badenian. 21×, p. 71.
Strobilops subconoidea (JOOSS) — Várpalota, Sarmatian. 33×, p. 71.
Strobilops costata (CLESSIN) — Várpalota, Sarmatian. 20×, p. 72.
Strobilops costata bilamellata (CLESSIN) — Várpalota, Sarmatian. 20×, p. 72.
Strobilops sandbergeri (LOMN.) — Várpalota, Sarmatian. 20×, p. 72.
Plates
159
160
JÓZSEF KÓKAY
Plate XXVII
1–3.
4–5.
6.
7.
8.
9.
10.
11.
12.
13.
14–16.
Strobilops pappi SCHLICK. — Várpalota, Sarmatian. 20×, p. 72.
Strobilops tiarula (SANDB.) — Tapolca, Upper Badenian. 20×, p. 73.
Strobilops sp. — Várpalota, Sarmatian. 32×, p. 73.
Strobilops sp. — Várpalota, Sarmatian. 28×, p. 73.
Mastus pupa sarmatica PAPP — Várpalota, Sarmatian. 4×, p. 73.
Ena sp. — Somlóvásárhely, Lower Miocene. 13×, p. 73.
Napaeus complanatus (REUSS) — Somlóvásárhely, Lower Miocene. 18×, p. 73.
Napaeus complanatus (REUSS) — Somlóvásárhely, Lower Miocene. 20×, p. 73.
Napaeus complanatus (REUSS) — Somlóvásárhely, Lower Miocene. 5×, p. 73.
Succinea minima KLEIN — Várpalota, Sarmatian. 10×, p. 74.
Helicodiscus subteres (CLESSIN) — Tapolca, Upper Badenian. 20×, p. 74.
Plates
161
162
JÓZSEF KÓKAY
Plate XXVIII
1.
2.
3.
4.
5.
6–7.
8–9.
10.
11.
12.
13–14.
15.
Discus euglyphoides (SAND.) — Somlóvásárhely, Lower Miocene. 6.7×, p. 74.
Discus euglyphoides (SAND.) — Somlóvásárhely, Lower Miocene. 5×, p. 74.
Discus pleuradrus (BOURG.) — Somlóvásárhely, Lower Miocene. 20×, p. 75.
Discus pleuradrus (BOURG.) — Somlóvásárhely, Lower Miocene. 14×, p. 75.
Semilimax intermedia (REUSS) — Várpalota, Sarmatian. 25×, p. 75.
Semilimax intermedia crassitesta (ANDREAE) — Pusztamiske, Sarmatian. ×12, p. 75.
Vitrea procrystallina (ANDREAE) — Tapolca, Upper Badenian. 34×, p. 75.
Vitrea procrystallina steinheimensis GOTT. — Várpalota, Sarmatian. 27×, p. 76.
Vitrea procrystallina steinheimensis GOTT. — Várpalota, Sarmatian. 24×, p. 76.
Vitrea procrystallina steinheimensis GOTT. — Várpalota, Sarmatian. 20×, p. 76.
Nesovitrea boettgeriana (CLESS.) — Somlóvásárhely, Lower Miocene. 26×, p. 76.
Nesovitrea boettgeriana (CLESS.) — Somlóvásárhely, Lower Miocene. 35×, p. 76.
Plates
163
164
JÓZSEF KÓKAY
Plate XXIX
1–3.
4–5.
6–7.
8–9.
10.
11.
12.
13–15.
Nesovitrea mendica (SLAVIK) — Herend, Upper Badenian. 19×, p. 76.
Nesovitrea mendica (SLAVIK) — Várpalota, Upper Badenian. 20×, p. 76.
Nesovitrea subhammonis (GOTT.) — Tapolca, Upper Badenian. 26×, p. 76.
Nesovitrea subhammonis (GOTT.) — Tapolca, Upper Badenian. 20×, p. 76.
Aegopis algiroides (REUSS) — Somlóvásárhely, Lower Miocene. 11×, p. 76.
Aegopis costatus (SANDB.) — Nagygörbő, Lower Badenian. 9×, p. 77.
Aegopis costatus (SANDB.) — Pusztamiske, Upper Badenian. 2×, p. 77.
Retinella applanata nov. sp. holotypus — MÁFI, Gy. 110., Várpalota, Sarmatian. 15×, p. 77.
Plates
165
166
JÓZSEF KÓKAY
Plate XXX
1–2.
3.
4.
5.
6.
7–8.
9–11.
12.
13–15.
16.
17–18.
19.
20.
Aegopinella denudata (REUSS) var. — Somlóvásárhely, Lower Miocene. 1.7×, p. 77.
Daudebardia praecursor ANDREAE — Várpalota, Sarmatian. 16×, p. 78.
