The stratigraphical importance of the Viviparus boeckhi Horizon of Serbia

Quaternary International 292 (2013) 101e112

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The stratigraphical importance of the Viviparus boeckhi Horizon of Serbia Tivadar Gaudenyi a, *, Dra zenko Nenadi c b, Mladjen Jovanovi c c, Katarina Bogi cevi cd a

Geographical Institute “Jovan Cvijic” of the Serbian Academy of Sciences and Arts, Djure Jaksica 9, 11000 Belgrade, Serbia Department of Geology, University of Belgrade, Faculty of Mining and Geology, Kamenicka 6, 11000 Belgrade, Serbia c Chair of Physical Geography, Faculty of Sciences, University of Novi Sad, Trg Dositeja Obradovica 3, 21000 Novi Sad, Serbia d Department of Paleontology, University of Belgrade, Faculty of Mining and Geology, Kamenicka 6, 11000 Belgrade, Serbia b

a r t i c l e i n f o

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Article history: Available online 6 December 2012

Fossils of non-marine molluscs are among the most prominent in Pleistocene deposits. With stabilization of taxonomies in the 19th century, numerous publications appeared with faunal lists of Quaternary warm stage molluscs, but few attempts were made to use those data for interpretation. The purpose of this work, along with reevaluation of sites and their molluscan assemblages described in the literature, is to make it possible for non-marine molluscan faunas to be used as biostratigraphic indicators. Biostratigraphic data obtained in this work are comparable with parallel investigation of other fossil records and lithostratigraphy, combined with geochronologic methods. The four most important localities with Viviparus boeckhi in Serbia are Subotica, Sombor, Zrenjanin, and the vicinity of Belgrade. According to the available data, the molluscan assemblage of the V. boeckhi Horizon is identified as fluvial warm stage assemblage which corresponds to the oldest part of the Pleistocene Corbicula Beds. It is generally considered that the age of the V. boeckhi Horizon(s) corresponds to the younger parts of the Early Pleistocene. However, according to the regional data, the V. boeckhi Horizon of Serbia is younger than the Paludina Beds (of PlioceneeEarly Lower Pleistocene age). According to the vertebrate fauna, it is older than the upper Biharian stage (younger part of the Middle Pleistocene). Ó 2012 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction Non-marine molluscs were among the earliest Pleistocene fossils (beside the vertebrate fauna) to be found in the territory of today’s Republic of Serbia (e.g. Halaváts, 1895; Adda, 1899; Koch, 1907). At that time, composition of faunas was generally presented in the form of species lists rather than by numbers of individual shells, although notes of “rare” or “abundant” taxa were sometimes added. These late 19th and early 20th century authors tried to use molluscs for biostratigraphic zonation as was done for older geological formations, “a task for which they are inherently rather badly fitted” (Keen, 2001), and perhaps less suitable than other fossil groups, notably pollen. Investigations of Quaternary molluscs after the 1950s, particularly those conducted by Sparks (1964) and Kerney (1977) in the United Kingdom, Puisségur (1976) in France and Lo zek (1964) in the former Czechoslovakia stressed the increasing trend of using molluscs for palaeoenvironmental reconstruction rather than for * Corresponding author. E-mail addresses: [email protected] (T. Gaudenyi), [email protected] (D. Nenadi c), [email protected] (M. Jovanovi c), k.bogicevic@ rgf.bg.ac.rs (K. Bogi cevi c). 1040-6182/$ e see front matter Ó 2012 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2012.11.034

biostratigraphical purposes. Krolopp in the 1960s and 1970s revised the existing data. He established the biostratigraphical subdivision for Hungary on the basis of the mollusc material from numerous boreholes. The analysis included the material from outcrops and more than 100 boreholes from the area of the Middle Danube Basin. These investigations included the results of previous works and supporting evidence from lithology, tectonics, and biostratigraphical proxies. Based on quantitative analysis, these data enable not only recognition of the former environment dynamics, but they also can be used for (bio)stratigraphical purposes (Krolopp, 1970). In the southeastern part of the Middle Danube Basin (nowadays Republic of Serbia) the most important fluvial polycyclic deposits were identified in artesian wells (Halaváts, 1895, 1914a, 1914b; Adda, 1899; Pálfy, 1912; Laskarev, 1922, 1938). The revision of the studies of Halaváts (1895, 1914a, 1914b) was done by Krolopp (1970, 1976a, 1976b, 1977, 1978a, 1978b) and Kretzoi and Krolopp (1972). These new concepts concerning the biostratigraphy of the Middle Danube Basin, were, however, not acknowledged by Serbian scientists (e.g. Stevanovic et al., 1992; Rabrenovi c et al., 2003), and were not even used for correlation during the geological surveying for the general geological map 1:100 000 (e.g. Trifunovi c, 1996). The aim of this work is to try, using the regional data, to clearly

