Taxonomical analysis of the Badenian oysters from the Moravian part of the Carpathian Foredeep (Czech Republic): a new revision

СПИСАНИЕ НА БЪЛГАРСКОТО ГЕОЛОГИЧЕСКО ДРУЖЕСТВО, год. 76, кн. 2 3, 2015, с. x x REVIEW OF THE BULGARIAN GEOLOGICAL SOCIETY, vol. 76, part 2 3, 2015, p. x x Taxonomical analysis of the Badenian oysters from the Moravian part of the Carpathian Foredeep (Czech Republic): a new revision Lorenzo De Bortoli, Šárka Hladilová PdF UP Katedra biologie Pedagogické fakulty, Univerzita Palackého v Olomouci, Purkrabská 2, 77140 Olomouc, Česká Republika; E-mail: lorenzo.de@upol.cz; sarka.hladilova@upol.cz. Таксономичен анализ на баденски бивалвии от моравската част на Карпатското предпланинско понижение (Чешка република): нова ревизия Лоренцо Де Бортоли, Шарка Хладилова Катедра по биология, Педагогически факултет, Палацки университет, Пуркрабска 2, Оломоуц 77140, Чешка република Abstract. The presented study is aimed at the Badenian oysters from the Carpathian Foredeep in Moravia, Czech Republic, deposited in collections of the Regional Museum (Vlastivědné Muzeum) in Olomouc (VMO), and Muzeum Prostějovska in Prostějov (MPP). The analysed samples come from 23 localities in Olomouc and Brno regions, the most important being Hluchov, Laškov, Luleč, Myslejovice, Přemyslovice, and the historical Slatinky. We made the systematic revision of the species reviewing the old nomenclature; first time for all the oyster samples in Moravian collections after several discontinuous works through time. The original ~22 species with old nomenclature have been revised to 4 species: Crassostrea gryphoides (Schlotheim, 1813), Hyotissa hyotis (Linné, 1758), Neopycnodonte navicularis (Brocchi, 1814), and Ostrea digitalina (Dubois, 1831). Key words: Oysters, Badenian, Miocene, Carpathian Foredeep, Czech Republic. Резюме. Настоящото изследване е фокусирано върху баденските бивалвии от Карпатското предпланинско понижение в Моравия, Чешка република, намиращи се в колекциите на Регионалния музей (Vlastivědné Muzeum) в Оломоуц (ВМО) и Muzeum Prostějovska в Простейов (МПП). Анализираните проби са от 23 локалитета в района на Оломоуц и Бърно, най-важните сред които са Хлучов, Лашков, Лулеч, Мислейовице, Пржемисловице и историческия Слатинки. За първи път е извършена таксономична ревизия на всички налични екземрляри от Моравските колекции. Оригиналните ~22 вида са ревизирани до 4: Crassostrea gryphoides (Schlotheim, 1813), Hyotissa hyotis (Linné, 1758), Neopycnodonte navicularis (Brocchi, 1814) и Ostrea digitalina (Dubois, 1831). Ключови думи: бивалвии, Баден, Миоцен, Карпатско предпланинско понижение, Чешка република. Introduction The aim of this paper is to document the research of Middle Miocene (Badenian) oyster samples from the Carpathian Foredeep (CF) in Moravia, Czech Republic. Our main objective is the systematic study and revision of older museum collections of oysters from the Badenian localities of this region. Although detailed geological investigations of the Miocene oysters in Europe, but not only, began during the last decades of the 19th century (Prestwich, 1850; Sacco, 1890 1904; Abel, 1898) and they have since been the subject of numerous studies (Jimenez et al., 1991; Dávid, 2001; Videt, Néreaudeau, 2002; Harzhauser et al., 2003; Zuschin et al., 2004; El-Hedeny, 2005; Mauna et al., 2005; Videt, 2007; Gramigna et al., 2008; Hoşgör, 2008; Lopes, 2011; Studencka, Jasionowski, 2011; D Amico et al., 2012; Górka et al., 2012; Wiedl et al., 2012, 2013; Hladilová, Fordinál, 2013; Hosseinipour, Dastanpour, 2013; Pekař, 2013), no taxonomic and palaeoecological joint study of Czech Republic oysters has ever been undertaken. The studied Badenian oyster samples (more than 700) are stored in Vlastivědné Muzeum in Olomouc 33

