СПИСАНИЕ НА БЪЛГАРСКОТО ГЕОЛОГИЧЕСКО ДРУЖЕСТВО, год. 76, кн. 2–3, 2015, с. 89–108 REVIEW OF THE BULGARIAN GEOLOGICAL SOCIETY, vol. 76, part 2–3, 2015, p. 89–108

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

Lorenzo De Bortoli, Šárka Hladilová

PdF UP – Katedra biologie Pedagogické fakulty, Univerzita Palackého v Olomouci, Purkrabská 2, 77140 , Česká Republika; E-mail: [email protected]; [email protected].

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, , 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), navicularis (Brocchi, 1814), and Ostrea digitalina (Dubois, 1831).

Key words: Oysters, Badenian, , 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 так- сономична 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 numerous studies (Jimenez et al., 1991; Dávid, 2001; Videt, Néreaudeau, 2002; Harzhauser et al., 2003; The aim of this paper is to document the research of Zuschin et al., 2004; El-Hedeny, 2005; Mauna et al., Middle Miocene (Badenian) oyster samples from the 2005; Videt, 2007; Gramigna et al., 2008; Hoşgör, Carpathian Foredeep (CF) in Moravia, Czech Republic. 2008; Lopes, 2011; Studencka, Jasionowski, 2011; Our main objective is the systematic study and revi- D’Amico et al., 2012; Górka et al., 2012; Wiedl et al., sion of older museum collections of oysters from the 2012, 2013; Hladilová, Fordinál, 2013; Hosseinipour, Badenian localities of this region. Although detailed Dastanpour, 2013; Pekař, 2013), no taxonomic and pa- geological investigations of the Miocene oysters in laeoecological joint study of Czech Republic oysters Europe, but not only, began during the last decades of has ever been undertaken. the 19th century (Prestwich, 1850; Sacco, 1890–1904; The studied Badenian oyster samples (more than Abel, 1898) and they have since been the subject of 700) are stored in Vlastivědné Muzeum in Olomouc

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

90 Fig. 1. Distribution of oyster localities and species: a, Crassostrea gryphoides; b, Hyotissa hyotis; c, Neopycnodonte navicu- laris; d, Ostrea digitalina (according to geological map of the Moravian part of the Carpathian Foredeep from http://mapy.geol- ogy.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 пред- планинско понижение, с изменения)

91 92 →

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 пред- планинско понижение (район Оломоуц) (по Kupková, 1981–1995; Jašková, 1992; Jašková, 1998; Jašková, Lehotský, 2009, с допълнения)