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summarize the developments and to justify the use of the Viviparus boeckhi Horizon as a biostratigraphical unit of the fluvial Pleistocene warm stage sediments of Serbia. 2. Principles Species-rich faunas from the warmest parts of the warm/ temperate stages (interglacials) are the only ones that are suitable for development of a molluscan biostratigraphical scheme in the current state of knowledge. The faunas from the early and late parts of warm stages usually comprise a restricted number of species, and are not directly comparable with the fluvial climatic optimum faunas, so it may be difficult to fit them into any general biostratigraphical scheme. The faunas that have been used in this work come from fluvial deposits which have the advantage of usually originating from similar facies, and thus are directly comparable. During floods, rivers accumulate the inhabitants of different ecological niches of the river and floodplain in particular locations, as was recognized by Sparks (1964). This homogenization of molluscs from different environments, although sometimes causing problems in interpretation, also allows comparison of faunas of the same or different ages (Keen, 2001). As became apparent to late 19th and early 20th century authors, fluvial assemblages of warm stages consisted overwhelmingly of very similar sets of species that retreated from the northern and central parts of the Middle Danube Basin in each cold stage and returned again during warm stages. However, the use of molluscan fauna for developing a biostratigraphic scheme is made possible by the fact that a small number of those species became extinct. The occurrence of extinct species allows the use of the last appearance datum (LAD) of a classical indicator. The presence/ absence of species now exotic to the Middle Danube Basin allows the possibility of characterising each temperate stage by a combination of data on extinctions and presence/absence. The utility of this approach (despite the dangers of using “absence” as a criterion) is improved by using only the faunal assemblages from fluvial deposits from climatic optima of warm stages. If conditions are closely comparable, but particular species are missing, the major differences between the faunas are not due to any environmental cause, but to the occurrence or lack of a species in the Middle Danube Basin during a particular time. One further problem in using fluvial assemblages is that molluscan faunas tend to comprise any available shell in a river catchment. This will include primarily specimens of the living fauna, and recently dead shells. However, given the cyclic nature of Quaternary sedimentation and the operation of the rivers within closely similar height ranges for much of the Early and Middle Pleistocene, reworking of molluscan material from earlier deposits cannot be discounted. Many species of fluvial gastropods have thin shells which are easily broken in post mortem transport, so an increased state of wear or fragmentation may not necessarily be an indication of recycling deposits of different ages. Reworking in bivalves can be more easily identified. Meijer and Preece (2000) gave examples of the Pleistocene Corbicula shells which were reworked and deposited into younger (MIS 5) sediments from the earlier deposits in both England and The Netherlands. However, the redeposition of warm climate species can be very difficult to detect (Keen, 2001). Also to be considered in establishing a biostratigraphic scheme is that presence/absence of species across the southern parts of the Middle Danube Basin may be controlled by biogeographic boundaries between the catchments. The modern distribution of species is to some extent artificial, and does not reflect the situation in the warm Pleistocene stages. Since the 19th century, the development of canals connecting together all major catchments (e.g. the Danubee TiszaeDanube canal system) in the “canal basin” in the southeastern part of the Middle Danube Basin has tended to homogenize fluvial

faunas. There is no evidence, either in earlier Holocene times or in Pleistocene warm stages, that there were any biogeographical boundaries which could cause restriction of species to individual catchments and so give a false indication of presence or absence. The uniformity of fauna across a relatively small area such as the Pannonian Plain (Alföld) is controlled by the rapid dispersal potential of many aquatic species, allowing a number of taxa to colonize the whole continent in the present century. Many examples of rapid rates of spread of freshwater molluscs include species introduced into new areas by human activity; for example the introduction of Potamopyrgus antipodarum (Gray) into Europe, probably from New Zealand since 1859 (Haynes et al., 1985; Kerney, 1999); and a comparably rapid spread of Corbicula fluminea (Müller) into the Rhine system since 1985 (bij de Vaate and Greijdanus-Klaas, 1990; Kinzelbach, 1991; Turner et al., 1998). However, despite of this element of artificiality in their spread, these examples show the potential of non-marine molluscs for the colonisation of new habitats. Thus, even without the linking effects of canals between river basins, the spread of freshwater species from catchment can be rapid, and would prevent the development of different biogeographic provinces in an area as small as the Pannonian Plain. 3. The main characteristics of the V. boeckhi Horizon 3.1. Defining the V. boeckhi Horizon After Krolopp’s analysis based on more than 100 sites of the Middle Danube Basin, it became evident that a unique freshwater fauna was formed in the fluvial environment. A total of 80% of the species present in Pleistocene fluvial fauna still inhabits the freshwater river network of the Middle Danube Basin. However, the most significant species were those which disappeared from the Middle- and/or Upper-Pleistocene deposits. They represent the main biostratigraphical characteristics of the Pleistocene fluvial fauna. Several species lived only in some phases of the Early Pleistocene. This knowledge enables establishment of biostratigraphical subdivisions based on the malacofauna. The Lower Pleistocene river network of the Middle Danube Basin (especially in the lowlands of Serbia, Croatia, and Hungary) was reconstructed on the basis of the presence of extinct species which have connections with the Pontic region (Bessarabia, Moldavia, and Ukraine) (Krolopp, 2002b). The research results suggest that a similar river network system existed then with climatic conditions as in the Middle Danube Basin. The origin of the Middle Danube Basin fluvial fauna along with the existing data and palaeogeography suggests the connection with the south and southeastern Slavonian s.l. and Pontic (Black Sea) region. The Lower Pleistocene stagnant freshwater (slow moving or lacustrine) fauna was very similar to the modern assemblages. Neogene taxa had a continent-wide distribution during the whole Neogene (Soós, 1943; Krolopp, 1973) with similar biotope conditions. Among the major influences were Neoalpine tectonics and climatic evolution, which were directly reflected in the development of the river network system of the Quaternary. At the end of the Neogene, palaeogeographical and climatic changes caused a significant transition in molluscan fauna (especially the Central European), which resulted in the appearance of the modern molluscan fauna. In the Lower Pleistocene fauna of the Middle Danube Basin, a significant number of extinct species or species which cannot be found in the area of modern distribution were still present. The disappearance of the fauna begun mainly in Middle Pleistocene due to climatic changes, with separation and decreased variety in the faunal composition. This new impact on fauna richness was caused by migrations, and started with the beginning of the last warm stage. Of great importance was the quantitative analysis of the palaeontological material from the boreholes collected during the

