Magyar, Imre

Magyar, I

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Palaeobiogeography of lacustrine cockles as a tool to reconstruct palaeogeography in the Late Neogene Paratethys

Palaeobiogeography of lacustrine cardiids (subfamily Lymnocardiinae) is an effective tool in the palaeogeographic reconstruction of the Paratethyan area throughout the Neogene. Increasing endemism reflects decreasing connectivity towards the sea (as in the case of the Sarmatian Sea). Extensive formation of endemic species flocks indicates long-lived lacustrine environment (e.g. Lake Pannon). Further taxonomic study is required to improve our palaeogeographic models for the one-way migration events, such as the early Pontian and end-Messinian dispersal of lacustrine cockles into the Eastern Paratethys and into the Mediterranean, respectivel

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IntroductionThe term “Paratethys” was coined by LASKAREV (1924) todesignate the restricted marine to lacustrine biogeographicprovince stretching from the North Alpine Foreland Basin toCentral Asia in the Early Oligocene to Recent. The Paratethysconsisted of a series of basins that have been separated fromthe Tethys (or later from the Mediterranean Sea) by the upliftof the Alpine-Caucasian mountain chain. Connections ofindividual basins with each other and with the ocean wererepeatedly confined or closed, triggering the evolution ofspecific, endemic taxa in molluscs, ostracods, fish, and othergroups of the biota. Borders between parts of the Paratethysand towards the Mediterranean were dynamic: Mediterra-nean waters occasionally flooded the Paratethys, or viceversa, Paratethyan waters drained into the Mediterraneanbasins. Thus the history of the Paratethyan basins was anintricate story of opening and closing connections, catas-trophic floods, and episodes of desiccation. A brief characterization of the Paratethys, similar to theabove paragraph, can be found in dozens if not hundreds ofpapers dealing with the geology of the region. What is thebasis of these statements though? How do we perceive theseisolation and connectivity events in the geological record?The question is relevant because in most cases there is nodirect geological evidence for the presence of gatewaysconnecting individual basins; most of these supposedgateways are located in subsequently elevated areas whereerosion destroyed the coeval sedimentary record.  Palaeobiogeography as a tool to reconstructisolation and connectivity eventsAlthough some recent studies experiment with theapplication of chemical and isotope signals to detect inflowof marine waters into isolated or semi-isolated Paratethyanbasins (VASILIEV et al. 2010, 2013), the most traditional and— so far — most effective principle to reconstruct thehistory of aquatic gateways in the Paratethys is palaeo-biogeography (HARZHAUSER & PILLER 2007). Large-scaleappearance of endemic forms within a given basin indicatesthat oceanic connections were confined and a restrictedmarine environment formed. Extinction of stenohalinemarine organisms in a basin is usually interpreted as a signof a change in water chemistry, a phenomenon caused byrestriction of oceanic connection. Appearance of aquaticspecies flocks (a monophyletic assemblage of species re-sulting from extraordinary endemic radiations; GREENWOOD(1984) indicates the formation of effective barriers to bioticdispersal and gene flow, i.e. isolation from the sea andformation of a lacustrine environment. Dispersal of origin-ally endemic species into another basin or appearance ofmarine species in a formerly lacustrine basin is usuallyPalaeobiogeography of lacustrine cockles as a tool to reconstruct palaeogeography in the Late Neogene ParatethysImre MAGYARMTA-MTM-ELTE Research Group for Paleontology, H–1431 Budapest, POB 137. (immagyar@mol.hu)AbstractPalaeobiogeography of lacustrine cardiids (subfamily Lymnocardiinae) is an effective tool in the palaeogeographicreconstruction of the Paratethyan area throughout the Neogene. Increasing  endemism reflects decreasing connectivitytowards the sea (as in the case of the Sarmatian Sea). Extensive formation of endemic species flocks indicates long-livedlacustrine environment (e.g. Lake Pannon). Further taxonomic study is required to improve our palaeogeographic modelsfor the one-way migration events, such as the early Pontian and end-Messinian dispersal of lacustrine cockles into theEastern Paratethys and into the Mediterranean, respectively.Keywords: palaeobiogeography, palaeogeography, Paratethys, Pannonian