Daudebardia praecursor ANDREAE — Várpalota, Sarmatian. 22×, p. 78.
Zonitoides schraieri SCHLICK. var. — Várpalota, Sarmatian. 14×, p. 78.
Zonitoides schraieri SCHLICK. var. — Várpalota, Sarmatian. 13×, p. 78.
Zonitoides suevicus (JOOSS) var. — Herend, Upper Badenian. 18×, p. 78.
Zonitoides suevicus (JOOSS) — Várpalota, Sarmatian. 25×, p. 78.
Janulus supracostatus (SANDB.) — Várpalota, Sarmatian. 6×, p. 78.
Janulus sp. — Várpalota, Sarmatian. 9×, p. 79.
Opeas minutum (KLEIN) — Herend, Upper Badenian. 8×, p. 80.
Fortuna tertia SCHLICK. et STRAUCH — Várpalota, Sarmatian. 16×, p. 80.
Fortuna tertia SCHLICK. et STRAUCH — Várpalota, Sarmatian. 14×, p. 80.
Fortuna varpalotensis nov. sp. holotypus — MÁFI, Gy. 111., Várpalota, Sarmatian. 12×, p. 80.
Plates
167
168
JÓZSEF KÓKAY
Plate XXXI
1.
2.
3.
4.
5.
6.
7–8.
9.
10.
11.
12–13.
14–16.
17.
18–19.
20.
Triptychia vulgata (REUSS) — Várpalota, Ottnangian. 4.6×, p. 81.
Triptychia grandis (KLEIN) — Somlóvásárhely, Lower Badenian. 2.2×, p. 81.
Triptychia grandis (KLEIN) — Somlóvásárhely, Lower Badenian. 6.5×, p. 81.
Triptychia grandis (KLEIN) — Somlóvásárhely, Lower Miocene. 5.5×, p. 81.
Triptychia cf. suturalis (SANDB.) — Pusztamiske, Sarmatian. 2.5×, p. 81.
Triptychia cf. suturalis (SANDB.) — Pusztamiske, Sarmatian. 10×, p. 81.
Triptychia leobersdorfensis sarmatica nov. sp. holotypus — MÁFI, Gy. 112., Várpalota, Sarmatian. 1.7×, p. 82.
Triptychia leobersdorfensis sarmatica nov. sp. — Várpalota, Sarmatian. 1.7×, p. 82.
Nordsieckia pontica LUEGER — Várpalota, Sarmatian. 12×, p. 82.
Nordsieckia pontica LUEGER — Várpalota, Sarmatian. 17×, p. 82.
Cochlodina varpalotensis nov. sp. holotypus — MÁFI, Gy. 113., Várpalota, Sarmatian. 12×, p. 82.
Cochlodina varpalotensis nov. sp. — Várpalota, Sarmatian. 12×, p. 82.
Cochlodina varpalotensis nov. sp. — Várpalota, Sarmatian. 14×, p. 82.
Cochlodina varpalotensis nov. sp. “clausilium” — Várpalota, Sarmatian. 22× , p. 82.
Pseudidyla boettgeri NORDSIECK — Várpalota, Sarmatian. 23×, p. 83.
Plates
169
170
JÓZSEF KÓKAY
Plate XXXII
1–2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
Pseudidyla tricarinata nov. sp. holotypus — MÁFI, Gy. 114., Várpalota, Sarmatian. 14×, p. 84.
Pseudidyla tricarinata nov. sp. — Várpalota, Sarmatian. 22×, p. 84.
Mactrogastra cf. voesendorfensis (PAPP et THENIUS) — Várpalota, Sarmatian. 25×, p. 84.
Mactrogastra cf. voesendorfensis (PAPP et THENIUS) — Várpalota, Sarmatian. 27×, p. 84.
Pseudoleacina producta (REUSS) — Somlóvásárhely, Lower Miocene. 2.6×, p. 85.
Pseudoleacina producta (REUSS) — Somlóvásárhely, Lower Miocene. 12×, p. 85.
Palaeoglandina gracilis porrecta (GOBANZ) — Várpalota, Sarmatian. 2×, p. 85.
Palaeoglandina gracilis porrecta (GOBANZ) — Várpalota, Sarmatian. 8×, p. 85.
Poiretia taurinensis (SACCO) — Várpalota, Sarmatian. 1.7×, p. 86.
Poiretia ex aff. taurinensis (SACCO) — Várpalota, Sarmatian. 1.7×, p. 86.
Pseudoleacina rakosdensis (GAÁL) — Várpalota, Sarmatian. 15×, p. 85.
Pseudoleacina rakosdensis (GAÁL) — Várpalota, Sarmatian. 7×, p. 85.