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systematic geological survey of the Great Hungarian Plain/Pannonian Plain (Alföld) in the 1960s. V. boeckhi (Halavats) was found in the explanatory drill cores of the Geological Institute of Hungary from numerous sites. Drilling used dry techniques with double walled coring, which prevented the mixing of material of different age during the explanatory drilling. After the revision of the old data and comparing them with the new results, Krolopp (1970) stated the following: 1) V. boeckhi is always found in the characteristic assemblage, which indicates a fluvial environment. 2) The recent results based on the assemblage in which V. boeckhi was found show that all the species have a Pleistocene record. 3) The conditions of the preserved material of V. boeckhi do not differ from the other shells found in the same assemblage. The V. boeckhi shells are preserved in excellent condition, which excludes the longer post mortem fluvial transport of the shells. 4) However, in many parts of the Great Hungarian Plain, the thin Quaternary sediments are underlain in relatively high positions by the blocks of the Pannonian Stage. According to the datasets, the Upper Pannonian facies were formed in a brackish environment (Sümeghy, 1929, 1939). To date, there are no Upper Pannonian deposits which contain shells of V. boeckhi. In the sites from the Transdanubian region of Hungary (Tengelic, Kisláng) where V. boeckhi was found, the vertebrate fauna remains also support the proposed Pleistocene age. 5) In the basins such as the Pannonian Basin System, where sinking movements are predominant (especially in the area of the Great Hungarian Plain), the older sediments are rapidly covered with the younger ones. The possible addition of material from the peribasinal area is excluded, because the palaeontological material is well preserved and shows no traces of transport or mixing with the material from the older strata. The deposits are clearly undisturbed, and no ex situ layers can be identified. The area of the palaeogeographical distribution of the assemblage of the V. boeckhi Horizon (described by Halaváts, 1888a, 1888b) comprises more than 40 000 km2 of the Middle Danube Basin, where the fluvial Lower Pleistocene facies were identified (Krolopp, 2002b). In the Pannonian Plain, the area of the Pleistocene Corbicula Beds (“Corbicula fluminalis Beds” in the literature; but as the status of the Pleistocene Corbicula is unclear, the term “Pleistocene Corbicula” is used, as in NW Europe and recommended in Meijer and Preece, 2000) stretches from the northernmost site of Tápiósüly (Hungary), to southwest sites Kalocsa, Püspökladány (in Hungary), the vicinity of Osijek (Croatia), and to Makis, Ada Ciganlija and Pan cevo (all in the vicinity of Belgrade, Serbia) (Fig. 1). These beds should contain the V. boeckhi Horizon, because V. boeckhi is a regional species of the Middle Danube Basin. The characteristic environment of this species was a fluvial environment in warm/temperate stages (e.g. Krolopp, 1970). In the molluscan assemblage of the V. boeckhi Horizon, besides the index species V. boeckhi, the most common species are: the Pleistocene Corbicula [“C. fluminalis (O.F. Müller)”], Fagotia acicularis (Férrusac), Fagotia esperi (Férrusac), Lithoglyphus naticoides (Pfeiffer), Amphimelania hollandri (Férrusac), Neumayria crassitesta (Brömme), Theodoxus danubialis (Pfeiffer), Theodoxus prevostianus (Pfeiffer) and Theodoxus fluviatilis (Linnaeus). 3.2. Most important taxa of the V. boeckhi Horizon Pisidium clessini Neumayr, 1875 This species was identified in Lower and Middle Pleistocene fluvial deposits at many places in the Western Europe. In the area of

Fig. 1. Viviparus boeckhi (Halaváts, 1888a, 1888b) from Makis (Belgrade, Serbia) borehole profile (Photo: Vladan Radulovic).

the Middle Danube Basin, the species P. clessini was abundant only in fluvial deposits. According to the accessible data, this species is missing from the Upper Pleistocene fauna (Krolopp, 2002b). Viviparus acerosus zsigmondyi (Halaváts, 1889) It was defined as an endemic Early Pleistocene (i.e. Lower Biharian, which is partly equivalent of the “Cromerian”, after Preece, 2001) subspecies of the recent V. acerosus (Bourguignat). According to the lithology, its biotope was the fluvial environment (Krolopp, 1980). Viviparus boeckhi (Halaváts, 1888) The first description of “Vivipara Böckhi” was made by Halaváts (1888a, p. 176). The revision of V. boeckhi (Halavats, 1888a) has been done by Krolopp (1976a, p. 200). The holotype and paratypes are stored at the Geological Institute of Hungary in Budapest (Fig. 2). Halaváts (1888a, 1888b) concluded that its age corresponds to the “Levantine Stage” (Pliocene). In the latest revision (Krolopp, 2002a), the age of this species is Early Pleistocene. It is the most abundant and most characteristic species from the Lower Pleistocene in the Pannonian Plain. The data from the Hungarian sites (more than 30) were of importance for understanding and defining the V. boeckhi species according to the palaeogeography and stratigraphical position. According to some well preserved individuals from the collection of the Geological Institute of Hungary, dimorphism was present in this species. The oval to circular shells belonged to females, and the slightly elongated ones to males. The bigger individuals of V. boeckhi Halaváts (1889a, 1889b), first ascribed to a “new species of Viviparus artesica (Halavats)”, are described in the Krolopp revision (1970, 1980) as a variety of the V. boeckhi male individual. The shells of these individuals are similar to the recent Viviparus species which have an ornamentation of 4 dark brown lines on the shell. In the excellently preserved V. boeckhi from Mindszent K-88 borehole, even the two yellowish brown lines (faded in other specimens) were preserved and visible (Krolopp, 1970). Krolopp (1970) suggested that Viviparus glacialis (S. Wood) and Viviparus diluvianus (Kunth) were regional synchronous species which were found in NW and W Europe. However, in the later studies of Dutch material by Meijer (1989), it was concluded that V. diluvianus ranged in age from Late Tiglian to Holsteinian, whereas for V. glacialis a Tiglian age has been proposed. The species are different but they show taxonomical connections, so it seems that the ancestor of both species inhabited Bessarabia (Mangikian, 1968; Motuz, 1968) and through the (palaeo)Danube and (palaeo)Rhine river network system reached to the north to southern Britain. Viviparus fauna from the Lower Pleistocene (and older) deposits of Moldavia (Viviparus kagarliticus (Lung), Viviparus tirasopitanus