basin, Neogene, molluscs, CardiidaeHantkeniana 10, 143–146., Budapest, 2015Barnabás GÉCZY Jubilee Volumeinterpreted to have taken place through a newly formedgateway between the respective basins. The palaeogeographical interpretation of a palaeobio-geographic pattern, however, is not always straightforward.Dispersal across a (former) barrier may also happen bymeans of wind or migrating animals, such as fish or water-fowl (e.g. WESSELINGH et al. 1999). Even when a barrier isdestroyed due to geological processes, such as erosion,water level rise, or tectonic subsidence (commonly adressedas “geodispersal”), the newly established connection be-tween aquatic basins may be either a strait providing theopportunity for two-way migrations, or a river flowing fromone basin into the other and functioning as an effective filteror even barrier for upstream migration.In addition, chronology is a key issue in understandingpalaeobiogeographic changes. Non-marine basins oftenhave uncertain chronostratigraphy, and this may make therecognition of palaeobiogeographic events difficult or im-possible, leading to unfounded theories and hypotheses.   The bivalve subfamily LymnocardiinaeOne of the groups whose palaeobiogeography is veryuseful in palaeogeographic reconstructions comprises thenon-marine cardiids (subfamily Lymnocardiinae). Cardiidaeincludes mostly stenohaline, normal marine bivalves thatenter the fossil record in the Late Triassic and has an extantdiversity of about 200 species (SCHNEIDER 1995). Lacustrinecardiids (Lymnocardiinae) include only a few extant speciesthat live in the Caspian and Black seas and in the Aral Sea.The highly euryhaline marine genus Cerastoderma, which isalso known to occur in Saharan lakes hundreds of kilometresaway from the sea (ROSE 1972), is also assigned into thissubfamily. Cerastoderma first appeared in the Early Oligo-cene and is believed to be the source of all laterlymnocardiid radiations in the Paratethys. Therewere four radiatons, resulting in the appearanceof about 70 genera and ca. 700 species, allendemic to the Paratethys or any of its subbasins(NEVESSKAJA et al. 2001; MAGYAR 2006).Examples from the Late NeogeneSarmatianThe most spectacular series of lymnocar-diid radiations started in the Sarmatian (12.8 to11.6 Ma) from two Badenian species, Obso-letiformes lithopodolicus and Plicatiformespraeplicatus (STUDENCKA et al. 1998). TheSarmatian cardiid fauna, evolving from thesetwo species, includes 4 additional endemicgenera and more than 40 endemic species(PARAMONOVA 1977), indicating that theSarmatian Sea, stretching from the Viennabasin to Aral Sea, formed an independent biogeographicunit. It is the Proto-Caspian Subprovince of HARZHAUSER &PILLER (2007). Based on the high degree of endemism, onlyvery restricted geographical connectivity can be supposedbetween the Sarmatian Sea and the Mediterranean. PannonianThe lymnocardiid fauna of the Paratethys went extinct inthe early late Miocene; only a subset of the species took refugein the Pannonian basin. This event indicates the splitting of theonce uniform Sarmatian Sea province into two palaeo-geographical units: the Eastern Paratethys without cardiids(probably due to strongly reduced salinity), and the Pannonianbasin, where lymnocardiids started to radiate again, thusindicating a brackish environment (HARZHAUSER & PILLER2007). In the Pannonian basin, at least 15 genus-level taxaevolved, with more than 200 species (MÜLLER et al. 1999).This intense speciation and formation of species flocks is acharacteristic feature of long-lived (“ancient”) lakes, bothfossil and recent (BROOKS 1950), and thus indicates that thePannonian basin functioned as a lake at that time (Lake Pan-non, Balatonian biogeographic province, HARZHAUSER &PILLER, 2007).Within the Pannonian basin system, the lymnocardiid-bearing Upper Miocene – Lower Pliocene sedimentarylayers are fairly continuous today, but there are patches,physically not connected to the main rock body (Figure 1).Were these disjunct patches deposited in Lake Pannon aswell, and it is due to subsequent erosion that they are notcontinuous any more, or they represent the deposits of other,smaller lakes? The palaeobiogeography of cardiids cananswer this question. The Upper Miocene patch of thesouthern Transylvanian basin lies 20-100 km east from themain sedimentary rock body of Lake Pannon. Its cockle144 MAGYAR, I.: Palaeobiogeography of lacustrine cockles as a tool to reconstruct palaeogeography in the Late Neogene ParatethysFigure 1. Late Neogene lymnocardiid-bearing lacustrine sediments in the intra-Carpathian area.Based