INTRODUCTION

In the Middle Miocene Vienna Basin, which represented an embayment located in the north-western periphery of the Central Paratethys Sea (Kováč et al. 2008, Lambert et al. 2008), the members of the family Turritellidae Lovén, 1847 represented significant elements in the gastropod assemblages (cf. Hörnes 1855, Handmann 1882, Sieber 1958a, 1960, Švagrovský 1981a, 1982, Harzhauser & Landau 2019, Biskupič 2023a, 2023c).

From several fossil-bearing localities in the Slovak part of the Vienna Basin, namely in the area of Devínska Nová Ves (formerly also known as Theben – Neudorf, Neudorf an der March, Dévény – Ujfalu), the borough of the city of Bratislava, Turritellidae gastropods were reported by numerous authors. The first mentions of the Middle Miocene turritellids from the study area were given by Kornhuber (1865) and Fuchs (1868). They were mentioned by Schaffer (1898) and Toula (1900) from a former clay pit of the locality Brickyard, with the description of a new species, Turritella neudorfensis Toula, 1900. Subsequently, Schaffer (1908) and Horusitzky (1917), from the famous fossiliferous site of Sandberg, and Sieber (1958b), from an unspecified locality, briefly reported on the molluscan faunas, including the Turritellidae. Much later, Švagrovský (1981a) studied in detail the molluscan assemblages across a wide area of Devínska Kobyla Hill and identified three turritellid species from Devínska Nová Ves. Three taxa belonging to this gastropod group were listed in the unpublished manuscript of Ondrejičková (1987). Later, the marine faunas collected from several localities of the Devínska Kobyla Hill were analysed by Hyžný et al. (2012), who recognised two Turritellidae species from Sandberg. In the list of the fauna, Ruman & Hudáčková (2015) mentioned six species from the Útočnice locality. Finally, the turritellids originating from Devínska Nová Ves, illustrated in Švagrovský (1981a), were revised by Harzhauser & Landau (2019).

The Turritellidae material reported by Schaffer (1898) and Toula (1900) from Brickyard is unavailable. No additional specimens were found there, as suggested by the research results of Toula (1915), Seneš & Cicha (1973), Švagrovský (1981a), Tomašových (1998) and Ruman & Hudáčková (2015), the author´s field works in the 90s, and the revision of the institutional collection of the Natural History Museum of the Slovak National Museum, Bratislava. Also, no shells are to be found in the Natural History Museum Vienna collections. A poorly known, almost forgotten species described from the locality, Turritella neudorfensis Toula, 1900, is only known from the literature. The species was not included in the comprehensive monograph on Miocene Turritellidae gastropods from the Paratethys Sea by Harzhauser & Landau (2019). Moreover, the shell description of Toula (1900) is very brief and insufficient. Since its discovery, the taxon has not been discussed or re-examined in detail, has remained poorly understood, and its taxonomic status has remained unclear and problematic. According to MolluscaBase (2025), the species is considered a taxon inquirendum. Thus, further re-examination and clarification would be needed. Since the type material seems to be lost, only the original description and illustrations provided by Toula (1900) serve as the basis for the revision presented herein, which limits its convincing genus-level placement.

As this study shows, many specimens stored in institutional collections and some shells published in previous papers have often been incorrectly determined. Although some misidentified species mentioned by Švagrovský (1981a) were clarified by Harzhauser & Landau (2019), an additional, comprehensive re-examination of turritellids from the studied area would be needed. Moreover, other paleontological localities were discovered more recently, yielding new yet unpublished conchological material.

GEOLOGICAL SETTINGS AND STRATIGRAPHY

The surveyed area is situated in the Slovak part of the Vienna Basin (Záhorská nížina Lowland), at Devínska Nová Ves (borough of the city of Bratislava) (Fig. 1). The examined material originates from the Middle Miocene marine deposits, that were stratigraphically ascribed to the Central Paratethyan Upper Badenian (Fig. 2) regional stage (Švagrovský 1981a, Baráth et al. 1994, Tomašových 1998, Holec 2001, Hyžný et al. 2012, Ruman & Hudáčková 2015), corresponding to the early Serravallian in the standard chronostratigraphy (e.g., Harzhauser & Piller 2007, Piller et al. 2007, Kováč et al. 2004, 2017, 2018, Pelech et al. 2021). The sediments are dated to the Bolivina-Bulimina Biozone (e.g., Hudáčková & Kováč 1993, Hudáčková & Spezzaferri 2002, Hyžný et al. 2012) and the NN6 nannoplankton Zone (e.g., Lehotayová 1977, Hudáčková & Spezzaferri 2002, Jamrich & Halásová 2010). The Upper Badenian sediments include the clayey Studienka Member belonging to the Hrušky Formation sensu Kováč et al. (2024), and coarse-grained marginal sediments of the clastic Stupava Formation and the limestone Sandberg Formation (Jamrich et al. 2024).

Fig. 1

Geographic position of the studied localities (red stars) at Devínska Nová Ves (Bratislava): 1 – Brickyard, 2 – Záveterná, 3 – Kasárne, 4 – Bačnegovice, 5 – Útočnice, 6 – Sandberg, 7 – Vodojem (Map source: Google Earth)

Fig. 2

Chronostratigraphic and biostratigraphic zonation of the Badenian (Middle Miocene) Vienna Basin (modified from Harzhauser et al. 2018)

Brickyard. The site is located in the northern part of Devínska Nová Ves (48°13'42.3"N, 16°58'24.4"E), and represents two former clay pits situated on the north margin of Devínska Nová Ves, of which the second was located about 300 m N from the first initial clay pit (Tomašových 1998). The first clay pit was opened in 1870 and was exploited probably until 1925, whilst the new clay pit was opened sometime after 1918 (Zlochová 1998). The first sub-locality no longer exists, and the second is currently abandoned, largely destroyed, and inaccessible. Marine deposits are mainly represented by massive and laminated grey calcareous clays, in places with intercalations of sands and silts (Tomašových 1998). Grey calcareous clays yielded rich invertebrate (e.g., foraminifers, bryozoans, gastropods, bivalves, cephalopods, echinoids, decapods), and vertebrate (bony fishes, sharks) marine faunas studied since the 19th century (e.g., Schaffer 1898, Toula 1900, 1915, Švagrovský 1981a, Činčurová 1990, Holec & Sabol 1996, Tomašových 1998, Chalupová 2001, Hudáčková & Spezzaferri 2002, Gregorová 2009, Ruman & Hudáčková 2015, Košťák et al. 2016, 2018, Harzhauser & Landau 2021, Harzhauser et al. 2022, Biskupič 2023b). Remnants of terrestrial plants were studied by Berger (1951) and Sitár & Kováčová-Slamková (1999).