Lower Badenian sands of varied colours, with abun- The smoothed steep to vertical walls of the Culmian dant foraminifers, bryozoans, molluscs (locally mas- rocks represent presumably cliffs formed by surf. sive oyster valves), cirripeds, echinoids, corals and The uppermost part of the section is represented by red-algae, are overlying the Culmian shales. The lo- loess (Spitzner, 1899, 1904, 1906, 1907; Barth et al., cality was studied in detail at first by a digged hole 1971). up to the depth of 2.4 m (Jašková, 1998), afterwards Laškov. Within 1991–1993 in the woody area be- by two shallow boreholes HL1 and HL2 that reached tween the hills Rmýz and Křemela (area of Náměšť na sandy clays with large clasts of Carboniferous rocks, Hané), about 1.8 km NE of the village Laškov, a small calcareous sands, sandy limestones, limestones and denudation relic of Miocene sediments preserved in clays of Lower Badenian age. In addition to bryo- the depression of the underlying rocks represented by zoans, the samples contain molluscs, foraminifers, the Lower Carboniferous (Culmian) shales has been fragments of echinoderms, cirripeds, serpulid tubes, found and documented by a digged hole. Under the exceptionally colonial corals. ostracods, otoliths, fish sediments (0.4 m) the grey-white to grey- teeth, fragments and spicules of sponges and brachio- yellow sand with fragments of massive oyster valves pods (Zágoršek et al., 2010; Nehyba, Jašková, 2012; mixed with rubble resulting from their disintegra- Hladilová et al., 2014). tion is cropping out. The amount of valve fragments Kelčice–Taliánská skála (Taliánka). In the grove is markedly increasing downwards, in 0.9–1.0 m the about 1 km SE of the Kelčice village there is an aban- oyster horizon has been ascertained. In the wider area, doned small quarry in a partly exhausted body of the about 1 km northeastwards of Laškov, fragments of Lower Badenian red-algal limestones (grey-yellow algal limestones have been ascertained spread about siliceous limestones with abundant bivalve shells, the fields (Novák, 1975; Jašková, 1998; Horčičko et especially of pectinids). Fragments of these rocks al., 2006). are spread even about the surrounding fields. In the Luleč. Lower Badenian oyster bank, its underlying western part of the quarry there occurs a layer of the rocks are represented by the Culmian sediments of the Lower Badenian basal gravels. In 2000, the extent of Drahany Upland (Spitzner, 1899, 1907; Čouka, 1925; the Miocene relic has been verified by 13 orientational Lang, 1933). holes in the fields around Taliánka; the thickness of Myslejovice. During the excavations of the fam- the limestone bench does not exceed 6 m. The underly- ily house foundations Lower Badenian grey clays with ing rocks are represented by Culmian greywackes and fragments of oyster valves have been ascertained. shales (the most eastern projections of the Culmian Section: 0.5 m variegated clays, 0.6 m grey-green clay rocks of the Drahany Upland) creating morphological- with oysters Crassostrea gryphoides, 0.6 m Quaternary ly conspicuous elevations (Jašková, 2002). In the sand – clay loam with tiny fragments of Culmian rocks and pit about 0.5 km north of Taliánka the base of Lower fragments of oyster valves (Jašková, Lehotský, 2009). Badenian limestones occurs, namely grey-yellow Olomouc–Neředín. By shallow boreholes there to grey calcareous sands, in places consolidated and were detected Lower Badenian pelitic facies overly- forming benches, consisting practically exclusively of ing basal clastic sediments of Early Badenian age: the Culmian rocks grains, and sporadically with fossils grey-green, yellow-grey, blue-grey to brown-grey cal- (bivalves – Jašková, 2002). careous and finely sandy clays (tegels), in places with Kobeřice near Prostějov. The abandoned sand pit ochre smudges, often with slight silty or sandy admix- is situated on the western periphery of Kobeřice, in the ture. These pelites contain abundant calcareous con- Hůrka Hill (277 m above sea level), near the road be- cretions and sometimes even macrofauna, first of all tween Hradčany and Kobeřice. The Lower Badenian oysters and other marine molluscs (Hrubeš, Čtyroká, basal clastic sediments – shallow-water marine grav- 1999; Čtyroká, Švábenická, 2000; Zapletal, 2004). els and sands (locally consolidated into conglomer- Přemyslovice. The Miocene (Lower Badenian) de- ates and sandstones with oysters) are deposited with posits near the village of Přemyslovice were mentioned discordance on the Lower Carboniferous sediments. by Spitzner (1899, 1907), and studied by Schwarz