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Fig. 2. Vivipara species from the collection of the Geological Institute of Hungary (from Krolopp, 1976a, 1976b) 1aeb “Vivipara zsigmondyi n. sp.” ¼ Viviparus acerosus zsigmondy }vásárhely, Hungary); 2aeb Viviparus acerosus zsigmondy (Halavats) (MÁV Railway Station well in Szeged, Hungary); 3aeb Viviparus acerosus (Bourg.) (Halavats) holotype (Hódmezo recent (Szeged, Hungary); 4aeb Viviparus contectus (Mill.) (Matészalka, Hungary); 5aec Viviparus contectus (Mill.) juvenile (Matészalka, Hungary); 6aec “Valvata levantica n. } vásárhely, Hungary); 7aeb Viviparus boeckhi (Halavats) holotype (Szentes artesian well, Hungary); 8aeb Viviparus boeckhi sp.” ¼ Viviparus contectus (Mill.) juvenile (Hódmezo }vásárhely, Hungary), 10aeb Viviparus boeckhi (Halavats) holotype (Szeged, artesian well-Tisza Lajos krt, Hungary); 9aeb “Viviparus artesica n. sp.” ¼ Viviparus boeckhi (Hódmezo (Szentes artesian well, Hungary).

(Pavl), Viviparus subcrassus (Lung)) seems to be connected with the Viviparus species of the Middle Danube Basin (Motuz, 1969). Further investigations on systematics are needed to solve the problems of evolution and taxonomy of Vivipara species. Theodoxus prevostianus (Pfeiffer, 1828) This is one of the characteristic species of Lower Pleistocene fluvial sediments. In its recent range it can be found in thermal springs. In earlier references it was mentioned as a relict species from the Tertiary. It seems that water temperatures of the rivers during the Early Pleistocene warm/temperate stages was sufficiently warm for this species, while after the climatic changes in the Middle and Late Pleistocene in Central Europe it could survive only in thermal springs (Krolopp, 1970, 1973).

In some specimens from the Lower Pleistocene material from Hungary, some morphological marks common to both T. prevostianus and T. danubialis were present. According to these marks, T. prevostianus and T. danubialis had a common ancestor with a mixture of common features (Theodoxus ex gr. prevostianuse danubialis) in the Early Pleistocene (Krolopp, 1970, 1973). Bithynia (¼Neumayria) crassitesta (Brömme, 1885) This species was in earlier works mentioned as Late Miocenee Pliocene, on the basis of the borehole shelled material. More recent data from NW Europe (e.g. The Netherlands) suggest Early and Middle Pleistocene age (Meijer, 1986). The shells of this species are quite rare in the assemblage, although their characteristic opercula were more abundant in the Lower Pleistocene deposits of

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the Middle Danube Basin. Their presence in younger deposits cannot be confirmed (Krolopp, 1970, 1977). Fagotia taxa In the modern freshwater fauna, two species of Fagotia (F. acicularis and F. esperi) are easily identified and separated according to their morphological marks. In some Lower Pleistocene deposits there are specimens of F. acicularis and F. esperi with mixed characteristics of both species (Fagotia ex gr. acicularis-esperi). As in the case of Theodoxus ex gr. prevostianusedanubialis, this proves common origin and great evolutionary importance of these two species in the Middle Danube Basin (Krolopp, 1970). Ferrissia pleistocaenica Krolopp, 1978 Ferrissia pleistocaenica is very similar to the recent Ferrissia wautieri (Mirolli) that was also found in the Upper Pliocene of France. An important fact regarding the autochthony of F. wautieri is the occurrence of the genus in the Pleistocene. Previously, the Ferrissia species were known only from Tertiary deposits (Wautier, 1975). The occurrence of F. pleistocaenica in the Lower Pleistocene (fluvial facies with gravels and sand) may be regarded, at least in part, as the consequence of migration of the genus first to southern Europe, before the cold impact of Pleistocene climate, and then back to the north (after Krolopp, 1969, 1973) (Krolopp, 1978a, 1978b). Parmacella kormosi Krolopp, 1978 Previously, the Parmacella genus was unknown from the Pleistocene of Europe, although some fragmentary specimens were found in the borehole material from the Lower Pleistocene of the Great Hungarian Plain (Kretzoi and Krolopp, 1972). The age of the assemblage in which Parmacella kormosi was found at Szabadhídvég is the Lower Pleistocene, and according to vertebrate fauna, the Lower Biharian stage (Krolopp, 1978a, 1980). The Pleistocene Corbicula [Corbicula fluminalis (Müller, 1774)] The systematics of Corbicula are in urgent need of revision, but it seems that only one polymorphic species (C. fluminalis) occurred in the Pleistocene of NW Europe (Meijer and Preece, 2000). A similar situation was observed in the Middle Danube Basin. As the systematic problems are unresolved, the compromised term “Pleistocene Corbicula” is used here. The Pleistocene Corbicula is one of the most famous Pleistocene warm stage (interglacial) guide fossils. C. fluminalis is recognized as a successful invasive species. In Hungary it seems to be confined to the Lower Pleistocene deposits (Krolopp, 1988), although Krolopp (1978b) earlier suggested the Early and the beginning of the Middle Pleistocene. The Serbian record also points to the Early and Middle Pleistocene interglacials (e.g. Rakic, 1977a, 1977b). For palaeoecological reconstruction, the most useful were the horizons with the Pleistocene Corbicula which, according to the modern analogues of C. fluminalis, suggests river temperature values between 22 and 23  C in JuneeAugust (Sinclair and Isom, 1963; Krolopp, 1970). 4. The stratigraphical position of the V. boeckhi Horizon 4.1. Historical background of the definition of the V. boeckhi Horizon in the Middle Danube Basin (before WWI) In Serbian part of the Pannonian Plain/Alföld, only a small number of outcrops with great thickness of fluvial polycyclic sediments have been found. The most valuable data came from