Záveterná. The site was discovered in 2024 during excavation work in a private backyard on Záveterná Street in Devínska Nová Ves (48°13'03"N, 16°58'29"E). Fine- to coarse-grained sands with sandstone concretions were revealed there. According to the available fossil material, only two molluscan species were recorded: the bivalve Cubitostrea digitata (Eichwald, 1830) and the gastropod Ptychidia vindobonensis (Handmann, 1882). The locality has not yet been surveyed, and the marine micro- and macrofauna have not been evaluated. The age of the sediments is estimated as Badenian, based on the occurrence of the turritellid Ptychidia vindobonensis, which was stratigraphically distributed in the Central Paratethys during the Badenian (Harzhauser & Landau 2019), and on the correlation with the nearby Upper Badenian shallow-water sandy strata revealed at Kasárne and Bačnegovice.

Kasárne. The locality was situated on the eastern margin of Devínska Nová Ves (48°12'58"N, 16°59'27"E), approximately 200 m N from the hill of Dlhý kopec (162 m) and represented several temporal outcrops of a former construction site during the 80s. The marine sediments comprise pale- to ochreous sands and sandstone concretions, and include abundant shallow-water assemblages of bivalves and gastropods. The Badenian age of the deposits was deduced from the presence of some molluscan species (e.g. Aequipecten elegans (Andrzejowski, 1830), Oopecten solarium (Lamarck, 1819), Pecten besseri Andrzejowski, 1830), which typically occurred during the Badenian. The locality description, age estimation and molluscan fauna found there were reported only in the unpublished manuscript of Ondrejičková (1987). Currently, the area is home to the Police Force Secondary Vocational School, and the locality is inaccessible.

Bačnegovice. The locality lies in the eastern part of Devínska Nová Ves (48°12'47"N, 16°59'21"E), and is situated about 150 m SW from the Dlhý kopec hillock (162 m). The site comprises a few fields on a slightly elevated knoll in Bačnegovice, previously known in the literature as Vinohrady (cf. Švagrovský 1981a, Holec 2001) and Útočnica (cf. Meszároš 1986, Ruman & Hudáčková 2015). These deposits are mainly represented by fine-grained yellow sands, silty sands, and white and ochreous, fine- to coarse-grained sands in places intercalated with sandstone concretions; to a small extent, scattered blocks of superjacent corallinacean limestones and discontinuous intercalations of organodetritic corallinacean marls also occur. Species-rich associations of gastropods and bivalves were analysed by Švagrovský (1981a), who recognised 36 molluscan taxa. Anomalies in bivalve shells were discussed by Meszároš (1986); the survey of polyplacophorans was provided by Ruman & Hudáčková (2015), and the Costellariidae gastropods were recently studied by Biskupič (2023d). Rare sharks and fishes were mentioned by Holec (2001).

Útočnice. This relatively little-known paleontological site is located on the eastern periphery of Devínska Nová Ves, approximately 700 m east of the locality Bačnegovice (48°12'51"N, 16°59'58"E), and represents a field on a slightly elevated knoll named Útočnice (152 m). The fossil-bearing marine strata are revealed at the top of the hill, comprising white, occasionally ochreous, fine- to coarse-grained sands, in places intercalated with massive sandstone concretions and intercalations of marly sands, and corallinacean limestones overlying the sandy deposits. The diverse and abundant macrofauna comprised gastropods, bivalves, scaphopods, chitons, scleractinian corals, echinoids, sharks and fishes. At present, only a few works have dealt with marine faunal assemblages, focusing on gastropods (see Biskupič 2023d, 2024b).

Sandberg. This well-known fossil-bearing locality (sand quarry) is situated in the southern periphery of Devínska Nová Ves, on the north-western edge of the Devínska Kobyla Hill (Section 1: 48°12'03"N, 16°58'29"E, Section 2: 48°11'59"N, 16°58'32"E, Section 3: 48°11'51"N, 16°58'38"E). Breccias, conglomerates, sands, organodetritic algal limestones and intercalations of marls were exposed in the section reaching almost 90 m in height (Fig. 3). The faunal assemblages have been studied from the 19th century onwards. These studies have been mainly on marine and terrestrial vertebrates (e.g., Boué 1830, Hauer 1837, Münster 1846, Meyer 1847, Holec & Schlögl 2000, Holec 2001, Sabol & Holec 2002, Holec & Emry 2003), but molluscs (e.g., Hörnes 1848, 1851, 1856, Kornhuber 1865, Horusitzky 1917, Švagrovský 1978, 1981a) have often been the object of paleontological research. Decapods (Lőrenthey & Beurlen 1929, Bachmayer 1962, Hyžný 2011), echinoids (Schaffer 1962, Kroh 2005), bryozoans and brachiopods (Bitner et al. 2014) were studied as well. Geological settings were sketched by several authors (e.g., Švagrovský 1978, 1981a, Seneš & Ondrejičková 1991, Baráth et al. 1994, Holec 2001, Fordinál et al. 2012). New data on the lithostratigraphy, biostratigraphy and faunal assemblages of the locality were provided by Hyžný et al. (2012), Bitner et al. (2014), and Jamrich et al. (2024).

Fig. 3

Simplified lithological scheme of the Upper Badenian section of the locality Sandberg (modified from Baráth et al. 1994 and Hyžný et al. 2012)

Vodojem. The site is located on the southern margin of Devínska Nová Ves and the north slopes of Devínska Kobyla Hill (48°11'57"N, 16°58'54"E). An outcrop is situated in the forest near the reservoir and is about 4 m high and 20 m long. Pale-yellow coarse- to fine-grained sands and sandstone beds in some places with intercalations of sandy-gravel mixture are exposed in the section and contain specimen-rich fossils of marine invertebrates. Gastropods (e.g., Haliotis, Diloma, Archimediella, Helminthia, Cerithiidae indet., Cypraeidae indet.), bivalves (e.g., Glycymeris, Atrina, Flabellipecten, Ostrea, Callista, Acanthocardia, Megacardita, Callista, Panopea, Pholadomya) and rare sand dollar echinoids (Parascutella) characterise the assemblage, indicating shallow-water marine conditions. The locality and its fauna have not previously been studied. Based on the associated marine fauna and lithology, the sediments can be correlated with the shallow-water sandy facies exposed at neighbouring locality Sandberg (SA 1, Sandberg facies 2 sensu Hyžný et al. 2012), indicating a late Badenian age.