93 (1946) and Novák (1975). New investigations were Terezské údolí. The natural park (since 1996) initiated after the findings of rich bryozoans-bearing (Horčičko et al., 2006). An accidental find of oyster sediments in the vicinity of Přemyslovice (Jašková, shells in 2003 (Čermák, 2007) in the slope above a 1998). The sedimentary succession rich in foraminif- sharp loop of the Šumice river about 3 km w. of the ers, molluscs, bryozoans, brachiopods and red algae railway stop Náměšť na Hané and 2 km NE of the bus was examined by 7 shallow boreholes (Zágoršek, stop in Laškov. During the field investigation (2005) Holcová 2009; Zágoršek et al., 2012). an accumulation of oyster shells in a small depres- Slatinky. The former sand pit, nowadays out sion within the Lower Carboniferous shales has been of existence and recultivated; Miocene (Badenian) confirmed under the Quaternary sediments (~0.25 m) sands and sandy clays 10–20 m in thickness, overly- – Jašková, Lehotský (2009). The oyster assemblage ing the Devonian limestones. Section from the base: is most probably redeposited and manifests relations fine-grained grey-green clayey sands and sandy clays with the close Laškov locality (Jašková, 1993, 1998). poor in fossils, in the upper part yellowish, in places Abundant boring traces of marine organisms (Entobia calcareous coarse-grained siliceous sands abundant in isp., ?Entobia isp., ?Trypanites isp., Gastrochaenolites fossils, in particular Foraminifera, – isp.) occur on the oyster shells (Hladík, 2009). (including oysters) and Gastropoda, Echinoidea, Cirri­ pedia, Ostracoda. Shallow-water sediments, littoral to neritic facies. The uppermost part of the section is loess Brno region (10 localities) (Pleistocene) – Wolf (1862), Toula (1893), Spitzner (1899, 1904, 1906, 1907), Schubert (1913), Remeš Bedřichovice. In the abandoned quarry west of the (1930), Kostka (1931), Kalabis (1934, 1937, 1938, 1949, village margin there occur Paleozoic limestones 1956, 1961a–d), Jedlitschka (1937), Roth et al. (1962), (Upper Frasnian) (Müller et al., 2000), transgressively Barth et al. (1971), Řehoř et al. (1978), Kupková (1981, overlapped by Lower Badenian calcareous clays with 1982, 1984, 1986, 1995), Kupková et al. (1987), Jaš­ oysters, nowadays preserved only in denudation relics. ková (1998), Zapletal (2004), Hladík (2009), Jašková, The oyster banks or limestone breccias rich in fora- Lehotský (2010), Kopecká (2012), Pekař, Lehotský minifers (Lenticulina) occur in the basal parts of the (2013). A relic of a typical oyster bank overlying the Lower Badenian calcareous clays even at some other Devonian limestones is situated about 50 m SW of the surrounding localities, for example in the Mokrá quar- former sand pit towards Kaple (Kupková, 1982). ry or on the Palacký Hill (Müller et al., 2000). Služín. About 1.5 km W of the village Služín on the Blažovice. Between the villages Blažovice and east margin of the wood Brus in the Stříbrný potok val- Holubice (S and SW of Holubice) the Lower Badenian ley near the lane towards Hluchov there were two relics rocks of the formation with algal limestones are crop- of Miocene sediments (in one of them formerly a sand ping out in two morphologically evident denudation pit). Nowadays, the locality is totally destroyed, as a relics, and unearthed in two pit quarries at the elevation pond has been excavated within the occurrence area of point 286.6. Sandy, medium- to coarse-grained orga- the Miocene sediments (Jašková, 1998). Sediments of nodetritic limestones and calcareous sandstones with the formation with the red-algal limestones with rela- abundant flora and fauna – Foraminifera, Anthozoa, tively abundant fauna and flora overlying sandy, mica- Mollusca, Vermes, Bryozoa, Echinoidea, Rhodophyta. ceous, calcareous claystones with shreds of shaly clay- The underlying rocks are represented by tegels (Paulík, stones (foraminifers, sponge spicules), higher ~2.5 m 1953; Novák, 1975; Nehyba et al., 2014). grey, fine-grained sandy calcareous claystone (sponge Borač. About 800 m NW of the village, to the left spicules, ostracods, foraminifers), ~0.70 m rusty of the road Borač–Doubravník, there are cropping out brown to greenish sedimentary breccia with a layer of small denudation relics of marine, dark green-grey, unconsolidated grey-white sandy limestone (0.2 m). grey-yellow or grey-blue, slightly sandy calcareous In the breccia sharply angular fragments of Culmian clays (tegels) extraordinarily rich both in macro- and shales dominate, large foraminifers (heterostegines), microfossils (Foraminifera, Anthozoa, Mollusca, Ostra­ sponge spicules, fragments of corals and bryozoans, coda, Vertebrata, nannoplankton) (Procházka, 1892, echinoids, molluscs, shark and fish teeth and frag- 1893; Tausch, 1895; Cicha et al., 1957; Cicha, Dornič, ments of red algae occur in the intersticial mass – a 1960; Hladil, 1976; Seitl, 1978). Tegels are deposited typical example of a surf zone. In the uppermost parts transgressively without basal clastic sediments direct- of the section white sandy limestone with recrystal- ly on the metamorphites of Moravicum. This locality lized foraminifers and grey, yellowish grey to rusty, was assigned s a regional faciostratotype of the Lower slightly sandy algal limestones with fauna and flora Badenian (Moravian) by Papp et al. (1978). – foraminifers, sponges, corals, bryozoans, molluscs Brno–Vinohrady. Old sand pits, partly recultivat- (bivalves – including oysters, gastropods), ostracods, ed, partly built in, north of the tram-line to the Líšeň echinoids, red-algae, ichnofossils – Spitzner (1899, Quarter of Brno (near Krásného Str.). Lower Badenian 1904, 1906, 1907), Blekta (1932), Kalabis (1934, basal clastic sediments, so-called Brno sands (fining- 1937, 1938, 1949, 1960), Novák (1975), Vysloužil upwards sequence, in places with benches of sand- (1981), Lehotský (2006), Zborník et al. (2013). stones and clay galls), and superposed calcareous

