Grass snakes are widely distributed across the Western Palearctic. Recent phylogeographic studies provided evidence that three distinct parapatric species exist. Two of these occur in Italy, Natrix helvetica and N. natrix, and a contact zone between both taxa has been suggested for north‐eastern Italy. Moreover, previous investigations revealed for the Italian Peninsula a complex phylogeographic structure. Using mtDNA sequences and microsatellite loci, we examined the situation for mainland Italy, Sicily, Sardinia, and Corsica. Our study confirmed the occurrence of N. natrix in north‐eastern Italy. Cline analyses revealed limited gene flow between N. helvetica and N. natrix across a narrow hybrid zone. Within N. helvetica, conflicting patterns of mitochondrial and nuclear genomic differentiation were revealed. Three nuclear genomic clusters were found; one of them corresponded to no fewer than five distinct and, in part, deeply divergent and ancient mitochondrial lineages from mainland Italy and Sicily. This cluster was paraphyletic with respect to the two remaining mitochondrial lineages, each of which matched with another nuclear genomic cluster (one from Corsica plus Sardinia and another one from western Europe north of the Alps). This unexpected pattern most likely results from mainly male‐mediated gene flow and female philopatry combined with population‐density‐dependent processes such as ‘high‐density blocking’. With respect to taxonomy, we propose to synonymize N. h. lanzai Kramer, 1970 with N. h. sicula (Cuvier, 1829), acknowledging their lacking nuclear genomic differentiation. The studied hybrid zone of N. h. helvetica and N. h. sicula in Italy is wide, with a smooth cline for nuclear markers, supporting their subspecies status. We found no evidence for the distinctiveness of the two subspecies from Corsica (N. h. corsa) and Sardinia (N. h. cetti), suggesting their synonymy, but refrain from taxonomic conclusions because of small sample sizes and the endangered status of the Sardinian taxon.

The distribution and hybridisation zone of the two grass snake species occurring in the German state of Baden-Württemberg are described, based on genetic data from maternally inherited mitochondrial DNA (mtDNA, up to 1983 bp) and biparentally inherited microsatellite DNA (13 loci). In agreement with previously published morphological evidence, the barred grass snake (Natrix helvetica) occurs in the Upper Rhine Valley and the Black Forest, while the common grass snake (N. natrix, ‘yellow lineage’) is distributed across the remaining, more eastern parts of Baden-Württemberg. Cline analyses across two transects running through the region of Karlsruhe and the Black Forest indicate that the hybrid zone is similarly narrow here as in the previously characterised stretch near Lake Constance. With respect to nuclear DNA, the Black Forest constitutes no impediment to gene flow in comparison with lowland regions (Karlsruhe, Lake Constance). However, on the eastern slope of the Black Forest, the abrupt replacement of mtDNA of N. helvetica by that of N. natrix indicates male-mediated gene flow and that the Black Forest represents a dispersal barrier for female grass snakes.

We examined the contact zone of two parapatric species of grass snake (Natrix astreptophora and Natrix helvetica) in southern France. To this end, we used comprehensive sampling, analysed mtDNA sequences and microsatellite loci, and built Species Distribution Models for current and past climatic conditions. The contact zone had established by the mid-Holocene during range expansions from glacial refuges in the Iberian Peninsula (N. astreptophora) and southern or western France (N. helvetica). The contact zone represents a narrow bimodal hybrid zone, with steep genetic transition from one taxon to the other and rare hybridization, supporting species status for N. astreptophora and N. helvetica. Our results suggest that the steepness of the clines is a more robust tool for species delimitation than cline width. In addition, we discovered in western France, beyond the hybrid zone, a remote population of N. helvetica with genetic signatures of hybridization with N. astreptophora, most likely the result of human-mediated long-distance dispersal. For N. helvetica, we identified a southern and a northern population cluster, connected by broad-scale gene flow in a unimodal hybrid zone running across France. This pattern either reflects genetic divergence caused by allopatry in two microrefuges and subsequent secondary contact or introgression of foreign alleles into the southern cluster.

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