MATERIAL AND METHODS

Altogether 1,007 specimens of Turritellidae from Devínska Nová Ves are re-examined. The majority of the studied shells originate from the personal collection of the author, collected over the last three decades. A minor part of the material originates from the institutional collection of the Natural History Museum of the Slovak National Museum (Bratislava), the State Geological Institute of Dionýz Štúr (Bratislava), and the Dúbravka Museum (Bratislava). Additional Turritellidae material was provided by Martin Vlačiky, Jozef Bumbera, Andrej Bernáth (all Bratislava), Jakub Borko (Kanianka), Samuel Veselý (Hrnčiarovce nad Parnou), and Róbert Hodal (Trenčín). The conchological material illustrated in this paper, initially collected by the author, is stored in the institutional collection of the Natural History Museum of the Slovak National Museum, Bratislava. The material from the Brickyard locality mentioned by Schaffer (1898) and Toula (1900) is not available, and, therefore, it is only discussed, not evaluated. The specimens of Turritella neudorfensis Toula, 1900 initially figured in Toula (1900: p. 15, fig. 6a, b) are re-illustrated herein, including the proposed notation of primary spiral cords; the original description in German and its translated version in English are given. Because Toula’s description is relatively brief and insufficient, a revised and more precise description of the shell is provided herein in terms of the shell morphology of Allmon (1996), Landau et al. (2013), Van Dingenen et al. (2016), Landau et al. (2018) and Harzhauser & Landau (2019). The higher systematics of Gastropoda Cuvier, 1795, proposed by Bouchet et al. (2017) and the taxonomic concept of turritellids suggested by Harzhauser & Landau (2019) are followed in this study.

Morphometric abbreviations: SL – shell length, MD – maximum diameter, AA – apical angle, PA – pleural angle. Institutional abbreviations: SNM-PM – Natural History Museum of the Slovak National Museum, Bratislava; ŠGÚDŠ – State Geological Institute of Dionýz Štúr, Bratislava.

SYSTEMATICS

Class Gastropoda Cuvier, 1795

Subclass Caenogastropoda Cox, 1960

Superfamily Cerithioidea Fleming, 1822

Family Turritellidae Lovén, 1847

Subfamily Turritellinae Lovén, 1847

Genus Archimediella Sacco, 1895

Type species: Turritella archimedis Brongniart, 1823, by original designation (Sacco 1895). Rupelian (Early Oligocene), Italy

Archimediella abundans (Handmann, 1882)

(Figs 4–9)

Figs 4–15

Turritellidae species, Devínska Nová Ves, Upper Badenian: 4–5 – Archimediella abundans (Handmann, 1882), Z 41949, Bačnegovice; 6 – Archimediella abundans (Handmann, 1882), Z 41950, Bačnegovice; 7 – Archimediella abundans (Handmann, 1882), Z 41951, Bačnegovice; 8–9 – Archimediella abundans (Handmann, 1882), Z 21177, Kasárne; 10–11 – Archimediella carpathica Harzhauser et Landau, 2019, Z 41952, Útočnice; 12–13 – Archimediella carpathica Harzhauser et Landau, 2019, Z 21177, Kasárne; 14 – Archimediella carpathica Harzhauser et Landau, 2019, Z 21177, Kasárne; 15 – Archimediella carpathica Harzhauser et Landau, 2019, Z 21177, Kasárne. Scale bars 10 mm

[Turritella (Oligodia) Archimedis Brong.] var. abundans – Handmann 1882: p. 217.

Archimediella abundans (Handmann, 1882) new comb. – Harzhauser & Landau 2019: p. 26, figs 6G, 7A–B, 10C, 10D–E, 10F (cum syn.).

Material. Kasárne: 71 specimens, Bačnegovice: 21 specimens, Sandberg: 16 specimens, Vodojem: 1 specimen.

Illustrated material and dimensions. Figs 4–5: Z 41949, SL: 24.4 mm, MD: 12.1 mm (fragment), Bačnegovice; Fig. 6: Z 41950, SL: 19.5 mm, MD: 8.2 mm, Bačnegovice; Fig. 7: Z 41951, SL: 36.3 mm, MD: 11.4 mm, Bačnegovice; Figs 8–9: Z 21177, SL: 31.3 mm, MD: 9.1 mm, Kasárne.

Remarks. Although mostly poorly preserved specimens are available, the overall morphological features of the shells are concordant with those figured in the revised study by Harzhauser & Landau (2019). The shells studied herein represent the first evidence of Archimediella abundans (Handmann, 1882) from Kasárne, Útočnice, and Vodojem. Ruman & Hudáčková (2015) mentioned this species from Bačnegovice and Hyžný et al. (2012) from Sandberg as Turritella (Archimediella) erronea Cossmann in Friedberg, which is a junior synonym of A. abundans (Handmann, 1882) (Harzhauser & Landau 2019). The species was most likely identified from Brickyard already by Schaffer (1898) and Toula (1900), but under the name Turritella archimedis Brong. However, the species-level affiliation of these specimens cannot be verified because the material was not illustrated in their papers, and is not available in any institutional collections.

Stratigraphic and geographic distribution. In the Slovak part of the Western Carpathians, this turritellid was found in the Badenian (Middle Miocene) deposits of the eastern Vienna Basin at Kúty (drill well Kúty-45) (Ruman & Hudáčková 2015), Rohožník – Konopiská (Fuksi 2015a), Pernek (own data), Devínska Nová Ves (unspecified locality) (Sieber 1958b), Brickyard? (Schaffer 1898, Toula 1900), Sandberg (Hyžný et al. 2012), Devín – northeast from Devín-Terasy (Švagrovský 1981a), Lomnická, Terasy, Šibeničník (Hyžný et al. 2012), from the Danube Basin at Želiezovce – Lontov (drill well ŽI-2) (Tejkal 1968, Brestenská 1978b) and also from the Novohrad-Nógrad Basin near Kosihovce (Hano 1950), Stredné Plachtince, and Horné Strháre (Čechovič & Seneš 1950).

The species was distributed from the Karpatian (Early Miocene) to the Badenian (Middle Miocene) in the Central Paratethys, and from the Late Miocene (Tortonian) to the Pliocene in the Proto-Mediterranean (Harzhauser & Landau 2019).

Archimediella carpathica Harzhauser et Landau, 2019

(Figs 10–15)

Archimediella carpathica Harzhauser & Landau 2019: p. 30, figs 6H, 7C, 11A–G (cum syn.)

Archimediella carpathica Harzhauser et Landau, 2019 – Koubová et al. 2024: fig. 8H

Material. Kasárne: 7 specimens, Bačnegovice: 1 specimen, Útočnice: 9 specimens, Sandberg: 5 specimens.

Illustrated material and dimensions. Figs 10–11: Z 41952, SL: 29.1 mm, MD: 10.1 mm, Útočnice; Figs 12–13: Z 21177, SL: 28.8 mm, MD: 9.1 mm, Kasárne; Fig. 14: Z 21177, SL: 29.2 mm, 9.3 mm, Kasárne; Fig. 15: Z 21177, SL: 31.3 mm, 9.1 mm, Kasárne.

Remarks. For a full list of synonyms and a comprehensive discussion on related species, see Harzhauser & Landau (2019). The available conchological material agrees with the morphological features of the specimens illustrated by those authors. From Kasárne, Bačnegovice, Útočnice, and Sandberg, the first records of Archimediella carpathica Harzhauser et Landau, 2019 are confirmed herein.