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

95 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 dium-large in size (≈ 5–10 cm), present a crenulated (De Serres, 1843), now determined as Hyotissa hyotis external surface, a resilifer zone rather boxy in shape (Linné, 1758). In our specimens the state of preserva- and a moderate thickness, therefore we refer them to tion is not good, so according to the previous literature Ostrea digitalina (Dubois, 1831). Ostrea leopolitana: we accepted it as Hyotissa hyotis. Ostrea postvesicu- Ostrea leopolitana Niedèwiedzki, 1909 has been de- laris: Taylor (1885) reports Ostrea vesicularis in the scribed by Friedberg (1936) and Radwánski (1973) as old collections from England, but it is not a Miocene Pycnodonta leopolitana from Poland, now in synony- species; according to the small size (5 cm) and the my with Neopycnodonte navicularis (Brocchi, 1814); main features of the valves (elongated outline, smooth our specimens fit perfectly with Neopycnodonte na- external surface, thickness and depth on the left valve) vicularis as for size than for morphological patterns. the specimen present in our collections could be re- ferred to Neopycnodonte navicularis (Brocchi, 1814). Ostrea cf. fimbriata: Ostrea cf. fimbriata is attributed 2. Revised oyster species in the collections to Ostrea digitalina (Dubois, 1831) by Pekař (2013). Ostrea crassicostata: Ostrea crassicostata is described Here we present the systematic descriptions of the 4 by Antunes et al. (1999) as var. gigantea (not Miocene updated species – Crassostrea gryphoides, Hyotissa species), from sediments of Portugal. In our collec- hyotis, Neopycnodonte navicularis and Ostrea digi- tions these specimens are characterised by a rounded talina – occurring in the collections. The determina- shape, large dimensions, crenulation on the external tions are also based on the literary data, mainly on surface, muscle scar (left valve) in central position of the publications by Stenzel (1971), Schultz (2001– the internal surface; based on these facts we refer them 2005), Mandic, Harzhauser (2003), D’Amico et al. as morphological variations of Hyotissa hyotis (Linné, (2012), etc. 1758). Ostrea eduliformis: Ostrea eduliformis is a taxon referred in literature to Bajocian, by Pockrandt Class Bi v a l v i a Linné, 1758 (1984), and should not to be considered in Badenian Subclass Pt e r i o m o r p h i a Beurlen, 1944 samples; as there are not other indications from the Order Pt e r i o i d a Newell, 1965 collector, we should think it has been confused with Suborder Os t r e i n a Ferussac, 1822 Ostrea edulis (Linné, 1758). Our specimens are me- Superfamily Os t r e o i d e a Rafinesque, 1815