Plates
171
172
JÓZSEF KÓKAY
Plate XXXIII
1.
2.
3.
4–5.
6.
7.
8.
9–10.
11.
Perforatella homalospira (REUSS) var. — Somlóvásárhely, Lower Miocene. 3×, p. 86.
Perforatella punctigera (THOMAE) — Várpalota, Sarmatian. 2.3×, p. 87.
Caracollina phacodes barreri (BOURG.) — Várpalota, Sarmatian. 11×, p. 87.
Canariella bakonyensis nov. sp. holotypus — MÁFI, Gy. 115., Várpalota, Sarmatian. 3×, p. 87.
Canariella bakonyensis nov. sp. — Várpalota, Sarmatian. 10×, p. 87.
Canariella nov. sp. — Somlóvásárhely, Lower Miocene. 9×, p. 87.
Canariella nov. sp. — Somlóvásárhely, Lower Miocene. 18×, p. 87.
Galactochilus mattiacum (STEININGER) — Somlóvásárhely, Lower Miocene. 4.8×, p. 88.
Galactochilus mattiacum (STEININGER) — Somlóvásárhely, Lower Miocene. 5×, p. 88.
Plates
173
174
JÓZSEF KÓKAY
Plate XXXIV
1–2.
3–4.
5–6.
7.
8–9.
10.
11.
12, 14.
13.
Galactochilus sarmaticum GAÁL — Várpalota, Sarmatian. 1×, p. 88.
Galactochilus leobersdorfensis (TROLL) — Várpalota, Sarmatian. 1×, p. 88.
Tropidomphalus robustus (REUSS) — Somlóvásárhely, Lower Miocene. 1.7×, p. 89.
Tropidomphalus extinctus (RAMBUR) — Somlóvásárhely, Lower Miocene. 3.4×, p. 89.
Tropidomphalus gigas PFEFFER — Herend (Bánd), Upper Badenian. 1.6×, p. 89.
Tropidomphalus zelli depressus WENZ — Várpalota, Sarmatian. 1.4×, p. 90.
Tropidomphalus zelli depressus WENZ — Várpalota, Sarmatian. 1.3×, p. 90.
Tropidomphalus cf. incrassatus (KLEIN) — Pusztamiske, Upper Badenian. 10×, p. 90.
Tropidomphalus cf. incrassatus (KLEIN) — Pusztamiske, Upper Badenian. 5.5×, p. 90.
Plates
175
176
JÓZSEF KÓKAY
Plate XXXV
1.
2–3.
4–7.
8.
9.
10.
11–15.
16.
17.
Helicigona wenzi SOÓS — Várpalota, Sarmatian. 3×, p. 90.
Klikia osculum tennis (KLIKA) — Somlóvásárhely, Lower Miocene. 4×, p. 91.
Klikia osculum tennis (KLIKA) — Somlóvásárhely, Lower Miocene. 4.5×, p. 91.
Klikia ex gr. osculum (THOMAE) — Somlóvásárhely, Lower Miocene. 7×, p. 91.
Klikia giengensis (KLEIN) — Várpalota, Sarmatian. 4.3×, p. 91.
Klikia kaeufeli WENZ — Várpalota, Sarmatian. 4×, p. 92.
Klikia cf. devexa (REUSS) — Somlóvásárhely, Lower Miocene. 2.5×, p. 92.
Klikia cf. coarctata steinheimensis JOOSS — Várpalota, Sarmatian. 3.5×, p. 93.
Klikia goniostoma (SANDB.) — Várpalota, Sarmatian. 2.3×, p. 92.
Plates
177
178
JÓZSEF KÓKAY
Plate XXXVI
1.
2–3.
4.
5.
6.
7.
8–10.
Cepaea bohemica (BOETTGER) — Somlóvásárhely, Lower Miocene. 2.8×, p. 93.
Cepaea silvana (KLEIN) — Pusztamiske, Upper Badenian. 2.2×, p. 93.
Dreissena dobrei (BRUS.) — Várpalota, Sarmatian. 6×, p. 94.
Congeria boeckhi WENZ — Várpalota, Upper Badenian. 1.8×, p. 94.
Congeria venusta KOCH.-DEVIDÉ var. — Várpalota, Upper Badenian. 3.3×, p. 94.
Congeria hercegovinensis KOCH.-DEVIDÉ — Várpalota, Upper Badenian. 2.4×, p. 94.
Margaritifera flabellata (GOLDFUSS) — Várpalota, Lower Badenian. 1.8×, p. 95.