(VMO), and Muzeum Prostějovska in Prostějov (MPP). The material comes from 23 localities of the CF in Moravia (Fig. 1 A D). The first group (13 localities, namely Čelechovice, Drysice, Hluchov, Kelčice, Kobeřice, Laškov, Luleč, Myslejovice, Olo mouc-neředín, Přemyslovice, Slatinky, Služín, Terezské údolí) is situated in the wider surroundings of Olomouc (middle part of the CF in Moravia), while the second one (10 localities: Bedřichovice, Blažovice, Borač, Brno Vinohrady, Černá Hora, Kroužek, Lomnička, Opatov, Prace, Újezd) is spread in the wider surroundings of Brno (southern part of the CF in Moravia and isolated relics of the CF sediments in Boskovice Graben). The oysters have been taxonomically determined by using a binocular microscope (Nikon SZP 11-BH) and camera (Canon Power Shot G12). All the older determinations have been revised and for each locality it was counted the number of specimens by species. For this study, the comparative materials from the paleontological collections of the Museum of Natural History in Vienna, the Institute of Paleobiology, and the Museum of the Earth, Polish Academy of Sciences in Warszawa, have been used. The determinations are also based on the literary data, mainly on the publications by Stenzel (1971), Schultz (2001 2005), Mandic, Harzhauser (2003), D Amico et al. (2012), etc. For each locality it was counted the number of specimens by species and the data were evaluated statistically. Geological settings The Miocene Paratethys was a typical epicontinental sea characterized by an extraordinarily rich and diversified mollusc record, including oysters. Among others, very abundant oyster accumulations with large shells of Crassostrea gryphoides (Schlotheim, 1813) are mentioned in the literature (Harzhauser et al., 2003; Schultz, 2001 2005; Hoşgör, 2008; Gramigna et al., 2008). From the regional point of view, the studied Neogene deposits belong to the Carpathian Foredeep (CF), a peripheral foreland basin of the Central Paratethys that developed due to subsurface loading of the Alpine-Carpathian orogenic belt on the passive margin of the Bohemian Massif. The basin continues southwards into the Austrian part of the Alpine Foredeep/Alpine Molasse Zone and northeastwards into the Polish part of the CF (Oszczypko, 1998; Nehyba et al., 2008). In the area of today s Moravia (Czech Republic) the deposition in the CF began in Egerian Early Eggenburgian and continued up to the Late Badenian. The location of the basin and the distribution and character of the deposits could be explained by the tectonic development of the Carpathian orogenic wedge. Both eustacy and tectonics influenced the depositional architecture of the basin during the Early Middle Badenian (Nehyba, Šikula, 2007; Nehyba et al., 2008). The Badenian transgression in the Moravian part of the CF could be correlated with global sea level cycle (14.8 13.6 Ma, Hardenbol et al., 1998), although local cycles were strongly controlled by tectonics (Kováč et al., 2001) and associated with basin subsidence and continuous mountain chain uplift. The CF in Moravia was profoundly differentiated: sand, gravel, biostromal and biohermal limestone or calcareous sandstone with a dominance of red algae and bryozoans (Doláková et al., 2008; Zágoršek, 2010a,b) were deposited in the shallow or elevated places, a hundred meter deep depression was filled with unlaminated calcareous clays Tegels (Brzobohatý, 1997; Kováč, 2000; Kováč et al., 2007). Isolated relics of Early Badenian deposits can be found far to the West of the present extent of the basin (Nehyba, Hladilová, 2004; Zágoršek et al., 2012). As previously mentioned the studied oysters come altogether from 23 localities (Fig. 1 A D; Fig. 2); some of them are known and mentioned in the literature (see in the following text) since 19th century (for example Čelechovice, Slatinky nowadays completely filled up, Borač, etc.), other ones were discovered recently (Myslejovice, Laškov). The following brief descriptions are generally arranged in alphabetical order and divided geographically (Olomouc and Brno regions). Olomouc region (13 localities) Čelechovice. In the area between Kaple near Čelechovice and Slatinice the Devonian limestones are superposed by the Lower Badenian sediments of the border facies: sands, clays, surf gravels with pebbles indicating the local origin and bearing bioerosion traces of the assemblage representing the Entobia ichnofacies (Pek et al., 1998). Numerous shells of oysters and oyster banks, oysters attached even to the Devonian limestones (Spitzner, 1899, 1907; Smyčka, 1906; Remeš, 1933; Kalabis, 1934; Jedlitschka, 1937; Chlupáč, Kalabis, 1966; Barth et al., 1971; Novák, 1975; Kupková, 1982; Zapletal, 2004; Jašková, Lehotský, 2009). Drysice. Lower Badenian calcareous clays (tegels) are infilling the surface depressions of the underlying beds represented by the Lower Carboniferous (Culmian) rocks of the Drahany Upland. Section (observable in the local quarry Bílá skála between Drysice and Želeč): at the base 40 cm greenish marl, 15 cm fine-grained yellow sand, 30 cm marl, 5 cm sand, ~40 cm marl, in the top loess of variable thickness. In marls abundant molluscs, in particular oysters (character of oyster bank, well-preserved both left and right valves, frequently accumulations of intergrown valves of variable sizes), foraminifers, ostracods, corals, to a lesser extent echinoids, teeth of vertebrates. The sands are lacking the fossils (Čouka, 1925; Lang, 1933). Hluchov. The locality is situated eastwards of the Hluchov village, westwards of the grove called Brus (also Trávník), it is known since 1996. Fine-grained 34

Fig. 1. Distribution of oyster localities and species: a, Crassostrea gryphoides; b, Hyotissa hyotis; c, Neopycnodonte navicularis; d, Ostrea digitalina (according to geological map of the Moravian part of the Carpathian Foredeep from http://mapy.geology.cz/geovedni_mapy500/, modified) Фиг. 1. Местоположение на локалитетите и най-характерните видове: a Crassostrea gryphoides, b Hyotissa hyotis, c Neopycnodonte navicularis, d Ostrea digitalina (според Геоложката карта на моравската част на Карпатското предпланинско понижение, с изменения) 35

36

Fig. 2. Geological sections of some of the most significant fossiliferous localities in Moravian part of the Carpathian Foredeep (Olomouc region) (modified from Kupková, 1981 1995; Jašková, 1992; Jašková, 1998; Jašková, Lehotský, 2009) Фиг. 2. Геоложки разрези на някой от най-значимите фосилни находища в моравската част на Карпатското предпланинско понижение (район Оломоуц) (по Kupková, 1981 1995; Jašková, 1992; Jašková, 1998; Jašková, Lehotský, 2009, с допълнения) Lower Badenian sands of varied colours, with abundant foraminifers, bryozoans, molluscs (locally massive oyster valves), cirripeds, echinoids, corals and red-algae, are overlying the Culmian shales. The locality was studied in detail at first by a digged hole up to the depth of 2.4 m (Jašková, 1998), afterwards by two shallow boreholes HL1 and HL2 that reached sandy clays with large clasts of Carboniferous rocks, calcareous sands, sandy limestones, limestones and clays of Lower Badenian age. In addition to bryozoans, the samples contain molluscs, foraminifers, fragments of echinoderms, cirripeds, serpulid tubes, exceptionally colonial corals. ostracods, otoliths, fish teeth, fragments and spicules of sponges and brachiopods (Zágoršek et al., 2010; Nehyba, Jašková, 2012; Hladilová et al., 2014). Kelčice Taliánská skála (Taliánka). In the grove about 1 km SE of the Kelčice village there is an abandoned small quarry in a partly exhausted body of the Lower Badenian red-algal limestones (grey-yellow siliceous limestones with abundant bivalve shells, especially of pectinids). Fragments of these rocks are spread even about the surrounding fields. In the western part of the quarry there occurs a layer of the Lower Badenian basal gravels. In 2000, the extent of the Miocene relic has been verified by 13 orientational holes in the fields around Taliánka; the thickness of the limestone bench does not exceed 6 m. The underlying rocks are represented by Culmian greywackes and shales (the most eastern projections of the Culmian rocks of the Drahany Upland) creating morphologically conspicuous elevations (Jašková, 2002). In the sand pit about 0.5 km north of Taliánka the base of Lower Badenian limestones occurs, namely grey-yellow to grey calcareous sands, in places consolidated and forming benches, consisting practically exclusively of the Culmian rocks grains, and sporadically with fossils (bivalves Jašková, 2002). Kobeřice near Prostějov. The abandoned sand pit is situated on the western periphery of Kobeřice, in the Hůrka Hill (277 m above sea level), near the road between Hradčany and Kobeřice. The Lower Badenian basal clastic sediments shallow-water marine gravels and sands (locally consolidated into conglomerates and sandstones with oysters) are deposited with discordance on the Lower Carboniferous sediments. The smoothed steep to vertical walls of the Culmian rocks represent presumably cliffs formed by surf. The uppermost part of the section is represented by loess (Spitzner, 1899, 1904, 1906, 1907; Barth et al., 1971). Laškov. Within 1991 1993 in the woody area between the hills Rmýz and Křemela (area of Náměšť na Hané), about 1.8 km NE of the village Laškov, a small denudation relic of Miocene sediments preserved in the depression of the underlying rocks represented by the Lower Carboniferous (Culmian) shales has been found and documented by a digged hole. Under the Quaternary sediments (0.4 m) the grey-white to greyyellow sand with fragments of massive oyster valves mixed with rubble resulting from their disintegration is cropping out. The amount of valve fragments is markedly increasing downwards, in 0.9 1.0 m the oyster horizon has been ascertained. In the wider area, about 1 km northeastwards of Laškov, fragments of algal limestones have been ascertained spread about the fields (Novák, 1975; Jašková, 1998; Horčičko et al., 2006). Luleč. Lower Badenian oyster bank, its underlying rocks are represented by the Culmian sediments of the Drahany Upland (Spitzner, 1899, 1907; Čouka, 1925; Lang, 1933). Myslejovice. During the excavations of the family house foundations Lower Badenian grey clays with fragments of oyster valves have been ascertained. Section: 0.5 m variegated clays, 0.6 m grey-green clay with oysters Crassostrea gryphoides, 0.6 m Quaternary clay loam with tiny fragments of Culmian rocks and fragments of oyster valves (Jašková, Lehotský, 2009). Olomouc Neředín. By shallow boreholes there were detected Lower Badenian pelitic facies overlying basal clastic sediments of Early Badenian age: grey-green, yellow-grey, blue-grey to brown-grey calcareous and finely sandy clays (tegels), in places with ochre smudges, often with slight silty or sandy admixture. These pelites contain abundant calcareous concretions and sometimes even macrofauna, first of all oysters and other marine molluscs (Hrubeš, Čtyroká, 1999; Čtyroká, Švábenická, 2000; Zapletal, 2004). Přemyslovice. The Miocene (Lower Badenian) deposits near the village of Přemyslovice were mentioned by Spitzner (1899, 1907), and studied by Schwarz 37