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artesian well boreholes. The artesian wells were drilled in search for stable and quality water supplies for the towns or municipalities. Zsigmondy, an engineer with Hungarian royal recommendation, conducted artesian well drilling in a great number of cities of the Austro-Hungarian Empire in the last decades of the 19th century (Zsigmondy, 1872). The material from these drilling cores was given to the Geological Institute of Hungary for geological, palaeontological, and stratigraphical analysis. The lithostratigraphy of polycyclic fluvial sediments is very important because of its applied potentials for water supply. The water yield capacity of coarser grained sediments is principally very good as opposed to clayey sediments. Using the lithology and hydrogeological features, the “lithostratigraphy” was created in the second part of the 19th century, according to which the Upper and Lower Pleistocene sequences were good aquifers, while the Middle Pleistocene ones have poor aquifer characteristics. The “Levantine stage” was also classified as a good aquifer. A thick series of sands and marls with lignites in southeastern Europe (Paludina Beds) with rich fauna of Viviparidae, Melanopsidae and Unionidae rests directly upon the Pontian formation. Because of their widespread distribution in the Levant, they were given the rank of a “Levantine stage” by F. von Hochstetter. According to recent developments and changes in stratigraphy, “Levantine stage” is not in official use anymore. This subdivision is suitable only for labeling the water supply potential of sequences. From a stratigraphical viewpoint it is not acceptable, because it is not connected and correlated with other stratigraphical proxies, especially those based on biostratigraphical subdivision. This subdivision cannot be connected with any European, or any other international stratigraphical standards (Krolopp, 1970). The first data on V. boeckhi shells came from the borehole material of artesian wells from Szentes (Hungary). This material was found by Zsigmondy and analysed by Halaváts in 1888. Halaváts found the V. boeckhi shells at the depth of 243 m. The age of this species, according to the state of knowledge of that time (based on lithology and the molluscan assemblage), was assigned to “Upper Levantine” Stage (Halaváts, 1888a, 1888b). Later, V. boeckhi was found in the material from numerous boreholes, e.g. in Szeged, Hód}Vásárhely (nowadays Hódmezo } vásárhely), Nagybecskerek Mezo (nowadays Zrenjanin in Serbia), Szarvas, Szabadka (nowadays Subotica in Serbia) from the Great Hungarian Plain (Alföld). The mentioned horizon was labelled “V. boeckhi Horizon of the Upper Levantine substage” (Halaváts, 1888a, 1888b, 1889a, 1889b, 1895, 1900, 1914a, 1914b). Since the end of the 19th century, V. boeckhi has been found at many locations in the Middle Danube Basin (Krolopp, 1970). 4.2. Sratigraphical position of the V. boeckhi Horizon in Hungary (after WWI) 4.2.1. Research status and stratigraphical viewpoints on the V. boeckhi Horizon in Hungary (before malacostratigraphical biozonation) The age of the V. boeckhi Horizon was defined as “Upper Levantine”; this statement was especially strongly supported by Sümeghy (1939, 1941, 1953). In the beginning of the 1950s, Sümeghy reported that there was no Early Pleistocene molluscan fauna in the area of the Great Hungarian Plain/Alföld, because the shell material which was earlier attributed to Early Pleistocene was actually older (Pliocene age) or younger (Late Pleistocene) (Sümeghy, 1953). Kretzoi (1954), in his subdivision based on the vertebrate biostratigraphy, placed the V. boeckhi Horizon in the Sicilian Stage of the Pleistocene. Bartha (1962), after the examination of the palaeontological material from the borehole of Makó, concluded that below the V. boeckhi Horizon were several hundred meters thick Pleistocene sequences. However, to assert that the V. boeckhi species is older

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than Pleistocene, he stated that in the Makó borehole the V. boeckhi specimens were inserted in the younger facies and the whole material is ex situ. Because of the use of old flushing drilling techniques, the possibility of mixing the material from different fossiliferous deposits cannot be excluded (Krolopp, 1970). The lower parts of the V. boeckhi Horizon have an age younger than the Paludina Beds. The vertebrate record from Öcsöd (Hungary) with Allophiomys pliocaenicus implies the Upper Villanyian stage. Krolopp (1970) in the conclusions on the age of the

and the upper Biharian to the youngest parts of Early Pleistocene and early Middle Pleistocene (e.g. Fejfar and Heinrich, 1990). In the V. boeckhi Horizon, palaeontological remains of Mimomys savini were found, which suggests that it cannot be younger than the upper Biharian stage (Krolopp, 1970, 2002b; Kretzoi and Krolopp, 1972). Among vertebrate fossil fauna, remains of small mammals (Pliomys episcopalis, Desmana thermalis, Talpa praeglacialis) were found at several sites and are also of great stratigraphic importance (Krolopp, 1970) (Table 1).

Table 1 The position of the Viviparus boeckhi Horizon after the analysis of the Great Hungarian Plain/Alföld in Hungary (after Kretzoi and Krolopp, 1972).

