Cryptic crested newt diversity at the Eurasian transition: the mitochondrial DNA phylogeography of near Eastern Triturus newts

Crested newts of the Triturus karelinii group occur in a phylogeographically understudied region: the Near East. Controversy surrounds the systematic position of these newts within the complete crested newt assemblage (the Triturus cristatus superspecies). We explore the situation using mitochondrial sequence data (ND2 and ND4, ≈1.7 kb) and employing different methods of phylogenetic inference (Bayesian inference and Maximum Likelihood using mixed models) and molecular dating (r8s and BEAST). The T. karelinii group is monophyletic and constitutes one of four main lineages in the T. cristatus superspecies. The separation of the T. karelinii group from the remaining crested newts around 9 Ma is related to the formation of the Mid-Aegean Trench, which separated the Balkan and Anatolian landmasses. The T. karelinii group comprises three geographically structured clades (eastern, central and western). The genetic divergence shown by these clades is comparable to that among recognized crested newt species. We suggest the uplift of the Armenian Plateau to be responsible for the separation of the eastern clade around 7 Ma, and the re-establishment of a marine connection between the Black Sea and the Mediterranean at the end of the Messinian Salinity Crisis to have caused the split between the central and western clade around 5.5 Ma. Genetic structuring within the three clades dates to the Quaternary Ice Age (<2.59 Ma) and is associated with alternating periods of isolation and reconnection caused by periodic changes in sea level and surface runoff.