(1946) and Novák (1975). New investigations were initiated after the findings of rich bryozoans-bearing sediments in the vicinity of Přemyslovice (Jašková, 1998). The sedimentary succession rich in foraminifers, molluscs, bryozoans, brachiopods and red algae was examined by 7 shallow boreholes (Zágoršek, Holcová 2009; Zágoršek et al., 2012). Slatinky. The former sand pit, nowadays out of existence and recultivated; Miocene (Badenian) sands and sandy clays 10 20 m in thickness, overlying the Devonian limestones. Section from the base: fine-grained grey-green clayey sands and sandy clays poor in fossils, in the upper part yellowish, in places calcareous coarse-grained siliceous sands abundant in fossils, in particular Foraminifera, Mollusca Bivalvia (including oysters) and Gastropoda, Echinoidea, Cirripedia, Ostracoda. Shallow-water sediments, littoral to neritic facies. The uppermost part of the section is loess (Pleistocene) Wolf (1862), Toula (1893), Spitzner (1899, 1904, 1906, 1907), Schubert (1913), Remeš (1930), Kostka (1931), Kalabis (1934, 1937, 1938, 1949, 1956, 1961a d), Jedlitschka (1937), Roth et al. (1962), Barth et al. (1971), Řehoř et al. (1978), Kupková (1981, 1982, 1984, 1986, 1995), Kupková et al. (1987), Jašková (1998), Zapletal (2004), Hladík (2009), Jašková, Lehotský (2010), Kopecká (2012), Pekař, Lehotský (2013). A relic of a typical oyster bank overlying the Devonian limestones is situated about 50 m SW of the former sand pit towards Kaple (Kupková, 1982). Služín. About 1.5 km W of the village Služín on the east margin of the wood Brus in the Stříbrný potok valley near the lane towards Hluchov there were two relics of Miocene sediments (in one of them formerly a sand pit). Nowadays, the locality is totally destroyed, as a pond has been excavated within the occurrence area of the Miocene sediments (Jašková, 1998). Sediments of the formation with the red-algal limestones with relatively abundant fauna and flora overlying sandy, micaceous, calcareous claystones with shreds of shaly claystones (foraminifers, sponge spicules), higher ~2.5 m grey, fine-grained sandy calcareous claystone (sponge spicules, ostracods, foraminifers), ~0.70 m rusty brown to greenish sedimentary breccia with a layer of unconsolidated grey-white sandy limestone (0.2 m). In the breccia sharply angular fragments of Culmian shales dominate, large foraminifers (heterostegines), sponge spicules, fragments of corals and bryozoans, echinoids, molluscs, shark and fish teeth and fragments of red algae occur in the intersticial mass a typical example of a surf zone. In the uppermost parts of the section white sandy limestone with recrystallized foraminifers and grey, yellowish grey to rusty, slightly sandy algal limestones with fauna and flora foraminifers, sponges, corals, bryozoans, molluscs (bivalves including oysters, gastropods), ostracods, echinoids, red-algae, ichnofossils Spitzner (1899, 1904, 1906, 1907), Blekta (1932), Kalabis (1934, 1937, 1938, 1949, 1960), Novák (1975), Vysloužil (1981), Lehotský (2006), Zborník et al. (2013). Terezské údolí. The natural park (since 1996) (Horčičko et al., 2006). An accidental find of oyster shells in 2003 (Čermák, 2007) in the slope above a sharp loop of the Šumice river about 3 km w. of the railway stop Náměšť na Hané and 2 km NE of the bus stop in Laškov. During the field investigation (2005) an accumulation of oyster shells in a small depression within the Lower Carboniferous shales has been confirmed under the Quaternary sediments (~0.25 m) Jašková, Lehotský (2009). The oyster assemblage is most probably redeposited and manifests relations with the close Laškov locality (Jašková, 1993, 1998). Abundant boring traces of marine organisms (Entobia isp.,?entobia isp.,?trypanites isp., Gastrochaenolites isp.) occur on the oyster shells (Hladík, 2009). Brno region (10 localities) Bedřichovice. In the abandoned quarry west of the village margin there occur Paleozoic limestones (Upper Frasnian) (Müller et al., 2000), transgressively overlapped by Lower Badenian calcareous clays with oysters, nowadays preserved only in denudation relics. The oyster banks or limestone breccias rich in foraminifers (Lenticulina) occur in the basal parts of the Lower Badenian calcareous clays even at some other surrounding localities, for example in the Mokrá quarry or on the Palacký Hill (Müller et al., 2000). Blažovice. Between the villages Blažovice and Holubice (S and SW of Holubice) the Lower Badenian rocks of the formation with algal limestones are cropping out in two morphologically evident denudation relics, and unearthed in two pit quarries at the elevation point 286.6. Sandy, medium- to coarse-grained organodetritic limestones and calcareous sandstones with abundant flora and fauna Foraminifera, Anthozoa, Mollusca, Vermes, Bryozoa, Echinoidea, Rhodophyta. The underlying rocks are represented by tegels (Paulík, 1953; Novák, 1975; Nehyba et al., 2014). Borač. About 800 m NW of the village, to the left of the road Borač Doubravník, there are cropping out small denudation relics of marine, dark green-grey, grey-yellow or grey-blue, slightly sandy calcareous clays (tegels) extraordinarily rich both in macro- and microfossils (Foraminifera, Anthozoa, Mollusca, Ostracoda, Vertebrata, nannoplankton) (Procházka, 1892, 1893; Tausch, 1895; Cicha et al., 1957; Cicha, Dornič, 1960; Hladil, 1976; Seitl, 1978). Tegels are deposited transgressively without basal clastic sediments directly on the metamorphites of Moravicum. This locality was assigned s a regional faciostratotype of the Lower Badenian (Moravian) by Papp et al. (1978). Brno Vinohrady. Old sand pits, partly recultivated, partly built in, north of the tram-line to the Líšeň Quarter of Brno (near Krásného Str.). Lower Badenian basal clastic sediments, so-called Brno sands (finingupwards sequence, in places with benches of sandstones and clay galls), and superposed calcareous 38