V. boeckhi Horizon based on the molluscs and ostracods suggests a Lower Biharian or Lower Pleistocene interglacial age. The upper parts of V. boeckhi Horizon belong to the Middle Pleistocene, the lower Biharian Stage (sensu Kretzoi, 1941, described in Jánossy,1986; Kretzoi, 1965,1969, 1983, 1985). In the context of the recent data, the lower Biharian corresponds to the Early Pleistocene,

However, the numerical ages of boundaries of the biozones and the age ranges of the biozones are still a matter of debate even in the case of smaller mammals, although it has been established that the species (M. savini) occurred in European faunas during the late early and the early Middle Pleistocene. The transition from populations of water voles with rooted molars (M. savini) to populations with more

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hypsodont, unrooted molars referred to the genus Arvicola, took place during the first half of the Middle Pleistocene (van Kolfschoten, 2007). After the revision of the molluscan collections of the Hungarian Geological Institute in Budapest and the campaign of the systematic geological survey of the Hungarian part of the Great Hungarian Plain, the Early Pleistocene age for numerous deposits (Fig. 3) was confirmed by the molluscan fauna and vertebrate palaeontological data. These datasets enabled establishment of the Quaternary malacostratigraphical subdivision in Hungary in the 1970s and 1980s. 4.2.2. Viviparus boeckhi “Oppel-zone” of Hungary and its subdivision Biostratigraphical units were separated on the basis of the development of Pleistocene malacofauna and the knowledge of the “stratotype-place locations of occurrence”. These stratigraphical units correspond to the fauna phases (“faunal waves”) of the vertebrate biostratigraphy, that is, practically, of biozones, namely Table 2 Quaternary molluscan biostratigraphy of Hungary (after Krolopp, 2004).

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of Oppel-zone value. Therefore the malacological phases are divided not on the basis of one or a few taxa, but rather after the characteristics of the whole faunal assemblages (Krolopp, 1995). The biostratigraphical subdivision based on Oppel-zones of Hungary was developed by Krolopp (1983, 1995), but there is some confusion. The attribution of Oppel-zones is based on the first appearance and the extinction of a species (V. boeckhi). In the description of the 1.b. N. crassitesta subzone, the lack of V. boeckhi is evident (e.g. Krolopp, 1983, p. 76). In the same paper, V. boeckhi was described as having a continuous record from Upper Villanyian to lower Biharian (Krolopp, 1983, p. 72). It seems that the appearance of V. boeckhi was more linked to regional climato-environmental changes, and the Oppel-zone biostratigraphy needs more supporting data. The biozones established on the basis of the development of the molluscan fauna in Pleistocene of Hungary (after Krolopp, 1983) are presented in Table 2 (with some timeframe changes). This paper will focus only on the description of the first biozone, the V. boeckhi Oppel-zone.

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Fig. 3. Lower and upper Middle Pleistocene sections of Hungary (after Krolopp, 1995).

4.2.2.1. V. boeckhi Oppel-zone. Type locality: Szabadhídvég (Fejér County, Hungary), Kavicshehegy (Krolopp, 1978b). The malacological assemblage of the biozone has the following features: representatives of Unionidae and Viviparidae with decorated shells, as well as the genera Melanopsis, Cochlostoma and Triptychia, characteristic for the final phase of Pliocene, are absent. The molluscan fauna is ecologically similar to the recent one in the same area. According to its origin, the fauna is divided into four groups. A significant part of fauna consists of species which lived in this region since the Neogene (41 species, i.e. 28% of total fauna). The species in the second group were not found in the older sediments but occur in the recent fauna (83 species, 58%). The most peculiar part of the fauna (12 species, 8%), however, is represented by species which are unknown from the Pliocene sediments, and are not found in younger biozones. These extinct species are exclusively fluviatile (V. boeckhi, Viviparus acerosus zsigmondyi, N. crassitesta, Bithynia sp., Prosostenia sp., F. pleistocaenica) or terrestrial (Gastrocopta serotina,

Gastrocopta moravica, Gastrocopta moravica oligodonta, Gastrocopta sacracoronae, P. kormosi, Spirodiscus sp.). The species from the fourth group are either extinct, but not confined to this biozone (P. clessini, Succinea schumacheri, Aegopis klemmi, Zonitoides sepultus) or can be considered extinct only in this faunal region (C. fluminalis, Melanoides tuberculata, Anisus strauchianus, Helicodiscus cf. singleyanus; i.e. 8 species, 6%; Krolopp, 1973) (Krolopp, 1983). The fluviatile facies of the sedimentary complex of the V. boeckhi Oppel-zone is best known. Its fauna is characterized, in addition to the high individual number of extinct species, with a great number of species which are present in the recent fauna: F. acicularis, F. esperi, Fagotia ex gr. aciculariseesperi and Theodoxus ex gr. prevostianuse danubialis (Krolopp, 1977). In its lacustrine fauna no characteristic features can be found. Its terrestrial deposits are characterized by the great number of Granaria frumentum, Chondrula tridens, Helicopsis striata, Cepaea vindobonensis, in addition to a restricted number of the extinct forms. According to faunal composition, the major part of sediments of this biozone was deposited during a period of mild, even warm climate (Krolopp, 1983). Within the V. boeckhi Oppel-zone three subzones were identified by Krolopp (1983): the G. serotina subzone, the N. crassitesta subzone and the G. sacracoronae subzone: 4.2.2.2. G. serotina subzone. It is characterized by deposits of fluvial facies with a great number of species. Among the extinct species, V. boeckhi, N. crassitesta, Prososthenia sp., P. clessini, Fagotia ex gr. aciculariseesperi and Theodoxus ex gr. prevostianus-danubialis occur with high percentages. In the aquatic fauna, the presence of the F. pleistocaenica, and in the terrestrial fauna the presence of G. serotina, G. moravica, G. moravica oligodonta, P. kormosi, Helicodonta cf. singleyanus, and Spirodiscus sp. are noteworthy. Among terrestrial forms, G. frumentum predominates in number of individuals (Krolopp, 1973, 1983, 1995). 4.2.2.3. N. crassitesta subzone. It has an impoverished fauna as compared to the previous subzone, which is reflected in smaller

Fig. 4. Location of the Serbian borehole profiles where Viviparus boeckhi was found.