96 Family Crassostreidae Scarlato, Starobogatov, 1979 Description. The shell is thin (vesicular structure) and Genus Crassostrea Sacco, 1897 of small size, normally 5 to 10 cm long; the shape is Ostrea (C.) virginica (Gmelin, 1791). ovoidal-elongated. The posterior and anterior mar- Type species: Ostrea virginica Gmelin, 1791 gins have an edge. The right valve is plano-concave; Crassostrea gryphoides (Schlotheim, 1813) its ligament area presents a median depression. The (Plate I, a–h; Plate III, c, f) external surface lacks the sculpture. The adductor in- sertion is wide and it is close to the hinge and to the Description. Shells inequivalve, elongated, sizes are ventral edge. The chomata are present in the zone of much greater than other species. A well-developed the ribs extension. umbonal cavity in the right valve; the left valve is more Notes. The name navicularis has for a long time been convex or quite flat. The adductor muscle imprint is in used at the level of either variety (Hoernes, 1870; postero-ventral position, outline crescentic with fair- Friedberg, 1934; Kojumdgieva, 1960), or subspe- ly sharp corners in the dorsal margin. The shells are cies (Caprotti, 1972) of N. cochlear (Studencka et al., broader next to the ventral margin and thicker in the 1998). It was also demonstrated that the resilifer was umbo zone. On the external surface there are folded too big in order to consider N. cochlear (Poli, 1795) ribs and growth lamellae. Chomata are absent along and its variety navicularis as two different species the entire margin of the both valves. (Freneix, 1975). Videt, Néraudeau (2002) compare Material. Bedřichovice: 6883 (VMO); Drysice: 30102 Poli, 1795 (sensu Freneix et (VMO); Laškov: 087175/1-72(MPP); (MPP); 30446, al., 1988) to Neopycnodonte navicularis (Poli, 1795), 30447, 30448 (VMO); Luleč: 190774/2, 190774/3, from Miocene of Spain according to the paleoecologi- 190774/8, 190774/9, 190774/10 (MPP); Myslejovice: cal point of view. 190442, 190443, 190444, 190445, 190447, 190448 Material. Borač: 4739, 5845 (VMO); Brno: 6540, (MPP); Slatinky: 7746, 7801, 7889–7891, 7853, 7923, 6689 (VMO); Čelechovice: 073275, 073762, 078729 7942–45, 7955, 7956, 7957–7959, 7960–7962, 7971, (MPP); Lomnička: 4595, 4596 (VMO); Prace: 1527 7979, 7992, 8000–8003, 8009–8013, 8014–8017, (MPP); Slatinky: 7736, 7737, 7760, 7761, 7778, 7779, 23188–23189–23190, 28227, 28233, 28712, 28713, 7812–7814, 7815–7818, 7901–7904, 7918–7922, 30101, 30168, 30186, 30270, 30327, 30420–30421 7963–7967, 26731, 26732, 30211–30215 (VMO); (VMO); Vinohrady (Brno): 051886 (MPP). Vinohrady (Brno): 073291 (MPP).

Family Vyalov, 1936 Family Os t r e i d a e Rafinesque, 1815 Subfamily PYCNODONTEINAE Stenzel, 1959 Subfamily Os t r e i n a e Rafinesque, 1815 Genus Hyotissa Stenzel, 1971 Genus Ostrea Linné, 1758 Hyotissa hyotis (Linné, 1758) Ostrea digitalina (Dubois, 1831) (Plate II, g–l; Plate III, d, g) (Plate II, a–f; Plate III, b, h)

