Vipera graeca Nilson & Andrén, 1988,

Mizsei, Edvárd, Jablonski, Daniel, Roussos, Stephanos A., Dimaki, Maria, Ioannidis, Yannis, Nilson, Göran & Nagy, Zoltán T., 2017, Nuclear markers support the mitochondrial phylogeny of Vipera ursinii – renardi complex (Squamata: Viperidae) and species status for the Greek meadow viper, Zootaxa 4227 (1), pp. 75-88 : 84

Vipera graeca Nilson & Andrén, 1988 stat. nov.

Greek meadow viper

Vipera ursinii graeca Nilson & Andrén, 1988 Vipera macrops graeca Welch, 1994: 123

Holotype. Göteborg Natural History Museum, GNM Re. ex. 4942. Leg. Nilson & Andrén 1988.

Paratypes. GNM Re. ex. 6823 (six newborn), GNM Re. ex. 6849 (ZIG 146), GNM Re. ex. 6850 (ZIG 147), GNM Re. ex. 6851 (ZIG 142) + GNM ZIG 145. Leg. Nilson & Andrén 1988.

Terra typica. Peristeri , Lakmos Mountains in the central Pindos mountain range, 1900 m altitude, Greece ( Nilson & Andrén 1988).

Morphological Diagnosis. This taxon differs from all other members of V. ursinii – renardi complex by having the following combination of morphological characters ( Nilson & Andrén 1988; Nilson & Andrén 2001; Mizsei et al. 2016): small body size (for males a snout to vent length (SVL) max. 40.6 cm, and tail length is 5.4 cm, and for females a SVL max. 44.3 cm, and tail length is 4.1 cm); non-bilineate body ground colour pattern; white or bright brownish-grey ventral colour; no dark spots on labial, lateral and dorsal sides of head except occipital and postorbital stripes; dorsal zigzag pattern tagged with pointed corners at windings, or consisting of a narrow vertebral line only; 45–58 dorsal windings; nasal divided into two plates or united with nasorostralia; rostral as high as broad; 2–8 loreals; 13–20 circumoculars; upper preocular not separated from nasal; 7–20 crown scales; more fragmented parietals; 12–15 supralabials (sum of right and left sides); first three supralabials two times larger than the following ones; third supralabial below orbit; 14–19 sublabials (sum of right and left sides); 3–5 mental scales; early dorsal scale row reduction; 120–129 ventrals for males, 119–133 ventrals for females; lowest number of subcaudals in the complex: 21–29 subcaudals for males, 13–26 subcaudals for females.

Molecular Diagnosis. The works of Ferchaud et al. (2012) and Zinenko et al. (2015) showed divergence of Vipera graeca stat. nov. based on mtDNA datasets. Its distinction is supported by phylogenetic position (basal taxon for all other species of the ursinii-renardi complex), time of divergence (the Middle Pliocene) and value of uncorrected pairwise p–distances (4.5% in view of ursinii clade) as is defined by Ferchaud et al. (2012). All our analyses based both on mitochondrial and nuclear loci support these results. Specimens in the study of Ferchaud et al. (2012) originated from Stavros area in the Vardoussia Mts., Greece), but in the present study we used samples from Albania . The specimens used here share the same CYT B and ND4 haplotypes as the previously analysed Greek samples of Ferchaud et al. (2012). A single ND4 haplotype is presented by the southernmost ( Stavros area , Vardoussia Mts. , Greece) and the northernmost populations ( Tomorr Mts. , Albania; Mizsei et al. 2016). Therefore , a single haplotype is very likely to be present throughout most of the distribution area of this species. Because the sequence variability of the nDNA regions is, in general, much less variable than mtDNA ( Townsend et al. 2008), we could use specimens originated from another locality than the type locality as they represent the same phylogenetic pattern and position.

Distribution. The species occurs in the subalpine meadows of the Hellenides mountain system of southern Albania and central Greece ( Dimitropoulos 1985; Nilson & Andrén 1988; Nilson & Andrén 2001; Korsós et al. 2008; Mizsei et al. 2016). These localities are Koziakas, Lakmos (Peristeri; type locality), Metsovon, Oiti, Tsouka Karali, Tzoumerka (Athamanika), Tymfristos, Vardoussia (including Stavros area) mountains in Greece, and Dhëmbel, Llofiz, Lunxhërisë, Griba, Nemerçkë (crossborder mountain, called Nemertzika in Greek), Shëndelli, Tomorr and Trebeshinë mountains in Albania. The entire distribution is extremely fragmented and each mountain population is completely isolated by a large matrix of unsuitable habitat for the taxon consisting of deep valleys and plains.

Ecology and habitats. A mosaic of open or closed grass and shrub communities formed on limestone characterizes the main habitats of the taxon. Annual mean temperatures are about ~6°C, and the meadows are partially covered by snow until early summer (May-June. South-facing slopes are usually more open and rocky than north-facing slopes. Different species of Festuca , Poa and Sesleria dominate the open grasslands, and characteristic shrubs are Juniperus sabina , Daphne oleoides and Astragalus creticus . Most of the observed vipers were found close to shrubs or stone piles in south-facing habitat patches. The diet of the species consists mainly of Orthoptera (97%) species, of which Stenobothrus rubicundulus , Platycleis sp., Decticus verrucivorus is the most frequent prey (Mizsei et al. in prep.). The abundance of Orthopterans is high from June to September (Lemonnier- Darcemont et al. 2015.). Known predators of the species are Vulpes vulpes, Falco tinnunculus and Circaetus gallicus .