Abstract
In this paper we review five cancellariid assemblages from the Hungarian part of the Pannonian Basin (Börzsöny, Bakony, and Mecsek Mts regions) which yielded 26 species. Ten species are recorded for the first time in Hungary. One species, Scalptia nemethi n. sp. is described as new. A revision of the Hungarian museum collections and historical Hungarian literature is also provided. Sveltia salbriacensis Peyrot 1928 is considered a junior subjective synonym of Petitina inermis (Pusch 1837).
Introduction
The aim of this paper is to review the early Badenian (early Middle Miocene) distribution of the family Cancellariidae in the Hungarian part of the Pannonian Basin (Central Paratethys) by investigation of five recently collected mollusk assemblages (Börzsöny Mts: Letkés, Bakony Mts: Bánd, and Devecser, Mecsek Mts: Mecsekpölöske and Tekeres, Fig. 1). In the previous literature the cancellariids were underrepresented; only three species were documented from Letkés (Csepreghy-Meznerics 1956; Strausz 1966), three from Bánd (Kókay 1966), and two from Devecser (Csepreghy-Meznerics 1958; Strausz 1966). For this study gastropod collections of the Hungarian Natural History Museum, Budapest (HNHM), and the Mining and Geological Survey of Hungary, Budapest (MGSH) were also revisited. Photos of the cancellariid assemblage from Letkés in the Senckenberg Forschungsinstitut, Frankfurt am Main (SFI) were studied; these specimens were collected by Helmut Krock, late German fossil collector.
Early Badenian fossiliferous localities in Hungary discussed in the text, and their locations in the Central Paratethys. (B – Börzsöny Mts, F – Făget Basin, H – Herend Basin, KB – Korytnica Basin, LB – Lom Basin, M− Mecsek Mts, NB – Nógrád Basin, TCR – Transdanubian Central Range, SCF – South Carpathian Foredeep, VB – Vienna Basin) (modified from Haas 2012 and Sant et al. 2019)
Citation: Central European Geology 64, 2; 10.1556/24.2020.00006
As a result of the research 26 species are described and illustrated in this paper; 21 species are recorded from Letkés, seven from Bánd, five from Devecser and Tekeres, and two from Mecsekpölöske. Ten taxa, Bonellitia austriaca (Hoernes and Auinger), Contortia callosa (Hörnes), Contortia tortoniana (Sacco), Merica obsoleta (Hörnes), Ovilia excassidea (Sacco), Perplicaria mioquadrata (Sacco), Scalptia dertoparva (Sacco), Scalptia polonica (Pusch), Trigonostoma exampullaceum (Sacco) and Tritonoharpa mariechristinae Lesport, Cluzaud and Verhecken are recorded for the first time in Hungary; a new species, Scalptia nemethi n. sp., is also designated.
Materials and methods
The specimens investigated in this paper are stored in museum collections of the HNHM, the MGSH, and the SFI, as well as in private collections of Anton Breitenberger (Austria), Tamás Hirmetzl (Fót), Zsolt Kovács (Gödöllő), László Nádai (Budapest), Tamás Németh (Balatonkenese), László Sövér (Bonyhád), Márton Zsoldos (Bakonynána) (Hungary), Gerhard Stein (Germany), and of the authors. The Cancellariidae taxonomy and morphological terminology follow Harzhauser and Landau (2012) and Landau et al. (2006, 2013). Comprehensive works of Bałuk (1997, 2006) and Cahuzac et al. (2004) were also used for taxonomical revision. Only Miocene geographical distribution of the Cancellariidae is cited; for Pliocene ranges see Landau et al. (2006) and Harzhauser and Landau (2012).
Abbreviation: shell length (SL) and shell width (SW) in mm.
Geologic settings and the gastropod fauna
Letkés (W Börzsöny Mts, N Hungary)
Letkés is a Middle Miocene site in the northern part of the Pannonian Basin. A comprehensive work of Badenian mollusks of Szob and Letkés was published by Csepreghy-Meznerics (1956), describing 280 gastropod and 86 bivalve taxa. The locality studied herein is a new excavation situated about 400 m eastward of the village of Letkés (47.888319° N, 18.784647° E). The deposits are characterized by resedimented beds consisting of limonitic marly sand with andesite rock fragments, andesitic tuff and eroded colonial coral blocks – the sediments represent the Lower Badenian Pécsszabolcs Member of the Lajta Formation (referred to as Sámsonháza Fm. in former literature, see Kercsmár 2015) that was deposited in littoral–sublittoral zones. The extremely rich macrofauna shows a mixture of taxa deriving from different biotopes. Five superfamilies or families have been revised within the gastropod assemblage: Conoidea (Kovács and Vicián 2014; Harzhauser and Landau 2016), Siphonarioidea (Harzhauser et al. 2017), Tonnoidea and Ficoidea (Kovács and Vicián 2018), and Muricidae (Kovács 2018; Kovács et al. 2018), as well as several uncommon species of special interest were described by Vicián et al. (2017).
Three cancellariid species were recorded at Letkés by Csepreghy-Meznerics (1956) and Strausz (1966). In the newly collected mollusk material, more than 400 cancellariid specimens were determined at species level, representing 21 species (Table 1).
Cancellariidae species of the investigated early Badenian assemblages
| Letkés | Bánd | Devecser | Tekeres | |
|---|---|---|---|---|
| Bonellitia austriaca | 3 | – | – | – |
| Pseudobabylonella nysti | – | – | 3 | – |
| Cancellicula dregeri | 1 | – | – | – |
| Bivetiella dertonensis | 25 | – | 1 | – |
| Calcarata vindobonensis | 3 | – | – | 7 |
| Contortia callosa | 51 | – | – | – |
| Contortia fenestrata | 6 | – | – | 1 |
| Contortia saccoi | 1 | – | – | – |
| Contortia tortoniana | – | 1 | 4 | – |
| Merica obsoleta | 7 | – | – | – |
| Ovilia excassidea | 7 | – | – | – |
| Perplicaria mioquadrata | – | – | – | 1 |
| Petitina inermis | 4 | – | – | – |
| Scalptia dertoparva | 1 | – | – | – |
| Scalptia gradata | 93 | 6 | – | – |
| Scalptia nemethi n. sp. | 1 | 58 | – | – |
| Scalptia neugeboreni | 6 | 1 | – | – |
| Scalptia polonica | 140 | – | – | – |
| Scalptia spinosa | 7 | – | – | 3 |
| Sveltia dertovaricosa | 8 | 1 | 5 | 100 |
| Sveltia lyrata | – | – | 150 | – |
| Trigonostoma exampullaceum | 10 | – | – | – |
| Trigonostoma exgeslini | 6 | 67 | – | – |
| Ventrilia imbricata | 22 | 55 | – | – |
| Tritonoharpa mariechristinae | 5 | – | – | – |
Mollusks of Letkés represent shallow to deeper-water biotopes (Kovács and Vicián 2014), so the gastropod alpha diversity is generally high. The Cancellariidae material is among the most highly diverse assemblages in the Central Paratethys. Three species, Contortia callosa, Scalptia gradata, and S. polonica are dominant in the cancellariid assemblage, representing altogether 70% of the specimens. The abundance of C. callosa is remarkable, as the species is very rare at other Middle Miocene localities of the Paratethys. Both the Letkés and the Bánd materials are of special interest for the occurrence of a new species: S. nemethi n. sp. Other uncommon species such as B. austriaca, Cancellicula dregeri, Contortia saccoi, M. obsoleta, Ovilia excassidea or Trigonostoma exampullaceum are also documented. The appearance of the genus Tritonoharpa in Hungary is recorded for the first time.
Bánd (Herend Basin, Bakony Mts, W Hungary)
Each gastropod material studied herein was collected from artificial trenches of Locality 28. The locality was described first as “Excavation 28” by Kókay (1966); it is located 250 m southeast of the village church of Bánd (47.121087° N, 17.786770° E). The deposits are characterized by mollusk and coral-bearing grey clay and yellowish clayey sand of approx. 2 m thickness; these shallow marine sediments belong to the Pécsszabolcs Member. The colonial coral, brachiopod, polyplacophoran, gastropod, and bivalve material indicate intertidal to infralittoral zones, and a patch reef paleoenvironment (Dulai 2007).
Three species were recorded from the locality by Kókay (1966). Based on revisions of museum collections and the newly collected private material, more than 190 specimens are recorded, representing seven species (Table 1).
Gastropods from this locality were recently reviewed by Vicián et al. (2017) and Kovács (2020). In the shallow-water patch reef environment, species of the nearshore are over-represented; e.g., within the turritellid material Archimediella carpathica Harzhauser and Landau 2019 [= Turritella (Archimediella) dertonensis in Kókay 1966 (non Mayer)] and Oligodia bicarinata (Eichwald 1830) are the most abundant. In the family Conidae Lautoconus kovacsi Harzhauser and Landau 2016 is the dominant species, while the muricid Janssenia echinulata (Pusch 1837) accounts for about 50% of the gastropods found.
