On the Leptocytheridae Ostracods of the Long-Lived Lake Ohrid: A Reappraisal of their Taxonomic Assignment and Biogeographic Origin.

Leptocythere karamani Klie, one of few non-marine species of the family Leptocytheridae (Ostracoda), is redescribed from specimens recently collected from the long-lived Lake Ohrid on the Albanian-Macedonian border. Detailed morphologies of valves and limbs of this species were compared with those of other Ohrid-Prespa leptocytherids, of some recent marine representatives of the genera Leptocythere Sars and Callistocythere Ruggieri from the Mediterranean, Irish and Baltic seas as well as with that of fossil non-marine species from the Miocene palaeo-Lake Pannon belonging to the genera Amnicythere Devoto and Euxinocythere Stancheva. Comparison with other species of Leptocytheridae inhabiting fresh to brackish waters of the Black-Azov, Caspian and Aral seas were also carried out using descriptions provided in the literature. Based on the comparative morphological studies it is shown that L. karamani and other Ohrid leptocytherids have a number of characters distinguishing them from other members of the genus Leptocythere but demonstrating a relationship with species of the genus Amnicythere. The most reliable of these characters are: a) anterior valve vestibulum from where mostly uni-ramified pore canals start, b) the entomodont hinge type with a strong anterior anti-slip tooth, a smooth posterior anti-slip bar on the left valve, and c) the hemipenis with underdeveloped lateral lobe and reduced clasping organ. From this strong evidence, the Ohrid leptocytherid species are allocated to the genus Amnicythere. Finally, a biogeographic scenario on the origin of the Ohrid leptocytherids is proposed which matches the "Lake Pannon derivate hypothesis". Close relationship of the Ohrid Amnicythere species with the non-marine leptocytherid taxa from the Neogene lakes of Central and Eastern Europe and with extant taxa from the Black and Caspian seas may indicate that the Ohrid Amnicythere derived from Lake Pannon species which were able to colonise lakes in Southern Europe through a stepping-stone process and subsequently to adapt to freshwater environment.