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Life and depositional environments in the sublittoral zone of Lake Pannon, a large, brackish Paratethyan lake from the Late Miocene, were reconstructed from fossils and facies of the Szák Formation. This formation is exposed in several, roughly coeval (9.4-8.9 Ma) outcrops, located along strike of the paleo-shelf-break in northwestern Hungary. The silty argillaceous marl of the formation was deposited below storm wave base, at 20-30 to 80-90 m water depth. The abundance of benthic organisms indicates that the bottom water was usually well oxygenated. Interstitial dysoxia, however, may have occurred immediately below the sediment-water interface, as evidenced by occasional preservation of trace fossils such as Diplocraterion. The fauna comprised endemic mollusks, including brackish cockles of the subfamily Lymnocardiinae, dreissenid mussels (Congeria), and highly adapted, uniquely large-sized deep-water pulmonate snails (planorbids and lymnaeids). Ostracods were dominated by endemic species and, in some cases, endemic genera of candonids, leptocytherids, cypridids, and loxoconchids. Fish remnants include a sciaenid otolith and the oldest skeletal occurrence of Perca in Europe. The phytoplankton comprised exclusively endemic coccolithophorids, mostly endemic dinoflagellates (prevailingly Spiniferites), and cosmopolitan green algae. The Late Miocene fauna and flora of Lake Pannon were in many ways similar to the modern Caspian biota, and in particular cases can be regarded as its precursor.

I. 1999 04 15: The endemic molluscs of the Late Miocene Lake Pannon: their origin, evolution, and family-level taxonomy. Lethaia, Vol. 32, pp. 47-60. Oslo. ISSN 0024-1 164.Long-lived lakes are often sites of spectacular endemic radiations. During the Oligocene to recent history of the Paratethys, large, long-lived (more than a million years) lakes with endemic faunas formed three times, in three different basins: the first in the Pannonian basin, the second in the Euxinian (Black Sea) basin, and the third in the Caspian basin. Because the Euxinian lake inherited much of the fauna of Lake Pannon, the three lakes together hosted two endemic radiations of molluscs. The most long-lived lake in the region was Lake Pannon, which persisted approximately seven million years from the late Middle Miocene to the Early Pliocene. Lake Pannon was formed by isolation from the sea. Changes in hydrological regime and/or water chemistry in addition to the relative lowstand which accompanied (or caused) the isolation almost completely exterminated the restricted marine fauna of the basin. A few highly euryhaline and marginal marine cardiids, dreissenids, and hydrobiids survived this environmental change. As in other fossil and extant long-lived lakes, the originally low-diversity fauna radiated into a high number of related endemic species ('species flocks') and genera in the expanding and ecologically vacated lake. Many originally freshwater taxa (unionids, sphaeriids, viviparids, valvatids, melanopsids, lymnaeids, planorbids) entered the lake as well, and some of them also gave rise to endemic clades. Evolution in both relict and freshwater immigrant groups led to the appearance of highly unusual shell shapes. Many lineages exhibit gradual morphological changes over one to several million years. More than 900 endemic mollusc species have been described from Lake Pannon, although this number includes junior synonyms, invalid species names, and highly similar chronospecies. Applying a conservative taxonomy, all these species belong to four bivalve and eight gastropod families. The high degree of endemism, however, is reflected by proposals of some authors to establish as many as five new families based on Lake Pannon endemics.

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