clays (tegels), Mollusca (mostly redeposited from the Rzehakia beds), including oysters (Krystek, 1974; Krystek, Tejkal, 1968). Černá Hora. Near the road NW of Černá Hora two denudation relics of the Lower Badenian calcareous clays tegel (white-grey, yellow-green or grey-green), with foraminifers, echinoids, bivalves (abundant oysters), gastropods, scaphopods, corals, sponge spicules; in the upper part loess loams or coarse-grained gravels (Dvořák, 1952; Hladil, 1976). Kroužek. Mostly overgrown outcrops on the slopes of the valley SE of Kroužek and on the elevation point 348.9, sediments of Early Badenian age greenish calcareous clays (tegels), underlying the formation with organodetritic algal limestones coarse-grained, slightly sandy limestones with abundant Foraminifera, Vermes, Bryozoa, Mollusca, Echinoidea, Ostracoda, Teleostei and Rhodophyta (Šob, 1940; Kalášek et al., 1963; Zapletal, 1934; Paulík, 1953; Novák, 1975; Hladilová, 1984; Holcová et al., 2015); http://lokality. geology.cz3097. Lomnička. Denudation relics of the Lower Badenian sediments in the Boskovice Furrow among Lomnička, Železné and Tišnov, transgressively deposited on the metamorphites of the Vranov-Olešnice series and geologically belonging to the sea-lochs in the western part of the Carpathian Foredeep in Moravia. Green-grey calcareous clays (tegels) with abundant oysters, furthermore gastropods and foraminifers, and red-algal limestones with foraminifers, ostracods, bryozoans and rare molluscs (Procházka, 1892; Tausch, 1895; Zapletal, 1933; Cicha, Dornič, 1959, 1960; Krystek, Tejkal, 1968; Novák, 1975; Hladil, 1976). Opatov. Artificial exposure to the right of the railway station Opatov Česká Třebová between 236.359 to 236.700 km, 2 6 m in width. Grey to blue-grey, in places rusty mottled, disaggregating calcareous clays of Early Badenian age, 2 3 m in thickness, in places slightly sandy, rich in macrofauna, particularly mollusks, including oysters, and ichnofossils, are overlying the Upper Cretaceous sediments (Procházka, 1895, 1900; Cicha, Dornič, 1959; Pek, Mikuláš, 1996; Hamršmíd et al., 1998). Prace. In the shelf quarry on the south slope of the Prace Hill between the villages Prace and Újezd red-algal limestones (Lower Badenian) are unearthed overlying mostly calcareous clays (Lower Badenian), only at the elevation point 307.6 schlier of Karpatian age. In the clays the amount of the sand fraction is increasing upwards and gradually benches of calcareous sandstones with red algae occur in them. In the uppermost parts, there occur mostly benches of organodetritic algal limestones of varying thickness, often tapering off and alternating with calcareous sands to sandstones. Nowadays, the locality does not exist (Novák, 1975; Zdražílková, 1985, 1987, 1988; http:// lokality.geology.cz3697). Újezd u Brna Stará Hora. Locality in the southern part of the Prace Hill massif, grey to brownish, bedded, micaceous, silty clays schliers (Karpatian), sandy limestones and calcareous sandstones with sandy layers (Lower Badenian) with rich micro- and macrofauna; the upper parts of the section are represented by loess (Novák, 1975; http://lokality.geology.cz3699). Obtained results 1. Oysters the original determinations and revisions Among the studied oysters there are many specimens with ancient nomenclature and other yet to be assessed; we revised these old determinations (Table 1). Ostrea cyathula. In particular Ostrea cyathula or Crassostrea cyathula (Lamarck, 1806) is described as the Oligocene typical specimen (Hosgör, Okan, 2009) in Turkey. Gümbel (1888) described it; also it has been determined as Gryphaea cyathula cf. cyathula (by Michalík, Zágoršek, 1986); the specimens in the collections are of a small size (~8 5 cm) and the same morphological features (outer surface, structure, texture and margin) attributable to Ostrea digitalina (Dubois, 1831). Ostrea cymbularis: Ostrea cymbularis Muenster in G. A. Goldfuss (1833) is mentioned in the collections of the Museum of Natural History in Paris and in the historical report of Hoernes (1845) and von Letocha (1863); it must be considered as clustered in the group of Ostrea digitalina (Dubois, 1831) according to the same remarks made for Ostrea cyathula, because the morphological features are very similar. Ostrea cochlear: Ostrea cochlear Poli, 1975 is in synonymy with Neopycnodonte navicularis (Poli, 1975), from Miocene of Spain (Videt, Néraudeau, 2002). Ostrea fimbriata: It is a determination not too frequent in our collections and we consider these specimens still as Ostrea digitalina in comparison with the morphologically very close Ostrea cyathula and Ostrea cymbularis ones. Ostrea gingensis: Ostrea gingensis Schlotheim, 1813 was formerly described by Rolle (1859) and recently recognized in synonymy with Crassostrea gryphoides (Schlotheim, 1813) by Vazzana, Cecalupo (2007) from Cessaniti, Italy. Ostrea hoernesi: Ostrea hoernesi is a taxon with ancient nomenclature, described by Kittl (1887) in the Austrian historical collections. It is of a medium-large size (15 cm), the general outline is quite rounded and the shape is concavo-flat (the muscle scare on the left valve is in central position with rounded shape); as for these features we refer the studied specimens at this moment of the study to Hyotissa hyotis. They are coming only from the Bedřichovice locality. Ostrea lamellosa: After morphological and morphometric analyses we consider these specimens as morphometric variation of Ostrea digitalina (Dubois, 1831). Ostrea plicata: Wozny (1977) refers Ostrea plicata (Solander, 1766) to Eocene of Poland, whereby dubious presence in the collections analyzed; Fuchs (1873) 39