Cancellariids are less diverse than at Letkés. Species of the deeper circalittoral zone, which are frequent at Letkés (e.g., Contortia callosa, S. polonica), are absent at Locality 28. The fauna is of special interest for its abundance of S. nemethi n. sp., the appearance of S. neugeboreni, and a new record in the Pannonian Basin: Contortia tortoniana. Three taxa: S. nemethi n. sp., Trigonostoma exgeslini, and Ventrilia imbricata predominate among the cancellariid material.
Devecser (Devecser–Nyirád Basin, Bakony Mts, W Hungary)
The geology of the area was treated by Csepreghy-Meznerics (1958) and Selmeczi (2003). Fauna lists of Badenian mollusks from the vicinity were published by the first author without illustrations; later two cancellariid specimens were illustrated by Strausz (1966). The locality studied herein is located on Tik Hill east of Devecser (46.10221° N, 17.46773° E), and was first recorded by Németh (2005) and Dulai (2005). Yellow marly sand of 190 cm thickness was excavated by artificial trenches; the deposit represents the Pécsszabolcs Member. The deposits are very rich in excellently preserved gastropods. One hundred and sixty-three cancellariid specimens were determined at species level, and five species are documented. We highlight the presence of Contortia tortoniana and the abundance of Sveltia lyrata (Table 1).
Tekeres (NW Mecsek Mts, SW Hungary)
The geology and stratigraphy of the Orfű–Tekeres region were dealt with by Báldi et al. (2002); gastropods were recorded from borehole Tekeres-1 by Bohn-Havas (1973). The Lower Badenian deposits belong to the Pécsszabolcs Member and the Tekeres Schlier Formation (the latter represents offshore marine environments). The locality studied herein is located 100 m east of Lake Herman Ottó in a forested area (46.17449° N, 18.13063° E); it was first recorded by Bosnakoff, 2013. Clayey sand of 180 cm thickness was excavated by artificial trenches; the deposit represents the Pécsszabolcs Fm. The macrofauna is very rich in Badenian invertebrate and vertebrate fossils. The analysis of the gastropod assemblage is in progress; the family Muricidae was investigated by Kovács (2020).
The gastropod material was collected by the staff of the HNHM (M. Szabó, A. Dulai) and T. Németh. Eighty-two specimens were determined at species level, representing five species (Table 1).
The deposit indicates nearshore to offshore paleoenvironment. The gastropod fauna is dominated by the turritellid Ptychidia vindobonensis (Handmann 1882) and the aporrhaid Aporrhais dactylifera (Boettger 1902); from the Conidae only one species, Conilithes exaltatus (Eichwald 1830) appears frequently. The cancellariid diversity is low. Of special interest are the presence of the rare genus Perplicaria, and the extended geographical range of the other species. The dominant cancellariid is Sveltia dertovaricosa, that otherwise occurs only sporadically in other Badenian localities of the Pannonian Basin; it forms 89% of the family. Bathymetric ranges of two abundant species, Calcarata vindobonensis and S. dertovaricosa can be inferred from the habitats of their Late Miocene–Pliocene descendants. Calcarata calcarata (Brocchi 1814) is abundant in the deep circalittoral zone, while Sveltia varicosa (Brocchi 1814) is typical of the sublittoral inner neritic settings (Brunetti et al. 2011). The faunal composition of the assemblage is unique in the Pannonian Basin, as a few gastropod species [e.g., P. mioquadrata or the muricid Pteropurpura friedbergi (Cossmann and Peyrot 1924)] appear only at Tekeres in Hungary (Kovács 2020).
Mecsekpölöske (N Mecsek Mts, SW Hungary)
The geology and stratigraphy of the Mecsekpölöske region were treated by Strausz (1928) and Chikán (1991). The Lower Badenian deposits of the area represent the Pécsszabolcs Member. The newly discovered locality is located 1 km east of the village in a ravine on the western side of the Nagy Hill (46.22311° N, 18.22904° E). Three collections were examined: the mollusk assemblage collected by the staff of the HNHM (M. Szabó, A. Dulai) and the private collections of T. Németh and L. Sövér.
Alternating layers of mollusk-bearing limestone and dark yellow sandy clay of approx. 3 m thickness were excavated in the locality. The analysis of the gastropod fauna is in progress (Kovács 2020). Two species, Bivetiella dertonensis (Bellardi 1841) (2 specimens) and Scalptia hidasensis (Hoernes and Auinger 1890) (20 specimens) represent the Cancellariidae; the latter is relatively common in the assemblage. S. hidasensis has been known only from Hidas in the Pannonian Basin.
Systematic paleontology
Class Gastropoda Cuvier, 1795.
Order Neogastropoda Wenz 1938.
Superfamily Cancellarioidea Forbes and Hanley 1851.
Family Cancellariidae Forbes and Hanley 1851.
Admetula-clade (sensu Harzhauser and Landau 2012).
Genus Bonellitia Jousseaume 1887.
Type species: Cancellaria bonellii Bellardi 1841
Bonellitia austriaca (Hoernes and Auinger 1890) (Fig. 2/A)
1890 Cancellaria Austriaca nov. form. – Hoernes and Auinger, p. 275, Pl. 33, Figs 15/A–C.
2012 Bonellitia austriaca (Hoernes and Auinger) – Harzhauser and Landau, p. 10, Figs 3/B, 4/B, C (cum syn.).
(A) Bonellitia austriaca (Hoernes and Auinger), SL 10, Letkés (HNHM: INV 2020.45). (B) Cancellicula dregeri (Hoernes and Auinger), SL 3, Letkés (444441, Coll. Krock, SFI). (C–D) Bivetiella dertonensis (Bellardi), SL 27, Letkés (Coll. Németh). (E–G) Calcarata vindobonensis (Hilber): (E–F) SL 17.5, Tekeres, (G) SL 9, Letkés (Coll. Németh). (H–L) Contortia callosa (Hörnes), Letkés: (H–I) SL 38 (Coll. Nádai), (J–K) SL 32.2 (Coll. Hirmetzl), (L) SL 38 (Coll. Németh). (M−N) Contortia fenestrata (Eichwald), SL 17, Letkés (Coll. Hirmetzl). (O) Contortia saccoi (Hoernes and Auinger), SL 22, Letkés (Coll. Hirmetzl). (P–S) Contortia tortoniana (Sacco), Devecser: (P–Q) SL 26.2, (R–S) SL 23.3 (Coll. Németh). Scale bars: 1 mm for A–B, G; 5 mm for C–D, H–S
Citation: Central European Geology 64, 2; 10.1556/24.2020.00006
Studied material: 3 specimens (max. SL 10 mm, SW 6 mm). HNHM: INV 2020.45. SFI: Krock Coll.: 444322, 444577.
Remarks: B. austriaca differs from the congeners by characteristic sculpture. The strong axial ribs are crossed by raised spiral cords producing a cancellate pattern of deep, subquadrate interspaces (Harzhauser and Landau 2012). The genus is known in the Badenian Pannonian Basin, Bonellitia bonellii (Bellardi) was recorded by Strausz (1966) and Csepreghy-Meznerics (1969). B. austriaca is newly recorded in the Pannonian Basin; it occurs in the gastropod assemblage of Letkés.
Distribution: Central Paratethys. ?Ottnangian: North Alpine Foreland Basin (S Germany), Badenian: Vienna Basin (Austria), Southern Moravia (Czechia), Pannonian Basin (Hungary: Börzsöny Mts: Letkés).
Subfamily Admetinae Troschel 1865.
Genus Pseudobabylonella Brunetti, Della Bella, Forli and Vecchi 2009
Type species: Cancellaria minima Reeve 1856.
Pseudobabylonella nysti (Hörnes 1854).
1854 Cancellaria nysti – Hörnes, p. 305, Pl. 34, Fig. 1.
1956 Admete (Babylonella) fusiformis nysti Hörnes – Csepreghy-Meznerics, p. 418, Pl. 9, Figs 11–12, 18–19, 28–29.
1966 Cancellaria (Admete) fusiformis nysti Hörnes – Strausz, p. 384, Pl. 12, Figs 13–14.
2012 Pseudobabylonella nysti (Hörnes) nov. comb. – Harzhauser and Landau, p. 13, Figs 3/C, 4/E (cum syn.).
Studied material: 3 specimens in private collections (max. SL 8.8 mm, SW 3 mm).
Remarks: The taxonomic arrangement of C. nysti was discussed by Harzhauser and Landau (2012), and it was assigned to the genus Pseudobabylonella. The species was the most frequent cancellariid in the locality of Szob (Csepreghy-Meznerics 1956), but it is uncommon at other sites of the Pannonian Basin. P. nysti is a new occurrence at Devecser.