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numbers both of species and individuals. The absence of V. boeckhi is especially apparent. According to the data, of the extinct species only N. crassitesta is found in great numbers. The lacustrine or terrestrial fauna is poorly known. The impoverishment of the fauna could be caused by a climate drier and cooler than in the previous phase, which is also reflected in decreased fluvial activity. This subzone can be best characterized by the absent forms (Krolopp, 1983, 1995). 4.2.2.4. G. sacracoronae subzone. In its fluvial facies, fauna with great number of species and individuals can be found. V. boeckhi and other extinct species of the G. serotina subzone are present in large numbers. In addition to extinct species (G. sacracoronae, A. klemmi, Z. sepultus) its terrestrial fauna is also characterized by en masse occurrence of G. frumentum and H. striata and the presence of Truncatellina claustralis, C. vindobonensis, Helix pomatia. The fauna from these locations indicates a mild and warm climate respectively (Krolopp, 1983, 1995). Although faunas of the G. sacracoronae and G. serotina subzones are rather similar, in the first subzone V. boeckhi usually appears in large numbers (the V. boeckhi level of the Great Hungarian Plain). From the Gastrocopta genus, only G. sacracoronae occurs, while

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G. serotina, G. moravica and G. moravica oligodonta are absent. Furthermore, the genera Ferrissia and Parmacella are missing while A. klemmi and Z. sepultus occur (Krolopp, 1983, 1995). Because of the lack of data and the traditional separation of land and freshwater fauna in different biozonation units, they cannot be included in the same unit/biozone. In spite of great similarities with the biozones in Hungary, this subdivision is not recommended for the Serbian Quaternary. However, the V. boeckhi biozone exists in fluvial record of Serbia and the data are comparable with the Hungarian record. In his latest works, Krolopp suggested an Early Pleistocene age for the V. boeckhi Horizon. In the list of Pleistocene molluscs species from Hungary (Krolopp, 2002a) V. boeckhi is confined to the Early Pleistocene. In his last subdivision, the V. boeckhi Oppel-zone has an Early Pleistocene age (Table 2) and V. boeckhi represents an index fossil for Early Pleistocene (Krolopp, 2004). 4.3. Position of V. boeckhi Horizon in Serbia and the region The mollusc species V. boeckhi was found in the core material in the Middle Danube Basin at more than 30 locations [the material is

Fig. 5. Profile of borehole Ada Ciganlija P-1 (after Nenadic et al., 2009).

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stored in several institutions e Geoinstitut (now Croatian Geological Survey, Zagreb, Croatia), Geozavod (now Geological Survey of Serbia, Beograd, Serbia) and Geological Institute of Hungary (Budapest, Hungary)]. Six of those locations are situated in the Republic of Serbia (Fig. 4.). The 77 species found in the association and deposits with V. boeckhi were of Quaternary age. The fauna is clearly connected with polycyclic fluvial freshwater environmental conditions (Krolopp, 1970; Kretzoi and Krolopp, 1972). The sites revision is mainly done in Hungary, so its results can only partly be applied to Serbian sections. The problem with applying Hungarian biostratigraphical division is that the V. boeckhi biozones and subzones cannot be clearly identified in Serbia. Laskarev (1938) determined the age of the assemblage of the V. boeckhi Horizon as the lowermost interglacial stage of the Middle Pleistocene. Later, with help of better datasets, Laskarev (1951) changed the position of the V. boeckhi Horizon and put it in the Lower Pleistocene (Markovic-Marjanovi c, 1977). The main sections were Zrenjanin and Ada Ciganlija (Fig. 5), where the V. boeckhi Horizon lies below the Pleistocene Corbicula Beds. The V. boeckhi Horizon could be identified with lower parts of the Pleistocene Corbicula Beds (the assemblages are overlapping) as in the borehole profile of Zrenjanin (Krolopp, 1977). After the extinction of some guide fossils (e.g. V. boeckhi, V. acerosus zsigmondyi, N. crassitesta), the molluscan assemblage of younger parts of the Pleistocene Corbicula Beds became poorer but still kept its warm stage(s) fluvial character. The assemblage of the V. boeckhi Horizon corresponds to the oldest assemblage of the Pleistocene Corbicula Beds which are easily separated from the other sediments. These two horizons are overlapping (only in the case of the oldest Pleistocene Corbicula Beds). The time range of the V. boeckhi Horizon should be from Lower Pleistocene to the lowermost parts of Middle Pleistocene. However, in Serbia its presence in the Middle Pleistocene is not confirmed and it is mainly held that its upper parts belong to the Lower Pleistocene, whereas the lower parts are the (Upper) Paludina Beds. 5. Conclusions The V. boeckhi Horizon was first described by Halaváts (1888a, 1888b) in the same paper in which he described a new species: Vivipara Böckhi (V. boeckhi). Later investigations of Krolopp (e.g. 1970) made a revision of the earlier data and put the new data in quantitative and stratigraphical context, and justified the use of term of V. boeckhi Horizon. In the geological formations of Serbia, the assemblages of the V. boeckhi Horizon were known from the Paludina Beds (“Levantine Deposits”) (e.g. Halaváts, 1895). Laskarev (1938) determined the age of the assemblage of the V. boeckhi Horizon as the lowermost interglacial stage of the Middle Pleistocene. Laskarev (1951) changed the position of the V. boeckhi Horizon and put it in the Lower Pleistocene. Kretzoi and Krolopp (1972) defined the V. boeckhi Horizon as the lower substage of the Lower Biharian, in the lowermost part of the Middle Pleistocene. The age of the V. boeckhi species (Krolopp, 1980) was defined as Lower Biharian (equivalent to the lower part of the “Cromerian complex”). Krolopp (1983, 1995) correlated the V. boeckhi biozone to the upper parts of the Villanyian and lower parts of the Biharian vertebrate stages (the time range of the lower parts of the Biharian corresponds to the upper parts of the Middle Pleistocene). The studies of the V. boeckhi assemblage support the investigations based on paleontological (small mammals, palynology, ostracods) and sedimentological investigations in the Great Hungarian Plain/Alföld (e.g. Krolopp, 1970; Kretzoi and Krolopp, 1972). The results from Hungary seem to be synchronous with

