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Phylogeography and cryptic variation within the Lacerta viridis complex (Lacertidae, Reptilia)


  • Manja U. Böhme,

  • Uwe Fritz,

  • Tatiana Kotenko,

  • Georg Džukić,

  • Katarina Ljubisavljević,

  • Nikolay Tzankov,

  • Thomas U. Berendonk

Manja U. Böhme, University of Leipzig, Biology II, Molecular Evolution and Animal Systematics, Talstrasse 33, 04103 Leipzig, Germany. E-mail:
Uwe Fritz, Museum of Zoology, Natural History State Collections Dresden, A.B. Meyer Building, 01109 Dresden, Germany. E-mail:
Tatiana Kotenko, Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Vul. B. Khmelnyts’kogo 15, 01601Kyiv, Ukraine. E-mail:
Georg Džukić and Katarina Ljubisavljević, Department of Evolutionary Biology, Institute of Biological Research ‘Sinisa Stankovic’, 29 Novembra 142, 11060 Belgrade, Serbia and Montenegro. E-mail:;
Nikolay Tzankov, National Museum of Natural History, Blvd Tzar Osvoboditel 1, 1000 Sofia, Bulgaria. E-mail:
Thomas U. Berendonk, University of Leipzig, Biology II, Molecular Evolution and Animal Systematics, Talstrasse 33, 04103 Leipzig, Germany. E-mail:


It is well known that the current genetic pattern of many European species has been highly influenced by climatic changes during the Pleistocene. While there are many well known vertebrate examples, knowledge about squamate reptiles is sparse. To obtain more data, a range-wide sampling of Lacerta viridis was conducted and phylogenetic relations within the L. viridis complex were analysed using an mtDNA fragment encompassing part of cytochrome b, the adjacent tRNA genes and the noncoding control region. Most genetic divergence was found in the south of the distribution range. The Carpathian Basin and the regions north of the Carpathians and Alps are inhabited by the same mitochondrial lineage, corresponding to Lacerta viridis viridis. Three distinct lineages occurred in the south-eastern Balkans — corresponding to L. v. viridis, L. v. meridionalis, L. v. guentherpetersi— as well as a fourth lineage for which no subspecies name is available. This distribution pattern suggests a rapid range expansion of L. v. viridis after the Holocene warming, leading to a colonization of the northern part of the species range. An unexpected finding was that a highly distinct genetic lineage occurs along the western Balkan coast. Phylogenetic analyses (Bayesian, maximum likelihood, maximum parsimony) suggested that this west Balkan lineage could represent the sister taxon of Lacerta bilineata. Due to the morphological similarity of taxa within the L. viridis complex this cryptic taxon was previously assigned to L. v. viridis. The distribution pattern of several parapatric, in part highly, distinct genetic lineages suggested the existence of several refuges in close proximity on the southern Balkans. Within L. bilineata sensu stricto a generally similar pattern emerged, with a high genetic diversity on the Apennine peninsula, arguing for two distinct refuges there, and a low genetic diversity in the northern part of the range. Close to the south-eastern Alps, three distinct lineages (L. b. bilineata, L. v. viridis, west Balkan taxon) occurred within close proximity. We suggest that the west Balkan lineage represents an early offshoot of L. bilineata that was isolated during a previous Pleistocene glacial from the more western L. bilineata populations, which survived in refuges on the Apennine peninsula.