Table 1 Revised list of the oyster species present in the VMO and MPP Museum collections Таблица 1 Списък на ревизираните видове от музейните колекции на ВМО и МПП Ancient determination Revision Crassostrea gryphoides (Schlotheim, 1813) Crassostrea gryphoides (Schlotheim, 1813) Neopycnodonte navicularis (Brocchi, 1814) Neopycnodonte navicularis (Brocchi, 1814) Hyotissa squarrosa (De Serres, 1843) Hyotissa hyotis (Linné, 1758) Ostrea cf. fimbriata Grat. in Raulin, Delbos (1855) Ostrea digitalina (Dubois, 1831) Ostrea crassicostata (Sowerby in Smith, 1847) Hyotissa hyotis (Linné, 1758) Ostrea cyathula (Lamarck, 1806) Ostrea digitalina (Dubois, 1831) Ostrea cymbularis Muenster in G. A. Goldfuss (1833) Ostrea digitalina (Dubois, 1831) Ostrea digitalina (Dubois, 1831) Ostrea digitalina (Dubois, 1831) Ostrea eduliformis (Schlotheim, 1820) Ostrea digitalina (Dubois, 1831 Ostrea edulis (Linné, 1758) Ostrea digitalina (Dubois, 1831) Ostrea fimbriata Grat. in Raulin, Delbos (1855) Ostrea digitalina (Dubois, 1831) Ostrea gigantea Sowerby, 1814 Hyotissa hyotis (Linné, 1758) Ostrea gingensis Schlotheim, 1813 Crassostrea gryphoides (Schlotheim, 1813) Ostrea hoernesi Reuss, 1870 Hyotissa hyotis (Linné, 1758) Ostrea lamellosa Brocchi, 1814 Ostrea digitalina (Dubois, 1831) Ostrea leopolitana Niedèwiedzki, 1909 Neopycnodonte navicularis (Brocchi, 1814) Ostrea plicata plicata (Solander in Brander, 1766) Hyotissa hyotis (Linné, 1758) Pycnodonte vesicularis Lamarck, 1806 Neopycnodonte navicularis (Brocchi, 1814) reports Ostrea plicata Gmelin as Hyotissa squarrosa (De Serres, 1843), now determined as Hyotissa hyotis (Linné, 1758). In our specimens the state of preservation is not good, so according to the previous literature we accepted it as Hyotissa hyotis. Ostrea postvesicularis: Taylor (1885) reports Ostrea vesicularis in the old collections from England, but it is not a Miocene species; according to the small size (5 cm) and the main features of the valves (elongated outline, smooth external surface, thickness and depth on the left valve) the specimen present in our collections could be referred to Neopycnodonte navicularis (Brocchi, 1814). Ostrea cf. fimbriata: Ostrea cf. fimbriata is attributed to Ostrea digitalina (Dubois, 1831) by Pekař (2013). Ostrea crassicostata: Ostrea crassicostata is described by Antunes et al. (1999) as var. gigantea (not Miocene species), from sediments of Portugal. In our collections these specimens are characterised by a rounded shape, large dimensions, crenulation on the external surface, muscle scar (left valve) in central position of the internal surface; based on these facts we refer them as morphological variations of Hyotissa hyotis (Linné, 1758). Ostrea eduliformis: Ostrea eduliformis is a taxon referred in literature to Bajocian, by Pockrandt (1984), and should not to be considered in Badenian samples; as there are not other indications from the collector, we should think it has been confused with Ostrea edulis (Linné, 1758). Our specimens are medium-large in size ( 5 10 cm), present a crenulated external surface, a resilifer zone rather boxy in shape and a moderate thickness, therefore we refer them to Ostrea digitalina (Dubois, 1831). Ostrea leopolitana: Ostrea leopolitana Niedèwiedzki, 1909 has been described by Friedberg (1936) and Radwánski (1973) as Pycnodonta leopolitana from Poland, now in synonymy with Neopycnodonte navicularis (Brocchi, 1814); our specimens fit perfectly with Neopycnodonte navicularis as for size than for morphological patterns. 2. Revised oyster species in the collections Here we present the systematic descriptions of the 4 updated species Crassostrea gryphoides, Hyotissa hyotis, Neopycnodonte navicularis and Ostrea digitalina occurring in the collections. The determinations are also based on the literary data, mainly on the publications by Stenzel (1971), Schultz (2001 2005), Mandic, Harzhauser (2003), D Amico et al. (2012), etc. Class Bi v a l v i a Linné, 1758 Subclass Pt e r i o m o r p h i a Beurlen, 1944 Order Pt e r i o i d a Newell, 1965 Suborder Os t r e i n a Ferussac, 1822 Superfamily Os t r e o i d e a Rafinesque, 1815 40