the results from Serbia and Croatia. Because of the insufficient data and traditional subdivision of Quaternary in Serbia, land and freshwater species (or different sedimentation domains) are generally not placed in the same biozone, but in separate biostratigraphical units. V. boeckhi Horizon represents the Pleistocene fluvial warm stage. The molluscan assemblage of the V. boeckhi Horizon identified it as a fluvial warm stage assemblage which represents the oldest Pleistocene Corbicula Beds. The age of the V. boeckhi Horizon is generally considered to correspond the younger parts of the Early Pleistocene, but according to the regional data the V. boeckhi Horizon of Serbia is younger than the Paludina Beds (of PlioceneEarly Lower Pleistocene age) and according to the vertebrate fauna it is older than the upper Biharian stage (younger part of the Middle Pleistocene). Acknowledgements The authors are grateful to Bálint Peterdi, the curator at the Museum of the Geological Institute of Hungary and to Ivan Stefanovi c, the curator of the Palaeontological collection at the University of Belgrade-Faculty of Mining and Geology for help and access to the collections. We received great support from Endre Kroloppy who, with his help and comments, had a significant impact in the early stage of the manuscript. The authors are also grateful to the reviewers (D. Esu and N. Limondin-Lozouet) for improving the quality of the manuscript. These investigations were supported by grants from Ministry of Education and Science, project No. 176015 and 177023. References Adda, K., 1899. Az újvidéki városi artézi kútról (About the artesian well in Novi Sad). Földtani Közlöny (Bulletin of the Hungarian Geological Society) XXIX, 13e15 (in Hungarian). } inek o } slénytani Bartha, F., 1962. A makói és gyulai vízkutató fúrások puhatestu vizsgálata (The palaeonological analysis of mollsucs from geohydrological drilling core material from Makó and Gyula). In: A Magyar Állami Földtani } l (Annual Report of the Geological Institute of Intézet Évi Jelentése 1959-ro Hungary of the year 1959), pp. 271e295 (in Hungarian). bij de Vaate, A., Greijdanus-Klaas, M., 1990. Asiatic clam, Corbicula fluminea (Müller, 1974) (Peleypoda, Corbiculidae), a new immigrant to the Netherlands. Bulletin Zoologisch Museum, Universiteit van Amsterdam 12 (12), 173e177. Fejfar, O., Heinrich, M., 1990. Muroid rodent biochronology of the Neogene and Quaternary in Europe. In: Lindsay (Ed.), Fahlbusch and Mein e European Neogene Mammal Chronology. Plenum Press, New York. Halaváts, GY., 1888a. A szentesi ártézi kút (The artesian well of Szentes). A Magyar Királyi Földtani Intézet Évkönyve (Annals of the Hungarian Geological Institute) 8, 203e222 (in Hungarian). Halaváts, J., 1888b. Der artesische Brunnen von Szentes. Mittelungen aus dem Jahrbuche der königlich ungarischen geologischen Anstalt 8 (3), 166e194. } -vásárhelyi két artézi kút (The artesian well of Halaváts, GY., 1889a. A hód-mezo }-Vásárhely). A Magyar Királyi Földtani Intézet Évkönyve (Annals of Hód-Mezo the Hungarian Geological Institute) 8, 203e222 (in Hungarian). }-Vásárhely. MittelunHalaváts, J., 1889b. Zwei artesischen Brunnen von Hód-Mezo gen aus dem Jahrbuche der königlich ungarischen geologischen Anstalt 8 (3), 214e231. Halaváts, GY., 1895. Az Alföld Duna Tisza közötti részének földtani viszonyai (The geology of the Great Hungarian Plain at the Danube-Tisza Interfluve). A Magyar Királyi Földtani Intézet Évkönyve (Annals of the Hungarian Geological Institute) XI (3), 104e173 (in Hungarian). Halaváts, GY., 1900. A szarvasi artézi kút (The artesian well of Szarvas). Magyar } léseinek Munkálatai 30, 585e Orvosok és Természetvizsgálók Vándorgyu 589 Halaváts, GY., 1914a. A nagybecskereki fúrólyuk (The borehole of Nagybecskerek). A Magyar Királyi Földtani Intézet Évkönyve (Annals of the Hungarian Geological Institute) 22, 171e202 (in Hungarian). Halaváts, GY., 1914b. Die Bohrung in Nagybecskerek. Mittelungen aus dem Jahrbuche der königlich ungarischen geologischen Anstalt 22 (3), 190e222. Haynes, A., Taylor, B.J.R., Varley, M.E., 1985. The influence of the mobility of Potamopyrgus jenkisi (Smith, E. A.) (Prosorbachia; Hydrobiidae) on its spread. Archiv für Hydrobiologie 103 (4), 497e508. Jánossy, D., 1986. Pleistocene Vertebrate Faunas of Hungary. In: Developments in Palaeontology and Stratigraphy, vol. 8. Elsevier, Amsterdam.

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