From Brickyard, the taxon Turritella pythagoraica Hilber was reported by Schaffer (1898). This species, considered a junior synonym of Archimediella indigena (Eichwald, 1830), known from the Middle Miocene of the Central and Eastern Paratethys, was often confused with A. carpathica in the malacological literature, as stated by Harzhauser & Landau (2019). I have not seen the specimens from Brickyard, but the shell characters given in Schaffer’s (1898) shell description could indicate a possible placement in Archimediella carpathica. Nevertheless, his description is brief, and no specimen is illustrated in his paper, which does not allow for verifying its species affiliation with certainty.

Stratigraphic and geographic distribution. In Slovakia, this taxon was identified from the Badenian (Middle Miocene) sediments of the Vienna Basin at Borský Mikuláš – Vinohrádky (Švagrovský 1982), Plavecký Mikuláš (Buday 1939), Kúty (drill well Kúty-45) (Ruman & Hudáčková 2015), and Devín – Zelené terasy (Madarás et al. 2014, Biskupič 2023a, Jamrich et al. 2024), in the Danube Basin at Modra – Kráľová (Hörnes 1855, Hladilová & Fordinál 2013), Dubová – Malý trávnik (Ruman & Hudáčková 2015, Koubová et al. 2024), Smolenice (Zlinská & Fordinál 1992), Želiezovce – Lontov (drill well ŽI-2) (Tejkal 1968, Brestenská 1978b), Salka (drill well K-5) (Brestenská 1978a), Salka – Vineyards, Bajtava, and Malá nad Hronom (Seneš 1949), in the Novohrad-Nógrad Basin at Kosihovce (Hano 1950), Pôtor, Ľuboriečka, and Horné Strháre (Čechovič & Seneš 1950) and also from the Eastern Slovakian Basin from Kuzmice (Švagrovský 1960, 1964).

This turritellid species was a common component in the molluscan assemblages of the Central Paratethys Sea during the Badenian (Middle Miocene). Its occurrence during the Tarkhanian (Middle Miocene) of the Eastern Paratethys is uncertain (Harzhauser & Landau 2019).

Genus Helminthia Handmann, 1882

Type species: Turbo vermicularis Brocchi, 1814, by subsequent designation (Landau et al. 2013). Miocene, Italy.

Helminthia tricincta (Borson, 1821)

(Figs 16–20)

Figs 16–27

Turritellidae species, Devínska Nová Ves, Upper Badenian: 16 – Helminthia tricincta (Borson, 1821), Z 17342, Sandberg; 17 – Helminthia tricincta (Borson, 1821), Z 17219, Sandberg; 18 – Helminthia tricincta (Borson, 1821), Z 17937, Sandberg; 19 – Helminthia tricincta (Borson, 1821) Z 16986, Sandberg; 20 – Helminthia tricincta (Borson, 1821) Z 16986, Sandberg; 21–22 – Helminthia vermicularis (Brocchi, 1814), Z 21705, Sandberg (illustrated as Turritella (Haustator) tricincta Borson, 1821 in Švagrovský 1981a, pl. 39, fig. 1); 23–24 – Helminthia vermicularis (Brocchi, 1814), Z 41953, Útočnice; 25 – Helminthia vermicularis (Brocchi, 1814), Z 41954, Útočnice; 26 – Helminthia vermicularis (Brocchi, 1814), Z 41955, Útočnice; 27 – Helminthia vermicularis (Brocchi, 1814), Z 41956, Útočnice. Scale bars 10 mm

Turritella tricincta Borson 1821: p. 342, pl. 6, fig. 11.

Helminthia tricincta (Borson, 1821) – Harzhauser & Landau 2019: p. 38, figs 6I, 13A–B, 13C (cum syn.)

Helminthia tricincta (Borson, 1821) – Biskupič 2023c: p. 14., figs 4A–H, 5A–F, 6A–F, 8A–B, 9A–D

Material. Sandberg: 41 specimens.

Illustrated material and dimensions. Fig. 16: Z 17342, SL: 50.8 mm, MD: 27.4 mm (fragment), Sandberg; Fig. 17: Z 17219, SL: 43.7 mm, MD: 17.2 mm, Sandberg; Fig. 18: Z 17937, SL: 29.9 mm, MD: 14.9 mm, Sandberg; Fig. 19: Z 16986, SL: 21.8 mm, MD: 11.7 mm, Sandberg; Fig 20: Z 16986, SL: 19.4 mm, MD: 7.6 mm (juvenile specimen), Sandberg.

Remarks. The material found at Sandberg is mostly poorly preserved; the aragonite shells are often dissolved or recrystallised, sometimes preserved only as inner moulds, or as imprints of the shells with sculpture in the sandstones. Nevertheless, their shell morphology (large-sized, robust, tricostate shells) is concordant with that reported by Hörnes (1855), Sieber (1960), Harzhauser & Landau (2019), and Biskupič (2023c). Švagrovský (1981a) mentioned the species from Vineyards (=Bačnegovice) and Sandberg. However, the material from Bačnegovice belongs to H. vermicularis (Brocchi, 1814) and only fossils from Sandberg agree with Helminthia tricincta (Borson, 1821), as suggested by the revision of his material stored in the collections of SNM-PM. This species most probably also occurred at Brickyard; it was mentioned under the name Turritella riepeli Partsch by Schaffer (1898) and Toula (1900). From the Upper Badenian locality Rohožník – Konopiská, located on the eastern Vienna Basin, Biskupič (2023c) distinguished two morphologically distinct morphs within the species based on conchological characters: the smooth-sculptured and the coarse-sculptured morphs. In the material from Devínska Nová Ves, only a typically coarse-sculptured morph was detected.

Stratigraphic and geographic distribution. As summarised by Biskupič (2023c), the species was found in Slovakia at only a few Middle Miocene (Badenian) sites in the eastern Vienna Basin.

Helminthia tricincta (Borson, 1821) was widespread in the Middle Miocene (Badenian) of the Central Paratethys and the Tarkhanian (Early to Middle Miocene) of the Eastern Paratethys. The species also occurred in the Tortonian (Late Miocene) and the Pliocene of the Proto-Mediterranean Sea (Harzhauser & Landau 2019).

Helminthia vermicularis (Brocchi, 1814)

(Figs 21–27)

Turbo vermicularis Brocchi 1814: p. 372, pl. 6, fig. 13.

Helminthia vermicularis (Brocchi, 1814) – Harzhauser & Landau 2019: p. 41, figs 6K, 7G–H, 13D–E, 13F (cum syn.)

Helminthia vermicularis (Brocchi, 1814) – Biskupič 2023c: p. 16, figs 7A–G, 8C–D, 9E–F

Material. Kasárne: 14 specimens, Bačnegovice: 66 specimens, Útočnice: 283 specimens, Sandberg: 18 specimens, Vodojem: 1 specimen.