Distribution: Central Paratethys. Badenian: Vienna Basin (Austria, Czechia), Pannonian Basin (Hungary: Börzsöny Mts: Szob, Cserhát Mts: Mátraverebély, Bakony Mts: Devecser, Mecsek Mts: Pécsvárad), Făget Basin (Romania).
Genus Cancellicula Tabanelli 2008.
Type species: Cancellaria (Narona) Dregeri Hoernes and Auinger 1890.
Cancellicula dregeri (Hoernes and Auinger 1890) (Fig. 2/B)
1890 Cancellaria (e. Narona?) Dregeri nov. form. – Hoernes and Auinger, p. 280, Pl. 33, Figs 18–19.
1969 Cancellaria dregeri Hoernes et Auinger – Csepreghy-Meznerics, p. 94, Pl. 5, Figs 20, 32.
2012 Cancellicula dregeri (Hoernes and Auinger) – Harzhauser and Landau, p. 16, Figs 3/F, 4/F–G (cum syn.).
Studied material: 1 specimen (SL 3 mm, SW 1.8 mm). SFI: Krock Coll.: 444441.
Remarks: The specimen figured here with flattened protoconch and teleoconch sculpture corresponds to the holotype and the syntype of Cancellicula dregeri reillustrated by Harzhauser and Landau (2012). The species is typical of deeper water environments of the Neogene Paratethys and the Proto-Mediterranean Sea. Although it is widely distributed in the Central Paratethys, C. dregeri is rare in Pannonian Basin localities. Only one fragmentary specimen appears in the gastropod assemblage of Letkés; it was collected by H. Krock. We were able to study photos by courtesy of Ronald Janssen and Sigrid Hof.
Distribution: Middle Miocene–Pleistocene. Central Paratethys. Badenian: Vienna Basin (Austria, Czechia), Eisenstadt–Sopron Basin (Austria), Pannonian Basin (Hungary: Börzsöny Mts: Letkés, Bükk Mts: Borsodbóta), Făget Basin (Romania). Proto-Mediterranean Sea. Tortonian–Messinian: Torino Hills (Italy).
Subfamily Cancellariinae Forbes and Hanley 1851.
Genus Bivetiella Wenz 1943.
Type species: Cancellaria similis Sowerby 1833.
Bivetiella dertonensis (Bellardi 1841) (Fig. 2/C–D)
1841 Cancellaria bonellii var. Dertonensis Bell. – Bellardi, p. 248, Pl. 3, Figs 11–12.
1890 Cancellaria subcancellata – Hoernes and Auinger, p. 273 (non d’Orbigny).
1954 Cancellaria subcancellata perscalarata – Csepreghy-Meznerics, p. 49, Pl. 6, Figs 19–21 (reillustrated by Strausz 1966, Pl. 44, Figs 2–3) (non Sacco).
1956 Cancellaria (Cancellaria) subcancellata – Csepreghy-Meznerics, p. 415, Pl. 9, Figs 32–35 (non d’Orbigny).
1966 Cancellaria cancellata dertonensis Bellardi – Strausz, p. 376, Pl. 45, Figs 5–6.
non 1966 Cancellaria cancellata dertonensis – Strausz, Pl. 45, Figs 2–4 [= Contortia fenestrata (Eichwald 1830)].
1971–1972 Cancellaria cancellata perscalarata – Csepreghy-Meznerics, Pl. 14, Fig. 26 (non Sacco).
1973 Cancellaria cancellata – Borza, Pl. 3, Fig. 4 (non Linnaeus).
2011 Bivetiella praecedens – Brunetti et al., Fig. 2/D, E (non Beyrich).
2012 Bivetiella dertonensis (Bellardi) – Harzhauser and Landau, p. 18, Figs 3/H, 5/B–D (cum syn.).
2013 Bivetiella dertonensis (Bellardi) – Landau et al., p. 224, Pl. 35, Fig. 3, Pl. 80, Fig. 11.
Studied material: 28 specimens (max. SL 27 mm, SW 19.7 mm). HNHM: M.62.885. Private collections: 27 specimens.
Remarks: The taxonomical revision of the species was arranged by Harzhauser and Landau (2012). B. dertonensis is widespread in the Paratethys, and appears in numerous Badenian localities in Hungary; however, it is usually a rare element of the gastropod assemblages. Based on morphology the specimen figured by Brunetti et al. (2011, Fig. 2/D, E) as B. praecedens from the Badenian of Szob is here considered as a representative of B. dertonensis.
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin, Eisenstadt–Sopron Basin, Styrian Basin (Austria), Vienna Basin (Austria, Czechia), Korytnica Basin (Poland), Pannonian Basin (Hungary: Börzsöny Mts: Szob, Letkés, Hont, Cserhát Mts: Sámsonháza, Bükk Mts: Borsodbóta, Bakony Mts: Devecser, Mecsek Mts: Hidas, Mecsekpölöske), Făget Basin (Romania). NE Atlantic. Serravallian: Aquitanian Basin (France), Montjuic Basin (Spain), Tortonian: Cacela Basin (Portugal). Proto-Mediterranean Sea. Serravallian–Tortonian: Karaman Basin (Turkey), Tortonian–Messinian: Po Basin (Italy).
Genus Calcarata Jousseaume 1887.
Type species: Voluta calcarata Brocchi 1814.
Calcarata vindobonensis (Hilber 1892) (Fig. 2/E–G)
1892 Cancellaria (Trigonostoma) calcarata Broc. var. Vindobonensis Hilb. – Hilber, p. 1020.
1956 Cancellaria (Calcarata) calcarata – Csepreghy-Meznerics, p. 417, Pl. 9, Figs 36–39 (non Brocchi).
1966 Cancellaria (Calcarata) calcarata – Strausz, p. 381, Text-fig. 166, Pl. 43, Figs 16–18 (non Brocchi).
2012 Calcarata vindobonensis (Hilber) nov. comb. – Harzhauser and Landau, p. 19, Figs 3/I, 5/A (cum syn.).
Studied material: 10 specimens (max. SL 17.5 mm, SW 10.5 mm). HNHM: INV 2020.54 (4 spec.). SFI: Krock Coll.: 444107A–B. Private collections: 4 specimens.
Remarks: Based on taxonomical revision the Cancellaria calcarata recorded in the Badenian Central Paratethys were assigned to vindobonensis by Harzhauser and Landau (2012). The latter species differs from the Late Miocene–Pliocene Calcarata calcarata (Brocchi) in morphology by presence of weakly developed spiral cords on the last whorl between the shoulder and the basal carina, and on the base, by shorter base and less developed basal spiral carina. Based on new collecting work the occurrence of the species is documented in the Letkés, Tekeres and Hidas gastropod assemblages; these records extend the geographical range of C. vindobonensis in the Pannonian Basin.
Distribution: Central Paratethys. Badenian: Vienna Basin (Austria, Czechia, Slovakia), Korytnica Basin (Poland), Pannonian Basin (Bosnia; Hungary: Börzsöny Mts: Szob, Letkés, Mecsek Mts: Bodolyabér, Hidas, Pécsvárad, Tekeres), Făget Basin, Caransebeș Basin, Transylvanian Basin (Romania), Krka Basin (Slovenia), South Carpathian Foredeep (Bulgaria).
Genus Contortia Sacco 1894.
Type species: Cancellaria contorta Basterot 1825.
Contortia callosa (Hörnes 1854) (Fig. 2/H–L)
1854 Cancellaria callosa Partsch – Hörnes, p. 314, Pl. 34, Figs 14–16.
2012 Contortia callosa (Hörnes) nov. comb. – Harzhauser and Landau, p. 21, Figs 3/J, 5/E, F, G (cum syn.).
2013 Contortia callosa (Hörnes) – Landau et al., p. 226, Pl. 35, Figs 5–6, Pl. 69, Fig. 6, Pl. 80, Fig. 12.
Studied material: 51 specimens in private collections (max. SL 38.2 mm, SW 27.8 mm).
Remarks: Contortia callosa differs from its congeners by a larger and more robust shell with fine spiral threads and broad, weakly developed axial ribs. The species displays moderate morphological variability in height of the spire. C. callosa is widely distributed in the Paratethys and the Proto-Mediterranean Sea but generally rare. It is a new occurrence in the Pannonian Basin, occurring in the recently discovered early Badenian fossiliferous locality near Kisbattyán (for detailed description of the site see Kovács 2020), and in the Letkés assemblage, where its relative abundance is notable.
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin, Vienna Basin, Eisenstadt–Sopron Basin (Austria), Korytnica Basin (Poland), Pannonian Basin (Hungary: Börzsöny Mts: Letkés, Mecsek Mts: Kisbattyán), Făget Basin (Romania). Proto-Mediterranean Sea. Langhian: Torino Hills (Italy), Serravallian–Tortonian: Karaman Basin (Turkey).