Plates
179
180
JÓZSEF KÓKAY
Plate XXXVII
1. Margaritifera flabellata bavarica MODELL — Herend, Lower Badenian. 1.1×, p. 95.
2. Unio jaccardi LOCARD — Várpalota, Upper Badenian. 1.3×, p. 96.
3. Unio eseri KRAUSS — Várpalota, Lower Badenian. 1.2×, p. 95.
4. Unio larteti NOULET — Nagygörbő, Lower Badenian. 3.2×, p. 96.
5. Unio larteti NOULET — Nagygörbő, Lower Badenian. 1.4×, p. 96.
Plates
181
182
JÓZSEF KÓKAY
Plate XXXVIII
1.
2–3.
4–5.
6–8.
Unio hinschi MODELL — Várpalota, Upper Badenian. 0.9×, p. 96.
Unio lorioli LOCARD — Várpalota, Lower Badenian. 2×, p. 96.
Pisidium bellardii BRUS. — Herend, Upper Badenian. 2.1×, p. 97.
Pisidium pseudosphaerium SCHLESCH var. — Várpalota, Sarmatian. 22×, p. 97.
Plates
183
184
JÓZSEF KÓKAY
Plate XXXIX
1–4, 6. Pisidium pseudosphaerium SCHLESCH var. — Várpalota, Sarmatian. 23×, p. 97.
5. Pisidium steinheimense GOTT. — Várpalota, Sarmatian. 24×, p. 97.
7. Pisidium annandalei (PRASHAD) — Herend (Bánd), Upper Badenian. 30×, p. 98.
Plates
185
186
JÓZSEF KÓKAY
Plate XL
1–2.
3–4.
5–8.
9–10.
11.
Pisidium bakonyensis nov. sp. holotypus — MÁFI, Gy. 116., Pusztamiske, Upper Badenian. 27×, p. 98.
Pisidium bakonyensis nov. sp. — Pusztamiske, Upper Badenian. 27×, p. 98.
Pisidium ex gr. conventus CLESSIN — Várpalota, Lower Badenian. 30×, p. 98.
Pisidium ex gr. nitidum JENYNS — Várpalota, Lower Badenian. 20×, p. 98.
Pisidium ex gr. casertanum (POLI) — Somlóvásárhely, Lower Miocene. 26×, p. 99.
Plates
187
188
JÓZSEF KÓKAY
Plate XLI
1. Pisidium ex gr. casertanum (POLI) — Somlóvásárhely, Lower Miocene. 28×, p. 99.
2. Pisidium ex gr. casertanum (POLI) — Somlóvásárhely, Lower Miocene. 26×, p. 99.
3. Pisidium ex gr. casertanum (POLI) — Somlóvásárhely, Lower Miocene. 46×, p. 99.
4. Pisidium ex gr. casertanum (POLI) — Somlóvásárhely, Lower Miocene. 33×, p. 99.
5. Argna oppoliensis (ANDREAE) — Várpalota, Sarmatian. 34×, p. 67.
6. Triptychia leobersdorfensis sarmatica nov. sp. — Várpalota, Sarmatian. 1.8×, p. 82.
7. Triptychia leobersdorfensis sarmatica nov. sp. — Várpalota, Sarmatian. 3×, p. 82.
Plates
189
Geologica Hungarica series Palaeontologica, fasciculus 56
APPENDIX
INVENTORY OF NONMARINE MOLLUSCS
FROM THE BAKONY MTS
191
192
JÓZSEF KÓKAY
1 — Ottnangian, Várpalota Basin,
2 — Karpatian, Várpalota and Polgárdi Basin,
3 — Lower Miocene, Somlóvásárhely,
4 — Lower Badenian, Nagygörbő,
5 — Lower Badenian, Herend Basin,
6 — Lower Badenian, Várpalota Basin,
7 — Middle Badenian, Várpalota Basin,
8 — Upper Badenian, Tapolca Basin,
9 — UpperBadenian, Nyirád Basin,
10 — Upper Badenian, Pusztamiske Basin,
11 — Upper Badenian, Herend Basin,
12 — Upper Badenian, Várpalota Basin,
13 — Upper Badenian, Fehérvárcsurgó,
14 — Sarmatian, Pusztamiske Basin,
15 — Sarmatian, Nyirád Basin,
16 — Sarmatian, Tapolca Basin,
17 — Sarmatian, Várpalota Basin,
M = North Alpine Molasse zone and its foreland,
a = pre-Badenian,
i = pre-Sarmatian,
f = post-Badenian,
s = post-Sarmatian
Appendix
193
194
JÓZSEF KÓKAY
Appendix
195
196
JÓZSEF KÓKAY

The load history shown in the Fig. 1 is repeatedly applied to

寸 法 Dimensions(mm) 特徴 Features 仕様 Specifications 金属磁性粉

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