Historical biogeography and systematics of yellow-bellied toads (Bombina variegata), with the description of a new subspecies from the Balkans by Christophe Dufresnes, Simeon Lukanov, Sven Gippner, Johanna Ambu, Ilias Strachinis, Dragan Arsovski, Benjamin Monod-Broca, Hugo Cayuela, Petros Lymberakis, Daniele Canestrelli, Dan Cogălniceanu, Nikolay A. Poyarkov, Spartak N. Litvinchuk, Tomasz Suchan, Mathieu Denoël, Daniel Jablonski

“…The analyses support four major lineages, one assigned to B. v. variegata (Carpathians and northwestern ranges), one assigned to B. v. pachypus (Apennine Peninsula), and two assigned to B. v. scabra (Dinarides, Hellenides and Balkanides vs. the Rhodope mountains). Spatiotemporal patterns of diversification suggest a role for a Late Miocene marine incursion in the Pannonian Plain (Paratethys) as the initial trigger of divergence, followed by a vicariance event in the Apennines and a “sky island” process of Pleistocene differentiation in the Balkan Peninsula. …”
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