Contortia fenestrata (Eichwald 1830) (Fig. 2/M−N)
1830 Cancellaria fenestrata – Eichwald, p. 222.
1956 Cancellaria (Merica) fenestrata Eichwald – Csepreghy-Meznerics, p. 417, Pl. 9, Figs 24–27.
1966 Cancellaria cancellata dertonensis – Strausz, Pl. 45, Figs. 2–4 (non Bellardi).
2008 Contortia contorta – Brunetti et al., Fig. 7/E–F (non Basterot).
2012 Contortia fenestrata (Eichwald) nov. comb. – Harzhauser and Landau, p. 23, Fig. 5/H, I (cum syn.).
Studied material: 7 specimens in private collections (max. SL 17 mm, SW 10.2 mm).
Remarks: Contortia fenestrata differs from C. callosa by its small size, slender shell, higher spire and presence of an umbilicus. The species is newly recorded at Letkés and Tekeres in the Pannonian Basin. The specimen illustrated by Brunetti et al. (2008, Fig. 7/E–F) as Contortia contorta from the Badenian deposits of Szob (Börzsöny Mts) in fact represents C. fenestrata.
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin, Vienna Basin (Austria), Polish–Carpathian Foredeep (Poland, Ukraine), Pannonian Basin (Bosnia; Hungary: Börzsöny Mts: Szob, Letkés, Mecsek Mts: Hidas, Tekeres), Făget Basin (Romania), South Carpathian Foredeep (Bulgaria).
Contortia saccoi (Hoernes and Auinger 1890) (Fig. 2/O)
1854 Cancellaria Bellardii – Hörnes, p. 314, Pl. 34, Fig. 17 (non Michelotti).
1890 Cancellaria Saccoi – Hoernes and Auinger, p. 274 pars.
1969 Cancellaria (Bonellitia) evulsa taurinia – Csepreghy-Meznerics, p. 93, Pl. 5, Figs 35, 36 (non Sacco).
1971–1972 Cancellaria fenestrata – Csepreghy-Meznerics, Pl. 14, Figs 17, 19 (non Eichwald).
2012 Contortia saccoi (Hoernes and Auinger) nov. comb. – Harzhauser and Landau, p. 25, Fig. 6/A, B (cum syn.).
Studied material: 1 specimen in private collection (SL 22 mm, SW 13.9 mm).
Remarks: The species differs from the similar C. tortoniana in having a somewhat broader shell, lower spire, strong varices on the last whorl, two columellar folds (as opposed to three in C. tortoniana) and lack of an umbilicus.
Distribution: Central Paratethys. Badenian: Vienna Basin, Eisenstadt–Sopron Basin, Styrian Basin (Austria), Pannonian Basin (Hungary: Börzsöny Mts: Letkés, Bükk Mts: Borsodbóta, Balaton, Csermely), Făget Basin (Romania).
Contortia tortoniana (Sacco 1894) (Fig. 2/P–S, Fig. 3/A–B)
1854 Cancellaria Bellardii – Hörnes, p. 314, Pl. 34, Fig. 18 (non Michelotti).
1894 Cancellaria (Bonellitia) evulsa var. tortoniana – Sacco, p. 46.
1966 Cancellaria (Merica) fenestrata – Strausz, Pl. 45, Fig. 7 (non Eichwald).
2012 Contortia tortoniana (Sacco) nov. comb. – Harzhauser and Landau, p. 25, Fig. 6/C, D (cum syn.).
(A–B) Contortia tortoniana (Sacco), SL 27, Bánd (Coll. Németh). (C–D) Merica obsoleta (Hörnes), SL 22, Letkés (Coll. Hirmetzl). (E–F) Ovilia excassidea (Sacco), SL 20, Letkés (Coll. Nádai). (G–H) Perplicaria mioquadrata (Sacco), SL 12, Tekeres (Coll. Németh). (I–J) Petitina inermis (Pusch), SL 41, Letkés (HNHM: INV 2020.46). (K–L) Scalptia dertoparva (Sacco), SL 16, Letkés (Coll. Hirmetzl). (M−P) Scalptia gradata (Hörnes), Letkés: (M−N) SL 20 (HNHM: INV 2020.47), (O–P) SL 24 (Coll. Hirmetzl). Scale bar: 5 mm
Citation: Central European Geology 64, 2; 10.1556/24.2020.00006
Studied material: 5 specimens in private collections (max. SL 27 mm, SW 17.3 mm).
Remarks: The taxonomical revision of the species was provided by Harzhauser and Landau (2012). Contortia tortoniana is characterized by moderate morphological variability in height of the spire and the strength of the sculpture. It differs from the similar C. saccoi by a somewhat higher spire, lack of varices on the last whorl and three columellar folds. C. tortoniana is newly recorded in the Pannonian Basin.
Distribution: Central Paratethys. Badenian: Vienna Basin, Eisenstadt–Sopron Basin (Austria), Korytnica Basin (Poland), Pannonian Basin (Hungary: Bakony Mts: Bánd, Devecser), Făget Basin (Romania), South Carpathian Foredeep (Bulgaria).
Genus Merica H. and A. Adams 1854.
Type species: Cancellaria melanostoma G. B. Sowerby II 1849.
Merica obsoleta (Hörnes 1856) (Fig. 3/C–D)
1856 Cancellaria obsoleta – Hörnes, 678, Pl. 52, Fig. 3.
2012 Merica obsoleta (Hörnes) – Harzhauser and Landau, p. 29, Fig. 6/I, J (cum syn.).
Studied material: 7 specimens (max. SL 22.6 mm, SW 16.8 mm). SFI: Krock Coll.: 444468, 444483. Private collections: 5 specimens.
Remarks: Three species of the genus: Merica crenata (Hörnes 1856), M. obsoleta (Hörnes 1856) and Merica succineiformis (Boettger 1906) were recognized by Harzhauser and Landau (2012); all three are very uncommon. M. crenata and succineiformis are locally endemic species; the first is known from Grund (Austria), the second from Coşteiu de Sus (Romania). M. obsoleta has been recorded from both sites; however, in the Badenian it is known only at Grund. The species is a new occurrence in the Pannonian Basin.
Distribution: Central Paratethys. Karpatian: Waschberg Unit (Austria), Badenian: North Alpine Foreland Basin (Austria), Pannonian Basin (Hungary: Börzsöny Mts: Letkés).
Genus Ovilia Jousseaume 1887.
Type species: Cancellaria doliolaris Basterot 1825.
Ovilia excassidea (Sacco 1894) (Fig. 3/E–F)
1856 Cancellaria cassidea – Hörnes, p. 682, Pl. 52, Fig. 8 (non Brocchi).
1894 Cancellaria (Trigonostoma) cassideum forma excassidea – Sacco, p. 8.
2012 Ovilia excassidea (Sacco) nov. comb. – Harzhauser and Landau, p. 32, Fig. 7/C (cum syn.).
2013 Ovilia excassidea (Sacco) – Landau et al., p. 232, Pl. 36, Fig. 9.
Studied material: 7 specimens in private collections (max. SL 20 mm, SW 17 mm).
Remarks: This uncommon species has been known only from the Vienna and the Karaman basins; it is recorded for the first time in the Pannonian Basin.
Distribution: Central Paratethys. Badenian: Vienna Basin (Austria), Pannonian Basin (Hungary: Börzsöny Mts: Letkés). Proto-Mediterranean Sea. Serravallian: Karaman Basin (Turkey).
Genus Perplicaria Dall 1890.
Type species: Perplicaria perplexa Dall 1890.
Perplicaria mioquadrata (Sacco 1894) (Fig. 3/G–H)
1890 Cancellaria (d. Merica) Laurensii – Hoernes and Auinger, p. 281, Pl. 33, Fig. 1–3 (non Grateloup).
1894 Merica mioquadrata – Sacco, p. 66.
2012 Perplicaria mioquadrata (Sacco) nov. comb. – Harzhauser and Landau, p. 34, Figs 3/K, 7/F, G (cum syn.).
Studied material: 1 specimen in private collection (SL 12 mm, SW 5.4 mm).
Remarks: The species is widespread in the Central Paratethys but recorded for the first time in the Pannonian Basin. Only a single P. mioquadrata specimen is known from the mollusk assemblage of Tekeres.
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin (Austria), Vienna Basin (Czechia), Korytnica Basin (Poland), Pannonian Basin (Hungary: Mecsek Mts: Tekeres), Făget Basin (Romania), South Carpathian Foredeep (Bulgaria).
Genus Petitina Harzhauser and Landau 2012.
Type species: Cancellaria inermis Pusch 1837.
Petitina inermis (Pusch 1837) (Fig. 3/I–J)
1837 Cancellaria inermis – Pusch, p. 129, Pl. 11, Fig. 22.
1928 Sveltia salbriacensis nov. sp. – Peyrot, p. 220, Pl. 13, Figs 25–26 only [27–28 = S. lyrata (Brocchi 1814)] (refigured by Cahuzac et al. 2004, p. 249, Fig. 9/H–K).