Illustrated material and dimensions. Figs 21–22: Z 21705, SL: 42.8 mm, MD: 17.8 mm, Sandberg (illustrated as Turritella (Haustator) tricincta Borson, 1821 in Švagrovský 1981a, pl. 39, fig. 1); Figs 23–24: Z 41953, SL: 38 mm, MD: 12.3 mm, Útočnice; Fig. 25: Z 41954, SL: 40.1 mm, MD: 12.8 mm, Útočnice; Fig. 26: Z 41955, SL: 35.6 mm, MD: 12.3 mm, Útočnice; Fig. 27: Z 41956, SL: 39.3 mm, MD: 12.1 mm, Útočnice.

Remarks. Even though Švagrovský (1981a) determined these shells as Turritella (Haustator) tricincta Borson, 1821 from Vineyards (=Bačnegovice) and Sandberg, a part of the material is ascribed to H. vermicularis (Brocchi, 1814) in this study based on the revision of his conchological material deposited in the SNM-PM collection. The species is among the most abundant Turritellidae species at Bačnegovice and Útočnice. H. vermicularis (Brocchi, 1814) was listed from Brickyard in the papers of Schaffer (1898) and Toula (1900).

Stratigraphic and geographic distribution. From the Slovak part of the Western Carpathians, the species is known from several Middle Miocene (Badenian) localities of the Vienna Basin and the Novohrad-Nógrád Basin, as summarised by Biskupič (2023c). As reported by Harzhauser & Landau (2019), Helminthia vermicularis (Brocchi, 1814) occurred abundantly during the Middle Miocene (Badenian) through the Central Paratethys Sea and was also distributed from the Early Miocene (Burdigalian) to the Pleistocene in the Proto-Mediterranean Sea and the northeastern Atlantic.

Genus Oligodia Handmann, 1882

Type species: Turritella bicarinata Eichwald, 1830, by subsequent designation (Landau et al. 2013). Middle Miocene, Paratethys Sea.

Oligodia bicarinata (Eichwald, 1830)

(Figs 28–34)

Figs 28–40

Turritellidae species, Devínska Nová Ves, Upper Badenian: 28–29 – Oligodia bicarinata (Eichwald, 1830), Z 21791, Sandberg; 30–31 – Oligodia bicarinata (Eichwald, 1830) Z41957, Útočnice; 32 – Oligodia bicarinata (Eichwald, 1830), Z 41958, Útočnice; 33–34 – Oligodia bicarinata (Eichwald, 1830), Z 41959, Útočnice; 35 – Ptychidia vindobonensis (Handmann, 1882), Z 40757, Záveterná; 36 – Ptychidia vindobonensis (Handmann, 1882), Z 40757, Záveterná; 37–38 – Viennella sp., Z 41960, Bačnegovice; 39 – “Turritella” neudorfensis Toula, 1900, Brickyard (the reproduction of the original illustration of the syntype figured by Toula (1900); SL, MD, and the scale bar are unknown); 40 – “Turritella” neudorfensis Toula, 1900, detail of the teleoconch whorl with the addition of the notation of primary spiral cords (B, C/d), Brickyard (the reproduction of the original illustration of the syntype figured by Toula (1900); SL, MD, and the scale bar are unknown). Scale bars 10 mm (Figs 28–36) and 5 mm (Figs 37–38)

Turritella bicarinata Eichwald 1830: p. 220.

Oligodia bicarinata (Eichwald, 1830) – Harzhauser & Landau 2019: p. 50, figs 6P, 7H, 16A–E (cum syn.)

Material. Bačnegovice: 50 specimens, Útočnice: 307 specimens, Sandberg: 80 specimens.

Illustrated material and dimensions. Figs 28–29: Z 21791, SL: 18.7 mm, MD: 6.5 mm, Sandberg; Figs 30–31: Z 41957, SL: 28.1 mm, MD: 10.6 mm, Útočnice; Fig. 32: Z 41958, SL: 23.5 mm, MD: 9.7 mm, Útočnice; Figs 33–34: Z 41959, SL: 22.9 mm, MD: 9.3 mm, Útočnice.

Remarks. Švagrovský (1981a) erroneously identified this species as Turritella (Zaria) spirata (Brocchi, 1814) from Vineyards (= Bačnegovice) based on a few small shell fragments. The revision of his material stored in SNM-PM clearly suggests that these specimens represent just spire fragments of O. bicarinata (Eichwald, 1830). The species was identified from Kasárne by Ondrejičková (1987). However, her conchological material deposited in the collections of ŠGÚDŠ comprised no specimens concordant with O. bicarinata (Eichwald, 1830), and thus the occurrence of the species at Kasárne is unconfirmed in this paper. These specimens were probably confused with the Archimediella species that occur abundantly at the locality.

Stratigraphic and geographic distribution. From Slovakia, this turritellid was mentioned from the Badenian (Middle Miocene) deposits of the Vienna Basin, and was identified from Borský Mikuláš (Švagrovský 1982), Kuchyňa (Buday 1939), Stupava – Vrchná hora (Buday 1939), Stupava – drill well HGP-3 (Fordinál et al. 2003), Devínska Nová Ves (unspecified locality) (Sieber 1958b), Devínska Nová Ves – railway sand pits (Fuchs 1868), Sandberg (Švagrovský 1981a, Hyžný et al. 2012), Bačnegovice (Švagrovský 1981a, Ruman & Hudáčková 2015), Devín – northeast from Devín-Terasy (Švagrovský 1981a), and Kúty (drill well Kúty-45) (Ruman & Hudáčková 2015), from the Danube Basin at Trstín (Fuksi 2015b), Chľaba (drill well ŠO-1) (Ondrejičková 1978), Salka – Vineyards, and Malá nad Hronom (Seneš 1949) and also from the Novohrad-Nógrad Basin near Kosihovce (Hano 1950) and Horné Strháre (Čechovič & Seneš 1950).

According to Harzhauser & Landau (2019), the Central Paratethyan records of this well-known species are known from the Karpatian (Early Miocene) and the Badenian (Middle Miocene); it was widespread in the Tschokrakian and Konkian (Middle Miocene) of the Eastern Paratethys and also from the Burdigalian (Early Miocene) to the Tortonian (Late Miocene) in the Proto-Mediterranean Sea and in the northeastern Atlantic during the Langhian to the Tortonian (Middle – Late Miocene).

Genus Ptychidia Handmann, 1882

Type species: Turritella (Ptychidia) vindobonensis Handmann, 1882, by monotypy (Handmann 1882). Middle Miocene, Paratethys Sea.

Ptychidia vindobonensis (Handmann, 1882)

(Figs 35–36)

Turritella (Ptychidia) vindobonensis Partsch forma typica Handmann 1882: p. 215 [new name for Turritella turris Bast. (Hörnes 1855: pl. 43, fig. 15)]

Ptychidia vindobonensis (Handmann, 1882) – Harzhauser & Landau 2019: p. 62, figs 6U, 7J, 18A–D, 18O (cum syn.)

Material. Záveterná: 14 specimens, Bačnegovice: 1 specimen.