Description. The valves are rather circular, with the Description. The left valve is more convex than the adductor muscle in the center; a thick shell not really right one (elevated marginal area and coarse radial concave with the ribs in the external surface. The mar- ribs) and it is used in cementing. The outline can gin is clearly crenulated. be oval or subquadrate. The size is medium and it Notes. Hoernes (1870) referred the Austrian speci- reaches 15 cm. mens to Ostrea plicatula Gmelin and O. crassi- Notes. According to Studencka (1986) it is possibly costata Sowerby, but it was attributed to Pycnodonte synonymous with the Atlantic and Mediterranean squarrosa (De Serres, 1843) by Studencka et al. Ostrea frondosa De Serres; it is considered a fully (1998). There were demonstrated the differences in marine oyster, settled on different kinds of hard the adductor muscle insertion in squarrosa and hyo- substrata. We can compare it with the Ostrea edu- tis, considered as intraspecific variability (Freneix lis (Schultz, 2001–2005), typical from Recent et al., 1988). Ostrea crassicostata is reported in the Mediterranean taxa, living in colonies in shallow wa- sediments of Portugal (Antunes et al., 1999) as var. ters about 10 meters (Milisic, 1991). It can be also gigantea [Pycnodonta squarrosa (De Serres, 1843), morphologically compared to Ostrea fimbriata, re- now Hyotissa hyotis]. ported as cf. Crassostrea fimbriata in the sediments Material. Bedřichovice: 1579, 1580, 5873, 6400, of Turkey (Hakyemez, 1989). 6519, 6946 (VMO); Blažovice: 6899 (VMO); Brno: Material. Borač: 4738, 6541 (VMO); Brno: 6516, 2417 (VMO); Hluchov: 139623/3(VMO); Slatinky: 6517, 6691 (VMO); Černá Hora: 6688 (VMO); 28225, 28226, 28234 (VMO); Terezské údolí: Kelčice: 078730 (MPP); Kobeřice: 8046, 8047 190797/7, 190797/10 (MPP). (VMO); Lomnička: 5874 (VMO); Luleč: 4737, 4743 (VMO); Olomouc: 26873, 26874 (VMO); Genus Neopycnodonte Stenzel, 1971 Opatov: 6412 (VMO); Slatinky: 1680, 1681, 6722, Neopycnodonte navicularis (Brocchi, 1814) 6723, 7017, 7743–7745, 7799, 7800, 7879, 7905– (Plate I, i–r; Plate III, a, e) 7907, 8011, 23187, 27759, 27774–27778, 27780,

97 PLATE I ТАБЛИЦА I

98 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: Discussion 6126 (VMO). In addition to the systematic review, where the old no- Summary. After the revision work we reduced the menclature has been updated in 4 species (Crassostrea number to 4 species (~22 original), namely Crassostrea gryphoides, Neopycnodonte navicularis, Ostrea digi- gryphoides, Hyotissa hyotis, Neopycnodonte navicular- talina and Hyotissa hyotis) (Fig. 1 A–D), we also tried is, and Ostrea digitalina more Ostrea sp.. Crassostrea to recognize the oyster percentage distribution accord- gryphoides and Ostrea digitalina are most abundant ing to the provenance locality and to the morpho-spe- (Olomouc); the most numerous collections come from cies kind. In spite of the relatively high numbers of the Slatinky (78%) and Laškov (6%). Ostrea digitalina studied Lower Badenian oysters, it was generally im- represents the most abundant taxon in Brno region possible to use them for more detailed paleoecological [for this region: Bedřichovice (20%), Brno (17%), interpretations. The main reason was the heterogene- Luleč (13%) and Borač (13%)]. Grouping all the ity of the material, namely the limited knowledge of samples from the Moravian part of the CF (excluding the individual collections histories and of the circum- Ostrea sp.) we obtain: Crassostrea gryphoides (38%), stances connected with their formation: for example Neopycnodonte navicularis (14%), Ostrea digitalina we could not verify whether the species composi- (36%) and Hyotissa hyotis (12%). tions or numbers of individuals are not significantly