1956 Cancellaria (Sveltia) inermis Pusch – Csepreghy-Meznerics, p. 416, Pl. 9, Figs 20–21.
1966 Cancellaria (Sveltia) inermis Pusch – Strausz, p. 380, Pl. 43, Figs 6–9.
2012 Petitina inermis (Pusch) – Harzhauser and Landau, p. 35, Figs 3/L, 7/H, I, J (cum syn.).
2013 Petitina inermis (Pusch) – Landau et al., p. 230, Pl. 35, Figs 12–13.
2017 Petitina inermis (Pusch) – Biskupič, Fig. 6/E.
Studied material: 4 specimens (max. SL 52.5 mm, SW 25 mm). HNHM: INV 2020.46. SFI: Krock Coll.: 444438. Private collections: 2 specimens.
Remarks: The species is widespread in the Middle Miocene of Europe, and is characterized by significant shell variability, mainly in the height of the spire and the development of the sculpture (Harzhauser and Landau 2012). The specimen illustrated herein with high, gradate spire is closely allied in morphology to the specimen that was described as S. salbriacensis by Peyrot (1928) from the Late Burdigalian (or Langhian) Aquitanian Basin. According to Cahuzac et al. (2004: 249) only one specimen, the holotype of salbriacensis is known. Based on size and morphology Peyrot's taxon is considered as a junior synonym of P. inermis.
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin, Eisenstadt–Sopron Basin (Austria), Vienna Basin (Austria, Czechia, Slovakia), Korytnica Basin (Poland), Pannonian Basin (Hungary: Börzsöny Mts: Szob, Letkés, Bükk Mts: ?Borsodbóta), Făget Basin (Romania), Krka Basin (Slovenia). NE Atlantic and Proto-Mediterranean Sea. Serravallian–Tortonian: Aquitanian Basin (France), Adiça (Portugal), Montjuic Basin (Spain), Karaman Basin (Turkey).
Genus Scalptia Jousseaume 1887b.
Type species: Cancellaria obliquata Lamarck 1822.
Scalptia dertoparva (Sacco 1894) (Fig. 3/K–L)
1856 Cancellaria scabra – Hörnes, p. 681, Pl. 52, Fig. 7 (non Deshayes).
1894 Cancellaria (Trigonostoma) scabrum var. dertoparva – Sacco, p. 7.
2012 Scalptia dertoparva (Sacco) nov. comb. – Harzhauser and Landau, p. 38, Figs 3/N, 8/C (cum syn.).
Studied material: 1 specimen in private collection (SL 16 mm, SW 13 mm).
Remarks: S. dertoparva differs in morphology from the more abundant S. polonica by its shorter and broader shell, wider aperture, as well as by the presence of a parietal denticle and a small umbilicus. The species is newly recorded in the Pannonian Basin.
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin, Vienna Basin (Austria), Pannonian Basin (Hungary: Börzsöny Mts: Letkés).
Scalptia gradata (Hörnes 1854) (Fig. 3/M−P)
1854 Cancellaria gradata – Hörnes, p. 319, Pl. 35, Fig. 2.
1956 Cancellaria (Trigonostoma) gradata Hörnes – Csepreghy-Meznerics, p. 416, Pl. 9, Figs 30–31.
1962 Cancellaria sp. – Kecskeméti-Körmendi, p. 91, Pl. 11, Fig. 16.
1966 Cancellaria (Trigonostoma) gradata Hörnes – Strausz, p. 378, Pl. 45, Figs 8–11.
?1966 Cancellaria (Trigonostoma) gradata Hörnes – Strausz, Text-fig. 165, Pl. 45, Figs 12–13.
1973 Cancellaria (Trigonostoma) gradata Hörnes – Bohn-Havas, p. 1061, Pl. 6, Figs 14–15 only (Figs 12-13: Scalptia nemethi n. sp.).
2012 Scalptia gradata (Hörnes) – Harzhauser and Landau, p. 42, Fig. 8/I (cum syn.).
Studied material: 99 specimens (max. SL 24 mm, SW 12 mm). HNHM: INV 2020.47. Private collections: 98 specimens.
Remarks: S. gradata is widespread in the Central Paratethys. It is a new occurrence at Letkés, where it is the second most abundant cancellariid (23%) in the cancellariid material. The S. gradata specimens in this assemblage are characterized by a moderate morphological variability in strength of the parietal denticle. On adult specimens the spire may attain more than 50% of the total shell length (Fig. 3/O–P). The specimen illustrated by Strausz (1966, Pl. 45, Figs 12–13) differs from the type of S. gradata, and needs to be revised.
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin, Vienna Basin, Eisenstadt–Sopron Basin (Austria), Polish–Carpathian Foredeep (Poland), Pannonian Basin (Bosnia; Hungary: Börzsöny Mts: Szob, Letkés, Cserhát Mts: Sámsonháza, Bakony Mts: Várpalota, Márkó, Bánd, Mecsek Mts: Hidas, Hosszúhetény, Pécsvárad), Făget Basin (Romania).
Scalptia hidasensis (Hoernes and Auinger 1890) (Fig. 5/A–D)
1890 Cancellaria (d. Trigonostoma) Hidasensis nov. form. – Hoernes and Auinger, p. 276, Pl. 33, Figs 13–14.
non 1950 (Trigonostoma) hidasensis – Csepreghy-Meznerics, Pl. 3, Fig. 15 [= Trigonostoma exgeslini (Sacco 1894)].
1966 Cancellaria (Trigonostoma) hidasensis Hoernes and Auinger – Strausz, p. 378, Pl. 45, Fig. 16 only.
2012 Scalptia hidasensis (Hoernes and Auinger) nov. comb. – Harzhauser and Landau, p. 43, Figs 3/P, 9/A, B (cum syn.).
Studied material: 20 specimens in private collections (max. SL 19 mm, SW 14 mm).
Remarks: The occurrence of S. hidasensis in the early Badenian deposits of Mecsekpölöske, and the neighboring Kisbattyán, extends the paleogeographic range of the species that has been known only at Hidas (Mecsek Mts) and Buitur (Transylvanian Basin).
Distribution: Central Paratethys. Badenian: Pannonian Basin (Hungary: Mecsek Mts: Hidas, Mecsekpölöske, Kisbattyán), Transylvanian Basin (Romania).
Scalptia nemethi n. sp. (Fig. 4/A–N)
?1966 Cancellaria (Trigonostoma) gradata Hörnes – Strausz, Text-fig. 165, Pl. 45, Figs 12–13 (only).
1973 Cancellaria (Trigonostoma) gradata – Bohn-Havas, Pl. 6, Figs 12–13 (only) (non Hörnes).
Holotype: PAL 2020.25.1., HNHM, Dep. Palaeontology and Geology, SL 30 mm, SW 17.4 mm, Bánd (Fig. 4/A–C).
Paratype 1: PAL 2020.26.1., HNHM, Dep. Pal. and Geol., SL 22 mm, SW 12.6 mm, Bánd (Fig. 4/H–I).
Paratype 2: PAL 2020.27.1., HNHM, Dep. Pal. and Geol., SL 18 mm, SW 10.5 mm, Bánd (Fig. 4/L–N).
Paratype 3: PAL 2020.28.1., HNHM, Dep. Pal. and Geol., SL 17 mm, SW 11 mm, Letkés (Fig. 4/J–K).
Paratype 4: 2020.01, Coll. Németh, SL 30 mm, SW 16.6 mm, Bánd (Fig. 4/D–E).
Paratype 5: 2019.Bánd.10, Coll. Vicián, SL 28 mm, SW 15.5 mm, Bánd (Fig. 4/F–G).
Type strata: Badenian (Middle Miocene) clayey sand (Pécsszabolcs Member of the Lajta Formation).
Type locality: Locality 28, Bánd, Bakony Mts, Hungary.
Derivation of name: In honor of Tamás Németh, Hungarian fossil shell collector.
Material: 59 specimens at Bánd, 1 specimen at Letkés, 1 specimen at Pécsszabolcs.
(A–N) Scalptia nemethi n. sp.: (A–B–C) Holotype, SL 30, Bánd (HNHM PAL 2020.25.1.), (D–E) Paratype 4, SL 30, Bánd (Coll. Németh), (F–G) Paratype 5, SL 28, Bánd (Coll. Vicián), (H–I) Paratype 1, SL 22, Bánd (HNHM PAL 2020.26.1.), (J–K) Paratype 3, SL 17, Letkés (HNHM PAL 2020.28.1.), (L–M–N) Paratype 2, SL 18, Bánd (HNHM PAL 2020.27.1.). Scale bars: 5 mm for A–B, D–M; 2 mm for C; 1 mm for N
Citation: Central European Geology 64, 2; 10.1556/24.2020.00006
(A–D) Scalptia hidasensis (Hoernes and Auinger), Mecsekpölöske: (A–B) SL 19, (C–D) SL 14 (Coll. Németh). (E–H) Scalptia neugeboreni (Hörnes): (E–F) SL 22, Bánd (Coll. Németh), (G–H) SL 18, Letkés (HNHM: INV 2020.48). (I–P) Scalptia polonica (Pusch), Letkés: (I–J) SL 37.4 (Coll. Németh), (K–L) SL 24 (HNHM: INV 2020.49), (M−N) SL 20 (HNHM: INV 2020.50), (O–P) SL 26 (Coll. Németh). Scale bar: 5 mm
Citation: Central European Geology 64, 2; 10.1556/24.2020.00006
Diagnosis: Scalptia with medium sized fusiform shell, paucispiral protoconch, five teleoconch whorls, gradate spire, last whorl with angulated shoulder, ovate aperture with lirate outer lip and well-developed parietal denticle, three columellar folds, sculpture of marked axial ribs and spiral cords.