Illustrated material and dimensions. Fig. 35: Z 40757, SL: 59.6 mm, MD: 13.8 mm, Záveterná; Fig. 36: Z 40757, SL: 53.3 mm, MD: 10.2 mm, Záveterná.

Remarks. Only a single shell from Bačnegovice is available, which points to the low abundance of the species at the locality. In contrast, the species seems to be the dominant turritellid at Záveterná. From Brickyard, Schaffer (1898) reported one fragmentary shell ascribed to Turritella turris Bast., which has often been confused with Ptychidia vindobonensis (Handmann, 1882) by the early authors (see discussion in Landau et al. 2013 and Harzhauser & Landau 2019). However, according to his short description, the assumed affiliation with Ptychidia vindobonensis is doubtful, as the specimen may also represent another similar Badenian species, Ptychidia erynella Harzhauser et Landau, 2019 (cf. Harzhauser & Landau 2019: figs 18E–H, J–K). Moreover, Schaffer´s specimen is not available for further re-examination, making its reliable placement in the particular species impossible. The species was also reported from Devínska Nová Ves by Sieber (1958b), but no exact locality was given in his paper. Although this taxon was documented from Kasárne by Ondrejičková (1987), the revision of her material stored in the collections of ŠGÚDŠ reveals no evidence of the species at the locality.

Tejkal (1968) was the first to clarify the taxonomic status of the species and consider Turritella turris var. badensis Sacco, 1895 as a junior synonym of Turritella (Ptychidia) vindobonensis Handmann, 1882. This opinion was much later confirmed by Landau et al. (2013) and Harzhauser & Landau (2019). The Handmann’s (1882) proposed new subgenus name, Ptychidia, was accepted and elevated to full genus rank by Landau et al. (2013). For a complete list of synonyms and a discussion on the taxonomy of the species, see the monographs of those authors.

Stratigraphic and geographic distribution. From the Slovak part of the Western Carpathians, this taxon was reported from the Badenian (Middle Miocene) deposits of the eastern Vienna Basin, namely from Rohožník – Konopiská (Fuksi 2015a), Devínska Nová Ves (unspecified locality) (Sieber 1958b), and Bačnegovice (Ruman & Hudáčková 2015), from the Danube Basin at Želiezovce – Lontov (drill well ŽI-2) (Tejkal 1968, Brestenská 1978b), Chľaba (drill well ŠO-1) (Ondrejičková 1978, Holcová et al. 2019), Malá nad Hronom, and Kamenica nad Hronom (Seneš 1949), and also from the Novohrad-Nógrad Basin near Ľuboriečka, Horné Plachtince, and Modrý Kameň (Čechovič & Seneš 1950).

P. vindobonensis is known from the Badenian (Middle Miocene) of the Central Paratethys Sea (Harzhauser & Landau 2019) and was also recorded in the Serravallian (Middle Miocene) of the Proto-Mediterranean Sea and northeastern Atlantic (Landau et al. 2013).

Genus Viennella Harzhauser et Landau, 2019

Type species: Turritella incisaeformis Csepreghy-Meznerics, 1956, by original designation (Harzhauser & Landau 2019). Middle Miocene (Badenian), Hungary, Central Paratethys Sea.

Viennella sp.

(Figs 37–38)

Material. Bačnegovice: 1 specimen.

Illustrated material and dimensions. Figs 37–38: Z 41960, SL: 13.2 mm, MD: 4.4 mm, Bačnegovice.

Remarks. The shell is almost complete, but the spiral sculpture is poorly preserved, mostly abraded, and only visible in places. Early teleoconch whorls seem flat-sided, later whorls are slightly convex and bear seven spiral cords with smooth interspaces between them. This specimen resembles in shell size, shape of teleooconch whorls, and spiral sculpture the species Viennella incisaeformis (Csepreghy-Meznerics, 1956) (cf. Harzhauser & Landau 2019: p. 78, figs 21A–D), known from the Lower Badenian (Middle Miocene) marine sediments of the Vienna Basin (Austria) and the Pannonian Basin (Hungary). However, due to poor preservation of the shell, reliable species-level identification is not possible, and thus it remains left in open nomenclature. Although another Upper Badenian congener, Viennella baluki Biskupič, 2024, is known from the Slovak part of the Vienna Basin (Rohožník – Konopiská), the specimen from Devínska Nová Ves has little in common with it. Viennella baluki clearly differs from the studied shell in its more convex-sided teleoconch whorls, the presence of tertiary spiral cords, prominent mid-whorl angulation and strongly raised spiral sculpture (cf. Biskupič 2024a: figs 3–5). Viennella sp. represents the first evidence for the genus from Devínska Nová Ves.

“Turritella” neudorfensis Toula, 1900, taxon inquirendum

(Figs 39–40)

Turritella neudorfensisToula1900: p. 15.

Material. The specimens illustrated by Toula (1900: fig. 6a, b). Repository data and the shell measurements are unknown.

Original description (Toula 1900: p. 15). “Die Umgänge sind flach und mit etwa 30 nicht ganz gleichen Spirallinien bedeckt. Gegen die Mündung zu lassen sich zwei wenig ausgeprägte Spiralkiele erkennen.” [The whorls are flat and covered by about 30 not-quite-equal spiral lines. Towards the aperture, two less pronounced spiral keels can be seen].

Emended description. Shell incomplete, robust, moderately slender, composed of five teleoconch whorls; protoconch and early teleoconch whorls are missing, order of appearance of spiral cords unknown. Shell dimensions (SL, MD) unknown. AA unknown, PA = 17°. First known teleoconch whorl abraded. Later teleoconch whorls stocky, broad, weakly convex in shape, with maximum diameter bellow mid-whorl. Whorl profile imbricate to slightly frustate, bicarinate, sutural ramp somewhat concave. Teleoconch whorls covered by two indistinct, low, suppressed primary spiral cords; medial primary B somewhat prominent, thin, forming little pronounced mid-whorl angulation, primary cord (C/d) close abapical suture slightly depressed. Primaries separated by broad, shallow, weakly concave to straight-sided interspaces. Suture linear, incised, well defined. Whorl surfaces covered by densely spaced growth lines and numerous, about 30, fine secondary and tertiary spiral threads, nonuniform in strength, separated by narrow furrows. Delicate, indistinct tubercles or nodules developed in intersections of growth lines and spiral threads. Smooth, flat, narrow spiral band running below adapical suture. Transition into base angulate based on last preserved teleoconch whorl. Last adult whorl and aperture destroyed, aperture shape and internal sculpture unknown. Lateral sinus probably in mid-whorl, inflection points not recognised, base and basal sinus unknown.