Family Cr a s s o s t r e i d a e Scarlato, Starobogatov, 1979 Genus Crassostrea Sacco, 1897 Ostrea (C.) virginica (Gmelin, 1791). Type species: Ostrea virginica Gmelin, 1791 Crassostrea gryphoides (Schlotheim, 1813) (Plate I, a h; Plate III, c, f) Description. Shells inequivalve, elongated, sizes are much greater than other species. A well-developed umbonal cavity in the right valve; the left valve is more convex or quite flat. The adductor muscle imprint is in postero-ventral position, outline crescentic with fairly sharp corners in the dorsal margin. The shells are broader next to the ventral margin and thicker in the umbo zone. On the external surface there are folded ribs and growth lamellae. Chomata are absent along the entire margin of the both valves. Material. Bedřichovice: 6883 (VMO); Drysice: 30102 (VMO); Laškov: 087175/1-72(MPP); (MPP); 30446, 30447, 30448 (VMO); Luleč: 190774/2, 190774/3, 190774/8, 190774/9, 190774/10 (MPP); Myslejovice: 190442, 190443, 190444, 190445, 190447, 190448 (MPP); Slatinky: 7746, 7801, 7889 7891, 7853, 7923, 7942 45, 7955, 7956, 7957 7959, 7960 7962, 7971, 7979, 7992, 8000 8003, 8009 8013, 8014 8017, 23188 23189 23190, 28227, 28233, 28712, 28713, 30101, 30168, 30186, 30270, 30327, 30420 30421 (VMO); Vinohrady (Brno): 051886 (MPP). Family GRYPHAEIDAE Vyalov, 1936 Subfamily PYCNODONTEINAE Stenzel, 1959 Genus Hyotissa Stenzel, 1971 Hyotissa hyotis (Linné, 1758) (Plate II, g l; Plate III, d, g) Description. The valves are rather circular, with the adductor muscle in the center; a thick shell not really concave with the ribs in the external surface. The margin is clearly crenulated. Notes. Hoernes (1870) referred the Austrian specimens to Ostrea plicatula Gmelin and O. crassicostata Sowerby, but it was attributed to Pycnodonte squarrosa (De Serres, 1843) by Studencka et al. (1998). There were demonstrated the differences in the adductor muscle insertion in squarrosa and hyotis, considered as intraspecific variability (Freneix et al., 1988). Ostrea crassicostata is reported in the sediments of Portugal (Antunes et al., 1999) as var. gigantea [Pycnodonta squarrosa (De Serres, 1843), now Hyotissa hyotis]. Material. Bedřichovice: 1579, 1580, 5873, 6400, 6519, 6946 (VMO); Blažovice: 6899 (VMO); Brno: 2417 (VMO); Hluchov: 139623/3(VMO); Slatinky: 28225, 28226, 28234 (VMO); Terezské údolí: 190797/7, 190797/10 (MPP). Genus Neopycnodonte Stenzel, 1971 Neopycnodonte navicularis (Brocchi, 1814) (Plate I, i r; Plate III, a, e) Description. The shell is thin (vesicular structure) and of small size, normally 5 to 10 cm long; the shape is ovoidal-elongated. The posterior and anterior margins have an edge. The right valve is plano-concave; its ligament area presents a median depression. The external surface lacks the sculpture. The adductor insertion is wide and it is close to the hinge and to the ventral edge. The chomata are present in the zone of the ribs extension. Notes. The name navicularis has for a long time been used at the level of either variety (Hoernes, 1870; Friedberg, 1934; Kojumdgieva, 1960), or subspecies (Caprotti, 1972) of N. cochlear (Studencka et al., 1998). It was also demonstrated that the resilifer was too big in order to consider N. cochlear (Poli, 1795) and its variety navicularis as two different species (Freneix, 1975). Videt, Néraudeau (2002) compare Neopycnodonte cochlear Poli, 1795 (sensu Freneix et al., 1988) to Neopycnodonte navicularis (Poli, 1795), from Miocene of Spain according to the paleoecological point of view. Material. Borač: 4739, 5845 (VMO); Brno: 6540, 6689 (VMO); Čelechovice: 073275, 073762, 078729 (MPP); Lomnička: 4595, 4596 (VMO); Prace: 1527 (MPP); Slatinky: 7736, 7737, 7760, 7761, 7778, 7779, 7812 7814, 7815 7818, 7901 7904, 7918 7922, 7963 7967, 26731, 26732, 30211 30215 (VMO); Vinohrady (Brno): 073291 (MPP). Family Os t r e i d a e Rafinesque, 1815 Subfamily Os t r e i n a e Rafinesque, 1815 Genus Ostrea Linné, 1758 Ostrea digitalina (Dubois, 1831) (Plate II, a f; Plate III, b, h) Description. The left valve is more convex than the right one (elevated marginal area and coarse radial ribs) and it is used in cementing. The outline can be oval or subquadrate. The size is medium and it reaches 15 cm. Notes. According to Studencka (1986) it is possibly synonymous with the Atlantic and Mediterranean Ostrea frondosa De Serres; it is considered a fully marine oyster, settled on different kinds of hard substrata. We can compare it with the Ostrea edulis (Schultz, 2001 2005), typical from Recent Mediterranean taxa, living in colonies in shallow waters about 10 meters (Milisic, 1991). It can be also morphologically compared to Ostrea fimbriata, reported as cf. Crassostrea fimbriata in the sediments of Turkey (Hakyemez, 1989). Material. Borač: 4738, 6541 (VMO); Brno: 6516, 6517, 6691 (VMO); Černá Hora: 6688 (VMO); Kelčice: 078730 (MPP); Kobeřice: 8046, 8047 (VMO); Lomnička: 5874 (VMO); Luleč: 4737, 4743 (VMO); Olomouc: 26873, 26874 (VMO); Opatov: 6412 (VMO); Slatinky: 1680, 1681, 6722, 6723, 7017, 7743 7745, 7799, 7800, 7879, 7905 7907, 8011, 23187, 27759, 27774 27778, 27780, 41