Description: Shell medium-sized (max. SL 30, SW 18), fusiform with a high, gradate spire. Small nucleus, eroded, low, paucispiral protoconch of approx. 1 3/4 smooth, convex whorls, junction with teleoconch delimited by weak prosocline scar. First teleoconch whorl rounded, cancellate sculpture with dense axial ribs and fine spiral cords. The second to fifth teleoconch whorls slightly convex, scalate and angulated, suture undulating, sutural ramp horizontal, narrow. Last whorl 73–76% of the total length, slightly convex with sloping subsutural ramp, sharply angled at shoulder. Aperture moderately narrow and ovate, outer lip thickened by labral varix, strongly lirate within. Strong parietal denticle, well-developed, thickened parietal callus. Columella weakly excavated, bearing three oblique columellar folds, columella callus thickened, tuberculate. Short and open siphonal canal, narrow umbilical chink. Axial sculpture of regularly spaced, rounded, orthocline to prosocline ribs on spire whorls, and 9–10 raised, prosocline ribs on the last whorl. Spiral sculpture of narrow and sharp primary cords (10 on the last whorl) and fine secondary and tertiary threads in the interspaces. The spiral cords and threads override the axial ribs.
Remarks: Based on morphology the new species is assigned to the genus Scalptia. S. gradata (Hörnes) resembles S. nemethi n. sp. in gradate and angulated spire but differs by its smaller size, more slender shell, higher protoconch, more strongly angulated first teleoconch whorl, rounded primary spiral cords, triangular aperture, thinner outer lip and weaker parietal denticle. Two morphotypes of S. gradata were described from N Italy, the Tortonian Trigonostoma gradatum var. dertumbilicata and the Langhian Trigonostoma gradatum? var. tauroconnectens (Sacco 1894: 10–11, Pl. 1, Figs 21–22); both are small forms with weakly developed spiral sculpture, thin lips, and lack of parietal denticle. S. dertocosticillata (Sacco 1894) – known in Poland in the Badenian Paratethys – is also a similar form but is distinguishable by its shorter last whorl, weaker axial sculpture and lack of parietal denticle. The Middle Miocene S. crassicosta (Bellardi 1841) (Sacco 1894, Pl. 1, Figs 33–35; Ferrero Mortara et al. 1984, Pl. 29, Fig. 7) is also similar in size and overall morphology but differs by its rounded axial ribs, weaker spiral cords, a broader parietal callus, and presence of a small umbilicus. The specimen illustrated by Bohn-Havas (1973, Pl. 6, Figs 12–13) as C. gradata from the Mecsek Mts differs from the type of S. gradata in shell proportion and aperture with parietal denticle; it is considered as a juvenile representative of S. nemethi n. sp. The S. gradata illustrated by Strausz (1966, Pl. 45, Figs 12–13) may also represent the new species. However, this specimen is not available in the museum collections; it is presumed to be lost.
S. nemethi n. sp. is known only in the Pannonian Basin at the moment, but as juvenile specimens are similar to S. gradata it is possible that more occurrences will be recorded in the Central Paratethys.
Distribution: Central Paratethys. Badenian: Pannonian Basin (Hungary: Bakony Mts: Bánd, Börzsöny Mts: Letkés, Mecsek Mts: Pécsszabolcs).
Scalptia neugeboreni (Hörnes 1856) (Fig. 5/E–H)
1856 Cancellaria Neugeboreni – Hörnes, p. 680, Pl. 52, Fig. 6.
2012 Scalptia neugeboreni (Hörnes) – Harzhauser and Landau, p. 46, Fig. 9/C (cum syn.).
2017 Scalptia neugeboreni (Hörnes) – Vicián et al., p. 271, Pl. 3, Fig. 10.
Studied material: 7 specimens (max. SL 22 mm, SW 15.8 mm). HNHM: INV 2020.48. SFI: Krock Coll.: 444611A–B, 444638. Private collections: 3 specimens.
Remarks: S. neugeboreni is distinguished from the similar S. hidasensis in morphology by stronger sculpture with broad primary cords without any secondaries.
Distribution: Central Paratethys. Badenian Vienna Basin (Austria, Czechia), Pannonian Basin (Hungary: Börzsöny Mts: Letkés, Bakony Mts: Bánd), Făget Basin (Romania).
Scalptia polonica (Pusch 1837) (Fig. 5/I–P, Fig. 6/A–B)
1837 Cancellaria acutangula Fauj. var. polonica – Pusch, p. 128, Pl. 11, Fig. 17.
2012 Scalptia polonica (Pusch) – Harzhauser and Landau, p. 39, Figs 3/M, 8/D, E (cum syn.).
2013 Scalptia polonica (Pusch) – Landau et al., p. 233, Pl. 36, Fig. 5.
(A–B) Scalptia polonica (Pusch, SL 19 (HNHM: INV 2020.51). (C–G) Scalptia spinosa (Grateloup), (C–D) SL 24, Letkés (Coll. Hirmetzl), (E–F) SL 24, Tekeres (Coll. Németh), (G) SL 17, Letkés (HNHM, M.62.883). (H–I) Sveltia dertovaricosa (Sacco), SL 20, Tekeres (HNHM: INV 2020.52). (J–M) Sveltia lyrata (Brocchi), Devecser: (J–K) SL 38, (L–M) SL 32 (Coll. Németh). Scale bar: 5 mm
Citation: Central European Geology 64, 2; 10.1556/24.2020.00006
Studied material: 140 specimens (max. SL 37.4 mm, SW 27 mm). HNHM: INV 2020.49–51. SFI: Krock Coll.: 444422 (13 spec.). Private collections: 125 specimens.
Remarks: S. polonica is a very common species in the Badenian Paratethys; however, its presence is recorded for the first time in the Pannonian Basin at Letkés, where it is the most abundant cancellariid (35%). The species shows moderate intraspecific morphological variability, and the large material makes it possible to distinguish two morphotypes. The dominant morphotype A agrees well with the typical form with elongated shell, ovate last whorl, narrow subsutural ramp and ovate aperture (see Harzhauser and Landau 2012, Fig. 8/D, E; Fig. 5/G–J). Morphotype B (13 specimens) is characterized by identical protoconch and sculpture but possesses a broader shell, a more deeply canaliculate infrasutural area, a more angulated shoulder on the last whorl, and wider aperture (Fig. 5/O-P, Fig. 6/A-B)Fig. 5/K–N. This morphology somehow resembles that of S. dertoparva but differs in the protoconch, in the strongly gradate spire and the sculpture. The worn fragmentary specimen illustrated by Kókay (1966, Pl. 9, Fig. 11) as Cancellaria (Trigonostoma) puschi at Bánd represents Ventrilia imbricata (Hörnes).
Distribution: Proto-Mediterranean Sea. Burdigalian: Torino Hills (Italy), Rhône Basin (France), Serravallian–Tortonian: Karaman Basin (Turkey). Central Paratethys. Ottnangian: North Alpine Foreland Basin (S Germany), Karpatian: Pannonian Basin (Hungary). Badenian: Korytnica Basin (Poland), North Alpine Foreland Basin, Eisenstadt–Sopron Basin, Styrian Basin (Austria), Vienna Basin (Austria, Czechia), Pannonian Basin (Hungary: Börzsöny Mts: Letkés, Bakony Mts: Bánd), Făget Basin (Romania), Krka Basin (Slovenia), South Carpathian Foredeep (Bulgaria).
Scalptia spinosa (Grateloup 1827) (Fig. 6/C–G)
1827 Cancellaria Spinosa – Grateloup, p. 21.
1832 Cancellaria spinifera – Grateloup, p. 342.
non 1950 Cancellaria (Trigonostoma) spinifera Grat. – Csepreghy-Meznerics, p. 58, Pl. 3, Fig. 13 (refigured by Strausz 1966, Pl. 45, Fig. 14) [= Scalptia exgeslini (Sacco 1894)].
?1966 Cancellaria (Trigonostoma) spinifera Grateloup – Strausz, p. 377, Text-fig. 164.
1966 Cancellaria (Trigonostoma) ampullacea – Strausz, p. 377, Pl. 45, Figs 17–18 (non Brocchi).
1971–1972 Cancellaria (Trigonostoma) spinifera Grat. – Csepreghy-Meznerics, Pl. 14, Fig. 28.