on their  mollusc fauna, the lighter gray patches in the Pannonian basin proper and in thesouthern part of the Transylvanian basin were deposited in Lake Pannon (Late Miocene to EarlyPliocene), and their disjunct distribution is due to subsequent erosion. The darker gray patches,however, represent the deposits of Lake Turiec (Late Miocene) and Lake Transylvania(Pliocene)fauna consists of the same taxa as as those living in thePannonian basin proper, indicating that the Upper Mioceneof the Transylvanian basin was deposited in Lake Pannon. Incontrast, the cardiids of the Turiec basin, lying some 50 kmdistance from the continuous Lake Pannon deposits, aredifferent from the Lake Pannon forms, and, together with theentire mollosc fauna, suggest independent origin in aseparate lake (NEUBAUER et al. 2015). Similarly, cardiidsfrom the Pliocene Brasov-Baraolt basins (JEKELIUS 1932)are different from those of Lake Pannon, indicating in-dependent origin in a separate lake (“Lake Transylvania”:HARZHAUSER & MANDIC 2008; Figure 1).    PontianAt the beginning of the Pontian age (6.1 Ma; VASILIEV etal. 2011), a uniform lymnocardiid fauna characterized theentire Paratethys. This palaeobiogeographic pattern led tothe interpretation of a large Pontian Paratethys, similar inextent to the Sarmatian Sea (NEVESSKAJA et al. 1987). Amore careful chronological analysis, however, revealed thatgenera and even species of this uniform Pontian lymno-cardiid fauna appeared in the Pannonian basin well beforethe beginning of the Pontian (Figure 2). The first represen-tatives of Euxinicardium, Pseudocatillus, and Paradacnaappeared in Lake Pannon before 9.5 Ma; Prosodacnomyaand Pontalmyra are also at least 8 Ma old in the Pannonianbasin (MAGYAR & SZTANÓ 2007; Figure 2). All these formssuddenly appeared in the Eastern Paratethys at thebeginning of the Pontian, and quickly dispersed from theDacian basin to the Caspian region (NEVESSKAJA 1990).Migration between Lake Pannon and the Eastern Paratethysat the beginning of the Pontian seems to have been uni-directional (west to east). The connection between LakePannon and the Eastern Paratethys functioned as a bio-geographic filter, facilitating migration in one direction andhindering it in the other. It could have been either a marine(or lacustrine) gateway (corridor) with unidirectional stream(POPOV et al. 2006), or, more probably, a river flowing fromthe Pannonian basin to the Eastern Paratethys (MÜLLER et al.1999).       MessinianIn the late Messinian, at 5.5 Ma, lymnocardiids suddenlyappeared all across the Mediterranean. In spite of their widegeographic range, the fossils of these animals are poorlyknown. Some of them appear to be identical with theParatethyan species even at species level (e.g. Paradacnaabichi, Euxinicardium subodessae), whereas others couldhave been the result of Mediterranean radiations (ESU2007). In the history of the Mediterranean this 0.2 millionyear long phase is called “lagomare” (ROVERI et al. 2014). Atthis time lymnocardiids reached their largest distribution:they thrived from the Iberian coast to Central Asia. Thenature of connection between the Paratethys and theMediterranean remains debated (KRIJGSMAN et al. 2010).The base levels of the two large basins were apparentlyindependent from the global sea level, thus a lacustrinegateway with Paratethyan outflow into the Mediterranean ora river connection are both conceivable.  AcknowledgementWith this brief essay I would like to pay a tribute ofrespect and recognition to dr. Barnabás GÉCZY, my belovedprofessor of palaeontology, for his 90th birthday. He mademe curious — a precious, lifelong gift for a student. This is MTA-MTM-ELTE Paleo contribution #214.Hantkeniana 10 (2015) 145Figure 2. The first appearance of some of the “Pontian” cockles in the stratigraphic column of Lake Pannon. These endemic generaoriginated in Lake Pannon, and dispersed into the Eastern Paratethys during the early Pontian migration event at 6.1 MaReferencesBROOKS, J. L. 1950: Speciation in ancient lakes. — Quarterly Review of Biology 25, 30–60 and 131–176.ESU, D. 2007: Latest Messinian “Lago-Mare” Lymnocardiinae from Italy: close relations with the Pontian fauna from the Dacic Basin.— Geobios 40, 291–302.GREENWOOD, P. H. 1984: What is a species flock? — In: ECHELLE, A. A. & KORNFIELD, I. (eds): Evolution of fish species flocks. 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Palaeobiogeography of lacustrine cockles as a tool to reconstruct palaeogeography in the Late Neogene Paratethys

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