PLATE I ТАБЛИЦА I 42

Plate І a. Crassostrea gryphoides (Schlotheim, 1813) (i.n.7992), Slatinky, right valve, internal side (VMO). b. Crassostrea gryphoides (Schlotheim, 1813) (i.n.7992), Slatinky, right valve, external side (VMO). c. Crassostrea gryphoides (Schlotheim, 1813) (i.n.23190), Slatinky, left valve internal side (VMO). d. Crassostrea gryphoides (Schlotheim, 1813) (i.n.23190), Slatinky, left valve, external side (VMO). e. Crassostrea gryphoides (Schlotheim, 1813) (i.n.087175/6), Laškov, right valve, internal side (VMO). f. Crassostrea gryphoides (Schlotheim, 1813) (i.n.087175/6), Laškov, right valve, external side (VMO). g. Crassostrea gryphoides (Schlotheim, 1813) (i.n.190447), Myslejovice, right valve, internal side (MPP). h. Crassostrea gryphoides (Schlotheim, 1813) (i.n.190447), Myslejovice, right valve, external side (MPP). i. Neopycnodonte navicularis (Brocchi, 1814) (i.n.30211), Slatinky, right valve, internal side (VMO). l. Neopycnodonte navicularis (Brocchi, 1814) (i.n.30211), Slatinky, right valve, external side (VMO). m. Neopycnodonte navicularis (Brocchi, 1814) (i.n.30214), Slatinky, right valve, internal side (VMO). n. Neopycnodonte navicularis (Brocchi, 1814) (i.n.30214), Slatinky, right valve, external side (VMO). o. Neopycnodonte navicularis (Brocchi, 1814) (i.n.6689), Brno right valve, internal side (VMO). p. Neopycnodonte navicularis (Brocchi, 1814) (i.n.6689), Brno, right valve, external side, (VMO). q. Neopycnodonte navicularis (Brocchi, 1814) (i.n.073291), Vinohrady, right valve, internal side (MPP). r. Neopycnodonte navicularis (Brocchi, 1814) (i.n.073291), Vinohrady, right valve, external side (MPP). scale bar = 5 cm Таблица I a. Crassostrea gryphoides (Schlotheim, 1813) (инв. 7992), Слатинки, дясна валва, вътрешна страна (ВМО). b. Crassostrea gryphoides (Schlotheim, 1813) (инв. 7992), Слатинки, дясна валва, външна страна (ВМО). c. Crassostrea gryphoides (Schlotheim, 1813) (инв. 23190), Слатинки, лява валва, вътрешна страна (ВМО). d. Crassostrea gryphoides (Schlotheim, 1813) (инв. 23190), Слатинки, лява валва, външна страна (ВМО). e. Crassostrea gryphoides (Schlotheim, 1813) (инв. 087175/6), Лашков, дясна валва, вътрешна страна (ВМО). f. Crassostrea gryphoides (Schlotheim, 1813) (инв. 087175/6), Лашков, дясна валва, външна страна (ВМО). g. Crassostrea gryphoides (Schlotheim, 1813) (инв. 190447), Мислейовице, дясна валва, вътрешна страна (МПП). h. Crassostrea gryphoides (Schlotheim, 1813) (инв. 190447), Мислейовице, дясна валва, външна страна (МПП). i. Neopycnodonte navicularis (Brocchi, 1814) (инв. 30211), Слатинки, дясна валва, вътрешна страна (ВМО). l. Neopycnodonte navicularis (Brocchi, 1814) (инв. 30211), Слатинки, дясна валва, външна страна (ВМО). m. Neopycnodonte navicularis (Brocchi, 1814) (инв. 30214), Слатинки, дясна валва, вътрешна страна (ВМО). n. Neopycnodonte navicularis (Brocchi, 1814) (инв. 30214), Слатинки, дясна валва, външна страна (ВМО). o. Neopycnodonte navicularis (Brocchi, 1814) (инв. 6689), Бърно, дясна валва, вътрешна страна (ВМО). p. Neopycnodonte navicularis (Brocchi, 1814) (инв. 6689), Бърно, дясна валва, външна страна, (ВМО). q. Neopycnodonte navicularis (Brocchi, 1814) (инв. 073291), Винохради, дясна валва, вътрешна страна (МПП). r. Neopycnodonte navicularis (Brocchi, 1814) (инв. 073291), Винохради, дясна валва, външна страна (МПП). Мащабна линия = 5 cm 28186, 30182 30185, 30427 30432 (VMO); Újezd: 6126 (VMO). Summary. After the revision work we reduced the number to 4 species (~22 original), namely Crassostrea gryphoides, Hyotissa hyotis, Neopycnodonte navicularis, and Ostrea digitalina more Ostrea sp.. Crassostrea gryphoides and Ostrea digitalina are most abundant (Olomouc); the most numerous collections come from Slatinky (78%) and Laškov (6%). Ostrea digitalina represents the most abundant taxon in Brno region [for this region: Bedřichovice (20%), Brno (17%), Luleč (13%) and Borač (13%)]. Grouping all the samples from the Moravian part of the CF (excluding Ostrea sp.) we obtain: Crassostrea gryphoides (38%), Neopycnodonte navicularis (14%), Ostrea digitalina (36%) and Hyotissa hyotis (12%). Discussion In addition to the systematic review, where the old nomenclature has been updated in 4 species (Crassostrea gryphoides, Neopycnodonte navicularis, Ostrea digitalina and Hyotissa hyotis) (Fig. 1 A D), we also tried to recognize the oyster percentage distribution according to the provenance locality and to the morpho-species kind. In spite of the relatively high numbers of the studied Lower Badenian oysters, it was generally impossible to use them for more detailed paleoecological interpretations. The main reason was the heterogeneity of the material, namely the limited knowledge of the individual collections histories and of the circumstances connected with their formation: for example we could not verify whether the species compositions or numbers of individuals are not significantly 43

PLATE II ТАБЛИЦА II 44

Plate ІІ a. Ostrea digitalina (Dubois, 1831) (i.n.30431), Slatinky, right valve, external side (VMO). b. Ostrea digitalina (Dubois, 1831) (i.n.30431), Slatinky, right valve, internal side (VMO). c. Ostrea digitalina (Dubois, 1831) (i.n.6412), Opatov, left valve, external side (VMO). d. Ostrea digitalina (Dubois, 1831) (i.n.6412), Opatov, left valve, internal side (VMO). e. Ostrea digitalina (Dubois, 1831) (i.n.26873), Neředin (Olomouc), left valve, external side (VMO). f. Ostrea digitalina (Dubois, 1831) (i.n.26873), Neředin (Olomouc), left valve, internal side (VMO). g. Hyotissa hyotis (Linné, 1758) (i.n.28255), Slatinky, right valve, internal side (VMO). h. Hyotissa hyotis (Linné, 1758) (i.n.28255), Slatinky, right valve, external side (VMO). i. Hyotissa hyotis (Linné, 1758) (i.n.139623), Hluchov, left valve, internal side (VMO). l. Hyotissa hyotis (Linné, 1758) (i.n.139623), Hluchov, left valve, external side (VMO). scale bar = 4 cm Таблица ІІ a. Ostrea digitalina (Dubois, 1831) (инв. 30431), Слатинки, дясна валва, външна страна (ВМО). b. Ostrea digitalina (Dubois, 1831) (инв. 30431), Слатинки, дясна валва, вътрешна страна (ВМО). c. Ostrea digitalina (Dubois, 1831) (инв. 6412), Опатов, лява валва, външна страна (ВМО). d. Ostrea digitalina (Dubois, 1831) (инв. 6412), Опатов, лява валва, вътрешна страна (ВМО). e. Ostrea digitalina (Dubois, 1831) (инв. 26873), Нержедин (Оломоуц), лява валва, външна страна (ВМО). f. Ostrea digitalina (Dubois, 1831) (инв. 26873), Нержедин (Оломоуц), лява валва, вътрешна страна (ВМО). g. Hyotissa hyotis (Linné, 1758) (инв. 28255), Слатинки, дясна валва, вътрешна страна (ВМО). h. Hyotissa hyotis (Linné, 1758) (инв. 28255), Слатинки, дясна валва, външна страна (ВМО). i. Hyotissa hyotis (Linné, 1758) (инв. 139623), Хлучов, лява валва, вътрешна страна (ВМО). l. Hyotissa hyotis (Linné, 1758) (инв. 139623), Хлучов, лява валва, външна страна (ВМО). Мащабна линия = 4 cm changed by the collectors themselves in comparison with the real situation at the localities. Even the data on exact original positions/orientations of individuals in the sediments are entirely missing. Crassostrea gryphoides occurs mostly in Bedřichovice, Laškov, Drysice, Luleč, Myslejovice, Slatinky and Vinohrady, namely in the area close to the present western margin of the Carpathian Foredeep (Fig. 1A); this could reflect paleogeographic conditions related to the depth and morphology of the Foredeep. According to Jimenez et al. (1991) for Almanzora Corridor (Spain), in the past, Crassostrea banks developed basinward of coral reefs, very different from actual style of life, in shallow-intertidal to shallow sea water with lowered salinity. Hyotissa hyotis is reported in two distinct zones: at Bedřichovice and Blažovice in the Brno area, and at Hluchov, Slatinky and Terezské údolí in the Olomouc area (Fig. 1B); it prefers shallow subtidal habitats down to 30 m with higher-energy seawater conditions (Bieler et al., 2004; Mikkelsen, Bieler, 2008; Pekař, 2013; Pekař, Lehotský, 2013; Hladilová et al., 2014). Neopycnodonte navicularis is prevalently present in Brno, Čelechovice, Prace, Slatinky, Vinohrady and to W in Borač and Lomnička (Fig.1 C); from paleogeographic point of view, the last two localities belong to denudation relics of the Carpathian Foredeep sediments situated in the Boskovice Furrow, as presented for instance by Doláková et al. (2008). In spite of Crassostrea, this taxon prefers greater depths below 40 m (Bagdasaryan et al., 1966; Zágoršek et al., 2009). Typically the oyster as Neopycnodonte forms dense overgrowths constituting oyster buildups, which are usually several meters thick and generally consist of articulated, massive shells, as seen, for example, in the Upper Badenian deposits of Roztocze in Poland (Pisera, 1985; Studencka et al., 2005), but the individual oyster buildups in the Moravian part of the Carpathian Foredeep have distinctly smaller areal extents and thicknesses in comparison with Poland, being related to the very different conditions of basin configuration. Ostrea digitalina mainly occurs in eleven localities (Fig. 1D), situated not only within the present-day extent of the CF in the Brno and Olomouc areas, but also in the isolated relics of CF sediments (Borač, Černá Hora, Lomnička and Opatov) described in literature from palaeontological point of view (Hladil, 1976; Seitl, 1978; Hudec, 1986; Pek, Mikuláš, 1996; Seko, 2011; Mihulková, 2013, etc.). Ostrea digitalina is typical for a rocky coastline environment with dominant wave action (highly exposed rocky medio/sublittoral, intertidal to shallow subtidal) (Zágoršek et al., 2009). Similar energy-rich rocky 45