2012 Scalptia spinosa (Grateloup) – Harzhauser and Landau, p. 41, Figs 3/O, 8/F, G, H (cum syn.).
2013 Scalptia spinosa (Grateloup) – Landau et al., p. 233, Pl. 36, Fig. 8.
2019 Scalptia spinosa (Grateloup) – Cárdenas et al., Fig. 7/B.
Studied material: 10 specimens (max. SL 24 mm, SW 17.6 mm). HNHM: M.62.883, M.62. 884 (2 spec.), INV 2020.55. Private collections: 6 specimens.
Remarks: A taxonomical revision of the species was provided by Cahuzac et al. (2004) and Harzhauser and Landau (2012), in which C. spinifera Grateloup was considered as a junior synonym of S. spinosa. The species is distinguished from Trigonostoma exampullaceum (Sacco) by a higher spire, rounded teleoconch whorls and widely spaced axial ribs. The specimen shown by Strausz (1966, Pl. 45, Figs 17–18) as C. (T.) ampullacea from Letkés represents S. spinosa. The species is uncommon but widespread in the Badenian Pannonian Basin; the appearance in the Tekeres assemblage extends its geographical range.
Miocene distribution: NE Atlantic and Proto-Mediterranean Sea. Burdigalian–Tortonian: Aquitanian Basin (France), Cacela Basin (Portugal), Torino Hills, Po Basin (Italy), Serravallian–Tortonian: Karaman Basin (Turkey), Tortonian: Guadalquivir Basin (Spain). Central Paratethys. Badenian: Korytnica Basin (Poland), North Alpine Foreland Basin, Eisenstadt–Sopron Basin, Styrian Basin (Austria), Vienna Basin (Austria, Czechia), Pannonian Basin (Hungary: Börzsöny Mts: Szob, Letkés, Bükk Mts: Balaton, Mecsek Mts: Tekeres), Făget Basin (Romania).
Genus Sveltia Jousseaume 1887.
Type species: Voluta varricosa Brocchi 1814.
Sveltia dertovaricosa (Sacco 1894) (Fig. 6/H–I)
1894 Cancellaria (Sveltia) dertovaricosa Sacco – Sacco, p. 57, Pl. 3, Fig. 49.
1950 Cancellaria (Sveltia) dertovaricosa Sacco – Csepreghy-Meznerics, p. 59, Pl. 3, Fig. 17.
1966 Cancellaria (Sveltia) varicosa dertovaricosa Sacco – Strausz, p. 380, Pl. 43, Figs 10–12.
1966 Cancellaria (Sveltia) ex aff. dertovaricosa Sacco – Kókay, Pl. 9, Fig. 12.
2012 Sveltia dertovaricosa (Sacco) – Harzhauser and Landau, p. 49, Figs 3/S, 9/H (cum syn.).
2013 Sveltia dertovaricosa (Sacco) – Landau et al., p. 228, Pl. 35, Fig. 9.
2019 Sveltia dertovaricosa (Sacco) – Cárdenas et al., Fig. 7/C.
Studied material: 114 specimens (max. SL 22 mm, SW 10 mm). HNHM: M.62.886; INV 2020.52, 2020.56 (39 spec.). SFI: Krock Coll.: 444516, 444591. Private collections: 71 specimens.
Remarks: A taxonomical revision of the species was carried out by Harzhauser and Landau (2012). The widespread Middle Miocene S. dertovaricosa was frequently synonymized in the literature with the very similar but two-folded Late Miocene–Pliocene S. varicosa (Brocchi 1814); however, dertovaricosa is distinguishable by having three columellar folds (Sacco 1894, Landau et al. 2006). The species is newly recorded at Tekeres, where it is unusually abundant.
Distribution: Proto–Mediterranean Sea. Burdigalian: Torino Hills (Italy), Serravallian–Tortonian: Karaman Basin (Turkey), Gavdos (Greece), Tortonian: Po Basin (Italy), Guadalquivir Basin (Spain). NE Atlantic. Langhian–Serravallian: Aquitanian Basin (France). Central Paratethys. Badenian: North Alpine Foreland Basin, Vienna Basin, Styrian Basin (Austria), Korytnica Basin (Poland), Pannonian Basin (Bosnia, Serbia; Hungary: Börzsöny Mts: Szob, Letkés, Bakony Mts: Bánd, Devecser, Mecsek Mts: Hidas, Tekeres), Făget Basin (Romania), Krka Basin (Slovenia), South Carpathian Foredeep (Bulgaria).
Sveltia lyrata (Brocchi 1814) (Fig. 6/J–M)
1814 Voluta lyrata – Brocchi, p. 311, Pl. 3, Fig. 6.
1956 Cancellaria (Sveltia) lyrata Brocchi – Csepreghy-Meznerics, p. 416, Pl. 9, Figs 22–23.
1966 Cancellaria (Calcarata) lyrata Brocchi – Strausz, p. 382, Pl. 43, Figs 19–24.
1966 Cancellaria (Calcarata) lyrata spinulosa Brocchi – Strausz, p. 382, Pl. 45, Fig. 1.
1971–1972 Cancellaria (Narona) lyrata Brocchi – Csepreghy-Meznerics, Pl. 14, Figs 21, 24.
1971–1972 Cancellaria (Narona) lyrata spinulosa (Brocchi) – Csepreghy-Meznerics, Pl. 14, Figs 25, 27, 29.
2012 Sveltia lyrata (Brocchi) – Harzhauser and Landau, p. 51, Figs 3/T–U, 10/A–B (cum syn.).
Studied material: 150 specimens in private collections (max. SL 38 mm, SW 18.6 mm).
Remarks: S. lyrata is characterized by moderate morphological variability in width of the shell, and development of small spines. The species is common in the Miocene of Europe, and also widespread in the Pannonian Basin.
Distribution: Proto-Mediterranean Sea. Miocene: Torino Hills, Montegibbio (Italy), Late Miocene: Spain. NE Atlantic. Langhian–Serravallian: Aquitanian Basin (France). North Sea Basin: Early–Middle Miocene (Belgium, the Netherlands, Germany, Denmark). Central Paratethys. ?Ottnangian: North Alpine Foreland Basin (Bavaria), Badenian: North Alpine Foreland Basin (Austria), Vienna Basin (Austria, Czechia), Eisenstadt–Sopron Basin (Austria, Hungary: Sopron), Pannonian Basin: Börzsöny Mts: Szob, Bükk Mts: Balaton, Borsodbóta, Csermely, Bakony Mts: Devecser), Făget Basin (Romania), South Carpathian Foredeep (Bulgaria).
Genus Trigonostoma Blainville 1827.
Type species: Delphinula trigonostoma Lamarck 1822.
Trigonostoma exampullaceum (Sacco 1894) (Fig. 7/A–D)
1854 Cancellaria ampullacea var. – Hörnes, p. 321, Pl. 35, Fig. 4.
1894 Cancellaria (Trigonostoma) gradatum forma exampullacea – Sacco, p. 10.
non 1966 Cancellaria (Trigonostoma) ampullacea Brocchi – Strausz, p. 377, Pl. 45, Figs 17–18 [= Scalptia spinosa (Grateloup 1827).
2012 Trigonostoma exampullaceum (Sacco) nov. comb. – Harzhauser and Landau, p. 56, Fig. 10/G (cum syn.).
(A–D) Trigonostoma exampullaceum (Sacco), Letkés: (A–B) SL 25, (Coll. Hirmetzl), (C–D) SL 24 (Coll. Németh). (E–I) Trigonostoma exgeslini (Sacco): (E–F) SL 21, Bánd (Coll. Németh), (G) SL 20, Letkés, (H–I) SL 17, Letkés (Coll. Hirmetzl). (J–L) Ventrilia imbricata (Hörnes): (J–K) SL 30, Bánd (HNHM: INV 2020.53), (L) SL 35, Letkés (Coll. Hirmetzl), (M−O) Tritonoharpa mariechristinae Lesport, Cluzaud and Verhecken, Letkés: (M) SL 8.8 (444202, Coll. Krock, SFI, Photo: Sigrid Hof), (N–O) SL 4 (444349, Coll. Krock, SFI, Photo: Sigrid Hof). Scale bars: 5 mm for A–L; 1 mm for M–O
Citation: Central European Geology 64, 2; 10.1556/24.2020.00006
Studied material: 10 specimens in private collections (max. SL 25.2 mm, SW 20.8 mm).
Remarks: This uncommon species is recorded for the first time in the Pannonian Basin at Letkés. The C. (T.) ampullacea specimen illustrated by Strausz (1966, Pl. 45, Figs 17–18) in fact represents S. spinosa (see above).
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin (Austria), Vienna Basin (Austria, Czechia), Pannonian Basin (Hungary: Letkés).