Remarks. According to the figure of Toula (1900: fig. 6a, b), the neanic teleoconch whorls and the last adult whorl are missing. Several morphological features are unavailable, such as the protoconch, the order of appearance of the spiral cords, the sculpture of early teleoconch whorls, the lateral and basal sinus, and the aperture shape; also, the presence/absence of internal sculpture cannot be confirmed. In the illustration, the growth lines are not so clearly sketched, so it is impossible to define the lateral sinus. In addition, the shell length, maximum diameter, and scale bar were not given. Another problem is that defining the primary spiral cord running close to the abapical suture is not reliably possible. It may represent spiral cord C if the species belongs to Archimediella or peribasal spiral cord d if it represents an Oligodia. On the other hand, the shape of teleoconch whorls and primary spiral cords are well-defined in the illustration and may allow its possible genus-level placement. The shell shows the imbricate, weakly bicarinate profile of teleoconch whorls and slightly concave sutural ramp, and its whorls bear two primary spiral cords. At first sight, the overall shell morphology seems concordant with the genus Oligodia Handmann, 1882, in the sense of Landau et al. (2013), Van Dingenen et al. (2016), and Harzhauser & Landau (2019). However, due to a lack of important diagnostic features, such as the order of appearance of the spiral cords, sculpture of early whorls, and the lateral and basal sinuses, its placement in the proposed genus is not formally possible.

Although this species was initially placed in the genus Turritella Lamarck, 1799 by Toula (1900), it has little in common with it. Representatives of Turritella featured their large-sized shells, a frustate to campanulate spire whorl profile, and well-defined primary and secondary spiral cords on early teleoconch whorls, which become faintly obsolete to attenuated in later whorls, and in having the order of appearance of primary spiral cords B-A-C (cf. Harzhauser & Landau 2019). Because the clear generic relation of the species is uncertain, it remains provisionally placed in “Turritella” herein for now.

It is evident that “Turritella” neudorfensis Toula, 1900 morphologically resembles the genus Oligodia Handmann, 1882. Oligodia bicarinata (Eichwald, 1830), which commonly occurred in the Paratethyan, Proto-Mediterranean, and northeastern Atlantic Miocene (cf. Landau et al. 2013, 2018, Harzhauser & Landau 2019), is a similar species that from “T.” neudorfensis differs in its typical bicarinate whorl profile and two prominent, more thickened and strongly raised primary spiral cords B and d, sometimes separated by deep, concave, and wide interspaces, making the overall sculpture appearance more coarse; also, the suture and sutural ramp are more deeply incised, and spiral threads are more delicate and uniform (cf. Harzhauser & Landau 2019: figs 16A–E). Oligodia spirata (Brocchi, 1814), a well-known Neogene European turritellid (cf. Landau et al. 2004, Van Dingenen et al. 2016, Harzhauser & Landau 2019), is another superficially similar species that has teleoconch whorls decorated by a very prominent carina forming strong mid-whorl angulation, the sutural ramp and abapical half of the whorls are more concave, and the tertiary spiral threads seem more delicate (cf. Harzhauser & Landau 2019: figs 16F–G), which clearly differs from the species from Devínska Nová Ves. Toula (1900) noticed some conchological similarities between Turritella neudorfensis and the shell from Steinebrunn (Vienna Basin, Austria) figured by Hörnes (1855: pl. 43, fig. 4) as Turritella marginalis Brocci var., which was later distinguished by Sacco (1895) as a new species named Turritella (Haustator) sulcomarginalis, and more recently transferred into Oligodia by Harzhauser & Landau (2019). O. sulcomarginalis (Sacco, 1895) from the Vienna Basin is a distinctly different species that has a much more slender shell with discernible medial primary B forming moderate mid-whorl angulation developed on nearly flat-sided teleoconch whorls (cf. Harzhauser & Landau 2019: figs 16H–J). The herein-discussed “Turritella” from Devínska Nová Ves is also slightly similar to O. guillaumei (Brébion in Lauriat-Rage et al. 1989) and O. palumbina Van Dingenen, Ceulemans et Landau, 2016, both known from the Lower Pliocene of the Loire Basin (France). O. guillaumei is a superficially similar species characterised by its slightly concave latest teleoconch whorls (cf. Van Dingenen et al. 2016: pl. 3, figs 3–5). O. palumbina has similar flat-sided whorls but has a much taller and slimmer shell, slightly higher whorls and considerably faint to obsolete primaries, which are weakly developed on post-neanic teleoconch whorls (cf. Van Dingenen et al. 2016: pl. 3, figs 6–7). O. chauvereauensis Landau, Ceulemans et Van Dingenen, 2018, described from the Tortonian (Upper Miocene) of northwestern France, is a quite different turritellid, featured by its relatively broad turriculate shell, strongly carinate teleoconch whorls with prominent medial primary B placed in mid-whorl, and impressed suture (cf. Landau et al. 2018: pl. 17, figs 1–3).

The genus Archimediella Sacco, 1895 is a partly similar turritellid whose members differ from “T.” neudorfensis in their coarse spiral sculpture on teleoconch whorls, by the presence of prominent spiral cords A, B and C and numerous nonuniform secondary and tertiary spiral cords of variable strength covering the surface of teleoconch whorls (see Landau et al. 2018, Harzhauser & Landau 2019). For a complete description of diagnostic features that define the genus, see Harzhauser & Landau (2019).

Stratigraphic and geographic distribution. This Late Badenian (Middle Miocene) Central Paratethyan species is known only from the type locality Devínska Nová Ves – Brickyard (former clay pit), Vienna Basin, Slovakia.

TAXONOMIC COMPOSITION

Each locality is characterised by a distinct and unique diversity of turritellids (Fig. 41). Their species richness and abundance may be related to distinct ecological factors and to changes in these factors in depositional environments during the Upper Badenian. Seven Turritellidae species were recognised from these localities. The taxonomic composition of Turritellidae assemblages, except for Brickyard, is evaluated herein. Although several species of turritellids were briefly mentioned by Schaffer (1898) and Toula (1900) from this site, they are not included in the statistical evaluation because the data reported by those authors are poor, they cannot be verified, and no further material is available for re-examination. It is worth noting that similar high-specimen-rich assemblages, comparable to those of Turritellidae, were observed in other gastropod families, e.g., Cerithiidae, Costellariidae, Ancillariidae, and Conidae at Útočnice, and in Turbinidae at Sandberg and Bačnegovice.

Fig. 41

Taxonomic composition and percentage representation of studied Turritellidae species at respective localities of Devínska Nová Ves

Záveterná locality is characterised by a monotypic Turritellidae association, with a single species, Ptychidia vindobonensis (Handmann, 1882) (14 specimens). The assemblage of Kasárne is characterised by the dominance of Archimediella abundans (Handmann, 1882) (71 specimens), accompanied by Helminthia vermicularis (Brocchi, 1814) (14 specimens) and rare A. carpathica Harzhauser et Landau, 2019 (7 specimens). At Bačnegovice, Helminthia vermicularis (Brocchi, 1814) (66 specimens) and Oligodia bicarinata (Eichwald, 1830) (50 specimens) represent the most common Turritellidae taxa, followed by Archimediella abundans (Handmann, 1882) (21 specimens), and rare A. carpathica Harzhauser et Landau, 2019 (1 specimen), Ptychidia vindobonensis (Handmann, 1882) (1 specimen), and Viennella sp. (1 specimen). Partly similar taxonomic composition was observed at the neighbourhood site Útočnice, where O. bicarinata (307 specimens) and H. vermicularis (283 specimens) prevailed in the turritellid assemblage, whereas A. carpathica (9 specimens) occurred rarely. The assemblages of Sandberg were characterised by the dominating O. bicarinata (80 specimens), followed by Helminthia tricincta (Borson, 1821) (41 specimens), H. vermicularis (Brocchi, 1814) (18 specimens), Archimediella abundans (Handmann, 1882) (16 specimens), and A. carpathica Harzhauser et Landau, 2019 (5 specimens). At Vodojem, two rare species occur: H. vermicularis (1 specimen) and A. abundans (Handmann, 1882) (1 specimen).

