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We used a multidisciplinary approach to infer the taxonomy and historical biogeography of Hierophis viridiflavus and H. gemonensis, performing molecular analyses of itochondrial (16S, Cyt-b, ND4) and nuclear markers (PRLR), a landmark-based morphometric study and a cytogenetic analysis. Our data distinguished three main groups in the studied species, corresponding to H. gemonensis and to two monophyletic clades (E and W) within H. viridiflavus. Clades E and W display a significant genetic (about 4% for Cyt-b and ND4) and morphological divergence and a different morphology of the W sex chromosome (submetacentric in clade E and telocentric in clade W). Taking into account the existing divergence, these clades appear to represent independent phylogenetic units, deserving elevation to species status. Specific names should be H. viridiflavus (Lacepede, 1789) and H. carbonarius (Bonaparte 1833) for cladesWand E, respectively. The phylogeography of the studied species is only partially concordant with a general pattern of ‘southern richness and northern purity’ of genetic diversity, whereas H. gemonensis exhibits high genetic diversity at low latitudes (especially in the Peloponnese), H. carbonarius shows a number of different haplotypes both at low (along the Southern Italian Apennines and in Sicily) and high latitudes in Italy. Furthermore, a relaxed clock model hypothesizes the differentiation between H. gemonensis and H. viridiflavus sensu lato at about 7 Mya, in the Messinian. Subsequently, the speciation involving H. viridiflavus sensu stricto and H. carbonarius took place in the Quaternary, probably as a result of Pleistocene climatic oscillations. Furthermore, our results are consistent with the existence of several ‘refugia within refugia’ in Italy and in the Balkans and depict the major cladogenesis as allopatric events, mainly driven by paleoclimatic and geographical factors

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Sex chromosomes were studied in eight species of lacertid lizards using C-banding, G-banding and restriction enzyme treatment. All of the species showed female heterogamety. The W chromosome was a microchromosome in Lacerta graeca and Ophisops elegans. Two types of W were found in Lacerta vivipara; in specimens from The Netherlands it was metacentric, whereas in specimens from Russia it was acrocentric or subtelocentric. The W chromosome was homomorphic or nearly homomorphic but completely C-banded and heterochromatic in Lacerta agilis, Podarcis hispanica, Algyroides moreoticus and A. nigropunctatus. In was only possible to find sex chromosomes using the G-banding method in Podarcis sicula. The results obtained, together with data in the literature, suggest that sex chromosomes are likely to be present in all Lacertidae and that their differentiation took place repeatedly and independently in different taxa within the family. A model for sex chromosome evolution in the family, in which the starting point was the heterochromatization of the W chromosome, is proposed.

Lizards represent unique model organisms in the study of sex determination and sex chromosome evolution. Among tetrapods, they are characterized by an unparalleled diversity of sex determination systems, including temperature-dependent sex determination (TSD) and genetic sex determination (GSD) under either male or female heterogamety. Sex chromosome systems are also extremely variable in lizards. They include simple (XY and ZW) and multiple (X1X2Y and Z1Z2W) sex chromosome systems and encompass all the different hypothesized stages of diversification of heterogametic chromosomes, from homomorphic to heteromorphic and completely heterochromatic sex chromosomes. The co-occurrence of TSD, GSD and different sex chromosome systems also characterizes different lizard taxa, which represent ideal models to study the emergence and the evolutionary drivers of sex reversal and sex chromosome turnover. In this review, we present a synthesis of general genome and karyotype features of non-snakes squamates and discuss the main theories and evidences on the evolution and diversification of their different sex determination and sex chromosome systems. We here provide a systematic assessment of the available data on lizard sex chromosome systems and an overview of the main cytogenetic and molecular methods used for their identification, using a qualitative and quantitative approach.