Trigonostoma exgeslini (Sacco 1894) (Fig. 7/E–I)
1854 Cancellaria Geslini – Hörnes, p. 320, Pl. 35, Fig. 3 (non Basterot).
1894 Cancellaria (Gulia) Geslini forma exgeslini – Sacco, p. 22.
1950 Cancellaria (Trigonostoma) spinifera – Csepreghy-Meznerics, p. 58, Pl. 3, Fig. 13 (refigured by Strausz 1966, Pl. 45, Fig. 14) (non Grateloup).
1966 Cancellaria (Trigonostoma) exgeslini (Sacco) – Kókay, p. 63, Pl. 9, Fig. 10 [non Scalptia gradata Hörnes 1854)].
2012 Trigonostoma exgeslini (Sacco) – Harzhauser and Landau, p. 58, Figs 3/W, 11/A, B (cum syn.).
2013 Trigonostoma exgeslini (Sacco) – Landau et al., p. 231, Pl. 36, Figs 1–2.
Studied material: 73 specimens (max. SL 29 mm, SW 26 mm). SFI: Krock Coll.: 444266, 444666. Private collections: 71 specimens.
Remarks: Trigonostoma exgeslini was recorded by Kókay (1966) in the gastropod assemblage of Bánd; however, Plate 9, Fig. 10 was erroneously marked as C. (T.) gradata Hörnes on the explanation. T. exgeslini forms 37% in the cancellariid fauna of Locality 28. The species is a new occurrence at Letkés. In this material the shells are somewhat variable; the specimens are characterized by strongly angulated to slightly rounded spire whorls.
Distribution: Central Paratethys. Badenian: North Alpine Foreland Basin, Eisenstadt–Sopron Basin, Vienna Basin (Austria), Korytnica Basin (Poland), Pannonian Basin (Hungary: Mecsek Mts: Hidas, Bakony Mts: Bánd, Börzsöny Mts: Letkés), Făget Basin (Romania). Proto-Mediterranean Sea. Serravallian–Tortonian: Karaman Basin (Turkey).
Genus Ventrilia Jousseaume 1887.
Type species: Cancellaria tenera Philippi 1848.
Ventrilia imbricata (Hörnes 1854) (Fig. 7/J–L)
1854 Cancellaria imbricata – Hörnes, p. 327, Pl. 35, Fig. 16.
1950 Cancellaria (Trigonostoma) exgeslini – Csepreghy-Meznerics, p. 58, Pl. 3, Fig. 16 (non Sacco).
v 1966 Cancellaria (Trigonostoma) puschi Hoernes et Auinger – Kókay, p. 63, Pl. 9, Fig. 11.
2012 Ventrilia imbricata (Hörnes) – Harzhauser and Landau, p. 61, Fig. 10/I (cum syn.).
2013 Ventrilia imbricata (Hörnes) – Landau et al., p. 231, Pl. 36, Figs 3–4.
Studied material: 77 specimens (max. SL 35 mm, SW 23.7 mm). HNHM: INV 2020.53. MGSH: 1 specimen without inv. number. SFI: Krock Coll.: 444503. Private collections: 74 specimens.
Remarks: The presence of the species in the Pannonian Basin was recorded by Harzhauser and Landau (2012). V. imbricata was common in the Badenian Central Paratethys; the new records presented herein verify its wide distribution in Hungary as well. The species is one of the most abundant cancellariids in the gastropod assemblage of Bánd, but uncommon at Letkés. The fragmentary specimen illustrated by Kókay (1966, Pl. 9, Fig. 11) as C. (T.) puschi represents V. imbricata.
Miocene distribution: Central Paratethys. Karpatian–Badenian: Korneuburg Basin, North Alpine Foreland Basin, Eisenstadt–Sopron Basin, Styrian Basin (Austria), Vienna Basin (Austria, Czechia), Korytnica Basin (Poland), Pannonian Basin (Hungary: Börzsöny Mts: Letkés, Bakony Mts: Bánd, Mecsek Mts: Hidas), Făget Basin (Romania). Proto-Mediterranean Sea. Serravallian–Tortonian: Karaman Basin (Turkey), Po Basin (Italy). NE Atlantic. Tortonian: Cacela Basin (Portugal).
Subfamily Plesiotritoninae Beu and Maxwell 1987.
Genus Tritonoharpa Dall 1908.
Type species: Tritonoharpa vexillata Dall 1908.
Tritonoharpa mariechristinae Lesport, Cluzaud and Verhecken 2015 (Fig. 7/M−O)
2015 Tritonoharpa mariechristinae nov. sp. – Lesport et al., p. 31, Fig. 10, Pl. 8, Figs 10–12, Pl. 9, Figs 1–5, 7–10.
Studied material: 5 specimens (max. SL 8.8 mm, SW 3.3 mm). SFI: Krock Collection: 444202, 444203, 444323, 444349, 444450.
Remarks: Miocene records of the genus Tritonoharpa are very rare in Europe, and only four species have been designated. The material presented herein was collected by H. Krock at Letkés, and stored in the collection of the Senckenberg Forschungsinstitut. We studied photos of the specimens, courtesy of Ronald Janssen and Sigrid Hof.
The fragmentary Badenian specimens from Poland (Bałuk 1997) probably represent the same taxon. The Chattian–Burdigalian T. aquitaniensis Lesport, Cluzaud and Verhecken (2015) from France is distinguishable by its broader shell, more convex teleoconch whorls and stronger axial and spiral sculpture. The Burdigalian T. speciosum Bellardi 1873 from northern Italy is characterized by its very fine sculpture, while the Serravallian T. alanbeui Landau et al. 2013 from Turkey differs mainly in sculpture. The presence of the NE Atlantic T. mariechristinae in the Central Paratethys remarkably extends the paleogeographic range of the species.
Distribution: NE Atlantic. Langhian: Aquitanian Basin (France). Central Paratethys. Badenian: ?Korytnica Basin (Poland), Pannonian Basin (Hungary: Börzsöny Mts: Letkés).
Conclusion
Twenty-six cancellariid species are recorded from five early Badenian (Middle Miocene) mollusk assemblages in the Hungarian part of the Pannonian Basin. The cancellariid material of Letkés (Börzsöny Mts, N Pannonian Basin) is the most diverse with 21 species, of which eight (Bonellitia austriaca, Contortia callosa, Merica obsoleta, Ovilia excassidea, Scalptia dertoparva, S. polonica, Trigonostoma exampullaceum, Tritonoharpa mariechristinae) are new occurrences for Hungary. Beside Letkés three other Central Paratethyan regions are characterized by similar high diversity of the family. Based on the comprehensive revision of Harzhauser and Landau (2012) and the gastropod collection of the HNHM, the highest diversity can be documented in the Făget Basin (SW Romania) at Coşteiu de Sus with 22 species, while 21 species are known from the mollusk assemblages of Grund (North Alpine Foreland Basin), and in the Vienna Basin (Austria) at Gainfarn and Steinebrunn. Distant parts of the Central Paratethys are characterized by almost identical cancellariid diversity.
In the Herend Basin (Bakony Mts) the cancellariid fauna of Locality 28 at Bánd consists of seven species; the material is of special interest because of the frequent occurrence of Scalptia nemethi n. sp. The material of Devecser (Bakony Mts) is characterized by low diversity (five species); it extends the paleogeographic range of Contortia tortoniana and Pseudobabylonella nysti. The fauna of Tekeres (Mecsek Mts) shows similar low diversity, but Perplicaria mioquadrata is recorded for the first time in the Pannonian Basin. The occurrence of Scalptia hidasensis at Mecsekpölöske extends the geographic range of the species in the Pannonian Basin.
Beside the species described or mentioned in the present paper, two other taxa are known from the early Badenian of Hungary: Solatia exwestiana (Sacco 1894) (= Cancellaria subcancellata infracosticillata in Csepreghy-Meznerics 1969, Pl. 5, Figs 20, 32), and Trigonostoma subsuturale (d’Orbigny 1852) (= C. subcancellata in Csepreghy-Meznerics 1971–1972, Pl. 14, Fig. 23). In the early Middle Miocene 43 Cancellariidae species were recorded from the Central Paratethys by Harzhauser and Landau (2012). Based on new field work, as well as on revisions of museum collections and the previous Hungarian literature, occurrences of 29 species are documented herein from the Hungarian part of the Pannonian Basin.
Acknowledgements
We thank Attila Ősi (Eötvös University, Budapest), Ronald Janssen and Sigrid Hof (Senckenberg Forschungsinstitut, Frankfurt am Main) for their professional help. Private fossil collectors mentioned above kindly offered their gastropod materials for study. Special thanks are due to Tamás Németh (Hungary); this paper would not have been possible without his selfless help. We are grateful to Alfréd Dulai (Hungarian Natural History Museum, Budapest) and Bernard M. Landau (Naturalis Biodiversity Center, Leiden), whose critical comments helped to improve the manuscript. Domonkos Verestói-Kovács (Budapest) contributed to the illustration work.
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