PALEOECOLOGY

The studied Turritellidae communities inhabited primarily shallow-water marine paleoenvironments with soft, sandy sea bottoms (Fig. 43). They were often associated with seagrass habitats, where they reached their highest abundance and diversity. Four paleoenvironments with Turritellidae occurrence were recognised at the respective sites, ranging from intertidal to infralittoral zones (Fig. 42). Deeper, middle neritic paleoenvironments characterised by muddy sea bottom and influenced by occasional hypoxic events were identified at the locality Brickyard. In these habitats, the herein discussed “Turritella” neudorfensis Toula, 1900, occurred. However, because the Turritellidae are known only from the literature and no further material is available from the locality, they are not statistically evaluated.

Fig. 42

Occurrence and percentage of respective Turritellidae species in the Upper Badenian marine paleoenvironments of Devínska Nová Ves

Fig. 43

Simplified, idealised reconstruction of the Upper Badenian shelf and its various paleoenvironments on the eastern margin of the Vienna Basin at Devínska Nová Ves, showing the occurrence of the respective Turritellidae species: 1 – sandy coast (intertidal to upper infralittoral zone), 2 – sandy seabottom with seagrass and corals (upper infralittoral zone), 3 – sandy seabottom with seagrass (infralittoral zone), 4 – sandy to sandy-silty seabottom with a huge seagrass meadows (infralittoral zone), 5 – sandy to clayey seabottom with corallinacean red algae and seagrass (infralittoral zone), 6 – muddy seabottom, paleoenvironment affected by sluggish water circulation and episodic hypoxic events (middle to outer neritic zone)

INTERTIDAL TO UPPER INFRALITTORAL ZONE – SANDY COASTAL ENVIRONMENT

Coarse-grained pale-grey sands with sandstone beds and lenses of gravel exposed at Sandberg (SA 2, Sandberg facies 2 and 3 sensu Hyžný et al. 2012) indicate coastal sandy environments. Marine invertebrates (gastropods, bivalves, echinoids, decapods) and vertebrates (sharks, rays, bony fishes) are only rarely present and sparsely distributed, and the assemblage shows a strong decrease in diversity and abundance. Some layers of the Sandberg facies 3 are in places almost sterile of fossils. The presence of the echinoid Parascutella points to shallow-water marine and/or mobile coastal habitats characterised by sandy substrate and higher water energy (cf. Kroh & Nebelsick 2003, Mikša 2009). Coastal sandy habitats in the intertidal to subtidal zone can also be deduced by the occurrence of the trace fossil Piscichnus waitemata Gregory, 1991 (cf. Šimo et al. 2014, Uchman et al. 2018). Finds of the limpet gastropods Patella suggest intertidal to shallow-water environments and exposed rocky shores (e.g., Poppe & Goto 1991, Forli et al. 2004, 2021, Alf et al. 2020). The co-occurrence of oyster Ostrea digitalina (Eichwald, 1830) demonstrates a possible rocky-coastline paleoenvironment with dominant wave action in the medio/sublittoral, intertidal to shallow subtidal zone (Zágoršek et al. 2009).

In summary, the sedimentology and macrofauna suggest nearshore environments with high hydrodynamic energy and a coarse-grained, sandy to sandy-gravely substrate; sedimentation took place in intertidal/upper infralittoral zones.

Turritellidae members occupied these habitats only rarely and marginally, and reached extremely low diversity and abundance, as evidenced by the very sporadic finds of Helminthia tricincta (Borson, 1821) and H. vermicularis (Brocchi, 1814).

DISCUSSION

CONCLUSIONS

The members of the Turritellidae gastropods originating from the Upper Badenian inner neritic deposits exposed at six paleontological localities situated at Devínska Nová Ves are re-examined. Eight species attributed to five genera are confirmed: Archimediella abundans (Handmann, 1882), Archimediella carpathica Harzhauser et Landau, 2019, Helminthia tricincta (Borson, 1821), Helminthia vermicularis (Brocchi, 1814), Oligodia bicarinata (Eichwald, 1830), Ptychidia vindobonensis (Handmann, 1882), Viennella sp., and “Turritella” neudorfensis Toula, 1900. Two taxa, Archimediella carpathica and Viennella sp., are recorded for the first time from the study area. Oligodia bicarinata and Helminthia vermicularis are the most abundant species in the assemblages, often found in most of the localities. In contrast, Helminthia tricincta occurs exclusively at Sandberg, and Viennella sp., known by a single specimen, was found only at Bačnegovice. The assemblage at Záveterná is monotypic, composed solely of the single species, Ptychidia vindobonensis. Members of the genus Archimediella, such as Archimediella abundans or Archimediella carpathica, were distributed in all localities except Záveterná.

The highest diversity of Turritellidae was detected at Bačnegovice (six species), whereas the highest abundance was recorded at the nearby site of Útočnice (599 specimens). The assemblages of turritellids show strong preferences for shallow-water marine inner neritic habitats with seagrass meadows or mixed coralline red algae and seagrass paleoenvironments. In coastal habitats (intertidal to subtidal zones), they reached significantly lower abundance and diversity.

The Turritellidae assemblage from Brickyard is not re-examined and is not taxonomically or paleoecologically evaluated herein. The material from the locality is unavailable, which does not allow its revision. “Turritella” neudorfensis Toula, 1900, an almost forgotten, enigmatic species known only from the literature, found in the pelitic deposits of Brickyard, is more thoroughly discussed. The overall shell morphology of the species suggests a possible relationship to the genus Oligodia Handmann, 1882. However, as demonstrated by the figured material of Toula, some important conchological features are not preserved or are not so clearly drawn in the illustration, and, unfortunately, do not allow its reliable genus-level placement. Therefore, its formal placement in Oligodia is avoided herein; the species remains provisionally assigned to “Turritella” and is considered taxon inquirendum for now. Only new material could clarify the uncertain taxonomic status of this turritellid. The species occupied deeper, middle neritic marine habitats and was adapted to unfavourable, unstable paleoenvironmental settings characterised by lowered water circulation or episodic hypoxic events near the soft, muddy sea floor.