PLATE III ТАБЛИЦА III 46

Plate ІІІ a. Neopycnodonte navicularis (Brocchi, 1814) (i.n.7737), Slatinky, right valve, internal side (VMO). b. Ostrea digitalina (Dubois, 1831) (i.n.4738), Borač, right valve, internal side (VMO). c. Crassostrea gryphoides (Schlotheim, 1813) (i.n.23190), Slatinky, right valve, external side (VMO). d. Hyotissa hyotis (Linné, 1758) (i.n.139623/4), Hluchov, right valve, internal side (VMO). e. Neopycnodonte navicularis (Brocchi, 1814) (i.n.30213), Slatinky, left valve, external side (VMO). f. Crassostrea gryphoides (Schlotheim, 1813) (i.n.28227), Slatinky, umbo zone and resilifer, left valve, internal side (VMO). g. Hyotissa hyotis (Linné, 1758) (i.n.6519), Bedřichovice, left valve, internal side (VMO). h. Ostrea digitalina (Dubois, 1831) (i.n.6723), Slatinky, left valve, internal side (VMO). scale bar = 5 cm Таблица ІІІ a. Neopycnodonte navicularis (Brocchi, 1814) (инв. 7737), Слатинки, дясна валва, вътрешна страна (ВМО). b. Ostrea digitalina (Dubois, 1831) (инв. 4738), Борач, дясна валва, вътрешна страна (ВМО). c. Crassostrea gryphoides (Schlotheim, 1813) (инв. 23190), Слатинки, дясна валва, външна страна (ВМО). d. Hyotissa hyotis (Linné, 1758) (инв. 139623/4), Хлучов, дясна валва, вътрешна страна (ВМО). e. Neopycnodonte navicularis (Brocchi, 1814) (инв. 30213), Слатинки, лява валва, външна страна (ВМО). f. Crassostrea gryphoides (Schlotheim, 1813) (инв. 28227), Слатинки, зона на върха и лигаментната ямичка, лява валва, вътрешна страна (ВМО). g. Hyotissa hyotis (Linné, 1758) (инв. 6519), Бедржиховице, лява валва, вътрешна страна (ВМО). h. Ostrea digitalina (Dubois, 1831) (инв. 6723), Слатинки, лява валва, вътрешна страна (ВМО). Мащабна линия = 5 cm sublittoral environments down to 10 m water depths are interpreted even in Austrian localities (Mandic, Harzhauser, 2003). The higher proportion of right valves (60%) in the collections may result from differential transport, as these valves are thinner and smaller than the left ones (40%) attached to the substrate, and are easily remobilized even by weak currents after death and decomposition of the soft parts (Lopes, 2011). Conclusions We have studied the Badenian oyster collections from 23 localities of the Carpathian Foredeep in Moravia deposited in Vlastivědné Muzeum in Olomouc and Muzeum Prostějovska in Prostějov (Czech Republic). The original ~22 species with old nomenclature have been revised to 4 species: Crassostrea gryphoides, Hyotissa hyotis, Neopycnodonte navicularis and Ostrea digitalina. In the analyzed material Crassostrea gryphoides is represented by ~38%, Neopycnodonte navicularis by ~14%, Ostrea digitalina by ~36% and Hyotissa hyotis by ~12%. Crassostrea specimens coming from the localities, which are situated today at the western margin of the CF (from Vinohrady to Laškov); Hyotissa specimens are clearly divided in two different groups of localities (both in Brno- Olomouc regions); the distribution of Neopycnodonte is similar to Hyotissa s one, with the presence in the isolated relics of Borač and Lomnička; O. digitalina specimens are decidedly more widespread in all the area (also in the N to Opatov), without a particular distribution range. Although taking into account the collection bias and their randomness and arbitrariness, these results represent a certain contribution to paleogeographic interpretations of the Carpathian Foredeep in Moravia during the Badenian. Acknowledgements: This work was supported by the ESF Research Project POST-UP II, reg. No. CZ.1.07/2.3.00/30.0041 of the Palacky University Olomouc (Czech Republic). The authors are grateful to Dr. Andrzej Pisera (Institute of Paleobiology, PAN, Warsaw) for his important help in the study of the Ostreoidea, and to Dr. Oleg Mandic (Naturhistorisches Museum in Vienna) who made valuable suggestions in the mollusc systematics and palaeontology. Acknowledgement is also referred to Dr. Tomáš Lehotský (Faculty of Science, Palacky University and Vlastivědné Muzeum of Olomouc) and Dr. Vladimíra Jašková (Muzeum Prostějovska v Prostějově) for their support during the preparation of this paper. The UPLIFT of Palacky University in Olomouc is thanked for helpful observations and remarks, facilitating the final English revision. 47

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