99 PLATE II ТАБЛИЦА II

100 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 Foredeep sediments situated in the Boskovice Furrow, with the real situation at the localities. Even the data as presented for instance by Doláková et al. (2008). In on exact original positions/orientations of individu- spite of Crassostrea, this taxon prefers greater depths als in the sediments are entirely missing. Crassostrea below 40 m (Bagdasaryan et al., 1966; Zágoršek et gryphoides occurs mostly in Bedřichovice, Laškov, al., 2009). Typically the oyster as Neopycnodonte Drysice, Luleč, Myslejovice, Slatinky and Vinohrady, forms dense overgrowths constituting oyster buildups, namely in the area close to the present western mar- which are usually several meters thick and generally gin of the Carpathian Foredeep (Fig. 1A); this could consist of articulated, massive shells, as seen, for ex- reflect paleogeographic conditions related to the ample, in the Upper Badenian deposits of Roztocze depth and morphology of the Foredeep. According to in Poland (Pisera, 1985; Studencka et al., 2005), but Jimenez et al. (1991) for Almanzora Corridor (Spain), the individual oyster buildups in the Moravian part of in the past, Crassostrea banks developed basinward the Carpathian Foredeep have distinctly smaller areal of coral reefs, very different from actual style of life, extents and thicknesses in comparison with Poland, in shallow-intertidal to shallow sea water with low- being related to the very different conditions of ba- ered salinity. Hyotissa hyotis is reported in two dis- sin configuration. Ostrea digitalina mainly occurs in tinct zones: at Bedřichovice and Blažovice in the Brno eleven localities (Fig. 1D), situated not only with- area, and at Hluchov, Slatinky and Terezské údolí in in the present-day extent of the CF in the Brno and the Olomouc area (Fig. 1B); it prefers shallow sub- Olomouc areas, but also in the isolated relics of CF tidal habitats down to 30 m with higher-energy seawa- sediments (Borač, Černá Hora, Lomnička and Opatov) ter conditions (Bieler et al., 2004; Mikkelsen, Bieler, described in literature from palaeontological point of 2008; Pekař, 2013; Pekař, Lehotský, 2013; Hladilová view (Hladil, 1976; Seitl, 1978; Hudec, 1986; Pek, et al., 2014). Neopycnodonte navicularis is preva- Mikuláš, 1996; Seko, 2011; Mihulková, 2013, etc.). lently present in Brno, Čelechovice, Prace, Slatinky, Ostrea digitalina is typical for a rocky coastline envi- Vinohrady and to W in Borač and Lomnička (Fig.1 ronment with dominant wave action (highly exposed C); from paleogeographic point of view, the last two rocky medio/sublittoral, intertidal to shallow subtidal) localities belong to denudation relics of the Carpathian (Zágoršek et al., 2009). Similar energy-rich rocky

101 PLATE III ТАБЛИЦА III

102 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 Olomouc regions); the distribution of Neopycnodonte are interpreted even in Austrian localities (Mandic, is similar to Hyotissa’s one, with the presence in the Harzhauser, 2003). isolated relics of Borač and Lomnička; O. digitalina The higher proportion of right valves (60%) in the specimens are decidedly more widespread in all the collections may result from differential transport, as area (also in the N to Opatov), without a particular these valves are thinner and smaller than the left ones distribution range. Although taking into account the (40%) attached to the substrate, and are easily remobi- collection bias and their randomness and arbitrariness, lized even by weak currents after death and decompo- these results represent a certain contribution to paleo- sition of the soft parts (Lopes, 2011). geographic interpretations of the Carpathian Foredeep in Moravia during the Badenian.

Conclusions Acknowledgements: This work was supported by the ESF Research Project POST-UP II, reg. No. We have studied the Badenian oyster collections from CZ.1.07/2.3.00/30.0041 of the Palacky University 23 localities of the Carpathian Foredeep in Moravia Olomouc (Czech Republic). The authors are grateful deposited in Vlastivědné Muzeum in Olomouc and to Dr. Andrzej Pisera (Institute of Paleobiology, PAN, Muzeum Prostějovska in Prostějov (Czech Republic). Warsaw) for his important help in the study of the The original ~22 species with old nomenclature have Ostreoidea, and to Dr. Oleg Mandic (Naturhistorisches been revised to 4 species: Crassostrea gryphoides, Museum in Vienna) who made valuable sugges- Hyotissa hyotis, Neopycnodonte navicularis and tions in the mollusc systematics and palaeontology. Ostrea digitalina. In the analyzed material Crassostrea Acknowledgement is also referred to Dr. Tomáš gryphoides is represented by ~38%, Neopycnodonte Lehotský (Faculty of Science, Palacky University and navicularis by ~14%, Ostrea digitalina by ~36% and Vlastivědné Muzeum of Olomouc) and Dr. Vladimíra Hyotissa hyotis by ~12%. Crassostrea specimens Jašková (Muzeum Prostějovska v Prostějově) for coming from the localities, which are situated today their support during the preparation of this paper. The at the western margin of the CF (from Vinohrady UPLIFT of Palacky University in Olomouc is thanked to Laškov); Hyotissa specimens are clearly divided for helpful observations and remarks, facilitating the in two different groups of localities (both in Brno- final English revision.

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